    [From the U.S. Government Printing Office, www.gpo.gov]

                Distribution and Abundance of Fishes and
                  Invertebrates in West Coast Estuaries
                Volume II: Species Life History Summaries

                                � o
    0

                                                      COASTAL ZONE
                                                   INFORMATION CENTER
             QL                   August 1991
            139
            .E4           U.S. Department of Commerce
            no.8     National Oceanic and Atmospheric Administration
                              National Ocean Service

    0

                 Distribution and Abundance of Fishes and
                    Invertebrates in West Coast Estuaries
                 Volume Ih Species Life History Summaries

                                        Project Team

                           Robert L. Emmett* and Susan A. Hinton
                              Point Adams Biological Field Station
                          Coastal Zone and Estuarine Studies Division
                                  Northwest Fisheries Center
                               National Marine Fisheries Service
                                    Hammond, OR 97121

                            Steven L. Stone* and Mark E. Monaco
                        Strategic Environmental Assessments Division**
                   Office of Ocean Resources Conservation and Assessment
                                    National Ocean Service
                                     Rockville, MD 20852

                                   ELMR Report Number 8

                                         August 1991

                                            ~,o,'/,~;:�    {J. S. DEPARTMENT OF COMMERCE NOA.
                                                           COASTAL SERVICES CENTER
                                                           2234 SOUTH HOBSON AVENUE
                                                           CHARLESTON , SC  29405-2413

                                               Property of CSC Library

    * Contact for copies of this report.
    ** Formerly Strategic Asessment Branch

    This report should be cited as:
    Emmett, R. L., S. L. Stone, S. A. Hinton, and M. E. Monaco. 1991. Distribution and abundance of fishes and
    invertebrates in west coast estuaries, Volume I1: species life history summaries. ELMR Rep. No. 8. NOAAINOS
    Strategic Environmental Asessments Division, Rockville, MD, 329 p.

    Data Collection and Organization                                                                     ....................................................2

    Data Content and Quality                                                                             ...........................................................5
    Analysis of Data Content and Quality                                                                  ...................................................6

    Concluding Comments                                           ..............................................................10

    Index to Species Life History Summaries...............................................13

    Figure 1: Location of ELM R regions                                                                  ....................................................1
    Figure 2: Location of the 32 west coast estuaries included in the ELMR program,
              and their salinity zones as identified by the National Estuarine Inventory.................2
    Figure 3: Example of a species/estuary data sheet: threespine stickleback in
              Central San Francisco, San Pablo, and Suisun Bays, California........................4
    Figure 4: Mean data reliability of fish and invertebrate data collected for 32 west coast estuaries .....6
    Figure 5: Mean data reliability of species data collected for 32 west coast estuaries................7
    Figure 6: Life history table headings used to develop the information in Appendix 5...............    9

    Table 1: ELM R species list                                                                           ...........................................................3
    Table 2: Occurrence of 47 species in 32 west coast estuaries................................          5
    Table 3: Format of species life history summaries..........................................8

    Appendix 1: Summary table example: Spatial distribution and relative abundance.................288
    Appendix 2: Summary table example: Temporal distribution..................................289
    Appendix 3: Summary table example: Data reliability .......................................290
    Appendix 4: Presence/absence of 47 species in west coast estuaries ..........................291
    Appendix 5: Life history characteristics of 47 species in west coast estuaries.....................305
      Table 5A. Biogeography ............................................................306
      Table 56. Habitat associations .......................................................312
      Table SC. Biological attributes and economic value .......................................318
      TalTabRerdutol...............................................................3......42
    Appendix 6: Terms used in life history tables ..............................................327

      Distribution and Abundance of Fishes and Invertebrates in West Coast Estuaries
                             Volume II: Species Life History Summaries

             ,_________________________________________ .estuary for three salinity zones (seawater, mixing, and
                                                        tidal fresh zones) identified in NOAA's National
    This is the second of two volumes that present    Estuarine nventory(NEI)DataAtlas-Volumel(NOAA
    information on the spatial and temporal distribution,   1985).  When completed, the entire data base will
    relative abundance, and life history characteristics of   contain information for approximately 150 fish and
    47 fish and invertebrate species in 32 estuaries along    invertebrate species found in approximately 120 U.S.
    the contiguous west coast of the U.S.  Information   estuaries.
    presented in this volume focuses on species life history
    summaries which were written to identify the critical life
    history characteristics that help define a species'
    occurrence in estuaries.  These summaries were    Estuaries are among the most productive natural
    developedto complement datapresented in Distribution   systems and are important nursery areas that provide
    and abundance of fishes and invertebrates in west   food, refuge from predation, and valuable habitat for
    coastestuaries, Volume: Datasummaries(Monacoet    many species (Gunter 1967, Joseph 1973, Weinstein
    al. 1990), hereafter referred to as Volume I. The life   1979, Mann 1982). Estuarine organisms that support
    history summaries are not a complete treatise on each    importantcommercial and recreational fisheries include
    species; however, they provide a concise account of   salmonids, crabs, and shrimp.  In spite of the well-
    the most important physical and biological factors   documented importance of estuaries to fishes and
    known to influence a species' occurrence.             invertebrates, few consistent and comprehensive data
                                                        bases exist which allow examinations of the
    This report is a product of the National Oceanic and    relationships between estuarine species found in or
    AtmosphericAdministration's (NOAA) Estuarine Living   among groups of estuaries. Furthermore, much of the
    MarineResources(ELMR) program (inside back cover),   distribution and abundance information for estuarine-
    a joint study by the National Ocean Service and the   dependent species (i.e., species that require estuaries
    National Marine Fisheries Service (NMFS).  The    during their life cycle) is for offshore life stages and
    objective of the ELMR program isto develop aconsistent    does not adequately describe estuarine distributions
    data base on the distribution, abundance, and life   (Darnell et al. 1983, NOAA 1988).
    history characteristics of important fishes and
    invertebrates in the Nation's estuaries. The nationwide    Only a few comprehensive sampling programs collect
    data base is divided into four study regions (Figure 1).   fishes and invertebrates with identical methods across
    This data base contains the relative abundance and    groups of estuaries within a region ( Hammerschmidt
    monthly occurrence of each species' life stage by    and McEachron 1986).  Therefore, most existing

           Pt. Adams,
          OR Lab,
      West Coast
                                        32 estXaries, / I  2034 estuaries,
                  ~~~~~~~~~~32 estuaries,
       47 species                                                                                60 species

                                                                                         Marine Sci.
                                                                                        Beaufort,
                                                                                       NC Lab

                                                Galveston,                         Southeast
                                                TX Lab~/  ~  i   i*~Z~Sd         \  20 estuaries,

                                                           31 estuaries,
    Figure 1. Location of ELMR regions.                      44 species

    estuarine fisheries data cannot be compared among    (NOAA 1985), identify information gaps, and assess
    estuaries because of the variable sampling strategies.    the content and quality of existing estuarine fisheries
    In addition, existing research programs do not focus on    data.
    how groups of estuaries may be important for regional
    fishery management, and few compile information for                 _       �
    species having little or no economic value.
                                                           Volume Icontains detailed distribution and abundance
    Becauselifestagesofmanyspeciesusebothestuarine    data for 47 fish and invertebrate species in 32 west
    and marine habitats, information on distribution,    coast estuaries, and a complete discussion of the
    abundance, temporal utilization, and life history    methods used to compile these data. However, a brief
    characteristics are needed to understand the coupling    description of methods from Volume I are presented
    of estuarine, nearshore, and offshore areas. To date,    here to aid interpretation of distribution and relative
    a national, comprehensive, and consistent data base    abundance tables included in the species life history
    of this type does not exist. Consequently, there is a    summaries presented in this report.  The following
    need todevelop a program which integrates fragments    sections provide an overview of the estuary/species
    of information on marine and estuarine species and    selection process, and development of the ELM R data
    theirassociated habitats into a useful, comprehensive,    base.
    and consistent format. The ELMR program was
    designed to help fulfill this need by developing a    SelectionofEstuaries. Nineteenestuariesandmarine
    uniform nationwide data base on selected estuarine    embayments of the west coast (Figure 2) were initially
    species.  Results will complement NOAA efforts to   selected from the National Estuarine Inventory Data
    develop a national estuarine assessment capability    Atlas: Volumel (NOAA 1985). However, 13 additional

                           sl-iM Puget Sound
                           s M r Hood Canal
                            TSiMT Skagit Bay
                           IS IM T Grays Harbor
                           sM'TI Willapa Bay                       Washington
                           IsMTi- Columbia River
                           rSMl TNehalem Bay
                          rS MTi Tillamook Bay
                           S Ml T Netarts Bay
                          S'M- T Siletz River                    Oregon
                           S MT1 Yaquina Bay
                          ITTMTI1 Alsea River
                             M T Siuslaw River
                           S MTl Umpqua River
                           SMTT Coos Bay
                           Is M IT Rogue River  Salinity Zones
                           I s I MI T IKlamath River  F/S  |               ] Seawater zone (>25X%)
                                  i s i Mi T i Humboldt Bay               E Mixing zone (0.5-25%,)
                                  HumboEel     Ba mr'Tidal fresh zone (0-0.5%,)
                           SI'sI M T TaIoEemales Bay                     [] Zone not present'
                                  ~i~ si M i XiTormales Bay  .'Freshwater inflow is relatively low
                           SrSMIj TCentral San Francisco I                   in many southern California
                                    Suisun / San Pablo Bays                  estuaries/embayments.
                           sIMI I South San Francisco Bay
                          sIE-  Elkhorn Slough             7  California
                          IrtEII Morro Bay
                          Is II ISanta Monica Bay
                          IED -IISan Pedro Bay
                          Is   I | Alamitos Bay
                          ZIsl   Anaheim Bay
                          lIEm  INewport Bay
                           Iso fl Mission Bay
                          ri1fl  San Diego Bay
                          Irn  Tijuana Estuary

    Figure 2. Location of the 32 west coast estuaries included in the ELMR program, and their salinity zones as
              identified by the National Estuarine Inventory (NOAA 1985).

                                                        2

    west coast estuaries were added tothe NEI (and ELM R      Table 1. ELMR species list.
    program) due to their importance as habitat for west
    coast fishes and invertebrates. Data on the spatial and
    temporal distributions of species were compiled and       Scientific Name            Common Name
    organized based on three salinity zones delineated for
    each estuary in the NEI; tidal fresh (0.0 to 0.5�/),      Mytilis edulis               blue mussel
    mixing (0.5 to 25.0%o), and seawater (>25.0%o). While     Crassostrea gigas            Pacific oyster
    some west coast estuaries do not contain all three        Tresus capax                 horseneck gaper1
    salinity zones (e.g., southern California embayments),    Tresus nuttali               California ackknife clam 2
                         theywereidbecause they provide important  Tagelus californianus   California jackknife clam 2
    they were included because they provide important         Protothaca staminea          Pacific littleneck clam
    habitat for many euryhaline species.                      Venerupis japonica           Manila clam 3
                                                                Mya arenaria                softshell
    Selection of Species. To ensure that important west       Panopea abrupta             geoduck 4
    coast estuarine species were included in the ELMR         Crangon franciscorum         bay shrimp 5
    study, a species list was developed and reviewed by       Cancer magister              Dungeness crab
    regional experts (Table 1). Four criteria were used to    Triakis semifasciata         leopard shark
    identify the 47 species entered into the data base:       Acipenser medirostris       green sturgeon
                                                                Acipenser transmontanus      white sturgeon
                                                                Alosa sapidissima            American shad
    1) Commer                  spcial value - a species that commercia Clupea pallasi      Pacific herring
    fishermen specifically try to catch (e.g., Pacific herring  Anchoa compressa           deepbody anchovy
    and Dungeness crab), as determined from catch and         Anchoa delicatissima         slough anchovy
    value statistics of the NMFS and state agencies.          Engraulis mordax             northern anchovy
                                                                Oncorhynchus clarki         cutthroat trout
    2) Recreational value - a species that recreational       Oncorhynchus gorbuscha       pink salmon
    fishermen specifically try to catch that may or may not   Oncorhynchus keta            chum salmon
    be of commercial importance.  Recreational species        Oncorhynchus kisutch         coho salmon
    (e.g., steelhead and California halibut) were determined  Oncorhynchus mykiss          steelhead 6 (3 races)
    by consulting regional experts and NMFS reports.          Oncorhynchus nerka           sockeye salmon
                                                                Oncorhynchus tshawytscha   chinook salmon (5 races)
                                                                Hypomesus pretiosus         surf smelt
    3) Indicator species of environmental stress - identified  Spirinchus thaleichthys     longfin smelt
    from the literature, discussions with fisheries experts,  Thaleichthys pacificus       eulachon
    and from monitoring programs such as NOAA's National      Microgadus proximus          Pacific tomcod
    Status and Trends Program (NOAA 1984). These              Atherinops affinis           topsmelt
    species (e.g., Pacific oyster and white croaker) are      Atherinopsis californiensis    jacksmelt
    molluscs or bottom fishes that consume benthic            Gasterosteus aculeatus       threespinestickleback
    invertebrates or have a strong association with bottom    Morone saxatilis             striped bass
                                                                Paralabrax clathratus        kelp bass
    sediments. Their physiological disorders, morphological   Paralabrax nebulifer         barred sand bass
    abnormalities,  and  ability to bioaccumulate             Genyonemuslineatus           whitecroaker
    contaminants indicate environmental pollution or stress.  Atractoscion nobilis         white seabass
                                                                Cymatogaster aggregata       shiner perch
    4) Ecological value - based on several species attributes,  Ammodytes hexapterus       Pacific sand lance
    including trophic level, relative abundance, and          Clevelandia ios              arrow goby
    importance of species as a key predator or prey           Ophiodon elongatus           lingcod
    organism (e.g., bay shrimp and topsmelt).                 Leptocottus armatus          Pacific staghorn sculpin
                                                                Paralichthys californicus    California halibut
    Data Sheets. A data sheet was developed for each          Hypsops etta guttulata       diamond turbot
                                                                Pleuronectes vetulus         English sole
    species in each estuary to enable quick compilation       Platichthys stellatus        starry flounder
    and data presentation. For example, Figure 3 shows
    the data sheet forthreespine stickleback in central San    1 Also known as fat gaper (Turgeon et al. 1988).
    Francisco, San Pablo, and Suisun bays. Data sheets         2Alsoknown as California tagelus (Turgeon et al. 1988).
    were developed by project staff and reviewed by local      3 so known as Japanese littleneck, Tapes phillippinarum (urgeon
                                                                  etal. 1988).
    experts. Data compiled for each species' life stage        4 Also known as Pacific geoduck (Turgeon et al. 1988).
    included: 1 ) the salinity zones it occupies, 2) its monthly  5 Also known as California bay shrimp (Williams et al. 1989).
    occurrence in the zones, and 3) its relative abundance     6 The name steelhead refers to sea-run rainbow trout (Robins et al. 1980).
    in the zones.

                                                             3

      Threespine sticikeback     Central San Francisco/San  and spawning adults as those releasing eggs or sperm.
      Gasterosteusaculeatus      Pablo/Suisun Bays          A few exceptions existed to these defined life stages,
                                  StReviewer ,. Aorn        such as mating of Dungeness crab, and parturition (live
                                                           birth) of the viviparous leopard shark and shiner perch.
        Salinity    Lif    Relatie  Abundance by Month      In addition, the following unique life history information
                         Zone   ~~Adults    vl .S. _  N    is provided to interpret the data: 1) for the Pacific
      Tidal Fresh  Spawdng  . :2                           oyster, spawning adults, larvae, and eggs are not
       C o - 0      ::.oko.5 r le- '.. ' Io".
                Larvae                 I                   shown because spawning is sporadic (most spat is
                Eggs, . o   in                      1      hatchery produced and placed on beds), 2) forthe pink,
        Mixing   Spawning    i>.jiiiii  l            3      chum, coho, and chinook salmon, the onset of sexual
      0.5-25.07  Juveniles      i                  __ i    maturation (accompanied by morphological changes,
                Eggs        Il1:ii::,                       homing behavior, and a reduction in feeding/growth)
           Seawater    nivng                       u3 wsed to define the beginning of the adult life stage,
       Seawater  Spawing3
       >25.0,  Juveniles II:i:       ::             3      and 3) because migrating juveniles of different races of
                Larvae                               3
                Eggs                                        chinook salmon are difficult to separate in the field, the
                                                           data for juveniles of the different races of chinook
      Legend:  Relative Abundance  Data Reliability (R)     salmon include all races. However, yearling juveniles
            EII-'- Not Present    1 - Highly Certain       (spring and winter races) usually migrate to the ocean
                  -No Data .2 - Moderately Certain         earlier than subyearling juveniles (fall race).
               - _Rare
             : .- Common          3 = Reasonable Inference
                  _- .Abundant                             For well-studied species such as salmon, quantitative
                  -Highly Abundant                         data were used to estimate abundance levels. For
                                                           many species, however, reliable quantitative data were
    Figure 3. Example of a species/estuary data sheet:     limited. Therefore, regional and local experts were
             threespine stickleback in Central San         consulted to estimate relative abundances based on
             Francisco, San Pablo, and Suisun Bays.        theabovecriteria. Several referenceor"guide"species
                                                           with abundance levels corresponding to the above
    The relative abundance of a species was defined using    criteria were identified for each estuary. These guide
    one of the following categories:                        species typified fishes and invertebrates belonging to
                                                           a particular life mode (e.g., pelagic, demersal) or
       * Highly abundant - species is numerically           occupying similar habitats. Once guide species were
        dominant relative to other species.                selected, other species were then placed into the
                                                           appropriate abundance categories relative to them.
       * Abundant - species is often encountered in         These data represent relative abundance levels within
        substantial numbers relative to other species.     aspecificestuaryonly; relativeabundancelevelsacross
                                                           west coast estuaries could not be determined.
       � Common - species is generally encountered but
        not in large numbers; does not imply an even       Information in Volumelwas compiled for each species
        distribution over a specific salinity zone.        and estuary combination, and organized into four data
                                                           summaries:
       � Rare- species is present but not frequently
        encountered.                                          * Spatial distribution and relative abundance
                                                              � Temporal distribution
       � Notpresent- species or life stage not found,          � Data reliability
        questionable data as to identification of the         * Presence/absence data
        species, or recent loss of habitat or environmental
        degradation suggests absence.                      When compiled in this manner, the data can be easily
                                                           translated into various tables, such as the overall
       * Noinformationavailable- nodata available, and      occurrence of ELMR west coast species depicted in
        after expert review it was determined that even    Table 2. Appendix tables 1-3 are examples of how the
        an educated guess would not be appropriate.        data were summarized and presented in Volume I.
                                                           Due to post-publication revisions of the presence/
    Information was compiled for each of five life stages.    absence information in Volume I, Appendix table 4
    Adults were defined as sexually mature individuals,    provides the revised west coast ELMR presence/
    juveniles as immature but otherwise similar to adults,    absence data.

                                                         4

    Data Verification. Approximately three years were
    required to develop the 1,760 data sheets and consult
    with regional and local experts. Each data sheet was    An important aspect of the ELMR program, especially
    carefully reviewed during consultations or by mail.    since it was based primarily on published and
    These important consultations complemented the    unpublished literature and consultations, was to
    published and unpublished literature and data sets    determine the quality of the data used.  For many
    compiled by NOAA.   Ninety-one scientists at 26    species, gear selectivity, difficulty in identifying larval
    institutions or agencies were consulted. Local experts    stages to species, and difficulty of sampling various
    were particularly helpful in providing estuary/species-    habitats has limited the amount of reliable information.
    specific information.  They also provided additional    Therefore, a deliberate effort was made to assess the
    references and contacts and identified additional    overall reliability of the data base so it could be used
    species to be included in the ELMR data base.            appropriately. Estimates of the reliability of distribution

    Table 2. Occurrence (a) of 47 species (adults or juveniles rated as "common" to "highly abundant") in 32 west
            coast estuaries.
                                                              Estuary

          Species                                                                             r   c
       blue mussel     � � � � � � � � �    �    � �  � � � � � � � � � � � � � � �
       Pacific oyster                    **                            *
       horseneck gaper  0 � � � �        � �    � � �    �
       Pacific gaper    0 0         0
       California jackknife clam
       Pacific littleneckclam   0    �    � �                 � �  � � �     � @                         0
       Manila clam     0 0 0�      �        �    �  �
       Eastem softshell clam  *� � � � � � � � �

       bay shrimp      * * * * * * * * *  *� � � � � 0                  � � � �
       Dungenesscrab   000���� 0 000
       leopard shark                                                *   * *
       green sturgeon          �      � �        * �
       white sturgeon         *� * *� **
       American shad        � � � �              * * - - -
       Pacific herring      00 0 0 0 *                                   0 *    *
       deepbody anchovy   *
       slbugh anchovy                                                                       *    * -
       northern anchovy  0 0 0 0 0000000 * * * * *
       cutthroattrout  � � � ������* *
       pink salmon        *
       chum salmon     ��*   � ****0�   �0
       cohosalmon      ��*       ****                * *� �*
       steelhead       * -0                            0- -0-
       sockeye salmon    �0
       chinooksalmon   0 0 0�         �        �0 0      0    0
       sud smelt       � � � � � -  -    0000
       Iongfinsmelt    0    0 0
       eulachon                0 �           �    �
       Pacific tomcod  0� � � � � � � �
       topsmelt                           *      0 **-
       jacksmelt                                                                            -
       threespine stickleback 0� �0 000000000 00                0 � �*
       striped bass                                            � � �        � � �
       kelp bass
       barred sand bass                                                                *o-    --
       white seabass
       white croaker0                                                             0   0   0
       shiner perch    * * * *                              *                               *
       Pacific sand lance    *                                      0    0**
       arroaw goby     * * *                                        0 *0                  00
       lingcod                                                            0
       Pacific staghornsculpin � � � ��                                                       - 0   0
       Califomia halibut                                                  0000000000000
       diamond turbot  __                        V0000                                           00
       English sole    * *�'                         0               0      0 0 0
       starry flounder  0 0 0 0 0 0 0 0 0   0 0 0 0 0 0 0 0 0 0 0  0

            Includes Central San Francisco, Suisun, and San Pablo Bays.

                                                     5

    and abundance information organized by species, life   estuaries have not. Developed estuaries (i.e., those
    stage, and estuary are presented in Volume I(p. 149-    subjected to dredging and filling, jetty and port
    184). Data reliability was rated numerically as:          construction, and nearby urbanization) and their
                                                              drainages typically have been the focus of numerous
    1= Highly certain.  Considerable sampling data    research studies.  In contrast, some of the least-
    available. Distribution, ecology, and preferred habitats    developed estuaries (Willapa Bay, Nehalem Bay, Siletz
    well-documented within an estuary.                        River, and Tomales Bay) appear to be the least-
                                                              studied. Hence, there appears to be a need to collect
    2= Moderately certain. Some sampling data available    baseline fish and invertebrate distribution and
    for an estuary.  Distribution, preferred habitats, and    abundance data from relatively undeveloped and
    ecology well-documented in similar estuaries.             unpolluted estuaries.

    3= Reasonable inference. Little or no sampling data                             Mean data reliability
                                                                           Less certain               Highly certain
    available. Informationon species distributions, ecology,                  3.0   2.5   2.0    1.5   1.0
    and preferred habitatsdocumented in similar estuaries.     Estuaries
                                                               Puget Sound
    Appendix table 3 is an example of how data reliability    Hood Canal
    estimates were summarized in Volume I, and the             Skagt Bay
    following section presents an analysis of that volume's    Grays Harbor
    data reliability estimates.                               Willapa Bay
                                                               Columbia River
    Analysis of Data Contentand Quality. To assess the    Nehalem Bay
    overall certainty of the ELMR west coast data, mean        Tillamook Bay
    data reliability was calculated by estuary, species, and   Netarls River
    life stage. Mean data reliability was calculated using    Slletz River
    data reliability values for only those species and life    Yaquina Bay
    stages that were known to occur within an estuary.    Alsea River
    This allowed accu rate comparisons between estuaries    Siuslaw River
    and species since species and life stages known to be      Umpqua River
    absent were always recorded as highly certain.             coos Bay
                                                               Rogue River
    This analysis identified estuaries, species, and life    Klamath River
    stages that have the most reliable information and         Humboldt Bay
    those with the poorest. This information, combined         Eel River
    with the data in Volume I, clearly defines the ELMR        Tomales Bay
    species, life stages, and estuaries which should be the    c. San Francisco Bay 's
    focus of research efforts.  Future research should         S. an FrandscoBay
    include a comprehensive and consistent sampling            Elkhorn Slough
    program to quantify species distributions and              MorroBay
    abundances within and across estuaries. In addition,
    life history data (like the information in this report)             San Ped ro Bay
    should be compiled, especially for those species that    Alamitos Bay
    may not have economic value, but are ecologically
    important.                                                       Bay
                                                               Newport Bay                  ]
                                                               Mission Bay
    Mean data reliability of fish and invertebrate data for    San DieoBay
    west coast estuaries ranged from 2.8 (poorly-studied       TanaRiver
    Nehalem Bay) to almost 1.2 (highly-studied Columbiana River
    River) (Figure 4), with an overall average of 2.0                         3.0   2'5   2'0   115   1.0
    (moderatelycertain). In general, the reliability estimates             Less crtain                Highly certain
    reflect the amount of fisheries research that has been     �      Includes Central San Francisco, Suisun, and San Pablo Bays.
    conducted within an estuary. These data reveal that
    large estuaries (Puget Sound, Hood Canal, Skagit    Figure 4. Mean data reliability of fish and
    Bay, Columbia River, and San Francisco Bay) have                     invertebrate data collected for 32 west
    been relativelywell-studied, while most small bays and               coast estuaries.

                                                           6

    When analyzed by species (Figure 5), the data show    these species and consequently the large number of
    that salmonids and Pacific oyster have the best data    research studies that have focused on them. Poorly-
    reliability (<1.6). This reflects the economic value of    studied species (data reliability 22.0) include California
                                                             Mean data reliability
                                  Less certain                                                   Highly certain
                                       3.0            2.5             2.0             1.5              1.0
                       Species
                           blue mussel
                          Pacific oyster
                       horseneck gaper
                          Pacific gaper
                California jackknife clam _              I
                  Pacific littleneck clam
                           Manila clam -      S
                          softshell clam
                              geoduck
                            bay shrimp
                       Dungeness crab  --......
                          leopard shark
                        green sturgeon
                         white sturgeon
                        American shad
                         Pacific herring
                     deepbody anchovy
                        slough anchovy
                      northern anchovy _
                  cutthroat trout (adults)
                   cutthroat trout (kelts)
                           pink salmon                ..              .
                          chum salmon
                           coho salmon -
                  steelhead (winter run)
                steelhead (summer run) -
                steelhead (half pounder)
                     steelhead (fall run)
                       sockeye salmon
                chinook salmon (fall run) ...
            chinook salmon (late fall run)
             chinook salmon (spring run)
             chinook salmon (winter run)    - M-
           chinook salmon (summer run)
                             surf smelt
                           longfin smelt
                              eulachon
                         Pacific tomcod
                              topsmelt
                              jacksmelt
                  threespine stickleback
                           striped bass- ------ -
                              kelp bass
                      barred sand bass
                         white seabass
                          white croaker
                           shiner perch
                            arrow goby
                      Pacific sand lance _
                                lingcod
                 Pacific staghorn sculpin
                       California halibut -
                         diamond turbot    -      ----   ------- --
                           English sole - .
                         starry flounder
                                                        l              I
                                       3.0            2.5              2.0             1.5             1.0
                                   Less certain                                                   Highly certain

    Figure 5. Mean data reliability of species data collected for 32 west coast estuaries.

                                                              7

    jackknife clam, Pacific gaper, bay shrimp, cutthroat              ale     :Format o scies life history summaries.
    trout, three smelt species, Pacific tomcod, topsmelt,
    jacksmelt, threespine stickleback, arrow goby, Pacific           Common Name:the most: often used common name.
    sand lance, and Pacific staghorn sculpin.  Most of              ScIentIfc Name: the most recenttaxonomic genus:and
                                                                              spedes name.
    these species have not been studied because they are             Other Common Name s: other names that are sometimes
    not commercially important.  However, some (e.g.,                      used fora species.
    Pacific sand  lance) have  potential for increased              Classification: the mostrecent taxonomic classification
    commercial harvest or as indicators of environmental                   (Phylumrnto Family).
    health, and should be the focus of future research.             Value      s
                                                                       commercia:I information on othe commercial catches.
    When analyzed by life stage, data for juvenile and adult         Recreational: information on recreational catches.
    life stages were most reliable (1.8 and 1 .7, respectively),     Indicator of Environmental Stress: identifies if a species is an
                                                                              Iindicator of environmental degradation.
    while data pertaining to spawning adults, larvae, and              dica: the role (eg.,key predator o prey) a species
    eggswere less certain (average >2.3). This reflectsthe                 plays in a marine/estuarine ecosystem.
    number of research studies which have concentrated
    on adult and juvenile life stages.  Species-specific             Range
                                                                       Qyvrall the complete range of a species.
    studies of spawning adults, larvae, and eggs, have not           Within Study Area the range of a species within west
    been conducted in most estuaries. Thus, some of the                    coast estuaries. In addition, each summary
    information for these life stages was inferred from life               contains a relative abundance table (from Volume I)
    history studies and data from similar estuaries.                       for the 32 ELMR west coast estuaries.
                                                                       Life Mode: the life history strategy of a species and its
                                                                              life stages (e.g., anadromous, estuarine resident).

    A concise life history summary was written for each              Habitat
                                                                       I=;: the habitats used by specific life stages (e.g.,
    species to provide an overview of how and when a                       riverine, neritic, epipelagic),
    species uses estuaries and what specific habitats it             Substrate: the substrate preferences of specific life stages.
    uses.  The summaries highlight species-specific life             Phvsical/Chemical haracteristics: the physical and
    history characteristics that relate directly to estuarine              water chemistry preferences of specific life stages
                           history-~~~~~~~~~~~ ~(elg., temperature, salinity, stream flows).
    spatial and temporal distribution and abundance (e.g.,           Micirations and Movements: the movements and migratory
    many molluscs have particular salinity and substrate                   behavior of a species/life stage between or within
    preferences).  Information for the species life history                habitats.
    summaries was gathered primarily from published and
    unpublished literature; individuals who had species-             Mode: type of reproductive strategy (e.g., oviparous
    specific knowledge were also consulted. Summaries                      viviparous) and fertilization (e.g., external, internal).
    were written using the format shown in Table 3.  A               Matina/Spawnin: timing of spawning and description of
    glossary of scientific terms used  in the species                       mating or spawning behavior.
                                                                       Fecunditv: the number of-eggs or young produced by an
    summaries is provided afterthe last summary (p. 273).                  individual.

    Included with each summary is a relative abundance               Growth andDevelopment
    table based on ELMR data from Volume I. This table               Eon size and Embrvonic Develooment: the size of an
                                                                               egg and length of time for embryonic development.
    provides a synopsis of the species' occurrence in 32             Aae and Size of Larvae: the age and size range of larvae.
    west coast estuaries. Information for each table was             Juveniles Size Ranae: the size range of juveniles.
    obtained by summarizing the ELMR  data for each                  Aae and Size of Adults: the age and size range of adults.
    month of the year and across all salinity zones to obtain
    the highest level of abundance for each life stage.              Trobhic mode:type of feeder (e.g., carnivorous, herbivorous).
    Hence,  these tables depict a species' highest                   Food Items: the types of prey eaten (e.g., copepods,
    abundance within an estuary, but lack the temporal and                  amphipods, larval fish).
    spatial definition provided in Volume               Biologica Interactions
                                                                        Biological Interactions
                                                                        Predation: the predators which consume a species.
    Life History Tables.  While the species life history             Factors Influencina Pooulations: biological and physical
    summaries provide brief accounts of important life                      parameters that are known to influence a species'
    history attributes, they do not permit a direct and simple              population abundance (e.g., overfishing, ocean
    assessment of characteristics that a species shares                     productivity spawning habitat).
    with others (or lacks altogether). Furthermore, many             References: alphabetical listing of literature cited.
    life history attributes are categorical (e.g., feeding    \.

    types  can  be  classified  as  carnivore,  herbivore,                  physical and biological criteria and condensed into four
    detritivore, etc.) and more easily viewed in a tabular                  life historytables (Appendixtables 5A-5D). Majortable
    format. Therefore, information found in the species life                headings  are: Biogeography,  Habitat Associations,
    history summaries was augmented with additional   Biological Attributes and Economic Value, and

       Figure 6. Life history table headings used to develop the information in Appendix 5.

                                                                            HABITAT ASSOCIATIONS
                                                    Habitats          Substrate preference                 Domain
                                                            Z//////////                   l       Benthic     I PelagicI  Estuarine
                                                                                        I Littorall Sublitoral IBathyal

                                                             4~~42
                                       -a~~
                   Threespine     lAllOl     l l    I  I I  I    I I I I 101   I I              I I I I IOI1OII1IAI
                   stickleback    ISIool 9Io    I           I I  I I I I   11  I 10101 1  I I I I  I I 10101  1SI
                    Gasterosteas   1J1l0101010        I I I   I    I I I   11  I I I I I               10 I     oII eI.IJI
                    acudea~t.s    ILI*1*IOIOI    I   I I I I I I I I 101  I I I I I   I   I I I0I 1   ILL
                                 IEII0101*41 I I  101  I I I I 1I1  I 1.1 *I  I  I I I I I I 1.101  I El

                                                                         REPRODUCTION
                                       I  Fertikzation  Spawning    Sawnng        TemporalSchedul   I             Domain
                                       IEgg development    type    -bt empvora Scedl Periodicity   Dmi

                   Threespine
                   stickleback
                    Gasterosteus
                    aculeatcus    Ji �                III �/ I�                   �              �!

                                                                     BIOGEOGRAPHY
                                              Marine  I                  Estuarine                  Riverine
                                                           Salinity Range  I .           Catiypn
                                                        SAB  I Venice System S.--.'--'1  Stratfi-

                  Threespine     I   Ai~        I 0 1 0 1 1   I   I        A �1�� 11� 1             I~
                  stickleback   Is                       I0II* I  01  �11�  1010181910161:1:1       I I
                    Gasterostetis  lJ10101 I  101010181010101 6101610161019                  1        I III
                    aculeaturs   ILI I    II 011  1010101 I   1010101010101011                        I IL
                                  E I           10101  1010101   I    I   1��11��                    I E I

                                                     BIOLOGICAL ATTRIBUTES              Eoonomic
                                            Feeding type  I Spatialstrategy  Longevity    Viklue

                  Threespine     IAI0I I   I    101  01 01 01 1I1@I      01   I I                I IA I
                  stickleback    Isi  I I  I I         I I  I 1.1.1  I     I I I I I lIlI I SI
                    Gaaterosteus   I iIlII    I 0 I  IlI IS   lolo lI I 1               I I I   I   I  i   I
                    aculeatus       Ll01                   I I I 01  II0101   I   I l.I     IlIIl I  IL
                                 IEI  I I  I 1..1  1 1.1.1                I IooIlIlI              I    I E
                                                I   BOLGIALATRIUTS                       Eon9i

    Reproduction (Figure 6). These tables present life             =      _
    history characteristics for each species along with
    behaviortraits and preferred habitats. They reflect the    Darnell, R. M., R. E. Defenbaugh, and D. Moore. 1983.
    most current information about a species as gathered    Northwestern Gulf shelf bio-atlas. Open File Rep. No.
    from published and unpublished literature and can be    82-04. Min. Manag. Serv., Gulfof MexicoOCSRegional
    used to quickly identify species with similar traits. For    Office, Metairie, LA, 438 p.
    example, a reader interested only in pelagic (as opposed
    to benthic) species can use Appendix table 5B, Habitat   Gunter, G. 1967. Some relationships of estuaries to
    Associations, to identify relevant species. In addition,    the fisheries of the Gulf of Mexico.  In G. H. Lauff
    terms used in the life history tables are defined in   (editor), Estuaries, p. 621-638. Am. Assoc. Adv. Sci.
    Appendix 6.                                             Special Publ. No. 83, Washington, D.C.

                          I___________________________ _ _   Hammerschmidt, P. C., and L. W. McEachron. 1986.
                                                           Trends in relative abundance of selected shellfishes
    As it becomes apparent that the cumulative effects of   along the Texas coast: January 1977 - March 1986.
    smallalterationsinmanyestuarieshaveatotalsystemic    Texas Parks Wildl. Dept., Coast. Fish. Branch, Mgmt.
    impact on coastal ocean resources, it is more important    Data Ser., No. 108, 149 p.
    than ever to compile consistent information on the
    Nation's estuarine fishes and invertebrates. Although    Joseph, E. B. 1973. Analysis of a nursery ground. In
    the knowledge available to effectively preserve and    A. L. Pacheco (editor), Proceedings of a workshop on
    manage estuarine resources is limited, the ELMR data    egg, larval, and juvenile stages of fish in Atlantic Coast
    base provides an important tool for assessing the    estuaries. p. 118-121. Mid. Atlantic coast. Fish. Cent.,
    status of estuarine fauna and examining their    Tech. Publ. No. 1, Beaufort, NC.
    relationships with other species and their environment.
    These life history summaries and life history tables    Mann, K. H. 1982. Ecology of coastal waters. Univ.
    highlight many of the biological and environmental    Calif. Press, Los Angeles, CA, 322 p.
    factors that play a role in determining each species'
    distribution and abundance. Together, the ELMR data    Monaco, M. E. 1986. National estuarine inventory:
    base and life history information will provide valuable    Living marine resources component, preliminary west
    baseline information on the biogeography and ecology    coast study.  Ocean Assessments Division, NOS/
    of estuarine fishes and invertebrates, and identify gaps    NOAA, Rockville, MD, 33 p.
    in our knowledge of these valuable national resources.
                                                           Monaco, M. E., T. E. Czapla, D. M. Nelson, and M. E.
    The ELMRprogramiscontinuingtocompileandassess    Pattillo. 1989. Distribution and abundance of fishes
    estuarine biological and physical data to improve the    and invertebrates in Texas estuaries.  Strategic
    Nation's ability to manage coastal ocean resources.    Assessment Branch, NOS/NOAA,  Rockville, MD,
    Forthcoming reports will help further define the    107 p.
    importance of west coast estuaries to fishes and
    invertebrates.  One of these reports will present    Monaco, M. E., R. L. Emmett, D. M. Nelson, and S. A.
    information on salmonid hatchery production and    Hinton. 1990. Distribution and abundance of fishes
    escapement for several west coast estuarine basins.    and invertebrates in west coast estuaries, Volume I:
    Another will present results of multivariate analyses of    Data summaries. Strategic Assessment Branch, NOS/
    the ELMR west coast fish data to identify the coupling    NOAA, Rockville, MD, 240 p.
    of species distributions and estuarine physical and
    hydrological characteristics.                           NOAA  (National  Oceanic  and  Atmospheric
                                                           Administration). 1984. The national status and trends
          ,_, .,* o.                   .                    program for marine environmental quality: Program
                                                           description (memo).  Ocean Assessments Division,
    The authors thank the many individuals who provided    NOSINOAA, Rockville, MD, 28 p.
    information forthis report, and the many other scientists
    and managers who provided contacts and references.    NOAA  (National  Oceanic  and  Atmospheric
    We appreciate the editorial assistance provided by    Administration). 1985. National estuarine inventory:
    Mitchell Katz, Kim Keeter-Scott, and Robert Wolotira.    Data atlas.  Volume 1.  Physical and hydrologic
    Special thanks is due to Ron Pitard, Nancy Nelson, and    characteristics. Strategic Assessment Branch, NOS/
    Sandy Noel for preparing the species illustrations.     NOAA, Rockville, MD, 103 p.

                                                        10

    NOAA  (National  Oceanic  and  Atmospheric    For additional copies or information contact:    .
    Administration). 1988. Bering, Chukchi, and Beaufort
    Seas strategic assessment: Data atlas. Volume 1.   Robert L. Emmett
    Physical and hydrologic characteristics.  Strategic    Point Adams Biological Field Station
    Assessment Branch, NOS/NOAA, Rockville, MD,            CoastalZone& Estuarine Studies Division
    135 p.                                                 Northwest Fisheries Center
                                                          National Marine Fisheries Service
                                                          Hammond, OR 97121
    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,   FrS/Comm. (503) 861-118
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the              or
    United States and Canada.  Am. Fish. Soc., Spec.
    Publ. No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.    Steven L. Stone
                                                          Strategic Environmental Assessments Division
    Turgeon, D. D., A. E. Bogan, E. V. Coan, W. K.   Office of Ocean Resources Conservation & Assessment
    Emerson, W. G. Lyons, W. L. Pratt, C. F. E. Roper, A.    National Ocean Service
    Scheltema, F. G. Thompson, and J. D. Williams. 1988.    FTS/Comm,(301) 443-0453
    Common and scientific names of aquatic invertebrates
    from the United States and Canada: mollusks. Am.
    Fish. Soc. Spec. Publ. No. 16, Am. Fish. Soc., Bethesda,
    MD, 277 p.

    Weinstein, M. P. 1979. Shallow marsh habitats as
    primary nurseries for fishes and shellfish, Cape Fear
    River, North Carolina. Fish. Bull., U.S. 77:339-357.

    Williams, A. B., L. G. Abele, D. L. Felder, H. H. Hobbs,
    Jr., R. B. Manning, P. A. McLaughlin, and I. P6rez
    Farfante. 1988. Common and scientific names of
    aquatic invertebrates from the United States and
    Canada: decapod crustaceans. Am. Fish. Soc. Spec.
    Publ. No. 17, Am. Fish. Soc., Bethesda, MD, 77 p.

    Williams, C. W., D. M. Nelson, M. E. Monaco, S. L.
    Stone, C. lancu, L. C. Clements, L. R. Settle, and E. A.
    Irlandi. 1990. Distribution and abundance of fishes
    and invertebrates in eastern Gulf of Mexico estuaries.
    Strategic Assessment Branch, NOS/NOAA, Rockville,
    MD, 240 p.

    12

    blue mussel (Mytilus edulis) ............................................................................................................... 14
    Pacific oyster (Crassostrea gigas) ..................................................................................................... 20
    horseneck gaper (Tresus capax) ....................................................................................................... 26
    Pacific gaper (Tresus nuttalli) ............................................................................................................ 30
    California jackknife clam ( Tagelus californianus) ........................................................................... 34
    Pacific littleneck clam (Protothaca staminea) ..................................................................................... 38
    Manila clam (Venerupisjaponica) ........................................                44
    softshell (Mya arenaria) ..................................................................................................................... 50
    geoduck (Panopea abrupta) ............................................................................................................... 56
    bay shrimp (Crangon franciscorum) ................................................................................................... 62
    Dungeness crab (Cancer magister) ................................................................................................... 68
    leopard shark ( Triakis semifasciata) .................................................................................................. 78
    green sturgeon (Acipenser medirostris) ............................................................................................. 82
    white sturgeon (Acipenser transmontanus) ....................................................................................... 86
    American shad (Alosa sapidissima) ................................................................................................... 90
    Pacific herring (Clupea pallas) .......................................................................................................... 94
    deepbody anchovy (Anchoa compressa) ........................................           100
    slough anchovy (Anchoa delicatissima) ........................................         104
    northern anchovy (Engraulis mordax) ........................................           108
    cutthroat trout (Oncorhynchus clarkr) ........................................         114
    pink salmon (Oncorhynchus gorbuscha) ........................................          120
    chum salmon (Oncorhynchus keta) ................................................................................................. 128
    coho salmon (Oncorhynchus kisutch) ........................................            136
    steelhead (Oncorhynchus mykiss) ........................................               146
    sockeye salmon (Oncorhynchus nerka) ........................................           152
    chinook salmon (Oncorhynchus tshawytscha) ........................................     160
    surf smelt (Hypomesus pretiosus) ........................................              170
    longfin smelt (Spirinchus thaleichthys) ........................................       174
    eulachon (Thaleichthys pacificus) ........................................             178
    Pacific tomcod (Microgadus proximus) ........................................          182
    topsmelt (Atherinops affinis) ........................................                 186
    jacksmelt (Atherinopsis californiensis) ........................................       190
    threespine stickleback (Gasterosteus aculeatus) ........................................  194
    striped bass (Morone saxatilis) ........................................               200
    kelp bass (Paralabrax clathratus) ........................................             208
    barred sand bass (Paralabrax nebulifer) ........................................       212
    white seabass (Atractoscion nobilis) ........................................          216
    white croaker (Genyonemus lineatus) ........................................           220
    shiner perch (Cymatogaster aggregata) ........................................         226
    Pacific sand lance (Ammodytes hexapterus) ........................................     232
    arrow goby (Clevelandia ios) ........................................                  238
    lingcod (Ophiodon elongatus) ........................................                  242
    Pacific staghorn sculpin (Leptocottus armatus) ........................................  246
    California halibut (Paralichthys californicus) ........................................  250
    diamond turbot (Hypsopsetta guttulata) ........................................        256
    English sole (Pleuronectes vetulus) ........................................           260
    starry flounder (Platichthys stellatus) ........................................       266

                                                        13

        Mytilus edulis
       Adult

                                          2cm

    Common Name: blue mussel                              However, California inland waters are closed to
    Scientific Name: Mytilus edulis                       harvesting from May 1 to October 31 (both sport and
    Other Common Names: bay mussel, edible mussel,    commercial) because of potential forparalyticshellfish
    black mussel, pile mussel (Gates and Frey 1974)       poisoning. Sixculturemethodsarecurrentlyemployed:
    Classification (Bernard 1983)                         raft, post, bottom, pole and line, long line, and rack.
    Phylum: Mollusca                                      Spain is currentlythe world's largest producerofcultured
    Class: Bivalvia                                       blue mussels (Oceanographic Institute of Washington
    Order: Mytiloida                                      1981). There appears to be an excellent opportunity
    Family: Mytilidae                                     for more U.S. aquaculture of this species (Lutz 1980).

    Recent research has shown that Pacific coast "Mytilus   Recreational: Estimates of blue mussels harvested by
    edulis" populations may actually be composed of two    sportsmen are presently unknown.  However, this
    distinct species: M. trossulus Gould, 1850, distributed    species is regularly used as bait and human food
    from northern California through Alaska and the Soviet   throughout its range.
    Union, and M. galloprovincialis Lamarck, 1819,
    distributed in Japan, Hong Kong, South Africa, the    Indicator of Environmental Stress: Since it readily
    Mediterranean Sea, the Atlantic coasts of Europe and    takes up and concentrates contaminants in the marine
    the British Isles, and southern California. In central   environment,thisspecies hasbeenused as a "sentinel"
    California, both species are present along with hybrids   of environmental quality (National Research Council
    (McDonald and Koehn 1988). However, this species    1980, Broman and Ganning 1986).  Increased
    summary presents information using the previous   temperatures can interact with zinc and salinity to
    nomenclature of M. edulis.                            accelerate toxic effects (Cotter et al. 1982). Even low
                                                          concentrations of tributyltin oxide (a paint additive)
    Value                                                 reduce mussel growth hyperbolically (Str0mgren and
    Commercial: Between 1942 and 1947, up to 1,350 t   Bongard 1987). A decline in the scope forgrowth of M.
    were harvested annually in the United States (Cheney    edulishas been correlated with increasing body burdens
    and Mumford 1986), but the harvest declined    of chromium, copper, mercury, silver, aluminum, zinc,
    dramatically after that period.  Since the 1960s,   total chlordanes, and dieldrin (Martin et al. 1984).
    cultivation and harvesting increased; in 1981, 7,500 t   Heavy metals, particularly mercury and copper, inhibit
    were landed with most cultivation and harvesting    byssal-thread formation. Lead is incorporated at a rate
    occurring on the east coast, primarily in New England    that is linear with seawater concentration, thus making
    (Cheney and Mumford 1986).  Cultivation of blue   this an ideal animal for monitoring lead pollution in
    mussels has recently been initiated in Oregon and    marine environments (Haderlie and Abbott 1980).
    California coastal waters, and in Puget Sound,    Mussel embryos are highly sensitive to trace metals
    Washington.  Presently, mussels are commercially    (Martin et al. 1981). Crudeoil is not highlytoxicto adult
    harvested from California offshore oil platforms.    and juvenile blue mussels (Roberts 1976).

                                                       14

                                                                                            Blue mussel continued
                                                            Range
      Table 1. Relative abundance of blue mussel             Overall: The blue mussel, cosmopolitan in temperate
               in 32 U.S. Pacific coast estuaries.          and cold seas (Bernard 1983), is very abundant in
                         Life Stage                         quiet-water locations from Puget Sound to Alaska
            Estuary    A S J L E                            (Ricketts et al. 1985). In the Pacific Ocean, it ranges
            PugetSound a s a     *    Relative Abundance:   fromAlaskatoCedrosisland,Mexico(Morris1966). It
             Hood Canal ( {3 {I t)     *   Highly abundant  is also found on the west coast of South America, and
              Skagit Bay *           � �   3   Abundant     in Japan, Australia, and the North Atlantic (Haderlie
                                       O   Common
            Grays Harbor0 0    C       �   Common     and Abbott 1980). On the east coast of North America,
             Willape Bay C O  0 0  Blank Not present        the blue mussel ranges from Cape Hatteras, North
           Columbia River O  0 0   O                         Carolina to Labrador (Newell 1989). In the western
            Nehalem Bay  ]  ) 1 3i                          Atlantic, it is found in Great Britain, Ireland, Scandinavia,
            Tillamook Bay 13 13  (9 I) (    Life Stage:      and the Baltic Sea.
             Netarts Bay i (t) 3 (   3   A - Adults
                                       S - Spawning adults
             Siletz River a 4 4 ': V   J -Juveniles         Within Studv Area: This species is found in nearly all
            Yaquina Bay   3       13I (3   L- Larvae        Pacific coast estuaries, but is most abundant in the
                    ~Alsea River v4 f  E - Eggs              northern part of its range (Table ). In many southern
             AlseaRiver  '4 'J- 4 nothr pato't4rne(al                                       1).I aysuhr
            Siuslaw River (i  13 i (&3                      California estuaries, this species is restricted to wharf
           UmpquaRiver * 6   �                              pilingsandtheundersidesoffloatingdocks(Rickettset
               CoosBay a3  3  (3 (�                         al. 1985).
             Rogue River '4    i
            Klamath River                                    Life Mode
            Humboldt Bay a �    � �                          Eggs and larvae are pelagic. Juveniles and adults are
               Eel River O C 0 O                            sessile and epibenthic, living on hard or rocky bottoms
            Tomales Bay (* i Ci 1 (                         orany relativelystable habitats (pilings, wharfs, hanging
       CentSanFran.Bay* ' S     �      * IncdudesCentralSan  ropes, etc.). Juvenilesandadultsdonotneed lightand
                                      Francisco, Suisun,
       South San Fran. Bay S U S    �   and San Pablo Bays.  are often found underneath floating objects.  They
           Elkhorn Slough (3 (3]I) (  ]                      attachthemselvestothesesubstratesbybyssalthreads.
              Morro Bay { 1 (3  13           }              All lifestagescanbefoundinestuariesandinnearshore
        SantaMonicaBay  3 (i 3 (3 {                         marine environments.  Juveniles and adults do not
           San Pedro Bay a (3 *     {3                       dominate exposed nearshore rocky marine habitats;
            AlamitosBay (S) C) (3 (                         theCaliforniamussel M. californianusappearstohave
            Anaheim Bay (13 (3 (3 (3                        a competitive advantage in these areas.
            Newport Bay O 0 0 0 0
             Mission Bay ({3 i } (3 (3                      Habitat
           San Diego Bay (3 s3  {3 () (3                    Type: All life stages inhabit marine and estuarine
          Tijuana Estuary C D O C   O                       environments. They are most often found in estuaries
                       A S J L E                            or protected bays, since they prefer quiet water. Blue
                                                            mussels occur primarily intertidally to 5 m depth, but
                                                            have been found to 36 m (Cheney and Mumford 1986).
                                                            In many northern locations, they are found only
    Ecological: Aggregations of this species often form a    sublittorally (Seed 1976). The upper tidal limit of blue
    distinct "band" on substrates (pilings, rocks, etc.) where    mussels is related to physical factors (e.g., exposure to
    environmental conditions are suitable. These bands    air and desiccation), while the lower limit is probably
    have a characteristic animal assemblage (i.e., mussel    determined by predation (Seed 1976).
    shells also provide substrates for barnacles, hydroids,
    bryozoans, and ascidians) (Kozloff 1976, Ricketts et al. Substrate: Plantigrades (late larval stages) appear to
    1985). This species is a common fouling organism.    use algae-covered substrates initially before finding
    Larvae  are  important  prey  for carnivorous    final attachment sites (Seed 1976).  Juveniles and
    zooplanktivores (Bayne 1976). Bbe mussel populations    adults can be found on a variety of substrates, ranging
    appearto be important in cycling nitrogen, phosphorus,    from coarse unconsolidated substrates to rocky
    and amino-nitrogen in some marine environments    outcrops. Almost any fairly stable substrate can be
    (Kautsky and Wallentinus 1980, Kautsky 1981, Kautsky    used for settlement; including many man-made objects
    and Evans 1987).  Genetic differences between    such as pilings, ropes, wharfs, boat bottoms, buoys,
    populations may enable them to invade suboptimal    etc. (Shaw et al. 1988).
    habitats (Koehn et al. 1984, Mallet et al. 1987).

                                                        15

    Blue mussel continued
    Physical/Chemical Characteristics: This species is   spawning occurs when water temperatures warm to
    found in waters that range in temperature from -4 to   18�C (late spring or summer) (D. Tufts, Willapa Bay
    30�C (Bernard 1983). Itcanwithstandtemperaturesof   Shellfish Lab., Washington Department of Fisheries,
    1.7-26.7�C (Cheney and Mumford  1986), but    P.O. Box 190, Ocean Park, WA, pers. comm.). Mussels
    temperatures above 20�C appear to be stressful (Hines   in British waters spawn when water temperatures rise
    1979). Trochophore development occurs best within a   from 9.5�C to 11 -12.50C (Chipperfield 1953). In Puget
    salinity range of 30 to 40%o and temperatures of 8-1 8�C    Sound, Washington spawning occurs from late spring
    (Bayne 1965). Larval survival at salinities from 15-   through midsummer, with the spawning duration being
    40/0o and temperatures of 5-200C is good, but drops    a few weeks in any location (Cheney and Mumford
    drastically at 25�C. Optimum larval growth occurs at   1986). Spawning begins in May in northern California,
    200C in salinities of 25-30%o (Brenko and Calabrese    with partially spent mussels found until November
    1969). Juveniles and adults tolerate salinities of 5-   (Edwards 1984). In southern California, some males
    37%/o andcanwithstand 0%oforashortperiod. Optimum    may be ripe all year-round, but females have mature
    temperature for juvenile and adult growth is 10-200C    ova from November-May (Moore and Reish 1969,
    (Haderlie and Abbott 1980) and optimum salinity is 10-   Haderlie and Abbott 1980). In British Columbia, most
    30%o; it can tolerate low oxygen for several days. The    blue mussels appearto spawn in spring, but some may
    blue mussel prefers areas with slow to medium water   also spawn again in fall (Emmett et al. 1987). Mussels
    currents and areas protected from surf. Limited data    are stimulated to spawn by increasing water
    suggest that environmental requirements may limit   temperature, mechanical action, strong wave action,
    embryonic development, especially in estuarine   lunar cycle, and various chemicals (Cheney and
    populations (Bayne 1976). It appears that when water    Mumford 1986).
    conditions become adverse, adult and juvenile mussels
    will isolate themselves from these conditions (close   Fecundity: Fecundities rangefrom3 millionto6 million
    shellandreducepumpingactivity)andrelyonanaerobic    eggs per female (Skidmore and Chew 1985).
    metabolism (Aunaas et al. 1988). Bay mussels are
    often infected with the parasitic copepod Mytilicola   Growth and Development
    orientalis (Bradley and Siebert 1978).                Eaoo Size and Embrvonic DeveloDment: Eggs are ovoid
                                                          and 0.068-0.070 mm in diameter (Bayne 1976).
    Miarations and Movements: Larvae swim freely for   Embryonic development is indirect and external, and
    approximately 4 weeks, settling mainly in the summer    takes about 48 hours.
    in southern California (Haderlie and Abbott 1980). In
    Puget Sound, peaksettlementvarieswidelybutusually    Aae and Size of Larvae: Fertilized eggs first form
    occurs from late April through early July. The period of   trochophore and then veliger larvae; these larval stages
    settlement appears todepend primarilyontemperature    do not have a shell. Once secretion of the shells has
    (Skidmore and Chew 1985).  Post-larval mussels    started, the larva is called a veliconcha. In this form,
    secrete long, single, unattached byssalthreads, which    locomotion is provided by the velum. As the larva nears
    increase drag and allow young mussels to be carried   metamorphosis, a pedal organ develops; when this is
    by weak currents (Haderlie and Abbott 1980).   functional, the larva is called a pediveliger.  After
    Plantigradesoften attach anddetachthemselves many    secretion of the adult shell (dissochonch) begins, the
    times before finally settling (Seed 1976). Larvae may    larva is called a plantigrade (Bayne 1976) and is ready
    undergo diurnal vertical migrations and "selective   to settle out of the water column. The length of the
    swimming" (swimming at different tide stages), thus   larval stages dependson food availability, temperature,
    aiding retention in estuaries (Bayne 1976). Juvenile   salinity, and other variables (Bayne 1976).  Larvae
    and adult blue mussels appearto be more mobile than    mature into spat in 3-4 weeks, but may remain planktonic
    M.califomianus. Blue mussels apparentlycancrawlto   for up to 10 weeks (Cheney and Mumford 1986).
    the edge of mixed colonies. This ability also permits    Veliger larvae are about 0.110-0.260 mm  wide;
    them to move when sedimentation threatens to bury   plantigrades are approximately 0.26-1.50 mm wide
    them (Haderlie and Abbott 1980).                      (Bayne 1976).

    Reproduction                                          Juvenile Size Ranae: The blue mussel is 1.0-1.5 mm
    Mode: The blue mussel is gonochoristic (but some    long at settlement (Newell 1989). Growth rates are
    hermaphroditism has been reported), oviparous, and    highly variable depending on area, temperature, food
    iteroparous.  It is a broadcast spawner; eggs are   availability, and other factors.
    fertilized externally.
                                                          Aae and Size of Adults: Most appearto mature in about
    Matina/SDawnina: In Willapa Bay, Washington,    ayear, dependingonfoodavailabilityandotherphysical

                                                       16

                                                                                          Blue mussel continued
    factors. The smallest adults may be 10 mm long and    related to predation. Above mean tide level, the blue
    they rarely grow more than 5 cm long.  However,    mussel competes with Balanus glandula (Ross and
    specimens upto 10 cm long have been found (Ricketts    Goodman 1974).
    et al. 1985). Cultured mussels can reach 50 mm long
    (marketable size) in 12-13 months in Puget Sound    References
    (Skidmore and Chew 1985). This size is reached in 2-
    3 years in natural California populations. The oldest    Aunaas, T., J. P. Denstad, and K. E. Zachariassen.
    recorded specimens (18-24 years old ) were from cool    1988.  Ecophysiological importance of the isolation
    northern climates (Seed 1976). Growth may be limited    response of hibernating blue mussels (Mytilus edulis).
    by immersion time which in turn may be a result of   Mar. Biol. 98:415-419.
    vertical distribution (Suchanek 1978).
                                                           Bayne, B. L. 1965. Growth and delay of metamorphosis
    Food and Feeding                                        of the larvae of Mytilus edulis (L.). Ophelia 2:1-47.
    Tro]hic Mode: Larvae, juveniles, and adults are
    planktivorous filter feeders; pelagic detritus and    Bayne, B. L. 1976. The biology of mussel larvae. In
    planktonic organisms are trapped by mucus sheets    B. L. Bayne (editor), Marine mussels:their ecology and
    that move over the gills. They can select food items    physiology, p. 81-410.   Cambridge Univ. Press,
    and reject non-food items.                              Cambridge, U.K.

    Food Items: Larvae feed on phytoplankton. Juveniles    Bernard, F. R. 1983. Catalogue of the living Bivalvia
    and adults feed on detritus, phytoplankton (such as    of the eastern Pacific Ocean: Bering Strait to Cape
    dinoflagellates) and organisms as small as 4-5 pm in   Horn. Can. Spec. Publ. Fish. Aquat. Sci. 61, 102 p.
    diameter (Incze et al. 1980). Organic detritus can be a
    majorfood source, and they also absorb dissolved and    Bradley, W., and A. E. Siebert, Jr. 1978. Infection of
    particulate organic compounds (Haderlie and Abbott    Ostrealuridaand Mytilusedulisbytheparasiticcopepod
    1980).                                                  Mytilicola orientalis in San Francisco Bay, California.
                                                           Veliger 21 (1):131 -134.
    Biological Interactions
    Predation: Predation has at times resu Ited in the loss of    Brenko, M. H., and A. Calabrese. 1969. The combined
    50% of the harvestable blue mussels in an area.    effects of salinity and temperature on larvae of the
    Important predators include perch (Embiotocalateralis    mussel Mytilus edulis. Mar. Biol 4(3):224-226.
    and Rhacochilus vacca), crabs (Cancer spp., and
    Pachygrapsuscrassipes),starfish(Pisasterochracea),    Broman, D., and B. Ganning.  1986.  Uptake and
    snails (Nucella spp.), and scoter ducks (Melanitta spp.    release of petroleum hydrocarbons by two brackish
    and  Oidemia nigra) (Waterstrat et al. 1980,    waterbivalves, Mytilusedulis(L.)andMacomabalthica
    Oceanographic  Institute of Washington  1981).    (L.). Ophelia25(1):49-57.
    Planktivorous fishes and invertebrates are important
    predators of blue mussel larvae.                        Cheney, D. P., and T. F. Mumford, Jr. 1986. Shellfish
                                                           and seaweed harvests of Puget Sound. Wash. Sea
    Factors Influencina PoDulations: Paralytic shellfish    Grant, Univ. Wash. Press, Seattle, WA, 164 p.
    poisoning can reduce mussel abundances (Reish 1963)
    and may result in unharvestable products. Diseases    Chipperfield, P. N. J.  1953.  Observations of the
    suchas hemocytic neoplasia mayalsocausesubstantial    breeding and settlement of Mytilus edulis (L.) in British
    mortality (Elston et al. 1988). Pollution (both industrial   waters. J. Mar. Biol. Ass. U.K. 32:449-476.
    and residential) is a major problem for mussel growers
    (Oceanographic Institute of Washington 1981). Other    Cotter, L. J. R., D. J. H. Phillips, and M. Ahsanullah.
    factors which reduce this species' abundance are    1982. The significance of temperature, salinity, and
    diseases, fouling, and storms.  The mortality rate   zinc as lethal factors for the mussel Mytilus edulis in a
    during the pelagic larval stage is probably as high as    polluted estuary. Mar. Biol. 68:135-141.
    99% (Bayne 1976). Causes of larval mortality include
    predation, excessive dispersal, andunsuitablephysical    Edwards, R. L. 1984. The reproductive percentage
    parameters. Adult mortality may also be caused by    solidscyclesofMytilusedulisand Mytiluscalifornianus
    spawning-related stress (Emmett et al. 1987). The    in Humboldt County, California. M.S.Thesis, Humboldt
    blue mussel's upper intertidal distribution appears to   State Univ., Arcata, CA, 57 p.
    be related to the survival of settling spat (Ross and
    Goodman 1974). Lower distribution is most often

                                                        17

    Blue mussel continued
    Elston, R. A., M. L. Kent, and A. S. Drum.  1988.    Mussels culture and harvest: a North American
    Progression, lethality and remission of hemic neoplasia    perspective, p. 1-17.  Elsevier Scientific Publ. Co.,
    in the bay mussel Mytilus edulis.  Diseases Aquat.    Amsterdam, Holland.
    Organ. 4:135-142.
                                                            Mallet, A. L., C. E. A. Carver, S. S. Coffen, and K. R.
    Emmett, B., K. Thompson, and J. D. Popham. 1987.    Freeman.  1987.  Mortality variations in natural
    The reproductive and energy storage cycles of two    populations of the blue mussel, Mytilus edulis. Can. J.
    populations of Mytilus edulis (L.) from British Columbia.    Fish. Aquat. Sci. 44:1589-1594.
    J. Shellfish Res. 6(1):29-36.
                                                            Martin, M., G. Ichikawa, J. Goetzl, M. de los Reyes, and
    Gates, D. E., and H. W. Frey.  1974.  Designated    M. D. Stephenson.  1984.  Relationships between
    common names of certain marine organisms of   physiological stress and trace toxic substances in the
    California. Calif. Fish Game, Fish Bull. 161:55-90.     bay mussel, Mytilus edulis, from San Francisco Bay,
                                                            California. Mar. Envir. Res. 11:91-110.
    Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: the
    clams and allies. In R. H. Morris, D. P. Abbott, and E.    Martin, M., K. E. Osborn, P. Billing, and N. Glickstein.
    C. Haderlie (editors),  Intertidal invertebrates of    1981. Toxicities often metals to Crassostreagigasand
    California, p. 355-411. Stanford Univ. Press, Stanford,    Mytilus edulis embryos and Cancer magister larvae.
    CA.                                                     Mar. Poll. Bull. 12(9):305-308.

    Hines, A. H. 1979. Effects of a thermal discharge on    McDonald, J. H., and R. K. Koehn. 1988. The mussels
    reproductive cycles in Mytilus edulis  and  Mytilus   Mytilusgalloprovincialisand M. trossuluson the Pacific
    californianus (Mollusca, Bivalvia). Fish. Bull., U.S.    coast of North America. Mar. Biol. 99:111-118.
    77(2):498-503.
                                                            Moore, D. R., and D. J. Reish. 1969. Studies on the
    Incze, L. S., R. A. Lutz, and L. Watling.   1980.    Mytilusedulis community in Alamitos Bay, California.-
    Relationships between effects of environmental    IV. Seasonal variation in gametes from different regions
    temperature and seston on growth and mortality of   in the bay. Veliger 11(3):250-255.
    Mytilus edulis in a temperate northern estuary. Mar.
    Biol. 57:147-156.                                       Morris, P.A. 1966. A field guide to Pacific coast shells.
                                                            Houghton Mifflin Company, Boston, MA, 297 p.
    Kautsky, N. 1981. On the trophic role of the blue
    mussel (Mytilus edulis L.) in a Baltic coastal ecosystem    National Research Council. 1980. The international
    and the fate of the organic matter produced by the    mussel watch.  Nat. Acad. Sci., Washington, D.C.,
    mussels. Kieler Meeresforsch., Sonderh. 5:454-461.    248 p.

    Kautsky, N.,and S. Evans. 1987. Roleofbiodeposition    Newell, R. I. E., 1989. Species profiles: life histories
    by Mytilus edulisin the circulationof matterand nutrients    and environmental requirements of coastal fishes and
    in a Baltic coastal ecosystem. Mar. Ecol. Prog. Ser.    invertebrates (North and Mid-Atlantic)-blue mussel.
    38:201-212.                                             U.S. Fish. Wildl. Serv. Biol. Rep. 82(11.102).  U.S.
                                                            Army Corps Eng., TR EL-82-4, 25 p.
    Kautsky, N., and I. Wallentinus. 1980. Nutrient release
    from a Baltic Mytilus -red algal community and its role    Oceanographic Institute of Washington. 1981. Clam
    in benthic and pelagic productivity. Ophelia (suppl).    and mussel harvesting industries in Washington State.
    1:17-30.                                                Oceanogr. Comm. Wash., Seattle, WA,   various
                                                            pagination.
    Koehn, R. K., J. G. Hall, D. J. Innes, and A. J. Zera.
    1984. Genetic differentiation of Mytilus edulis in eastem    Reish, D. J. 1963. Mass mortality of marine organisms
    North America. Mar. Biol. 79:117-126.                   attributed to the "red tide" in southern California. Calif.
                                                            Fish Game 49:265-270.
    Kozloff, E. N. 1976. Seashore life of Puget Sound, the
    Strait of Georgia, andthe San Juan Archipelago. Univ.    Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W.
    Wash. Press, Seattle, WA, 282 p.                        Phillips. 1985. Between Pacific tides. Stanford Univ.
                                                            Press, Stanford, CA, 652 p.
    Lutz, R. A. 1980. Introduction: mussel culture and
    harvest in North America. In R. A. Lutz (editor),

                                                         18

                                                                                          Blue mussel continued
    Roberts, D. 1976. Mussels and pollution. In B. L.
    Bayne (editor), Marine mussels: their ecology and
    physiology, p. 67-80, Cambridge Univ. Press,
    Cambridge, U.K.

    Ross, J. R. P., and D. Goodman. 1974. Vertical
    intertidal distribution of Mytilusedulis. Veliger 16(4):388-
    395.

    Seed, R. 1976. Ecology. In B. L. Bayne (editor),
    Marine mussels: their ecology and physiology, p. 13-
    65, Cambridge Univ. Press, Cambridge, U.K.

    Shaw, W. N., T. J. Hassler, and D. P. Moran. 1988.
    Species profiles: life histories and environmental
    requirements of coastal fishes and invertebrates (Pacific
    Southwest)-California sea mussel and bay mussel.
    U.S. Fish Wildl. Serv. Biol. Rep. 82(11.84), U.S. Army
    Corps Eng., TR EL-82-4, 16 p.

    Skidmore, D., and K. K. Chew. 1985. Mussel
    aquaculture in Puget Sound. Wash. Sea Grant, Univ.
    Wash., Seattle, WA, 57 p.

    Str0mgren, T., and T. Bongard. 1987. The effect of
    tributyltin oxide on growth of Mytilus edulis. Mar. Poll.
    Bull. 18(1):30-31.

    Suchanek, T. H. 1978. The ecology of Mytilus edulis
    L. in exposed rocky intertidal communities. J. Exp.
    Mar. Biol. Ecol. 31:105-120.

    Waterstrat, P., K. Chew, K. Johnson, and J. H. Beattie.
    1980. Mussel culture: a west coast perspective. In R.
    A. Lutz (editor), Mussel culture and harvest: a North
    American perspective, p. 141-165, Elsevier Scientific
    Publ. Co., Amsterdam, Holland.

                                                       19

       Crassostrea gigas
       Adult

                                          5cm
    Common Name: Pacific oyster                           Bolinas Lagoon, and Morro Bay (Barrett 1963, Pauley
    Scientific Name: Crassostreagigas                     et al. 1988, Wolotira et al. 1989). Nearly all Pacific
    Other Common Names: Japanese oyster, Miyagi   oysters are cultivated on "oyster farms" in protected
    oyster, giant oyster, immigrant oyster, giant Pacific   coastal estuaries. Since successful spawning in many
    oyster (Fitch 1953, Gates and Frey 1974, Wolotira et al.   estuaries is erratic, Pacific coast hatcheries have been
    1989)                                                 developed to produce spat, which is then sold to oyster
    Classification (Bernard 1983a)                        growers who use this to "seed" their oyster beds. Prior
    Phylum: Mollusca                                     to the development of these hatcheries, all seed was
    Class: Bivalvia                                       imported from Japan (Conte and Dupuy 1981, Ricketts
    Order: Pterioida                                      et al. 1985, Pauley et al. 1988). The seed is allowed to
    Family: Ostreidae                                     grow, but clusters may have to be broken up and the
                                                          oysters moved to fattening grounds before harvest
    Value                                                 (Beattie et al. 1981). Pacific oysters are harvested
    Commercial: The Pacific oyster is a highly valuable   primarily by hydraulic dredge, tongs, and hand-picking
    estuarine speciesthat is cultured in appropriate habitats   (Frey 1971, Cheney and Mumford 1986). Most oysters
    all over the world, including Australia, Japan, Hawaii,    are sold fresh-shucked and frozen, while some are
    Palau, southwest Europe, and the Pacific coast of   canned or sold fresh in the shell. The Japanese have
    North America (Haro et al. 1981, Lee et al. 1981,    cultured Pacific oysters for over 300 years, and have
    Menzel 1974, Quayle 1988). It was introduced to the   developed numerous raft, line, and pole mariculture
    United States from Japan in the early 1900s and has    methods instead of on-bottom methods used primarily
    been cultured ever since (Quayle 1988).  In North   in the U.S. and British Columbia (Bardach et al. 1972,
    America, they are harvested from southeast Alaska to   Haderlie and Abbott 1980, Gunn and Saxby 1981,
    northern Baja California, with most produced in   Pauley et al. 1988).
    Washington and southwest British Columbia waters
    (Wolotiraetal. 1989). It is Washington's mostvaluable    Recreational: Although most oysters are cultivated,
    shellfish resource (Pauley et al. 1988).  In 1982,    some wild beds do exist in Washington and British
    Washington alone harvested over 2,700 t of meat,    Columbia. In Puget Sound and Hood Canal, the daily
    worth $20.4 million, and representing over 70% of all   limit is 18/person,withtheseasonopenfromSeptember
    Pacific coast harvests (Cheney and Mumford 1986).    16 to July 14, except for a couple of state parks
    AbouthalfofWashington'slandingscomefromWillapa    (Washington Department of Fisheries 1986, Wolotira
    Bay (Hedgpeth and Obrebski 1981, Washington    et al. 1989). Oysters are primarily taken in intertidal
    Department of Fisheries and Washington Department    regions to depths of <1.6 m (Wolotira et al. 1989).
    of Ecology 1985). Other important western U.S. areas
    include the southern waters of Puget Sound, Hood    IndicatorofEnvironmentalStress:Becauseofitsrelative
    Canal, Grays Harbor, Tillamook Bay, Yaquina Bay,    hardiness and abilitytoconcentrate contaminates, the
    Coos Bay, Humboldt Bay, Tomales Bay, Drakes Estero,   Pacific oyster has been used to indicate water quality

                                                       20

                                                                                         Pacific oystercontinued
                                                           cnidarians, polychaetes, molluscs, crustaceans, and
     Table 1. Relative abundance of Pacific oyster          bryozoans; many of these introduced species are
               in 32 U.S. Pacific coast estuaries.          predators or competitors with native species or are
                        Life Stage                         mariculture pests (Smith and Carlton 1975, Ricketts et
           Estuary    A S J L E                             al. 1985, Quayle 1988).  Pacific oysters appear to
            PugetSound �    �        Relative abundance:    successfully compete with the native oyster (Ostrea
             Hood Canal *�    a            Highly abundant  lurida), which is now restricted to typically deep low
              SkagltBay O    O         3  Abundant          salinity areas (Sayce 1976).
                                      O   Common
            Grays Harbor     a    o        Rare�
             Willapa Bay 6 �         Blank Not present      Range
          Columbia River                                   Overall: The Pacific oyster is a temperate species that
            Nehalem Bay                                     iS now found in southern Australia to New Zealand,
           Tillamook Bay *    *       Life stage:           Hawaii, Palau, along the Asian coast from China to the
             Netarts Bay  3   IN       A - Adults           southern Kuril Islands, and the North American coast
             Slletz River                Spawning adults   from southeast Alaska to northern Mexico (Morris
                                       J - Juveniles
            Yaqulna Bay *              L- Larvae            1966, Young 1966, Haro et al. 1981, Lee et al. 1981,
             Alsea River               E - Eggs             Quayle 1988, Wolotira et al. 1989). The Portuguese
            Siuslaw River                                   oyster (C. angulatus), which ranges from Portugal,
           Umpqua River O    O                              England, and southwest Europe, may be the same
              coos Bay �    �                              species (Menzel 1974, Wolotira et al. 1989).
             Rogue River
           Klamath River                                   Within Study Area: The Pacific oyster is found in most
           Humboldt Bay �    �                              Pacific coast estuaries from Morro Bay, California, to
               EelRiver                                     Skagit Bay, Washington, where estuarine physical
            Tomales Bay �    �                              conditions are appropriate and water pollution is not a
       Cent San Fran. Bay*   * Includes Central San         problem (Table 1). Pacific oysters were once cultured
       South San Fran. Bay   ~ a   Franciso, Suisun.        in San Francisco Bay and Elkhorn Slough, California,
           Elkhom Slough  '                                 but high pollution levels now make oysters from these
              MorroBay �    �                               areas unhealthy to consume (Frey 1971).  The
        Santa Monica Bay                                    Columbia, Rogue, Klamath, and Eel Riverestuaries do
           San Pedro Bay                                    not have oysters because salinities are not appropriate.
            Alamitos Bay
            Anaheim Bay                                     Life Mode
            Newport Bay                                     Eggs and early larval stages are pelagic. Late larval
             Mission Bay                                    stages are sedentary.  Juveniles and adults are
           San Diego Bay                                    sedentary and benthic/epibenthic (Quayle 1988).
          Tijuana Estuary
                       A S J L E                            Habitat
                                                           IyW: Eggs and larvae are estuarine/neritic, occurring
                                                            in the upper warmer waters of the watercolumn (Quayle
    problems in many estuaries. For example, antifouling    1988). Juveniles and adults are found in bays and
    paintscontainingcopperandtri-n-butyltincauseoyster    estuaries in lower intertidal areas to depths of 7 m
    shell thickening, alter growth rates, increase oxygen    below mean lower low water (Haderlie and Abbott
    consumption, and may affect larvae viability (Paul and    1980).
    Davies 1986, His and Robert 1987, Lawler and Aldrich
    1987, Quayle 1988). Presently, many estuarine areas    Substrate: Firm bottoms appear to be preferred;
    are closed to oyster culture and harvest because of    however, this species can be found on mud or mud-
    bacterial contamination commonly associated with    sandbottoms. Pacificoystersareusuallyfoundattached
    urban centers, marinas, and sewage outfalls (Cheney    to rocks, debris, or other oyster shells (Barrett 1963,
    and Mumford 1986).                                       Quayle 1988).

    Ecological: The Pacific oyster is the dominant bivalve    Physical/Chemical Characteristics: The Pacific oyster
    species in many estuarine areas where it is cultured.   is found in mesohaline-euhaline waters (usually 10-
    Many other "exotic" organisms were introduced in   35%,) (Barrett 1963, Berg 1971, Quayle 1988).  It
    Pacific coast estuaries along with Pacific and Virginia   tolerates air temperatures to -4�C during low tides and
    oysters (C. virginica). These exotics include sponges,    watertemperatures of 4-36�C (Quayle 1988, Wolotira

                                                        21

    Pacific oystercontinued
    et al. 1989), and spawns at water temperatures of 14-   notoccur annually. Therefore, spawning is sporadic or
    30�C, but only rarely below 18�C (Haderlie and Abbott    nonexistent in most estuaries (Span 1978, Ricketts et
    1980). Optimum spawning temperatures are probably   al. 1985, Quayle 1988). In California and other areas,
    21-23�C (Quayle 1988).  Larvae can survive water   Pacific oysters may spawn but the larvae may not
    temperatures of 17.5-35.0oC (Berg 1971), and 150C for   survive (Berg 1971, Haderlie and Abbott 1980, Ricketts
    a short time (Pauley et al. 1988). Larval setting is best   et al. 1985). Areas where successful reproduction
    at temperatures of 25 to 300C, salinities of 19 to 27%o,   does occur include: Pendrell Sound and the Strait of
    and on oyster shells that were first dipped in an aqueous    Georgia to Tofino Inlet on the west coast of Vancouver
    extract of oystertissue (Carlson 1981, Nell and Holliday   Island, Dabob Bay in Hood Canal, Washington, and
    1988). Adults will continue to feed down to 3�C, but   occasionally in Willapa Bay, Washington (Quayle 1988,
    growth stops when temperatures drop below 10�C    Wolotira et al. 1989). Eggs are not released into the
    (Barrett 1963, Quayle 1988). Best conditions for somatic   exhalant siphon like manyother bivalves, but discharged
    growth are 17�C (ranges 15-18�C), salinities >24'%   into the suprabranchial chambers, passedthrough the
    (ranges10-35%0),foodsuspensionsof120 mg/l(ranges   gills into the mantle chamber, and then expelled by
    24-550 mg/l), oxygen levels above 70%, suspended    contraction of the adductor mussel. Eggs may travel
    sediments between 0.0 and 8.0 mg/I, and pH levels   30 cm or more when discharged. Females release
    above 7.8 (Bernard 1983b, Brown and Hartwick 1988a).   eggs 5-10 times/minute, while the males release a
    Growth rates correlate primarily with suspended    continuous stream of sperm through their exhalant
    particulate organic material levels and secondarily with   siphons (Quayle 1988).
    temperature, but are mediated by salinity (Malouf and
    Bresse 1977, Brown 1988, Brown and Hartwick 1988b).    Fecundity: Fecundity ranges from 10 million to 200
    Paralytic shellfish poisoning can be a problem when    million eggs per female, with fecundity increasing with
    oysters feed on the dinoflagellate Protogonyaulax   age (Frey 1971, Wolotira et al. 1989). The average
    acatanella, but they quickly lose theirtoxicity when the   market-sized oyster produces 50-100 million eggs/
    dinoflagellate bloom is gone. (Haderlie and Abbott    year(Quayle 1988). Individuals mayspawnrepeatedly
    1980, Quayle 1988). Embryos are very sensitive to   during a spawning season (Haderlie and Abbott 1980,
    zinc and other metals (Boyden et al. 1975).           Quayle 1988).

    Miarations and Movements: Planktonic eggs and larvae   Growth and Development
    are moved bywatercurrents. Late-stage larvae settle    Eaa Size and Embryonic Development: Eggs are
    out of the water column and crawl on the bottom    spherical and 0.05 mm in diameter (Quayle 1988).
    searching for suitable substrates before finally setting    Embryonic development is indirect and external.
    (Quayle 1988). Juveniles and adults are sedentary
    and usually become firmly attached to materials on the    Aae and Size of Larvae: Fertilized eggs develop into
    bottom (Quayle 1988).                                 veliger larvae in 24-48 hours depending on temperature
                                                           (Cahn 1950, Quayle 1988). Larvae arefree-swimming
    Reproduction                                         for 2-4 weeks depending on temperature (Haderlie and
    Mode: The Pacific oyster is gonochoristic (some    Abbott 1980, Strathmannetal. 1987). Then they settle
    hermaphroditism occurs) and a batch spawner,    on to substrates and metamorphose into spat (Quayle
    broadcasting its gametes and relying on external   1988). Larvae range in size from 0.06 to 1.32 mm
    fertilization (Berg 1969, Haderlie and Abbott 1980).    (Wolotira et al. 1989); they are 0.27-0.31 mm long at
    Thisspecies isaprotandrichermaphrodite, developing    settlement (Strathmann et al. 1987). They will grow
    first as a male and later changing to a female (Quayle   from 0.075 mm to about 0.3 mm in about a month at 18
    1988). Sex appearsto be influenced byenvironmental    to 24�C (Quayle 1988).
    conditions, with some females becoming males when
    the food supply is low and males becoming females    Juvenile Size Ranae: Juvenile sizes range from about
    when food is abundant (Quayle 1988).                  0.30 mm to 40.0 mm. Size depends on tidal height,
                                                           area of settlement, and other factors (Quayle 1988).
    Matina/SDawnina: Spawning is initiated by a rise in
    water temperatures (usually above 18�C) or by    Aae and Sizeof Adults: The Pacificoystermaymature
    hormones released from the sperm of other oysters   in 1 year and may be as small as 30 mm shell length
    (Quayle 1988, Wolotira et al. 1989). This species    (Wolotiraetal. 1989). Adults growto 10-1 2cm (market
    spawns from June to September (primarily July to   size) in2to3 years in California'swaters, but maygrow
    August) during high tide (Quayle 1988).  Minimum    for 20 years or more (Haderlie and Abbott 1980). In
    threshold spawningtemperatures are notoften reached    Oregon and southern Washington, 2-4 years are
    in many Pacific coast estuaries, or if they are, they do    required to grow to market size; 4-6 years' growth is

                                                        22

                                                                                        Pacific oyster continued
    required in northern Washington, British Columbia,    Siltation and increased turbidities of oyster beds
    and Alaska (Pauley et al. 1988). This species may    resulting from logging, upland alterations, and natural
    grow to 25.4 cm in shell length, but most are 10.2-12.7    causes can result in high mortalities (Pauley et al.
    cm (Pauley et al. 1988). Shell growth and shape are    1988, Quayle 1988). In northern latitudes, icecan push
    highlyvariable, dependingontemperature, food supply,    them into sediments.  In areas of high population
    culture method, and other factors (Cahn 1950, Quayle    densities, food may be a limiting factor (Pauley et al.
    1988).                                                  1988). Diseases, algal blooms that inhibit feeding, bay
                                                          ghost shrimp (Callianassa californiensis), and blue
    Food and Feeding                                        mud shrimp (Upogebia pugettensis) can also reduce
    Trophic Mode: Juveniles and adults are detritivores,    population sizes.  In the 1960s and 1970s, mass
    nannoplanktivores, and suspension feeders (Haderlie    mortalities of older (>2 years old) Pacific oysters
    and Abbott 1980, Quayle 1988). Food is taken in the    occurred in Washington and California during late
    inhalant siphon, filtered and collected by mucus on the    summer when water temperatures approached or
    gills, sorted on the palps, and transferred to the mouth.    exceeded 200C. The cause of this mortality was never
                                                           positively identified, but infection by Vibrio spp. and
    Food Items: Larvae feed on naked flagellates (Berg    variability in the oyster's carbohydrate cycle were
    1971).   Juveniles and  adults eat primarily    implicated (Beattie et al. 1981, Elston et al. 1987,
    nannoplankton, such as bacteria, dinoflagellates,    Pauley et al. 1988). However, environmental stresses
    flagellates, diatoms, and algal and invertebrategametes    such as prolonged  air exposure times, warm
    (Barrett1963,Quayle1988). Theyalsoconsumeplant    temperatures, and dinoflagellate blooms may have
    and animal detritus, but the importance of this material    promoted mortality of already stressed oysters (Pauley
    to their diet is unknown (Barrett 1963, Quayle 1988).    et al. 1988). Other estuarine species reduce Pacific
                                                           oyster growth or indirectly affect oyster viability. Mud
    Biological Interactions                                 and ghost shrimp cause serious damage to oyster
    Predation: Larvae are eaten by numerous predators    beds by making grounds too soft for culture or by
    including: Tintinnidae and other ciliates, ctenophores,    smothering them. This has required the controversial
    jellyfish (Aurelia aurita and Chrysaora melanaster),    use of the insecticide SEVIN (carbaryl) to reduce
    oysters, barnacles, Pacific herring (Clupea pallasi),    shrimp populations (Washington  Department of
    and smelt (Berg 1971).  The introduced flatworm    Fisheriesand Washington Department of Ecology 1985,
    (Pseudostylochus ostreophagus) can be a major    Quayle 1988).  Other harmful organisms include
    predator of oyster spat (Quayle 1988). Predators of   protozoa, bacterial diseases, sponges, flatworms,
    juveniles and adults include crabs (C. magister, C    polychaetes, and a parasitic copepod (Mytilicola
    productus, and C. gracilis), oyster drills (Ceratostoma    orientalis) (Dungan and Elston 1988, Quayle 1988).
    inornatum and Urosalpinx cinerea), starfish (Pisaster   Fouling organisms such as mussels, tunicates, algae,
    ochraceus, P. brevispinus, Evasterias troschelii, and    sponges, anemones, hydroids, and bryozoans may
    Pycnopodia helianthoides), and ducks (Aythya affinis),    compete with oysters for food, reduce oyster growth
    and surf and white winged scoters (Mellanita spp.).   rates, and affect spat settlement (Quayle 1988). The
    Important fish predators of juvenile and adult oysters in   Pacific oyster's chief enemy is man, who by dredging
    California include the bat ray (Myliobatis californica)   activities and pollution, reduces areas where viable
    and angel shark (Squatina californica) (Haderlie and    oysterproductioncan occur (Wallace 1966, Ricketts et
    Abbott 1980, Ricketts et al. 1985).                     al. 1985). Forexample, sulfite liquor effluentfrom pulp
                                                           mills in the Pacific Northwest appears to affect survival
    Factors Influencina PoDulations: Probably the most    and growthof alloyster life stages (Cheneyand Mumford
    important factor limiting Pacific oyster populations on    1986). Because of pollution, many bays and estuaries
    the Pacific coast is low water temperatures which    once used for oystering are now closed or restricted
    inhibit spawning.  In areas where they do spawn,    (Gunn and Saxby 1981, Qualman 1981, Cheney and
    Pacific oyster larvae often do not survive and set,    Mumford 1986).
    except in a few warm bays whenconditions are optimal.
    Mortality of larvae may be due to low temperatures,    References
    excessive turbidity, lack of food, toxins from
    dinoflagellateblooms, predation, andbacterialorfungal    Bardach, J. E., J. H. Ryther, and W. O. McLarney.
    diseases (Berg 1971).  Juveniles may be killed by    1972.  Aquaculture: The farming and husbandry of
    abruptchangesinsalinityandtemperature. Adults and    freshwater and marine organisms.  John Wiley and
    juvenile populations are affected by storms and    Sons, Inc., New York, NY, 868 p.
    associated waves that can displace individuals and
    bury them in sediments (Cheney and Mumford 1986).

                                                       23

    Pacific oyster continued
    Barrett, E. M.  1963. The California oyster industry.    and seaweed harvests of Puget Sound. Wash. Sea
    Calif. Fish Game, Fish Bull. 123, 103 p.                Grant, Univ. Wash. Press, Seattle, WA, 164 p.

    Beattie, J. H., D. McMillin, and L. Wiegardt. 1981. The    Conte, F. S., and J. L. Dupuy. 1981. The California
    Washington State oyster industry: a brief overview. In   oyster industry. In K. K. Chew (editor), Proceedings of
    K.K. Chew(editor),ProceedingsoftheNorthAmerican    the North American oyster workshop, p. 43-63.
    oyster workshop, p. 28-38.  Louisiana State Univ.,    Louisiana State Univ., Baton Rouge, LA.
    Baton Rouge, LA.
                                                            Dungan,  C.  R.,  and  R.  A.  Elston.    1988.
    Berg, C. J., Jr. 1969. Seasonal gonadal changes of    Histopathological and ultrastructural characteristics of
    adult oviparous oysters in Tomales Bay, California.   bacterial destruction of the hinge ligaments of cultured
    Veliger 12:27-36.                                       juvenile Pacific oysters, Crassostrea gigas. Aquacul.
                                                            72:1-14.
    Berg, C. J., Jr. 1971. A review of possible causes of
    mortality of oyster larvae of the genus Crassostrea in   Elston, R. A., J. H. Beattie, C. Friedman, R. Hedrick,
    Tomales Bay, California. Calif. Fish Game 57(1):69-    and M. L. Kent. 1987. Pathology and significance of
    75.                                                     fatal inflammatory bacteraemia in the Pacific oyster,
                                                            CrassostreagigasThunberg. J. Fish. Diseases 10:121 -
    Bernard, F. R. 1983a. Catalogue of the living Bivalvia    132.
    of the eastern Pacific Ocean: Bering Strait to Cape
    Horn. Can. Spec. Publ. Fish. Aquat. Sci. 61, 102 p.      Fitch, J. E. 1953. Common marine bivalves of California.
                                                            Calif. Fish Game, Fish Bull. 90, 102 p.
    Bernard, F. R. 1983b. Physiology and the mariculture
    of some northeastern Pacific bivalve molluscs. Can.    Frey, H. W. 1971. California living marine resources
    Spec. Publ. Fish. Aquat. Sci. 63, 24 p.                  and their utilization.  Calif. Dept. Fish Game,
                                                            Sacramento, CA, 148 p.
    Boyden, C. R., H. Watling, and I. Thorton. 1975. Effect
    of zinc on the settlement of the oyster Crassostrea    Gates, D. E., and H. W. Frey.  1974.  Designated
    gigas. Mar. Biol. 31:227-234.                            common  names of certain marine organisms of
                                                            California. Calif. Fish Game, Fish Bull. 161:55-90.
    Brown, J. R. 1988. Multivariate analyses of the role of
    environmental factors in seasonal and site-related    Gunn, C. R., and D. J. Saxby. 1981. A brief history of
    growth variation inthe Pacific oyster Crassostreagigas.    the oyster industry in British Columbia. In K. K. Chew
    Mar. Ecol. Prog. Ser. 45:225-236.                        (editor), Proceedings of the North American oyster
                                                            workshop, p. 17-27. Louisiana State Univ., Baton
    Brown, J. R., and E. B. Hartwick. 1988a. A habitat    Rouge, LA.
    suitability index model for suspended tray culture of the
    Pacific oyster, Crassostreagigas Thunberg. Aquacul.    Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: The
    Fish. Manag. 19:109-126.                                clams and allies. In R. H. Morris, D. P. Abbott, and E.
                                                            C. Haderlie (editors), Intertidal invertebrates of
    Brown, J. R., and E. B. Hartwick. 1988b. Influence of   California, p. 355-411. Stanford Univ. Press, Stanford,
    temperature, salinity and available food upon    CA.
    suspended culture of the Pacific oyster, Crassostrea
    gigas:l1. Absolute and allometric growth.  Aquacul.    Haro, B. H., E. P. Nunez, A. F. Mattus, and M. A.
    70:231-251.                                              Landin. 1981. The development and perspective of
                                                            oyster culture in Mexico. In K. K. Chew (editor),
    Cahn, A. R. 1950. Oysterculture in Japan. Fish. Wildl.    Proceedings of the North American oyster workshop,
    Serv., Fish. Leafit. 383, 80 p.                          p. 64-69. Louisiana State Univ., Baton Rouge, LA.

    Carlson, B. L. K. 1981 Effects of temperature, salinity,    Hedgpeth, J. W., and S. Obrebski. 1981. Willapa Bay:
    feeding, substrate, and storage on the setting and    a historical perspective and a rationale for research.
    survival of commercially-reared eyed larvae of the    U.S. Fish Wildl. Serv., FWS/OBS-81/03, 52 p.
    Pacific oyster, Crassostreagigas. M.S. Thesis, Oregon
    State Univ., Corvallis, OR, 90 p.                        His, E., and R. Robert. 1987. Comparative effects of
                                                           two antifouling paints on the oyster Crassostrea gigas.
    Cheney, D. P., and T. F. Mumford, Jr. 1986. Shellfish    Mar. Biol. 95:(1):83-86.

                                                        24

                                                                                         Pacific oyster continued

    Lawler, I. F., and J. C. Aldrich. 1987. Sublethal effects    Smith, R. I., and J. T. Carlton (editors). 1975. Light's
    of Bis (tri-n-butyltin) Oxide on Crassostrea gigas spat.    manual: Intertidal invertebratesof the central California
    Mar. Poll. Bull. 18(6):274-278.                         coast. Univ. Calif. Press, Berkeley, CA, 716 p.

    Lee, K. W. F., J. S. Corbin, and W. A. Brewer. 1981.    Span, J. A. 1978. Successful reproduction of giant
    Overview of oyster culture in Hawaii and various U.S.    Pacific oysters in Humboldt Bay and Tomales Bay,
    Pacific island territories.  In K. K. Chew (editor),   California. Calif. Fish Game 64(2):123-124.
    Proceedings of the North American oyster workshop,
    p. 70-85. Louisiana State Univ., Baton Rouge, LA.       Strathmann, M. F., A. R. Kabat, and D. O'Foighil. 1987.
                                                           Phylum Mollusca, class Bivalvia. In M. F. Strathmann
    Malouf, R. E., and W. P. Breese.  1977.  Food    (editor), Reproduction and development of marine
    consumption and growth of larvae of the Pacific oyster,    invertebrates of the northern Pacific coast, p. 309-353.
    Crassostrea gigas  (Thunberg), in a constant flow    Univ. Wash. Press, Seattle, WA.
    rearing system. Proc. Natl. Shellfish. Assoc. 67:7-16.
                                                           Wallace, D. H. 1966. Oysters in the estuarine
    Menzel, R. W.  1974.  Portuguese and Japanese    environment. In Symposium on estuarinefisheries, p.
    oysters are the same species. J. Fish. Res. Board Can.    68-73. Am. Fish. Soc. Spec. Publ. No. 3., Am. Fish.
    31:453-456.                                             Soc., Bethesda, MD.

    Morris, P. A. 1966. Afield guideto Pacificcoast shells.   Washington Department of Fisheries.  1986.  1986-
    Houghton-Mifflin Co., Boston, MA, 297 p.                1987 salmon, shellfish, bottom fish sport fishing guide.
                                                           Wash. Dept. Fish., Olympia, WA, 20 p.
    Nell, J. A., and J. E. Holliday. 1988. Effects of salinity
    and the growth and survival of Sydney rock oyster    Washington Department of Fisheries and Washington
    (Saccostrea commercialis) and Pacific oyster    Department of Ecology. 1985. Use of the insecticide
    (Crassostrea gigas) larvae and spat. Aquacul. 68:39-    SEVIN to control ghost and mud shrimp in oyster beds
    44.                                                     of Willapa Bay and Grays Harbor. Final Env. Impact
                                                            Statament, Wash. Dept. Fish. and Wash. Dept. Ecol.,
    Paul, J. D., and I. M. Davies. 1986. Effects of copper    Olympia, WA, 64 p plus appendices
    and tin-based anti-fouling compounds on the growth of
    scallops (Pecten maximus) and oysters (Crassostrea    Wolotira, R. J., Jr., M. J. Allen, T. M. Sample, C. R. Iten,
    gigas). Aquacult. 54:191-203.                           S. F. Noel, and R. L. Henry. 1989. Life history and
                                                            harvest summaries for selected invertebrate species
    Pauley, G. B., B. Van Der Raay, and D. Troutt. 1988.    occurring off the west coast of North America. Volume
    Species profiles: life histories and environmental    1: shelled molluscs. NOAA Tech. Memo,  NMFS F/
    requirementsofcoastalfishesand invertebrates (Pacific    NWC-160,177 p.
    Northwest)-Pacific oyster. U.S. Fish Wildl. Serv. Biol.
    Rep. 82(11.85), U.S. Army Corps Eng., TR EL-82.4,    Younge, C. M. 1966. Oysters, 2nd edition. Collins,
    28 p.                                                   London, 209 p.

    Qualman, J. L. 1981. Oregon's oyster industry. In K.
    K. Chew (editor), Proceedings of the North American
    oyster workshop, p. 39-42. Louisiana State Univ.,
    Baton Rouge, LA.

    Quayle, D. B. 1988. Pacific oyster culture in British
    Columbia. Can. Bull. Fish. Aquat. Sci. 218, 214 p.

    Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W.
    Phillips. 1985. Between Pacific tides. Stanford Univ.
    Press, Stanford, CA, 652 p.

    Sayce, C. S. 1976. The oyster industry of Willapa Bay.
    In Proceedings of the symposium on terrestrial and
    aquatic ecological studies of the Northwest, p. 347-
    356. Eastern Wash. State College, Cheney, WA.

                                                        25

        Tresus capax
       Adult

                                         5cm

    Common Name: horseneck gaper                          et al. 1989).
    Scientific Name: Tresus capax
    Other Common Names: Alaskan gaper, fat gaper,    Recreational: The horseneck gaper is harvested
    blue clam, empire clam, gaper, gaper clam, greyneck    recreationally from Humboldt Bay, California, to Puget
    clam, horseneckclam, horseclam, bigneckclam,giant    Sound, Washington (Machell and DeMartini 1971,
    rockdweller, butter clam, money shell, giant saxidome    Wolotira et al. 1989). No more than 10/day can be
    (Morris 1966, Gates and Frey 1971, Haderlie and    taken in California (Ricketts et al. 1985),12/day in
    Abbott 1980, Wolotira et al. 1989)                    Oregon (Oregon Department of Fish and Wildlife 1976),
    Classification (Bernard 1983a)                        and 7/day in Washington (Washington Department of
    Phylum: Mollusca                                      Fisheries 1986).  It is harvested primarily by hand
    Class: Bivalvia                                       (using shovels, rakes, etc.) during low tides.
    Order: Veneroida
    Family: Mactridae                                     Indicator of Environmental Stress: Clam beds are
                                                           sometimes closed to harvest because of paralytic
    Value                                                 shellfish poisoning or coliform bacterial contamination.
    Commercial: This species andthe Pacific gaper ( Tresus   As a result of pollution in Washington waters, over 25%
    nuttallil) are harvested commercially from northern   of the potential areas for subtidal hardshell clam
    California to British Columbia (landings are not   harvesting are closed (Schink et al. 1983).
    separated by species) (Wolotira et al. 1989). It istaken
    both subtidally and intertidally using hydraulic pumps,    Ecological: The horseneck gaper is often the largest
    mechanical dredges, potato forks, shovels, and clam    subtidal and intertidal suspension/filterfeeding bivalve
    rakes (Frey 1971, Wolotira et al. 1989). Recent harvests   in many Pacific coast estuaries (Hancock et al. 1979).
    have averaged about 225 t annually, placing them fifth
    in volume for the entire U.S. and Canada Pacific coast    Range
    clam harvest (Wolotira et al. 1989). This species is   Overall: This species' overall range is from Monterey,
    taken year-round, but most are harvested from July to   California, to Kodiak, Alaska and the mouth of Prince
    December in British Columbia and Oregon (Wolotira et   William Sound, Alaska.  It is uncommon south of
    al. 1989). Althoughthehorseneckgaperisalargeclam    Humboldt Bay, where it is replaced by T. nuttallii
    that provides excellent meat for chowder or clam    (Bernard 1983a, Rudy and Rudy 1983, Wolotira et al.
    steaks, it is not often sold fresh. Instead, it is usually    1989).
    canned because it has a fragile shell that breaks easily
    and its valves gape, reducing shelf life and allowing   Within Study Area: The horseneck gaper is found from
    water loss. Also, a tough outer covering on its neck    Humboldt Bay to Puget Sound, reaching highest
    increases processing/packaging time and meat yield   abundances in Coos and Siuslaw Bays, Oregon (Table
    during processing is low (25-30% of total body weight)    1). It is rare from Humboldt Bay south to San Francisco
    (Quayleand Bourne 1972, Ricketts et al. 1985, Wolotira    Bay, California, and is not found in any estuaries further

                                                        26

                                                                                     horseneck gaper continued

                                                          MLLW (Wendell et al. 1976, Goodwin and Shaul 1978,
     Table 1. Relative abundance of horseneck gaper      Cheney and Mumford 1986).
              in 32 U.S. Pacific coast estuaries.
                        Life Stage                        Substrate: The horseneck gaper is found primarily in
           Estuary    A S J L E                          substrates consisting of shell fragments and dense
            PugetSound   ) 0  ) O O   Relative abundance:  sand, as well as silty-sand and gravel (Bourne and
             Hood Canal di 3 l  (3   *   Highly abundant  Smith 1972b,Wendelletal. 1976, Cheneyand Mumford
              SkagitBay  C O    0i    0     Abundant      1986). InHumboldtBay,clamdensitiesaregreatestin
            Grays Harbor     0 a o    O 0   Common        silty-sand substrates covered with eelgrass (Zostera
                          Graysror  (~] 0  ORare
             WillapaBay O O   0     0   Blank Not present  spp.) (Wendell 1973).  Sediment structure affects
          Columbia River                                 burrowing depth; clams burrow deeper in mud and
            Nehalem Bay  i    i                           sand substratesthan in clay substrates (Oceanographic
           Tillamook Bay  3 �  ] 13 �   Life stage:      Institute of Washington 1981).
             Netarts Bay 3  13  � �   X    A-Adults
                                      S - Spawning adults
             Siletz River             - JuveSpawning adults  Phvsical/Chemical Characteristics: Juveniles and adults
            Yaquina Bay  i,          L- Larvae           arefound in polyhaline-euhalinewaters, attemperatures
                                      E - Eggs
             AlseaRiver O CO  O C                         of 2-20�C (Bernard 1983a). Larvae do not survive at
            SiuslawRiver � � � � �                        20�C (Bourne and Smith 1972a). Optimum conditions
           UmpquaRiver i                ' / i i          forsomaticgrowthare13�Cwatertemperatures (range
               Coos Bay *  �       � �                    11-180C), 28%0 salinities (range 26-31%o), and food
             Rogue River                                  suspension density of 95 mg/I (range 15-200 mg/l)
            Klamath River                                 (Bernard 1983b).
            Humboldt Bay  i 3     )
               Eel River                                  Miarations and Movements: Eggs and larvae are
            Tomales Bay                                   dispersed by currents. Juveniles and adults do not
       Cent San Fran. Bay'         ' Includes Central San  move laterally once they become established. Clams
       South San Fran. Bay Francisco, Suisun.
           SlougthSanFran.ay        and San Pablo bays.   older than two years (77 mm shell length) lose the
           Elkhom Slough                                  ability to reburrow (Wendell et al. 1976).
              Morro Bay
         Santa Monica Bay                                 Reproduction
           San Pedro Bay                                  Mode: The horseneckgaper is gonochoristic, oviparous,
            Alamitos Bay                                  and iteroparous. It is a broadcast spawner, hence eggs
            Anaheim Bay          E                        are fertilized externally (Bourne and Smith 1972b).
            Newport Bay
             Mission Bay
             San Diego Bay                                Matina/Soawnina: Spawning begins when waters warm
           San Diego Bay
           Tijuana Estuary                                afterthe seasonal minimum (Bourne and Smith 1972b,
                       A S J L E                          Cheney and Mumford 1986), usually late winterto early
                                                           spring. In British Columbia and Puget Sound, spawning
                                                          occurs from February-May, peaking primarily in March
    south than San Francisco Bay. It is not found in many    (Bourne and Smith 1972b). In California and Oregon,
    small estuaries or estuaries with relatively high river   spawning occurs from January-March, peaking in
    flows (e.g., Oregon's Columbia, Siletz, and Rogue    February (Machell1968, Machelland DeMartini 1971,
    Rivers, and California's Klamath and Eel Rivers).     Breed-Willeke and Hancock 1980, Robinson and
                                                           Breese 1982). The horseneck gaper may spawn more
    Life Mode                                             than once during the spawning season (Bourne and
    Eggs and larvae are pelagic. Juveniles and adults are    Smith 1972b)
    benthic infau na, burrowing into sediments to depths 1l
    m, but usually 25-50 cm (Cheney and Mumford 1986,    Fecundity: Unknown.
    Wolotira et al. 1989).
                                                           Growth and Development
    Habitat                                                Eaa Size and Embrvonic DeveloDment: Eggs are
    Type: Eggs and larvae are neritic. Juveniles and adults   spherical and 0.06-0.07 mm in diameter (Bourne and
    are found primarily in bays and estuaries, occurring    Smith 1972a). Embryonic development is indirect and
    from mid-tide levels (+2 m) down to 30 m below mean    external; after fertilization, polar bodies form within 40
    lower lowwater (MLLW). In Puget Sound and Humboldt    minutes, trochophoresformwithin 24 hours, and veligers
    Bay, they are most abundant at depths 1-5 m below    by 48 hours.

                                                       27

    horseneckgapercontinued
    Aae and Size of Larvae: Larvae range from 0.06-0.07    horseneck gapers burrow deeper, escaping many
    mm to 0.26-0.27 mm in diameter (Boume and Smith    physical and biological stresses. Recruitment may be
    1972a). Metamorphosisto spattakes 24days at 15�C,    highly variable on some clam beds, resulting in beds
    26 days at 10�C, and 34 days at 5�C (Bourne and Smith    dominated by only one or two age classes (Wendell et
    1972a). Larval settlement occurs primarily between    al. 1976, T. Gaumer, Oregon Department of Fisheries,
    early spring and summer.                              Newport, OR, pers. comm.).  In general, intertidal
                                                           populations of this species are affected by numerous
    Juvenile Size Ranoe: Juveniles range in size from    afterationsanddisturbances,including:siltation,storms,
    0.26-0.28 mmto about 70 mm shell length (Bourne and    freshwater runoff, floods, erosion, dredging, and marina
    Smith 1972a, 1972b). They may grow to 2.54 cm after   development (Schink et al. 1983). Diseases may also
    1 winter (Quayle and Bourne 1972).  Most growth    affect horseneck gaper populations (Wendell 1973,
    occurs during the spring and summer when    Armstrong and Armstrong 1974); it is often infected
    phytoplankton is abundant (Wendell et al. 1976, Haderlie   with a haplosporidan parasite (43% in Yaquina Bay,
    and Abbott 1980).                                     Oregon) (Armstrong and Armstrong 1974). Two species
                                                          of pinnotherid crabs (Pinnixa faba and P. fittoralis) are
    Aae and Size of Adults: Size appears to determine    known to inhabitthe mantlecavityofhorseneckgapers
    maturity; most horseneck gapers mature at about 70    (Pearce 1965, Stout 1967), but apparently cause little
    mm shell length (SL) (Bourne and Smith 1972b). In   harm to the clam (Haderlie and Abbott 1980).
    British Columbia, this takes four years, but only three
    years in California and Oregon (Bourne and Smith    References
    1972b, Wendell et al. 1976, Hancock et al. 1979). In
    Oregon, subtidal clams between the ages of four and    Armstrong, D. A., and J. L. Armstrong.  1974.  A
    seven years grow fasterthan intertidal clams of similar   haplosporidan infection in gaper clams, Tresus capax
    ages(Hancocketal. 1979). The horseneck gaper can    (Gould), from Yaquina Bay, Oregon.  Proc. Natl.
    live to 16 years and can reach 254 mm SL (Morris 1966,    Shellfish. Assoc. 64:68-72.
    Bourne and Smith 1972b). The oldest clams found in
    Oregon were 10-12 years old (Hancock et al. 1979).    Bernard, F. R. 1983a. Catalogue of the living Bivalvia
                                                           of the eastern Pacific Ocean: Bering Strait to Cape
    Food and Feeding                                      Horn. Can. Spec. Fish. Aquat. Sci. 61,102 p.
    Trophic Mode: Juveniles and adults are suspension/
    filterfeeders (HaderlieandAbbott 1980). Food particles    Bernard, F. R. 1983b. Physiology and the mariculture
    travel in water through the inhalant siphon and are   of some northeastern Pacific bivalve molluscs. Can.
    collected on the gills, sorted by the palps, and passed    Spec. Publ. Fish. Aquat. Sci. 63, 24 p.
    to the mouth. Energy reserves are stored as glycogen
    in the gonads and as fat (Reid 1969).                 Bourne, N., and D. W. Smith. 1972a. The effect of
                                                           temperature on the larval development of the horse
    Food Items: Juveniles and adults feed on suspended    clam, Tresus capax (Gould). Proc. Natl. Shellfish.
    diatoms, flagellates, dinoflagellates, and fine detritus,   Assoc. 62:35-37.
    including small eelgrass (Z. marina) particles (Stout
    1967, Haderlie and Abbott 1980).                      Bourne, N., and D. W. Smith. 1972b. Breeding and
                                                           growth of the horse clam, Tresus capax (Gould), in
    Biological Interactions                               southern British Columbia. Proc. Natl. Shellfish. Assoc.
    Predation: Eggs and larvae are probably preyed on by    62:38-46.
    many planktivorous organisms. Predators of juveniles
    include:worms,snails, crustaceans, andcopperrockfish   Breed-Willeke, G. M., and D. R. Hancock.  1980.
    (Sebastes caurinus) (Wolotira et al. 1989). Common    Growth and reproduction of subtidal population of the
    predators of juveniles and adults include moon snails   gaper clam Tresus capax (Gould) from Yaquina Bay,
    (Polinices spp.), Dungeness crab (Cancermagister),    Oregon. Proc. Natl. Shellfish. Assoc. 70:1-13.
    bat ray (Myliobatis californica), and sea stars (Pisaster
    spp.) (Haderlie and Abbott 1980).                     Cheney, D. P., and T. F. Mumford, Jr. 1986. Shellfish
                                                           and seaweed harvests of Puget Sound. Wash. Sea
    Factors Influencino PoDulations: Predation can cause    Grant, Univ. Wash. Press, Seattle, WA, 164 p.
    very high mortalities on some clam beds (Haderlie and
    Abbott 1980).  High mortality of small juveniles is   Frey, H. W. 1971. California's living marine resources
    probably due to low salinities, temperature stress and    and their utilization. Calif. Fish Game, Sacramento,
    predation (Wendell et al. 1976).  As they grow,    CA, 148 p.

                                                       28

                                                                                     horseneck gaper continued
    Gates, D. E., and H. W. Frey.  1971.  Designated    Robinson, A. M., and W. P. Breese.  1982.  The
    common  names of certain marine organisms of    spawning season of four species of clams in Oregon.
    California. Calif. Fish Game, Fish Bull. 161:55-90.     J. Shellfish Res. 2(1):55-57.

    Goodwin, L., and W. Shaul.  1978.  Puget Sound    Rudy, P.,Jr., and L. H. Rudy. 1983. Oregon estuarine
    subtidal hardshell clam survey data. Prog. Rep. 44,    invertebrates - An illustrated guide to the common and
    Wash. Dept. Fish., Olympia, WA, 92 p.                   important invertebrate animals. U.S. Fish Wildl. Serv.,
                                                          Biol. Serv. Prog., FWS/OBS-83/16, Portland, OR,
    Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: The    225 p.
    clams and allies. In R. H. Morris, D. P. Abbott, and E.
    C. Haderlie (editors), Intertidal invertebrates of    Schink, T. D., K. A. McGraw, and K. K. Chew. 1983.
    California, p. 355-411. Stanford Univ. Press, Stanford,    Pacific coast clam fisheries. Wash. Sea Grant, Univ.
    CA.                                                     Wash., Seattle, WA, 72 p.

    Hancock, D. R., T. F. Gaumer, G. B. Willeke, G. P.   Stout, W. E. 1967. A study of the autecology of the
    Robart, and J. Flynn. 1979. Subtidal clam populations:    horse neck clams Tresus capax and Tresus nuttallii in
    distribution, abundance, and ecology.  Oregon Sea    South Humboldt Bay, California. M.A. Thesis, Humboldt
    Grant Publ. No. ORESU-T-79-002.  Oregon State    State Univ., Arcata, CA, 51 p.
    Univ., Corvallis, OR, 243 p.
                                                          Washington Department of Fisheries. 1986. 1986-
    Machell, J. R. 1968. The reproductive cycle oftheclam    1987 (April 1 thru March 31) salmon, shellfish, bottom
    Tresus capax (Gould, 1850), Family Mactridae, in   fish sport fishing guide. Wash. Dept. Fish., Olympia,
    south Humboldt Bay, California. M.A. Thesis, Humboldt    WA, 20 p.
    State Univ., Arcata, CA, 28 p.
                                                           Wendell, F. E. 1973. Ecology of the gaper clam,
    Machell, J. R., and J. D. DeMartini. 1971. An annual    Tresus capax  (Gould, 1850) in Humboldt Bay,
    reproductive cycle of the gaper clam, Tresus capax    California. M.S. Thesis, Humboldt State Univ., Arcata,
    (Gould), in southern Humboldt Bay, California. Calif.    CA, 37 p.
    Fish Game 57(4):274-282.
                                                           Wendell, F., J. D. DeMartini, P. Dinnel, and J. Siecke.
    Morris, P. A. 1966. Afield guideto Pacificcoast shells.    1976. The ecology of the gaper or horse clam, Tresus
    Houghton-Mifflin Co., Boston, MA, 297 p.                capax (Gould 1850) (Bivalvia: Mactridae) in Humboldt
                                                           Bay, California. Calif. Fish Game 62(1):41-64.
    Oceanographic Institute of Washington. 1981. Clam
    and mussel harvesting industries in Washington State.   Wolotira, R. J., Jr., M. J. Allen, T. M. Sample, C. R. Iten,
    Oceanog. Comm.  Wash., Seattle, WA, various    S. F. Noel, and R. L. Henry. 1989. Life history and
    pagination.                                             harvest summaries for selected invertebrate species
                                                           occurring off thewestcoastof NorthAmerica. Volume 1:
    Oregon Department of Fish and Wildlife.  1976.    Shelled molluscs. NOAATech. Memo. NMFS F/NWC-
    Oregon's captivating clams. Corvallis, OR.              160, 177 p.

    Pearce, J. B. 1965. On the distribution of Tresus
    nuttalliland Tresuscapaxinthe waters of Puget Sound
    and the San Juan Archipelago. Veliger 7(3):166-170.

    Quayle, D. B., and N. Bourne. 1972. The clam
    fisheries in British Columbia. Fish. Res. Board Can.,
    Bull. No. 179, 70 p.

    Reid, R. G. B. 1969. Seasonal observations on diet,
    and stored glycogen and lipids in the horse clam,
    Tresus capax (Gould, 1850). Veliger 11 (4):378-381.

    Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W.
    Phillips. 1985. Between Pacific tides. Stanford Univ.
    Press, Stanford, CA, 652 p.

                                                       29

        Tresus nuttallii
       Adult

                                        5cm

    Common Name: Pacific gaper                            1971, Wolotira et al. 1989). It is taken year-round, but
    Scientific Name: Tresus nuttallii                     most are harvested from July to December in British
    Other Common Names: Washington clam, big-neck    Columbia (Wolotira et al. 1989).
    clam, blue clam, empire clam, gaper clam, great
    horseneck clam, otter-shell clam, rubberneck clam,    Recreational: The Pacific gaper is an important
    summer clam (Wolotira et al. 1989)                    recreational species in Puget Sound, Washington, and
    Classification (Bernard 1983)                         in California estuaries, including Humboldt Bay,
    Phylum: Mollusca                                      Tomales Bay, Bodega Bay, Drakes Estero, Bolinas
    Class: Bivalvia                                       Lagoon, Elkhorn Slough, and Morro Bay. It is rarely
    Order: Veneroida                                      found in the estuaries of coastal Washington and
    Family: Mactridae                                     Oregon except for Netarts Bay, Oregon, where >50%
                                                          of the gapers are T. nuttallii (T. Gaumer, Oregon
    Value                                                 Department of Fish and Wildllife, Newport, OR, pers.
    Commercial: The Pacific gaper is harvested with the   comm.).  It is particularly abundant in Tomales Bay
    similar horseneck clam, Tresus capax. Landings are   where up to 35,000 have been taken annually at one
    not identified to species, but instead reported together    location (Frey 1971 ). This species is dug at low tide by
    as"horseclams". From 1981 -1983, horseclamlandings    hand or with hand tools (Frey 1971). It is one of the
    from the U.S. and Canadian Pacific coast averaged    most common bay clams along the California coast.
    about 225 t annually, and ranked fifth in volume of all   Not more than ten Pacific gapers per person per day
    clams harvested (Wolotira et al. 1989). Much of the   can be taken in most areas of California (Schultze
    commercial harvest in British Columbia has been by    1986). This species is often made into chowder (Frey
    geoduck (Panopea abrupta) divers after they have    1971).
    reached their geoduck quota (Wolotira et al. 1989).
    The Pacific gaper is relatively large and has many    Indicator of Environmental Stress: Clam beds are
    biological  characteristics  which  discourage    sometimes closed to harvest because of paralytic
    commercialization. It burrows deep into soft sediments,   shellfish poisoning. Other beds are permanently closed
    making hand harvest difficult. The shells are relatively   to harvesting because of contamination by coliform
    fragile and tend to break; once harvested, the shells   bacteria. As a result of pollution in Washington waters,
    gape, causing water loss and reducing shelf life. Meat    over 25% of the potential areas for subtidal clam
    yield per clam is relatively low, usually <30%, and the    harvesting are closed (Schink et al. 1983). In California,
    large siphon (often 60% of its shucked weight) has a    clams in estuaries such as San Francisco Bay are not
    tough, leathery skin that requires extra effort to remove    commonly harvested because of pollution. Embryos
    (Quayle and Bourne 1972, Ricketts et al. 1985, Wolotira   are good bioassay organisms (Woelke et al. 1971).
    et al. 1989). This species is harvested both subtidally
    and intertidally using hydraulic pumps, mechanical    Ecological: This species is a large, subtidal and lower
    dredges, potato forks, shovels, and clam rakes (Frey   intertidal suspension/filter feeding bivalve and is

                                                       30

                                                                                         Pacific gaper continued
                                                           Within Study Area: The Pacific gaper is found in Pacific
      Table 1. Relative abundance of Pacific gaper          coast estuaries from Puget Sound, Washington, to
               in 32 U.S. Pacific coast estuaries.         Tomales Bay (Table 1). However, it is rarely found in
                        Life Stage                         the coastal estuaries of Washington and Oregon (except
           Estuary    A S J L E                            Netarts Bay), and is not common in most bays and
            PugetSound i( (3 3 (  3    Relative abundance:  lagoons south of Pt. Conception, California.
             Hood Canal (3 3 (  3  (   �*  Highly abundant
              Skagit Bay 0 0     0 0 I      Abundant        Life Mode
                  Grays Harbor        0   Common
            Grays Harbor l    ii4  Rare                     Eggs and larvae are pelagic. Juveniles and adults are
             Willapa Bay              Blank Not present     benthic infauna; adults may burrow to depths of 1 m
          Columbia River                                   (usually found 25-50 cm deep) (Cheney and Mumford
            Nehalem Bay                                     1986, Wolotira et al. 1989).
           Tillamook Bay             Life stage:
             NetarisBay (g )           A -Adults
                                      S - Spawning adults  Habitat
             Siletz River             J - Juveniles         Hype: Eggs and larvae are neritic. Juveniles and adults
            YaquinaBay     - J ' ,      .Larvae,
                                      E - Eggs             are found primarily in bays and estuaries, but may also
             Alsea River                                   occur in protected coastal waters (Frey 1971, Wolotira
           Siuslaw River                                   et al. 1989). Juveniles and adults occur from the lower
           Umpqua River                                    intertidal zone to 30 m below mean lower low water
              Coos Bay                                     (MLLW). In Puget Sound, they are most abundant from
            Rogue River                                    1-5 m below MLLW (Goodwin and Shaul 1978, Cheney
           Klamath River                                   and Mumford 1986).
           Humboldt Bay 0 0 O 0 O
               Eel River
               TmaEel~es Bay (3 (3(3(3(3Substrate: The Pacific gaper is most abundant in
            Tormales Bay 00000 13 <3 <3                    sediments consisting of fine sand or firm sandy mud.
       Cent. San Fran. Bay '     '4 ' '  Includes Central San
                                     Frarisco. Suisun,     But, it is also found in relatively firm sediments consisting
           EkhmSout h San Fran. Bay    and San Pablo bays.  of sand, silty-sand, sandy-clay, and gravel (Swan and
           ElkhormnSlough 0003Q (3 61 tFinucane 1951, Bourne and Smith 1972, Cheney and
              Morro Bay  3       I < 1                      Mumford1986,Wolotiraetal. 1989). Sedimentstructure
           Santa Monica Bay  '4 '4 '4 '4 '4                 affects burrowing depth; clams burrow deeper in mud
           SanaPeroBay        d �                           and  sand  substrates  than  clay  substrates

            Anaheim Bay '4 '41 '4                           (Oceanographic Institute of Washington 1981).
            NewportBay  '4   '4 '4 '4
                       Missior Say   '4 i i i iPhvsical/Chemical Characteristics: It occurs in
             Mission Bay V-V  ,
           SanDiego Bay     '4 ' '4                         polyhaline-euhaline waters, and temperatures of 1-
          TijuanaEstuary  i v i v '                        21 C (Bernard 1983). Freezing temperatures on mud
                      A S J L E                            flats may limit this species' northern distribution (Pearce
                                                           1965).

    important in Puget Sound and many California estuaries,    Miarations and Movements: Eggs and larvae are
    bays, and lagoons (Frey 1971). Pea crabs (Pinnixa    dispersed by currents. Juveniles and adults do not
    faba and occasionally P. littoralis) can be found in the    move laterally once they become established. Small
    Pacific gaper's mantle cavity (Ricketts et al. 1985). The    Pacific gapers have the ability to reburrow after being
    hard, leatherytips areoften covered with many different    disturbed, but like T. capax, older, larger clams (>60
    species of plants and animals (Haderlie and Abbott    mm shell length) lose the ability to reburrow (Pholo
    1980). The Pacific gaper appears to harbor pea crabs    1964, Wendell et al. 1976). However, since most larger
    only in the southern part of its range (Pearce 1965).    clamslivedeepwithinthesediment(upto1 m)theyare
    This species is an intermediate hostforthetapeworm,    protected from most natural disturbances.   Peak
    Echeneibothrium sp., whose definitive host is the bat    settlement for spat occurs in May in central California
    ray (Myliobatis californica) (Haderlie and Abbott 1980).   and probably July in Puget Sound (Woelke et al. 1971,
                                                           Clark et al. 1975).
    Range
    Overall: The Pacific gaper is a temperate, amphi-North    Reproduction
    Pacific species (Bernard 1983,Wolotiraet al. 1989). In   Mode: The Pacific gaper is gonochoristic, oviparous,
    North America, it is found from Scammons Lagoon,    and iteroparous. It is a broadcast spawner; eggs are
    Baja California, to British Columbia (Fitch 1953).      fertilized externally (Quayle and Bourne 1972).

                                                        31

    Pacific gaper continued
    Matina/Soawnina: Spawning occurs year-round,    (Polinices spp.), Dungeness crab (Cancer magister),
    dependingongeographical location. Spawningoccurs    bat ray (Myliobatus californica), leopard shark (Triakis
    during summer in northern regions such as British   semifasciata), starry flounder (Platichthys stellatus),
    Columbia and Puget Sound (Quayle and Bourne 1972,   sea stars (Pisaster spp.), and sea otters (Enhydra
    Cheney and Mumford 1986). Spawning occurs from    lutris) (Talent 1976, Haderlie and Abbott 1980, Kvitek
    spring to fall for much of California (Frey 1971), and    et al. 1988). Many planktivorous organisms prey on
    year-round in central California, with a peak from    Pacific gaper eggs and larvae.
    Februaryto April when temperatures are lowest (Laurent
    1971, Clark et al. 1975, Haderlie and Abbott 1980,    Factors Influencina Populations: Sea otters prefer to
    Ricketts etal. 1985). Thewidedailywatertemperature    feed in areas where Pacific gaper densities are high
    fluctuations in central California may explain the    and composed of small individuals unable to burrow
    occurrence of year-round spawning (Clark et al. 1975).   deeply because of sediment characteristics (Kvitek et
                                                           al. 1988); large Pacific gapers in soft sediments are
    Fecundity: Unknown.                                     resistant to sea otter predation. The Pacific gaper may
                                                           compete with T. capax, however T. capax is more
    Growth and Development                                  common in gravel-shell soils whereas T. nuttallii is
    Eaa Size and Embrvonic Development: Egg size is   more common in pure sand substrates (Swan and
    unknown, however, embryonic development is indirect    Finucane 1951, Quayle and Bourne 1972, Wolotira et
    and external (Wolotira et al. 1989).                    al. 1989). The Pacific gaper also burrows deeper than
                                                           T. capaxand thus avoidstemporaryfreezingconditions
    Aae and Size of Larvae: Larvae are probably 0.06-0.28    (Quayle and Bourne 1972, Haderlie and Abbott 1980).
    mm in diameter (Bourne and Smith 1972). In Elkhorn    No information is available concerning mortality rates,
    Slough, California, the duration of the larval stage is   but very high mortality rates probably occur during
    estimated to be 21-30 days (Clark et al. 1975). Spat    larval and early juvenile stages, becoming lower as
    require ten days to grow to 2 mm, and 25 days to grow    clams mature (Wolotira et al. 1989). Annual juvenile
    to 5 mm (Clark et al. 1975).                            recruitment varies widely and probably has a major
                                                           effect on the population structure (Clark et al. 1975).
    Juvenile Size Ranae: Juveniles are 0.26 mm to 51.0-
    71.0 mm in diameter; small clams (4 mm) grow 0.25    References
    mm/day (Frey 1971, Bourne and Smith 1972, Haderlie
    and Abbott 1980). One-year-old clams average 50 mm    Bernard, F. R. 1983. Catalogue of the living Bivalvia
    in shell length (Clark et al. 1975, Haderlie and Abbott    of the eastern Pacific Ocean: Bering Strait to Cape
    1980).                                                  Horn. Can. Spec. Fish. Aquat. Sci. 61,102 p.

    Aae and Size of Adults: This species matures in about    Bourne, N., and D. W. Smith.  1972. The effect of
    two years and between 51.0-70.0 mm shell length    temperature on the larval development of the horse
    (Frey 1971, Clark et al. 1975, Haderlie and Abbott    clam, Tresus capax  (Gould).  Proc. Natl. Shellfish.
    1980). The Pacific gaper may live to 17 years, with a    Assoc. 62:35-46.
    shell length as great as 200 mm (Frey 1971, Wolotira
    et al. 1989).                                           Cheney, D. P., and T. F. Mumford, Jr. 1986. Shellfish
                                                           and seaweed harvests of Puget Sound. Wash. Sea
    Food and Feeding                                        Grant, Univ. Wash. Press, Seattle, WA, 164 p.
    TroDhic Mode: This species is a suspension/filterfeeder.
    Food particles are transported via the inhalant siphon    Clark, P., J. Nybakken, and L. Laurent. 1975. Aspects
    and are filtered fromthewaterbythe gills, sorted bythe    of the life history of Tresus nuttallii in Elkhorn Slough.
    palps, and passed to the mouth.                         Calif. Fish Game 6(4):215-227.

    Food Items: Food items include suspended diatoms,    Fitch,J. E. 1953. Common marine bivalvesof California.
    flagellates, dinoflagellates, and detritus. Detritus may    Calif. Fish Game, Fish Bull. 90, 102 p.
    include particles of eelgrass (Zostera marina) (Stout
    1967, Haderlie and Abbott 1980).                        Frey, H. W. 1971. California's living marine resources
                                                           and their utilization.  Calif. Dept. Fish Game,
    Biological Interactions                                 Sacramento, CA, 148 p.
    Predation: Predators include those that prey on T.
    capax, especiallyworms, snails, crustaceans, fish, and
    mammals. Common predators include moon snails

                                                        32

                                                                                         Pacific gaper continued
    Goodwin, L., and W. Shaul.  1978.  Puget Sound    Swan, E. F., and J. H. Finucane. 1951. Observations
    subtidal hardshell clam survey data. Prog. Rep. 44,    on the genus Schizothaerus. Nautilus 66(1):19-26.
    Wash. Dept. Fish., Olympia, WA, 92 p.
                                                           Talent, L. G. 1976. Food habits of the leopard shark,
    Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: The    Triakis semifasciata, in Elkhorn Slough, Monterey Bay,
    clams and allies. In R. H. Morris, D. P. Abbott, and E.   California. Calif. Fish Game 62(4):286-298.
    C. Haderlie (editors), Intertidal invertebrates of
    California, p. 355-411. Stanford Univ. Press, Stanford,    Wendell, F., J. D. DeMartini, P. Dinnel, and J. Siecke.
    CA.                                                     1976. The ecology of the gaper or horse clam, Tresus
                                                           capax (Gould 1850) (Bivalvia: Mactridae) in Humboldt
    Kvitek, R. G., A. K. Fukayama, B. S. Anderson, and B.    Bay, California. Calif. Fish Game 62(1):41-64.
    K. Grimm. 1988. Sea otter foraging on deep-burrowing
    bivalves in a California coastal lagoon.  Mar. Biol.   Woelke, C., T. Schink, E. Sanborn, and W. Hoffman.
    98:157-167.                                             1971.  Bivalve embryo bioassays of marine waters
                                                           from Drayton Harbor to Hale Passage. Wash. Dept.
    Laurent, L. L. 1971. The spawning cycle and juvenile    Fish., Unpubl. Rep. to Atlantic Richfield Co., Olympia,
    growth rate of the gaper clam, Tresus nuttallii, of   WA, 18 p.
    Elkhorn Slough, California. M.A. Thesis, San Francisco
    State College, San Francisco, CA, 55 p.                 Wolotira, R. J., Jr., M. J. Allen, T. M. Sample, C. R. Iten,
                                                           S. F. Noel, and R. L. Henry. 1989. Life history and
    Oceanographic Institute of Washington. 1981. Clam    harvest summaries for selected invertebrate species
    and mussel harvesting industries in Washington State.    occurring off the west coast of North America. Volume
    Oceanog. Comm.  Wash., Seattle, WA,  various    1: Shelled molluscs. NOAA Tech. Memo. NMFS F/
    pagination.                                             NWC-1 60, 177 p.

    Pearce, J. B.  1965.  On the distribution of Tresus
    nuttallii and Tresus capax in the waters of Puget
    Sound and the San Juan Archipelago. Veliger7(3):166-
    170.

    Pohlo, R. H. 1964. Ontogenetic changes of form and
    mode of life in Tresus nuttallii (Bivalvia: Mactridae).
    Malacologia 1 (3):321-330.

    Quayle, D. B., and N. Bourne. 1972. The clam
    fisheries in British Columbia. Fish. Res. Board Can.,
    Bull. No. 179, 70 p.

    Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W.
    Phillips. 1985. Between Pacific tides. Stanford Univ.
    Press, Stanford, CA, 652 p.

    Schink, T. D., K. A. McGraw, and K. K. Chew. 1983.
    Pacific coast clam fisheries. Wash. Sea Grant, Univ.
    Wash., Seattle, WA, 72 p.

    Schultze, D. L. 1986. Digest of California commercial
    fish laws, January 1, 1986. Calif. Dept. Fish Game,
    Sacramento, CA, 40 p.

    Stout, W. E. 1967. A study of the autecology of the
    horse neck clams Tresus capax and Tresus nuttallii in
    South Humboldt Bay, California. M.A. Thesis, Humboldt
    State Univ., Arcata, CA, 51 p.

                                                        33

       Tagelus californianus
       Adult

                                               5cm

    Common Name: California jackknife clam                Tijuana estuary to Morro Bay, California; it is not
    Scientific Name: Tagelus californianus                common north of Monterey Bay, California (Table 1)
    Other Common Names: California short razor, short   (Fitch 1953, Haderlie and Abbott 1980, Seapy 1981).
    razor clam, jackknife clam, razorclam (Gates and Frey
    1974)                                                 Life Mode
    Classification (Bernard 1983)                         Eggs and larvae are planktonic. Juveniles and adults
    Phylum: Mollusca                                      are benthic infauna of bays, estuaries, or lagoons.
    Class: Bivalvia                                       Juveniles and adults live in a permanent, nonmucous-
    Order: Veneroida                                      lined, vertical burrow 10-50 cm deep in which they can
    Family: Psammobiidae                                  readily move up and down (Fitch 1953, Meinkoth
                                                          1981).
    Value
    Commercial: This species is commercially dug for use    Habitat
    as fish bait (Fitch 1953). Harvest began in 1962 and    Iype: Eggs and larvae are estuarine-neritic. Adults
    during the mid-1970s harvests averaged about 6t/year   and juveniles are common near mean low tide where
    (Wolotira et al. 1989).                               sediments are appropriate (Seapy and Kitting 1978,
                                                          Merino 1981). Adults and juveniles inhabit sand, mud,
    Recreational: Although edible, it is most often used as    or muddy sand flats near the low tide level in bays,
    fish bait (Fitch 1953, Meinkoth 1981).                sloughs, and estuaries (Fitch 1953, Smith and Carlton
                                                          1975, Meinkoth 1981). This species reportedly occurs
    Indicator of Environmental Stress: High temperatures    from +0.2 to -0.5 m mean tide level (Wolotira et al.
    (e.g.,thermal effluent from power plants) can adversely    1989), but does not occur above mean sea level in San
    affect populations (Merino 1981).                     Diego Bay (Merino 1981). The bays and lagoons this
                                                           species inhabits are euhaline on an annual basis. In
    Ecological:TheCaliforniajackknifeclamisa numerically    low intertidal substrates, it is commonly associated
    important bivalve species in southern California bays    withthe rosy jackknife (Solen rosaceus) (Merino 1981).
    and lagoons.
                                                           Substrate: The California jackknife clam prefers
    Range                                                 sediments having some silts and clays (2-15%), and
    Overall: This species' overall range is from Cape San    cannot burrow into sediments that are composed
    Lucas, Baja California to Cape Blanco, Oregon (Fitch   primarily of sand (Merino 1981).
    1953, Meinkoth 1981, Wolotira et al. 1989). Its recorded
    presenceoff Panama is probably not accurate (Wolotira    Phvsical/Chemical Characteristics: This species is
    et al. 1989).                                         found in mesohaline-euhaline waters where water
                                                           temperatures range from 9 to 300C (Bernard 1983).
    Within Study Area: It is common to abundant from    Temperatures >35�C cause adult mortality. In San

                                                       34

                                                                              California jackknife clam continued

                                                          species is unknown, however, spawning occurs
      Table 1. Relative abundance of California            intertidally during high tide.  Eggs and sperm are
               jackknife clam in 32 U.S. Pacific coast     released through the exhalant siphon. Based on the
               estuaries.                                  settlement of young, a peak spawning probably occurs
                        Life Stage                        in early spring (May-June recruitment), with some
            Estuary    A S J L E                           spawning occurring year-round (Merino 1981).
             Puget Sound             Relative abundance:
              Hood Canal  :                Highly abundant  Fecundity: Unknown.
              Skagit Bay                  Abundant
            Grays Harbor                   ommon          Growth and Development
                                      V   lRare
             Willapa Bay            Blank Not present      Eaa Size and Embrvonic Develooment: Unknown, but
           Columbia River                                  embryonic development is probably indirect and
            Nehalem Bay                                   external.
            Tillamook Bay             Ufe stage:
              Netarts Bay              A- Adults           Aae and Size of Larvae: Unknown.
                                       S - Spawning adults
              Siletz River             J - Juveniles
             Yaquina Bay               E- Eggs             Juvenile Size Ranae: The stout tagelus (Tagelus
              Alsea River                                  plebius) is a congener, and has spat that settle out of
            Siuslaw River                                 the water column at 155-175 pLm in shell length (SL)
            Umpqua River                                   (Merino 1981). Clams average about 46 mm SL at 2.5
               Coos Bay                                    years (Merino 1981).
             Rogue River
            Klamath River
            uKlamath River   ;Aae and Size of Adults:The California jackknife reaches
               EeHumboldt Bay                              maturity between 60 and 120 mm SL (Merino 1981).
               TomaEel River  I i -J                      Ageandgrowthof this species has notbeendetermined,
             Cent.SanFran.ls Bay  - i V    i J    IncludesCentralSan  but it appears to reach reproductive size in 2-3 years
       Cent. San Fran. Bay *   -4   i V   -] -4 / * Includes Central San
       South San Fran. Bay          Francisco, Suisun.     (Merino 1981). Ultimate age is unknown. Clams in San
           Elkhorn Slough F  Ba     and San Pablo bays.    Diego Bay average 72 mm SL and appearto be 5 years
              MorroBay 00000                              old (Merino 1981).
         Santa MonicaBay { * 3]
           SanPedroBay    ) i) Q    ci                     Food and Feeding
             Alamitos Bay O0 0    O :  Trophic Mode: This species is a suspension feeder,
            Anaheim Bay O OO 0 0                          although originally it was thoughtto be a depositfeeder
             NewportBay O C O  O O                         (Pohlo 1966, Haderlie and Abbott 1980). When feeding,
             Mission Bay i  3   c  13                     it is located about 10 cm below the substratum surface
           San Diego Bay ( c     i                         and extends its two siphons into the water through
           Tijuana Estuary  ]  ] O c c                     separate openings (Haderlie and Abbott 1980). The
                       A S J L E                           siphon openings lay at the sediment-water interface.

    Diego Bay, the clam's upper lethal tolerance limit   Food Items: The California jackknife clam feeds on
    (LT50) was 35.5�C in December and 37.6�C in May    phytoplankton,  probably  including  diatoms,
    (Merino 1981). Smaller sizes (23-46 mm) are more    dinoflagellates, and other types of phytoplankton. Its
    resistant to elevated temperatures (Merino 1981).      diet may include suspended detrital particles and their
                                                          associated epifauna (Wolotira et al. 1989).
    Miarations and Movements: Eggs and larvae are
    dispersed bycurrents. Juveniles and adults migrate up    Biological Interactions
    and down in their burrow as the tide rises and falls   Predation:Larvaeprobablyareeatenbyplanktivorous
    (Meinkoth 1981) and will rapidlydescend intheirburrows    fishes and invertebrates.  Newly-settled individuals
    when disturbed.                                        and juveniles are eaten by numerous fishes, including
                                                          diamond turbot (Hypsopsetta guttulata) (Lane 1975),
    Reproduction                                           stingrays (Dasyatis spp.), and other rays. Birds such
    Mode: This species is gonochoristic, oviparous, and    as stilts (Himantopus spp.), godwits (Limosa spp.),
    iteroparous.  It is a broadcast spawner; eggs are    curlews ( Numenius  spp.),  and  dowitchers
    fertilized externally.                                 (Limnodromus spp.), also prey on the California
                                                          jackknife clam (Merino 1981).
    Matina/SDawnina: The exact spawning time for this

                                                       35

    California jackknife clam continued
    Factors Influencino PoDulations: Population densities    Wolotira, R. J., Jr., M. J. Allen, T. M. Sample, C. R. Iten,
    are influenced by tidal elevation, water temperature,    S. F. Noel, and R. L. Henry. 1989. Life history and
    sediment characteristics, recruitment, and mortality.    harvest summaries for selected invertebrate species
    There are no indications that populations are controlled    occurring off the west coast of North America. Volume
    by density-dependent interactions (Merino 1981).         1: shelled molluscs.  NOAA Tech. Memo. NMFS F/
                                                            NWC-160, 177 p.
    References

    Bernard, F. R. 1983. Catalogue of the living Bivalvia
    of the eastern Pacific Ocean: Bering Strait to Cape
    Horn. Can. Spec. Publ. Fish. Aquat. Sci. 61,102 p.

    Fitch, J. E. 1953. Common marine bivalves of California.
    Calif. Fish Game, Fish Bull. 90,102 p.

    Gates, D. E., and H. W. Frey. 1974. Designated
    common names of certain marine organisms of
    California. Calif. Fish Game, Fish Bull. 161:55-90.

    Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: The
    clams and allies. In R. H. Morris, D. P. Abbott, and E.
    C. Haderlie, Intertidal invertebrates of California, p.
    355-411. Stanford Univ. Press, Stanford, CA.

    Lane, E. D. 1975. Quantitative aspects of the life
    history of the diamond turbot, Hypsopsetta guttulata
    (Girard), in Anaheim Bay. In E. D. Lane and C. W. Hill
    (editors), The marine resources of Anaheim Bay. Calif.
    Fish Game, Fish Bull. 165:153-173.

    Meinkoth, N. A. 1981. The Audubon Society field
    guide to North American seashore creatures. Alfred A.
    Knopf, Inc., New York, NY, 799 p.

    Merino, J.-M. 1981. A study of the temperature
    tolerances of adult Solen rosaceus and Tagelus
    californianus in south San Diego Bay: the effects of
    power plant cooling water discharge. Ph.D. Diss., San
    Diego State Univ., San Diego, CA, 140 p.

    Pohlo, R. 1966. A note on the feeding behavior in
    Tagelus califomianus. Veliger 8(4):225.

    Seapy, R. R. 1981. Structure, distribution, and seasonal
    dynamics of the benthic community in the upper Newport
    Bay, California. Mar. Res. Tech. Rep. 46, Calif. Dept.
    Fish Game, Sacramento, CA, 74 p.

    Seapy, R. R., and C. L. Kitting. 1978. Spatial structure
    of an intertidal molluscan assemblage on a sheltered
    sandy beach. Mar. Biol. 46:137-145.

    Smith, R. I, and J. T. Carlton (editors). 1975. Lights
    manual: Intertidal invertebrates of the central California
    coast. Univ. Calif. Press, Berkeley, CA, 716 p.

                                                         36

    37

       Protothaca staminea
       Adult

                                         2cm

    Common Name: Pacific littleneck clam                    1953). In California, upto 50 clams/day over 3.8 cm in
    Scientific Name: Protothaca staminea                    diameter are allowed (California Department of Fish
    OtherCommonNames:TomalesBaycockle, common    and Game 1987), while Oregon limits recreational
    littleneck, littleneck clam, ribbed carpet shell, common    harvest to only 36/day. The Washington limit varies
    Pacific littleneck, native littleneck, rock cockle, hardshell,    depending on the area (60/day or 10 lb, 40/day or 7 lb,
    rock clam, steamer, butterclam (Fitch 1953, Gates and    5 lb/day) (Washington Department of Fisheries 1986).
    Frey 1974, Hancock et al. 1979)                         Clam diggers usually harvest this species at low tide
    Classification (Bernard 1983a)                          during daylight using rakes, trowels, and shovels (Frey
    Phylum: Mollusca                                        1971).
    Class: Bivalvia
    Order: Veneroida                                        Indicator of Environmental Stress: Habitat alterations
    Family: Veneridae                                       (water pollution, marina construction, loss of habitat,
                                                           etc.) directly affect the abundance of this species.
    Value                                                   Paralytic shellfish poisoning often closes clam beds to
    Commercial: The Pacific littleneck clam is usually sold    harvest for temporary periods and contamination by
    fresh in the shell (Wolotira et al. 1989), but it is also sold    coliform bacteria has permanently closed many areas
    frozen and canned (Paul and Feder 1976).  It is   (Cheney and Mumford 1986). Commercial landings
    harvested using rakes, shovels, and by mechanical    from the U.S. Pacific Northwest (excluding Alaska)
    and hydraulic devices (Frey 1971, Schink et al. 1983,    have decreased in recent years, while effort has
    Cheney and Mumford 1986). Harvested from Prince    increased (Chew and Ma 1987). This species is highly
    William Sound, Alaska to southern California, this   sensitive to copper and tri-n-butyltin (a paint additive)
    speciesconstitutes about8%of theentireclam harvest    (Roesijadi 1980).  Crude oil reduces this species'
    alongthePacificcoastofthe United States and Canada    growth rate, but does not appear to be highly toxic.
    (Wolotira et al. 1989). Most of this harvest comes from    However, the addition of oil dispersants can alter clam
    Washington and British Columbia. Most Pacific coast    behavior deleteriously (Chew and Ma 1987).
    waters are open year-round, but California waters are
    closed to littleneck harvest from April to August in Marin    Ecological: This species is common to highly abundant
    County and from May to August for much of northern    in many Pacific coast estuaries (Table 1).  It is an
    California (Schultze 1986).  Because California    important suspension feeder along protected gravel-
    commercial clammers are allowed only 50 clams/day    mud beaches (Wolotira et al. 1989) and the most
    over3.8 cm diameter, the California commercial harvest    important lower intertidal clam in Puget Sound (Kozloff
    is limited. New aquaculture programs may increase    1983).
    the production and harvest of this species.
                                                            Range
    Recreational: The Pacific littleneck clam is highly    Overall: This species may be distributed from Socorro
    esteemed for its good taste and ease of capture (Fitch   Island, Mexico, around the North Pacific rim to the

                                                        38

                                                                             Pacific littleneck clam continued
                                                          individuals are often found deeper than smaller ones
      Table 1. Relative abundance of Pacific littleneck    (Fitch 1953, Quayle and Bourne 1972, Paul and Feder
               clam in 32 U.S. Pacific coast estuaries.    1973, Abbott 1974, Meinkoth 1981, Wolotira et al.
                        Life Stage                         1989).
           Estuary    A S J L E
            Puget Sound * 6 � 6  �   Relative abundance:   Habitat
            Hood Canal � � �  � �   Highly abundant       Type: Eggs and larvae are estuarine-neritic. Adults
                                     O Common
             skagit Bay *���� @ @ @       Abumndant        and juveniles are found in coarse, sandy-rocky muds of
           Grays Harbor O     C O O O     Rare            bays, sloughs, and estuaries, and on the open coast
            Willapa Bay 0 O O O O   Blank Not present     where there is appropriate substrate and protection
          Columbia River                                   (Fitch 1953). It is often associated with butter clams
           Nehalem Bay *     0 9  6 6                      (Saxidomus giganteus) (Paul and Feder 1976). The
           Tillamook Bay      3 t3 ( ~]    Life stage:     Pacific littleneckclam is found intertidallydownto37 m
            Netarts Bay & ci tj      A-Adults
                                      S -Spawning adults  (usually <10 m), but normally from -1.0 to 1.3 m mean
             Siletz River             J J-Juveniles        lower low water (MLLW) (Chew and Ma 1987). It is
            YaquinaBay 00000          L-Larvae
                                     EYauina  - Eggs     most abundant from the lower intertidal zone to 0.4 m
             Alseiaw River  i a} 5 ivabove MLLW (Goodwin and Shaul 1978, Bernard
           Siuslaw River  i no 71 i i1983a, Wolotira et al. 1989).
           Umpqua River
              Coos Bay * S -                  ~Substrate: The Pacific littleneck clam prefers firm,
            Rogue River                                    gravel orclay-gravel sediments, butoccurs in sediments
           Klamath River
           HuKlamath River ~ ~ ~ranging from mud to cobble (Quayle and Bourne 1972,
              EeHumboldt Bay (Goodwin and Shaul 1978). Along the open coast it is
              EeTomalesray O O 8     l                    found in coarse sand, gravel, and cobble near rock
            Cent.omSanFran. Bay* i            InudesCentralSan  points and reefs or under large rocks (Fitch 1953).
      Cent. San Fran. Bay * ",    ' Includes Central San
      South San Fran. Bay           Francisco, Suisun,
                                    and SanPablobays.     Phvsical/Chemical Characteristics: It is found in
          Elkhorn Slough 0 0    0 0C
             Morro Bay O O O O  mesohaline to euhaline waters and temperatures of
        SantaMonicaBay  O C  O O                           just below freezing to 25�C (Glude 1978, Bernard
          San Pedro Bay O O O O 0                          1983a). Water temperatures above 25�C are lethal to
            Alamitos Bay ( r (3 Q s                        larvae, andtheycan withstand 200C onlywhen salinity
           Anaheim Bay i &     t   is near 32%0o (Strathmann et al. 1987). This species
            NewportBay O C O O O                           may tolerate salinities as low as 20%o for extended
            Mission Bay 'i i i i 'i                       periods (Quayle and Bourne 1972); however, it closes
          San Diego Bay  1 i i i ,                         its shell at very low salinities. Optimum conditions for
          Tijuana Estuary O O 6 Q 3                        growth appearto be 12-18�C, 24-31%o salinity, and 15-
                      A S J L E                            150 mg/I suspended food particles (Bernard 1983b).
                                                          Also, areas near strong tidal currents may enhance
                                                          growth (Chew and Ma 1987). Burial by decomposing
    northern Sea of Japan (Wolotira et al. 1989). However,    bark has been shown to reduce survival (likely due to
    most authors show it distributed from Cape San Lucas,    elevated levels of hydrogen sulfide and ammonia along
    Baja California, to the Aleutian Islands, Alaska (Fitch   with decreases in dissolved oxygen) (Freese and O'Clair
    1953, Schinketal. 1983, Cheney and Mumford 1986).    1987).  High turbidities (>2 g/l) may reduce larval
                                                          survival (Glude 1978).
    Within Studv Area: It is found in most Pacific coast
    estuaries where appropriate substrates and salinities    Miaration and Movements: Eggs and larvae are pelagic
    exist. It is not found in the Columbia, Siletz, Umpqua,    and dispersed by water currents. Veliger larvae move
    and Rogue River estuaries of Oregon, or the Klamath,    to the bottom after developing a foot. Herethey search
    and Eel River estuaries in California (Table 1) (Monaco    for an appropriate surface on which to settle, then
    et al. 1990).                                          undergo metamorphosis, and attach themselves to the
                                                          sediment surface by secreting byssal threads (Chew
    Life Mode                                              and Ma 1987). Very young clams probably first attach
    Eggs and larvae are pelagic, while very small clams are   in deeper waters and then move to shallow waters as
    epifaunal (Paul and Feder 1973). Juveniles and adults   they grow (Chew and Ma 1987). Adults are sedentary
    are benthic infauna and found in the upper 15-20 cm of   and remain in the same area for life, but a small juvenile
    sediments (rarely deeper than 5-7 cm).  Larger    clam can use its foot to crawl to new areas (Shaw

                                                      39

    Pacific littleneck clam continued
    1986). Adults and juveniles can reburrow if they have    et al. 1985, Cheney and Mumford 1986).  British
    been disturbed (Quayle and Bourne 1972).               Columbia and Alaska clams are often not mature until
                                                          their second or third year (Fraser and Smith 1928,
    Reproduction                                           Quayle 1943, Nickerson 1977). This species may live
    Mode: The Pacific littleneck clam is gonochoristic    13-16 years (Fraser and Smith 1928, Abbott 1974,
    (although some hermaphroditism occurs), oviparous,    Chew and Ma 1987). In California, many die before
    iteroparous, and a broadcast spawner; eggs are    reaching sexual maturity and rarely do they reach 7
    fertilized externally (Fraserand Smith 1928, Frey 1971).   years old (Schmidt and Warme 1969). Maximum size
    Females may spawn several times during a season    is about 8 cm SL (Quayle and Bourne 1972,
    (Quayle and Bourne 1972).                              Oceanographic Institute of Washington 1981). Growth
                                                          rates vary widely, depending on substrate, clam
    Matina/lSawnina: Spawning occurs during spring and    densities, tidal level, and geographic location (Chew
    summerdependingon the region: from March toAugust    and Ma 1987). For example, they may grow to 37 mm
    and sometimes later in Oregon estuaries (Robinson    SL in 3.5-4 years in the Strait of Georgia (Cheney and
    and Breese 1982); April to September in British    Mumford 1986), and 6-8 years to reach 32 mm SL in
    Columbia; late spring to summer (April-July) in Puget    Alaska (Paul and Feder 1973, 1976, Ricketts et al.
    Sound; late May to mid-June in Prince William Sound,    1985).
    Alaska (Fraser and Smith 1928, Haderlie and Abbott
    1980, Cheney and Mumford 1986, Strathmann et al.   Food and Feeding
    1987, Wolotiraetal. 1989). It spawns attemperatures    Trophic Mode: The Pacific littleneck clam is a
    of 5.6-13.60C in Prince William Sound (Wolotira et al.   nonselective suspension/filter feeder. It gathers food
    1989), and begins spawning in south-central Alaska    by sucking in water and food particles through the
    when water temperatures are about 8�C (Chew and    inhalant siphon. Particles are then filtered through the
    Ma 1987). Dense algal suspensions may stimulate   gills (ctenidia), and sorted by the palps before being
    spawning (Robinson and Breese 1982).  Optimum    brought to the mouth (Wolotira et al. 1989).
    temperatures for rearing are 15-200C (Strathmann et
    al. 1987).                                             Food Items: Larvae, juveniles, and adults feed on
                                                          phytoplankton, benthic diatoms, and detritus. The role
    Fecundity: Unknown.                                    of detritus in its diet is not well understood, but thought
                                                          to be important (Peterson 1982, Chew and Ma 1987,
    Growth and Development                                 Wolotira et al. 1989).
    Eao Size and Embrvonic Develooment: Eggs are
    spherical and 0.06 mm in diameter (Wolotira et al.   Biological Interactions
    1989). Embryonic development is indirect and external.    Predation: Important predators of the Pacific littleneck
    Fertilized eggs hatch to become free-swimming    clam include: oyster drills (Ceratostoma spp. and
    trochophore larvae in 10-12 hours; these transform    Urosalpinx spp.), moon snails (Polinices spp.), and
    intoveligerlarvae approximately24 hours later (Quayle    othergastropods, sea stars (Pycnopodiahelianthoides,
    and Bourne 1972, Schink et al. 1983, Chew and Ma    Evasterias troschelli, and Pisaster brevispinis), two-
    1987).                                                 spotted octopus (Octopus bimaculatus), rock crabs
                                                          (Cancerspp.), and fishes (Chew and Ma 1987, Wolotira
    Aae and Size of Larvae: Larvae range from 0.06-0.25   et al. 1989). Rock crabs have the ability to identify
    mm long (Quayle and Bourne 1972, Wolotira et al.   foraging areas with high littleneck clam densities
    1989). The larval period lasts about three weeks, but    (Boulding and Hay 1984). In California lagoons, siphons
    may be longer depending on water temperatures    are nipped off by Pacific staghorn sculpin (Leptocottus
    (Quayle and Bourne 1972, Cheneyand Mumford 1986).    armatus), diamond turbot (Hypsopsettaguttulata), and
                                                          California halibut (Paralichthys californicus) (Peterson
    Juvenile Size Ranae: At settlement, juveniles are 0.26-    and Quammen 1982). Sea otters (Enhydra lutris) are
    0.28 mm in shell length (SL) (Quayle and Bourne    majorpredatorsinPrinceWilliamSound, Alaska (Chew
    1972) and grow to 15-35 mm SL before maturity.    and Ma 1987), and the Pacific littleneck clam is also
    Growth varies depending on the region.  In Prince    eaten by ducks and other birds (Schink et al. 1983,
    William Sound, clams are2 mm SL attheend of thefirst   Cheney and Mumford 1986).
    growing season (Paul and Feder 1973).
                                                           Factors Influencino Pooulations: Recruitment (i.e.,
    Aaeand Sizeof Adults: Thisspecies is usually sexually   survival of the settling spat) is highly variable and is a
    mature after 1.5 years (and at 15-35 mm SL), but this   dominant factordetermining population size (Paul and
    depends upon location (Paul and Feder 1976, Ricketts    Feder 1973, 1976). Many environmental conditions

                                                       40

                                                                                Pacific littleneck clam continued

    affect successful settlement, such as temperature,    Fraser, C. M., and G. M. Smith. 1928. Notes on the
    adequate food supply, predation, currents, beach    ecology of the littleneck clam, Paphiastaminea Conrad.
    topography, and appropriate substrate (Paul and Feder    Trans. Roy. Soc. Can. 3(22):249-269.
    1973, Peterson 1982). High siltation caused by upland
    development and construction of marinas can cause    Freese, J. L., and C. E. O'Clair.  1987.  Reduced
    problems (Schink et al. 1983).  Dredging has been    survival and condition of the bivalves Protothaca
    shown to affect subtidal populations.  For example,    staminea and Mytilus edulis buried by decomposing
    mechanical clam harvesters may adversely affect   bark. Mar. Env. Res. 23:49-64.
    populations by suspending and depositing fine
    sedimentsthat can smotherclams (Schink et al.1983).    Frey, H. W. 1971. California living marine resources
    Similarly, severe weather often affects intertidal    and their utilization.  Calif. Dept. Fish Game,
    populations by producing high freshwater run-off that    Sacramento, CA, 148 p.
    kills clams by covering them with sediment or washing
    away sediments and exposing them (Cheney and    Gates, D. E., and H. W. Frey.  1974.  Designated
    Mumford 1986). "Winterkills"causedbylowsalinities,    common names of certain marine organisms of
    lowtemperatures, and microbial diseases mayoccurin    California. Calif. Fish Game, Fish Bull. 161:55-90.
    northern latitudes (Schink et al. 1983, Cheney and
    Mumford 1986).                                          Glude, J. B. 1978. The clams genera Mercenaria,
                                                           Saxidomus, Protothaca, Tapes, Mya, Panopea, and
    References                                              Spisula: A literature review and analysis of the use of
                                                           thermal effluent in the culture of clams. Unpubl. Rep.
    Abbott, R. T. 1974. American seashells: The marine    to Tenn. Valley Authority, J. B. Glude, Seattle, WA,
    mollusca of the Atlantic and Pacific coasts of North    74 p.
    America, 2nd edition. Van Nostrand Reinhold Co., NY,
    663 p.                                                  Goodwin, L., and W. Shaul.  1978.  Puget Sound
                                                           subtidal hardshell clam survey data. Prog. Rep. 44,
    Bernard, F. R. 1983a. Catalogue of the living bivalvia    Wash. Dept. Fish., Olympia, WA, 92 p.
    of the eastern Pacific Ocean: Bering Strait to Cape
    Horn. Can. Spec. Publ. Fish. Aquat. Sci. No. 61,        Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: The
    102 p.                                                  clams and allies. In R. H. Morris, D. P. Abbott, and E.
                                                           C. Haderlie (editors), Intertidal invertebrates of
    Bernard, F. R. 1983b. Physiology and the mariculture    California, p. 355-411. Stanford Univ. Press, Stanford,
    of some northeastern Pacific bivalve molluscs. Can.    CA.
    Spec. Publ. Fish. Aquat. Sci. No. 63, 24 p.
                                                           Hancock, D. R., T. F. Gaumer, G. B. Willeke, G. P.
    Boulding, E. G., and T.K. Hay. 1984. Crab response    Robart,and J. Flynn. 1979. Subtidalclam populations:
    to prey density can result in density-dependent mortality   distribution, abundance, and ecology. Sea Grant Coll.
    of clams. Can. J. Fish. Aquat. Sci. 41:521-525.         Prog. Publ. No. ORESU-T-79-002, Oregon State Univ.,
                                                           Corvallis, OR, 243 p.
    California Department of Fish and Game. 1987. 1987
    California sport fishing regulations. Calif. Dept. Fish    Kozloff, E. N. 1983.  Seashore life of the northern
    Game, Sacramento, CA, 12 p.                             Pacific coast. Univ. Wash. Press, Seattle, WA, 370 p.

    Cheney, D. P., and T. F. Mumford, Jr. 1986. Shellfish    Meinkoth, N. A.  1981.  The Audubon Society field
    and seaweed harvests of Puget Sound. Wash. Sea    guidetoNorthAmericanseashorecreatures. AlfredA.
    Grant, Univ. Wash. Press, Seattle, WA, 164 p.           Knopf, New York, NY, 799 p.

    Chew, K. K., and A. P. Ma. 1987. Species profiles: life   Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    histories and environmental requirements of coastal    Nelson. 1990. Distribution and abundance of fishes
    fishes and invertebrates (Pacific Northwest) -common    and invertebrates in west coast estuaries, Volume I:
    littleneck clam.  U.S. Fish Wildl. Serv. Biol. Rep.    data summaries.   ELMR  Rep. No. 4. Strategic
    82(11.78). U.S. Army Corps Eng., TR EL-82-4, 22 p.    Assessment Branch, NOS/NOAA, Rockville, MD,
                                                           240 p.
    Fitch, J. E. 1953. Common marinebivalves of California.
    Calif. Fish Game, Fish Bull. 90, 102 p.                 Nickerson, R. B. 1977. A study of the littleneck clam
                                                           (Protothaca staminea Conrad) and the butter clam

                                                        41

    Pacific littleneck clam continued
    (Saxidomusgianteus Deshayes)ina habitat permitting   Schultze, D. L. 1986. Digest of California commercial
    coexistence, Prince William Sound, Alaska.  Proc.   fish laws. Calif. Dept. Fish Game, Sacramento, CA,
    Natl. Shellfish Assoc. 67:85-102.                      40 p.

    Oceanographic Institute of Washington. 1981. Clam    Shaw, W. N. 1986. Species profiles: life histories and
    and mussel harvesting industries of Washington State.    environmental requirements of coastal fishes and
    Oceanog. Comm.  Wash., Seattle, WA, various    invertebrates. (PacificSouthwest)-commonlittleneck
    pagination.                                             clam. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.46), U.S.
                                                           Army Corps Eng., TR EL-82-4, 11 p.
    Paul, A. J., and H. M. Feder. 1973. Growth, recruitment,
    and distribution of the littleneck clam, Protothaca    Strathmann, M. F.,A. R. Kabat, and D. O'Foighil. 1987.
    staminea, in Galena Bay, Prince William Sound, Alaska.    Phylum Mollusca, class Bivalvia. In M. F. Strathmann
    Fish. Bull., U.S. 71(3):665-677.                        (editor), Reproduction and development of marine
                                                           invertebrates of the northern Pacific coast, p. 309-353.
    Paul, A. J., and H. M. Feder. 1976. Clam, mussel, and    Univ. Wash. Press, Seattle, WA.
    oyster resources of Alaska. Sea Grant Rep. 76-6,
    Univ. Alaska, Fairbanks, AK, 41 p.                     Washington Department of Fisheries. 1986. 1986-
                                                           1987 salmon, shellfish, bottomfish sport fishing guide.
    Peterson, C. H. 1982. The importance of predation    Wash. Dept. Fish., Olympia, WA, 20 p.
    and intra- and interspecific competition in the population
    biology of two infaunal suspension-feeding bivalves,    Wolotira, R. J., Jr., M. J. Allen, T. M. Sample, C. R. Iten,
    Protothaca staminea  and Chione undatella.  Ecol.   S. F. Noel, and R. L. Henry. 1989. Life history and
    Monog. 52(4):437-475.                                   harvest summaries for selected invertebrate species
                                                           occurring off the west coast of North America. Volume
    Peterson, C. H., and M. L. Quammen. 1982. Siphon    1: shelled molluscs. NOAA Tech. Memo.  NMFS F/
    nipping: its importance to small fishes and its impact on    NWC-160,177 p.
    growth of the bivalve Protothaca staminea (Conrad).
    J. Exp. Mar. Biol. Ecol. 63:249-268.

    Quayle, D. B. 1943. Sex, gonad development and
    seasonal gonad changes in Paphia staminea Conrad.
    J. Fish. Res. Board Can. 6(2):140-151.

    Quayle, D. B., and N. Bourne. 1972. The clam
    fisheries in British Columbia. Fish. Res. Board Can.,
    Bull. No. 179, 71 p.

    Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W.
    Phillips. 1985. Between Pacific tides. Stanford Univ.
    Press, Stanford, CA, 652 p.

    Robinson, A. M., and W. P. Breese. 1982. The
    spawning season of four species of clams in Oregon.
    J. Shellfish Res. 2(1):55-57.

    Roesijadi, G. 1980. Influence of copper on the clam
    Protothaca staminea: Effects on gills and occurrence
    of copper-binding proteins. Biol. Bull. 158:233-247.

    Schink, T. D., K. A. McGraw, and K. K. Chew. 1983.
    Pacific coast clam fisheries. Wash. Sea Grant Prog.,
    Tech. Rep. WSG 83-1, Univ. Wash., Seattle, WA, 72p.

    Schmidt, R. R., and J. E. Warme. 1969. Population
    characteristics of Protothaca staminea (Conrad) from
    Magu Lagoon, California. Veliger 12(2):193-199.

                                                        42

    43

        Venerupis japonica
       Adult

                                        2cm
    Common Name: Manila clam                              recreational diggers because of its good taste and
    Scientific Name: Venerupisjaponica                    ease of capture (Chew 1989). It is one of the most
    Other Common Names: Japanese cockle, Japanese    important recreationally dug clams on the Pacific coast
    littleneck, Manila cockle, Manila littleneck, Philippine    (Wolotira et al. 1989). Clammers harvest Manila dams
    cockle, steamer, asari (in Japan) (Cahn 1951, Chew    year-round during low tide periods by hand or using a
    1989)                                                 fork, pick, rake, shovel, or garden trowel (Frey 1971,
    Classification (Bernard 1983a)                        Wolotira et al. 1989). It is so heavily harvested in some
    Phylum: Mollusca                                      areas of Puget Sound, Washington, that it has been
    Class: Bivalvia                                       almost eliminated (Williams 1980a). Sport harvesting
    Order: Veneroida                                      of this species does occur in San Francisco Bay,
    Family: Veneridae                                     California, despite the possibility of harvesting clams
                                                          contaminated by urban wastes and the lack of official
    Value                                                 authorization (Nichols and Pamatmat 1988).
    Commercial: The Manila clam is the second-most
    important commercial clam species on the Pacific   Indicator of Environmental Stress: The Manila clam is
    coast of North America. It is primarily sold as a fresh   highly tolerant of pollution (Fitch 1953) and it may
    product. About 500 t have been landed annually in   accumulatelargeamountsofpollutantsthatareharmful
    Washington since 1975 (Schink et al. 1983, Chew    to humans.  Hence, many waters are closed to the
    1989).  Presently, only a limited commercial Manila    harvest of this species due to urban waste water and
    clam harvest exists in California or Oregon. Nearly all   industrial contamination (primarily coliform bacteria).
    Pacific coastcommercial harvest of this species comes    Only recently have limited areas in San Francisco Bay
    fromWashington and British Columbia. InWashington,    been open for Manila clam harvest.
    it is harvested year-round by diggers using forks,
    rakes, clam hacks, and hydraulic dredges (Wolotira et   Ecological:The Manila clam was introduced accidentally
    al. 1989). This harvest occurs on private and state tide   to the Pacific coast of North America probably around
    lands, for which diggers pay a royalty or "stumpage    the 1930s with Pacific oysters (Crassostrea gigas)
    fee" according to the weight landed (Chew 1989).    imported from Japan. It was first reported from British
    Harvest of this species is often aligned with oyster   Columbia in 1936 (Quayle 1938). It is often one of the
    growers, who also participate in a Manila clam fishery   most abundant bivalves in estuarine intertidal habitats,
    (Chew 1989). Minimum commercial size is 38 mm    and the dominant intertidal bivalve in San Francisco
    shell length (SL) (Frey 1971, Wolotira et al. 1989).    Bay (Frey 1971). Because its preferred distribution is
    Becauseof strong marketdemands andgood biological   in the upper tidal zone, it is not believed to have
    attributes, aquacultureofthisspecieshasbeeninitiated   displaced any native species (Bourne 1982).  The
    (Anderson et al. 1982).                               Manila clam often occurs with Pacific littleneck clam
                                                           (Protothaca staminea), butter clam (Saxidomus
    Recreational: This species is highly prized by    giganteus), softshell (Mya arenaria), Macoma spp.

                                                       44

                                                                                         Manila clam continued

                                                          in some areas of San Francisco Bay, but not in other
     Table 1. Relative abundance of Manila clam
              Table in 32 U.S. Pacific coast estuariesae oCalifornia estuaries. Oregon has had little success
                                                          with establishing and increasing Manila clam
                       Life Stage                        populations in the state's estuaries. Aquaculture of this
          Estuary    A S J L E                           species is presently being conducted in Humboldt Bay,
           PugetSound *� � �  �6   Relative abundance:   California, Puget Sound, and other estuaries.
            Hood Canal 1] t 9 1 i    �*  Highly abundant
             Skagit Bay O 0 0 0 O        Abundant        Life Mode
           Grays Harbor     X     RareCommon              Eggs and larvae are pelagic. Juveniles and adults are
                                      4   Rare
            WillapaBay � � � � �   Blank Not present      benthic infauna, occurring just below the sediment
          Columbia River                                  surface down to about 5 cm (sometimes to 10 cm)
           Nehalem Bay                                    (Bourne 1982, Wolotira et al. 1989).
           Tillamook Bay O O O O O    Life stage:
            Netarts Bay 0:  0 0    0 0     A-Adults       Habitat
                                      S - Spawning adults
            Siletz River             J-Juveniles        ITY: It is found from the intertidal zone to depths of
            Yaquina Bay               L - Larvae          about 10 m (Wolotira et al. 1989), but is primarily found
            Alsea River                                  at 0.9-2.4 m above mean lower low water (MLLW)
           Siuslaw River                                  (Quayle and Bourne 1972). It is not found subtidally in
           Umpqua River                                   British Columbia (Bourne 1982).
              Coos Bay  0 0 0 00
            Rogue River                                   Substrate: An ideal substrate appears to consist of
           Klamath River                                  gravel (much of which is <25 mm in diameter), sand,
           Humboldt Bay ci O    ii 0                      some mud (4-5%), and shell (Anderson et al. 1982).
              Eel River                                   Beaches having this type of substrate are often relatively
           Tomales Bay 0 0 0 0 o                         stable, and occur in many protected areas of Pacific
      CentSanFran. Bay  * �  �  �    �   IncdudesCentralSan  Northwest inlets and bays (Chew 1989). However,
                                    SFrancisco, Suisun,
      South San Fran. Bay �*. � �   and San Pablo bys.    Manila clams can inhabit a wide range of substrates.
          Elkhorn Slough  '    '                          Dense concentrations of Manila clams have been
             Morro Bay                                    found in substrates ranging from primarily sand (Cahn
        Santa Monica Bay                                  1951, Ohba 1959) to mud. Additions of pea gravel and
          San Pedro Bay                                   small rock on Manila clam beds can enhance settlement
           Alamitos Bay                                   (Chew 1989).
           Anaheim Bay
            Newport Bay                                   Physical/Chemical Characteristics: The Manila clam is
            Mission Bay                                   found in mesohaline-euhaline waters (Haderlie and
          San Diego Bay                                   Abbott 1980). Optimum salinities for larval development
          Tijuana Estuary                                 are 20-30%o (Robinson and Breese 1984). Optimum
                      A S J L E                           temperatures for larval development are 23-25�C, but
                                                          they can withstand temperatures of 0-36�C (Cahn
    clams, and other estuarine infauna (Wolotira et al.   1951, Robinson and Breese 1984). Optimum conditions
    1989). Pinnotherid crabs (Pinnixa fabaand P. littoralis)   for adult and juvenile growth are 28%� salinity (range of
    are common commensals within the mantle cavity of   24-31%o), 16�C temperature (range of 13-21 �C), and a
    Manila clams (Haderlie and Abbott 1980).              food suspension density of 55 mg/I (ranges 10-135 mg/
                                                           I) (Bernard 1983b). Prolonged salinities below 10%o
    Range                                                 are lethal (Bardach et al. 1972). Optimum tidal level
    Overall:TheManilaclamisatropical-temperatewestern    appears to be 1.5-2.5 m above MLLW (Quayle and
    Pacific species, originally found from the Philippines    Bourne 1972, Glock and Chew 1979). Small clams do
    and China north along Japan to the southern Sea of   notappeartogrowduringthewinterwhentemperatures
    Okhotsk (Wolotira et al. 1989).  It now occurs on    are <10�C (Bardach et al. 1972, Glock 1978, Williams
    eastern Pacific shores from Elkhorn Slough, California    1980a). The Manila clam requires temperatures >14-
    to British Columbia (Fitch 1953), and is also found in   15�Cformaturation, spawning, and larval development
    Hawaii (Morris 1966).                                  (Holland and Chew 1974, Mann 1979, Bourne 1982).
                                                           Juvenile and adult clams require maximum summer
    Within Studv Area: The Manila clam is abundant in   temperatures greater than about 12�C to survive
    Washington estuaries, but is not commonly found in   (Bourne 1982). Steeply-sloped beaches are not good
    many Oregon estuaries (Table 1). It is highly abundant    Manila clam habitat (Miller 1982, Chew 1989). Waves

                                                       45

    Manila clam continued
    and water currents playa major role in regulating clam    mature at 15 mm SL (Ko 1957, Holland and Chew
    productivity. Currents removewaste, supply food and    1974).  Growth rates vary considerably among
    oxygen, distribute spat, and may redistribute young    geographic locations. One-year-oldclams are reported
    clams (Miller 1982, Chew 1989).                       to be 8 mm SL in Hokkaido, 18 mm SL in the Inland Sea
                                                          (Ohba 1959), 27 mm SL in southern Japan (Tanaka
    Miarations and Movements: Larvae are carried by    1954), 24 mm SL in Hood Canal, Washington (Nosho
    currents into appropriate areas for settlement.    and Chew 1972), and 10-15 mm SL in the Strait of
    Convergences and eddies often concentrate larvae.   Georgia, British Columbia (Quayle and Bourne 1972).
    Larvae attach a byssus thread to a pebble or shell   Growth is also dependent upon the tidal level clams
    during settlement (Cahn 1951, Nosho 1971, Quayle    inhabit, with growth often lower at higher tidal levels
    and Bourne 1972).                                     (Chew 1989).  Clams take 16-22 months to reach
                                                          market size in Washington (Glock 1978), and about 24
    Reproduction                                          months in California (Frey 1971). However, they may
    Mode: The Manila clam is gonochoristic, oviparous,    need 3-4 years before reaching legal size in British
    and iteroparous. It is a broadcast spawner, expelling   Columbia (Bourne 1982).  Manila clams also grow
    gametes from the exhalant siphon; eggs are fertilized    more slowly in overcrowded conditions (Haderlie and
    externally.                                           Abbott 1980). The maximum age is probably 7-10
                                                          years (Frey 1971).
    Matina/Spawning: In Japan, spawning occurs both in
    the spring and autumn (Chew 1989).  In Kasaoka,    Food and Feeding
    Japan the Manila clam spawns from early May to July   Trophic Mode: The Manila clam is a nonselective
    and then again between early November and late   suspension/filter feeder.  Food particles are inhaled
    December (Chew 1989).  Other Japanese studies   with water through the inhalant siphon, trapped by the
    reveal spawning times from early March to mid-May   gill, sorted by the palps, and passed to the mouth
    and from late October to early November (Yasuda et al.   (Wolotira et al. 1989).
    1945, Ko 1957). In Washington's waters, the Manila
    clam spawns once per year, usually between May and    Food Items: Food consists of suspended detritus and
    September (typically peaking during June and July)   phytoplankton.
    (Nosho and Chew 1972, Holland and Chew 1974).
    Spawning apparently does not take place at water    Biological Interactions
    temperatures below 15�0C (Mann 1979).                 Predation: Important predators include:the moonsnails
                                                          (Polinicesspp.), rock crabs (Cancerspp.), shore crabs,
    Fecundity: Unknown.                                   rock sole (Lepidopsetta bilineata), English sole
                                                          (Pleuronectes vetulus), starry flounder (Platichthys
    Growth and Development                                stellatus), pile perch (Rhacochilus vacca), shiner perch
    Eaa Size and Embrvonic DeveloDment: Eggs are    (Cymatogaster aggregata), starfish (Pisaster spp.),
    spherical and 0.06 mm in diameter (Wolotira et al.   ducks, and scoters (Cahn 1951, Glude 1964, Bardach
    1989). Embryonicdevelopmentisindirectandextemal.    et al. 1972, Quayle and Bourne 1972, Anderson et al.
                                                          1982, Chew 1989). Nematodes and other meiofaunal
    Aae and Size of Larvae: Larvae rangefrom 0.06 mmto    predators may prey heavily on newly-setting spat
    0.19-0.24 mm  in length (Wolotira et al. 1989).  A    (Williams 1980a).
    ciliated, motile, trochophore larvae forms within 24-48
    hours after fertilization at 13-16�C. The veliger needs    Factors Influencina PoDulations: Spat settlement areas
    about 3-4 weeks before metamorphosing to spat (setting   are dependent on currents and substrates (Chew 1989).
    juveniles) (Cahn 1951, Quayle and Bourne 1972,    Wave damage, extreme temperatures, and siltation
    Bourne 1982).  The duration of larval stages is   can adversely affect population sizes (Bardach et al.
    dependent ontemperature and food availability (Chew    1972, Chew 1989). Extreme substrate temperatures
    1989).                                                during winter and summer are potentially lethal (Chew
                                                          1989). High densities of adult clams may decreasethe
    Juvenile Size Ranae: At settlement, clams range from   ability of spat to settle (Williams 1980a, 1980b). Most
    0.190-0.235 mm SL (Williams 1978, 1980a), and reach    mortality appears to occur within the first two months
    15mmSL(range:12-20mm)beforebecomingsexually    after settlement (Williams 1980a, 1980b). Losses of
    mature (Ko 1957, Nosho and Chew 1972, Holland and    newly settled spat are probably a result of predation,
    Chew 1974, Wolotira et al. 1989).                     starvation, and climatic conditions. Because of good
                                                          market conditions, numerous aquaculture ventures
    Aae and Size of Adults: Some Manila clams may    are being established or considered (Anderson et al.

                                                       46

                                                                                          Manila clam continued

    1982). This species' northern distribution is probably    (Deshayes) at Squaxin Island, Washington.  Proc.
    limited by cold water temperatures (Bourne 1982). Its   Natl. Shellfish. Assoc. 69:15-20.
    southern distribution may be limited bythe high salinities
    and substrate structure of southern California bays and    Glude, J. B. 1964. The effect of scoter duck predation
    estuaries. Plastic netting placed on beaches improves    on a clam population in Dabob Bay, Washington. Proc.
    settlement and growth (Glock 1978, Glock and Chew    Natl. Shellfish. Assoc. 55:73-86.
    1979).
                                                           Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: The
    References                                              clams and allies. In R. H. Morris, D. P. Abbott, and E.
                                                           C. Haderlie (editors), Intertidal invertebrates of
    Anderson, G. J., M. B. Miller, and K. K. Chew. 1982. A    California, p. 355-411. Stanford Univ. Press, Stanford,
    guide to Manila clam aquaculture in Puget Sound.    CA.
    Wash. Sea Grant, Univ. Wash., Seattle, WA, 45 p.
                                                           Holland, D. A., and K. K. Chew. 1974. Reproductive
    Bardach, J. E., J. H. Ryther, and W. O. McLarney.    cycle of the Manila clam (Venerupis japonica) from
    1972. Aquaculture - the farming and husbandry of    Hood Canal, Washington. Proc. NatI. Shellfish. Assoc.
    freshwater and marine organisms.  John Wiley and    64:53-58.
    Sons, New York, NY, 868 p.
                                                           Ko, Y. 1957. Some histological notes on the gonads
    Bernard, F. R. 1983a. Catalogue of the living Bivalvia   of Tapes japonica Deshayes. [in Japanese, English
    of the eastern Pacific Ocean: Bering Strait to Cape    summary]. Bull. Jap. Soc. Sci. Fish. 23(7/8):394-399.
    Horn. Can. Spec. Publ. Fish. Aquat. Sci. 61, 102 p.
                                                           Mann, R. 1979. The effect of temperature on growth,
    Bernard, F. R. 1983b. Physiology and the mariculture    physiology and gametogenesis in the manila clam,
    of some northeastern Pacific bivalve molluscs. Can.    Tapesphilippinarum Adams and Reeve 1850. J. Exp.
    Spec. Publ. Fish. Aquat. Sci. 63, 24 p.                 Mar. Biol. Ecol. 38:122-133.

    Bourne, N.  1982.  Distribution, reproduction, and    Miller, M. B. 1982. Recovery and growth of hatchery-
    growth of Manila clam, Tapes philippinarum (Adams    produced juvenile Manila clams, Venerupis japonica
    and Reeves) in British Columbia.  J. Shellfish Res.    (Deshayes)planted on several beaches in Puget Sound.
    2(1):47-54.                                             Ph.D. Thesis, Univ. Wash, Seattle, WA, 250 p.

    Cahn, A. R. 1951. Clam culture in Japan. U.S. Fish    Morris, P. A. 1966. Afield guideto Pacificcoast shells.
    Wildl. Serv., Fish Leafl. No. 299, 103 p.               Houghton-Mifflin Co., Boston, MA, 297 p.

    Chew, K. K. 1989. Manila clam biology and fishery    Nichols, F. H., and M. M. Pamatmat.  1988.  The
    development in western North America. InJ. J. Manzi    ecology of the soft-bottom benthos of San Francisco
    and M. Castagna (editors), Clam mariculture in North    Bay: a community profile. U.S. Fish Wildl. Serv. Biol.
    America, p. 243-261. Dev. Aquat. Fish. Sci., Vol. 19.    Rep. 85(7.19), 73 p.
    Elsevier Press, New York, NY.
                                                           Nosho, T. Y. 1971. The setting and growth of the
    Fitch, J. E. 1953. Common marinebivalves of California.    Manila clam, Venerupisjaponica (Deshayes) in Hood
    Calif. Fish Game, Fish Bull. 90, 102 p.                 Canal, Washington. M.S. Thesis, Univ. Wash., Seattle,
                                                           WA, 67 p.
    Frey, H. W. 1971. California's living marine resources
    and their utilization.  Calif. Dept. Fish Game,    Nosho, T. Y., and K. K. Chew. 1972. The setting and
    Sacramento, CA, 48 p.                                   growth of the Manila clam,  Venerupis japonica
                                                           (Deshayes) in Hood Canal, Washington. Proc. Natl.
    Glock, J. W. 1978. Growth, recovery, and movement    Shellfish. Assoc. 62:50-58.
    of Manila clams, Venerupis japonica planted under
    protective devices and on open beaches at Squaxin    Ohba, S.  1959.  Ecological studies in the natural
    Island, Washington. M.S. Thesis, Univ. Wash., Seattle,    population of a clam, Tapes japonica, with special
    WA, 66 p.                                               referenceto seasonal variations in the size and structure
                                                           ofthe population and individual growth. Biol. J. Okayama
    Glock, J. W., and K. K. Chew. 1979. Growth, recovery,    Univ. 5(1/2):13-42.
    and movement of Manila clams, Venerupis japonica

                                                        47

    Manila clam continued
    Quayle, D. B. 1938. Paphia bifurcata, a new molluscan
    species from Ladysmith Harbor, B.C. J. Fish. Res.
    Board Can. 4:53-54.

    Quayle, D. B., and N. Bourne. 1972. The clam
    fisheries in British Columbia. Fish. Res. Board Can.
    Bull. No. 179, 70 p.

    Robinson, A. M., and W. P. Breese. 1984. Gonadal
    development and hatchery rearing techniques for the
    Manilaclam Tapesphilippinarum (Adams and Reeve).
    J. Shellfish Res. 4(2):161-163.

    Schink, T. D., K. A. McGraw, and K. K. Chew. 1983.
    Pacific coast clam fisheries. Wash. Sea Grant, Univ.
    Wash., Seattle, WA, 72 p.

    Tanaka, Y. 1954. Spawning season of important
    bivalves in Ariake Bay - Venerupis semidecussata
    (Reeve). [In Japanese, English summary]. Bull. Jap.
    Soc. Sci. Fish 19(12):1165-1167.

    Williams, J. G. 1978. The influence of adults on the
    settlement, growth, and survivals of spat in the
    commercially important clam, Tapes japonica
    Deshayes. Ph.D. Thesis, Univ. Wash., Seattle, WA,
    60 p.

    Williams, J. G. 1980a. Growth and survival in newly
    settled spat of the Manila clam, Tapes japonica. Fish.
    Bull., U.S. 891-900.

    Williams, J. G. 1980b. The influence of adults on the
    settlement of spat of the clam, Tapes japonica. J. Mar.
    Res. 38(4):729-741.

    Wolotira, R. J., Jr., M. J. Allen, T. M. Sample, C. R. Iten,
    S. F. Noel, and R. L. Henry. 1989. Life history and
    harvest summaries for selected invertebrate species
    occurring offthewest coast of North America. Volume
    1: shelled molluscs. NOAA Tech. Memo. NMFS F/
    NWC-1 60, 177 p.

    Yasuda, J., I. Hamai, and H. Hotta. 1945. A note on the
    spawning season in Venerupis philippinarum. Bull.
    Jap. Soc. Sci. Fish. 290(4):277-279.

                                                       48

    49

       Mya arenaria
       Adult

                                  2 cm

    Common Name: softshell                                Extension Service et al. 1976). In general, this species
    Scientific Name: Mya arenaria                         is underutilized by sport diggers because of the
    Other Common Names: soft clam, long clam, mud    abundance of more desirable species.
    clam, sand clam, common mya, nanninose, eastern
    softshell clam, softshell clam, steamer clam, long-   Indicator of Environmental Stress: The softshell often
    necked clam, sand gaper (Fitch 1953, Gates and Frey   occurs in estuarine areas where industrial and domestic
    1974, Newell and Hidu 1986)                          pollution problems first occur and theclams then become
    Classification (Bernard 1983)                         unsafe to consume. Many areas (e.g., San Francisco
    Phylum: Mollusca                                      Bay, California) that have harvestable numbers of M.
    Class: Bivalvia                                       arenaria are presently closed to harvesting due to
    Order: Myoida                                         pollution. However, this species is relatively tolerant of
    Family: Myidae                                        pollution. The softshell accumulates crude oil into its
                                                          lipid-containing tissues when oil is in low concentrations
    Value                                                 (90-380 jag oil/liter) (Fong 1976). It also concentrates
    Commercial: The softshell is not as valuable as some    heavy metals in its tissues.  However, at water
    other bivalves along the Pacific coast, but may be   temperatures of 22.0�C and salinities of 30.0%oo, the
    underutilized in Washington.  Over 181 t were    following concentrations caused death in 50% of the
    commercially harvested in Washington in 1985    testclamswithin 96 hours:copper, 0.039mg/l; cadmium,
    (Washington Department of Fisheries 1985). It has    0.850 mg/l;zinc, 5.2 mg/l; lead, 27.0 mg/I; manganese,
    been estimated that 900 t could be harvested annually    >300.0 mg/I; and nickel, >50.0 mg/I (Eisler 1977).
    in Skagit Bay and Port Susan, Washington (Cheney
    and Mumford 1986). About 34 t were harvested in   Ecological: The softshell was probably introduced to
    Oregon in 1980, but in California this species has not   the Pacific coast before 1874, perhaps in 1869 when
    been harvested since about 1948 (Skinner 1962, Schink    the first eastern oysters were introduced. However,
    et al. 1983). The limited commercial harvest of this   there is some evidence that softshell clams were once
    species in Oregon and California occurs because of   native to the Pacific coast (Porter 1974). This species
    small population sizes (Oregon) and pollution   is common in estuaries from Elkhorn Slough, California,
    (California) (Schink et al. 1983).  This species is   to Alaska (Ricketts et al. 1985), and may have crowded
    harvested primarilyby hydraulicescalatordredge(Kyte    out the native Macoma species in some areas of the
    and Chew 1975 ).                                      Pacific coast (Rudy and Rudy 1983).

    Recreational:This is an important clam forsport diggers.    Range
    In some areas of Washington over 9.1 kg/day are   Overall: In the Atlantic, it is found along the coast of
    allowed to be dug perperson (Washington Department    North America from Labrador to Cape Hatteras, North
    of Fisheries 1986). Oregon permits sport diggers to   Carolina, and less commonly to South Carolina. In
    harvest 36 clams/day (Oregon State University    Europe, it occurs from northern Norway to the Bay of

                                                        50

                                                                                          Softshellcontinued

                                                           adults are benthic infauna.
     Table 1. Relative abundance of softshell in
              32 U.S. Pacific coast estuaries.             Habitat
                       Life Stage                        ITye: The softshell is a true estuarine organism, with
          Estuary    A S J L E                            all life stages occurring there. A euryhaline species, it
           Puget Sound e1   s  ]   Relative abundance:     is found primarily in mesohaline and polyhaline water.
            Hood Canal O 0  0  0     �*  Highly abundant   Eggs and larvae are found in the estuarine and
             Skagit Bay *� � � � �  Abundant               nearshore marine plankton, while juveniles and adults
           Grays Harbor &9 W (        0 '4� Common         occur primarily in quiet estuarine mud flats that are
            WillapaBay S  3 5 (3      Blank Notpresent     near river mouths where  low salinity occurs
         Columbia River O O O O O                          (Oceanographic Institute of Washington 1981, Newell
           Nehalem Bay �  �  �  � a                        and Hidu 1986). Adults and juveniles are often most
          Tillamook Bay * O    1 O O    Life stage:        abundant in the upper mid-tidal zone [+1.8 to 0.6 feet
            NetarnsBay O O   �           Spawning  adults  meanlower low water(MLLW)] (Cheney and Mumford
            SilelzRiver     V ' ' 4  J - JJuveniles        1986), but they can occur down to approximately -5.5
           YaquinaBay       �        La� �  rvaggse       to -9 m MLLW (Filice 1958, Meinkoth 1981). Adults
            Alsea River     o  Eg g s                      may be found buried in sediments down to 25-30 cm
           Siuslaw River  *-56U                            (Haderlie and Abbott 1980, Abraham and Dillon 1986).
          Umpqua River *    ...
             coos Bay � � � � �                            Substrate: Adults and juveniles prefer medium to soft
            Rogue River                                     substrates, consisting primarily of sand, compact clays,
          Klamath River                                    coarse gravel, a mixture of sand and mud, and gravel
          HumboldtBay i) S S { S                           and mud (Cheneyand Mumford 1986, Newell and Hidu
              EelRiver O O O 0 0                           1986). However, they are often found in thick, dark
           Tomales Bay O O O CO                            mud (Haderlie and Abbott 1980) that may consist of up
      Cent. San Fran. Bay  O    X (0 5       '   IncludesCentralSan  to 50% silt (Abraham and Dillon 1986). Adults and
      SoulthSan Fran. Bay  O S S a S   and ann Pablo bays.  juveniles cannot burrow or maintain themselves in
          Elkhom Slough 0D 0 0 C0                          shifting substrates (Ricketts et al. 1985). Growth rates
             Morro Bay  i i  4 i '4                        andshellformaredependentonthesubstrateproperties
        Santa Monica Bay                                    (Newell and Hidu 1982).
          San Pedro Bay
           Alamitos Bay ::                                 Phvsical/Chemical Characteristics: The softshell is a
           Anaheim Bay                                     euryhaline species. Adults can tolerate salinities down
           Newport Bay                                     to 5%o, but larvae are more sensitive to low salinities
            Mission Bay                                    (Newell and Hidu 1986). Adult clams on the Atlantic
          San Diego Bay                                    coast have preferred salinities that decrease north to
          Tijuana Estuary                                   south (Newell and Hidu 1986); it is not known if this is
                      A S J L E                            trueforPacificcoastpopulations. Juvenileclamsalinity
                                                           tolerances are related to size; larger juveniles can
    Biscay, France. In the eastern Pacific, it occurs from    withstand lower salinities.  The ability to withstand
    Monterey Bay (maybe San Diego), California, through    extremely low salinities is inversely related to
    Alaska (Gross 1967, Paul and Feder 1976, Rudy and    temperature.  Temperature also controls timing of
    Rudy 1983, Abraham and Dillon 1986), and is also    spawning and influences distribution. The northern
    found along the western Pacific coast from the    rangeof M. arenaria is limited bytemperaturestoo low
    Kamchatka PeninsulatothesouthernJapaneseislands    for spawning, while southern distribution is limited by
    (Hanks 1963). It is apparently still extending its range    high temperatures (Laursen 1966).  Temperatures
    as seen by its expansion into the Black Sea (Ivanov    above 280C can affect its distribution and abundance
    1969, Porter 1974).                                     (Newell and Hidu 1986). However, it can withstand
                                                           temperatures down to at least -1.7�C (Newell and Hidu
    Within Studv Area: The softshell is commonly found    1986). The softshell clam can function as a facultative
    from Elkhorn Slough, California, north through    anaerobe at lowtide (Collip 1920), surviving anaerobic
    Washington's estuaries (Table 1) (Haderlie and Abbott    conditions longer at lower temperatures (Newell and
    1980, Kozloff 1983, Ricketts et al. 1985).              Hidu 1986). Spawningtemperaturesdependon latitude
                                                           and location, ranging from about 4�C to22�C. Spawning
    Life Mode                                               on the Pacific coast appears to occur at temperatures
    Eggs and larval stages are planktonic; juveniles and    between 10 and 15�C (Simel 1980).  This species

                                                        51

    Softshellcontinued
    prefers to orient its siphon perpendiculartothe principal   on temperature before transforming into a spat, which
    component of water currents (Vincent et al. 1988).    has a muscular foot, byssal gland, no velum, and
                                                          settles out of the water column (Abraham and Dillon
    MiarationsandMovements: Planktoniceggs and larvae    1986).  Initially, veliger larvae are about 80 I.m in
    are dispersed by waves and currents.  Newly-    diameter and most metamorphose to spat soon after
    metamorphosed spat may spend 2-5 weeks floating   reaching 200 I.m (Loosanoff et al. 1966).
    and crawling. During this time, the spat uses a byssal
    thread to hold on to various substrates, such as eelgrass    Juvenile Size Ranae: Juveniles grow from 0.2 mm shell
    (Zostera spp.), filamentous algae, and other objects.    length (SL) (newly-settled spat) upto 25.0-45.0 mm SL
    Eventually the spat finds a favorable location where it  before maturing (Porter 1974).
    drops to the bottom and burrows into the sediment.
    Initially spat settle primarily in lower intertidal and    Aae and Size of Adults: The softshell may reach
    subtidal areas, but as they grow they may move    maturity at one year and 27-34 mm SL (Brosseau and
    shoreward. This shoreward movement is believed to   Baglivo 1988); adults may reach commercial size (50-
    be caused primarily by shoaling wave sorting    75 mm SL) in 2-3 years in Washington (Oceanographic
    (Matthiessen 1961, Newell and Hidu 1986). Clams up    Institute of Washington 1981, Cheney and Mumford
    to 12-13 mm in diameter will wander (Smith 1955),    1986), but may reach this size earlier in Oregon and
    while larger clams are sedentary.                     California. Growth is slower during winter and faster
                                                          during early spring and summer, but is modified by
    Reproduction                                          sediment type, tidal level, population densities, and
    Mode: The softshell clam is gonochoristic (but some    food abundance (Newell and Hidu 1986, Brousseau
    hermaphroditism has been reported), oviparous, and    and Baglivo 1987). Softshells have been reported to
    iteroparous.  It is a broadcast spawner; eggs are    liveupto28years (MacDonaldandThomas 1980), but
    fertilized externally (Porter 1974, Brousseau 1978,    10-1 2years is morelikelythe maximum age (Brousseau
    Brousseau 1987).                                      1978, Brousseau and Baglivo 1987).

    Matina/SDawnina:There areonlytwopublished records    Food and Feeding
    of softshell spawning times on the Pacific coast; one    Trophic Mode: Larvae, juveniles, and adults are
    from Skagit Bay, Washington (Porter 1974) and the   planktivorous filter feeders, trapping and ingesting
    other from Humboldt Bay, California (Simel 1980).    food particles via mucus on the gill tissues.
    Similar to northern Atlantic coast populations (Ropes
    and Stickney 1965, Brousseau 1987), M. arenaria in   Food Items: Trochophores feed on various suspended
    Skagit Bay spawns one time between May and    particles,whileveligersfeedprimarilyonphytoplankton.
    September, peaking in June or July (Porter 1974). In   Adultsand juveniles preferflagellates anddiatoms, but
    Humboldt Bay, it appears to spawn at the peak of   bacteria,dissolvedorganicmaterial,andorganicdetritus
    phytoplankton abundance from late March through    are also fed upon (Abraham and Dillon 1986, Newell
    April (Simel 1980).  Males normally spawn first,   and Hidu 1986).
    producing both pheromones and sperm which stimulate
    females to spawn (Newell and Hidu 1986).              Biological Interactions
                                                           Predation: Veligers are important prey for many species
    Fecundity: Fecundity has been reported to be 3 million   of larval fish.  Jellyfish, combjellies (Holland et al.
    eggs per female per year, but may actually be 120,000    1980), and fish are efficient predators of softshell
    to 1,000,000 (Brousseau 1978, Newelland Hidu 1986).    larvae.  Important predators of spat and juveniles
                                                           include birds, fish, shrimp, polychaetes, crabs, snails,
    Growth and Development                                and flatworms. Important predators of adults include
    Eoan Size and Embrvonic DeveloDment:When released    raccoons (Procyon lotor) and otters (Enhydra lutris).
    into seawater, eggs are spherical and about 66 gim in
    diameter (Newell and Hidu 1986).   Embryonic    Factors Influencino Pooulations: Lessthan 0.1 %of the
    development is indirect and external. Fertilized eggs    eggs produced during a spawning season successfully
    may take 12 hours to develop into the trocophore (a   settle, but only 1 % of the settled spat need to mature to
    top-shapedciliatedlarvae).                             maintain populations (Newell and Hidu 1986).
                                                           Extremely high densities of spat settlement have been
    Ace and Size of Larvae: The trochophore takes 24-36    observed, but densities are quickly reduced, probably
    hours to develop into a veliger, which has calcareous    due to predation. First year survivorship rates ranged
    valves and stays in the water column by its ciliated   from 24 million to 420 million at two Atlantic coast sites
    velum. The veligerstage may last 2-6 weeks, depending    (Brousseau and Baglivo 1988). Alterations of estuarine

                                                       52

                                                                                          Softshellcontinued

    habitats adversely affect populations.  Municipal    Fong, W. C. 1976. Uptake and retention of Kuwait
    sewage, industrial effluent, and estuarine development    crude oil and its effects on oxygen uptake by the soft-
    projects (e.g., dredging, pier and jetty construction)    shell clam, Mya arenaria. J. Fish. Res. Board Can.
    may all reduce softshell clam populations.              33:2774-2780.

    References                                              Gates, D. E., and H. W. Frey.  1974.  Designated
                                                           common names of certain marine organisms of
    Abraham, B. J., and P. L. Dillon.  1986.  Species    California. Calif. Fish Game, Fish Bull. 161:55-90.
    profiles: life histories and environmental requirements
    of coastal fishes and invertebrates (mid-Atlantic)-    Gross, J. B. 1967. Note on the northward spreading
    softshell clam.  U.S. Fish Wildl. Serv. Biol. Rep.    of Mya arenaria Linnaeus in Alaska. Veliger 10:203.
    82(11.68), U.S. Army Corps Eng., TR EL-82-4, 18 p.
                                                           Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: the
    Bernard, F. R. 1983. Catalogue of the living Bivalvia    clams and allies. In R. H. Morris, D. P. Abbott, and E.
    of the eastern Pacific Ocean: Bering Strait to Cape    C. Haderlie (editors), Intertidal invertebrates of
    Horn. Can. Spec. Publ. Fish. Aquatic Sci. 61,102 p.    California, p. 355-411. Stanford Univ. Press, Stanford,
                                                           CA.
    Brousseau, D.J. 1978. Spawning cycle, fecundity and
    recruitment in a population of soft-shell clam, Mya    Hanks, R. W.  1963. The soft-shell clam. U.S. Fish
    arenaria, from Cape Ann, Massachusetts. Fish. Bull.,   Wildl. Serv., Bureau Comm. Fish. Circular 162,16 p.
    U.S. 76:155-166.
                                                           Holland, A. F., N. K. Mountford, M. H. Hiegel, K. R.
    Brousseau, D. J. 1987. A comparative study of the    Kaumeyer, and J. A. Mihursky.  1980.  Influence of
    reproductive cycle of the soft-shell clam, Mya arenaria    predation on infaunal abundance in upperChesapeake
    in Long Island Sound. J. Shellfish Res. 6:7-15.         Bay, USA. Mar. Biol. 57:221-235.

    Brousseau, D. J., and J. A. Baglivo. 1987. A comparative    Ivanov, A. I. 1969. Immigration of Mya arenaria L. to
    study of age and growth in Mya arenaria (soft-shell   the Black Sea, its distribution and quantity. [In Russ.
    clam) from three populations in Long island Sound. J.   with Engl. summary], Okawnologiya 9:341-347.
    Shellfish Res. 6:17-24.
                                                           Kozloff, E. N. 1983. Seashore life of the northern
    Brosseau, D. J., and J. A. Baglivo. 1988. Life tables for   Pacific coast. Univ. Wash. Press, Seattle, WA, 370 p.
    two field populations of soft-shell clam, Mya arenaria,
    (Mollusca: Pelecypoda)from Long Island Sound. Fish.    Kyte, M. A., and K. K. Chew. 1975. A review of the
    Bull., U.S. 86(3):567-579.                              hydraulic escalator shellfish harvester and its known
                                                           effects in relation to the soft-shell clam, Mya arenaria.
    Cheney D. P., and T. F. Mumford. 1986. Shellfish and    Wash. Sea Grant, Univ. Wash., Seattle, WA, 32 p.
    seaweed harvests of Puget Sound. Wash. Sea Grant,
    Univ. Wash. Press, Seattle, WA, 164 p.                  Laursen, D. 1966. The genus Mya in the Arctic region.
                                                            Malacologia 3: 399-418.
    Collip, J. B. 1920. Studies on molluscan celomic fluid.
    Effect of change in environment on the carbon dioxide    Loosanoff, V. L., H. C. Davis, and P. E. Chanley. 1966.
    content of the celomic fluid. Anaerobic respiration in   Dimensions and shapes of larvae of some marine
    Mya arenaria. J. Biol. Chem. 45:23-39.                  bivalve mollusks. Malacologia 4:351-435.

    Eisler, R. 1977. Acute toxicities of selected heavy    MacDonald, B. A., and M. L. H. Thomas. 1980. Age
    metals to the softshell clam, Mya arenaria.  Bull.   determinationofthesoft-shellclam Myaarenariausing
    Environm. Contam. Toxicol. 17(2):137-145.               shell internal growth lines. Mar. Biol. 58:105-109.

    Filice, F. P. 1958. Invertebrates from the estuarine    Matthiessen, G. C.  1961.  Intertidal zonation in
    portion of San Francisco Bay and some factors    populations of Mya arenaria. Limnol. Ocean.5:381-
    influencingtheirdistributions. Wasmann J. Biol. 16:159-    388.
    211.
                                                            Meinkoth, N. A. 1981. The Audubon Society field
    Fitch, J. E. 1953. Common marine bivalves of California.    guide to North American seashore creatures. Alfred A.
    Calif. Fish Game, Fish Bull. 90, 98 p.                  Knopf, New York, NY, 799 p.

                                                        53

    Softshellcontinued
    Newell, C., and H. Hidu. 1982. The effects of sediment    Water Proj. Branch Rep. No. 1, Calif. Dept. Fish Game,
    type on growth rate and shell allometry in the soft    Sacramento, CA, 226 p.
    shelled clam Mya arenaria L. J. Exp. Mar. Biol. Ecol.
    65:285-295.                                              Smith, O. R.  1955.  Movements of small soft-shell
                                                            clams, (Mya arenaria). U.S. Fish Wildl. Serv., Special
    Newell, C. R., and H. Hidu. 1986. Species profiles: life   Sci. Rep. Fish. 159, 9 p.
    histories and environmental requirements of coastal
    fishes and invertebrates (North Atlantic)-softshell    Vincent, B., G. Desrosiers, and Yves Gratton. 1988.
    clam. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.53), U.S.    Orientation of the infaunal bivalve Mya arenaria L. in
    Army Corps Eng. TR EL-82-4, 17 p.                        relation to local current direction on a tidal flat. J. Exp.
                                                            Mar. Biol. Ecol. 124:205-214.
    Oceanographic Institute of Washington. 1981. Clam
    and mussel harvesting industries in Washington state.    Washington Department of Fisheries.  1985.  1985
    Oceanogr. Comm.  Wash., Seattle, WA, various    fisheries statistical report. Wash. Dept. Fish., Olympia,
    pagination.                                              WA, 101 p.

    Oregon State University Extension Service, Sea Grant    Washington Department of Fisheries.  1986. 1986-
    Marine Advisory Program, and Oregon Department of    1987 (April 1 thru March 31) salmon, shellfish, bottom
    Fish and Wildlife. 1976. Oregon's captivating clams.    fish sport fishing guide. Wash. Dept. Fish., Olympia,
    Oregon State Univ. Ext. Serv. , Sea Grant Marine    WA, 20 p.
    Advis. Prog., and Oregon Fish Wildl., Corvallis, OR,
    2p.

    Paul, A. J., and H. M. Feder. 1976. Clam, mussel, and
    oyster resources of Alaska. Inst. Marine Res. Rep. 76-
    4, Sea Grant Rep. 76-6, Univ. Alaska, Fairbanks, AK,
    40 p.

    Porter, R. G. 1974. Reproductive cycle of the soft-shell
    clam, Mya arenaria at Skagit Bay, Washington. Fish.
    Bull., U.S. 72:648-656.

    Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W.
    Phillips. 1985. Between Pacific tides. Stanford Univ.
    Press, Stanford, CA, 652 p.

    Ropes, J. W., and A. P. Stickney. 1965. Reproductive
    cycle of Mya arenaria in New England. Biol. Bull.
    (Woods Hole) 128:315-327.

    Rudy, P., Jr., and L. H. Rudy. 1983. Oregon estuarine
    invertebrates. An illustrated guide to the common and
    important invertebrate animals. U.S. Fish Wildl. Serv.,
    Biol. Serv. Prog. FWS/OBS-83/16, Portland, OR,
    225 p.

    Schink, T. D., K. A. McGraw, and K. K. Chew. 1983.
    Pacific coast clam fisheries. Wash. Sea Grant, Univ.
    Wash., Seattle, WA, 72 p.

    Simel, N. R. 1980. Aspects of the ecology of Mya
    arenaria L. in Humboldt Bay, California. M.A. Thesis,
    Humboldt State Univ., Arcata, CA, 90 p.

    Skinner, J. E. 1962. An historical reviewof the fish and
    wildlife resources of the San Francisco Bay area.

                                                         54

    55

                           Panopea abrupta
                           Adult

                                                            5cm

    Common Name: geoduck                                  are harvested year-round, but primarily during spring
    Scientific Name: Panopea abrupta                      and summer (Wolotira et al. 1989).  Meat quality
    Other Common  Names: Pacific geoduck, giant   appearstobecorrelatedwithsubstratetype;geoducks
    panopaea, geoduc, gweduc, king clam, gooey-duck    growing in coarse substrates produce a better quality
    (Gates and Frey 1974, Wolotira et al. 1989)           product (Goodwin and Pease 1987). The Washington
    Classification (Bernard 1983)                         commercial geoduck industry pays a royalty fee which
    Phylum: Mollusca                                      supports a geoduck hatchery that raises cultured
    Class: Bivalvia                                      juveniles to seed harvested beds. Geoducks must be
    Order: Myoida                                         processed within 24 hours after harvesting or they
    Family: Hiatellidae                                   gape, lose water and body fluids, die, and the meat
                                                          dries out (Schink et al. 1983).
    Value
    Commercial: The geoduck was not commercially    Recreational:This species is recreationally harvested
    harvested until 1970 (Wolotira et al. 1989), but it now    from British Columbia to California, but is particularly
    supports the largest clam fisheryonthe Pacific coast of   important in Washington (Schink et al. 1983). Because
    North America (Schink et al. 1983). It is commercially    the geoduck lives deep within the sediment, shovels
    harvested from Alaska to Washington, but primarily   and open-ended tubes are used to dig them. It is
    from southern British Columbia, Puget Sound, and    harvested year-round, usually during very low tides on
    Hood Canal, Washington.  In 1977, 3,900 t were    intertidal flats. However, a small numberare harvested
    harvested from Washington State's subtidal areas.    by sport divers (Goodwin and Shaul 1984).
    The industry is now limited to below the optimum
    sustained yield quota of about 2.25 t per year (Schink   Indicator of Environmental Stress: Geoduck beds may
    et al. 1983, Goodwin and Shaul 1984, Cheney and    be closed to harvesting because of coliform bacteria
    Mumford 1986). Geoduck neck meat is sold in Japan,    contamination. Beds may also be temporarily closed
    Taiwan, and withinthe U.S.; body meat is sold primarily   because of paralytic shellfish poisoning, however, this
    in California and on the U.S. Atlantic coast (Cheney    has not been a significant problem in Puget Sound.
    and Mumford 1986).  Geoduck harvests are worth    Many productive subtidal clam beds in Puget Sound
    about $2.4 million annually to U.S. fishermen (Wolotira   are closed to shellfish harvesting because of industrial
    et al. 1989). This species is harvested by divers during    and municipal pollution (Schink et al. 1983). Little is
    daylight using hand-held, high-pressure water jets.   known about this species' ability to concentrate heavy
    Most harvesting is in depths <18.3 m because diving   metals, pesticides, and other chemicals (Goodwin and
    time is limited in deeper water (Schink et al. 1983). In   Pease 1989).
    Washington, subtidal tracts are leased from the state.
    Tracts are required to be >182 m away from the mean    Ecological: This is the largest burrowing bivalve on the
    high-water line and have depths >5.5 m below mean    Pacific coast of North America. The geoduck is very
    lower lowwater (MLLW) (Schink et al. 1983). Geoducks    abundant in subtidal areas of Puget Sound and British

                                                       56

                                                                                         Geoduck continued
                                                          Washington (Table 1). It is not found in coastal estuaries
     Table 1. Relative abundance of geoduck in            of Washington and Oregon except for Netarts Bay,
              32 U.S. Pacific coast estuaries.            Oregon, where some are harvested. It is not found or
                        Life Stage                       is rare in California's estuaries, except for Morro Bay
           Estuary    A S J L E                           where it is common (Marriage 1954, Haderlie and
            Puget Sound  3    (3 ��  Relative abundance:  Abbott 1980, Maclntyre et al. 1986).
             Hood Canal  1 �3 (   3   * �   Highly abundant
              Skagit Bay O O   0  0 0     Abundant        Life Mode
            Grays Harbor              /  Rare             Eggs and larvae are pelagic. Juveniles and adults are
             WillapaBay             Blank Notpresent      benthic infauna, burrowing to depths of 100 cm
           Columbia River                                  (Goodwin et al. 1979, Haderlie and Abbott 1980).
            Nehalem Bay
           Tillamook Bay             Life stage:          Habitat
             Netarts Bay      .    /  A - Adults
                                      S - Spawning adults  Type: The   geoduck is found   intertidally to depths of at
             Siletz River            J - Juveniles        least 110 m in bays, sloughs, and estuaries (Goodwin
            Yaquina Bay L - Larvae
            Yaquina Bay              L - ELggs            1973a, Bernard 1983, Goodwin and Pease 1987,
             Alsea River                                  Wolotira et al. 1989). In Alaska, geoducks are found
            Siuslaw River                                 only subtidally at depths from 4.5-12.0 m (Wolotira et
           Umpqua River                                   al. 1989). This species is most abundant between 9.1
              Coos Bay                                    and 18.2 m below MLLW (Goodwin 1973a). The length
             Rogue River                                  and weight of geoducks decreases with depths between
           Klamath River                                  3 and 20 m (Goodwin and Pease 1987).
           Humboldt Bay    4  'i                 "o
               EelRiver                                   Substrate: The geoduck is found in a variety of
            Tomales Bay                                   substrates ranging from soft mud to pea gravel, but
       Cent San Fran. Bay*  * Includes Central San
       Conut San Fran. Bay            Francudeoa Suisun,  primarily in stable mud or sand bottoms (Goodwin
       South San Fran. Bay  Pand San ablo bays.           1973a, Goodwin and Pease 1987). It is often associated
           Elkhorn Slough                                 with the sea pen (Ptilosarcus gurnmeyl) and polychaete
              MorroBay 0 0 0   0                          tubes  (Cox  1979).    Polychaete  tubes  of
         SantaMonicaBay                                    Spiochaetopeterus costarum, Phyllochaetopeterus
            S&anPedro Bay  '  "/ '    / '                 prolifica, and Diopatraornata, are preferred attachment
            Alamitos Bay                                  areas for juveniles (Strathmann et al. 1987).
            Anaheim Bay
            Newport Bay                                   Physical/Chemical Characteristics: This species is
             Mission Bay
                                                          found in areaswhere watertemperatures range from 3-
           San DiegosBay                                  20�C (Bernard 1983). Eggs and larvae are found in
                                                          polyhaline-euhaline waters ranging from 22.0-35.0?0o;
                      A S J L E                           optimum is 27.5-32.5%o (Goodwin 1973b). Juveniles
                                                           and adults occur in mesohaline-euhaline waters (5.0-
    Columbia and it often dominatesthe biomassof benthic   35.0%o), but prefer salinities above 25.0?0/ (Andersen
    infaunacommunitiesthere(Cheneyand Mumford 1986,    1971,  Goodwin  1976).   Optimum  spawning
    Goodwin and Pease 1989). A conservative population    temperatures are 12-14�C, but spawning occurs in
    estimate of 117.6 million geoducks was made for   temperatures from 8-160�C (Goodwin 1976). The best
    33,799 acres of subtidal beds surveyed in Puget Sound    temperature for larval survival is between 6 and 16�C
    in 1977 (Cheney and Mumford 1986).                     (Goodwin 1973b).  Although juveniles and adults
                                                          withstand air temperatures of 0-250C, they are only
    Range                                                 found in areas where water temperatures during the
    Overall: This is a temperate amphi-North Pacific   spawning period (April to July) are not above 16�C
    species, foundfrom Kyushuto Hokkaido Islands,Japan,    (Andersen 1971, Goodwin 1973b, 1976).
    and in the northeast Pacific from southeast Alaska to
    Baja California (Scammons Lagoon), and also in the    Miarations and Movements: Planktonic eggs and larvae
    northern Gulf of California (Fitch 1953, Haderlie and    are dispersed by water currents.  Bottom-dwelling
    Abbott 1980, Bernard 1983, Wolotira et al. 1989).      post-larvae are active crawlers (Goodwin et al. 1979).
                                                           Newly-settled juveniles remain at or nearthe sediment
    Within StudvArea:Thegeoduckis commonto abundant    surface until they grow to 15 mm shell length (SL), then
    in Skagit Bay, Puget Sound, and Hood Canal,    their siphons begin to lengthen. Once siphons are

                                                       57

    Geoduck continued
    elongated and well-developed, juveniles begintoburrow    Aae and Size of Adults: In Puget Sound, most males
    deeply (Strathmann et al. 1987). Juvenile and adults   mature in three years at 60-100 mm SL; females
    are sedentary infauna, remaining in the area where    mature in four years at 100-120 mm SL (Andersen
    they initially burrowed.                             1971). In British Columbia, maturity may be reached in
                                                          5-7 years (Sloan and Robinson 1984, Wolotira et al.
    Reproduction                                         1989). During the first four years they grow rapidly, but
    Mode: The geoduck is gonochoristic, oviparous, and a   older, large clams (>100 mm SL) grow little if at all
    broadcast spawner; eggs are fertilized externally. It is   (Andersen 1971, Goodwin 1973a, 1976, Shaul and
    iteroparous and a batch spawner with one spawning    Goodwin 1982, Breen and Shields 1983). In general,
    period per year (Andersen 1971, Goodwin 1973a,   this is a very long-lived and slow-growing species, but
    Goodwin et al. 1979).                                growth can be highly variable. Depending on geographic
                                                          area, geoducks may reach 75 mm SL in 2-8 years
    Matina/Snawnina: In Hood Canal and Puget Sound,    (Goodwin and Shaul 1984). In most areas in Puget
    spawning occurs from April to July (primarily from May    Sound and British Columbia, it reaches 0.9 kg (market
    to June) (Goodwin 1973a, 1976, Strathmann et al.   size) in 8 to 10 years (Cheney and Mumford 1986,
    1987).  In British Columbia, the geoduck spawns    Wolotira et al. 1989). The oldest individuals are about
    primarily from Juneto July (Sloan and Robinson 1984).    146 years old. Maximum size and weight is 230 mm SL
    It is stimulated to spawn by increasing water   and 9.1 kg, but most weigh <4.5 kg (Oceanographic
    temperatures, the presence of geoduck sperm in the   Instituteof Washington 1981, Kozloff 1983, Wolotira et
    water, and (at least in hatchery situations) by increased   al. 1989).
    algae concentrations (Goodwin 1973b, Wolotira et al.
    1989). When it spawns, both eggs and sperm are   Food and Feeding
    expelled from the exhalant siphon continuously for   Trophic Mode: This species is a suspension/filter-
    several minutes or upto an hour (Goodwin et al. 1979).   feeding planktivore. Larvae, juveniles, and adults feed
                                                          by filtering food particles from seawater with their gills.
    Fecundity: A female can release 7.5-20.0 million eggs    Post-larval geoducks may also feed on substrate
    during a single spawning; hatchery stock have been    deposits (Goodwin and Pease 1989).
    induced to spawn again if returned to cooler water.
    (Goodwin et al. 1979). Although reproductive output is   Food Items: Larvae have been successfully reared on
    high, recruitment (i.e., settlement of larvae and survival   the following algae species: Pavlova lutheri, Isochrysis
    of young) is usually erratic or low (Goodwin et al. 1979).   galbana, Pseudoisochrysisparadoxa, Phaeodactylum
                                                          tricornutum, Monochrysis lutheri, Chaetoceros
    Growth and Development                               calcitrans, and Thalassiosira pseudonona (Goodwin
    Eaa Size and Embrvonic Develonment: Eggs are   1973a, Goodwin et al. 1979, Strathmann et al. 1987).
    spherical and 0.082 mm  (Goodwin et al. 1979).    Larvae, juveniles, and adults feed on various
    Embryonic development is indirect and external.      phytoplankton and suspended detritus.

    Aae and Size of Larvae: Larval size ranges are 0.11-   Biological Interactions
    0.40 mm (pelagic larvae) and 0.40-0.80 mm (epibenthic   Predation: Important predators of small juveniles include
    post-larvae) (Goodwin et al. 1979). At 140C, larval   northern moon snail (Polinices lewisil), coonstriped
    growth is as follows: at 48 hr, straight-hinge larvae   shrimp (Pandalus danae), rock crabs (Cancer spp.),
    develops; at 6 days, veligers are 0.120 x 0.105 mm; at   English sole (Parophrys vetulus), rock sole
    10 days, veligers are 0.150 x 0.125 mm. Settlement   (Lepidopsetta bilineata), sand sole (Psettichthys
    occurs at 30 days at 17.60�C, and 47 days at 14-15�C    melanostictus), pile perch (Rhacochilus vacca), spiny
    (Goodwin 1973a, 1973b). The largest veligers (before   dogfish (Squalusacanthias), starry flounder (Platichthys
    metamorphosis to benthic juveniles) are 0.350-   stellatus), and other flatfish. Seastars (Pisaster spp.)
    0.400 mmindiameter(Goodwinetal. 1979). Settlement   and sunstar (Pycnopodia helianthoides) feed on
    is usually from April to August, peaking in mid-July   juveniles and adults (Sloan and Robinson 1983, Wolotira
    (Andersen 1971).                                     et al. 1989). Rock crabs will feed on any dislodged
                                                          individuals (Wolotira et al. 1989). The tips of geoduck
    Juvenile Size Ranoe: Juveniles range in size from 0.8-   siphons are eaten by the Pacific staghorn sculpin
    100.0 mm SL (Andersen 1971). When 1.5-2.0 mmSL,    (Leptocottus armatus) (Andersen 1971). Adults are
    they start to burrow into the substrate (Goodwin and    also excavated and eaten by sea otters (Enhydra
    Pease 1989). Juveniles <5 mm SL still have the ability   lutris). Geoducks reduce predation rates by burrowing
    to move, while largerjuveniles simply burythemselves    deeply into sediments as they grow.  Siphons are
    as they grow (Goodwin and Shaul 1984).               protected by retracting them when inactive and allowing

                                                       58

                                                                                          Geoduck continued
    the siphon hole to be buried (Wolotira et al. 1989).    Columbia. British Columbia Ministry Env., Mar. Res.
    Predation is probably highest in areas where a hard    Branch, Fish. Man. Rep. No. 15, 25 p.
    layer of rock or clay does not permit geoducks to
    burrow deeply.                                         Fitch, J. E. 1953. Common marine bivalves of California.
                                                           Calif. Fish Game, Fish Bull. 90, 102 p.
    Factors Influencina Pooulations: Larvae and small
    juveniles appear to suffer extremely high mortality   Gates, D. E., and H. W. Frey.  1974.  Designated
    which results in low recruitment (Goodwin et al. 1979).    common names of certain marine organisms of
    However, mortality rates for older juveniles (2+ years)   California. Calif. Fish Game, Fish Bull. 161:55-90.
    and adults are very low (Andersen 1971, Goodwin et al.
    1979). Recruitment of juveniles appears to be highest    Goodwin, C. L. 1973a. Subtidal geoducks of Puget
    in areas containing adults, indicating that commercial    Sound, Washington.  Tech. Rep. 14, Wash. Dept.
    harvest may adversely affect recruitment (Goodwin    Fish., Olympia, WA, 81 p.
    and Shaul 1984). To assist reestablishment of geoducks
    in areas where they have recently been harvested, the    Goodwin, C. L.  1973b.  Effects of salinity and
    Washington Department of Fisheries has developed a   temperature on the  embryos of the geoduck clam
    geoduck hatchery and "seeds" these areas (Goodwin    (Panopea generosa  Gould).  Proc. Natl. Shellfish.
    and Shaul 1984). Some adult mortalities result from    Assoc. 63:93-95.
    anoxicconditions arising from vegetation accumulation
    and decomposition, dredging operations, sediment    Goodwin, C. L. 1976. Observations on spawning and
    slumping and earthquakes (which may crack their   growth of subtidal geoducks (Panopea generosa
    shells) (Andersen 1971, Wolotira et al. 1989). Other    Gould). Proc. Natl. Shellfish. Assoc. 65:49-58.
    factors possibly affecting populations include disease,
    siltation (especially intertidal and shallow water subtidal   Goodwin, C. L., and B. Pease. 1989. Species profiles:
    beds), and illegal harvest (Andersen 1971, Schink et al.   life histories and environmental requirements of coastal
    1983). Somegeoduck beds in Puget Sound areclosed    fishes and invertebrates (Pacific Northwest)-Pacific
    to harvest because of industrial and municipal pollution.    geoduck clam.  U.S. Fish. Wildl. Serv. Biol. Rep.
    Other beds have been lost because of pier, jetty,   82(11.120), U.S. Army Corps Eng., TR EL-82-4, 14 p.
    marina, and pipeline development projects. Aquaculture
    of other species (primarily salmonid net pens) has    Goodwin, L., and B. Pease. 1987. The distribution of
    altered and reduced geoduck harvest in some areas    geoduck (Panopea abrupta) size, density, and quality
    (Goodwin and Pease 1989).                             in relation to habitat characteristics such as geographic
                                                           area, water depth, sediment type, and associated flora
    References                                            and fauna in Puget Sound, Washington. Tech. Rep.
                                                           102, Wash. Dept. Fish., Olympia, WA, 44 p.
    Andersen, A. M., Jr. 1971. Spawning, growth, and
    spatial distribution of the geoduck clam, Panopea    Goodwin, L.,and W. Shaul. 1984. Age recruitment and
    generosa Gould, in Hood Canal, Washington. Ph.D.    growth of the geoduck clam (Panopeagenerosa, Gould)
    Thesis, Univ. Wash., Seattle, WA, 118 p.               in Puget Sound Washington. Prog. Rep. 215, Wash.
                                                           Dept. Fish., Olympia, WA, 30 p.
    Bernard, F. R. 1983. Catalogue of the living Bivalvia
    of the eastern Pacific Ocean: Bering Strait to Cape    Goodwin, L., W. Shaul, and C. Budd. 1979. Larval
    Horn. Can. Spec. Publ. Fish. Aquat. Sci. 61, 102 p.    developmentofthegeoduckclam (Panopeagenerosa,
                                                           Gould). Proc. Natl. Shellfish. Assoc. 69:73-76.
    Breen, P. A., and T. L. Shields. 1983. Age and size
    structureinfivepopulationsofgeoducclams(Panopea    Haderlie, E. C., and D. P. Abbott. 1980. Bivalvia: The
    generosa) in British Columbia. Can. Tech. Rep. Fish.    clams and allies. In R. H. Morris, D. P. Abbott, and E.
    Aquat. Sci. No. 1169, 62 p.                            C. Haderlie (editors), Intertidal invertebrates of
                                                           California, p. 355-411. Stanford Univ. Press, Stanford,
    Cheney, D. P., and T. F. Mumford, Jr. 1986. Shellfish    CA.
    and seaweed harvests of Puget Sound. Wash. Sea
    Grant, Univ. Wash. Press, Seattle, WA, 164 p.          Kozloff, E. N.  1983. Seashore life of the northern
                                                           Pacific coast. Univ. Wash. Press, Seattle, WA, 370 p.
    Cox, R. K. 1979. The geoduck, Panopea generosa:
    some general information on distribution, life history,   Maclntyre, J., S. R. Sparling, M. Faustini, T. L. Richards,
    harvesting, marketing and management in British   R. Nakamura, and B. F. Putman.  1986.  Resource

                                                       59

    Geoduck continued
    inventory: Marine life: Cayucos State Beach, Morrow
    Strand State Beach, Atascadero State Beach, Morro
    Bay State Park, Montana De Oro State Park. Calif.
    Polytech. State Univ., San Luis Obispo, CA, various
    pagination.

    Marriage, L. D. 1954. The bay clams of Oregon: their
    economic importance, relative abundance, and general
    distribution. Cont. No. 20, Fish Comm. Oregon,
    Portland, OR, 47 p.

    Oceanographic Institute of Washington. 1981. Clam
    and mussel harvesting industries in Washington state.
    Oceanog. Comm. Wash., Seattle, WA, various
    pagination.

    Schink, T. D., K. A. McGraw, and K. K. Chew. 1983.
    Pacific coast clam fisheries. Wash. Sea Grant Prog.,
    Univ. Wash., Seattle, WA, 72 p.

    Shaul, W., and L. Goodwin. 1982. Geoduck (Panopea
    generosa: Bivalvia) age as determined by internal
    growth lines in the shell. Can. J. Fish. Aquat. Sci.
    39:632-636.

    Sloan, N.A., and S. M. C. Robinson. 1983. Winter
    feeding by asteroids on a subtidal sandbed in British
    Columbia. Ophelia 22(2):125-140.

    Sloan, N. A., and S. M. C. Robinson. 1984. Age and
    gonad development in the geoduck clam, Panopea
    abrupta (Conrad) from southern British Columbia,
    Canada. J. Shellfish Res. 4(2):131-137.

    Strathmann, M. F., A. R. Kabat, and D. O'Foighil. 1987.
    Phylum Mollusca, class Bivalvia. In M. F. Strathmann
    (editor), Reproduction and development of marine
    invertebrates of the northern Pacific coast, p. 309-353.
    Univ. Wash. Press, Seattle, WA.

    Wolotira, R. J., Jr., M. J. Allen, T. M. Sample, C. R. Iten,
    S. F. Noel, and R. L. Henry. 1989. Life history and
    harvest summaries for selected invertebrate species
    occurring off the west coast of North America. Volume
    1: shelled molluscs. NOAA Tech. Memo. NMFS F/
    NWC-160, 177 p.

                                                       60

    61

       Crangon franciscorum
       Adult

                                              2cm

    Common Name: bay shrimp                                changes in estuarine temperature and salinity regimes
    Scientific Name: Crangon franciscorum                  (Khorram and Knight 1977).  River discharge and
    Other Common  Names: Franciscan bay shrimp,    subsequent changes to estuarine salinity regimes
    California shrimp, grass shrimp (Gates and Frey 1974,    appear to determine distribution, recruitment levels,
    Khorram and Knight 1977)                               survival, and growth (Hatfield 1985, California
    Classification (Bowman and Abele 1982)                 Department of Fish and Game 1987). Alicyclic hexanes
    Phylum: Crustacea                                      at concentrations ranging of 1.5-10.9 ppm are acutely
    Class: Malacostraca                                    toxic to bay shrimp; these chemicals can be
    Order: Decapoda                                        bioaccumulated by a factor of 13 (Benville et al. 1985).
    Family: Crangonidae
                                                           Ecological: The bay shrimp is the dominant decapod
    Two subspecies are defined, C. franciscorum    shrimp in most Pacific coast estuaries (Krygier and
    franciscorum and C. franciscorumangustimana. The    Horton 1975, Hoeman 1982, Rudy and Rudy 1983,
    latter differs from C. f. franciscorum by having a long   Hatfield 1985). It is an important prey for many Pacific
    chela, with tip of dactylus crossing under basal part of   coast fish and crab species (Haertel and Osterberg
    fixed finger (Butler 1980).                            1966, Stevens et al. 1982), and an important estuarine
                                                           benthic and epibenthic predator (Sitts and Knight 1979,
    Value                                                  Siegfried 1980, Hatfield 1985). The agitation of bottom
    Commercial: The bay shrimp is commercially fished    sediments (caused by this species as it searches for
    (primarily with trawls) only in San Francisco Bay,    food) may contribute to nutrient cycling (Krygier and
    California (Smith and Kato 1979, Chace and Abbott    Horton 1975). Estuaries are used as nursery areas by
    1980). It once supported a larger fishery that utilized   this species, with lower salinity areas particularly
    trawls, fyke nets, and seines (Butler 1980). It is fished    important to young shrimp (Krygier and Horton 1975).
    mainly for use as bait, but some is used for human
    consumption.  Recently, annual landings for three    Range
    Crangon species (C. franciscorum, C. nigricauda, and    Overall: The bay shrimp's overall range is from San
    C. nigromaculata) captured in San Francisco Bay have    Diego, California, to Alaska (Butler 1980, Chace and
    ranged from 2.3 to 25.0 t (Chace and Abbott 1980).     Abbott 1980). C. f. angustimana is apparently only
                                                           found in deeper waters (18-183 m) from Tillamook
    Recreational: This species is used as bait for striped    Rock, Oregon to Kachemak Bay, Alaska (Butler 1980).
    bass (Morone saxatilis) and sturgeon (Acipenserspp.).
                                                           Within Study Area: This species is abundant to common
    Indicator of Environmental Stress: Because estuaries   in all Pacific coast estuaries from San Francisco Bayto
    play a critical role in the bay shrimp's life history,    Puget Sound, Washington, but it is not normally found
    alterationsof these habitatsdirectlyaffectitspopulations    in estuaries south of San Francisco Bay (Table 1)
    (Frey 1971).  This species is a good indicator of    (Monaco et al. 1990).

                                                        62

                                                                                           Bay shrimp continued

                                                            muddy substrates (Kuris and Carlton 1977).
      Table 1. Relative abundance of bay shrimp
               in 32 U.S. Pacific coast estuaries.          Phvsical/Chemical Characteristics: The bay shrimp is
                         Life Stage                         a euryhaline species. Juveniles and adults are found
            Estuary    A S J L E                            in euhaline to oligohaline waters in Prince William
             Puget Sound c  i X3 c c   Relative abundance:  Sound, Alaska (2.2-28.3%o) (Butler 1980). In San
              Hood Canal O O O O O    0   Highly abundant    Francisco Bay and Delta, highest densities are found
              Skagit Bay   � �  � 6    �   Abundant
              ySkagitBay /i (. *           A0 n Common      atsalinitiesof 1-7%0(Siegfried1980). Juvenilesappear
            Grays Harbor ci ci S c" 8c    'i  Rare          to prefer lower salinities (<32.0%0), while ovigerous
              Willapa Bay     ( *   (   Blank Not present    females prefer salinities >14.6%/o (Krygier and Horton
           Columbia River it it-3 S 3                       1975).  Juveniles and nonovigerous adults tolerate
            Nehalem Bay i i * i  temperatures of 5.2-21.3�C; ovigerous females prefer
            Tillamook Bay �  � � �    Life stage:
             Netamkrtsay C 0  *    (  UfA stagedults        temperaturesof6.8-19.2�C(Krygierand Horton 1975).
                                      e   -Spawningadults  Salinity and temperature influence this species'
              SYaquina Bay     L0      L J-Juveniles        distribution significantly.  High salinities retard the
                                       E - Eggs             movements of juveniles to lower estuarine areas, while
              AlseaRiver    � � � � �
              Siuslaw River  3 O * O O                      hightemperatures in the summerincrease movements
            Siuslaw River �i �i � �i �i
            UmpquaRiver ci c3 *   O l                       to upper estuarine areas (Krygier and Horton 1975).
               coos Bay                                     Low salinities probably retard egg development (Krygier
             Rogue River                                    and Horton 1975), and salinities <12%o may reduce
            Klamath River                                   larval survival (Siegfried 1980). Optimum conditions
            Humboldt Bay  C D  O                            for adults are salinities of 1 8-20%0 and temperatures of
               Eel River cIS (O i ci                        4.5-17.0�C (Khorram and Knight 1977, Siegfried 1980).
             Tomales Bay O 0    C   0
       Cent. SanFran. Bay'  3 c � � .    * IncludesCentral San  Miarations and Movements: A "spawning migration"
       South San Fran. Bay      O         O   Francisco. Suisun,  occurs during the reproductive periods; adult females
           Elkhorn Slough                                   and males move to lower, more saline areas of estuaries
              Morro Bay                                     (primarily March to July) (Krygier and Horton 1975).
         Santa Monica Bay                                   Juveniles move up estuaries during the summerto rear
           San Pedro Bay                                    in lower salinity, highertemperature areas (Israel 1936,
             Alamitos Bay                                   Armstrong et al. 1981, Hatfield 1985). As they grow
            Anaheim Bay                                     and mature, bay shrimp move to lower, more saline
             Newport Bay                                    areas (Krygierand Horton 1975). In the fall and winter,
             Mission Bay                                    many adults move to near the mouth of estuaries and
           San Diego Bay                                    nearshore areas outside estuaries (Hatfield 1985).
           TijuanaEstuary                                   Juveniles and adults undergo nocturnal vertical
                       A S J L E                            migrations to feed (Sitts and Knight 1979). Larvae
                                                            appear to be advected seaward by river flow (Hatfield
                                                            1985).
    Life Mode
    Eggs are brooded on the female's body, carried under    Reproduction
    the abdomen, attached to and between the basal joints    Mode: The bay shrimp is gonochoristic and oviparous.
    and inner rami of the pleopods or abdominal legs    Sperm is stored internally in the female; eggs are
    (Israel 1936). The larvae are epipelagic, and juveniles   fertilized when extruded and brooded externally on the
    and adults are epibenthic.                               female's body.

    Habitat                                                  Matina/SDawnina: Although gravid females may be
    Type: Adults are found in estuaries and offshore,    found year-round, usually only two spawning periods
    intertidally down to 183 m (Butler 1980). Ovigerous    exist (sometimes only one depending on the estuary)
    femalesarefoundinthelowerportionsofestuariesand    (Israel 1936, Krygier and Horton 1975). In Yaquina
    adjacent offshore waters (Krygier and Horton 1975).    Bay, Oregon, spawning occurs from December to
    Juveniles primarily inhabit channels and flats in the low    March (older females), and from April to August (first-
    salinity areas of estuaries.                             time and repeat spawners). The second spawning is
                                                            usually larger (more spawners present for a longer
    Substrate: Larvae are found overa varietyof substrates.    period) than the first (Krygier and Horton 1975). In San
    Juveniles and adults occur primarily over sandy to    Francisco Bay,only a single extended spawning period

                                                        63

    Bay shrimp continued
    was thought to exist, with a peak from March to   reported is 110 mm TL off the Columbia River (Durkin
    September (Israel 1936).  However, a bimodal    and Lipovsky 1977). Females may live 2-2.5 years,
    reproductive schedule appears to occur here also;   and males about 1.5 years (Stevenson et al. 1987).
    during the first period, gravid females reside primarily
    off the mouth of San Francisco Bay (Hatfield 1985). A    Food and Feeding
    "spawning migration" occurs, with females and males    TrolhicMode: Larvae, juveniles, and adults are primarily
    moving to deeper, higher salinity areas (usually >21 'oo,   carnivorous (occasionally detritivorous), feeding on
    depending on water temperature) when they become    benthic and epibenthic prey. Food habits depend on
    reproductively active (Krygier and Horton 1975,   the shrimp's size, temperature-salinity preferences,
    Siegfried 1980). Nearshore areas outside of estuaries    and prey availability (Wahle 1985).
    are often used by spawning adults during the winter
    and spring (Durkin and Lipovsky 1977, Hatfield 1985).    Food Itms:The bay shrimpfeedson mysids (Neomysis
                                                          mercedis), amphipods (primarily Corophium spp.,
    Fecundity: Females from 47.8-67.4 mm total length   Ampelisca abdita, and Grandidierellajaponica), bivalves
    (TL) carried 1,923-4,764 eggs perfemale, with a mean    (primarily Mya arenaria, Gemma gemma, and
    of 3,528 (Krygier and Horton 1975). Fecundity of bay    Venerupis japonica), foraminiferans, isopods,
    shrimp ranged from 1,977-3,103 in Grays Harbor,   copepods, ostracods, gastropods, and plant material
    Washington (Hoeman 1982), and from 2,499-8,840 in   (Wahle 1985).
    south San Francisco Bay (Stevenson et al. 1987).
    Fecundity  (Y)  was  calculated  to  be  Y=-    Biological Interactions
    5338.7+156.1 (TL) for shrimp in Yaquina Bay (Krygier   Predation: The bay shrimp is an important prey for the
    and Horton 1975), and log Y=-3.66+4.091og(TL) for   striped bass (Morone saxatilis), brown smoothhound
    shrimp in San Francisco Bay (Siegfried 1980).         (Mustelus henlei), green sturgeon (A. medirostris),
                                                          white sturgeon (A. transmontanus), Pacific staghorn
    Growth and Development                                sculpin (Leptocottus armatus), Pacific tomcod
    Eaa Size and Embrvonic DeveloDment: Eggs are    (Microgadusproximus), prickly sculpin (Cottus asper),
    spherical and 0.60 mm in diameter (Mondo 1980).    sand sole (Psettichthys melanostictus), waterfowl,
    Embryonic development is indirect and external; eggs    harbor seal (Phoca vitulina), and the Dungeness crab
    remain in the female's brood pouch until hatching.    (Cancer magister) (Ganssle 1966, Hoeman 1982,
    Eggs appear to take 8-1 2 weeks to mature, depending    Stevens et al. 1982). The bay shrimp is also susceptible
    ontemperature. Larvae hatched in early spring develop   to cannibalism (Mondo 1980).
    into juveniles by Mayto July (Krygier and Horton 1975).
                                                          Factors Influencino PoDulations: This species may
    Aae and Size of Larvae: Larvae range from 6.0-7.4 mm    compete with the introduced oriental shrimp (Palaemon
    TL (Israel 1936, Krygier and Horton 1975). Larvae    macrodactylus) for food and resources, especially
    undergo seven larval stages in 21 days at 17.5�C    during drought years (Sitts and Knight 1979, Siegfried
    (Mondo 1980).                                         1980). The bay shrimp is one of the most abundant
                                                          organisms entrained during dredging operations in
    Juvenile Size Ranae: Juvenile bay shrimp range from    Pacific Northwest estuaries (Armstrong et al. 1981,
    6.0-7.4 mm to about 34 mm TL for males, 48 mm TL for   Hoeman 1982). Its distribution is also influenced bythe
    females (Israel 1936, Krygier and Horton 1975),   availability and abundance of the mysid Neomysis
    however, this may differ between estuaries (Israel   mercedis (Siegfried 1980).  Freshwater inflow into
    1936). After reaching 30 mm TL, growth is estimated    estuaries strongly influences this species' distribution
    to be 2.0 mm/month (Stevenson et al. 1987).           and abundance (Hatfield 1985, California Department
                                                          of Fish and Game 1987). Abiotic conditions during
    Aae and Size of Adults: Both sexes mature in about 1 -   winter and spring off the mouths of estuaries may also
    1.5 years, with most males reaching maturity at 34 mm    influence populations (Hatfield 1985). The bay shrimp
    TLandfemales at 48 mmTL(KrygierandHorton1975,   is a short-lived species that shows large annual
    Butler 1980, Stevenson et al. 1987) or 55-60 mm TL in   fluctuations in abundance and may be highly sensitive
    San Francisco Bay (Hatfield 1985, Stevenson et al.   to effects of short-term estuarine pollution (Frey 1971).
    1987). Males appearto spawn onlyonce, whilefemales    Parasitism by the branchial isopod Argeiapugettensis
    may produce two broods (Butler 1980). Females are   inhibits female reproduction (Butler 1980, Hoeman
    60 mmTLin 1.5years, males50-52mmTLafterl year;   1982). Necrotic shell lesions may affect populations,
    females >62 mm TL are rare in Yaquina Bay, but are   but little information is available (Stevenson et al.
    common off the Columbia River (Krygier and Horton    1987). Predation may also significantly control year
    1975, Durkin and Lipovsky 1977). The largest size   class strength (Stevenson et al. 1987).

                                                       64

                                                                                         Bay shrimp continued
    References                                              common  names of certain marine organisms of
                                                           California. Calif. Fish Game, Fish Bull. 161:55-90.
    Armstrong, D. A., B. G. Stevens, and J. C. Hoeman.
    1981. Distribution and abundance of Dungeness crab    Haertel, L., and C. Osterberg.  1966.  Ecology of
    and Crangon shrimp, and dredging-related mortality of   zooplankton, benthos and fishes in the Columbia River
    invertebrates and fish in Gray's Harbor, Washington.    estuary. Ecology 48(3):459-472.
    Tech. Rep. to Wash. Dept. Fish. and U.S. Army Corps
    Eng., School Fish., Univ. Wash., Seattle, WA, 349 p.    Hatfield, S. E. 1985. Seasonal and interannualvariation
                                                           in distribution and population abundance of the shrimp
    Benville, P. E., J. A. Whipple, and M. B. Eldridge. 1985.    Crangon franciscorumin San Francisco Bay. Hydrobiol.
    Acute toxicity of seven alicyclic hexanes to striped    129:199-210.
    bass, Morone saxatilis, and bay shrimp, Crangon
    franciscorum, in seawater. Calif. Fish Game 71 (3):132-    Hoeman, J. C. 1982. The distribution and ecology of
    140.                                                    three species of crangonid shrimp in Grays Harbor,
                                                           Washington, and their susceptibility to entrainment by
    Bowman, T. E., and L. G. Abele. 1982. Classification    dredges. M.S. Thesis, Univ. Wash., Seattle, WA,
    of the recent crustacea. In L. G. Abele (editor), D. E.    135 p.
    Bliss (editor-in-chief), The biology of Crustacea, Volume
    1. Systematics, the fossil record, and biogeography, p.   Israel, H. R.  1936.  A contribution toward the life
    1-25. Academic Press, New York, NY.                     histories oftwo California shrimps, Crago franciscorum
                                                           (Stimpson) and Crago nigricauda (Stimpson). Calif.
    Butler, T. H. 1980. Shrimps of the Pacific coast of    Fish Game, Fish Bull. 46, 28 p.
    Canada. Can. Bull. Fish Aquat. Sci., Bull. No. 202,
    280 p.                                                  Khorram, S., and A. W. Knight.  1977.  Combined
                                                           temperature-salinity effects on grass shrimp. J. Environ.
    California Department of Fish and Game. 1987. Delta    Engin. Div., Am. Soc. Civil Engin. 103:381-388.
    outflow effects on the abundance and distribution of
    San Francisco Bay fish and invertebrates, 1980-1985.    Krygier, E. E., and H. Horton.  1975.  Distribution,
    Exhibit 60, entered by the California Department of   reproduction, and growth of Crangon nigricauda and
    Fish and Game forthe State Water Resources Control    Crangon franciscorum  in Yaquina Bay, Oregon.
    Board 1987 Water Quality/Water Rights Proceeding    Northw. Sci. 49(4):216-240.
    on the San Francisco Bay/Sacramento-San Joaquin
    Delta. Calif. Dept. Fish Game, Stockton, CA, 345 p.    Kuris, A. M., and J. T. Carlton. 1977. Description of a
                                                           new species, Crangon handi, and new genus,
    Chace, F. A., Jr., and D. P. Abbott. 1980. Caridea: the    Lissocrangon, of crangonid shrimps (Crustacea:
    shrimps.  In R. H. Morris, D. P. Abbott, and E. C.    Caridea) from the California coast, with notes on
    Haderlie (editors), Intertidal invertebrates of California,    adaptation in body shape and coloration. Biol. Bull.
    p. 567-593. Stanford Univ. Press, Stanford, CA.         153:540-559.

    Durkin, J. T., and S. J. Lipovsky.  1977.  Aquatic    Mondo, G.S. 1980. The larvaldevelopmentof thebay
    disposal field investigations Columbia River disposal    shrimp Crangon franciscorum.  M.A. Thesis, San
    site, Oregon. Appendix E: Demersal fish and decapod    Francisco State Univ., San Francisco, CA, 120 p.
    shellfish studies. Tech. Rep. D-77-30, U.S. Army
    Corps Eng., Waterways Exper. Sta., Vicksburg, Ml,    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    184 p.                                                  Nelson. 1990. Distribution and abundance of fishes
                                                           and invertebrates in west coast estuaries, Volume I:
    Frey, H. W. 1971. California's living marine resources    data summaries.   ELMR  Rep. No. 4. Strategic
    and their utilization. Calif. Dept. Fish Game,    Assessment Branch, NOS/NOAA, Rockville, MD,
    Sacramento, CA, 148 p.                                  240 p.

    Ganssle, D. 1966. Fishes and decapods of San Pablo    Rudy, P., Jr., and L. H. Rudy. 1983. Oregon estuarine
    and Suisun Bays. In D. W. Kelley (compiler), Ecological    invertebrates. An illustrated guide to the common and
    studiesofthe Sacramento-SanJoaquin estuary. Calif.   important invertebrate animals. U.S. Fish Wildl. Serv.
    Fish Game, Fish Bull. 133:64-94.                        Biol. Serv. Prog., FWS/OBS-83/16, Portland, OR,
                                                           225 p.
    Gates, D. E., and H. W. Frey. 1974. Designated

                                                        65

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    Siegfried, C. A. 1980. Seasonal abundance and
    distribution of Crangon franciscorum and Palaemon
    macrodactylus (Decapod, Caridea) inthe San Francisco
    Bay-Delta. Biol. Bull. 159:177-192.

    Sitts, R. M., and A. W. Knight. 1979. Predation by the
    estuarine shrimps Crangon franciscorum Stimpson
    and Palaemon macrodactylus Rathbun. Biol. Bull.
    156:356-368.

    Smith, S. E., and S. Kato. 1979. The fisheries of San
    Francisco Bay: past, present and future. In T. J.
    Conomos (editor), San Francisco Bay: the urbanized
    estuary, p. 445-468. Pac. Div. Am. Assoc. Adv. Sci.,
    and Calif. Acad. Sci., San Francisco, CA.

    Stevens, B. G., D. A. Armstrong, and R. Cusimano.
    1982. Feeding habits of the Dungeness crab Cancer
    magister as determined by the index of relative
    importance. Mar. Biol. (Berl.) 72(1):135-145.

    Stevenson, M. L., T. C. Goddard, L. M. Kiguchi, and P.
    J. Kinney. 1987. South Bay Discharges Authority
    water quality monitoring program. Final Report to
    South Bay Discharges Authority, San Jose, CA.
    Kennetic Lab. Inc., Santa Cruz, CA, and Larry Walker
    Assoc., Inc., Davis, CA, 467 p.

    Wahle, R. A. 1985. The feeding ecology of Crangon
    franciscorum and Crangon nigricauda in San Francisco
    Bay, California. J. Crust. Biol. 5(2):311-326.

                                                       66

    67

       Cancer magister
       Adult

                                                               5cm

    Common Name: Dungeness crab                           Dungeness crab is sold as cooked whole or shelled
    Scientific Name: Cancer magister                      (and frozen or vacuum-packed ) in cans.
    Other Common Names: Pacific edible crab, edible
    crab, market crab, commercial crab (Hart 1982, Pauley    Recreational: Limited data are available onthe numbers
    et al. 1986)                                          of Dungeness crab captured by sport fishermen. It is
    Classification (Bowman and Abele 1982)                primarily caught in bays and estuaries, captured either
    Phylum: Crustacea                                     intertidally by hand or subtidally by baited crab pots,
    Class: Malacostraca                                   ring nets, dip nets, and hook and line (Pauley et al.
    Order: Decapoda                                       1986). Legal crabs for recreational fishermen must be
    Family: Cancridae                                     male and >146 mm CW in Oregon, >152 mm CW in
                                                          Washington, and >165 mm CW in California (where
    Value                                                 males and females can be taken) (Dahlstrom and Wild
    Commercial: The Dungeness crab is an important    1983).
    commercial shellfish that is harvested from the waters
    of Alaska to California. In 1985, more than 12,700 t   IndicatorofEnvironmentalStress:Theeffectsofurban
    worth over $39 million were landed (National Marine    pollution including chlorine residuals, heavy metals,
    Fisheries Service 1986). The abundance of this species    chlorinated pesticides, polychlorinated biphenyls, and
    fluctuates considerably, but long-term average annual    hydrocarbons, on Dungeness crab are not clear.
    landings are near 17,000 t (Pacific Marine Fisheries    However, sublethal effects are indicated for some
    Commission 1987). Baited crab pots are used to catch    pollutants at concentrations presently occurring in San
    this species in nearshore marinewaters normally<120    Francisco Bay, California (Guard et al. 1983, Haugen
    m deep (Dahlstrom and Wild 1983, Barry 1985). In the    1983a, 1983b, Horne et al. 1983, Cheney and Mumford
    study area, major commercial landings occur north    1986). Exposuretooiledsedimentslowersthisspecies'
    from Fort Bragg, California (Garth and Abbott 1980).    reproductive activity and larval survival (Karinen et al.
    The commercial season occurs primarily when males    1985). Crabs are intolerant of low dissolved oxygen
    are hard-shelled. Off northern California, Oregon, and    (optimal is>5 ppm),and lowconcentrations of ammonia
    Washington the season usually opens December 1   aretoxic (Cheney and Mumford 1986). Theinsecticide
    and only male crabs 2159 mm carapace width (CW)    SEVIN (carbaryl) is sometimes used to control ghost
    are legal (Barry 1985, Demory 1985, Warner 1985). In   shrimp (Callianassa spp.) in Pacific oyster( Crassostrea
    Alaska, the commercial season in the Southeast opens    gigas) beds, but is also very toxic to Dungeness crabs
    July 1, Yakutat opens May 1,and Kodiak opens May 1.   (Buchanan et al. 1985). Zoeae of C. magister are
    In Washington, the season in Prince William Sound    among the most sensitive life stages to insecticides
    opens April 1. Only male crabs .165 mm CW are legal   and fungicides (Buchanan et al. 1970, Armstrong et al.
    in these areas (Eaton 1985, Kimker 1985a, Koeneman    1976, Caldwell et al. 1979).
    1985). The commercial season may last 9 months, but
    most crabs are captured within the first 2 months. The

                                                        68

                                                                                     Dungeness crab continued
                                                          coastal waters and probably all bays and estuaries
      Table 1. Relative abundance of Dungeness crab        from Morro Bay, California (Soule and Tasto 1983), to
               in 32 U.S. Pacific coast estuaries.         Puget Sound, Washington (Table 1).
                        Life Stage
            Estuary    A M J L E                           Life Mode
             Puget Sound (i  0   O O   Relative abundance:  Eggs adhere to pleopods of the epibenthic-living adult
             Hood Canal O O �  0  0    �   Highly abundant  female. Larvae (zoeae) are planktonic. Post-larvae
              Skagit Bay (3 0 6 o 0 I     Abundant        (megalopae) are primarily planktonic, but become
            Grays Harbor 0  O             Common          mostly benthic when close to molting (Reilly 1983a).
             Willapa Bay 0    a O   Blank  iot present    Megalopae can actively swim and sometimes form
           Columbia River O    O                           "swarms"nearthesurface(Lough 1976, Hatfield 1983).
            Nehalem Bay       e� �                         Megalopae are often found on the hydrozoan Velella
            Tillamook Bay             Life stage:          velella (Wickham 1979, Stevens and Armstrong 1985).
             Netarts Bay DO  * �      M- Mating           Juveniles and adults are epibenthic.
              Siletz River 0    (3 3   J - Juveniles
                   ~Yaquina Bay (     ~L- Larvae
             Yaquina Bay 1 :  a�       E- Eggs             Habitat
             AlsesaRiver  a a                             Iype: Eggs adhere to pleopods of female crabs in
            Siuslaw River  � �  euhaline (30-40%0) waters. Females with eggs can be
               UmouBayRiver  0 a             a             found intertidally and in deeper nearshore waters
               CoRogue River  0 C                          (MacKay 1942). Larvae initially occur in nearshore
            Klamath River O    O                           euhaline waters (5-16 km from shore) (Lough 1976,
            KHumboldthBay   o                              Orcutt 1977, Reilly 1983a), with offshore movement
               EelRiver 0     0                           and distribution influenced by depth, latitude,
            Tomales Bay O  a  C                           temperature, salinity, and currents (Reilly 1983a, 1985).
       Cent. San Fran. Bay *    /   a .   * tIndudes Central San  Larvae are found near the surface at night and 15-25
       South San Fran. Bay  -   0   Fr an bbaysu Sun,     mdeepduringdaylight (Reilly 1983a, 1985). Megalopae
           Elkhorn Slough  4 a  ,i                        are primarily found in shallow nearshore areas (Lough
              Morro Bay i  1976, Hatfield 1983, Reilly 1983a). Megalopaeoccupy
         Santa Monica Bay                                  theupper 15 m both day and night (Reilly 1983a, 1985),
           San Pedro Bay                                   but they also have diel migrations (Booth et al. 1985).
            Alamitos Bay                                  Juveniles occur primarily in shallow coastal waters and
            Anaheim Bay                                   estuaries (Butler 1956, Orcutt et al. 1975, Stevens and
             Newport Bay                                   Armstrong 1984, 1985). Adults are found primarily
             Mission Bay                                  intertidally to 90 m depths in marine (euhaline) waters,
           San Diego Bay                                  but sizable numbers occur in the lower reaches of
           Tijuana Estuary                                 estuaries.
                       AM J LE
                                                          Substrate: The Dungeness crab is found over various
                                                          substrates. Juveniles are often found intertidally in
    Ecological: The Dungeness crab is important as both a    estuarine areas of soft substrate containing eelgrass
    predator (on Crangon spp. shrimp and bivalves) and    (Zostera spp.) and bivalve shells (Armstrong and
    prey species in nearshore and estuarine habitats.    Gunderson 1985). Adults can be found on mud, rock,
    Estuaries are very important to early life stages (Tasto    and gravel bottoms, but they prefer sand (Frey 1971,
    1983, Armstrong and Gunderson 1985, Emmett and    Karpov 1983, Rudy and Rudy 1983).
    Durkin 1985).
                                                          Phvsical/Chemical Characteristics: Salinity tolerance
    Range                                                  varies with life stage (Pauley et al. 1986), but small
    Overall: This species occurs from Santa Barbara,    juvenilesdonot appeartobemoretolerantthanadults
    California in the south, to the Pribilof Islands    (Stevens and Armstrong 1985). Eggs hatch over a
    (southeastern Bering Sea) in the north (Schmitt 1921,    wide range of salinities, but survival is best in euhaline
    MacKay 1942, Pauley et al. 1986). It does not occuroff    waters (Pauley et al. 1986). Larvae are highly sensitive
    Baja California (Garth and Abbott 1980). It is found    tosalinityvariationsandarefoundprimarilyineuhaline
    along the Pacific coast in intertidal waters down to 420    waters (Buchanan and Milleman 1969, Lough 1976,
    m, but is not abundant at depths below 90 m.           Reilly 1983a). The interaction between salinity and
                                                          temperature can significantly affect larval survival. At
    Within Studv Area: The Dungeness crab occurs in   lowertemperatures (<10�C) eggs take longerto hatch

                                                       69

    Dungeness crab continued
    and have lower hatching mortality rates (Mayer 1973,    Mating usually occurs when the female is soft-shelled.
    Wild 1983). Larval survival is best when temperatures    To accomplish this, the male may hold the female in a
    are 10.0-14.0�Candsalinities are25-30%0(Reed 1969,    premating embrace for up to 7 days before she molts
    Pauleyetal. 1986);larvaewillnotsuccessfullydevelop    (Snow and Neilsen 1966). After she molts, the male
    to megalopae at 200C (Sulkin and McKeen 1989).   inserts his gonopods into the spermathecae of the
    Juvenile and adult crabs in estuaries are exposed to   female and deposits spermatophores. The male may
    rapidly changing salinities which they respond to by    remain with the female for two days to insure her
    pulsing, closure(Surgarmanetal. 1983), and movement    protection (Snow and  Neilsen  1966).   The
    (Stevens et al. 1984).  Mating takes place at   spermatophores remain viable in the female for many
    temperatures of 8.0-17.00C (Pauley et al 1986). Water    months and fertilize the eggs when they are extruded
    temperatures >20.0-25.0�C may cause juvenile and    (MacKay 1942, Wild 1983). Males can mate with more
    adult mortalities, depending on other environmental   than one female.
    factors (Wild 1983, Pauley et al. 1986).
                                                          Fecundity: Eggs are extruded in the fall and winter;
    Miarationsand Movements:Beforespreadingoffshore,    from September to February in British Columbia
    larvae initially appear in nearshore waters 5-16 km    (MacKay 1942, Butler 1956), October to December in
    fromshorein December(off California) and lateJanuary    Washington (Cleaver 1949), October to March in
    (off Oregon). Megalopae appear in early March to mid-    Oregon (Waldron 1958), and September to November
    April in California and Apriloff Oregon and Washington    in California (Orcutt et al. 1975, Wild 1983). A female
    (Lough 1976, Reilly 1983a, Pauley et al. 1986). Both    mayhave3or4broodsinalifetime(MacKay1942)and
    larvae and megalopae undertake daily vertical   can carry up to 2.5 million eggs (Wickham 1980), but
    migrations, being at the surface at night (Reilly 1983a,    the actual number that hatch is much less (Wild 1980,
    Booth et al. 1985, Shenker 1988). Tidal currents and    1983). Females have to be buried in sand for eggs to
    self-propulsion bring megalopae within 1 km of shore    adhereproperlytopleopods(Wild 1983). Eggsforman
    and into estuaries in Oregon (Lough 1976). Megalopae    orange "sponge" that gets darker as the eggs mature.
    may also "ride"the hydrozoan Velella velellato inshore
    waters (Wickham 1979). Early juveniles settle out in   Growth and Development
    shallow water estuarine areas or adjacent marine    Eaa size and Embryonic Develooment: Eggs are 0.4-
    waters (Tasto 1983, Stevens and Armstrong 1985),   0.6 mm in diameter, and smaller at higher incubation
    and also settle on tidal flats at high tide (Stevens and    temperatures (Wild 1983). Embryonic development is
    Armstrong 1984, Armstrong and Gunderson 1985).   indirect and external. Egg incubation takes 64-128
    Adult crabs move out of estuaries to mate, but there are   days depending on temperature (Cleaver 1949, Orcutt
    always some adults in estuaries. Whiletagging studies    1978, Wild 1983). Upon hatching, crabs emerge as
    have shown that adult Dungeness crabs can move    prezoeae and moltto zoeae within one hour. (Buchanan
    over a wide area, most exhibit limited random    and Milleman 1969).
    movements (Waldron 1958, Diamond and Hankin
    1985). However, there is some evidence that male    Aae and Size of Larvae: Larvae are 2.5-11.0 mm in
    crabs move northward and into shallow waters during   length (Poole 1966). The larvae moltthrough five zoeal
    winter and southward and deeper during summer    stages before metamorphosing into megalopae (Poole
    (Gotshall 1978). Diel movements to intertidal habitats    1966, Lough 1976). The megalopaisthefinalplanktonic
    may be a result of food availability (Stevens et al.   stage; it molts to become the initial juvenile instar
    1984).                                                (Reilly 1983a, 1985).

    Reproduction                                         Juvenile Size Ranae: Juveniles range in size from 5.0
    Mode:TheDungenesscrabisgonochoristic, oviparous,    mm CW to about 100 mm CW (larger for males)
    anditeroparous. Eggsarefertilizedwhilebeingextruded    (Cleaver 1949, Waldron 1958,  Butler 1960, 1961,
    by the female.                                        Poole 1967). Crabs may molt 11 or 12 times before
                                                          reaching sexual maturity (Butler 1961). Juveniles in
    Matina/Soawnina: Mating occurs from April to   estuariesgrowfasterthanjuvenilesresidingincoastal
    September in British Columbia (MacKay 1942, Butler   waters. Subyearling crabs in Grays Harbor and Willapa
    1956), primarily from March to April (but sometimes to   Bay, Washington, grew to 40 mm CW by Septemberof
    June) in Washington (Cleaver 1949, Pauley et al.   their first year (Gunderson et al. 1990).
    1986), and from March to July in California (Pauley et
    al. 1986). Mating takes place in non-estuarine locations,    Aae and Size of Adults: The Dungeness crab matures
    with males finding females via the possible aid of   after approximately two years when 116 mm CW
    pheromones (Knudsen 1964, Pauley et al. 1986).    (males) or 100 mm CW  (females) (Butler 1960, 1961).

                                                       70

                                                                                     Dungeness crab continued
    Some male crabs reach harvestable size three years   cyclic natureof crab abundance. The successof a year
    after settlement, and most males reach this size after   class is probably determined by larval survival to
    four years (Warner 1987, Smith and Jamieson 1989).    metamorphosis, thus factors which influence egg,
    This species can live up to 8-10 years and reach a size   larvae, and megalopae survival are very important
    of 218 mm CW (males), and 160 mm CW (females)    (Peterson 1973, Lough 1976, Pauley et al. 1986).
    (MacKay 1942, Butler 1961).                           Factors which affect larval survival include predation,
                                                          extreme water temperatures, currents, and food
    Food and Feeding                                      availability (Lough 1976). Other causes of mortality
    TroDhic Mode: Larvae are planktivorous. Juveniles    which may influence population abundance include
    and adults are carnivorous.                           egg predation by C. errans (Wickham 1980), megalopae
                                                          predation by salmon (Reilly 1983b), and diseases
    Food Items: Larvae and megalopae eat phytoplankton    (Stevens and Armstrong 1981). Commercial trawling
    and zooplankton, but primarily zooplankton (Lough   kills approximately 53 crabs pertrawling hour (males)
    1976, Ebert et al. 1983).  Juvenile crabs eat fish,   in California (Reilly 1983c). Finally, estuaries play a
    molluscs, and crustaceans (Butler 1954, Gotshall 1977,   vital role in Dungeness crab abundance. Estimates of
    Stevens et al. 1982). Shrimp (Crangonspp.) appearto    juvenile crab populations in Willapa Bay and Grays
    be a preferred prey for juveniles that are 61-100 mm    Harbor showed that these two systems contribute
    CW in Grays Harbor (Stevens et al. 1982).  Larger   substantially to future crab catches (Stevens and
    juveniles often cannibalize smaller crabs (MacKay    Armstrong 1984,1985). Estuariesare important nursery
    1942, Butler 1954, Gotshall 1977, Stevens et al. 1982).    habitats for subyearling and yearling crabs (Gunderson
    Adults also eat fish, molluscs, and crustaceans, and    et al. 1990). Hence, dredging and habitat modification
    are nonspecific feeders that alter their food habits as    projects in estuaries should consider the potential
    prey abundances fluctuate (Gotshall 1977). In general,    impacts on crab populations (Armstrong and Gunderson
    crabs eat bivalves their first year, Crangon spp. their   1985, Emmett and Durkin 1985, Pauley et al. 1986,
    second year, and fish their third year (Stevens et al.   McGraw et al. 1988).
    1982).
                                                          References
    Biological Interactions
    Predation: Dungeness crab eggs are consumed by a    Armstrong, D. A., and D. R. Gunderson. 1985. The
    nemertean (Carcinonemertes errans) which can cause    role of estuaries in Dungeness crab early life history: a
    large losses in egg production (Wickham 1980). Larvae    case study in Grays Harbor, Washington.  In B. R.
    are eaten by planktivorous fishes such as Pacific   Melteff (coordinator), Proceedings of the symposium
    herring ( Clupea pallasi), Pacific sardine (Sardinops    on Dungeness crab biology and management, p. 145-
    sagax), and others (Garth and Abbott 1980, Pauley et   170.  Lowell Wakefield Fisheries Symposia Series,
    al. 1986). Megalopae are eaten by rockfish (Sebastes    Univ. Alaska, Alaska Sea Grant Rep. No. 85-3,
    spp.), coho salmon (Oncorhynchus kisutch), and    Fairbanks, AK.
    chinook salmon (Oncorhynchus tshawytscha), and
    probablyotherfishes (Prince and Gotshall 1976, Emmett    Armstrong, D. A., D. V. Buchanan, and R. S. Caldwell.
    et al. 1986). Juveniles are eaten by many species of   1976.  A mycosis caused by Lagenidium sp. in
    fish, including starry flounder (Platichthys stellatus),   laboratory-reared larvaeof the Dungenesscrab, Cancer
    English sole ( Pleuronectes vetulus), rock sole    magister, and possible chemicaltreatments. J. Invert.
    (Lepidopsetta bilineata), lingcod (Ophiodon elongatus),    Pathol. 28: 329-336.
    cabezon (Scorpaenichthys marmoratus), wolf-eel
    (Anarrhichthysocellatus), rockfish, sturgeon (Acipenser   Barry, S. 1985. Overview of the Washington coastal
    spp.), sharks, skates, Pacific halibut (Hippoglossus    Dungeness crab fishery. In B. R. Melteff (coordinator).
    stenolepis), and others (Waldron 1958, Orcutt 1977,    Proceedings of the symposium on Dungeness crab
    Reilly 1983b).  Other important predators include    biology and management, p. 33-36. Lowell Wakefield
    Octopus spp. and sea otters (Enhydra lutris) (Kimker    Fisheries Symposia Series, Univ. Alaska, Alaska Sea
    1985b).  Adults are consumed by humans, harbor    Grant Rep. No. 85-3, Fairbanks, AK.
    seal (Phoca vitulina), sea lions, and gulls.
                                                          Booth, J., A. Phillips, and G. S. Jamieson. 1985. Fine
    Factors Influencina Porulations: Upwelling (Peterson    scale spatial distribution of Cancermagistermegalopae
    1973), cannibalism (Botsford and Wickham 1978), sea    and its relevance to sampling methodology. In B. R.
    surface temperature (Wild 1980), sunspot number    Melteff (coordinator), Proceedings of the symposium
    (Love and Westphal 1981), and wind stress (Johnson    on Dungeness crab biology and management, p. 273-
    et al. 1986) have been proposed as causes for the    286  Lowell Wakefield Fisheries Symposia Series,

                                                       71

    Dungeness crab continued
    Univ. Alaska, Alaska Sea Grant Rep. No. 85-3,    and seaweed harvests of Puget Sound. Wash. Sea
    Fairbanks, AK.                                          Grant, Univ. Wash. Press, Seattle, WA, 164 p.

    Botsford, L. W., and D. E. Wickham. 1978. Behavior    Cleaver, F. C. 1949. Preliminary results of the coastal
    of age-specific, density-dependent models and the    crab (Cancermagister) investigation. Biol. Rep. 49A:74-
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    fishery. J. Fish. Res. Board Can. 35(6):833-843.
                                                           Dahlstrom, W. A., and P. W. Wild. 1983. A history of
    Bowman, T. E., and L. G. Abele. 1982. Classification    Dungeness crab fisheries in California. In P. W. Wild
    of the recent crustacea. In L. G. Abele (editor), D. E.    and R. N. Tasto (editors), Life history, environment,
    Bliss (editor-in-chief), The biologyof Crustacea, Volume    and mariculture studies ofthe Dungeness crab, Cancer
    1. Systematics, the fossil record, and biogeography, p.   magister, with emphasis on the central California fishery
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    Buchanan, D. W., and R. E. Milleman.  1969. The    Demory, D. 1985. An overview of Oregon Dungeness
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    Fairbanks, AK.                                          Lowell Wakefield Fisheries Symposia Series, Univ.
                                                           Alaska, Alaska Sea Grant Rep. No. 85-3, Fairbanks,
    Butler, T. H. 1954. Food of the commercial crab in the    AK.
    Queen Charlotte Islands region. Fish. Res. Board Can.
    Prog. Rep. 99:3-5.                                      Ebert, E. E., A. W. Hazeltine, J. L. Houk, and R. O.
                                                           Kelly. 1983. Laboratory cultivation of the Dungeness
    Butler, T. H. 1956. The distribution and abundance of   crab, Cancer magister. In P. W. Wild and R. N. Tasto
    early larval stages of the British Columbia commercial    (editors), Life history, environment, and mariculture
    crab. Fish. Res. Board Can. Prog. Rep. 107:22-23.       studies of the Dungeness crab, Cancermagister, with
                                                           emphasis on the central California fishery resource.
    Butler, T. H. 1960. Maturity and breeding ofthe Pacific   Calif. Fish Game, Fish Bull. 172:259-309.
    edible crab, Cancer magister Dana. J. Fish. Res.
    Board Can. 17(5):641-646.                               Emmett, R. L., and J. T. Durkin. 1985. The Columbia
                                                           River estuary: an important nursery for Dungeness
    Butler, T. H. 1961. Growth and age determination of   crabs, Cancermagister. Mar. Fish. Rev. 47(3):21-25.
    the Pacific edible crab, CancermagisterDana. J. Fish.
    Res. Board Can. 18(5):873-889.                          Emmett, R. L., D. R. Miller, and T. H. Blahm. 1986.
                                                           Food of juvenile chinook, Oncorhynchus tshawytscha,
    Caldwell, R. S., D. V. Buchanan, D. A. Armstrong, M.    and coho, 0. kisutch, salmon off the northern Oregon
    H. Mallon, and R. E. Millemann. 1979. Toxicity of the    and southern Washington coasts, May-September
    herbicides 2,4-D, DEF, Propanil and Trifluralin to the    1980. Calif. Fish Game 72(1):38-46.
    Dungeness crab, Cancer magister. Arch. Environ.
    Contam. Toxicol. 8:383-396.                             Frey, H. W. 1971. California's living marine resources
                                                           and their utilization. Calif. Dept. Fish Game,
    Cheney, D. P., and T. F. Mumford, Jr. 1986. Shellfish    Sacramento, CA, 148 p.

                                                        72

                                                                                    Dungeness crab continued
    Garth, J. S., and D. P. Abbott. 1980. Brachyura: the    P. Russell, and P. W. Wild.  1983.  The effects of
    true crabs. In R. H. Morris, D. P. Abbott, and E. C.    chlorinationofwastewateronjuvenileDungenesscrabs
    Haderlie (editors). Intertidal invertebrates of California,   in San Francisco Bay waters. In P. W. Wild and R. N.
    p. 594-630. Stanford Univ. Press. Stanford, CA.         Tasto (editors), Life history, environment,  and
                                                          mariculture studies of the Dungeness crab, Cancer
    Gotshall, D. W. 1977. Stomach contents of northern    magister, withemphasisonthecentralCaliforniafishery
    California Dungeness crabs (Cancermagister). Calif.   resource. Calif. Fish Game, Fish Bull. 172:215-225.
    Fish Game 63(1):43-51.
                                                          Johnson, D. F., L. W. Botsford, R. D. Methot, Jr., and
    Gotshall, D. W. 1978. Northern California Dungeness    T. C. Wainwright. 1986. Wind stress and cycles in
    crab, Cancer magister, movements as shown by    Dungeness crab (Cancermagister) catch off California,
    tagging. Calif. Fish Game 64(4):234-254.                Oregon and Washington.  Can. J. Fish. Aquat. Sci.
                                                          43:838-845.
    Guard, H. E., L. H. DiSalvo, J. Ng, and P. W. Wild.
    1983.  Hydrocarbons in Dungeness crabs, Cancer    Karinen, J. F., S. D. Rice, and M. M. Babcock. 1985.
    magister, and estuarine sediments. In P. W. Wild and    Reproductive success in Dungeness crab (Cancer
    R. N. Tasto (editors), Life history, environment, and    magister) during long-term  exposures to oil-
    mariculture studies of the Dungeness crab, Cancer    contaminateed sediments. In Final Reports of Principal
    magister, with emphasison thecentral California fishery    Investigators, Vol. 67, p. 435-461. Outer Cont. Shelf
    resource. Calif. Fish Game, Fish Bull. 172:243-257.     Environ. Assm. Prog., U.S. Dept. Comm., Nat. Ocean.
                                                          Atmos. Adm., Ocean Asses. Div., U.S. Dept. Int., Min.
    Gunderson, D. R., D. A. Armstrong, Y.-B. Shi., and R.    Man. Serv., MMS 90-0044.
    A. McConnaughey. 1990. Patterns of estuarine use by
    juvenile English sole (Parophrys vetulus) and    Karpov, K.A. 1983. Effectofsubstratetypeonsurvival
    Dungeness crab (Cancermagister). Estuaries 13(1):59-    and growth in high densitycommunal culturesof juvenile
    71.                                                     Dungeness crabs, Cancermagister. In P. W. Wild and
                                                          R. N. Tasto (editors), Life history, environment, and
    Hart, J. F. L. 1982. Crabs and their relatives of British    mariculture studies of the Dungeness crab, Cancer
    Columbia.   British Columbia Provincial Museum    magister, with emphasisonthecentralCalifornia fishery
    HandbookNo. 40. BritishColumbia Provincial Museum,    resource. Calif. Fish Game, Fish Bull. 172:311-318.
    Victoria, B.C., 267 p.
                                                          Kimker, A. 1985a. Overview of the Prince William
    Hatfield, S. E. 1983. Intermolt staging and distribution    Sound management area Dungeness crab fishery. In
    of Dungeness crab, Cancer magister, megalopae. In   B. R. Melteff (coordinator),   Proceedings of the
    P. W. Wild and R. N. Tasto (editors), Life history,    symposium  on  Dungeness crab biology and
    environment, and mariculturestudiesofthe Dungeness    management, p. 77-83. Lowell Wakefield Fisheries
    crab, Cancer magister, with emphasis on the central    Symposia Series, Univ. Alaska, Alaska Sea Grant
    California fishery resource. Calif. Fish. Game, Fish    Rep. No. 85-3, Fairbanks, AK.
    Bull. 172:85-96.
                                                          Kimker, A. 1985b. A recent history of the Orca Inlet,
    Haugen, C. W. 1983a. Field and laboratory studies of    Prince William Sound Dungeness crab fishery with
    toxictraceelementsin Dungenesscrabs. InP.W.Wild    specific reference to sea otter predation.  In B. R.
    and R. N. Tasto (editors), Life history, environment,    Melteff (coordinator), Proceedings of the symposium
    and mariculture studiesof the Dungenesscrab, Cancer   on Dungeness crab biology and management, p. 231 -
    magister, with emphasison thecentral California fishery    241.  Lowell Wakefield Fisheries Symposia Series,
    resource. Calif. Fish Game, Fish Bull. 172:227-238.     Univ. Alaska, Alaska Sea Grant Rep. No. 85-3,
                                                          Fairbanks, AK.
    Haugen, C. W. 1983b. Chlorinated hydrocarbon
    pesticides and polychlorinated biphenyls in Dungeness    Knudsen, J. W. 1964. Observationsofthe reproductive
    crabs. In P. W. Wild and R. N. Tasto (editors), Life   cycles and ecology of the common Brachyura and
    history, environment, and mariculture studies of the    crablike Anomura of Puget Sound, Washington. Pac.
    Dungeness crab, Cancer magister, with emphasis on    Sci. 18(1):3-33.
    the central California fishery resource. Calif. Fish
    Game, Fish Bull. 172:239-241.                           Koeneman, T. M. 1985. Abrief reviewofthecommercial
                                                          fisheries for Cancer magisterin southeast Alaska and
    Horne, A. J., M. Bennett, R. Valentine, R. E. Selleck, P.    Yakutat waters, with emphasis on recent seasons. In

                                                       73

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    B. R. Melteff (coordinator),  Proceedings of the   82-4, Washington, D.C, 20 p.
    symposium on Dungeness crab biology and
    management, p. 61-76. Lowell Wakefield Fisheries    Peterson, W. T. 1973. Upwelling indices and annual
    Symposia Series, Univ. Alaska, Alaska Sea Grant    catches of Dungeness crab, Cancer magister, along
    Rep. No. 85-3, Fairbanks, AK.                         the west coast of the United States. Fish. Bull., U.S.
                                                          22(3):902-910.
    Lough, R. G. 1976. Larval dynamics of the Dungeness
    crab, Cancer magister, off the central Oregon coast,    Poole, R. L. 1966. A description of the laboratory-
    1970-71. Fish. Bull., U.S. 74(2):353-375.             reared zoeae of CancermagisterDana, and megalopae
                                                          taken under natural conditions (Decapoda, Brachyura).
    Love, M. S., and M. V. Westphal. 1981. A correlation   Crustac. 11(1):83-97.
    between annual catches of Dungeness crab, Cancer
    magister, along the west coast of North America and    Poole, R. L. 1967. Preliminary results of the age and
    mean annual sunspot number. Fish. Bull., U.S. 79:794-    growth study of the market crab, Cancer magister, in
    796.                                                  California: the age and growth of Cancer magister in
                                                          Bodega Bay. In Proceedings of the symposium on
    MacKay, D. C. G.  1942. The Pacific edible crab,   crustacea. Mar. Biol. Assoc. India, Ernakulam, Part
    Cancer magister. Fish. Res. Board Can., Bull. 621:1-   11:553-567.
    32.
                                                          Prince, E. D., and D. W. Gotshall. 1976. Food of the
    Mayer, D. L. 1973. Theecologyandthermalsensitivity   copper rockfish, Sebastes caurinus Richardson,
    of the Dungeness crab, Cancermagister, and related    associated with an artificial reef in south Humboldt Bay,
    species of its benthic community in Similk Bay,    California. Calif. Fish Game 64(4):274-285.
    Washington. Ph.D. Thesis. Univ. Wash., Seattle, WA,
    188 p.                                                Reed, P. H.  1969. Culture methods and effect of
                                                          temperature and salinity on survival and growth of
    McGraw, K. A., L. L. Conquiest, J. O. Waller, P. A.    Dungeness crab (Cancer magister) larvae in the
    Dinnel, and D. A. Armstrong. 1988. Entrainment of   laboratory. J. Fish. Res. Board Can. 26(2):389-397.
    Dungeness crabs, Cancermagister, by hopper dredge
    in Grays Harbor, Washington. J. Shellfish Res. 7(2):219-   Reilly, P. N. 1983a. Dynamics of the Dungeness crab,
    231.                                                  Cancer magister, larvae off central and northern
                                                          California. In P. W. Wild and R. N. Tasto (editors), Life
    National Marine Fisheries Service. 1986. Fisheriesof   history, environment, and mariculture studies of
    the United States, 1985. Current Fishery Statistics No.    Dungeness crab, Cancer magister, with emphasis on
    8368. U.S. Dept. Comm., NOAA, Nat. Mar. Fish Serv.,   the central California fishery resource.  Calif. Fish
    Nat. Fish. Stat. Prog., Washington, D.C., 122 p.      Game, Fish Bull. 172:57-84.

    Orcutt, H. G. 1977. Dungeness crab research program.    Reilly, P. N. 1983b. Predation on Dungeness crabs,
    Marine Res. Rep. No. 76-16. Calif. Dept. Fish Game,    Cancer magister, in central California. In P. W. Wild
    Sacramento, CA, 55 p.                                 and R. N. Tasto (editors), Life history, environment,
                                                          and mariculture studies of Dungeness crab, Cancer
    Orcutt, H. G., R. N. Tasto, P. W. Wild, C. W. Haugen,    magister,withemphasisonthecentral Californiafishery
    and P. C. Collier. 1975. Dungeness crab research    resource. Calif. Fish Game, Fish Bull. 172:155-164.
    program. Marine Res. Rep. No. 75-16. Calif. Dept.
    Fish Game, Sacramento, CA, 77 p.                      Reilly, P. N. 1983c. Effects of commercial trawling on
                                                           Dungeness crab survival. In P. W. Wild and R. N. Tasto
    Pacific Marine Fisheries Commission.  1987. 39'th   (editors), Life history, environment, and mariculture
    annual report of the Pacific Marine Fisheries    studies of Dungeness crab, Cancer magister, with
    Commissionfortheyear 1986. Pac. Mar. Fish. Comm.,    emphasis on the central California fishery resource.
    Portland, OR, 29 p.                                   Calif. Fish Game, Fish Bull. 172:165-169.

    Pauley, G. B., D. A. Armstrong, and T. W. Heun. 1986.    Reilly, P. N. 1985. Dynamics of the Dungeness crab,
    Species profiles: life histories and environmental    Cancer magister, larvae off central and northern
    requirementsofcoastal fishes and invertebrates (Pacific   California. In B. R. Melteff (coordinator), Proceedings
    Northwest)-Dungeness crab. U.S. Fish Wildl. Serv.   of the symposium on Dungeness crab biology and
    Biol. Rep. 82(11.63). U.S. Army Corps Eng., TR EL-    management, p. 245-272. Lowell Wakefield Fisheries

                                                        74

                                                                                    Dungeness crab continued
    Symposia Series, Univ. Alaska, Alaska Sea Grant    importance. Mar. Biol. (Berlin) 72(1):135-145.
    Rep. No. 85-3, Fairbanks, AK.
                                                          Stevens, B. G., D. A. Armstrong, and J. C. Hoeman.
    Rudy, P., Jr., and L. H. Rudy. 1983. Oregon estuarine    1984.  Diel activity of an estuarine population of
    invertebrates - an illustrated guide to the common and    Dungeness crabs, Cancer magister, in relation to
    important invertebrate animals. U.S. Fish Wildl. Serv.,   feeding and environmental factors.  J. Crust. Biol.
    Biol. Serv. Prog. FWS/OBS-83/16, Portland, OR,        4(3):390-403.
    225 p.
                                                          Sulkin, S. D., and G. L. McKeen. 1989. Laboratory
    Schmitt, W. L. 1921. The marine decapod crustacea    studyof survival anddurationof individual zoeal stages
    of California. Univ. Calif. Publ. Zool., No. 23, 470 p.    as a function of temperature in the brachyuran crab
                                                          Cancer magister. Mar. Biol. 103:31-37.
    Shenker, J. M. 1988. Oceanic associations of neustonic
    larval and juvenile fishes and Dungeness crab    Sugarman, P. C., W. H. Pearson, and D. L. Woodruff.
    megalopae off Oregon.  Fish. Bull., U.S. 86(2):299-    1983. Salinitydetectionandassociatedbehaviorinthe
    317.                                                  Dungeness crab, Cancermagister. Estuaries 6(4):380-
                                                          386.
    Smith, B. D., and G. S. Jamieson. 1989. Growth of
    male and female Dungeness crabs nearTofino, British   Tasto, R. N. 1983. Juvenile Dungeness crab, Cancer
    Columbia. Trans. Am. Fish. Soc. 118:556-563.          magister, studies in the San Francisco Bay area. In P.
                                                          W. Wild and R. N. Tasto (editors), Life history,
    Snow, C. D., and J. R. Neilsen. 1966. Premating and    environment, and mariculturestudiesofthe Dungeness
    mating behavior of the Dungeness crab (Cancer   crab, Cancer magister, with emphasis on the central
    magisterDana). J. Fish. Res. Board Can. 23(9):1319-   California fishery resource.  Calif. Fish Game, Fish
    1323.                                                 Bull. 172:135-154.

    Soule, M., and R. N. Tasto. 1983. Stock identification   Waldron, K. D. 1958. The fishery and biology of the
    studiesonthe Dungenesscrab, Cancermagister. InP.    Dungeness crab (Cancer magister Dana) in Oregon
    W. Wild and R. N. Tasto (editors), Life history,   waters. Oregon Fish Comm., Contrib. No. 24:1-43.
    environment, and mariculture studies of the Dungeness
    crab, Cancer magister, with emphasis on the central   Warner, R. W.  1985.  Overview of the California
    California fishery resource. Calif. Fish Game, Fish    Dungeness crab, Cancer magister, fisheries. In B. R.
    Bull. 172:39-42.                                      Melteff (coordinator), Proceedings of the symposium
                                                          on Dungeness crab biology and management, p. 11-
    Stevens, B. G., and D. A. Armstrong.  1981. Mass    26.  Lowell Wakefield Fisheries Symposia Series,
    mortalityof female Dungenesscrab, Cancermagister,    Univ. Alaska, Alaska Sea Grant Rep. No. 85-3,
    on the southern Washington coast. Fish. Bull., U.S.    Fairbanks, AK.
    79(2):349-352.
                                                          Warner, R. W. 1987. Age and growth of male
    Stevens, B. G.,and D. A. Armstrong. 1984. Distribution,    Dungeness crabs, Cancer magister, in northern
    abundance and growth of juvenile Dungeness crabs,   California. Calif. Fish Game 73:4-20.
    Cancermagister, in Grays Harbor estuary, Washington.
    Fish. Bull., U.S. 82(3):469-483.                      Wickham, D. E.  1979.  The relationship between
                                                          megalopae of the Dungeness crab, Cancer magister,
    Stevens, B. G., and D. A. Armstrong. 1985. Ecology,    and the hydroid, Velella velella, and its influence on
    growth, and population dynamics of juvenile Dungeness    abundance estimates of C. magistermegalopae. Calif.
    crab, Cancer magister Dana, in Grays Harbor,    Fish Game 65(3):184-186.
    Washington 1980-1981. In B. R. Melteff (coordinator),
    Proceedings of the symposium on Dungeness crab   Wickham, D. E. 1980. Aspects of the life history of
    biology and management, p. 119-134. Lowell Wakefield    Carcinonemertes        errans       (Nemertea:
    Fisheries Symposia Series, Univ. Alaska, Alaska Sea    Carcinonemertidae), an egg predatorof thecrab Cancer
    Grant Rep. No. 85-3, Fairbanks, AK.                   magister. Biol. Bull. (Woods Hole) 159:247-257.

    Stevens, B. G., D. A. Armstrong, and R. Cusimano.    Wild, P. W. 1980. Effects of seawatertemperature on
    1982. Feeding habits of the Dungeness crab Cancer   spawning, egg development, hatching success, and
    magister as determined by the index of relative   populationfluctuationsoftheDungenesscrab, Cancer

                                                      75

    Dungeness crab continued
    magister. Calif. Coop. Ocean. Fish. Invest. Rep. 21:115-
    120.

    Wild, P. W. 1983. The influence of seawater
    temperature on spawning, egg development, and
    hatching success of the Dungeness crab, Cancer
    magister. In P. W. Wild and R. N. Tasto (editors), Life
    history, environment, and mariculture studies of the
    Dungeness crab, Cancer magister, with emphasis on
    the central California fishery resource. Calif. Fish
    Game, Fish Bull. 172:197-214.

                                                      76

    77

        Triakis semifasciata
       Adult

                25 cm

    Common Name: leopard shark                              California beaches (Miller and Lea 1972). It is the most
    Scientific Name: Triakis semifasciata                   abundant shark in San Francisco Bay (Ebert 1986) and
    Other Common Names: cat shark, sand tiger               is common near jetties and piers (Talent 1976).
    Classification (Robins et al. 1980)
    Phylum: Chordata                                        Range
    Class: Osteichthyes                                     Overall: Overall range of this species is from Baja
    Order: Carcharhiniformes                                Mexico, to southern Oregon.  It is also found in the
    Family: Triakidae                                       northern Gulf of California (Miller and Lea 1972,
                                                           Eschmeyer et al. 1983).
    Value
    Commercial: The leopard shark is caught and sold    Within Studv Area: The leopard shark inhabits most
    commerciallyyear-round, but it is not normallytargeted    California estuaries and bays, but is primarily found
    by commercial fishermen. However, a limited longline    south of Tomales Bay (Table 1) (Monaco et al. 1990).
    fishery exists in San Francisco Bay, California (S.
    Smith, National Marine Fisheries Service, La Jolla,   Life Mode
    California, unpubl. manuscr.). The meat is considered    The leopard shark is a live-bearer; eggs are fertilized
    excellent and is sold fresh and fresh-frozen (Compagno    internally and embryogenesis occurs within the female.
    1984). This species was not sought during early shark    Juveniles and adults are demersal, sometimes resting
    fisheries because its liver does not contain high    on the bottom (Feder et al. 1974).
    concentrations of vitamin A (Roedel and Ripley 1950).
                                                           Habitat
    Recreational: This species is a valuable sport fish in  Tipe: This shark is a neritic species found primarily in
    nearshore shallow waters of central and southern    polyhaline to euhaline waters. It is most common in
    California.  Important sport fisheries exist in San    waters <3.7 m deep, but may occur down to 91 m
    Francisco Bay and Elkhorn Slough, California (Herald    (Eschmeyer et al. 1983, Compagno 1984). Estuaries
    and Ripley 1951, Smith and Kato 1979).                  appear to be used as pupping and feeding/rearing
                                                           areas (Ackerman 1971, Talent 1973, Barry and Cailliet
    Indicator of Environmental Stress: Concentrations of    1981).
    polychlorinated biphenyls of 46.9 ppm have beenfound
    in leopard sharks in San Francisco Bay (Russo 1975).   Substrate: Juveniles and adults prefer sandy or muddy
    However, it is not known how or at what levels   flats, but they may also be found over cobble bottoms,
    contaminants affect leopard shark biology.              and near rocky reefs and kelp beds (Feder et al. 1974)

    Ecological: The leopard shark is one of the most    Phvsical/Chemical Characteristics: The leopard shark
    common sharks in California bays and estuaries (Talent   is a marine species, but no information is available
    1973, de Wit 1975, Ebert 1986) and along southern    concerning salinity tolerances.  However, sharks

                                                        78

                                                                                      Leopard shark continued
                                                          nomadic, spending a few hours in one location and
      Table 1. Relative abundance of leopard shark        then moving to another area (Compagno 1984).
               in 32 U.S. Pacific coast estuaries.        Leopard sharks often enter shallow bays and onto
                        Life Stage                        intertidal flats during high tide, retreating during ebb
            Estuary    A P J M                            tide (Compagno 1984). Unlike many sharks which are
             Puget Sound            Relative abundance:   nocturnal, leopard sharks appear to be active during
              Hood Canal             �   Highly abundant  daylight (Dubsky 1974).
              Skagit Bay                Abundant
                                     O   Common
             Grays Harbor            0 m  Rare             Reproduction
              WillapaBay            Blank Notpresent      Mode: The  leopard  shark  is gonochoristic,
           Columbia River                                 ovoviviparous, and iteroparous. Fertilization is internal
            Nehalem Bay                                   and there is no yolk-sac placenta.
            Tillamook Bay           Life stage:
              Netarlsay              A - Adults
                                     Netar- Parturition   Matina/SDawnina: Mating appears to occur soon after
              Siletz River           J -Juveniles         females give birth, primarily during April and May.
             Yaquina Bay            M- Mating             Mating (as observed in the Steinhart Aquarium in San
              Alsea River                                 Francisco, California) is preceded by the male and
            Siuslaw River                                 female swimming rapidly together andthe male holding
            Umpqua River                                  the female's left pectoral fin in his mouth. By twisting
               Coos Bay i                                 his body under hers, the male is able to insert his left
             Rogue River                                  clasper into the female's cloaca. Hence, coitus occurs
            Klamath River                                 while swimming, with the male retaining the female's
            Humboldt Bay � � 3                            pectoral fin in his mouth the entire time (Ackerman
                Eel River                                  1971). Females give birth from March through August,
             Tomales Bay 0 0 *3                           with an April or May peak (Ackerman 1971, Talent
       Cent SanFran.Bay '   O    0O    * Includes Central San  1973, S. Smith unpubl. manuscr.).
       South San Fran. Bay  � � 0  0     FandSanPablo bays.
           Elkhom Slough C C  Fecundity: Litter size is 4-29 pups (Compagno 1984).
              Morro Bay 0 0 0
         Santa Monica Bay  O 0 0 0                        Growth and Development
           San Pedro Bay  O  O 0 C                        Eaa Size and Embrvonic Develooment: Eggs develop
             Alamitos Bay 4    I                          within the female, but do not receive nourishment from
             Anaheim Bay  q    4                          the female (Jones and Stokes Associates, Inc. 1981).
             Newport Bay  q    I                          Embryonic development is direct and internal. The
              Mission Bay 'j    I                          required developmental period for embryos appears to
           San Diego Bay  4    I                          be 10-12 months (Ackerman 1971).
           Tijuana Estuary
                       A P J M                            Aae and Size of Larvae: There is no larval stage;
                                                          embryonic development is direct and internal.
    disperse in fall and winter in San Francisco Bay during
    months of high freshwater outflows (S. Smith, unpubl.   Juvenile Size Ranae: Young are 18-20 cm long at birth
    manuscr.).                                             (S. Smith unpubl. manuscr.).

    Miarations and Movements: Most adult leopard sharks    Aae and Size of Adults: Females may take 12-14 years
    leave Elkhorn Slough by June, but begin to return by    and be 110-129 cm long before reaching maturity.
    October (Talent 1973); juveniles have their highest    Malesmatureearlierandatsmallersizesthanfemales
    abundance in Elkhorn Slough inthesummer. Tagging    (Ackerman 1971, Compagno 1984). The maximum
    studies in San Francisco Bay showed that most sharks    recorded length is 1.8 m. Growth is apparently slow,
    resided in the Bay from March to September, but   tagged fish grew only 1.4 cm/yr (S. Smith unpubl.
    dispersed both inside and outsidethe Bayfrom October    manuscr.). Calcified rings (useful for aging a fish) are
    through February. One tagged shark was recovered in   laid down in vertebral centra sometime between May
    Elkhorn Slough, 140 km south of San Francisco Bay    and September each year (Smith 1984).
    (S. Smith unpubl. manuscr.). Leopard sharks may
    form large schools mixed with gray or brown    Food and Feeding
    smoothhound sharks (Mustelus californicus and M.    Trophic Mode: Juveniles and adults are carnivorous,
    henlel) (Compagno 1984).  Schools appear to be    feeding primarily on benthic and epibenthiccrustacea.

                                                       79

    Leopard shark continued
    However, large adults also feed on pelagic fishes such    levels of fishes in Morro Bay, California as determined
    as northern anchovy (Engraulismordax) (Russo 1975).    by ultrasonic tagging. M.S. Thesis, Calif. Polytech.
                                                           State Univ., San Luis Obispo, CA, 51 p.
    Food Items: Young, smaller leopard sharks feed heavily
    on crabs (e.g., yellow shore crab, Hemigrapsus    Ebert, D. A. 1986. Observations on the elasmobranch
    oregonensis) and other crustacea. As leopard sharks    assemblage of San Francisco Bay. Calif. Fish Game
    grow (80-130 cm long), echinuroid worms (Urechis    72(4):244-249.
    caupo), fish eggs, and dam siphons become important
    prey. Larger adults (>130 cm in length) feed primarily    Eschmeyer, W. N., W. S. Herald, and H. Hammann.
    on fish (Ackerman 1971, Russo 1975, Talent 1976).    1983. A field guide to Pacific coast fishes of North
    Common prey includeghost shrimp (Callianassaspp.),    America. Houghton Mifflin Co., Boston, MA, 336 p.
    rock crabs (Cancer spp.), octopus (Octopus spp.),
    shiner perch (Cymatogaster aggregata), arrow goby    Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    (Clevelandia ios), Pacific herring (Clupea pallasi),   Observations on fishes associated with kelp beds in
    topsmelt (Atherinops affinis), and northern anchovy    southern California. Calif. Fish Game, Fish Bull. 160,
    (Talent 1973, Russo 1975, Talent 1976).                 144 p.

    Biological Interactions                                 Herald, E. S., and W. E. Ripley. 1951. The relative
    Predation: The leopard shark probably has no major    abundanceofsharks andbatstingrays inSan Francisco
    predators except man.                                   Bay. Calif. Fish Game 37(3):315-329.

    Factors Influencina PoDulations: Recent reductions in   Jones and Stokes Associates, Inc. 1981. Ecological
    shark numbers in San Francisco Bay may be due to   characterization of the central and northern California
    reducedsalinity, warmwater, orover-harvesting (Ebert    coastal region. Volume II, Part 2, Species. U.S. Fish
    1986). Populations may also be adversely affected by    Wildl. Serv., Off. Biol. Serv., and Bureau Land Manag.,
    pollutants (Russo 1975). High pesticideconcentrations    Pacific Outer Contin. Shelf Off., Washington, D.C.,
    in the livers of leopard sharks may relate to its benthic    FWS/OBS-80146.2, various pagination.
    feeding habits and preference for nearshore habitat. A
    large shark die-off of unknown origin occurred in San    Miller, D. J., and R. N. Lea. 1972. Guidetothe coastal
    Francisco Bay in 1967 (Russo and Herald 1968).    marinefishesof California. Calif. Fish Game, Fish Bull.
    However, a connection between pollutant loads and    157, 235 p.
    die-offs has not been established.
                                                           Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    References                                              E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
                                                           of common and scientific names of fishes from the
    Ackerman, L. T. 1971. Contributions to the biology of    United States and Canada. Am. Fish. Soc. Spec. Publ.
    the leopard shark, Triakis semifasciata (Girard) in   No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Elkhorn Slough, Monterey Bay, California. M.A. Thesis,
    Sacramento State Coill., Sacramento, CA, 54 p.          Roedel, P. M., and W. E. Ripley.  1950.  California
                                                           sharks and rays. Calif. Fish Game, Fish Bull. 75:1-85.
    Barry, J. P., and G. M. Cailliet. 1981. The utilization of
    shallow marsh habitats by commercially important    Russo, R. A. 1975. Observations on the food habits of
    fishes in Elkhorn Slough, California. Cal.-Nev. Wildl.   leopard sharks (Triakis semifasciata) and brown
    Trans. 1981:38-47.                                      smoothhounds (Mustelus henlel). Calif. Fish Game
                                                           61 (2):95-103.
    Compagno, L. J. V. 1984. FAO species catalogue.
    Vol. 4.  Sharks of the world.  An annotated and    Russo, R. A., and E. S. Herald. 1968. The 1967 shark
    illustrated catalogue of shark species known to date.   kill in San Francisco Bay. Calif. Fish Game 54(3):215-
    Part 2.  Carcharhiniformes.   FAO  Fish. Synop.    216.
    125(4):433-434
                                                           Smith, S. E. 1984. Timing of vertebral-band deposition
    deWit,L.A. 1975. Change inthespeciescomposition    in tetracycline-injected leopard sharks. Trans. Am.
    of sharks in south San Francisco Bay. Calif. Fish Game    Fish. Soc. 113:308-313.
    61 (2):106-111.
                                                            Smith, S. E., and S. Kato. 1979. The fisheries of San
    Dubsky, P. A. 1974. Movement patterns and activity    Francisco Bay: past, present and future.  In San

                                                        80

                                                                                       Leopard shark continued
    Francisco Bay: the urbanized estuary, p. 445-468.
    Pac. Div. Am. Adv. Sci. and Calif. Acad. Sci., San
    Francisco, CA.

    Talent, L. G. 1973. The seasonal abundance and food
    of elasmobranchs occurring in Elkhorn Slough,
    Monterey Bay, California. M.A. Thesis, Calif. State
    Univ., Fresno, CA, 58 p.

    Talent, L. G. 1976. Food habits of the leopard shark,
    Triakissemifasciata, in Elkhomrn slough, Monterey Bay,
    California. Calif. Fish Game 62(4):286-298.

                                                       81

       Acipenser medirostris
       Adult

             25 cm

    Common Name: green sturgeon                          Indicator of Environmental Stress: Since the green
    Scientific Name: Acipensermedirostris                sturgeon is long-lived, it mayconcentratecontaminants.
    OtherCommonNames:Sakhalinsturgeonorsterlyad   However, no chemical body burden information is
    in USSR (Scott and Crossman 1973)                    presently available.
    Classification (Robins et al. 1980)
    Phylum: Chordata                                     Ecological: This species is not highly abundant in any
    Class: Osteichthyes                                  Pacific coast estuary, and very little is known about its
    Order: Acipenseriformes                              life history (spawning areas, marine distributions,
    Family: Acipenseridae                                migrations, etc.). The green sturgeon is more marine-
                                                          oriented than white sturgeon and spends limited time in
    Value                                                fresh water (except perhaps early juveniles and
    Commercial: The green sturgeon is commercially    spawning adults).
    targeted with white sturgeon (A. transmontanus) in the
    Columbia River estuary, Grays Harbor, and Willapa   Range:
    Bay, Washington. It is not as valuable as the white   Overall: The green sturgeon's overall range is along
    sturgeon because its meat is considered inferior. The   the Pacific coast from Ensenada, Mexico (Moyle 1976)
    green sturgeon is often captured while gillnetting for   to southeast Alaska. It is also found in Asia (north
    salmon (Oncorhynchus spp.) in estuaries. The green    Japan, Korea, and Sakhalin) (Wydoski and Whitney
    sturgeon is rarely captured in the trawl fishery. In   1979).
    Washington, an average of 4.7 and 15.9 t are annually
    landed in Grays Harbor and Willapa Bay, respectively   Within Studv Area:This species occurs in lower reaches
    (G. Kreitman, Washington Department of Fisheries,   of larger rivers. It appears to be the most common
    Battle Ground, WA, pers. comm.).  It is the primary    sturgeon in the Klamath River (Fry 1973, Tuss et al.
    bottomfish landed in Willapa Bay. In 1986, during a 4-   1987) and Willapa Bay (Table 1).
    day commercial sturgeon season in the Columbia
    River estuary, 5,000 green sturgeon were captured (S.   Life Mode
    King, Oregon Department of Fish and Wildlife,   Eggs, juveniles, and adults are alldemersal. Eggs are
    Clackamas, OR, pers. comm.). The green sturgeon is   probably similar to the white sturgeon's, being slightly
    also gillnetted by Native Americans in Grays Harbor   adhesive to substrates after fertilization.  Larvae,
    and the Klamath River, California.                   juveniles, and adults are benthic feeders.

    Recreational: The green sturgeon is incidentally   Habitat
    captured during the white sturgeon sport fishery in   Iype: Green sturgeon larvae have not been positively
    manyestuaries. However,thisspeciesdoesnotappear   identified, but they probably inhabit similar benthic
    to take a hook as readily as the white sturgeon.     freshwaterareas as do white sturgeon larvae (Stevens
                                                          and Miller 1970). Juveniles may occur in shallow water

                                                                                      Green sturgeon continued
                                                          summer and fall. Juvenile emigration through the
     Table 1. Relative abundance of green sturgeon        lower Klamath River may peak in September (CH2M
              in 32 U.S. Pacific coast estuaries.         Hill 1985). Juveniles appear to remain near estuaries
                         Life Stage                       at first, but as they grow, they can become highly
            Estuary    A S J L E                          migratory and move out to nearshore waters. Adults
             Puget Sound i           Relative abundance:  appear to move into estuaries and rivers to feed and
             Hood Canal              �   Highly abundant  spawn (riverine areas) in spring and early summer.
              Skagit Bay              6   Abundant        The green sturgeon seldom migrates far up rivers or
            Grays Harbor C           0   common           estuaries in Oregon or Washington, but may migrate
             Willapa Bay             Blank Not present    extensively upthe Klamath andTrinity Rivers, California.
           Columbia River O                               Some travel long distances in the ocean; fish tagged in
            Nehalem Bay                                   the Sacramento-San Joaquin estuary have been
            Tillamook Bay             Life stage:         collected fromthe Columbia River and in Grays Harbor
             Netarts By                  pawnin adults    1-3 years later (Miller 1972). Adult immigration to the
              Siletz River             J-Juveniles         Klamath River occurs between late February and late
             Yaquina Bay               -  ELarvae         July (CH2M Hill 1985). Adults appearto migrate back
             Alsea River              E - Eggs           to the ocean during summer and fall.
            Siuslaw River /    v
            Umpqua River o0                                Reproduction
               Coos Bay O    O                            Mode: The green sturgeon is gonochoristic, oviparous,
             Rogue River 0    o                           and iteroparous. It is a broadcast spawner; eggs are
            Klamath River 0    0                          fertilized externally.
            Humboldt Bay O    O
               EelRiver o    O                            Matina/SDawnina: Spawning occurs in the Klamath
            Tomales CBay                                  River and perhaps in the lower reaches of other rivers.
       Cent. San Fran. Bay* O    O O   Includes Central San
                                     Francisco, Suisun,   The only known spawning site in the U.S.S.R. is the
       South SanFran.Bay O    0     and San Pablo bays.   Tumnin River (Artyukhin and Andronov 1990). Adults
           Elkhorn Slough                                 spawn in spring and early summer in California, and
              Morro Bay                                   between March and July (with a peak from mid-April to
           SantaeMonica Bay                               mid-June) in the Klamath River (CH2M Hill 1985).
            San Pedro Bay                                 However, three gravid females were captured during
            Anamitos Bay                                  fall in the Columbia River estuary (G. Kreitman,
             Newportim Bay                                 Washington Department of Fisheries, Battle Ground,
             Mission Bay                                  WA, Pers. commun.). Females broadcast spawn near
           San Diego Bay                                  appropriate substrate (believed to range from clean
           Tijuana Estuary                                sand to bedrock) and at relatively fast water flows.
                       A S J L E                          Water depths in spawning areas are probably greater
                                                          than 3 m.

    (Radtke 1966), and probably movetodeeperand more    Fecundity: Fecundity ranges from 60,000 to 140,000
    saline areas as they grow. Adults are euryhaline and    eggs per female (Artyukhin and Andronov 1990).
    reside in subtidal areas.
                                                           Growth and Development
    Substrate: Spawning substrate is probably similar to   Because eggs and larvae have not been described,the
    that preferred by other sturgeon, (i.e., large cobble).    following information is inferred from what is known for
    Adults and juveniles are found primarily on clean sand.    white sturgeon, a very similar species.

    Phvsical/ChemicalCharacteristics:Juvenilesarefound    Eaa Size and Embrvonic Develooment: Eggs are
    in marine, estuarine, and freshwater habitats (Radtke    probably 4 mm in diameter and darkly pigmented
    1966). Adults are primarily marine.                    (Wanget.al. 1985). Embryonicdevelopmentisindirect
                                                           and external. Time to hatching is 196 hours at 12.7�C
    Miarations and Movements: Juveniles are common in   (Artyukhin and Andronov 1990).
    freshwater areas of the San Joaquin Delta, California,
    insummer(Radtke 1966), andalsointhelowerKlamath    Aae and Size of Larvae: Larval development has not
    River (Tuss et al. 1987). Juveniles migrate out to sea    been described, but larvae in the U.S. may be 8 to 19
    before they are two years old and primarily during    mm (Kohlhorst 1976). Larvae in the U.S.S.R. are about

                                                       83

    Green sturgeon continued
    12.3 mm long at hatching (Artyukhin and Andronov    Khoroshko, P. N. 1972. The amount of water in the
    1990).                                                 Volga Basin and its effect on the reproduction of
                                                           sturgeon (Acipenseridae) under conditions of normal
    JuvenileSize Ranae: Minimum juvenilesize isunknown,    and regulated discharge. J. Ichthyol. 12: 608-615.
    but is probably 2.0 cm; maximum juvenile size is
    probably about 1.5 m.                                   Kohlhorst , D. W.  1976. Sturgeon spawning in the
                                                           Sacramento River in 1973, as determined by distribution
    Aae and Size of Adults:Adults can reach a lengthof 2.1    of larvae. Calif. Fish Game 62(1):32-40.
    m and weigh 136 kg (Hart 1973). Very little age data
    exists, but the estimated maximum age for Klamath    Kohlhorst, D. W.  1980. Recent trends in the white
    River green sturgeon is 60 years (CH2M Hill 1985).      sturgeon population in California's Sacramento-San
                                                           Joaquin estuary. Calif. Fish Game 66(4):210-219.
    Food and Feeding
    Trophic Mode: Larvae initially feed on their yolk sac.    Miller,L.W. 1972. Migrationsofsturgeontaggedinthe
    Juveniles and adults are primarily carnivorous benthic    Sacramento-San Joaquin estuary. Calif. Fish Game
    feeders.                                                58(2):102-106.

    Food items: Young feed on benthic invertebrates.    Moyle, P. B. 1976. Inland fishes of California. Univ.
    Adults andlargerjuvenilesfeedon benthicinvertebrates,    Calif. Press, Berkeley, CA, 405 p.
    epibenthic invertebrates, and small fish (Radtke 1966).
                                                           Radtke, L. D. 1966. Distribution of smelt, juvenile
    Biological Interactions                                 sturgeon, and starry flounder in the Sacramento-San
    Predation: Eggs, larvae, and small juveniles are    Joaquin delta with observations on food of sturgeon. In
    probablypreyeduponbynumerousfishspecies. Large    J. L. Turner and D. W. Kelley (compilers), Ecological
    green sturgeon have few known predators except for   studies of the Sacramento-San Joaquin delta, Part II,
    man and some large marine mammals.                      Fishes of the delta.  Calif. Fish Game, Fish Bull.
                                                           136:115-129,.
    Factors Influencino Populations: Riverflow (Khoroshko
    1972, Kohlhorst 1980), watertemperature, and salinity   Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    may  affect survival of larvae and juveniles.    E.A. Lachner, R. N. Lea, and W. B. Scott. 1980. Alist
    Bioaccumulation of polychlorinated biphenyls or other    of common and scientific names of fishes from the
    contaminants may reduce sturgeon survival.  The    United States and Canada. Am. Fish. Soc. Spec. Publ.
    overall number of adult females in the population may    No. 12. Am. Fish. Soc., Bethesda, MD, 174 p.
    be important because they mature late in life and
    probably not all females spawn every year. Very little   Scott, W. B., and E. J. Crossman. 1973. Freshwater
    is known about this species and there is need for more    fishes of Canada. Fish. Res. Board Can., Bull. No. 184,
    research into all aspects of its biology and ecology.   966 p.

    References                                              Stevens, D. E., and L. W. Miller. 1970. Distribution of
                                                           sturgeon larvae in the Sacramento-San Joaquin River
    Artyukhin, E. N., and A. E. Andronov.  1990.  A    system. Calif. Fish Game 56 (2):80-86.
    morphological study of the green sturgeon, Acipenser
    medirostris (Chondrostei, Acipenseridae), from the    Tuss, D., T. Kisanuki, J. Larson, J. Polos, and T.
    Tumnin (Datta) Riverand some aspects of the ecology    Frazer. 1987. Klamath River fisheries investigation
    and zoogeography of the Acipenseridae. J. Ichthyol.    program. Annual Rep. 1986. U.S. Fish Wildl. Serv.,
    30(7):11-22.                                            Arcata, CA, 93 p.

    CH2M  Hill.  1985.  Klamath River basin fisheries    Wang,Y. L., E. P. Binkowski, and S.l. Doroshov. 1985.
    resource plan. U.S. Dept. Inter., various pagination.    Effect of temperature on early development of white
                                                           sturgeon and lake sturgeon, Acipensertransmontanus
    Fry, D. H., Jr. 1973. Anadromous fishes of California.    and A. fulvescens. Env. Biol. Fish. 14 (1) 43-51.
    Calif. Dept. Fish Game, Sacramento, CA, 41 p.
                                                           Wydoski, R. S., and R. R. Whitney. 1979. Inland fishes
    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    of Washington. Univ. Wash. Press, Seattle, WA,
    Board Can., Bull. No. 180, 740 p.                       220 p.

                                                        84

    85

       Acipenser transmontanus
       Adult

               25 cm

    Common name: white sturgeon                           found in the bile of white sturgeon identified their
    Scientific Name: Acipenser transmontanus              exposure to petroleum hydrocarbons from an oil spill
    Other Common Names: Pacific sturgeon, Oregon    (Krahn et al. 1986).
    sturgeon, Columbia sturgeon, Sacramento sturgeon
    Classification (Robins et al. 1980)                   Ecological: Although the white sturgeon is anadromous,
    Phylum: Chordata                                      it is capable of completing its entire life cycle in fresh
    Class: Osteichthyes                                   water. It generally spawns in large rivers and spends
    Order: Acipensiformes                                 time in both marine and fresh water. However, dams
    Family: Acipenseridae                                 have created landlocked populations because the
                                                          species does not normally use fish ladders.
    Value
    Commercial: The white sturgeon is primarily captured    Range
    incidentallywhile gillnetting for salmon (Oncorhynchus    Overall: The white sturgeon's overall range is from
    spp.), but has recently become a target fishery. In the    Ensenada, Mexico (Moyle 1976) to Cook Inlet in
    Columbia River, 199t were landed in 1985. Washington    northwestern Alaska (Wydoski and Whitney 1979).
    State total landings were nearly 46 t in 1985 (G.
    Kreitman, Washington Department of Fisheries, Battle   Within Studv Area: This species is found in most
    Ground, WA, pers. comm.).  Roe is valuable caviar.   estuaries on the Pacific coast from San Francisco Bay,
    Columbia River sturgeon production is second only to   California, north to Grays Harbor, Washington, but is
    the total Soviet Union production. This species is an    rare in Puget Sound and Hood Canal, Washington
    important fish for Native American fishermen in the    (Table 1). It is most common in estuaries of large
    Columbia River and Klamath River, California. Private   rivers.
    aquaculture operations in California are capable of
    producing a 4.5 kg fish in 30 months (Anderson 1988).    Life Mode
                                                           It is principally an anadromous species. Adults,
    Recreational: The white sturgeon is the focus of an    juveniles, and eggs are demersal. Eggs are adhesive
    intense sport fishery in the lower Columbia River;   after fertilization.
    62,400 were landed in this fishery during 1987 (Bohn
    and Mclsaac 1988). Sport fisheries also exist in the    Habitat
    Sacramento-San Joaquin Delta of California, Willapa    Type: Larvae and very young juveniles are riverine.
    Bay, Washington, and other estuaries.                 Older juveniles and adults are found in riverine,
                                                          estuarine, and marine waters. However, the older life
    Indicator of Environmental Stress: River flow may    stages are primarily found in riverine and estuarine
    affect larval dispersal and survival. Because of its long    areas.  Young-of-the-year white sturgeon may be
    life span, the white sturgeon mayconcentrate pollutants    associated with structures such as pile jetties, rocks,
    in its flesh. Metabolites from aromatic hydrocarbons    and submerged logs (McCabe and McConnell 1988).

                                                                                       White sturgeon continued

                                                           The white sturgeon is a euryhaline species, although
      Table 1. Relative abundance of white sturgeon         younger and smaller fish do not osmoregulate as well
               in 32 U.S. Pacific coast estuaries.          as larger, older individuals (McEnroe and Cech 1985).
                        Life Stage                         Eggs, larvae, and small juveniles are found only in
           Estuary    A S J  L E                           freshwater. Olderjuveniles are common in freshwater
            PugetSound  i            Relative abundance:   areas of the Columbia River estuary.
             Hood Canal i             �   Highly abundant
              Skagit Bay               6}  Abundant         Miarations and Movements: Initially after hatching, fry
                                      0   Common
            Grays Harbor 0 C               Rare             are found throughout the water column. Within 5 to 6
             Willapa Bay C    (       Blank Not present     days, fry become negatively phototaxic and primarily
          Columbia River 0    O                            benthic (Conte et al. 1988). General movements for
            Nehalem Bay C    O                              juveniles and adults exist, but no "migration" has been
           Tillamook Bay C    O       Life stage:          established.  Large white sturgeon appear to move
             Netarts Bay                A - Adults
                                       S - Spawning adults  upstream to spawning grounds in late winter and spring
             Silez JRiver              J-Juveniles         and downstream in fall and winter (Miller 1972).
            Yaquina Bay 0    0         1- LLarvae          Movement is probably related to both spawning and
             Alsea River V4    V    :                      feeding conditions (Bajkov 1951). Some individuals
            Siuslaw River  V    V                           move extensively (between California and Oregon or
           Umpqua River 0                                  Washington), but most do not (Stockley 1981). The
              Coos Bay O    O                              creation of dams/impoundments has created isolated
            Rogue River 0    0                             populations. Estuarine residing sturgeon may move
           Klamath River  I 0 oonto intertidal flats to feed during high tide.
           Humboldt Bay     J
               Eel River    V                               Reproduction
            Tomales Bay                                     Mode: The white sturgeon is gonochoristic, oviparous,
       Cent San Fran. Bay   a  O      Includes Central San
                           Cent San FFrancisco. Suisun     and iteroparous. It is a broadcast spawner; eggs are
       South San Fran. Bay  0    0    and San Pablo bays.   fertilized externally.
          Elkhom Slough
              Morro Bay                                     Matina/Soawnina: Spawning occurs during the spring
        Santa Monica Bay                                   in areas with swift currents and large cobble. Peak
          San Pedro Bay                                    spawning in the Sacramento River occurs at 14.40C
            Alamitos Bay                                    (Kohlhorst 1976). In the Columbia River, spawning
            Anaheim Bay                                     apparently occurs at temperatures of 13-200C (end of
            Newport Bay                                    May to early July) below John Day Dam (Palmer et al.
             Mission Bay                                    1988), and 10-16�C below Bonneville Dam (late April
           San Diego Bay                                    to early July) (McCabe and McConnell 1988). Females
          Tijuana Estuary                                  do not spawn annually, but every 3-5 years.  They
                      A S J L E                            broadcastspawnnearappropriatesubstrateandwater
                                                           flow; no nest is built.
    The white sturgeon is not usually found in intertidal
    areas, although it may feed on intertidal flats at high    Fecundity: The white sturgeon is very fecund; a 2.7 m
    tide.  Water flow is important to the downstream    long female in California contained 4.7 million eggs
    movement of larvae. Subyearlings are common during    (Moyle 1976).
    the summer in shallow freshwater areas of the San
    Joaquin Delta in summer (Radtke 1966).  In the    Growth and Development
    Columbia River, small juveniles appear to prefer deep-    Ean Size and Embryonic DeveloDment: White sturgeon
    water channel habitat.                                  eggs are 4.0 mm in diameter, and darkly pigmented
                                                           (Wang et al. 1985). Eggs hatch in approximately seven
    Substrate: Adults and juveniles occur on a wide range    days (depending on temperature) (Conte et al. 1988).
    of sediment types, ranging from sandy-mud and coarse
    sand to cobble. Spawning substrate is large smooth    Aae and Size of Larvae: Captured larvae ranged from
    cobble.                                                 8-19 mm in total length (Kohlhorst 1976), while cultured
                                                           larvae averaged 12.6 mm (Wang et al. 1985). Fry yolk
    Phvsical/Chemical  Characteristics: Best egg    sacs are depleted and active feeding begins
    development and survival is 14-16�C, although    approximately12 daysafterhatching (Anderson 1988).
    incubation is possible from 10-18�0C (Wang et al. 1985).

                                                        87

    White sturgeon continued
    Juvenile Size Ranae: Newly-metamorphosed juveniles    References
    are about 20 cm long. Olderjuveniles may be 1.2 m or
    longer before maturing.                               Anderson, R. S.  1988.  Columbia River sturgeon.
                                                          Wash. Sea Grant, Seattle, WA, 19 p. (WSG-AS 88-
    Aae and Size of Adults: The white sturgeon is a very   14).
    slow-growing, late-maturing fish. Growth and maturity
    are highly variable. In California, females mature at   Bajkov. A. D. 1951.  Migration of white sturgeon
    approximately 11 years and 1.2 m long (Moyle 1976).    (Acipenser transmontanus) in the Columbia River.
    In Oregon, female white sturgeon mature at about 15    Fish Comm. Oreg. Res. Briefs 3(2):8-21.
    years and 1.7 m long (Stockley 1981). Males mature
    earlier and at a shorter length. The life span of white    Bohn, B. R., and D. Mclsaac. 1988. Columbia River
    sturgeon is unknown, but probablyexceeds 1 00years.    fish runs and fisheries 1960-1987. Oreg. Dept. Fish
    Thereare reports of some fish weighing morethan 816    Wildl. and Wash. Dept. Fish., Clackamas, OR, 83 p.
    kg and almost 6 m long (Anderson 1988). White
    sturgeon are North America's largest freshwater fish.   Conte, F. S., S. I. Doroshov, P. B. Lutes, and E. M.
                                                          Strange. 1988. Hatchery manual forthe white sturgeon
    Food and Feeding                                      Acipensertransmontanus Richardson with application
    TrophicMode:Larvaefeedontheiryolksac. Juveniles,   to other North American Acipenseridae.  Publ. No.
    and adults are primarily benthic carnivores.          3322, Coop. Extension, Div. Agricul. Nat. Res., Univ.
                                                          Calif., Oakland, CA, 104 p.
    Food items: Very small juveniles probably feed on
    benthic algae and small invertebrates.  Juveniles    Hung, S. S. O., P. B. Lutes, F. S. Conte, and T.
    consume benthic and epibenthic invertebrates,    Storebakken.  1989. Growth and feed efficiency of
    including amphipods, shrimp, mysids, bivalves, and    white sturgeon (Acipenser transmontanus) sub-
    insect larvae (Radtke 1966).  Larger juveniles and    yearlings at different feeding rates. Aquacult. 80:147-
    adults feed on benthic invertebrates and fish such as    153.
    eulachon ( Thaleichthyspacificus) and northern anchovy
    (Engraulis mordax).  They also feed on clams,    Khoroshko, P. N. 1972. The amount of water in the
    amphipods, Crangonshrimp,ghost shrimp(Callianasa    Volga Basin and its effect on the reproduction of
    spp.), mud shrimp (Upogebia spp.), and other benthic    sturgeon (Acipenseridae) under conditions of normal
    invertebrates (Semakula and Larkin 1968, Muir et al.   and regulated discharge. J. Ichthy. 12:608-615.
    1988). Optimum growth of hatchery juveniles occurs
    whenfedadietconsistingof40%crudeprotein(Moore    Kohlhorst , D. W. 1976. Sturgeon spawning in the
    et al. 1988). The optimal feeding rate for subyearlings    Sacramento River in 1973, asdetermined bydistribution
    at 18�C is between 1.5 and 2.0% of their body weight   of larvae. Calif. Fish. Game 62(1):32-40.
    per day (Hung et al. 1989).
                                                          Krahn, M. M., L. J. Kittle, Jr., and W. D. MacLeod, Jr.
    Biological Interactions                               1986. Evidence for exposure of fish to oil spilled into
    Predation: Eggs, larvae, and small juveniles are   the Columbia River. Mar. Envir. Res. 20:291-298.
    probably preyed upon by numerous fish species. Larger
    juveniles and adult white sturgeon are primarily taken    McCabe, G. T. , Jr., and R. J. McConnell.  1988.
    by man, however, some may be eaten by marine    Appendix D. InA. A. Nigro (editor), Status and habitat
    mammals.                                              requirements of white sturgeon populations in the
                                                          Columbia River downstream from McNary Dam, p.
    Factors Influencina Ponulations: Dams have created    114-139. Annual Prog. Rep., July 1987 - March 1988.
    land-locked populations and destroyed spawning    Bonneville Power Admin., Portland, OR.
    grounds. Bioaccumulation of contaminants such as
    polychlorinated biphenyls may inhibit growth and impair    McEnroe, M., and J. J. Cech, Jr. 1985. Osmoregulation
    egg and larval survival (Parsley et al. 1989).  High    in juvenile and adult white sturgeon, Acipenser
    temperatures (>200C) may reduce larval viability (Wang    transmontanus. Env. Biol. Fish. 14(1):23-30.
    et al. 1985). Overfishing could reduce the adult
    spawning stock, although present regulations prohibit   Miller, L. W. 1972. Migration of sturgeon tagged in the
    taking fish longerthan 6 ft (1.8 mtotal length) in Oregon    Sacramento-San Joaquin estuary. Calif. Fish Game
    and Washington. Reduced river flows may also hinder   58(2):102-106.
    sturgeon production (Khoroshko 1972).

                                                                                        White sturgeon continued

    Moore, B. J., S. S. O. Hung, and J. F. Medrano. 1988.
    Protein requirement of hatchery-produced juvenile white
    sturgeon (Acipensertransmontanus). Aquacult. 71:235-
    245.

    Moyle, P. B. 1976. Inland fishes of California. Univ.
    Calif. Press, Berkeley, CA, 405 p.

    Muir, W. D., R. L. Emmett, R. J. McConnell. 1988. Diet
    of juvenile white sturgeon in the lower Columbia River
    and its estuary. Calif. Fish Game. 74(1):49-54.

    Palmer, D. E., M.J. Parsley, and L. G. Beckman. 1988.
    Appendix C. InA. A. Nigro (editor), Status and habitat
    requirements of white sturgeon populations in the
    Columbia River downstream from McNary Dam, p. 89-
    113. Annual Prog. Rep., July 1987 - March 1988,
    Bonneville Power Admin., Portland, OR.

    Parsley, M. J., S. D. Duke, T. J. Underwood, and L. G.
    Beckman. 1989. Report C. In A. A. Nigro (editor),
    Status and habitat requirements of white sturgeon
    populations in the Columbia River downstream from
    McNary Dam, p. 101-166. Annual Prog. Rep., April
    1988 - March 1989, Bonneville Power Admin., Portland,
    OR.

    Radtke, L. D. 1966. Distribution of smelt, juvenile
    sturgeon, and starry flounder in the Sacramento-San
    Joaquin delta with observations on food of sturgeon. In
    J. L. Turner and D. W. Kelley (compilers), Ecological
    studies of the Sacramento-San Joaquin delta, Part II,
    Fishes of the delta. Calif. Fish Game, Fish. Bull,
    136:115-129.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Semakula, S. N., and P. A. Larkin. 1968. Age, growth,
    food, and yield of white sturgeon (Acipenser
    transmontanus) of the Fraser River, British Columbia.
    J. Fish. Res. Board Can. 25:2589-2602.

    Stockley, C. 1981. Columbia River sturgeon. Prog.
    Rep. No. 150, Wash. Dept. Fish., Olympia, WA, 28 p.

    Wang, Y. L., E. P. Binkowski, and S. I. Doroshov. 1985.
    Effect of temperature on early development of white
    sturgeon and lake sturgeon, Acipensertransmontanus
    and A. fulvescens. Env. Biol. Fish. 14 (1):43-51.

    Wydoski, R. S., and R. R. Whitney. 1979. Inland fishes
    of Washington, Univ. Wash. Press, Seattle, WA, 220 p.

        Alosa sapidissima

                       25 cm
    Common Name: American shad                             sport fishery (such as for salmonids) has not developed.
    Scientific Name: Alosa sapidissima                     The Sacramento River harvest is all recreational (Moyle
    Other Common Names: Atlantic shad, Potomac shad,    1976).
    shad, whiteshad, common shad, North River shad,
    Connecticut River shad, Alose (Scott and Crossman    Indicator of Environmental Stress: This species is very
    1973)                                                  temperature-sensitive and many aspects of its life
    Classification (Robins et al. 1980)                    cycle are cued by specific temperatures.
    Phylum: Chordata
    Class: Osteichthyes                                    Ecological: The introduction of American shad to the
    Order: Clupeiformes                                    Pacific coast does not appear to have displaced native
    Family: Clupeidae                                      species, but competition may occur. Juvenile shad in
                                                           fresh water and estuaries are prey for salmonids and
    Value                                                  many other fish and birds.
    Commercial: The American shad was introduced to the
    Pacific coast in 1871, 1885, and 1886 (Craig and    Range
    Hacker 1940). It has since proliferated and now is   Overall: The American shad is found along the east
    highly abundant in many western rivers and estuaries.    coast of North America from Florida to Newfoundland.
    Average minimum run size for the Columbia River is   It also ranges along the Pacific coast from San Pedro,
    >1.4 million fish/year for the past five years (Bohn and    California, to Cooks Inlet, Alaska, and the Kamchatka
    Mclsaac 1988). In the Sacramento-San Joaquin River,    Peninsula on the Asiatic side of the North Pacific (Scott
    California, run sizes range from 0.7 to 4.0 million fish/   and Crossman 1973).
    year. Commercial fishermen primarily use gill nets for
    this species.  The commercial harvest of shad in   Within Studvy Area:This species is found in all estuaries
    California rivers was terminated in 1957 (Stevens et al.   that have rivers with appropriate spawning habitat, but
    1987) due to conflicts with salmonid (Oncorhynchus    primarilyoccursfrom San Francisco Bay, California, to
    spp.) resources and sport anglers. Large Pacific coast    Puget Sound, Washington (Table 1).
    commercial catches were once common, but only
    small catches presently occur because of poor market    Life Mode: Eggs are semibuoyant and float downstream
    demand and conflicts with the incidental catch of   near the bottom in slow currents. Larvae, juveniles,
    salmonids. In Oregon, it can only be commercially    and adults are nektonic and pelagic.
    caught in the Columbia River. In 1987, 159t (121,000
    fish) were caught in the Columbia River (Bohn and    Habitat
    Mclsaac 1988).                                         Tye: Eggs are demersal. Larvae are pelagic, but are
                                                           found in shallow water, primarily along river bank
    Recreational: The American shad is considered a good    areas. Juveniles and adu Its are also pelagic. Juveniles
    sport fish for light tackle, but an intense Pacific coast    rear in rivers and estuaries before moving offshore.

                                                                                      American shad continued

                                                          while in the ocean (Neves and Depres 1979), and their
     Table 1. Relative abundance of American shad         migration patterns are closely linked with water
              in 32 U.S. Pacific coast estuaries.         temperature. Optimum temperatures for egg survival
                        Life Stage                        are 15.5-26.6�C (Leggett and Whitney 1972). Dissolved
           Estuary    A S J L E                           oxygen (DO) levels above 4.0 mg/I are needed for
           PugetSound  a:ni        Relative abundance:   spawning (Facey and Van Den Avyle 1986) and DO
            HoodCanal B a            �   Highly abundant  levels above 2.5-3.0 mg/I (perhaps 5.0 mg/I) are
             SkagitBay 0    0        O   Common           necessaryforall life stages (Facey and Van Den Avyle
           Grays Harbor     i            Rare             1986, Weiss-Glanz et al. 1986). Spawning occurs in
            Willapa Bay  1   61     Blank Not present     water flows of 30.5 to 91.0 cm/sec.
          Columbia River I* 0 �
           Nehalem Bay  4    i                            Miarations and Movements: Juveniles begin their
           Tillamook Bay   I         Life stage:          downstream migration in late summer and fall when
            Netarts Bay              A -Adults
                                      S - Spawning adults  watertemperature approaches 15.50C. Most juveniles
             Siletz River             J -Juveniles        will migrate out to sea before winter, but some may
            Yaquina Bay               L - Larvae          reside morethan a year in rivers and estuaries (Stevens
            Alsea River O    a                           gset al. 1987). A schooling species, adults return primarily
           Siuslaw River  3                               to their natal river, but there is some straying. Adults
           Umpqua River                                   begin entering estuaries when water temperatures are
              coos Bay S O S O O                          10-15�C, and typically remain there for two or three
            Rogue River <    1                            days before moving upstream (Leggett and O'Boyle
           Klamat River CO                                1976).  Adult upstream migration typically peaks in
           Humboldt Bay    E                              spring when water temperature is near 18.5�C, usually
              Eel River  3                                May to June on the Pacific coast (Leggett and Whitney
           Tomales Bay                                    1972). In theocean, adults appearto migrate vertically,
      Cent San Fran. Bay  � 5        Includes Central San
                                    Francisco. Suisun.    following the diel movements of zooplankton (Neves
      South San Fran. Bay  C        and San Pablo bays.   and Depres 1979). Adults and ocean-dwelling juveniles
          Elkhom Slough                                   may be found down to 340 m depth, but most reside
             Morro Bay                                    within the 50-100 m isobath (Neves and Depres 1979).
        Santa MonicaBSay                                  The American shad is highly migratory; for example,
          San Pedro Bay                                   individuals have been caught 3,000 km from where
           Alamitm Bay                                    they were tagged (Whitehead 1985).
           Anaheim Bay
            Newport Bay                                   Reproduction
            Mission Bay                                   Mode: The American shad is gonochoristic, oviparous,
          San Diego Bay                                   and iteroparous (although many die after spawning). It
          Tijuana Estuary                                 is a broadcast spawner; eggs are fertilized externally.
                      A S J L E
                                                           Matina/Soawnina: This species returnsto its natal river
    Reservoirs appear to be ideal rearing habitat for   tospawn. Spawning usuallyoccursattemperaturesof
    juveniles, therefore, the development of reservoirs on    14-21cC during spring and early summer in the
    the Columbia and other rivers appears to have benefitted    mainstem of rivers. Many shad die soon after spawning,
    this species.                                          with post-spawning survival highest in northern
                                                           estuaries. Spawners prefer shallow water in gently
    Substrate: Larvae, juveniles and adults are not substrate    sloping areas with sand or gravel substrates.  Most
    selective. Spawning occurs over various substrates,    spawning probably occurs during late afternoon and
    but primarily over clean sand and gravel.              evening (Facey and Van Den Avyle 1986). Before
                                                           spawning, males may chase females into a tight circle
    Phvsical/ChemicalCharacteristics:TheAmerican shad    and spawning is often indicated by splashing at the
    is a euryhaline anadromous species. Eggs cantolerate   surface.
    moderate salinities (7.5-15%o), depending on water
    temperatures (Facey and Van Den Avyle 1986).    Fecundity:Spawningfemalesrelease30,000-300,000
    Juveniles rearin both freshwater and estuarine habitats.   eggs, depending on their body size (Moyle 1976). On
    Adults apparently need two or three days in estuaries    the Atlantic coast, American shad fecundity is reported
    to acclimate to fresh water (Weiss-Glanz et al. 1986).    to rangefrom 100,000-600,000 eggs perfemale(Facey
    Adults reside within a temperature range of 3-15�C    and Van Den Avyle 1986).

                                                        91

    American shad continued
    Growth and Development                                Shad year-class strength appears to be determined by
    EaaSizeandEmbrvonicDeveloDment: Egg diameters    river flow and water temperatures during and
    are 2.5-3.8mm afterfertilization (Walburg and Nichols    immediately after spawning (Leggett 1976).  Larval
    1967). Eggs are nonadhesive and slightly heavierthan    survival ultimately determines year-class strength
    water. Eggs need adequate water circulation during    (Crecco and Savoy 1985).  High river flows during
    incubation (Facey and Van Den Avyle 1986). Embryonic    spawning and early life stages positively affect
    development is indirect, and eggs hatch in 4-5 days at   population abundances intheSacramento-SanJoaquin
    15-18�C (depending on temperature).                   river systems (Stevens et al. 1987).  Probably the
                                                           largest factor influencing populations on the Pacific
    Aae and Size of Larvae: Larvae are 7-10 mm long at   coast has been the creation of dams and reservoirs,
    hatching and develop into juveniles in 4-5 weeks at   which has both created and destroyed habitat. Water
    about 25 mm in length (Walburg and Nichols 1967).     irrigation projects can also have an adverse affect on
                                                           shad populations (Stevens et al. 1987) and proper dam
    Juvenile Size Ranae: The minimum size of juveniles is   bypass systems for adults and juveniles are necessary.
    about 2.5 cm. Sexual maturity is reached when this   On the Pacific coast, commercial fishing is presently
    species is about 30-40 cm long.                       limited due to limited markets and the incidental catch
                                                           of depressed salmonid stocks.
    Aae and Size of Adults: Mature shad range from 30-76
    cm total length, with males typically being shorter and    References
    younger than females. Males are usually three years
    old and females four years old when they first mature    Bohn, B. R., and D. Mclsaac. 1988. Columbia River
    (Moyle 1976). Shad may live for seven years (Clemens    fish runs and fisheries 1960-1987. Oregon Dept. Fish
    and Wilby 1961).                                      Wildl. and Wash. Dept. Fish., Clackamas, OR, 83 p.

    Food and Feeding                                      Brodeur, R. D., H. V. Lorz, and W. G. Pearcy. 1987.
    Trophic Mode:  Larvae, juveniles and adults are    Food habits and diet variations of pelagic nekton off
    planktivorous.                                        Oregon and Washington, 1979-1984.  U.S. Dept.
                                                           Commer., NOAA, Tech. Rep. NMFS 57, 32 p.
    Food items: American shad larvae eat small
    zooplankton (copepods and cladocerans) and midge    Clemens, W. A., and G. V. Wilby. 1961. Fishes of the
    larvae and pupae (Facey and Van Den Avyle 1986).    Pacific coast of Canada. Fish. Res. Board Can., Bull.
    Riverine- and estuarine-dwelling juveniles consume    No. 68. 443 p.
    primarily zooplankton, such as copepods, cladocerans
    (Daphnia spp.), amphipods (Corophium spp.), mysids    Craig, J. A., and R. L. Hacker. 1940. The history and
    (Neomysisspp.), and shrimp (Crangonspp.) (Stevens    development of the fisheries of the Columbia River.
    1966, Hammann 1982). Juveniles also eat aquaticand    Fish. Bull., U.S. 32:133-216.
    terrestrial insects. The diet of American shad in Pacific
    coast marine waters is not well-studied, but likely   Crecco, V. A., and T. F. Savoy. 1985. Effects of biotic
    consists of euphausiids, copepods, decapod larvae,   and abiotic factors on growth and relative survival of
    cephalopod larvae, and probably small fishes (Hart   young American shad, Alosa sapidissima, in the
    1973, Brodeur et al. 1987).                           Connecticut River. Can. J. Fish. Aquat. Sci. 42:1640-
                                                           1648.
    Biological Interactions
    Predation: Young shad in rivers and estuaries are    Facey, D. E., and M. J. Van Den Avyle. 1986. Species
    eaten by white sturgeon (Acipenser transmontanus),    profiles: life histories and environmental requirements
    juvenile salmonids, walleye (Sizostedian vitreum), bass    of coastal fishes and invertebrates (South Atlantic)-
    (Micropterus spp.), striped bass (Morone saxatilis),   American shad. U.S. Fish Wildl. Serv. Biol. Rep. 82
    gulls,osprey(Pandion haliaetus), bald eagles (Haliaetus   (11.45). U.S. Army Corps Eng., TR EL-82-4, 18 p.
    leucocephalus), harbor seals (Phoca vitulina), and
    other large predators. After moving offshore, they are    Hammann, M. G. 1982. Utilization of the Columbia
    probably prey for sharks, tuna, porpoises, sea lions,   River estuary by American shad, Alosa sapidissima
    salmonids, and other piscivorous fishes.              (Wilson). M.S. Thesis, Oregon State Univ., Corvallis,
                                                          OR, 48 p.
    Factors Influencina PoDulations: Alteration of
    temperature regimes can affect all life stages (Leggett   Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    and Whitney 1972, Facey and Van Den Avyle 1986).    Board Can., Bull. No. 180, 740 p.

                                                                                       American shad continued
    Leggett, W. C.  1976.  The American shad (Alosa    Rep. 82(11.59). U.S. Army Corps Eng., TR EL-82-4,
    sapidissima), with special reference to its migration    16 p.
    and population dynamics in the Connecticut River. In
    D. Merriman and L. M. Thorpe (editors),  The    Whitehead, P. J. P. 1985. Clupeoidfishesoftheworld.
    Connecticut River ecological study, p. 169-225. Am.    An annotated and illustrated catalogue of herrings,
    Fish. Soc. Monog. No. 1, Am. Fish. Soc., Bethesda,    sardines, pilchards, sprats, shads, anchovies and wolf-
    MD.                                                     herrings, Part 1-Chirocentridae, Clupeidae and
                                                           Pristigasteridae. FAO Fish. Synop. 125(7):1-303.
    Leggett, W. L., and R. N. O'Boyle. 1976. Osmotic
    stress and mortality in adult American shad during
    transfer from saltwater to freshwater. J. Fish Biol.
    8:459-469.

    Leggett, W. C., and R. R. Whitney. 1972. Water
    temperature and the migrations of American shad.
    Fish. Bull., U.S. 79(3):659-670.

    Moyle, P. B. 1976. Inland fishes of California. Univ.
    Calif. Press, Berkeley, CA, 405 p.

    Neves, R. J., and L. Depres. 1979. The oceanic
    migration of American shad, Alosa sapidissima, along
    the Atlantic coast. Fish. Bull., U.S. 77(1):199-212.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No.12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Scott, W. B., and E. J. Crossman. 1973. Freshwater
    fishesof Canada. Fish. Res. Board Can., Bull. No. 184,
    966 p.

    Stevens, D. E. 1966. Distribution and food habits of the
    American shad, Alosa sapidissima, in the Sacramento-
    San Joaquin delta. In J. L. Turner and D. W. Kelley
    (compilers), Ecological studies of the Sacramento-San
    Joaquin delta. Calif. Fish Game, Fish Bull. 136:97-114.

    Stevens, D. E., H. K. Chadwick, and R. E. Painter.
    1987. American shad and striped bass in California's
    Sacramento-San Joaquin River system. Am. Fish.
    Soc. Symp. 1:66-78.

    Walburg, C. H., and P. R. Nichols. 1967. Biology and
    management of the American shad and status of the
    fisheries, Atlantic coast of the United States, 1960.
    U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. No. 550,
    105 p.

    Weiss-Glanz, L. S., J. G. Stanley, and J. R. Moring.
    1986. Species profiles: life histories and environmental
    requirements of coastal fishes and invertebrates (North
    Atlantic)-American shad. U.S. Fish Wildl. Serv. Biol.

                                                        g3

        Clupea pallasi
       Adult

                    5 cm
    Common Name: Pacific herring                            Indicatorof Environmental Stress: Herring larvae appear
    Scientific Name: Clupea pallasi                         to have high mortality rates in oil-contaminated water
    Other Common Names: California herring, Ches-    (Nelson-Smith 1973). The water-soluble fraction of
    Pechora herring, eastern herring, herring, Kara herring,    crude oil reduces larval feeding and growth at low
    Pacific Ocean herring, seld, white sea herring          concentrations and mortalities at high levels (Lassuy
    Classification (Robins et al. 1980)                     1989).   Populations show wide fluctuations in
    Phylum: Chordata                                        abundance, apparently related to environmental
    Class: Osteichthyes                                     conditions (see "Factors Influencing Populations"), and
    Order: Clupeiformes                                     are affected by alterations of bays and estuaries
    Family: Clupeidae                                       (spawning habitats).

    Value                                                   Ecological: Seasonally, C. pallasi is one of the most
    Commercial: The Pacific herring has a long history of   abundant species in Pacificcoast marine and estuarine
    exploitation. It has been sold fresh or salted and also   neritic zones. Juveniles are highly abundant in many
    used for fish meal.  Since 1965, the fishery has    Pacific coast estuaries in summer. They are important
    concentrated on gravid females for roe (eggs), which    prey for many marine species (e.g., Pacific salmon,
    are exported primarily to Japan. Presently, over 90%    seals, and gulls).
    of the Pacific herring caught are in the roe fishery.
    Fishermen take advantage of the Pacific herring's    Range
    natural spawning cycle by fishing in nearshore areas    Overall: The Pacific herring is Arctic-circumboreal. In
    when it spawns. They are primarily caught by purse    the eastern Pacific it ranges from Ensenada, Baja
    seine and gill net. Recent U.S. annual harvests have    California, to St. Michael Island and to Cape Bathurst
    been 52,600 t, worth $47 million (National Marine    in the Beaufort Sea(Hart 1973). It is alsofound inArctic
    Fisheries Service 1986).  The San Francisco and    waters from Coronation Gulf, Canada, to the Chukchi
    Tomales Bay, California, fishery alone is worth $11    Sea and the USSR arctic. In the western Pacific, it is
    million (Suer 1987). Most U.S. harvest comes from    found to Toyama Bay, Japan, west to Korea, and the
    Alaska, California, and Washington. Since spawning    Yellow Sea (Haegele and Schweigert 1985, Wang
    adults are highly vulnerable to overfishing, the fishery    1986).
    is strictly regulated (Grosse and Hay 1989). Commercial
    bait fisheries (which harvest juveniles) exist in Puget    Within StudyArea:This species is found in most Pacific
    Sound, Washington, and other Pacific coast estuaries    coast estuaries north of San Diego, California, but
    (Trumble 1983).                                         occurs primarily north of Point Conception, California
                                                           (Table 1).
    Recreational: The Pacific herring is used as bait for
    Pacific salmon (Oncorhynchusspp.) and other fishes.    Life Mode
    However, some are caught for human consumption.         Eggs are adhesive after fertilization and attach to

                                                                                        Pacific herring continued
                                                            temperatures of 5-14�C and salinities of 3-33%o
     Table 1. Relative abundance of Pacific herring         (Haegele and Schweigert 1985). Larvae aretolerantof
               in 32 U.S. Pacific coast estuaries.          salinities ranging from 2-28o/0  (Alderdice and Velsen
                          Life Stage                        1971, Alderdice and Hourston 1985). Best spawning
             Estuary    A S J  L E                          salinities in British Columbia are 27.0-28.7%o. (Alderdice
              Puget Sound � � � S    �   Relative abundan   and Hourston 1985).  Optimum temperatures and
               Hood Canal    �)  � * -    �   Highly abun4  salinities for egg and larval survival appear to be 5.5-
                SkagitBay * a i a g    �   Abundant          8.7�C and 13-19%o (Alderdice and Velsen 1971).
                                         '4  Rare
              Grays Harbor   : o         i   Rae             However, spawning temperatures in California are
               Willapa Bay O O a 0 0   Blank Notpresent     normallyabove9�C (Barnhart 1988). Salinitytolerances
            Columbia River O O � O O                        of larvae are affected by temperature and salinity
             Nehalem Bay      xi �   i R   during egg incubation (Alderdice and Hourston 1985).
             Tillamook Bay Li � Li � �    Life stage:       Turbidity in estuaries may increase larval survival
               Netarts Bay 13�  O O      S - spawning adul  (Boehlert and Morgan 1985).
               Siletz River  O           J -   Juveniles
              Yaqulna Bay    i �    g *  L- LaE sae         Miarations and Movements: The Pacific herring does
               AlseaRiver O O O O O                         not make extensive coastal migrations, but moves
             Siuslaw River O O O O 0                        onshore and offshore in schools as it spawns and feeds
             UmpquaRiver � L    � S  �                      (Morrow 1980). Adults typically move onshore during
                Coos Bay �  �  �                            winter and early spring, residing in "holding" areas
              Rogue River    0                              before moving to adjacent spawning grounds.  The
             Klamath River  i 0                             Pacific herring population consists of many discrete
             Humboldt Bay *     g g S S                     stocks (Grosse and Hay 1989).  However, offshore
                 Eel River O  (O 1i �                       distributions of adults for many Pacific coast stocks are
              TomalesBay � � �     �*                        unknown (Barnhart 1988).  Pacific herring return to
        Cent. San Fran. Bay' * L       Includes Central San
                                       Francisoo. Suisun,   natal spawning grounds to spawn. Larvae are easily
        South San Fran. Bay  * *  0  i      andSanPablobays.  dispersed by currents, but their behavior and local
            Elkhorn Slough L L L L                          currents often retain them in specific areas. Juveniles
                Morro Bay 0    0                             usually stay in nearshore shallow-water areas until fall
          SantaMonicaBay  Ad   -1                           when they disperse to deeper offshore waters.
            San PedroBay  ,4  ax                            However, they may reside year-round in some estuaries
              Alamitos Bay                                   (San Francisco Bay) (Wang 1986). Adult Pacific herring
             Anaheim Bay
                                                            are found down to 100-150 m, with vertical distribution
              Newport Bay                                   apparently controlled bytemperature (Grosse and Hay
               Mission Bay
               Mission Bay                                   1989). Larvae, juveniles, and adults move toward the
            TSiuanaeEstBary                                 surface to feed at dawn and dusk (Grosse and Hay
            Tijuana Estuary                                 1989).
                                                            1989).
                        A S J L E
                                                            Reproduction
    benthic substrates. Larvae, juveniles, and adults are    Mode: This species is gonochoristic, oviparous, and
    pelagic, schooling nekton.                               iteroparous; eggs are fertilized externally. It spawns
                                                            annually after reaching maturity.
    Habitat
    Type: Eggs are laid in intertidal (3.7 m above mean    Matina/Soawnin: Spawning occurs from Novemberin
    lower low water) and subtidal areas (to 20 m depth), but    the southern part of its range to August in the far north.
    normally occur in +1 to -2 m depth.  Larvae and    SpawningpeaksinDecemberandJanuaryinCalifornia
    juveniles are neritic and adults are neritic-oceanic    (Spratt 1981) and February and March in Puget Sound
    (Eldridge and Kaill 1973, Suer 1987).                    (Trumble 1983). Herring spawn in the same areas
                                                            every year. These areas are high-energy areas, located
    Substrate: Eggs are found on eelg rass (Zostera spp.),    in protected coastal habitats or bays and estuaries, and
    algae, tubeworms, Pacificoysters(Crassostreagigas),    are usually influenced by fresh water.  Spawning
    hydroids, driftwood, pilings, brush, rocks, and rocky-    apparently does not occur until a tactile stimulus (e.g.,
    sandy bottoms (Garrison and Miller 1982). Larvae,    a storm, contact with bottom orotherfish) causes some
    juveniles, and adults occurthroughoutthe watercolumn.    males to extrude milt, which in turn stimulates the entire
                                                            school to spawn. During spawning both sexes come in
    Physical/Chemical Characteristics: Eggs can tolerate    contact with the spawning substrate (Haegele and

                                                         95

    Pacific herring continued
    Schweigert 1985).  Most spawning occurs at night    amphipods, chaetognaths, and various fishes.
    (Eldridge and Kaill 1973, Suer 1987).                 Juveniles and adults are consumed by squid, sharks,
                                                          salmonids, gadids, sculpins (Cottus spp.), lingcod
    Fecundity: Fecundity increases with female size and    (Ophiodon elongatus), sand sole (Psettichthys
    ranges from 4,000-134,000 eggs per female (Hart   melanostictus), and other fishes. They are also eaten
    1973). Fecundity is 227 and 220 eggs/gram of female    by many species of birds and marine mammals, such
    weight in Tomales Bay and San Francisco Bay,    as seals and sperm whales (Physetercatodon) (Hart
    respectively(Hardwick 1973, Rabin and Barnhart 1977).    1973, Simenstad et al. 1979, Grosse and Hay 1989).
    Size-specific fecundity is inversely related to latitude
    (Hay 1985).                                           Factors Influencina PoPulations: No relation exists
                                                          between numberof eggs spawned and adult population
    Growth and Development                                size (Pacific Fishery Management Council 1981). Egg
    Eaa Size and Embryonic DeveloDment: Unfertilized    and larval mortalities are thought to be the major
    eggs are 1.0 mm in diameter (Outram 1955); 1.2-1.5   events influencing population sizes. Eggs and larvae
    mmindiameterafterfertilization (Hart 1973). Hatching    suffer natural mortalities due to tidal fluctuations,
    occurs in 11-12 days at 10.7�C, 14-15 days at 8.5�C,   desiccation, freezing, low oxygen, wave action, and
    and 28-40 days at 4.4�C (Outram 1955). Most eggs    predation.  Approximately 98-99% of all larvae are
    hatch at night (Alderdice and Velsen 1971).           killed by predation, competition, andoffshoretransport.
                                                           In general, a clupeoid year-class' strength appears to
    Aae and Size of Larvae: Larvae range from 5 mm to   be determined within the first 6 months (Smith 1985).
    about 26 mm total length (TL).  Metamorphosis to   Other studies indicate that onshoretransport, density-
    juvenile begins at about 26 mm TL and is completed by   dependent mechanisms, upwelling, sea temperatures,
    35 mm  TL  (Fraser 1922, Stevenson  1962);    predation, climate fluctuations, initial feeding period of
    metamorphosis takes about 2 to 3 months (Hay 1985).   larvae, and larvaldispersal patterns may all be important
                                                          in determining population abundances (Lasker 1985,
    Juvenile Size Ranae: Juveniles are 35-150 mm TL,    Grosse and Hay 1989).  Juveniles and adults are
    depending on region. Growth of juveniles is dependent    affected by competition, predation, disease, spawning
    on population size and environmental conditions (Reilly   stress, and fishing. Human and natural alterations of
    1988).                                                water quality, prey species, migration rates, spawning
                                                          substrate and habitat can also impact populations
    Aae and Size of Adults: Adult lengths are from 13-26    (Alaska Department of Fish and Game 1985).
    cm TL, depending on region. The Pacific herring
    matures in 2 to 3 years in California and 3 to 4 years in   References
    Washington. It lives up to 19 years and grows to a
    maximum length of 38 cm TL (Hart 1973). Northern    Alaska Department of Fish and Game. 1985. Alaska
    stocks live longer than southern stocks (Wang 1986,    habitat management guide. South central region, Vol.
    Grosse and Hay 1989).                                  1: life histories and habitat requirements of fish and
                                                          wildlife. Alaska Dept. Fish Game, Juneau, AK, 429 p.
    Food and Feeding
    Trophic Mode: Larvae, juveniles, and adults are    Alderdice, D. F., and A. S. Hourston. 1985. Factors
    selective pelagic plankton feeders, although filter   influencing development and survival of Pacific herring
    feeding has been observed.                             (Clupea harenguspallast) eggs and larvaeto beginning
                                                          of exogenous feeding. Can. J. Fish. Aquat. Sci. 42
    Food Items: Larvae consume diatoms, tintinnids,   (Suppl. 1):56-68.
    invertebrate and fish eggs, crustacean larvae, mollusc
    larvae, and copepods.  Juveniles eat primarily   Alderdice, D. F., and F. P. J. Velsen. 1971. Some
    crustaceans (copepods, cladocerans, euphausiids,    effects of salinity and temperature on earlydevelopment
    mysids, amphipods, and decapod larvae). They also   of Pacific herring (Clupeapallasi). J. Fish. Res. Board
    consume mollusc and fish larvae. Adults eat planktonic    Can. 28(10):1545-1562.
    crustaceans (copepods, euphausiids, and amphipods)
    and fish larvae (Hart 1973, Simenstad et al. 1979,    Barnhart, R. A. 1988. Species profiles: life histories
    Miller et al. 1980, McCabe et al. 1983).              and environmental requirements of coastal fishes and
                                                          invertebrates (Pacific Southwest) - Pacific herring.
    Biological Interactions                               U.S. Fish Wildl. Serv. Biol. Rep. 82(11.79). U.S. Army
    Predation: Eggsare eaten by manyfish species, ducks,    Corps Eng., TR EL-82-4, 14 p.
    and gulls, while larvae are prey forctenophores, jellyfish,

                                                                                        Pacific herring continued
    Boehlert, G. W., and J. B. Morgan. 1985. Turbidity    macroinvertebrate assemblages along the Strait of
    enhances feeding abilities of larval Pacific herring,    Juan de Fuca including food habits of the common
    Clupeaharenguspallasi. Hydrobiol. 123(2):161-170.    nearshore fish.  Interagency (NOAA, EPA) Energy/
                                                           Environ. Res. Dev. Prog. Rep., EPA-600/7-80-027,
    Eldridge, M. B., and W. M. Kaill. 1973. San Francisco    Washington, D.C., 211 p.
    Bay area's herring resource - a colorful past and a
    controversial future. Mar. Fish. Rev. 25:25-31.         Morrow, J. E. 1980. The freshwater fishes of Alaska.
                                                           Alaska Northw. Publ. Co., Anchorage, AK, 248 p.
    Fraser, C. M. 1922. The Pacific herring. Biol. Board
    Can., Contrib. Can. Biol. Fish. 1921 (6):103-111.       National Marine Fisheries Service. 1986. Fisheries of
                                                           the United States, 1985. Current Fishery Statistics No.
    Garrison, K. J. and B. S. Miller. 1982. Review of the    8368. U.S. Dept. Comm., NOAA, Nat. Mar. Fish Serv.,
    early life history of Puget Sound fishes. Fish. Res. Inst.,   Nat. Fish. Stat. Prog., Washington, D.C., 122 p.
    Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).
                                                           Nelson-Smith, A. 1973. Oil pollution and marine
    Grosse, D. J., and D. E. Hay. 1989. Pacific herring,    ecology. Plenum Press, New York, NY, 260 p.
    Clupea harengus pallasi, in the Northeast Pacific and
    Bering Sea. In N. J. Wilimovsky, L. S. Incze, and S. J.   Outram, D. N. 1955. The development of the Pacific
    Westerheim (editors), Species synopses, life histories   herring egg and its use in estimating age of spawn.
    of selected fish and shellfish of the Northeast Pacific   Fish. Res. Board Can., Pac. Biol. Sta. Circ. 40, 9 p.
    and Bering Sea, p. 34-54. Wash. Sea Grant Prog. and
    Fish. Res. Inst., Univ. Wash., Seattle, WA.             Pacific Fishery Management Council. 1981. Pacific
                                                           herring fishery management plan. Pac. Fish. Manag.
    Haegele, C.W.,and J. F. Schweigert. 1985. Distribution    Council, Portland, OR, 127 p.
    and characteristics of herring spawning grounds and
    description of spawning behavior. Can.J. Fish. Aquat.   Rabin, D. J., and R. A. Barnhart. 1977. Fecundity of
    Sci. 42(Suppl. 1):39-55.                                Pacific herring, Clupea harenguspallasi, in Humboldt
                                                           Bay. Calif. Fish Game 63(3):193-196.
    Hardwick, J. E. 1973. Biomass estimates of spawning
    herring, Clupea harengus pallasi, herring eggs, and    Reilly, P. N. 1988. Growth of young-of-the-year and
    associated vegetation in Tomales Bay.  Calif. Fish    juvenile Pacific herring from San Francisco Bay,
    Game 59(1):36-61.                                       California. Calif. Fish Game 74(1):38-48.

    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Board Can., Bull. No. 180, 740 p.                       E. A. Lachner, Robert N. Lea, and W. B. Scott. 1980.
                                                           A list of common and scientific names of fishes from the
    Hay, D. E.  1985.  Reproductive biology of Pacific    United States and Canada. Am. Fish. Soc. Spec. Publ.
    herring(Clupea harenguspallasi). Can. J. Fish. Aquat.    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Sci. 42(Suppl. 1):111-126.
                                                           Simenstad, C. A., B. S. Miller, C. F. Nyblad, K.
    Lasker, R. 1985. What limits clupeoid production?    Thornburgh, and L. J. Bledsoe.  1979.  Food web
    Can. J. Fish. Aquat. Sci. 42(Suppl. 1):31-38.           relationships of northern Puget Sound and the Strait of
                                                           Juan de Fuca. Interagency (NOAA/EPA) Energy/
    Lassuy, D. R. 1989. Species profiles: life histories and    Environ. Res. Dev. Prog. Rep., EPA-600/7-79-259,
    environmental requirements of coastal fishes and    Washington, D.C., 335 p.
    invertebrates (Pacific Northwest)-Pacific herring. U.S.
    Fish Wildl. Serv. Biol. Rep. 82(11.126), U.S. Army    Smith, P. E. 1985. Year-class strength and survival of
    Corps Eng., TR EL-82-4. 18 p.                           0-group clupeoids. Can. J. Fish. Aquat. Sci. 42(Suppl.
                                                            1):69-82.
    McCabe, G. T. Jr., W. D. Muir, R. L. Emmett, and J. T.
    Durkin.  1983.  Interrelationships between juvenile    Spratt, J. D. 1981. Statusofthe Pacific herring, Clupea
    salmonids and nonsalmonid fish in the Columbia River    harengus pallasi, in California to 1980.  Calif. Fish
    estuary. Fish. Bull., U.S. 81(4):815-826.               Game, Fish Bull. 171:1-104.

    Miller, B. S., C. A. Simenstad, J. N. Cross, K. L. Fresh,    Stevenson, J. C. 1962. Distribution and survival of
    and S. N. Steinfort.  1980.  Nearshore fish and    herring larvae (ClupeapallasiValenciennes) in British

                                                        97

    Pacific herring continued
    Columbia waters. J. Fish. Res. Board Can. 19(5):735-
    810.

    Suer, A. L. 1987. The herring of San Francisco and
    Tomales Bays. The Ocean Res. Inst., San Francisco,
    CA, 64 p.

    Trumble, R. J. 1983. Management plan for baitfish
    species in Washington State. Prog. Rep. No. 195,
    Wash. Dept. Fish., Olympia, WA, 106 p.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: A
    guide to the early life histories. Tech. Rep. 9.
    Interagency ecological study program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    and U.S. Fish Wildl. Serv., various pagination.

                                                      98

    99

       Anchoa compressa
       Adult

            2cm

    Common Name: deepbody anchovy                         Life Mode
    Scientific Name: Anchoa compressa                     Eggs and larvae are planktonic, while juveniles and
    Other Common Names: Californiadeepbodyanchovy,    adults are pelagic.
    sprat, deep-bodied anchovy, sardinus (Walford 1931,
    Gates and Frey 1974)                                  Habitat
    Classification (Robins et al. 1980)                   Type: All life stages live primarily in estuaries, bays,
    Phylum: Chordata                                      and lagoons, but schools of juveniles and adults are
    Class: Osteichthyes                                   occasionally found along coastal shorelines (Miller and
    Order: Clupeiformes                                   Lea 1972).
    Family: Engraulidae
                                                          Substrate: Because this is a pelagic species, all life
    Value                                                 stages are found over various substrates.
    Commercial: The deepbody anchovy is of little
    commercial value.                                      Phvsical/Chemical Characteristics: Population
                                                          abundances of this species were significantly correlated
    Recreational: This species is occasionally used as live   with temperature and dissolved oxygen (Allen 1982,
    bait for other fishes (Roedel 1953).                  Horn and Allen 1985). However, thermal and salinity
                                                          tolerances have not been identified.
    Indicator of Environmental Stress: The deepbody
    anchovy uses estuaries during all life stages and may    Miarations and Movements: Adults move from the
    be a good indicator of environmental stress. However,    lower portions of bays and estuaries to upper portions
    little ecological research has been done forthis species.    during the spawning season (spring and summer).
                                                          Adults show post-spawning movements away from
    Ecological: This is an abundant pelagic fish in many    spawning areas, while juveniles reside in the upper
    southern California estuaries (Klingbeil et al. 1975,    portions of bays until late fall and winter (Heath 1980).
    Heath 1980, Horn and Allen 1985).
                                                          Reproduction
    Range                                                 Mode: The deepbody anchovy is gonochoristic,
    Overall: The deepbody anchovy's overall range is from    oviparous, and iteroparous. It is a broadcast spawner;
    Todos Santos Bay, Baja California, to Morro Bay,    eggs are fertilized externally.
    California (Miller and Lea 1972, Eschmeyer et al.
    1983).                                                Matina/SDawnina: Spawning occurs from March to
                                                          August, with most spawning activity occurring at night
    WithinStudvArea:ltismostcommoninCalifomiabays    from April to June (McGowan 1977, Heath 1980,
    and estuaries south of Alamitos Bay (Table 1) (Horn    Edmands 1983). The upper reaches of bays and
    and Allen 1976).                                       estuaries are the usual spawning areas (Heath 1980,

                                                      100

                                                                                    Deepbody anchovy continued

                                                           juvenile characteristics (Caddell 1988), probably in
    Table 1. Relative abundance of deepbody anchovy        about 30 days (Heath 1980).
              in 32 U.S. Pacific coast estuaries.
                        Life Stage                         Juvenile Size Ranae: Juveniles grow from 20-25 mm to
           Estuary    A S J L E                            approximately7ommstandardlength(minimum)before
            Puget Sound              Relative abundance:   reaching maturity.
             Hood Canal    :  :            Highly abundant
             SkagitBay            V       Abundant         Aae and Size of Adults: This species may live to 6
                                      0   Common
           Grays Harbor       :       '   Rare             years, but most die before 5 years.  One-year-olds
             Willapa BaY             Blank Not present      range from 70 mm to about 90 mm in length (Heath
          Columbia River                                   1980). The largest reported deepbody anchovy was
           Nehalem Bay                                     165 mm (Miller and Lea 1972).
           Tillamook Bay              Life stage:
             Netarts Bay               A - Adults
                                        S - Spawning adults  Food and Feeding
             Sllez River               J - Juveniles       Trophic Mode: All feeding life stages are planktivorous.
            Yaquina Bay                L - Larvae
                                        E - Eggs
             Alsea River                                   Food Items: Larvae probably feed on phytoplankton
           Siuslaw River                                   and small zooplankton. Primary prey for juveniles and
           Umpqua River                                    adults are small crustaceans. Major prey taxa include
              Coos Bay                                     calanoid, harpacticoid, and cyclopoid copepods,
            Rogue River                                    ostracods, cumaceans, amphipods, and Callianassa
           Klamamt River                                   spp. larvae.  Minor taxa eaten are polychaetes,
           Humboldt Bay                                    oligochaetes, small gastropods, mysids, tanaidaceans,
               Eel River                                   isopods, crab zoea, dipterans, small gobiids, and plant
            Tomales Bay                                    material (Klingbeil et al. 1975, Horn and Allen 1985).
      Cent San Fran. Bay           * Indcludes Central San
                                     Francisco, Suisun,    This species utilizes the entire water column when
      South San Fran. Bay           and San Pablo bays.    searching for prey (Klingbeil et al. 1975).
          Elkhorn Slough
              Morro Bay                                     Biological Interactions
        Santa Monica Bay  t                                           I      c
                       San t a M onica Bay o  cPredation: The deepbody anchovy is probably eaten
            Alamitos Bay 0 0 0 0                           by many species of birds and piscivorous fishes.
            AlamitosBay      )b
            Anaheim Bay     I * 0 0C                        Factors Influencina PoDulations: The abundance of
            Newport Bay O 0  is   eggs and larvae (and probably juveniles and adults) of
             MSasion Bay CD 0o    C O                       this species appears to cycle widely. The dominant
           San DiesoBay    0 0 3 00                         Anchoa species in southern California estuaries
          Tijuana Estuary
                                                           appears to fluctuate year to year.  Some years A.
                                                            compressa may dominate in ichthyoplankton surveys,
                                                            while in otheryears A. delicatissima prevails. Reasons
    Edmands 1983). This species reduces competition    forthesewidefluctuations areunknown (Heath 1980).
    with the slough anchovy (A. delicatissima) by spawning    Since all life stages reside in estuaries, any estuarine
    in different areas of bays (Edmands 1983).              modifications or pollution directly affects this species.

    Fecundity: Average fecundity is about 15,000 eggs per    References
    female (Heath 1980). Fecundity is significantly related
    to size (1,268 eggs/g female weight) (Heath 1980).    Allen, L. G. 1982. Seasonal abundance, composition,
    Large females may lay over 28,000 eggs (Heath 1980).   and productivity of the littoral fish assemblage in upper
                                                            Newport Bay. Fish. Bull., U.S. 80(4):769-790.
    Growth and Development
    Eaa Size and Embrvonic Develooment: Eggs are    Caddell, S. M. 1988. Early life history descriptions of
    spherical and 0.8 mm in diameter (White 1977, Caddell    the deepbody and slough anchovies with comparisons
    1988). Embryonicdevelopmentisindirectandexternal.   to the northern anchovy (family Engraulidae).  Bull.
    Time to hatching is probably less than 4 days.          Mar. Sci. 42(2):273-291.

    Aoe and Size of Larvae: Larvae are 1.5-2.5mm long at    Edmands, F. A., 11. 1983. The diel distribution and
    hatching and growto about 20-25mm before taking on    transport of ichthyoplankton collected by stationary

                                                        101

    Deepbody anchovy continued
    nets in Newport Bay, Calif., July 1979. M.A. Thesis,    and game fishes of California. Calif. Fish Game, Fish
    Calif. State Univ., Fullerton, CA, 112 p.                Bull. 28,183 p.

    Eschmeyer, W. N., W. S. Herald, and H. Hammann.    White, W. S.   1977.   Taxonomic composition,
    1983.  A field guide to Pacific coast fishes of North    abundance, distribution and seasonality of fish eggs
    America. Houghton Mifflin Co., Boston, MA, 336 p.        and larvae in Newport Bay, California. M.A. Thesis.
                                                           Calif. State Univ., Fullerton, CA, 107 p.
    Gates, D. E., and H. W. Frey. 1974. Designated
    common names of certain marine organisms of
    California. California Fish Game, Fish Bull. 161:55-88

    Heath, K. L. 1980. Comparative life histories of two
    species of anchovies, Anchoa delicatissima and A.
    compressa (F. Engraulidae) from Newport Bay,
    California. M.A. Thesis, Calif. State Univ., Fullerton,
    CA, 71 p.

    Horn, M. H., and L. G. Allen. 1976. Numbers of species
    and faunal resemblance of marine fishes in California
    bays and estuaries. Bull. South. Calif. Acad. Sci.
    75(2):159-170.

    Horn, M. H., and L. G. Allen. 1985. Fish community
    ecology in southern California bays and estuaries.
    Chapter 8. In A. Yanez-Arancibia (editor), Fish
    community ecology in estuaries and coastal lagoons:
    towards an ecosystem integration, p. 169-190. DR (R)
    UNAM Press, Mexico.

    Klingbeil, R. A., R. D. Sandell, and A. W. Wells. 1975.
    An annotated checklist of the elasmobranchs and
    teleosts of Anaheim Bay. In E. D. Lane and C. W. Hill
    (editors), The marine resources of Anaheim Bay. Calif.
    Fish Game, Fish Bull. 165: 79-90.

    McGowan, G. E. 1977. Ichthyoplankton populations in
    south San Diego Bay and related effects of an electricity
    generating station. M.S. Thesis, San Diego State
    Univ., San Diego, CA, 157 p.

    Miller, D. J., and R. N. Lea. 1972. Guidetothe coastal
    marine fishesof California. Calif. Fish Game, Fish Bull.
    No. 157, 235 p.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Roedel, P. M. 1953. Common ocean fishes of the
    California coast. Calif. Fish Game, Fish Bull. 91,
    184 p.

    Walford, L.A. 1931. Handbookof common commercial

                                                       102

    103

       Anchoa delicatissima
       Adult

             2cm

    Common Name: slough anchovy                           Life Mode
    Scientific Name: Anchoa delicatissima                 Eggs and larvae are planktonic, while juveniles and
    Other Common Names: southern anchovy (Gates    adults are pelagic.
    and Frey 1974)
    Classification (Robins et al. 1980)                   Habitat
    Phylum: Chordata                                      jype: All life stages reside primarily in estuaries, bays,
    Class: Osteichthyes                                   and lagoons.  Juveniles and adults are found
    Order: Clupeiformes                                   occasionally in neritic environments (Miller and Lea
    Family: Engraulidae                                   1972, Heath 1980).

    Value                                                 Substrate: All life stages are pelagic and thus found
    Commercial: The slough anchovy is not of commercial    over various substrates.
    value.
                                                          PhvsicaVChemical Characteristics: The slough anchovy
    Recreational: It is occasionally used as live bait for   will avoid temperatures >25�C (San Diego Gas and
    other fishes (Roedel 1953).                           Electric 1980).  Salinity tolerance and tolerance to
                                                          other physical factors have not been identified. The
    Indicator of Environmental Stress: Since this species   estuaries, bays, and lagoons inhabited by this species
    uses estuaries during all life stages it may be an good    are primarily euhaline with salinities rarely <25%oo,
    indicator of environmental stress, however, little   except during the winter rainy period.
    ecological research has been done for this species.
                                                          Miarations and Movements: During spring and early
    Ecological: The slough anchovy is a highly abundant    summer, adults move to spawning areas and then
    pelagic fish in many southern California estuaries    show post-spawning movements to other bay areas
    (Allen and Horn 1975, Heath 1980, San Diego Gas and    (Heath 1980). Schools are sometimes found along the
    Electric 1980, Horn and Allen 1985).                  coast (Eschmeyer et al. 1983, Love et al. 1986).
                                                          Larvae undertake nocturnal vertical migrations
    Range                                                 (Edmands 1983).
    Overall: This species' overall range is from southern
    Baja Californiato Long Beach Harbor, California (Miller   Reproduction
    and Lea 1972, Eschmeyer et al. 1983).                 Mode:The slough anchovy is gonochoristic, oviparous,
                                                          and iteroparous. It is a broadcast spawner; eggs are
    Within Studv Area: It is found in all estuaries and   fertilized externally.
    lagoons from Alamitos Bay, California, south through
    Tijuana Estuary (Table 1) (Horn and Allen 1976).      Matina/SDawnina: Spawning occurs from May to
                                                          September, with most spawning probably occurring in

                                                      104

                                                                                     Slough anchovycontinued

                                                         but probably less than 4 days.
     Table 1. Relative abundance of slough anchovy
              in 32 U.S. Pacific coast estuaries.        Aae and Size of Larvae: Larvae are approximately 1.5-
                        Life Stage                        2.5 mm long at hatching (White 1977, Caddell 1988).
           Estuary    A S J L E                           Upperlength limitof larval stage has not been identified,
            PugetSound              Relative abundance:   but is probably about 20-25 mm.  Metamorphosis to
             Hood Canal              �   Highly abundant  juvenile probably begins after about 30 days (Heath
             Skagit Bay                  Abundant        1980).
                                     O   Common
           Grays Harbor                  Rare
             Willapa Bay            Blank Not present     Juvenile Size Ranoe: Juveniles range from about 25 to
          Columbia River                                  50 mm in length.
           Nehalem Bay
           Tillamook Bay             Life stage:          Aae and Size of Adults: The slough anchovy matures
             Netarts Bay                Saningadults      in one year at a minimum length of about 50 mm
                                      S - Spawning adults
             Siletz River             J - Juveniles       (standard length). Maximum age appears to be 3 years
            Yaqulna Bay               L-Larvae            (Heath 1980), with maximum length about 94 mm
             Alsea River              E-gg                (Miller and Lea 1972). Females tend to grow larger
           Siuslaw, River                                than males (Heath 1980).
           Umpqua River
              Coos Bay                                    Food and Feeding
            Rogue River                                  Trophic Mode: Larvae, juveniles, and adults are
           Klamath River                                  planktivorous.
           Humboldt Bay
               Eel River                                  Food Iems: Calanoid copepods appearto be the major
            Tomales Bay                                   prey for juveniles and adults. Otherprey items include
       Cent. San Fran. Bay'          ncludes Central s    plant material, polychaetes, oligochaetes, gammarid
       South San Fran. Bay          and San Pablobays.    amphipods, harpacticoid and cyclopoid copepods,
          Elkhorn Slough                                  cumaceans, ostracods, and cladocerans (Horn and
              Morro Bay                                   Allen 1985).
        Santa Monica Bay
          SanPedro Bay  ~i~ 70 i 7Biological Interactions
            Alamitos Bay *  *t  tu h                      Predation: The slough anchovy is probably preyed on
            Anaheim Bay                                   by many piscivorous birds and fishes.
            NewportBay ( @ (3 * �
             Mission Bay ( t    S *I 3                    Factors Influencina PoDulations: This species is often
           San Diego Bay� � �  �                          impinged on power plant intake screens during July
          Tijuana Estuary  iI                             and August in San Diego Bay (San Diego Gas and
                      A S J L E                           Electric 1980). Modification and pollution of bays and
                                                          estuaries can significantly affect this species because
    July (White 1977). Spawning takes place in bays and   it spends its entire life within these habitats (Horn and
    estuaries at night (Heath 1980, Edmands 1983). This    Allen 1985).  Abundance of this species appears to
    speciesappearstospawnprimarilyinthelowerreaches    cycle widely; some years the slough anchovy is the
    of bays and estuaries, whereas the deepbody anchovy    dominant Anchoa species in California bays and other
    (A. compressa) utilizes the upper reaches of bays for   years A. compressadominates (Heath 1980). Reasons
    spawning (Edmands 1983).                              for the wide fluctuations are unknown, however the
                                                          slough anchovy may prefer cooler temperatures and
    Fecundity: Meanfecundity is approximately 7,000 eggs    more oceanic conditions for spawning than A.
    per female (or 1,418 eggs/g of female weight), with    compressa (Edmands 1983).
    larger fish producing more eggs (Heath 1980).
                                                          References
    Growth and Development
    Eggaa Size and Embrvonic Development: Eggs are   Allen, L. G., and M. H. Horn.  1975. Abundance,
    ellipsoid, similarto northern anchovy (Engraulismordax)   diversity and seasonality of fishes in Colorado Lagoon,
    eggs (Heath 1980), and are 0.94-1.10 mm maximum    Alamitos Bay, California. Est. Coast. Mar. Sci. 3:371-
    width (White 1977, Caddell 1988). Larval development    380.
    is indirect and external. Time to hatching is unknown,

                                                       105

    Slough anchovycontinued
    Caddell, S. M. 1988. Early life history descriptions of    San Diego Gas and Electric. 1980. Silvergate power
    the deepbodyand slough anchovieswithcomparisons    plant cooling water intake system demonstration [in
    to the northern anchovy (family Engraulidae).  Bull.    accordancewithsection316(b) FederalWater Pollution
    Mar. Sci. 42(2):273-291.                                 Control Act Amendment of 1972]. Rep. to Calif. Reg.
                                                           Water Qual. Control Board, San Diego Gas and Electric,
    Edmands, F. A., 11. 1983. The diel distribution and    San Diego, CA, various pagination.
    transport of ichthyoplankton collected by stationary
    nets in Newport Bay, California, July 1979.  M.A.    White,   W.  S. 1977.   Taxonomic composition,
    Thesis, Calif. State Univ., Fullerton, CA,112 p.         abundance, distribution and seasonality of fish eggs
                                                           and larvae in Newport Bay, California. M.A. Thesis,
    Eschmeyer, W. N., W. S. Herald, and H. Hammann.    Calif. State Univ., Fullerton, CA, 107 p.
    1983. A field guide to Pacific coast fishes of North
    America. Houghton Mifflin Co., Boston, MA, 336 p.

    Gates, D. E., and H. W. Frey. 1974. Designated
    common names of certain marine organisms of
    California. Calif. Fish Game, Fish Bull. 161:55-88.

    Heath, K. L. 1980. Comparative life histories of two
    species of anchovies, Anchoa delicatissima and A.
    compressa (F. Engraulidae) from Newport Bay,
    California. M.A. Thesis, Calif. State Univ., Fullerton,
    CA, 71 p

    Horn, M. H., and L. G. Allen. 1976. Numbers of species
    and faunal resemblance of marine fishes in California
    bays and estuaries. Bull. South. Calif. Acad. Sci.
    75(2):159-170.

    Horn, M. H., and L. G. Allen. 1985. Fish community
    ecology in southern California bays and estuaries.
    Chapter 8. In A. Yanez-Arancibia (editor), Fish
    community ecology in estuaries and coastal lagoons:
    towards an ecosystem integration, p. 169-190. DR (R)
    UNAM Press, Mexico.

    Love, M. S., J. S. Stephens, Jr., P. A. Morris, M. M.
    Singer, M. Sandhu, and T. C. Sciarrotta. 1986. Inshore
    soft substrata fishes in the southern California bight: an
    overview. Calif. Coop. Ocean. Fish. Invest. Rep.
    27:84-104.

    Miller, D. J., and R. N. Lea. 1972. Guidetothecoastal
    marine fishes of California. Calif. Fish Game, Fish Bull.
    157, 235 p.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Roedel, P. M. 1953. Common ocean fishes of the
    California coast. Calif. Fish Game, Fish Bull. 91,184
    p.

                                                       106

    107

       Engraulis mordax
       Adult

                        5cm

    Common Name: northern anchovy                          survival, juvenile feeding, and growth are reduced
    Scientific Name: Engraulis mordax                      when exposed to water-soluble fractions of crude oil
    OtherCommon Names: California anchovy, pinhead,    (MBC Applied Environmental Sciences 1987).
    anchoa, anchoveta, anchovy, bay anchovy, North
    American anchovy, plain anchovy                         Ecological: The northern anchovy is one of the most
    Classification (Robins etal. 1980)                      abundant fish in the California Current and is an
    Phylum: Chordata                                       important prey for many species of fishes, seabirds,
    Class: Osteichthyes                                    and marine mammals (Frey 1971, Eschmeyer et al.
    Order: Clupeiformes                                     1983). It is highly abundant in many Pacific coast bays
    Family: Engraulidae                                    and estuaries during spring, summer, and fall. Elegant
                                                           tern (Thalasseus elegans) and California brown pelican
    Value                                                   (Pelecanus occidentalis) production is strongly
    Commercial: The northern anchovy is commercially    correlated with anchovy abundance (Anderson et al.
    fished from British Columbiato northern Baja California,    1980, Schaffner 1986). The northern anchovy occupies
    Mexico, butprimarily from San Francisco, California,to    an ecological niche similarto the Pacific sardine's and
    Bahia San Ramon, Baja California.  It was not    may be inhibiting its comeback (Frey 1971).
    commercially important until after the collapse of the
    Pacific sardine (Sardinops sagax) fishery in the 1940s.    Range
    In 1981, over400,000 t were landed, representing the    Overall: The northern anchovy was distributed from
    25th largest species catch in the world (Food and    Cape San Lucas, Baja California, to Queen Charlotte
    Agriculture Organization 1984).   The California    Islands, Canada, but has recently moved into the Gulf
    commercial catch in 1981 was estimated to be worth    of California, Mexico (Hammann and Cisneros-Mata
    $3.2 million (Pacific Fishery Management Council    1989). Three geneticallydistinctsubpopulationsexist
    1983). This species is commercially fished for reduction    (Vrooman and Smith 1971). One rangesfrom northern
    (i.e., fish meal and paste) and live bait, however, the    California to British Columbia.  The second is off
    reductionfisheryhasdeclineddramaticallysince 1981.    southern California and the northern Baja California
                                                           peninsula in Mexico. The third occurs off central and
    Recreational: It isthe most important bait fishfornearly   southern Baja California (Vrooman and Smith 1971).
    all marine recreational fisheries off southern California.
    It is also used as bait in Oregon and Washington for   Within Studv Area: This species can be found in all
    sturgeon (Acipenserspp.), salmonids (Oncorhynchus    estuaries within the study area (Table 1). A subspecies
    spp.), and other fishes.                                (E. mordax nanus) is restricted to San Francisco Bay
                                                           (Hubbs 1925).
    Indicator of Environmental Stress: Low dissolved
    oxygen can cause die-offs (Pacific Fishery Management    Life Mode
    Council 1983).  Anchovy hatching success, larval    Eggs and larvae are planktonic, while juveniles and

                                                       108

                                                                                     Northern anchovy continued
                                                           stages are found over various substrates.
     Table 1. Relative abundance of northern anchovy
               in 32 U.S. Pacific coast estuaries.          Phvsical/Chemical Characteristics: Eggs are found in
                         Life Stage                        euhaline waters (32-35%o), while adults, juveniles, and
            Estuary    A S J L E                           larvae can be found in estuarine and marine waters
             PugetSound O O O O 0   Relative abundance:    (Simenstad 1983).  Spawning occurs at water
              HoodCanal O  O  O  O  O    �   Highly abundant  temperaturesof12-15�Candusuallywithin10mofthe
               SkagitBay 0 0 00    O i   Abundant           surface (Ahlstrom  1959).   Eggs are found in
             Grays Harbor           0      Common          temperatures of 10.0-23.30C, larvae at 10.0-19.7�C
                            mras o  O    O      Rare
              Willapa Bay �    � O  Blank Not present      (mostly 14.0-17.40C), and juveniles and adults at 5.0-
           Columbia River �    � O O                       25.0�C.  The lower lethal temperature for juveniles
             Nehalem Bay O    a O                           appears to be 7�C, but at 1 0.0�C larvae do not develop
            Tillamook Bay O    � O    Ufe stage:           properly.  Temperatures above 25�C are actively
              Netarts Bay O    O      A-Adults             avoided by juveniles and adults. (Brewer 1974).
                                      S - Spawning adults
              Siletz River 0    O     J - Juveniles
             YaqulnaBay O     0      L-Larvae             Miarations and Movements: The northern anchovy
                                      E-Eggs
              AlseaRiver                Eggs               does not make extensive migrations (Pacific Fishery
             SiuslawRiver  i                                Management Council 1983), but it does undertake
            Umpqua River                                   inshore-offshore movements as well as movements
               CoosBay    0                                alongtheshore. InthePacificNorthwest,juvenilesand
             Rogue River 0    �  adults move into estuaries during spring and summer
            Klamath River i � *                            and then out during fall (Waldvogel 1977, National
            Humboldt Bay *    *                            Marine Fisheries Service 1981, Simenstad and Eggers
                Eel River O    0                            1981). In southern California, young-of-the year and
             Tomales Bay  9   @3 0    t                    yearling anchovies utilize shallow inshore areas (Parrish
        Cent San Fr ran, .               Francc. Sunisan    et al. 1985). Adult and juvenile anchovies show some
        South San Fran.Bay  a     and San Pabb bays.        diel movements during the summer, staying at depths
            Elkhom Slough *    *  3                         of 1 10-183 m during the day and coming to the surface
               Morro Bay *    * 1                          at night (Hart 1973). Larvae swim to the surface at
         Santa Monica Bay  *- 6 � �                        night to gulp air and inflate their swim bladder (Hunter
            SanPedroBay  � � � �  �
             SanPedros Bay *- *     0and Sanchez 1976). Larvae, juveniles, and adults form
             Alamitos Bay  *:  a 0 0                       small low density schools during the day and disperse
             Anaheim Bay  0)   0 CD 0                       into athin surface scattering layerat night (Mais 1974).
             Newport Bay  3    3                           Juveniles and adults may also form dense schools or
              Mission Bay
            San Diego Bay 0    0 0 0                        "balls" when being attacked by predatory fishes.
           Tijuana Estuary                                 Reproduction
                       A S J L E                           Reproduction
                                                           ModA   J L E  : This species is gonochoristic, oviparous, and
                                                           iteroparous; eggs are fertilized externally. It is a
    adults are pelagic nekton (Garrison and Miller 1982).    broadcast spawner that spawns in batches annually
                                                           after reaching maturity.
    Habitat
    Tyje: Eggs are neritic and epipelagic (fromthesurface    Matina/SDawnina: Spawning is reported from Barkley
    to 50 m depth, but primarily in the upper 20 m). Larvae    Sound and the Strait of Georgia, British Columbia, to
    are also neritic and epipelagic, occurring from the    south of Magdalena Bay, Baja California, and in the
    surface to 75 m depth, but usually in the upper 50 m.    Gulf of California. Spawning can occurthroughoutthe
    Juveniles are epipelagic and often highly abundant in   yeardepending on region (i.e., subpopulation). Times
    shallow nearshore areas and estuaries. Adults are    for spawning are July to August in British Columbia
    oceanic-neritic, occurring from the surface to 300 m    waters, June to August off Oregon, Decemberto June
    deep. Adults can also be abundant in shallow nearshore    in central California waters, May to September in San
    areas and estuaries. Eggs and larvaecan be foundout    Francisco Bay, and January to May off southern
    to 480 km offshore (Hart 1973, Garrison and Miller   California (McGowan 1986).  Most spawning takes
    1982), while adults occur out to 157 km offshore    place within 100 km of the coast in the upper mixed
    (Pacific Fishery Management Council 1983).              layer (sometimes surface) at night (Baxter 1967, Hunter
                                                           and Macewicz 1980). The majority of spawning in
    Substrate: Because this is a pelagic species, all life   California waters occurs at depths less than 10 m and

                                                        109

    Northern anchovy continued
    water temperatures between 12 and 15�C. However,    Food Items: Larvae consume copepods (primarily eggs
    spawning has been recorded up to 482 km offshore    and nauplii), naked dinoflagellates, rotifers, ciliates,
    (Ahlstrom 1959).  In the northern subpopulation,    and foraminiferans (Baxter 1967, Arthur 1976, Hunter
    spawning appears to be associated with the Columbia    1977). Larvae, juveniles, and adults are often found in
    River plume, which may provide a stable and productive    areas of plankton blooms. Adults and juveniles prey on
    environment for egg and larval survival (Richardson    phytoplankton, planktonic crustaceans, and fish larvae
    1981). The timing of reproduction nearSan Pedro Bay,    (Loukashkin 1970, Frey 1971, Hart 1973, Pacific Fishery
    California, may be constrained bydietary requirements    Management Council 1983).
    (Brewer 1978). This species is a batch spawner
    (Hunter and Goldberg 1980) and may spawn about 20    Biological Interactions
    times per spawning season (Hunter and Leong 1981).    Predation: Northern anchovy eggs and larvae are
                                                          eaten by adult anchovies (Hunter 1977) and probably
    Fecundity: Females lay eggs in batches and can    many other fishes. In the California Current, juveniles
    produce up to 130,000 eggs per year (20 spawnings)    and adults are consumed by most species of predatory
    in southern California (Hunter and Macewicz 1980,   fishes, including California halibut (Paralichthys
    Hunter and Leong 1981).  Females in the northern   californicus), chinook (0. tshawytscha) and coho
    subpopulation are apparently limited to only a few    salmon (0. kisutch), rockfishes, yellowtail (Seriola
    batches and a totalfecundity of 35,000 eggs perfemale    lalandei), tunas, and sharks. Other predators include
    per year (Laroche and Richardson 1980).  Batch    harbor seal (Phoca vitulina), northern fur seal
    fecundities are estimated to be 2,794-16,662 eggs per    (Callorhinus ursinus), California sea lions (Zalophus
    female (Hunter and Macewicz 1980).                    califomianus), common murre (Uria aalge), sooty
                                                          shearwater   (Puffinus  griseus),  cormorant
    Growth and Development                                (Phalacrocorax spp.), gulls, and tems (Kucas 1986).
    Eaa Size and Embrvonic DeveloDment: Eggs are   The northern anchovy is the primary prey for the
    ellipsoidal with dimensions of 1.23-1.55 mm x 0.65-   California brown pelican, an endangered species
    0.82 mm  (Garrison and Miller 1982).  Embryonic    (Huppert et al. 1980).
    development is indirect and external. Eggs hatch in 2-
    4 days, depending on temperature.                     Factors Influencino Populations: Egg and larval survival
                                                          probably determines subsequent year-class strength
    Aae and Size of Larvae: The yolk sac is absorbed    (Smith 1985). However, egg and larval abundance are
    within36hoursofhatching(Laskeretal. 1970). Larvae    not correlated with age-1 recruits (Peterman et al.
    range from 2.5 mm to 25.0 mm in length (Hart 1973).    1988).  Anchovy spawning biomass is presently
    Larvae begin schooling at 11-12 mm standard length    estimated from egg production (Lasker 1985). Good
    (SL) (Hunter and Coyne 1982), and transform into   larval survival appears to depend on many factors,
    juveniles in approximately 70 days (Hart 1973).       including the availability and density of appropriate
                                                          phytoplankton species (Lasker 1975, Lasker and Smith
    Juvenile Size Ranae: Juveniles range in sizefrom 2.5-   1976, Lasker 1981, Peterman and Bradford 1987).
    14.0 cm SL (Clark and Phillips 1952).                 Larval food availability is reduced by storms and strong
                                                          upwelling. Strong upwelling may also transport larvae
    Aae and Size of Adults: Some fish mature at less than    out of the Southern California Bight (Power 1986),
    one year of age (7.1-10.0 cm) and all are mature at 4    however, upwelling may benefit later life stages. El
    years, dependingon location and populationsize (Clark   Nihfo events affect populations both positively and
    and Phillips 1952, Hart 1973, Hunter and Macewicz    negatively, depending on subpopulation and life stage
    1980, Laroche and Richardson 1980).  Larger fish   (Brodeuretal. 1985, Fiedleretal. 1986). High rates of
    mature earlier than smaller fish in the same age group    predation and commercial harvest also impact
    (Huppert et al. 1980). The maximum age reported for   populations. Northern anchovy populations increased
    this species is 7 years (Frey 1971).                  dramatically during the collapse of the Pacific sardine
                                                          populations, suggesting competition between these
    Food and Feeding                                      species (Smith 1972, Kucas 1986).
    Trophic Mode: Juveniles and adults are random filtering
    or particulate (i.e., biting) planktivores, depending on    References
    food concentrations (O'Connell 1972). Anchovies
    apparentlyfeed primarily duringthe day (Kucas 1986).    Ahlstrom, E. H. 1959. Vertical distribution of pelagic
    Females need to eat approximately 4-5% of their wet   fish eggs and larvae off California and Baja California.
    weight per day for growth and reproduction (Hunter   Fish. Bull., U.S. 60:107-146.
    and Leong 1981).

                                                       110

                                                                                     Northern anchovy continued
    Anderson, D. W., F. Gress, K. F. Mais, and P. R. Kelly.   northern anchovy, Engraulismordax Girard, in the Gulf
    1980. Brown pelicans as anchovy stock indicators and    of California, Mexico. Calif. Fish Game 75(1):49-53.
    their relationship to commercial fishing. Calif. Coop.
    Ocean. Fish. Invest. Rep. 21:54-61.                     Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
                                                           Board Can., Bull. No. 180, 740 p.
    Arthur, D. K. 1976. Food and feeding of larvae of three
    fishes occurring in the California Current, Sardinops    Hubbs, C. L. 1925. Racial and seasonal variation in
    sagax, Engraulis mordax, and Trachurus symmetricus.    the Pacific herring, California sardine, and California
    Fish. Bull., U.S. 74:517-530.                           anchovy. Calif. Fish Game, Fish Bull. 8:1-23.

    Baxter, L. L. 1967. Summary of biological information    Hunter, J. R. 1977. Behavior and survival of northern
    on the northern anchovy, Engraulis mordax Girard.    anchovy, Engraulis mordax, larvae.  Calif.  Coop.
    Calif. Coop. Ocean. Fish. Invest. Rept. 11:110-116.     Ocean. Fish. Invest. Rep. 19:138-146.

    Brewer, G. 1974. Thermal tolerance and sediment    Hunter, J. R., and K. M. Coyne. 1982. The onset of
    toxicity studies. In D. F. Soule and M. Oguri (editors),   schooling in northern anchovy larvae, Engraulismordax.
    Marine studies of San Pedro Bay, California, p. 21-43.    Calif. Coop. Ocean. Fish. Invest. Rep. 23:246-251.
    Allan Hancock Found., Off. Sea Grant Publ., Univ. S.
    Calif., Los Angeles, CA.                                Hunter, J. R., and S. R. Goldberg. 1980. Spawning
                                                           incidence and batch fecundity in northern anchovy,
    Brewer, G. D. 1978. Reproduction and spawning of    Engraulis mordax. Fish. Bull., U.S. 77(3):641-652.
    the northern anchovy, Engraulis mordaxin San Pedro
    Bay, California. Calif. Fish Game 64(3):175-184.        Hunter, J. R., and R. Leong.  1981. The spawning
                                                           energetics of female northern anchovy, Engraulis
    Brodeur, R. D., D. M. Gadomski, W. G. Pearcy, G. P.    mordax. Fish. Bull., U.S. 79(2):215-230.
    Batchelder, and C. B. Miller. 1985. Abundance and
    distributionofichthyoplankton intheupwellingzoneoff    Hunter, J. R., and B. J. Macewicz. 1980.  Sexual
    Oregon during anomalous El Nihio conditions. Estuar.    maturity, batch fecundity, spawning frequency, and
    Coast. Shelf Sci. 21:365-378.                           temporal pattern of spawning forthe northern anchovy,
                                                           Engraulis mordax, during the 1979 spawning season.
    Clark, F. N., and J. B. Phillips. 1952. The northern    Calif. Coop. Ocean. Fish. Invest. Rep. 21: 139-149.
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    Calif. Fish Game 38(2):189-207.                         Hunter, J. R., and C. Sanchez. 1976. Diel changes in
                                                           swim bladder inflation of the larvae of the northern
    Eschmeyer, W. N., W. S. Herald, and H. Hammann.    anchovy, Engraulismordax. Fish. Bull., U.S. 74(4):847-
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                                                           Huppert, D. D., A. D. MacCall, G. D. Stauffer, K. R.
    Fiedler, P. C., R. D. Methot, and R. P. Hewitt. 1986.    Parker, J. A. McMillan, and H. W. Frey.  1980.
    Effects of California El N iio 1982-1984 onthe northern    California's northern anchovy fishery: biological and
    anchovy. J. Mar. Res. 44:317-338.                       economic basis for fishery management. NOAA Tech.
                                                           Mem. NMFS, Nat. Mar. Fish. Serv., Southwest Fish.
    Food and Agriculture Organization. 1984. Yearbook    Cent., La Jolla, CA, 121 p. plus appendices.
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    U.N., Rome, 393 p.                                      Kucas, S. T., Jr. 1986. Species profiles: life histories
                                                           and environmental requirements of coastal fishes and
    Frey, H. W. 1971. California's living marine resources    invertebrates (Pacific Southwest)-northern anchovy.
    and their utilization.  Calif. Dept. Fish Game,    U.S. Fish Wildl. Serv. Biol. Rep. 82(11.50). U.S. Army
    Sacramento, CA, 148 p.                                  Corps Eng., TR EL-82-4, 11 p.

    Garrison, K. J. and B. S. Miller. 1982. Review of the    Laroche, J. L. , and S. L. Richardson   1980.
    early life historyof Puget Sound fishes. Fish. Res. Inst.,   Reproduction of northern anchovy, Engraulis mordax,
    Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).         off Oregon and Washington. Fish. Bull., U.S. 78(3):603-
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    Hammann, M. G., and M. A. Cisneros-Mata. 1989.
    Range extension and commercial capture of the

                                                        111

    Northern anchovy continued
    Lasker, R. 1975. Field criteria for survival of anchovy    anchovyfishery management plan, fourth draft revision.
    larvae: the relation between inshore chlorophyll    Pac. Fish. Manag. Council, Portland, OR, various
    maximum layers and successful first feeding. Fish.   pagination.
    Bull., U.S. 73:453-462.
                                                           Parrish, R. H., D. L. Mallicoate, and K. F. Mais. 1985.
    Lasker, R.  1981. Factors contributing to variable    Regional variations in the growth and age composition
    recruitmentof the northern anchovy (Engraulismordax)    of northern anchovy, Engraulis mordax.  Fish. Bull.,
    in the Califomia current: contrasting years, 1975 through    U.S. 83(4):483-496.
    1978. Rapp. P.-v. Reun. Cons. Int. Explor. Mer.
    178:375-388.                                           Peterman, R. M., and M. J. Bradford.  1987. Wind
                                                           speed and mortality rate of a marine fish, the northern
    Lasker, R. (editor). 1985. An egg production method    anchovy (Engraulis mordax). Science 235:354-356.
    for estimating spawning biomass of pelagic fish:
    applicationtothe northern anchovy, Engraulismordax.    Peterman, R. M., M. J. Bradford, N. C. H. Lo, and R. D.
    U.S. Dept. Commer., NOAA Tech. Rep. NMFS 36,           Methot.  1988.  Contribution of early life stages to
    99 p.                                                  interannual variability in recruitment of northern anchovy
                                                           (Engraulis mordax). Can. J. Fish. Aquat. Sci. 45(1):8-
    Lasker, R., H. M. Feder, G. H. Theilacker, and R. C.    46.
    May. 1970. Feeding, growth, and survivalof Engraulis
    mordaxreared in the laboratory. Mar. Biol. 5:345-353.    Power, J. H. 1986. A model of the drift of northern
                                                           anchovy, Engraulis mordax larvae in the California
    Lasker, R., and P. Smith. 1976.  Estimation of the   current. Fish. Bull., U.S. 84(3):585-603.
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    larvae of the northern anchovy. Calif. Coop. Ocean.    Richardson, S. L. 1981. Spawning biomass and early
    Fish. Invest. Rep. 19:128-137.                          life of northern anchovy, Engraulis mordax, in the
                                                           northern subpopulation off Oregon and Washington.
    Loukashkin, A. S.  1970.  On the diet and feeding    Fish. Bull., U.S. 78(4):855-876.
    behavior of the northern anchovy, Engraulis mordax
    (Girard). Proc. Calif. Acad. Sci. 37:419-458.           Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
                                                           E. A. Lachner, Robert N. Lea, and W. B. Scott. 1980.
    Mais, K. F. 1974. Pelagic fish surveys in the California    A listofcommon and scientific namesof fishesfromthe
    Current. Calif. Fish Game, Fish Bull. No. 162, 721 p.    United States and Canada. Amer. Fish. Soc. Spec.
                                                           Publ. No. 12, Am. Fish. Soc., Bethesda, MD,174 p.
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    of importantfisheriesspeciesoffshoreCalifornia. Miner.    Schaffner, F. C.  1986. Trends in elegant tern and
    Manag. Serv. Study 86-0093, MBC Appl. Envir. Sci.,   northern anchovy populations in California. Condor
    Costa Mesa, CA, 251 p.                                 88:347-354.

    McGowan, M. F. 1986. Northern anchovy, Engraulis   Simenstad, C. A. 1983.  The ecology of estuarine
    mordax, spawning in San Francisco Bay, California,    channels of the Pacific Northwest coast: a community
    1978-79, relativeto hydrography and zooplankton prey   profile. U.S. Fish Wildl. Serv., FWS/OBS-83/05, 181 p.
    of adults and larvae. Fish. Bull., U.S. 84(4):879-894.
                                                           Simenstad, C. A., and D. M. Eggers, (editors). 1981.
    National Marine Fisheries Service. 1981. Salmonid    Juvenile salmonidandbaitfishdistribution, abundance,
    and non-salmonid fishes. Annual Data Rep., Second    and prey resources in selected areas of Grays Harbor,
    Year, to Pacific NW River Basins Comm., CREDDP    Washington.  Final Rep. to Seattle Dist., U.S. Army
    Tasks A-2.8 and A-2.9, 139 p. Northwest and Alaska    Corps Eng., Fish. Res. Inst., Coill. Fish., Univ. Wash.,
    Fish. Cent., Natl. Mar. Fish. Serv., NOAA, 2725 Montlake    Seattle, WA, 205 p. (FRI-UW-8116).
    Blvd. E., Seattle, WA 98112.
                                                           Smith, P. E. 1972. The increase in spawning biomass
    O'Connell, C. P. 1972. The interrelation of biting and    of northern anchovy, Engraulis mordax. Fish. Bull.,
    filtering in the feeding activity of the northern anchovy    U.S. 70:849-874.
    (Engraulis mordax). J. Fish. Res. Board. Can. 29:285-
    293.                                                    Smith, P. E. 1985. Year-class strength and survival of
                                                            0-group clupeoids. Can. J. Fish. Aquat. Sci. 42 (Suppl.
    Pacific Fishery Management Council. 1983. Northern    1):69-82.

                                                        112

                                                                                     Northern anchovy continued
    Vrooman, A. M., and P. E. Smith. 1971. Biomass of the
    subpopulations of the northern anchovy Engraulis
    mordax Girard. Calif. Coop. Ocean. Fish. Invest. Rep.
    15:49-51.

    Waldvogel, J. B. 1977. Age, maturity and distribution
    of northern anchovy, Engraulis mordax, in Humboldt
    Bay, California. M.S. Thesis, Humboldt State Univ.,
    Arcata, CA, 36 p.

                                                        113

        Oncorhynchus clarki
        Juvenile

               5cm

    Common Name: cutthroat trout                            Range
    Scientific Name: Oncorhynchus clarki                    Overall: The overall rangeof this species' anadromous
    OtherCommonNames:Clark'strout, coastalcutthroat,    form is from the Eel River, California, to Seward,
    coastal cut-throat trout, sea-run cutthroat trout, red-   southeastern Alaska (Scott and Crossman 1973).
    throated trout, sea trout, short-tailed trout, harvest trout
    Classification (Smith and Stearley 1989)                Within Study Area:This species is common in nearly all
    Phylum: Chordata                                       estuaries along the Pacific coast from the Eel River to
    Class: Osteichthyes                                     Puget Sound, Washington (Table 1) (Monaco et al.
    Order: Salmoniformes                                    1990).
    Family: Salmonidae
                                                           Life Mode
    Value                                                   The cutthroat trout has four life histories: 1) an
    Commercial: The cutthroat trout is not commercially    anadromous form, 2) a form that migrates between
    fished, but isincidentallycapturedduringgillnettingfor    lakes and small streams, 3) a form that migrates
    Pacific salmon (Oncorhynchus spp.) (Tipping 1982).      between small tributaries and main rivers, and 4) a
                                                           form that lives its entire life in small streams (Trotter
    Recreational: It is the third most popular gamefish in   1987). This life history summary focuses primarily on
    the Pacific Northwest (Washington 1977).  In   the anadromous variety, 0. clarki clarki. Eggs and
    Washington, the Cowlitz Riverrecreational fishery was    larvae (alevins) are benthic and infaunal.  Young
    estimatedto be worth $290,000 recently (Tipping 1982).    juveniles (fry and parr) are benthopelagic; parr become
    Hatcheries in Oregon and Washington stockthis species    pelagic when they transform into smolts (juveniles that
    into numerous streams.                                  migrate to the ocean).  Smolts, ocean-dwelling and
                                                           maturing juveniles (subadults), and adults are primarily
    Indicatorof Environmental Stress: Thesea-run cutthroat    pelagic. Subadults and adults in rivers and streams are
    trout is sensitive to temperature changes and stream    benthopelagic.
    alterations resulting from logging practices (Moring
    and Lantz 1975). It has been compared to the "canary    Habitat
    in the mine", being one of the first species to sufferfrom    Iype: Eggs, alevins, fry, and parr are riverine. Smolts
    environmental degradation (Behnke 1987).                are riverine and estuarine.  Young-of-the-year are
                                                           often found only in small coastal streams; many of
    Ecological: The cutthroat trout is a minor predator in   these streams will have low summer flows. Subadults
    nearshore coastal waters (Loch and Miller 1988) and    and adults are found in coastal neritic waters during
    an important resident of many streams and rivers. It   ocean residence (spring and summer), and in riverine
    has been displaced by introduced salmonids and non-    habitats during the spawning migration.  Smolts,
    native fishes in many rivers and streams.               subadults, adults, and "kelts" (spent adults) migrate
                                                           through estuaries. Some individuals are permanent

                                                        114

                                                                                       Cutthroat trout continued

                                                           can be found in streams with flows as low as 0.01 -0.03
     Table 1. Relative abundance of cutthroat trout       m3/s (DeWitt 1954). Spawning occurs in stream flows
               in 32 U.S. Pacific coast estuaries.         ranging from 0.11-0.90 m/s and depths of 10-100 cm
                         Life Stage                        (Pauley et al. 1989).  While in fresh water, adults
           Estuary    A S K J L E                          typically reside in pools, while fry reside in riffles.
           Puget Sound 0    O 0O     Relative abundance:
            Hood Canal O    O O        �   Highly abundant  Miarations and Movements: Parr in fresh water often
             Skagit Bay    O  )  Abundant                  move upstream and downstream (Moring and Lantz
           Grays Harbor O    0 0       I  Rare             1975). Parr remain in streams for at least 1 year, but
            Willapa Bay O    O O      Blank Not present    may stay up to 9 years. Parr become smolts as they
          Columbia River 0    O O                          migratetoestuaries. In Oregon and Washington, most
           Nehalem Bay (O   0 a                            smoltsmigrateduringspringintheirthirdyear(Wydoski
           Tillamook Bay O    O 0      Life stage:         and Whitney 1979).  However, the juvenile's size
            Notarts Bay 0    00       A- Adults
                                       S - Spawning adults  appears to determine its year of migration; larger fish
             Siletz River O    O       K - Kelts           migratetoseawhile smallerfish remain (Tipping 1986).
            Yaquina Bay                L 00  J- Juveniles
                                       L -Larvae           In Oregon, immature fish moved downstream from
            Alsea River @3   0        E - Eggs            February through May, with April being a peak month
           Siuslaw River '3   0                                               Wo
           Umpqua River O I   for outmigration. In Washington, outmigration occurs
              Coos Bay O     O                             from March to July (peaking in May) (Michael 1989).
            Rogue River O    O O                           Few juveniles remain in the ocean for more than one
           Klamath River C    O O                          summer and most migrate back to natal streams in late
           Humboldt Bay "V   / -I                          summer and fall of the same year (Johnston 1982).
              Eel River O    O O                           Dependingonthestock, aproportion of thefish returning
           Tomales Bay                                    to fresh water after their first summer in the ocean are
      Cent. San Fran. Bay *  Includes Central San         still not reproductively mature (Johnston 1982). Prior
                                       Francisco, Suisun.
      South San Fran. Bay             and an Pablo Suisun.  to their spawning migration, adult cutthroat trout often
          Elkhom Slough                                    reside in tidal freshwater areas of estuaries, awaiting
             Morro Bay                                     increased stream flows and decreased water
        Santa Monica Bay                                   temperatures before proceeding upstream. InOregon,
          San Pedro Bay                                    adults move upstream from October to March, with
           Alamitos Bay                                   most movement during November through January;
           Anaheim Bay                                     kelts move downstream from January to April, with
            Newport Bay                                    most moving in January and February (Lowry 1965).
            Mission Bay                                    Some streams are used for overwintering only and
          San Diego Bay                                    others for spawning (Michael 1989). Afteroverwintering
          Tijuana Estuary                                  (or spawning), sea-run cutthroat trout migrate to the
                      A S K J L E                          ocean again in spring. Information concerning ocean
                                                           movements and migrations are limited, but some fish
                                                           do not migrate far from where they entered the ocean
    residents of estuaries (Levy and Levings 1978).        (Johnston 1982). However, some have been found out
                                                           to 31 km offshore (Loch and Miller 1988). The cutthroat
    Substrate: Eggs are found beneath gravel (0.6-10.2   trout may school while in estuarine and marine
    cm in diameter) in shallow riffle areas at the tail end of   environments (Giger 1972). When returning to their
    pools (Reiser and Bjornn 1979). Juveniles and adults    natal stream, wild fish rarely stray. However, straying
    occur over various substrates depending on life stage    of hatchery fish (from streams in which they were
    and habitat.                                           stocked) may be 30% (Pauley et al. 1989).

    Phvsical/Chemical Characteristics: The cutthroat trout    Reproduction
    prefers water temperatures of 9-120C (Bell 1984). It   Mode: The cutthroat trout is gonochoristic and
    can tolerate 260�C, but is not usually found where    oviparous; eggs are fertilized externally. This species
    stream temperatures are consistently greater than    differs from all other members of the genus
    220C (Pauley et al. 1989).  The best spawning    Oncorhynchus (except steelhead trout, 0. mykiss) in
    temperature appears to be 10�C, but spawning occurs    being iteroparous.
    over a range from 6-17�C (Scott and Crossman 1973).
    Waters withdissolvedoxygen concentrations less than    Matina/SDawnina: Sea-run cutthroat trout return to
    5 mg/I are avoided (Pauley et al. 1989). This species   their natal streams to spawn from late fall to late winter

                                                        115

    Cutthroat trout continued
    (Johnston 1982, Pauley et al. 1989), however, only 41 -   salmonids, euphausiids, mysids, and crab megalopae
    61% of a "run" may actually be sexually mature (Jones    (Brodeur et al. 1987, Loch and Miller 1988).
    1977). Spawning occurs primarily in gravel riffles of
    small tributary coastal streams at the tail of pools in   Biological Interactions
    water that is 10-15 cm deep (Jones 1977). Like other    Predation: Little is known about predation on this
    salmonids, the female digs a redd in thegravel and lays   species, but 58% of the adults and subadults returning
    her eggs while the male fertilizes them with his milt.   to the Alsea River, Oregon, had marks indicating
    The female then covers the eggs with more gravel.   predator attacks (Giger 1972). Marine mammals prey
    Although this species is iteroparous, substantial post-   on this species at sea, while belted kingfishers
    spawning mortality can occur.  The best spawning    (Megaceryle alcyon) and other piscivorous birds are
    conditions include incubation temperatures from 6.1-   probably major predators in streams and estuaries.
    17.20C, depths >6 cm, water velocities from 11-72 cml    Sculpins and salmonids may also be predators of
    sec, and gravel that is 0.6-10.2cm2 in diameter (Reiser   alevins and fry in streams.
    and Bjornn 1979).
                                                          Factors Influencino Populations: This species is very
    Fecundity: Fecundity ranges from 226-4,420 eggs per   sensitive to changes in its freshwater habitat. The
    female (depending on female size), averaging 1,000-    amount of cover, water quality, and substrate
    1,700 (Scott and Crossman 1973).                      characteristics determine stream population densities
                                                          (Reiser and Bjornn 1979). Forestry practices influence
    Growth and Development                                stream carrying capacity and can affect spawning
    Eaa Size and Embrvonic Develooment: Eggs are 4.3-   success. Increases intemperatureandturbidity reduces
    5.1 mm in diameter, orange-red in color, and demersal    cutthroat trout production (Behnke 1979) and predation,
    (Pauley et al. 1989). Embryonic development is indirect   disease, residualism, and straying, affect the number
    and external.  Eggs usually hatch in 28-40 days    of returning adults (Tipping 1982). The myxosporidean
    (depending on temperature) (Scott and Crossman    protozoan Ceratomyxashastacan cause severe larval/
    1973).                                                juvenile mortalities in hatcheries (Tipping 1988). Natural
                                                          production of the sea-run cutthroat appears to be
    Aae and Size of Larvae: Alevins are 15 mm long at   severely depressed in many rivers and watersheds. In
    hatching and spend 1 to 2 weeks in the redd before    someareas, urbanizationhasadverselyaffectedstream
    emerging. Fry (small young juveniles) are approximately    environments and subsequently cutthroat trout
    35 mm in length.                                      populations (Trotter 1987). Ocean survival of first-year
                                                           smolts reportedly ranges from 1.8-21.7% in Washington
    Juvenile Size Ranae: Juveniles range from 35-200 mm    (Michael 1989) and 20-40% in Oregon (Giger 1972).
    in length.                                            Survival of subadults and adults in fresh water ranges
                                                          from 22.2-76.9% (Michael 1989). Because sea-run
    Aae and Size of Adults: Wild sea-run cutthroat mature    cutthroat trout are accessible to many anglers and
    after 2-10 years, ranging in length from 131 to 450 mm    relatively easy to catch, populations are easily
    (Summer 1962, Scott and Crossman 1973, Jones    overfished (Jones 1977, Tipping 1982). As a result,
    1977). However, hatchery fish grow quicker than wild    strict harvest restrictions have been implemented in
    fish and may return to spawn as one-year-old fish   British Columbia and Washington (Pauley et al. 1989).
    (Tipping 1982).                                       The genetic integrity of some stocks is threatened
                                                           because there are very few adults in the spawning
    Food and Feeding                                      population (Michael 1989).  By selecting the small
    Trophic Mode: Larvae feed on theiryolk. Juveniles and    tributaries of rivers and streams for spawning, sea-run
    adults are carnivorous.                               cutthroat avoid competition with rainbow trout and
                                                          coho salmon (Johnston 1982, Pauley et al. 1989).
    Food Items: Fry feed on insects, crustaceans, and    Although stream-dwelling juveniles eat similar foods
    somefish. Largecutthroattroutmaypreyonthreespine    as juvenile coho salmon, competition is reduced by
    stickleback (Gasterosteus aculeatus) and young    habitat partitioning. Juvenile cutthroat trout are often
    sockeye(O. nerka)andcoho(O. kisutch) salmon while    forced to reside in riffle areas until falling water
    in fresh water(Lowry 1966, Pauley et al. 1989). Large   temperatures reduce the aggressive behavior of other
    juveniles (migrants) and adults are highly piscivorous    salmonids (Glova 1986, 1987, Pauley et al. 1989).
    when in estuaries and marine waters (Behnke 1979,
    Loch 1982). In the ocean, cutthroat trout feed on
    northern anchovy (Engraulis mordax), kelp greenling
    (Hexagrammos decagrammus), scorpaenids,

                                                       116

                                                                                      Cutthroat trout continued

    References                                            Mar. Serv. Manuscr. Rep. 1475, 63 p.

    Behnke, R.J. 1979. Monograph of the native trouts of    Loch, J. J. 1982. Juvenile and adult steelhead and
    the genus Salmoof western North America. U.S. Fish   sea-run cutthroattrout withinthe Columbia Riverestuary
    and Wildl. Serv., and U.S. Forest Serv., Lakewood,    1980. 1982 Ann. Rep., Wash. Dept. Game, Olympia,
    CO, 163 p.                                            WA, 47 p. plus appendices.

    Behnke, R. J.  1987.  Forward.  In P. C. Trotter,   Loch, J.J.,andD. R. Miller. 1988. Distribution and diet
    Cutthroat: native trout of the west. Colorado Assoc.    of sea-run cutthroat trout captured in and adjacent to
    Univ. Press, Boulder, CO.                             the Columbia River plume, May-July 1980. Northw.
                                                          Sci. 62(1):41-48.
    Bell, M. C. 1984. Fisheries handbook of engineering
    requirements and biological criteria. Fish Passage    Lowry, G. R. 1965. Movementofcutthroattrout, Salmo
    Development and Evaluation Program, U.S. Army    clarki clarki (Richardson) in three Oregon coastal
    Corps Eng., North Pac. Div., Portland, OR, 290 p.   streams. Trans. Am. Fish. Soc. 94(4):334-338.
    (Contract No. DACW57-79-M-1 594 and DACW57-80-
    M-0567).                                              Lowry, G. R. 1966. Production and food of cutthroat
                                                          trout in three Oregon coastal streams. J. Wildl. Manag.
    Brodeur, R. D., H. V. Lorz, and W. G. Pearcy. 1987.    30(4):754-767.
    Food habits and diet variations of pelagic nekton off
    Oregon and Washington, 1979-1984. NOAA, Tech.    Michael, J. H., Jr. 1989. Life history of anadromous
    Rep. NMFS 57, 32 p.                                   coastal cutthroat trout in Snow and Salmon Creeks,
                                                          Jefferson County, Washington, with implications for
    DeWitt, J. W., Jr. 1954. Asurveyofthecoastcutthroat    management. Calif. Fish Game 75(4):188-203.
    trout, Salmo clarki clarki Richardson, in California.
    Calif. Fish Game 40:329-335.                          Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
                                                          Nelson. 1990. Distribution and abundance of fishes
    Giger, R. D.  1972.  Ecology and management of   and invertebrates in west coast estuaries, Volume I:
    coastal cutthroat trout in Oregon. Fish. Res. Rep. No.    data summaries. ELM R Rep. No. 4. Strategic Assess-
    6, Oregon State Game Comm., Portland, OR, 61 p.       ment Branch, NOS/NOAA, Rockville, MD, 240 p.

    Glova, G. J. 1986. Interaction for food and space    Moring, J. R., and R. L. Lantz. 1975. Alsea watershed
    between experimental populations of juvenile coho    study: effects of logging on the aquatic resources of
    salmon (Oncorhynchus kisutch) and coastal cutthroat   three headwater streams on the Alsea River, Oregon.
    trout (Salmoclarki) in a laboratory stream. Hydrobiol.    Part I-Biological studies.  Fish. Res. Rep. No. 9,
    131:155-168.                                          Oregon Dept. Fish Wildl., Corvallis, OR, 66 p.

    Glova, G. J. 1987. Comparison of allopatric cutthroat    Pauley, G. B., K. Oshima, K. L. Bowers, and G. L.
    troutstockswiththosesympatricwithcohosalmonand    Thomas.  1989. Species profiles: life histories and
    sculpins in small streams. Env. Biol. Fish. 20(4):275-    environmental requirements of coastal fishes and
    284.                                                  invertebrates (Pacific Northwest)-sea-run cutthroat
                                                          trout. U.S. Fish Wild. Serv. Biol. Rep. 82(11.86), U.S.
    Johnston, J.  1982.  Life histories of anadromous    Army Corps Eng. TR EL-82-4, 21 p.
    cutthroat with emphasis on migratory behavior. In E. L.
    Brannon and E. O. Salo (editors), Salmon and trout   Reiser, D. W., and T. C. Bjornn.  1979.  1. Habitat
    migratory behavior symposium, p. 123-127. School    requirements of anadromous salmonids.  In W. R.
    Fish., Univ. Wash., Seattle, WA.                      Meehan (editor), Influence of forest and rangeland
                                                          management on anadromous fish habitat in the western
    Jones, D. E. 1977. Life history of steelhead trout and    United States and Canada, p. 1-54.  USDA Forest
    life history of sea-run cutthroat trout. Alaska Dept. Fish    Service, Gen. Tech. Rep. PNW-96, Pac. Northw. Forest
    Game, Compl. Rep. AFS-42, 18:52-105.                   Range Exp. Sta., Portland, OR.

    Levy, D. A., and C. D. Levings. 1978. A description of   Scott, W. B., and E. J. Crossman. 1973. Freshwater
    the fish community of the Squamish River estuary,   fishes of Canada. Fish. Res. Board Can., Bull. No. 84,
    British Columbia: relative abundance, seasonal    966 p.
    changes and feeding habits of salmonids. Can. Fish.

                                                       117

    Cutthroat trout continued
    Smith, G. R., and R. F. Stearley. 1989. Theclassification
    and scientific names of rainbow and cutthroat trouts.
    Fisheries 14(1) :4-10.

    Summer, F. H. 1962. Migration and growth of coastal
    cutthroat trout in Tillamook County, Oregon. Trans.
    Am. Fish. Soc. 91:71-83.

    Tipping, J. 1982. Cowlitz Riversea-run cutthroat 1979-
    1981. Wash. Dept. Game, Olympia, WA, 24 p.

    Tipping, J. 1986. Effect of release size on return rates
    of hatchery sea-run cutthroat trout. Prog. Fish-Cult.
    48:195-197.

    Tipping, J. 1988. Ozone control of ceratomyxosis:
    survival and growth benefits to steelhead and cutthroat
    trout. Prog. Fish-Cult. 50:202-210.

    Trotter. P.C. 1987. Cutthroat: native trout of the west.
    Colorado Assoc. Univ. Press., Boulder, CO, 219 p.

    Washington, P. 1977. The sea-run cutthroat trout
    resource and sport fishery. Mar. Fish. Rev.39(12):20-
    22.

    Wydoski, R. S., and R. R. Whitney. 1979. Inland fishes
    of Washington, Univ. Wash. Press, Seattle, WA, 220 p.

                                                         118

    119

                        25 cm
    Common Name: pink salmon                              salmon occur in Alaska, although in odd years they are
    Scientific Name: Oncorhynchus gorbuscha               caught in Oregon and Washington (21,000 in 1983)
    Other Common Names: humpy salmon, dog salmon,    (Pacific Marine Fisheries Commission 1985,1987).
    hone salmon, humpback salmon, lost salmon (Shiino   This species is primarily captured when fishing for
    1976, Alaska Department of Fish and Game 1985)        othersalmon species, although it is regionally abundant
    Classification (Robins et al. 1980)                   at times. The pink salmon is caught by trolling in
    Phylum: Chordata                                      nearshore marine waters and by spincasting in streams
    Class: Osteichthyes                                   and along beaches (Squire and Smith 1977).
    Order: Salmoniformes
    Family: Salmonidae                                    Indicator of Environmental Stress: As with other
                                                          salmonids, destruction of spawning habitat reduces
    Value                                                 run sizes.
    Commercial: The pink salmon is the smallest Pacific
    salmon and fishermen receivethe lowestprice/lbforit.   Ecological: The pink salmon is the most abundant
    However, it isthe most abundant salmon species inthe   epipelagic fish in the subarctic oceanic North Pacific
    North Pacific. Annual harvest is over 84 million fish,   (Fredin et al. 1977).  See "Factors Influencing
    with over 95% of the U.S. catch coming from Puget    Populations".
    Sound, Washington, through Alaska (Forrester
    1981a,1981b, Takehama 1983). In 1985, landings of   Range
    pink salmon (144.7 t) were worth $75 million to U.S.   Overall: Overall, the pink salmon is found in oceanic
    fishermen. (National Marine Fisheries Service 1986).    and coastal areas of the North Pacific Ocean, north of
    Since virtually all pink salmon mature in their second    about 40�N latitude, in the Bering Sea, and along the
    year, commercial catches in a particular area fluctuate   southern coastline of the Polar Sea (Neave 1962). In
    markedly from one year to the next. In Puget Sound,    North America, occasional runs occur in the Russian
    odd-year runs predominate, but in the Gulf of Alaska    River, California, and along the Oregon coast. Regular
    and Bristol Bay, even-year runs are largest (Fredin et   spawning runs occur from the Puyallup River,
    al. 1977). Most Puget Sound pink salmon are captured    Washington, north to central Alaska, west to Attu
    from July to September (Washington Department of   Island, north to northern Alaska, and east to the
    Fisheries and Northwest Indian Fisheries Commission    Mackenzie River in Canada's Northwest Territories. In
    1986). This species is harvested primarily by purse    Asia, this species is distributed from the Tumen and
    seines, but also by trolling, stationary and drift gill nets,    North Nandai Rivers, North Korea, and Hokkaido,
    and reef nets.                                        Japan, to the Yana and Lena Rivers that flow into the
                                                          Arctic Ocean (Takagi et al. 1981). The pink salmon has
    Recreational: The pink salmon is not as important as    also been successfully introduced intothe Great Lakes
    coho (0. kisutch) andchinook salmon (0. tshawytscha)    (Scott and Crossman 1973).
    to coastal sport fisheries. Most sport harvests of pink

                                                       120

                                                                                         Pink salmon continued
                                                           Whitney 1979, Takagi et al. 1981).
      Table 1. Relative abundance of pink salmon
               in 32 U.S. Pacific coast estuaries.         Habitat
                         Life Stage                       Iye: Eggs and alevins occur primarily in the lower
             Estuary    A S J L E                          reaches of rivers, but can also occur in intertidal
              PugetSound �    �       Relative abundance:  estuarine areas (Helle et al. 1964, McNeil 1966). Fry
              Hood Canal �  1   0     �   Highly abundant    are riverine initially, but soon move downstream and
               Skagit Bay a�    �   i      Abundant        utilize estuaries and nearshore shallow water marine
             Grays Harbor             0 0   Common         environments (Healey 1980, 1982, Simenstad et al.
             Grays                         Rare
              Willapa Bay            Blank Notpresent      1982). Juveniles are initially neritic, but become oceanic
            Columbia River  i                              as they mature.  Adults are primarily estuarine and
             Nehalem Bay                                   riverine.
             Tillamook Bay             Life stage:
              Netarts Bay             A-Adults            Substrate: Eggs and alevins are normally found in
               Siletz River            S - Spawning adults  gravel that is 1.3-10.2 cm in diameter (Reiser and
                                       J - Juveniles
              Yaquina Bay              L - Larvae          Bjornn 1979). Gravel cover protects eggs and alevins
              Alsea River             E - Eggs            from predation, mechanical injury, and ultraviolet light
             Siuslaw River                                 (Raleigh and Nelson 1985). Fry, juveniles, and adults
             Umpqua River                                  are found in the water column over various substrates.
                Coos Bay
              Rogue River i                                Phvysical/Chemical Characteristics: Eggs and alevins
             Klamath River i'                              are found primarily in fresh water, but can withstand
             Humboldt Bay                                  constant salinities of 18%o and brief periods of higher
                Eel River                                  salinities (33%o) (McNeil 1966, Takagi et al. 1981). Fry
             Tomanes Bay                   er             adapt very quickly to high salinities (Takagi et al. 1981)
        Cent San Fran. Bay'  -       Includes Central San
                                      Francisco. Suisun.   and the species was originallythought to require marine
        South San Fran. Bay          and San Pablo bays.   waters for survival (Baggerman 1960). However, the
            Elkhorn Slough                                 successful introduction of pink salmon into the Great
               Morro Bay                                   Lakes demonstrates that this species can complete its
            SantaeMonica Bay                               entire life cycle in fresh water.  The pink salmon
             San Pedro Bay                                generally spawns at temperatures of 7.2-12.80�C, with
             Alamitos Bay  0incubation temperatures of 4.4-13.3�C providing the
             Anaheimwport Bay                             best hatching (Bell 1984). Optimum temperatures for
              Mission Bay                                  pink salmon are 5.6-14.40C, with 0.0�C and 25.6�C
              Mission Bay
            San Diego Bay                                  being lower and upper lethal limits, respectively (Bell
            Tijuana Estuary                                1984). Low pH impairs embryo and alevin development
                        A S J L E                          (Rombough 1983). Embryos and alevins need fast-
                                                           flowing (21-101 cm/sec) and well-oxygenated (>6 mg/
                                                           I) water for proper development and survival (Bailey et
    Within Studv Area: Although there are reports of pink   al. 1980).  Spawning gravel must be permeable to
    salmon occurring in many California rivers (Hallock    water flow for proper egg and alevin development
    and Fry 1967), probably only the Russian River and    (Wickett 1962, McNeil 1969). Adults cannot migrate
    possibly the Sacramento River have any spawning    upstream in velocities greater than about 2.13 m/sec
    runs (Fry 1973). Only very limited spawning runs occur    (Reiser and Bjornn 1979).
    along the Oregon and Washington coasts, but strong
    spawning runs occur in Puget Sound (Atkinson et al.   Miarations and Movements: The pink salmon is a
    1967) (Table 1).                                       highly-migratory anadromous species. Downstream
                                                           movement begins immediately upon emergence from
    Life Mode                                              the gravel (Neave 1966), and normally at night
    The pink salmon is an anadromous species. Eggs and    (McDonald 1960, Neave 1966). Fry are about 30 mm
    larvae (alevins) are benthic and infaunal.  Young    long at emergence.  Peak out-migration from rivers
    juveniles (fry) are benthopelagic and live in shallow    occurs between late March and mid-May in southern
    waters.  Ocean-dwelling and maturing juveniles    British Columbia, Washington, and Oregon (Healey
    (subadults) and adults are epipelagic, occurring possibly    1982, Simenstad et al. 1982). Most pink salmon spend
    down to depths of 36 m, but usually within the top 10 m    little time residing in estuaries (Levy et al. 1979, Healey
    (Hart 1973, Scott and Crossman 1973, Wydoski and    1982, Simenstad et al. 1982), but move and disperse

                                                        121

    Pink salmon continued
    rapidly into shallow marine waters and nearshore    Growth and Development
    nursery areas (Healey 1980). However, they may be    Eaa Size and Embrvonic DeveloDment: Eggs are 6.0-
    abundant in estuarine tidal channels for a short time   7.0 mm and orange-red in color (Scott and Crossman
    (Levy and Northcote 1982). As juveniles grow to about    1973, Bell 1984). Embryonic development is indirect
    60-80 mm in length (May and June), they move to   andexternal. Incubationtimeisaffectedbytemperature,
    offshore waters (Healey 1980), with larger individuals   but hatching occurs primarily in December and January
    moving first. During their first summer and fall, migrating   (McPhail and Lindsey 1970, Scott and Crossman 1973).
    pink salmon move north in coastal waters. By late fall/
    early winter, many turn south, dispersing to the high   Aae and Size of Larvae: Alevins are 6.0 mm to 30-45
    seas (Takagi et al. 1981, Hartt and Dell 1986). Pink   mm in length (Morrow 1980) and remain in the gravel
    salmon return to their natal streams after about 18   until most of the yolk is absorbed. Peak emergence is
    months at sea. Some pink salmon apparently never   in April and May, but may begin as early as late
    leave Puget Sound (Wydoski and Whitney 1979). A    February (Neave 1966).
    combined map-compass-calendar system probably
    guides this species on the high seas, but olfaction   JuvenileSizeRanae:Juvenilesareapproximately3.0-
    dominates riverine orientation as adults return to their   45.0 cm long and weigh up to 1.8 kg (Bell 1984). Pink
    natal stream (Brannon 1982, Quinn 1982). Upstream    salmon move to the open ocean when they are 6.0-8.0
    (i.e., spawning) migration may be disrupted if adults   cm long (central British Colu mbia) or 9.0-1 0.0 cm long
    encounter hydrocarbon concentrations above 1-10   (Strait of Georgia) (Healey 1980).
    ppb (Martin et al. 1990).
                                                          Aae and Size of Adults: Adults are two years old with
    Reproduction                                         rare reports of three-year-olds (Scott and Crossman
    Mode: The pink salmon is gonochoristic, oviparous,    1973). Adults can reach 76.0 cm in length and weigh
    and semelparous (all adults die soon after spawning).    5.5 kg, however most are 1.4-2.3 kg (Hart 1973).
    Eggs are fertilized externally.
                                                           Food and Feeding
    Matina/SDawnina: Spawning generally occurs from    TrophicMode:Larvaefeedontheiryolk. Juvenile and
    June (north) to October (south), and primarily August    adult pink salmon are carnivorous, opportunistic
    through October in Washington (Atkinson et al. 1967,   feeders.
    Wydoski and Whitney 1979). Most spawning takes
    place in the lower reaches of coastal rivers and can    Food Items: Fry will feed sparingly on nymphal and
    include intertidal areas (Helle et al. 1964). However,    larval insects if their migration to the ocean is lengthy
    pink salmon may spawn far upstream in large rivers   (Scott and Crossman 1973).  In nearshore nursery
    such as the Skagit River, Washington (Wydoski and    areas,juvenilepink salmon eat mainlyepibenthicprey,
    Whitney1979). Spawningusuallyoccursinriffleareas   particularly harpacticoid copepods (Gerke 1972,
    >15 cm deep, with water velocities of 12-101 cm/s, in   Kaczynski et al. 1973, Godin 1981). However, juveniles
    gravel that is 1.3-10.2 cm in diameter, and at   will also eat pelagic zooplankton such as Cirripedae
    temperatures of 7.2-12.80C (Reiser and Bjornn 1979).   larvae, calanoid copepods, amphipods, crustacean
    In Alaska, preferred spawning velocities are 35-47 cm/   larvae, and other invertebrate larvae (Kaczynski et al.
    s (Bonar et al. 1989). Females build the redd (nest) by    1973, Bailey et al. 1975, Fresh et al. 1979, Godin
    digging up the substrate with the caudal fin. During    1981). When juvenile pink salmon first enter offshore
    spawning, the female and male move to the bottom of   habitats,theyfeed on zooplankton, primarily copepods,
    the redd and release eggs and sperm while vibrating,   amphipods, chaetognaths, larvaceans, decapodlarvae,
    gaping their mouths, and erecting their fins.  The    andlarvalandjuvenilefishes(Healey1980, Brodeuret
    femalewillthendepositgravelovertheeggsbydigging   al. 1987).  Later in life, they feed on euphausiids,
    upstream of the redd. Males may spawn with more    decapod larvae, fishes, amphipods, squids, copepods,
    than one female, and females with more than one    pteropods, and other invertebrates (Allen and Aron
    male. Females may dig more than one nest (Scott and    1958, Andrievskaya 1958, Ito 1964, LeBrasseur 1966,
    Grossman 1973). Males develop enlarged teeth, a    Hart 1973, Fresh et al. 1981, Takagi et al. 1981). Pink
    large hump on their back, a hooked snout, and when    salmon are usually crepuscular feeders (Godin 1981,
    mature, are aggressive toward other males (Scott and    Takagi et al. 1981), however, they are known to feed on
    Crossman 1973).                                       euphausiids at night (Pearcy et al. 1984).

    Fecundity: Fecundity ranges from 800-2,000 eggs per    Biological Interactions
    female, averaging 1500-1900 (depending on size of   Predation: Eggs, alevins, andfry are eatenbycutthroat
    female) (Scott and Crossman 1973).                   trout (0. dark/), rainbow trout (0. mykiss), ooho salmon,

                                                       122

                                                                                     Pink salmon continued
    Dolly Varden (Salvelinus malma), northern squawfish    help maintain and rehabilitate pink salmon stocks and
    (Ptychocheilus oregonensis), and various sculpins   millions of pink salmon are released annually (Wahle
    (Cottusspp.) (Hunter 1959, Scott and Crossman 1973).    and Smith 1979). However, increased fishing pressure
    Belted kingfisher (Megaceryle alcyon), mergansers,    due to hatchery runs can destroy wild populations
    other predatory birds and small mammals also eat fry   (McNeil 1980).
    (Scott and Crossman 1973). Mammals (e.g., bears)
    and large avian predators (e.g., bald eagles, Haliaeetus    References
    leucocephalus) feed on adult pink salmon in fresh
    water. Marine and estuarine fish predators include    Alaska Department of Fish and Game. 1985. Alaska
    lamprey (Lampetra spp.), spiny dogfish (Squalus    habitat managementguide. Southcentral Region, Vol.
    acanthias), coho salmon, chinook salmon, rainbow   I: Life histories and habitat requirements of fish and
    trout, cutthroat trout and Pacific staghorn sculpin   wildlife. Alaska Dept. Fish Game, Juneau, AK, 429 p.
    (Leptocottus armatus). Predatory birds such as
    common murre (Uria aalge), common merganser    Allen, G. H., and W. Aron. 1958. Food of salmonid
    (Mergus merganser), bald eagle, and Caspian    fishes of the western North Pacific Ocean. U.S. Fish
    tern(Hydroprogne caspia), and mammals such as   Wildl. Serv., Spec. Sci. Rep. Fish. 237,11 p.
    harbor seal (Phoca vitulina), northern fur seal
    (Callorhinus ursinus), killer whale (Orcinus orca), and    Ames, J. 1983. Salmon stock interactions in Puget
    sea lions also prey on the pink salmon (Fresh 1984).    Sound: a preliminary look.  In M. A. Miller (editor),
    Smalljuvenilepinksalmonapparentlyaltertheirhabitat    Southeast Alaska coho salmon research and
    preferences depending on predation risk (Magnhagen    management review and planning workshop, May 18-
    1988).                                                19,1982, p. 84-95. Alaska Dept. Fish Game, Juneau,
                                                          AK.
    Factors Influencina PoDulations: Chum (0. keta) and
    pink salmon have similar feeding habits during their   Andrievskaya, L. D. 1958. Pitanie tikhookeanskikh
    early marine life; thus, competition may be occurring in   lososei v severo-zapadnoi chasti tikhovo okeana (The
    the shallow marine habitats (Ames 1983, Fresh 1984).    food of Pacific salmon in the northwestern Pacific
    A chum escapement variable is used inthe Washington    Ocean). [In Russ.] From: Materialy po biologii morskovo
    Department of Fisheries' model for forecasting pink    perioda zhizni dalnevostochnykh lososei, p. 64-75.
    salmon abundance/returns (Washington Department    Publ. by: Vses Nauchno-lssled. Inst. Morsk. Rybn.
    of Fisheries 1983).  One of the primary factors    Khoz. Okeanogr. (VNIRO), Moscow. [Fish. Res. Board
    determining recruitment appears to be survival from    Can., Trans. Ser. No. 182.]
    egg to fry stage (McNeil 1966, 1969, 1980), which is
    typically around 10% (Merrell 1962, McNeil 1980).    Atkinson, C. E., J. H. Rose, and T. O. Duncan. 1967.
    Mortality can result from low dissolved oxygen    Pacific salmon in the United States. Internat. North
    concentrations, high temperatures, high stream    Pac. Fish. Comm., Bull. No. 23:43-223.
    discharges, and unsuitable gravel structure (McNeil
    1966). Average marine survival from fry to adult is   Baggerman, B.  1960.  Salinity preference, thyroid
    about 4% (McNeil 1980), with much of the mortality   activity and seaward migration offourspecies of Pacific
    believed to occur as a result of predation during early   salmon (Oncorhynchus).  J. Fish. Res. Board Can.
    marine residency (Parker 1971). There also appears    17(3):295-322.
    to be density-dependent marine mortality and growth
    (Peterman 1980). Suitablecoastalwatertemperatures    Bailey, J. E., B. L. Wing, and C. R. Mattson. 1975.
    and salinities are also considered important to juvenile   Zooplankton abundance and feeding habits of fry of
    survival (Tabata 1983).  Besides natural mortality,    pink salmon, Oncorhynchus gorbuscha, and chum
    there is fishing and incidental fishing mortality (Ricker   salmon, Oncorhynchusketa in Traitor's Cove, Alaska,
    1976). Although the U.S. harvest of pink salmon has    with speculations on the carrying capacity of the area.
    declined since the 1930s, the Canadian harvest has    Fish. Bull., U.S. 73:846-861.
    not (Fredin 1980). Some pink salmon originating from
    North America are taken by the Japanese salmon    Bailey, J. E., S. Rice, J. Pella, and S. Taylor. 1980.
    fishery (Fredin et al. 1977). Man-made alterations to   Effectsofseedingdensityofpinksalmon, Oncorhynchus
    streams, estuaries, and shallow marine environments    gorbuscha, eggs on water chemistry, frycharacteristics
    caused by improper road and rail construction, logging    and fry survival in gravel incubators. Fish. Bull., U.S.
    practices, dredging, bulkheading, dam and irrigation   78(3):649-658.
    development, and pollution can adversely affect pink
    salmon populations. Hatcheries have been built to

                                                       123

    Pink salmon continued
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    development and evaluation program, U. S. Army    studies in the Nisqually Reach area of southern Puget
    Corps Eng., North Pac. Div., Portland, OR, 290 p.   Sound, Washington. Fish. Res. Inst., Coll. Fish., Univ.
    (Contract No. DACW57-79-M-1594 AND DACW57-    Wash., Seattle, WA, 229 p.
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                                                           Fry, D. H., Jr. 1973. Anadromous fishes of California.
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    Oregon and Washington, 1979-1984.  U.S. Dept.    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
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                                                           D. C. Himsworth (editors), Salmonid ecosystems of the
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    Distribution and origin of pink salmon (Oncorhynchus

                                                       126

    127

       Oncorhynchus keta
        Adults

    Common Name: chum salmon                              oil content of other salmon species.
    Scientific Name: Oncorhynchus keta
    Other Common Names: dog salmon, calico salmon,    Indicator of Environmental Stress: The freshwater,
    chub, fall salmon, keta salmon, le kai salmon (Shiino   estuarine, and early marine life stages are the most
    1976)                                                 sensitive to habitat alterations and pollution (Shepard
    Classification (Robins et al. 1980)                   1981).
    Phylum: Chordata
    Class: Osteichthyes                                   Ecological: The chum salmon is the second most
    Order: Salmoniformes                                  abundant salmonid in the North Pacific region (Forrester
    Family: Salmonidae                                    1981), and has the widest distribution of any Pacific
                                                          salmon (Bakkala 1970).
    Value
    Commercial: The chum salmon is the most important    Range
    Pacific salmon to Japanese commercial fishermen    Overall: In North America, the chum salmon inhabits
    (Forrester 1981), but third in importance to U.S.   coastal streamsfromthe Sacramento River, California
    fishermen (National Marine Fisheries Service 1986).   [occasionally as far south as the San Lorenzo River
    From 1980-84, nearly 43,000 t were landed by U.S.    (Moyle 1976)], northward to the Arctic shore of Alaska
    fishermen and the 1985 catch was worth over $36    (Aro and Shepard 1967, Atkinson et al. 1967, Hallock
    million. This species is commercially fished in North    and Fry 1967). It is found as far east as the Mackenzie
    American waters from Oregon to Alaska. However,    River in Canada. In Asia, the chum salmon is found
    most (75%) are landed in Alaskan waters, with only   south to the Tone River of Chiba Prefecture on the
    Puget Sound, Washington, producing any sizable   Pacific side of Honshu, in Nagasaki Prefecture of
    landings outside of Alaska (Forrester 1981). The chum    Kyushu in the Sea of Japan, and in the Nakdong River
    salmon is captured primarily by fixed or drift gill nets   of the Republic of Korea (Sano 1967, Bakkala 1970).
    and purse seines. It is primarily caught from June to   In Asia most spawning occurs in the lower 100 km of
    September in Alaska, and September to December in   coastal streams.  However, some spawn 2,500 km
    Washington (Forrester 1981 ).                         from the sea in both the Amur River of the U.S.S.R. and
                                                          the Yukon River of Alaska and Canada (Sano 1966,
    Recreational: The chum salmon is not a target sport   Bakkala 1970).  This species' oceanic distribution
    fish in marine waters (Scott and Crossman 1973), but    ranges from the Bering Sea to about lat. 40�N in the
    it is sometimes fished in rivers that have large runs.   western Pacific Ocean and approximately lat. 44�N in
    The marine sport catch is low and is grouped with   the eastern Pacific Ocean (Neave et al. 1976, Fredin et
    sockeye salmon in the reported marine sport catches   al. 1977).
    (Pacific Marine Fisheries Commission 1985, 1986).
    This species does not strike lures or baits as readily as    Within Study Area: The chum salmon is primarily found
    other salmonids and its flesh does not have the desired   in Oregon and Washington, north of the Rogue River,

                                                       128

                                                                                       Chum salmon continued
                                                           from intertidal areas to 2,500 km upriver in large river
      Table 1. Relative abundance of chum salmon           systems (Bakkala 1970), buttheyare normallyfound in
               in 32 U.S. Pacific coast estuaries.         riverine areas less than 200 km from the ocean (Sano
                         Life Stage                        1966). Fry are found in rivers, estuaries, and marine
            Estuary    A S J L E                           waters.  Fry prefer shallow waters (nearshore and
             PugetSound a    :       Relative abundance:   intertidal areas <1.0 m  deep) during their initial
              Hood Canal a   :         �   Highly abundant  outmigration (Bakkala 1970, Healey 1980). Once at
               Skagit Bay *    6           Abundant        sea juveniles are primarily epipelagic (surface to 60 m
             Grays H-arbor I    ( 11  Rare                 depth) (Manzer 1964), but may be found to depths of
              WillapaBay �    *       Blank Notpresent     95 m (LeBrasseur and Barner 1964).  Adults are
           Columbia River O    O                           estuarine and riverine (Bakkala 1970, Fredin et al.
             Nehalem Bay 0    0                            1977).
            Tillamook Bay I S         Life stage:
              NelartsBay 13   �        A-Adults            Substrate: Eggs and alevins are found primarily in
              SiletzRiver0    C- Spawning adults           medium-sized gravel (about 2-4 cm in diameter)
             Yaquina Bay O    O       L - Larvae           (Bakkala 1970, Alaska Department of Fish and Game
              AlseaRiver O    0       E-Eggs               1985) and are buried down to 40 cm (Moyle 1976).
             Siuslaw River 0    0                          Recommended spawning gravel diameters range from
            Umpqua River i                                 1.3-10.2cm (Reiser and Bjornn 1979). Burner(1951)
               Coos Bay 0    o                             found Columbia River redds were composed of 81%
             Rogue River i                                 medium and small gravel (< 15 cm diameter), 13%
            Klamath River n                                large gravel (> 15 cm) and 6% mud-silt-sand. Juveniles
            Humboldt Bay -i                                and adults occur over a variety of substrates.
                Eel River   V/
             Tomales Bay                                   Phvsical/Chemical Characteristics: Best spawning
        Cent. San Fran. Bay '       * Includes Central San
                                     Cent. San Fran. Bay  ncues Centro. Suiansu  temperatures range from 7.2-12.80C, and incubation
        South San Fran. Bay           and San Pabb bays.   temperatures range from4.4-13.3�C (Bell 1984). Eggs
           Elkhorn Slough                                 can survive lower temperatures provided initial
               Morro Bay                                   development has progressed to a stage that is cold-
         Santa Monica Bay                                  water tolerant (Reiser and Bjornn 1979). Incubation
           San Pedro Bay                                  temperatures affect alevin length at hatching (Beacham
             Alamitos Bay                                  and Murray 1987). Optimum temperatures for fry to
             Anaheim Bay
                                                           outmigrate from rivers range from 6.7-13.30C (Bell
             Newport Bay                                   1984). Ocean-dwelling juveniles occur in waters of
             Mission Bay                                  1.0-15.0�C, but prefer2.0-11.0�C. Duringthespawning
            TSuanDiego uBay                                migration, adults migrate upstream at temperatures
                                                           from just above freezing to 21.10C, but optimum
                       A S J L E                           temperatures are 8.3-15.6�C.  The upper lethal
                                                           temperature is 25.6�C, and the lower lethal temperature
    Oregon (Table 1) (Atkinson et al. 1967, Ratti 1979).   is 0.0�C (Bell 1984).  Adults migrate upstream in
    OccasionallysomearefoundintheSacramento River,   velocities up to 2.44 m/sec and successfully spawn in
    California (Hallock and Fry 1967). In the ocean, this   velocities of 46-101 cm/sec (Reiser and Bjornn 1979).
    species can occasionally be found as far south as San    Dissolved oxygen levels below saturation can adversely
    Diego, California (Eschmeyer et al. 1983).             affect swimming performance of adults. Oxygen levels
                                                           above 80% saturation with temporary levels no lower
    Life Mode                                              than 5.0 mg/I are recommended for spawning (Reiser
    The chum salmon is an anadromous species. Eggs    and Bjornn 1979). High concentrations of suspended
    and larvae (alevins) are benthic and infaunal. Young    sediments (1 5.8-54.9g/l) can kill juvenilechum salmon
    juveniles (fry) are benthopelagic, while ocean-dwelling    (Hale et al. 1985). Eggs and alevins are found primarily
    and maturing juveniles (subadults) and adults are   in fresh water, but can tolerate euhaline conditions for
    epipelagic (Sano 1966, Bakkala 1970, Fredin et al.   shortperiods(McNeil1966). Fry showapreferencefor
    1977). Subadults and adults in rivers and streams are   salt water soon after their yolk sac is absorbed and
    bottom-oriented.                                       cannot live for extended periods in fresh water
                                                           (Baggerman 1960, Iwata et al. 1986). A limited
    Habitat                                                residence in a mesohaline (10-15%o) estuarine
    Type: Eggs and alevins occur in rivers and streams,    environment may be needed for complete adaptation

                                                       129

    Chum salmon continued
    to sea water (Iwata and Komatsu 1984). Alevins with   more fang-like teeth, than females (Bakkala 1970). As
    completely-absorbed yolk sacs show abnormal    with other salmonids, the female builds the nest by
    behavior in waters with a pH <6.0 (Rombough 1983).   turning on her side and excavating the nest by fanning
                                                          the streambed with her caudal fin (Bakkala 1970).
    Miarations and Movements: The chum salmon is highly   During spawning, the male and female will settle into
    migratory.  Fry migrate seaward immediately after   the nest and quiver with mouths agape as they release
    emerging from the redd, although some may reside in   eggs and milt (Scott and Crossman 1973). After laying
    fresh waterfor several months (Simenstad et al. 1982).   the eggs, the female covers them by digging upstream.
    They migrate primarily at night in small rivers and    Thisprocesscontinuesuntilthefemaleisspent. Males
    sometimes during daylight in larger rivers (Bakkala    may spawn with more than one female; both sexes are
    1970). Juveniles are typically 30-55 cm long when    aggressiveonthespawninggrounds. An average redd
    they enter estuaries (March to mid-May), however    is2.8m2(ReiserandBjornn 1979). Afemalewillguard
    some may be larger if the migration is long (Moyle    her redd as long as she is able before dying. Some
    1976). Once juveniles enter estuaries, their migration   adults may spend less than a week in fresh water if they
    typically slows and many will rear for up to several   arrive sexually mature (Scott and Crossman 1973).
    months inthe estuary (Healey 1982, Levy and Northcote
    1982, Simenstad et al. 1982).  Increasing salinities   Fecundity:Largefemalescanreleaseover4,000eggs,
    prompt schooling behavior(Shelboun 1966). Juveniles   but on average 2,400-3,000 eggs are laid per female
    occur in Washington estuaries from January to July,   (Scott and Crossman 1973). Late-run southern stocks
    peaking from late March to mid-May.  Most chum    are more fecund than early-run stocks (Sano 1966,
    salmon leave Oregon estuaries by mid-May (Myers    Bakkala 1970). This may be a function of different body
    1980). Chum salmon juveniles move in and out of tidal   sizes between the stocks.
    creeks, sloughs, marsh habitats, and intertidal areas
    as the tide fluctuates (Mason 1974, Healey 1982).    Growth and Development
    Besides this daily tidal movement, there is a general    Eaa Size and Embrvonic Develooment: Eggs are
    movement seaward as the juveniles grow (Healey    reportedtobe6.0-9.5mmindiameterafterfertilization
    1982). Individuals may spend 4-32 days in estuaries;   (Bakkala 1970, Bell 1984). Embryonic development is
    residency varies seasonally.  In some stocks, early   indirect and external.  Eggs require from 0.5 to 4.5
    migrants may reside longerthan later migrants while in   monthsto hatch (depending on temperature). Hatching
    other stocks, the opposite is true (Healey 1979,   usually occurs from December to February (McPhail
    Simenstad et al. 1982, Kaeriyama 1986). Most chum    and Lindsey 1970, Scott and Crossman 1973, Pauley
    salmon move offshore from April to June when they are   et al. 1988).
    80-100 mm in fork length (Healey 1982). Once in the
    ocean, migrating chum salmon head north, but stay   Aae and Size of Larvae: Alevins absorb their yolk-sac
    alongthecontinentalshelf until fall, whentheydisperse   in 30-50 days, depending on temperatures (Wydoski
    out into the Gulf of Alaska (Hartt and Dell 1986) and mix    and Whitney 1979). Alevins are 20.0-24.0 mm long at
    with other salmon species and other age groups of   hatching (Bakkala 1970, Kaeriyama 1986, Beacham
    chum salmon. Some chum salmon do not appear to   and Murray 1987) and grow to 30.0-35.0 mm before
    migrate out of Puget Sound (Hartt and Dell 1986).    leaving the gravel (Moyle 1976, Wydoski and Whitney
    Immature fish move about 28 km/day, while maturing    1979).
    fish average 35 km/day (Neave et al. 1976). Immature
    fish aretemperature sensitive and move south in winter   Juvenile Size Ranae: Fry in fresh water are 30.0-70.0
    and north in summer (Neave et al. 1976).              mm long, depending on the distance between the
                                                          estuary and spawning grounds (Scott and Crossman
    Reproduction                                          1973). Growth in estuaries and the ocean is rapid; by
    Mode: The chum salmon is gonochoristic, oviparous,   the end of their first year at sea juveniles will average
    and semelparous (all adults die soon after spawning)    over 30.0 cm in length and after five years will be 50.0
    (Bakkala 1970). Eggs are fertilized externally.       cm long (Fredin et al. 1977).

    Matina/SDawnina: Two spawning populations exist; a   Aae and Size of Adults: Adults return to spawn at 2-7
    northern stock that spawns from June to September,    years of age (primarily 3-5 years) (Scott and Crossman
    and a southern (late-run) stock that spawns from    1973).  Bell (1984) determined that chum salmon
    August to January (Sano 1966, Bakkala 1970).    average 63.5 cm in length and 4.0 kg at maturity, but
    Washington, Oregon, and California stocks are all late-   Squire and Smith (1977) reported that they can grow
    run fish. Chum salmon are sexually dimorphic when    up to 107 cm in length and their average weight is 4.5-
    mature; males have a hooked snout, a slight hump, and    5.3 kg at maturity.

                                                       130

                                                                                       Chum salmon continued
    Food and Feeding                                      Factors Influencina Pooulations: To augment natural
    TroDhic Mode: Larvae feed on theiryolk. Juveniles and    production, chum salmon are produced by hatcheries
    adults are carnivores and "opportunistic" feeders.    in Oregon, Washington, Alaska, Canada, U.S.S.R.,
                                                          and Japan (Atkinson et al. 1967, Sano 1967). Over
    Food Items: Fry may not feed in fresh water if their   23.7 million juveniles were released from hatcheries
    migration to estuaries is short. However, if freshwater    along the Pacific coast in 1976 (Wahle and Smith
    residency is lengthy, fry will feed on aquatic and    1979). However, in1987, over90 millionchumfrywere
    terrestrial insects and small crustaceans. Chironomid    released just in Washington (Abrahamson 1988). In
    larvae appearto be particularly important to fry in fresh   Japan, over 2 billion fry are released from hatcheries
    water (Sano 1966, Bakkala 1970,Scott and Crossman    annually (Kaeriyama 1989). Most natural mortality
    1973). Feeding innearshore marine areas and estuaries    occurs in fresh water during the embryonic stage as a
    by fry and fingerlings appears to be an important   resultofpoorenvironmentalconditionssuchassiltation,
    component of chum salmon life history (Healey 1980,    low dissolved oxygen, spawning gravel disruptions,
    Simenstad 1983).  Initially juveniles feed in shallow    and freezing (McNeil 1966, Wydoski and Whitney
    waters and concentrate on epibenthic prey such as    1979).  Beacham and Starr (1982) concluded that
    harpacticoid copepods and gammarid amphipods, but   freshwater survival in Canada's Fraser Riverwas mostly
    they may also eat terrestrial insects and other small   a function of interactions among temperature, rainfall,
    crustacea (Sibert et al. 1977, Healey 1979, Simenstad    and egg abundance. Human alterations of freshwater
    and Salo 1982, Kaeriyama 1986). Young chum salmon    habitat caused by improper logging practices,
    are size-selectivefeeders (Fellerand Kaczynski 1975).    hydroelectric  and  irrigation  developments,
    Food limitation in shallow waters may induce movement    channelization, chemical and pollutant introductions,
    to deeper waters (Healey 1980, Simenstad and Salo    and other factors, can lower chum salmon production
    1982) where juvenile chum salmon shift their diets to   (Bottom et al. 1985, Holtby and Scrivener 1989). High
    include more pelagic prey, such as calanoid copepods,    rivertemperatures affect chum salmon migrations, rate
    hyperiid amphipods, crustacean larvae, and larvaceans   of maturation, cause direct mortality, and increase the
    (Freshetal. 1981,SimenstadandSalo1982, Kaeriyama    incidence of diseases (Hale et al. 1985). Survival of
    1986). In the ocean, juveniles and subadults feed on    chum salmon eggs iscorrelatedwiththepermeabiltyof
    euphausiids,  squids,  pteropods,  and  fishes   the redd to water flow (Pauley et al. 1988). Besides
    (Andrievskaya 1957, Allen and Aron 1958, LeBrasseur   their initial freshwater residency, early estuarine and
    1966, Peterson et al. 1982, Pearcy et al. 1984).      marine residence appears to be a critical period for
                                                          chum salmon and can affect the eventual number of
    Biological Interactions                               returning adults (Bakkala 1970, Bax 1983). Bax (1983)
    Predation: In freshwaterand estuarine environments,    showed that chum salmon in Puget Sound can have
    this species' primary predators are probably other   high early marine mortality. Parker (1971) suggested
    salmonids. Chum salmon fry are reportedly eaten by   that chum salmon fry must "outgrow" their marine
    juvenile coho (0. kisutch), sockeye (0. nerka), and    predators. Streamtemperaturesaffectfryemergence
    chinook salmon (0. tshawytscha), cutthroat (0. clark,)   and migration, and maypromptsynchronizedemigration
    and rainbow trout (0. mykiss), Dolly Varden (Salvelinus   during "windows of opportunity" (Holtby et al. 1989).
    malma), scu Ipins, Pacific cod (Gadus macrocephalus),    There also appears to be adverse interactions between
    and birds [belted kingfisher (Megaceryle alcyon),   pink salmon (0. gorbuscha) and chum salmon, based
    merganser (Merginae), and others] (Bakkala 1970,    on fewer chum salmon returning to spawn in years
    Scott and Crossman 1973, Bax et al. 1980, Fresh    when pink salmon are abundant (Ames 1983, Fresh
    1984, Nagata and Miyamota 1986). Predation rates   1984).  Beacham and Starr (1982) suggested that
    are variable, depending on such factors as predator   competition between chum and pink salmon in the
    and prey size, the alevin's amount of yolk, abundance    Fraser River estuary or Strait of Georgia reduces
    of fry, and composition of other prey (Hunter 1959,    eventualadultchum salmon abundance. Andrievskaya
    Fresh and Schroeder 1987). At sea, juveniles are    (1970) found that in years of low pink salmon abundance,
    preyed on by lamprey, shark, and probably other large   chum and pink salmon in the ocean eat similar prey.
    predatoryfishes. Subadultandadultchumsalmonare    But in years of high pink salmon abundance, chum
    eaten by killer whales (Orcinus orca), harbor seals    salmon consume different prey. Fishing pressure also
    (Phoca vitulina), and other marine mammals (Fiscus   affects abundance. The Japanese high seas salmon
    1980). Bears and large predatorybirds such asosprey    fishing fleets and an unrestricted squid gillnet fishery
    (Pandion haliaetus)  and bald eagles (Haliaeetus   take an unknown bycatch of chum salmon from North
    leucocephalus) prey on spawning adults (Scott and    America.
    Crossman 1973).

                                                       131

    Chum salmon continued
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                                                       132

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    North Pacific, p. 203-229, Oregon State Univ. Press,    Manzer, J. I. 1964. Preliminary observations on the
    Corvallis, OR.                                        vertical distribution of Pacific salmon (genus
                                                          Oncorhynchus) in the Gulf of Alaska. J. Fish. Res.
    Healey, M. C.  1982.  Juvenile Pacific salmon in   Board Can. 21(5):891-903.
    estuaries: the life support system. In V. S. Kennedy

                                                       133

    Chum salmon continued
    Mason, J. L. 1974. Behavior ecology of chum salmon    Biol. Rep. 82(11.81). U.S. Army Corps Engin., TR EL-
    fry (Oncorhynchus keta) in a small estuary. J. Fish.   82-4, 17 p.
    Res. Board Can. 31:83-92.
                                                           Pearcy, W., T. Nishiyama, T. Fujii, and K. Masuda.
    McNeil, W. J.  1966.  Effect of the spawning bed    1984. Diel variations in the feeding habits of Pacific
    environment on reproduction of pink and chum salmon.    salmon caught in gill nets during a 24-hour period in the
    Fish. Bull., U.S. 65(2):495-523.                       Gulf of Alaska. Fish. Bull., U.S. 82(2):391-399.

    McPhail, J. D., and C. C. Lindsey. 1970. Freshwater    Peterson, W. T., R. D. Brodeur, and W. G. Pearcy.
    fishes of northwestern Canada and Alaska. Fish. Res.    1982. Food habits of juvenile salmon in the Oregon
    Board Can., Bull. No. 173, 381 p.                       coastal zone, June 1979. Fish. Bull., U.S. 80(4):841-
                                                           851.
    Moyle, P. B. 1976. Inland fishes of California. Univ.
    Calif. Press, Berkeley, CA, 405 p.                      Ratti, F. 1979. Natural resources of Rogue estuary.
                                                           Oregon Dept. Fish Wildl., Portland, OR, 33 p.
    Myers, K. W. W. 1980. An investigation of the
    utilizationof fourstudy areas in Yaquina Bay, Oregon,    Reiser, D. W., and T. C. Bjornn.  1979.  1. Habitat
    by hatchery and wild juvenile salmonids. MS. Thesis,    requirements of anadromous salmonids.  In W. R.
    Oregon State Univ., Corvallis, OR, 234 p.               Meehan (editor), Influence of forest and rangeland
                                                           management on anadromous fish habitat in the western
    Nagata, M.,and M. Miyamoto. 1986. The downstream    United States and Canada, p. 1-54.  USDA Forest
    migration of chum salmon fry, Oncorhynchus keta,   Service, Gen. Tech. Rep. PNW-96, Pacific Northw.
    released into the Utabetsu River of eastern Hidaka in   Forest Range Exp. Sta., Portland, Oregon.
    Hokkaido, and the estimation of predation amount of
    the fry byfreshwatersculpin, Cottus nozawae, Synder.    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Sci. Rep. Hokkaido Fish Hatchery 41:13-22.              E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
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    National Marine Fisheries Service. 1986. Fisheries of    United States and Canada. Am. Fish. Soc. Spec. Publ.
    the United States, 1985. Current Fishery Statistics No.    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    8368. U.S. Dept. Comm., NOAA, Nat. Mar. Fish Serv.,
    Nat. Fish. Stat. Prog., Washington, D.C., 122 p.        Rombough, P. J. 1983. Effects of low pH on eyed
                                                           embryos and alevins of Pacific salmon. Can. J. Fish.
    Neave, F., T. Yonemori, and R. G. Bakkala. 1976.    Aquat. Sci. 40(10):1575-1582.
    Distribution and origin of chum salmon in offshore
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    Pacific Marine Fisheries Commission. 1985.  37th    Sano, S. 1967. Spawning populations of North Pacific
    annual report of the Pacific Marine Fisheries    salmon, 3. Chum salmon in the Far East. Internat.
    Commissionfortheyear 1984. Pac. Mar. Fish. Comm.,    North Pac. Fish. Comm. Bull. No. 23: 23-41.
    Portland, OR, 35 p.
                                                           Scott, W. B., and E. J. Crossman. 1973. Freshwater
    Pacific Marine Fisheries Commission. 1986.  38th    fishes of Canada. Fish. Res. Board Can., Bull. No. 84,
    annual report of the Pacific Marine Fisheries    966 p.
    Commission forthe year 1985. Pac. Mar. Fish. Comm.,
    Portland, OR, 36 p.                                     Shelboun, J. E.  1966.  Influence of temperature,
                                                           salinity, and photoperiod on the aggregations of chum
    Parker, R. R. 1971. Size selective predation among    salmon fry. J. Fish. Res. Board Can. 23:293-304.
    juvenile salmonid fishes in a British Columbia inlet. J.
    Fish. Res. Board Can. 28(10):1503-1510.                 Shepard, M. F.  1981.  Status and review of the
                                                           knowledge pertaining to the estuarine habitat
    Pauley, G. G., K. L. Bowers, and G. L. Thomas. 1988.    requirements and life history of chum and chinook
    Species profiles: life histories and environmental    salmon juveniles in Puget Sound. Final Rep. to Wash.
    requirements of coastal fishes and invertebrates (Pacific    Coop. Fish. Res. Unit, College Fish., Univ. Wash.,
    Northwest) -  chum salmon.  U.S. Fish Wildl. Serv.    Seattle, WA, 113 p.

                                                       134

                                                                                       Chum salmon continued
    Shiino, S. 1976. List of common names of fishes of the
    world, those prevailing among English-speaking
    Nations. Sci. Rep. Shima Marineland No. 4,
    Kashikojima, Shima, Mie, Japan, 262 p.

    Sibert, J., T. J. Brown, M. C. Healey, and B. A. Kask.
    1977. Detritus-based food webs: exploitation by juvenile
    chum salmon (Oncorhynchus keta). Science
    196(4290) :649-650.

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    Simenstad, C. A., and E. O. Salo. 1982. Foraging
    success as a determinant of estuarine and nearshore
    carrying capacity of juvenile chum salmon
    (Oncorhynchus keta) in Hood Canal, Washington. In
    B. R. Melteff and R. A. Neve (editors), Proceedings of
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    descriptive account of Pacific coast anadromous
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    Wydoski, R. S. and R. R. Whitney. 1979. Inland fishes
    of Washington, Univ. Wash. Press, Seattle, WA, 220 p.

                                                       135

       Oncorhynchus kisutch
        Adults

    Common Name: coho salmon                              Recreational:The coho salmon isthe primarytargetfor
    Scientific Name: Oncorhynchus kisutch                many marine and freshwater sport fishermen on the
    Other Common Names: silver salmon, blueback    Pacific coast. A total of 674,000 fish (not including
    salmon, hookbill, hooknose salmon, hoopid salmon,    freshwater catch) were caught by sport anglers off
    jack salmon, medium red salmon, salmon trout,   California, Oregon, Washington, and Alaska in 1984
    siverside salmon, white salmon (Scott and Crossman    (Pacific Marine Fisheries Commission 1986). Sport
    1973, Shiino 1976, Laufle et al. 1986)               caught coho salmon originating from the Columbia
    Classification (Robins et al. 1980)                   River were estimated to be worth over $30 million
    Phylum: Chordata                                     (Richards 1986). Most coho salmon are caught by
    Class: Osteichthyes                                  trolling (in ocean and estuaries), but they are also taken
    Order: Salmoniformes                                 by spin casting and fly-fishing. It is a highly-esteemed
    Family: Salmonidae                                   sport fish because of its abundance, availability, size,
                                                         fighting ability, and excellent taste. This species was
    Value                                                 introduced into the Great Lakes and is now very
    Commercial: The coho salmon is fished commercially    abundant there (Morrow 1980).
    from Norton Sound, Alaska, south to northern Japan,
    andalongwestem NorthAmericato northern California.   Indicator of Environmental Stress: Reduced run sizes
    It is also fished on the high seas (International North   are often the result of adverse environmental and
    Pacific Fishery Mangement Council 1979).  Coho    habitat changes.  Coho salmon exposed to low
    salmon make up 8-11% of the total Pacific salmonid    concentrations of aromatic hydrocarbons decrease
    catch (Forrester 1982, Takehama 1983). This species   feeding, while fish exposed to high concentrations may
    is usuallyrankedfourth in commercial catches (numbers    stop feeding for days (Purdy 1989).  See "Factors
    and weight) of salmonids [behind pink (Oncorhynchus    Influencing Populations".
    gorbuscha), chum (O. keta), and sockeye salmon (O.
    nerka)]. An average of 19,500 t were landed in the    Ecological: The coho salmon is a common species in
    United States from 1980-1984 (National Marine    many coastal streams (Atkinson et al. 1967). Stream-
    Fisheries Service 1986). The 1985 commercial catch   dwelling juveniles are territorial (Shapovalov and Taft
    was worth approximately $46 million (National Marine    1954, Steine et al. 1972) and sometimes prey on other
    Fisheries Service 1986). It is commerciallycaughtwith   salmonids (Fresh and Schroeder 1987). Adults and
    gill nets (drift and set), purse seines, reef nets, and   juveniles are common in neritic waters off Oregon and
    trolling (primary method). Some fish are canned, but   Washington (Fisher et al. 1983, Fisher and Pearcy
    most are sold fresh or fresh-frozen for human    1985).
    consumption. About 75% of the U.S. catch comesfrom
    Alaska and is harvested primarily during July and    Range
    August. Native Americans are allocated 50% of the   Overall: The coho salmon spawns in coastal streams
    coho salmon harvest in Washington (Clark 1985).       from northern Japan to the Anadyr River in Siberia and

                                                      136

                                                                                         Coho salmon continued
                                                            into smolts (juveniles that migrate to the ocean). Smolts
      Table 1. Relative abundance of coho salmon            and ocean-dwelling and maturing juveniles (subadults)
               in 32 U.S. Pacific coast estuaries.          and adults are primarily pelagic (Shapovalov and Taft
                         Life Stage                        1954). Subadults and adults in rivers and streams are
            Estuary    A S J L E                           bottom-oriented.
             Puget Sound IV   I S    Relative abundance:
              Hood Canal �3 �  0           Highly abundant  Habitat
               SkagitBay �  �          @   Abundant        Iype: Eggs, alevins, fry, and parr are riverine. Eggs
             Grays Harbor Xi   0       0   Common          and alevins occur primarily in riffle areas of streams.
              Willapa Bay   1         Blank Not present    Fry inhabit shallow stream areas adjacent to pools, but
            Columbia River f1    a                          move into deeper waters as they grow (Shapovalov
             NehalemBay � I                                 and Taft 1954, Moyle 1976). Smolts are found in rivers,
             Tillamook Bay IS   I      Life stage:          estuaries, and nearshore coastal waters. In estuaries,
              NetansBay C    O         A-Adults            smolts occur in intertidal and pelagic habitats
              Siletz River            SSpawning adults     (Simenstad and Eggers 1981, Durkin 1982, Myers and
                               � tivJ - Juveniles
              YaquinaBay �    *        L-Larvae             Horton 1982), with deep, marine-influenced habitats
              Alsea River             E-Eggs               often preferred (Macdonald et al. 1987). Smolts are
             Siuslaw River  ] �                             epipelagic in offshore marine waters (Milleret al. 1983).
            UmpquaRiver    �                               Subadults range from neritic to oceanic (Hartt and Dell
                Coos Bay (                                  1986). Adults are estuarine and riverine.
              Rogue River �  �
             Klamath River O  Substrate: Eggs are buried in areas that are composed
             Humboldt Bay 0    0                            of gravel ranging from 1.3-10.2 cm in diameter (Reiser
                Eel River 0     0                          and Bjornn 1979, Bell 1984). Coho salmon arethe only
             Tomales Bay 0    O                            salmon whose redd can contain upto 10% mud (Burner
        Cent. San Fran. Bay  *        Includes Central San
                                      Francisco. Suisun,   1951 ). Juveniles in streams are not substrate selective,
        South San Fran. Bay           and San Pablo bays.   but prefer areas with good cover and food availability.
            Elkhorn Slough                                  Smolts, subadults, and adults can be found migrating
               Morro Bay                                   over a wide range of substrates (mudflats to rocks).
          Santa Monica Bay
             San Pedro Bay                                 Physical/Chemical Characteristics: The coho salmon
                     Alaheito Bay                          is found in fresh water to euhaline waters.  Eggs,
             ANewpom Bay                                   alevins, fry, and parr occur in fresh water. Smolts and
              Mission Bay                                  adults are euryhaline. Eggs and alevins are found in
            San Diego Bay                                   waters ranging from 4.4-21.0�C (Bell 1984), but 4.4-
            Tijuana Estuary                                 13.3�C is best for egg incubation (Reiser and Bjornn
                               A S J L E                    1979). Juveniles prefer stream temperatures of 11.8-
                                                            14.6�C, with 25.1�C the upper lethal limit (Brett 1952).
                                                            Growth ceases above 20.3�C because of increased
    from northern Monterey Bay, California, to Point Hope,    metabolic rate (Bell 1984).  However, other water
    Alaska (Moyle 1976). In the ocean, it occurs in coastal    quality parameters can lower this upper thermal limit
    waters from Baja California to the Bering Sea (Hart    (Ebel et al. 1971). Water temperature can also affect
    1973, Hartt and Dell 1986).                             juvenile osmoregulatory ability (Zaugg and McLain
                                                            1976). At sea, most coho salmon are found in waters
    Within Study Area: This species occurs in all estuaries    that are 4.0-15.2�C. (Godfrey et al. 1975, Fredin et al.
    north of Monterey Bay, California, to Puget Sound,    1977). Adults can migrate upstream in velocities up to
    Washington (Table 1). It is very rare in San Francisco    2.44 m/sec; juveniles prefer stream velocities of 0.09-
    Bay(strays). Major U.S. spawninggrounds(otherthan    0.46 m/sec depending on the habitat (Reiser and
    Alaska) are in Washington and Oregon (Atkinson et al.   Bjornn 1979). Adequate stream cover is important to
    1967).                                                  freshwater life stages.  Juveniles and eggs require
                                                            well-oxygenatedwaters. Dissolvedoxygen (DO) levels
    Life Mode                                               below 8 mg/I sharply reduce embryo survival (Phillips
    Thecohosalmonisananadromousspecies. Eggsand    and Campbell 1968) and DO levels below 4 mg/I
    larvae (alevins) are benthic and infaunal.  Young    reduce juvenile food consumption, food conversion,
    juveniles (fryand parr) are benthopelagic. Parrbecome    and growth (Herrmann et al. 1962). Low pH (below
    pelagic and acquire a silver color when they transform    5.01)can be lethal to newly-hatched alevins (Rombough

                                                        137

    Coho salmon continued
    1983). Adults need a minimum depth of 18 cm to   salmon can migrate up to 30 km/day (Godfrey et al.
    migrate and spawn (Thompson 1972).  Short-term    1975). Ocean migration appears to involve the use of
    pulses of suspended sediment in streams can cause a    magnetic information, celestial cues, and polarized
    breakdown of social organization, a change in   light. Olfaction appears to be the dominant guidance
    aggressive behavior, an increase in activity, and a    mechanism during the riverine (spawning) migration
    decrease in feeding ability (Berg 1982). High turbidity   (Brannon 1982, Quinn 1982, Hasler and Scholz 1983).
    can affect emergence and growth of young coho salmon
    (Sigler et al. 1984) and also alters feeding habits   Reproduction
    (Reiser and Bjornn 1979).                            Mode: The coho salmon is gonochoristic, oviparous,
                                                          and semelparous (all adults die after spawning). Eggs
    Miarations and Movements: Over their range, adult   are fertilized externally.
    coho salmon can be found to migrate into their natal
    streams from June to February and spawn from    Matina/Snawnina: Spawning occurs from September
    September through March (Washington 1982). Fry   to March (depending on location).  Peak spawning
    initially live and school in shallow gravel areas, but   occurs from September to February in the Columbia
    soondisperseupstreamanddownstreamandtodeeper    River (Netboy 1980) and November to January in
    waters as they grow. Fry may be displaced downstream    California (Moyle 1976). This species typically spawns
    by fall freshets. Fry may entertributaries, sloughs, and    in small streams (sometimes in large rivers) within 240
    side channels to overwinter, and return to the    km of the river mouth (Laufle et al. 1986). Although
    mainstream in spring (Tschaplinski and Hartman 1982).    coho salmon may spawn in the same habitats as
    After residing approximately one year in fresh water   chinook salmon (Burner 1951), it normally spawns in
    (two or more in northern streams) most juveniles will   areas that have lower stream velocities, shallower
    migrate to the ocean (outmigration) (Gribanov 1948,    depths, and smaller gravel (Fraser et al. 1982). The
    Godfrey 1965). Mostjuvenilesoutmigratefrom Aprilto   coho salmon typically spawns in riffle areas where
    August, peaking in May (Shapovalov and Taft 1954,   water velocities are 0.08-0.70 m/sec, stream depths
    Deschamps et al. 1971, Simenstad and Eggers 1981,   are 0.05-0.66 m, substrate gravel ranges from 2-15 cm
    Myers and Horton 1982, Dawley et al. 1986).   in diameter, and water temperatures are 4-140�C
    Outmigration has been reported to occur at night   (Schmidt et al. 1979). Spawning adults are dimorphic.
    (McDonald 1960) and day (Durkin 1982, Dawley et al.   Males have a thick, hooked snout, exposed teeth, and
    1986). Migrating smolts are approximately 8.8-13.8   change color, while females change little (Scott and
    cm long (Salo and Bayliff 1958, Durkin 1982), with   Crossman 1973). Females select and build the redds
    larger smolts migrating sooner than smaller smolts    and both sexes areterritorial. Adominant (larger) male
    (Durkin 1982). Limited estuarine rearing occurs in the    moves into the nest and spawns with the female when
    Columbia River estuary (Dawley et al. 1986). However,    ready. At this time subdominant males may dart in and
    in Puget Sound, residency forcoho salmon smolts was    release sperm (Scott and Crossman 1973). Females
    estimated to be 6-40 days, with 3-5% of the naturally-   will spawn in uptofourdifferent nests and with different
    produced yearling coho salmon residing inside the    males.   Eggs are covered by the digging and
    Strait of Juan de Fuca until maturity (Simenstad et al.   displacement of gravel upstream (Scott and Crossman
    1982).  In Yaquina Bay, Oregon, a few overwinter   1973). Redds average 2.9 m2 (Burner 1951),with eggs
    within and nearthe bay, but most juveniles migrate out   buried an average of 22.0 cm deep (Gribanov 1948).
    of the bay in 2-9 days (Myers and Horton 1982). Some
    coho salmon fry in Canada may rear in estuaries from    Fecundity: In North America, a coho salmon female
    March to October or November (Tschaplinski 1982).    can lay 1,000-5,700 eggs (depending on size) (Scott
    Once in the ocean, smolts from Oregon and coastal   and Crossman 1973, Moyle 1976). Average fecundity
    Washington rivers appear to initially head south, but   is about 2,500-3,500 eggs per female (Rounsefell
    later head north (Pearcy 1984). Most Oregon coho    1957, Crone and Bond 1976, Wydoski and Whitney
    salmon probably remain in coastal waters off California,    1979). In Kamchatka, U.S.S.R., the average is about
    Oregon,and Washington (Parmenter and Bailey 1985,    5,000 eggs per female (Gribanov 1948).
    Pearcy and Fisher 1988). However, during the first
    summer some may make extensive migrations to the   Growth and Development
    Gulf of Alaska  (Hartt and Dell 1986), but by their   Eca Size and Embrvonic Develooment: This species'
    second summer, many will be captured by sport and    egg is relatively large and second only to the chinook
    commercial fisheries near their river of origin (Wright   salmon's in size (Rounsefell 1957). In Canada, coho
    1968). Both juveniles and adults stay nearthe surface   salmon eggs have a diameter of 4.5-6.0 mm (McPhail
    (within 10 m), except when the sea is covered by a layer   and Lindsey 1970), but are reported to be 6.6-7.9 mm
    of warm water (Fredin et al. 1977). Maturing coho    in diameter in the U.S. (Bell 1984).  Embryonic

                                                      138

                                                                                      Coho salmon continued
    development is indirect and external. Eggs hatch in 38    1941, Ito 1964, Scott and Crossman 1973, Fresh et al.
    days at 11�C, 48 days at 90C, and 86-101 days at 4.5�C    1981). An opportunistic feeder, the coho salmon's diet
    (Laufle et al. 1986).                                 differs spatially and temporally, and probably reflects
                                                          relative prey availability (Prakash 1962, Brodeur et al.
    Aae and Size of Larvae: Larvae (alevins) are 17-19 mm    1987).
    long at hatching and growto 27-30 mm in length before
    the yolk sac is absorbed (Gribanov 1948). It takes    Biological Interactions
    about 2-5 weeks (depending on temperature) before    Predation: In fresh water, juveniles are eaten by other
    larvae absorb the yolk sac and leave the gravel   fishes, including coho salmon smolts, cutthroat trout
    (Gribanov 1948, Laufle et al. 1986).                  (0. clarki), rainbow trout (0. mykiss), Dolly Varden
                                                          (Salvelinus malma), squawfish (Ptychocheilus
    Juvenile Size Ranae: Juveniles range from 3 cm to at   oregonensis), and sculpins (Scott and Crossman 1973).
    least 40 cm long (Gribanov 1948).                     Marine fish predators include spiny dogfish (Squalus
                                                          acanthias) and other sharks. Juveniles are also eaten
    Aae and Size of Adults: Most coho salmon mature and    by birds such as mergansers, belted kingfishers
    spawn during their 3rd year, but some mature as 2-5    (Megaceryle alcyon), loons (Gavia spp.), gulls, and
    year-olds (Scott and Crossman 1973, Moyle 1976).    common murres (Uria aalge) (Scott and Crossman
    Two-year-old mature males that have spent only one    1973, Varoujean and Matthews 1983).  Marine
    year in the ocean are call "jacks". Off Oregon and    mammals such as harbor seals (Phoca vitulina),
    Washington, "jack" abundance is a good predictor of   northern and California sea lions (Eumetropias tubata
    nextyear'sthree-year-oldcohosalmonabundance. In   and Zalophus californianus, respectively), and killer
    the Fraser River, Canada,  the coho salmon run is   whales(Orcusorcinus) will also eatcoho salmon. Most
    usually composed of 92% three-year-olds, 4% four-   marine mammal predation occurs in nearshore,
    year-olds, and 4% "jacks" (Fraser et al. 1982). Adults    estuarine and river areas (Fiscus 1980, Beach et al.
    range from 40-99 cm in length (Gribanov 1948, Kessler    1981). On their spawning run, coho salmon are taken
    1985).                                                by bears and other mammals, bald eagles (Haliaeetus
                                                          leucocephalus), and osprey ( Pandion haliaetus).
    Food and Feeding
    Trophic Mode: Larvae feed on their yolk. Juveniles and    Factors Influencino Populations: Freshwater mortality
    subadults are carnivorous, "opportunistic" feeders.   is high, with only 0.13-12.0% survival from egg to age
                                                           1 smolt expected (Fredin et al. 1977). This mortality is
    Food Items: Once fry emerge they begin feeding on a    related to habitat suitability and alteration, disease,
    variety of terrestrial and aquatic invertebrates (spiders,   predation, disruption of eggs and larvae, siltation, food
    mites, insects, snails, etc.) (Shapovalovand Taft 1954,    abundance, and competition with other fishes
    Scott and Crossman 1973). Parr may eat invertebrates    (Chapman 1966, Steine et al. 1972, Fredin et al. 1977,
    and other salmon (Roos 1960, Fresh and Schroeder    Reiser and Bjornn 1979). Man-induced changes to
    1987). In reservoirs, parr feed on zooplankton (e.g.,   streams by improper logging, road construction,
    Daphnia), insects, and amphipods (Wydoski and    irrigation, pollutants, dams and reservoirconstruction,
    Whitney 1979, Muir and Emmett 1988). In estuaries,    channelization, residential development,  and
    they feed primarily on large planktonic or small nektonic    agricultural practices can cause physical and chemical
    animals, such as amphipods ( Corophium spp.,   alterations which may be detrimental to coho salmon
    Eogammarus spp.), insects, mysids, decapod larvae,   production (Reiser and Bjornn 1979, Laufle et al. 1986,
    and larval and juvenilefishes (including othersalmonids)    Scrivener and Brownlee 1989). Summer streamflow
    (Levy and Levings 1978, Fresh et al. 1979, Simenstad    affects survival and is an important determinant of
    and Eggers 1981, Durkin 1982, Pearce et al. 1982).    Puget Sound coho salmon runs (Mathews and Olson
    Initially, ocean-dwelling coho salmon eat decapod    1980). Valley tributaries and sloughs may be important
    larvae, gammarid and hyperid amphipods, euphausiids,    for winter survival for many coho salmon juveniles
    terrestrial insects, copepods, cephalopods, Cnideria,    (Tschaplinski and Hartman 1982). Marine mortality
    gastropods (Limacina helicina), planktonic annelids,    can also be high; Lander and Henry (1973) estimated
    and larval and juvenile fishes (Peterson et al. 1983,    that only 5-6% of Columbia River smolts survived after
    Emmett et al. 1986, Brodeur et al. 1987, Brodeur    13.5monthsatsea. Year-classstrengthappearstobe
    1989).  As they grow, juveniles become more    determined very early in ocean residence and may be
    piscivorous, eating northern anchovy (Engraulis   related to predation rates (Fisher and Pearcy 1988).
    mordax), Pacific herring (Clupeapallasl), Pacific sardine    Ricker (1976) estimated that the offshore troll fishery
    (Sardinops sagax), juvenile scorpaenids, capelin   kills one coho salmon (below legal size) for every two
    (Mallotus villosus), and other fish species (Silliman   landed. Coho salmon abundance has beencorrelated

                                                       139

    Coho salmon continued
    with ocean "upwelling" one year earlier (Gonsolus    Bilton, H. T., D. F. Alderdice, and J. T. Schnute. 1982.
    1978). The Oregon Production Area coho salmon    Influence of time and size at release of juvenile coho
    population has gone from predominantly high-survival    salmon (Oncorhynchus kisutch) on returns at maturity.
    wild fish to predominantly low-survival hatchery fish   Can. J. Fish. Aquat. Sci. 39:426-447.
    (Nickelson 1986). Over 62 million hatchery smolts
    werereleasedintheOregon Production Area (Monterey    Brannon, E. L.  1982.  Orientation mechanisms of
    Bay, California to Leadbetter Point, Washington) in   homing salmonids. In E. L. Brannon and E. O. Salo
    1981, including 24 million from private hatcheries   (editors), Proceedingsofthesalmonandtroutmigratory
    (Nickelson 1986). Hatcheries (private and public) play   behavior symposium, p. 219-227. School Fish., Univ.
    a dominant role in the abundance of this species in the   Wash, Seattle, WA.
    Pacific Northwest. However, the introduction of hatchery
    coho salmon presmolts into streams appears to reduce    Brett. J. R. 1952. Temperature tolerance in young
    wild coho salmon populations (Nickelson et al. 1986).   Pacific salmon, genus Oncorhynchus. J. Fish. Res.
    Hatcheries may also precipitate overharvest of wild   Board Can. 9(6):265-323.
    stocks and cause density-dependent mortality in both
    freshwater and marine environments (Lichatowich and    Brodeur, R. D. 1989. Neustonic feeding by juvenile
    McIntyre 1987). Coho salmon smolts may need to   salmonids in coastal waters of the northeast Pacific.
    reach a "critical size" for proper smoltification and    Can. J. Zool. 67:1995-2007.
    marine survival. Hence, growth and time of release are
    importantattributesforhatcheryfish(Biltonetal. 1982,    Brodeur, R. D., H. V. Lorz, and W. G. Pearcy. 1987.
    Mahnken et al. 1982).  Thomas (1985) found a    Food habits and diet variations of pelagic nekton off
    correlation between coho salmon hatchery production    Oregon and Washington, 1979-1984. NOAA Tech.
    and a decline in central California Dungeness crab    Rep. NMFS 57, 32 p.
    (Cancer magister) abundance, probably related to
    coho salmon feeding on crab megalopae. El Niio also   Burner, C. J. 1951. Characteristics of spawning nests
    affects coho salmon abundance (Hayes and Henry   of Columbia River salmon. Fish. Bull., U.S. 61(52):97-
    1985). Finally, Japanese high-seas fishing fleets take    110.
    unknown numbers of coho salmon andthesquid gillnet
    fisheries may also take coho salmon incidentally.     Chapman, D. W. 1966. Food and space as regulators
                                                          of salmonid populations in streams. Am. Nat. 100:345-
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                                                       143

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                                                       144

    145

       Oncorhynchus mykiss
       Adults

                10 cm

    Common Name: steelhead (rainbow trout)                suspended sediment (Reiser and Bjornn 1979, Bell
    Scientific Name: Oncorhynchus mykiss, previously    1984).
    known as Salmo gairdneri (Smith and Stearley 1989)
    Other Common Names: Kamchatka salmon-trout,    Ecological: The steelhead is a dominant fish in many
    coastal rainbow trout, silvertrout, salmon trout, ironhead,   coastal and inland streams/rivers.
    chromer, hardhead, steelie, sea-run rainbow trout,
    seatrout, silversides, or summer salmon (Pauley et al.   Range
    1986)                                                 Overall: This species was originally found from
    Classification                                        northwestern Mexico to Kuskokwim River, Alaska.
    Phylum: Chordata                                      Now it is rarely found south of the Ventura River,
    Class: Osteichthyes                                   California (Wydoski and Whitney 1979, Barnhart 1986).
    Order: Salmoniformes                                  It is alsofound in Kamchatka and Okhotsk Sea drainages
    Family: Salmonidae                                    (McPhail and Lindsey 1970).

    Value                                                 Within Studv Area: The steelhead is found in all Pacific
    Commercial: The peak commercial catch (3,900 t) of   coast estuaries north of San Francisco Bay, California
    steelheadoccurredin 1945(Sheppard 1972). Presently,    (Table 1) (Monaco et al. 1990). A small run occurs in
    onlyNativeAmericansareallowedto fish commercially    Morro Bay, California (Horn 1980).
    for steelhead in Oregon and Washington. In 1985,342
    t were landed in the Columbia River, caught primarily   Life Mode
    with gillnets (Bohn and Mclsaac 1986).                The steelhead is the anadromous form of the rainbow
                                                          trout. Eggs and larvae (alevins) are benthic and
    Recreational: The steelhead is a highly-prized sport   infaunal.  Young juveniles (fry and parr) are
    fish because of its size, fighting abilities, and excellent   benthopelagic.  Parr become pelagic and acquire a
    taste. Nearly all recreational fishing occurs in streams    silver color when they transform into smolts (juveniles
    and rivers.  In Washington, steelhead allocation is   thatmigratetotheocean). Steelheadparrareterritorial
    divided 50:50 between Native American and non-    and reside in streams and rivers from 1 to 4 years
    treaty fishermen (Clark 1985). Although much natural   before transforming into smolts (Pauley et al. 1986).
    reproduction occurs, steelhead abundance has been    Smolts and ocean-dwelling and maturing juveniles
    augmented by hatchery production (Larson and Ward    (subadults), and adults are epipelagic (to depths of 23
    1954);approximately 17 million steelhead smolts were    m) (Okazaki 1983, Alaska Department of Fish and
    planted in the Columbia River basin in 1987.          Game 1985).  Subadults and adults in rivers and
                                                          streams are bottom-oriented.
    Indicator of Environmental Stress: This species is
    susceptible to changes in temperature, dissolved    Habitat
    oxygen, substrate, water depth, water velocities, and  TIye: Eggs, alevins, fry, and parr are riverine. Smolts

                                                       146

                                                                                         Steelheadcontinued

     Table 1. Relative abundance of steelhead in 32 U.S. Pacific coast estuaries.

                               Winter       Summer       Half-pounder       Fall
                              Life Stage    Life Stage     Life Stage     Life Stage
                 Estuary    IA S J L E  A S J L E   A S J L E   A S  J L E
                  Puget�Sound C                 j 0  0    0                             Relative abundance:
                                                                                      �   Highly abundant
                   Hood Canal C    0     0  00  O    O                                       Highly abundant
                                                                                      3   Abundant
                    SkagltBay 0    0       O                                             O   Common
                  Grays Harbor (3  -_ -               -__  Rare
                   Willapa Bay (3    Blank Not present
                Columbia River     (      (3   (
                  Nehalem Bay (3   (3
                 Tillamook Bay O      O         0                                       Life stage:
                   Netarts Bay q           -                                             A-Adults
                                                        Bay        _      __ -  -    -   - - - ---------S - Spawning adults
                   Siletz River O    O    :0                                            J - Juveniles
                                                                                      L - Larvae
                                                     - - - -  - -  - -_  -_ -_ -_--    E - Eggs
                   Alsea River  3   0   :
                  Siuslaw River O    O
                 Umpqua River 0    0       0    0
                    Coos Bay (3
                   Rogue River (    (      ( _                 (
                  Klamath River0   3  0(3 _0    _O 0
                  Humboldt Bay0    0
                     Eel River 0  3        0  O                0
                  Tomales Bay 0    0
             Cent. San Fran. Bay ' 0    0  Includes Central San
                                                                                       Francisco, Suisun.
             South San Fran. Bay                                                          Francisco, Suisun.
                                                                                       and San Pablo Bays.
                 Elkhorn Slough
                    Morro Bay  '    'i
               Santa Monica Bay
                 San Pedro Bay
                  Alamitos Bay
                  Anaheim Bay
                  Newport Bay
                   Mission Bay
                 San Diego Bay
                 Tijuana Estuary
                             A S J L E  A S J L E   A S J L E   A S J  L E

    are riverine and estuarine. Fry and parr reside in areas    probably not substrate-dependent.
    that have cover and move to deeper water (such as
    pools) astheygrow. Subadults and adults are found in    Phvsical/Chemical Characteristics: The steelhead
    coastal neritic waters during ocean residence and in   survives temperatures from 0-28�C, but at the upper
    riverinehabitatsduringthespawning migration. Smolts,    limit water must be saturated with dissolved oxygen.
    subadults, and "kelts" (spent adults) migrate through    The best temperatures for growth and development
    estuaries, but this species does not spend much time    are 13-210C (Moyle 1976).  Freshwater life stages
    rearing in estuaries (Dawley et al. 1986).               prefer temperatures of 10.0-12.80C (Bell 1984);
                                                          spawning occurs at 8.0-15.5�C (Wang 1986). The
    Substrate: Eggs are found in redds made in areas    steelhead appears to grow best in slightly alkaline (pH
    containing medium and small gravel (<85 mm  in    = 7.0-8.0) waters (Moyle 1976). Eggs, alevins, fry, and
    diameter) (Shapovalov and Taft 1954,  Alaska    parr are only found in fresh water. Juvenile salinity
    Department of Fish and Game 1985). Fry overwinter    tolerance is determined by fish size and water
    in stream areas where rubble is present. Sport-caught    temperature (Johnsson and Clarke 1988). Successful
    adults are often captured below spawning tributaries in   smoltification appears to be temperature-dependent
    swift-flowing water containing boulders (Scott and    (Zaugg et al. 1972,  Adams et al. 1975).  Smolts,
    Crossman 1973). Oceanic juveniles and adults are    subadults, and adults are found in fresh to marine

                                                       147

    Steelhead continued
    waters. This species' ocean distribution is influenced    the ocean and return a year or more later to their natal
    by sea surface temperatures (Sutherland 1973).         stream as "repeat spawners".  The percentage of
                                                           repeat spawners appears to vary according to stock,
    Miarations and Movements: The steelhead has    habitat quality, fishing intensity, and management
    excellent homing abilities, so unique stocks or races    practices (Shapovalov and Taft 1954, Withler 1966,
    have developed in specific drainage areas or streams    Jones 1977, Barnhart 1986). Females survive spawning
    (Moyle 1976). At least two races exist, as defined by    more often than males (Withler 1966); up to five times
    when adult fish enter fresh water to spawn (Smith    has been documented (Jones 1984).
    1960). The winter run migrates upstream during fall,
    winter and early spring, whilethe summer run migrates    Fecundity: Fecundity varies with female size and
    during spring, summer, and earlyfall (Bell 1984). Inthe    geographic origin (Buckley 1967).  Most females
    Columbia River and other large rivers with many    produce an average of 1,500-5,000 eggs (Bell 1984),
    tributaries, there are probably some steelhead entering    although large females may produce over 12,000 eggs
    year round. Adults appear to enter spawning streams    (Moyle 1976).
    during freshets (Pautzke and Meigs 1940). Juvenile
    steelhead normally rear in fresh water for 1-4 years    Growth and Development
    (usually2or3). Theythenmigratetotheocean(during    Eaa Size and Embryonic Development: Eggs are
    spring-early summer) where they spend 1-5 years    spherical, non-adhesive, and 3.0-6.2 mm in diameter
    (usually 2 or 3) before returning to their natal river. In   (Scott and Crossman 1973, Wang 1986). Embryonic
    some northern California and southern Oregon Rivers    development is indirect, external, and has an alevin
    (e.g., Klamath, Eel, and Rogue rivers), a "half-pounder"    (prolarval) stage. Eggs hatch in 18-101 days, depending
    run exists.  These are immature fish (weighing    onwatertemperatureandoxygenconcentrations(Silver
    approximately one-half pound) that return to rivers and    et al. 1963, Carlander 1969).
    streams after just a few months in the ocean. They
    overwinter in streams and then migrate back to sea in   Aae and Size of Larvae: Alevins are 14.0 mm long at
    the spring (Kesner and Barnhart 1972). Virtually all   hatching, and grow to a length of 28.0 mm before
    summersteelheadfromthese rivers make half-pounder    becoming juveniles (Wang 1986).
    migrations, but only a small percentage of winter
    steelhead do (Satterthwaite 1988).  Half-pounders    Juvenile Size Ranoe: Juvenile lengths are extremely
    appear to stray significantly more than adults    variable (2.8-40.6 cm), depending on age and
    (Satterthwaite 1988). Smolts and adults spend little   environmental conditions (Scott and Crossman 1973).
    time in estuaries (Dawley et al. 1986). In the ocean, the
    steelhead is most abundant in the Gulf of Alaska and    Aae and Size of Adults: Wild fish usually spend 2-4
    the eastern North Pacific (Sutherland 1973). In some    years in fresh water and 1-5 years at sea.  Most
    California coastal streams, it may return only in the fall   hatchery fish spend only one year in fresh water. Most
    because river mouths are not open (i.e., of sufficient   returning wild fish are 2/2, 2/3, 3/2, and 3/3 (years in
    depth) until after heavy rains (Fry 1973).              freshwater/years in ocean),while hatcheryfish are 1/1,
                                                           1/2, or 1/3 (Pauley et al. 1986). The more time spent
    Reproduction                                            in the ocean (during the initial ocean residency), usually
    Mode: The steelhead is gonochoristic and oviparous;    the larger the fish is at maturity (Maher and Larkin
    eggs arefertilized externally. This species differs from    1954).  Mature steelhead range from 45-70 cm in
    all other members of the genus Oncorhynchus (except    length and usually 2-5 kg (Shapovalov and Taft 1954,
    cutthroat trout, 0. clarkl) in that it is iteroparous.  Wydoski and Whitney 1979, Jones 1984). However,
                                                           steelhead can reach nine years (Washington 1970),
    Matina/Soawnina: Winter-run steelheadtypically spawn    122 cm in length (Scott and Crossman 1973), and 19.5
    from December to June (Bell 1984), while summer    kg (Hart 1973). Fish in the southern part of the range
    steelhead (which return to fresh water in spring and    aresmallerandspendlesstimeatseathanthosetothe
    summer)donotspawnuntilthefollowingspring(Everest    north (Withler 1966).  Adults averaged 58.1 cm in
    1973). Spawning periods vary from north to south and    length in California, 66.7 cm in Oregon, and 71.0 cm in
    by river system (Leider et al. 1984).  Females build   southern British Columbia (Withler 1966).
    redds (up to 5.5 m2)in areas with appropriate gravel
    and water flows. The mating male defends the female    Food and Feeding
    and redd from intruders and fertilizes the eggs as the    TroDhic Mode: Larvaefeed ontheiryolk. Juveniles and
    female extrudes them (Shapovalov and Taft 1954).    adults are carnivorous.
    Spawning occurs day and night. Spent adults (kelts)
    may not die after spawning, but instead move back to    Food Items: In freshwaterand estuarine areas, primary

                                                       148

                                                                                           Steelhead continued
    food items include gammarid amphipods, small    Barnhart, R. A. 1986. Species profiles: life histories
    crustaceans, insects, and small fishes (Moyle 1976,    and environmental requirements of coastal fishes and
    Wydoski and Whitney 1979, Loch 1982, Dawley et al.   invertebrates (Pacific Southwest) - steelhead. U.S.
    1986).  In the ocean, juveniles and adults eat    Fish Wildl. Serv. Biol. Rep. 82(11.60),  U.S. Army
    crustaceans, insects, squid, and fishes (LeBrasseur    Corps Eng., TR EL-82-4, 21 p.
    1966, Wydoski and Whitney 1979).
                                                           Bell, M. C. 1984. Fisheries handbook of engineering
    Biological Interactions                                 requirements and biological criteria.  Fish Passage
    Predation: In fresh water, this species is eaten by coho    Development and Evaluation Program, Corps Eng.,
    salmon(O. kisutch),char(Salvelinusspp.),mergansers,    North Pac. Div., Portland, OR, 290 p. (Contract No.
    gulls, belted kingfisher (Megaceryle alcyon), bears,    DACW57-79-M-1594 and DACW57-80-M-0567).
    marten (Martes americana), otter (Loutra canadensis),
    and other steelhead.  In the ocean, Pacific lamprey    Bohn, B. R., and D. Mclsaac. 1986. Columbia River
    (Lampetratridentata),seals, sea lions, and killerwhale    fish runs and fisheries 1960-1985. Oreg. Dept. Fish
    (Orcinus orca) prey upon this species (Scott and    Wildl. and Wash. Dep. Fish., Clackamas, OR, 77 p.
    Crossman 1973, Simenstad et al. 1979).
                                                           Buckley, R. V. 1967. Fecundity of steelhead trout,
    Factors Influencina Pooulations: Freshwater life stages    Salmo gairdneri from Alsea River, Oregon. J. Fish.
    are often adversely affected by natural and human-    Res. Board Can. 24(4):917-926.
    induced habitat alterations. Most natural mortality
    occurs in the egg and larval stages (97%) (Shapovalov    Carlander, K. D. 1969. Handbookoffreshwaterfishery
    and Taft 1954). Factors which influence freshwater    biology, Vol. 1. Iowa State Univ. Press, Ames, IA,
    mortality include the numberof eggs deposited, siltation,    752 p.
    dissolved oxygen, watervelocity, temperature, turbidity,
    depth, barriers, pollution, and competition with other    Chilcote, M. W., S. A. Leider, and J. J. Loch. 1986.
    fishes (Pauley et al. 1986). Survival of migrating smolts    Differential reproductive success of hatchery and wild
    is size-dependent, with larger and older fish having    summer-run steelhead under naturalconditions. Trans.
    higher survival rates (Pauley et al. 1986, Ward et al.   Am. Fish. Soc. 115:726-735.
    1989). "El Nino" (i.e., abnormally warm ocean
    conditions) also affects survival and growth (Pearcy et   Clark, W. G. 1985. Fishing in a sea of court orders:
    al. 1985). Overfishing has reduced some populations    Puget Sound salmon management ten years after the
    and the proliferation of hatchery smolts can adversely    Boldt decision. N. Am. J. Fish. Manag. 5(3B):417-434.
    affectwildfish populations (Pauley etal. 1986). Hatchery
    fish do not have survival rates as high as wild fish nor    Dawley, E. M., R. D. Ledgerwood, T. H. Blahm, C. W.
    are they as successful in producing smolted offspring    Sims, J. T. Durkin, R. A. Kirn, A. E. Rankis, G. E.
    (Chilcote et al. 1986). Manywild stocks in Washington    Monan, and F. J. Ossiander.   1986.  Migrational
    appear to have reduced genetic diversity because of   characteristics, biological observations, and relative
    interbreeding   with   hatchery-produced   fish    survival of juvenile salmonids entering the Columbia
    (Reisenbichler and Phelps 1989). Some stocks are    River estuary,  1966-1983.  Final Rep. to Bonneville
    more resistant to disease than others (Wade 1986).    Power Adm., Contract DE-A179-84BP39652, 256 p.
    Hence, interbreeding between wild and hatchery fish   Available Northwest and Alaska Fish. Center, 2725
    may produce fish with lower resistance to disease.      Montlake Blvd. E., Seattle, WA.

    References                                              Everest, F. H. 1973.  Ecology and management of
                                                           summer steelhead in the Rogue River. Fish Res. Rep.
    Adams, B. L., W. S. Zaugg, and L. R. McLain. 1975.    7, Oreg. State Game Comm., Portland, OR, 48 p.
    Inhibition of salt water survival and Na-K-ATPase
    elevation in steelhead trout (Salmo gairdnern) by    Fry, D. H., Jr. 1973. Anadromous fishes of California.
    moderate water temperatures. Trans. Am. Fish. Soc.    Calif. Dept. Fish Game, Sacramento, CA, 111 p.
    104(4):766-769.
                                                           Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    Alaska Department of Fish and Game. 1985. Alaska    Board Can., Bull. No. 180, 740 p.
    habitat management guide, southcentral region, Vol. I:
    Life histories and habitat requirements of fish and    Horn, M. H.  1980.  Diel and seasonal variation in
    wildlife. Alaska Dept. Fish Game, Juneau, AK, 429 p.    abundance and diversity of shallow-water fish

                                                       149

    Steelhead continued
    populations in Morro Bay, California. Fish. Bull., U.S.    Moyle, P. B. 1976. Inland fishes of California. Univ.
    78(3):759-770.                                          Calif. Press, Berkeley, CA, 405 p.

    Johnsson, J., and W. C. Clarke. 1988. Development    Okazaki, T. 1983. Distribution andseasonalabundance
    of seawater adaptation in juvenile steelhead trout   of Salmo gairdneri and Salmo mykiss in the North
    (Salmo gairdnert) and domesticated rainbow trout   Pacific Ocean. Japan. J. Ichthyol. 30(3):235-246.
    (Salmo gairdnen)-effects of size, temperature and
    photoperiod. Aquacult. 71:247-263.                      Pauley, G. B., B. M. Bortz, and M. F. Shepard. 1986.
                                                           Species profiles: life histories and environmental
    Jones, D. E. 1977. Life history of steelhead trout and    requirementsof coastalfishesandinvertebrates (Pacific
    life history of sea-run cutthroat trout. Alaska Dept. Fish    Northwest) - steelhead trout. U.S. Fish Wildl. Serv.
    Game, Compl. Rep. AFS-42, 18:52-105.                    Biol. Rep. 82(11.62). U.S. Army Corps Eng., TR EL-
                                                           82-4, 24 p.
    Jones, D. E. 1984. A study of cutthroat-steelhead in
    Alaska. Alaska Dept. Fish Game, Anad. Fish Studies.    Pautzke, C. F., and R. C. Meigs. 1940. Studies on the
    Annual Performance Rep. 1983-84. Project AFS-42-    life history of the Puget Sound steelhead trout (Salmo
    11, 25:73-87..                                         gairdneril). Trans. Am. Fish. Soc. 70:209-220.

    KesnerW. D., and R. A. Barnhart. 1972. Characteristics    Pearcy, W., J. Fisher, R. Brodeur, and S. Johnson.
    ofthefall-runsteelheadtrout(Salmogairdnerigairdnen)    1985. Effects of El Nino on coastal nekton off Oregon
    of the Klamath River system with emphasis on the half-   and Washington, In W. S. Wooster and D. L. Fluharty
    pounder. Calif. Fish Game 58(3):204-220.                (editors),  El Nino North - El Nino effects in the
                                                           subartic Pacific Ocean, p. 186-204. Wash. Sea Grant
    Larson, R. W., and J. M. Ward. 1954. Management of   Publ. WSG-WO 85-3, Univ. Wash., Seattle, WA.
    steelhead trout in the state of Washington. Trans. Am.
    Fish. Soc. 84:261-274.                                  Reisenbichler, R. R., and S. R. Phelps. 1989. Genetic
                                                           variation in steelhead (Salmo gairdnen) from the north
    LeBrasseur, R. J. 1966. Stomach contents of salmon    coast of Washington. Can. J. Fish. Aquat. Sci. 46:66-
    and steelhead trout in the northeastern Pacific Ocean.    73.
    J. Fish. Res. Board Can. 23(1):85-100.
                                                           Reiser, D. W., and T. C. Bjornn. 1979. 1. Habitat
    Leider, S. A., M. W. Chilcote, and J. J. Loch. 1984.    requirements of anadromous salmonids.  In W. R.
    Spawning characteristics of sympatric populations of    Meehan (editor), Influence of forest and rangeland
    steelhead trout (Salmo gairdnen): evidence for partial    management on anadromousfish habitat in the western
    reproductive isolation.  Can. J. Fish. Aquat. Sci.    United States and Canada, p. 1-54. U.S. Forest Serv.
    41(10):1454-1462.                                       Gen. Tech. Rep. PNW-96, Pac. Northw. Forest Range
                                                           Exp. Sta., Portland, OR.
    Loch, J. J. 1982. Juvenile and adult steelhead and
    sea-run cutthroat trout within the Columbia River    Satterthwaite, T. D. 1988.  Influence of maturity on
    estuary, 1980. 1982 Ann. Rep., Wash. Dept. Game,    straying rates of summer steelhead into the Rogue
    Olympia, WA, 47 p. plus appendices.                     River, Oregon. Calif. Fish Game 74(4):203-207.

    Maher, F. P., and P. A. Larkin 1954. Life history of   Scott, W. B., and E. J. Crossman. 1973. Freshwater
    steelhead trout of the Cilliwack River, British Columbia.    fishes of Canada. Fish. Res. Board Can., Bull. No. 184,
    Trans. Am. Fish. Soc. 84:27-38                          966 p.

    McPhail, J. D., and C. C. Lindsey. 1970. Freshwater    Shapovalov, L., and A. C. Taft. 1954. The life histories
    fishes of northwestern Canada and Alaska. Fish. Res.    of the steelhead rainbow trout (Salmo gairdneri
    Board Can., Bull. No. 173, 381 p.                       gairdnen) and silver salmon (Oncorhynchus kisutch)
                                                           with special reference to Waddell Creek, California,
    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.    and recommendations regarding their management.
    Nelson. 1990. Distribution and abundance of fishes   Calif. Fish. Game, Fish Bull. 98, 375 p.
    and invertebrates in west coast estuaries, Volume I:
    data summaries.   ELMR  Rep. No. 4. Strategic    Sheppard, D. 1972. Thepresentstatusofthesteelhead
    Assessment Branch, NOS/NOAA, Rockville, MD,             trout stocks along the Pacific coast. In D. H. Rosenberg
    240 p.                                                  (editor). A review of the oceanography and renewable

                                                        150

                                                                                           Steelhead continued
    resources of the northern Gulf of Alaska, p. 519-556.    Board Can. 23(3):365-393.
    Univ. Alaska Inst. Mar. Sci. Rep. R72-73, Sea Grant
    Rep. 73-3, Fairbanks, AK.                               Wydoski, R. S., and R. R. Whitney. 1979. Inland fishes
                                                           of Washington. Univ. Wash. Press, Seattle, WA,
    Silver, S. J., C. E. Warren, and P. Doudoroff. 1963.    220 p.
    Dissolved oxygen requirements of developing steelhead
    trout and chinook salmon embryos at different water    Zaugg, W. S., B. L. Adams, and L. R. McLain. 1972.
    velocities. Trans. Am. Fish. Soc. 92(4):327-341.        Steelhead migration: potential temperature effects as
                                                           indicated by gill adenosine triphosphatease activities.
    Simenstad, C. A., B. S. Miller, C. F. Nyblade, K.    Science 176:415-416.
    Thornburgh, and L. J. Bledsoe. 1979. Food web
    relationships of northern Puget Sound and the Strait of
    Juan de Fuca. U.S. Interagency (NOAA, EPA) Energy/
    Environ. Res. Dev. Prog. Rep., EPA-600/7-79-259,
    Wash., D.C., 335 p.

    Smith, G. R., and R. F. Stearley. 1989. The classification
    and scientific names of rainbow and cutthroat trouts.
    Fisheries 14(1):4-10.

    Smith, S. B. 1960. A note on two stocks of steelhead
    trout (Salmo gairdneri) in Capilano River, British
    Columbia. J. Fish. Res. Board Can. 17:739-742.

    Sutherland, D. F. 1973. Distribution, seasonal
    abundance, and some biological features of steelhead
    trout, Salmo gairdneri, in the North Pacific Ocean.
    Fish. Bull., U.S. 73(3):787-826.

    Wade, M. 1986. The relative effects of Ceratomyxa
    shasta on crosses of resistant and susceptible stocks
    of summer steelhead. Info. Rep. 86-6, Oreg. Dept.
    Fish Wildl., Corvallis, OR, 16 p.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: A
    guide to the early life histories. Tech. Rep. No. 9.
    Interagency ecological study program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    and U.S. Fish Wildl. Serv., various pagination.

    Ward, B. R., P. A. Slaney, A. R. Facchin, and R. W.
    Land. 1989. Size-biased survival in steelhead trout
    (Oncorhynchus mykiss): back-calculated lengths from
    adults' scales compared to migrating smolts at the
    Keogh River, British Columbia. Can. J. Fish. Aquat.
    Sci. 46(11):1853-1858.

    Washington, P. 1970. Occurrence on the high seas of
    a steelhead trout in its ninth year. Calif. Fish Game
    56(4):312-314.

    Withler, I. L. 1966. Variability in life history
    characteristics of steelhead trout (Salmo gairdnen)
    along the Pacific coast of North America. J. Fish. Res.

                                                       151

      Oncorhynchus nerka
      Adults

    Common Name: sockeye salmon                           catches do occur in Alaska) (Pacific Marine Fisheries
    Scientific Name: Oncorhynchus nerka                   Commission 1987). However, the landlocked variety
    Other Common  Names: red salmon, kokanee    (kokanee) is a very important freshwater sport fish in
    (landlocked populations), blueback, redfish, Fraser   California, Oregon, Washington, Idaho, and Alaska
    River salmon, nerka, sau-aui salmon, sukkegh salmon,    (Scott and Crossman 1973, Moyle 1976).
    Kennerly's salmon, kootenary salmon, silvertrout, little
    redfish, princess trout (Shiino 1976)                 Indicatorof Environmental Stress: Upstream migrations
    Classification (Robins et al. 1980)                   may be disrupted when waters have hydrocarbon
    Phylum: Chordata                                      concentrations of 1-10 ppb (or greater) (Martin et al.
    Class: Osteichthyes                                   1990). See "Factors Influencing Populations".
    Order: Salmoniformes
    Family: Salmonidae                                    Ecological: This species is the third most abundant
                                                           salmonid in the North Pacific [behind pink and chum
    Value                                                 salmon (0. keta)] (Fredin et al. 1977).
    Commercial: The sockeye salmon is a prized
    commercialfish because of its excellent flesh color and    Range
    flavor (Scott and Crossman 1973). It is second only to   Overall: This is a boreal Pacific species. In Asia, it is
    pink salmon (0. gorbuscha) in U.S. salmonid landings,   found from the southern Kurile Islands to the northern
    but first in value. In 1985, U.S. fishermen received over   sea coast of the U.S.S.R. In North America, important
    $239 million for their sockeye salmon catch (National   spawning populations occur from the Columbia River
    Marine FisheriesService 1986). In 1978, U.S.fishermen   in the south to northern Alaska in the north (French et
    caught over 19 million sockeye salmon, primarily in   al. 1976). The oceanic distribution ranges from the
    Alaska (Forrester 1981).  The sockeye salmon is   eastern Bering Sea south to lat. 450N, and is associated
    caught throughout the North Pacific (Japan to Oregon),    with the California Current as far south as Los Angeles
    with U.S. fisheries catching most (Fredin 1980). U.S.    Harbor (French et al. 1976, Eschmeyer et al. 1983).
    commercial catches of sockeye salmon have fluctuated
    dramatically in the past, primarily due to fluctuations in   Within Studv Area: The Columbia River is the southern
    the important Bristol Bay fishery in Alaska (Fredin et al.   limit of all sizable runs (Table 1) (Foerster 1968). The
    1977). The sockeye is primarily captured by gill net and    sockeye salmon is abundant in Puget Sound (Wydoski
    purse seine (occasionally by trolling), primarily during    and Whitney 1979). Two runs also exist on the northern
    June to August (peak in July).                        coast of Washington in Lake Quinault and Lake Ozette
                                                           (Pauley et al. 1989).
    Recreational: The sockeye salmon (anadromous
    variety) does not take a hook as readily as other   Life Mode
    salmonids. Hence, it is not considered an important   This is an anadromous species with a landlocked
    recreational salmonid in the study area (although large   variety (kokanee).  Eggs and larvae (alevins) are

                                                        152

                                                                                       Sockeye salmon continued

                                                            without going to sea (Moyle 1976). Smolts are riverine
      Table 1. Relative abundance of sockeye salmon         and estuarine. Ocean-dwelling juveniles stay in neritic
               in 32 U.S. Pacific coast estuaries.          and epipelagic areas until fall and early winter, then
                         Life Stage                         move to oceanic areas (Hartt and Dell 1986). While in
            Estuary    A S J L E                           the ocean, they reside in the upper 61 m (French et al.
             Puget Sound i;: :3       Relative abundance:   1976). Adults are primarily estuarine and riverine.
              Hood Canal      :            Highly abundant
              Skagit Bay  4   '1           Abundant        Substrate: Eggs and alevins reside beneath fine gravel/
             Grays Harbor          c        ommon           cobble. Fry and adults occur in the water column, but
             Grays Harbor              ~   Rare
              Willapa Bay             Blank Not present     are associated with gravel bottoms. Parr, smolts, and
           Columbia River O    O                           juveniles live in the water column (Foerster 1968, Hart
            Nehalem Bay                                    1973).
            Tillamook Bay             Life stage:
              Netars Bay               A - Adults           Phvsical/Chemical Characteristics: Eggs, alevins, fry,
                                       J-JvS Spaning adults  and parr live in fresh water, while smolts and adults
             YaquinaBay               L-Larvae             inhabit fresh to euhaline waters.  Ocean-dwelling
              Alsea River              E-Eggs               juveniles do not appear to be affected by salinity
             Siuslaw River                                  changes, but are sensitive to temperature variations
            Umpqua River                                    (French et al. 1976). Normal spawning temperatures
               Coos Bay                                    range from 3-70C (Ricker 1966, Foerster 1968). Adult
             Rogue River                                   sockeye salmon migrate in river temperatures of 7.2-
            Klamath River                                   15.60C (Reiser and Bjornn 1979).  Recommended
            Humboldt Bay                                    incubation guidelines are: dissolved oxygen at or near
                Eel River                                   saturation (lower level of 5.0 mg/I); watertemperatures
             Tomales Bay                u   na d            of 4-14�C; apparent velocity (within the redd) more
        Cent. San Fran. Bay  '        Includes Central San
                                      nt. San Fran. Bay  Francisco. Suisun.  than 20 cm/hr; and spawning sediment composed of
       South San Fran. say           and San Pablo bays.   less than 25% (by volume) fines (<6.4 mm) (Reiser and
           Elkhorn Slough                                  Bjornn 1979). The upper lethal water temperature is
               Morro Bay
               Santa Moirro Bay                             24.4�C (Brett 1952), but growth ceases at temperatures
           San Pedro Bay                                   above 20.30C (Bell 1984). Ocean-dwelling juveniles
             SAlamitos Bay                                  reside in temperatures of 1.0-13.0�C (French et al.
             Anaheim Bay                                    1976). Low pH can affect the viability of embryos and
             Newport Bay                                    alevins (Rombough 1983), and nitrogen supersaturation
              Mission Bay                                   can adversely affect outmigrating smolts (Ebel et al.
            San Diego Bay                                   1971).
           Tijuana Estuary
                                                            Miarations and Movements: Kokanee do not migrateto
                                    A S J L El              sea, but anadromous stocks migrate extensively.
                                                            Sockeye salmon generally spend 1-2 years rearing in
    benthic and infaunal. Young juveniles (fry and parr) are    freshwater lakes and 2-3 years in the ocean. However,
    benthopelagic.   Parr become pelagic before they    depending on geographic area, they may spend 0-4
    transform into smolts (juveniles that migrate to the    yearsinfreshwaterbeforemigrating, and up to 4 years
    ocean).  Smolts and ocean-dwelling and maturing    in the ocean (Foerster 1968, Fredin et al. 1977). After
    juveniles (subadults), and adults are pelagic. Subadults    emerging from the redd (January-June), fry typically
    and adults in rivers and streams are bottom-oriented.    move upstream or downstream into a nursery lake,
                                                            although some may move directlyto estuaries (Foerster
    Habitat                                                 1968). Once in lakes, young sockeye salmon live for
    iyp.: Eggs, alevins and fry are primarily riverine (some    approximately 1 month in the littoral zone before moving
    lacustrine); if in lacustrine environments they occur    out into open lake waters, where they reside until they
    wherethereisfreshwaterflowthroughtheredd(Wydoski    migrate to sea (McCart 1966, Foerster 1968). While
    and Whitney 1979). Parr normally rear in lakes for 1 -   residing in lakes, juvenilesundertakevertical migrations,
    2 years, feeding primarily in the upper 20 m. However,    probably related to food availability and predation risks
    in some populations parr do not rear in lakes, but move    (Clark and Levy 1988). Smolts begin to migrate out of
    downstream after emerging from the gravel (Foerster    lakes when temperatures riseto4-7�C (usually March-
    1968). Anadromous stocks usually smoltify after 1-2   July) and normally at night (Hart 1973). One exception
    years,butkokaneeremainandcompletetheirlifecycle    is in Lake Washington, Washington, where smolts

                                                         153

    Sockeye salmon continued
    migrate both day and night (Simenstad et al. 1982).   (Fredin et al. 1977). Males and females may spawn
    Sockeye salmon smolts in the Pacific Northwest   with several different fish. Females defend the nest
    outmigrate primarily between April and early June    site after spawning until they tire and die. During their
    (Anas and Gauley 1956, Simenstad et al. 1982). Smolts    spawning migration, sockeye salmon undergo sexually
    are 40-130mm in length when they enterestuaries and    dimorphic changes; both sexes developing bright red
    are guided to ocean waters by salinity gradients (Healey    bodies and green heads, while males develop a humped
    1980, Straty and Jaenicke 1980). Residence time in   back, hooked snout, and large teeth (Foerster 1968).
    estuaries is shorterthan other salmonid species (Healey
    1982, Simenstad et al. 1982).  Upon entering the   Fecundity: Fecunditydependsonthesizeof thefemale
    ocean, juvenile sockeye salmon (not including Bristol   and the stock (Rounsefell 1957, Manzer and Miki
    Bay stocks) move north, staying within the coastal belt   1986). The anadromous sockeye salmon has from
    of the Gulf of Alaska until late-fall or early-winter when    2,200-4,300 eggs per female with 3,500-3,600 eggs
    theydisperseoffshore, moving westand south (French    per female being average (Hart 1973, Fredin et al.
    et al. 1976, Hartt and Dell 1986).  In spring and    1977, Bell 1984).
    summer, they move north, but turn south and west
    again in winter (French et al. 1976). Migrants initially   Growth and Development
    travel 3.9-30.2 km/day (Hartt and Dell 1986) and older    Eaa Size and Embryonic Develooment: Scott and
    fish normally travel 13-33 km/day. Maturing fish may    Crossman (1973) and McPhail and Lindsey (1970)
    travel 46-56 km/day (French et al. 1976). Sockeye    reported sockeye salmon egg diameters of 4.5-5.0
    salmon show some diel migrations, moving to the   mm, whereas Bell (1984) reported eggs 5.5-6.0 mm in
    surface at night and deeper during the day (French et   diameter.  Embryonic development is indirect and
    al. 1976). North American sockeye salmon populations   external. Hatching can take slightly less than 50 days
    have a single spawning run, occurring from May to   or more than 5 months, depending on temperature
    December(dependingongeographiclocation). Pacific   (Hart 1973, Scott and Crossman 1973).
    Northwest adult sockeye salmon migrate into fresh
    water during June to August (peaking in early July)   Aae and Size of Larvae: Size at hatching is not reported
    (Simenstad et al. 1982, Bohn and Mclsaac 1986).    butprobably20-25mmtotallength(TL). Afterhatching,
    Oceanic migration is thought to be guided by a map-    alevins stay in the gravel for 2-3 weeks (or up to 4
    compass-calendar system (Quinn 1982), but the natal   months, depending on temperature) and emerge from
    stream is located by olfaction (Brannon 1982).       March to June (Hanamura 1966, Ricker 1966, Hart
                                                          1973, Scott and Crossman 1973, Wydoski and Whitney
    Reproduction                                         1979). At approximately 30 mm TL, alevins become fry
    Mode: The sockeye salmon is gonochoristic, oviparous,    (Hanamura 1966, Alaska Department Fish and Game
    and semelparous (all adults die soon after spawning).    1985).
    Eggs are fertilized externally.
                                                          Juvenile Size Ranae:Juveniles range in size from3 cm
    Matina/SDawnina: PacificNorthweststocksspawnfrom    to at least 46 cm TL.
    August to December, with an October peak (Wydoski
    and Whitney 1979, Bell 1984).  Except for a few   Aae and Size of Adults: Adults average 63.5 cm TL
    instances, the sockeye salmon spawns in rivers and    (50.0-84.0 cm), weighing an average of 3.0-4.0 kg
    streamsthatconnecttolakes. Spawningoccursmostly    (Fredin et al. 1977, Bell 1984, Kessler 1985) and 3-8
    in riffle areas in streams, but also in some lakes down    years old (average of 4 years) at spawning (Foerster
    to 30 m (Ricker 1966); spawning usually at depths <8    1968).
    m (Moyle 1976). Like other salmonids, the female
    builds the redd by facing upstream and thrashing her   Food and Feeding
    caudal fin against the substrate. Males may also make    Trophic Mode: Larvae feed on theiryolk. Juveniles and
    digging movements (McCart 1969). Males and females    adults are carnivorous (primarily planktivorous).
    are territorial, defending the nest site against members
    of the same sex.  During spawning, the male and    Food Items: Spawning adults typically do not feed,
    female place themselves in the redd with vents close   however, some will feed when held in net pens. All
    togetherand extrudeeggs and spermwiththeirmouths    free-swimming life stages are principally plankton
    agape and bodies quivering (Foerster 1968). Females    feeders. Planktonic Crustacea, cladocerans (Daphnia
    will repeat the digging slightly upstream, burying the   spp., Bosmina spp., etc.), and copepods (Epischura
    previous eggs in the process and creating a new    spp., Cyclopsspp., etc.) are eaten, along with a variety
    "pocket". A redd typically has 3-10 pockets (usually 5)   of terrestrial and aquatic insects (Ricker 1966, Foerster
    (Hart 1973) and averages in size at about 1.8 m2    1968, Hart1973,ScottandCrossman1973, Dobleand

                                                       154

                                                                                      Sockeye salmon continued
    Eggers 1978). During their downstream migration,   pollutants, etc.) which can be a result of poor forest
    smolts may feed heavily on gammarid amphipods    practices, industrial waste, mining and refining effluents,
    (Muir and Emmett 1988). In estuaries, euphausiids,    agriculture practices,and urban development. Physical
    fish larvae, juvenile shrimp, insects, amphipods, and    disturbance of the redd (by erosion, subsequent
    mysids are eaten (Levy and Yesaki 1982, Simenstad et   spawners, ice scour) and predation can also diminish
    al. 1982). In the ocean, juvenile sockeye salmon feed    freshwaterproduction (Foerster 1968, Hart 1973). River
    on euphausiids, hyperiid amphipods, copepods,    obstructions such as dams (manmade and natural,
    decapod larvae, pteropods, juvenile and larval fish,   such as Hell's Gate and the Fraser River rock slide of
    squid, and other invertebrates.  The primary prey    1913) can affect upstream and downstream migrations
    consumed depends on the location, time of day, and    (Foerster 1968). Columbia River sockeye salmon runs
    fish's age (Andrievskaya 1957, Allen and Aron 1958,    have diminished primarily as a result of dams and
    Ito 1964, LeBrasseur 1966, Foerster 1968, Pearcy et   irrigation diversions of spawning rivers (Mullan 1986).
    al. 1984). In lakes and in the ocean, juvenile sockeye    The abundance of food relative to parr numbers in
    salmon appear to feed primarily at dusk or at night   reservoirs and lakes also affects production; when
    (Doble and Eggers 1978, Pearcy et al. 1984). Parr may    sockeye parr densities are high, food may limit their
    not feed during the winter in lakes (Doble and Eggers    growth which in turn can reduce smolt size and marine
    1978). Juveniles (ocean- and lake-dwelling) feed near   survival (Foerster 1954,1968, Kyle et al. 1988). Nutrient
    the surface, except in lakes when surface temperatures    fertilization of lakes has been attempted to increase
    are high (Foerster 1968).                             lake primary production and zooplankton standing
                                                          crop and thus juvenile sockeye salmon growth and
    Biological Interactions                               survival (LeBrasseur et al. 1978, Hyatt and Stockner
    Predation: Primary fish predators of fry and parr in   1985).  Predators and competition can reduce
    fresh water are coho salmon (0. kisutch), cutthroat   populations in reservoirs (Foerster 1968).  Ocean
    trout (0. clark,), char (Salvelinus spp.), rainbow trout   conditions may also reduce production as a result of
    (0. mykiss), Dolly Varden (Salvelinus malma), lake   density-dependent mortality (Peterman 1980). The
    trout (Salvelinus namaycush),  lake whitefish    Japanese high seas fishery (located west of long.
    (Coregonus clupeaformis), mountain whitefish    174�W) intercepts many North American sockeye
    (Prosopium williamsonm), northern squawfish    salmon (Fredin et al. 1977). This fishery took over 46
    (Ptychocheilus oregonensis), burbot (Lota Iota), and    million North American sockeye over a 20 year period.
    sculpins (Foerster 1968, Fresh 1984). Gulls, common    This catch, togetherwith the accidental mortalities and
    loon (Gavia immer), red-necked grebe (Podiceps   lost additional weight gain before North American
    grisegena), common merganser (Mergus merganser),    harvest, represents a substantial loss to U.S. fishermen
    belted kingfisher (Megaceryle alcyon), terns, and large   (Ricker 1976, Fredin et al. 1977). Hatchery releases of
    predatory birds [osprey (Pandion haliaetus) and bald    sockeye salmon are used to maintain this species'
    eagle (Haliaeetus leucocephalus)] are important avian    abundance in some areas (Wahle and Smith 1979).
    predators (Fresh 1984). Marine predators include
    lamprey (Lampetra spp.), spiny dogfish (Squalus    References
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    estuaries in the life history of Pacific salmon: an
    unappreciated function. In V. S. Kennedy (editor),
    Estuarine comparisons, p. 343-364. Academic Press,
    New York, NY.

    Simenstad, C. A., B. S. Miller, C. F. Nyblade, K.
    Thornburgh, and L. J. Bledsoe. 1979. Food web
    relationships of northern Puget Sound and the Strait of
    Juan de Fuca. U.S. Interagency (NOAA, EPA) Energy/
    Environ. Res. Dev. Prog. Rep., EPA-600/7-79-259,
    Washington, D.C., 335 p.

    Straty, R. R., and H. W. Jaenicke. 1980. Estuarine
    influence of salinity, temperature, and food on the
    behavior, growth, and dynamics of Bristol Bay sockeye
    salmon. InW. J. McNeiland D. C. Himsworth (editors),
    Salmonid ecosystems of the North Pacific, p. 247-265.
    Oregon State Univ. Press, Corvallis, OR.

                                                      158

    159

      Oncorhynchus tshawytscha
      Adults

    Common Name: chinook salmon                          Canada Pacific Salmon Interception Treaty of 1985
    Scientific Name: Oncorhynchus tshawytscha            reduced the ocean take of chinook salmon off British
    OtherCommon Names:Columbia Riversalmon, king    Columbia and Alaska by 25% of 1984 catch levels
    salmon, black salmon, blackmouth salmon, chub    (Phinney1986).
    salmon, hookbill, quinnat salmon, Sacramento River
    salmon, saw-keivey, spring salmon, tchaviche, tule or   Recreational: This species is a prized sport fish because
    tyee salmon, winter salmon (Allen et al. in press)    of its size, fighting ability, availability, and excellent
    Classification (Robins et al. 1980)                  taste. Along with cohosalmon (O. kisutch),thechinook
    Phylum: Chordata                                     salmon supports a sport and charter boat fishery from
    Class: Osteichthyes                                   San Francisco, California, to Alaska. It is sport-caught
    Order: Salmoniformes                                 primarily in marine and estuarine waters, but many are
    Family: Salmonidae                                   also caught in fresh water.  Over 438,000 chinook
                                                          salmon were sport caught in the United States in 1984
    Value                                                 (not including California, Washington, and Oregon
    Commercial: The chinook salmon isthe least-abundant    freshwatercatch) (Pacific Marine Fisheries Commission
    Pacificsalmon,butitgrowsthelargestandcommands    1986).  The value of the recreational fishery is
    the highest price. In 1985, over 12,200 t worth $43    undetermined, butthevalueperkg is much higherthan
    million were landed on the Pacific coast (National   for commercial fish (Beauchamp et al. 1983). This
    Marine Fisheries Service 1986).  From 1875 to the   species is fished almost year-round in Puget Sound,
    1920s, the Columbia River had the largest chinook    but primarily fished from summerto fall in other areas.
    salmon run in theworld, with annual landings averaging
    9,100-18,100 t (Van Hyning 1973). In North America,    Indicator of Environmental Stress: Copper adversely
    the chinook salmon is commercially fished from    affects proper smoltification (Beckman and Zaugg
    Kotzebue Sound, Alaska, to Santa Barbara, California.   1988), and smolts in sea water are more sensitive to oil
    It is also commercialy fished along the Kamchatka    than when in freshwater. Reduced riverflows, increased
    Peninsula, U.S.S.R., to northern Japan. In California,   water temperatures, and many other man-induced
    only ocean trolling is allowed (Frey 1971). In Oregon    alterations to the environment can affect this species
    and Washington, it is captured by gill net, ocean    (see "Factors Affecting Populations").
    trolling, purse seine, and reef net. It is the most
    abundant salmon in California (McGinnis 1984).    Ecological: Juveniles are important due to their
    Chinook are often captured far from their place of   abundance in many Pacific coast rivers and streams
    origin, with large numbersof chinooksalmonoriginating    and are one of the most abundant neritic fish in Puget
    from the Columbia River caught off British Columbia,    Sound (Simenstad et al. 1979). Adults and juveniles
    Canada, and Alaska (Wright 1968). In Puget Sound,    are common in neritic waters off Oregon and
    Washington, half of the chinook salmon are harvested    Washington (Fisher et al. 1983, Fisher and Pearcy
    by Native Americans (Clark 1985). The United States/   1985).

                                                      160

                                                                                       Chinook salmon continued

     Table 1. Relative abundance of five races of chinook salmon in 32 U.S. Pacific coast estuaries.

                        Winter        Spring         Summer           Fall         Late-fall
                       Life Stage    Life Stage      Life Stage     Life Stage     Life Stage
          Estuary    A S J L E   A S J  L E   A S J L E   A S J L E   A S J  L E
           Puget Sound -                  (3-         - -  -   Relative abundance:
            Hood Canal      0:      O                                             _    ___         �   Highly abundant
             Skagit Bay              (3  :3                        fi 1                            (I)  Abundant
                                                                                              O   Common
           Grays Harbor              13              m3                                            o
                                                                                              - Rare
            Willapa Bay� _____   _____ - - -                               __   ___          _ -Blank Not present
          Columbia River     a            6         0  a�          3
           Nehalem Bay                                              3    3
          Tillamook Bay             o__   _____a -                                                Life stage:
            Netarts Bay     A- Adults
            Siletz River            :    9    : a          :                                       S - Spawning adults
                                             ~- - - -                                             - -- J - Juveniles
           Yaquina Bay              O   a�  :                                                     L - Larvae
            Alsea River             0      E- Eggs
           Siuslaw River                                           (    (X
          Umpqua River3   6                                        3
             Coos Bay               03 a
            Rogue River                                            (3   6
           Klamath River                   *  :                    *    *
          Humboldt Bay                                             0    0
              Eel River                                            <    �
           Tomales Bay
      Cent. San Fran. Bay *    C)    O   a�  �                                    O    � *        Includes Central San
                                                                                             Francisco, Suisun,
      South San Fran. Bay  (3        i   (3  (_ _____   w _ 43 __   Y _  3 __   and San Pablo bays.
          Elkhorn Slough
             Morro Bay
        Santa Monica Bay                        -_
          San Pedro Bay
           Alamitos Bay
           Anaheim Bay
           Newport Bay
            Mission Bay
          San Diego Bay
          Tijuana Esluary
                      A S J L E   A S J L E   A S J L E   A S J  L E  A S J L E

    Range                                                     infaunal.   Young  juveniles (fry and parr) are
    Overall: This species is recorded as far north as the    benthopelagic.  Parr become pelagic and acquire a
    Coppermine River in Arctic Canada, and south to    silver color when they transform into smolts (juveniles
    northeastern Hokkaido, Japan, and southern California    that migrate to the ocean). Smolts and ocean-dwelling
    (Ventura River) (Hart 1973, Scott andCrossman 1973).    and maturing juveniles (subadults), and adults are
    It is rarely found in fresh watersouthof the Sacramento-    pelagic (Alaska Department of Fish and Game 1985).
    San Joaquin river system of California (Eschmeyer et    Subadults and adults in rivers and streams are bottom-
    al. 1983). This species has been successfully introduced    oriented.
    to New Zealand and the Great Lakes (Scott and
    Crossman 1973).                                           Habitat
                                                           Type: The chinook salmon is an anadromous species.
    Within Studv Area: The chinook salmon is found in all    Eggs, alevins, fry, and parroccur in riverine areas from
    estuaries north of San Francisco Bay, except Tomales    just above the intertidal zone to altitudes of 2,268 m
    Bay, California (Table 1) (Monaco et al. 1990).           above sea level (Allen et al. in press).  Smolts are
                                                           riverine and estuarine. Ocean-dwelling juveniles are
    Life Mode                                                 neritic and epipelagic, and found within 128 m of the
    Eggs and alevins (yolk-sac larvae) are benthic and    surface (Fredin et al. 1977). Adults may be neritic and

                                                       161

    Chinook salmon continued
    estuarine, but are primarily riverine and may travel    Phinney 1986). Within these races are different "stocks"
    upstream over 4,700 km from the ocean.                which separate as they reach their natal streams
                                                           (Phinney 1986). In California, spring, fall, and winter
    Substrate: Eggs and alevins occur in spawning gravel    (December to February) runs exist, while the summer
    or cobble that is 1.3-10.2 cm in diameter (Reiser and    run is now extinct (Frey 1971, Moyle 1976). Fry and
    Bjornn 1979). Juveniles in fresh water are found over    smolts stay in fresh water from 1 to 18 months
    various substrates, ranging from silt bottoms to large   (Beauchamp et al. 1983).  Three types of juvenile
    boulders (Chapman and Bjornn 1968). Juveniles in   migrants have been defined according to their use of
    estuaries occur over mud, sand, gravel, and eelgrass   rivers and estuaries.  The first type, "subyearling
    (Zostera spp.) (Healey 1980a). Adults in marine waters    estuarine smolts", moves into estuaries early after
    show no sediment preference, but may be associated    hatching and rears there until late-spring or summer
    with gravel-cobble bottoms in rivers and streams (Alaska   when it moves to the ocean (Healey 1980a, 1982, Levy
    Department of Fish and Game 1985).                    and Northcote 1982, Levy 1984). The second type,
                                                           "subyearling riverine smolts", rears for less than one
    Phvsical/Chemical Characteristics: Eggs onlydevelop    year in the river before smolting and migrating to the
    in fresh water, but larvae can tolerate 15%o at hatching    estuary and spends only a little time in the estuary
    (Wagner et al. 1969). Three months after hatchingthey    (Reimers 1973, Healey 1982). Thethird type, "yearling
    can tolerate full seawater, with fastergrowing individuals   riverine smolts", rears for a year in the river and smolts
    better able to handle salinity changes (Wagner et al.   and migrates the spring after hatching (Healey 1982).
    1969). Juveniles and adults occur in fresh water to   Reimers (1973) also found two other life history types:
    euhaline waters.  Subadults (i.e., those that have    emergent fry that move directly downstream and into
    migrated to the marine environment), are found in   the ocean, and juveniles that stay in streams or rivers
    polyhaline to euhaline waters.  Successful egg    until fall, when they migrate directly to the ocean.
    incubation occurs from just above freezing to 20.0�C    Juvenile migration into estuaries has been reported to
    (Olsen and Foster 1955), however, best incubation   occur at night (Seiler et al. 1981) and during daylight
    temperatures are 5.0-14.4�C (Bell 1984). The upper    (Dawley et al. 1986). Juvenile chinook salmon may
    lethal temperature for the chinook salmon is 25.10C    move quickly through estuaries (Dawley et al. 1986) or
    (Brett 1952), but may be lower depending on other   residethereforupto 189 days (Simenstad et al. 1982).
    water quality factors (Ebel et al. 1971).  Eggs and    Peak estuarine outmigration usually occurs in spring
    alevins are found in areas with flows of 20-150 cm/sec    and summer, depending on life history (Healey 1982,
    and juveniles where flows are 0.5-60.0 cm/sec (at pool    Kjelson et al. 1982, Simenstad et al. 1982, Myers and
    edges). Adults can migrate upstream in flows up to   Horton 1982, Dawley et al. 1986, McCabe et al. 1986).
    2.44 m/sec (Thompson 1972).  Successful egg    Chinook salmon spend from 1-8 years (usually 3-4) in
    developmentrequiresreddstohaveadequatedissolved    the ocean before they return to their natal stream
    oxygen (.5.0 mg/I), water temperatures (4-140C),    (Wydoski and Whitney 1979). Some maystay in Puget
    substrate permeability, sediment composition (<25%    Sound until maturity (Simenstad et al. 1982). Upon
    fines,<6.4mmindiameter),surfaceflowsandvelocities,   entering the ocean, most stocks appear to migrate
    and low biochemical oxygen demand (Reiser and    north (Wright 1968) and many move into the Gulf of
    Bjornn 1979). Freshwater juveniles avoid waters with   Alaska (Hartt and Dell 1986). Chinook salmon produced
    <4.5 mg/I dissolved oxygen at 20�C (Whitmore et al.   in streams from the Rogue River (Oregon) and south
    1960). Migrating adults will pass through water with    appear to rear in the ocean off northern California-
    dissolved oxygen levels as low as 3.5-4.0 mg/I (Fujioka   southern Oregon, while chinook salmon produced in
    1970, Alabaster 1988, 1989). Excessive silt loads   streams from the Elk River (Oregon) and north rear
    (>4,000 mg/I) may halt chinook salmon movements or   primarily off British Columbia and Alaska (Cramer
    migrations (Reiser and Bjornn 1979). Silt can also    1987).  During its migrations, the chinook salmon
    hinder fry emergence, and limit benthic invertebrate    appears to use electromagnetic, olfactory, and visual
    (food) production (Reiser and Bjornn 1979). Low pH    cues for guidance (Hasler and Scholz 1983, Quinn
    decreases egg and alevin survival (Rombough 1983).    1984). Straying to spawning streams other than its
                                                           natal stream is very limited (Quinn and Fresh 1984).
    Miarations and Movements: Races of chinook salmon
    have been defined by when the adults migrate from the    Reproduction
    ocean to fresh water (Mason 1965). In the Columbia    Mode: This species is gonochoristic, oviparous, and
    River, spring chinook salmon enter from January    semelparous.  All adults die after spawning except
    through May, summer chinook salmon from June    some "jacks" (i.e., precocious males that mature early
    through mid-August, and fall chinook salmon during   in fresh water) (Miller and Brannon 1982). Eggs are
    August to November (Galbreath 1966, Netboy 1980,   fertilized externally.

                                                       162

                                                                                      Chinook salmon continued
    Matina/SDawnina: The spawning period is specific for   1976, Eschmeyer et al. 1983). Northern populations
    each run and/or stock, but can occur from April to   mature later, and spend more time in fresh water and
    February. For example, the Columbia River spring run   at sea (Scott and Crossman 1973). The largest chinook
    spawns from July to late September, the summer run   salmon recorded was 147cm in length and weighed 57
    from August to mid-November, and the fall run from    kg (Scott and Crossman 1973), but most are under
    SeptembertoJanuary (Fulton 1968, Netboy 1980, Bell   22.7 kg (Squire and Smith 1977).
    1984). In the Sacramento River, the winter run spawns
    duringApriltoJulyand otherruns fromJulyto December    Food and Feeding
    (Moyle 1976).  Chinook salmon normally spawn in   Trophic Mode: Larvae feed on their yolk. Juveniles,
    larger rivers and tributaries and in deeper water (10 m)    and adults are carnivorous, "opportunistic" feeders.
    and larger gravel than other Pacific salmon (Scott and
    Crossman 1973). Females make the redd by lying   Food Items: Juveniles in fresh water eat primarily
    sideways to the bottom and thrashing their tails. The    terrestrial and aquatic insects, Cladocera, amphipods
    redd can be 1.2-10.7 m in diameter (Chapman 1943).   and othercrustacea, and sometimes fish (Becker 1973,
    During spawning, a female will be attended by one    Higley and Bond 1973, Scott and Crossman 1973,
    dominant male and occasionally other subdominant    Craddock et al. 1976, Muir and Emmett 1988, Sagar
    males. Eggs and sperm are extruded simultaneously,    and Glova 1988).  In estuaries, juveniles consume
    after which the female will bury the eggs and move    gammarid amphipods, insects, harpacticoid copepods,
    upstream and repeat the process until spent.          mysids, decapod larvae and fish (Levy and Levings
                                                          1978, Levy et al. 1979, Healey 1980a, 1982, Kjelson et
    Fecundity:From2,000-14,000eggsarelaidperfemale,   al. 1982, Simenstad et al. 1982, Simenstad 1983,
    with 5,000 eggs per female being average (Rounsefell    McCabe et al. 1986). In the neritic zone, small chinook
    1957, Moyle 1976, Bell 1984). Fecundity depends on    salmon (those having recently migrated) feed on small
    female size, stream latitude, and subpopulation (Alaska   (larval and juvenile) fishes, decapod larvae, amphipods,
    Department of Fish and Game 1985).                    euphausiids, terrestrial insects, and other invertebrates
                                                          (Healey 1980b, Peterson et al. 1983, Emmett et al.
    Growth and Development                                1986). Largerchinook salmon (captured by sport and
    EaaSizeand Embrvonic Develooment:Chinooksalmon    commercial fishing) feed primarily on fishes [e.g.,
    eggs are spherical, nonadhesive, and the largest of all   northern anchovy (Engraulis mordax), scorpaenids,
    the salmonids (6.0-8.5 mm in diameter) (Rounsefell    Pacific herring (Clupeapallasl), and Pacific sand lance
    1957, Scott and Crossman 1973, Wang 1986).    (Ammodytes hexapterus)], euphausiids, decapod
    Embryonic development is indirect and external. The    larvae, squid, and other invertebrates (Silliman 1941,
    duration of incubation ranges from 33 to 178 days,    Merkell11957, Prakash 1962, Ito 1964, Hart 1973, Fresh
    depending on levels of dissolved oxygen, water   et al. 1981). Adults do not actively feed in fresh water.
    temperature, biochemical oxygen demand, substrate,
    channel gradient and configuration, waterdepth, water    Biological Interactions
    velocityanddischarge(Reiserand Bjornn 1979, Alaska    Predation: In fresh water, juveniles are eaten by many
    Department of Fish and Game 1985). Time of hatching    fishes [e.g., northern squawfish (Ptychocheilus
    is dependent on the spawning period, with fall-spawned    oregonensis), channel catfish (Ictalurus punctatus),
    eggs usually hatching in March and April (Columbia    coho salmon, Dolly Varden (Salvelinus malma), rainbow
    River) and eggs from winter-run fish hatching from May    trout (O. mykiss), cutthroat trout (0. clarke), smallmouth
    to August (Sacramento River) (Moyle 1976).            bass (Micropterus dolomieul), walleye (Stizostedian
                                                          vitreum), and sculpins] and birds [e.g., mergansers,
    Aae and Size of Larvae: Larval sizes range from 20-35   terns,osprey (Pandion haliaetus), and belted kingfisher
    mm total length (Wang 1986). Alevins remain in the   (Megacerylealcyon)] (Buchanan etal. 1981,Grayetal.
    gravel until the yolk sac is absorbed (usually 2-3    1982, Beauchamp et al. 1983, Maule and Horton
    weeks) (Scott and Crossman 1973, Wydoski and    1984). Intheoceanandestuaries,chinooksalmonare
    Whitney 1979).                                        prey for pelagic fishes, Pacific lamprey (Lampetra
                                                          tridentata), birds [e.g., common murre (Uria aalge),
    Juvenile Size Ranae: Juveniles are 2-152 cm (usually   and shearwaters (Puffinusspp.)], and marine mammals
    lessthan91 cm) in length, andfromafewgramsto61.4   [e.g., harbor seal (Phoca vitulina), sea lions, killer
    kg (usually less than 11.3 kg) (Wydoski and Whitney    whale (Orcinus orca)] (Simenstad et al. 1979, Fiscus
    1979, Allen et al. in press).                         1980, Beach et al. 1981, Alaska Department of Fish
                                                          and Game 1985). Adults in fresh water are eaten by
    Aae and Size of Adults: Maturity is reached between 1   bald eagle (Haliaeetus leucocephalus), bears, and
    and 9 years, with most maturing in 3-6 years (Moyle   other mammals (Scott and Crossman 1973).

                                                       163

    Chinook salmon continued
    Factors Influencina Ponulations: High mortality occurs    Allen, M.J., R. J. Wolotira, Jr., T. M. Sample, S. F. Noel,
    during the early freshwater life stages (eggs, fry, parr).   and C. R. Iten. (in press).  Salmonids: life history
    This mortality is caused by redd destruction, siltation   descriptions and brief harvest summaries for salmonid
    and destruction of spawning grounds, extremely high   species of the northeast Pacific Ocean and eastern
    or low watertemperatures, low dissolved oxygen, loss   Bering Sea. Tech. Memo., NOAA, NMFS, Northwest
    of cover, disease, and predation (Reiser and Bjornn    Alaska Fish. Cent., Seattle, WA.
    1979). Besides the above factors, man-made changes
    such as river flow reductions, the creation of dams and    Ames, J. 1983. Salmon stock interactions in Puget
    reservoirs, pollution, and logging practices, have    Sound: a preliminary look.  In M. A. Miller (editor),
    affected population abundances (Raymond 1979,    Southeast Alaska coho salmon research  and
    Netboy 1980, Stevens and Miller 1983). Estuaries    management review and planning workshop, May 18-
    appearto play a vital role in chinook salmon life history   19,1982, p. 84-95. Alaska Dept. Fish Game, Juneau,
    (MacDonald et al. 1988). In the ocean, this species is   AK, 109 p.
    affected by disease, predation, food availability, and
    oceanographicconditions. Overfishing has not allowed    Beach, R. J., A. C. Geiger, S. J. Jeffries, and S. D.
    optimal spawning escapement and has reduced the   Tracy. 1981. Marine mammal- fishery interactions on
    age and size structure ofsome populations (Fraidenburg   the Columbia River and adjacent waters, 1981. Second
    and Lincoln 1985). Also, the high-seas gill net fishery   Annual Rep. to NOAA, NMFS, Northwest and Alaska
    for squid is taking an unknown number of chinook    Fish. Cent., Seattle, WA. Wash. Dept. Game, Olympia,
    salmon. The release of millions of juvenile chinook    WA, 186 p.
    salmon by public and private hatcheries has helped
    maintain some runs (Phinney 1986), and the United    Beauchamp, D. A., M. F. Shepard, and G. B. Pauley.
    States-Canada Salmon Interception Treatyshould allow    1983. Species profiles: life histories and environmental
    more escapement in the future. The survival of hatchery    requirements of coastal fishes and invertebrates (Pacific
    smolts to maturity is influenced by time of release, size   Northwest) - chinook salmon. U.S. Fish Wildl. Serv.,
    of release, health of fish, degree of smoltification at   Div. Biol. Serv., FWS/OBS-82/11.6. U.S. Army Corps
    release, release location, and ocean conditions    Eng., TR EL-82-4, 15 p.
    (Vreeland 1988). In rivers and streams, juveniles are
    not as aggressive as coho salmon and steelhead    Becker, C. D. 1973. Food and growth parameters of
    juveniles (Wydoski and Whitney 1979).  However,    juvenile chinook salmon, Oncorhynchus tshawytscha,
    adults typically spawn in deeper water and use larger   in central Columbia River. Fish. Bull., U.S. 71:387-400.
    gravel than other salmonids (Scott and Crossman
    1973). The chinook salmon may compete with other    Beckman, B. J., and W. S. Zaugg. 1988.  Copper
    salmonid species in the marine environment (Ames    intoxication in chinook salmon (Oncorhynchus
    1983) and it is known to feed on the same food as coho    tshawytscha) induced by natural spring water: effects
    salmon (Emmett et al. 1986). In estuaries, juveniles   ofgillNa+-K+ATPase, hematocrit, andplasmaglucose.
    are associated with many other fish species that often   Can. J. Fish. Aquat. Sci. 45:1430-1435.
    feed on similar prey items (McCabe et al. 1983).
                                                          Bell, M. C. 1984. Fisheries handbook of engineering
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    Alabaster, J. S.  1988.  The dissolved oxygen    Corps Eng., North Pac. Div., Portland, OR,  290 p.
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                                                       164

                                                                                     Chinook salmon continued

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                                                          oxygen content in the Duwamish River estuary on
    Ebel, W. J., E. M. Dawley, B. H. Monk. 1971. Thermal    migrating adult chinook salmon. M.S. Thesis. Univ.
    tolerance of juvenile Pacific salmon and steelhead    Wash., Seattle, WA, 77 p.
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    Fish. Bull., U.S. 69(4):833-843.                       Fulton, L. A. 1968. Spawning areas and abundance of
                                                          chinook salmon (Oncorhynchus tshawytscha) in the
    Emmett, R. L., D. R. Miller, T. H. Blahm. 1986. Food    Columbia Riverbasin - past and present. U.S. Fish and
    of juvenile chinook, Oncorhynchus tshawytscha, and    Wildl. Serv., SSRF 571, 26 p.
    coho, 0. kisutch, salmon off the northern Oregon and
    southern Washington coasts, May-September 1980.    Galbreath, J. L. 1966.  Timing of tributary races of
    Calif. Fish Game 72(1):38-46.                          chinook salmon through the lower Columbia River
                                                          based on analysis of tag recoveries. Fish. Comm.
    Eschmeyer, W. N., W. S. Herald, and H. Hammann.    Oregon, Res. Briefs 12(1):58-80.
    1983. A field guide to Pacific coast fishes of North
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                                                          Huntington, and D. E. Palmer. 1982. Feeding activity,
    Fiscus. C. H.  1980.  Marine mammal-salmonid    rate of consumption, daily ration, and prey selection of
    interactions: a review.  In W. J. McNeil and D. C.    major predators in the John Day pool. Annual Rep. to
    Himsworth (editors), Salmonid ecosystems of the North    Bonneville PowerAdm., Portland, OR, 81 p. Available
    Pacific, p. 121-132. Oregon State Univ. Press, Corvallis,    U.S. Fish and Wildl. Serv., Cook, WA.
    OR.
                                                          Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    Fisher, J. P., and W. G. Pearcy. 1985. Studies of    Board Can., Bull. No. 180, 740 p.

                                                       165

    Chinook salmon continued
    Hartt, A. C., and M. B. Dell.  1986.  Early oceanic    Levy, D. A., and T. G. Northcote.  1982.  Juvenile
    migrations and growth of juvenile Pacific salmon and    salmon residency in a marsh area of the Fraser River
    steelhead trout.  Internat. North Pac. Fish. Comm.,    estuary. Can. J. Fish. Aquat. Sci. 39:270-276.
    Bull. No. 46:1-105.
                                                           Levy, D. A., T. G. Northcote, and G. J. Birch. 1979.
    Hasler, A. D., and A. T. Scholz.  1983.  Olfactory    Juvenile salmon utilization of tidal channels in the
    imprinting and homing in salmon.  Springer-Verlag,    Fraser River estuary, British Columbia.  Westwater
    Berlin, 134 p.                                          Res. Cent. Tech. Rep. 23, Univ. British Columbia,
                                                           Vancouver, B.C., 70 p.
    Healey, M. C. 1980a. Utilization of the Nanaimo River
    estuary by juvenile chinook salmon, Oncorhynchus    Macdonald, J. S., C. D. Levings, C. D. McAllister, U. H.
    tshawytscha. Fish. Bull., U.S. 77(3):653-668.           M. Fagerlund, and J. R. McBride.  1988.  A field
                                                           experiment to test the importance of estuaries for
    Healey, M. C. 1980b. The ecology of juvenile salmon    chinook salmon (Oncorhynchus tshawytscha) survival:
    in Georgia Strait, British Columbia. InW. J. McNeil and    short-term results. Can. J. Fish. Aquat. Sci. 45(8):1366-
    D. C. Himsworth (editors), Salmonid ecosystems of the    1377.
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    Corvallis, OR.                                          Mason, J. E. 1965. Salmon of the north Pacific Ocean
                                                           - Part IX. Coho, chinook and masu salmon in offshore
    Healey, M.C. 1982. JuvenilePacificsalmoninestuaries:    waters. Internat. North Pac. Fish. Comm., Bull. No. 6,
    the life support system.  In V. S. Kennedy (editor),    135.
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    New York, NY.                                           Maule, A. G., and H. F. Horton. 1984. Feeding ecology
                                                           of walleye, Stizostedion vitreum vitreum, in the mid-
    Higley, D. L., and C. E. Bond.  1973. Ecology and    Columbia River, with emphasis on the interactions
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                                                           McCabe, G. T., Jr., R. L. Emmett, W. D. Muir, and T. H.
    Ito, J. 1964. Food and feeding habits of Pacific salmon    Blahm. 1986. Utilization ofthe Columbia River estuary
    (genus Oncorhynchus) intheir ocean life. Bull Hokkaido    by subyearling chinook salmon. Northw. Sci. 60(2):113-
    Reg. Fish. Res. Lab. 29:85-97. (Fish. Res. Board Can.    124.
    Transl. Ser. 1309).
                                                           McCabe, G. T., Jr., W. D. Muir, R. L. Emmett, and J. T.
    Kjelson, M. A., P. F. Raquel, F. W. Fisher. 1982. Life   Durkin.  1983.  Interrelationships between juvenile
    history of fall-run chinook salmon, Oncorhynchus    salmonids and nonsalmonid fish in the Columbia River
    tshawytscha in the Sacramento-San Joaquin estuary,    estuary. Fish. Bull., U.S. 81(4):815-826.
    California. In V. S. Kennedy (editor), Estuarine
    comparisons, p. 393-411 . Academic Press, New York,    McGinnis, S. M. 1984. Freshwaterfishes of California.
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    Levy, D. A. 1984. Commentary: variations in estuary    Merkel, T. J.  1957.  Food habits of king salmon,
    utilization among juvenile chinook salmon populations.    Oncorhynchus tshawytscha (Walbaum), in the vicinity
    In W. G. Pearcy (editor), The influence of ocean    of San Francisco, California.  Calif. Fish. Game
    conditions on the production of salmonids in the North    43(4):249-270.
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    001, Oregon State Univ., Corvallis, OR.                 Miller, R. J., and E. L. Brannon. 1982. The origin and
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    Levy, D. A., and C. D. Levings. 1978. A description of   In E. L. Brannon and E. O. Salo (editors). Salmon and
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    British Columbia: relative abundance, seasonal    Fish., Univ. Wash., Seattle, WA.
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                                                          impoundments on migrations of juvenile chinook salmon
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    migrating salmonid smolts passing Bonneville Dam in   Am. Fish. Soc. 108(6):505-529.
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    comparisons, p. 377-392. Academic Press, NewYork,    requirements of anadromous salmonids.  In W. R.
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                                                       168

    169

       Hypomesus pretiosus
       Adult

                  5cm

    Common Name: surf smelt                                 Range
    Scientific Name: Hypomesus pretiosus                    Overall: The surf smelt's overall range is from Long
    Other Common Names: Pacific surf smelt, silver   Beach, California, to southeast Alaska (Frey 1971).
    smelt
    Classification (Robins et al. 1980)                     Within Study Area: This species is occasionally found
    Phylum: Chordata                                        in California estuaries (Moyle 1976), but is seasonally
    Class: Osteichthyes                                     common to abundant in Oregon and Washington
    Order: Salmoniformes                                    estuaries (Table 1) (Monaco et al. 1990).
    Family: Osmeridae
                                                            Life Mode:
    Value                                                   Eggs are benthic. Larvae, juveniles, and adults are
    Commercial: The surf smelt is commercially fished in   pelagic but remain principally inshore. Except in Puget
    California and Washington. More than 4 million were    Sound and adjacent areas, this is a nearshore coastal
    taken in California in 1958 (Frey 1971). An average of   species which does nottypically spawn in estuaries but
    51 t are taken annually in Washington, most of which    utilizesthem forfeeding and rearing. Itdoes not appear
    are caught in Puget Sound (Trumble 1983).               to form large pelagic schools likethe northern anchovy
                                                            (Engraulis mordax). However, schools of surf smelt
    Recreational: This species is considered an excellent    are often common in Northwest estuaries.
    food fish and is captured by recreational fishermen in
    Washington, Oregon, and California.  It is taken by    Habitat
    jump net (in California), jig, and dip net. The numbers    Iype: Eggs are laid intertidally on beaches. Larvae,
    taken by recreational anglers areunknown, butthought    juveniles, and adults live in neritic waters.
    to be substantial.
                                                            Substrate: Spawning adults select substratesof coarse
    Indicator of Environmental Stress: The surf smelt    sandwithfinegravel(Trumble1983). Larvae, juveniles,
    spawns at specific beach sites where appropriate    and adults can be found over a variety of substrates.
    physical conditions for spawning exist. Hence, loss or
    alteration of these spawning sites can be very    Physical/Chemical Characteristics: All life stages are
    detrimental to populations of this species.             found in estuarine and marine waters. Beaches used
                                                            for spawning typically have some freshwater seepage
    Ecological: This species is important prey for many    and are usually shaded by trees or bluffs (Schaefer
    fishes, birds, and mammals. Puget Sound stocks are    1936). Watertemperature and salinity of the spawning
    genetically different from coastal stocks (Kilambi 1965,    areas do not appea rto affect spawning activity, but tide
    Kilambi et al. 1965).                                   stage and time of day do. Survival of embryos does not
                                                            appearto be significantly different at salinities of 20, 25,
                                                            or 30'o (Middaugh et al. 1987).

                                                        170

                                                                                           Surf smelt continued
                                                           Matina/SDawnina: Spawning populations can be found
     Table 1. Relative abundance of surf smelt in         nearly year-round along the Pacific coast. However,
              32 U.S. Pacific coast estuaries.            they spawn at specific beaches at specific times of the
                         Life Stage                        year (Penttila 1978). Spawning occurs primarily at high
            Estuary    A S J L E                          tide and early ebb, from late afternoon to evening
             Puget Sound         6 6: 1    �   Relative abundance:  (Schaefer 1936, Thompson et al. 1936, Yap-Chiongco
              HoodCanal a6 * 1 *      6*    *   Highlyabundant  1941). Before a spawning "run", schools appear in the
               Skagit Bay (   � a          Abundant        water 0.9-1.2 m from the edge of the beach. During
             Grays Harbor C    O O    0   Common           spawning, a female (usually accompanied by 2 to 5
                                       '4  Rare
              Willapa Bay O    O O   Blank Not present     males) moves to the highest point reached by a wave.
           Columbia River O      O0                       As they reach the shore, the fishes release their
             Nehalem Bay �  a   O                          gametes. This process occurs in 2.5-5.0 cm of water
            Tillamook Bay �    � O    Life stage:         and takes about 5 to 10 seconds (Loosanoff 1937).
              NetartsBay "4    � O    A-Adults             Eggs are usually concentrated at the 2.1-3.4 m tidal
              Siletz River  C  J J nin  adults            levels (upper intertidal zone) (Penttila 1978, Middaugh
                                      J - Juveniles
             YaquinaBay O    � O      L-Larvae            et al. 1987).  Eggs are adhesive and stick to sand
              AlseaRiver 0     O      E - Eggs             grains and wave action covers them with a thin layer of
             SiuslawRlver    6 O0                          sand. Adultsusuallyeatverylittleduringspawning,but
            Umpqua River �    � O                         do not die after spawning (Loosanoff 1937).
               CoosBay i    � O
             Rogue River v4      0                        Fecundity: Females release eggs in batches and
            Klamath River      O0                         spawning can last for several days. Females usually
            Humboldt Bay O    O O                         produce 15,000-20,000 eggs, but can produce from
                Eel River 0    0 0                         1,300-37,000 eggs (depending on body size) (Leong
             Tomales Bay O   0  1967).
        Cent. San Fran. Bay *  '4 i  Includes Central San
                                      Francisco, Suisun,
        SouthSanFran. Bay  "4   '    and San Pablo bays.   Growth and Development
            Elkhom Slough   '4                             Eca Size and Embrvonic DeveloPment: Fertilized eggs
               Morro Bay                                   are spherical and about 1.0-1.2 mm  in diameter
         Santa Monica Bay
            San~ Pedro Bay                                 (Schaefer 1936). Eggs adhere to gravel substrates by
             SAamitos n Bay                               the adhesive zona radiata membrane which ruptures
                     Anaheim Say                           and turns inside out at the time the eggs are fertilized.
             Newport Bay                                  Embryonic development is indirect and external
              Mission Bay                                  (Garrison and Miller 1982). After several days embryos
            San Diego Bay                                  detach from the spawning substrates and are washed
           Tijuana Estuary                                seaward and down into the gravel substrate in the
                       A S J L E                          intertidal zone (Middaugh et al. 1987). Hatching occurs
                                                           from 8.5 to 30 days after incubation (depending on
                                                           temperature) and may be initiated by mechanical or
    MiarationsandMovements:Migrationsandmovements    chemical stimuli.  Eggs are stimulated to hatch by
    have not been studied. Although specific spawning    immersion in water (high tide) (Loosanoff 1937). At
    sites are used,there is no information regardingwhether    extremely low temperatures (e.g., during winter) the
    fish return to their natal spawning sites. The seasonal    incubation period may be 90 days or more (Middaugh
    utilization of estuaries byjuveniles and adults probably   et al. 1987).
    relates to food abundance and refuge from predators.
    At the beginning of a spawning run, schools are usually    Aae and Size of Larvae: Larvae are 5.0-6.5 mm long at
    composed of individuals of the same sex; female    hatching. Postlarvae are 17-35mm in total length (TL)
    schools usually arrive before male schools (Loosanoff    (Yap-chiongco 1941).
    1937). Later, as more schools arrive, the unisexual
    character of the schools is lost.                      Juvenile Size Ranae: Juveniles range from 35 mm to at
                                                           least 85 mm TL. Scales first appear when fish are 55-
    Reproduction                                           68 mm TL.
    Mode: The surf smelt is gonochoristic, oviparous, and
    iteroparous. It has external egg fertilization and probably    Aae and Size of Adults: Adults range from 81-178 mm
    spawns annually after reaching maturity.               TL. Most mature in their second year but some are
                                                           gravid in their first. Individuals older than three years

                                                       171

    Surf smelt continued

    are rare, but they may reach 5 years old. Females are    methods. J. Mar. Biol. Assoc. India 7(2):364-368.
    typically largerthan similarly-aged males (Yapchiongco
    1949). Both sexes have asymmetrical gonads, with the    Leong, Choon-Chiang. 1967. Fecundity of surf smelt,
    left gonad being much more developed (Yap-chiongco    Hypomesus pretiosus (Girard) in the state of
    1941). Males have pearl organs (small protuberances    Washington. M.S. Thesis, Univ. Wash., Seattle, WA,
    on their snouts) during the breeding season while    99 p.
    females do not (Yapchiongco 1949). Males are dull
    olive green on their back while females are bright    Loosanoff, V. L. 1937. The spawning run of the Pacific
    metallic green (Yap-chiongco 1941).                      surf smelt, Hypomesus pretiosus (Girard).  Internat.
                                                           Rev. Hydrobiol. Hydrogr. 36(1-2):170-183.
    Food and Feeding
    Troohic Mode: Larvae, juveniles, and adults are    Middaugh, D. P., M. J. Hemmer, and D. E. Penttila.
    planktivorous carnivores (typically zooplanktivorous).    1987.  Embryo ecology of the Pacific surf smelt,
                                                           Hypomesuspretiosus (Pisces: Osmeridae). Pac. Sci.
    Food Items: The surf smelt feeds primarily on planktonic    41 (1-4):44-53.
    crustacea, including amphipods, euphausiids,
    copepods, cladocerans, crustacean larvae, and some    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    larval fish (Hart 1973).                                 Nelson. 1990. Distribution and abundance of fishes
                                                           and invertebrates in west coast estuaries, Volume I:
    Biological Interactions                                  data summaries.   ELMR  Rep. No. 4. Strategic
    Predation: This species is eaten by many fishes,    Assessment Branch, NOS/NOAA, Rockville, MD,
    including Pacific salmon (Oncorhynchus spp.), lingcod    240 p.
    (Ophiodon elongatus), and striped bass (Morone
    saxatilis) (Frey 1971).  It is also commonly eaten by    Moyle, P. B. 1976. Inland fishes of California. Univ.
    birds and marine mammals.                                Calif. Press, Berkeley, CA, 405 p.

    Factors Influencina Pooulations: Eggand larval mortality    Penttila, D. 1978. Studies of surf smelt (Hypomesus
    can result from thermal stress, and desiccation.    pretiosus) in Puget Sound.  Tech. Rep. 42, Wash.
    Predation can be high (Penttila 1978, Garrison and    Dept. Fish., Olympia, WA, 47 p.
    Miller 1982) and probably plays a large role in
    determining population size.  The specific beaches    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    used for spawning can be ruined by pollution,    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    bulkheading, and other habitat alterations.              of common and scientific names of fishes from the
                                                           United States and Canada. Am. Fish. Soc. Spec. Publ.
    References                                               No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Frey, H. W. 1971. California's living marine resources    Schaefer, M. B. 1936. Contribution to the life history of
    and their utilization.  Calif. Dept. Fish Game,    the surf smelt Hypomesus pretiosus in Puget Sound.
    Sacramento, CA, 148 p.                                   Wash. Dept. Fish., Biol. Rep. 35B:1-45.

    Garrison, K. J., and B. S. Miller. 1982. Review of the    Thompson, W. F., F. H. Bell, L. P. Schultz, H. A.
    early life historyof PugetSoundfishes. Fish. Res. Inst.,    Dunlop, and R. Van Cleve. 1936. The spawning of the
    School Fish., Univ. Wash., Seattle, WA, 729 p, (FRI-    silver smelt, Hypomesus pretiosus. Ecology 17:148-
    UW-8216).                                                168.

    Hart, J. L. 1973. Pacific Fishes of Canada. Fish. Res.    Trumble, R. J. 1983. Management plan for baitfish
    Board Can., Bull. No. 180, 740 p.                        species in Washington State. Prog. Rep. 195, Wash.
                                                           Dept. Fish., Olympia, WA, 106 p.
    Kilambi, R. V. 1965. Heterogeneity among three
    spawning populations of the surf smelt, Hypomesus    Yap-Chiongco, J. V. 1941. Hypomesus pretiosus: Its
    pretiosus (Girard) in the state of Washington.  Ph.D.    development and early life history. Ph.D. Thesis, Univ.
    Thesis, Univ. Wash., Seattle, WA, 154 p.                 Wash., Seattle, WA, 123 p.

    Kilambi, R. V., F. M. Utter, and A. C. DeLacy. 1965.    Yap-Chiongco, J. V. 1949. Hypomesus pretiosus: Its
    Differentiation of spawning populations of the surf    development and early life history. Nat. Appl. Sci. Bull.
    smelt, Hypomesus pretiosus (Girard) by serological    9(1):3-108..

                                                       172

    173

        Spirinchus thaleichthys
       Adult

             2cm

    Common Name: longfin smelt                             Within Study Area: It is found in most Pacific coast
    Scientific Name: Spirinchus thaleichthys                estuaries from San Francisco Bay (Moyle 1976) north
    Other Common Names: Pacific smelt, long-finned    to Puget Sound, Washington (Garrison and Miller
    smelt, Sacramento smelt                                 1980) (Table 1).
    Classification (Robins et al. 1980)
    Phylum: Chordata                                        Life Mode
    Class: Osteichthyes                                     Eggs are benthic and adhesive. Larvae and juveniles
    Order: Salmoniformes                                    are primarily pelagic, while adults are both pelagic and
    Family: Osmeridae                                      demersal.

    Value                                                   Habitat
    Commercial:The longfin smelt is occasionally captured    Iype: Eggs are benthic and riverine or upper estuarine.
    incidentally with other smelt species, and marketed    Larvae are pelagic and occur in riverine-marine waters,
    with these species as "smelt" (Skinner 1962). The    but are most often found in estuarine environments.
    longfin smelt is seasonally sold at markets in California's    Juveniles are primarily pelagic and estuarine. Adults
    San Francisco Bay area (Wang 1986).                     are pelagic but are often found near the bottom in
                                                           estuarine and marine waters.
    Recreational: Presently, only a very limited recreational
    fishery exists.                                         Substrate: Type of spawning substrate has not been
                                                           positively identified, but is thought to be sandy-gravel
    Indicatorof Environmental Stress: Information regarding    areas with sand oraquatic plants (Wang 1986). Nektonic
    population sizes and fluctuations are limited. However,    life stages occur over a variety of substrates.
    since all life stages use estuaries, any estuarine
    alterations potentially affect this species. Freshwater    Physical/Chemical Characteristics: The longfin smelt
    flow into estuaries is important forthis species (Stevens    is an anadromous, euryhaline species. However, the
    and Miller 1983, California Department of Fish and    existence of landlocked freshwater populations
    Game 1987).                                             indicates that this species does not need marine/
                                                           estuarine waters to complete its life cycle. Most early
    Ecological: The longfin smelt is abundant in many    life history information pertains to landlocked
    Pacific coast estuaries and is consumed by numerous    populations, thus very littledatais available for estuarine/
    marine and estuarine vertebrates.                       marine populations (Garrison and Miller 1980).

    Range                                                   Miarations and Movements: Juveniles and adults
    Overall: This species' overall range is from Monterey    appear to move to lower estuarine/marine areas in
    Bay, California, to Prince William Sound, Alaska    spring and summer, and to upper estuarine areas in
    (Eschmeyer et al. 1983).                               fall. In winter, adults move to freshwater spawning

                                                        174

                                                                                         Longfin smelt continued

                                                           Fecundity: A femalecan produce an average of 18,000-
     Table 1. Relative abundance of longfin smelt           24,000 eggs (Hart 1973 Moyle 1976), although fish
              in 32 U.S. Pacific coast estuaries.          from landlocked populations may produce much fewer
                         Life Stage                         (Wydoski and Whitney 1979).
            Estuary    A S J L E
             PugetSound O    o00      Relative abundance:   Growth and Development
              Hood Canal               �   Highly abundant  Eao Size and Embrvonic DeveloPment: Fertilized eggs
              Skagi Bay 0         0   C   Abundant        are spherical, 1.2 mm  in diameter, and adhesive
            Grays Harbor C  a   . a   0   Common           (Dryfoos 1965). Eggs incubated at 70C hatch in 40
             WiliapaBay    C   *  Blank Not Rare           days(Dryfoos1965). Embryonicdevelopmentisindirect
           ColumbiaRiver              ank Not present       and external.
            Nehalem Bay
            Tillamook Bay  i          Life stage:           Aae and Size of Larvae: At hatching, larvae are reported
              Netarts Bay              A-Adults             to be 5.3-9.8 mm long (Dryfoos 1965, Moulton 1970,
              Siletz River            S-Spawning adults     Wang 1986).  Metamorphosis to juvenile probably
                                      J - Juveniles
             Yaqulna Bay O 0 0 o 0    L- Larvae             begins in 30-60 days, depending on temperature.
              Alsea River              E - Eggs
            Siuslaw River   'v                             Juvenile Size Ranae: Juveniles range from 22 mm to
            UmpquaRiver i    i4                             approximately 88 mm long (Moulton 1970, 1974).
               CoosBay 00 0 0 0
             Rogue River                                   Aae and Size of Adults: Spawning occurs at age 2, with
            Klamnath River o    O                           adults being 8.8-15.2 cm in total length, but averaging
            Humboldt Bay o    0   0           a             10.0 cm. (Moulton 1974).  Size, age, and possibly
                Eel River o O O O o                         watertemperature influence age at maturation (Moulton
             Tomales Bay                                    1974).
       Cent. San Fran. Bay  (*  � � . (   Includes Central San
                                      Francisco, Suisun.
       South San Fran. Bay  O    0 0  and San Pablo bays.   Food and Feeding
           Elkhom Slough                                   Trophic Mode: Larvae, juveniles, and adults are
               Morro Bay                                    carnivorous planktivores.
         Santa Monica Bay
           San Pedro Bay                                    Food Items: Larvae probably consume zooplankton
             Alamitos Bay
                                                           and some phytoplankton.  Juveniles and adults eat
             Anaheim Bay
             Newport Bay                                    calanoid copepods, cladocerans, amphipods, and other
              MissioneBay                                   small crustaceans (Moyle 1976).  Adults also prey
             Mission Bay
            San Diego Bay                                   heavily on the mysid Neomysis mercedis.
           Tijuana Estuary
                       A S J L E                            Biological Interactions
                                                            Predation: Larvae, juveniles, and adults are eaten by
                                                            predatory fishes, birds, and marine mammals. The
    areas (Ganssle 1966).  Adults show diel vertical    longfinsmeltis an importantyear-roundpreyforharbor
    movements, being found deep during the day and in   seals (Phoca vitulina) in the Columbia River estuary
    theupperwatercolumnatnight(WydoskiandWhitney    (Jeffries 1984).  It is probably an important prey for
    1979).                                                  piscivorous birds such as gulls and terns.

    Reproduction                                             Factors Influencina PoDulations: Larval and juvenile
    Mode: The longfin smelt is gonochoristic, oviparous,    survival appears to be the major determinant of adult
    and iteroparous. It has external egg fertilization and    population size. In San Francisco Bay, juvenilesurvival
    spawns in batches.                                       appears to correlate directly with freshwater inflow
                                                            (California Department of Fish and Game 1987). Pulses
    Matino/SDawnina: Spawning occurs in freshwater areas    of freshwater inflow can alter the estuarine distribution
    at night during winter (October-March), when river   and abundance of this species. In San Francisco Bay,
    temperatures are 4.4-7.2�C (Wydoski and Whitney    there is a positive association between spring riverflow
    1979), 5.6-6.70C (Moulton 1974), and 7.0-14.50C (Wang    and longfin smelt abundance (Stevens and Miller 1983,
    1986). During spawning, eggs and sperm are released    Armor and Herrgesell 1985, California Department of
    near the substrate. Once fertilized, the eggs become    Fish and Game 1987).
    adhesive. Almost all adults die after spawning.

                                                        175

    Longfin smelt continued
    References                                            Skinner, J. E. 1962. An historical review of the fish and
                                                           wildlife resources of the San Francisco Bay area.
    Armor, C., and P. L. Herrgesell. 1985. Distribution and   Water Proj. Bureau Rep. 1, Calif. Dept. Fish Game,
    abundance of fishes in the San Francisco Bay estuary    Sacramento, CA, 255 p.
    between 1980 and 1982. Hydrobiol. 129:211-227.
                                                           Stevens, D. E., and L. W. Miller. 1983. Effects of flow
    California Department of Fish and Game. 1987. Delta   on abundance of young chinook salmon, American
    outflow effects on the abundance and distribution of   shad, longfin smelt, and delta smelt in the Sacramento-
    San Francisco Bay fish in invertebrates, 1980-1985.    San Joaquin River system. N. Am. J. Fish. Manag.
    Exhibit 60, entered by the Calif. Dept. Fish Game for   3:425-437.
    the State Water Resources Control Board 1987 Water
    Quality/Water Rights ProceedingontheSan Francisco   Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Bay/Sacramento-San Joaquin Delta. Calif. Dept. Fish   Joaquin estuary and adjacent waters, California: A
    Game, Stockton, CA, 345 p.                            guide to the early life histories. Tech. Rep. No. 9.
                                                           Interagency ecological study program for the
    Dryfoos, R. L. 1965. The life history and ecologyof the    Sacramento-San Joaquin estuary. Calif. Dept. Water
    longfin smelt in Lake Washington. Ph.D. Thesis, Univ.    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    Wash., Seattle, WA, 159 p.                            and U.S. Fish Wildl. Serv. various pagination.

    Eschmeyer, W. N., W. S. Herald, and H. Hammann.    Wydoski, R.S.,and R. R.Whitney. 1979. Inland fishes
    1983. A field guide to Pacific coast fishes of North   of Washington.  Univ. Wash. Press, Seattle, WA,
    America. Houghton Mifflin Co., Boston, MA, 336 p.     220 p.

    Ganssle, D. 1966. Fishes and decapods of the San
    Pablo and Suisun Bays. In D. W. Kelley (compiler),
    Ecological studies of the Sacramento-San Joaquin
    estuary. Calif. Fish Game, Fish Bull. 133:64-94.

    Garrison, K. J., and B. S. Miller. 1980. Review of the
    early life historyof Puget Sound fishes. Fish. Res. Inst.,
    Univ. Wash., Seattle, WA., 729 p. (FRI-UW-8216).

    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    Board Can., Bull. No.180, 740 p.

    Jeffries, S. 1984. Marine mammals of the Columbia
    River estuary. Col. Riv. Est. Data Dev. Prog., CREST,
    Astoria, OR, 62 p. plus appendices.

    Moulton, L. L. 1970. The 1970 longfin smelt spawning
    run in Lake Washington with notes on egg development
    and changes in the population since 1964. M.S.
    Thesis, Univ. Wash., Seattle, WA, 84 p.

    Moulton, L. L. 1974. Abundance, growth, and spawning
    of the longfin smelt in Lake Washington. Trans. Am.
    Fish. Soc. 103(1):46-52.

    Moyle, P. B. 1976. Inland fishes of California. Univ.
    Calif. Press, Berkeley, CA, 405 p.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

                                                        176

    177

        Thaleichthys pacificus
       Adult

                 5cm

    Common Name: eulachon                                Ecological: The eulachon is the largest smelt along the
    Scientific Name: Thaleichthys pacificus               Pacific coast of North America and a prey species for
    Other Common Names: candlefish, oilfish, small fish,   many marine vertebrates.
    salvation fish, fathom fish (Scott and Crossman 1973)
    Classification (Robins et al. 1980)                   Range
    Phylum: Chordata                                     Overall: This species is found from the Klamath River,
    Class: Osteichthyes                                  California, along the Pacific coast to the eastern Bering
    Order: Salmoniformes                                 Sea in Bristol Bay, Alaska, and the Pribilof Islands
    Family: Osmeridae                                    (Scott and Crossman 1973). A few have been found
                                                          down to Bodega Head, California (Odemar 1964).
    Value
    Commercial: Major commercial runs occur in the   Within Studv Area: Major runs occur in the Columbia
    Columbia River and its tributaries, and the Klamath    and Klamath Rivers (Table 1), while many othercoastal
    River, California(Moyle1976). Thisspeciesiscaptured   rivers support small runs (Monaco et al. 1990).
    by gill net, trawl, and dip net. The 1968-69 lower
    Columbia River fishery (454 t) was estimated to be    Life Mode
    worth more than $280,000 (Snyder 1969). In 1985,   The eulachon is an anadromous species. Eggs are
    over 907 t were landed in the Columbia River (Bohn    demersal and attach to substrate. Larvae, juveniles,
    and Mclsaac 1986). Almost 862 t were landed in the   and adults are pelagic.
    lower Columbia River in 1987 (Mclsaac and Bohn
    1988).                                               Habitat
                                                          Type: Eggs occur in fresh water. Larvae are found in
    Recreational: The eulachon's annual spawning run   rivers, estuaries, and the marine neritic zone. Juveniles
    supportsapopularrecreationaldipnetfishery. Twenty    and adults are found in the marine neritic zone at
    years ago, the sport fishery of the Columbia River and    various depths (Barraclough 1964).  During their
    its tributaries had an estimated economic value of   spawning migration, adults are found near the bottom
    $570,000 (Snyder 1969). In manyyears the numberof    of estuarine and riverine channels.
    smelt harvested by the recreational fishery on the
    Columbia River and its tributaries equals the number    Substrate: Eggs are deposited in areas of pea-sized
    harvested commercially.                              gravel and/or semi-sandy areas with sticks and debris
                                                          (Smith and Saalfeld 1955).
    Indicator of Environmental Stress: All life stages are
    very sensitive to changes in temperature (Blahm and    Phvsical/Chemical Characteristics: Spawning occurs
    McConnell 1971).  However, information regarding   in riverine areas with moderate water velocities and at
    tolerances to chemical pollution is limited.          temperatures from 4-10�C. Water temperatures colder
                                                          than 40C appear to slow or stop adult migrations.

                                                       178

                                                                                             Eulachon continued

                                                            at night and do not build nests (Parente and Snyder
     Table 1. Relative abundance of eulachon in            1970, Garrison and Miller 1980).
               32 U.S. Pacific coast estuaries.
                         Life Stage                         Fecundity: Approximately 7,000-31,000 eggs are laid,
            Estuary    A S J L El                          depending on female size (Parente and Snyder 1970).
             Puget Sound 5         I Relative abundance:
              Hood Canal               *   Highly abundant  Growth and Development
              Skagit Bay          I |     Abundant         Eca  Size and Embrvonic Develorment: Eggs are
            Grays Harbor C    C       O0  Common           spherical and approximately 1 mmindiameter(Parente
             WillapaBay O     O       q   Rare             and Snyder 1970).  Mature eggs have double
           Columbia River 8 3        Blank Notpresent      membranes.  After fertilization, the outer membrane
            Nehalem Bay                                    ruptures and turns inside out with the outer membrane
            Tillamook Bay             Life stage:          remaining attached to the inner membrane at a small
              Netarts Bay             A -      Adults       spot. The adhesive edges of the outer membrane stick
                                      S - Spawning adults
              Silelz River           J -Juveniles          to sand or other particles, hence the egg is supported
             Yaquina Bay             L- Larvae             on a peduncle (Hart and McHugh 1944). Embryonic
              Alsea River                                   development is indirect and external. Eggs hatch in 19
            Siuslaw River V                                days at water temperatures of 8.5-11.50C, and 30-40
            Umpqua River O    0                            days at temperatures of 4.4-7.2�C (Garrison and Miller
               Coos Bay V                                  1980).
             Rogue River i
            Klamath River     C O0                         Ace and Size of Larvae: Larvae are 4-7 mm at hatching.
            Humboldt Bay  4                                 Postlarvae length is unknown, but probably about 35
               Eel River                                   mm (Barraclough 1964). Transformation to juvenile
             Tomales Bay                                    stage probably occurs at 30-35 mm  in length
       Cent. San Fran. Bay'        * Indudes Central San
                                     Francism. Suisun.     (Barraclough 1964).
       South San Fran. Bay           and San Pabo bays.
           Elkhorn Slough                                  Juvenile Size Ranae: Juveniles range from 30-140 mm
              Morro Bay                                    in length.
         Santa Monica Bay
           San Pedro Bay
             SanmPedro Bay                                  Aae and Size of Adults: Spawning usually occurs at 3
                    Alaheito By                            years of age. Spawning adult lengths range from 14.0-
             Anahewm Bay                                    20.0 cm, averaging 17.0 cm (Smith and Saalfeld 1955).
             Newpon Bay
             Mission Bay
             Mission Bay                                   The eulachon can live to 5 years.
           San Diego Bay
           Tijuana Estuary                                       F     eeding
                       A S J L E                           Trophic Mode: Larvae, juveniles, and adults are
                                                            planktivorous.

    Miarations and Movements: Larvae are apparently    Food Items: Larvae and postlarvae eat phytoplankton,
    swept quickly out to sea, spending little time in rivers or    copepod eggs, copepods, mysids, ostracods, barnacle
    estuaries. Adults migrate to spawning grounds from    larvae, cladocerans, worm larvae, and larvae of their
    December to April, but usually peak in February and    own species (Hart 1973). Juveniles and adultsconsume
    March.  Spawning grounds range from just above    primarily euphausiids,copepods, and otherplanktonic
    estuaries to many miles above, but no extensive    crustacea.  Adults do not usually feed during their
    migrations exist. Ocean movements are unknown, but    spawning migration.
    they are found in the echo scattering layers (Barraclough
    1964).                                                  Biological Interactions
                                                            Predation: Many predatory species follow and feed on
    Reproduction                                            eulachon during its spawning migration.  The spiny
    Mode: The eulachon is gonochoristic and iteroparous,    dogfish shark (Squalusacanthias), sturgeon (Acipenser
    however, most die after spawning. It is oviparous; eggs    spp.), Pacific halibut (Hippoglossus stenolepis), gadids,
    are fertilized externally.                              porpoise, finback whale (Balaenopteraphysalus), killer
                                                            whale (Orcinus orca), sea lions, seals, and gulls follow
    Matina/Soawnina: Spawning usually occurs inthe lower    eulachon runs (Hart 1973).  Harbor seals (Phoca
    reaches of rivers ortributaries. Eulachon mass spawn    vitulina) feed intensively on eulachon in the Columbia

                                                        179

    Eulachon continued
    River(Jeffries 1984), and salmon (C7corhynchusspp.)    Moyle, P. B. 1976. Inland fishes of California. Univ.
    and other fishes eat them at sea (Hart 1973).          Calif. Press, Berkeley, CA, 405 p.

    Factors Influencina PoDulations: Temperature changes    Odemar, M. W. 1964. Southern range extension of the
    (Blahm and McConnell 1971) and industrial pollution    eulachon, Thaleichthyspacificus. Calif. Fish. Game,
    (Smith and Saalfeld 1955) can have lethal and sublethal    50(4) :305-307.
    effects. Complete failure (i.e., disappearance) of the
    Cowlitz River run (a Columbia River tributary) from    Parente, W. D., and G. R. Snyder. 1970. A pictorial
    1949-1952 may have been due to industrial pollution.    record of the hatching and early development of the
    River flows can also alter migration patterns.  The    eulachon (Thaleichthys pacificus).  Northw. Sci.
    drought year of 1977 caused eulachon to bypass the    44(1):50-57.
    Cowlitz River and spawn in other rivers (J. Galbreath,
    Oregon Department of Fish and Wildlife, Clackamas,    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    OR, pers. comm.).                                       E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
                                                           of common and scientific names of fishes from the
    References                                              United States and Canada. Am. Fish. Soc. Spec. Publ.
                                                           No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Barraclough, W. E. 1964. Contribution to the marine
    life history of the eulachon, Thaleichthys pacificus. J.   Scott, W. B. and E. J. Crossman. 1973. Freshwater
    Fish. Res. Board Can. 21(5):1333-1337.                 fishes of Canada. Fish. Res. Board Can., Bull. No. 184,
                                                           966 p.
    Blahm, T. H., and R. J. McConnell. 1971. Mortality of
    adult eulachon (Thaleichthys pacificus) subjected to   Smith, W. E. and R. W. Saalfeld. 1955. Studies on
    sudden increases in water temperature. Northw. Sci.    Columbia River smelt,  Thaleichthys pacificus
    45(3):178-182.                                          (Richardson).  Wash. Dept. Fish., Fish. Res. Pap.
                                                           1 (3):3-26.
    Bohn, B. R., and D. Mclsaac. 1986. Columbia River
    fish runs and fisheries 1960-1985. Oreg. Dept. Fish    Snyder, G. R. 1969. Thermal pollution of Columbia
    Wildl. and Wash. Dept. Fish., Clackamas, OR, 77 p.    River might threaten smelt. Comm. Fish. Rev. 899:58-
                                                           64.
    Garrison, K. J., and B. S. Miller. 1980. Review of the
    early life historyof Puget Sound fishes. Fish. Res. Inst.,
    Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).

    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    Board Can., Bull. No. 180, 740 p.

    Hart, J. L., and J. L. McHugh. 1944. The smelts
    (Osmeridae) of British Columbia. Fish. Res. Board
    Can., Bull. No. 64, 27 p.

    Jeffries, S. 1984. Marine mammals of the Columbia
    Riverestuary. Col. Riv. Est. Data Dev. Prog., CREST,
    Astoria, OR, 62 p. plus appendices.

    Mclsaac, D., and B. Bohn. 1988. Columbia River fish
    runs and Fisheries, 1960-1987. Wash. Dept. Fish.,
    and Oreg. Dept. Fish Wildl., Olympia, WA, 83 p.

    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    Nelson. 1990. Distribution and abundance of fishes
    and invertebrates in west coast estuaries, Volume I:
    data summaries. ELMR Rep. No. 4. Strategic
    Assessment Branch, NOS/NOAA, Rockville, MD,
    240 p.

                                                       180

    181

       Microgadus proximus
       Adult

              5cm

    Common Name: Pacific tomcod                            However, it has not been collected in the Bering Sea
    Scientific Name: Microgadus proximus                   recently (Matarese et al. 1981).
    Other Common Names: California tomcod, tomcod,
    piciata (Gates and Frey 1974)                          Within Studv Area: The Pacific tomcod occurs in all
    Classification (Robins et al. 1980)                    estuariesfrom Elkhorn Slough, California, north through
    Phylum: Chordata                                       Puget Sound (Table 1) (Ganssle 1966, Aplin 1967,
    Class: Osteichthyes                                    Beardsley and Bond 1970, Bane and Bane 1971, Miller
    Order: Gadiformes                                      and Borton 1980, Wang 1986).
    Family: Gadidae
                                                           Life Mode
    Value                                                  Eggs have not been found, but are probably demersal
    Commercial: The Pacific tomcod is not a targeted    and adhesive (Walters 1984, Dunn and Matarese
    commercial fish, although some fishermen catch them    1987).  Larvae and small juveniles (<50 mm) are
    for personal use (Hart 1973).                          pelagic, while juveniles and adults are demersal
                                                           (Richardson and Pearcy 1977, Matarese et al. 1981,
    Recreational: Although not often targeted, this species    Walters 1984).
    is esteemed as a food fish by some anglers and should
    receive more fishing pressure (Roedel 1953, Beardsley    Habitat
    and Bond 1970).                                        Iype: Eggs apparently are released in marine (euhaline)
                                                           water. Larvae and small juveniles are found in nearshore
    Indicator of Environmental Stress: This is a useful    marine waters (Matarese et al. 1981) and estuaries
    indicator species because it is a demersal fish often    (Blackburn 1973, Misitano 1977). Adults and juveniles
    found in estuarine and marine areas containing    are common in polyhaline to euhaline waters (National
    contaminants.  Lesions appear more frequently in   Marine Fisheries Service 1981, Bottom et al. 1984,
    populations near pollution sources (Malins et al. 1980).    Emmett et al. 1987) and occur primarily in depths <92
                                                           m (Hart 1973).
    Ecological: The Pacific tomcod is an important prey for
    harbor seals (Phoca vitulina) (Beach et al. 1981) and    Substrate: Juveniles and adults are found primarily
    probably other marine mammals (Simenstad et al.   oversoftbottomsof mud, silt, and fine sand (Washington
    1979). It is an important predator of shrimp (Crangon    1977, Emmett et al. 1987).
    spp.) (Armstrong et al. 1981, Bottom et al. 1984).
                                                           Phvsical/Chemical Characteristics: The Pacific tomcod
    Range                                                  is primarily a marine species that utilizes estuaries.
    Overall: The Pacific tomcod's overall range is from    Specific salinity and temperature tolerances for each
    central California (Isaacson 1965) north to the Gulf of   life stage are not available.
    Alaska, Unalaska Island, and Bering Sea (Hart 1973).

                                                       182

                                                                                      Pacific tomcod continued

                                                          Matina/SDawnina: The Pacific tomcod apparently has
     Table 1. Relative abundance of Pacific tomcod        an extended spawning period (Dunn and Matarese
              in 32 U.S. Pacific coast estuaries.         1987). Spawning occurs in marine (euhaline) coastal
                           Life Stage                     waters (Waldron 1972, Pearcy and Myers 1974,
              Estuary    A S J L E                        Misitano 1977) from JanuarytoJune off San Francisco
               Puget Sound  3     (0    Relative abundance    Bay, California (Wang 1986), winterto spring off Oregon
                Hood Canal O O C O O        Highly abundan:   (Richardson and Pearcy 1977, Matarese et al. 1981),
                 Skagit Bay  i O C    0 0    �   Abundant  and FebruarytoMayin PortTownsend Bay, Washington
               Grays Harbor 0    * 0    0   Common        (Walters 1984).
                Willapa Bay O    * 0        Rare
              ColumbiaRiver     *      Blank Not present   Fecu ndit: Fecundity is estimated to be 1,200 eggs per
               Nehalem Bay O    O O                       female (Bane and Bane 1971).
              Tillamook Bay 0    I 0   Life stage:
                Netarls Bay   O O        A-Adults         Growth and Development
                                        S - Spawning adults
                Siletz River O    O    J - Juveniles      Eaa Size and Embryonic Development: Mature, non-
               Yaquina Bay O    O v    L-Larvae          fertilized eggs are spherical and 0.96 mm in diameter
                AlseaRiver    0        E - Eggs           (Walters 1984). Embryonic development is indirect
               Siuslaw River0  C_   _                     and external (Matarese et al. 1981, Walters 1984). No
              Umpqua River C    0 C                       information exists for length of embryogenesis.
                 CoosBay i    t
                Rogue River0    0                         Aae and Size of Larvae: Larvae range from 2.7-26.3
              Klarnath River  i                           mm in length. The yolk-sac is absorbed by 3.0 mm
              Humboldt Bay 0                              (Matarese et al. 1981).
                  Eel River  '
               TomalesBay O    O                          Juvenile Size Ranae: Juveniles are 26.3 mmtoprobably
          Cent. San Fran. Bay'    * Includes Central San
                                       Franysc * o. Suisun  200.0 mm in total length (TL) (Matarese et al. 1981,
          SouthSan Fran. Bay    i      and San Pablo bays.  National Marine Fisheries Service 1981).
              Elkhorn Slough  4
                 Morro Bay                                Aae and Size of Adults: Size and age of adults have not
            Santa Monica Bay
              Sana MPedronaBay                            been studied, but maturity is probably reached in 2
               SAlamitos Bay                              years and >200 mm TL (National Marine Fisheries
                                                          Service 1981). Adults can reach lengths of 310 mm TL
               Newport Say                                (Bane and Bane 1971).
                Mission Bay
              San Diego Bay                               Food and Feeding
             TijuanaEstuary                               Trophic Mode: Larvae are planktonic carnivores.
                         A S J L E                        Juveniles and adults are epibenthic, planktonic, and
                                                          benthic carnivores (depending on fish size and food
                                                          availability).
    Miarations and Movements: This species' movements
    are not well-studied.  Large, older fish move out of   Food Items: Larvae eat calanoid and harpacticoid
    estuaries in the early winter and return in the early   copepods, mysids, and juvenile crangonid shrimp
    spring (National Marine Fisheries Service 1981, Walters    (Walters 1984). Juveniles consume crangonid shrimp,
    1984). This is probably not an active migration, but   crab megalops, fish larvae, polychaetes, isopods,
    movement related to prey availability, spawning    gammaridamphipods, andcalanoidcopepods. Adults
    behavior, and temperature preferences. Larvae can    eat fish [e.g., northern anchovy (Engraulis mordax)l,
    be abundant in some bays (Walters 1984), but most    gammarid amphipods, crangonid shrimp, crab
    appear in nearshore waters along the open coast    megalops, polychaetes, mysids, andotherinvertebrates
    (Matarese et al. 1981). Juveniles appear to move to   (Bane and Bane 1971, Armstrong et al. 1981, Bottom
    shallow nearshore waters and estuarine areas after   et al. 1984).
    their pelagic phase.
                                                          Biological Interactions
    Reproduction                                          Predation: Larvae are probably consumed by many
    Mode: The Pacific tomcod is gonochoristic, oviparous,   fishes. Juveniles and adults are eaten by white sturgeon
    and iteroparous; eggs are fertilized externally.       (Acipenser transmontanus) (Robert Emmett, pers.

                                                       183

    Pacific tomcod continued
    observation) and other large fishes and marine    Emmett, R. L., T. C. Coley, G. T. McCabe, Jr., and R.
    mammals (Simenstad et al. 1979).                       J. McConnell.  1987.  Demersal fishes and benthic
                                                           invertebrates at four interim dredge disposal sites off
    Factors Influencinao PoDulations: Successful recruitment    the Oregon coast. Proc. Rep., Northwest Alaska Fish.
    of larvae and early juvenile stages is probably related    Cent., Nat. Mar. Fish. Serv., Coastal Zone Est. Stud.
    to predation and adequate prey availability. The Pacific   Div., NOAA, 2725 Montlake Blvd. E., Seattle, WA, 69
    tomcod appears to be a fast-growing, early-maturing    p. plus appendices.
    fish that has a high natural mortality rate (Walters
    1984).                                                  Ganssle, D. 1966. Fishes and decapods of San Pablo
                                                           and Suisun Bays. In D. W. Kelley (compiler), Ecological
    References                                              studies of the Sacramento-San Joaquin estuary. Calif.
                                                           Fish Game., Fish Bull. 133:64-94.
    Aplin, J. A. 1967. Biological survey of San Francisco
    Bay, 1963-1966. Calif. Dept. Fish Game, Mar. Res.    Gates, D. E., and H. W. Frey.  1974.  Designated
    Oper., MRO Ref. 67-4, 131 p.                            common names of certain marine organisms of
                                                           California. Calif. Fish Game, Fish. Bull. 161:55-90.
    Armstrong, D. A., B. G. Stevens, and J. C. Hoeman.
    1981. Distribution and abundance of Dungeness crab    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    and Crangonshrimp and dredging-related mortality of    Board Can., Bull. No. 180, 740 p.
    invertebrates and fish in Grays Harbor, Washington.
    Final Rep. to U.S. Army Corps Eng., Seattle, WA,    Isaacson, P.A. 1965. Southern range extension of the
    Contract No. DACW67-80-C-0086, School Fish., Univ.   tomcod, Microgadus proximus.  Calif. Fish. Game
    Wash., Seattle, WA, 349 p.                              51:58.

    Bane, G. W., and A. W. Bane. 1971. Bay fishes of   Malins, D. C., B. B. McCain, D. W. Brown, A. K. Sparks,
    northern California. Mariscos Publ., Hampton Bays,    and H. O. Hodgins. 1980. Chemical contaminants and
    NY, 143 p.                                              biological abnormalities in central and southern Puget
                                                           Sound. Tech. Memo. OMPA-2, Northwest Alaska
    Beach, R. J., A. C. Geiger, S. J. Jeffries, and S. D.    Fish. Cent., Nat. Mar. Fish. Serv., NOAA, 2725Montlake
    Treacy. 1981. Marine mammal-fishery interactions on    Blvd. E. Seattle, WA, 295 p.
    the Columbia River and adjacent waters, 1981. NWAFC
    Proc. Rep. 82-04, Northwest Alaska Fish. Cent., Nat.    Matarese, A. C., S. L. Richardson, and J. R. Dunn.
    Mar. Fish. Serv., NOAA, 2725 Montlake Blvd. E. Seattle,    1981.  Larval development of Pacific tomcod,
    WA, 186 p.                                              Microgadus proximus, in the northeast Pacific Ocean
                                                           with comparative notes on larvae of walleye pollack,
    Beardsley, A.J., and C. E. Bond. 1970. Field guideto    Theragra chalcogramma, and Pacific cod, Gadus
    common marine and bay fishes of Oregon. Agr. Exp.    macrocephalus (Gadidae). Fish. Bull., U.S. 78(4):923-
    Sta. Bull No. 607, Oregon State Univ., Corvallis, OR,    940.
    27 p.
                                                           Miller, B. S., and S. F. Borton. 1980. Geographical
    Blackburn, J. E. 1973. Pelagic eggs and larval fish of   distribution of Puget Sound fishes: maps and data
    Skagit Bay. In Q. J. Stober and E. O. Salo (editors),    source sheets. 3 vol. Fish. Res. Inst., Coll. Fish., Univ.
    Ecological studies ofthe proposed Kiket Island nuclear    Wash., Seattle, WA, various pagination.
    power site, p. 71-118, Fish. Res. Inst. Coll. Fish., Univ.
    Wash., Seattle, WA (FRI-UN-7304).                       Misitano, D. A.  1977.  Species composition and
                                                           relative abundance of larval and post-larval fishes in
    Bottom, D. L., K. K. Jones, and M. J. Herring. 1984.    the Columbia River estuary. Fish. Bull., U.S. 75:218-
    Fishes of the Columbia River estuary. Col. Riv. Est.   222.
    Data Dev. Prog., CREST, Astoria, OR, 113 p. plus
    appendices.                                             National Marine Fisheries Service. 1981. Columbia
                                                           River estuary data development program report:
    Dunn, J. R., and A. C. Matarese. 1987. A review of the    salmonid and non-salmonid fish 1981. Unpubl. Rep.,
    early life history of northeast Pacific gadoid fishes.   Pt. Adams Biol. Field Sta., Northwest and Alaska Fish
    Fish. Res. 5:163-184.                                   Cent., P.O Box 155, Hammond, OR, various pagination.

                                                        184

                                                                                      Pacific tomcod continued
    Pearcy, W. G., and S. S. Myers. 1974. Larval fish of
    Yaquina Bay, Oregon: A nursery ground for marine
    fishes? Fish. Bull., U.S. 72:201-213.

    Richardson, S. L., and W. G. Pearcy. 1977. Coastal
    and oceanic fish larvae in an area of upwelling off
    Yaquina Bay, Oregon. Fish. Bull., U.S. 75(1 ):125-146.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Roedel, P. M. 1953. Common ocean fishes of the
    California coast. Calif. Fish Game, Fish Bull. 91:1-184.

    Simenstad, C. A., B. S. Miller, C. F. Nyblade, D.
    Thornburgh, and L. J. Bledsoe. 1979. Food web
    relationships of northern Puget Sound and the Strait of
    Juan de Fuca: a synthesis of the available knowledge.
    U.S. Interagency (NOAA/EPA) Energy/Environ. Res.
    Dev. Prog. Rep., EPS-600/7-79-259, Washington, D.C.,
    335 p.

    Waldron, K. D. 1972. Fish larvae collected from the
    northeastern Pacific Ocean and Puget Sound during
    April and May 1967. Tech. Rep. NM FS SSR F-663, 16
    p. Northwest Alaska Fish. Cent., Nat. Mar. Fish. Serv.,
    NOAA, 2725 Montlake Blvd. E., Seattle, WA, 16p.

    Walters, G. E. 1984. Ecological aspects of larval and
    juvenile Pacific cod (Gadus macrocephalus), walleye
    pollock ( Theragra chalcogramma), and Pacific tomcod
    (Microgadusproximus) in Port Townsend, Washington.
    M.S. Thesis, Univ. Wash., Seattle, WA, 129 p.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: a
    guide to the early life histories. Tech. Rep. No. 9.
    Interagency ecological study program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    and U. S. Fish Wildl. Serv., various pagination.

    Washington, P. M. 1977. Recreationally important
    marine fishes of Puget Sound, Washington. Proc.
    Rep., Northwest Alaska Fish. Cent., Nat. Mar. Fish.
    Serv., NOAA, 2725 Montlake Blvd. E. Seattle, WA,
    122 p.

                                                       185

       Atherinops affinis
        Adult

               5cm

    Common Name: topsmelt                                  from San Francisco Bay (and surrounding areas) to
    Scientific Name: Atherinops affinis                     Monterey, California, 3) A. affinis littoralis ranges from
    Other Common Names: bay smelt, rainbow smelt,    Monterey down to San Diego Bay, California, 4) A.
    panzarotto, little smelt, least smelt, silverside, capron,    affinis cedroscensis is called the kelp topsmelt, and 5)
    jack pescadillo (Walford 1931, Gates and Frey 1974)    A. affinis insularium is the "island topsmelt", being
    Classification (Robins et al. 1980)                     found around the Santa Barbara Islands, California,
    Phylum: Chordata                                        (Schultz 1933, Federet al. 1974). When not in estuaries,
    Class: Osteichthyes                                     it appears to stay in shallow water along the shore line
    Order: Atheriniformes                                   (Hubbs 1918).
    Family: Atherinidae
                                                            Range
    Value                                                   Overall:The topsmelt is found from the Gulf of California
    Commercial: Although the topsmelt is an excellent   to Vancouver Island, British Columbia, Canada (one
    food fish (Bane and Bane 1971), there is a very limited   record) (Miller and Lea 1972, Hart 1973, Eschmeyer et
    commercialcatch. The topsmelt represents only about   al. 1983). However, it is not usually found north of
    15-25% of the California "smelt" catch (Bane and Bane    Tillamook Bay, Oregon.
    1971). It is usually taken in association with jacksmelt
    (Atherinopsis californiensis) (Frey 1971).              Within Studv Area: This species is found in most
                                                            estuaries of the study area south of Tillamook Bay,
    Recreational: It is taken by recreational anglers year    Oregon (Table 1) (Schultz 1933, Myers 1980).
    round and is one of the most commonly caught fishes
    from piers in California. Since this species is abundant    Life Mode
    and can be easily captured by light tackle, it is an    Eggs are benthic, larvae are planktonic, and juveniles
    important recreational fish for children (Frey 1971).   andadultsareschoolingpelagicfish. However, juvenile
                                                            and adults will apparently move into shallow waters
    Indicator of Environmental Stress: The topsmelt can    and feed on the bottom.
    withstand extreme salinities (80%o) (Carpelan 1955),
    and is an excellent bioassay organism (Reish and    Habitat
    Lemay 1988).                                            Type: Eggs are benthic and found in estuaries, bays,
                                                            and lagoons. Larvae are also found in embayments.
    Ecological: This species is one of the most abundant    Larvae are planktonic but school near the surface in
    pelagic fishes in many Pacific coast estuaries (Allen    shallow and open water (Wang 1986). Juveniles and
    and Horn 1975, Horn 1980, Allen 1982, Horn and Allen    adults are pelagic but are found over a wide range of
    1985). Five subspecies are presently recognized: 1) A.    habitats depending on time of year (Feder et al. 1974).
    affinisoregonia is a northern varietyfoundfrom Oregon    The topsmelt is primarily a marine fish that prefers
    to Humboldt Bay, California, 2) A. affinis affinis occurs    estuaries, bays, sloughs, and lagoons (Moyle 1976).

                                                        186

                                                                                          Topsmelt continued
                                                          at salinities of 30%o (Middaugh and Shenker 1988).
      Table 1. Relative abundance of topsmelt             The topsmelt is often found in waters of high turbidity.
               in 32 U.S. Pacific coast estuaries.         Maximum temperature for proper egg development is
                        Life Stage                        between 27.0-28.50C, and the minimum temperature
            Estuary    A S J L E                          foreggdevelopmentappearstobenear12.8�C(Hubbs
             PugetSound             Relativeabundance:     1965).  Juvenile and adult topsmelt appear to be
             Hood Canal              6   Highly abundant  eurythermal (Carpelan 1955), but temperatures of 26-
              Skagit Bay             @3  Abundant         27�C appear to cause stress (Ehrlich et al. 1979). The
            Grays Harbor '4           O   Common         upper and lower lethal temperatures for juvenile fish
             Willapa  Bay V     il  Rare                  were found to be 31.7�C and 10.40C, respectively
           Columbia River           Blank Not present     (Doudoroff 1945).
            Nehalem Bay i    i
            TillamookBay 0    O      Ufe stage:           Miarations and Movements: Larvae appear to stay
             NetartsBay Q   �  A   dS- snAingadults       near the surface in slow-moving waters.  Although
              Siletz River           J - Juveniles        some adults and juveniles will stay in the open waters
             YaquinaSay  0 a  013   L -LEaarsve           of some estuaries and bays year-round, most move to
              AlseaRiver O O O O O                         neritic areas and coastal kelp beds during fall and
            SiuslawRiver  3 @ (3 0 O                      winter (Wang 1986).  During spring, they are often
            Umpqua River O    O                           found nearthe entrance of bays (Schultz 1933). Adults
               Coos Bay � � � O �                         move into shallow water sloughs and mud flats in late
             Rogue River                                  spring and summerto spawn, and follow the salt wedge
            Klamath River                                 to upperestuarine areas during summer and fall (Wang
            Humboldt Bay O 0 0 0 0                        1986).
               Eel River  0  0 0 0
             TomalesBay @ @3� 6 @3 @
                        Toales Baya   * 13iiReproduction
       Cent. San Fran. Bay' O    O O   I ncludes Central San
                                     Frandsco. Suisun,.   Mode: The topsmelt is gonochoristic, iteroparous, and
       South San Fran. Bay  3      3      ad San Pabo bays.  oviparous; eggs are fertilized externally.
           Elkhorn Slough    *  3*
              MorroBay  666@3
              SantaMonicarro Bay                          Matina/SDawnina: Spawning occurs in Newport Bay,
         Santa Monica Bay   C0 0  )0
           San Pedro Bay    C )   0                       California, as early as February but most occurs during
            SanamitosBay      0  3 0 0                    May and June (Fronk 1969). Spawning occurs from
            Alnaheimm ay ISO  13 * 6> zApril to October in San Francisco Bay, with peaks in
             Newport Bay      0  � 6 0                     May and June (Wang 1986). Spawning takes place at
             Mission Bay a1  * 0 3                        temperatures of 10-25�C and in shallow water habitats
           San Diego Bay  3   *3           a 0that have appropriate submerged aquatic vegetation
           Tijuana Estuary @gle   O          a            (Schultz 1933). Most spawning may occur at night
                       Tijuan A S J L E                   (Fronk 1969).  The topsmelt appears to spawn in
                                                          batches, laying eggs morethan once during a spawning
                                                          season (Fronk 1969, Wang 1986).
    Substrate: Eggs are laid primarily on eelgrass (Zostera
    spp.) and adhere to macroalgae on tidal flats (Schultz   Fecundity: Fecundities range from 200 eggs per female
    1933). Larvae are often found over soft, unconsolidated   (of length 110-120 mm) to about 1,000 eggs per female
    sediments andothersubstrates (Wang 1986). Juveniles   (of length 160 mm and over) (Fronk 1969).
    and adults occur along sandy beaches, in kelp beds,
    over rocky reefs, and around piers (Feder et al. 1974).    Growth and Development
                                                          Eaa Size and Embrvonic Develooment: Eggs are
    Phvsical/Chemical Characteristics: The topsmelt is a    spherical and 1.5-1.7 mm in diameter (Wang 1986).
    euryhaline species (Fronk 1969).  Eggs develop    Eggs have athickchorionbearing 2-8filaments attached
    successfully in salinities up to 72/%o (Carpelan 1955).   in a random pattern. These filaments cause the eggs
    Smaller fish may tolerate high salinities better than    tobecomeentangledwitheelgrassandothervegetation
    larger fish (Carpelan 1955). Young-of-the-year are    (Wang 1986). Embryonic development is indirect and
    common in mesohaline and oligohaline areas of   external. Hatching time varies from 35 days at 13�C to
    southern San Francisco Bay (Wang 1986).  While    <9 days at 27�C (Hubbs 1965).
    juveniles can tolerate salinities ranging from 2-80%0,
    growth is reduced at higher salinities (Middaugh and    Aae and Size of Larvae: Larvae are 4.3-4.9 mm long
    Shenker 1988). Optimum survival and growth occurs    (total length) at hatching and about 0.0011 g (wet

                                                      187

    Topsmeltcontinued
    weight). They are also reported to be 5.1-5.4 mm long    References
    (standard length) at hatching (Middaugh et al. 1990).
    They are 9.5-10.0 mm  long when the yolk-sac is   Allen, L. G. 1982. Seasonalabundance, composition,
    absorbed.  Juvenile characteristics are formed at    and productivity of the littoral fish assemblage in upper
    approximately 18.5 mm (Wang 1986).                      Newport Bay, California. Fish. Bull., U.S. 80(4):769-
                                                           790.
    Juvenile Size Ranae: Juveniles are approximately
    18.5-120.0 mm long (Schultz 1933, Fronk 1969).         Allen, L. G., and M. H. Horn.  1975.  Abundance,
                                                           diversity and seasonality of fishes in Colorado Lagoon,
    Aae and Size of Adults: Northern varieties grow larger   Alamitos Bay, California. Est. Coast. Mar. Sci. 3:371 -
    than southern subspecies (Schultz 1933). Maturity is   380.
    reached in two years at about 120 mm in length by A.
    affinislittoralis (Schultz 1933, Fronk 1969). In Oregon,    Bane, G. W., and A. W. Bane. 1971. Bay fishes of
    only 5% mature in their second year; most mature in   northern California with emphasis on the Bodega
    their third year when >200 mm long (Schultz 1933).    Tomales Bay area. Mariscos Publ., Hampton Bays,
    This species can live up to 8 years and reach lengths    NY, 143 p.
    up to 37 cm (Schultz 1933, Eschmeyer et al. 1983).
                                                           California Department of Fish and Game. 1987. Delta
    Food and Feeding                                       outflow effects on the abundance and distribution of
    Trophic Mode: The topsmelt is omnivorous (Quast    San Francisco Bay fish and invertebrates, 1980-1985.
    1968, Horn andAllen 1985). Juveniles and adultsoften    Exhibit 60, entered by the Calif. Dept. Fish Game for
    feed near the water surface, but feed on the bottom    the State Water Resources Control Board 1987 Water
    when in shallow water (about 2 m or less). They feed    Quality/WaterRights Proceeding onthe San Francisco
    primarily during the day (Hobson et al. 1981).          Bay/Sacramento-San Joaquin Delta. Calif. Dept. Fish
                                                           Game, Stockton, CA, 345 p.
    Food Items: Estuary and bay inhabitants feed primarily
    on plant material, including Melosira moniliformis,    Carpelan, L. H. 1955. Tolerance ofthe San Francisco
    Entermorpha spp., andotheralgaeanddiatoms(Fronk    topsmelt, Atherinops affinis affinis, to conditions in
    1969, Moyle 1976, Ruagh 1976). They also consume    salt-producing ponds bordering San Francisco Bay.
    small crustacea (amphipods, copepods, insects, and    Calif. Fish Game 41(4):279-284.
    cumaceans) and some benthic invertebrates
    (polychaetes and gastropods) (Horn and Allen 1985).    Doudoroff, P. 1945. The resistanceand acclimatization
    Oceanicinhabitantsareprimarilyplanktoniccrustacean    of marine fishes to temperature changes.  II.
    carnivores.  Primary prey include gammarid and    Experiments with Fundulus and Atherinops. Biol. Bull.
    caprellid amphipods, mysids, ostracods, copepods,    88(2):197-206.
    and crustacean larvae (Quast 1968, Fronk 1969).
                                                           Ehrlich, K. F., J. M. Hood, G. Muszynski, and G. E.
    Biological Interactions                                 McGowen. 1979. Thermal behavioral responses of
    Predation: The topsmelt is an important prey for many    selected California littoral fishes.  Fish. Bull., U.S.
    piscivorous birds and fishes, including yellowtail (Seriola   76(4):837-849.
    lalandel) and other large fishes (Feder et al. 1974).
                                                           Eschmeyer, W. N., W. S. Herald, and H. Hammann.
    Factors Influencina Populations: Population abundance    1983. A field guide to Pacific coast fishes of North
    was significantlycorrelated totemperature and salinity   America. Houghton Mifflin Co., Boston, MA, 336 p.
    in Newport Bay, California (Allen 1982). No relation
    was found between abundance indices and river flow    Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    in San Francisco Bay (California Department of Fish    Observations of the fishes associated with kelp beds in
    and Game 1987). This species is commonly impinged    southern California. Calif. Fish Game, Fish Bull. 160,
    on power plant intake screens, but this may not be a    138 p.
    significant cause of mortality for the bay population
    (San Diego Gas and Electric 1980). Since this species    Frey, H. W. 1971. California's living marine resources
    uses shallow-water eelgrass areas for spawning, the    and their utilization.  Calif. Dept. Fish Game,
    removal or destruction of this habitat adversely affects    Sacramento, CA, 148 p.
    topsmelt abundance.

                                                       188

                                                                                           Topsmelt continued

    Fronk, R. H.  1969.  Biology of Atherinops affinis   Myers, K. W. W.   1980.  An investigation of the
    littoralis Hubbs in Newport Bay. M.S. Thesis, Univ.    utilization of four study areas in Yaquina Bay, Oregon,
    Calif., Irvine, CA, 106 p.                              by hatchery and wild juvenile salmonids. M.S. Thesis,
                                                           Oregon State Univ., Corvallis, OR, 234 p.
    Gates, D. E., and H. W. Frey. 1974. Designated
    common names of certain marine organisms of   Quast, J. C. 1968. Observations on the food of the
    California. Calif. Fish Game, Fish Bull. 161:55-90.     kelp-bed fishes. Calif. Fish Game, Fish Bull. 139:109-
                                                           142.
    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    Board Can., Bull. No. 180, 740 p.                       Reish, D. J., and J. A. Lemay. 1988. Bioassay manual
                                                           fordredged sediments. Research Rep., various pages.
    Hobson, E. W. N. McFarland, and J. R. Chess. 1981.   Available, U.S. Army Corps Eng., Los Angeles District,
    Crepuscular and nocturnal activities of Californian    Los Angeles, CA, (Contract Number DACW-09-83R-
    nearshore fishes, with consideration of their scotopic    005).
    visual pigments and the photic environment. Fish.
    Bull., U.S. 79(1):1-30.                                 Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
                                                           E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    Horn, M. H.  1980.  Diel and seasonal variation in   of common and scientific names of fishes from the
    abundance and diversity of shallow-water fish    United States and Canada. Am. Fish. Soc. Spec. Publ.
    populations in Morro Bay, California. Fish. Bull., U.S.    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    78(3):759-770.
                                                           Ruagh, A. A. 1976. Feeding habits of silversides
    Horn, M. H., and L. G. Allen. 1985. Fish community    (Family Atherinidae) in Elkhorn Slough, Monterey Bay,
    ecology in southern California bays and estuaries.    California. M.S. Thesis, Calif. State Univ., Fresno, CA,
    Chapter 8.  In A. Yanez-Arancibia (editor),  Fish    60 p.
    community ecology in estuaries and coastal lagoons:
    towards an ecosystem integration, p. 169-190 DR (R)    San Diego Gas and Electric. 1980. Silvergate power
    UN AM Press, Mexico.                                    plant cooling water intake system demonstration (in
                                                           accordance with section 316(b) Federal Water Pollution
    Hubbs, C. 1918. The fishes of the genus Atherinops,    Control Act Amendment of 1972). San Diego Gas and
    their variation, distribution, relationships, and history.   Electric, San Diego, CA, various pagination.
    Bull. Amer. Mus. Nat. Hist. 38(13):409-440.
                                                           Schultz, L. P. 1933. The age and growth of Atherinops
    Hubbs, C. 1965. Developmentaltemperaturetolerance    affinis oregonia  Jordan and Snyder and other
    and rates of four southern California fishes, Fundulus    subspecies of baysmelt along the Pacific coast of the
    parvipinnis, Atherinops affinis, Leuresthes tenuis, and    United States. Wash. State Univ. Publ. Biol. 2(3):45-
    Hypsoblennius sp. Calif. Fish Game 51(2):113-122.    102.

    Middaugh, D. P., M. J. Hemmer, J. M. Shenker, and T.   Walford, L. A. 1931. Handbook of common commercial
    Takita.   1990.   Laboratory culture of jacksmelt,    and game fishes of California. Calif. Fish Game, Fish
    Atherinopsis californiensis, and topsmelt, Atherinops    Bull. 28, 181 p.
    affinis(Pisces:Atherinidae), with a description of larvae.
    Calif. Fish Game 76(1):4-43.                            Wang, J. C. S. 1986. Fishes of the Sacramento-San
                                                           Joaquin estuary and adjacent waters, California: A
    Middaugh, D. P., and J. M. Shenker. 1988. Salinity   guide to the early life histories.  Tech. Rep. No. 9.
    toleranceof youngtopsmelt, Atherinopsaffinis, cultured    Interagency ecological study program for the
    in the laboratory. Calif. Fish Game 74(4):232-235.      Sacramento-San Joaquin estuary. Calif. Dept. Water
                                                            Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    Miller, D. J., and R. N. Lea. 1972. Guide to the coastal    U.S. Fish Wildl. Serv., various pagination.
    marine fishes of California. Calif. Fish Game, Fish Bull.
    157, 235 p.

    Moyle, P. B. 1976. Inland fishes of California. Univ.
    Calif. Press, Berkeley, CA, 405 p.

                                                        189

       Atherinopsis californiensis
       Adult

                    10cm

    Common Name: jacksmelt                                 caught within 5 km of shore (Ruagh 1976).
    Scientific Name: Atherinopsis californiensis
    Other Common Names: California smelt, silverside,   Range
    horse smelt, blue smelt, pescado del rey, peixe rey,   Overall: Overall range is from Santa Maria Bay, Baja
    pesce rey, jack smelt (Gates and Frey 1974)            California, to Yaquina Bay, Oregon (Miller and Lea
    Classification (Robins et al. 1980)                    1972, Eschmeyer et al. 1983).
    Phylum: Chordata
    Class: Osteichthyes                                    With in Studv Area: Thejacksmelt is commonly found in
    Order: Atheriniformes                                  most bays and estuaries that have appropriate habitat
    Family: Atherinidae                                    south of Coos Bay, Oregon (Table 1).

    Value                                                  Life Mode
    Commercial: In 1945, over 907 kg of jacksmelt were    Eggs are demersal and adhesive (Clark 1929). Larvae
    landed, primarily from Newport, Monterey, San    school and are pelagic (Wang 1986). Juveniles and
    Francisco, Tomales and Humboldt Bays, California   adults are surface-oriented pelagic schooling fishes
    (Frey 1971). Presently, it forms the largest portion of   (Allen and DeMartini 1983).
    the "smelt" captures in California, but is not considered
    an important commercial fish. It is primarily caught    Habitat
    incidentally during other fisheries.                   Iype: Eggs are usually found on vegetation in shallow-
                                                           water nearshore marine habitats as well as estuaries
    Recreational: The jacksmelt is commonly captured    and bays (Wang 1986).  Larvae are also found in
    from California piers (Frey 1971) and is easily caught    estuarine, bay, and kelp bed habitats and actively
    using light hook and line fishing gear (Frey 1971). In   school near the surface.  Juveniles and adults are
    California, there are no recreational catch limits   found in neritic, estuarine, and bay environments.
    (California Department of Fish and Game 1987a).        Juveniles and adults are most often found in murky
                                                           water from the surface down to 29 m, but tend to
    Indicator of Environmental Stress: No information is   concentrate between 1.5and 15m(Federetal. 1974).
    presently available. However, because the jacksmelt
    uses estuaries for spawning and rearing, degradation    Substrate: Eggs are laid on substrates/vegetation that
    of estuarine habitats can affectthis species'population.    allow them to become entangled (Zostera spp.,
                                                           Gracillaria spp., and hydroids, etc.) (Frey 1971, Wang
    EcolQgical: The jacksmelt is an important member of    1986). Larvae are found over a variety of substrates,
    the California nearshore coastal, bay, and estuary    but mostly sandy and muddy bottoms and in the kelp
    fauna (Clark 1929, Allen and DeMartini 1983, California   canopy (Frey 1971). Juveniles and adults prefer sandy
    Department of Fish and Game 1987b). It is often found    bottoms (Feder et al. 1974).
    schooling with topsmelt (Atherinops affinis) and usually

                                                       190

                                                                                           Jacksmelt continued
                                                           During summer, large schools of juveniles and some
      Table 1. Relative abundance of jacksmelt in          adults reside in bays and estuaries, moving out to
               32 U.S. Pacific coast estuaries.            coastal waters in the fall.

                        Life Stage                         Reproduction
           Estuary    A S J L E                           Mode:Thejacksmelt isgonochoristic, iteroparous, and
            Puget Sound  0         Relative abundance:     oviparous. It is a batch spawner and eggs are fertilized
             Hood Canal :                Highly abundant   externally (Clark 1929).
              Skagit Bay    :            : Abundant
            Grays Harbor             O   Common            Matina/SDawnino: Spawning may occur from October
             ]  Willapa Bay         Bnk Notpresent         to March with a peak during November-March (Clark
           Columbia River               Not preset         1929), and reportedlyyear-round in southern California
            Nehalem Bay                                    (Feder et al. 1974) In San Francisco Bay, spawning
            illarooktsBay           Life stage:            occurs from October to early August (Wang 1986).
                                    A - Adults
                                     -a S-Spawning adults  Spawning in San Pablo Bay is reported to occur from
             Siletz River           J -Juveniles           September to April (Ganssle 1966). In Tomales Bay,
             Yaquina Bay     4      L - Larvae
             Alsea River            E-Eggs                 spawning occurs from January to March (Banerjee
            Siuslaw River                                  1966). Spawning occurs over marine vegetation in
           Umpqua River                                   shallow coastal waters and in bays and estuaries
              Coos Bay O    O                             where appropriate substrate is available.
             Rogue River
            Klamath River   :                              Fecundity: Fecundity is not documented, but probably
            Humboldt Bay O O O O O                         over 2,000 eggs per female.
               Eel River
            Tomales Bay *� �   a a  Growth and Development
       Cent. San Fran. Bay � �  � � �  IncludesCentralSan  Eaa Size and Embryonic DeveloDment: Unfertilized
                                    Francisco, Suisun,
       South San Fran. Bay � � � � �  and San Pablo bays   eggs are spherical and 0.9-2.2 mm in diameter (Clark
           ElkhomrnSlough ( 1 0 CD i1                      1929); fertilized eggs are 1.9-2.5 mm  in diameter
              Morro Bay (1 Q  0  (                         (Wang 1986). Eggshaveathick, hard chorion that has
         SantaMonicaBay 0i IM  l 0 5                       15 or 16, 1-2 mm-long filaments attached.  These
           San Pedro Bay a   0  I O                        filaments entangle eggs on substrates to form large
            Alamitos Bay                                   egg masses (Wang 1986). Embryonic development is
            Anaheim Bay
                                                           indirect and external.  The yellowish-orange eggs
            Newport Bay -o    a                            hatch within seven days at 10-120C (Wang 1986).
             Mission Bay 5( ]    (  I
           San Diego Bay O C C) O O0
           Tijuana Estuary                                 Aae and Size of Larvae: After hatching, larvae remain
                      A S J L E                            on the bottom for a moment and then actively swim
                                                           near the surface (Wang 1986). Larvae live on their
                                                           yolk-sac until it is absorbed (about 48 hours after
    Phvsical/Chemical Characteristics: Temperature and    hatching) (Middaugh et al. 1990). The larval size range
    salinity tolerances of this species are not known.    is 7.5-8.6 mm long at hatching to about 25 mm long at
    However, the distribution of juvenile and adult jacksmelt   transformation to juvenile (Clark 1929, Wang 1986). At
    in San Francisco Bay shows they occur primarily in   8days, theyare10.5-11.7mmlong;at24daystheyare
    polyhalineandeuhalinewaters(Califomia Department    17.6-20.3 mm long (Middaugh et al. 1990).
    of Fish and Game 1987b). Eggs may hatch in salinities
    as low as 5?O/ (Wang 1986).  Optimum larval and    Juvenile Size Rance: Juvenile jacksmelt average 110
    juvenile survival and growth appears to be within    mm long at the end of their first year, and 180-190 mm
    salinities of 10 to 20%o, indicating larvae may prefer   at the end of two years (Clark 1929).
    mesohaline salinities (Middaugh and Shenker 1988,
    Middaughetal. 1990). The jacksmelt appearstoprefer    Aae and Size of Adults: Individuals that grow quickly
    turbid waters (Feder et al. 1974).                     (>200 mm  long) will mature in their second year.
                                                          However, all individuals mature by their third year
    Miarations and Movements: This species is seldom    (Clark 1929). The largest jacksmelt reported was 78
    found far from shore (Baxter 1960). Jacksmelt move    cm long, but the largest actually measured was 62 cm
    inshore and into bays and estuaries to spawn during    (Miller and Lea 1972). Maximum age may be 1 1 years
    late winter and early spring (Clark 1929, Wang 1986).    (Frey 1971).

                                                       191

    Jacksmeft continued
    Food and Feeding                                       Bane, G. W., and A. W. Bane. 1971. Bay fishes of
    TrophicMode:Thejacksmeltisomnivorous(Baneand    northern California with emphasis on the Bodega
    Bane 1971, Ruagh 1976).                                Tomales Bay area. Mariscos Publ., Hampton Bays,
                                                           NY, 143 p.
    Food Items: Primary preyforthis species include algae
    (Ulothrix spp., Melosira moniliformis, Enteromorpha    Banerjee, T. 1966. Survey of the fishes of Tomales
    spp., and otherfilamentous algae, and benthic diatoms),    Bay with notes on the life history of the white seaperch,
    crustaceans (mysids, copepods, decapod larvae), and    Phanerodon furcatusGirard. M.S. Thesis, Univ. Pacific,
    detritus (Bane and Bane 1971, Ruagh 1976).              Stockton, CA, 81 p.

    Biological Interactions                                Baxter, J. L. 1960. Inshore fishes of California. Calif.
    Predation: The jacksmelt is eaten byyellowtail (Seriola   Dept. Fish Game, Sacramento, CA, 80 p.
    lalandet), kelp bass (Paralabrax clathratus), sharks,
    and other piscivorous fishes (Baxter 1960, Feder et al.   California Department of Fish and Game. 1987a. 1987
    1974). It is probably also eaten by piscivorous birds   California sport fishing regulations. Calif. Dept. Fish
    [e.g.,brown pelican (Pelecanus occidentalis) and gu Ils]   Game, Sacramento, CA, 12 p.
    and marine mammals.
                                                           California Department of Fish and Game. 1987b.
    Factors Influencina PoDulations: Because this species    Delta outflow effects on the abundance and distribution
    utilizes embayments and estuaries for spawning, it is   of San Francisco Bay fish and invertebrates, 1980-
    highly susceptible to adverse effects from pollution and    1985. Exhibit 60, entered by the California Department
    habitat modification. Interestingly, jacksmelt are not   of Fish and Game for the State Water Resources
    commonly found in Anaheim Bay, Alamitos Bay, or   Control Board 1987 Water Quality/Water Rights
    Newport Bay, California (Klingbeil et al. 1974, Allen    Proceeding on the San Francisco Bay/Sacramento-
    and Horn 1975, Allen 1982), whereas topsmelt are    San Joaquin Delta. Calif. Dept. Fish Game, Stockton,
    abundant in these bays.  Apparently jacksmelt are    CA, 345 p.
    much more sensitive to salinity and temperature
    fluctuations than topsmelt. A parasitic nematode often    Clark, F. N. 1929. The life history of the California jack
    infests the flesh of jacksmelt, thus reducing its   smelt, Atherinopsis californiensis. Calif. Fish Game,
    commercial and recreational value (Frey 1971). The    Fish Bull. 16, 22 p.
    final host forthis parasite is perhaps sharks or pelicans
    (Frey 1971).  Freshwater inflow affects jacksmelt    Eschmeyer, W. N., W. S. Herald, and H. Hammann.
    distributions in San Francisco Bay; during years of low    1983. A field guide to Pacific coast fishes of North
    freshwater inflow, jacksmelt use San Pablo Bay and    America. Houghton Mifflin Co., Boston, MA, 336 p.
    Carquinez strait, but in high-flow years they are more
    abundant in South and Central San Francisco Bay    Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    (California Department of Fish and Game 1987b).         Observations on fishes associated with kelp beds in
                                                           southern California. Calif. Fish Game, Fish Bull. 160,
    References                                              144 p.

    Allen, L. G. 1982. Seasonal abundance, composition,    Frey, H. W. 1971. California's living marine resources
    and productivity ofthe littoral fish assemblage in upper    and their utilization.  Calif. Dept. Fish Game,
    Newport Bay, California. Fish. Bull., U.S. 80(4):769-    Sacramento, CA, 148 p.
    790.
                                                           Ganssle, D. 1966. Fishes and decapods of San Pablo
    Allen, L. G., and E. E. DeMartini. 1983. Temporal and    and Suisun Bays. In D. W. Kelley (compiler), Ecological
    spatial patterns of nearshore distribution and abundance    studies of the Sacramento-San Joaquin estuary. Calif.
    of the pelagic fishes off San Onofre-Oceanside,    Fish Game, Fish Bull. 133:64-94.
    California. Fish. Bull., U.S. 81(3):569-586.
                                                           Gates, D. E., and H. W. Frey. 1974. Designated
    Allen, L. G., and M. H. Horn.  1975.  Abundance,    common names of certain marine organisms of
    diversityandseasonalityof fishesin ColoradoLagoon,    California. Calif. Fish Game, Fish Bull. 161:55-90.
    Alamitos Bay, California. Est. Coast. Mar. Sci. 3:371-
    380.                                                    Klingbeil, R. A., R. D. Sandell, and A. W. Wells. 1974.
                                                           An annotated checklist of the elasmobranchs and
                                                           teleosts of Anaheim Bay. In E. D. Lane and C. W. Hill

                                                       192

                                                                                        Jacksmelt continued
    (editors), The marine resources of Anaheim Bay.
    Calif. Fish Game, Fish Bull. 165:79-90.

    Middaugh, D. P., M. J. Hemmer, J. M. Shenker, and T.
    Takita. 1990. Laboratory culture of jacksmelt,
    Atherinopsis californiensis, and topsmelt, Atherinops
    affinis(Pisces:Atherinidae), with a description of larvae.
    Calif. Fish Game 76(1):4-43.

    Middaugh, D. P., and J. M. Shenker. 1988. Salinity
    tolerance of young topsmelt, Atherinopsaffinis, cultured
    in the laboratory. Calif. Fish Game 74(4):232-235.

    Miller, D. J., and R. N. Lea. 1972. Guidetothecoastal
    marine fishes of California. Calif. Fish Game, Fish Bull.
    157, 235 p.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Ruagh, A. A. 1976. Feeding habits of silversides
    (Family Atherinidae) in Elkhorn Slough, Monterey Bay,
    California. M.S. Thesis, Calif. State Univ. Fresno, CA,
    60 p.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: A
    guide to the early life histories. Tech. Rep. No. 9.
    Interagency ecological study program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    U.S. Fish Wildl. Serv., various pagination.

                                                        193

       Gasterosteus aculeatus
       Adult

                    2cm

    Common Name: threespine stickleback                    Wootton 1976). Trophic phenotypes have also been
    Scientific Name: Gasterosteus aculeatus                identified (Lavin and McPhail 1986).
    Other Common Names: common stickleback, two-
    spined stickleback, stickleback, thornfish, thornback,    Range
    needle stickleback (Bigelow and Schroeder 1953,    Overall: Overalldistribution is amphiboreal (interrupted
    Okada 1955, Gates and Frey 1974)                       northern circumpolar range), found between lat. 35�N
    Classification (Robins et al. 1980)                    and 700N in Europe (Wootton 1976). In eastern North
    Phylum: Chordata                                       America it is found from Chesapeake Bay north to
    Class: Osteichthyes                                    Baffin Island, while in western North America it occurs
    Order: Gasterosteiformes                               from Baja California, Mexico, to St. Lawrence Island,
    Family: Gasterosteidae                                 Alaska(McPhail and Lindsey 1970, Scott and Crossman
                                                          1973, Wootton 1976, Wydoski and Whitney 1979). In
    Value                                                  the western North Pacific, it is found from the Bering
    Commercial: This species is not commercially    SeasouthtonorthemJapan(Andriyashev1954, Okada
    harvested.                                             1955).

    Recreational: The threespine stickleback is a good    Within Studv Area: The anadromous plated form
    aquarium fish and commonly used for studying fish   (trachurus) is found in all Pacific coast estuaries from
    behavior and physiology (Carlander 1969, Wootton    theSanLorenzoRiverin north MontereyBay, Califomia,
    1976).                                                 through Washington (Table 1) (Millerand Hubbs 1969,
                                                          Wootton 1976). The southern distribution of the
    Indicator of Environmental Stress: Because the    anadromous form appears to be limited by high
    threespine stickleback is easy to collect and hold in   temperatures (Bell 1976). The non-anadromous form
    laboratory conditions, it has often been used as an    has a wider distribution (Wooton 1976).
    experimental animal fortesting water pollution (Wootton
    1976). For example, heavy metals have been found to   Life Mode
    be highly toxic to this species (Wootton 1976).        Eggs are demersal and are laid by the female in a nest
                                                          built by a male. Larvae are free-swimming, but stay
    Ecological: The threespine stickleback is prey for many    with the nest, which is guarded by the male. Juveniles
    species of fishes and birds, and is an important resident    and adults are pelagic, but typically do not travel far
    of shallow-water estuarine habitats and lakes. It also   from shore. However, some have been captured far
    colonizes irrigation canals and reservoirs (Moyle 1976,    out at sea (Clemens and Wilby 1961, Wootton 1976).
    Simenstad 1983). Different morphological forms exist   Within the study area, at least two morphological
    (each having distinct habitats with little hybridization)   varieties occur. The trachurus form is anadromous,
    leading scientists to describe many subspecies (see    migrating from marine waters to brackish and fresh
    "Life Mode") (Hagen 1967, Miller and Hubbs 1969,    waters to breed. It possesses a complete set of lateral

                                                       194

                                                                                Threespine stickleback continued

                                                          Substrate: Although adults and juveniles are found
     Table 1. Relative abundance of threespine             over a variety of substrates, breeding male sticklebacks
              stickleback in 32 U.S. Pacific coast         normally attempt to build their nests over soft mud or
              estuaries.                                   sand bottoms that have vegetation nearby (Scott and
                        Life Stage                         Crossman 1973, Wootton 1976, Wydoski and Whitney
           Estuary    A S J L E                            1979)
            Puget Sound 0  ) :0   �0 0   Relative abundance:
             Hood Canal 0 C0 (0  0 0    *   Highlyabundant  Phvsical/Chemical Characteristics: The threespine
              Skagit Bay a*   a a  O    (1  Abundant        stickleback can tolerate minimum dissolved oxygen
            Grays Harbor 0 { � O (j    O   Common
             Grllaparbayor  13 3'   0 0 0   Rare           concentrations as low as 0.25-0.50 mg/I (Wootton
             ColumbiaRiver I     * � �   Blank Notpresent  1976). Maximumtemperature before mortality is 26�C
            Nehalem Bay CD   CDI  0  CD e::(Blahm and Snyder 1975). This species can withstand
            NehalemBay O O i) O O
           Tiilamook Bay C C O O OC) C   Life stage:       a wide range of salinities, but this depends on water
             NetairtsBay a a:o a     A -eAdults            temperature, degree of sexual maturity, and
                                     S-Spawning adults     morphological form (leiurus or trachurus) (Wootton
             SietaquRiver 0 0 0 00   J-Luvenies            1976). The migratory trachurus form loses its ability to
            Yaquina~aO O C) C)    O C] Larvae
             AlseaaRiver o  0     IN 0a   E-Eggs           tolerate fresh water during fall (Wootton 1976).
            Suslaw River 0 0  t 0 o                         Spawning occurs at temperatures of 15.8-18.5�C (Vrat
           UmpquaRiver ( 0 a 0 0                           1949)inveryshallowfreshtopolyhalinewaters (Morrow
              Coos Bay  0BaC                               1980, Wang 1986).
             Rogue River 00)  0  O 0
                                                          Miarations and Movements:Thefreshwaterformwinters
           HumboldtBay R a  a O O                          in deep water and moves to shallow water in spring
               EealRiver y    00a  0 0  (Morrow 1980). The anadromous form migrates into
            Tomales Bay O O 0 : 0 0                        shallow fresh and brackish waters of coastal estuaries
            T.Fmalesay 0C0n 0a00
       Cent. San Fran. Bay' *      0    Includes Central San  in the spring to spawn (Wydoski and Whitney 1979,
       SouthSanFran.Bay  S0(3   0 0   Francisco, Suisun.    Whoriskey and FitzGerald 1989). Surviving spawners
       South Son Fran. Bay    C O    0 CD  and San Pablo beys.
           Elkhom Slough O    O                             (massive post-spawning mortality can occur) and
              Morro Bay O    0                             juveniles move back to sea in the fall (Wang 1986).
        Santa Monica Bay                                   Anadromous juveniles may start moving to sea at
           San Pedro Bay                                    about 5 weeks of age (Bakker and Feuth-De Bruijn
            Alamitos Bay                                   1988). Sticklebacks have been found far out to sea, but
            Anaheim Bay                                     these individuals may be lost from the population
            Newport Bay                                    (Quinn and Light 1989); most sticklebacks stay close to
             Mission Bay                                    shore (Bigelow and Schroeder 1953, McPhail and
           San Diego Bay                                    Lindsey 1970).  Juveniles and non-breeding adults
          Tijuana Estuary                                  form loose schools, probably to aid in finding food and
                      A S J L E                            protection from predators. During the breeding season
                                                           in estuaries (spring and early summer), adults breed in
    bony plates, and is silver in color. The leiurus form    shallow water. After the breeding season, adults and
    spends its entire life in fresh water, has few lateral bony    juveniles move into deeper open waters.
    plates, and is olive-brown in color (Scott and Crossman
    1973, Moyle 1976, Garrison and Miller 1980).            Reproduction
                                                          Mode: The threespine stickleback is gonochoristic,
    Habitat                                                 polygamous, oviparous, and iteroparous; eggs are
    jype: All life stages are typically found associated with   fertilized externally (Vrat 1949).
    vegetation in shallow water bays, lakes, and slow-
    moving rivers. This species occurs primarily in low-    Matina/SDawnina: Spawning occurs from early spring
    lyingcoastalstreamsandlakes(Moyle1976). However,    (March) to fall (October), depending on location.
    threespine sticklebacks have been found up to 500    However, the anadromous form spawns primarily in
    miles out to sea (McPhail and Lindsey1970). Breeding    June and July in the U.S. (Vrat 1949, Moyle 1976,
    andnestbuildingoccursonthebottominshallowwater    Wootton 1976,  Wydoski and Whitney 1979, Wang
    areas in both freshwater and marine habitats, but the    1986).  In the Mediterranean, sticklebacks begin
    success of reproduction in marine environments is   breeding in March, when watertemperatures are 1 0�C
    uncertain (Vrat 1949, Hart 1973).                       and the spawning season lasts about 50 days (Crivelli
                                                           and Britton 1987). During the breeding season the

                                                       195

    Threespine stickleback continued
    male becomes territorial (McPhail and Lindsey 1970,    Food and Feeding
    Wootton 1976), its body develops green and orange-    Trophic Mode: Larvae are planktonic carnivores.
    red spawning colors, and the eyes become blue. The    Juveniles and adults are opportunistic carnivores that
    male builds a nest out of available material (sand,   willfeedonbenthicandplanktonicorganismsdepending
    algae, etc.). The nest can be an irregular cocoon with   on prey availability (Hart 1973, Scott and Crossman
    two openings or a hollow sandy pit below a pad of   1973, Wydoski and Whitney 1979). Sticklebacks prefer
    material (Wang 1986). The male performs a zig-zag   planktonic prey, but will switch to benthic prey as
    dance to entice the female to his nest to deposit her   zooplankton densities are reduced (Ibrahim and
    eggs. After she has deposited her eggs and left, the   Huntingford 1989). Sticklebacks may not feed on the
    male fertilizes them.  Males may spawn with many    most abundant zooplankton if it is too large to be
    different females, and females with different males.    ingested (Williams and Delbeek 1988), and may be
    After rearing one clutch, the male may rebuild his nest   slow in exploiting new food resources (Moyle 1976). In
    and starts again (Moyle 1976, Wootton 1976, Morrow    areas where sympatric stickleback species occur,
    1980).  Depending on food supply, a female may    competition for food is not thought to occur because of
    spawn up to 20 times during a spawning season    abundance of prey and morphological constraints on
    (Wootton 1976). Highly aggressive males appear to   feeding behavior (Delbeek and Williams 1988).
    have lower breeding success than less aggressive
    males (Ward and FitzGerald 1987).                    Food Items:While in freshwaterand estuarine habitats,
                                                          the threespine stickleback consumes calanoid
    Fecundity: Females lay about 20-300 eggs per   copepods, cyclopoid copepods, cladocerans (e.g.,
    spawning (depending on female size) (Wootton 1976);    Daphnia spp.), ostracods, aquatic insect larvae, snails,
    average fecundity is probably near 200 (Bolduc and   terrestrial insects, annelids, spiders, larval fish, and
    FitzGerald 1989). Overall seasonal fecundity appears    amphipods (e.g., Corophium spp.) (Manzer 1976). In
    to be related to the amount of time spent on the   marine environments,calanoid copepods (Centropages
    breeding grounds (Bolduc and FitzGerald 1989).   typicus, Eurytemoraspp.,andothers),copepodnauplii,
    Trachurus forms are more fecund than leiurus forms    euphausiid larvae, decapod larvae, and clam larvae
    (Wootton 1976, Mori 1990).                           are eaten (Maitland 1965, Hart 1973, Moyle 1976,
                                                          Wydoski and Whitney 1979, Worgan and FitzGerald
    Growth and Development                                1981, Bottom et al. 1984, Snyder 1984).  Female
    Eaa Size and Embryonic DeveloDment: Eggs are   sticklebacks will cannibalize eggs if a nest is left
    spherical and 1.1-1.9 mm in diameter (Vrat 1949,    unguarded by a male (Smith and Whoriskey 1988).
    Wootton 1976, Wang 1986). Embryonic development
    is indirect and external. Eggs take 7 or8 days to hatch    Biological Interactions
    at 18-190C (Wootton 1976, Wang 1986). However,    Predation: The threespine stickleback is an important
    time to hatching can range from 6-40 days depending    preyfor manyfishes[e.g., cutthroattrout (Oncorhynchus
    on temperature (Wootton 1976).                        clark/), rainbowtrout (0. mykiss), laketrout (Salvelinus
                                                          namaycush), Dolly Varden (Salvelinus malma), northem
    Aae and Size of Larvae: Larvae are 3.0-5.5 mm at   pike (Esoxlucius), northern squawfish (Ptychocheilus
    hatching, depending on location. Metamorphosis to   oregonensis), yellow perch (Perca flavescens)], birds
    juvenile begins in about 30 days at approximately 10   (e.g., herons, gulls, terns, diving ducks, and
    mm total length (TL) (Vrat 1949, Bigelow and Schroeder    mergansers), and some mammals (Hart 1973, Wootton
    1953, Wootton 1976, Wang 1986).                       1976, Morrow 1980).  Adult sticklebacks also eat
                                                          stickleback eggs and larvae.
    Juvenile Size Ranae: Juveniles are probably 11-30
    mm TL, depending on location and availability of food    Factors Influencina Populations: In lakes, thethreespine
    (Wootton 1976).                                      stickleback may compete with sockeye salmon (O.
                                                          nerka) for food (Foerster 1968). However, sticklebacks
    Aae and Size of Adults: Most populations of sticklebacks   usually do not inhabit the limnetic zone (where sockeye
    mature within one year and at approximately 30 mm TL    typically reside), so food competition is probably minimal
    (Jones and Hynes 1950, Wootton 1976). Theycan live   (Manzer 1976). A variety of parasites are believed to
    to4years and 76-85 mm long (Wootton 1976, Wydoski    affect the stickleback's feeding behavior and
    and Whitney 1979). Some are reported to have grown    susceptibilityto predation (Wootton 1976, Milinski 1986).
    larger than 102 mm (Scott and Crossman 1973). In   Temperature, food availability, predation, competition,
    California, the maximum age is probably 2 or 3 years    and parasitism play a role in determining population
    (Moyle 1976, Wang 1986).                             size, but which factor has the greatest influence is
                                                          unknown (Wootton 1976). The number of lateral plates

                                                       196

                                                                                Threespine stickleback continued

    appears to be directly related to predation pressure    Biol. 32:41-62.
    (Morrow 1980). Population abundances are also
    influenced by harsh environmental conditions during    Foerster, R. E. 1968. Thesockeye salmon. Fish. Res.
    breeding and overwintering (Whoriskey et al. 1986).    Board Can., Bull. No. 162, 422 p.
    Spawners using brackish-water pools appear to suffer
    greater egg cannibalism and bird predation than    Garrison, K. J., and B. S. Miller. 1980. Review of the
    freshwater spawners (Kedney et al. 1987).               early life historyof Puget Sound fishes. Fish. Res. Inst.,
                                                           Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).
    References
                                                           Gates, D. E., and H. W. Frey. 1974. Designated
    Andriyashev, A. P. 1954. Fishes of the northern seas    common  names of certain marine organisms of
    of the U.S.S.R. Acad. Sci. Union Soviet Soc. Rep. No.    California. Calif. Fish Game, Fish Bull. 161:55-90.
    53 (In Russian). Transl. by Israel Prog. Sci. Transl.
    Ltd., 1964, 617 p.                                      Hagen, D. W. 1967. Isolating mechanisms in threespine
                                                           sticklebacks. J. Fish. Res. Board Can., 24(8):1637-
    Bakker, T. C. M., and E. Feuth-De Bruijn.  1988.    1691.
    Juvenile territoriality in stickleback Gasterosteus
    aculeatus L., Anim. Behav. 36(5):1556-1559.             Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
                                                           Board Can., Bull. No.180, 740 p.
    Bell, M. A. 1976. Evolution of phenotypic diversity in
    Gasterosteus aculeatus superspecies on the Pacific    Ibrahim, A. A., and F. A. Huntingford. 1989. Laboratory
    coast of North America. Sys. Zool. 25(3):211-227.       and field studies on diet choice in three-spined
                                                           sticklebacks, Gasterosteus aculeatus L., in relation to
    Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of   profitability and visual features of prey. J. Fish Biol.
    the Gulf of Maine. Fish. Bull., U. S. 74(53):1-577.     34:245-257.

    Blahm, T. H., and G. R. Snyder.  1975.  Effect of   Jones, J. W., and H. B. N. Hynes. 1950. The age and
    increased water temperature on survival of adult    growth of Gasterosteus aculeatus, Pygosteuspungitius
    threespine stickleback and juvenile yellow perch inthe    and Spinachia vulgaris, as shown by their otoliths. J.
    Columbia River. Northw. Sci. 49(4):267-270.             Anim. Ecol. 19:59-73.

    Bolduc, F., and G. J. FitzGerald. 1989. The role of    Kedney, G. I., V. Boule, and G. J. FitzGerald. 1987.
    selected environmental factors and sex ratio upon egg    The reproductive ecology of threespine sticklebacks
    production in threespine sticklebacks, Gasterosteus    breeding in fresh and brackish water. Am. Fish. Soc.
    aculeatus. Can. J. Zool. 67:2013-2020.                  Symp. 1:151-161.

    Bottom, D. L., K. K. Jones, and M. J. Herring. 1984.    Lavin, P. A., and J. D. McPhail.  1986.  Adaptive
    Fishes of the Columbia River estuary. Col. Riv. Data    divergence of trophic phenotype among freshwater
    Dev. Prog., CREST,  Astoria, OR,  113 p. plus    populationsofthethreespinestickleback (Gasterosteus
    appendices.                                             aculeatus).  Can. J. Fish. Aquat. Sci. 43(12):2455-
                                                           2463.
    Carlander, K. D. 1969. Handbookof freshwaterfishery
    biology. Iowa State Univ. Press, Ames, IA, 752 p.       Maitland, P. S.  1965. The feeding relationships of
                                                           salmon, trout, minnows, stone loach and three-spined
    Clemens, W. A., and G. V. Wilby. 1961. Fishes of the    sticklebacks in the River Endrick, Scotland. J. Anim.
    Pacific coast of Canada. Fish. Res. Board Can., Bull.   Ecol. 34(1):109-133.
    No. 68, 443 p.
                                                           Manzer, J. I. 1976. Distribution, food, and feeding of
    Crivelli, A. J., and R. H. Britton. 1987. Life history   thethreespinestickleback, Gasterosteusaculeatus, in
    adaptations of Gasterosteus aculeatus in a    Great Central Lake, Vancouver Island, with comments
    Mediterranean wetland. Envir. Biol. Fish. 18(2):109-    on competition for food with juvenile sockeye salmon,
    125.                                                    Oncorhynchus nerka. Fish. Bull., U.S. 74(3):647-668.

    Delbeek, J. C., and D. D. Williams. 1988. Feeding    McPhail, J. D., and C. C. Lindsey. 1970. Freshwater
    selectivity of four species of sympatric stickleback in   fishes of northwestern Canada and Alaska. Fish. Res.
    brackish-water habitats in eastern Canada.  J. Fish    Board Can., Bull. No. 173, 381 p.

                                                        197

    Threespine stickleback continued
    Milinski, M. 1986. A review of competitive resource    aculeatus) of California. Copeia 4:252-260.
    sharing under constraints in sticklebacks. J. Fish Biol.
    29(suppl. A):1 -14.                                    Wang, J. C. S. 1986. Fishes of the Sacramento-San
                                                           Joaquin estuary and adjacent waters, California: A
    Miller, R. R., and C. L. Hubbs. 1969. Systematics of   guide to the early life histories.  Tech. Rep. No. 9.
    Gasterosteus aculeatus with particular reference to    Interagency ecological study program for the
    intergradationandintrogressionalongthePacificcoast    Sacramento-San Joaquin estuary. Calif. Dept. Water
    of North America: a commentary on a recent    Res., Calif. Dept. Fish Game, U.S. Bureau Recl., and
    contribution. Copeia 1969(1):52-69.                     U.S. Fish Wildl. Serv., various pagination.

    Mori, S.  1990.  Two morphological types in the    Ward, G., and G.J. FitzGerald. 1987. Male aggression
    reproductive stock of three-spined stickleback,    and female mate choice in the threespine stickleback,
    Gasterosteus aculeatus, in Lake Harutori, Hokkaido    Gasterosteus aculeatus L. J. Fish. Biol. 30:679-690.
    Island. Env. Biol. Fish. 27:21-31.
                                                           Whoriskey, F. G., and G. J. FitzGerald. 1989. Breeding-
    Morrow, J. E. 1980. The freshwater fishes of Alaska.    season  habitat use by sticklebacks (Pisces:
    Alaska Northw. Publ. Co., Anchorage, AK, 248 p.         Gasterosteidae) at Isle Verte, Quebec. Can. J. Zool.
                                                           67:2126-2130.
    Moyle, P. B. 1976. Inland fishes of California. Univ.
    Calif. Press, Berkeley, CA, 405 p.                      Whoriskey, F. G., G. J. FitzGerald, and S. G. Reebs.
                                                           1986. The breeding-season population structure of
    Okada, Y. 1955. Fishesof Japan. MaruzenCo., Ltd.,   three sympatric territorial sticklebacks (Pisces:
    Tokyo, Japan, 434 p.                                    Gasterosteidae). J. Fish. Biol. 29:635-648.

    Quinn, T. P., and J. T. Light. 1989. Occurrence of   Williams, D. D., and J.C. Delbeek. 1988. Biologyof the
    threespine sticklebacks (Gasterosteus aculeatus) in   threespine stickleback, Gasterosteus aculeatus, and
    the open North Pacific Ocean: migration or drift? Can.    the blackspotted stickleback, G. wheatlandi, during
    J. Zool. 67:2850-2852.                                  their marine pelagic phase in the Bay of Fundy, Canada.
                                                           Env. Biol. Fish. 24(1):33-41.
    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list   Wootton, R. J. 1976. The biology of the sticklebacks.
    of common and scientific names of fishes from the    Academic Press, New York, NY, 387 p.
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.            Worgan, J. P., and G. J. FitzGerald. 1981. Diel activity
                                                           and diet of three sympatric sticklebacks in tidal salt
    Scott, W. B., and E. J. Crossman. 1973. Freshwater    marsh pools. Can. J. Zool. 59:2375-2379.
    fishes of Canada. Fish. Res. Board Can., Bull. No. 184,
    966 p.

    Simenstad, C. A. 1983. The ecology of estuarine
    channels of the Pacific Northwest coast: a community
    profile. U.S. Fish Wildl. Serv., FWS/OBS-83/05,
    181 p.

    Smith, R. S., and F. G. Whoriskey, Jr. 1988. Multiple
    clutches: femalethreespine sticklebacks lose the ability
    to recognize their own eggs. Anim. Behav. 36(6):1838-
    1839.

    Snyder, R.J. 1984. Seasonal variation in the diet of the
    threespine stickleback, Gasterosteus aculeatus, in
    Contra Costa County, California. Calif. Fish Game
    70(3) :167-172.

    Vrat, V. 1949. Reproductivebehavioranddevelopment
    of eggs of the three-spined stickleback (Gasterosteus

                                                        198

    199

       Morone saxatilis
       Adult

                  10 cm

    Common Name: striped bass                               The San Francisco Bay striped bass fishery was one of
    Scientific Name: Morone saxatilis                       the most important recreational fisheries on the Pacific
    Other Common  Names: striper, streaked bass,    coast, with annual landings ranging from 107,000 to
    squidhound, rock, rock bass, rock fish, greenhead,    403,000 fish in 1978 and 1975, respectively (White
    linesider, roller (Gates and Frey 1974, Fay et al. 1983)    1986). The value of this fishery was estimated to be
    Classification (Robins et al. 1980)                     over $45 million (Meyer Resources 1985, cited by
    Phylum: Chordata                                        Stevens et al. 1987). However, stock size has dropped
    Class: Osteichthyes                                     dramatically;only slightly morethan 72,000 were caught
    Order: Perciformes                                      along the Pacific Coast in 1985 (National Marine
    Family: Percichthyidae                                  Fisheries Service 1986).

    Value                                                   Indicatorof Environmental Stress: It appears that certain
    Commercial: Small numbers (135 yearlings) of striped    petrochemicals  interact with other pollutants
    bass were introduced to California's San Francisco    (polychlorinated biphenyls and heavy metals) to
    Bay in 1879, and 300 were released in 1882. In 1899,    adversely affect striped bass populations in San
    560 t were landed in San Francisco Bay (Hassler    Francisco Bay (Whipple 1984). High concentrations of
    1988).  Historically, this species was commercially    organochlorines, metals, and petrochemicals have been
    caught on the Pacific coast in San Francisco Bay and    found in striped bass tissues (Whipple et al. 1983).
    Coos Bay, Oregon.   Until 1915, the annual San    Correlations exist between pollutants and parasite
    Francisco Baycatch usuallyexceeded 454t; thereafter    burdens, body condition, liver condition, and egg and
    only twice did harvest exceed this value (Smith and    gonad conditions. Fish exposed to a chronic pollutant
    Kato 1979). In 1935, commercial fishing for striped    stress have significant reductions in reproductive
    bass in the San Francisco Bay system was prohibited    capacity, fecundity, and gametic viability  (Whipple
    because of demands by sport anglers (Smith and Kato    1984).
    1979, Stevens et al. 1987). Oregon has prohibited
    commercial fishing for this species since1976 (Parks    Ecological: Inthe estuaries where it occurs, M. saxatilis
    1978).                                                 is one of the most important predators of estuarine
                                                           fishes and invertebrates.
    Recreational: The striped bass is an important sport
    fish from north/central California to southemrn Oregon. It   Range
    is highly sought because of its fighting ability, large    Overall: On the Pacific coast, the striped bass is found
    size, easy accessibility, and excellent taste. Most are    from about 40 km south of California-Mexico border to
    taken by hook and line using artificial or natural baits.    Barkley Sound, British Columbia (Millerand Lea 1972),
    In the San Francisco Bay system, most sport fishing   but is not common south of Monterey, California, or
    took place in San Pablo Bay and the Delta, but now    north of the Siuslaw River, Oregon (Parks 1978). On
    occurs in San Francisco Bay proper (Stevens 1977).    the Atlantic coast, itoccursfromtheSt. Lawrence River

                                                        200

                                                                                        Striped bass continued

                                                          1964, Scott and Crossman 1973, Wang 1986). Larvae
     Table 1. Relative abundance of striped bass          are initially feeble swimmers- if they encounter still
              in 32 U.S. Pacific coast estuaries.         waterthey settle to the bottom and die (Skinner 1962).
                        Life Stage                        Postlarval stages ("fry") inhabit lower river channels
           Estuary    A S J L E                           and upper estuarine shallow-water bays and sloughs
            Puget Sound :1       I  Relative abundance:   (Skinner 1962, Sasaki 1966a, Wang 1986). Juveniles,
             Hood Canal         :         Highly abundant  subadults, and adults are pelagic but are somewhat
              Skagit Bay i'           I   Abundant        bottom-oriented (Skinner 1962, Sasaki 1966b), as are
            Grays Harbor              O   Common          the eggs and early larvae (Turner 1976). Juveniles and
             Willapa Bay             iJ  Rare             adults are anadromous and form small separate (by
           Columbia River            Blank Not present     size or age) schools or feeding groups (Raney 1952).
            Nehalem Bay
            Tillamook Bay            Life stage:           Habitat
             NetartsBay              A wnngAdults         Tje: Eggs and larvae are found in lower riverine
             Siletz River 1         J-Juveniles           (freshwater) areas and upper estuarine (oligohaline)
             rYaquina  Bay'i         E - Eggs              areas. Young-of-the-year also occur in these areas,
             Alsea River                                  with many moving to more saline environments
            Siuslaw River o0    0 o o                     (mesohaline and polyhaline) in the fall (Calhoun 1953,
            Umpqua River 0 0 0 0 0                         Sasaki 1966a). Juveniles may also move into rivers
               Coos Bay 0 0 0 0 0                          upstream of estuaries (Turner 1972). Older juveniles
             Rogue River                                   may be found in all estuarine areas, but appear to
            Klamath River '                                prefer certain areas (Skinner 1962), perhaps because
            Humboldt Bay                                   of food availability and temperature. Young striped
               Eel River                                  bass can be highly abundant in mixing areas of estuaries
            TonmaiesBSay O         ,clude     Swhere fresh water and salt water mix (Turner 1972).
       Cent. San~tan BaY- 8 � 8 (3 �Francisco, Suisun.    This area is often referred to as the "null zone" or
       SouthSan Fran.Bay     0       and San Pablo bays.  "critical zone". Adults are found in the lower estuary
           ElkhhOm Slough                                 (polyhaline and euhaline waters) from late spring to
              Morro Bay                                   early fall, in the upper (mesohaline and oligohaline)
           Sana Pedro Bay                                 areas in late fall and winter, and in freshwater and
             SAramitos Bay                                 oligohaline areas during spawning.   Temperature
                                                           appears to be an important determinant of the estuarine
            Anaheim Bay
             Newpo nBay I                                  distributions of juveniles and adults (Coutant 1986,
             Mission Bay J                                1987).
           San Diego Bay
           Tijuana Estuary                                 Substrate: Eggs and larvae are swept over various
                       A S J L E                           sediments. Juveniles appear to prefer clean sandy
                                                           bottoms, but have been found over gravel beaches,
                                                           rock, mud, and mixed sand and silt bottoms (Setzler et
    down to the St. Johns River, Florida, and into streams    al. 1980). Adults and subadults are also found over
    thatflowintotheGulfofMexicofromFloridatoLouisiana    various substrates, such as sandy beaches, rocky
    (Moyle 1976). Stocking into reservoirs has established    shores, and mussel beds (Setzler et al. 1980).
    some self-sustaining freshwater populations (Moyle
    1976).                                                 Phvsical/Chemical Characteristics: Striped bass eggs
                                                           are found in fresh water to 11%o salinity (Rulifson et al.
    Within Studv Area: M. saxatilis was introduced to the    1982).  Optimum salinities for egg survival are 1.5-
    Sacramento-San Joaquin River system during the    3.0%0 (Mansueti 1958, cited by Fay et al. 1983). Eggs
    1870s. ItisfoundmainlyinestuariesfromSanFrancisco    can withstand temperatures of 12-24�C (Fay et al.
    Bay north to the SiuslawRiver(Tablel)(Monacoetal.    1983), with the optimum being 180C (Morgan et al.
    1990). It has been stocked in some southern California   1981). Larvae tolerate temperatures of 10-250C, but
    bays, but these populations are not self-sustaining    optimal temperatures for survival are 15-220C (Fay et
    (Horn et al. 1984).                                    al. 1983). Preferred temperatures change as the fish
                                                           grow older (Coutant 1986). Adults can withstand
    Life Mode                                              temperatures as high as 35�C, but become stressed at
    Eggs are non-adhesive, slightly heavier than fresh    temperatures above 25�C (Moyle 1976). They tolerate
    water, and are swept along with currents (Albrecht    temperatures of 0-32�C, but prefer 20-240C (Fay et al.

                                                       201

    Striped bass continued
    1983, Coutant 1986). Adults can also withstand low    after eggs are extruded from the female.   High
    dissolved oxygen (4.0 ppm) and high turbidity, but this   concentrations of total dissolved solids (>180 ppm)
    will inhibit reproduction (Moyle 1976).  Optimum    may block spawning migrations (Farley 1966, Radtke
    spawningtemperaturesare15.6-20.0�C,with spawning    1966).  Cooler water temperatures in spring allow
    ceasing at 21.1 C (Moyle 1976). Dissolved oxygen    striped bass to move further upriver to spawn (Farley
    levels below 4.0 ppm with temperatures of 22.2�C    1966). Successful spawning requires the following: 1)
    reduced egg survival by more than 50% (Turner and    a large river, 2) water velocities sufficient to keep eggs
    Farley 1971). Low oxygen levels (2.0-3.5 ppm) may    and larvae suspended off the bottom but not so fast that
    have eliminated some spawning areas inthe Delaware    it washes them to calm waters before the larvae can
    River, New Jersey (Setzler et al. 1980).               swim, and 3) an estuary where young can feed and
                                                          grow (Moyle 1976). Striped bass have a tendency to
    Miarations and Movements: Atlantic population    returntothesamespawningareaeachyear(Chadwick
    prespawners may travel long distances upriver in fresh    1967).
    water (Scott and Crossman 1973), however Pacific
    populationsdo not. Unlike some east coast populations    Fecundity: Fecundity depends on the age and size of
    that make extensive coastal migrations, Sacramento-    the female.  In San Francisco Bay, mean fecundity
    San Joaquin River populations and other Pacific coast    ranges from 243,000 (for 4 year-olds) to 1,427,000 for
    populations appearto spend most of their lives in bays    8 year-olds and older (Stevens et al. 1985). Up to
    and estuaries. This may be related to the cool oceanic    5,300,000 eggs may be produced by very large females
    temperatures found off the Pacific coast (Radovich    (Skinner 1962, Wang 1986).
    1963). San Francisco Bay adults move into bays
    (some into the Delta) in the fall, overwinter in the Bay    Growth and Development
    and Delta, and then after spawning in spring, move    Eaa Size and Embrvonic DeveloDment: Eggs are 3.3-
    back to salt water (Calhoun 1952, Moyle 1976). Eggs    4.2 mm in diameter, averaging 3.3 mm in California
    and larvae are transported downstream by river flow   populations (Woodhull 1947, Doroshev 1970). Eggs
    into lower river and estuarine areas or may stay in the    are spherical, nonadhesive, slightly heavierthan fresh
    general spawning area if this is an area where outflow   water, and nearly transparent when developing (Wang
    is balanced by tidal currents (Moyle 1976). Larvae    1986). Embryonicdevelopmentisindirectandextemal.
    school within 4 or 5 days of hatching and are found    Eggs hatch in about 1.5-3.5 days (temperature
    primarily in shallow water shore zones of fresh and    dependent), 2 'days at optimum temperatures (16-
    brackish waters (Rulifson et al. 1982). Although there    19�C) (Doroshev 1970). Hatching times range from 48
    is some straying, each Pacific coast river system    hours (at 17.8-19.4�C) to 70-74 hours (at 14.4-15.6�C)
    appears to have a distinct stock (McGie and Mullen    (Scott and Crossman 1973).
    1979).
                                                           Aae and Size of Larvae: Larvae are 2.0-3.7 mm total
    Reproduction                                           length (TL) at hatching, averaging 2.9-5.0 mm TL on
    Mode: The striped bass is gonochoristic (occasionally    the Pacific coast (Wang 1986). Absorption of the yolk
    hermaphroditic), polygamous, and oviparous; eggs    sac is highly variable and dependent on temperature;
    are fertilized externally. It is iteroparous, but mature    from 3 days at 24�C to 9 days at 120�C (Setzler et al.
    females maynotspawneveryyear(Raney 1952, Scott    1980).  Development  from the  finfold stage
    and Crossman 1973).                                    (metamorphose) to juvenile varies with temperature,
                                                           reportedly taking 23 days at 240�C to 68 days at 150C
    Matina/SDawnina: Spawning occurs in riverine    (Rogers et al. 1977, cited by Hassler 1988).  Final
    (freshwater) or slightly brackish waters in the upper    length of larvae before the development of the second
    portionsofestuaries (Hart 1973). In California, spawning    dorsal fin ranges from 25.0 to 36.0 mm (Hardy 1978).
    begins in April, and peaks in May and early June,
    depending on temperature, river flow, and salinity   Juvenile Size Rance: Juveniles are typically 2-3 cm
    (Turner 1972, 1976). Striped bass are mass spawners.    fork length (FL) in their first year, 23-35 cm FL in their
    During spawning runs they will gather close to shore    second, 38-39 cm FL in their third, and 48-50 cm FL in
    with groups (5-30 fish) breaking off to spawn in the   their fourth year. Thereafter, growth is only 1-3 cm/
    main river channel. Actual spawning occurs near the    year (Moyle 1976). Striped bass in Oregon tend to
    surface, with individuals frequently turning on their   grow larger than California stocks (McGie and Mullen
    sides and splashing at the surface (Woodhull 1947,    1979).
    Moyle 1976). Spawning activity usually peaks during
    late afternoon or early evening (Moyle 1976).    Ace and Size of Adults:Some males may mature atthe
    Fertilization is external, andmustoccurwithinonehour    end of their first year, but most mature during their

                                                       202

                                                                                         Striped bass continued
    second and third year; all are mature by the fifth year   system indicates that production of young bass has
    (Moyle 1976). Most females matureduringtheirfourth    been exceptionally low since 1977 (even considering
    or fifth year (87%) and all are mature by their seventh    river flows). Reasons forthis decline include increased
    (Hart 1973; Moyle 1976).  At first spawning, males    adult mortality, inadequate egg production, reduced
    average 25 cm FL, while females average 45 cm FL.   plankton food for young striped bass as a result of
    The maximum size is 122 cm long and 41 kg, but fish   water diversions, large numbers of eggs and young
    in Pacific populations are usually less than 4.5 kg    bass being entrained by freshwater diversions, and
    (Eschmeyer et al. 1983). The maximum age of the    high levels of contaminants (Stevens et al. 1985,
    striped bass is >30 yr, and these are usually females    California Department of Fish and Game 1987). Adult
    (Moyle 1976).                                         mortality may also be increasing because changes in
                                                           water flow have "squeezed" (i.e., limited its preferred
    Food and Feeding                                      habitat) this species between its thermal and dissolved
    Trophic Mode: Striped bass larvae are pelagic   oxygen preferences or requirements (Coutant 1985,
    carnivores. Juveniles and adults are opportunistic,    1986,1987). An overall decrease in the San Francisco
    top-level epibenthic and pelagic carnivores that feed    Bay population appears to be due to the interactive
    on invertebrates and fish (depending on the striped   effects of reduced freshwater flows, increased
    bass' size and food availability) (Moyle 1976). They    freshwater diversions, decreased bay flushing, and
    are reported to not feed continuously, but gorge    increased body burdens of pollutants which have
    themselves and then wait until digestion is complete    reduced egg production and egg and larval survival
    (Scott and Crossman 1973). They feed most intensively   (Setzler-Hamilton et al. 1988). High rates of infestation
    from after spawning through October.                  by ectoparasites (e.g., Nerocila californiensis) in some
                                                           bays may be detrimental (Horn et al. 1984). Successful
    Food Items: On the Pacific coast, the food habits of   reproduction in Oregon appears to depend on optimal
    striped bass in the Sacramento-San Joaquin Delta   conditions of temperature and riverf low, often resulting
    have been well-studied. Large juveniles and adults   in the striped bass population being dominated by one
    feed on fishes and large invertebrates such as Crangon    year-class (McGie and Mullen 1979).
    spp., while smaller juveniles are primarily invertebrate
    feeders; Neomysis mercedis, Corophiumspp., Crangon    References
    spp., and copepods and cladocera, are primary prey
    (Ganssle 1966, Turner 1972). Important fish prey for   Albrecht, A. B. 1964. Some observations on factors
    larger juveniles and adults include threadfin shad    associated with survival of striped bass eggs and
    (Dorosoma petenense), threespine stickleback    larvae. Calif. Fish Game 50(2):100-113.
    (Gasterosteus aculeatus), American shad (Alosa
    sapidissima), pond smelt (Hypomesus olidus), juvenile   Calhoun, A. J. 1952. Annual migrations of California
    chinook salmon (Oncorhynchustshawytscha), northern    striped bass. Calif. Fish Game 38(3):391 -403.
    anchovy (Engraulis mordax), Pacific staghorn sculpin
    (Leptocottus armatus), various smelt species, and    Calhoun, A. J. 1953. Distribution of striped bass fry in
    young-of-the-yearstriped bass (Johnson and Calhoun    relation to major water diversions. Calif. Fish Game
    1952, Stevens 1966).                                  39(3):279-299.

    Biological Interactions                               Califomia Department of Fish and Game. 1987. Factors
    Predation: Man and large marine mammals [e.g., harbor   affecting striped bass abundance in the Sacramento-
    seals (Phoca vitulina) and sea lions] are probably the   San Joaquin River system. Exhibit 25, entered by the
    onlypredatorsofadultstripedbass. Juveniles areprey    California Department of Fish and Game for the State
    for large striped bass and other piscivorous fishes.  Water Resources Control Board 1987 Water Quality/
                                                          Water Rights Proceeding on the San Francisco Bay/
    Factors Influencina PoDulations: Survival of larvae    Sacramento-San Joaquin Delta.  Calif. Dept. Fish
    appears to strongly determine recruitment to the adult   Game, Stockton, CA, 149 p. plus appendices.
    life stage. Factors which affect larval survival are
    temperature, salinity, predation, food availability   Chadwick, H. K.  1967.  Recent migrations of the
    (Eldridge et al. 1981), and pollution. One of the major    Sacramento-SanJoaquin Riverstripedbasspopulation.
    determinants is the amount of freshwater discharge    Trans. Am. Fish. Soc. 96(3):327-342.
    during summer. Normally, the higher the summer
    flows, the higher the larval survival rate (Sommani    Coutant, C. C. 1985. Striped bass, temperature, and
    1972, Turner 1972, Turner and Chadwick 1972).    dissolved oxygen: a speculative hypothesis for
    However, recent research in the San Francisco Bay    environmental risk. Trans. Am. Fish. Soc. 114:31-61.

                                                       203

    Striped bass continued
    Coutant, C. C. 1986. Thermal niches of striped bass.    Fish. Wildl. Serv. Biol. Rep. 82(11.82).  U.S. Army
    Sci. Am. 255(2):98-104.                                 Corps. Eng., TR EL-82-4, 29 p.

    Coutant, C. C. 1987. Thermal preference: when does    Horn, M. H., L. G. Allen, and F. D. Hagner.  1984.
    an asset become a liability? Envir. Biol. Fish. 18(3):1 61-    Ecological status of striped bass, Morone saxatilis, in
    172.                                                   upper Newport Bay, California.  Calif. Fish Game
                                                           70(3):180-1 82.
    Doroshev, S. I. 1970. Biological features of the eggs,
    larvae and young of the striped bass [Roccus saxatilis   Johnson, W. C., and A. J. Calhoun. 1952. Food habits
    (Walbaum)] in connection with the problem of its   of California striped bass. Calif. Fish Game38(4):531-
    acclimatization in the USSR. J. Ichthyol. 10:235-248.    534.

    Eldridge, M. B., J. A. Whipple, D. Eng, M. J. Bowers,    Mansueti, R. J. 1958. Eggs, larvae, and young of the
    and B. M. Jarvis. 1981. Effects of food and feeding    striped bass. Chesapeake Lab. Biol. Contr. No. 112,
    factors on laboratory-reared striped bass larvae. Trans.    35 p.
    Am. Fish. Soc. 110:111-120.
                                                           McGie, A. M., and R. E. Mullen. 1979. Age, growth,
    Eschmeyer, W. N., W. S. Herald, and H. Hammann.    and population trends of striped bass, Moronesaxatilis,
    1983. A field guide to Pacific coast fishes of North    in Oregon. Info. Rep. Ser., Fish. No. 79-8. Oregon
    America. Houghton Mifflin Co., Boston, MA, 336 p.       Dept. Fish Wildl., Corvallis, OR, 57 p.

    Farley, T. C. 1966. Striped bass, Roccus saxatilis,    Meyer Resources.  1985.  The economic value of
    spawning in the Sacramento-San Joaquin River    striped bass, Morone saxatilis, chinook salmon,
    systems during 1963 and 1964. InJ. L. Turner and D.    Oncorhynchustshawytscha, andsteelheadtrout, Salmo
    W. Kelley (compilers), Ecological studies of the    gairdneri, of the Sacramento and San Joaquin river
    Sacramento-San Joaquin Delta.  Calif. Fish Game,    systems. Admin. Rep. 85-3, Anad. Fish. Branch, Calif.
    Fish Bull. 136:28-43.                                   Dept. Fish Game, Sacramento, CA.

    Fay, C. W., R. J. Neves, and G. B. Pardue.  1983.    Miller,D.J.,andR. N. Lea. 1972. Guidetothecoastal
    Species profiles: life histories and environmental    marinefishesof California. Calif. Fish Game, Fish Bull.
    requirements of coastal fishes and invertebrates (Mid-    157, 235 p.
    Atlantic) - striped bass. U.S. Fish Wildl., Div. Biol.
    Serv., FWS/OBS-82/11.8. U.S. Army Corps Eng., TR    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    EL-82-4, 36 p.                                          Nelson. 1990. Distribution and abundance of fishes
                                                           and invertebrates in west coast estuaries, Volume I:
    Ganssle, D. 1966. Fishes and decapods of the San    data summaries.   ELMR  Rep. No. 4. Strategic
    Pablo and Suisun Bays. In D. W. Kelley (compiler),    Assessment Branch, NOS/NOAA, Rockville, MD,
    Ecological studies of the Sacramento-San Joaquin    240 p.
    estuary. Calif. Fish Game, Fish Bull. 133:64-94.
                                                           Morgan, R. P., II, V. J. Rasin, Jr., and R. L. Copp. 1981.
    Gates, D. E., and H. W. Frey.  1974.  Designated    Temperature and salinity effects on development of
    common names of certain marine organisms of   striped bass eggs and larvae. Trans. Am. Fish. Soc.
    California. Calif. Fish Game, Fish Bull. 161:55-90.     110:95-99.

    Hardy, J. P., Jr. 1978. Development of fishes of the    Moyle, P. B. 1976. Inland fishes of California. Univ.
    mid-Atlantic Bight. Vol. III. Aphredoderidae through    Calif. Press, Berkeley, CA, 405 p.
    Rachycentridae. U.S. Dept Int., U.S. Fish Wildl. Serv.,
    FWS/OBS-78/12.                                          National Marine Fisheries Service.  1986.  Marine
                                                           recreational fishery statistics survey, Pacific coast,
    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    1985. Current Fishery Statistics, No. 8328. Nat. Mar.
    Board Can., Bull. No. 180, 740 p.                       Fish. Serv., NOAA, Washington, D.C., 109 p.

    Hassler, T.J. 1988. Species profiles: life histories and    Parks, N. B. 1978. The Pacific Northwest commercial
    environmental requirements of coastal fishes and    fishery for striped bass, 1922-1974. Mar. Fish. Rev.
    invertebrates (Pacific Southwest) - striped bass. U.S.    40(1 ):18-20.

                                                       204

                                                                                         Striped bass continued
    Radovich, J. 1963. Effectofoceantemperatureonthe    environmentally impacted estuaries. Mar. Poll. Bull.
    seaward movements of striped bass, Roccus saxatifis,    19(9):466-477.
    on the Pacific coast. Calif. Fish Game 49(3):191-206.
                                                           Skinner, J. E. 1962. An historical review of the fish and
    Radtke, L. D. 1966. Distribution of adult and subadult    wildlife resources of the San Francisco Bay Area.
    striped bass, Roccussaxatilis, in the Sacramento-San    Water Proj. Br. Rep. No. 1, Calif. Dept. Fish Game,
    Joaquin Delta.  In J. L. Turner and D. W. Kelley    Sacramento, CA, 226 p.
    (compilers), Ecological studies of the Sacramento-San
    Joaquin Delta. Calif. Fish Game, Fish Bull. 136:15-27.    Smith, S. E., and S. Kato. 1979. The fisheries of San
                                                           Francisco Bay: past, present and future. In T.J.
    Raney, E. C. 1952. The life history of the striped bass,    Conomos (editor), San Francisco Bay: the urbanized
    Roccussaxatilis(Walbaum). Bull. BinghamOceanogr.    estuary, p. 445-468. Am. Assoc. Adv. Sci, and Calif.
    Coil. 4(1):1-95.                                        Acad. Sci., San Francisco, CA.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,    Sommani, P. 1972. Astudyonthe population dynamics
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list   of striped bass (Morone saxatilis Walbaum) in the San
    of common and scientific names of fishes from the    Francisco Bay estuary. Ph.D. Thesis., Univ. Wash.,
    United States and Canada. Am. Fish. Soc. Spec. Publ.    Seattle, WA, 1 14 p.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
                                                           Stevens, D. E. 1966. Food habits of striped bass,
    Rogers, B. A., D. T. Westin, and S. B. Saila. 1977. Life   Roccussaxatilis, intheSacramento-SanJoaquin Delta.
    stage duration in Hudson River striped bass.  Mar.    InJ. L. Turner and D. W. Kelley (compilers), Ecological
    Tech. Rep. No. 3, Univ. Rhode Island, Kingston, RI,   studies of the Sacramento-San Joaquin Delta. Calif.
    111 p.                                                  Fish Game, Fish Bull. 136:68-96.

    Rulifson, R. A., M. T. Huish, and R. W. Thoesen. 1982.    Stevens, D. E. 1977. Striped bass (Morone saxatilis)
    Status of anadromous fishes in southeastern U.S.    year class strength in relation to river flow in the
    estuaries.  In V. S. Kennedy (editor), Estuarine    Sacramento-San Joaquin estuary, California. Trans.
    comparisons, p. 413-425, Academic Press, New York,    Am. Fish. Soc. 106(1 ):34-42.
    NY.
                                                           Stevens, D. E., D. K. Kohlhorst, L. W. Miller, and D. W.
    Sasaki, S. 1966a. Distribution of young striped bass,    Kelley.  1985.  The decline of striped bass in the
    Roccussaxatilis, intheSacramento-SanJoaquin Delta.    Sacramento-San Joaquin estuary, California. Trans.
    InJ. L. Turner and D. W. Kelley (compilers), Ecological    Am. Fish. Soc. 114:12-30.
    studies of the Sacramento-San Joaquin Delta. Calif.
    Fish Game, Fish Bull. 136:44-58.                        Stevens, D. E., H. K. Chadwick, and R. E. Painter.
                                                           1987. American shad and striped bass in California's
    Sasaki, S. 1966b. Distribution of juvenile striped bass,    Sacramento-San Joaquin river system. Am. Fish. Soc.
    Roccussaxatilis, in the Sacramento-SanJoaquin Delta.    Symp. 1:66-78.
    InJ. L. Turnerand D. W. Kelley (compilers), Ecological
    studies of the Sacramento-San Joaquin Delta. Calif.   Turner, J. L. 1972. Striped bass.  In J. E. Skinner
    Fish Game, Fish Bull. 136:59-67.                        (editor), Ecological studies of the Sacramento-San
                                                           Joaquin estuary. Delta Fish Wildl. Protection Study
    Scott, W. B., and E. J. Crossman. 1973. Freshwater    Rep. No. 8, p. 36-43.  Calif. Dept. Fish Game,
    fishes of Canada. Fish. Res. Board Can., Bull. No. 184,    Sacramento, CA.
    966 p.
                                                           Turner, J. L. 1976. Striped bass spawning in the
    Setzler, E. M., W. R. Boynton, K. V. Wood., H. H. Zion,    Sacramento and San Joaquin rivers in central California
    L. Lubbers, N. K. Mountford, P. Frere, L. Tucker, and J.   from 1963 to 1972. Calif. Fish Game 62(2):106-118.
    A. Mihursky. 1980. Synopsis of biological data on
    striped bass, Morone saxatilis (Walbaum).   FAO    Turner, J. L., and H. K. Chadwick. 1972. Distribution
    Synopsis No. 121, 69 p.                                 and abundance of young-of-the-year striped bass,
                                                           Morone saxatilis, in relation to river flow in the
    Setzler-Hamilton, E. M., J. A. Whipple, and R. B.    Sacramento-San Joaquin estuary. Trans. Am. Fish.
    MacFarlane.   1988.  Striped bass populations in   Soc. 101(3):442-452.
    Chesapeake and San Francisco Bays: Two

                                                       205

    Striped bass continued
    Turner, J. L. and T. C. Farley. 1971. Effects of
    temperature, salinity, and dissolved oxygen on survival
    of striped bass eggs and larvae. Calif. Fish Game
    57:268-273.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: A
    guide to the early life histories. Tech. Rep. No. 9.
    Interagency ecological study program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    and U.S. Fish Wildl. Serv., various pagination.

    Whipple, J. A. 1984. The impact of estuarine
    degradation and chronic pollution on populations of
    anadromous striped bass (Moronesaxatilis) in the San
    Francisco Bay-Delta, California. A summary for
    managers and regulators. SWAFC Adm. Rep. T-84-
    01,47p. Southw. Fish. Center, Tiburon Fish. Lab., Nat.
    Mar. Fish. Serv., NOAA, 3150 Paradise Drive, Tiburon,
    CA 94920.

    Whipple, J. A., D. G. Crosby, and M. Jung. 1983. Third
    progress report, Cooperative striped bass study.
    CaliforniaStateWater Resources Control Board, Toxic
    Substances Control Program, Spec. Proj. Rep. No. 83-
    3sp, 208 p.

    White, J. R. 1986. The striped bass sport fishery in the
    Sacramento-San Joaquin estuary, 1969-1979. Calif.
    Fish Game 72(1):17-37.

    Woodhull, C. 1947. Spawning habits of the striped
    bass (Roccus saxatilis). Calif. Fish Game 33(2):97-
    102.

                                                       206

    207

       Paralabrax clathratus
       Adult

                    10cm

    Common Name: kelp bass                                 Industrial and domestic wastes are released in large
    Scientific Name: Paralabrax clathratus                 quantities near some kelp bass habitat, but the effects
    Other Common Names: California kelp bass, rock   of these pollutants on kelp bass survival is unclear.
    bass, sand bass, cabrilla, calico bass, bull bass, kelp
    salmon, lockee cod (Gates and Frey 1974)               Ecological: It is an abundant top-level predator in kelp
    Classification (Robins et al. 1980)                    beds off southern California, with juveniles and small
    Phylum: Chordata                                       adults often abundant in the surf zone (but not
    Class: Osteichthyes                                    intertidally) (Quast 1968a).
    Order: Perciformes
    Family: Serranidae                                     Range
                                                           Overall: The kelp bass' overall range is from Magdalena
    Value                                                  Bay, Baja California (including Guadalupe Island,
    Commercial: Since 1953, it has been illegal to sell kelp    Mexico) to the Columbia River (Miller and Lea 1972).
    bass harvested in California waters. A limited
    commercial catch may occur in Mexican waters (Frey    Within Study Area: This species is commonly found
    1971).                                                 south of Point Conception, but is rare in shallow water
                                                           bays and lagoons such as Newport Bay (Bane 1968),
    Recreational: The kelp bass is an important sport fish   Anaheim Bay, Alamitos Bay, and San Diego Bay
    in southern California, prized for its excellent taste,   (Table 1).  Juveniles can be common at times in
    good fighting ability, year-round availability, and    Mission Bay, California (Noah 1985). It is abundant in
    relatively high abundance.  It is caught from about    Santa Monica and San Pedro Bays, California (Quast
    Tomales Bay, California, to central Baja California, but    1968a, Carlisle 1969, Fay et al. 1978), and may be
    most effort occurs from Point Conception south to San    found in developed areas (e.g., marinas and harbors)
    Diego, California. Over 2.5 million were captured in   (Horn and Allen 1981, Stephens and Zerba 1981, Allen
    1985, the second highest catch of recreational fish in   et al. 1983).
    southern California (U.S. Department of Commerce
    1986). It is usually caught by party and private boats    Life Mode
    fishing over kelp beds and trolling with bait. Some are    Eggs and larvae are pelagic, while juveniles and adults
    also caught by spearfishing and shore and pier   are benthopelagic (Young 1963, Quast 1968a, Feder
    fishermen using hook and line (Young 1963, Quast    et al. 1974).
    1968a, Frey 1971).
                                                           Habitat
    Indicator of Environmental Stress: This species is   Type: Eggs and larvae are neritic-epipelagic and occur
    dependent on healthykelp beds. Temperatures above    near the surface. Juveniles are distributed from the
    24�C (e.g., wastedischarges from metropolitan centers)    surf zone out to depths of 30 m, but occur primarily
    appear to be detrimental to kelp beds (Quast 1968b).    inshore at depths of 8-20 m (Quast 1968a, Feder et al.

                                                       208

                                                                                          Kelp bass continued

                                                          adults often live in deep rocky areas containing little or
     Table 1. Relative abundance of kelp bass in          no algae (Feder et al. 1974).
              32 U.S. Pacific coast estuaries.
                         Life Stage                       Phvsical/Chemical Characteristics: A euhaline species,
             Estuary    A S J L E                          it is primarily found in waters of 33.5-34.5%0 and
              Puget Sound             Relative abundance:  temperatures of 13-280C (Quast 1968c, MBC Applied
              Hood Canal              *   Highly abundant    Environmental Sciences 1987). This species will avoid
               Skagit Bay                Abundant          areas with high turbidity (Quast 1968a).
             Grays Harbor              O   Common
              Willapa Bay              i   Rare           Miorations and Movements: No migrations are known
            Columbia River            Blank Not present    to occur. Adult home ranges appearto be up to 40 ha,
             Nehalem Bay                                  depending on habitat structure (Quast 1968a). Very
             Tillamook Bay            Life stage:          few kelp bass will move more than 16 km (Young
              Netarts Bay             A -  wninAdults     1963). As adult kelp bass are harvested from areas
                                      S - Spawning adults
               Siletz River           J - Juveniles        with good habitat, bass from adjacent areas will move
              Yaquina Bay             L-  Eggs             in to replace them (Quast 1968a).
               Alsea River
             Siuslaw River                                 Reproduction
             Umpqua River
                                                          Mode: The kelp bass is gonochoristic, oviparous, and
                Coos Bay                                   iteroparous.  It is a broadcast spawner; eggs are
              Rogue River                                  fertilized externally (Quast 1968a, Feder et al. 1974).
             Klamath River
             Humboldt Bay                                  Matina/SDawnina: Spawning takes place in relatively
                Eel River                                 deep water (to 46 m) over rough bottom in or near kelp.
              Gent. San Fran. Bay' Includes Central San    Spawning occurs from April to November, probably
                                      FranciscoSuisun,    peaking during June and July (Quast 1968a, Frey
        South San Fran. Bay           and San Pablo bays.  1971, Feder et al. 1974). Successful spawning probably
            Elkhorn Slough                                only occurs from Point Conception to Magdalena Bay,
               Morro Bay
               Santa Morroa Bay      0(I0Baja California (Quast 1968a).  Larger individuals
            San Pedro Bay    (D 0     0                    mature earlier and remain reproductively active longer.
              San PedroBayC0)   0 C)                      Hundreds of kelp bass may aggregate in a small area
             AnaheimBay  V    1                           during spawning (Feder et al. 1974).  Males often
              Newport Bay   develop a yellowcolorontheirsnoutduring the breeding
              Mission Bay 01   0 0                        season (Quast 1968a).
            San Diego Bay  ' |I     ' J
            Tijuana Estuary   I  Fecundity: Unknown.

                        A S J L E                          Growth and Development
                                                          Eaa Size and Embrvonic DeveloDment: Eggs are
    1974) Adultsarefoundfromthesurfzoneouttodepths    spherical and range from 0.94-0.97 mm in diameter
    of 183 m, but are most common between 2 and 21 m    (Butler et al. 1982). Embryonic development is indirect
    (Quast 1968a, Feder et al. 1974). Juveniles and adults    and external. At 19�C, eggs hatch in 36.0-40.5 hours.
    can be found throughout the water column depending
    on habitat complexity (Quast 1968a). This species is   Aae and Size of Larvae: Larval lengths range from 2.2-
    considered a kelp bed "cosmopolite", occurring    16.5 mm (Butler et al. 1982).  Yolk-sac absorption
    throughout the water column (Larson and DeMartini   takes about 5 days at 190C. At 21 0C, larvae transform
    1984).                                                 to juveniles 28 days after hatching (Butler et al. 1982).
                                                          Yolk-sac larvae of three Paralabrax species are
    Substrate: Eggs and larvae are notsubstratedependent.    indistinguishable (Butler et al. 1982).
    Juveniles are found among inshore seaweeds such as
    eelgrass (Zostera spp.), as well as in clumps of feather    Juvenile Size Ranae: Juveniles range in size from 1.6-
    boa kelp, in the kelp canopy, algae holdfast regions,    35.0 cm (Quast 1968a, Butler et al. 1982), and are
    and in rocky areas below kelp beds (Feder et al. 1974).    about 10 cm after 1 year.
    Adults also prefer areas containing habitat relief. This
    relief can be kelp beds or rocky bottoms, including    Aae and Size of Adults: Some may mature in 2 or 3
    submarine canyons and cliffs (Quast 1968a). Larger    years at 18 cm, with most males maturing at 25 cm, and

                                                       209

    Kelp bass continued
    females at 35 cm in length (Quast 1968a, Frey 1971,    1981 to 1984, but whether this was a result of reduced
    Feder et al. 1974). The kelp bass is a relatively slow-    population sizes, reduced fishing effort, or related to El
    growing fish; a 31 cm long fish may be 4-6 years old.   Nio is unclear(MBC Applied Environmental Sciences
    Maximum age may be 31 years, and maximum size is   1987).  Isolated populations do not appear to be
    reportedly 72 cm and 6.6 kg (Young 1963, Eschmeyer    genetically different (Beckwitt 1983).
    et al. 1983).
                                                           References
    Food and Feeding
    Trophic Mode: Larvae, juveniles, and adults are    Allen, L. G., M. H. Horn, F. A. Edmands II, and C. A.
    opportunistic, generalized carnivores.                  Usui. 1983. Structure and seasonal dynamics of the
                                                           fish assemblage in the Cabrillo Beach area of Los
    Food Items: Oncetheiryolk sac is used, larvae probably    Angeles Harbor, California. Bull. S. Calif. Acad. Sci.
    feed on small pelagic crustacea and other plankton.    82(2):47-70.
    Juveniles consume primarily invertebrates such as
    crabs (Pleuronocodes planipes and others), isopods,    Bane, G. W. 1968. Fishes of the upper Newport Bay.
    gammarid and caprellid amphipods, pistol shrimp    Univ. Calif. Irvine Res. Ser. 3:1-114.
    (Alphaeus spp.), caridean shrimps, euphausiids,
    mysids, polychaetes, coelenterates, but also small fish   Beckwitt, R. 1983. Genetic structure of Genyonemus
    and algae (Quast 1968a, Diaz and Hammann 1987).    lineatus, Seriphuspolitus (Sciaenidae) and Paralabrax
    Adults feed on similar organisms as juveniles, but shift   clathratus (Serranidae) in southern California. Copeia
    to eating primarily larger taxa such as pipefish    1983(3):691-696.
    (Syngnathus spp.), giant kelp fish (Heterostichus
    rostratus),topsmelt (Atherinopsaffinis), pleuronectids,    Butler, J. L., H. G. Moser, G. S. Hageman, and L. E.
    engraulids, embiotocids, cottids, serranids, gobiids,    Nordgren.  1982.  Developmental stages of three
    and cephalopods.  Fish and cephalopods (primarily   California sea bass (Paralabrax, Pisces, Serranidae).
    Octopusspp.) arethedominant prey of largeadultkelp   Calif. Coop. Ocean. Fish. Invest. Rep. 23:252-268.
    bass (Young 1963, Quast 1968a, 1968d, Feder et al.
    1974). The kelp bass appears to have two general    Carlisle, J. G., Jr. 1969. Results of a six-year trawl
    feeding peaks during the year, one in the spring and    study in an area of heavy waste discharge: Santa
    one in the fall (Quast 1968a). While normally a solitary    Monica Bay, California. Calif. Fish Game 55(1 ):26-46.
    feeder, it may assemble to feed on schooling bait fish,
    and even leap from the water if actively pursuing prey    Diaz, M. E. D, and M. G. Hammann. 1987. Trophic
    (Feder et al. 1974). The kelp bass typically feeds by    relations among fishes associated to a kelp forest,
    searching substrates and kelp stipes, and foraging into   Macrocystis pyrifera, in Baha de Todos Santos, Baja
    crevices.  It appears to prefer prey from the water    California, Mexico. Ciencias Mar. 13(4):81-96.
    column (Diaz and Hammann 1987) and only rarely
    forages near the surface (Quast 1968a).  It feeds    Ebling, A. W., and R. N. Bray. 1976. Day versus night
    primarily during the day and retreats into cover at night   activity of reef fishes in a kelp forest off Santa Barbara,
    (Hobson et al. 1981, Hobson and Chess 1986)             California. Fish. Bull., U.S. 74(4):703-717.

    Biological Interactions                                 Eschmeyer, W. N., W. S. Herald, and H. Hammann.
    Predation: The kelp bass is a cannibalistic species    1983. A field guide to Pacific coast fishes of North
    (Quast 1968a) that avoids predation by hiding at night    America. Houghton Mifflin Co., Boston, MA, 336 p.
    (Ebling and Bray 1976). Other predators of small kelp
    bass may include giant sea bass (Stereolepis gigas)    Fay, R. C., J. A. Vallee, and P. Brophy.  1978. An
    and broomtail grouper(Mycteropercaxenarcha) (MBC    analysis of fish catches obtained with an otter trawl in
    Applied Environmental Sciences 1987). Large kelp    Santa Monica Bay, 1969-73.  Calif. Fish Game
    bass probably have few predators other than man.        64(2):104-116.

    Factors Influencina Populations: This species may    Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    compete with the barred sand bass (P. nebulifer)    Observations on fishes associated with kelp beds in
    wherethey co-occur, howeverbarred sand bass prefer    southern California. Calif. Fish Game, Fish Bull. 160:1 -
    slightly different habitat (Turner et al. 1969). Because    144.
    of the kelp bass' slow growth and nonmigratorybehavior,
    intense sport fishing may have a detrimental effect on
    populations. Recreational landings decreased from

                                                        210

                                                                                             Kelp bass continued

    Frey, H. W. 1971. California's living marine resources    (editors), Utilization of kelp-bed resources in southern
    and their utilization.  Calif. Dept. Fish Game,    California. Calif. Fish Game, Fish Bull. 139:143-212.
    Sacramento, CA, 148 p.
                                                           Quast, J. C. 1968c. Some physical aspects of the
    Gates, D. E., and H. W. Frey.  1974.  Designated    inshore environment, particularly as it affects kelp-bed
    common  names of certain marine organisms of   fishes.  In W. J. North, and C. L. Hubbs (editors),
    California. Calif. Fish Game, Fish Bull. 161:55-90.      Utilization of kelp-bed resources in southern California.
                                                           Calif. Fish Game, Fish Bull. 139:25-34.
    Hobson, E. S., and J. R. Chess. 1981. Relationships
    among fishes and their prey in a nearshore sand    Quast, J. C. 1968d. Observations on the food of the
    community off southern California.  Env. Biol. Fish    kelp-bed fishes.  In W. J. North, and C. L. Hubbs
    17(3):201-226.                                           (editors), Utilization of kelp-bed resources in southern
                                                            California. Calif. Fish Game, Fish Bull. 139:109-142.
    Hobson, E. S., W. N. McFarland, and J. R. Chess.
    1981. Crepuscularand nocturnal activities of Californian    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    nearshore fishes, with consideration of their scotopic    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    visual pigments and the photic environment.  Fish.    of common and scientific names of fishes from the
    Bull., U.S. 79(1):1-30.                                  United States and Canada. Am. Fish. Soc. Spec. Publ.
                                                            No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Horn, M. H., and L. G. Allen. 1981. A review and
    synthesis of ichthyofaunal studies in the vicinity of Los    Stephens, J. S., Jr., and K. E. Zerba. 1981. Factors
    Angeles and Long Beach Harbors, LosAngelesCounty,    affecting fish diversity on a temperate reef. Env. Biol.
    California. Final Rep. to U.S. Fish Wildl. Serv., Dept.    Fish. 6(1):111-121.
    Biol. Sci., Calif. State Univ., Fullerton, CA, 96 p.
                                                            Turner, C. H., E. E. Ebert, and R. R. Given. 1969. Man-
    Larson, R. J., and E. E. DeMartini. 1984. Abundance    made reef ecology. Calif. Fish Game, Fish Bull. 146,
    and vertical distribution of fishes in a cobble-bottom    221 p.
    kelp forest off San Onofre, California. Fish. Bull., U.S.
    82(1):37-53.                                             U.S. Department of Commerce.   1986.  Marine
                                                            recreational fishery statistics survey, Pacific coast.
    MBC Applied Environmental Sciences. 1987. Ecology    U.S. Dept. Comm., NOAA,  Current Fish. Stat. No.
    of important fisheries species offshore California. Min.    8328, 109 p.
    Man. Serv., U.S. Dept. Int., Washington, D.C., 251 p.
                                                            Young, P. H. 1963. The kelp bass (Paralabrax
    Miller,D.J.,andR. N. Lea. 1972. Guidetothecoastal    clathratus) and its fishery,  1947-1958.  Calif. Fish
    marinefishesofCalifornia. Calif. FishGame, Fish Bull.   Game, Fish Bull. 122:1-67.
    157, 235 p.

    Noah, M.D. 1985. Appendix A. Structure, abundance
    and distribution of the fish and macroinvertebrate
    communities inhabiting Mission Bay, California between
    November 1979 and February 1981. In E. A. Weirich,
    M. D. Noah, and S. J. Schwarz (preparers), San Diego
    River and Mission Bay improvements, Draft suppl.
    environ. assess., 37 p. plus appendices. U.S. Army
    Corps Eng., Los Angeles, CA.

    Quast, J. C. 1968a. Observations on the food and
    biology of the kelp bass, Paralabrax clathratus with
    noteson its sportfisheryat San Diego, California. InW.
    J. North, and C. L. Hubbs (editors), Utilization of kelp-
    bed resources in southern California. Calif. Fish Game,
    Fish Bull. 139:81-108.

    Quast, J. C. 1968b. Effects of kelp harvesting on the
    fishes of the kelp beds. InW. J. North, and C. L. Hubbs

                                                        211

       Paralabrax nebulifer
       Adult

                        10cm

    Common Name: barred sand bass                         human populations (Valentine et al. 1973).
    Scientific Name: Paralabrax nebulifer
    Other Common Names: California rock bass, rock    Ecological: This is an important fish in California reef
    bass, Johnny verde, kelp bass, sand bass, ground    communities.  The greatest abundance of adults
    bass, sugar bass, cabrilla, California sand bass (Gates    appears to be near "edge" habitats where rocky and
    and Frey 1974)                                        sandy areas meet (Quast 1968).
    Classification (Robins et al. 1980)
    Phylum: Chordata                                      Range
    Class: Osteichthyes                                   Overall: The barred sand bass' overall range is from
    Order: Perciformes                                    Magdelana Bay, Baja California, to Santa Cruz,
    Family: Serranidae                                    California (including Guadalupe Island) (Miller and Lea
                                                           1972). It is not common north of Pt. Conception,
    Value                                                 California, but is occasionally taken in Monterey Bay,
    Commercial: Nocommercial fisheryexists inthe United   California (Roedel 1953).
    States for the barred sand bass, but this species is
    harvested in Mexico (Frey 1971).                      Within Studvy Area: This species is found in all bays and
                                                          estuaries from the Tijuana Estuary to Santa Monica
    Recreational: The barred sand bass is an important    Bay, California (Table 1) (Monaco et al. 1990).
    sport fish in southern California. It is highlysought after
    because of its good taste, fighting ability, availability,   Life Mode
    and relatively high abundance. It is often captured with    Eggs and larvae are pelagic, while juveniles and adults
    the kelp bass (Paralabrax clathratus) and regularly   are benthopelagic. Adults usually remain within a few
    seen byskin divers, snorkelers, and glass-bottom-boat    meters over the substrate. (Feder et al. 1974). This
    sightseers (Frey 1971).  It is usually caught by    species is more bottom-oriented than kelp bass.
    spearfishing and shore and pier fisherman using hook
    and line.  Over 1.7 million barred sand bass were    Habitat
    captured in 1985 (U.S. Department of Commerce    Type: The barred sand bass inhabits shallow neritic
    1986).                                                environments down to depths of 183 m (Miller and Lea
                                                           1972). Adults and subadults are most numerous
    Indicator of Environmental Stress: Industrial and    between depths of 5.2 and 26 m (Feder et al. 1974). It
    domestic wastes may be affecting barred sand bass    iscommonovernearshoresandyflats, nearkelp beds,
    habitat, but adverse effects have not been documented.    rocky areas, and bays (Squire and Smith 1977), and
    However,  a morphological anomaly (bilateral   can be the dominant fish on rocky reefs (Turner et al.
    asymmetry) has become more prevalent in fish from    1969). Small, immature sand bass prefer sheltered
    southern California populations. This condition may be    bays or harbors, especially around breakwaters.
    a result of sublethalpollution effects related to increasing   Juveniles are often found in mouths of bays in eelgrass

                                                       212

                                                                                      Barred sand bass continued

                                                            euhaline species. It may be more sensitive to cool
      Table 1. Relative abundance of barred sand            water temperatures than the kelp bass (Frey 1971).
               bass in 32 U.S. Pacific coast estuaries.
                         Life Stage                         Miarations and Movements: The barred sand bass
            Estuary    A S J L E                            moves to sandy flat bottoms to spawn, and then back
             Puget Sound  Relative abundance:               to rocky reefs (Turner et al. 1969). Like the kelp bass,
             Hood Canal                 Highly abundant     it appears to be nonmigratory (Turneret al. 1969). The
              Skagit Bay      :  i        Abundant          barred sand bass seeks cover in caves and holes if
            Grays Harbor              O   Common            frightened (Feder et al. 1974) and may feed actively at
             Wlllapa Bay                  Rare              night.
                 Columbia River      Blank Not present
           Columbia River
            Nehalem Bay                                     Reproduction
            Tillamook Bay             Life stage:           Mode: This species is gonochoristic, oviparous, and
             NetansBay                - Spultingadults      iteroparous.  It is a broadcast spawner; eggs are
              Silelz River            J - Juveniles         fertilized externally (Feder et al. 1974).
             Yaquina Bay             L - Larvae
                                     E-Eggs
             Alsea River                                    Matina/Soawnina: Spawning occurs from April to fall.
            Siuslaw River                                   This species forms spawning "schools" over sandy flat
            Umpqua River                                    bottoms (Frey 1971). The age, size, and frequency of
               Coos Bay                                     adult spawning is not documented.
             Rogue River
            Klamath River
                      Kamath Riv  Fecundity: Unknown
            Humboldt Bay
               Eel River
               Tamalaiver fis Bay  f off                    Growth and Development
       Cent San Fran. Bay�    Indcudes Ce:nral San          Eaa Size and Embryonic DeveloDment: Eggs are 0.94-
                        utS  Frn.  -  Francisc.Suisun,     0.97 mm in diameter and indistinguishable from kelp
       South San Fran. Bay           and San Pabo bays.     bass eggs (Butler et al. 1982). Embryonic development
              EMkho BaySlough                               is indirect and external. Eggs hatch in 36.0-40.5 h at
              Morro Bay19C.
         Santa Monica Bay      O    O  O                    19�C
           San PedroBay    C) O  X  0 0
            Alamitos Bay o    O                            Aae and Size of Larvae: Yolk-sac larvae are not
            Anaheim Bay 0    0                              distinguishable  from  P.  clathratus  or  P.
            NewportBay O    O                              maculotofasciatus (Butler et al. 1982). Larvae range
             MissionBay 3 a   0                             in length from 2.2-11.0 mm (Butler et al. 1982). Larval
           anDiego Bay O    O 01                            development is probably the same as P. clathratus-
          Tijuana Estuoary 0                               larval yolk-sac is absorbed in 5 days (at 19�C), and
                      A S J L E                            larvaltransformationoccurswhentheyare 11 mmlong
                                                           (Butler et al. 1982).

    (Zostera spp.) beds during fall and winter (Feder et al.   Juvenile Size Ranae: Minimum juvenile size is 12 mm.
    1974). It is the most common trawl-caught fish in
    Mission Bay (Noah 1985), and is also common in San    Aoe and Size of Adults: Age and size when mature is
    Diego Bay (Lockheed Ocean Science Laboratories    not known. This species reaches a maximum length of
    1983), and lower Newport Bay, California (Allen 1976).    65 cm (Miller and Lea 1972) and probably lives as long
    Bays and estuaries appear to play an important role in   as the kelp bass (31 years). A 20 year-old fish was 63
    this species early life history (Kramer and Hunter    cm (Turner et al. 1969).
    1987).
                                                           Food and Feeding
    Substrate: Preferred substrates range from sandy-    Trophic Mode: Larvae, juveniles, and adults are
    bottom flats to rocky areas and kelp beds. Spawning    carnivorous.
    occurs over flat sandy bottoms (Turner et al. 1969).
    Young juveniles are often found in and near eelgrass    Food Items: Larvae probably feed on small pelagic
    beds (Feder et al. 1974).                               crustaceans and other plankton once their yolk sac is
                                                           depleted. Small sand bass prefer a variety of
    Phvsical/Chemical Characteristics: No information is   crustaceans (shrimp, amphipods, crabs), molluscs
    available, but the barred sand bass is probably a    (octopus, squid), polychaetes, ophiuroids, and fish

                                                       213

    Barred sand bass continued
    (engraulidsandembiotocids)(Federetal. 1974). Crabs    Lockheed Ocean Science Laboratories.   1983.
    eaten are primarily spider and cancroid types (Quast    Distribution and abundance of fishes in central San
    1968). Large bass preferfish such as northern anchovy    Diego Bay, California: a study of fish habitat utilization.
    (Engraulis mordax) (Frey 1971) and other perciform    Rep. to Dept. of Navy, Contract No. N62474-82-C-
    fishes (Artedius spp., and Runula spp.) (Quast 1968).    1068, San Diego, CA, 38 p. plus appendices.

    Biological Interactions                                Miller, D. J., and R. N. Lea. 1972. Guidetothecoastal
    Predation: The barred sand bass is probably    marinefishesof California. Calif. FishGame, FishBull.
    cannibalistic and may have similar predators as kelp    157, 235 p.
    bass [e.g., giant sea bass (Stereolepis gigas) and
    broomtail grouper (Mycteroperca xenarcha)]. Large    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    barred sand bass probably have few predators except    Nelson. 1990. Distribution and abundance of fishes
    man.                                                    and invertebrates in west coast estuaries, Volume I:
                                                           data summaries. ELMR Rep. No.4. Strategic Assess-
    Factors Influencina PoDulations: Barred sand bass    ment Branch, NOS/NOAA, Rockville, MD,
    and kelp bass are often found in the same habitat, but    240 p.
    barred sand bass prefer sandy-rocky areas more than
    the kelp beds that the kelp bass prefers. As such, the    Noah, M. D. 1985. Appendix A. Structure, abundance
    barred sand bass is more abundant on manmade reefs    and distribution of the fish and macroinvertebrate
    (Turner et al. 1969). Large numbers of barred sand    communities inhabiting Mission Bay, California between
    bass have only been in southern California waters    November 1979 and February 1981. In E. A. Weirich,
    since 1957.  Before this period, sand bass were    M.D. Noah, and S. J. Schwarz (preparers), San Diego
    insignificant in the sport catch. Its higher abundance    River and Mission Bay improvements, Draft suppl.
    nowmayrelatetoincreasedcoastalwatertemperatures    environ. assess., 37 p. plus appendices, U.S. Army
    (Frey 1971).  Because of its slow growth and    Corps Eng., Los Angeles, CA.
    nonmigratory behavior, intense sport fishing may have
    a detrimental effect on the abundance of this species.    Quast, J. C. 1968. Observations on the food of the
                                                           kelp-bed fishes. In W. J. North, and C. L. Hubbs
    References                                              (editors), Utilization of kelp-bed resources in southern
                                                           California. Calif. Fish Game, Fish Bull. 139:109-142.
    Allen, L. G. 1976. Abundance, diversity, seasonality
    and community structure of the fish populations of    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Newport Bay, California. M.A. Thesis, Calif. State    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    Univ., Fullerton, CA, 107 p.                            of common and scientific names of fishes from the
                                                           United States and Canada. Am. Fish. Soc. Spec. Publ.
    Butler, J. L., H. G. Moser, G. S. Hageman, and L. E.    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Nordgren. 1982. Developmental stages of three
    California sea bass (Paralabrax, Pisces, Serranidae).    Roedel, P. M.  1953. Common ocean fishes of the
    Calif. Coop. Ocean. Fish. Invest. Rep. 23:252-268.      California Coast.  Calif. Fish Game, Fish Bull. 91,
                                                           184 p.
    Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    Observations on fishes associated with kelp beds in   Squire, J. L. Jr., and S. E. Smith. 1977. Anglers' guide
    southern California. Calif. Fish Game, Fish Bull. 160:1-    to the United States Pacific coast. Marine fish, fishing
    144.                                                    grounds and facilities.  NOAA, U.S. Dept. Comm.,
                                                           Seattle, WA, 139 p.
    Frey, H. W. 1971. California's living marine resources
    and their utilization.  Calif. Dept. Fish Game,    Turner, C. H., E. E. Ebert, and R. R. Given. 1969. Man-
    Sacramento, CA, 148 p.                                  made reef ecology. Calif. Fish Game, Fish Bull. 146,
                                                           221 p.
    Gates, D. E., and H. W. Frey. 1974. Designated
    common names of certain marine organisms of    U.S. Department of Commerce.   1986.  Marine
    California. Calif. Fish Game, Fish Bull. 161:55-90.     recreational fishery statistics survey, Pacific coast.
                                                           U.S. Dept. Comm., NOAA, Current Fish. Stat. No.
    Kramer, S. H., and J. R. Hunter.  1987.  Southern    8328, 109 p.
    California wetland/shallow water habitat investigation.
    Ann. Rep., Nat. Mar. Fish. Serv., La Jolla, CA, 12 p.

                                                       214

                                                                                       Barred sand bass continued
    Valentine, D. W., M. E. Soule, and P. Samollow. 1973.
    Asymmetry analysis in fishes: a possible statistical
    indicator of environmental stress. Fish. Bull., U.S.
    (2):357-370.

                                                        215

       Atractoscion nobilis
       Adult

                     25 cm

    Common Name: white seabass                             these waters since then (Vojkovich and Reed 1983).
    Scientific Name: Atractoscion nobilis
    Other Common Names: California white fish, sea    Recreational: In California, there is a limit of 3 fish per
    trout, weakfish, king croaker, whitecroaker(Gates and    day per person and fish must be >71 cm in length
    Frey 1974)                                              (Vojkovich and Reed 1983). The white seabass has
    Classification (Robins et al. 1980)                     been caught by hook and line (using live bait or lures)
    Phylum: Chordata                                       from piers, jetties, and private and party boats for the
    Class: Osteichthyes                                     past 100 years (Frey 1971, Vojkovich and Reed 1983).
    Order: Perciformes                                      Some are also taken by skindivers. This is a prized
    Family: Sciaenidae                                     sport fish because it is excellent eating, difficult to
                                                          capture, and may reach trophy size (Frey 1971). The
    Value                                                   sport catch peaked in 1949 (64,000 fish) and has
    Commercial:Thewhiteseabass iscommerciallyfished    declined since (Vojkovich and Reed 1983). Many of
    in California and Mexico (Frey 1971). The commercial    the white seabass hooked by sportsmen are below
    season in California is closed from March 14- May 16    legal size, but kept because fisherman cannot separate
    (during part of the spawning period). Legally, fish must    them from other croakers (or they are ignorant of the
    be at least 71 cm in length (Schultze 1986).  This    regulations) (Vojkovich and Reed 1983). Historically,
    species was historically caught by lampera, purse    coastal Native Americans used white seabass otoliths
    seine, hook and line, and drift and set gill nets (Frey    as jewelry (Fitch and Lavenberg 1971).
    1971). Now it is almost exclusively captured by set gill
    nets. Gill net mesh sizes must be 8.9 cm or larger   Indicator of Environmental Stress: Larvae and small
    (Schultze 1986). Set nets are typically set near rocky    juveniles appear to heavily utilize nearshore areas.
    headlands. From 1957 to 1961, much of the California    Therefore, human-caused environmental degradation
    catch occurred north of Point Conception, apparently    may be affecting recruitment (Vojkovich and Reed
    reflecting a period of warmer ocean temperatures.    1983). Juveniles may be easily affected by industrial
    After ocean temperatures returned to normal, catch    and domestic pollution (Fitch and Lavenberg 1971).
    levels dropped in this region, and have remained low    This pollution can cause hemorrhages of the eyes,
    (<1% of U.S. catch) (Vojkovich and Reed 1983).    blindness, and perhaps stimulate increased rates of
    Although landings have fluctuated widely, they have    parasitism by external parasites (Fitch 1958).
    dropped markedly since 1971 (Vojkovich and Reed
    1983). The five-year average from 1980 to 1984 was    Ecological: The white seabass is a major predator in
    159 t landed. However, in 1985, only 56 t of white    southern California nearshore waters. Fossil otoliths
    seabass were landed, but it was worth $241,000    have been found in California marine Pleistocene
    (National Marine Fisheries Service 1986).  Prior to   deposits that are 10-12 million years old (Fitch and
    1982, most of the U.S. catch (80%) was taken in   Lavenberg 1971).
    Mexican waters, but fishing has not been allowed in

                                                       216

                                                                                       White seabass continued

                                                           appears to have been once common in Newport Bay,
     Table 1. Relative abundance of white seabass         California (Skogsberg 1939).
               in 32 U.S. Pacific coast estuaries.
                         Life Stage                        Life Mode
           Estuary    A S J L E                           Eggs, larvae, juveniles and adults are all pelagic.
            Puget Sound              Relative abundance:   Juveniles may utilize the kelp canopy for cover (Feder
             Hood Canal               6   Highly abundant  et al. 1974). Adults may form loose schools (Fitch
              Skagit Bay 'J           1   Abundant         1958, Feder et al. 1974).
            Grays Harbor              O   Common
             Willapa Bay              i   Rare             Habitat
          Columbia River             Blank Not present     ype: Newly-metamorphosed white seabass occur in
            Nehalem Bay                                    open coastalwaters justoutsidethebreakerline (Kramer
           Tillamook Bay             Life stage:          and Hunter 1988). This habitat is often less than 8 m
             Netanrs Bay             A -   adults          deep. Juveniles and adults occur from the surface to
             Siletz River            J - Juveniles         depths of 122 m, with adults primarily found from 3-46
            YaquinaBay i             L-Larvae             m (Fitch and Lavenberg 1971). Very small fish are
             Alsea River               Egg                 found in bays and shallow nearshore waters near the
            Siuslaw River                                  surf zone, mid-sized fish are found in the mainland kelp
           Umpqua River                                    beds close to shore, and larger fish are often caught
              Coos Bay  /                                 near rocky headlands and offshore islands (Frey 1971).
            Rogue River
           Klamath River                                  Substrate: It is most often found over sandy bottoms or
           Humboldt Bay  i    '1                          along the edges of kelp beds (Squire and Smith 1977).
               Eel River                                   Schools can be found over rocky bottoms and among
            Tomales Bay  J   '1                            giant kelp just below the canopy (Feder et al. 1974).
       Cent San Fran. Bay'   /1    *    Includes Central San
                                     Francisco, Suisun,
       South San Fran. Bay  V  and San Pabo bays.         Physical/Chemical Characteristics: White seabass
           ElkhomrnSlough                                  occur in waters with salinities of 32-34%o and
              Morro Bay                                    temperatures of 13-30�C (Vojkovich and Reed 1983).
        Santa Monica Bay  C  0 0    0                      Larvae have been successfully reared attemperatures
           SanPedroBay 000 0    0                          of 18.7-21.70C (Moser et al. 1983).
            Alamitos Bay
            Anaheim Bay    -4                              Miarations and Movements: Some data indicate that
            Newport Bay                                    they migrate north in the spring and southward in the
             Mission Bay                                   fall,wintering off Baja California. This migration appears
           San Diego Bay  n1                               to correlate with spawning (Frey 1971). This species
          Tijuana Estuary   j                              may feed more actively at night than day (Skogsberg
                      A S J L E                            1939, Fitch and Lavenberg 1971)

    Range                                                  Reproduction
    Overall: This species has been recorded in coastal    Mode: The white seabass is gonochoristic, oviparous,
    waters from Magdalena Bay, Baja Californiato Juneau,    and iteroparous; eggs are fertilized externally.
    Alaska. There is also an isolated population occurring
    in the northern section of the Gulf of California (Frey    Matino/Soawnina: Spawning occurs from March to
    1971, Miller and Lea 1972).  It is most abundant    August, peaking from April to June (Thomas 1968,
    between Point Conception and Ballenas Bay, Baja    Frey 1971, Vojkovich and Reed 1983).  During the
    California (Frey 1971), but this range shifts with water    spawning period, spawners appear to congregate
    temperature fluctuations (Skogsberg 1939, Thomas    nearshore in certain areas (e.g., Long Point and Palos
    1968, Frey 1971).                                      Verdes Peninsula, California ), but specific spawning
                                                           sites have not been identified (Thomas 1968, Frey
    Within Study Area: Although it is possible to find white    1971).  Successful spawning probably occurs from
    seabass throughout the study area, it is very rare north    Santa Rosa Island, California to Santa Maria Bay, Baja
    of Point Conception. This species is common in San    California (based on larval distributions) (Moser et al.
    Pedro and Santa Monica Bays, but rare in othersouthern   1983).
    California bays and estuaries (Table 1) (Horn 1974,
    Horn and Allen 1981, Allen et al. 1983). However, it   Fecundity: Unknown.

                                                        217

    White seabass continued
    Growth and Development                                fishery by rearing juveniles in hatcheries and then
    Eaa Size and Embryonic DeveloDment: Eggs are   releasing them into the ocean (Crooke and Taucher
    spherical and 1.24-1.32 mm in diameter (Moser et al.   1988).
    1983). Embryonic development is indirect and external.
    Eggs hatch in about 3 days at temperatures of 16.5-   References
    20.00C (Moser et al. 1983).
                                                          Allen, L. G., M. H. Horn, F. A. Edmands II, and C. A.
    Ace and Size of Larvae: Larvae are 2.8-15.5 mm in   Usui. 1983. Structure and seasonal dynamics of the
    length (Moser et al. 1983). Metamorphosis to juvenile   fish assemblage in the Cabrillo Beach area of Los
    begins at about 33.0 mm standard length (SL), and 72    Angeles Harbor, California. Bull. S. Calif. Acad. Sci.
    days after hatching (Moser et al. 1983).              82(2):47-70.

    Juvenile Size Ranoe: Juveniles range in length from    Crooke, S., and C. Taucher. 1988. Ocean hatcheries
    33.0 mm SL to probably 50 cm SL for males and 60 cm    - wave of the future? Outdoor Calif. 49(3):10-13.
    SL for females (Frey 1971, Moser et al. 1983).
                                                           Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    Aae and Size of Adults: Some males mature at about    Observations on fishes associated with kelp beds in
    51 cm total length, and some females at 61 cm long   southern California. Calif. Fish Game, Fish Bull. 160:1-
    (one year later) (Frey 1971 ). However, all white seabass    144.
    are mature at 80 cm (Vojkovich and Reed 1983). Many
    females mature at age three, and most all are mature    Fitch, J. E. 1958. Offshore fishes of California. Calif.
    by age four (Fitch and Lavenberg 1971). This is the    Fish Game, Sacramento, CA, 80 p.
    largest member of the Sciaenidae family in California
    and may reach sizes over 1.2 m and 36 kg (individuals   Fitch, J. E., and R. J. Lavenberg. 1971. Marine food
    weighing over 27 kg are rare).  The largest white    and game fishes of California.  Calif. Nat. History
    seabass reportedwas 1.7 mandweighed38kg(Squire    Guides 28, Univ. Calif. Press, Berkeley, CA, 179 p.
    and Smith 1977). Most commercially-caught fish are 9-
    18 kg (Frey 1971). Scale analyses indicate that these    Frey, H. W. 1971. California's living marine resources
    are 3-20 year-old fish, but many may actually be older   and their utilization.  Calif. Dept. Fish Game,
    (Frey 1971). The 18 kg fish are often 20 years old or   Sacramento, CA, 148 p.
    older (Fitch and Lavenberg 1971).
                                                           Gates, D. E., and H. W. Frey. 1974. Designated
    Food and Feeding                                      common names of certain marine organisms of
    Trophic Mode: Larvae, juveniles, and adults are   California. Calif. Fish Game, Fish Bull. 161:55-90.
    carnivorous.
                                                           Horn, M. H. 1974. Fishes. InAsummaryofknowledge
    Food Items: Larvae feed on planktonic crustaceans    of the southern California coastal zone and offshore
    and other plankton (Moser et al. 1983). Juveniles eat   areas, Chapter 11. S. Calif. Ocean Stud. Consort.,
    fish, such as northern anchovy (Engraulis mordax),    Fullerton, CA, 124 p.
    Pacific sardine (Sardinops sagax), chub mackerel
    (Scomberjaponicus), and squid (Loligo opalescens),    Horn, M. H., and L. G. Allen. 1981. A review and
    and pelagic red crabs (Pleuroncodes planipes) when    synthesis of ichthyofaunal studies in the vicinity of Los
    available (Thomas 1968, Fitch 1958).                  Angeles and Long Beach Harbors, Los Angeles County,
                                                           California. Final Rep. to U.S. Fish Wildl. Serv., Dept.
    Biological Interactions                               Biol. Sci., Calif. State Univ., Fullerton, CA, 96 p.
    Predation: Eggs, larvae, and juveniles are probably
    eaten by many predators. Adults probably have few    Kramer, S. H., and J. R. Hunter.  1988.  Southern
    predators except man, but marine mammals and sharks   California wetland/shallow water habitat investigation.
    will feed on gill-netted fish (Fitch and Lavenberg 1971).    Ann. Rep., Nat. Mar. Fish. Serv., La Jolla, CA, 15p.

    Factors Influencina PoDulations: Historically, this   Miller,D.J.,and R. N. Lea. 1972. Guidetothecoastal
    species' population size has fluctuated widely.    marinefishesofCalifornia. Calif. FishGame, FishBull.
    Oceanographic conditions and changes in forage    157, 235 p.
    species may affect its distribution (Skogsberg 1939,
    Vojkovich and Reed 1983).  In southern California,    Moser, H. G., D. A. Ambrose, M. S. Busby, J. L. Butler,
    attemptsarebeingmadetoenhancethewhiteseabass    E. M. Sandknop, B. Y. Sumida, and E. G. Stevens.

                                                       218

                                                                                          White seabass continued
    1983. Description of early stages of white seabass,
    Atractoscion nobilis, with notes on distribution. Calif.
    Coop. Ocean. Fish. Invest. Rep. 24:182-193.

    National Marine Fisheries Service. 1986. Fisheries of
    the United States, 1985. Current Fishery Statistics No.
    8368. U.S. Dept. Comm., Nat. Ocean. Atm. Adm., Nat.
    Mar. Fish Serv., Nat. Fish. Stat. Prog., Washington,
    D.C., 122 p.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Schultze, D. L. 1986. Digest of California commercial
    fish laws, January 1, 1986. Calif. Dept. Fish Game,
    Sacramento, CA, 40 p.

    Skogsberg, T. 1939. The fishes of the family Sciaenidae
    (croakers) of California. Calif. Fish Game, Fish Bull.
    54:1-62.

    Squire, J. L., Jr., and S. E. Smith. 1977. Anglers' guide
    tothe United States Pacific coast. NOAA, Seattle, WA,
    139 p.

    Thomas, J. C. 1968. Management of the white
    seabass (Cynoscion nobilis) in California waters. Calif.
    Fish Game, Fish. Bull. 142:1-34.

    Vojkovich, M.,and R. J. Reed. 1983. White seabass,
    Atractoscion nobilis, in California-Mexican waters:
    status of the fishery. Calif. Coop. Ocean. Fish. Invest.
    Rep. 24:79-83.

                                                         219

       Genyonemus lineatus
       Adult

                       5cm

    Common Name: white croaker                            so easily caught in some localities that it is consider a
    Scientific Name: Genyonemus lineatus                  nuisance (Baxter 1960). This species can be caught
    Other Common Names: California white seabass,    year-round and is especially popular with some ethnic
    seatrout, weakfish, king croaker, whitecroaker, kingfish,   groups. Over 249,000 white croakers were caught by
    tomcod, tommy, roncky (Roedel 1953, Frey 1971,    anglers in 1985 (U.S. Department of Commerce 1986).
    Gates and Frey 1974, Squire and Smith 1977)           Most white croaker kept by anglers are 21-25 cm total
    Classification (Robins et al. 1980)                   length (TL) and 5-7 years old (Love et al. 1984).
    Phylum: Chordata
    Class: Osteichthyes                                   Indicatorof Environmental Stress: High concentrations
    Order: Perciformes                                    of polychlorinated biphenyls (PCBs) and other
    Family: Sciaenidae                                    contaminants in the tissues of white croaker pose a
                                                           potential health threat to humans, resulting in the
    Value                                                 closure of some fishing areas (Puffer et al. 1982).
    Commercial: The white croaker is sold in fresh-fish   White croakers found near southern California sewer
    markets, however, it is not a prime market fish because    outfalls are often malformed and diseased. Diseases
    of its soft flesh (Bane and Bane 1971, Eschmeyer et al.   include cancerous growths on lips (neoplasia), bulging
    1983). It is also caught and sold for bait (Hart 1973). In   and missing eyes, warped bodies, and high parasitism
    the southern California Bight, it is now primarily caught    rates. These conditions are probably a result of toxic
    by bottom set gill nets (7.0 cm stretch), but was once    effluents (Baxter 1960, Frey 1971, Phillips et al. 1972).
    caught by ottertrawl, round haul net, and hook and line   Since the white croaker accumulates contaminants
    (Love et al. 1984). Over 453 t were landed in 1952,    (Castle and Woods 1972) it is a good indicator species
    1953,1960, and 1965 (Baxter1960, Frey 1971). About    for pollution and is a target species of the National
    200 t/year are now landed, with the largest catches    Status and Trends Program (Ocean Assessments
    occurring in January and February (spawning season)    Division 1984).
    (Love et al. 1984). In 1982, fishermen received 13-18�/
    kg fortheir catch. Vietnamese fishermen have recently   Ecological: This is an abundant (often dominant) species
    started fishing for this species in Monterey Bay,    in nearshore shallow waters with sandy substratum in
    California, receiving 33-11 0�/kg fortheircatch (Love et   southern California, both within bays and estuaries,
    al. 1984).                                            and just outside the surf zone (Roedel 1953, Squire
                                                            and Smith 1977, Love et al. 1984). White croaker
    Recreational: The white croaker is an important sport   larvae are often second in abundance only to northern
    fish in California. Although small (and wronglythought    anchovy (Engraulis mordax) in the southern California
    of as wormy), it is a good food fish (Skogsberg 1939,    ichthyoplankton (Love et al. 1984), and this species
    Squire and Smith 1977, Love et al. 1984). It is commonly    often occurs with queenfish (Seriphuspolitus) (Roedel
    caught from piers and boats with hook and line using    1953). Fossil otoliths have been found in Pliocene
    various baits and lures (Eschmeyer et al. 1983). It is   deposits 12 to 20 million years old (Baxter 1960).

                                                        220

                                                                                       White croaker continued

                                                           al. 1984). Juveniles and adults are primarily epibenthic
      Table 1. Relative abundance of white croaker         schooling fishes (Eschmeyer et al. 1983, Wang 1986),
               in 32 U.S. Pacific coast estuaries.         but they may occur in midwater or at times near the
                        Life Stage                         surface (Skogsberg 1939, Love et al. 1984).
           Estuary    A S J L E
            Puget Sound  ;   -       Relative abundance:   Habitat
            Hood Canal :                  Highly abundant  Type: The white croaker is neritic and normally found
             Skagit Bay              i)  Abundant          inshore in waters less than 30 m deep, but it occurs to
           Grays Harbor              O   Common            depths of 183 m (Eschmeyer et al. 1983, Love et al.
            Willapa Bay             Bl Rare                1984).  It is common in bays and estuaries (Wang
          Columbia River             Blank Notpresent      1986). Juveniles occur in waters <27 m deep; large
           Nehalem Bay                                     croakers inhabit greaterdepths (Love et al. 1984). The
           Tillamook Bay             Life stage:           highest larval densities in southern California are found
            Netarts Bay             A     ning dults      in a narrow band along the coast at depths between 15
             Silelz River            J-Juveniles           and 22 m (Watson 1982, Love et al. 1984) and within
            Yaquina Bay              L- LEarvae            5 km of shore (Barnett et al. 1984). Juveniles occur
            Alsea River   :                               primarily in a narrow coastal band between the 18 and
           Siuslaw River                                   27 m isobaths (Love et al. 1984).
           Umpqua River
              Coos Bay                                     Substrate: Eggs and larvae are found over sand and
            Rogue River                                    gravel bottoms (Wang 1986). Adults and juveniles are
           Klamath River                                   found mostly oversandy bottoms, but mayoccasionally
           Humboldt Bay 0      :                           be found in kelp beds (Roedel 1953, Love et al. 1984).
              Eel River
           Tomales Bay 0 0  O 0                           Phvsical/Chemical Characteristics: The white croaker
      Cent.San Fran. Bay* O O O O O 0   Indudes Central San  iS found in euhaline to mesohaline waters (Wang
                                     Francisco, Suisun,
      South San Fran. Bay  O0 0 O O  O  O and Sa Pablo bays.  1986). The optimal temperature range for metabolism
          Elkhom Slough      (                             is broad (11 -17�C), and may account for this species'
             Morro Bay                                     wide depth and latitudinal distributions (Love et al.
        Santa Monica Bay   �  �                            1984).
          San Pedro Bay  g g
           Alamitos Bay  _ I   o                          Minrations and Movements: Adults appear to move
           Anaheim Bay O    O O                            shorewardto spawn in shallow waters. Eggs and early
           Newport Bay  D   O OC                          larvae apparently remain within this shallow "band".
            Mission Bay      ,   O0                        Larvae appear to drift into bays and estuaries on
          SanDiegoBay  O    O 4                            incoming tides (Wang 1986) and migrate tothe bottom
         Tijuana Estuary    4  / O                        after hatching (Schlotterbeck and Connally 1982, Jahn
                      A S J L E                            et al. 1988). Early juveniles initially reside in waters 3-
                                                           6 m deep, but move to deeper waters as they grow
    Range                                                  (Love et al. 1984).
    Overall: The white croaker's overall range is from
    Magdalena Bay, Baja California, to Vancouver Island,    Reproduction
    British Columbia (Miller and Lea 1972, Hart 1973,    Mode: The white croaker is gonochoristic, oviparous,
    Eschmeyer et al. 1983). It is generally not abundant    and iteroparous. It is a broadcast spawner; eggs are
    north of San Francisco Bay, and is rare north of   fertilized externally.
    California (Frey 1971).
                                                           Matina/SDawnina: Spawning occurs in shallow
    Within Studv Area: This species is found in almost all   nearshore waters essentially year-round in California,
    bays and estuaries south of Humboldt Bay, California,   with specific spawning times dependent on location
    but is extremely rare north of Humboldt Bay (Table 1)   (Skogsberg 1939, Bane and Bane 1971, Hart 1973,
    (Reish 1968, Bane and Bane 1971, Allen 1976, Horn    Goldberg 1976, Eldridge 1977, Love et al. 1984). It
    and Allen 1981, Allen et al. 1983).                    spawns primarily from November to April in southern
                                                           California, often peaking during February and March
    Life Mode                                              (Goldberg 1976, Schlotterbeck and Connally 1982,
    Eggs are pelagic, and larvae are benthopelagic to    Love et al. 1984). It is also known to spawn in San
    epibenthic (Schlotterbeck and Connally 1982, Love et   Francisco Bay, Tomales Bay, and Elkhorn Slough,

                                                       221

    White croaker continued
    California, and coastal waters of northern Mexico (Love    polychaetes, cumaceans, chaetognaths, cyprids,
    et al. 1984, Wang 1986). The white croaker may have    copepods, and fish larvae (Phillips et al. 1972). Larger
    a protracted spawning season off Monterey, California,   juveniles and adults switch from zooplankton to benthic
    because of cooler water temperatures there (Love et   and epibenthic organisms, consuming a wide variety of
    al. 1984). During spawning, watertemperatures range   fish [northern anchovy (Engraulis mordax) and others],
    from 8.0-19.0�C, with surface waters of 13-140C at   squid, shrimp, octopus, polychaetes, crabs, clams,
    peak spawning (Love et al. 1984, Wang 1986). A batch    and other living and dead organisms (Skogsberg 1939,
    spawner, the white croaker spawns 18-24 times per   Baxter 1960, Allen 1982).
    season, with large females spawning earlierand longer
    than small individuals (Love et al. 1984). This species    Biological Interactions
    appears to utilize two spawning centers from south of   Predation: The white croaker is eaten by sea lions,
    Point Conception to the Mexican border: one center    Pacific bottlenose dolphin (Tursiops truncatus),
    north and south of the Palos Verdes Peninsula (from   California halibut (Paralichthys californicus), black sea
    Redondo Beach to Laguna Beach), and a smaller   bass (Stereolepis gigas), bluefin tuna (Thunnus
    center around Ventura (Love et al. 1984).             thynnus), and probably otherpiscivorous animals (Fitch
                                                           1958, Baxter 1960)
    Fecundity: Batch fecundity is estimated to be 800 to
    37,200 eggs per female (Love et al. 1984).            Factors Influencina PoDulations: High levels of
                                                           contaminants apparently can impair reproduction (Cross
    Growth and Development                                and Hose 1988). Concentrations of PCBs and DDT in
    Eaa Size and Embrvonic DeveloDment: Eggs are 0.5-   this species are directly related to its reproductive state
    0.9 mm in diameter, averaging 0.85 mm (Watson    (Cross 1986). Pollutants may cause tail rot and liver
    1982). Embryonicdevelopmentisindirectandextemal.    damage (Phillips et al. 1972).  Because the white
    In one study, all eggs hatched in 52 hr at 20�C (Watson    croaker utilizes nearshore coastal habitats for spawning
    1982).                                                and rearing, it is directly affected by man's activities in
                                                           these areas.
    Aae and Size of Larvae: Larvae range from 1.8-2.8 mm
    standard length at hatching (Watson 1982, Wang 1986).    References

    Juvenile Size Ranae: Juveniles are 1.3 to about 13 cm    Allen, L. G. 1976. Abundance, diversity, seasonality
    total length (TL) (Love et al. 1984).                 and community structure of the fish populations of
                                                           Newport Bay, California. M.A. Thesis, Calif. State
    Aae and Size of Adults: Maturity is reached in 1 to 4    Univ., Fullerton, CA, 107 p.
    years, with about 50% maturing in 1 year; all are mature
    at 19 cmTL (Love et al. 1984). Males appearto mature    Allen, M. J. 1982. Functional structure of soft-bottom
    at about 12 cm and females at 13 cm TL (Love et al.   fishcommunitiesofthesouthernCaliforniashelf. Ph.D.
    1984). Females growfasterthan males, and bothgrow    Diss., Univ. Calif., San Diego, CA, 577 p.
    at fairly constant rates throughout their lives (Love et al.
    1984). The largest specimen recorded was 39 cm and    Allen, L. G., M. H. Horn, F. A. Edmands II, and C. A.
    0.7kg (Squireand Smith 1977). Whitecroakermay live   Usui. 1983. Structure and seasonal dynamics of the
    for 12 to 15 years (Love et al. 1984).                fish assemblage in the Cabrillo Beach area of Los
                                                           Angeles Harbor, California. Bull. S. Calif. Acad. Sci.
    Food and Feeding                                      82(2):47-70.
    Trophic Mode: Larvae, juveniles, and adults are
    omnivorous bottom feeders, feeding primarily at night.    Bane, G. W., and A. W. Bane. 1971. Bay fishes of
    However, juveniles may feed in midwater during the   northern California with emphasis on the Bodega
    day (Allen 1982).                                     Tomales Bay area. Mariscos Publ., Hampton Bays,
                                                           NY, 143 p.
    Food Items: Larvae eat rotifers, tintinnids,
    dinoflagellates, polychaete larvae, lamellibranch larvae,   Barnett, A. M., A. E. Jahn, P. D. Sertic, and W. Watson.
    copepods, amphipods, and invertebrate eggs. Very    1984. Distribution of ichthyoplankton off San Onofre,
    small larvae eat primarily rotifers, while larger larvae   California, and methods for sampling very shallow
    prey on copepods (Jahn et al. 1988). Small juveniles   coastal waters. Fish. Bull., U.S. 82(1):97-111.
    (<87 mm TL) eat mainly zooplankton, including
    cladocerans, amphipods, ostracods, mysids,    Baxter, J. L. 1960. Inshore fishes of California. Calif.
    euphausiids, crab zoea and megalopae, larval    Dept. Fish Game, Sacramento, CA, 80 p.

                                                       222

                                                                                        White croaker continued

    Castle, W. T., and L. A. Woods, Jr.  1972.  DDT    82(1):179-198.
    residues in white croakers. Calif. Fish Game 58:(3):198-
    203.                                                     Miller, D. J., and R. N. Lea. 1972. Guide to the coastal
                                                           marinefishes of California. Calif. Fish Game, Fish Bull.
    Cross, J. N. 1986. Seasonal changes in DDT and PCB    157, 235 p.
    concentrations in white croaker are related to the
    reproductive cycle. Coastal Water Res. News 1(2):2.    Ocean Assessments Division.  1984.  The national
                                                           status and trends program for marine environmental
    Cross, J. N., and J. E. Hose.  1988.  Evidence for   quality: program description (mimeo). Ocean Assess.
    impaired reproduction in white croaker (Genyonemus    Div., Nat. Ocean Serv., Nat. Ocean. Atm. Adm.,
    lineatus) from contaminated areas of southern    Rockville, MD, 28 p.
    California. Mar. Env. Res. 24(1-4):185-188.
                                                           Phillips, L., C. Terry, and J. Stephens. 1972. Status
    Eldridge, M. B. 1977. Factors influencing distribution    of the white croaker (Genyonemus lineatus) in the San
    of fish eggs and larvae over eight 24-hr samplings in    Pedro Region. Rep. to Southern Calif. Coast. Water
    Richardson Bay, California. Calif. Fish Game 63(2) :101 -    Res. Proj., Longbeach, CA, 47 p.
    116.
                                                           Puffer, H. W., M. J. Duda, and S. P. Azen. 1982.
    Eschmeyer, W. N., E. S. Herald, and H. Hammann.    Potential health hazards from consumption of fish
    1983. A field guide to Pacific coast fishes of North    caughtinpollutedcoastalwatersofLosAngelesCounty.
    America. Houghton Mifflin Company, Boston, MA,           N. Am. J. Fish. Man. 2:74-79.
    336 p.
                                                           Reish, D. J. 1968. Marine life of Alamitos Bay. Forty-
    Fitch, J. E. 1958. Offshore fishes of California. Calif.    Niner Shops, Inc., Long Beach, CA, 92 p.
    Dept. Fish Game, Sacramento, CA, 80 p.
                                                           Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Frey, H. W. 1971. California's living marine resources    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    and their utilization.  Calif. Dept. Fish Game,    of common and scientific names of fishes from the
    Sacramento, CA, 148 p.                                   United States and Canada. Am. Fish. Soc. Spec. Publ.
                                                           No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Gates, D. E., and H. W. Frey. 1974. Designated
    common  names of certain marine organisms of    Roedel, P. M.  1953. Common ocean fishes of the
    California. Calif. Fish Game, Fish Bull. 161:55-90.      California coast. Calif. Fish Game, Fish Bull. 91,
                                                           184 p.
    Goldberg, S. R. 1976. Seasonal spawning cycles of
    the Sciaenidfishes Genyonemuslineatusand Seriphus    Schlotterbeck, R. E., and D. W. Connally.  1982.
    politus. Fish. Bull., U.S. 74(4):983-984.                Vertical stratification of three nearshore southern
                                                           California larvalfishes (Engraulis mordax, Genyonemus
    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    lineatus, and Seriphus politus).  Fish. Bull., U.S.
    Board Can., Bull. No. 180, 740 p.                        80(4):895-902.

    Horn, M. H., and L. G. Allen.  1981.  A review and    Skogsberg,T. 1939. ThefishesofthefamilySciaenidae
    synthesis of ichthyofaunal studies in the vicinity of Los    (croakers) of California. Calif. Fish Game, Fish Bull.
    Angeles and Long Beach Harbors, Los Angeles County,    54, 62 p.
    California. Final Rep. to U.S. Fish Wildl. Serv., Dept.
    Biol. Sci., Calif. State Univ., Fullerton, CA, 96 p.     Squire, J. L., Jr., and S. E. Smith. 1977. Anglers' guide
                                                           to the United States Pacific Coast. NOAA, Seattle,
    Jahn, A. E., D. M. Gadomski, and M. L. Sowby. 1988.    WA, 139 p.
    On the role of food-seeking in the suprabenthic habit of
    larval white croaker, Genyonemus lineatus (Pisces:    U.S. Department of Commerce.   1986.  Marine
    Sciaenidae). Fish. Bull., U.S. 86(2):251-262.             recreational fishery statistics survey, Pacific coast.
                                                           U.S. Dept. Comm., Nat. Ocean. Atm. Adm., Current
    Love, M. S., G. E. McGowen, W. Westphal, R. J.   Fish. Stat. No. 8328, 109 p.
    Lavenberg, and L. Martin. 1984. Aspects of the life
    history and fishery of the white croaker, Genyonemus    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    lineatus (Sciaenidae), off California. Fish. Bull., U.S.    Joaquin estuary and adjacent waters, California: a

                                                        223

    White croaker continued
    guide to the early life histories. Tech. Rep. No. 9.
    Interagency Ecological Study Program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res, Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    and U.S. Fish Wildl. Serv., various pagination.

    Watson,W. 1982. Development ofeggsandlarvaeof
    the white croaker, Genyonemus lineatus Ayres (Pisces:
    Sciaenidae), off the southern California coast. Fish.
    Bull., U.S. 80(3):403-416.

                                                       224

    225

      Cymatogaster aggregata
      Adult

                   2cm

    Common Name: shiner perch                            et al. 1979, Wydoski and Whitney 1979).
    Scientific Name: Cymatogasteraggregata
    Other Common Names: shiner seaperch, shiner   Range
    surfperch, yellow shiner, shiner, bayperch, poggie,   Overall: Overall range is from Todos Santos Bay, Baja
    sparada, minny, bayperch, seven-eleven perch (Roedel   California, to Port Wrangell, Alaska (Roedel 1953,
    1953, Gates and Frey 1974, Washington 1977,    Bane and Bane 1971). The shiner perch is scarce at
    Eschmeyer et al. 1983)                               the northern and southern ends of its range, but
    Classification (Robins et al. 1980)                   abundant from San Diego, California, to Ketchikan,
    Phylum: Chordata                                     Alaska (Morrow 1980).
    Class: Osteichthyes
    Order: Perciformes                                   Within Studvy Area:This species is common to abundant
    Family: Embiotocidae                                 in all Pacific coast estuaries and bays from San Diego
                                                          Bay, California, through Puget Sound, Washington
    Value                                                 (Table 1) (Horn 1974, Morrow 1980, Proctor et al.
    Commercial: The shiner perch is not commercially    1980).
    important, although some are landed for use as bait
    (Frey 1971) and human consumption (Roedel 1953).   Life Mode
    This species is considered to be a delicacy by some    The shiner perch is a live-bearer; eggs are retained
    (Washington 1977, Wydoski and Whitney 1979).         within the female and juveniles are born fully developed.
                                                          Juveniles and adults are primarily neritic and pelagic
    Recreational:The shinerperch is commonlycaught by    (Garrison and Miller 1982).
    children fishing with small hooks in estuaries and bays
    (Baxter 1960, Eschmeyer et al. 1983). It is occasionally   Habitat
    used for bait in California's San Francisco Bay striped   Type: This species occurs primarily in nearshore
    bass fishery (Smith and Kato 1979).                   shallow-water marine, bay, and estuarine habitats,
                                                          both intertidally and subtidally. It is commonly
    Indicator of Environmental Stress: The shiner perch   associated with aquatic vegetation (eelgrass, Zostera
    has been used to assess the toxicity of some common    spp.) and docks and pilings (Bane 1968).  During
    organochlorine insecticides (Earnest and Benville   spring and summer, juveniles prefer intertidal and
    1972). Becausethisspecies utilizes nearshorepolluted   shallow-water subtidal habitats in bays and estuaries
    environments, it may have body burden pesticide levels    (Shaw et al. 1974, Moyle 1976). In winter, they occur
    higher than other fishes (Earnest and Benville 1971).   primarily in neritic marine habitats, occasionally as
                                                          deep as 70 m (Hart 1973, Wydoski and Whitney 1979).
    Ecological: The shiner perch is a small yet abundant
    species in many estuaries and bays. It is preyed upon    Substrate: The shiner perch prefers sandy and muddy
    by numerous birds, mammals, and fishes (Simenstad    bottoms (Bane and Bane 1971), but may be found over

                                                      226

                                                                                        Shiner perch continued
                                                          1975). It is not normally found at depths >30 m in
     Table 1. Relative abundance of shiner perch         California (Bane 1968), but is commonly captured at
              in 32 U.S. Pacific coast estuaries.        depths between 18 and 73 m in Puget Sound in winter
                       Life Stage                         (Wydoski and Whitney 1979). This species has been
             Estuary    A P J                             taken as deep as 128 m (Clemens and Wilby 1961).
                   Puget Sound            Relative abundance:
              Hood Canal �0       9 �   Highly abundant  Miarations and Movements: The shiner perch forms
               Skagit Bay *� i �   3   Abundant           loose schools that move seasonally- onshore and into
             Grays Harbor (3 %    �    O    Common        shallow water marine areas, estuaries, and bays in the
              Willapa Bay  3 3 �    Rare                 spring, and offshore into deeper marine waters in the
            Columbia River 1 (X    *  Blank Not present   fall and winter (Bane and Robinson 1970, Stober et al.
             Nehalem Bay      13 �*                       1973, Wydoski and Whitney 1979). No coastal (north-
             Tillamook Bay  O O �    Life stage:          south) migrations are known to occur.  During the
              NetartnsBay * � �    A-Adults               prespawning period, adults stay in shallow waters
               SiletzRiver �   � �      JaJuveniles       during daylight and move to deeper waters at night.
              Yaquina Bay � � �                           Afterthis period, most adults reversethis movement by
               Alsea River � �     �                      schooling in deeper water during the day and moving
             Siuslaw River �   *            �             to shallow water at night (Gordon 1965 as cited by
             Umpqua River �    � *                        Wiebe 1968). Adults and juveniles appearto school in
                Coos Bay * *                              separate areas (Shaw et al. 1974). The shiner perch
              Rogue River � � �                           may use intertidal eelgrass beds significantly more at
             Klamath River (_3 1 3                        night than day (Bayer 1981).
             Humboldt Bay � �
                Eel River � �                             Reproduction
              Tomales Bay 13 Ci S                         Mode: This species is gonochoristic and iteroparous. It
        Cent. San Fran. Bay' d i �   * IncludesCentralSan  iS ovoviviparous; eggs are fertilized internally (Wiebe
        South San Fran. Bay  * (  and San Pablo bays.     1968, Garrison and Miller 1982).
            Elkhom Slough *  1
               Morro Bay         �                        Matina/SDawnina: The shinerperch performs elaborate
          Santa Monica Bay      O                         courtship and mating behavior.  This behavior has
            San Pedro Bay0 0  O                           been broken down into six phases: (1) male(s) will
              Alamitos Bay 0O C I)                        chase females, (2) one male will isolate one female
             Anaheim Bay O CO                            from other females, (3) the male will aggressively
              Newport Bay 0 0 O                           protect his female from other male shiner perch, (4)
              Mission Bay  3 1            a               with his dorsal fin raised, the male will swim in a figure-
            San Diego Bay  �                              eight interspersed with wide circular sweeps in front of
            Tijuana Estuary    i                          and around the female; this may continue for many
                        A P J                             minutes and be interrupted periodically by aggressive
                                                          attacks against other males, (5) the male becomes limp
    substrates ranging from silt-claytoboulders (Simenstad    and quivers near the female, this is associated with
    1983). In Yaquina Bay, Oregon, 95% were collected    rapid jaw and dorsal fin movement, (6) the male turns
    on eelgrass beds (Bayer 1979, 1981).                  on its side and applies his anal fin appendages to the
                                                          urogenital region to copulate with the female (Wiebe
    Phvsical/ChemicalCharacteristics:Juvenilesandadults   1968). The courtship behavior can be lengthy, but
    occur in oligohaline to euhaline waters (Moyle 1976,    copulation may last only a fraction of a second (Wiebe
    Simenstad 1983) and occasionally in fresh water    1968). Matingoccursprimarilyinthespring-summerin
    (Beardsley and Bond 1970, Moyle 1976). While in   California (Bane and Robinson 1970, Shaw 1971),
    estuaries they are normally found in salinities >8-1 0%o   April-July in British Columbia (Hart 1973), and probably
    (Moyle 1976). During the spring and summer when    summer in Oregon and Washington.   Sperm is
    adults are giving birth, large schools are found in   apparently stored in the female for several months
    mesohaline and polyhaline waters (Ganssle 1966,    before fertilization occurs in the winter (Eigenmann
    Moyle 1976). The upper lethal temperature is 26.5-   1892, Wiebe 1968). Females give birth during April
    30.00C (Stober 1973). The shiner perch is reported to    and May in California (Odenweller 1975), June and
    occur in temperatures ranging from 4 to 21�C (Tarp   July in British Columbia (Wiebe 1968),July and August
    1952), but shiner perch left Anaheim Bay, California,   in Puget Sound (Wydoski and Whitney 1979) and
    when temperatures exceeded 18.5�C (Odenweller   spring in Oregon (Beardsley and Bond 1970).

                                                       227

    Shiner perch continued
    Fecundity: The reproductive capacityof this species is   amphipods, algae, mussels, barnacle appendages,
    directly related to female size; small young females    polychaetes, bivalves, crab larvae, cladocera, isopods,
    produce as few as five young, while largerolderfemales    and mysids (Bane and Robinson 1970, Bane and Bane
    can produce over 20 (Wilson and Millemann 1969). A    1971, Hart 1973, Odenweller 1975, Bottom et al. 1984).
    female may produce up to 36 young (Clemens and
    Wilby 1961).                                          Biological Interactions
                                                          Predation: The shiner perch is eaten by many species
    Growth and Development                               of large marine fishes [e.g., sturgeon (Acipenserspp.),
    Eaa Size and Embryonic Develooment: Embryonic    salmon (Oncorhynchus spp.), and barred sand bass
    development is direct and internal. Eggs are 0.3 mm in   (Paralabraxnebulifeo] (Wydoski and Whitney 1979). It
    diameter (Eigenmann 1892). Embryos are initially   is a seasonally important prey for harbor seal (Phoca
    0.45 mm in sagittal section (Wang 1986). Embryos    vitulina) (Simenstad et al. 1979, Jeffries et al. 1984)
    develop spatulate vascular expansions of tissue at the   and piscivorous birds such as cormorant (Phalacrocorax
    margins of the dorsal and anal fins to aid in oxygen and    spp.), great blue heron (Ardia herodias), and bald
    carbon dioxide exchange (Turner 1952). During later   eagles (Haliaeetus leucocephalus) (Bayer 1979, M. G.
    stages of development, a fold of ovarian tissue may    Garrett 1985, Pacific Power and Light, Portland, OR,
    invadethe opercularopening of some embryos (Turner   pers. comm.).
    1952).
                                                          Factors Influencina PoDulations: There is little
    Aae and Size of Larvae: There is no larval stage;    information available regarding the factors influencing
    embryonic development is direct and internal.        shiner perch populations. High water temperatures
                                                          may reduce the length of estuarine residence
    Juvenile Size Ranae: At birth, the fully-developed    (Odenweller 1975).  The availability and quality of
    shiner perch averages 34.0-43.7 mm long (Wilson and    estuarine areas for giving birth and rearing may also
    Millemann 1969, Wang 1986). Juveniles are less than   limit shiner perch abundance.  The shiner perch
    5.0 cm long (Shaw 1971).                             populations in San Pedro Bay and adjacent areas have
                                                          been declining since 1974, but it is not known why
    Aae and Size of Adults: The shiner perch can live for 8    (Stephens et al. 1983).
    years and growto 20 cm in length (Beardsley and Bond
    1970, Wydoski and Whitney 1979). However, fish over   References
    6 years old are rare and most are under 16.5 cm in
    length (Anderson and Bryan 1970). Males are smaller   Anderson, R. D., and C. F. Bryan. 1970. Age and
    than females and are rarely longer than 13.0 cm    growth of three surfperches (Embiotocidae) from
    (Anderson and Bryan 1970). Growth is very rapid the   Humboldt Bay California. Trans. Am. Fish. Soc. 3:475-
    first year and then slows considerably (Anderson and    482.
    Bryan 1970, Bane and Robinson 1970, Odenweller
    1975).  Males mature soon after birth, but are not   Bane, G. W. 1968. Fishes of the upper Newport Bay.
    mature at birth as earlierthought (Shaw 1971, Garrison    Univ. Calif. Irvine Res. Ser. 3:1-114.
    and Miller 1982). Most females mature their first year
    (Wilson and Millemann 1969, Shaw 1971, Shaw et al.   Bane, G. W., and A. W. Bane. 1971. Bay fishes of
    1974), except in British Columbia (Gordon 1965 as    northern California with emphasis on the Bodega
    cited in Garrison and Miller 1982).                   Tomales Bay area. Mariscos Publ., Hampton Bays,
                                                          New York, NY, 143 p.
    Food and Feeding
    Trophic Mode: Embryos receive oxygen and nutrition   Bane, G. W., and M. Robinson. 1970. Studies on the
    from highly-developed ovarian cavitytissues and fluids   shiner perch, Cymatogaster aggregata Gibbons, in
    (Wiebe 1968). Juveniles and adults are omnivorous    upper Newport Bay, California.  Wasmann J. Biol.
    (Bane and Bane 1971). Food eaten depends on sex,   28(2):259-268.
    age, and season (Hart 1973). Juveniles and adults will
    feed on benthos or plankton, depending on prey   Baxter, J. L. 1960. Inshore fishes of California. Calif.
    availability (Odenweller 1975). Juveniles and adults   Dept. Fish Game, Sacramento, CA, 80 p.
    can be nocturnal or day feeders (Hobson et al. 1981,
    Hobson and Chess 1986).                               Bayer, R. D. 1979. Intertidal shallow-waterfishes and
                                                          selected macroinvertebrates in the Yaquina estuary,
    Food Items: Juveniles and small adults eat primarily   Oregon.  Unpubl. Rep., 134 p. Oregon State Univ.
    copepods (Hart 1973). Other prey include gammarid    Marine Sci. Cent. Library, Newport, OR.

                                                       223

                                                                                         Shinerperch continued

    Bayer, R. D.   1981.   Shallow-water intertidal    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    ichthyofaunaoftheYaquinaestuary, Oregon. Northw.    Board Can., Bull. No. 180, 740 p.
    Sci. 55(3):182-193.
                                                           Hobson, E. S., and J. R. Chess. 1986. Relationships
    Beardsley, A. J., and C. E. Bond. 1970. Field guide to    among fishes and their prey in a nearshore sand
    common marine and bay fishes of Oregon. Agr. Exp.    community off southern California.  Env. Biol. Fish.
    Sta. Bull. No. 607, Oregon State Univ., Corvallis, OR,    17(3):201-226.
    27 p.
                                                           Hobson, E. S., W. N. McFarland, and J. R. Chess.
    Bottom, D. L., K. K. Jones, and M. J. Herring. 1984.    1981. CrepuscularandnocturnalactivitiesofCalifornia
    Fishes of the Columbia River estuary. Columbia River    nearshore fishes, with consideration of their scotopic
    Data Dev. Prog., CREST, Astoria, OR, 113 p. plus   visual pigments and the photic environment.  Fish.
    appendices.                                             Bull., U.S. 79(1):1-30.

    Clemens, W. A., and G. V. Wilby. 1961. Fishes of the    Horn, M. H. 1974. Fishes. InA summary of knowledge
    Pacific coast of Canada. Fish. Res. Board Can., Bull.   of the southern California coastal zone and offshore
    No. 68, 443 p.                                          areas, Chapter 11. South. Calif. Ocean Stud. Consort.,
                                                           Fullerton, CA, 124 p.
    Earnest, R. D., and P. E. Benville, Jr. 1971. Correlation
    of DDT and lipid levels for certain San Francisco Bay    Jeffries, S. J., S. D. Treacy, and A. C. Geiger. 1984.
    fish. Pest. Monitor. J. 5(3):235-241.                   Marine mammals of the Columbia River estuary.
                                                           Columbia River Estuary Data Dev. Prog., CREST,
    Earnest, R. D., and P. E. Benville, Jr. 1972. Acute    Astoria, OR, 62 p. plus appendices.
    toxicity of four organochlorine insecticides totwo species
    of surf perch. Calif. Fish Game 58(2):127-132.          Morrow, J. E. 1980. The freshwater fishes of Alaska.
                                                           Alaska Northw. Publ. Co., Anchorage, AK, 248 p.
    Eigenmann, C. L. 1892. Cymatogaster aggregata
    Gibbons; a contribution to the ontogeny of viviparous    Moyle, P. B. 1976. Inland fishes of California. Univ.
    fishes. Bull. U.S. Fish Comm. Vol. XII:401-478.         Calif. Press, Berkeley, CA, 405 p.

    Eschmeyer, W. N., W. S. Herald, and H. Hammann.    Odenweller, D. B. 1975. The life history of the shiner
    1983. A field guide to Pacific coast fishes of North    surfperch, Cymatogaster aggregata  Gibbons, in
    America. Houghton Mifflin Co., Boston, MA, 336 p.       Anaheim Bay, California. In E. D. Lane and C. W. Hill
                                                           (editors), The marine resources of Anaheim Bay. Calif.
    Frey, H. W. 1971. California's living marine resources    Fish Game, Fish Bull. 165:107-115.
    and their utilization. Calif. Dept. Fish Game,
    Sacramento, CA, 148 p.                                  Proctor, C. M., J. C. Garcia, D. V. Galvin, G. B. Lewis,
                                                           and L. C. Loehr. 1980. An ecological characterization
    Ganssle, D. 1966. Fishes and decapods of San Pablo    of the Pacific Northwestcoastal region. 5vol. U.S. Fish
    and Suisun Bays. In D. W. Kelley (compiler), Ecological    Wildl. Serv., Biol. Serv. Prog. (FWS/OBS-79/11 through
    studies of the Sacramento-San Joaquin estuary. Calif.   79/15), various pagination.
    Fish Game, Fish Bull. 133:64-94,
                                                            Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Garrison, K. J., and B. S. Miller. 1982. Review of the    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    earlylifehistoryofPugetSoundfishes. Fish. Res. Inst.,   of common and scientific names of fishes from the
    Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).         United States and Canada. Am. Fish. Soc. Spec. Publ.
                                                            No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Gates, D. E., and H. W. Frey.  1974.  Designated
    common  names of certain marine organisms of    Roedel, P. M.  1953.  Common ocean fishes of the
    California. Calif. Fish Game, Fish Bull. 161:55-90.     California coast. Calif. Fish Game, Fish Bull. 91,
                                                            184 p.
    Gordon, C. D. 1965. Aspects of the age and growth of
    Cymatogasteraggregata Gibbons. M.S. Thesis, Univ.    Shaw, E.  1971.  Evidence of sexual maturation in
    British Columbia, Vancouver, B.C., 90 p.                young adult shiner perch, Cymatogaster aggregata
                                                            Gibbons (Perciformes, Embiotocidae). Am. Mus. Nov.
                                                            2479:1-10.

                                                        229

    Shiner perch continued
    Shaw, E., J. Allen, and R. Stone.  1974. Notes on    Interagency ecological study program for the
    collection of shiner perch, Cymatogasteraggregata in   Sacramento-San Joaquin estuary. Calif. Dept. Water
    Bodega Harbor, California. Calif. Fish Game 60(1):15-    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    22.                                                     and U.S. Fish Wildl. Serv., various pagination.

    Simenstad, C. A.  1983. The ecology of estuarine    Washington, P. M.  1977.  Recreationally important
    channels of the Pacific Northwest coast: a community    marine fishes of Puget Sound, Washington.  Proc.
    profile. U.S. Fish Wildl. Serv., FWS/OBS-83/05,181 p.   Rep., Northwest Alaska Fish. Cent., Nat. Mar. Fish.
                                                           Serv., NOAA, 2725 Montlake Blvd. E., Seattle, WA,
    Simenstad, C. A., B. S. Miller, C. F. Nyblade, D.    122 p.
    Thornburgh, and L. J. Bledsoe. 1979. Food web
    relationshipsofnorthernPugetSoundandtheStraitof    Wiebe, J. P.  1968.  The reproductive cycle of the
    Juan de Fuca: a synthesis of the available knowledge.    viviparous seaperch, Cymatogaster aggregata
    U.S. Interagency (NOAA, EPA) Enery/Environ. Res.    Gibbons. Can. J. Zool. 46:1221-1234.
    Dev. Prog. Rep. EPA-60017-79-259, Washington, D.C.,
    335 p.                                                  Wilson, D. C., and R. E. Millemann. 1969. Relationships
                                                           of female age and size to embryo number and size in
    Smith, S. E., and S. Kato. 1979. The fisheries of San    the shiner perch, Cymatogaster aggregata. J. Fish.
    Francisco Bay: past, present and future.  In T.J.    Res. Board Can. 267:2339-2344.
    Conomos (editor), San Francisco Bay: the urbanized
    estuary, p. 445-468. Am. Assoc. Adv. Sci, and Calif.   Wydoski, R.S.,and R. R.Whitney. 1979. Inland fishes
    Acad. Sci., San Francisco, CA.                          of Washington, Univ. Wash. Press, Seattle, WA, 220 p.

    Stephens, J. S., Jr., P. A. Morris, and W. Westphal.
    1983. Assessing the effects of a coastal steam electric
    generating station on fishes occupying its receiving
    waters. In D. F. Soule and D. Walsh (editors), Waste
    disposal in the oceans; minimizing impact, maximizing
    benefits, p. 194-208. Westview Press, Boulder, CO.

    Stober, Q. J. 1973. Summary and overview of
    experimental thermal effects studies. In Q. J. Stober
    and E. O. Salo (editors), Ecological studies of the
    proposed Kiket Island nuclear power site, p. 441 -448.
    Fish. Res. Inst., Coill. Fish., Univ. Wash., Seattle, WA
    (FRI-UW-7304).

    Stober, Q. J., D. T. Griggs, and D. L. Mayer. 1973.
    Species diversity of the marine fish community in north
    Skagit Bay. In Q., J. Stober and E. O. Salo (editors),
    Ecological studies of the proposed Kiket Island nuclear
    power site, p. 373-400. Fish. Res. Inst., Coill. Fish.,
    Univ. Wash., Seattle, WA (FRI-UW-7304).

    Tarp, F. H. 1952. A revision of the family Embiotocidae
    (the surfperches). Calif. Fish Game, Fish Bull. 88,
    99 p.

    Turner, C. L. 1952. An accessory respiratory device in
    embryos of the embiotocid fish, Cymatogaster
    aggregata, during gestation. Copeia 1952(3):146-
    147.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: A
    guide to the early life histories. Tech. Rep. 9.

                                                       230

    231

       Ammodytes hexapterus
       Adult

                         5cm

    Common Name: Pacific sand lance                        will stay burrowed (Pearson et al. 1984). Contaminated
    Scientific Name: Ammodytes hexapterus                  sediments (300 ppm and 3,000 ppm oil) may also
    Other Common Names: sandlance, sand launce,    cause hemorrhaging in the head and gill regions of
    sand eel                                               Pacific sand lance (Pearson et al. 1984).
    Classification (Robins et al. 1980)
    Phylum: Chordata                                       Range
    Class: Osteichthyes                                    Overall: Overall range is from southern California to
    Order: Perciformes                                     Alaska and the Bering Sea, from Arctic Alaska to the
    Family: Ammodytidae                                    Sea of Japan (Eschmeyer et al. 1983). The center of
                                                           abundance appears to be in the Gulf of Alaska (Trumble
    Value                                                  1973). It is also found in Arctic waters, Hudson Bay, the
    Commercial: The Pacific sand lance is not commercially    northwest Atlantic Ocean, and Europe (Leim and Scott
    fished in the U.S. and Canada except for a limited   1966).
    amount for use as bait. Commercial fisheries exist in
    Japan and Europe; the Japanese Pacific sand lance    Within Studv Area: The Pacific sand lance is common
    fishery takes about 100,000 t/year (Field 1987).       to highly abundant in Puget Sound, but has highly
                                                           patchy distributions in marine areas of many other
    Recreational: This species is not generally used for   Pacific coast estuaries (Table 1) (Monaco et al. 1990).
    human consumption, but is reported to be delicious
    (Clemens and Wilby 1961). It is mostly used as bait for   Life Mode
    larger fishes.                                         Eggs are demersal and adhesive. Larvae, juveniles,
                                                           and adults are pelagic and schooling, but juveniles and
    Ecological: The Pacific sand lance is an important prey    adults are occasionally demersal (Garrison and Miller
    for many different species of marine vertebrates (Hart    1982).
    1973) and some invertebrates. It is the main prey for
    many seabirds in the northern Gulf of Alaska (Sanger    Habitat
    1987)andwasthedominantfishcapturedinanearshore    Tye: Adults and juveniles rest and escape from
    habitat (<30 m deep) in Alaska (Houghton 1987).    predators by burrowing into clean, unconsolidated
    Because of its life history characteristics, it is not often    substrates. A neritic species, it is usually associated
    sampled by normal trawl gear. Two Ammodytesspecies    with clean sand bottoms in areas <100 m deep (Trumble
    occur off Japan that are morphologically very similar   1973). However, it may be found to depths of 275 m
    but probably distinct species: A. hexapterus and A.    (Allen and Smith 1988).  Since it needs clean,
    personatus (Okamoto 1989).                             unconsolidated sand to burrow into and still have
                                                           sufficient oxygen, these burrow areas typically have
    Indicator of Environmental Stress: Oil-contaminated    high bottom current velocities.  Hence, areas with
    sediment reduces the amount of time that this species    suitable current velocities and substrate types are

                                                       232

                                                                                     Pacific sand lance continued
                                                           Miarations and Movements: No migration has been
      Table 1. Relative abundance of Pacific sand          documented, but juveniles and adults probably move
               lance in 32 U.S. Pacific coast estuaries.   into coastal and estuarine waters during spring and
                         Life Stage                        summerto feed and escape from predators. In summer,
            Estuary    A S J L E                           they are most abundant in nearshore habitats (Craig
            PugetSound 0   �a � 6 0�   Relative abundance:  1987). In Alaska, 1- and 2-year-old sand lance appear
             Hood Canal 0 � * � �    �   Highly abundant   to move inshore in early summer and then offshore
              SkagitBay O *�  �   @  3  Abundant          beginning in late August (Houghton 1987). On the
            Grays Harbor O       bO   0   Common           Atlantic coast, newly-hatched Ammodytesspp. larvae
             WillapaBay O      0           Rare            are found throughout the water column in well-mixed
           ColumbiaRiver O    O O     Blank Notpresent     shelf waters, with most larvaefound inwaters lessthan
            Nehalem Bay  O    t) O                         10-20 m deep. Larger larvae appear to spend the day
            Tillamook Bay 1    ta O   Life stage:          near the bottom and move up into the water column at
             NetartsBay  ]   i CD    A -Adults             night.  By April and May, most pre-metamorphosis
                                      S - Spawning adults
             Siletz River O    O O   J -Juveniles         juveniles were captured at night, indicating they were
             Yaquina Bay C    0�      L - Larvae           nearthebottomorburrowed inthe substrateduringthe
             Alsea River O       O     Eggs                day (Potter and Lough 1987). At night, A. hexapterus
            Siuslaw River O    O O                         juveniles and adults appear to burrow into the bottom
           UmpquaRiver (    (3 0                           (GirsaandDanilov1976, Hobson1986). During winter,
               Coos Bay O    0 0                           adults are relatively inactive and remain buried in clean
             Rogue River                                   sand except when spawning (Pinto 1984). Juveniles
            Klamath River                                  and adults often form mixed feeding schools with
           Humboldt Bay 0  0 0                            Pacific herring (Clupeapallasi), but they may also form
               Eel River :  I,                             dense "balls" or tight monospecific schools during the
            Tomales Bay 0 0 0 0:0                          day.
       Cent San Fran. Bay *   * Includes Central San
                                     Francisco, Suisun.
       South San Fran. Bay  ' , \    and San Pabo bays.    Reproduction
           Elkhom Slough  q    1                           Mode: The Pacific sand lance is gonochoristic,
              Morro Bay                                    oviparous, and iteroparous; eggs are fertilized
        Santa Monica Bay                                  externally.
           San Pedro Bay
            Alamitos Bay                                   M atina/Soawnina: The spawning biology of this species
            Anaheim Bay                                    is not well-studied, but is assumed to be similar to that
            Newport Bay                                    of the Atlantic sand lance (A. americanus). The Pacific
             Mission Bay                                   sand lance spawns in marine waters during the winter
           San Diego Bay                                   (November-March) (Andriyashev 1954, Fitch and
          Tijuana Estuary                                  Lavenberg 1975, Wang 1986) in varying depths of
                      A S J L E                           water, and probably in strong currents (Andriyashev
                                                           1954). Along Kodiak Island, Alaska, spawning occurs
    critical for defining proper habitat (Auster and Stewart    intertidally at high tide in October (Dick and Warner
    1986). This type of habitat is often found at the mouths    1982).
    of estuaries and may be the reason these fish are often
    found there.                                           FecundJit: This species' fecundity is unknown, but
                                                           other Ammodytes species have been found to have
    Substrate: Larvae arefoundoveravarietyofsubstrates.    3,300-22,100 eggs per female, averaging 6,800 per
    When pelagic, juveniles and adults are found over    female (Andriyashev 1954).
    various substrates. When they burrow, they choose
    clean, unconsolidated sand (perhaps with some small    Growth and Development:
    gravel). Eggs are also found in these substrates.      Ean Size and Embrvonic Develooment: Fertilized eggs
                                                           are spherical and 0.88-1.20 mm in diameter (Pinto
    Phvsical/Chemical Characteristics: The Pacific sand    1984). They also have an oil globule and adhere to
    lance is primarily a marine species; larvae are found in   sand grains (Williams et al. 1964).  Embryonic
    full seawater to mesohaline waters (Wang 1986).    development is indirect and external. Near Japan,
    However, it is often found in sandy areas near freshwater    eggs hatch in 33 days at 6.2�C, with optimaltemperature
    seeps.                                                 being 8.20C (Inoue et al. 1967). At 90C, eggs hatch in
                                                           24 days (Pinto 1984).

                                                       233

    Pacific sand lance continued
    Ace and Sizeof Larvae: Larvae apparently stay in sand    Factors Influencina Pooulations: Little is known
    until they are 4-5 mm  standard length (SL) (Reay    concemingfactorsthatinfluencepopulations,butlarval
    1970). At hatching, they are 4.9-5.7 mm SL (noue et al.   survival and predation on all life stages are believed to
    1967, Pinto 1984) and grow to 30-40 mm long before    be most important. Major spawning areas have not
    metamorphosis.                                         been positively identified, butthe areas where prolarvae
                                                           have been found indicate spawning occurs in and at
    Juvenile Size Rance: The juvenile size range is   the mouths of bays and estuaries (Wang 1986). Larval
    unknown, but probably from 0.4 cm up to 10.0 cm total   fish surveys in the northwestern Atlantic showed a 20-
    length.                                                fold increase in abundance of Ammodytes species
                                                           from 1974 to 1979, reflecting a 50-fold change in adult
    Aae and Size of Adults: This species may become    spawning biomass (Field 1987).  Studies of other
    sexually mature after 1 to 3 years (approximately 10 cm    Ammodytes species indicate watertemperature during
    long). In Alaska, juveniles appear to mature at 2 or 3    spawning season may affect recruitment, and some
    years(DickandWarner1982). FewalongtheCalifornia    density-dependent effects of recruitment and growth
    coast reach 20 cm long, butthis speciescangrowto 28    have been noted.  Increases in populations of the
    cm in length (Hart 1973). The Pacific sand lance may    Newfoundland and North Seas may be related to
    live to be 8 years old (Fitch and Lavenberg 1975).     decreases in predator populations (cod and mackerel)
                                                           (Field 1987). In the lower Columbia River estuary, the
    Food and Feeding                                        Pacific sand lance is the dominant fish captured during
    Trophic Mode: Larvae, juveniles, and adults are    annual hopper dredging operations (K. Larson, U.S.
    planktivorous carnivores.                               Corps of Engineers, Portland District, Portland, OR,
                                                           pers. comm.).
    Food  Items:  Small  larvae eat diatoms  and
    dinoflagellates, while larger larvae consume copepods    References
    and copepod nauplii (Garrison and Miller 1982).
    Juveniles and adults feed primarily on copepods    Allen, M. J., and G. B. Smith.  1988.  Atlas of
    (Simenstad et al. 1979), with other plankton being    zoogeographyofcommonfishesintheBeringSeaand
    supplementary (Hart 1973). In Alaska, juveniles and    northeastern Pacific. NOAA Tech. Rep. NMFS 66,
    adults feed on zooplankton (primarily euphausiids in   151 p.
    winter and copepods in summer), but their diet varies
    greatly between years (Craig 1987).                     Andriyashev, A. P. 1954. Fishes of the northern seas
                                                           of the USSR. Akad. Nauk SSR, Opred. po. FauneSSR
    Biological Interactions                                 53,556 p. (1964 transl. available, Nat. Tech. Int. Serv.,
    Predation: The Pacific sand lance is eaten by crabs,    Springfield, VA).
    seals, whales, and many species of fish, including
    Pacific cod (Gadus macrocephalus), Pacific halibut   Auster, P. J., and L. L. Stewart. 1986. Species profiles:
    (Hippoglossus stenolepis), Pacific hake (Merluccius   life histories and environmental requirements of coastal
    productus), sole, lingcod (Ophiodon elongatus),    fishes and invertebrates (North Atlantic)-sand lance.
    scorpaenids, salmonids, and sculpins.  Many birds    US. Fish Wildl. Serv. Biol. Rep. 82(11.66), U.S. Army
    also prey on the sand lance, including kittiwake (Rissa    Corps Eng., TR EL-82-4, 11 p.
    spp.), common murres (Uria aalge), puffins, rhinoceros
    auklet (Cerorhinca monocerata), ancient murrelet    Beacham, T. D. 1986. Type, quantity, and size of food
    (Synthliboramphus antiquum), sooty shearwater    of Pacific salmon (Oncorhynchus) in the Strait of Juan
    (Puffinus griseus), cormorants (Phalacrocorax spp.),    de Fuca, in British Columbia. Fish. Bull., U.S. 84(1):77-
    red-throated loon (Gavia stellata), and gulls (Field   89.
    1987). It is an important prey for juvenile salmonids off
    Oregon and Washington (Peterson et al. 1983, Emmett    Clemens, W. A., and G. V. Wilby. 1961. Fishes of the
    et al. 1986), and the primary fish prey for salmonids in   Pacific coast of Canada. Fish. Res. Board Can., Bull.
    the Strait of Juan de Fuca (Beacham 1986).  This    No. 68, 443 p.
    species is also a primary forage fish along the northern
    shore of the Alaska Peninsula (Craig 1987). Intense    Craig, P. 1987. Forage fishes in the shallow waters of
    predation often occurs when the Pacific sand lance    the north Aleutian Shelf. In M. J. Allen and R. R. Ware
    undertakes the transition from sediment burrows to life   (editors), Forage fishes of the southeastern Bering
    in the water column (Hobson 1986).                      Sea, Conference proceedings, p. 49-54. U.S. Dept.
                                                           Int., Min. Manag. Serv., Anchorage, AK.

                                                       234

                                                                                    Pacific sand lance continued
    Dick, M. H., and I. M. Warner.  1982.  Pacific sand    Leim, A. H., and W. B. Scott.  1966.  Fishes of the
    lance, Ammodytes hexapterus Pallas, in the Kodiak    Atlantic coast of Canada. Fish. Res. Board Can. Bull.
    Island group. Syesis 15:43-50.                          No. 155, 485 p.

    Emmett, R. L., D. R. Miller, and T. H. Blahm. 1986.    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    Foodof juvenilechinook, Oncorhynchustshawytscha,    Nelson. 1990. Distribution and abundance of fishes
    and coho, 0. kisutch, salmon off the northern Oregon    and invertebrates in west coast estuaries, Volume I:
    and southern Washington coasts, May-September    data summaries. ELMR Rep. No.4. Strategic Assess-
    1980. Calif. Fish Game 72(1): 38-46.                    ment Branch, NOS/NOAA, Rockville, MD,
                                                           240 p.
    Eschmeyer, W. N., W. S. Herald, and H. Hammann.
    1983. A field guide to Pacific coast fishes of North    Okamoto, H. 1989. A genetic comparison of sympatric
    America. Houghton Mifflin Co., Boston, MA, 336 p.       populations of sand lance (Genus Ammodytes) from
                                                           the region east of Cape Soya, Japan. Can. J. Fish.
    Field, L. J.  1987.  Pacific sand lance, Ammodytes    Aquat. Sci. 46:1945-1951.
    hexapterus, with notes on related Ammodytesspecies.
    In N. J. Wilimovsky, L. S. Incze, and S. J. Westrheim    Pearson, W. H., D. L. Doodruff, P. C. Sugarman, and
    (editors), Species synopses, life histories of selected    B. L. Olla.  1984. The burrowing behavior of sand
    fish and shellfish of the northeast Pacific and Bering    lance, Ammodytes  hexapterus: effects of oil-
    Sea, p. 15-33. Wash. Sea Grant Prog., and Fish. Res.    contaminated sediment. Mar. Environ. Res. 11:17-32.
    Inst., Univ. Wash, Seattle, WA.
                                                           Peterson, W. T., R. D. Brodeur, and W. A. Pearcy.
    Fitch, J. E., and R. J. Lavenberg. 1975. Tidepool and    1983.  Feeding habits of juvenile salmonids in the
    nearshore fishes of California. Calif. Nat. Hist. Guides:    Oregon coastal zone in June 1979. Fish Bull., U.S.
    38, Univ. Calif. Press, Berkeley, CA, 156 p.            80(4):841-851.

    Garrison, K. J., and B. S. Miller. 1982. Review of the    Pinto, J. 1984. Laboratory spawning of Ammodytes
    earlylifehistoryofPugetSoundfishes. Fish. Res. Inst.,   hexapterus from the Pacific coast of North America
    Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).         with a description of its eggs and early larvae. Copeia
                                                           1984:242-244.
    Girsa, I. I., and A. N. Danilov. 1976. The defensive
    behavior of the white sea sand lance Ammodytes    Potter, D. C., and R. G. Lough. 1987. Verticaldistribution
    hexapterus. J. Ichthyol. 16:862-865.                    and sampling variability of larval and juvenile sand
                                                           lance (Ammodytes sp.) on Nantucket Shoals and
    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    Georges Bank. J. Northw. Atl. Fish. Sci. 7:107-116.
    Board Can., Bull. No. 180, 740 p.
                                                           Reay, P. J. 1970. Synopsis of biological data on North
    Hobson, E. S. 1986. Predation on the Pacific sand    Atlanticsand-eelsofthegenusAmmodytes(A. tobianus,
    lance, Ammodytes hexapterus (Pisces: Ammodytidae),    A. dubius, A. americanus, and A. marinus). FAO Fish.
    during the transition between day and night in   Synop. 82, various pagination.
    southeastern Alaska. Copeia 1986:223-226.
                                                           Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Houghton, J. P. 1987.  Forage fish use of inshore    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    habitats north of the Alaska peninsula. In M. J. Allen,   of common and scientific names of fishes from the
    and R. R. Ware (editors), Forage fishes of the    United States and Canada. Am. Fish. Soc. Spec. Publ.
    southeastern Bering Sea, Conference proceedings, p.    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    39-47. U.S. Dept. Int., Min. Manag. Serv., Anchorage,
    AK.                                                     Sanger, G. A. 1987. Trophic interactions between
                                                           forage fish and seabirds in the southeastern Bering
    Inoue, A., S. Takamori, K. Kuniyaki, S. Kobayashi, and    Sea. In M. J. Allen and R. R. Ware (editors), Forage
    S. Nishina. 1967.  Studies on fishery biology of the    fishes of the southeastern Bering Sea, Conference
    sand-lance, Ammodytes personatus (Girard).  Bull.   proceedings, p. 19-28. U.S. Dept. Int., Min. Manag.
    Naikai Reg. Fish. Res. Lab. 25(121):1-335 (InJapanese,    Serv., Anchorage, AK.
    English summary).
                                                           Simenstad, C. A., B. S. Miller, C. F. Nyblade, D.
                                                           Thornburgh, and L. J. Bledsoe. 1979. Food web

                                                       235

    Pacific sand lance continued
    relationships of northern Puget Sound and the Strait of
    Juan de Fuca: a synthesis of the available knowledge.
    U.S. Interagency (NOAA, EPA) Energy/Environ. Res.
    Dev. Prog. Rep., EPA-600/7-79-259, Washington, D.C.,
    335 p.

    Trumble, R.J. 1973. Distribution, relative abundance,
    and general biology of selected underutilized fishery
    resources of the eastern north Pacific Ocean. M.S.
    Thesis, Univ. Wash., Seattle, WA, 178 p.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: A
    guide to the early life histories. Tech. Rep. No. 9.
    Interagency ecological study program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    and U.S. Fish Wildl. Serv., various pagination.

    Williams, G. C., S. W. Richards, and E. G. Farnworth.
    1964. Eggs of Ammodytes hexapterus from Long
    Island, New York. Copeia 1964:242-243.

                                                       236

    237

       Clevelandia ios
       Adult

                             2cm

    Common Name: arrow goby                               Range
    Scientific Name: Clevelandia lios                    Overall: The arrow goby is found from the Gulf of
    Other Common Names: mud goby (Gates and Frey   California, Baja California, to Vancouver Island, British
    1974)                                                Columbia (Miller and Lea 1972).
    Classification (Robins et al. 1980)
    Phylum: Chordata                                     Within Studvy Area: This species is found in most Pacific
    Class: Osteichthyes                                  coast estuaries, but is most abundant in southern
    Order: Perciformes                                   California bays, estuaries, and lagoons (Table 1)
    Family: Gobiidae                                     (Monaco et al. 1990).

    Value                                                 Life Mode
    Commercial: This species has no commercial value.    Eggs are semi-adhesive and demersal. Larvae are
                                                          pelagic, while juveniles and adults are demersal and
    Recreational: This species has no recreational value.   live freely or commensally in the burrows of the
                                                           innkeeper worm (Ricketts et al. 1985), and mud and
    Indicator of Environmental Stress: The arrow goby is   ghost shrimps (Prasad 1948).
    easy to keep in aquaria and is an excellent bioassay
    organism (Reish and Lemay 1988). However, very   Habitat
    little is known about this species' pollution tolerances.   IType: All life stages are found in intertidal and subtidal
                                                           areas of bays, estuaries, and lagoons (Prasad 1948,
    Ecological: The arrow goby is an importantcomponent    Carter 1965, Brothers 1975, Wang 1986). Larvae are
    of the ichthyofauna in many California estuaries, where    most abundant in areas of high salinity in San Francisco
    it plays a critical role in the food webs. It is the most    Bay, California (Wang 1986, California Department of
    abundant goby in Elkhorn Slough (Cailliet et al. 1977),    Fish and Game 1987). Juveniles and adults are found
    Anaheim Bay (Macdonald 1975), and Newport Bay,   inoligohalineto euhalinewaters (California Department
    California (Allen 1982). The arrow goby is commonly    of Fish and Game 1987).
    associated with the ghost shrimp (Callianassa spp.),
    but the shrimp probably derives no direct benefits from   Substrate: Eggs are laid on mud, sand, and sometimes
    the use of its burrows by arrow gobies (Hoffman 1981).   gravel (Wang 1986). Larvae can be found over a wide
    However, the arrow goby benefits from this association   rangeof substrates. Juveniles and adults prefer bottoms
    by having a refuge from predation and a residence   of mixed sand and mud, but they can also be found on
    during low tide. The arrow goby also uses the burrows   clay/sand (Prasad 1948) and other substrates.
    of the innkeeper worm (Urechis spp.) and mud shrimp
    (Upogebia spp.). However, goby abundance may not   Phvsical/Chemical Characteristics: Eggs are found at
    correlate with the density of any of these species'   temperatures >10�C (Wang 1986).  Juveniles and
    burrows (Macdonald 1975).                             adults are eurythermal, withstanding temperatures from

                                                       238

                                                                                         Arrowgoby continued
                                                          refuge within invertebrate burrows and intertidal pools
     Table 1. Relative abundance of arrow goby            (MacGinitie 1935, Prasad 1948, Macdonald 1975).
              in 32 U.S. Pacific coast estuaries.        Arrow gobies are most active at low light levels
                         Life Stage                       (Macdonald 1975). Light reflected from the silver belly
            Estuary    A S J L E                          of a threatened goby can stimulate other gobies to
             Puget Sound O 0 0   �   Relative abundance:  search for cover, thus causing gobies in an entire area
              Hood Canal 0 0 0 C 0    �   Highly abundanl    to retreat into burrows (Macdonald 1975). In some
               SkagitBay 0 0 0 O 0 0       Abundant       northern estuaries they mayonly use Callianassaspp.
             Grays Harbor 0 0 0 0 0    0   Common         burrows during spring and summer (Hoffman 1981).
              Wllapa Bay 0O O   0 0       Rare
           Columbia River            Blank Not present    Reproduction
             Nehalem Bay                                  M      ode:The arrow goby is gonochoristic, oviparous, and
            Tillamook SBay    i      Life stage:          possibly iteroparous; eggs are fertilized externally.
              Netarts Bay ~/   '     A - Adults
                                      S - Spawning adults
              Siletz River           J-Juveniles          Matina/SDawnina: Spawning occurs on intertidal mud
             Yaquina Bay O O o o o   E  LEargae          or sand flats of estuaries, bays, or lagoons (Wang
              Alsea River                                 1986). It may spawn year-round, depending on estuary
             Siuslaw River i    i                         (Brothers 1975). The principal spawning period is from
            Umpqua River i    i                           December to September. Peak spawning activity in
               Coos Bay O O O O O                         many southern California estuaries is from Februaryto
              Rogue River                                 June (Prasad 1948, Macdonald 1975), and from
            Klamath River                                 November to April in Mission Bay, California (Brothers
            Humboldt Bay O 0   0 0                       1975). The female's abdomen becomes swollen near
                Eel River                                 spawning time and the yellow color of the eggs shows
             Tomales Bay  0            3   de C           through the abdominalwall. Females may also develop
        Cent. San Fran. Bay' 0  ( 0  ' Includes Central San
                                     Franciso,. Suisun,   a streak of black pigment on the anal fin. Males show
        SouthSanFran.Bay  0 i c (3 (9  and San Pabo bays.  a considerable increase in pigmentation during the
            Elkhorn Slough lii (3i Si cii cii             spawning season; dorsal fins and the upper half of the
               Morro Bay (I    3 3  pectoral finsbecomedarkeranda black streakisfound
                                                          on the anal fin (Prasad 1948).  Females become
            San Pedro Bay         Q 0   0 C               lethargic near spawning time, while males are very
             Alamitos Bay cii iii cii ci l3               active.  Male breeding behavior includes fighting,
             Anaheim Bay ( {  ] (  ]
             NewportBay 0                                 chasing, nipping, and belly-flashing (Macdonald 1975).
              Mission Bay *  a g i 3                      No nest is built, eggs are deposited singly or in small
            San Diego Bay  3                              groups (Prasad 1948), with 15-25 eggs laid at a time
           Tijuana Estuary ( 0 ( ci c]                    (MacGinitie 1935). Eggs are laid on walls of a burrow
                       A S J L E                          which is about 10 cm deep (Wang 1986).

                                                          Fecundity: Fecundity ranges from 300-1,200 eggs per
    4-260C (Prasad 1948). Arrow gobies may inhabit the   female, depending on body size (Brothers 1975).
    cooler waters in invertebrate burrows when intertidal
    bay waters reach hightemperatures (Macdonald 1972).   Growth and Development
    The arrow goby spawns in polyhaline to euhaline    Eaa Size and Embrvonic DeveloDment: Eggs are
    waters(Wang1986). Juvenile andadultsareeuryhaline,   elliptical, club-shaped (Prasad 1948, Brothers 1975,
    tolerating fresh water and salinities greater than    Wang 1986), and 0.735 mm long and 0.645 mm wide
    seawater (Carter 1965). However, prolonged exposure    (MacGinitie 1935, Brothers 1975). They are adhesive
    to fresh wateror low salinities can result in death (L. G.    only at the anchoring point (Prasad 1948). Embryonic
    Allen, Calif. State Univ., Northridge, CA, pers. comm.).    development is indirect and external. At 15-15.50C,
    This species is also tolerant of low oxygen    hatchingtakes10-12days. Noparentalcareisprovided
    concentrations (Carter 1965).                          (Macdonald 1975).

    Minrations and Movements: Pelagic larvae are widely    Aae and Size of Larvae: Larval lengths range from
    transported within bays and lagoons and probably to   2.75-3.20mm at hatching (Prasad 1948, Brothers
    offshorewaters(Nordby1982, Wang1986). Intertidal-   1975).  Transformation to juvenile occurs at about
    dwelling juveniles and adults do not appearto migrate    14.0 mm  after the larvae develop the external
    down to subtidal habitats during low tide, but take   characteristics of adults (Prasad 1948).

                                                       239

    Arrowgoby continued
    Juvenile Size Ranae: Juveniles are from 14.0 mm to at   from spawning (Brothers 1975). The arrow goby is an
    least 29.0 mm long (Prasad 1948). Juveniles are less    estuary-dependent species, hence, any factor which
    than one year old (Prasad 1948, Brothers 1975).         impacts tidal flats and invertebrate burrows probably
                                                          directly affects arrow goby abundance. However,
    Aoe and Size of Adults: The arrow goby matures in at   annual freshwater inflow was not found to influence
    least one year, when it is longer than 29 mm.  All   arrow goby populations in San Francisco Bay (California
    females are mature by a length of 34 mm (Prasad    Department of Fish and Game 1987).
    1948, Brothers 1975). Some gobies may mature after
    one summer if they settled in spring (Brothers 1975).    References
    The maximum size reported is 52 mm (Carter 1965).
    Most live for only 1 year, but a few will live 2-3 years    Allen, L. G. 1982. Seasonal abundance, composition,
    (Prasad 1948, Brothers 1975). The sex of individuals    and productivity of the littoral fish assemblage in upper
    >19mm long can be distinguished bythe shapeof their    Newport Bay, California. Fish Bull., U.S. 80(4):769-
    anal papillae (Prasad 1948).                            790.

    Food and Feeding                                        Brothers, E. B. 1975. The comparative ecology and
    Trophic Mode: This species is primarily carnivorous    behavior of three sympatric California gobies. Ph. D.
    (Macdonald 1975).  Larvae are planktonic feeders,    Thesis, Univ. Calif., San Diego, CA, 365 p.
    while juveniles and adults are epibenthic/benthicfeeders
    (Prasad 1948, Brothers 1975, Macdonald 1975).           Cailliet, G. M., B. Antrim, D. Ambrose, S. Pace, and M.
                                                          Stevenson. 1977. Species composition, abundance
    Food Items: Larvae feed primarily on the copepod    and ecological studies of fishes, larval fishes, and
    Acartiatonsaandprobablyotherzooplankton. Juveniles    zooplankton in Elkhorn slough. In J. Nybakken, G.
    and adults consume harpacticoid and cyclopoid    Cailliet, and W. Broenkow (editors). Ecological and
    copepods, ostracods, nematodes, and oligochaetes.    hydrographic studies of Elkhorn Slough Moss Landing
    Gammarid and caprellid amphipods, and large    Harbor and nearshore coastal waters July 1974 to
    oligochaetes are important prey for larger gobies    June 1976, p. 216-386, Moss Landing Marine Lab.,
    (Prasad 1948, Macdonald 1975).  Other food may    Moss Landing, CA.
    include isopods, filamentous algae, crustacean nauplii
    and zoeae, diatoms, and tintinnids (Prasad 1948).    California Department of Fish and Game. 1987. Delta
    However, these items may only be eaten incidentally   outflow effects on the abundance and distribution of
    with other prey (Macdonald 1975). Besides the above    San Francisco Bay fish and invertebrates, 1980-1985.
    prey, pieces of food released by a ghost shrimp (while    Exhibit 60, entered by the California Department of
    ittears its food) maybe snatched and eaten (MacGinitie    Fish and Game forthe State Water Resources Control
    1934, cited by Carter 1965).                            Board 1987 Water Quality/Water Rights Proceeding
                                                           on the San Francisco Bay/Sacramento-San Joaquin
    Biological Interactions                                 Delta. Calif. Dept. Fish Game, Stockton, CA, 345 p.
    Predation: This species is consumed by many predators,
    including: California halibut (Paralichthys californicus)   Carter, W. R., il. 1965. Racial variations of the arrow
    (Haaker 1975), walleye surfperch (Hyperprosopon    goby, Clevelandia ios (Jordan and Gilbert) 1882 in
    argenteum), California corbina (Menticirrhusundulatus),    Puget Sound and on the coast of Washington State.
    whitecroaker (Genyonemus lineatus), Pacific staghorn    M.S. Thesis, Univ. Wash., Seattle, WA, 88 p.
    sculpin (Leptocottus armatus), diamond turbot
    (Hypsopsettaguttulata), queenfish (Seriphuspolitus),    Gates, D. E., and H. W. Frey.  1974.  Designated
    specklefin midshipman (Porichthys myriaster), round    common names of certain marine organisms of
    stingray (Urolophis hallern), shovelnose guitarfish    California. Calif. Fish Game, Fish Bull. 161:55-90.
    (Rhinobatos productus), California killifish (Fundulus
    parvipinnis), and probably many species of piscivorous    Haaker, P. L.  1975.  The biology of the California
    birds [gulls, greateryellowleg ( Totanusmelanoleucos),    halibut, Paralichthys californicus (Ayres). In E. D. Lane
    and short-billed dowitcher (Limnodromus griseus)]   and C. W. Hill (editors), The marine resources of
    (Prasad 1948, Brothers 1975, Macdonald 1975).           Anaheim Bay. Calif. Fish Game, Fish Bull. 165:137-
                                                           151.
    Factors Influencina Populations: Predation probably
    plays a major role in determining population size    Hoffman, C. J. 1981. Associations between the arrow
    (Macdonald 1975). Other important factors include    goby Clevelandia ios (Jordan and Gilbert) and the
    parasites, competition with other fishes, and stress

                                                       240

                                                                                          Arrowgoby continued
    ghost shrimp Callianassacaliforniensis Dana in natural    Press, Stanford, CA, 652 p.
    and artificial burrows. Pac. Sci. 35(3):211-216.
                                                           Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Horn, M. H., and L. G. Allen. 1985. Fish community    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    ecology in southern California bays and estuaries.    of common and scientific names of fishes from the
    Chapter 8.  In A. Yanez-Arancibia (editor), Fish    United States and Canada. Amer. Fish. Soc. Spec.
    community ecology in estuaries and coastal lagoons:    Publ., No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    towards an ecosystem integration, p. 169-190. DR (R)
    UNAM Press, Mexico.                                     Wang, J. C. S. 1986. Fishes of the Sacramento-San
                                                           Joaquin estuary and adjacent waters, California: A
    Macdonald, C. K. 1972. Aspects of the life history of   guide to the early life histories.  Tech. Rep. No. 9.
    the arrow goby, Clevelandia ios (Jordan and Gilbert),    Interagency ecological study program for the
    in Anaheim Bay, California, with comments on the    Sacramento-San Joaquin estuary. Calif. Dept. Water
    cephalic-lateralis system in the fish family Gobiidae.    Res., Calif. Dept. Fish Game, U.S. Bureau of Reclam.,
    M.S. Thesis, Calif. State Univ. Long Beach, CA, 157 p.    and U.S. Fish Wildl. Serv. various pagination.

    Macdonald, C. K. 1975. Notes on the family Gobiidae
    from Anaheim Bay. In E. D. Lane and C. W. Hill
    (editors), The marine resources on Anaheim Bay.
    Calif. Fish Game, Fish Bull. 165:117-121.

    MacGinitie, G. E. 1934. The natural history of
    Callianassacaliforniensis Dana. Am. Midl. Nat. 15:166-
    177.

    MacGinitie, G. E. 1935. Ecological aspects of a
    California marine estuary. Am. Midi. Nat. 16(5):629-
    765.

    Miller, D. J., and R. N. Lea. 1972. Guide to the coastal
    marine fishesof California. Calif. Fish Game, Fish Bull.
    157, 235 p.

    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    Nelson. 1990. Distribution and abundance of fishes
    and invertebrates in west coast estuaries, Volume I:
    data summaries. ELMR Rep. No.4. Strategic Assess-
    ment Branch, NOS/NOAA, Rockville, MD,
    240 p.

    Nordby, C. S. 1982. The comparative ecology of
    ichthyoplankton within Tijuana estuary and in adjacent
    nearshore waters. M.S. Thesis, San Diego State
    Univ., San Diego, CA, 101 p.

    Prasad, R. R. 1948. Life history of Clevelandia ios
    (Jordan and Gilbert). Ph.D. Thesis, Stanford Univ.,
    Stanford, CA, 141 p.

    Reish, D. J., and J. A. Lemay. 1988. Bioassay manual
    fordredged sediments. Res. Rep. to U.S. Army Corps
    Eng., Los Angeles District, Los Angeles, CA, 36 p. plus
    appendices.

    Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W.
    Phillips. 1985. Between Pacific tides. Stanford Univ.

                                                       241

       Ophiodon elongatus

                      25 cm

    Common Name: lingcod                                 concentrate heavy metals (Shaw and Hassler 1989).
    Scientific Name: Ophiodon elongatus
    Other Common Names: cultus cod (McClane 1978)    Ecological: This species is a major predator of smaller
    Classification (Robins et al. 1980)                  fishes and crustaceans in rocky reef habitats and kelp
    Phylum: Chordata                                     beds.
    Class: Osteichthyes
    Order: Perciformes                                    Range
    Family: Hexagrammidae                                Overall: The lingcod is found along coastal areas from
                                                          Baja California to Kodiak and Shumigan Islands in the
    Value                                                Gulf of Alaska (Hart 1973). It is most abundant from
    Commercial: The lingcod is an important commercial    Point Conception, California, to Cape Spencer, Alaska
    species, with over 4,000 t landed in 1985, worth $2.9    (MBC Applied Environmental Sciences 1987).
    million (National Marine Fisheries Service 1986). It is
    harvested from California to Alaska using trawls, long   Within Studv Area: The lingcod is common in Puget
    lines, and gill nets. Since the 1960s, there has been a    Sound and present in many otherestuaries of the study
    general reduction of commercial catches in both    area (Table 1) (Monaco et al. 1990). Small coastal
    Canadian and American waters (Cass 1981, Bargmann    estuaries are used primarily by juveniles.
    1981). It is the eighth most important commercial
    species in Puget Sound, Washington (by dollar value)   Life Mode
    (Bargmann 1981).  In Washington coastal waters,    Eggs are demersal and adhesive. Larvae and small
    most commercial catches occur between 40 and 100   juveniles (<70 mm long) are epipelagic, while larger
    fathoms (80-200 m) (Jagielo 1988).                    juveniles and adults are demersal (Miller and Geibel
                                                           1973). Adults are found in marine waters, intertidally
    Recreational: This is a prized sport fish because of its   and deeper (down to approximately 475 m), but are
    size and excellent taste (Eschmeyer et al. 1983). It is   most abundant at depths between 100-150 m (Allen
    the top California sport fish (by poundage) between Pt.   and Smith 1988) Juveniles settle out of the plankton
    Arguello and the Oregon border (Frey 1971), and the   into nearshore shallow-water areas (<20 m deep),
    seventh most important sport fish in Puget Sound (by   often where there is some freshwater runoff and lower
    number) (Bargmann 1981). This species is taken by   salinities (Day et al. 1986).
    anglers using hook and line from boats, piers, and
    shore, and also by spearfishing divers.               Habitat
                                                          Type: Eggs are laid in marine, rocky subtidal areas (to
    Indicator of Environmental Stress: Eggs require well-   at least 19 m below low tide) where adults reside. The
    oxygenated water (Giorgi and Congleton 1984). Oil   pelagic larvae occur in the near-surface waters in
    and otherpetrochemical spills may reduce populations    marine and estuarine areas (Hart 1973). Juveniles are
    (Shaw and Hassler 1989).  The lingcod may also   found in intertidal areas of shallow estuarine bays and

                                                       242

                                                                                              Lingcodcontinued
                                                            (MBC Applied Environmental Sciences 1987).
      Table 1. Relative abundance of lingcod in 32
               U.S. Pacific coast estuaries.                Miarations and Movements: Adults apparently move
                         Life Stage                         into shallow-water habitats during the spawning season
            Estuary    A S J  L E|                          (winter) (Miller and Geibel 1973), but in general, adults
             Puget Sound 0 O 0  Ol Relative abundance:     are relatively sedentary.  In spring, pelagic larvae
              Hood Canal O O  O  O  O    �   Highly abundant  (approximately 20 mm  in length) are transported
              SkagitBay O O O 0 0    @   Abundant           inshore. In late spring, (May and June) juveniles settle
            Grays Harbor    O O        O   Common           out or move into shallow-water coastal areas and
              Willapa Bay    0 0        I  Rare              estuaries (Phillips and Barraclough 1977). Juveniles
           Columbia River             Blank Nopresent       appearto move away from shallow-water sandy habitats
            Nehalem Bay     O                               in the fall and early winter, but like adults, do not appear
            Tillamook Bay    O         Life stage:          to show extensive migrations.
              Netarts Bay    O         A - Adults
                                      S - Spawning adults
              Siletz River            J - Juveniles         Reproduction
             YaquinaBay 0    0         L-Larvae             MQde: The lingcod is gonochoristic, oviparous, and
              Alsea River   gg                              iteroparous; eggs are fertilized externally.
            Siuslaw River
            Umpqua River     O                               Matina/SDawnina: Spawning occurs from November
               Coos Bay  i   o O                            to March off California, and Decemberto March/April in
             Rogue River                                    Puget Sound (LaRiviere et al. 1981). Peak spawning
            Klamath River                                   takes place in December and January in California
            Humboldt Bay  OO O O OC                         (Miller and Geibel 1973), and February and March in
               Eel River                                    Washington (LaRiviere et al. 1981). Females extrude
             Tomales Bay     O O                             eggs, along with a yellow secretion, directly onto the
       Cent San Fran. Bay  O O    'Indudes Central San      spawning site. The eggs adhere to the rocks and each
                                      Francjsoo, Suisun,
       South San Fran. Bay            and San Pablo bays.   other. The male then swims over the egg mass and
           Elkhom Slough     /                              fertilizes them with his milt.  The egg laying and
              Morro Bay                                    fertilization continues until the female leaves the nest
         Santa Monica Bay  i                                site (Wilby 1937). The male stays andguardsthe eggs
           San Pedro Bay  .    i                            and may fan the eggs with his pectoral fins (Garrison
            Alamitos Bay                                    and Miller 1982).  Males may be monogamous or
            Anaheim Bay       i                             polygamous and are commonly found guarding more
             Newport Bay      l                             than one egg mass (Garrison and Miller 1982). Larger
             Mission Bay      l                             fish often spawn earlier than smaller fish.
           San Diego Bay      l
           Tijuana Estuary      I                            Fecundity: From 6,000-500,000 eggs can be laid,
                       A S J  L E                           depending on the size of the female (Phillips 1959).

    to at least 61 m depth in the ocean (Miller and Geibel    Growth and Development
    1973). Thisspeciesis commonlyfoundonsteeprocky    Eaa Size and Embryonic Development: Eggs are
    reefs, near algae and seagrass beds, and in areas with    spherical and 2.8 mm in diameter when laid, and 3.5
    strong tidalcurrents. Malesareusuallyfoundinwaters    mm in diameter after fertilized and water hardened
    <185 m deep.                                             (Wilby 1937). The egg mass can be large, up to 33 kg
                                                           (Forrester 1969). Embryonic development is indirect
    Substrate: Eggs are laid in rocky crevices and    and external. Eggs hatch in about 6 weeks, with eggs
    overhangs.  Juveniles are found on sandy bottoms,    on the outside of the mass hatch first (Jewell 1968).
    while adults prefer rocky reefs or kelp beds.
                                                           Aae and Size of Larvae: Larvae are approximately 7
    Physical/Chemical Characteristics: Currents may    mm long at hatching and grow to 55 mm in length
    influence spawning site selection and eggs are usually    before metamorphosis (MBC Applied Environmental
    laid in euhaline areas having swift currents (Giorgi    Sciences 1987).
    1981, Giorgi and Congleton 1984). Juveniles are
    found in marine and mixing zones of estuaries, buttheir    Juvenile Size Rance: Juveniles grow from 5.5 to 60.0
    salinity tolerances are unknown. Adults are typically    cm long (female) or 50.0 cm long (male) in California
    found in marine waters at temperatures of 5-15�C    before reaching maturity (Millerand Geibel 1973). Fish

                                                        243

    Lingcod continued
    in more northerly populations tend to grow larger    References
    before reaching maturity.
                                                           Allen, M. J., and G. G. Smith. 1988. Atlas and
    Ane and Size of Adults: In California, most females    zoogeographyofcommonfishesintheBeringSeaand
    mature at 60.0 cm total length (TL) (3 years), and most    northeastern Pacific. NOAA Tech. Rep. NMFS 66,
    males at 50.0 cm TL (some 2 years) (Miller and Geibel    151 p.
    1973). The lingcod matures at slightly larger sizes
    north of California (Hart 1973), but grows faster in the    Bargmann, G. 1981. Management of lingcod in Puget
    southern part of their range, where both males and    Sound.  In A. Cass (chairman), Proceedings of the
    females average 50.0 cm after 3 years.  Female    February 25-26,1981 international lingcod workshop,
    lingcod can growto morethan 152 cm long (Eschmeyer    p. 103-115. Unpubl. Rep., Pacific Biol. Sta., Nanaimo,
    et al. 1983), 32 kg, and 20 years old (Miller and Geibel    B.C., Canada.
    1973). However, males usually nevergrow longerthan
    90 cm (MBC Applied Environmental Sciences 1987).    Bargmann, G. G. 1982. The biology and fisheries for
                                                           lingcod (Ophiodon elongatus) in Puget Sound. Tech.
    Food and Feeding                                        Rep. 66, Wash. Dept. Fish., Olympia, WA, 69 p.
    Troghic Mode: Larvae are carnivorous zooplanktivores.
    Juveniles and adults are carnivorous.                   Buckley, R., G. Hueckel, B. Benson, S. Quinnell, and
                                                           M. Canfield. 1984. Enhancement research on lingcod
    Food Items: Larvae eat copepods, copepod nauplii    (Ophiodon elongatus) in Puget Sound.  Prog. Rep.
    andeggs,andothercrustaceans. Small juveniles feed    216, Wash. Dept. Fish., Olympia, WA, 93 p.
    on crustaceans, but as they grow they concentrate
    their feeding on small fishes.  Adults are top-level    Cass, A. 1981. Juvenile lingcod purse seine survey
    carnivores and feed on Pacific herring (Clupea    and its application to estimate densities during pelagic
    harengus), sand lance (Ammodytes hexapterus),    development. In A. Cass (chairman), Proceedings of
    flounders, Pacific hake (Merluccius productus),    the February 25-26, 1981  international lingcod
    rockfishes (Sebastes spp.), and large crustaceans.    workshop, p. 73-102. Unpubl. Rep., Pacific Biol. Sta.,
    They are also cannibalistic (Hart 1973).  However,    Nanaimo, B.C. Canada.
    females do not eat during spawning (MBC Applied
    Environmental Sciences 1987).                           Day, M. E., C. A. Coomes, P. L. Striplin, and D. Grosse.
                                                           1986. Review and annotated bibliography of juvenile
    Biological Interactions                                 lingcod and flatfish populations inhabiting Grays Harbor
    Predation: Invertebrates (gastropods, crabs, starfishes,   with reference to potential adverse impacts caused by
    sea urchins) and vertebrates [spiny dogfish (Squalus    dredging. Final Rep. to U.S. Corps of Eng., Seattle
    acanthias) and Pacific staghorn sculpin (Leptocottus   District, Seattle, WA, 140 p.
    armatus)] prey on eggs (LaRiviere et al. 1981, MBC
    Applied Environmental Sciences 1987). Larvae and    Eschmeyer, W. N., W. S. Herald, and H. Hammann.
    juveniles are eaten by other fishes, including adult    1983. A field guide to Pacific coast fishes of North
    lingcod.  Besides humans, probably only marine    America. Houghton Mifflin Co., Boston, MA, 336 p.
    mammals and large sharks are predators on adults.
                                                           Forrester, C. R. 1969. Life history information on some
    Factors Influencina Populations: Overfishing can be a    groundfish species. Fish. Res. Board Can. Tech. Rep.
    problem because of this species' slow growth and    No. 105, 17 p.
    limited mobility (Bargmann 1982). Poor watercirculation
    reduces embryo survival (Giorgi and Congleton 1984).    Frey, H. W. 1971. California's living marine resources
    Estuarinedredging mayalter naturalopen-sand rearing    and their utilization.  Calif. Dept. Fish  Game,
    areas (Buckley et al. 1984). Predation, cannibalism,    Sacramento, CA, 148 p.
    disease, and poor larval survival may limit recruitment.
    Year-class strength apparently varies widely due to   Garrison, K. J., and B. S. Miller. 1982. Review of the
    many factors (Cass 1981, Day et al. 1986).              early life historyof Puget Sound fishes. Fish. Res. Inst.,
                                                           Univ. Wash., Seattle, WA, 729 p (FRI-UW-8216).

                                                           Giorgi, A. E. 1981. The environmental biology of the
                                                           embryos, egg masses and nesting sites of the lingcod,
                                                           Ophiodon elongatus. NWAFC Proc. Rep. 81-06, 107

                                                       244

                                                                                                Lingcodcontinued
    p. NorthwestAlaskaFish. Cent.,Natl. Mar. Fish. Serv.,    Can. Fish. Mar. Serv. Tech. Rep. No. 756, 35p.
    NOAA, 2725 Montlake Blvd. E., Seattle, WA, 98112.
                                                             Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    Giorgi, A. E., and J. L. Congleton.  1984. Effects of    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    currentvelocityondevelopmentandsurvivaloflingcod,    of common and scientific names of fishes from the
    Ophiodon elongatus, embryos. Env. Biol. Fish. 10(1/    United States and Canada. Am. Fish. Soc. Spec. Publ.
    2):15-27.                                                No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    Shaw, W. N., and T. J. Hassler. 1989. Species profiles:
    Board Can., Bull. No. 180. 740 p.                        life histories and environmental requirements of coastal
                                                            fishes and invertebrates (Pacific Northwest)-lingcod.
    Jagielo, T. 1988. The spatial, temporal, and bathymetric    U.S. Fish Wildl. Serv. Biol. Rep. 82(11.119), U.S. Army
    distribution of coastal lingcod trawl landings and effort   Corps Eng., TR EL-82-4, 10 p.
    in 1986. Prog. Rep. No. 268, Wash. Dept. Fish,
    Olympia, WA, 46 p.                                       Wilby, G. V. 1937. The lingcod, Ophiodon elongatus,
                                                            Girard. Bull. Biol. Board Can. 54:1-24.
    Jewell, E. D. 1968.  SCUBA diving observations on
    lingcod spawning at a Seattle breakwater. Wash.
    Dept. Fish., Fish Res. Pap. 3:27-34.

    LaRiviere, M. G., D. D. Jessup, and S. B. Mathews.
    1981. Lingcod, Ophiodon elongatus, spawning and
    nesting in San Juan Channel, Washington. Calif. Fish
    Game 67(4):231-239.

    MBC Applied Environmental Sciences. 1987. Ecology
    of important fisheries species offshore California. Min.
    Man. Serv., U.S. Dept. Int., Wash., D.C., 251 p.

    McLane, A. J. 1978. McClanes field guide to saltwater
    fishes of North America. Holt, Rinehart, and Winston,
    Inc., New York, NY, 283 p.

    Miller, D. J., and J. J. Geibel. 1973. Summary of blue
    rockfish and lingcod life histories; a reef ecology study;
    and giant kelp, Macrocystis pyrifera, experiments in
    Monterey Bay, California. Calif. Fish Game, Fish Bull.
    158, 137 p.

    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.
    Nelson. 1990. Distribution and abundance of fishes
    and invertebrates in west coast estuaries, Volume I:
    data summaries. ELMR Rep. No. 4. Strategic Assess-
    ment Branch, NOS/NOAA, Rockville, MD,
    240 p.

    National Marine Fisheries Service. 1986. Fisheries of
    the United States, 1985. Current Fishery Statistics No.
    8368. U.S. Dept. Comm., NOAA, Nat. Mar. Fish. Serv.,
    Nat. Fish. Stat. Prog., Washington, D.C., 122 p.

    Phillips, J. B. 1959. A review of the lingcod, Ophiodon
    elongatus. Calif. Fish Game 45(1):19-27.

    Phillips, A. C., and W. W. Barraclough. 1977. On the
    early life history of the lingcod (Ophiodon elongatus).

                                                         245

       Leptocottus armatus
       Adult

                   5cm

    Common Name: Pacific staghorn sculpin                   Within Studv Area: It occurs in all estuaries within the
    Scientific Name: Leptocottus armatus                    study area (Table 1) (Monaco et al. 1990).
    Other Common Names: staghorn sculpin, bullhead,
    cabezon, buffalo sculpin, smooth cabezon (Gates and    Life Mode
    Frey 1974)                                              Eggs are demersal, adhesive, and are probably laid in
    Classification (Robins et al. 1980)                     marine waters.  Larvae are planktonic (marine and
    Phylum: Chordata                                        estuarine), and juveniles and adults are demersal.
    Class: Osteichthyes
    Order: Perciformes                                      Habitat
    Family: Cottidae                                        Type: This is a eu ryhali ne species. Juveniles are found
                                                           in shallow water, riverine, estuarine, and marine
    Value                                                   habitats. Older and larger Pacific staghorn sculpins
    Commercial: This species has no commercial value.    reside in marine and highly saline estuarine areas
                                                           (Wydoski and Whitney 1979).
    Recreational: The Pacific staghorn sculpin is usually
    capturedincidentallywithotherfisheries, such asthose    Substrate: Newly-settled juveniles prefer clean sand
    for sturgeon  (Acipenser  spp.)  and  salmon    (Marliave 1975, cited by Garrison and Miller 1980).
    (Oncorhynchusspp.), and isthusconsidered anuisance    Older juveniles and adults are also found primarily in
    by some. It is not usually consumed by anglers, but is   sandy habitats. Planktonic larvae and benthic living
    easily captured in shallow waters and sometimes used    juveniles and adults can be found over substrates
    as bait (Reish 1968).                                   ranging from soft mud to rock (Wang 1986).

    Indicatorof Environmental Stress: Sincethis species is   Phvsical/Chemical Characteristics: The location of egg
    distributed throughout most Pacific coast estuaries    masses has not been discovered (Garrison and Miller
    and may spend its entire life within estuaries, it is a    1980).   However,  optimum  egg survival and
    target species of the National Status and Trends    development in the laboratory occurs in salinities of
    Program (Ocean Assessments Division 1984).              26%o, while best larval survival occurs in salinities of
                                                           10.2-17.6%0 (Jones 1962). Juveniles withstand larger
    Ecological:ThePacificstaghornsculpinisanimportant    fluctuations in salinity and are more tolerant of low
    predator of ghost shrimp, Callianassa californiensis   salinity than eggs, larvae, or adults (Jones 1962).
    (Posey 1986). It is a common estuarine fish that is   Small juveniles are found intertidally, while larger
    eaten by various fishes, birds, and mammals.            juveniles and adults are found subtidally. This species
                                                           is not normally found below 50 m depth. Juveniles
    Range                                                   have wide salinity and temperature tolerances,
    Overall: This species is found from southern California    withstanding salinities near 67.5%o at 25�C, 37.5%o at
    to the Gulf of Alaska (Wydoski and Whitney 1979).       290C, and 0.0%o at 10�C (Morris 1960).

                                                       246

                                                                                Pacific staghorn sculpin continued

                                                           Matino/Soawnina: Spawning occurs from October to
     Table 1. Relative abundance of Pacific staghorn        March or April, peaking in January and February (Jones
              sculpin in 32 U.S. Pacific coast estuaries.   1962, Wang 1986).
                        Life Stage
           Estuary    A S J L E                            Fecundit: Fecundity averages 5,000 eggs per female
      I     Puget Sound (3 0  3 0 0   Relative abundance:   (Jones 1962) and ranges from 2,000-1 1,000 eggs per
             Hood Canal   ] 0 CO   0 0    �   Highly abundant  female (Moyle 1976).
             Skagit Bay (C 0 i ) 0 0i     Abundant
           Grays Harbor C} o i10  b   0   Common           Growth and Development
             Willapa Bay C) 0 I    N C 0    i   Rare        Eaa Size and Embryonic DeveloDment: Eggs are 1.36-
          Columbia River  3 0 � C0 C0   Blank Not present  1.50 mm in diameter (average 1.43 mm). Embryonic
           Nehalem Bay C    ai 0                           development is indirect and external. Eggs hatch in 9-
           Tillamook Bay  O o O  0    Life stage:          14 days after fertilization at 15�50C.
             NetartsBay i  06  a       A-Adults
                                     S - Spawning adults
             Siletz River CO 0 a 0 O    J -Juveniles       Aoe and Size of Larvae: At hatching, larvae range from
            YaquinaBay :� O �:O C    L-Larvae              3.9-4.8 mm  total length (TL) (Jones  1962).
             AlseaRiver � O  D O       O     Eggs          Metamorphosis to juvenile begins after about 2months,
           Siuslaw River * O �  O O                        when larvae are 15-20 mm standard length (Matarese
           Umpqua River Q O �      C0  et al. 1989).
              Coos Bay �        0 �  0 O
            Rogue River ( 0 ( O O0                         Juvenile Size Ranae: The juvenile size range is from
           KlamathRiver      O O C  0O                     about 20 mm to approximately 120 mm TL (Jones
           Humboldt Bay a  0 O     0 0  1962).
               Eel River 0     C (C  00
            Tomales Bay  ] O  IC ]   0                      Aae and Size of Adults: The Pacific staghorn sculpin
      CentSanFran.Bay* (6 O �    * IncludesCentral San    matures in 1 year and usually >12.0 cm TL.  This
                                     Francisco, Suisun,
      South San Fran. Bay  ( 3  �   Frand San Pablo bays,  species can live as long as 3 years and grow to 20.3 cm
          Elkhorn Slough O     �e                          in length in California (Jones 1962), and upto 10 years
              Morro Bay CO    � C                           and 22.9 cm in length in Washington (Wydoski and
        Santa Monica Bay  i,1 4 4 4                        Whitney 1979).
          San Pedro Bay  i    4
            Alamitos Bay O CO Ci    O                       Food and Feeding
           Anaheim Bay 0 0  OC  0                          Trophic Mode: Larvae are planktivorous, while juveniles
            Newport Bay     V                              and adults are carnivorous.
             Mission Bay       0
          SanDiego Bay O    i O                            Food Items: Juveniles feed primarily on benthic and
          Tijuana Estuary O    O O                         epibenthic organisms, including the amphipod
                      A S J L E                            Corophium   spp., other gammarids, decapod
                                                           crustaceans, and the polychaete Neanthes spp. Large
    Miorations and Movements: Although no true "migration"   juveniles and adults consume fish and large crustaceans
    exists, the Pacific staghorn sculpin shows seasonal    (Crangon  spp.) (Jones 1962, Tasto 1975, Conley
    movements within estuaries. Smalljuveniles settle-out    1977, Smith 1980, Posey 1986).
    in the lower marine areas of estuaries in winter and
    then move up into freshwater areas in spring and early    Biological Interactions
    summer (Conley 1977). There is a tendency to move    Predation: This species is eaten by large fishes, ducks,
    down into estuarine and then marine waters as they    loons (Gavia spp.), cormorants (Phalacrocorax spp.),
    grow (Jones 1962). After spawning, adults may leave   gulls, and marine mammals (Tasto 1975, Treacy 1984).
    shallowspawninggroundsand movetodeeperoffshore    To reduce predation, the Pacific staghorn sculpin will
    waters (Tasto 1975). However, many appearto spend    try to partially bury itself in the sediment. It will also
    their entire life in estuaries.                         erect its opercular spines laterally with the sharp
                                                           recurved hooks facing upward to deter predators (Tasto
    Reproduction                                            1975).
    Mode: The Pacific staghorn sculpin is gonochoristic,
    oviparous, and iteroparous; eggs are fertilized    Factors Influencina Ponulations: Larval success
    externally.                                             probably determines overall recruitment. Newly-settling
                                                           juveniles use shallow tidal flats and pools, hence

                                                       247

    Pacific staghorn sculpin continued
    destruction of this habitat will affect this life stage. The    Mar. Biol. Ecol. 103:143-161.
    Pacific staghorn sculpin may compete with the
    introduced yellowfin goby (Acanthogobius flavimanus)    Reish, D. J. 1968. Marine life of Alamitos Bay. Forty-
    in estuaries where both species exist (Usui 1981).     Niner Shops, Inc., Long Beach, CA, 92 p.

    References                                              Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
                                                           E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    Conley, R. L. 1977. Distribution, relative abundance,    of common and scientific names of fishes from the
    and feeding habits of marine and juvenile anadromous    United States and Canada. Am. Fish. Soc. Spec. Publ.
    fishes of Everett Bay, Washington. M.S. Thesis, Univ.    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    Wash., Seattle, WA, 57 p.
                                                           Smith, J. E. 1980. Seasonality, spatial dispersion
    Garrison, K. J., and B. S. Miller. 1980. Review of the    patterns and migration of benthic invertebrates in an
    earlylifehistoryofPugetSoundfishes. Fish. Res. Inst.,   intertidal marsh-sandflat system of Puget Sound,
    Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).        Washington, and their relation to waterfowl foraging
                                                           and the feeding ecologyof staghorn sculpin, Leptocottus
    Gates, D. E., and H. W. Frey.  1974.  Designated    armatus. Ph.D. Thesis, Univ. Wash., Seattle, WA,
    common names of certain marine organisms of    176p.
    California. Cal. Fish Game, Fish Bull. 161:55-90.
                                                           Tasto, R. N. 1975. Aspects of the biology of Pacific
    Jones, A. C. 1962. The biology of the euryhaline fish   staghorn sculpin, Leptocottus armatus  (Girard), in
    Leptocottus armatus armatus Girard (Cottidae). Univ.    Anaheim Bay. In E. E. Lane and C. W. Hill (editors), The
    Calif. Publ. Zool. 67(4):321-368.                       marine resources of Anaheim Bay. Calif. Fish. Game,
                                                           Fish Bull. 165:123-135.
    Marliave, J. B. 1975. The behavioral transformation
    from the planktonic larval stage of some marine fishes    Treacy, S. 1984. Marine mammals of the Columbia
    reared in the laboratory. Ph. D. Thesis, Univ. British    River estuary.   Columbia  River Estuary  Data
    Columbia, Vancouver, B.C., Canada, 231 p.               Development Program, CREST, Astoria, OR, 62 p.
                                                           plus appendices.
    Matarese, A. C., A. W. Kendall, Jr., D. M. Blood, and B.
    M. Vinter. 1989. Laboratory guide to early life history   Usui, C. A. 1981. Behavioral, metabolic, and seasonal
    stages of Northeast Pacific fishes. NOAA Tech. Rep.,    size comparisons of an introduced gobiid fish,
    NMFS80, 652p.                                           Acanthogobius flavimanus  and a native cottid,
                                                           Leptocottus armatus, from upper Newport Bay,
    Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.    California. M.S. Thesis, Cal. State Univ, Fullerton, CA,
    Nelson. 1990. Distribution and abundance of fishes    52 p.
    and invertebrates in west coast estuaries, Volume I:
    data summaries. ELMR Rep. No. 4. Strategic Assess-    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    ment Branch, NOS/NOAA, Rockville, MD,                   Joaquin estuary and adjacent waters, California: A
    240 p.                                                  guide to the early life histories.  Tech. Rep. No. 9.
                                                           Interagency ecological study program for the
    Morris, R.W. 1960. Temperature, salinity, and southern    Sacramento-San Joaquin estuary. Calif. Dept. Water
    limits of three species of Pacific cottid fishes. Limnol.    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    Oceanogr. 5(2):175-179                                  and U.S. Fish Wildl. Serv., various pagination.

    Moyle, P. B. 1976. Inland fishes of California. Univ.    Wydoski, R.S.,and R. R.Whitney. 1979. Inland fishes
    Calif. Press, Berkeley, CA, 405 p.                      of Washington.  Univ. Wash. Press, Seattle, WA,
                                                           220 p.
    Ocean Assessments Division 1984. The national
    status and trends program for marine environmental
    quality: program description (mimeo). Ocean Assess.
    Div., Nat. Ocean Surv., Nat. Ocean. Atm. Adm.,
    Rockville, MD, 28 p.

    Posey, M. H. 1986. Predation on a burrowing shrimp:
    distribution and community consequences. J. Exp.

                                                        248

    249

      Paralichthys californicus
      Adult

                     10cm

    Common Name: California halibut                       (Reed and MacCall 1988).  Incidental catches of
    Scientific Name: Paralichthys californicus            seabirds in gill nets set for California halibut and white
    Other Common Names: Monterey halibut, bastard   croaker (Genyonemus lineatus) are a problem.
    halibut, chicken halibut, southern halibut, alabato
    (Ginsburg 1952, Roedel 1953)                          Recreational:The California halibut is a highly desirable
    Classification (Robins et al. 1980)                   species because of its excellent taste and large size
    Phylum: Chordata                                      (Frey 1971). Over 916,000 were caught by anglers in
    Class: Osteichthyes                                   1985 (U.S. Department of Commerce 1986). Average
    Order: Pleuronectiformes                              size caught is 2.7-3.2 kg, but pier-caught fish are
    Family: Bothidae                                      usually much smaller (Squire and Smith 1977). This
                                                          species is rarely caught in waters >18.3-27.4 m deep
    Value                                                 (Squire and Smith 1977). From Morro Bay to Tomales
    Commercial: The California halibut is commercially    Bay, California, fishing is best from summerto early fall
    fished from Eureka to San Diego, California, withmost    (Squire and Smith 1977).  This species is caught
    caught between San Francisco and San Diego (MBC    primarily from piers and boats using hook and line and
    Applied Environmental Sciences 1987). The center of   live bait (Roedel 1953). In California, only fish >56 cm
    the fishery was originally southern California to Baja    long are legal to keep (Reed and MacCall 1988);
    California, but it has shifted northward (Frey 1971).    anglers are allowed totake 5/day except in the Bodega
    This species is harvested by set gill net, trammel net,   and Tomales Bay areas (California Department of Fish
    and trawl nets (Schultze 1986). Fish must be >56 cm    and Game 1987)
    or at least 1.8 kg (in round) or 1.6 kg dressed weight.
    Moreover, no morethan 4 less than 56cm in length can    Indicator of Environmental Stress: The size and health
    be kept for noncommercial uses when caught    of California halibut populations probably reflects the
    incidentally in trawls.  Open season for California   health of southern California shallow waters because
    halibuttrawlinggrounds(Point ArguellotoPoint Magu)    this species depends on these areas for its early life
    isJune 16through March 14 (Schultze 1986). California   stages (see "Factors Influencing Populations").
    fisherman landed an average of 534 t per year from
    1983 to 1987, receiving $0.64-1.59/kg in 1987    Ecological: This is the largest Paralichthys species in
    (California Departmentof Fish andGame 1988). Since    U.S. waters (Ginsburg 1952). It is common along
    1973, catches have steadily increased (California   sandy nearshoreareas and atoppredatorin nearshore
    Department of Fish and Game 1988). In 1987, most    sandy bottom environments in southern California.
    were caught in March and the fewest in September
    (California Department of Fish and Game 1988).    Range
    Mexican catches are highest during summer and fall   Overall: The California halibut's overall range is from
    (Roedel 1953).  The commercial fishery is biased    Magdalena Bay, Baja California, to the Quillayute
    toward females because they grow faster than males    River, Washington; an isolated population exists in the

                                                       250

                                                                                      California halibut continued

                                                           Juveniles and adults are benthicordemersal, however
      Table 1. Relative abundance of California halibut    they often will pursue food well off the bottom (Frey
               in 32 U.S. Pacific coast estuaries.         1971). Eggs occur primarily between the 6 and 20 m
                         Life Stage                        isobaths; larvae between 12 and 45 m isobaths (Haaker
            Estuary    A S J L E                           1975, Plummeretal. 1983). Small juveniles are found
             PugetSound              Relative abundance:   primarily in coastal embayments and estuaries, but
              Hood Canal               �   Highly abundant    they also occur in very shallow open coastal waters
              Skagit Bay              0   Abundant        (Clark 1930, Fierstine et al. 1973, Haaker 1975, Barry
            Grays Harbor              O   Common          and Cailliet 1981, Horn and Allen 1981, Plummer et al.
              Willapa Bay              V   Rare            1983, Noah 1985, Kramer and Hunter 1987, 1988).
                                      Blank Not present
           Columbia River
            Nehalem Bay                                   Habitat
            Tillamook Bay             Life stage:          Type: Eggs and larvae are found primarily along a
              Netarts Bay             A - Adults           shallow water "band" in nearshore open coastal waters
                                      S - Spawning adults
              Siletz River            J -Juveniles         (Ahlstrom and Moser 1975). Larvae <10 mm long are
             Yaquina Bay              L - Larvae           found throughout the water column, primarily between
              Alsea River               Eggs               the 12 and 45 m isobaths and within 2-5 km of shore
            Siuslaw River                                 (Barnett et al. 1984). Larvae are found in bays and
            Umpqua River                                   estuaries, but are not abundant there (Leithiser 1977,
               Coos Bay                                    McGowen 1977, Nordby 1982, Wang 1986). Small
             Rogue River                                  juveniles are found just outside the surf zone and in
            Klamath River                                  estuaries and bays (Haaker 1975, Plummer et al. 1983,
            Humboldt Bay                                   Kramer and Hunter 1987, 1988).  Adults and older
               EelRiver                                   juveniles occur nearshore, with larger and older
             Tomales Bay    O O                            individuals occurring deeper (to about 60 m depth)
       Cent San Fran. Bay     0    Indudes Central San    (Haaker 1975, Plummer et al. 1983).  Adults are
                                      Francisco. Suisun,
       South San Fran. Bay  "   0 0  and San Pablo bays.  normally found at 6-40 m depths (Ginsburg 1952), but
           Elkhorn Slough  i1   O  H                      can be found to 183 m (Eschmeyer et al. 1983). Adults
              Morro Bay  v   O  'I                        may be abundant in the surf zone during the spring as
         Santa Monica Bay    O O   0                       they prey on spawning California grunion (Leuresthes
           San Pedro Bay CO 0     0C tenuis) (Fitch 1958).
             Alamitos Bay   (
            Anaheim Bay O    )                            Substrate: Juveniles and adults prefer sandy bottoms
             Newport Bay i 0 o                             (Eschmeyer et al. 1983), but are also common near
             Mission Bay CO    t                          rocks, sand dollar beds, and in channels entering
           San Diego Bay 0   )                            coastal embayments (Fitch and Lavenberg 1971).
           Tijuana Estuary O    J
                       A S J L E                           Phvsical/Chemical Characteristics: The California
                                                           halibut is found in watertemperatures of 10-25�C, with
    Gulf of California (Ginsburg 1952, Miller and Lea 1972,    a preference for 20.8�C (Ehrlich et al. 1979). Young
    Eschmeyer et al. 1983, Allen et al. in prep.).         halibut (subyearlings and yearlings) are eurythermal,
                                                           but older halibut appearto be stenothermal (Kucas and
    Within Studv Area: This species is common in all bays    Hassler 1986). Eggs, larvae, and adults are found in
    and estuaries south of Tomales Bay, California, and    euhalinewaters, butjuvenilesoftenoccurinoligohaline
    abundant in most estuaries south of Point Conception.    to euhaline conditions (Haaker 1975, Allen et al. in
    It is rare or absent in estuaries north of Tomales Bay    prep.). Juveniles are relatively tolerant of reduced
    (Table 1) (Chapman 1963, Bane 1968, Bane and Bane    dissolved oxygen and increased water temperatures
    1971, Millerand Lea 1972, Fierstine et al. 1973, Haaker    (Waggoner and Feldmeth 1971).
    1975, Cailliet et al. 1977, Horn and Allen 1981, Lockheed
    Ocean Science Laboratories 1983, Wang 1986).           Miarations and Movements: Larvae occur in a coastal
                                                           band from San Francisco to southern Baja California
    Life Mode                                              (Ahlstrom and Moser 1975). They apparently settle out
    Eggs and larvae are pelagic (Ahlstrom and Moser    in shallow water areas on the open coast and also in
    1975, Ahlstrom et al. 1984). Larvae are most abundant    bays and estuaries, placing the newly-settled juveniles
    in coastal waters during March through September    in or near their rearing habitat (Frey 1971, Haaker
    (Ahlstrom and Moser 1975, Walker et al. 1987).    1975, Plummer et al. 1983, Kramer and Hunter 1988).

                                                       251

    California halibut continued
    Primary settlement times are from February to August    maturing at 37.5 cm (4-6 years) (Roedel 1953, Fitch
    (Kramer and Hunter 1988). Juveniles reside in bays    and Lavenberg 1971, Frey 1971, Haaker 1975). This
    and estuaries for about 2-3 years and then emigrate    species is estimated to grow 3.8-8.8 cm/year and live
    out to shallow open coastal waters. Males are about 20    to 30 years, with females growing faster and larger than
    cm and females 25 cm in length when they migrate    males (Frey 1971, Haaker 1975, MBC  Applied
    (Haaker 1975). Subadults and adults generally show    Environmental Sciences 1987, Reed and MacCall
    very limited along-shore movements (Ginsburg 1952,    1988). The largest California halibut reported was 1.5
    Haaker 1975); onlya few individuals have shown large    m total length (TL) and 33.6 kg (Miller and Lea 1972,
    migrations (Fitch and Lavenberg 1971). Adults move    Squire and Smith 1977, MBC Applied Environmental
    into shallow coastal waters (4-6 m deep) in early spring    Sciences 1987).
    to spawn (Ginsburg 1952, Haaker 1975). Juveniles
    and adults lie partially buried in the sediments when    Food and Feeding
    inactive (Allen 1982).                                 Trophic Mode: Larvae, juveniles, and adults are
                                                          carnivorous, probably feeding primarily during the
    Reproduction                                           daytime. Initially, the California halibut feeds on small
    Mode: The California halibut is gonochoristic, oviparous,    invertebrates, then switches to feed almost exclusively
    and iteroparous. It is a broadcast spawner and eggs    on fish as it grows (Haaker 1975). This species is an
    are fertilized externally.                             ambush feeder that locates prey by sight and possibly
                                                          via the lateral line (Haaker 1975, Allen 1982, Hobson
    Matina/SDawnina: From larval abundance information    and Chess 1987).
    it appears that some spawning may occuryear-round,
    with most spawning from January to August (Ahlstrom    Food Items: Larvae most likely feed on plankton. Small
    and Moser 1975, Wang 1986). In southern California,   juveniles feed on crustaceans (mysids, shrimp,
    spawning occurs from February to July, peaking in   gammarid amphipods, harpacticoidcopepods), squids,
    May. The actual depth of spawning is uncertain (Allen   octopus, and fish (gobies, killifish, and others). Large
    1988), but is known to occur over sandy substrates    juveniles and adults consume primarily fish (Haaker
    (Ginsburg 1952, Frey 1971, Feder et al. 1974, Haaker    1975, Allen 1982, Roberts et al. 1982, Plummer et al.
    1975 ). Successful spawning likely occurs along the    1983, Allen 1988) and the northern anchovy (Engraulis
    coastal zone from San Francisco Bay to Magdalena    mordax) is the most common fish eaten. Other fishes
    Bay, California, and probably in the Gulf of California    eaten by the California halibut include sardines,
    (Ahlstrom and Moser 1975, MBC Applied Environmental    atherinids, sciaenids, gobies, embiotocids, and other
    Sciences 1987).                                         flatfishes (Quast 1968, Allen 1982).  Arrow gobies
                                                           (Clevelandia ios) are particularly important prey for
    Fecundity: Unknown.                                    juvenile halibut rearing in estuaries and bays (Haaker
                                                           1975).
    Growth and Development
    Eaa Size and Embrvonic Develooment: California    Biological Interactions
    halibut eggs are 0.74-0.84 mm in diameter (Ahlstrom et    Predation: Sea lions eat California halibut caught in
    al. 1984).  Embryonic development is indirect and    trammel nets (Fitch and Lavenberg 1971).  Other
    external; eggs hatch approximately 2 days after   predators include Pacific angel shark (Squatina
    fertilization at 16�C.                                  californica), Pacific electric ray (Torpedo californica),
                                                           large California halibut, and bottlenose dolphin (Tursiops
    Aae and Size of Larvae: Larvae are 2.0 mm long at   truncatus) (Fitch and Lavenberg 1971, Frey 1971,
    hatching (Ahlstrom and Moser 1975, Ahlstrom et al.   Feder et al. 1974).  Parasites (both external and
    1984). The yolk-sac is depleted about 6 days after   internal) commonlyattackthis species; infestation rates
    hatching (Gadomski and Petersen 1988).  Time to   increase with age and size of fish (Haaker 1975).
    settlement is 5-6 weeks at 16�C (Gadomski and    Parasites include isopods, copepods, nematodes,
    Petersen 1988), or 20-29 days at 18.3-21.9�C (Allen   trematodes, and cestodes (Haaker 1975).
    1982). Metamorphosis occurs at a length of 7.5-
    9.4 mm.                                                 Factors Influencina Populations: Although landings
                                                           have increased since 1972, historical records indicate
    Juvenile Size Ranae: Juveniles range in length from    an overall decline in thepopulationof California halibut
    0.8-43.0 cm.                                            (Plummer et al. 1983).  Landings have fluctuated
                                                           widely, but are presently about 25% of those of 1920
    Ace and Size of Adults: Some males mature as small    (Frey 1971, MBC Applied Environmental Sciences
    as 20 cm in length (2-3 years), while females begin    1987). Thepopulationdeclinemaybearesultof large-

                                                       252

                                                                                      California halibut continued
    scale changes in the marine environment, overfishing,    Barnett, A.M., A. E. Hahn, P. D. Sertic, and W. Watson.
    alterations and destruction of estuarine habitat, or a    1984. Distribution of ichthyoplankton off San Onofre,
    shift in population centers (Plummer et al. 1983).    California, and methods for sampling shallow coastal
    Pollution, (e.g., watersoluble fractions of crude oil) can    waters. Fish. Bull., U.S. 82(1):97-111.
    reduce hatching success, reduce size of larvae at
    hatching, produce morphological and anatomical    Barry, J. P., and G. M. Cailliet. 1981. The utilization of
    abnormalities, and reduce feeding and growth rates    shallow marsh habitats by commercially important
    (MBC Applied EnvironmentalSciences 1987). Initiation   fishes in Elkhorn Slough, California. Cal.-Nev. Wildl.
    of feeding by larvae appears critical for larval survival    Trans. 1981:38-47.
    (Gadomski and Petersen 1988). Natural production
    has recently been augmented by hatchery production    Cailliet, G. M., B. Antrim, D. Ambrose, S. Pace, and M.
    (Crooke and Taucher 1988).  Substantial genetic    Stevenson. 1977. Species composition, abundance
    variation between two populations of California halibut    and ecological studies of fishes, larval fishes, and
    in the southern California Bight, suggests that the    zooplankton in Elkhorn Slough.   In Ecologic and
    natural population is subdivided (Hedgecock and Bartley    hydrographic studies of Elkhorn Slough, Moss Landing
    1988).   Wide fluctuations in young-of-the-year    Harbor and nearshore coastal waters, p. 216-386.
    recruitmentexist, but noexact causehasbeenidentified    Moss Landing Marine Lab., Moss Landing, CA.
    (Allen 1988). Southern California estuaries and
    protected shallow water habitats play a critical role in   California Department of Fish and Game. 1987. 1987
    the life history of this species.                        California sport fishing regulations. Calif. Dept. Fish
                                                           Game, Sacramento, CA, 12 p.
    References
                                                           California Department of Fish and Game. 1988. Review
    Ahlstrom, E. H., and H. G. Moser. 1975. Distributional    of some California fisheries for 1987.  Calif. Coop.
    atlas of fish larvae in the California current region:    Ocean. Fish. Invest. Rep. 29:11-20.
    flatfishes, 1955 through 1960. Calif. Coop. Ocean.
    Fish. Invest., Atlas No. 23., 207 p.                     Chapman, G. A.  1963.  Mission Bay, a review of
                                                           previous studies and the status of the sport fishery.
    Ahlstrom, E. H., K. Amaoka, D. A. Hensley, H. G.    Calif. Fish Game 49(1):30-43.
    Moser, and B. Y. Sumida. 1984. Pleuronectiformes;
    development. In H. G. Moser (chief editor), Ontogeny    Clark, G. H. 1930. California halibut. Calif. Fish Game
    and systematics of fishes, p. 640-670. Allen Press,    16(4):315-317.
    Inc., Lawrence, KS.
                                                           Crooke, S., and C. Taucher. 1988. Ocean hatcheries
    Allen, L. G.  1988.  Recruitment, distribution, and    - wave of the future? Outdoor Calif. 49(3):10-13.
    feeding habits of young-of-the-year California halibut
    (Paralichthys californicus) in the vicinity of Alamitos    Ehrlich, K. F., J. H. Hood, S. Muszynski, and G. E.
    Bay-Long Beach Harbor, California, 1983-1985. Bull.    McGowen.  1979.  Thermal behavior responses of
    So. Calif. Acad. Sci. 87(1):19-30.                       selected California littoral fishes.  Fish. Bull., U.S.
                                                           76(4) :837-849.
    Allen, M. J. 1982. Functional structure of soft-bottom
    fishcommunitiesofthesouthernCaliforniashelf. Ph.D.    Eschmeyer, W. N., E. S. Herald, and H. Hammann.
    Diss., Univ. Calif., San Diego, CA, 577 p.               1983. A field guide to Pacific coast fishes of North
                                                           America. Houghton Mifflin Co., Boston, MA, 336 p.
    Allen, M. J., R. J. Wolotira, Jr., T. M. Sample, S. F. Noel,
    and C. R. Iten.  In prep.  Living resources of the    Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    northeastern Pacific. Unpubl. mansc., Alaska Fish.    Observations on fishes associated with kelp beds in
    Cent., Nat. Mar. Fish. Serv., NOAA, Seattle, WA.         southern California. Calif. Fish Game, Fish Bull. 160:1-
                                                           144.
    Bane, G. W. 1968. Fishes of the upper Newport Bay.
    Univ. Calif. Irvine Res. Ser. 3:1-114.                   Fierstine, H. L., K. F. Kline, and G. R. Garman. 1973.
                                                           Fishes collected in Morro Bay, California between
    Bane, G. W., and A. W. Bane.  1971. Bay fishes of    January, 1968 and December, 1970. Calif. Fish Game
    northern California with emphasis on the Bodega    59(1):73-88.
    Tomales Bay area. Mariscos Publ., Hampton Bays,
    NY, 143 p.

                                                       253

    California halibut continued
    Fitch, J. E. 1958. Offshore fishes of California. Calif.   Leithiser, R. M. 1977. The seasonal abundance and
    Dept. Fish Game, Sacramento, CA, 80 p.                   distribution of larval fishes in Anaheim Bay, California.
                                                           M.S. Thesis, Calif. State Univ., Long Beach, CA, 132 p.
    Fitch, J. E., and R. J. Lavenberg. 1971. Marine food
    and game fishes of California. Calif. Nat. Hist. Guide    Lockheed Ocean Science Laboratories.   1983.
    28, Univ. Calif. Press, Berkeley, CA, 179 p.             Distribution and abundance of fishes in central San
                                                           Diego Bay, California: a study of fish habitat utilization.
    Frey, H. W. 1971. California's living marine resources    Rep. to Dept. of Navy, San Diego, CA.  38 p. plus
    and their utilization.  Calif. Dept. Fish Game,    appendices (Contract No. N62474-82-C-1068).
    Sacramento, CA, 148 p.
                                                           MBCApplied Environmental Sciences. 1987. Ecology
    Gadomski, D. M., and J. H. Petersen. 1988. Effects of   of important fisheries species offshore California. Rep.
    food deprivation on the larvae of two flatfishes. Mar.    to Min. Manag. Serv., U.S. Dept. Int., Washington,
    Ecol. Prog. Ser. 44:103-111.                             D.C., 251 p. (Contract No. MMS 14-12-0001-30294).

    Ginsburg, I. 1952. Floundersofthegenus Paralichthys    McGowen, G. E. 1977. Ichthyoplankton populations in
    and related genera in American waters. Fish. Bull.,    southSanDiegoBayandrelatedeffectsofanelectricity
    U.S. 71:1-351.                                           generating station.  M.S. Thesis, San Diego State
                                                            Univ., San Diego, CA, 88 p.
    Haaker, P. L.  1975.  The biology of the California
    halibut, Paralichthys californicus (Ayres), in Anaheim    Miller, D. J., and R. N. Lea. 1972. Guide to the coastal
    Bay, California. In E. D. Lane and C. W. Hill (editors),    marine fishes of California. Calif. Fish Game, Fish Bull.
    The marine resources of Anaheim Bay.  Calif. Fish    157, 235 p.
    Game, Fish Bull. 165:137-151.
                                                            Noah, M. D. 1985. Structure, abundance and
    Hedgecock, D., and D. M. Bartley. 1988. Allozyme    distribution of the fish and  macroinvertebrate
    variation in the California halibut, Paralichthys    communitiesinhabitingMissionBay, California between
    californicus. Calif. Fish Game 74(2):119-127.            November 1979 and February 1981. Appendix A. In
                                                            San Diego Riverand Mission Bay improvements, Draft
    Hobson, E. S., and J. R. Chess. 1987. Relationships    suppl. environ. assess., U.S. Army Corps Eng., Los
    among fishes and their prey in a nearshore sand    Angeles, CA. 37 p.
    community off southern California. Env. Biol. Fish.
    17(3):201-226.                                           Nordby, C. S.  1982.  A comparative ecology of
                                                            ichthyoplankton within Tijuana estuary and in adjacent
    Horn, M. H., and L. G. Allen. 1981. Ecology of fishes    nearshorewaters. M.S. Thesis, San Diego State Univ.,
    in upper Newport Bay, California: seasonal dynamics    San Diego, CA, 101 p.
    and community structure. Mar. Res. Tech. Rep. No.
    45, Calif. Fish Game, Long Beach, CA, 102 p.             Plummer, K. M., E. E. DeMartini, and D. A. Roberts.
                                                            1983. The feeding habits and distribution of juvenile-
    Kramer, S. H., and J. R. Hunter.  1987.  Southern    small adult California halibut (Paralichthyscalifornicus)
    California wetland/shallow water habitat investigation.    in coastal waters off northern San Diego county. Calif.
    Ann. Rep., Nat. Mar. Fish. Serv., Southw. Fish. Cent.,    Coop. Ocean. Fish. Invest. Rep. 24:194-201.
    La Jolla, CA, 12 p.
                                                            Quast, J. C. 1968. Observations on the food of the
    Kramer, S. H., and J. R. Hunter.  1988.  Southern    kelp-bed fishes.  In W. J. North and C. L. Hubbs
    California wetland/shallow water habitat investigation.    (editors), Utilization of kelp-bed resources in southern
    Ann. Rep., Nat. Mar. Fish. Serv., Southw. Fish. Cent.,    California. Calif. Fish Game, Fish. Bull. 139:109-142.
    La Jolla, CA, 15 p.
                                                            Reed, R. J., and A. D. MacCall. 1988. Changing the
    Kucas, S. T., and T. J. Hassler. 1986. Speciesprofiles:    size limit: how it could affect California halibut fisheries.
    life histories and environmental requirementsof coastal    Calif. Coop. Ocean. Fish. Invest. Rep. 29:158-166.
    fishes and invertebrates (Pacific Southwest)-California
    halibut. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.44).    Roberts, D., E. DeMartini, C. Engel, and K. Plummer.
    U.S. Army Corps Eng., TR EL-82-4, 8 p.                   1982.  A preliminary evaluation of prey selection by
                                                            juvenile-small adult California halibut (Paralichthys

                                                        254

                                                                                      California halibut continued
    californicus) in nearshore coastal waters off southern
    California. In G. M. Cailliet and C. A. Simenstad
    (editors), Gutshop 81, fish food habits studies,
    Proceedings of the third Pacific workshop, p. 214-223.
    Wash. Sea Grant Publ., Seattle, WA.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Roedel, P. M. 1953. Common ocean fishes of the
    California coast. Calif. Fish Game, Fish Bull. 91,
    184 p.

    Schultze, D. L. 1986. Digest of California commercial
    fish laws, January 1, 1986. Calif. Dept. Fish Game,
    Sacramento, CA, 40 p.

    Squire, J. L., Jr., and S. E. Smith. 1977. Anglers'guide
    tothe United States Pacificcoast. NOAA, Seattle, WA,
    139 p.

    U.S. Department of Commerce. 1986. Marine
    recreational fishery statistics survey, Pacific coast.
    U.S. Dept. Comm., Nat. Ocean. Atm. Adm., Current
    Fish. Stat. No. 8328,109 p.

    Waggoner, J. P., III, and C. R. Feldmeth. 1971.
    Sequential mortality of the fish fauna impounded in
    construction of a marina at Dana Point, CA. Calif. Fish
    Game 57(3):167-176.

    Walker, H. J., Jr., W. Watson, and A. Barnett. 1987.
    Seasonal occurrence of larval fishes in the nearshore
    southern California Bight off San Onofre, California.
    Est. Coast. Shelf Sci. 25:91-109.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: a
    guide to the early life histories. Tech. Rep. No. 9,
    prepared forthe interagency ecological study program
    for the Sacramento-San Joaquin estuary. Calif. Dept.
    Water Res, Calif. Dept. Fish Game, U.S. Bureau
    Reclam., and U.S. Fish Wildl. Serv., various pagination.

                                                       255

       Hypsopsetta guttulata
       Adult

                              5 cm

    Common Name: diamond turbot                            There is also an isolated population in the Gulf of
    Scientific Name: Hypsopsetta guttulata                  California (Miller and Lea 1972).
    Other Common Names: diamond flounder, turbot,
    halibut, sole (Gates and Frey 1974)                    Within Study Area:This species is common to abundant
    Classification (Robins et al. 1980)                     in nearshore coastal bays and estuaries from the
    Phylum: Chordata                                       Tijuana estuary to Tomales Bay, California (Table 1)
    Class: Osteichthyes                                     (Chapman 1963, Aplin 1967, Bane and Bane 1971,
    Order: Pleuronectiformes                                Fierstine et al. 1973, Lane 1975, Allen 1976, Cailliet et
    Family: Pleuronectidae                                 al. 1977, Horn and Allen 1981, Noah 1985, Zedler and
                                                           Nordby 1986). It is also found adjacent to kelp beds
    Value                                                   (usually buried in sand or near solid objects) between
    Commercial: The diamond turbot is of little commercial    the 1.2-18.2 m isobaths (Feder et al. 1974).
    value because of its small size. It is usually included
    with otherturbots when reporting catch (Baxter 1960,    Life Mode
    Bane and Bane 1971). It has a slight iodine flavor, but    Eggs and larvae are pelagic (McGowen 1977, Wang
    is excellent eating (Baxter 1960, Feder et al. 1974).   1986). Juveniles and adults are benthic or demersal
                                                           (Lane 1975).
    Recreational: The average weight of a sport-caught
    fish is 0.6 kg. It is caught year-round, with bays and    Habitat
    estuaries (e.g., Newport and Mission Bays in California)    Type: Eggs and larvae occur in estuaries (Eldridge
    providing the best fishing (Squire and Smith 1977).     1977, McGowen 1977, Wang 1986) and shallowcoastal
                                                           waters, usually within 2 km of shore (Barnett et al.
    Indicatorof Environmental Stress:This species appears    1984). Juveniles and adu Its are found in bays, estuaries
    to be dependent on bays and estuaries, thus population    and sloughs, and nearshore coastal waters down to
    sizes and fish health may reflect the condition of these    152.4 m, but prefer depths <4.6 m (Roedel, 1953,
    systems. It is a target species of the National Status    Miller and Lea 1972, Fitch and Lavenberg 1975, Squire
    and Trends Program (Ocean Assessments Division    and Smith 1977, Eschmeyer et al. 1983).
    1984).
                                                           Substrate: Eggs and larvae are found over various
    Ecological: The diamond turbot is often the dominant    substrates, and juveniles and adults are found on sand
    flatfish in southern California bays and estuaries (Lane    and mud bottoms (Federet al. 1974, Lane 1975, Squire
    1975).                                                  and Smith 1977).

    Range                                                   Phvsical/Chemical Characteristics: Eggs and larvae
    Overall: The diamond turbot is found from Magdalena    are found in euhaline-polyhaline waters, while juveniles
    Bay, Baja California to Cape Mendocino, California.    and adult occur in euhaline-mesohaline conditions.

                                                        256

                                                                                      Diamond turbot continued
                                                          Reproduction
     Table 1. Relative abundance of diamond turbot       Mode: The diamond turbot is gonochoristic, oviparous,
              in 32 U.S. Pacific coast estuaries.        and iteroparous; eggs are fertilized externally. It is
                         Life Stage                       probably a broadcast spawner.
            Estuary    A S J L E
             Puget Sound             R                    Matina/Soawnina: Larval distributions and abundances
                                      Relative abundance:
              Hood Canal     :             Highly abndant    indicate that spawning occurs all year with a winter
               Skagit Bay              3   Abundant       peak (depending on area) (Fitch and Lavenberg 1975,
             Grays Harbor              O   Common         McGowen 1977, Wang 1986). Spawning has been
              Willapa Bay              i   Rare           recorded during September-February in Anaheim Bay
           Columbia River            Blank Notpresent     (Lane 1975, Gadomski and Petersen 1988), and June-
             Nehalem Bay                                  October in Richardson Bay (Eldridge 1975, Eldridge
            Tillamook Bay            Life stage:          1977).  The diamond turbot may have a specific
              Netarts Bay             A- Adults           temperature preference for spawning. Thistemperature
              Siletz River           S -eSpawning adults probably occurs in winter in southern California (14-
              SileJ River            J-Juvenlles
             Yaquina Bay             L - Larvae           1 60C) (McGowen 1977, Walker et al. 1987), and spring
              Alsea River             E - Eggs            and summer near San Francisco Bay.
             Siuslaw River
            Umpqua River                                  Fecundity: Unknown.
               Coos Bay
              Rogue River                                 Growth and Development
            Klamath River                                 Eaa Size and Embryonic Development: Eggs are
            Humboldt Bay                                  spherical, ranging in diameter from 0.78-0.90 mm,
                Eel River                                 averaging 0.84 mm (Eldridge 1975, Su mida et al. 1979,
             Tomales Bay - i   O �                        Wang 1986). Embryonic development is indirect and
        Cent. San Fran. Bay *       *Includes Central San  external.
                                      Francisco, Suisun,
        South San Fran. Bay  0 0 0 0   and San Pablo bays.
            Elkhorn Slough   O                           Larval Size Ranae: The yolk-sac is depleted in 5 days
               Morro Bay      O0                          at 17�C (Gadomski and Petersen 1988). The larval life
         Santa Monica Bay OO 0    o 0O                    stage lasts at least 5-6 weeks at 16�C (Gadomski and
            SanPedroBay O O O O O                          Petersen 1988). Larvae average 1.6 mm standard
             AlamitosBay i    (9 O                        length (SL) at hatching, and grow 7-8 mm before
             Anaheim Bay (j    j 0                        metamorphosis (about 11.0 mm long) (Eldridge 1975,
             Newport Bay  3 '  3                          Sumida et al. 1979, Gadomski and Petersen 1988).
              Mission Bay 0    1 CD 0
            San Diego Bay O    O O t                     Juvenile Size Ranae: Juveniles settle out of the water
           Tijuana Estuary q    O  I                      column at metamorphosis (about 11.0 mm SL) (Eldridge
                       A S J L E                          1975, Sumida et al. 1979, Gadomski and Petersen
                                                          1988).
    The maximum salinity tolerated by juveniles and adults
    is 60%0 (Carpelan 1961). Juveniles and adults are   Aae and Size of Adults: Females mature in 2-3 years
    probably eurythermal; upper temperature limits are    (about 180 mm TL).  The largest diamond turbot
    unknown. Densities of eggs and larvae were positively    reported was 46 cm TL and the heaviest was a
    correlated with distance from thermal plant discharge    approximately 0.9 kg  (Baxter 1960, Miller and Lea
    and dissolved oxygen, and were negatively correlated    1972, Fitch and Lavenberg 1975). Individuals 30.5-
    with temperature and light extinction coefficients   38.1 cm long are probably 8-9 years old (Fitch and
    (McGowen 1977).                                       Lavenberg 1975).

    Miarations and Movements: Larvae appearto settleon    Food and Feeding
    sandy sediments in the shallow waters in or near bays    Trophic Mode: Larvae are planktivorous and juveniles
    and estuaries (Lane 1975). Once individuals are in a    and adults are carnivorous.  Juveniles and adults
    bay, they do not appear to move widely. However, a    appearto feed diurnally, foraging on or in the substrate
    general movement of larger fish to lower portions of   (Lane 1975).  Adult and juvenile diamond turbot in
    bays and estuaries is indicated and adults appear to   Anaheim Bay, California, consumed 3.76% of their
    move out of bays and estuaries to spawn (Lane 1975).    body weight each day (Lane et al. 1979).

                                                      257

    Diamond turbot continued
    Food Items: Larvae probably eat zooplankton and    Baxter, J. L. 1960. Inshore fishes of California. Calif.
    phytoplankton.   Juveniles and adults consume  . Dept. Fish Game, Sacramento, CA, 80 p.
    polychaetes, clams and clam siphons, gastropods,
    ghost shrimp (Callianassa spp.), amphipods,    Cailliet, G. M., B. Antrim, D. Ambrose, S. Pace, and M.
    cumaceans, various crustaceans, and small fish (Fitch    Stevenson. 1977. Species composition, abundance
    and Lavenberg 1975, Lane 1975).  Large diamond    and ecological studies of fishes, larval fishes, and
    turbot (>25g) eat more molluscs, fish, and large    zooplankton in Elkhorn Slough.  In Ecologic and
    crustaceans than smaller turbot (Lane 1975).            hydrographic studies of Elkhorn Slough, Moss Landing
                                                           Harbor and nearshore coastal waters, p. 216-386.
    Biological Interactions                                 Moss Landing Marine Lab., Moss Landing, CA.
    Predation: Predators probably include the Pacific
    electric ray (Torpedo californica), Pacific angel shark    California Department of Fish and Game. 1987. Delta
    (Squatina californica), andotherlargepiscivorousfishes    outflow effects on the abundance and distribution of
    (Fitch and Lavenberg 1975). Birds (such as herons)    San Francisco Bay fish and invertebrates, 1980-1985.
    and cormorants (Phalocrocorax spp.) are also    Exhibit 60, entered by the Calif. Dept. Fish Game for
    predators.                                              the State Water Resources Control Board 1987 Water
                                                           Quality/Water Rights Proceeding on the San Francisco
    Factors Influencino PoDulations: The diamond turbot    Bay/Sacramento-San Joaquin Delta. Calif. Dept. Fish
    population in San Francisco Bay increases in   Game, Stockton, CA, 345 p.
    abundance during wet years (Armor and Herrgesell
    1985, California Department of Fish and Game 1987).    Carpelan, L. H.  1961. Salinity tolerances of some
    Mortality rates for 1- and 2-year-old fish are very high    fishes of a southern California coastal lagoon. Copeia
    (Lane 1975), and many adults apparently die after   1961(1):32-39.
    spawning (Lane 1975). Few adu Its live beyond 2 years
    in Anaheim Bay (Lane 1975). For larvae, the onset of    Chapman, G. A.  1963.  Mission Bay, a review of
    initial feeding is important for their survival (Gadomski    previous studies and the status of the sportfishery.
    and Petersen 1988). The diamond turbot is dependent    Calif. Fish Game 49(1):30-43.
    on bays and estuaries, hence the health of these
    habitats is critical to this species' survival.         Eldridge, M. B. 1975.  Early larvae of the diamond
                                                          turbot, Hypsopsetta guttulata. Calif. Fish Game
    References                                              61(1):26-34.

    Allen, L. G. 1976. Abundance, diversity, seasonality    Eldridge, M. B. 1977. Factors influencing distribution
    and community structure of the fish populations of   of fish eggs and larvae over eight 24-hr samplings in
    Newport Bay, California.  M.A. Thesis, Calif. State    Richardson Bay, California. Calif. Fish Game63(2):1 01-
    Univ., Fullerton, CA, 107 p.                            116.

    Aplin, J. A. 1967. Biological survey of San Francisco    Eschmeyer, W. N., E. S. Herald, and H. Hammann.
    Bay. 1963-1966. MRO Ref. No. 67-4. Calif. Dept. Fish    1983. A field guide to Pacific coast fishes of North
    Game, Menlo Park, CA, 131 p.                            America. Houghton Mifflin Co., Boston, MA, 336 p.

    Armor, C., and P. L. Herrgesell. 1985. Distribution and    Feder, H. M., C. H. Turner, and C. Limbaugh. 1974.
    abundance of fishes in the San Francisco Bay estuary    Observations on fishes associated with kelp beds in
    between 1980 and 1982. Hydrobiol. 129:211-227.          southern California. Calif. Fish Game, Fish Bull. 160:1-
                                                           144.
    Bane, G. W., and A. W. Bane. 1971. Bay fishes of
    northern California with emphasis on the Bodega    Fierstine, H. L., K. F. Kline, and G., R. Garman. 1973.
    Tomales Bay area. Mariscos Publ., Hampton Bays,    Fishes collected in Morro Bay, California between
    NY, 143 p.                                              January 1968 and December 1970. Calif. Fish Game
                                                           59(1):73-88.
    Barnett, A. M., A. E. Jahn, P. D. Sertic, and W. Watson.
    1984. Distribution of ichthyoplankton off San Onofre,    Fitch, J. E., and R. J. Lavenberg. 1975. Tidepool and
    California, and methods for sampling very shallow    nearshore fishes of California. Calif. Nat. Hist. Guides
    coastal waters. Fish. Bull., U.S. 82(1):97-111.         38, Univ. Calif. Press, Berkeley, CA, 156 p.

                                                       258

                                                                                       Diamond turbot continued
    Gadomski, D. M., and J. H. Petersen. 1988. Effects of    Squire, J. L.,Jr., and S. E. Smith. 1977. Anglers'guide
    food deprivation on the larvae of two flatfishes. Mar.    to the United States Pacific Coast. U.S. Dept. Comm.,
    Ecol. Prog. Ser. 44:103-111.                             NOAA, Seattle, WA, 139 p.

    Gates, D. E., and H. W. Frey.  1974.  Designated    Sumida, B. Y., E. H. Ahlstrom, and H. G. Moser. 1979.
    common  names of certain marine organisms of    Early development of seven flatfishes of the eastern
    California. Calif. Fish Game, Fish Bull. 161:55-88.      North Pacific with heavily pigmented larvae (Pisces,
                                                           Pleuronectiformes). Fish. Bull., U.S. 77(1):105-145.
    Horn, M. H., and L. G. Allen. 1981. Ecology of fishes
    in upper Newport Bay, California: seasonal dynamics    Walker, H. J., Jr., W. Watson, and A. Barnett. 1987.
    and community structure. Mar. Res. Tech. Rep. No.    Seasonal occurrence of larval fishes in the nearshore
    45, Calif. Dept. Fish Game, Long Beach, CA, 102 p.    southern California Bight off San Onofre, California.
                                                           Est. Coast. Shelf Sci. 25:91-109.
    Lane, E. D. 1975. Quantitative aspects of the life
    history of the diamond turbot, Hypsopsetta guttulata    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    (Girard), in Anaheim Bay. In E. D. Lane and C. W. Hill   Joaquin estuary and adjacent waters, California: a
    (editors),ThemarineresourcesofAnaheimBay. Calif.    guide to the early life histories.  Tech. Rep. No. 9,
    Fish Game, Fish Bull.1 65:153-173.                       prepared forthe interagency ecological study program
                                                           for the Sacramento-San Joaquin estuary. Calif. Dept.
    Lane, E. D., M. C. S. Kingsley, and D. E. Thorton. 1979.    Water Res, Calif. Dept. Fish Game,  U.S. Bureau
    Daily feeding and food conversion efficiency of the    Reclam., and U.S. Fish Wildl. Serv., various pagination.
    diamond turbot: an analysis based on field data. Trans.
    Am. Fish. Soc. 108:530-535.                              Zedler, J. B., and C. S. Nordby. 1986. The ecology of
                                                           Tijuana estuary, California: an estuarine profile. U.S.
    McGowen,G. E. 1977. Ichthyoplankton populations in   Fish Wildl. Serv. Biol. Rep 85(7.5), 104 p.
    south San Diego Bay and related effects of an electricity
    generating station. M.S. Thesis, San Diego State Univ,
    San Diego, CA, 88 p.

    Miller, D. J., and R. N. Lea. 1972. Guide to the coastal
    marine fishes of California. Calif. Fish Game, Fish Bull.
    157, 235 p.

    Noah, M. D. 1985. Structure, abundance and
    distribution of the fish and macroinvertebrate
    communities inhabiting Mission Bay, California between
    November 1979 and February 1981. Appendix A. San
    Diego River and Mission Bay improvements, Draft
    suppl. environ. assess. U.S. Army Corps Eng., Los
    Angeles, CA, 37 p. plus appendices.

    Ocean Assessments Division. 1984. The national
    status and trends program for marine environmental
    quality: program description (mimeo). Ocean
    Assessments Division, NOS/NOAA, Rockville, MD,
    28 p.

    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    of common and scientific names of fishes from the
    United States and Canada. Am. Fish. Soc. Spec. Publ.
    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.

    Roedel, P. M. 1953. Common ocean fishes of the
    California coast. Calif. Fish Game, Fish Bull. 91,
    184 p.

                                                       259

       Pleuronectes vetulus
       Juvenile

    Common Name: English sole                            accumulates contaminants and is a target species for
    Scientific Name: Pleuronectes(or Parophrys) vetulus   the National Status and Trends Program (Ocean
    A recent review of the family Pleuronectidae indicates   Assessments Division 1984).  The English sole
    thatthis species maybelongtothegenus Pleuronectes   apparently develops cancerous tumors as a result of
    (Sakamoto 1984)                                      exposure to contaminants (Malins et al. 1983). Three
    Other Common Names: California sole, lemon sole,   types of superficial papillomas have been identified
    common sole, pointed nose sole, sharp nose sole   from subyearling English sole; all three appear to
    (Washington 1977)                                    cause substantial mortality. Tumors and liver lesions
    Classification (Sakamoto 1984)                       may be caused by exposure to contaminants such as
    Phylum: Chordata                                     aromatic hydrocarbons (Krahn et al. 1986,1987).
    Class: Osteichthyes
    Order: Pleuronectiformes                             Ecological: The English sole is a very important flatfish
    Family: Pleuronectidae                               in shallow-water, soft-bottom marine and estuarine
                                                          environments along the Pacific Coast (Westrheim 1955,
    Value                                                Washington 1977, Hogue and Carey 1982, Krygier and
    Commercial: The English sole is a moderately important    Pearcy 1986).
    commercial fish, captured primarily by trawls. Over
    2,500 t were landed in the U.S. in 1986, primarily in   Range
    Washington and California (Pacific Marine Fisheries   Overall:Thisspecies'overall range isfromcentral Baja
    Commission 1987). It is the most abundant flatfish   California, Mexicoto Unimak Island, Alaska(Hart 1973).
    species in Puget Sound, Washington (Pedersen and   It is most abundant northfrom Pt. Conception, California.
    DiDonato 1982). Females dominate the commercial
    catch because males rarely grow to marketable size   Within Study Area: Juveniles are found in nearly all
    (Pedersen and DiDonato 1982). The English sole has    Pacificcoast estuariesfrom San Pedro Bay, California,
    an "iodine" taste which some people prefer and is   to Puget Sound (Table 1). However, Elkhorn Slough,
    marketed as fillets of sole (Clemens and Wilby 1961,   California appears to be the most southern estuary
    Hart 1973). Itissecondonlyto Doversole(Microstomus   where they are abundant.
    pacificus) in flatfish pounds landed on the Pacific coast
    (Pacific Marine Fisheries Commission 1987).          Life Mode
                                                           Eggs and larvae are pelagic, while juveniles and adults
    Recreational: This is not an important recreational fish,   are demersal (Budd 1940, Forrester 1969, Hart 1973).
    although it is caught using hook and line by boat, shore,
    and pier anglers. Boat anglers caught over 1,400 in   Habitat
    Washington waters in 1984 (Hoines et al. 1984).      Type: Eggs are neritic and pelagic, but sink just before
                                                          hatching (Hart 1973). Larvae are also pelagic and are
    Indicator of Environmental Stress: This species often   found primarily in waters <200 m deep (Laroche and

                                                       260

                                                                                        English sole continued

                                                          preference for fine sediments (Becker 1988).
     Table 1. Relative abundance of English sole
              in 32 U.S. Pacific coast estuaries.         Phvsical/Chemical Characteristics: Adults are found
                         Life Stage                       primarily in marine (euhaline) waters. Juveniles and
            Estuary    A S J L E                          larvae occur in polyhaline and euhaline waters.
             PugetSound 130 � � 6�   Relative abundance:  Optimumconditionsforlarvalsurvivalwerefoundtobe
              Hood Canal   ::    �    �a 0  S   Highly abundant    salinities of 25-28%0 and temperatures of 8-9�C
              Skagit Bay I   ] 13   It S:  I3  Abundant   (Alderdice and Forrester 1968). No spawning occurs
            Grays Harbor   IS 0       0   Common          at temperatures below approximately 7.8�C (Jackson
              Willapa Bay    I     0   i  Rare            1981). Temperatures >18�C appear to be the upper
           Columbia River     0      Blank Notpresent     thermal tolerance (reduced daily ration and growth) for
            Nehalem Bay    (3 0                           juvenile English sole (Yoklavich 1982). The upper
            Tillamook Bay   � O       Life stage:         lethal limit forthis species is 26.1 �C (Ames et al. 1978).
              NetartsBay     0 O  A-Adults
                                      S - Spawning adults
              Siletz River  ( 3 0    J-Juveniles          Miarations and Movements: Adults make limited
             Yaquina Bay    * O      L - Larvae           migrations/movements.  Those off Washington and
              AlseaRiver               - Eggs             British Columbia show a northward post-spawning
            Siuslaw River   ( 0                           migration in the spring on their way to summer feeding
            Umpqua River    9 O                           grounds, and a southerly movement in the fall (Garrison
               Coos Bay     � O                           and Miller 1982).  Tagging studies have identified
             Rogue River                                  separate stocks based on this species' limited
            Klamath River                                 movements and meristic characteristics (Jow 1969).
            Humboldt Bay    a 0                           Tidal currents appear to be the mechanism by which
                Eel River   0 0                            English sole move into estuaries (Boehlert and Mundy
             TomalesBay     i 0                           1987); larvae are transported to nearshore nursery
       Cent San Fran. Bay *    O    * Indudes Central San  areas (i.e., shallow coastal waters and estuaries) by
                                      Francisco, Suisun,
       South San Fran. Bay   a    i  and San Pabb bays.   currents. Larvae metamorphose into juveniles in spring
           Elkhorn Slough  O(  V                          and early summer and rear until fall/winter at which
               Morro Bay   QO                             time most emigrate to deeper waters (Olson and Pratt
         Santa Monica Bay  O O 0 0 0                      1973). Although many postlarvae may settle outside of
           San Pedro Bay CO    ' "                        estuaries, apparently most will enter estuaries during
             Alamitos Bay                                 some part of their first year of life (Gunderson et al.
            Anaheim Bay                                   1990). Early- and late-stage larvae undergo diel vertical
             Newport Bay                                  migrations (Misitano 1970, 1976). There is a general
             Mission Bay                                  movement to deeper waters as fish grow (Ketchen
           San Diego Bay                                  1956). Smaller fish tend to be restricted to shallow
           Tijuana Estuary                                waters, with larger fish more abundant in deeper water
                       A S J L E                          (English 1967, Misitano 1970, Sopher 1974).

    Richardson 1979).  Adults are found in nearshore    Reproduction
    coastal waters down to 550 m depth, but primarily in   Mode: The English sole is gonochoristic, oviparous,
    depths <250 m (Allen and Smith 1988). In Canadian    and iteroparous;eggs are fertilized externally (Garrison
    waters, this species is commercially abundant between    and Miller 1982).
    36 and 128 m depths (Forrester 1969). Juveniles
    reside primarily in shallow-water coastal, bay, and    Matina/Soawnina: Spawning occurs over soft-bottom
    estuarine areas (Westrheim 1955, Ketchen 1956, Van    mud substrates at depths of 50-70 m (Ketchen 1956).
    Cleve and El-Sayed 1969, Olson and Pratt 1973,    Spawning occurs fromwintertoearlyspring depending
    Pearcy and Myers 1974, Laroche and Holton 1979,    on the stock: from January to May in Monterey Bay
    Toole 1980, National Marine Fisheries Service 1981,   stocks, peaking in March or April (Budd 1940); in
    Krygier and Pearcy 1986, Rogers et al. 1988).         Bodega Bay-Point Monterey stocks,from Decemberto
                                                          April, peaking January or February (Villadolid 1927,
    Substrate: Eggs are buoyant and larvae are pelagic.   cited in Garrison and Miller 1982); Santa Monica Bay-
    Adults and juveniles prefer soft bottoms composed of   Santa Barbara Channel stocks from December to
    fine sands and mud (Ketchen 1956). In Puget Sound,    April; in Eureka-Oregon border stocks during October
    juveniles and adults prefer shallow (<12 m deep)    to May (Jow 1969); in Oregon stocks from January to
    muddy substrates (Becker 1984).  Males show a    April, peaking in February or March (Harry 1959); in

                                                       261

    English sole continued
    Puget Sound stocks, from January to April, peaking in   sole feed on avarietyof benthicorganisms, but primarily
    February or March (Smith 1936); in Hecata Strait,   polychaetes, amphipods, molluscs, ophiouroids, and
    British Columbia stocks, from late December to early    crustaceans (Kravitz et al. 1976). English sole feed
    April, peaking in February (Ketchen 1956).              primarily by day, using sight and smell, and sometimes
                                                           dig for prey (Allen 1982, Hulberg and Oliver 1979).
    Fecundity: Five- to six-year-old females (36-38 cm in
    length) can produce about 1 million eggs, while large    Biological Interactions
    fish (43 cm) may produce nearly 2 million eggs (Ketchen    Predation: Larvae are probably eaten by larger fishes.
    1947, Harry 1959, Forrester 1969).                     A juvenile English sole's main predators are probably
                                                           piscivorous birds such as great blue heron (Ardia
    Growth and Development                                  herodias), larger fishes, and marine mammals. Adults
    Eaa Size and Embrvonic Development: Fertilized eggs    may be eaten by marine mammals, sharks, and other
    are spherical and average 0.98 mm in diameter (Orsi   large fishes. The English sole's sharp anterior anal
    1968). Embryonic development is indirect and external.    spine may provide a defense against predators (Allen
    The planktonic eggs hatch in 3.5 days at 12�C, or 11.8    1982).
    days at 40C (Alderdice and Forrester 1968).
                                                           Factors Influencina PoDulations: Upwelling (and thus
    Aoeand Sizeof Larvae:Afterhatching, larvaefloatwith    water temperatures) during the larval and spawning
    theiryolksacup. Theyolksacisabsorbedin9-10days    period affects eventual recruitment (Ketchen 1956,
    (Orsi 1968), with the planktonic larvae taking from 8-10    Kruse and Tyler 1983). Growth appears to be affected
    weeks to metamorphose to benthic living juveniles    by upwelling (Kreuz et al. 1982) and cohort abundance
    (Laroche et al. 1982). Larvae are 2.0-2.8 mm total   of age-1 fish (Peterman and Bradford 1987). Models
    length (TL) at hatching (Orsi 1968) and grow to 18-26    have been developed to identify oceanographic
    mm before becoming juveniles (Misitano 1976, Garrison    conditions that influence English sole recruitment (Kruse
    and Miller 1982).                                       and Tyler 1983), but it appears that numerous physical
                                                           and biological parameters combine to control year-
    Juvenile Size Rance: Juveniles range in size from 18    class strength (Botsford et al. 1989).  Important
    mm to about 26 cm long (depending on sex) (Harry    recruitment processes includethetiming of spawning,
    1959).                                                  surface temperatures during larval development,
                                                           onshore transport of larvae, and age- and density-
    Ace and Size of Adults: Some females mature as 3-    dependentgrowth and mortalityofjuvenilesandyoung
    year-olds and 26 cm long, but all females over 35 cm    adults (Botsford et al. 1989).  At high population
    long are mature. Males mature earlier, beginning at 2    densities, a myxosporidian disease can infect this
    years and 21 cm in length. All males are mature at   speciesandmakeitsflesh"milky"(Hart1973). Because
    lengths >29 cm (Harry 1959). In Puget Sound, all 2-   the English sole uses nearshore coastal and estuarine
    year-old males are mature, but most females do not    waters as nursery areas (Krygier and Pearcy 1986,
    mature until they are 4 years old (Smith 1936).         Rogers et al. 1988), it is exposed to numerous toxic
                                                           materials which can result in a high incidence of diseased
    Food and Feeding                                        fish in some estuaries. Sincethis species relies heavily
    Trophic Mode: Larvae are planktivorous. Juveniles,    on estuaries for rearing, the alteration and pollution of
    and adults arecarnivorous, apparentlyfeeding primarily    estuarine habitats adversely affects this species
    during daylight hours (Becker 1984).                    (Gunderson et al. 1990).

    Food Items: Larvae probably eat different life stages of    References
    copepods and othersmall planktonic organisms. Larvae
    appeartohaveastrongpreferenceforappendicularians    Alderdice, D. F., and C. R. Forrester.  1968. Some
    (Botsford et al. 1989). Juveniles feed on harpacticoid    effectsof salinityandtemperatureonearlydevelopment
    copepods, gammarid amphipods, cumaceans, mysids,    and survival of the English sole (Parophrys vetulus). J.
    polychaetes, small bivalves, clam siphons, and other    Fish. Res. Board Can. 25(3):495-521.
    benthic invertebrates (Simenstad et al. 1979, Allen
    1982, Hogue and Carey 1982, Becker 1984, Bottom et   Allen, M. J. 1982. Functional structure of soft-bottom
    al. 1984). Small juvenile English sole concentratetheir   fish communities ofthe southern California shelf. Ph.D.
    feeding on harpacticoid copepods and other epibenthic    Diss. Univ. Calif., San Diego, CA, 577 p.
    crustaceans until they reach approximately 50-65 mm
    in length, then they switch to feeding primarily on    Allen, M. J., and G. B. Smith.  1988.  Atlas and
    polychaetes (Toole 1980). Off Oregon, adult English    zoogeography of common and marine fishes in the

                                                        262

                                                                                        English sole continued
    northeast Pacific Ocean and Bering Sea. NOAATech.    juvenile English sole (Parophrys vetulus) and
    Rep. NMFS 66,151 p.                                     Dungeness crab (Cancermagister). Estuaries 13(1 ):59-
                                                           71.
    Ames, W. E., J. R. Hughes, and G. F. Slusser. 1978.
    Upper lethal water temperature levels for English sole    Harry, G. Y.  1959.  Time of spawning, length of
    (Parophrys vetulus) and rock sole (Lepidopsetta    maturity, and fecundity of the English, petrale, and
    bilineata) subjected to gradual thermal increases.    Dover soles (Parophrys vetulus, Eopsettajordani, and
    Northw. Sci. 52(3):285-291.                             Microstomus pacificus, respectively).  Fish. Comm.
                                                           Oreg. Res. Briefs 7(1):5-13.
    Becker, D. S. 1984. Resource partitioning by small-
    mouthed pleuronectids in Puget Sound, Washington.    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
    Ph.D. Diss. Univ. Wash., Seattle, WA, 138 p.            Board Can., Bull. No. 180, 740 p.

    Becker, D. S. 1988. Relationships between sediment    Hogue, E. W., and A. G. Carey, Jr. 1982. Feeding
    character and sex segregation in English sole,    ecology of 0-age flatfishes at a nursery ground on the
    Parophrys vetulus. Fish. Bull., U.S. 86(3):517-524.     Oregon coast. Fish. Bull., U.S. 80(3): 555-565.

    Boehlert, G. W.,and B. C. Mundy. 1987. Recruitment    Hoines, L. J., W. D. Ward., and C. Smitch.  1984.
    dynamics of metamorphosing English sole, Parophrys    Washington State sport catch report 1984.  Wash.
    vetulus, to Yaquina Bay, Oregon.  Estuar. Coastal    Dept. Fish., Olympia, WA, 58 p.
    Shelf Sci. 25:261-281.
                                                           Hulberg, L. W., and J. S. Oliver. 1979. Prey availability
    Botsford, L. W., D. A. Armstrong, and J. M. Shenker.    and the diets of two co-occurring flatfish. In S. J.
    1989. Oceanographic influences on the dynamics of    Lipovsky and C. A. Simenstad (editors), Fish food
    commercially fished populations. In M. R. Landry and    habits studies, proceedings of the second Pacific
    B. M. Hickey (editors), Coastal oceanography of    Northwest technical workshop, p. 29-36. Wash. Sea
    Washington and Oregon, p. 511-565.  Elsevier Sci.    Grant, Univ. Wash, Seattle, WA, (WSG-WO-79-1).
    Publ., B.V., Amsterdam.
                                                           Jackson, C. 1981. Flatfishes:A systematic study of the
    Bottom, D. L., K. K. Jones, and M. J. Herring. 1984.    Oregon pleuronectid production system and its fishery.
    Fishes of the Columbia Riverestuary. Col. Riv. Estuary    Sea Grant College Prog., Oregon State Univ., Corvallis,
    Data Devel. Prog., CREST, Astoria, OR, 113 p. plus    OR, 40 p. (ORESU-T-81-001).
    appendices.
                                                           Jow, T. 1969. Results of English sole tagging off
    Budd, P. L. 1940. Development of the eggs and early   California. Pac. Mar. Fish. Comm., Bull. 7:16-33.
    larvae of six California fishes. Calif. Fish Game, Fish
    Bull. 56:1-50.                                          Ketchen, K. S. 1947.   Studies on lemon sole
                                                           development and egg production. Fish. Res. Board
    Clemens, W. A., and G. V. Wilby. 1961. Fishes of the    Can., Prog. Rep. Pac. Coast Sta. 73:68-70.
    Pacific coast of Canada. Fish. Res. Board Can., Bull.
    No. 68, 443 p.                                          Ketchen, K. S. 1956. Factors influencing the survival
                                                           of the lemon sole (Parophrys vetulus) in Hecata Strait,
    English, T. S. 1967. Preliminary assessment of the    British Columbia. J. Fish Res. Board Can. 13:647-694.
    English sole in Port Gardner, Washington. J. Water
    Pollution Control Fed. 39(3):1337-1350.                 Krahn, M. M., L. D. Rhodes, M. S. Myers, L. K. Moore,
                                                           W. D. MacLeod, Jr., and D. C. Malins. 1986.
    Forrester, C. R. 1969. Lifehistoryinformationonsome    Associations between metabolites of aromatic
    groundfishspecies. Fish. Res. Board Can., Tech. Rep.    compounds in bile and the occurrence of hepatic
    No. 105, 17 p.                                          lesions in English sole (Parophrys vetulus) from Puget
                                                           Sound, Washington. Arch. Environ. Contam. Toxicol.
    Garrison, K. J., and B. S. Miller. 1982. Review of the    15:61-67.
    early life historyof Puget Sound fishes. Fish. Res. Inst.,
    Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).         Krahn, M. M, D. G. Burrows, W. D. MacLeod, Jr., and
                                                           D. C. Malins. 1987. Determination of individual
    Gunderson, D. R., D. A. Armstrong, Y. B. Shi, and R. A.    metabolites of aromatic compounds in hydrolyzed bile
    McConnaughey. 1990. Patterns of estuarine use by    of English sole (Parophrys vetulus) from polluted sites

                                                       263

    English sole continued
    in Puget Sound, Washington. Arch. Environ. Contam.    National Marine Fisheries Service. 1981. Columbia
    Toxicol. 16:511-522.                                    River estuary data development program report,
                                                          salmonid and non-salmonid fish. Unpubl. manuscr.,
    Kravitz, M. J., W. G. Pearcy, and M. P. Guin. 1976.    various pagination, Northwest Alaska Fish. Cent., Nat.
    Food of five species of cooccurring flatfishes on    Mar. Fish.Serv., P.O. Box 155, Hammond, OR,97121.
    Oregon's continental shelf. Fish. Bull., U.S. 74:984-
    990.                                                    Ocean Assessments Division.  1984.  The national
                                                          status and trends program for marine environmental
    Kreuz, K. F, A. V. Tyler, G. H. Kruse, and R. L. Demory.    quality: Program description (mimeo).   Ocean
    1982. Variation in growth of Dover soles and English    Assessments Division, NOS/NOAA, Rockville, MD,
    soles as related to upwelling. Trans. Am. Fish. Soc.    28 p.
    111 (2):180-192.
                                                           Olson, R. E., and I. Pratt. 1973. Parasites as indicators
    Kruse, G. H., and A. V. Tyler. 1983. Simulation of   of English sole (Parophrys vetulus) nursery grounds.
    temperature and upwelling effects on the English sole    Trans. Am. Fish. Soc. 102: 405-411.
    (Parophrys vetulus) spawning season. Can. J. Fish.
    Aquat. Sci. 40:230-237.                                 Orsi, J. J. 1968. The embryology of the English sole,
                                                           Parophrys vetulus. Calif. Fish Game 54(3):133-155.
    Krygier, E. E., and W. G. Pearcy. 1986. The role of
    estuarineandoffshorenurseryareasforyoung English    Pacific Marine Fisheries Commission.  1987.  39th
    sole, ParophrysvetulusGirard, of Oregon. Fish. Bull.,    annual report of the Pacific Marine Fisheries
    U.S. 84(1):119-132.                                    Commission forthe year 1986. Pac. Fish. Man. Comm.,
                                                           Portland, OR, 29 p.
    Laroche, W. A., and R. L. Holton. 1979. Occurrence
    of 0-age English sole, Parophrys vetulus, along the    Pearcy, W. G., and S. S. Myers. 1974. Larval fishes of
    Oregon coast: an open coast nursery area? Northw.    Yaquina Bay, Oregon: A nursery ground for marine
    Sci. 53:94-96.                                          fishes? Fish. Bull., U.S. 72:201-213.

    Laroche, J. L., and S. L. Richardson. 1979. Winter-    Pedersen, M., and G. DiDonato. 1982. Groundfish
    spring abundance of larval English sole, Parophrys    management plan for Washington's inside waters.
    vetulus, betweenthe Columbia River and Cape Blanco,    Prog. Rep. No. 170, Wash. Dept. Fish., Olympia, WA,
    Oregon during 1972-1975 with notes on occurrences    123 p.
    ofthreeotherpleuronectids. Estuar. Coastal Mar. Sci.
    8:455-476.                                              Peterman, R. M. and M. J. Bradford. 1987. Density-
                                                           dependent growth of age 1 English sole (Parophrys
    Laroche, J. L, S. L. Richardson, and A. Rosenberg.    vetulus) in Oregon and Washington coastal waters.
    1982. Age and growth of a pleuronectid, Parophrys    Can. J. Fish. Aquat. Sci. 44:48-53.
    vetulus, during the pelagic larval period in Oregon
    coastal waters. Fish. Bull. U.S. 80(1):93-104.          Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
                                                           E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
    Malins, D. C., M. S. Myers, and W. T. Roubal. 1983.    of common and scientific names of fishes from the
    Organic free radicals associated with idiopathic liver   United States and Canada. Am. Fish. Soc. Spec. Publ.
    lesions of English sole (Parophrys vetulus) from polluted    No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
    marine environments. Envir. Sci. Tech. 17(11):679-
    685.                                                    Rogers, C. W., D. R. Gunderson, and D. A. Armstrong.
                                                           1988. Utilization of a Washington estuary by juvenile
    Misitano, D. A. 1970. Aspects of the early life history    English sole, Parophrys vetulus.  Fish. Bull., U. S.
    of English sole (Parophrys vetulus) in Humboldt Bay,    86(4):823-831.
    California. M.S. Thesis, Humboldt State College,
    Eureka, CA, 54 p.                                       Sakamoto, K. 1984. Interrelationships of the family
                                                           Pleuronectidae (Pisces: Pleuronectiformes). Mem.
    Misitano, D. A. 1976. Size and stage of development    Fac. Fish. Hokkaido Univ. 31(1/2):85-215.
    of larval English sole, Parophrys vetulus, at time of
    entry into Humboldt Bay. Calif. Fish Game 62(1):93-    Simenstad, C. A., B. S. Miller, C. F. Nyblade, K.
    98.                                                     Thornburgh, and L. J. Bledsoe.  1979.  Food web
                                                           relationships of northern Puget Sound and the Strait of

                                                       264

                                                                                         English sole continued
    Juan de Fuca. U.S. Interagency (NOAA, EPA) Energy/
    Environ. Res. Dev. Prog. Rep., EPA-600/7-79-259.
    Washington, D.C., 335 p.

    Smith, R. T. 1936. Report on the Puget Sound otter
    trawl investigations. Wash. Dept. Fish. Biol. Rep.
    36B:1-61.

    Sopher, T. R. 1974. A trawl survey of the fishes of
    Arcata Bay, California. M.S. thesis, Humboldt State
    Univ., Arcata, CA, 103 p.

    Toole, C. L. 1980. Intertidal recruitment and feeding in
    relationto optimal utilization of nursery areas by juvenile
    English sole (Parophrys vetulus: Pleuronectidae).
    Environ. Biol. Fish. 5:383-390.

    Van Cleve, R., and S. Z. El-Sayed. 1969. Age, growth,
    and productivity of an English sole (Parophrys vetulus)
    population in Puget Sound, Washington. Pac. Mar.
    Fish. Comm. Bull. 7:51-71.

    Villadolid, D. V. 1927. The flatfish (Heterosomata) of
    the Pacific coast of the United States. Ph.D. Thesis,
    Stanford Univ., Palo Alto, CA, 332 p.

    Washington, P. M. 1977. Recreationally important
    marine fishes of Puget Sound, Washington. Proc.
    Rep., Northwest Alaska Fish. Cent., NOAA, Nat. Mar.
    Fish. Serv., Seattle, WA, 122 p.

    Westrheim, S. J. 1955. Size composition, growth, and
    seasonal abundance of juvenile English sole (Parophrys
    vetulus) in Yaquina Bay. Fish Comm. Oregon, Res.
    Briefs 6(2):4-9.

    Yoklavich, M. 1982. Growth, food consumption, and
    conversion efficiency of juvenile English sole (Parophrys
    vetulus). In G. M. Cailliet and C. M. Simenstad (editors),
    Gutshop 81, Fish food habits studies, Proceedings of
    the third Pacific workshop, p. 97-105. Wash. Sea
    Grant, Univ. Wash., Seattle, WA.

                                                        265

       Platichthys stellatus
       Adult                                             ,

                                  10cm

    Common Name: starry flounder                          accumulates contaminants (Ocean Assessments
    Scientific Name: Platichthys stellatus                Division 1984).
    Other Common Names: California flounder, grindstone
    flounder, greatflounder, roughjacket, diamond flounder,   Ecological: The starry flounder is the most abundant
    sole, flounder, emery flounder (Gates and Frey 1974,   flatfish in many Pacific coast estuaries north of San
    Washington 1977)                                      Francisco Bay, California (National Marine Fisheries
    Classification (Robins et al. 1980)                   Service 1981, Bottom et al. 1984, Pedersen and
    Phylum: Chordata                                      DiDonato 1982). It is prey for marine mammals (Jeffries
    Class: Osteichthyes                                   et al. 1984) and piscivorous birds.
    Order: Pleuronectiformes
    Family: Pleuronectidae                                Range
                                                          Overall: The starry flounder is distributed Arctic-
    Value                                                 circumboreal and found in the eastern Pacific Ocean
    Commercial: The starry flounder is a moderately    from Santa Ynez River, California, north through the
    important flatfish species landed by the Pacific coast    Bering and Chukchi Seas to Bathurst Inlet in Arctic
    trawlfisheryfromtheBeringSeatoSouthernCalifomia.    Canada. In the western Pacific, it is found along the
    From 1981 to 1983, an average of over 1,300 t were    Kamchatka Peninsula south to Tokyo Bay, Japan
    landed, of which 90% were taken by U.S. fishermen.    (Orcutt 1950, Okada 1955, Wilimovsky 1964, Allen and
    Most ofthe catch comes from Puget Sound, Washington    Smith 1988).
    (Pedersen and DiDonato 1982), and coastal areas of
    Oregon and Washington (Washington Department of   Within Study Area: This species is found in all study
    Fisheries 1985, Lukas and Carter 1987).               areaestuariesfrom Morro Bay, California (Orcutt 1950),
                                                          north through Washington (Table 1) (Monaco et al.
    Recreational: This species is a fairly important sport   1990).
    fish for anglers from central California to Alaska. It is
    fished year-round from boats, piers, and shore (Frey   Life Mode
    1971) and is captured primarily in estuaries and adjacent    Eggs and larvae are pelagic, while juveniles and adults
    near-shore shallow waters (Beardsley and Bond 1970,    are demersal (Orcutt 1950, Garrison and Miller 1982,
    Squire and Smith 1977, Wydoski and Whitney 1979).    Wang 1986). The starry flounder is unusual in that
    Sport fishermen caught approximately 43,000 starry   alongthecoastsofCalifornia, Oregon, andWashington,
    flounders in 1985 (National Marine Fisheries Service    50% are right-eyed and 50% are left-eyed; in Alaska
    1986).                                                70% are left-eyed, and in Japan nearly 100% are left-
                                                          eyed (Orcutt 1950, Miller 1965, Policansky 1982a).
    Indicator of Environmental Stress: This is a target
    species for the National Status and Trends Program    Habitat
    because it is common  in estuaries and often   IType: Eggs are buoyant and found at the surface in

                                                       266

                                                                                     Starry flounder continued

                                                          euhaline, but may be found in polyhaline waters.
      Table 1. Relative abundance of starry flounder      Juveniles preferbrackish bays (mesohaline) (Pedersen
              in 32 U.S. Pacific coast estuaries.        and DiDonato 1982, Simenstad 1983), but also occur
                                                          in fresh water. Adults occur primarily in euhaline and
                                                          mesohaline waters, but are sometimes found in fresh
            PugEstuary    A S J L E                      water (Hart 1973, Garrison and Miller 1982). This
             PugetSound  Canal         0  Relativeabundance:  species is found at water temperatures from 0.0 to
              SkagitnBay ((  3 0 0    6   ihly Abundant   21.5�C.  Temperatures >28.00C are lethal (Stober
            Grays Harbor O   �0O     O  Common            1973).
             WillapaBay 0   0          Rare
           Columbia River C    li0     Blank Not present  Miarations and Movements: The starry flounder does
            Nehalem Bay O  0                              not migrate extensively (Pedersen and DiDonato 1982).
            Tillamook Bay O    (I O  Life stage:          However, tagging studies have shown that there is
             Netarts Bay   O O      A - Adults            some movement along the coast (Westrheim 1955). It
              letriver B      a     y C D aS - Spawning adults
              SiletzRiver  0           SpawJuveniadults   also has seasonal bathymetric migrations probably
             YaquinaBay O   J3 0C   L-LLarvae             related to spawning.  Adults move inshore during
             Alsea River  13 0      E-Eggs               winter and early spring and offshore during summer
            Siuslaw River 0    0 0                        and fall. Juveniles move far up into rivers, but as they
            Umpqua River O �    O                         mature they tend to reside in estuaries (Morrow 1980).
               Coos Bay O    ( 0
             Rogue River   � I O                          Reproduction
            Klamath River  0 O  Mode: The starry flounder is gonochoristic, oviparous,
            Humboldt Bay 0    O 0                         and iteroparous; eggs are fertilized externally (Orcutt
               Eel River 0 O    0   1950).
            Tomales Bay O     O0
       Cent. San Fran. Bay  i   � O I * IndcudesCentralSan  Matinc/Soawnina:Spawningoccursnearrivermouths
                                    Francisco. Suisun.
       SouthSanFran.Bay     O O     and San Pab bays.    and sloughs in shallow water (<45 m deep) (Orcutt
           Elkhorn Slough o   (1                          1950, Garrison and Miller 1982), apparently at water
              Morro Bay    0                              temperatures of 11 �C (Alaska Department of Fish and
         Santa Monica Bay                                 Game 1986). Spawning may occur in and outside of
           San Pedro Bay                                  San Francisco Bay (Eldridge 1977, Wang 1986).
            Alamitos Bay                                  Spawning takes place primarily from winter to early
            Anaheim Bay                                   spring, depending on area: Novemberto January near
             Newport Bay                                  Elkhorn Slough (Orcutt 1950), and February to April in
             Mission Bay                                  Puget Sound and British Columbia (Smith 1936, Hart
           San Diego Bay                                  1973)
           Tijuana Estuary
                      A S  J L E                          Fecundity: Fecundities range from 900,000 to over 11
                                                          million eggs per female, depending on female size
    nearshore marine waters (Orcutt 1950, Yusa 1957).   (Orcutt 1950, Garrison and Miller 1982).
    Larvae are planktonic and found primarily nearshore
    (within 37 km) and in estuaries (Eldridge and Bryan    Growth and Development
    1972, Waldron 1972, Misitano 1977, Richardson and    Eaa Size and Embryonic Development: Eggs are
    Pearcy 1977).  Juveniles commonly invade far up    spherical and 0.89-1.28 mm in diameter (Orcutt 1950,
    rivers (Moyle 1976), but appear to be estuarine-   Yusa 1957, Garrison and Miller 1982).  Embryonic
    dependent. Adults have been found in marine waters    development is indirect and external. Eggs hatch in
    to 375 m depth, but most are captured at depths    2.8-14.7days,dependingontemperature(Orcutt 1950,
    <150 m (Frey 1971, Allen and Smith 1988).             Yusa 1957).

    Substrate: Eggs and larvae have no substrate   Aae and Size of Larvae: Newly hatched larvae are
    preference. Juveniles and adults prefer soft bottom    1.93-2.08 mm long (Orcutt 1950) or2.58-3.36 mm long
    types (mud, sand, gravel) but not rock (Orcutt 1950,    (Yusa 1957). Larvaetake39-75daystometamorphose
    Pedersen and DiDonato 1982).                          to bottom-dwelling postlarvae (Policansky 1982b).
                                                          Metamorphosis occurs when larvae are 6.6-7.7 mm
    Physical/Chemical Characteristics: Eggs are found in   long (Policansky 1982b).
    euhaline to polyhaline waters. Larvae are primarily

                                                      267

    Starry flounder continued
    Juvenile Size Ranae: Juveniles range in size from    References
    approximately 7 mm (Policansky 1982b) to 17-30 cm
    long, depending on sex and location (Orcutt 1950,    Alaska Department of Fish and Game. 1985. Alaska
    Campana 1984).                                         habitat management guide. Southcentral Region, Vol.
                                                          I: Life histories and habitat requirements of fish and
    Aae and Size of Adults: Males mature in 2 or 3 years at   wildlife. Alaska Dept. Fish Game, Juneau, AK, 429 p.
    17-30 cm in length, while some females mature in 3 or
    4 years at 23-35 cm; all females are mature after 4    Allen, M. J., and G. B. Smith.  1988.  Atlas and
    years (Orcutt 1950, Campana 1984). The maximum    zoogeographyofcommonmarinefishesinthenortheast
    ages reported for males and females are 24 and 17    Pacific Ocean and Bering Sea.  NOAA Tech. Rep.
    years, respectively (Campana 1984), andthe maximum    NMFS 66,151 p.
    size is 91 cm (17 kg) (Orcutt 1950, Hart 1973).
                                                          Bane, G. W., and A. W. Bane. 1971. Bay fishes of
    Food and Feeding                                       northern California. Mariscos Publ., Hampton Bays,
    TroDhic Mode: Larvae are planktivores. Juveniles and    NY, 143 p.
    adults are benthically-oriented carnivores (Orcutt 1950).
    Adults do not feed during the spawning period and    Beardsley, A. J., and C. E. Bond. 1970. Field guide to
    juveniles and adults apparently cease feeding in cold    common marine and bay fishes of Oregon. Agr. Exp.
    temperatures (probably <5�C) (Orcutt 1950, Miller   Sta., Sta. Bull. 607, Oregon State Univ., Corvallis, OR,
    1965).                                                 27 p.

    Eood Items:Larvaeeatphytoplanktonandzooplankton.    Bottom, D. L., K. K. Jones, and M. J. Herring. 1984.
    Small juveniles (<100 mm long) eat copepods and    Fishes of the Columbia River estuary. Col. Riv. Est.
    other small crustaceans. Larger juveniles and adults    Data Dev. Prog., CREST, Astoria, OR,  113 p. plus
    eat amphipods (Corophium spp. and Eogammarus    appendices.
    spp.), isopods, decapods (Crangon spp. and Cancer
    spp.), polychaetes, bivalves (Sliqua spp., Mya arenaria,    Campana, S. E. 1983. Mortality of starry flounders
    Macoma spp., and Yoldiaspp.), echinoderms (Ophiura    (Platichthys stellatus) with skin tumors. Can. J. Fish.
    spp. and Diamphiodia craterodmeta) and occasionally    Aquat. Sci. 40(2):200-207.
    fish, e.g., northern anchovy (Engraulis mordax) (Orcutt
    1950, Miller 1965, Bane and Bane 1971, Jewett and    Campana,  S. E.   1984.   Comparison of age
    Feder 1980, McCabe et al. 1983).                       determination methods for the starry flounder. Trans.
                                                           Am. Fish. Soc. 113:365-369.
    Biological Interactions
    Predation: The starry flounder is eaten by birds [great    Eldridge, M. B. 1977. Factors influencing distribution
    blue heron (Ardea herodias) and cormorants    of fish eggs and larvae over eight 24-hour samplings in
    (Phalacrocorax spp.)] and marine mammals [harbor    Richardson Bay, California. Calif. Fish Game 63(2):101-
    seal (Phoca vitulina) and sea lions] (Simenstad et al.   116.
    1979, Jeffries et al. 1984). To reduce predation,
    juvenilesandadultswillcoverthemselveswithsandor    Eldridge, M. B., and C. F. Bryan. 1972. Larval fish
    mud and change theircolorto match the bottom (Orcutt    surveyof Humboldt Bay, California. NOAA Tech. Rep.
    1950).                                                 NMFS SSRF-665, 8 p.

    Factors Influencina Populations: Contaminants can    Frey, H. W. 1971. California's living marine resources
    impairreproductivesuccess (Whipple etal 1978, Spies    and their utilization.  Calif. Dept. Fish Game,
    et al. 1985) and may cause fin erosion disease and    Sacramento, CA, 148 p.
    lethal skin tumors (Wellings et al. 1976, Campana
    1983).   Endoparasitic flukes and monogenetic    Garrison, K. J., and B. S. Miller. 1982. Review of the
    trematodes have been found on the gills (Bane and    earlylifehistoryofPugetSoundfishes. Fish. Res. Inst.,
    Bane 1971).  Population sizes are probably greatly   Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).
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    Shaw1984). Harvestingbycommercial andrecreational    Gates, D. E., and H. W. Frey.  1974.  Designated
    fishermenmayaffectpopulationsizes. Sincejuveniles    common names of certain marine organisms of
    are found almost exclusively in estuaries, alteration   California. Calif. Fish Game, Fish Bull. 61:55-90.
    and destruction of estuarine habitat undoubtedly affects
    this species population.

                                                       268

                                                                                       Starry flounder continued

    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    Norcross, B. L., and R. F. Shaw. 1984. Oceanic and
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                                                           Simenstad, C. A. 1983. The ecology of estuarine
    Morrow, J. E. 1980. The freshwater fishes of Alaska.    channels of the Pacific Northwest coast: a community
    Alaska Northw. Publ. Co., Anchorage, AK, 248 p.         profile. U.S. Fish Wildl. Serv. FWS/OBS-83/05. 181 p.

    Moyle, P. B. 1976. Inland fishes of California. Univ.    Simenstad, C. A., B. S. Miller, C. F. Nyblade, D.
    Calif. Press, Berkeley, CA, 405 p.                      Thornburgh, and L. J. Bledsoe.  1979.  Food web
                                                           relationships of northern Puget Sound and the Strait of
    National Marine Fisheries Service. 1981. Columbia    Juan de Fuca: a synthesis of the available knowledge.
    River estuary data development program report,    U. S. Interagency (NOAA, EPA) Energy/Environ. Res.
    salmonid and non-salmonid fish. Unpubl. manuscr.,    Dev. Prog. Rep. EPA-600/7-79-259, Washington, D.C.,
    various pagination, Northw. Alaska Fish. Cent., P.O.    335 p.
    Box 155, Hammond, OR, 97121.
                                                           Smith, R. T. 1936. Report on the Puget Sound otter
    National Marine Fisheries Service.  1986.  Marine    trawl investigations.  Wash. Dept. Fish. Biol. Rep.
    recreational fishery statistics survey, Pacific coast,    368:1-61.
    1985. U.S. Dept. Comm., NOAA, Nat. Mar. Fish. Serv.,
    Washington, D.C., 109 p.

                                                        269

    Starry flounder continued
    Spies, R. B., D. W. Rice, Jr., P. A. Montagna, R. R.    ecological impacts of oil spills, p. 757-806. Amer. Inst.
    Ireland, J. S. Felton, S. K. Healy, and P. R. Lewis.    Biol. Sci., Keystone, CO.
    1985. Pollutant body burdens and reproduction in
    PlatichthysstellatusfromSanFranciscoBay. Lawrence    Wilimovsky, N. J. 1964.  Inshore fish fauna of the
    Livermore Nat. Lab., Livermore, CA, 95 p.              Aleutian archipelago. Proc. Alaska Sci. Conf. 14:172-
                                                           190.
    Squire, J. L., and S. E. Smith. 1977. Anglers'guideto
    the United States Pacific coast: Marine fish, fishing    Wydoski, R. S. and R. R. Whitney. 1979. Inland fishes
    grounds, and facilities. NOAA, Seattle, WA, 139 p.    of Washington, Univ. Wash. Press, Seattle, WA,
                                                           220 p.
    Stober, Q. J. 1973. Summary and overview of
    experimental thermal effects studies. In Q. J. Stober    Yusa, T. 1957. Eggs and larvae of flatfishes in the
    and E. O. Salo (editors), Ecological studies of the    coastal waters of Hokkaido, Embryonic development
    proposed Kiket Island nuclear power site, p. 441-448.    of the starry flounder Platichthys stellatus (Pallas).
    Final Rep. to Snohomish County P.U.D. and Seattle   Bull. Hokkaido Reg. Fish. Res. Lab, 15:1-14.
    City Light. Coill. Fish., Fish. Res. Inst., Univ. Wash.,
    Seattle, WA (FRI-UW-7304).

    Waldron, K. D. 1972. Fish larvae collected from the
    northeastern Pacific Ocean and Puget Sound during
    April and May 1967. NOAA Tech. Rep. NMFS SSRF-
    663, 16 p.

    Wang, J. C. S. 1986. Fishes of the Sacramento-San
    Joaquin estuary and adjacent waters, California: A
    guide to the early life histories. Tech. Rep. No. 9.
    Interagency ecological study program for the
    Sacramento-San Joaquin estuary. Calif. Dept. Water
    Res., Calif. Dept. Fish Game, U.S. Bureau Reclam.,
    and U.S. Fish Wildl. Serv., various pagination.

    Washington Department of Fisheries. 1985. 1985
    Fisheries Statistical Report. Wash. Dept. Fish., Olympia,
    WA, 101 p.

    Washington, P. M. 1977. Recreationally important
    marine fishes of Puget Sound, Washington. Proc.
    Rep., Northwest Alaska Fish. Cent., Nat. Mar. Fish.
    Serv., NOAA, Seattle, WA, 122 p.

    Wellings, S. R., C. E. Alpers, B. B. McCain, and B. S.
    Miller. 1976. Fin erosion disease of starry flounder
    (Platichthys stellatus) and English sole (Parophrys
    vetulus) in the estuary of the Duwamish River, Seattle,
    Washington. J. Fish. Res. Board Can. 33:2577-2586.

    Westrheim, S. J. 1955. Migrations of starry flounder
    (Platichthys stellatus) tagged in the Columbia River.
    Oregon Fish. Comm. Res. Briefs 6(1):33-37.

    Whipple, J. A., T. G. Yocom, D. R. Smart, and M. H.
    Cohen. 1978. Effects of chronic concentrations of
    petroleum hydrocarbons on gonadal maturation in
    starry flounder (Platichthys stellatus) [Pallas]. In The
    proceedings of the conference of assessment of

                                                       270

    271

    272

    ABYSSAL ZONE-Ocean bottom at depths between 4,000 and 6,000 m.

    ABYSSOPELAGIC-Living in the water column at depths between 4,000 and 6,000 m; the abyssopelagic zone.

    ADDUCTOR MUSCLE-A muscle that pulls a part of the body toward the median axis of the body. In bivalve
    molluscs, this muscle is used to close the shell halves and hold them together.

    ALEUTIAN PROVINCE-A zoogeographic designation for the area of coastal faunal distributions that, based on
    minimum temperature requirements, extends from Puget Sound, Washington, to the Bering Strait, Alaska.

    ALEVIN-The larval stage of trout and salmon that feeds on its yolk sac and lives under gravel.

    ALGAE-A collective, or general name, applied to a number of primarily aquatic, photosynthetic groups (taxa) of
    plants and plant-like protists. They range in size from single cells to large, multicellularforms like the giant kelps.
    They are the food base for almost all marine animals. Important taxa are the dinoflagellates (division Pyrrophyta),
    diatoms (div. Chrysophyta), green algae (div. Chlorophyta), brown algae (div. Phaeophyta), and red algae (div.
    Rhodophyta). Cyanobacteria are often called blue-green algae, although blue-green bacteria is a preferable term.

    AMPHI-NORTH PACIFIC-A population distribution where a species is distributed on the east and west rims of
    the Pacific Ocean, but not on the northern rim.

    AMPHIPODA-An order of laterally compressed crustaceans with thoracic gills, no carapace, and similar body
    segments. Although most are <1 cm long, they are an important component of zooplankton and benthic
    invertebrate communities. A few species are parasitic.

    ANADROMOUS-Life cycle where an organism spends most of its life in the sea, and migrates to freshwater to
    spawn.

    ANTHROPOGENIC-Refers to the effects of human activities.

    ARCTIC REGION-The oceans north of the 0�C winter isotherm. Along the Pacific coast, this corresponds to 60�
    N in the Bering Sea.

    AREAL-Refers to a measure of area.

    ASCIDIAN-A tunicate (class Ascidiacea) that has a generalized sac-like, cellulose body and is usually attached
    to the substratum.

    BATHYAL-The zone of ocean bottom at depths of 200 to 4,000 m, primarily on the continental slope and rise.

    BATHYMETRIC-A depth measurement. Also refers to a migration from waters of one depth to another.

    BATHYPELAGIC-Ocean depths from 1,000 to 4,000 m.

    BENTHIC-Pertaining to the bottom of an ocean, lake, or river. Also refers to sessile and crawling animals which
    reside in or on the bottom.

    BIGHT-An inward bend or bow in the coastline.

    BIOMASS-The total mass of living tissues (wet or dried) of an organism or collection of organisms of a species
    or trophic level, from a defined area or volume.

                                                       273

    Glossary continued

    BIVALVIA-Bilaterally symmetrical molluscs (also referred to as Pelecypoda) that have two lateral calcareous
    shells (valves) connected by a hinge ligament. They are mostly sedentary filterfeeders. This class includes clams,
    oysters, scallops, and mussels.

    BOREAL REGION-The oceans of the northern hemisphere between the 0 and1 3�C winter isotherms. In neritic
    waters of western North America, it extends from Point Conception, California, to the southern Bering Sea, Alaska.

    BRANCHIAL-A structure or location on an organism associated with the gills.

    BRYOZOA-Minute, moss-like colonial animals of the phylum Bryozoa.

    BYSSAL THREAD-A tuft of filament, chemically similar to silk, that attaches certain molluscs to substrates.

    CALCAREOUS-Composed of calcium or calcium carbonate.

    CARNIVORE-An animal that feeds on the flesh of other animals. See PARASITISM and PREDATION.

    CESTODE-A parasitic, ribbon-like worm having no intestinal canal; class Cestoda (e.g., tapeworms).

    CHEMOTAXIS-A response movement by an animal either toward or away from a specific chemical stimulus.

    CHORDATA-A phylum of animals which includes the subphyla Vertebrata, Cephalochordata, and Urochordata.
    At some stage of their life cycles, these organisms have pharyngeal gill slits, a notochord, and a dorsal, hollow
    nerve cord.

    CILIA-Hair-like processes of certain cells, often capable of rhythmic beating that can produce locomotion or
    facilitate the movement of fluids.

    CIRRI-Flexible, thread-like tentacles or appendages of certain organisms.

    CLINE-A series of differing physical characteristics within a species or population, reflecting gradients or
    changes in the environment (e.g., body size or color).

    COLONY-A group of organisms living in close proximity. An invertebrate colony is a close association of
    individuals of a species which are often mutually dependent and in physical contact with each other. A vertebrate
    colony is usually a group of individuals brought together for breeding and rearing young.

    COMMENSALISM-A relationship between two species, where one species benefits without adversely affecting
    the other.

    COMMUNITY-A group of plants and animals living in a specific region under relatively similarconditions. Further
    restrictions are often used, such as the algal community, the invertebrate community, the benthic gastropod
    community, etc.

    COMPETITION-Two types exist - interspecific and intraspecific. Interspecific competition exists when two or
    more species use one or more limited resources such as food, attachment sites, protective cover, or dissolved
    ions. Intraspecific competition exist when individuals of a single species compete for limited resources needed
    for survival and reproduction. This form of competition includes the same resources involved in interspecific
    competition as well as mates and territories. It is generally more intense than interspecific competition because
    resource needs are essentially identical among conspecifics. See NICHE.

    CONGENER-Referring to members of the same genus.

    CONTINENTAL SHELF-The submerged continental land mass, not usually deeper than 200 m. The shelf may
    extend from a few miles off the coastline to several hundred miles.

                                                        274

                                                                                           Glossary continued

    CONTINENTAL SLOPE-The steeply sloping seabed that connects the continental shelf and continental rise.

    COPEPODA-A subclass of crustaceans with about 4,500 species, including several specialized parasitic orders.
    The free-living species are small (one to several mm) and have cylindrical bodies, one median eye, and two long
    antennae. One order is planktonic (Calanoida), one is benthic (Harpacticoida), and one has both planktonic and
    benthic species (Cyclopoida). In most species, the head appendages form a complex apparatus used to sweep
    in and possibly filter prey (especially algae). Thoracic appendages are used for swimming or crawling on the
    bottom. One of the most abundant group of animals on earth, they are a major link in aquatic food webs.

    CREPUSCULAR-Relates to animals whose peak activity is during the twilight hours of dawn and dusk.

    CRUSTACEA-A large class of over 26,000 species of mostly aquatic arthropods having five pairs of head
    appendages, including laterally opposed jaw-like mandibles and two pairs of antennae. Most have well-developed
    compound eyes and variously modified two-branched body appendages. The body segments are often
    differentiated into a thorax and an abdomen. Some common members are crabs, shrimp, lobsters, copepods,
    amphipods, isopods, and barnacles.

    CTENIDIA-The comblike respiratory apparatus of molluscs.

    CTENOPHORA-A phylum of mostly marine animals that have oval, jellylike bodies bearing eight rows of comb-
    like plates that aid swimming (e.g., ctenophores and comb jellies).

    DECOM POSERS-Bacteria and fungi that break down dead organisms of all types to simple molecules and ions.

    DEMERSAL-Refers to swimming animals that live near the bottom of an ocean, river, or lake. Often refers to
    eggs that are denser than water and sink to the bottom after being laid.

    DEPOSIT FEEDER-An animal that ingest small organisms, organic particles, and detritus from soft sediments,
    or filters organisms and detritus from such substrates.

    DESICCATE-To dry completely.

    DETRITIVORE-An organism that eats small fragments of partially decomposed organic material (detritus) and
    its associated microflora. See DECOMPOSER.

    DIATOMS-Single-celled protistan algae of the class Bacillariophyceae that have intricate siliceous shells
    composed of two halves. They range in size from about 10 to 200 microns. Diatoms sometimes remain attached
    after cellular divisions, forming chains or colonies. These are the most numerous and important group of
    phytoplankters in the oceans, and form the primary food base for marine ecosystems.

    DIEL-Refers to a 24-hour activity cycle based on daily periods of light and dark.

    DIMORPHISM-A condition where a population has two distinct physical forms (morphs). In sexual dimorphism,
    secondary sexual characteristics are markedly different (e.g., size, color, and behavior).

    DINOFLAGELLATE-A planktonic, photosynthetic, unicellular algae that typically has two flagella, one being in
    a groove around the cell and the other extending from the center of the cell.

    DIRECT DEVELOPMENT-See EMBRYONIC DEVELOPMENT.

    DISPERSAL-The spreading of individuals throughout suitable habitat within or outside the population range. In
    a more restricted sense, the movement of young animals away from their point of origin to locations where they
    will live at maturity.

    DISSOCHONCH-The adult shell secreted by newly-settled clam larvae or plantigrades.

                                                       275

    Glossary continued

    DISTRIBUTION-(1) A species distribution is the spatial pattern of its population or populations over its
    geographic range. See RANGE. (2) A population depth distribution is the proportion or number of all individuals,
    or those of various sizes or ages, at different depth strata. (3) A population age distribution is the proportions of
    individuals in various age classes. (4) Within a population, individuals may be distributed evenly, randomly, or in
    groups throughout suitable habitat.

    DIURNAL-Refers to daylight activities, or organisms most active during daylight. See DIEL.

    ECHINODERMATA-A phylum of radially-symmetrical marine animals, possessing a watervascular system, and
    a hard, spiny skeleton (e.g., sea stars, sea urchins, and sand dollars).

    ECTOPARASITE-A parasite that attacks (and usually attaches to) a host animal or plant on the outside. Feeding
    periods and/or attachment time may be brief compared to internal (endo-) parasites.

    EELGRASS-Vascular flowering plants of the genus Zostera that are adapted to living under water while rooted
    in shallow sediments of bays and estuaries.

    EL NINO10 CURRENT-An intermittent warm watercurrent fromthe tropics that overrides the opposing cold current
    along the Pacific coasts of North and South America (see GYRE). This raises near-surface temperatures,
    depresses the thermocline, and often suppresses upwelling, resulting in drastic drops in primary productivity and
    reduced recruitment of marine animals. This is most pronounced on the coast of Peru. Effects are not as severe
    in North America, but northward shifts in distributions of "southern" species are common in El Niflo years.

    EMBRYONIC DEVELOPMENT-The increase in cell number, body size, and complexity of organ systems as an
    individual develops from a fertilized egg until hatching or birth. In direct development, individuals at birth or
    hatching are essentially miniatures of the adults. In indirect development, newly hatched individuals differ greatly
    from the adult, and go through periodic, major morphological changes (larval stages and metamorphosis) before
    becoming a juvenile.

    EMIGRATION-A movement out of an area by members of a population. See IMMIGRATION.

    ENDEMIC-Refers to a species or taxonomic group that is native to a particular geographical region.

    EPIBENTHIC-Located on the bottom, as opposed to in the bottom.

    EPIDERMAL-Refers to an animal's surface or outer layer of skin.

    EPIFAUNA-Animals living on the surface of the bottom.

    EPIPELAGIC-The upper sunlit zone of oceanic water where phytoplankton live and organic production takes
    place (approximately the top 200 m). See EUPHOTIC.

    EPIPHYTIC-Refers to organisms which live on the surface of a plant (e.g., mosses growing on trees).

    EPIPODAL-A structure or location associated with the leg or foot; typically refers to arthropod anatomy.

    ESCARPMENT-A steep slope in topography, as in a cliff or along the continental slope.

    ESTUARY-A semi-enclosed body of water with an open connection to the sea. Typically there is a mixing of sea
    and fresh water, and the influx of nutrients from both sources results in high productivity.

    EUHALIN E-Water with salt concentrations of 30-40%,.

    EUPHOTIC-Refers to the upper surface zone of a water body where light penetrates and phytoplankton (algae)
    carry out photosynthesis. See EPIPELAGIC.

                                                        276

                                                                                           Glossary continued

    EURYHALINE-Refers to an organism that is tolerant of a wide range of salinities.

    EURYTHERMAL-Refers to an organism that is tolerant of a wide range of temperatures.

    EXTANT-Existing or living at the present time; not extinct.

    FAUNA-All of the animal species in a specified region.

    FECUNDITY-The potential of an organism to produce offspring (measured as the number of gametes). See
    REPRODUCTIVE POTENTIAL.

    FILTER FEEDER-Any organism that filters small animals, plants, and detritus from water or fine sediments for
    food. Organs used for filtering include gills in clams and oysters, baleen in whales, and specialized appendages
    in crustaceans and marine worms.

    FINGERLING-Refers to a small juvenile fish (often a salmonid) that is about 100 mm long.

    FLAGELLATE-Refers to cells that have motility organelles or microorganisms that possess one or more
    flagellum used for locomotion.

    FLORA-All of the plant species in a specified region

    FOOD WEB (CHAIN)-The feeding relationships of several to many species within a community in a given area
    during a particulartime period. Two broad types are recognized: 1 ) grazing webs involving producers (e.g., algae),
    herbivores (e.g., copepods), and various combinations of carnivores and omnivores, and 2) detritus webs
    involving scavengers, detritivores, and decomposers that feed onthe dead remains ororganisms from the grazing
    webs, as well as on their own dead. A food chain refers to organisms on different trophic levels, while a food web
    refers to a network of interconnected food chains. See TROPHIC LEVEL.

    FOULING--Occurs when large numbers of plants or animals attach and grow on various structures (floats, pipes,
    and pilings), often interfering with their use. Fouling organisms include barnacles, mussels, bryozoans, and
    sponges.

    FRESH WATER-Water that has a salt concentration of 0.0-0.5%0.

    FRY-Very young fish. For trout and salmon, they are young that have just emerged from the gravel and are
    actively feeding.

    GAMETE-A reproductive cell. When two gametes unite they form an embryonic cell (zygote).

    GASTROPODA-The largest class of the Phylum Mollusca. This group includes terrestrial snails and slugs as
    well as aquatic species such as whelks, turbans, limpets, conchs, abalones, and nudibranchs. Most have external
    shells that are often spiraled (but this has been lost or is reduced in some), and move on a flat, undulating foot.
    They are mostly herbivorous and scrape food with a radula, an organ analogous to a tongue.

    GONOCHORISTIC-Refers to a species that has separate sexes (i.e., male and female individuals).

    GROUNDFISH-Fish species that live on or near the bottom, often called bottomfish.

    GYRE-An ocean current that follows a circular or spiral path around an ocean basin, clockwise in the northern
    hemisphere and counterclockwise in the southern hemisphere.

    HABITAT-The particular type of place where an organism lives within a more extensive area or range. The
    habitat is characterized by its biological components and/or physical features (e.g., sandy bottom of the littoral
    zone, or on kelp blades within 10 m of the water surface).

                                                       277

    Glossary continued

    HAPLOSPORIDIAN-A unicellular protozoan occurring in vertebrate and invertebrate hosts, often causing
    disease.

    HERBIVORE-An animal that feeds on plants (phytoplankton, large algae, or higher plants).

    HERMAPHRODITIC-Refers to an organism having both male and female sex organs on the same individual.

    HOLARCTIC-The entire Arctic, including the Paleoarctic (Europe and Asia) and the Nearctic (North America).
    Also, the entire arctic region in oceanography.

    HYDROZOA-A class of the phylum Cnidaria. The primary life stage is nonmotile and has a sac-like body
    composed of two layers of cells and a mouth that opens directly into the body cavity. A second life stage, the free-
    living medusa, often resembles the common jellyfish.

    HYPERSALINE-Water with a salt concentration over 40%0.

    HYPOLIMNION-The cold bottom water zone of a lake below the thermocline.

    IMMIGRATION-A movement of individuals into a new population or region. See EMIGRATION, MIGRATION,
    and RECRUITMENT.

    INCIDENTAL CATCH-Catch of a species that was not the focus of a fishery, but taken along with the species
    being sought.

    INDIRECT DEVELOPMENT-See EMBRYONIC DEVELOPMENT.

    INFAUNA-Animals living in bottom substrates.

    INNER SHELF-The continental shelf extending from the mean low tide line to a depth of 20 m.

    INSTAR- The intermolt stage of a young arthropod.

    INSULAR-Of or pertaining to an island or its characteristics (i.e., isolated).

    INTERTIDAL-The ocean or estuarine shore zone exposed between high and low tides.

    ISOBATH-A contour mapping line that indicates a specified constant depth.

    ISOPODA-An order of about 4,000 species of dorsoventrally compressed crustaceansthat have abdominal gills
    and similar abdominal and thoracic segments. Terrestrial pillbugs and thousands of benthic marine species are
    included. Most species are scavengers and/or omnivores; a few are parasitic.

    ISOTHERM-A contour line connecting points of equal mean temperature for a given sampling period.

    ITEROPAROUS-Refers to an organism that reproduces several times during its lifespan (i.e., does not die after
    spawning).

    KELT-A spent (i.e., spawned out) trout.

    KINESIS-A randomly directed movement by an animal in response to a sensory stimulus such as light, heat, or
    touch. When the response is directed, it is called a taxis. See CHEMOTAXIS.

    LACUSTRINE-Pertaining to, or living in, lakes or ponds.

    LAGOON-A shallow pond or channel linked to the ocean, but often separated by a reef or sandbar.

                                                      278

                                                                                          Glossary continued

    LARVAE-An early developmental stage of an organism that is morphologically different from the juvenile or adult
    form. See EMBRYONIC DEVELOPMENT.

    LATERAL LINE-A pressure sensory system located in a line of pores underthe skin on both sides of most fishes.
    The system is connected indirectly with the inner ear and senses water pressure changes due to water movement
    (including sound waves).

    LITTORAL-The shore area between the mean low and high tide levels. Water zones in this area include the
    littoral pelagic zone and the littoral benthic zone.

    MANTLE-The upper fold of skin in molluscs that encloses the gills and most of the body in a cavity above the
    muscular foot. In squids and allies, the mantle is below the body and behind the tentacles (derived from the foot)
    due to the shift in the dorsal-ventral axis. The mantle produces the shell in species having them.

    MEAN LOWER LOW WATER (MLLW)-The arithmetic mean of the lower low water heights of a mixed tide over
    a specific 19-year Metonic cycle (the National Tidal Datum Epoch). Only the lower low water of each tidal day is
    included in the mean.

    MEGALOPAE-The larval stage of a crab characterized by an adult-like abdomen, thoracic appendages, and a
    developed carapace.

    MEIOFAUNA-Very small animals, usually < 0.5 mm in diameter.

    MERISTIC-Refers to countable measurements of segments or features such as vertebrae, fin rays, and scale
    rows. Counts of these are used in population comparisons and classifications.

    MESOHALINE-Water with a salt concentration of 5-18%o.

    MESOPELAGIC-Ocean zone of intermediate depths from about 200-1,000 m below the surface, where light
    penetration drops rapidly and ceases.

    METAMORPHOSIS-Process of transforming from one body form to another form during development (e.g.,
    tadpole changing to a frog). See EMBRYONIC DEVELOPMENT.

    METRIC TON (t)-A unit of mass or weight equal to 2,204.6 lb.

    MIGRATION-Movement by a population or subpopulation from one location to another (often periodic or
    seasonal, and over long distances). Vertical migrations in the water column may be daily or seasonal within the
    same area. Migrations between deep and shallow areas are usually seasonal and related to breeding. Many
    marine birds and mammals have seasonal latitudinal migrations associated with breeding. See EMIGRATION,
    IMMIGRATION, RANGE, and RECRUITMENT.

    MILT-The seminal fluid and sperm of male fish.

    MOLT-The process of shedding and regrowing an outer skeleton or covering at periodic intervals. Crustaceans
    and other arthropods molt their exoskeletons, grow rapidly, and produce larger exoskeletons. Most reptiles, birds,
    and mammals, molt skin, feathers, and fur, respectively.

    MORPHOLOGY-The appearance, form, and structure of an organism.

    MORPHOMETRICS-The study of comparative morphological measurements.

    MORTALITY-Death rate expressed as a proportion of a population or community of organisms. Mortality is
    caused by a variety of sources, including predation, disease, environmental conditions, etc.

                                                      279

    Glossary continued

    MOTILE-Capable of or exhibiting movement or locomotion.

    M UTUALISM-An interaction between two species where both benefit. Some authorities conside rtrue mutual ism
    to be obligatory for both species, while mutually beneficial relationships that are not essential for either species
    are classified as protocooperative (e.g., the blacksmith cleaning fish eating external parasites from sea basses).

    NACREOUS MATERIAL-A calcareous, lustrous secretion in the inner surface of the shell of many molluscs.
    Foreign particles lodging between the inner shell surface and the mantle are covered by nacre, sometimes forming
    pearls.

    NANOPLANKTON-Microscopic, planktonic organisms smaller than 20 microns in diameter.

    NATAL-Pertaining to birth or hatching.

    NEKTONIC-Refers to pelagic animals that are strong swimmers, live above the substrate in the water column,
    and can move independently of currents.

    NEMERTEA-A phylum of unsegmented, elongate marine worms having a protrusible proboscis and no body
    cavity, and live mostly in coastal mud or sand; nemerteans.

    NERITIC-An oceanic zone extending from the mean low tide level to the edge of the continental shelf. See
    INNER SHELF, LITTORAL, and OCEANIC ZONES.

    NEUSTON-Organisms that live on or under the water surface, often dependent on surface tension for support.

    NICHE-The fundamental niche is the full range of abiotic and biotic factors under which a species can live and
    reproduce. The realized niche is the set of actual conditions under which a species or a population of a species
    exists, and is largely determined by interactions with other species.

    NOCTURNAL-Refers to night, or animals that are active during night.

    OCEANIC-Living in or produced by the ocean.

    OCEANIC ZONE-Pelagic waters of the open ocean beyond the continental shelf. See BATHYPELAGIC,
    EPIPELAGIC, ABYSSOPELAGIC, MESOPELAGIC, and NERITIC.

    OLIGOHALINE-Water with a salt concentration of 0.5-5.0%0.

    OMNIVORE-An animal that eats both plants and animals.

    OOCYTES-The cells in ovaries that will mature into eggs.

    OREGON PROVINCE-A zoogeographical designation for faunal distributions that extends from Cape Flattery,
    Washington, to Point Conception, California.

    OTOLITHS-Small calcareous nodules located in the inner ear of fishes used for sound reception and
    equilibration. They are often used by biologists to assess daily or seasonal growth increments.

    OUT-M IGRATION-Movement of animals out of or awayfrom an area (e.g., juvenile salmonids moving from rivers
    to the ocean).

    OVIGEROUS-The condition of being ready to release mature eggs; egg-bearing.

    OVIPAROUS-Refers to animals that produce eggs that are laid and hatch externally. See OVOVIVIPAROUS
    and VIVIPAROUS.

                                                      280

                                                                                          Glossary continued

    OVIPOSITION-The process of placing eggs on or in specific places, as opposed to randomly dropping or
    broadcasting them.

    OVOVIVIPAROUS-Refers to animals whose eggs are fertilized, developed, and hatched inside the female, but
    receive no nourishment from her. See OVIPAROUS and VIVIPAROUS.

    PALP-An organ attached to the head appendages of various invertebrates; usually associated with feeding
    functions.

    PARASITISM-An obligatory association where one species (parasite) feeds on, or uses the metabolic
    mechanisms of the second (host). Unlike predators, parasites usually do not kill their hosts, although hosts may
    later die from secondary causes that are related to a weakened condition produced by the parasite. Parasitism
    may also be fatal when high parasite densities develop on or in the host.

    PARR-The freshwater life stage of juvenile salmon and trout that has a series of dark, vertical bars on its sides
    (parr marks).

    PARTURITION-The act of giving birth. See SPAWN.

    PATHOGEN-A microorganism or virus that produces disease and can cause death.

    PEDIVELIGER-The larval stage of bivalves during which a functional pedal (footlike) organ develops.

    PELAGIC-Pertaining to the water column, or to organisms that live in the water column.

    PELAGIVORE-A carnivore that feeds in the water column.

    PHYLOGENY-Refers to evolutionary relationships and lines of descent.

    PHYTOPLANKTON-Microscopic plants and plant-like protists (algae) of the epipelagic and neritic zones that are
    the base of offshore food webs. They drift with currents, but usually have some ability to control their level in the
    water column. See ALGAE and DIATOMS.

    PISCIVOROUS-Refers to a carnivorous animal that eats fish.

    PLANKTIVOROUS-Refers to an animal that eats phytoplankton and/or zooplankton.

    PLANKTON-See PHYTOPLANKTON and ZOOPLANKTON.

    PLANTIGRADE-A young, newly settled post-larval clam.

    PLEOPODS-Paired swimming appendages on the abdomen of crustaceans.

    POLYCHAETA-A class of segmented, mostly marine, annelid worms that bear bristles and fleshy appendages
    on most segments.

    POLYHALINE-Water with a salt concentration between 18 and 307/o.

    POPULATION-All individuals of the same species occupying a defined area during a given time. Environmental
    barriers may divide the population into local breeding units (demes) with restricted immigration and interbreeding
    between the localized units. See SPECIES, SUBSPECIES, and SUBPOPULATION.

    PREDATION-An interspecific interaction where one animal species (predator) feeds on another animal or plant
    species (prey) while the prey is alive or after killing it. The relationship tends to be positive (increasing) for the

                                                      281

    Glossary continued

    predator population and negative (decreasing) for the prey population. See PARASITISM, SYMBIOTIC,
    CARNIVORE, and TROPHIC LEVEL.

    PRODUCTION-Gross primary production is the amount of light energy converted to chemical energy in the form
    of organic compounds by autotrophs like algae. The amount left after respiration is net primary production and
    is usually expressed as biomass or calories/unit area/unit time. Net production for herbivores and carnivores is
    based on the same concept, except that chemical energy from food, not light, is used and partially stored for life
    processes. Efficiency of energy transfers between trophic levels ranges from 10-65% (depending on the organism
    and trophic level). Organisms at high trophic levels have only a fraction of the energy available to them that was
    stored in plant biomass. After respiration loss, net production goes into growth and reproduction, and some is
    passed to the next trophic level. See FOOD WEB and TROPHIC LEVEL.

    PROKARYOTIC-Organisms that have nuclear bodies, but lack chromosomes, nucleoli, and nuclear mem-
    branes.

    PROTANDRY-A type of hermaphroditism in which an individual initially develops as a male, then reverses to
    function as a female. Common for some species of shrimp.

    PROTISTAN-Pertaining to the eukaryotic unicellular organisms of the kingdom Protista, including such groups
    as algae, fungi, and protozoans.

    PROTOZOA-A varied group of either free-living or parasitic unicellular flagellate and amoeboid organisms.

    PYCNOCLINE--A zone of marked water density gradient that is usually associated with depth.

    RACE-An intraspecific group or subpopulation characterized by a distinctive combination of physiological,
    biological, geographical, or ecological traits. In salmonids, a race is determined by when it returns to its natal
    stream.

    RADULA-A toothed belt or tongue in the buccal cavity of most molluscs that is used to scrape food particles from
    a surface, or modified otherwise to serve a variety of feeding habits.

    RANGE-(1) The geographic range is the entire area where a species is known to occur or to have occurred
    (historical range). The range of a species may be continuous, or it may have unoccupied gaps between
    populations (discontinuous distribution). (2) Some populations, orthe entire species, may have different seasonal
    ranges. These may be overlapping, orthey may bewidely separated with intervening areas that are at most briefly
    occupied during passage on relatively narrow migration routes. (3) Home range refers to the local area that an
    individual or group uses for a long period or life. See DISTRIBUTION and TERRITORY.

    RECRUITMENT-The addition of new members to a population or stock through successful reproduction and
    immigration.

    RED TIDE-A reddish coloration of sea waters caused by a large bloom of red flagellates. The accumulation of
    metabolic by-products from these organisms is toxic to fish and many other marine species. The accumulation
    of these metabolites in shellfish makes shellfish toxic to humans.

    REDD-A gravel nest dug by spawning female salmon and trout. After eggs are released and fertilized by the
    male, the female covers them with gravel by sweeping movements of the tail.

    REPRODUCTIVE POTENTIAL-The total number of offspring possible for a female of a given species to produce
    if she lives to the maximum reproductive age. This is found by multiplying the number of possible reproductive
    periods by the average number of eggs or offspring produced by females of each age class. This potential is
    seldom realized, but this and the age of first reproduction, or generation time, determine the maximum rate of
    population increase under ideal conditions.

                                                        282

                                                                                          Glossary continued

    RESIDUALISM-Occurswhen juvenile salmon smolts do not migrate to sea but revert backto parr, usually loosing
    their ability to osmoregulate in seawater.

    RHEOTAXIS-A response movement by an animal toward or away from stimulation by a water current.

    RIVERINE-Pertaining to a river or formed by a river or stream.

    ROE-The egg-laden ovary of fish, or the egg mass of certain crustaceans.

    RUN-A group of migrating fish (e.g., a salmon run).

    SALT WEDGE-A wedge-shaped layer of salt water that intrudes upstream beneath a low-density freshwater lens
    that has "thinned" while flowing seaward.

    SAN DIEGO PROVINCE-A zoogeographical designation for faunal distributions that, based on minimum
    temperature requirements, extends from Point Conception, California, to Magdalena Bay, Baja California Sur.

    SCAVENGER-Any animal that feeds on dead animals and remains of animals killed by predators. See
    DECOMPOSER and DETRITIVORE.

    SEAMOUNT-An undersea mountain rising more than 3000 feet (914 m) from the sea floor, but having a summit
    at least 1000 feet (305 m) below sea level (in contrast to an island).

    SEDENTARY-Refers to animals that are attached to a substrate or confined to a very restricted area (or those
    that do not move or move very little). See SESSILE.

    SEMELPAROUS-Animals that have a single reproductive period during their lifespan.

    SESSILE-Refers to an organisms that is permanently attached to the substrate. See SEDENTARY.

    SETTLEMENT-The act of or state of making a permanent residency. Often refers to the period when fish and
    invertebrate larvae change from a planktonic to a benthic existence.

    SHOAL-(1I) A sand bar in a body of water that is exposed at low tide. (2) An area of shallow water. (3) A group
    of fish (school). (4) As a verb, to collect in a crowd or school.

    SIPHONS-The "necks" or tubes of clams and other bivalves that carry water containing food and oxygen into
    the gills, and then expels water containing waste products (exhalent siphon).

    SLOUGH--Ashallow inlet orbackwaterwhose bottom may be exposed at lowtide. Sloughs often borderestuaries
    and typically have a stream passing through them.

    SMOLT-A juvenile salmon or anadromoustrout that is in the process of migrating to the ocean and physiologically
    adapting to seawater. Smolts are usually very silvery and have very faint parr marks. See PARR.

    SPAT-Juvenile bivalve molluscs which have settled from the water column to the substrate to begin a benthic
    existence.

    SPAWN-The release of eggs and sperm during mating. Also, the bearing of offspring by species with internal
    fertilization. See PARTURITION.

    SPECIES-(1) A fundamental taxonomic group ranking after a genus. (2) A group of organisms recognized as
    distinct from other groups, whose members can interbreed and produce fertile offspring. See POPULATION,
    SUBPOPULATION, and SUBSPECIES.

                                                       283

    Glossary continued

    SPERMATOPHORE-A capsule orgelatinous packet (extruded by a male) containing sperm and used to transfer
    sperm to females. Spermatophores are produced by certain invertebrates and some primitive vertebrates.

    SPIROCHETE-A spiral-shaped, non-flagellated bacterium of the order Spirochaetales. This group can be free-
    living or parasitic. Some members cause diseases.

    SPIT-A long, narrow sand bar or peninsula extending into a body of water which is at least partly connected to
    the shore. See SHOAL.

    SPOROCYST-A simple larval stage of parasitic trematode worms. Contact with the host causes a metamorpho-
    sis from an earlier stage to this stage.

    STENOHALINE-Pertaining to organisms that are restricted to a narrow range of salinities, in contrast to
    EURYHALINE.

    STIPE-A thickened, stalk-like structure in kelps that bears other structures, such as blades. Also, the basal
    portion of the thallus or plant body of alga.

    STOCK-A related group or subpopulation. See POPULATION and SUBPOPULATION.

    SUBADULTS-Maturing individuals that are not yet sexually mature.

    SUBLITTORAL-The benthic zone along a coast, or lake that extends from mean low tide to depths of about
    200 m.

    SUBPOPULATION-A breeding unit (deme) of a larger population. These units may differ little genetically and
    taxonomically. See SUBSPECIES. Subpopulations may intergrade with some interbreeding, orthey may occupy
    a common seasonal range prior to the mating season. The units may have different reproduction times and be
    separated spatially or temporally. See RACE, STOCK, and POPULATION.

    SUBSPECIES-A taxonomic class assigned to populations and/or subpopulations when interbreeding (gene
    flow) between populations is limited, and there are significant differences in some combination of characteristics
    between subspecies (e.g., appearance, anatomy, ecology, physiology, and behavior). While successful
    interbreeding can occurwhen the groups are in contact, undernatural conditions reproductive isolation is complete
    and the groups are considered distinct. Classification of such groups is based on the comparative study and
    judgement of phylogenists. A second epithet for each subspecies is added to the binomial for the species (e.g.,
    Oncorhynchus clarki clarki). See SPECIES, POPULATION, and SUBPOPULATION.

    SUBTIDAL-See SUBLITTORAL.

    SUPRALITTORAL-The splash zone of land (adjacent to the sea) that is above the mean high tide level.

    SUSPENSION FEEDER-An animal that feeds directly or by filtration on minute organisms and organic debris
    that is suspended in the water column.

    SYMBIOSIS-The relationship between two interacting organisms that is positive, negative, or neutral in its
    effects on each species. See COMPETITION, MUTUALISM, PARASITISM, and PREDATION.

    TAXONOMY-A system of describing, naming, and classifying animals and plants into related groups based on
    common features (e.g., structure, embryology, and biochemistry).

    TEMPERATE REGION-Oceanic waters between the 13 and 200C winter isotherms. The temperate region of
    the neritic zone on the Pacific coast of North America extends from Point Conception, California, to Magdalena
    Bay, Baja California Sur.

                                                      284

                                                                                           Glossary continued

    TEMPORAL-Pertaining to time. Used to describe organism activities, developmental stages, and distributions
    as they relate to daily, seasonal, or geologic time periods.

    TERRITORY-An area occupied and used by an individual, pair, or larger social group, and from which other
    individuals orgroups of the species are excluded, often with the aid of auditory, olfactory, and visual signals, threat
    displays, and outright combat.

    TEST-A rigid calcareous exoskeleton produced by some echinoderms in the class Echinoidea (e.g., sea urchins
    and sand dollars).

    THERMOCLINE-A relatively narrow boundary layer of water where temperature decreases rapidly with depth.
    Little water or solute exchange occurs across the thermocline, which is maintained by solar heating of the upper
    water layers.

    TREMATODA-A class of parasitic flatworms of the phylum Platyhelminthes. Trematodes have one or more
    muscular, external suckers and are also known as flukes.

    TRIPLOIDY-The occurrence of threetimes the haploid numberof chromosomes. When genetically engineered,
    randomly occurring traits may be selected for commercial applications. For example, the Pacific oyster
    experiences a degradation in flesh quality associated with spawning. Non-reproducing triploid cultures avoid this
    seasonal problem.

    TROCHOPHORE-A molluscan larval stage (except in Cephalopoda) following gastrulation (embryonic stage
    characterized bythe development of a simple gut). It is commonly ciliated, biconically shaped, and free-swimming;
    it establishes an evolutionary link between annelids and molluscs, since both groups display a similar life stage.

    TROPHIC LEVEL-The feeding level in an ecosystem food chain characterized by organisms that occupy a
    similar functional position. At the first level are autotrophs or producers (e.g., kelps and diatoms); at the second
    level are herbivores (e.g., copepods and snails); at the third level and above are carnivores (e.g., salmon and
    seals). Omnivores feed atthe second and third levels. Decomposers and detritivores may feed at all trophic levels.
    See FOOD WEB and PRODUCTION.

    TROPICAL REGION-Oceanic waters between the 20�C winter isotherms in the southern and northern
    hemispheres. Tropical neritic waters along the west coasts of North and South America extend from the southern
    tip of Baja California, Mexico, to about lat. 50S along the coast of Peru.

    TURBELLARIA-A class of mostly aquatic, non-parasitic flatworms that are leaf-shaped and covered with cilia.

    UPWELLING-The process whereby prevailing seasonal winds create surface currents that allow nutrient rich
    cold water from the ocean depths to move into the euphotic or epipelagic zone. This process breaks down the
    thermocline and increases primary productivity, and ultimately fish abundance.

    VELICONCHA-A bivalve larval stage. A veliconcha has two larval shells and moves by using its velum.

    VELIGER-A ciliated larval stage common in molluscs. This stage forms afterthe trochophore larva and has some
    adult features, such as a shell and foot.

    VELUM-The ciliated swimming organ of a larval mollusc.

    VIVIPAROUS-Refers to animals that produce live offspring; eggs are retained and fertilized in the female (as
    compared to OVIPAROUS).

    WATER COLUMN-The water mass between the surface and the bottom.

                                                       285

    Glossary continued

    YEAR-CLASS-Refers to animals of a species population hatched or born in the same year at about the same
    time; also known as a cohort. Strong year-classes result when there is high larval and juvenile survival; the reverse
    is true forweak year-classes. The effects of strong and weak year-classes on population size and structure may
    persist for years in species with long lives. Variation in year-class strength often affects fisheries. See
    DISTRIBUTION and STOCK.

    ZOEA-An early larval stage of various marine crabs and shrimp; zoea have many appendages and long dorsal
    and anterior spines.

    ZOOPLANKTON-Animal members of the plankton. Most range in size from microscopic to about 2.54 cm in
    length. They reside primarily in the epipelagic zone and feed on phytoplankton and each other. Although they
    have only a limited ability to swim against currents, many undertake diel migrations. Taxa include protozoa,
    jellyfish, comb jellies, arrowworms, lowerchordates, copepods, water fleas, krill, and the larvae of many fish and
    invertebrates that are not planktonic as adults.

                                                        286

                                       Appendices

    Appendix 1: Summary table example: Spatial distribution and relative abundance

    Appendix 2: Summary table example: Temporal distribution

    Appendix 3: Summary table example: Data reliability

    Appendix 4: Presence/absence of 47 species in west coast estuaries

    Appendix 5: Life history tables: Life history characteristics of 47 west coast species
      Table 5Ak Biogeography
      Table 5B. Habitat Associations
      Table 50. Biological Attributes and Economic Value
      Table 5D. Reproduction

    Appendix 6: Definitions of terms used in life history tables

                                              287

                                                       West Coast Estuaries

                           Puget     Hood       Skagit     Grays   Willapa  Columbia Nehalem  Tillamook
                           Sound     Canal       Bay       Harbor      Bay       River      Bay        Bay
     Species/Life Stage    T  M   S T   M   S T   M   S T  M   S T   M   S T  M   S T   M   S T   M   S
     Blue mussel        A    0    a               00  00  0
      Mytilis           S  �           a     00    0 0                                      0
      edulis           J   O *   O                       00    00                      0
                       L a* *            i 1S   0 0   00                              0    1i
                       E  I             �    tWJ   0 0   00                           0    ( O
     Pacific oyster     A     0        0                      a         a  S
      Crassostrea      S
      gigas            J      *          U    0 �             *
                       L
                       E
     Horseneck gaper   A      0  1       1  1      0             0          0                           0  0
      Tresus           S         0          0          0         0                                      0
      capax            J   O0            �         0 C           0          0                           0
                       L  00   �1  00                           0    0                                 0�
                       E    0                          0       0          0                           0
     Pacific gaper      A     O  �       0         0  0
      Tresus           S         �    0
      nuttallii        J      O� O                 0 0
                       L  �                       0I 00
                       E   ��   01                    0
     California jackknife A
     clam               S
      Tagelus          J
      californianus     L
                       E
     Pacific littleneck  A   le -                  a    00    00                        a                0o
     clam               S �   ��  �  0 0  00    a                                                        00
      Protothaca �            �          �� �a                00   00                             �   0
      staminea          L a                                      0    00 a                               C
                       L I* -              I I -                  O0  O0                  �             OO
                       E  U    U   a   00  00                                           a               0C
                          T M ST M ST M ST M ST M S T M S T M ST M S

                           Puget     Hood       Skagit     Grays    Willapa  Columbia Nehalem  Tillamook
                           Sound     Canal       Bay       Harbor      Bay       River      Bay        Bay

                                                       West Coast Estuaries

    Relative Abundance                                 Salinity Zone                        Life Stage/Activity

    *       Highly Abundant                            T - Tidal Fresh                      A - Adults
    i       Abundant                                   M - Mixing                           S - Spawning
    O       Common                                     S - Seawater                         J -Juveniles
    Blank   Not Present, Rare, or                                                            L - Larvae
           No Data Available                                                               E - Eggs

                                                     288

                                                  West Coast Estuaries
                                Puget Sound          Hood Canal            Skagit Bay
     Month                   JFMAMJJASOND JFMAMJJASOND JFMAMJJASOND
     Species/Life Stage
     Blue mussel         A                         l
      Mytilis
      edulis             J   ............ .'........................ji.. ..............

                         E                             i i

     Pacific oyster      A
      Crassostrea        S
      gigas              J
                         L
                         E
     Horseneck gaper    A            .iii iiiiiiiii ....ii ....... ......"j.f ......
      Tresus             S                        : I :I.
           ~capax            I             I        i:                    I       I
                         E
     Pacific   gaper     A   !!ii iii i ii iiiii ii  ii i.iiii iiiiiiii !!!!!!!iiiiiii iiii ii
      Tresus             S                            "
      nuttallii          J  . iiiiiiiiiiiiiiiiiiiiiiiii iiiiiiii . .iii .iiiiiiiiiiiiiiiiiii.  I
                         L         1  5_iiiiiiiiiii!i:i iiiii!iiiiml  liiiiiiiiiiiiiii iiii i  I  I

     California jackknife   A
     clam
      Tagelus            J
      californianus      L
                         E
     Pacific littleneck  A
     clam                 S
      Protothaca         J
      staminea           L
                         E
                            JFMAMJJASOND JFMAMJJASOND JFMAMJJASOND
                                  Puget Sound          Hood Canal            Skagit Bay
                                                   West Coast Estuaries

    Relative Abundance                             Life Stage/Activity
             Highly Abundant                      A - Adults
             ~iiiiiii~l   Abundant                S - Spawning
                                                 J - Juveniles
    [I |   Common                                  L - Larvae
                                                  E - Eggs
    Blank    Not present, Rare, or
             No Data Available

                                                289

           h..                *     *         *   0      3*T

                                                                  West Coast Estuaries

                                Puget      Hood      Skagit      Grays    Willapa  Columbia  Nehalem  Tillamook
                               Sound      Canal       Bay       Harbor      Bay       River       Bay        Bay
    Species/Life Stage
     Blue mussel         A     0                                                     NEl
       MONilS            S    *IliU                                                   El                   171E
       edulis            i      U1 n            o               UU
                           L     U          j         li                                l          lE

     Pacific oyster      A     0         0          0          U          U          U
       Crassostrea       S     0
       gigas            'J      U
                           L  U            U          U          U          U          0
                           E  0            N          0
     Horseneck gaper   A       0          U         9          U          N          0
       Tresus            S                                                 E
       capax             i'     UUUU
                           L                                     r-                               1-U1lI]UElE
                           E
     Pacific gaper       A     *                                                                UU
       Tresus            S                l1E                   UUU
       nuttallii        'J      U
                           L     El        13         El      a             U          U

     California jackknife A    a          0         U          U
     clam                S *                                   U                     UUU
       Tagelus           J      UUUU                                                  UUU
       californianus     L      UUU                                        UUUU

      Pacific littleneck    A   *UUE
     clam                S     I!    F I!           IN]        13         13         U          0          0
       Prolothaca        J      U      a             U          U          El         U          U          U
       staminea          L      El        13         9~         17-        El         U          13         U
                           E     El        El   nol              E          El      a             El         U

                                Puget      Hood      Skagit      Grays    Willapa  Columbia  Nehalem  Tillamook
                                Sound      Canal       Bay       Harbor      Bay        River      Bay        Bay

                                                                  West Coast Estuaries

     Reliability                                           Life Stage/Activity

    N     Highly Certain                                  A - Adults
                   FRI Moderately Certain ~S - Spawning
                   iii   Moderately Certain                j~~~ - Juveniles
    El    Reasonable Inference                            L - Larvae
                                                               E - Eggs

                                                               290

    Note: Due to post-publication revisions of the presence/absence information in VolumeI
    (Table 5, pp. 185-197), data in this appendix has been updated and supersedes that presented in volume I

    Index to Appendix 4: Page location of presencelabsence table for each species and estuary

                                                                                                       Estuary

    Common and Scientific Name                                                                                              .9
     Blue mussel (Mytilu eduli)
     Pacific oyster (Crassosfrea gigas)
    Horseneack gaper (Tresus capax)
     Pacific gaper (Tresus nutralliO
     California jackknife clam (Tagalus californianus)
     Pacific littleneck clam (Protothaca starminea)
     Manila clam (Venerupisjaponica)
     Softshell (AMya aranaria)
     Geoduck (Panopea abrupra)
     Bay shrimp (Crangon franis corum)                        292                    293                       294                      295
     Dungeness crab (Cancer magjste,)
     Leopard shark (Thakis semifasciata)
    Green sturgeon (Acipenser madirosrins)
    White sturgeon (Acipenser transmonf anus)
    AmerIcan shad (Alasa sapidissima)
     Pacific herring (Clupea pallast)
    Deepbody anchovy (Anchaa compressa)
    Slough anchovy (Anchoa delicatissima)
     Northern anchovy (Engraufis mordax)
    Cunthroat trout (Oncortryncthus clarki)
     Pink salmon (Oncorhynchus gorbuscha)
    Chum salmon (Oncorhynchus kaea)
    Coho salmon (Oncorhynchus kisutch)
    Steelhead (Oncorhynchus mykiss)
    Sockeye salmon (Qncortrynchus nerka)                     296                    297                       298                      299
    Chinook salmon (Oncortrynchus Ishawytscha)
    Surf smell (l-lypomesus prefiosus)
    Longtin smelt (Spirinchus thaleichthys)
     Eulachon (Thaleichthys pacificus)
    Pacific tomrcod (Microgadus proximus)
    Topsmelt (Athhannops at finis)
    Jacksmell (Atherinopsis califomniensis)
    Threespine stickleback (Gasterosteus aculeaf us)
    Striped bass (Momone saxatlfis)
     Kelp bass (Paralabrax clathratus)
     Barred sand bass (Paralabrax nebuliler)
    White seabass (At tact oscion nobffis)
    White croaker (Genyonemus fineatus)
    Shiner parch (Cyrmalogastrer ggregala)                     (303233
    Pacific sand lance (Amnmodytes haxaplerus)30313233
    Arrow goby (Clevelandia ios)
    LUngcod (Ophiodon elongatus)
    Pacific staghorn sculpin (Leptocorus arrnatus)
    California halibut (Paralichthys califomricus)
    Diamond turbot (Hypsopserta gutfulata)
    English sole (Pleuronectes vefulus)
    Starry flounder (Platidrihys stellatus)

                                                                              291

    Appendix 4 continued

                                      Puget      Hood        Skagit      Grays       Willapa    Columbia   Nehalem   Tillamook
                                      Sound      Canal       Bay         Harbor      Bay         River       Bay        Bay
         Species                       T  M  S   T  M  S   T  M  S   T  M  S   T  M  S   T  M  S   T  M  S   T  M  S
          Blue mussel               A       44       : 1   V 4                                             / i         I 4         I
            Mytilus                 J       '14          I         '14         44           1              ' 1        4       3 4
            edulis                  L       1          4'              4         4 74      4               4          '4 3    4
         Pacific oyster            A       4   4       4   4           4      4   /       4  4                                 /
            Crassostrea             J       4  '        1  4           4       '4  '       ' I
            gigas                   L
         Horseneck gaper           A       4 4             J       J '4  4 4J     /1           1                                  V
            Tresus                  J    4 4 :   44                4               4 ']                               '4    4 4 J
            ca:ax                   LJ            J 4
         Pacific gaper             A    /      4       '4  4      '/  4           '4          '4
            Tresus                 J       '4  4       3   4           3                       '
            nuttallii               L       4   4 3        4       4  4
         California jackknife clam   A
            Tagelus                J
            californianus           L
          Pacific littleneck clam   A       44          4          '4          '4           44                         4       4 4/
            Protothaca              J       ' 4 41 3J 3                        ' 4         J  3                       3            I
            staminea                L         4 3      3  3        3  J 3                                              4      4 4
          Manila clam               A       44          1 /  /  I 4   44          4         44J                                4
            Venerupis              J       4J   3      34 3        i   J '     44 3  l
            iaponica                L    44            44           J          V 44J    4                                     4 3
         Softshell                 A    '4             44          4   '4  V                          4/    '                 4 4
            Mya                     J       '       3              4     3                 '  3
            arenaria                L       4j         4   '3 4 4                I           V 4       4           '4              J
         Geoduck                   A       4                           /                                          :
            Panopea                 J        4J '      4 4             4
            abrupta                 L       4/  '4     4   4
          Bay shrimp                A    '4    4                    4    V 4                                           V 4     44
            Crangon                 J  *   V 4                        4          V          4   4 4 4 '  4 4 4   4
            franciscorum            L    ' /              J 44         I  J 4  4  4    44                4           4        4 4
          Dungeness crab            A    4   V                     44                                                          -4 -,    V  J 4  4J   44  '4  4 4
            Cancer                  J       '1         44          ' 44        4 I J       4          4 4          4     i   J
            magister                L       '  4  J 3 J '                      4                       4 '             V          4
          Leopard shark             A
            Triakis                 J
            semifasciata            P
         Green sturgeon            A           4                              4  4        4   4/          4J
            Acipenser               J                                                                                          J  3
            medirostris             L
         White sturgeon            A          4            4               4                                                     4 x/ J  . . t/  q   '  �J   /
            Acipenser               J                                                                               4
            transmontanus           L
          American shad             A           4           V4  - 'J        /  V  V  ", " ,    V 44  '4 44  4  4  44
            Alosa                   J                          4J   4   J -
            sapidissima             L
          Pacific herring           A                               V 44  44                   4        44         4
            Clupea                  J    4 4             V  V                  '4 V                    4 4    V               4
            pallasi                 L    44 3                      44    4                 4           '4          '
          Deepbody anchovy          A
            Anchoa                  J
            compressa               L
         Slough anchovy            A
            Anchoa                  J
            delicatissima           L
         Northern anchovy          A "    "J 44            4          4    '4 4J    '                 44          4              4
            Engraulis               J       3       I              ' I  3      3                                   4            4 3
            mordax                  L         4  4    44                                      1            4    44             4
                                       T MS T MS T MS TM ST MS TM ST MS T MS
                                      Puget      Hood        Skagit      Grays       Willapa     Columbia   Nehalem   Tillamook
                                      Sound       Canal      Bay         Harbor      Bay         River       Bay         Bay
         Legend:
         T = Tidal fresh zone                      A = Adults                         3 = Species / lifestage is present
         M = Mixing zone                          J = Juveniles                       Blank = Species / lifestage is not present
         S = Seawater zone                         L = Larvae
                                                  P = Parturition

                                                                 292

                                                                                                  Appendix 4 continued

                               Netarts    Siletz     Yaquina    Alsea       Siuslaw    Umpqua    Coos         Rogue
                               Bay        River      Bay        River       River      River      Bay         River
    Species                      T  M  S  T  M  S  T  M  S  T  M  S   T  M  S   T  M  S  T  M  S   T  M  S
    Blue mussel              A                      -4 '4     '4          '4         '4 -'                  '
      Mytilus                J          '4         '4         '4          '          '4         '4         '4          '
      edulis                 L                     4
    Pacific oyster           A          '4                 '4                                           '4
      Crassostrea            J          '4                 '                                    '4      '4  '4
      .0igas                 L
    Horseneck gaper          A          '                  '  '4          '                      '       4
      Tresus                 J                             '4             '          '           '      '4'4
      capax                  L          '                  '              '          '           '4     '4'4
    Pacific gaper            A          '
      Tresus                 J          '
      nuttalliN              L          '
    California jackknife clam   A
      Tagelus                J
      californianus          L
    Pacific littleneckclam   A          '4                 '  '           '4         '                  ''
      Protothaca             J          '4                 '  '4          '          '
      staminea               L
    Manila clam              A          '                      '                                            '
      Venerupis             J           '                     '                                             '4
      japonica               L          '                                                                   '4
    Softshell                A       '4         '4         '           '4         '          '           '4'4
      Mya                    J      ''          4''                    44        '           4''
      arenaria               L      '4          '4         '4         '4         '           '          '4'4
    Geoduck                  A          '
      Panopea                J
      abrupta                L          '4
    Bayshrimp                A       '4         '4 -   4              ' 4'        '4         '4'4       ' 4'
      Crangon                J   '4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4
      franciscorum           L      '4'4        '4'4        4'        '4          '          '          '4'4
    Dungeness crab           A       '4         '          '          '           '          '4         '           '4'4
      Cancer                 j      '4         '             '        '4         '4          '4         '4         '4'4
      maoister               L          '4         '           '4         '          '
    Leopard shark            A
      Ttiakis                J
      semifasciata           P
    Green sturgeon           A                             '              '4         '4      '4          '4'4'4'4'4
      Acipenser              J                             '              '4         '4      '4         '4'4'4'4'4
      medirostris            L
    White sturgeon           A                          '      4          '           '                  '4'4'4'4'4
      Acipenser              J                         '                  '          '       4          '4'4'4'4'4
      transmontanus          L
    American shad            A                          '4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4
      Alosa                  J                         '4'4'4'4'4'4'4'4'4'4'4'J'4'4'4'4'4'4
      sapidissima            L                                                                       '
    Pacific herring          A       '4                    '4          '          '4         '4          '4             '4
      Clupea                 J      ''4         ''         '44        ''            '4       ''         '44          44
      pall1asi               L      ''4''                             '44         '''4                    44
    Deepbody anchovy         A
      Anchoa                 i
      compressa              L
    Slough anchovy           A
      Anchoa                 J
      delicatissima          L
    Northern anchovy         A          '       '          '           '          '4         '4          '4         '4,4
      Engraulis              J          '                                 4         '4       '44        '4          '4'4
      mordax                 L                             '4                             4              q          4  4
                                T MS TM ST MS T MS T MS T MS T MS TM S
                               Netarts    Siletz     Yaquina    Alsea       Siuslaw    Umpqua    Coos         Rogue
                               Bay        River      Bay        River       River      River       Bay        River
    Legend:                                 A   Adults
    T = Tidal fresh zone                    J  Juveniles                      '4= Species I lifestage is present
    M = Mixing zone                         L  Larvae                         Blank = Species / lifestage is not present
    S = Seawater zone
                                          P =2Parturition

                                                            293

    Appendix 4 continued

                                            Klamath    Humboldt  Eel          Tamales   CentralSan  South San    Elkhorn    Morro
                                            River      Bay        River       Bay        Francisou Bayt Francisco Bay Slough  Bay
           Species                      TM ST M STMST M ST M S*M S                                                  S    S*
           Blue mussel              A'1.                               4'                 4.         .44.44
             Mytilus                J                  4                                    4        4
              edulis                 L444'                                                44444
           Pacific oyster           A                  .   4.                                 4'                     4.
              Crassostrea            J4444444

           Horseneck gaper          A'44
              Tresus                 i
              capoax                 L44
           Pacific gaper            A4   4..44'44
              Tresus                 j                    444
             nuttalIl               L                   4
           California jackknife clam   A'1..44
              Tagelus                                                             4444
              californianus          L4444
           Pacific littleneck clam  A                  .4  '4                    -4          .4          .           4
              Protothaca             J                  4
              staminea               L                  4
           Manila clam              A.4..4                                                '   4       4.             4
              Venerypis              J444                                                 44          444
              ia,~onica              L444                                                 44          44
           Softshell                A                   4.         4.                     4.         .44.44
              Mya                    J                  4                                    4        4
              arenaria               L                  4                                    4        4
           Geoduck                  A                      .
              Panopea                J44
              abrupta                L44
           Bay shrimp               A                      -4      .   4.                 4.          4
              Crangon                J44 44 44                                                          4
              franciscorum           L44                           444                    44          44
           Dungeness crab           A       44.4                   44                                    4.                     4
              Cancer                 j   4                4        4                         4        4
              mapister               L444                                                 44          444
           Leopard shark            A                  4.                          4      .           4.             4.
              Triakis                j                    44                              44          4
              semnifasciata          P                      444                                       4444
           Green sturgeon           A...44                         44444                             44
              Acipenser              i4 4                4 4 444                                      4
              medirostris            L4
           White sturgeon           A.44.44                                                          '44
              Acipenser              J44444444                                                        44
              transmontanus          L4
           American shad            A  .   4.                   4.   4.   /.
              Alosa                   j4444444                                                        4
              sapidissima            L
           Pacific herring          A.4                ..4         .   44.                                4.44
              Clupea                 J         4        4             44                     4        4
              vallasi                L44                            44                    44          44
           Deepbody anchovy         A
              Anchoa                 J
              compressa              L
           Slough anchovy           A
              Anchoa                 J
              delicatissima          L
           Northern anchtovy        A       4             4        4               4      4             44.
              Engraulis              J         4        4             44                     4        4
              mordlax                L           4'                                          4        4
                                             T MST M ST MS T MS T MS   M MS                                              S *5**
                                             Klamath    Humboldt  Eel          Tomales  Coentral San   South San    Elkhorn    Morro
                                             River      By         River       Bay        Francisco Bayl Franciso Bay Slown   a

          Legend:                       1Includes San Pablo and Suisun Says.
          T =Tidal fresh zone                       A = Adults                       4=Species I lifestage is present
          M =Mixing zone                            J = Juveniles                    Blank = Species I lifestage is not present
          S =Seawater zone                          L = Larvae
             =Salinity zone is not present           P = Parturition

                                                                           294

                                                                                                  Appendix 4 continued

                               Santa      San Pedro Alamitos   Anaheim   Newport   Mission    San Diego Tijuana
                               Monica Bay Bay         Bay        Bay        Bay         Bay        Bay        Estuary
    Species                      *  *  S   *                          *              S   *  * S      * *  S     *  * S *  *  S   *  *  S   *  *  S
    Blue mussel              A    :                 : /1 J                           '4          ' :                    4
      Mytilus                J  /                          :         :: :4  4        4                      '
       edulis                 L          4          4           4J         4          4          4                      '
    Pacific oyster           A
       Crassostrea            J
      gigas                  L
    Horseneck gaper          A
       Tresus                 J
      capax                  L
    Pacific gaper            A          '4          V          '          '4         '4          i          '4         '/
       Tresus                J           4          4          4                      'J 4  4    4                       J
      nuttailli              L          4           J         41          4          4
    California jackknife clam   A       V    :        :: .                           '           V          '4         '/
       Tagelus               J           4          4     :                           4          ::          4 :
       californianus          L          4     J               4                                 :l  4  lJ lJ
    Pacific littleneck clam  A           4 /        V          ',         '4         '           q          '           V
       Protothaca             J          4          4              1 4                           44
       staminea               L          4                      4          J 4'                   4          4          4
    Manila clam              A
       Venerupis             J
      japonica               L
    Softshell                A
      Mya                    J
       arenaria               L
    Geoduck                  A                      '4'
       Panopea                J: :
      abrupta                L
    Bay shrimp               A
       Crangon                J
       franciscorum           L
    Dungeness crab           A
       Cancer                 J
       magister               L
    Leopard shark            A          '           '  V '4                                      V 'J
       Triakis                J          4          4          4                                             '
      semifasciata           P          4          l
    Green sturgeon           A
      Acipenser              J
      medirostris            L
    White sturgeon           A
      Acipenser              J
       transmontanus          L
    American shad            A
      Alosa                  J
      sapidissima            L
    Pacific herring          A           4/
       Clupea                 J          4
      pallasi                L
    Deepbody anchovy         A          '4          '          '4         '           '4        '4 ',
      Anchoa                 J          '
      compressa              L                                 4
    Slough anchovy           A                      '4        '4          '           '4         '4         '
      Anchoa                 J                     4           4                                 J 4  4
      delicatissima          L                                 l                     4
    Northern anchovy         A          '           ',         4          '          '                      '
      Engraulis              J          4          '4          4          4          4                      'J
      mordax                 L           1                     J 4  4     4          4                      44
                                  * S                                        *  * S *  *  S *  *           S *  *     S *  *  S   *  *  S
                               Santa      San Pedro Alamitos   Anaheim   Newport   Mission    San Diego Tijuana
                               Monica Bay Bay         Bay        Bay        Bay         Bay        Bay         Estuary
    Legend:
    T = Tidal fresh zone                   A = Adults                         4 = Species / lifestage is present
    M = Mixing zone                        J = Juveniles                      Blank = Species / lifestage is not present
    S = Seawater zone                      L = Larvae
     = Salinity zone is not present        P = Parturition

                                                            295

    Appendix 4 continued

                                           Puget      Hood        Skagit      Grays       Willapa     Columbia   Nehalem   Tillamook
                                           Sound      Canal       Bay         Harbor      Bay         River      Bay         Bay
          Species                       T MST M ST M STM ST M STM ST M STM S
          Cutthroat trout           A 44             4       44                  4       4       4       4       4       4       4
             Oncorhynchus            i -444 4 4 4 4 44 4 4 4
             clarki                  L
         Pink salmon               A  4 4 4 4 4 4                                                     4
             Oncorhynchus            J44444 4444
             gorbuscha               L4
         Chum salmon               A   4  -4  4  -4  4  -4  4  4 V 4 4  -4  4i 4  -4  4  4  4   4  4  -4  4   q  -4  -4
             Oncorhynchus,           J       4 4 4 4 4 44 4 4 4 4 4
             keta                    L
          Coho salmon               A 4444444444444 444444444 44
             Oncorhynchus            J i
             kisutch                 L
          Steelhead - fall          A
             Oncorhynchus            J
             mykiss (F)              L
          Steelhead - half pounder   A
             Oncorhynchus            j
             mykiss (H)              L
          Steelhead -summer    A44 4444                                                                  4 4              -4 4
             Oncorhynchus            J4 4 4 4 4                                                        4                       4
             mykiss (B)              L
          Steelhead -winter         A 444 4444 44444 44 444 444 4 444
             Oncorhynchus            J i
             mykiss (W)              L
          Sockeye salmon            A 4 444 4 44 44
             Oncorhynchus            J i
             nerka                   L
          Chinook salmon -fall       A 44 444444 4444 444 44 444 444 4
             OncorhynchusJ44 44                             4       4 44            4   '           44                     ''
             tshawytscha (F)         L
          Chinook salmon - late fall   A
             Oncorhynchus            J
             tshawytscha (LF)        L
          Chinook salmon - winter    A
             Oncorhynchus            J
             tshawvtscha (W)         L
          Chinook salmon -spring    A 4 4 4444 444444444444
             Oncorhynchus,   j4 4 4 4                                      4 444                                               4
             Ishawyfscha (Sp)        L
          Chinook salmon - summer  A44
             Oncorhynchus            i                                                                 4
             Ishawytscha (Su)        L
          Surf smelt                 A 44                4          44          4           44            4         44            4
             Hypomesus               i    4             4              4        4             4         4             4        4
             pretiosus ,    L 44                        4           4           44             4        44          44
           Longfin smelt             A       4                          4        4             4         4
             Spirinchus      ,       i
             thaleichthys            L       4                           4      4      4       4      4
           Eulachon                  A           44                              4      4       4      4
             Thaleichthys            J
             pacificus               L4444 4444 4
           Pacific tomood            A       44          4444                                444                     444
             Microgadus              J         4         4             4        4           4             4         4            '
             proximus                L 44 4                         44 4                    44            4 44                    4
          Topsmelt                  A                                               4444                                        4
             Atherinops              J                                                                      44                  4
             affinis                 L
                                            T M ST MS TMS T MS T MS T MS T MS TM S
                                            Puget      Hood        Skagit      Grays       Willapa     Columbia   Nehalem   Tillamook
                                           Sound      Canal       Bay         Harbor      Bay         River       Bay         Bay
          Legend:
          T =Tidal fresh zone                       A =Adults                          4=Species / lifestage is present
          M   Mixing zone                           J =Juveniles                       Blank = Species / lifestage is not present
          S   Seawater zone                         L =Larvae

                                                                          296

                                                                                                                 Appendix 4 continued

                                    Netarts    Siletz     Yaquina.-  Alsea      Siuslaw    Umpqua    coos        Rogue
                                    Say        River   Say           River      River      River      Bay        River
     Species                     T M STM ST M STM ST MS TM ST MS T MS
     Cutthroat trout         A  '4  '4~ '4  '4  '4  '4  '4  '4  '4  '4  '4 '-4  '4  '~ '4  '~ '4  '4  '4  '4  '4  '4  '4  '4
       Oncorhynchus          J '4             4 '4444444444444  '4  44 '4444
       clarki                L
    Pink salmon              A4'4''4''44'44
       Oncorhynchus          J
       gorbuscha              L
    Chum salmon              A'4'44''444'''                                                       4444'
       Oncorhynchus          J''i              4444 -  -   j ' 4'4  -4'q4' 4'       4                  44
       keta                  L
     Coho salmon             A  '4  '4'4' 4    '4  '4  '4 '4  '4  '4  4 4  '4  '4  '4  '4  '4  '4  4  '4  '4  '4'
       Oncorhynchus          J   4'          4'   4'   4'   4'   4'                  4'   4'   44'                 4'
       kisutch               L
    Steelhead - fall         A
       Oncorhyrnchus         J
       mykisqs (F)            L
    Steelhead - half pounder   A
       Oncorhynchus          J                                                                                  4''
       mykiss (H)            L
    Steelhead -summer       A              '''                     44                    444'''
       Oncorhynchus          i                44'4                                        444'4
       mykiss (8)            L
    Steelhead -winter        A'4''4''4''44''4''4'4'4''4''44'44'44
       Oncorhynchus          J   4'   44'                      44'   4'4'   4'   4'   4'   4'   4'
       mykiss (14/           L
    Sockeye salmon          A
       Oncorhynchus          J
       nerka                  L
    Chinook salmon -fall     A  '   4'   4'             4'   4'   44'                4'   4'   4'   4'   4'
       Oncorhynchus          J   4'   4'   4'   4'   4'   4'4'   4'   4'   4'   4'                                 4'
       tshawytscha (F)       L
    Chinook salmon - late fall  A
       Oncorhynchus          J
       tshawytscha (LF)      L
    Chinook salmon - winter    A
       Oncorhynchus          J
       tshawytscha (14'      L
    Chinook salmon -spring    A             q  V       -V -4 -q  q    4                  44' 4' '4 4'
       Oncorhynchus          i                  4 4 4 4 4'''''''''
       tshawytscha (SP)      L
    Chinook salmon - summer  A
       Oncorhynchus          J
       tshanwvtscha (Su)     L
    Surf smelt               A       '44        '44        '4         '4           4          4'         44'
       Hyporresus            J       '4'4       '4         '4 4       '4           44         44        ''4         44
       pretiosus              L          4          4'                     4'4'                             4'
     Longfin smelt           A                          '   444'                                 4'   4'
       Spirinchus            i                              44'                       4'                4'
       thaleichthys           L                        '                                                4
    Eulachon                A                                                       '''4''''
       Thaleichthys          J
       pacilicus              L4                                                               4
    Pacific tomood          A                  '4         '4          '4         4            4        44         4
       Microgadus            J        4           4        44         '4          ''          44        ''           4
       proximus               L'4''4                                  ''4         '4         '44
    Topsmelt                 A      ''                      4          44        '4'                     4
       Atherinops             J        4                     4          44        '4'                     4
       affinis                L      '4'4                             444'4
                                     T M STM ST MS TM ST M STM ST M STM S
                                    Netarts    Siletz     Yaquina    Alsea      Siuslaw    Umpqua    Coos         Rogue
                                    Bay        River      Bay        River      River      River      Bay         River
    Legend:
    T =Tidal fresh zone                    A = Adults                        '=Species / lifestage is present
    M =Mixing zone                         J = Juveniles                     Blank = Species / lifestage is not present
    S  Seawater zone                       L = Larvae

                                                                     297

    Appendix 4 continued

                                         Klamath    Humboldt  Eel         TaMaleS  Central San  South San    Elkhomn    Morro
                                         River   s ay          River'     Bay       Francisou Say' Francisco Bay Slough  Bay
          Species                     T MST M ST M S T M ST M S  M S*S   *5
          Cutthroat trout         A  4                    4q 4  4  4
            Oncorhynchus          j  44 4
            clarki                L
         Pink salmon              A444
            Oncorhynchus          J
            gorbuscha             L
         Churnsalmon              A  44-                  1
            Oncorhyrnchus             444
            keta                  L
          Coho salmon             A444444444444
             Oncorhynchus          J i
            kisutch               L
          Steeihead -fall         A444
             Oncorhynchus          J       4.
            rnvkiss (F)           L
          Steelhead - half pounder   A
             Oricorhynchus         J  4        44
            mykiss (H)            L
          Steelhead -summer       A  4  4  4
             Oncorhyrichus         J i
            mykiss (S)            L
          Steellhead -winter      A  4         i                      4144444
             Oncorhynchus          J i         ~
             mykiss 1W)            L
          Sockeye salmon          A
             OncorhynchusJ
             nerka                 L
          Chinook salmon -fall     A4444444444444
             Oncorhynchus           j      4      4      4     4                       4
             tshawytscha (F)
          Chinook salmon - late fall  A1                                              44
             Oncorhynchuvs                                                             44
             tshawytscha (LF)
          Chinook salmon - winter   A                                                  44
             OncorhynchuVS         J4                                                     44
             tshawytscha (W)      1L
          Chinook salmon -spring    A  4  -444444
             Oncorhynehus          J  4  4    V
             tshawiflscha fSp)     L
          Chinook salmon - summer  A
             Onco~rhynchus         J
             Ishawytscha (Su)      L
          Surf smelt               A          4       44         44
             Hypomeasus            J         4        4            44                  4
             pretiosus             L                               44444
          Longfin smelt           A           4      4      4     44                     4        4
             Spininchus            J     i
             thaleichthys          L4                    4      44444
          Eulachon                Al
             Thaleichthys          J
             Pacificurs            L444
          Pacific tomcod          A444                                                44         44
             Microgadus     ,      J                     444                              4        4
             proximus              L
          Topsmelt                A             14                                                 1

             Atherinops            j                               4                       4       4
                                          T M TMS  T MS  T S  T M1S    M MS    *                                               S
                                         Klamath    HmbolIdt  Eel         Tmls CnrlSn SuhSn                Eko         or
                                         River      Bay        River   S ay          Frlanciac Bay1YFrancisco Bay Slough  Say

          Legend:                     1Includes San Pablo and Suisun Bays.
          T  Tidal fresh zone                    A =Adults                        4 = Species / lifestage is present
          M =Mixing zone                         J =Juveniles                     Blank = Species I lifestage is not present
          S =Seawater zone                       L =Larvae
            =Salinity zone is not present

                                                                     298

                                                                                                      Appendix 4 continued

                                Santa      San Pedro Alamitos   Anaheim   Newport   Mission    San Diego Tijuana
                                Monica Bay Bay        Bay        Bay        Bay        Bay    S ay           Estuary
    Species                            S 5   *               S          S          5      *  5           S           S
    Cutthroat trout         A
       Oncorhynchus          J
       clark.i               L
    Pink salmon             A
       Oncorhynchus          J
      gorbuscha             L
    chum salmon              A
       Oncorhynchus          J
       kera                  L
    Coho salmon             A
       Oncorhynchus          i
       kisutch               L
    Steelhead - fall        A
       Oncorhynchus          J
       mvkiss (F)            L
    Steelhead - half pounder   A
       Oncorhynchus          J
       mykiss (H)            L
    Steelhead - summer      A
       Oncorhynchus          J
       mykiss (S)            L
    Steelhead - winter      A
       Oncorhynchus          J
       mykiss (W)            L
    Sockeye salmon          A
       Oncorhynchus          J
       nerka                 L
    Chinook salmon - fall    A         '
       Oncorhynchus          J
       tshaw-ytscha (F)      L
    Chinook salmon - late fall  A
       Oncorhynchus          U
       tshawytscha (LF)      L
    Chinook salmon - winter   A
       Oncorhynchus          J
       tshawytscha (Wi       L
    Chinook salmon - spring   A
       Oncorh~ynchus         J
       tshawytscha (SP)      L
    Chinook salmon - summer  A
       Oncorhynchus          J
       tshaw-ytscha (Su)     L
    Surf smelt               A
       Hypomesus             J
       pretiosus             L
    Longfin smelt           A
       Spirinchus            J
       thaleichthys          L
    Eulachon                A
       Thaleichthys          J
      pacificus             L
    Pacific tomcod          A
       Micro gadus           J
      pro ximus             L
    Topsmelt                A'4''4''44'44
      Atherinops            J          '
      affinis               L          '           4444'
                                     S *                      S *                   S 5  *                           S
                                Santa      San Pedro Alamitos   Anaheim   Newport   Mission    San Diego Tijuana
                                Monica Bay Bay        Bay        Bay        Bay        Bay        Bay        Estuary
    Legend:
    T =Tidal fresh zone                    A =Adults                        4=Species / lifestage is present
    M =Mixing zone                         J  Juveniles                     Blank = Species / lifestage is not present
    S =Seawater zone                       L =Larvae
      =Salinity zone is not present

                                                               299

    Appendix 4 continued

                                         Puget      Hood       Skagit     Grays      Willapa    Columbia   Nehalem   Tillarnook
                                         Sound      Canal      Bay        Harbor     Bay        River      Bay        Bay
         Species                     T MST M S TM STM ST M ST M STM STM S
         Jacksmelt               A
           Atherinopsis          J
           calitomiensis         L
         Threespine stickleback    A  '   4'   4'   4'   4'   4'   4'   4'   4'                          4'   4'   4'
           Gasterosteus          J'   4'   4'   4'   4'   4'   4'   4'   4'                             4'   4'   4'
           aculeatus             L'44           ''          44        ''          44         ''          44        ''
         Striped bass            A          '    .'
           Morone
           saxati/is             L
         Kelp bass               A
           Paralabrax            i
           clathratus            L
         Barred sand bass        A
           ParalabraxJ
           nebuliler             L
         White seabass           A                                 '
           Atracloscion          J
           nobilis               L
         White croaker           A
            Genyonemus            J
           lineatus              L
         Shiner perch            A       '44        ''         '44        ''         '44        ',          '4          44
            Cymatogaster          J   4'   4'   4'   4'   4'   4'   4'   4'   4'   4'   4'   4'
           aggregate             p       '4''4''4'''4                                                                   44
         Pacific sand lance      A       '44        '.4        ',             4'                '4 .       ',           44
           Ammodytes             j        4          44        ''           4W       'W         'W         ''           4
            hexapterus            L       'W         'W         ''          44        ''           4             4'
         Arrow goby              A       '44        ''         '44        '4         ''44
            Clevelandia           j        44        ''          4'         4          44'
            ios                   L       ''          44        ''          44        ''
         Ungcod                  A           '4        '4         '4
            Ophiodlon             J        44        ''          4'         4           '          4            '4'
            eloaqatus             L       '4'        '4'.       ''4        4'4        4 4
         Pacific staghorn sculpin   A    '44        ''          44        'W         '''''                    4,4      ',
            Leptocottus                4'   4'   4'   4'   4'4'   4'   4'   4'   4'  '4'44'44
            armalus               L       '44        ''          44'                      4      ''              4         '
         California halibut      A
            Paralichthys          J
            califomicus           L
         Diamond turbot          A
            Hypsopsefla           J
           _quttulata            L
         English sole            A           '          4'
            Pleuronectes          J       '4         '4         '4         W'         '4         ''4          44        'W
            vetulus               L       'W         ''          4           4         44        '4'                     4
         Starry flounder         A       ',          44        ''           44       'W          ',          44        ''
            Platichthys           J    4  '   4  '   4   '   4  '   4  '   4   '   4  '   4   '   4  '   4   '   4  '   4  '
            stellatus             L       '4         '44        ''         W'         '44        '''4                    44
                                          T MS8 T M S T M S T M S T M S T M S T M S T M S
                                         Puget      Hood       Skagit     Grays      Willapa    Columbia   Nehalern Tillamook
                                         Sound      Canal      Bay        Harbor     Bay        River      Bay         Bay

        Legend:
        T =Tidal fresh zone                    A =Adults                         '=Species / lifestage is present
        M =Mixing zone                         J =Juveniles                      Blank = Species/I lifestage is not present
        S =Seawater zone                       L =Larvae
                                                      P =Parturition

                                                                         300

                                                                                                                       Appendix 4 continue,

                                      Netarts     Siletz      Yaquina    Alsea        Siuslaw    Umpqua    Coos           Rogue
                                      Bay         River    S ay           River       River       River       Bay         River
     Species                        T M STM ST M STM STM ST M ST M STM S
     Jacksmelt                  A                                    4'4
       Atherinopsis            J                                                                                V
        californiensis          L
     Threespine stickleback     A       '                                4V VV '   4'   4V V '
       Gasterosteus            i
        aculealus               L    V V                       V         V             V         V              V        V
     Striped bass               A                                   V'4' '4 4'   4
       MoronejV                                                                                         V      V
       saxatilis                LVVV
     Kelp bass                 A
       Paralabrax              J
       clathratus              L
     Barred sand bass          A
       Paralabrax              J
       nebulifer               L
     White seabass              AVV
       Atractoscion            J
       nobilis                  L
     White croaker              A
       Genyonemus              J
       fineatus,                L
     Shiner perch               A          V        '44         V4          V4V                      VVV                     V4
        cymatogaster            i          V         V V V V                               V         V             V         V
       aggregate               P VV                 V           VV          V           VV             V VV                    V
     Pacific sand lance        AVVV'44VV
       Ammodytes               i
       hexapterus              LVVVV                                                                    VV
     Arrow goby                 AVV'4V4V                                                                         V V
       Clevelandia             J i
       ios                      L                               VV
     Lingcod                   AV
       Ophiodon                i                                                                                  f
       eloniqatus              LV
     Pacific staghorn sculpin    A      V            44         V              V         V             V        'V           V
       Leptocottus             JV V V V VV V VV V V V V V V V V V V V V                                                     V V
       armatus                  LVVVVVVVV
     California halibut         A
       Parafichthys            J
       cafifornicus            L
     Diamond turbot             A
       Hypsopsetta             J
       guttulata                L
     English sole              A
       Pleuronectes            i   VV    VV    VV    VV    VV                                       V           V
       vetulus                 LVVVVV
    Starry flounder            A                   -4VV                     VVV                     VVV
       Platichthys             i        V  V
       stellatus                L       VV          V             VV
                                      T M S T M S T M S T M S T M S T M S T M S T M S
                                      Netarts    Siletz       Yaquina    Alsea        Siuslaw     Umpqua    Coos          Rogue
                                      Bay         River       Bay         River       River       R iver      Bay         River

    Legend:                                 A =Adults
    T   Tidal fresh zone                    J =Juveniles                           V=Species / lifestage is present
    M =Mixing zone                          L =Larvae                              Blank = Species I lifestage is not present
    S =Seawater zone                        P =Parturition

                                                                         301

    Appendix 4 continued

                                            Klamath    Humboldt   Eel         Tomrales   Contra] San  South San    Elkhorn    Morro
                                            River   s ay          River       Bay        Francisco Bay' Francisco Say Slough  Bay
           Species                      T M ST MST M STM ST M S*M S*S   *
           Jacksmelt                A444                                                  44         4444
             Atherinopsis           i
             californiensis         L444                                                 44          4444
          Threespine stickleback   A 44           4       4      4                   4      4        4
             Gasterosteus           J44 4 4 4                                         4 4            4
             aculeatus              L44            44          444444
           Stnped bass              A4
             Morone                 J
             saxatilis              L                                                 4
           Kelp bass                A
             Paralabrax             J
             clathralus             L
           Barred sand bass         A
             Paralabrax
             nebulifer              L
           White seabass            A44                                                      44
             Atra ctos cion         J i
             nobilis                L
           White croaker            A444                                                              44
             Genyonemus             i
             lineatus               L4                                                    4444
           Shiner perch             A        44         44         -4  44                  44         444
              Cymatogaster           i    4 4    44q    444                                4   44    4
             a!aqreqata             p       4             4        4                        4        4
           Pacific sand lance       A444444
             Ammodytes              J i
             hexapterus             L44
           Arrow goby               A444                                                  44          4444
              Clevelandia            j444                                                 44          4444
             iOs                    L                  4                                    4        4
           Lingood                  A                      -4
              Ophiodon               J444                                                  44444
              elongatus              L444
           Pacific staghorn sculpin    A    4             4        4                         4        4
              Leptocottus            J         4        4             444 4                           4
             armatus                L4444                                                 44         444
           California halibut       A44444
              Paralichthys           i
              calitornicus           L44444
           Diamond turbot           A                                             44                  4
              Hypsopsetta            i                                                        4        4
             piuttulata             L                                             44
           English sole             A
              Pleuroftectes          J          4        4 4        44             4          4          4
              vetulus                L4444                                                   4                       44
           Starry flounder          A4444                                                 44          444
              Platichthys            J       4      4      4      4                    4     4        4
             stellatus              L           4       4                                    4        4
                                             T MS   TM   T MS T Mr' .Tn.MS *  S  *  5* *
                                            Klamath    Humboldt  Eel          Toae        etrlSn Suh Sa.Ekor                 or
                                            River      Bay         River      Bay        Francisco Bay1 Francisco Bay Slough  Bay
                                            1Includes San Pablo and Suisun Bays.

          Legend:                                 A = Adults
          T =Tidal fresh zone                     J = Juveniles                      4=Species / lifestage is present
          M =Mixing zone                          L = Larvae                         Blank = Species / lifestage is not present
          S =Seawater zone                        P = Parturition
             =Salinity zone not present

                                                                          302

                                                                                                       Appendix 4 continued

                                Santa      San Pedro  Alamitos   Anaheim   Newport    Mission    San Diego Tijuana
                                Monica Bay Bay         Bay        Bay         Bay         Bay        Bay         Estuary
    Species                      *  *  S   *  *  S          * *   S   *    S   *       S   *  *  S   *  *  S   *  *  S
    Jacksmelt                A                       H'                                ' :                    4
      Atherinopsis           J           41  ::                                       . :
      califomiensis          L   : '4                              :
    Threespine stickleback    A
      Gasterosteus           J
      aculeatus              L
    Striped bass             A           '4            ::                                          '4
      Morone                 J
      saxatilis              L
    Kelp bass                A           '           4          4          '4          '           /          '4          '
      Paralabrax             J           '          '           4          '4         '4          '           '
      clathratus             L           '4
    Barred sand bass         A           '4    :                '           / il                              ' i
      Paralabrax             J                         : '4     'J '4 .                           l           l          '4
      nebulifer              L           '          'J                                                        4
    White seabass            A           '          '4                                                        I
      Atractoscion           J           '
      nobilis                L
    White croaker            A                      V           V '4       '           '           '          ''
      Genyonemus             J                      ' l         l                      '4         4           'j        '4
      lineatus               L                      '4 l                   4           4          4           4          '
    Shiner perch             A           '4                                            '4 I'                  '4         '4
      Cymatogaster           J          '4          '                      '4         '4          '4         'j
      aq.qreqata             P           4          '                       4          4          '
    Pacific sand lance       A
      Ammodytes              J
      hexapterus             L
    Arrow goby               A           '4                                 '4        '4           '          '4          '
      Clevelandia            J          '4          '                                  '4         '           '          '4
      ios                    L                      '           l          l           4           4          '          '
    Lingcod                  A        :     ::
      Ophiodon               J           '          '4
      elongatus              L
    Pacific staghorn sculpin    A        '4         V           4          '4                      4          '4          '
      Leptocottus            J           '          '           '          '           l          '4         '4          V
      armatus                L           l                                                        '4         '4          'I
    California halibut       A           4V          4                      '          4           '4J        '
      Paralichthys           J           4           4          '4         '           '4          J          J 'j
      califomicus            L           '           4
    Diamond turbot           A           '           '4         '4         '4          '           '4         '           '
      Hypsopsetta            J           '4J '                                                    '4         '4          '
      Quttulata              L           'J '                              '                       4          '4         l
    English sole             A           '4
      Pleuronectes           J           4          l
       vetulus                L           '4         'J
    Starry flounder          A
      Platichthys            J
      stellatus              L
                                  *  *  S   *  *  S   *  *  S   *  * S   *  *  S   *  *                 S *  *    S *  *  S
                                 Santa      San Pedro Alamitos   Anaheim   Newport    Mission    San Diego Tijuana
                                Monica Bay Bay         Bay         Bay        Bay         Bay        Bay         Estuary

    Legend:                                         A = Adults
    T = Tidal fresh zone                             J = Juveniles             ' = Species / lifestage is present
    M = Mixing zone                                  L = Larvae                Blank = Species / lifestage is not present
    S = Seawater zone
     = Salinity zone is not present

                                                                303

    304

    Index to Appendix tables 5A-5D: Page location of Biogeography, Habitat Associations, Biological Attributes
    and Economic Value, and Reproduction tables for each species.

    Common and Scientific Name                                                      Cmmon and Scientific Name
    Blue mussel (My/i/us edufi6)                                                    Blue mussel (M'y/i/us edulis)
    Pacific oyster (Crassos/rea gigas)                                              Pacific oyster (Crassostrea gigas)
    Horseneck gaper (Tresus capax)                                                  Horseneck gaper (Tresus capax)
    Pacific gaper (Tresus nuttaffl~                  306        1       1           Pacific gaper (Tresus nflu/a//i
    California jackknife clam (Tagelus cafifornianus)                               California jackknife clam (Tagelus ca/iforniarnus)
    Pacific littleneck clanm (Proto/haca stamninea)                                 Pacific littleneck clam (Protiofhaca s/aminea)
    Manila clam (Venerupis japonica)                                                Manila clam (Venerupisjaponica)
    Softahell (Mya arenania)                                                        Softshell (Mya arenaria)
    Geoduck (Panaopea abru p/a)                                                     Geoduck (Panopea abrupla)
    Bay shrimp (Crangon franciscorum)                                               Bay shrimp (Crangon franciscorum)
    Dungeness crab (Cancer magistei)                                                Dungeness crab (Cancer nragistei)
    Leopard shark (Triakis semdiasciata)             307    313    319              Leopard shark (Triakis semifasciata)
    Green sturgeon (Acipenser medirostris)                                          Green sturgeon (Acipenser medirostris)            324
    White sturgeon (Acipenser transmontanus)                                        White sturgeon (Acipenser /ransmontanus)
    American shad (Alasa sapidissima)                                               American shad (Alosa sapidissirfa)
    Pacific herring (C/upea pa//asi)                                                Pacific herring (C/upea pal/asi)
    Deepbody anchovy (Anchoa compressa)                                             Deepbody anchovy (Anchoa compressa)
    Slough anchovy (Anchoa deficalissimna)                                          Slough anchovy (Anchoa de/icatssima)
    Northern anchovy (Engraulis mnorday)                                            Northern anchovy (Engrau//s mwrdax)
    Cutthroat trout (Oncorhynchus c/ark,)            308    314    320              Cutthroat trout (Oncorhynchus clarki)
    Pink salmon (Oncorfhynchus gorbuscha)                                           Pink salmon (Oncorhynchus gorbuscha)
    Chum salmon (Oncorhynchus keta)                                                 Chum salmon (Oncorhynchus keta)
    Coho salmon (Oncorhynchus kis utch)                                             Coho salmon (Oncorhynchus k/s utch)
    Steelhead (Oncorhynchus mykiss)                                                 Steelhead (Oncorhynchus mykiss)
    Sockeye salmon (Oncorhynchus, nerka)                                            Sockeye salmon (Oncorhynchus nerka)
    Chinook salmon (Oncorhynchus tshawy/scha)                                       Chinook salmon (Oncorhynchus tshawy/scha)
    Surf smelt (Hypamesaus pretiosus)                                               Surf smelt (Hyporresus pretiosus)
    Longfin smelt (Spirinchus tha/eichthys)            0        1       2           Longfin smelt (Spirinchus fthaeichthys)
    Eulachon (Tha/eicifhys pacificus)                       391521Eulachon (Tha/eichthys pacificus)
    Pacific tomcod (Microgadus proximus)                                            Pacific tomrcod (Micro gadus proximus)
    Topsmelt (Athernnops affinis)                                                   Topamnelt (Athermnops affinis)
    Jacksmelt (A/herinoposis ca/ifomniensis)                                        Jackamelt (Afherinopsis ca/ifomniensis)
    Threespine atickleback (Gas/erG steus aculeatus)                                Threespine stickleback (Gasterosteus acu/eatua)
    Striped bass (Mfororne saxa/ilis)                                               Striped bass (Morone sairafi/is)
    Kelp bass (Para/abrax c/athratus)                                               Kelp bass (Paralabrax c/athra/us)
    Barred sand bass (Par/alabras nebu/ifer)         30         16       22         Barred sand bass (Para/abrax nebu/ife,)32
    White seabass (Atractoscion nobi/is)                      31 1632White seabass (Atractoscion nobi/is)32
    White croaker (Gern'onemus lineatus)                                            White croaker (Genyonemus lines/us)
    Shiner perch (Cyrnatogaster aggregata)                                          Shiner perch (Cymnatogaster aggregata)
    Pacific sand lance (Ammodytes hexapterus)                                       Pacific sand lance (Ammodytes hexapterus)
    Arrow goby (C/eve/and/a ios)                                                    Arrow goby (C/eve/andia ibs)
    Lingcod (Ophiodon e/onga/us)                                                    Lingcod (Ophiodon elongatus)
    Pacific staghom sculpin (Leptocottus arma/us)                                   Pacific staghorn sculpin (Leptoco//us armatus)
    California halibut (Para/ich/hys ca/ifornicus)   311    317    323              California halibut (Para/ich/hys ca/ifom/icus)
    Diamond turbot (Hypsopset/a gu/tu/ata)                                          Diamond turbot (Hypsopsetta gut/u/a/a)
    English sole (Pleuronectes ve/u/us)                                             English sole (P/euronectes ye/u/us)
    Starry flounder (P/a tich/hys s/el/a/us)                                        Starry flounder (P/a tichthys ste//a/us)

                                                                             305

     L**.  *   **. I         *-.

                                                           BIOGEOGRAPHY
                                 IMarine                       Estuarine                 IRiverine
    Li     f sIg/ctv                               Salinity Range   EsuryTp Stratifi-
    A - Adults                              ISAB   I Venice System caio
    S -Spawning adults                       -       ~      -'~

    L -Larvae                           ~4
    E -Eggs                                               0

                       J      q0  0                  0000   0                 0           0
                           LC. 0         0  0       0 .* Q.    0  0     0~- .0    . 0L

     Pacific oyster   A                   55          000   SSSS@S0S@                          A
      Crassostrea     S                  55           SS       S     SS                        S
      gigas           J                   0*           *.      S0S                             J
                      L                                                          0S* S @ O @   L
                      E                  550SS S O                                            E
     Horseneck gaper  A  55               55              5.      5 5       5                  A
      Tresus capax    55                 5  **S*                        ***                    S
                      JO                 55              ** .S SS                             J
                      LO                 So              *S e e e                             L
                      ES                 55 0OSSO                                             E
     Pacific gaper    A     S             55              *S       S    S     S                A
      Tresus nuttalIN    55              SS              SS       S     S     S                S
                      J 55               55              So . . .
                      LS                 OS              SO S S S                             L
                      ES                 55              SS S S S                             E
     California jackknife A              SSSSSSSSSA
     clam             5                     5OSS*                                              S
      Tagelus         J                     SSS                  S55J
      californianus   L                   55 0S                S               55 00           L
                      E                  55 0SO                                55 00          E
     Pacific littleneck   AS              55           *SSS          SSS                       A
     clam              55                 55*                *     *     *     *               S
      Protothaca      J  55               55@               5     5     5     5                J
      staminea        LO                  55*                S     S    S     S                L
                      ES                 55           *S S SS                                 E
     Manila clam      A                   55                       0 SSSSSSSS                  A
      Venerupis       5                     5*sS                        S     *                S
      japonica        J                   55*                *iS               S
                      L                  5               O     S     S        SSL
                      E                  O                  SS       S        eeE
     Softshell        A                   5               O     SSO                            A
      Mya arenaria    S                   5               *     SS            *                S
                      J                  50           SSSSS4S00  00  00                       J
                      L                  50           SSSSSSSSSS0  0                          L
                      E                    5SO O S S SE

                                                306

                                                                 Appendix 5A continued

                                               BIOGEOGRAPHY
          A - Adults      I ~~~~Marine IEstuarine                        IRiverine
    S - Spawning adults            I     aiiyRne       Estuary Type I tatioi-
    M - Mating                     ISAB  IVenice System IIcto
    J -Juveniles                        ' e0'
    L -Larvae
    E -Eggs          C
    P -Parturition                                                   Q-

     Geoduck      AS  S00                    0 0    00 *      OS          A
     Panopea abrupta S 00        0          0@0    0 0    *000            S
                     i 0 0     0O           0 0    050   0 00            J
                  LOS           0 0         0 0    @ 0    0 00           L
                  ES 0          00          00    00        000          E
     Bay shrimp   A   0            0     0 0 @ 0 0 0                      A
      Crangon     S                         0 00S00 e o
      franciscorum   J   0    0 00    *@O@@@Se@@ee                        j 0
                  L   0         0 0       *0  0 0   0 0 00               L
                  E   S          0    0     0 0 0*00 0000                E
     Dungeness crab  ASS         0 0      0 0    0 0 0 0                  A
      Cancer magister M   0        0           0       0 0M

                  L   0         000 0 0 0 0L
                  E O              0                  0                  E
     Leopard shark   AS     0                00 0        0    0           A
      Triakis     M   500 0                              0      0M
      semifasciata    JO    0                00 0        0    0
                  P   0                        0 0      0    0           P
     Green sturgeon   AS 0 0SO O OS O O                                   A
      Acipenser   S                                                  *   S
      medirostris  J 0 0       @0 0 000                    05 50          J
                  L           0                                    0     L
                  E                                                  S E
     White sturgeon   A 0O  00 0 0 0                      O O O OA
     Acipenser   5                                                    005
      transmontanus  J 0 0    0 00 00 00000              0 0    0 00 0J
                  L           0                      0      0      OS L
                  E                                                    0 E
     American shad   A   0 0 000 000000000             0 0SOSOOS *A
     Alosa       S            0      0                    0        0 005
      sapidissima    J   0     0 0 00*0 0000    0 OSOSOOSJ  *
                  L                             0   0       0      OS L
                  E                             0   0       0      OS E
     Pacific herring   A 0        0 0      0 0  0 00 00 000               A
      C/upea         0 0         0 0         0050 0 00 00 0S
     pallasi      JOS00           S       O      @0 0   0 0   0           J
                   0 *0         0 00   SO S @S O                         L
                  E 0           0 0    050&060 000 05                     E

                                         307

    Appendix 5A continued

        Life -stage/activity     IBIOGEOGRAPHY
       A - Adults               IMarine  IEstuarine                                IRiverine
        S- Spawning adults                I     Salinity Range  'Estuary Type  Stratifi-
       J - Juveniles                      ISAS     Venice System    c     aion

        E -Eggs

         Deepbody       A    0             0             0    0             0          A
         anchovy        S                                      0             0         S
          Anchoa        J    0             0             0    0             0          J
          compressa     L                  0                   0            0          L
                       E                  0                                0          E
         Slough anchovy   A    0           0             S     S0                      A
          Anchoa        S                  S                                0          5
          delicatissimna    J    0         0             5     00J
                       L                  00                 0L
                       E                  05                 00                       E
         Northern       A    @ 0          05        *SSOSSOSOO                         A
         anchovy        S    0             0             00      0500                  S
          Engraulis     J    OS          @5 0       0 0S@SS@O                          J
          mordax        L    0           @    0*S0    0  0    0                         L
                       E    0             0@                    @5 0E
         Cutthroat trout    A    0    *@OS@OSO@OS0                     @0 0 00         A
          Oncorhynchus   S            S                                        0 00    S
          clarki        J   0            0  0 0 0 0 00 0               050 00   J
                       L            0                                         0       L
                       E            0                                           0    E
         Pink salmon    A    0        @00 *0 0  *0 0   050   0   S SOO A
          Oncorhynchus   S            0       0             0                  0  0SSS
          gorbuscha    J 0 0  00 0 0 0 0 00    S   *0 0 00 *0
                       L0                                     0        0 S      S L
                       E                                  0S           0 S      S E
         Chum salmon   A  S    0 0 0 0 0 60 0 0    00 S   OOSA
          Oncorhynchus   5            0       0             0    S
          keta          J  *0  0 0 0 00 0 0  0                         *6 0 00 0 J
                       L0                    0                       0   0   OS L
                       E0                                      0       0 S      O E
         Coho salmon   A  S                   *OS00 0 0 050 0 0 0A
          Oncorhynchus   S            0       0              0000                0 005
          kisutchJ**                *    *    *     *    *    *    *       *     **
                       L                                    0 000               S L
                       E                                    0 5 I5 5*O E
         Steelhead      A    0                    0**0000S SO SS 0S 0  0   0  0 0A
          Oncorhynchus   S            0       I   I         0 00*0             0 005
          mykiss        J                  *                     **    *    *D '* *********
                       L                                                        0  00   *@ L
                       E            5                           5             *O E

                                             308

                                                                             Appendix 5A continued

                 Life stagelacti~~ity                    BIOGEOGRAPHY
    Lif- sAdulatsvt               Marine                     Estuarine                  Riverine
    S - Spawning adults                  I      Salinity Range  E     s    ur Typ Stratifi-
    J - Juveniles                           SAB  I Venice System  EstaryTp     tion
    L - Larvae                ~       ~     ~
    E -Eggs                         '

           Oncor y n ch u S                  0                                              0
                      L                     6                                  0 00
                     E-0 0C.                   0                               * 0 0  *.    E
          ChinookC. sa0nA0          0 0       0   .0 0. 0        00  0    0 0  00.  A -
      Oncorhynchus   5                              *                           *  00   0
      nerkysca       J     **       *    **       *    * 0             *S@SSSJ   000        000
                     L                  0       0                      S000              L
                     E                                         0                  *0@E
     Chinoosmelto  A 0 0            *S00  0 0  0 0 0  0 0 0A
      Oncorhyncus    5               0    0                @000    0 00s

                     L            0                        000                  *      L
                     E  0                                     00                *0     E
     Lourfinsmelt    AO   0            @0          **S@@00 0 *  0      SOS               A
      Hypomescus     SO0  SO                          @0         0    *000  0S
      prthliosuhs    J    0            00       0       0    0         @ 0              J
                     L                00 00                                00           L
                     EO                                         0   0 00                E
     ELaongfnml   A   0            * 00 00 00 00 00 00 0 00    A
      Spirinchuhs    S              0                      0                                 0  S
      thaceichths    J     0           00       *@iOO                    @
                     L              *O@SS@@@O@OO@@@O0 0     0                           L
                     E                         0 0           0           0      0       E
     Euacihconco     A     SO       *@       OO      SO             0    0 SO            A
      ThMichthyds    5         0                                  S * **
      pacximius      J     S                         0                                   J
                     L              ***** 000                                           L
                     E             0          0                           0    0        E
     Tpacmfltomo     A     *                          000                  05            A
      Micognadus     5                              0 0
      proxinus       J   0             00          00      0     00                      J
                     L    0000                                  0        00             L
                     E    0                                                             E
     Jacksmelt       AO                @ 0         50      50               0            A
      Atherinopsi    5                 0 0            * S00                 0            5
      calffminis     J   00            OS0              0    0       0    0
                     L                 @50              0    0      0    0              L
                     E                 500                0            0    0           E

                                                309

    Appendix 5A continued

                                          I                  ~~~~BIOGEOGRAPHY
              Life stage/activety    Marine  IEstuarine                                  IRiverine
       A - Adults                           I      Salinity Range   I             Staii
       S - Spawning adults                     SBIVncSytmIEstuary Type  cattion
       M -Mating                               SBIVnc  ytmIcto
       J -Juveniles                            -            '~
       L -Larvae       ~       ~             o~
       E -Eggs
       P -Parturition               z     �

              Threspie,           0 00   0                   0        0        0 00    A~

         stickleback     S              55       55 e 0         0          SSOS*             S
          Gasterosteus    J  550        0  *SS      S     *    *     S     S    S            J
          aculeatus      L              550    05 0                SSSSSSS0 00               L
                         E              55 *       5  0           S0    0     0 S 00         E
         Striped bass    A    S          SO      S    S     O 05                    S                      A
          Morone         S              0               0               0 0O        S        S
          saxails        J              0O                                          S        J
                          L              55      5     5    5              @ ~ SL
                         E              55       S S5S S                            SE
         Kelp bass       ASS                                                                 A
          Paralabrax     S     S                             S                               S
          clathratus        55                S 0                                            i
                          L    055                                 55L
                         E    0                                                              E
         Barred sand bass  A S 0                                                             A
          Paralabrax     5    0                                                              S
          nebulifer         55 0 0            0              0                   5           J
                          L    S                                                             L
                         E    0                                                              E
         White seabass    A    S                0                                            A
          Atractoscion   S    6                                                              S
          nobilis           55 0                             SO 0                            J
                          LO 0  0                                                            L
                         E    0                                                              E
         White croaker   AS                55          SO      S 0             55  000       A
          Genyonemus    5             0                      0 55       0                    S
          lineatus        JO               55          55      5         5       5
                          LO                05          05     5         S                    L
                         E    S                             5                                E
         Shiner perch    AS                55           50      5    55                      A
          Cymatogaster   M                 5*                SS         SS      SM
          aggregata      J  SO 0S                S     S     S    S     S     S              J
                         P       D         S           *S      S     SS      S               P      00 0I
         Pacific sand lance AS             *S          * 00   0SOSSSSS0 00                   A
          Ammodytes       5550                5           55          @0 0 0                 5
          hexapterus     J  55             55           SS      *    S        *              J
                          L    S              S         SS      0     0S      S               L
                         ES                  5           55000                               E

                                                    310

                                                                           Appendix 5A continued

                                                      BIOGEOGRAPHY
    A - Adults               I Marine                   Estuarine               Riverine
    S- Spawning adults                      Salinity Range  E           ca
    J - Juveniles                               Venice System   stuaryTypI Stiti
    L-Larvae                                �
    E-Eggs

                         a                                                 P

                        a ~     $ 0`~u0                         C)   b     i~

     Arrow goby     A    0            S      *O@@SSOOOSO                          A
      Clevelandia   S               0          O @ 0 s   0 0 0OSSO a              S
      los           J    0          0        OSSOOSSOOSO                          J
                    L    S          OS         SOOSSOOSSO                         L
                    E               O          OSOOSSOOSS                         E
     Lingcod        A    5          55            00000           005             A
      Ophiodon      S    0             0                                          s 05    0    5  5
      elongatus     J    S          SO         OSOSOSSOOS                         J
                    L    0            0              05005500                     L
                    E    0            00                  0            0          E
     Pacific staghorn   A    0      0          *SOOOOOSOO                         A
     sculpin        5                             0       005050 0                S
      Leptocottus   J    S       *o 0          o    .ooo.....                     J
      armatus       L    0          00            @55000505                       L
                    E                  S            SOSOOSOS                      E
     California halibut  A    0        5            5 0 0    0    0 5             A
      Paralichthys    S    S                                                      S
      californicus     00i           05           0000         0    5             J
                    LOS             OS              *oo  S OS                     L
                    E    *                          S                             E
     Diamond turbot   A    0        0          0               5    *             A
     Hypsopsetta    S    0            0            0                  0          S
     gutfulata     J    0          00         00000           0    0             J
                    L    0            0             0    0    0    00             L
                    E    0             0            0                0            E
    English sole   A    0             0            0       0    0                A
     Pleuronectes    S    0                                                      s
     vetulus          00i          0     0    * 0    0 0 00                      J
                    LOS             OS            0505000 0                       L
                    E    0             5            0       0    000              E
    Starry flounder   A    0       5                5050 0050                    A
     Platichthys   S    0                                  0     0               s
     stellatus     J    0        *SOOOSOOOO    ees    o                          J
                    L    5          00         OOSOOOOO0o                         L
                    E    0                          0       5                     E

                                               311

    IAff      0      .-o:lm�

                                         I                     ~~~~~~~HABITAT ASSOCIATIONS
                                        Habitats          Substrate preference                Domain
                                                                                I  Benthic      I elagic I EstuarineI

                       Life staoe/a~~~~~~~~~~~~~~~~~~jy&  ~~~~Littorall Sublittoral I13athyall

      S -Spawning adults
      J -Juveniles
      L -Larvaea
      E- -Eggs                                                                     aa

     Blue mussel       A        0                      C  00000
      Mytikis edulis   S        *SO 0                                                              COOC  0*S
                      J        CC@ 0. 0                                   ..0
                       L        OS C                                                                       * C C L
                      E        C@ C                                                        C      C      CE
     Pacific oyster    A    a *C00   :0 * C A
      Crassostra       S                                                   0000S
      gigas            J        0                                          0C0
                      L       0                                                                  COOC0L
                      E       * 0                                   S                      S     C    S     E
     Horseneck gaper    A       *@ 0          9  CC0                                                       0O S S A
      Tresus capax     S        SC 0                                                               *55*500
                      J        Ce      S     @5                        5**0
                      L        *                                    0               0                  0  * S L
                           OC  S                                                                       0 *  S C E
     Pacific gaper     A        00    0       *S                          0C *                          C A
      Tresus nuttafli  S       @0      S                                                                     S5 0
                      J       0 0    0       OS 00                                                  0      46J
                      L        @ 00                                  *                                 0 0 O S L
                      E        @0                                                                   0 0 0 * S E
     California jackknife   A   * *            0@                        0@*C                                 A
    clam              S        SC 9                                       000    S*                         S
      Tagelus          J        CS            Ce                        OS 000
      califomianus     L        0 0                                                                     0 *      L
                      E        ge                                                          *      S O       E
     Pacific littleneck  A      *g0 *          * *                        0* *                          S A
    clam              S        em 0                                  see                          *e0  60S
      Protothaca       J        0 0     S      *@*  00  CO                                         C *       J
     staminea         L        *@0     0                                                          0 C  C  CO L
                      E        *@0    q                                                                0 *  * @ E
    Manila clam       A        0 @0                                   *5 CC                             O A
      Venerupis        S        0 00                                                                    *0 0
     japonica         J        0 CC0                                    0                               00
                      L       0 0                                                                      0 0 S OL
                      E       0                                                            C           0 CCE
    Softshell         A        0 CC             0                       CC                        *    C A
      Mya arenania    S         Ce 0                                       000   O C                         S
                                         J ~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 0C0  @0                .      .J
                                          L  @6~~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~ 0 * C                        L
                      E        OS0                   t              C                      C      * C       E

                                                       312

                                                                                           Appendix 5B continued

                                                              HABITAT ASSOCIATIONS
                                      Habitats          Substrate preference   IDomain
                                                                              I  Benthic      I1Pelagic I Estuarine
                      Life stacielactivety ~ ~ ~ ~ ~     ~     ~     ~    ~ ~~~~~~~~~~~~ Littoral ISublittoral lBathyall
    A -Adults
    S -Spawning adults
    M -Mating
    J -Juveniles
    L -Larvae
     - Eggs
    P -Parturition

    Geoduck           A        0 SO          60                         50      000 0S A
     Panopea abrupla   S       0 @    0   S                          @0        0                  0 0o  0s

                     L        @0 0         0    5                                                * O       L

    Say shrimp        A        06          00                                                     9 e        A
     Crangon          S        @6          000               a                                     000       S
     franciscorum     J        @ 0         @0 00                       0 0  060
                     L        @   0                       0         O                     0 0S  0 * 0 L
                     E        @ 0        0                          @00                         0 0         E
    Dungeness crab    A        @ 0        0 00   000500                                           @550       A
     Cancer magister   M             0    0                             0066                                 M

                     L       0 00   060                                     0        000L
                     E        @ 0        0                                                                  E

    Leopard shark    A         0 05       0 0 0   0                         0@0000                    0 0 A
     Triakis          M        ** 0                                                                          M
    semifasciata     J        @ 0      0    0 0   0           0560 0555                          000
                     P                                                                                      P
    Green sturgeon    A    0 @               0000 0@0 .   0 0        05 00500 *0 0A
    Acipenser        S    0                          0                                                      S
    medimostris      J   00  0   0   00 00 0 00   0 0 0S*0
                     L    0 0                        * oL

    White sturgeon    A        @                                 0 0 60 6   @00                   *      0  A
    Acipenser        S    0                                                                                 S0
     transmontanus   J         @ 0        0 00 0 0 00   0  0 0 0                                  0 0 0 0
                     L     @5@                    000    0                                                  L
                     E    0@02
    American shad     A     S      0              0                                            0000  0    A
    Alosa            S     55 0                                                                  @0         S
    sapidissirna     J  0 00           00                                                           0 S0 0 0
                     L     @5 0                                                                 0           L
                     E     06               00                                                   @5         E
    Pacific herrnng   A        @ 0        0                         6                       00      00       A
     Clupea           S        06                              60       00
    pallasi          J        @0          5       0               0                            0 0 0 0
                     L        @60                                     0              0            0 * S    L
                                         2   00              55      @0~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ *E

                                                          313

    Appendix 58 continued

                                                                    HABITAT ASSOCIATIONS
                                           Habitats           Substrate preference    I             Domain
                                                                                  I       Benthic      I Pelagic  Estuarine
                                                                                  [LItlorall Sublittoral I1athyall
      Life stage/agjjy&t
      A- Adults
      S - Spawning adults
      J-Juveniles                            -_X
      L -Larvae
      2- Eggs                                                                                              aa

     Deepbody           A           *     0                             6                       0       0 0 0  A
     anchovy            S           0                                   0                             0            S
       Anchoa            J           0     0                             0                       0      *00  J
       compressa         L           0                                   S                             S  000   L
                        2           0                                   0                              @0         E
     Slough anchovy     A           0     0                             0                        0      0@  0      A
       Anchoa            S           0                                                  0                           S
       delicatissima     J           0                                                                              i 0  S  @   U
                        L      a                                        0                              000         L
                        E           00                                                                            2
      Northern           A        00       00                                                                 0 @0000  A
      anchovy            S        0        @                             0                        00                S
       Engraulis         J        00       00                            0                        @0000             J
       mordax            L        *0                                                      0             00000       L
                        E        0        00                            0                        @0                E
      Cutthroat trout    A    0 0          0         00                                          0                  A
       Oncorhynchus      5     0                    00
       clarki f                            0        000                                          0      0000J
                        L    0                     00                                                              L
                        E                                                                                         E -(    ::::
      Pink salmon        A     0  0        0        0                                            *      000         A
       Oncorhynchus      S     0  000                                                                               S
      gorbuscha         J         0       00                                              0 aOOOOOJ
                         L     @0                   *0                                                              L
                        E   00                                                                                     E
      Chum salmon        A      0                    00                                                 0 00 00 A
       Oncorhynchus      S     00                   00                                                              S
       keta              J    0    00                                    0                       00 0000J
                        L     @0                   @0                                                              L
                        E  @0                      @                                                               2
     Coho salmon        A     00          0         0                                            0                A
       Oncorhynchus      5     0                    00                                                              S
       kisutch           J     00          0 0      0                                             000000 J
                        L    0                     00                                                              L
                        E     0                    00                                                              E
     Steelhead          A     @0 0                          0                                    0 0  0  0  0    A
       Oncorhynchus      S     0                    0                                                               S
       mykiss            i     00          00         0 0.                                        006600   J
                        L     0                    00                                                              L
                        E     0                    00                                                              E

                                                            314

                                                                                                   Appendix 5B continued

                                                                    HABITAT ASSOCIATIONS

                                          Habitats           Substrate preference  I                 Domain
                                                                                 I       Benthic       I Pelagic I Estuarine  I
                                                                                 ILittoral I Sublittoral IBathyall

    A -Adults
    S - Spawning adults                                                44
    J -Juveniles                                                                                            e
     L -Larvae
      - Eggs

    Sockeye salmon  A

     Oncorhynchus      S  0  C                                                                                       S
     nerka             JC                 CC 0  0                                                                    iee
                       LC 0 0                                                                                       L
                       2CC                         CC                                                               E
    Chinook salmon     AC   @0         C           C  C  C                                        C      C       CA
     Oncorhynchus      S     C                     C                                                                 S
     tshawytscha       J  0@           a@ e0  0       0  0                                                           i
                       L     C                        C                                                             L
                       2                                                                                            E C
    Surf Smelt         A        0C       O 0                                                       *CC*CC0   0A
     Hypomesus         S        C      CC          C
     pretiosus         J        0  0       a
                       L        C      C    C      CCL
                       2-                                                                                           E 0C
    Longfin smelt      A     0C           CC                             C                        C 4  6            A
     Spirinchus        S     0C                    C                                                    C            S
     thaleichthys     J         C  C     C                              0 C                      C      C  C        J
                       L           0C              C                                             C         *  0     L
                       E                                                                                0C          E
    Eulachon           A      CC6            0                           C                         00 0C             A
     Thaleichthys      S     C                     C
     pacificus         J                     C                           0                                           J
                       L     CC             C                                                        C     @        L
                       2-                                                                                            E
    Pacific tomcod     A         C     400   4C                                                          CCC         A
     Microgadus        SC                                                                                            S
     proximus          J         C     OCC   0C  0C                                                                  J
                       L        C         CCC                                                              C         L
                       -                               IF2
    Topsmelt           A         CC       000  CS                        C                        C      C     C A
     Atherinops        S         C  C     C                        C  C     C                                     C  S
     affinis           J                 C  CC                           C                        C      C       CJ
                       L        CC 0     0   C                          C                               C        CL
                       E        CS       C                           C      C                                     CE
    Jacksmelt          A         C    1  SI   C  C  C                                             C      C       CA
     Atherinopsis      S            000 0C CC                               C                                  C    S
     califomniensis    J        a  CI    CC        C                                              C            0  4CC
                       L    CC0 CSL
                       2          C                                  C                                           C

                                                               315

    Appendix 5B continued

                                                I                        ~~~~~~~HABITAT ASSOCIATIONS
                                                   Habitats      ~~Substrate preference                   Domain
                                                                                          I   Benthic       IPelagic IEstuarine
                            ~~~~~~~~~~~~~~~~~~~~~~~~~~~~ILifesaeatlyLtorall Sublittoral IBathyall
        A -Adults
        S - Spawning adults
        M -Mating
        J -Juveniles
        L -Larvae
          - Eggs
         P -Parturition                                          e            a-o                         u

        Threespine          A  0      0                                                                             0  * 0 @ A
        stickleback         S@       O     0         0                            @0                             OS       S
         Gasterosteus      i    0 0                        0                                                        0 *0@O
         aculeatus          LOO0       e                                      S                                  00        L
                           20000                    00                           @                                 0 r-
        Striped bass        A     *e       * 0       00                                                 *     SOS         A
         Marone            S      OS                                                                          S           S
         saxatifis          J     **0                   0                        0 0                          *S*OJ

                           L   @0                   O@0                                                         0         L

        Kelp bass          A             *0      @0                 0      0      0    00                     SA
         Paralabrax        S                  0                               0        @                                  S
         cla thra tus      J             ***O                0                                05       *      **           J
                           L           0                                              00L

        Barred sand bass    A         O*      O            *0             0       S    *0                     @     0      A
         Paralabrax        S                            0                                                                 S
         nebulifer          J        0  *     0         0               *0         0e                  0       000         J
                            L        Se                                                                *      0    0       L
                           2                                                                          0                   E
        White seabass       A                           0         0          0 *                        0                  A
         Atracloscion       S                                                                                              S
         nobifis            J        00              00              *@           Se 0                        000         J
                           L        0 0          *                                                    *         0         L
                           2n          a                            r-S
        White croaker       A         e@       **       * 0                       00      0                             0 O SA
         Genyonemus        S                            S                         0                                       S
         lineatus           J        SO0       0 0      0                  0                                           0  * S J
                            L        @        0                                  000                          @    0       L
                           2                                                     00
        Shiner perch        A         @0    0   *    0       0                0                                *0 0  A
         Cymatogaster      M         @0      0       0 0                                                            0 0 0  M
         aggregata          J        0  00        0   05                                      0                 0 *@i
                            P        00   0   0 0                                              0              e        eP
        Pacific sand lance    A       @0    0        0                            S4S                                      A
         Ammodtytes         S                  0                  0S05                                  0     0            S
         hexapterus         J          000        0     0                         .000                  40    0            J
                           L        0     0      0     00                                                    0L
                           2        S                                                                                     E S

                                                              316

                                                                                          Appendix 5B continued

                                        I                       ~~~~~~~HABITAT ASSOCIATIONS
                                       Habitats           Substrate preference                  Domnain
                                                                                  Benthic      I Pelagic IEstuarine
                                                                           ILittorall Sublittorall I1athyall

     A -Adults                          ~-
      - Spawning adults
    J -Juveniles                             CD~'
     L -Larvaea                                                                       2 Z-s a
      - Eggs

    Arrow goby         A    I   @0             @0                          @000
     Clevelandia      S         @0             @                             00 @00 *                      e     s
     jos               J         0             @0                         @0                             000
                     L   I 0                     0                                                   0 0    0L
                     E        @0             @0                         @0                          00         2
    Ungcod             A              0060                 @ 0   0   0 00000                                      A
     Ophiodon          S              0     0              00*            0 00                                    S
     elongatus         J        0 0 0 0 4            0     0        0     000                         0           J
                     L              0                                                           0L
                     2              0                      0               0
    Pacific staghorn   A        0 0   0 0 *00 0  0                        0@0 00                      0   40 0 A
    sculpin            S        0 0      0                                   40                       0           S
     Leptocottus       J     000         0         0 00 0 00              0 0                         000
     armatus           L        **4      *                                                      00 00             L
                     E        @0 0        0                                                        0            E
    Califomnia halibut    A           0  @0       S5                               00                 0           A
     Paralichthys      S                 0        0                          0                                    S
     califomnicus      J        SO       0 0040                           0 0   0 0                  0   0   *
                     L        @0       @ 0                           0                       0   0              L
                     2     0                                                                 0 F-
    Diamond turbot     A        @0       S     000                              0                     O       SA
     Hypsopsetta       S                 0                                   0S
     gutut~lata        J        @0       0        0@ 0                                                   O
                     L        00       S00                                                         0            L
                     E                 6                             00
    English sole       A                 0 0@                                         0               0           A
     Pleu~ronectes     S                 0     0                                0  0S
     vetuilus          J        0  .0 000 0                                00             0   0       S*0  0J
                     L        @0       @                                                      @00               L
                     2-                 E000
    Starry flounder    A        @0        @00                              00      00                             A
     Pla ach thys      S                 0        50S
     stellatus         J      000        0     0 0                          00000
                     L        @0       a                             000                                        L
                     E                 0                             0

                                                         317

         Life sage/atiylI                 BIOLOGICAL ATTRIBUTES                  IEconlomic
    A - Adults                jFeeding type    Spatial strategy     Longevity        Value
    S - Spawning adults
    J -Juveniles                              c    ~ bt
    L -Larvae

                   E~~~~~~~~~~~~                                                      - Eg

    Blue mussel       A                                                       0    0           A
     Mytilus edlulis  S                            **S
                    J            @5              5@              0                    J
                    L           0                  50L
                    E                            @0       0E
    Pacific oyster    A            06  0                                  0   0    0   A
     Crassostrea      S                            0                                    S
     gigas            J            0 0             0               0J
                    L           0                    ~L
                    E                           0         0E
    Horseneck gaper   A            0                                                    A
     Tresus capax     5                            0
                    J            0@0
                    L           000L
                    E                           0      0E
    Pacific gaper     A            0                  0          0               0      A
     Tresus nuttailli   S                          0
                    J            S@                 0                                 J
                    L           0               00                                    L
                    E                                                                 E
    California jackknife AS 0                0                         0  0   S    S   A
    clam              5                                                                  S
     Tagelus          J            50 0                            S                    J
     californianus    L            0  *                                                  L
                    E                            S     SE
    Pacific littleneck   A         0  0                                0      0 S       A
    clam              S                             @6
     Protothaca       J            0  0                            0  0                 J
     stamninea        L            5                  0                                  L
                    E.                              0                                 E
    Manila clam       A            SS 0                                S       S    S  A
     Venerupis        5                            0                                    S
    japonica         J            0  0            0                0                    J
                    L           SSSL
                    E                            5     5E
    Softshell         A            5O *                                0            0    A
     Mya arenaria     5                                                                 S
                                  J  05       0               5                    ~~~~~~~~~~~~~~~J
                    L           @500L
                    E                            S     SE

                                       318

                                                                              Appendix 5C continued

                                          BIOLOGICAL ATTRIBUTES               IEconomic
    AL- Adults                  Feeding type  jSpatialstrategy   Longevity       Value
    S - Spawning adults
    M- Mating
    J-Juveniles -                                 '-            0~
    L -Larvae
    E-Eggs
    P- Parturition

    Geoduck          A            @0             00                   0      0 0   A
     Panopea abrupta S                           0  0                               5
                      J           00                           i0                    J
                      L          0               00       0                          L
                     E                           000                                E
    Bay shrimp       A              0             0            0                0   A
     Crangon         S                           0  0                               5
     franciscorurm   J 0            0            0                                  J
                      LO                         0                                   L
                      E                          00                                  E
    Dungeness crab   A 0                         00                0       0    0   A
     Cancer magister M                              0                               M
                      J0                                        o                    J
                      LO         0               00          0                       L
                     E                             0        0                       E
    Leopard shark    A 0                0          0               00           0   A
     Triakis         M                           00                                 M
     semifasciata    J 0               0         0                                  J
                     P                          00                                  P
    Green sturgeon    A                 0     00                      0 0       0   A
     Acipenser       S                        0                                     5
     medirostris     J0                       0          0                      0   J
                      L                       0           0                          L
                     E                       0           0                          E
    White sturgeon    A 0               0     0                       0   0    0   A
     Acipenser       5                        0                                     S
     transmontanus   J 0               0      6  0  00                     0        J
                      L                       0           0                          L
                     E                       0           0                          E
    American shad    A            0     0                          0          0 0   A
     Alosa           S                        0                                     5
     sapidissima     J           0            000               0                   J
                      L                       0              0                       L
                     E                       0                                      E
    Pacific herring  A           0      0  0     0                 0         0 0   A
     Clupea          S                           00                             0   5
     pallasi         J           0      00       00             0          0    0   J
                      L          0               0           0                       L
                     E                          00       0                     0   E

                                               319

    Appendix 5C continued

              Life stage/activity     I            BIOLOGICAL ATTRIBUTES               IEconomic
             A-Adults                   Feeding type  j Spatial strategy  Longevity      Value
              S - Spawning adults
             J-Juveniles
              L- Larvae

                                                         8~ e
              E                                                  - Eggs0

              Deepbody         A           0        0             0                 0        A
              anchovy          S                          0                                  S
               Anchoa          J           0              0           S             0        J
               compressa       L           0              S 0                                L
                               E                          S                                  E
              Slough anchovy   A           0              0              0                   A
               Anchoa          S                                                             S
               delicatissima   J                          0           0             0        J
                               L          0              0            0                      L
                               E                          0        0                         E
              Northern         A                 SS0    0   A
              anchovy          S                          00                                 S
               Engraulis       J           0     5        0       0    0            S 0      J
               mordax          L           S              00                                 L
                               E                          @0       5                         E
              Cutthroat        AO                       @00               00                 A
              trout            5                                                             S
               Oncorhynchus   Ji                 0     0  0*             0  *      0         J
               clarki          L                       5           0                         L
                               E                       S              0                      E
              Pink salmon      A 0                     S  S  S           S               0   A
               Oncorhynchus    S                       S                                     S
              gorbuscha      Ji                  55500                  0                   J
                               L                      S               S                      L
                               E                       0                                     E
              Chum salmon      A 0                      00                5  0           0   A
               Oncorhynchus    S                       0                                     5
               keta            JO                 00055                   00                 J
                               L                      0               5                      L
                               E                       5              0                      E
             Coho salmon      A                        5                 5              0 S   A
               Oncorhynchus    S                       S                                     S
              kisutch         JO                                        0               0   J
                               L                      S                                      L
                               E                       S              S                      E
             Steelhead        A                       0                    0 0          0   A
               Oncorhynchus    S                       0                                     S
               mykiss          JO                    0000                0                   J
                               L                      0            0                         L
                               E                       5              0                      E

                                                   320

                                                                             Appendix 5C continued

    Life stage/activity      I           BIOLOGICAL ATTRIBUTES                IEconomic
    A - Adults                  Feeding type  ISpadal strategy   Longevity       Value
    S - Spawning adults
    J-Juveniles
    L-Larvae
             E        ~ ~ F-  Eggs

    Sockeye salmon   A0                       000               0 0        0    0   A
     Oncorhynchus    S                        0                                     5
     nerka          Ji           0      00000                    @                  J
                     L                       0              0                       L
                     E                       S                                      E
    Chinook salmon   A 0                      0  0   0             0       0    0   A
     Oncorhynchus    S                        0                                     S
     tshawytscha     J                  *                                       * 0  ' 0  0   0  0  .0    0   i
                     L                       S              0                       L
                     E                       0              0                       E
    Surf smelt       A 0         0      0        0              0          0    0   A
     Hypomesus       S                           0  0
     pretiosus       JO          S      0        @0             0                    J
                     L 0        000                                                 L
                     E                          00       S                          E
    Longfin smelt    A 0         0      0     00        0      0                    A
     Spirinchus      S                        0@0                                   S
     thaleichthys    JO          0     0       000              0
                     LO 0              0                    0                       L
                     E                       0              0                       E
    Eulachon         AS          0      0     @00         0           0         0   A
     Thaleichthys    S                       0                                      S
     pacificus       JO          0      0           0           0                    J
                     LO         0            @00            0                       L
                     E                       S            @0                        E
    Pacific tomcod   A 0                                                   0 0  0  5  A
     Microgadus      S                              0                               S
     proximus        J 0                         0 a            0                    J
                     LO0                           0                                L
                     E                             S                                E
    Topsmelt         A        0         0        00              00           0 S   A
     Atherinops      S                           *                                  S
     affinis         J         000         00                                        J
                     L        00                00          0                       L
                     E                           00       00                        E
    Jacksmelt        A        0         0        00              0         0    0   A
     Atherinopsis    5                           0@                                 S
     californiensis  J        0  0      0        0  0           0
                     L        @0                00        00                        L
                     E                          00       0                          E

                                              321

    Appendix 5C continued

              Life stage/activity      j            BIOLOGICAL ATTRIBUTES                 IEconomic
              A- Adults                    Feeding type   Spatial strategy   Longevity       Value
              S - Spawning adults
              M- Mating
              J -Juveniles
              L - Larvae
              E-Eggs
              P- Parturition

              Threespine        A *               0      0  0@ 0                                A
              stickleback       S                                                               S 0
               Gasterosteus     J 0         0           0                                       J
               aculeatus        L 0         0            0           0                          L
                                E                        00   0 0                               E
              Striped bass      A         0           0    0           0                        A
               Morone           S                       0                                       S
               saxatilis        JO                       00                 00                  J
                                LO                       00          0                          L
                                E                                                               E
              Kelp bass         A  S                                       0                0   A
               Paralabrax      S                               0                                S
               clathratus       JO                          00              0                   J
                                LO          0                        0                          L
                                E                              000                              E
              Barred sand bass  A *                         4 1            0                0   A
               Paralabrax       S                              0                                S
               nebulifer        J                           00                                  J
                                L           0                                                   L
                                E                              0     0                          E
              White seabass     A 0                0           0                  0 0       0   A
               Atractoscion     S                              0                                S
               nobills          JO                          00 0                                J
                                L 0         0               0                                   L
                                E                              0                                E
              White croaker     A 0                0           0               0 0          0   A
               Genyonemus      S                               0                                S
               lineatus         J *                           0             0                   J
                                LO          0               00   0                              L
                                E                              0     0                          E
              Shinerperch       A 0                         0  0               0       0        A
               Cymatogaster    M                            0                                   M
               aggregata       Ji 0                         00                                  J
                                P           0               0                                   P
              Pacificsand lance  A 0              0   0                     000            0 0   A
               Ammodytes        S                           0 0                                 s
               hexapterus       J0          0     0         00              0                   J
                                LO          00    0                                             L
                                E                           00    00                            E

                                                      322

                                                                                  Appendix 5C continued

    Life stagie/activitv       I             BIOLOGICAL ATTRIBUTES                  IEconomic
    A - Adults                    Feeding type   Spatial strategy     Longevity         Value
    S - Spawning adults
    J -Juveniles
    L- Larvae                                B
    E-Eggs                                                                               4

    Arrow goby         A  4                          0               0                     A
     Clevelandia      S                             0                                      S
     ios               JO                           0             0                        J
                       L           0                0 0                                    L
                      E                            0         0                            E
    Lingcod            A                                S 0                      S    0   A
     Ophiodon         S                                 S                                  S
     elongatus         JO                           0 0              0                     J
                       LO          0                             0                         L
                       E                                S                                  E
    Pacific staghorn    A                                            00          0         A
    sculpin            S                             0                                     S
     Leptocottus       J                               0             0          0          J
     armatus           L0           0                @0           0                        L
                       E                             00                                    E
    California halibut   A  00    0   A
     Paralichthys      S                                                                   S
     californicus      JS              0                      0      S                     J
                       LO          0                   0         S                         L
                       E                                S                                  E
    Diamond turbot    A  0                           0@                                    A
     Hypsopsetta      S                                 0                                  S
     guttulata        Ji 0                          0                0
                       L 0         0                                                       L
                       E                                0     0                            E
    English sole       A  S                                              0            0   A
     Pleuronectes      5                                0                                  S
     vetulus           JO                           0 0              0                     J
                       LO          0                   0          S                        L
                       E                                0                                  E
    Starry flounder    A  0                0         0  0                0         0 0   A
     Platichthys       S                                0                                  S
     stellatus         J  0                         0                                      J
                       L 0         0                                                       L
                       E                                0     0                            F

                                                 323

             I.,-   *  **~~    I   *.I.  ..0         .

                                                                           REPRODUCTION
                                       Fertilization!   I Spawning  Spawning         TPeriodicity Domain
                                     Egg Developmentl   type         behavior          mpora  ceue

                                                                 r

    Blue mussel
     Mytilus edulis
    Pacific oyster              *      *                                           *  *  *
     Crassostrea gigas
    Horseneck gaper              *     *             *     *  *                                                      *
     Tresus capax
    Pacific gaper               *      *             *      **
     Tresus nuttaffli
    California jackknife clam   *      *             *     *  *         a                    0 *  *  *  *
     Tagelus califomianus
    Pacific littleneck clam
     Pratothaca staminea
    Manila clam
     Venerupisjaponica
    Softshell                    *            a
     Mya arenaria
    Geoduck
     Panopea abrupta
    Bay shrimp                                                                                                       0
     Crangon franciscorumr
    Dungeness crab                  0  0             0                   0  41111 0  0  6                            0
     Cancer magister
    Leopard shark
     Triakis semifasciata
    Green sturgeon               *     *         *  *                       *  *  *
     Acipenser medirostris
    White sturgeon               *     *         *  *                       *  *  *                    *
     Acipenser transmontanus
    American shad
     Alosa sapidissima                                                                                                     0
    Pacific herring
     Clupea pallasi .
    Deepbody anchovy             *     *             *      *  *                *  *
     Anchoa compressa
    Slough anchovy               *     *             *      *  *                *  *
     Anchoa delicatissima
    Northern anchovy             0     0             0      0  0                   0  0  :400  0  0  0               0  0
     Engraulis mordax
    Cutthroat trout              *     * a                  * a
     Oncorhynchus clark,
    Pink salmon
     Oncorhynchus gorbuscha   0           0  a
    Chum salmon
     Oncorhynchus keta
    Coho salmon                  0     0                                                         0  0                       4
     Oncorhynchus kisutch
    Steelhead                    *     *          *         *
     Oncorhynchus mykiss
    Sockeye salmon
     Oncorhynchusnerka                 5    0                                             *
    Chinook salmon
     Oncorhynchus tshawytscha

                                                               324

                                                                                                                            Appendix 5D continued

                                                     I                             ~~~~~~REPRODUCTION

                                           Egg Development tySpewin    behavnior           Temporal Schedule        Periodicity    Domain

                       Hypomesu pretious                                           0  0  0a   6:4 a0  0                     0.

    Longfin smelt                                                                                 0  0
     Spirinchus thaleichthys            00
    Eulachon
     Thaleichthys pacificus                       C5            065
    Pacific tomcod               *      *                                     06  0                ...
     Microgadus proximus
    Topsmielt                                                                                                           @
     Atherinops affinis          C      Se                                              .
    Jacksmelt                        0                              0                   0 CS0

    Threespine stickleback              00000000
     Gasterosteus aculeatus             550                                  5CC0565
    Striped bass                 *      *
     Morone saxatilis
    Kelp bass                                                                    0a80
     Paralabrax clathralus       0                    S         SS5
    Barred sand bass                    000
     Paralabrax nebulifer        0      00                      60                   @     6
    White seabass                                     00
     Atractoscion nobilis        S      500                                         0
    White croaker                *      *             *      *  *             *  *  *  *  *  a
     Genyonemus lineatus
    Shiner perch
     Cymnatogasteraggregata         S          S      5S                            C
    Pacific sand lance*****
     Ammodytes hexapterus
    Arrow goby                   0      0411
     Clevelandia ios0                                                         se        .     .         .S
    Lingood                      *      *                           **        a**000
     Ophiodon elongatuis
    Pacific staghorn sculpin     *      *                                                      *  *  *  *              *  *
     Leptocattus armatus
    California halibut
     Paralichthys califomnicus   0         0                    0             50      0               0S 0
    Diamond turbot                                                                      000000
     Hypsopsetta guttulata       0      S0                                                                             0
    English sole                 *      *                                     *  a             *  *  *  *
     Pleuronectes vetulus
    Starry flounder*****
     Platichthys stellatus                            0 S

                                                                                 325

    326

                                                 BIOGEOGRAPHY
    Marine- Distribution of life stages in seawater areas.
       *Beach- Exposed shore areas receiving ocean waves and wash.
       *Continental shelf- Waters over the gradually-sloping continental seabed from shore to a depth of
                          about 200 m.
       *Continental slope- Waters over the steeply-sloping seabed from continental shelf to 1000 m.
       *Oceanic- The open ocean waters beyond the continental shelf. Defined here as the ocean beyond
                 the continental slope.

    Estuarine- Distribution of life stages in estuarine areas.
       Salinity Ranae: SAB (Strateaic Assessment Branch classification)
       * Tidal fresh- Salinities of 0.0-0.5%oo.
       *Mixing- Salinities of 0.5-25.0?/0.
       *Seawater- Salinities >25%0.

       Salinity Ranae: Venice classification
       *Limnetic- Salinities of 0.0-0.5%0.
       * Oligohaline-Salinities of 0.5-5.0%/o.
       *Mesohaline- Salinities of 5-18%o.
       *Polyhaline- Salinities of 18-30%o.
       *Euhaline- Salinities >30%0.

       Estuary type
       *Drowned river- Estuaries resulting from valleys being inundated by rising sea levels (e.g., Grays
                       Harbor and Columbia River estuary).
       *Bar-built- Estuaries resulting from the building of barrier islands or spits (e.g., Netarts Bay and
                  Humboldt Bay).
       *Fjord- Glacier-formed estuaries with deeply-carved, steep-sided channels (e.g., Puget Sound and
              Hood Canal).
       * Tectonic- Estuaries formed by faulting orsinking of the earth's crust (e.g., Tomales Bay and South San
                  Francisco Bay).

       Stratification
       *Highly-Very little mixing between surface and bottom layers, resulting in marked differences between
               surface and bottom salinities.
       *Moderately- Moderate mixing between surface and bottom layers primarily due to tidal-induced
                    turbulence. Surface salinities are usually lower than bottom salinities.
      *Homogeneous- High mixing of surface and bottom layers resulting in equivalent salinities.

    Riverine- Distribution of life stages in freshwater areas.
      *Coastalplain- River portions in the relatively flat land along a coast.
      -Piedmont- River portions at the base of mountains.
      * Upland- River portions in mountainous areas.

                                             HABITAT ASSOCIATIONS

    Habitats- General habitat of life stages.
      *Lake- Freshwater non-flowing areas with riverine connections to the sea.
      *River/stream- Areas with flowing fresh water.
      *Estuarine- Embayment with tidal fresh, mixing, and seawater zones.
      *Bay- Semi-enclosed water body that has predominantly seawater salinities.
      *Coastal (high energy)- Nearshore areas subject to significant wave or current action.
      *Coastal (low energy)- Nearshore areas subject to only minor wave or current action.
      *Offshore- Offshore areas beyond the coastal high or low energy areas.

                                                         327

    Appendix 6 continued

    Substrate preference- Size of substrate that life stages reside on or in.
      *Mud/clay/silt- Fine substrates <0.0625 mm in diameter.
      *Sand/granule- Substrates 0.0625-4.0 mm in diameter.
      *Pebble- Substrates 4-64 mm in diameter.
      PCobble- Substrates 64-256 mm in diameter.
      -Boulder- Large substrate >256 mm in diameter.
      �Rocky outcrop (bedrock)- Exposed solid rock.
      *Estuarine vegetation- Aquatic plants within an estuary.
      �Marine vegetation- Aquatic plants in marine waters.
      'None- No reported preference.

    Domain- Specific habitat where life stages occur.
      Benthic-Littoral
      �Intertidal (0-3 m)- On the bottom from the high tide mark to depths of 3 m.
      �Subtidal (3-10 m)- On the bottom at depths of 3-10 m.

       Benthic-Sublittoral
      �Inner shelf (10-50 m)- On the bottom of the continental shelf at depths of 10-50 m.
      �Middle shelf (50-100 m)- On the bottom of the continental shelf at depths of 50-100 m.
      *Outer shelf (100-200 m)- On the bottom to the edge of the continental shelf at depths of 100-200 m.

       Benthic-Bathval
      *Mesobenthal (200-500 m)- On the bottom of the continental slope at depths of 200-500 m.
      *Bathyobenthal (>500 m)- On the bottom of and beyond the continental slope at depths >500 m.

       Pelaiic
      -Neritic- Residing within the water column from the shore to the edge of the continental shelf.
       Oceanic- Residing within the water column beyond the edge of the continental shelf.

       Estuarine
       'Mainstem channel- The deep, drowned river channel of an estuary
       -Subsidiary channel- Small tributary channels emptying into the mainstem channel of an estuary.
       -Channel edge- Rim of an estuarine channel where the bottom slopes upward and meets shallow flats.
       'Intertidal flat- Shallow, often almost level estuarine areas alternately covered and left bare by tidal waters.

                               BIOLOGICAL ATTRIBUTES AND ECONOMIC VALUE
    Feeding type- Trophic role of life stages.
       'Carnivore- A flesh-eating organism.
       'Herbivore- A plant-eating organism.
       'Omnivore- An organism that eats both plants and animals.
       'Planktivore- A plankton-eating organism.
      �Detritivore- A detritus-eating organism.

    Spatial strategy- Use of habitats by life stages.
       -Coastal migrant- An organism which migrates within nearshore waters of the continental shelf.
       �Ocean migrant- An organism which migrates in ocean waters beyond the continental shelf.
       -Freshwater resident- An organism which resides primarily in freshwater habitats.
       �Estuarine resident- An organism which resides primarily in estuarine habitats (salinities 20.5 and <25%0).
       -Marine resident- An organism which resides primarily in seawater habitats (salinities >25%o).

    Longevity- Average lifespan of a particular life stage (  day to >20 years).

    Economic Value- Monetary worth (direct and indirect) from harvesting a species.
       �Recreational- Harvested by sport anglers.
       'Commercial- Harvested by professional fishermen for sale in markets.

                                                        328

                                                                                         Appendix 6 continued
                                                REPRODUCTION
    Fertilization/Egg Development- Method of egg fertilization and development.
       *External- Egg fertilization occurs after eggs and sperm are shed into the water.
       -Internal- Egg fertilization occurs when a male inseminates an egg within a female.
       -Oviparous- Eggs are laid and fertilized externally.
       �Ovoviviparous- Eggs are fertilized and incubated internally, and usually released as larvae. Little or
                       no maternal nourishment is provided.
       * Viviparous- Eggs are fertilized, incubated, and develop internally until birth. Maternal nourishment is
                       provided.

    Mating Type- Mate selection strategy.
      *Monogamous- A single male and a single female pair for a prolonged and exclusive relationship.
       *Polygamous- A male mates with numerous females or vice-versa.
       *Broadcast spawner- Numerous males and females release gametes during mass spawning.

    Spawning- Spawning mode.
      -Anadromous- Species spends most of its life at sea but migrates to fresh water to spawn.
      *Iteroparous- Species reproduces repeatedly during a lifetime.
      *Semelparous- Species reproduces only once during a lifetime.
       -Batch spawn- Species spawns (releases gametes) several times during a reproductive period.

    Parental Care- Type of egg protection.
      *Protected- Eggs are protected by parent(s); eggs are buoyant or attached to substrates, but not buried.
      *Nests- Eggs develop in the shelter of a nest.

    Temporal Schedule- Period when spawning typically occurs.
      *Early spring- From mid-March through April.
      *Late spring- From May to mid-June.
      *Early summer- From mid-June through July.
      -Late summer- From August to mid-September.
      -Early fall- From mid-September through October.
      -Late fall- From November to mid-December.
      *Early winter- From mid-December through January.
      *Late winter- From February to mid-March.

    Periodicity- Frequency of spawning events.
      -Annual spawning- Spawning once each year, usually during a restricted season.
      -2 or more per year- Spawning more than once each year (more than one spawning season).
      *2 or more years- Spawning events separated by at least two years.
      *Undescribed- Spawning frequency not documented.

    Domain- Location of spawning.
      -Oceans- Spawning occurs primarily in open marine waters.
      -Estuaries- Spawning occurs primarily in estuarine waters (to head of tide).
      -Rivers- Spawning occurs primarily in fresh water, above head of tide.

                                                       329

    The Strategic Environmental Assessments (SEA) Division of NOAA's Office of Ocean Resources Conservation
    and Assessment (ORCA) was created in response to'the need for comprehensive information on the effects of
    human activities on the Nation's coastal ocean. The SEA Division performs assessments of the estuarine and
    coastal environments and of the resources of the U.S. Exclusive Economic Zone (EEZ).

    In June 1985, the National Oceanic and Atmospheric Administration (NOAA) began a program to develop a
    comprehensive information base on the life history, relative abundance and distribution of selected fishes and
    invertebrates in estuaries throughout the Nation (Monaco 1986). This program, the Estuarine Living Marine
    Resources (ELMR) program, is conducted jointly by the Strategic Environmental Assessments Division of the
    Office of Ocean Resources Conservation and Assessment and laboratories of the National Marine Fisheries
    Service (NMFS). Currently, the Point Adams (Hammond), OR; Galveston, TX; Beaufort, NC; and Oxford, MD
    laboratories are compiling information for the contiguous West Coast, Gulf of Mexico, Southeast, and Northeast
    regions. Also, the Virginia Institute of Marine Science is compiling data for the Chesapeake Bay area.

    Three salinity zones, as defined in Volume 1 of NOAA's National Estuarine Inventory Data Atlas (NOAA 1985),
    provided the spatial framework for organizing information on species distribution and abundance within each
    estuary. These salinity zones are tidal fresh (0.0 to 0.5%o), mixing (0.5 to 25.0%o), and seawater (25.0%o and
    greater). The primary data developed for each species for each salinity zone include spatial and temporal
    distributions and relative abundance by life stage (i.e., adult, spawning or mating adults, juvenile, larva, and egg).
    In addition, a detailed estuarine life history summary is written for each species.

    Additional information on this or other programs of NOAA's Strategic Environmental Assessments Division is
    available from:
                                    Strategic Environmental Assessments Division
                              Office of Ocean Resources Conservation and Assessment
                                  National Oceanic and Atmospheric Administration
                                            6001 Executive Blvd., Rm. 220
                                              Rockville, Maryland 20852
                                          FTS/Comm. (301) 443-0453/8921

    Reports available from NOAA's Estuarine Living Marine Resources program include:

    Monaco, M. E., T. E. Czapla, D. M. Nelson, and M. E. Pattilo. 1989. Distribution and abundance of fishes and invertebrates
    in Texas estuaries. ELMR Rep. No. 3. Strategic Assessment Branch, NOS/NOAA, Rockville, MD, 107 p.

    Monaco, M. E., D. M. Nelson, R. L. Emmett, and S. A. Hinton. 1990. Distribution and abundance of fishes and invertebrates
    in west coast estuaries, Volume I: Data summaries. ELMR Rep. No. 4. Strategic Assessment Branch, NOS/NOAA, Rockville,
    MD, 240 p.

    Bulger, A. J., B. P. Hayden, M. E. Monaco, D. M. Nelson, and M. G. McCormick-Ray. 1990. Aproposed estuarine classification:
    analysis of species salinity ranges. ELMR Rep. No. 5. Strategic Assessment Branch, NOS/NOAA, Rockville, MD, 28 p.

    Williams, C. W., D. M. Nelson, M. E. Monaco, L. C. Clements, S. L. Stone, L. R. Settle, C. lancu, and E. A. Irlandi. 1990.
    Distribution and abundance of fishes and invertebrates in eastern Gulf of Mexico estuaries. ELMR Rep. No. 6. Strategic
    Assessment Branch, NOS/NOAA, Rockville, MD, 105 p.

    Czapla, T. E., M. E. Pattillo, D. M. Nelson, and M. E. Monaco. 1991. Distribution and abundance of fishes and invertebrates
    in central Gulf of Mexico estuaries. ELMR Rep. No. 7. NOAA/NOS Strategic Environmental Assessments Division, Rockville,
    MD, 82p.

    Emmett, R. L., S. L. Stone, S. A. Hinton, and M. E. Monaco. 1991. Distribution and abundance of fishes and invertebrates
    in west coast estuaries, Volume II: Species life history summaries. ELMR Rep. No. 8. NOAA/NOS Strategic Environmental
    Assessments Division, Rockville, MD, 329 p.

    D. M. Nelson, E. A. Irlandi, L. R. Settle, L. C. Coston-Clements, and M. E. Monaco. 1991. Distribution and abundance of fishes
    and invertebrates in southeast estuaries. ELMR Rep. No. 9. NOAA/NOS Strategic Environmental Assessments Division,
    Rockville, MD, 167 p.