[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

















                            ï¿½ 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,'/,~;:ï¿½    {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                 _       ï¿½
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 0.5ï¿½/),      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.
   ï¿½ 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:
   ï¿½ Rare- species is present but not frequently
    encountered.                                          * Spatial distribution and relative abundance
                                                          ï¿½ Temporal distribution
   ï¿½ Notpresent- species or life stage not found,          ï¿½ 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     ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½    ï¿½    ï¿½ ï¿½  ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½
   Pacific oyster                    **                            * 
   horseneck gaper  0 ï¿½ ï¿½ ï¿½ ï¿½        ï¿½ ï¿½    ï¿½ ï¿½ ï¿½    ï¿½
   Pacific gaper    0 0         0          
   California jackknife clam
   Pacific littleneckclam   0    ï¿½    ï¿½ ï¿½                 ï¿½ ï¿½  ï¿½ ï¿½ ï¿½     ï¿½ @                         0
   Manila clam     0 0 0ï¿½      ï¿½        ï¿½    ï¿½  ï¿½  
   Eastem softshell clam  *ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½       

   bay shrimp      * * * * * * * * *  *ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ 0                  ï¿½ ï¿½ ï¿½ ï¿½
   Dungenesscrab   000ï¿½ï¿½ï¿½ï¿½ 0 000
   leopard shark                                                *   * *  
   green sturgeon          ï¿½      ï¿½ ï¿½        * ï¿½  
   white sturgeon         *ï¿½ * *ï¿½ **
   American shad        ï¿½ ï¿½ ï¿½ ï¿½              * * - - -  
   Pacific herring      00 0 0 0 *                                   0 *    * 
   deepbody anchovy   *   
   slbugh anchovy                                                                       *    * -
   northern anchovy  0 0 0 0 0000000 * * * * *   
   cutthroattrout  ï¿½ ï¿½ ï¿½ ï¿½ï¿½ï¿½ï¿½ï¿½ï¿½* *
   pink salmon        * 
   chum salmon     ï¿½ï¿½*   ï¿½ ****0ï¿½   ï¿½0
   cohosalmon      ï¿½ï¿½*       ****                * *ï¿½ ï¿½*
   steelhead       * -0                            0- -0-
   sockeye salmon    ï¿½0 
   chinooksalmon   0 0 0ï¿½         ï¿½        ï¿½0 0      0    0 
   sud smelt       ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ -  -    0000
   Iongfinsmelt    0    0 0 
   eulachon                0 ï¿½           ï¿½    ï¿½
   Pacific tomcod  0ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½ ï¿½  
   topsmelt                           *      0 **-
   jacksmelt                                                                            -
   threespine stickleback 0ï¿½ ï¿½0 000000000 00                0 ï¿½ ï¿½*   
   striped bass                                            ï¿½ ï¿½ ï¿½        ï¿½ ï¿½ ï¿½
   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 ï¿½ ï¿½ ï¿½ ï¿½ï¿½                                                       - 0   0
   Califomia halibut                                                  0000000000000
   diamond turbot  __                        V0000                                           00
   English sole    * *ï¿½'                         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     ï¿½      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 ï¿½                III ï¿½/ Iï¿½                   ï¿½              ï¿½!

                                                                 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 ï¿½1ï¿½ï¿½ 11ï¿½ 1             I~
              stickleback   Is                       I0II* I  01  ï¿½11ï¿½  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   1ï¿½ï¿½11ï¿½ï¿½                    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 *           ï¿½ ï¿½   3   Abundant     in Japan, Australia, and the North Atlantic (Haderlie
                                   O   Common
        Grays Harbor0 0    C       ï¿½   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   ï¿½                              pilingsandtheundersidesoffloatingdocks(Rickettset
           CoosBay a3  3  (3 (ï¿½                         al. 1985).
         Rogue River '4    i
        Klamath River                                    Life Mode
        Humboldt Bay a ï¿½    ï¿½ ï¿½                          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     ï¿½      * IncdudesCentralSan  ropes, etc.). Juvenilesandadultsdonotneed lightand
                                  Francisco, Suisun,
   South San Fran. Bay S U S    ï¿½   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   18ï¿½C (late spring or summer) (D. Tufts, Willapa Bay
30ï¿½C (Bernard 1983). Itcanwithstandtemperaturesof   Shellfish Lab., Washington Department of Fisheries,
1.7-26.7ï¿½C (Cheney and Mumford  1986), but    P.O. Box 190, Ocean Park, WA, pers. comm.). Mussels
temperatures above 20ï¿½C appear to be stressful (Hines   in British waters spawn when water temperatures rise
1979). Trochophore development occurs best within a   from 9.5ï¿½C to 11 -12.50C (Chipperfield 1953). In Puget
salinity range of 30 to 40%o and temperatures of 8-1 8ï¿½C    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 25ï¿½C. 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 ï¿½    ï¿½        Relative abundance:    successfully compete with the native oyster (Ostrea
         Hood Canal *ï¿½    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        Rareï¿½
         Willapa Bay 6 ï¿½         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 ï¿½    ï¿½                              species (Menzel 1974, Wolotira et al. 1989).
         Rogue River
       Klamath River                                   Within Study Area: The Pacific oyster is found in most
       Humboldt Bay ï¿½    ï¿½                              Pacific coast estuaries from Morro Bay, California, to
           EelRiver                                     Skagit Bay, Washington, where estuarine physical
        Tomales Bay ï¿½    ï¿½                              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 ï¿½    ï¿½                               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 -4ï¿½C during low tides and
oysters (C. virginica). These exotics include sponges,    watertemperatures of 4-36ï¿½C (Quayle 1988, Wolotira


                                                    21






Pacific oystercontinued
et al. 1989), and spawns at water temperatures of 14-   notoccur annually. Therefore, spawning is sporadic or
30ï¿½C, but only rarely below 18ï¿½C (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,
21-23ï¿½C (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 3ï¿½C, but   occasionally in Willapa Bay, Washington (Quayle 1988,
growth stops when temperatures drop below 10ï¿½C    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 17ï¿½C (ranges 15-18ï¿½C), 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 24ï¿½C (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 18ï¿½C) 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 ï¿½  ] 13 ï¿½   Life stage:      Institute of Washington 1981).
         Netarts Bay 3  13  ï¿½ ï¿½   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 2-20ï¿½C (Bernard 1983a). Larvae do not survive at
        SiuslawRiver ï¿½ ï¿½ ï¿½ ï¿½ ï¿½                        20ï¿½C (Bourne and Smith 1972a). Optimum conditions
       UmpquaRiver i                ' / i i          forsomaticgrowthare13ï¿½Cwatertemperatures (range
           Coos Bay *  ï¿½       ï¿½ ï¿½                    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 15ï¿½C,    highly variable on some clam beds, resulting in beds
26 days at 10ï¿½C, and 34 days at 5ï¿½C (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  (   ï¿½*  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 ï¿½                           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 >35ï¿½C 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 35.5ï¿½C in December and 37.6ï¿½C 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 ï¿½ 6  ï¿½   Relative abundance:   Habitat
        Hood Canal ï¿½ ï¿½ ï¿½  ï¿½ ï¿½   Highly abundant       Type: Eggs and larvae are estuarine-neritic. Adults
                                 O Common
         skagit Bay *ï¿½ï¿½ï¿½ï¿½ @ @ @       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 25ï¿½C (Glude 1978, Bernard
      San Pedro Bay O O O O 0                          1983a). Water temperatures above 25ï¿½C 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 12-18ï¿½C, 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 8ï¿½C (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 *ï¿½ ï¿½ ï¿½  ï¿½6   Relative abundance:   California, Puget Sound, and other estuaries.
        Hood Canal 1] t 9 1 i    ï¿½*  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 ï¿½ ï¿½ ï¿½ ï¿½ ï¿½   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  * ï¿½  ï¿½  ï¿½    ï¿½   IncdudesCentralSan  Northwest inlets and bays (Chew 1989). However,
                                SFrancisco, Suisun,
  South San Fran. Bay ï¿½*. ï¿½ ï¿½   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 23-25ï¿½C, but
                                                      they can withstand temperatures of 0-36ï¿½C (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%ï¿½ salinity (range of
are common commensals within the mantle cavity of   24-31%o), 16ï¿½C temperature (range of 13-21 ï¿½C), 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 <10ï¿½C (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   15ï¿½Cformaturation, 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 12ï¿½C 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 15ï¿½0C (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 13-16ï¿½C. 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 22.0ï¿½C 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     ï¿½*  Highly abundant   Eggs and larvae are found in the estuarine and
         Skagit Bay *ï¿½ ï¿½ ï¿½ ï¿½ ï¿½  Abundant               nearshore marine plankton, while juveniles and adults
       Grays Harbor &9 W (        0 '4ï¿½ 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 ï¿½  ï¿½  ï¿½  ï¿½ 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   ï¿½           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       ï¿½        Laï¿½ ï¿½  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 ï¿½ ï¿½ ï¿½ ï¿½ ï¿½                            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 -1.7ï¿½C (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 4ï¿½C to22ï¿½C. Spawning
Life Mode                                               on the Pacific coast appears to occur at temperatures
Eggs and larval stages are planktonic; juveniles and    between 10 and 15ï¿½C (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 ï¿½ï¿½  Relative abundance:  Abbott 1980, Maclntyre et al. 1986).
         Hood Canal  1 ï¿½3 (   3   * ï¿½   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                                  20ï¿½C (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 12-14ï¿½C, but spawning occurs in
estimate of 117.6 million geoducks was made for   temperatures from 8-160ï¿½C (Goodwin 1976). The best
33,799 acres of subtidal beds surveyed in Puget Sound    temperature for larval survival is between 6 and 16ï¿½C
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 16ï¿½C
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 17.60ï¿½C, and 47 days at 14-15ï¿½C    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.












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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   ï¿½ ï¿½  ï¿½ 6    ï¿½   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 5.2-21.3ï¿½C; ovigerous females prefer
        Tillamook Bay ï¿½  ï¿½ ï¿½ ï¿½    Life stage:
         Netamkrtsay C 0  *    (  UfA stagedults        temperaturesof6.8-19.2ï¿½C(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    ï¿½ ï¿½ ï¿½ ï¿½ ï¿½
          Siuslaw River  3 O * O O                      hightemperatures in the summerincrease movements
        Siuslaw River ï¿½i ï¿½i ï¿½ ï¿½i ï¿½i
        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                        4.5-17.0ï¿½C (Khorram and Knight 1977, Siegfried 1980).
         Tomales Bay O 0    C   0 
   Cent. SanFran. Bay'  3 c ï¿½ ï¿½ .    * 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 17.5ï¿½C    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






Bay shrimp continued
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.




























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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

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                                                                                 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 ï¿½  0  0    ï¿½   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       eï¿½ ï¿½                         Megalopae are often found on the hydrozoan Velella
        Tillamook Bay             Life stage:          velella (Wickham 1979, Stevens and Armstrong 1985).
         Netarts Bay DO  * ï¿½      M- Mating           Juveniles and adults are epibenthic.
          Siletz River 0    (3 3   J - Juveniles
               ~Yaquina Bay (     ~L- Larvae
         Yaquina Bay 1 :  aï¿½       E- Eggs             Habitat
         AlsesaRiver  a a                             Iype: Eggs adhere to pleopods of female crabs in
        Siuslaw River  ï¿½ ï¿½  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 (<10ï¿½C) 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 10.0-14.0ï¿½Candsalinities 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 >20.0-25.0ï¿½C 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-
northern California Dungeness crab (Cancermagister)    82, Wash. Dept. Fish., Olympia, WA.
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
1-25. Academic Press, NY.                               resource. Calif. Fish Game, Fish Bull. 172:7-24.

Buchanan, D. W., and R. E. Milleman.  1969. The    Demory, D. 1985. An overview of Oregon Dungeness
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Dana. Biol. Bull. (Woods Hole) 137(2):250-255.          In B. R. Melteff (coordinator),  Proceedings of the
                                                       symposium on Dungeness crab biology and
Buchanan, D. V., R. E. Millemann, and N. E. Stewart.    management, p. 27-32. Lowell Wakefield Fisheries
1970.  Effects of the insecticide Sevin on various    Symposia Series, Univ. Alaska, Alaska Sea Grant
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                                                       Kelly. 1983. Laboratory cultivation of the Dungeness
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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
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H. Mallon, and R. E. Millemann. 1979. Toxicity of the    and southern Washington coasts, May-September
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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
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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.
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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
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                                                      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






Dungeness crab continued
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             ï¿½   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 ï¿½ ï¿½ 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  ï¿½ ï¿½ 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              ï¿½   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 12.7ï¿½C
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             ï¿½   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 10-16ï¿½C 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 14-16ï¿½C, although    approximately12 daysafterhatching (Anderson 1988).
incubation is possible from 10-18ï¿½0C (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 18ï¿½C 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 15.5-26.6ï¿½C (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            ï¿½   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 ï¿½
       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                          10-15ï¿½C, 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 18.5ï¿½C, 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  ï¿½ 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 3-15ï¿½C    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
15-18ï¿½C (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 5-14ï¿½C 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 ï¿½ ï¿½ ï¿½ S    ï¿½   Relative abundan   and Hourston 1985).  Optimum temperatures and
           Hood Canal    ï¿½)  ï¿½ * -    ï¿½   Highly abun4  salinities for egg and larval survival appear to be 5.5-
            SkagitBay * a i a g    ï¿½   Abundant          8.7ï¿½C 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     normallyabove9ï¿½C (Barnhart 1988). Salinitytolerances
        Columbia River O O ï¿½ O O                        of larvae are affected by temperature and salinity
         Nehalem Bay      xi ï¿½   i R   during egg incubation (Alderdice and Hourston 1985).
         Tillamook Bay Li ï¿½ Li ï¿½ ï¿½    Life stage:       Turbidity in estuaries may increase larval survival
           Netarts Bay 13ï¿½  O O      S - spawning adul  (Boehlert and Morgan 1985).
           Siletz River  O           J -   Juveniles
          Yaqulna Bay    i ï¿½    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 ï¿½ L    ï¿½ S  ï¿½                      (Morrow 1980). Adults typically move onshore during
            Coos Bay ï¿½  ï¿½  ï¿½                            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 ï¿½                       distributions of adults for many Pacific coast stocks are
          TomalesBay ï¿½ ï¿½ ï¿½     ï¿½*                        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 10.7ï¿½C, 14-15 days at 8.5ï¿½C,   desiccation, freezing, low oxygen, wave action, and
and 28-40 days at 4.4ï¿½C (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 >25ï¿½C (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              ï¿½   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 * ï¿½
         Mission Bay ( t    S *I 3                    Factors Influencina PoDulations: This species is often
       San Diego Bayï¿½ ï¿½ ï¿½  ï¿½                          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    ï¿½   Highly abundant  temperaturesof12-15ï¿½Candusuallywithin10mofthe
           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 10.0-19.7ï¿½C
                        mras o  O    O      Rare
          Willapa Bay ï¿½    ï¿½ O  Blank Not present      (mostly 14.0-17.40C), and juveniles and adults at 5.0-
       Columbia River ï¿½    ï¿½ O O                       25.0ï¿½C.  The lower lethal temperature for juveniles
         Nehalem Bay O    a O                           appears to be 7ï¿½C, but at 1 0.0ï¿½C larvae do not develop
        Tillamook Bay O    ï¿½ O    Ufe stage:           properly.  Temperatures above 25ï¿½C 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    ï¿½  adults move into estuaries during spring and summer
        Klamath River i ï¿½ *                            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 ï¿½ ï¿½                        night to gulp air and inflate their swim bladder (Hunter
        SanPedroBay  ï¿½ ï¿½ ï¿½ ï¿½  ï¿½
         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 15ï¿½C. 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.
anchovy (Engraulis mordax) in the California fishery.
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-
1983. A field guide to Pacific coast fishes of North    855.
America. Houghton Mifflin Co., Boston, MA, 336 p.
                                                       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.
of fishery statistics, 1983: catches and landings. FAO,
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-
                                                       618.
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.
effects of environmental variations on the eggs and
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.
MBC Applied Environmental Sciences. 1987. Ecology
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.

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                                                                                 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        ï¿½   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 260ï¿½C, 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 10ï¿½C, but spawning occurs    being iteroparous.
over a range from 6-17ï¿½C (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.






























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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 40ï¿½N 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 ï¿½    ï¿½       Relative abundance:  estuarine areas (Helle et al. 1964, McNeil 1966). Fry
          Hood Canal ï¿½  1   0     ï¿½   Highly abundant    are riverine initially, but soon move downstream and
           Skagit Bay aï¿½    ï¿½   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 7.2-12.80ï¿½C, with
         Alamitos Bay  0incubation temperatures of 4.4-13.3ï¿½C providing the
         Anaheimwport Bay                             best hatching (Bell 1984). Optimum temperatures for
          Mission Bay                                  pink salmon are 5.6-14.40C, with 0.0ï¿½C and 25.6ï¿½C
          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
Bell, M. C. 1984. Fisheries handbook of engineering    Fresh, K. L., D. Rabin, C. Simenstad, E. O. Salo, K.
requirements and biological criteria.  Fish passage    Garrison, and L. Matheson.  1979.  Fish ecology
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.
80-M-0567).
                                                       Fry, D. H., Jr. 1973. Anadromous fishes of California.
Bonar, S. A., G. B. Pauley, and G. L. Thomas. 1989.    Calif. Dept. Fish. Game, Sacramento, CA, 112 p.
Species profiles: life histories and environmental
requirements of coastal fishes and invertebrates (Pacific    Gerke, R. J. 1972. Food of juvenile pink and chum
Northwest)-pink salmon. U.S. Fish Wildl. Serv. Biol.    salmon in Puget Sound, Washington. Tech. Rep. No.
Rep. 82(11.88). U.S. Army Corps Eng., TR EL-82-4,    10, Wash. Dept. Fish., Olympia, WA, 7 p.
18 p.
                                                       Godin, J.-G. J. 1981. Daily patterns of feeding behavior,
Brannon, E. L.  1982.  Orientation mechanisms of   daily rations, and diets of juvenile pink salmon
homing salmonids. In E. L. Brannon and E. O. Salo    (Oncorhynchus gorbuscha) in two marine bays of
(editors), Proceedingsof the salmon and trout migratory    British Columbia. Can. J. Fish. Aquat. Sci. 38:10-15.
behavior symposium, p. 219-227. School Fish., Univ.
Wash., Seattle, WA.                                     Hallock, R. J., and D. H. Fry. 1967. Five species of
                                                       salmon, Oncorhynchus, in the Sacramento River,
Brodeur, R. D., H. V. Lorz, and W. G. Pearcy. 1987.    California. Calif. Fish Game 53(1):5-22.
Food habits and diet variations of pelagic nekton off
Oregon and Washington, 1979-1984.  U.S. Dept.    Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.
Comm., NOAA, Tech. Rep. NMFS 57, 32 p.                  Board Can., Bull. No. 180, 740 p.

Forrester, C. R. (compiler). 1981a. Statistical yearbook    Hartt, A. C., and M. B. Dell.  1986.  Early oceanic
1977. Internat. North Pac. Fish. Comm., Vancouver,    migrations and growth of juvenile Pacific salmon and
B.C., Canada, 96 p.                                     steelhead trout. Internat. North Pac. Fish. Comm.,
                                                       Bull. No. 46:1-105.
Forrester, C. R. (compiler). 1981 b. Statistical yearbook
1978. Internat. North Pac. Fish. Comm., Vancouver,    Healey, M. C. 1980. The ecology of juvenile salmon in
B.C., Canada, 113 p.                                    Georgia Strait, British Columbia. In W. J. McNeil and
                                                       D. C. Himsworth (editors), Salmonid ecosystems of the
Fredin, R. A. 1980. Trends in North Pacific salmon    North Pacific, p. 203-229, Oregon State Univ. Press,
fisheries. In W. J. McNeil and D. C. Himsworth (editors),   Corvallis, OR.
Salmonid ecosystems of the North Pacific, p. 59-119,
Oregon State Univ. Press, Corvallis, OR.                Healey, M. C.  1982.  Juvenile Pacific salmon in
                                                       estuaries: the life support system. In V. S. Kennedy
Fredin, R. A., R. L. Major, R. G. Bakkala, and G. K.   (editor), Estuarinecomparisons, p. 315-341, Academic
Tanonaka. 1977. Pacific salmon and the high seas    Press, New York, NY.
salmon fisheries of Japan. NWAFC Proc. Rep., 324 p.
Northwest and Alaska Fish. Cent., Nat. Mar. Fish.   Helle, J. H., R. S. Williamson, and J. E. Bailey. 1964.
Serv., NOAA, Seattle, WA.                               Intertidal ecology and life history of pink salmon at
                                                       Olsen Creek, Prince William Sound, Alaska. U.S. Fish
Fresh, K. L. 1984.  Evaluation of potential species    Wildl. Serv., Spec. Sci. Rep. Fish. No. 483, 26 p.
interaction effects in the planning and selection of
salmonid enhancement projects. Report prepared by    Hunter, J. G. 1959. Survival and production of pink and
the species interaction workgroup ofthe enhancement    chum salmon in a coastal stream. J. Fish. Res. Board
planning team. NOAA, Nat. Mar. Fish. Serv., Seattle,    Can. 16(6):835-886.
WA, 80 p.
                                                       Ito, J. 1964. Food and feeding habits of Pacific salmon
Fresh, K. L., R. D. Cardwell, and R. R. Koons. 1981.    (genus Oncorhynchus) in their oceanic life.  Bull.
Food habitsof Pacificsalmon, baitfish, andtheirpotential    Hokkaido Reg. Fish. Lab. 29:85-97. (Fish. Res. Board
competitors and predators in marine waters of    Can., Trans. Ser. No. 1309).
Washington, August 1978 to September 1979. Prog.
Rep. No. 145, Wash. Dept. Fish., Olympia, WA, 58 p.

                                                   124






                                                                                  Pink salmon continued
Kaczynski, V. W., R. J. Feller, J. Clayton, and R. J.   Merrell, T. R., Jr. 1962. Freshwater survival of pink
Gerke. 1973. Trophic analysis of juvenile pink and    salmon at Sashin Creek, Alaska. In N. J. Wilimovsky
chum salmon (Oncorhynchusgorbuscha and 0. keta)    (editor), Symposium on pink salmon. H. R. MacMillan
in Puget Sound. J. Fish. Res. Board Can. 30:1003-    lectures in fisheries, p. 59-72. Univ. British Columbia,
1008.                                                   Vancouver, B.C., Canada.

LeBrasseur, R. J. 1966. Stomach contents of salmon    Morrow, J. E. 1980. The freshwater fishes of Alaska.
and steelhead trout in the northeastern Pacific Ocean.    Alaska Northw. Publ. Co., Anchorage, AK, 248 p.
J. Fish. Res. Board Can. 23(1):85-100.
                                                       National Marine Fisheries Service. 1986. Fisheries of
Levy, D. A., and T. G. Northcote.  1982. Juvenile    the United States, 1985. Current Fishery Statistics No.
salmon residency in a marsh area of the Fraser River    8368. U.S. Dept. Comm., NOAA, Nat. Mar. Fish Serv.,
estuary. Can. J. Fish. Aquat. Sci. 39:270-276.          Nat. Fish. Stat. Prog., Washington, D.C., 122 p.

Levy, D. A., T. G. Northcote, and G. J. Birch. 1979.    Neave, F. 1962. The observed fluctuations of pink
Juvenile salmon utilization of tidal channels in the    salmoninBritishColumbia. InN.J.Wilimovsky(editor),
Fraser River estuary, British Columbia. Tech. Rep. 23,    Symposium on pink salmon. H. R. MacMillan lectures
Westwater Res. Cent., Univ. British Columbia,    in fisheries, p.3-14. Univ. British Columbia, Vancouver,
Vancouver, B.C., Canada, 70 p.                          B.C., Canada.

Magnhagen, C. 1988. Predation risk and foraging in   Neave, F. 1966.  Pink salmon in British Columbia.
juvenile pink (Oncorhynchus gorbuscha) and chum    Internat. North Pac. Fish. Comm., Bull. No. 18:71-79.
salmon (0. keta). Can. J. Fish. Aquat. Sci. 45(4):592-
596.                                                    Pacific Marine Fisheries Commission.  1985. 37th
                                                       annual report of the Pacific Marine Fisheries
Martin, D. J., C. J. Whitmus, L. A. Brocklehurst, A. E.    Commission - for the year 1984.  Pac. Mar. Fish.
Nevissi, J. M Cox, and K. Kurrus. 1990. Effects of    Comm., Portland, OR, 35 p.
petroleum contaminated waterways on migratory
behavior of adult pink salmon.  Outer Contin. Shelf    Pacific Marine Fisheries Commission.  1987  39th
Envir. Asses. Prog., FinalRep. Principallnvest. 66:281-    annual report of the Pacific Marine  Fisheries
529                                                     Commission - for the year 1986.  Pac. Mar. Fish.
                                                       Comm., Portland, OR, 29 p.
McDonald, J. 1960. The behavior of Pacific salmon fry
during their downstream migration to freshwater and    Parker, R. R. 1971. Size selective predation among
saltwater nursery areas.  J. Fish. Res. Board Can.    juvenile salmonid fishes in a British Columbia inlet. J.
17(5):655-676.                                          Fish. Res. Board Can. 28:1503-1510.

McNeil, M. J.  1966.  Effect of the spawning bed    Pearcy, W., T. Nishiyama, T. Fujii, and K. Masuda.
environmenton reproductionof pinkandchumsalmon.    1984. Diel variations in the feeding habits of Pacific
Fish. Bull., U.S. 65(2):495-523.                        salmon caught in gill nets during a 24-hour period in the
                                                       Gulf of Alaska. Fish. Bull., U.S. 82(2):391-399.
McNeil, M. J. 1969. Survival of pink and chum salmon
eggs and alevins.  In T. G. Northcote (editor),    Peterman, R. M. 1980. Testing for density-dependent
Symposium on salmon and trout in streams.  H. R.    marine survival in Pacific salmonids. In W. J. McNeil
MacMillan lectures in fisheries, p. 101 -117. Univ. British   and D. C. Himsworth (editors), Salmonid ecosystems
Columbia, Vancouver, B.C., Canada.                      of the North Pacific, p. 1-24. Oregon State Univ. Press,
                                                       Corvallis, OR.
McNeil, M. J. 1980. Vulnerability of pink salmon
populations to natural and fishing mortality. In W. J.   Quinn, T. P. 1982. A model for salmon navigation on
McNeil and D. C. Himsworth (editors), Salmonid    the high seas. InE. L. Brannon and E. O. Salo (editors),
ecosystems of the North Pacific, p. 147-151. Oregon    Proceedingsofthe salmon andtrout migratory behavior
State Univ. Press, Corvallis, OR.                       symposium, p. 229-237. School Fish., Univ. Wash.,
                                                       Seattle, WA.
McPhail, J. D., and C. C. Lindsey. 1970. Freshwater
fishes of northwestern Canada and Alaska. Fish. Res.    Raleigh, R. F., and P. C. Nelson.  1985.  Habitat
Board Can., Bull. No. 73. 381 p.                        suitability index models and instream flow suitability

                                                   125






Pink salmon continued
curves: Pink salmon. U.S. Fish Wildl. Serv. Biol. Rep.    gorbuscha) in offshore waters of the North Pacific
82(10.109), 36 p.                                       Ocean. Internat. North Pac. Fish. Comm., Bull. No. 40,
                                                       195 p.
Reiser, D. W., and T. C. Bjornn. 1979. 1. Habitat
requirements of anadromous salmonids.  In W. R.    Takehama, S. (compiler). 1983. Statistical yearbook
Meehan (editor), Influence of forest and rangeland    1980. Internat. North Pac. Fish. Comm., Vancouver,
managementonanadromousfishhabitatinthewestern    B.C., Canada, 115 p.
United States and Canada, p. 1-54. USDA Forest
Service, Gen. Tech. Rep. PNW-96, Pacific Northw.    Wahle, R. J., and R. Z. Smith. 1979. A historical and
Forest Range Exp. Sta., Portland, OR.                  descriptive account of Pacific coast anadromous
                                                       salmonid rearing facilities and summaryof their releases
Ricker, W. E. 1976. Review of the rate of growth and    by region, 1960-76. U.S. Dept. Comm., NOAA Tech.
mortality of Pacific salmon in salt water, and noncatch    Rep. NMFS, Spec. Sci. Rep. Fish. No. 736, 35 p.
mortality caused by fishing. J. Fish. Res. Board Can.
33:1483-1524.                                           Washington Department of Fisheries.  1983.  Prog.
                                                       Rep. No. 184, Wash. Dept. Fish., Olympia, WA, 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   Washington Department of Fisheries and Northwest
of common and scientific names of fishes from the    Indian Fisheries Commission  1986.  Puget Sound
United States and Canada. Am. Fish. Soc. Spec. Publ.    salmon management periods and their derivations.
No. 12, Am. Fish. Soc., Bethesda, MD.                   Wash. Dept. Fish, Olympia, WA, 6 p.

Rombough, R. J. 1983. Effects of low pH on eyed    Wickett, W. P. 1962. Environmental variability and
embryos and alevins of Pacific salmon. Can. J. Fish.    reproduction potentials of pink salmon in British
Aquat. Sci. 40:1575-1582.                               Columbia. InN. J. Wilimovsky (editor.), Symposium on
                                                       pink salmon. H. R. MacMillan lectures in fisheries, p.
Scott, W. B., and E. J. Crossman. 1973. Freshwater    73-86.  Univ. British Columbia, Vancouver, B.C.,
fishes of Canada. Fish. Res. Board Can., Bull. No. 184,    Canada.
966 p.
                                                       Wydoski, R. S., and R. R.Whitney. 1979. Inland fishes
Shiino, S. 1976. List of common names of fishes of the    of Washington, Univ. Wash. Press, Seattle, WA, 220 p.
world, those prevailing among English-speaking
nations. Sci. Rep. Shima Marineland No. 4,
Kashikojima, Shima, Mie, Japan, 262 p.

Simenstad, C. A., K. L. Fresh, and E. O. Salo. 1982.
The role of Puget Sound and Washington coastal
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.

Squire, J. L., Jr., and S. E. Smith. 1977. Anglers' guide
to the United States Pacific coast - marine fish, fishing
grounds & facilities. Nat. Mar. Fish. Serv., NOAA,
Seattle, WA, 139 p.

Tabata, S. 1983. Oceanographic factors influencing
the distribution, migration and survival of salmonids in
the northeast Pacific Ocean-a review. InW. J. McNeil
and D. C. Himsworth (editors), Salmonid ecosystems
of the North Pacific, p. 128-160. Oregon State Univ.
Press, Corvallis, OR.

Takagi, K., K. V. Aro, A. C. Hartt, and M. B. Dell. 1981.
Distribution and origin of pink salmon (Oncorhynchus

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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. 40ï¿½N in the
it is sometimes fished in rivers that have large runs.   western Pacific Ocean and approximately lat. 44ï¿½N 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   :         ï¿½   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 ï¿½    *       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   ï¿½        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 from4.4-13.3ï¿½C (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                                  1.0-15.0ï¿½C, but prefer2.0-11.0ï¿½C. 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 8.3-15.6ï¿½C.  The upper lethal
                                                       temperature is 25.6ï¿½C, and the lower lethal temperature
Oregon (Table 1) (Atkinson et al. 1967, Ratti 1979).   is 0.0ï¿½C (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
References                                            the chum salmon, Oncorhynchus keta (Walbaum) 1792.
                                                      FAO Species Synopsis No. 41. U.S. Fish Wildl. Circ.
Abrahamson, P. 1988. Adetailedlistingoftheliberations    No. 315, 89 p.
of salmon intotheopen waters of the state of Washington
during 1987. Prog. Rep. No. 267, Wash. Dept. Fish.,   Bax, N. J. 1983. Early marine mortality of marked
Olympia, WA, 447 p.                                   juvenile chum salmon (Oncorhynchus keta) released
                                                      into Hood Canal, Puget Sound, Washington, in 1980.
Alaska Department of Fish and Game. 1985. Alaska    Can. J. Fish. Aquat. Sci. 40:426-435.
habitat management guide. Southcentral Region, Vol.
I: Life histories and habitat requirements of fish and    Bax, N. J., E. O. Salo, B. P. Snyder, C. A. Simenstad,
wildlife. Alaska Dept. Fish Game, Juneau, AK, 429 p.   and W. J. Kinney. 1980. Salmon outmigration studies
                                                      in Hood Canal: a summary-1977. InW. J. McNeil and
Allen, G. H., and W. Aron. 1958. Food of salmonid    D. C. Himsworth (editors), Salmonid ecosystemsofthe
fishes of the western North Pacific Ocean. U.S. Fish    North Pacific, p. 171-201 Oregon State Univ. Press,
Wildl. Serv., Spec. Sci. Rep. Fish. 237,11 p.         Corvallis, OR.

Ames, J. 1983. Salmon stock interactions in Puget    Beacham, T. D., and C. B. Murray. 1987. Adaptive
Sound: a preliminary look.  In M. A. Miller (editor),   variation in body size, age, morphology, egg size, and
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Andrievskaya, L. D., 1957. Pitanie Tikhookeanskikh    of chum salmon, Oncorhynchus keta, from the Fraser
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Pacific salmon juveniles in the Sea of Okhotsk). [In   The effects of stream alterations on salmon and trout
Russ.] From: Izvestiya Tikhookeanskogo Nauchno-    habitat in Oregon. Oregon Dept. Fish Wildl., Portland,
Issledovatel'skogo.  Instituta rybnogo Knozyaistva i   OR, 70 p.
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Scientific Research Institute of Marine Fisheries and    Burner, C. J. 1951. Characteristics of spawning nests
Oceanography) 78:105-115 (Fish. Res. Board Can.,   of Columbia Riversalmon. Fish. Bull., U.S. 61(52):97-
Transl. Ser. No. 2441).                               110.

Aro, K. V., and M. P. Shepard. 1967. Pacific salmon in   Eschmeyer, W. N., W. S. Herald, and H. Hammann.
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23: 225-327.                                          America. Houghton Mifflin Co., Boston, MA, 336 p.

Atkinson, C. E., J. H. Rose, and T. O. Duncan. 1967.    Feller, R.J., and V. W. Kaczynski. 1975. Sizeselective
Pacific salmon in the United States. Internat. North    predation by juvenile chum salmon (Oncorhynchus
Pac. Fish. Comm., Bull. No. 23:43-223.                keta) on epibenthic prey in Puget Sound. J. Fish. Res.
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17(3):295-322.                                        Himsworth (editors), Salmonid ecosystems of the North
                                                       Pacific, p. 121-131. Oregon State Univ. Press, Corvallis,
Bakkala, R. G. 1970. Synopsis of biological data on    OR.

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Forrester, C. R.  1981.  Statistical yearbook 1978.    (editor), Estuarinecomparisons, p. 315-341. Academic
Internat. North Pac. Fish. Comm., Vancouver, B.C.,    Press, New York, NY.
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                                                      Holtby, L. B., T. E. McMahon, and J. C. Scrivener.
Fredin, R. A., R. L. Major, R. G. Bakkala, and G. K.    1989. Streamtemperaturesandinter-annualvariability
Tanonaka. 1977. Pacific salmon and the high seas    intheemigrationtimingofcohosalmon(Oncorhynchus
salmon fisheries of Japan. Proc. Rep., Northwest and    kisutch) smolts and fry and chum salmon (0. keta) fry
Alaska Fish. Cent., Natl. Mar. Fish. Serv., NOAA,    from Carnation Creek, British Columbia. Can. J. Fish.
Seattle, WA, 324 p.                                   Aquat. Sci. 46:1396-1405.

Fresh, K. L. 1984.  Evaluation of potential species    Holtby, L. B.,and J. C.Scrivener. 1989. Observed and
interaction effects in the planning and selection of   simulated effectsof climaticvariability, clear-cut logging,
salmonid enhancement projects. Report prepared by    and fishing on the numbers of chum salmon
thespeciesinteractionworkgroupoftheenhancement    (Oncorhynchus keta) and (0. kisutch) returning to
planning team. NOAA, Nat. Mar. Fish. Serv., Seattle,   Carnation Creek, British Columbia. Can. Spec. Publ.
WA, 80 p.                                             Fish. Aquat. Sci. 105:62-81.

Fresh, K. L., R. D. Cardwell, and R. R. Koons. 1981.    Hunter, J.G. 1959. Survivalandproductionofpinkand
Food habitsof Pacific salmon, baitfish, andtheirpotential    chum salmon in a coastal stream. J. Fish. Res. Board
competitors and predators in marine waters of   Can. 16(6):835-886.
Washington, August 1978 to September 1979. Prog.
Rep. No. 145, Wash. Dept. Fish., Olympia, WA, 58 p.   Iwata, M., and S. Komatsu.  1984.  Importance of
                                                      estuarine residence for adaptation of chum salmon
Fresh, K. L., and S. L. Schroeder. 1987. Influence of   (Oncorhynchus keta) fry to seawater. Can. J. Fish.
the abundance, size, and yolk reserves of juvenile   Aquat. Sci. 41:747-749.
chum salmon (Oncorhynchus keta) on predation by
freshwater fishes in a small coastal stream. Can. J.   Iwata, M., H. Ogura, S. Komatsu, and K. Suzuki. 1986.
Fish. Aquat. Sci. 44(22):236-243.                     Loss of seawater preference in chum salmon
                                                      (Oncorhynchus keta) fry retained in fresh water after
Hale, S. S., T. E. McMahon, and P. C. Nelson. 1985.    migration season. J. Exp. Zool. 240:369-376.
Habitat suitability index models and instream flow
suitability curves: chum salmon. U.S. Fish Wildl. Serv.    Kaeriyama, M. 1986. Ecological study of the early life
Biol. Rep. 82(10.108), 48 p.                          of the chum salmon, Oncorhynchus keta (Walbaum).
                                                      Sci. Rep. Hokkaido Salmon Hatchery 40:31-92.
Hallock, R. J., and D. H. Fry, Jr. 1967. Five species of
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California. Calif. Fish Game 53:5-22.                 Japan. Physiol. Ecol. Japan 1:625-638.

Hartt, A. C., and M. B. Dell.  1986.  Early oceanic    LeBrasseur, R.J. 1966. Stomach contents of salmon
migrations and growth of juvenile Pacific salmon and    and steelhead trout in the northeastern Pacific Ocean.
steelhead trout.  Internat. North Pac. Fish. Comm.    J. Fish. Res. Board Can. 23(1):85-100.
Bull., No. 46:1-105
                                                      LeBrasseur, R. J., and L. W. Barner. 1964. Midwater
Healey, M. C. 1979.  Detritus and juvenile salmon    trawl salmon catches in northern Hecata Strait,
production in the Nanaimo estuary: 1. Production and    November 1963. Fish. Res. Board Can., MS Rep. Ser.
feeding rates of juvenile chum salmon (Oncorhynchus    No. 176, 11 p.
keta). J. Fish. Res. Board Can. 36(5):488-496.
                                                      Levy, D. A., and T. G. Northcote. 1982. Juvenile
Healey, M. C. 1980. The ecology of juvenile salmon in   salmon residency in a marsh area of the Fraser River
Georgia Strait, British Columbia. In W. J. McNeil and    estuary. Can. J. Fish. Aquat. Sci. 39:270-276.
D. C. Himsworth (editors), Salmonid ecosystems of the
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
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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.
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                                                       Pearcy, W., T. Nishiyama, T. Fujii, and K. Masuda.
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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.
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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.
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                                                       embryos and alevins of Pacific salmon. Can. J. Fish.
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                                                       knowledge pertaining to the estuarine habitat
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                                                                                   Chum salmon continued
Shiino, S. 1976. List of common names of fishes of the
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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 ï¿½3 ï¿½  0           Highly abundant  Habitat
           SkagitBay ï¿½  ï¿½          @   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 ï¿½ 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
                           ï¿½ tivJ - Juveniles
          YaquinaBay ï¿½    *        L-Larvae             Horton 1982), with deep, marine-influenced habitats
          Alsea River             E-Eggs               often preferred (Macdonald et al. 1987). Smolts are
         Siuslaw River  ] ï¿½                             epipelagic in offshore marine waters (Milleret al. 1983).
        UmpquaRiver    ï¿½                               Subadults range from neritic to oceanic (Hartt and Dell
            Coos Bay (                                  1986). Adults are estuarine and riverine.
          Rogue River ï¿½  ï¿½
         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 4.4-21.0ï¿½C (Bell 1984), but 4.4-
        Tijuana Estuary                                 13.3ï¿½C is best for egg incubation (Reiser and Bjornn
                           A S J L E                    1979). Juveniles prefer stream temperatures of 11.8-
                                                        14.6ï¿½C, with 25.1ï¿½C the upper lethal limit (Brett 1952).
                                                        Growth ceases above 20.3ï¿½C 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 4.0-15.2ï¿½C. (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 4-140ï¿½C
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 11ï¿½C, 48 days at 90C, and 86-101 days at 4.5ï¿½C    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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                                                       summer 1980. Fish. Res. (Amsterdam) 2:1-17.
Kessler, D. W. 1985. Alaska's saltwater fishes and
other sea life. Alaska Northw. Publ. Co., Anchorage,    Morrow, J. E. 1980. The freshwater fishes of Alaska.
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Lander, R. H., and K. A. Henry.  1973.  Survival,    Moyle, P. B. 1976. Inland fishes of California. Univ.
maturity, abundance and marine distribution of 1965-    Calif. Press, Berkeley, CA, 405 p.
66 brood coho salmon, Oncorhynchus kisutch, from
the Columbia River hatcheries. Fish. Bull., U.S. 71:679-    Muir, W. D., and R. L. Emmett. 1988. Food habits of
695.                                                    migration salmonid smolts passing Bonneville Dam in
                                                       the Columbia River, 1984. Reg. Riv. Res. Man. 2:1-10.
Laufle, J. C., G. B. Pauley, and M. F. Shepard. 1986.
Species profiles: life histories and environmental    Myers, K. W., and H. F. Horton. 1982. Temporal use
requirementsofcoastalfishesandinvertebrates(Pacific   of an Oregon estuary by hatchery and wild juvenile
Northwest) -coho salmon. U.S. Fish Wildl. Serv. Biol.    salmon.   In V.S. Kennedy  (editor), Estuarine
Rep. 82(11.48), U.S. Army Corps Eng., TR EL-82-4,    comparisons, p. 377-392. Academic Press, New York,
18 p.                                                   NY.

Levy, D. A., and C. D. Levings. 1978. A description of   National Marine Fisheries Service. 1986. Fisheries of
the fish community of the Squamish River estuary,    the United States, 1985. Current Fishery Statistics No.
British Columbia: relative abundance, seasonal    8368. U.S. Dept. Comm., NOAA, Nat. Mar. Fish Serv.,
changes, and feeding habits of salmonids. Fish. Env.    Nat. Fish. Stat. Prog., Washington, D.C., 122 p.
Canada, Fish. Mar. Serv. Manusc. Rep. No. 1475,
63 p.                                                   Netboy, A. 1980. The Columbia. River salmon and
                                                       steelhead trout, their fight for survival. Univ. Wash.
Lichatowich, J. A., and J. D. Mcintyre. 1987. Use of   Press, Seattle, WA, 180 p.
hatcheries in the management of Pacific anadromous
salmonids. Am. Fish. Soc. Sympos. 1:131-136.            Nickelson, T. E. 1986. Influences of upwelling, ocean
                                                       temperature, and smolt abundance on marine survival
Macdonald, J. S., I. K. Birtwell, and G. M. Kruzynski.    of coho salmon (Oncorhynchus kisutch) in the Oregon
1987. Food and habitat utilization by juvenile salmonids    Production Area. Can. J. Fish. Aquat. Sci. 43:527-535.
in the Campbell River estuary. Can J. Fish. Aquat. Sci.
44:1233-1246.                                           Nickelson, T. E., M. F. Solazzi, and S. L. Johnson.
                                                       1986. Use of hatchery coho salmon (Oncorhynchus
Mahnken, C., E. Prentice, W. Waknitz, G. Monan, C.    kisutch) presmoltsto rebuild wild populations in Oregon
Sims, and J. Williams. 1982. The application of recent    coastal streams. Can. J. Fish. Aquat. Sci. 43:2443-
smoltification research to public hatchery releases: an    2449.
assessment of size/time requirements for Columbia
River hatchery coho salmon (Oncorhynchus kisutch).   Pacific Marine Fisheries Commission.  1986.  38th
Aquaculture 28:251-268.                                 annual report of the Pacific Marine  Fisheries
                                                       Commission fo rthe year 1985. Pac. Mar. Fish. Comm.,
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affecting Puget Sound coho salmon (Oncorhynchus
kisutch) runs. Can. J. Fish. Aquat. Sci. 37:1373-1378.

                                                   142






                                                                                  Coho salmon continued
Parmenter, T., and R. Bailey.  1985. The Oregon    North America, p. 1-54. U.S. Forest Serv. Gen. Tech.
oceanbook - an introduction to the Pacific Ocean off   Rep. PNW-96, Pac. Northw. Forest Range Exp. Sta.,
Oregon including its physical setting and living marine    Portland, OR.
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                                                       Columbia River salmon and steelhead during 1985.
Pearce, T. A., J. H. Meyer, and R. S. Boomer. 1982.    Unpubl. manuscr., NOAA, NMFS, Env.Tech. Ser. Div.,
Distribution and food habits of juvenile salmon in the    Portland, OR.
Nisqually estuary, Washington, 1979-1980. U.S. Dept.
Int., Fish. Assis. Off., U.S. Fish. Wildl. Serv., Olympia,    Ricker, W. E. 1976. Review of the rate of growth and
WA, 77 p.                                             mortality of Pacific salmon in salt water, and noncatch
                                                       mortality caused by fishing. J. Fish. Res. Board Can.
Pearcy, W. G. 1984. Where do all the coho go? The    33:1483-1524.
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influence of ocean conditions on the production of   E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
salmonids in the North Pacific, a workshop, p. 50-60.   of common and scientific names of fishes from the
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Peterson, W. T., R. D. Brodeur, and W. A. Pearcy.    Roos,J. F. 1960. Predation of young coho salmon on
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                                                       Rounsefell, G. A. 1957. Fecundity of North American
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beds.  In 14th Annual Report of the Pacific Marine    natural reproduction of silver salmon, Oncorhynchus
Fisheries Commission, p. 60-73.  Pac. Mar. Fish.   kisutch, a Minter Creek, Washington.  Wash. Dept.
Comm., Portland, OR.                                  Fish., Res. Bull. No. 4, 75 p.

Prakash, A. 1962. Seasonal changes in feeding of   Schmidt, A. H., C. C. Grahm, and J. E. McDonald.
coho and chinook (spring) salmon in southern British   1979. Summary of literature on fourfactors associated
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Purdy, J. E. 1989. The effects of brief exposure to   Scott, W. B., and E. J. Crossman. 1973. Freshwater
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                                                       Scrivener, J. C., and M. J. Brownlee. 1989. Effects of
Quinn, T. P. 1982. A model for salmon navigation on    forest harvesting on spawning gravel and incubation
the high seas. In E. L. Brannon and E. O. Salo (editors),   survival of chum (Oncorhynchus keta) and coho salmon
Proceedings of the salmon and trout migratory behavior    (0. kisutch) in Carnation Creek, British Columbia. Can.
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                                                       Shapovalov, L., and A. C. Taft. 1954. The life histories
Reiser, D. W., and T. C. Bjornn.  1979.  Habitat   of the steelhead rainbow trout (Salmo gairdneri
requirements of anadromous salmonids.  In W. R.   gairdnen) and silver salmon (Oncorhynchus kisutch)
Meehan (editor), Influence of forest and rangeland    with special reference to Waddel Creek, California,
management on anadromous fish habitat in western

                                                   143






Coho salmon continued
and recommendations regarding their management.    305. Dept. Fish. Oceans, Pacific Biol. Sta., Nanaimo,
Cal. Fish Game, Fish Bull. No. 98, 375 p.             B.C., Canada.

Shiino, S. M. 1976. List of common names of fishes of   Tschaplinski, P. J., and G. F. Hartman. 1982. Winter
the world, those prevailing among English-speaking    distribution of juvenile coho salmon (Oncorhynchus
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Kashikojima, Japan, 262 p.                            overwintersurvival. InG. Hartman (editor), Proceedings
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Effects of chronic turbidity on density and growth of   Nanaimo, B.C., Canada.
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                                                      Washington, P. M. 1982. An analysis of factors
Simenstad, C. A., and D. M. Eggers. 1981. Juvenile    affectingtheproductionofcohosalmon(Oncorhynchus
salmonid and baitfish distribution, abundance, and    kisutch) in the Columbia River. Ph.D. Thesis, Univ.
prey resources in selected areas of Grays Harbor,    Wash., Seattle, WA, 227 p.
Washington. Fish. Res. Inst., Univ. Wash., Seattle,
WA, 205 p. (FRI-UW-8816).                             Wright, S. G.  1968.  The origin and migration of
                                                      Washington's chinook and coho salmon. Info. Bookl.
Simenstad, C. A., K. L. Fresh, and E. O. Salo. 1982.    No. 1., Wash. Dept. Fish., Res. Div., Olympia, WA,
The role of Puget Sound and Washington coastal    25 p.
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Estuarine comparisons, p. 343-364. Academic Press,   of Washington, Univ. Wash. Press, Seattle, WA, 220 p.
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                                                      Zaugg, W. S., and L. R. McLain. 1976. Influence of
Steine, R. A., P. A. Reimers, and J. D. Hall. 1972.    watertemperatureongillsodium,potassium-stimulated
Social interaction between juvenile coho (Oncorhynchus    ATPase activity in juvenile coho salmon (Oncorhynchus
kisutch) and fall chinook salmon (0. tshawytscha) in   kisutch). Comp. Biochem. Physiol. 54A:419-421.
Sixes River, Oregon. J. Fish. Res. Board Can. 29:1737-
1748.

Takehama, S. (compiler). 1983. Statistical yearbook
1980. Internat. North Pac. Fish. Comm., Vancouver,
B.C., Canada, 115 p.

Thomas, D. H. 1985. A possible link between coho
(silver) salmon enhancement and a decline in central
California Dungeness crab abundance. Fish. Bull.,
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Thompson, K. 1972. Determining stream flows forfish
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Tschaplinski, P. J. 1982. Aspects of the population
biology of estuary-reared and stream-reared juvenile
coho salmon in Carnation Creek: a summary of current
research. In G. Hartman (editor), Proceedings of the
Carnation Creek workshop, a 10 year review, p. 289-

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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
              Pugetï¿½Sound C                 j 0  0    0                             Relative abundance:
                                                                                  ï¿½   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 0-28ï¿½C, 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 8.0-15.5ï¿½C (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 4-14ï¿½C; 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                             24.4ï¿½C (Brett 1952), but growth ceases at temperatures
       San Pedro Bay                                   above 20.30C (Bell 1984). Ocean-dwelling juveniles
         SAlamitos Bay                                  reside in temperatures of 1.0-13.0ï¿½C (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 riseto4-7ï¿½C (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    174ï¿½W) 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
acanthias), salmon shark (Lamna ditropis), other
salmonids, harbor seal (Phoca vitulina), beluga whale    Alaska Department of Fish and Game. 1985. Alaska
(Delphinapterus leucas), killer whale (Orcus orcinus),   habitat management guide, southcentral region, Vol.
and Dall's porpoise (Phocoenoides dalli) (Simenstad    1: Life histories and habitat requirements of fish and
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prey on adults during the spawning migration (Foerster
1968).                                                Allen, G. H., and W. Aron. 1958. Food of salmonid
                                                       fishes of the western North Pacific Ocean. U.S. Fish.
Factors Influencina PoDulations: Primary factors   Wildl., Spec. Sci. Rep. Fish. No. 237, 11 p.
influencing populations appear to be (1) overfishing,
(2) reduced production in freshwater environments,    Anas, R. E., andJ. R. Gauley. 1956. Blueback salmon,
and (3) reduced production in marine environments    Oncorhynchus nerka, age and length at seaward
(Foerster 1968, Peterman 1980). Overfishing reduces    migration past Bonneville Dam. U.S. Fish Wildl. Serv.,
freshwaterescapement andthus limits egg production    Spec. Sci. Rep. Fish. No. 185, 46 p.
(Foerster 1968). Mortality in fresh water during early
life stages is usually high. Foerster (1968) reported    Andrievskaya, L. D. 1957. Pitanie tikhookeanskikh
that egg to smolt survival ranged from 0.40-8.52%.    lososei v severo-zapadnoi chasti tikhovo okeana (The
This mortality is a result of poor waterquality (high and    food of Pacific salmon in the northwestern Pacific
low temperatures, turbidity, sedimentation, velocities,    Ocean). [In Russ.] From: Materialy po biologii morskovo

                                                   155






Sockeye salmon continued
perioda zhizni dalnevostochnykh lososei, p. 64-75.    Foerster, R. E.  1968.  The sockeye salmon,
Publ. by: Vses Nauchno-lssled. Inst. Morsk. Rybn.    Oncorhynchus nerka. Fish. Res. Board Can., Bull No.
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Bohn, B., and D. Mclsaac. 1986. Status report,
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antipredation window. Am. Nat. 131(2):271-290.          planning team). NOAA, Nat. Mar. Fish. Serv., Seattle,
                                                      WA, 80 p.
Doble, B. D., and D. M. Eggers. 1978. Diel feeding
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Thermal tolerance of juvenile Pacific salmon and    Hartt, A. C., and M. B. Dell.  1986.  Early oceanic
steelheadtroutinrelationtosupersaturationofnitrogen    migrations and growth of juvenile Pacific salmon and
gas. Fish. Bull., U.S. 69(4):833-843.                   steelhead trout.  Internat. North Pac. Fish. Comm.,
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OR.                                                     estuaries: the life support system. In V. S. Kennedy
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350.                                                    sockeye salmon (Oncorhynchus nerka) to fertilization

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                                                       Gulf of Alaska. Fish. Bull., U.S. 82(2):391-399.
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Gen. Tech. Rep. PNW-96, Pac. Northw. Forest Range    salmonid rearing facilities and a summary of their
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Salmonid ecosystems of the North Pacific, p. 247-265.
Oregon State Univ. Press, Corvallis, OR.

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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).

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                                                                                   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                                             _    ___         ï¿½   Highly abundant
         Skagit Bay              (3  :3                        fi 1                            (I)  Abundant
                                                                                          O   Common
       Grays Harbor              13              m3                                            o
                                                                                          - Rare
        Willapa Bayï¿½ _____   _____ - - -                               __   ___          _ -Blank Not present
      Columbia River     a            6         0  aï¿½          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   aï¿½  :                                                     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                                            <    ï¿½
       Tomales Bay
  Cent. San Fran. Bay *    C)    O   aï¿½  ï¿½                                    O    ï¿½ *        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 20.0ï¿½C    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 5.0-14.4ï¿½C (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 20ï¿½C (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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Washington D.C., 335 p.

Squire, J. L., Jr., and S. E. Smith. 1977. Anglers' guide
to the United States Pacific coast. Nati. Mar. Fish.
Serv., NOAA, Seattle, WA, 139 p.

Stevens, D. E., and L. W. Miller. 1983. Effects of river
flow on abundance of young chinook salmon, American
shad, longfin smelt, and delta smelt in the Sacramento-
San Joaquin River system. N. Am. J. Fish. Manag.
3:425-437.

Thompson, K. 1972. Determining stream flows for fish
life. In Proceedings, Instream flow requirements
workshop, p. 31-50. Pac. Northw. River Basin Comm.,
Vancouver, WA.

Van Hyning, J. M. 1973. Factors affecting the
abundance of fall chinook salmon in the Columbia
River. Oregon Fish Comm. Res. Rep. 4(1):1-87.

Vreeland, R. R. 1988. Evaluation of the contribution of
chinook salmon reared at Columbia River hatcheries to
the Pacific salmon fisheries. Ann. Rep. FY 1987, Nat.
Mar Fish. Serv., Env. Tech. Serv. Div., Portland, OR,
113 p.

Wagner, H. H., F. P. Conte, and J. L. Fessler. 1969.
Development of osmotic and ionic regulation in two
races of chinook salmon (Oncorhynchus tshawytscha).
Comp. Biochem. Physiol. 29:325-341.

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.



                                                   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    ï¿½   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 (   ï¿½ 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 ï¿½  a   O                          gametes. This process occurs in 2.5-5.0 cm of water
        Tillamook Bay ï¿½    ï¿½ O    Life stage:         and takes about 5 to 10 seconds (Loosanoff 1937).
          NetartsBay "4    ï¿½ 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    ï¿½ 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 ï¿½    ï¿½ O                         do not die after spawning (Loosanoff 1937).
           CoosBay i    ï¿½ 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               ï¿½   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  (*  ï¿½ ï¿½ . (   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 4.4-7.2ï¿½C (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 4-10ï¿½C. 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 4.4-7.2ï¿½C (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    ï¿½   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                          foreggdevelopmentappearstobenear12.8ï¿½C(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         27ï¿½C 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 31.7ï¿½C 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   ï¿½  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 ï¿½ ï¿½ ï¿½ O ï¿½                         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 @ @3ï¿½ 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  ï¿½ 6 0                     May and June (Wang 1986). Spawning takes place at
         Mission Bay a1  * 0 3                        temperatures of 10-25ï¿½C 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 13ï¿½C to
southern San Francisco Bay (Wang 1986).  While    <9 days at 27ï¿½C (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 *ï¿½ ï¿½   a a  Growth and Development
   Cent. San Fran. Bay ï¿½ ï¿½  ï¿½ ï¿½ ï¿½  IncludesCentralSan  Eaa Size and Embryonic DeveloDment: Unfertilized
                                Francisco, Suisun,
   South San Fran. Bay ï¿½ ï¿½ ï¿½ ï¿½ ï¿½  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. 35ï¿½N
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   ï¿½0 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 { ï¿½ O (j    O   Common
         Grllaparbayor  13 3'   0 0 0   Rare           concentrations as low as 0.25-0.50 mg/I (Wootton
         ColumbiaRiver I     * ï¿½ ï¿½   Blank Notpresent  1976). Maximumtemperature before mortality is 26ï¿½C
        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 15.8-18.5ï¿½C (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 0ï¿½C
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 ï¿½ 8 (3 ï¿½Francisco, 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 12-24ï¿½C (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 35ï¿½C, but become stressed at
Eggs are non-adhesive, slightly heavier than fresh    temperatures above 25ï¿½C (Moyle 1976). They tolerate
water, and are swept along with currents (Albrecht    temperatures of 0-32ï¿½C, 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
spawningtemperaturesare15.6-20.0ï¿½C,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 22.2ï¿½C    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    19ï¿½C) (Doroshev 1970). Hatching times range from 48
is some straying, each Pacific coast river system    hours (at 17.8-19.4ï¿½C) to 70-74 hours (at 14.4-15.6ï¿½C)
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 24ï¿½C to 9 days at 120ï¿½C (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 240ï¿½C 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


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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.























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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
24ï¿½C (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 19ï¿½C, 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. Bayï¿½    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                    19ï¿½C
       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 19ï¿½C), 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 13-30ï¿½C (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 13-18ï¿½/
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 0ï¿½/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 -17ï¿½C), and may account for this species'
         Morro Bay                                     wide depth and latitudinal distributions (Love et al.
    Santa Monica Bay   ï¿½  ï¿½                            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 8.0-19.0ï¿½C, 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 20ï¿½C (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 ï¿½0       9 ï¿½   Highly abundant  Miarations and Movements: The shiner perch forms
           Skagit Bay *ï¿½ i ï¿½   3   Abundant           loose schools that move seasonally- onshore and into
         Grays Harbor (3 %    ï¿½    O    Common        shallow water marine areas, estuaries, and bays in the
          Willapa Bay  3 3 ï¿½    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 ï¿½*                       1973, Wydoski and Whitney 1979). No coastal (north-
         Tillamook Bay  O O ï¿½    Life stage:          south) migrations are known to occur.  During the
          NetartnsBay * ï¿½ ï¿½    A-Adults               prespawning period, adults stay in shallow waters
           SiletzRiver ï¿½   ï¿½ ï¿½      JaJuveniles       during daylight and move to deeper waters at night.
          Yaquina Bay ï¿½ ï¿½ ï¿½                           Afterthis period, most adults reversethis movement by
           Alsea River ï¿½ ï¿½     ï¿½                      schooling in deeper water during the day and moving
         Siuslaw River ï¿½   *            ï¿½             to shallow water at night (Gordon 1965 as cited by
         Umpqua River ï¿½    ï¿½ *                        Wiebe 1968). Adults and juveniles appearto school in
            Coos Bay * *                              separate areas (Shaw et al. 1974). The shiner perch
          Rogue River ï¿½ ï¿½ ï¿½                           may use intertidal eelgrass beds significantly more at
         Klamath River (_3 1 3                        night than day (Bayer 1981).
         Humboldt Bay ï¿½ ï¿½
            Eel River ï¿½ ï¿½                             Reproduction
          Tomales Bay 13 Ci S                         Mode: This species is gonochoristic and iteroparous. It
    Cent. San Fran. Bay' d i ï¿½   * 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         ï¿½                        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  ï¿½                              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 21ï¿½C (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 18.5ï¿½C (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.

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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   ï¿½a ï¿½ 6 0ï¿½   Relative abundance:  1987). In Alaska, 1- and 2-year-old sand lance appear
         Hood Canal 0 ï¿½ * ï¿½ ï¿½    ï¿½   Highly abundant   to move inshore in early summer and then offshore
          SkagitBay O *ï¿½  ï¿½   @  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    0ï¿½      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 6.2ï¿½C, 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.





































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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 >10ï¿½C (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   ï¿½   Relative abundance:  search for cover, thus causing gobies in an entire area
          Hood Canal 0 0 0 C 0    ï¿½   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    ï¿½   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 5-15ï¿½C    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 25ï¿½C, 37.5%o at
to the Gulf of Alaska (Wydoski and Whitney 1979).       290C, and 0.0%o at 10ï¿½C (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    ï¿½   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 ï¿½ 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 15ï¿½50C.
         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 :ï¿½ O ï¿½:O C    L-Larvae              3.9-4.8 mm  total length (TL) (Jones  1962).
         AlseaRiver ï¿½ O  D O       O     Eggs          Metamorphosis to juvenile begins after about 2months,
       Siuslaw River * O ï¿½  O O                        when larvae are 15-20 mm standard length (Matarese
       Umpqua River Q O ï¿½      C0  et al. 1989).
          Coos Bay ï¿½        0 ï¿½  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 ï¿½    * IncludesCentral San    matures in 1 year and usually >12.0 cm TL.  This
                                 Francisco, Suisun,
  South San Fran. Bay  ( 3  ï¿½   Frand San Pablo bays,  species can live as long as 3 years and grow to 20.3 cm
      Elkhorn Slough O     ï¿½e                          in length in California (Jones 1962), and upto 10 years
          Morro Bay CO    ï¿½ 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.

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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               ï¿½   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 10-25ï¿½C, with
Gulf of California (Ginsburg 1952, Miller and Lea 1972,    a preference for 20.8ï¿½C (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 16ï¿½C.                                  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 16ï¿½C (Gadomski and    Parasites include isopods, copepods, nematodes,
Petersen 1988), or 20-29 days at 18.3-21.9ï¿½C (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 ï¿½                        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 17ï¿½C (Gadomski and Petersen 1988). The larval life
     Santa Monica Bay OO 0    o 0O                    stage lasts at least 5-6 weeks at 16ï¿½C (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 ï¿½ ï¿½ 6ï¿½   Relative abundance:  Optimumconditionsforlarvalsurvivalwerefoundtobe
          Hood Canal   ::    ï¿½    ï¿½a 0  S   Highly abundant    salinities of 25-28%0 and temperatures of 8-9ï¿½C
          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 7.8ï¿½C (Jackson
          Willapa Bay    I     0   i  Rare            1981). Temperatures >18ï¿½C 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   ï¿½ O       Life stage:         lethal limit forthis species is 26.1 ï¿½C (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     ï¿½ 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 12ï¿½C, 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

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                                                                                     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.
















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   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   21.5ï¿½C.  Temperatures >28.00C are lethal (Stober
        Grays Harbor O   ï¿½0O     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 ï¿½    O                         mature they tend to reside in estuaries (Morrow 1980).
           Coos Bay O    ( 0
         Rogue River   ï¿½ 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   ï¿½ 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 ï¿½C (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 <5ï¿½C) (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).
influenced by egg and larvae survival (Norcross and
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.

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                                                                                   Starry flounder continued

Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res.    Norcross, B. L., and R. F. Shaw. 1984. Oceanic and
Board Can., Bull. No. 180, 740 p.                       estuarine transport of fish eggs and larvae: a review.
                                                       Trans. Am. Fish. Soc. 113:153-165.
Jeffries, S. J., S. D. Treacy, and A. C. Geiger. 1984.
Marine mammals of the Columbia River estuary. Col.    Ocean Assessments Division.  1984.  The national
Riv. Estuary Data Dev. Prog., CREST, Astoria, OR, 62    status and trends program for marine environmental
p. plus appendices.                                     quality: Program description (mimeo). NOAA, NOS,
                                                       Ocean Assessments Division, Rockville, MD, 28 p.
Jewett, S. C., and H. M. Feder. 1980. Autumn food of
adult starry flounders, Platichthys stellatus, from the    Okada, Y. 1955. Fishes of Japan. Maruzen Co., Ltd.,
northeastern Bering Sea andthe southeastern Chukchi    Tokyo, Japan, 434 p.
Sea. J. Cons. Int. Explor. Mer. 39(1):7-14.
                                                       Orcutt, H. G. 1950. The life history of the starry
Lukas, J.,and C. Carter. 1987. 1985 pounds and value    flounder Platichthys stellatus  (Pallas).  Calif. Fish
of commercially caught fish and shellfish landed in   Game, Fish. Bull. 78:1-64.
Oregon. Oregon Dept. Fish Wildl., Portland, OR, 79 p.
                                                       Pedersen, M., and G. DiDonato. 1982. Groundfish
McCabe, G. T., Jr., W. D. Muir, R. L. Emmett, and J. T.    management plan for Washington's inside waters.
Durkin.  1983.  Interrelationships between juvenile    Prog. Rep. No. 170, Wash. Dept. Fish., Olympia, WA,
salmonids and nonsalmonid fish in the Columbia River    123 p.
estuary. Fish. Bull., U. S. 81(4):815-826.
                                                       Policansky, D. 1982a. The asymmetry of flounders.
Miller, B. S. 1965. Food and feeding studies on adults    Sci. Am. 246(5):116-122.
of two species of pleuronectids (Platichthys stellatus
and Psettichthys melanostictus) in East Sound, Orcas    Policansky, D.  1982b.  Influence of age, size, and
Island (Washington). M.S. thesis, Univ. Wash., Seattle,    temperature on metamorphosis in the starry flounder,
WA, 131 p.                                              Platichthys stellatus.  Can. J. Fish. Aquat. Sci.
                                                       39(3):514-517.
Misitano, D. A. 1977. Species composition and
relative abundance of larval and post-larval fishes in   Richardson, S. L., and W. G. Pearcy. 1977. Coastal
the Columbia River estuary, 1973. Fish. Bull., U.S.    and oceanic fish larvae in an area of upwelling off
75:218-222.                                             Yaquina Bay, Oregon. Fish. Bull., U.S. 75:125-146.

Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M.    Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
Nelson. 1990. Distribution and abundance of fishes    E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
and invertebrates in west coast estuaries, Volume I:  of common and scientific names of fishes from the
data summaries. ELMR Rep. No.4. Strategic Assess-    United States and Canada. Am. Fish. Soc. Spec. Publ.
ment Branch, NOS/NOAA, Rockville, MD,                   No. 12, Am. Fish. Soc., Bethesda, MD, 174 p.
240 p.
                                                       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.
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                                                       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.

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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


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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 0ï¿½C winter isotherm. Along the Pacific coast, this corresponds to 60ï¿½
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.




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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 3ï¿½C 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.


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                                                                                       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.


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                                                                                       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.


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                                                                                      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 20ï¿½C 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  ï¿½           a     00    0 0                                      0 
  edulis           J   O *   O                       00    00                      0 
                   L a* *            i 1S   0 0   00                              0    1i
                   E  I             ï¿½    tWJ   0 0   00                           0    ( O
 Pacific oyster     A     0        0                      a         a  S  
  Crassostrea      S
  gigas            J      *          U    0 ï¿½             * 
                   L
                   E
 Horseneck gaper   A      0  1       1  1      0             0          0                           0  0
  Tresus           S         0          0          0         0                                      0
  capax            J   O0            ï¿½         0 C           0          0                           0
                   L  00   ï¿½1  00                           0    0                                 0ï¿½
                   E    0                          0       0          0                           0 
 Pacific gaper      A     O  ï¿½       0         0  0
  Tresus           S         ï¿½    0
  nuttallii        J      Oï¿½ O                 0 0
                   L  ï¿½                       0I 00
                   E   ï¿½ï¿½   01                    0
 California jackknife A
 clam               S
  Tagelus          J
  californianus     L
                   E
 Pacific littleneck  A   le -                  a    00    00                        a                0o
 clam               S ï¿½   ï¿½ï¿½  ï¿½  0 0  00    a                                                        00
  Protothaca ï¿½            ï¿½          ï¿½ï¿½ ï¿½a                00   00                             ï¿½   0
  staminea          L a                                      0    00 a                               C
                   L I* -              I I -                  O0  O0                  ï¿½             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   '  ï¿½J   / 
        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     ï¿½


          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                                ï¿½
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:lmï¿½ 







                                     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.
  ï¿½Rocky outcrop (bedrock)- Exposed solid rock.
  *Estuarine vegetation- Aquatic plants within an estuary.
  ï¿½Marine vegetation- Aquatic plants in marine waters.
  'None- No reported preference.

Domain- Specific habitat where life stages occur.
  Benthic-Littoral
  ï¿½Intertidal (0-3 m)- On the bottom from the high tide mark to depths of 3 m.
  ï¿½Subtidal (3-10 m)- On the bottom at depths of 3-10 m.

   Benthic-Sublittoral
  ï¿½Inner shelf (10-50 m)- On the bottom of the continental shelf at depths of 10-50 m.
  ï¿½Middle 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.
  ï¿½Detritivore- 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.
   ï¿½Ocean migrant- An organism which migrates in ocean waters beyond the continental shelf.
   -Freshwater resident- An organism which resides primarily in freshwater habitats.
   ï¿½Estuarine 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.
   ï¿½Recreational- 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.
   ï¿½Ovoviviparous- 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.













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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.