Distribution and abundance of fishes and invertebrates in West Coast estuaries

[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

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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

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

Project Team

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

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

ELMR Report Number 8

August 1991

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

Property of CSC Library

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

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

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

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

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

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

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

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

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

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

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

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

Marine Sci.
Beaufort,
NC Lab

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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.

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

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

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

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

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

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

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

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

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

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

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

BIOLOGICAL ATTRIBUTES Eoonomic
Feeding type I Spatialstrategy Longevity Viklue

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

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

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

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.

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

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).

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

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

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

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

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.,
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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.
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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),

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.

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

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

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

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

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

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).

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.
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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
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Tomales Bay, California. Calif. Fish Game 57(1):69- and M. L. Kent. 1987. Pathology and significance of
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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,
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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.

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.
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Menzel, R. W. 1974. Portuguese and Japanese environment. In Symposium on estuarinefisheries, p.
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31:453-456. Soc., Bethesda, MD.

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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
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Paul, J. D., and I. M. Davies. 1986. Effects of copper Olympia, WA, 64 p plus appendices
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356. Eastern Wash. State College, Cheney, WA.

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

horseneck gaper continued

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

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

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.

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

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

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

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

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

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.

Tagelus californianus
Adult

5cm

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

California jackknife clam continued

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

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

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.

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

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

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

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

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.

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.

Manila clam continued

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

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

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

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.

Mya arenaria
Adult

2 cm

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

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

Softshellcontinued

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

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

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.

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

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

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

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

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

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

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.

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).

Bay shrimp continued

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

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

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

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.
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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

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

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

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

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

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

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
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magister as determined by the index of relative populationfluctuationsoftheDungenesscrab, Cancer

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
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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.

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

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

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

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

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.

Acipenser medirostris
Adult

25 cm

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

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

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

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

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

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.

Acipenser transmontanus
Adult

25 cm

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

White sturgeon continued

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

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

White sturgeon continued

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

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

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

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

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

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

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

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

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

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

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

Alosa sapidissima

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

American shad continued

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

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

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

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

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

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

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

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.

Clupea pallasi
Adult

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

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

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

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

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

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

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

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.

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.

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

Walford, L.A. 1931. Handbookof common commercial

102

103

Anchoa delicatissima
Adult

2cm

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

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

104

Slough anchovycontinued

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

105

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

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

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.

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.

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

106

107

Engraulis mordax
Adult

5cm

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

108

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

109

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

110

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

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

Brewer, G. 1974. Thermal tolerance and sediment Hunter, J. R., and K. M. Coyne. 1982. The onset of
toxicity studies. In D. F. Soule and M. Oguri (editors), schooling in northern anchovy larvae, Engraulismordax.
Marine studies of San Pedro Bay, California, p. 21-43. Calif. Coop. Ocean. Fish. Invest. Rep. 23:246-251.
Allan Hancock Found., Off. Sea Grant Publ., Univ. S.
Calif., Los Angeles, CA. Hunter, J. R., and S. R. Goldberg. 1980. Spawning
incidence and batch fecundity in northern anchovy,
Brewer, G. D. 1978. Reproduction and spawning of Engraulis mordax. Fish. Bull., U.S. 77(3):641-652.
the northern anchovy, Engraulis mordaxin San Pedro
Bay, California. Calif. Fish Game 64(3):175-184. Hunter, J. R., and R. Leong. 1981. The spawning
energetics of female northern anchovy, Engraulis
Brodeur, R. D., D. M. Gadomski, W. G. Pearcy, G. P. mordax. Fish. Bull., U.S. 79(2):215-230.
Batchelder, and C. B. Miller. 1985. Abundance and
distributionofichthyoplankton intheupwellingzoneoff Hunter, J. R., and B. J. Macewicz. 1980. Sexual
Oregon during anomalous El Nihio conditions. Estuar. maturity, batch fecundity, spawning frequency, and
Coast. Shelf Sci. 21:365-378. temporal pattern of spawning forthe northern anchovy,
Engraulis mordax, during the 1979 spawning season.
Clark, F. N., and J. B. Phillips. 1952. The northern Calif. Coop. Ocean. Fish. Invest. Rep. 21: 139-149.
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.

112

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

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

113

Oncorhynchus clarki
Juvenile

5cm

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

114

Cutthroat trout continued

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

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

115

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

116

Cutthroat trout continued

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117

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22.

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

118

119

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

120

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

121

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

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

122

Pink salmon continued
Dolly Varden (Salvelinus malma), northern squawfish help maintain and rehabilitate pink salmon stocks and
(Ptychocheilus oregonensis), and various sculpins millions of pink salmon are released annually (Wahle
(Cottusspp.) (Hunter 1959, Scott and Crossman 1973). and Smith 1979). However, increased fishing pressure
Belted kingfisher (Megaceryle alcyon), mergansers, due to hatchery runs can destroy wild populations
other predatory birds and small mammals also eat fry (McNeil 1980).
(Scott and Crossman 1973). Mammals (e.g., bears)
and large avian predators (e.g., bald eagles, Haliaeetus References
leucocephalus) feed on adult pink salmon in fresh
water. Marine and estuarine fish predators include Alaska Department of Fish and Game. 1985. Alaska
lamprey (Lampetra spp.), spiny dogfish (Squalus habitat managementguide. Southcentral Region, Vol.
acanthias), coho salmon, chinook salmon, rainbow I: Life histories and habitat requirements of fish and
trout, cutthroat trout and Pacific staghorn sculpin wildlife. Alaska Dept. Fish Game, Juneau, AK, 429 p.
(Leptocottus armatus). Predatory birds such as
common murre (Uria aalge), common merganser Allen, G. H., and W. Aron. 1958. Food of salmonid
(Mergus merganser), bald eagle, and Caspian fishes of the western North Pacific Ocean. U.S. Fish
tern(Hydroprogne caspia), and mammals such as Wildl. Serv., Spec. Sci. Rep. Fish. 237,11 p.
harbor seal (Phoca vitulina), northern fur seal
(Callorhinus ursinus), killer whale (Orcinus orca), and Ames, J. 1983. Salmon stock interactions in Puget
sea lions also prey on the pink salmon (Fresh 1984). Sound: a preliminary look. In M. A. Miller (editor),
Smalljuvenilepinksalmonapparentlyaltertheirhabitat Southeast Alaska coho salmon research and
preferences depending on predation risk (Magnhagen management review and planning workshop, May 18-
1988). 19,1982, p. 84-95. Alaska Dept. Fish Game, Juneau,
AK.
Factors Influencina PoDulations: Chum (0. keta) and
pink salmon have similar feeding habits during their Andrievskaya, L. D. 1958. Pitanie tikhookeanskikh
early marine life; thus, competition may be occurring in lososei v severo-zapadnoi chasti tikhovo okeana (The
the shallow marine habitats (Ames 1983, Fresh 1984). food of Pacific salmon in the northwestern Pacific
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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
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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.
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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.
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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,
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salmon populations. Hatcheries have been built to

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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

126

127

Oncorhynchus keta
Adults

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

128

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

129

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

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

130

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

131

Chum salmon continued
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135

Oncorhynchus kisutch
Adults

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

136

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

137

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

138

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

139

Coho salmon continued
with ocean "upwelling" one year earlier (Gonsolus Bilton, H. T., D. F. Alderdice, and J. T. Schnute. 1982.
1978). The Oregon Production Area coho salmon Influence of time and size at release of juvenile coho
population has gone from predominantly high-survival salmon (Oncorhynchus kisutch) on returns at maturity.
wild fish to predominantly low-survival hatchery fish Can. J. Fish. Aquat. Sci. 39:426-447.
(Nickelson 1986). Over 62 million hatchery smolts
werereleasedintheOregon Production Area (Monterey Brannon, E. L. 1982. Orientation mechanisms of
Bay, California to Leadbetter Point, Washington) in homing salmonids. In E. L. Brannon and E. O. Salo
1981, including 24 million from private hatcheries (editors), Proceedingsofthesalmonandtroutmigratory
(Nickelson 1986). Hatcheries (private and public) play behavior symposium, p. 219-227. School Fish., Univ.
a dominant role in the abundance of this species in the Wash, Seattle, WA.
Pacific Northwest. However, the introduction of hatchery
coho salmon presmolts into streams appears to reduce Brett. J. R. 1952. Temperature tolerance in young
wild coho salmon populations (Nickelson et al. 1986). Pacific salmon, genus Oncorhynchus. J. Fish. Res.
Hatcheries may also precipitate overharvest of wild Board Can. 9(6):265-323.
stocks and cause density-dependent mortality in both
freshwater and marine environments (Lichatowich and Brodeur, R. D. 1989. Neustonic feeding by juvenile
McIntyre 1987). Coho salmon smolts may need to salmonids in coastal waters of the northeast Pacific.
reach a "critical size" for proper smoltification and Can. J. Zool. 67:1995-2007.
marine survival. Hence, growth and time of release are
importantattributesforhatcheryfish(Biltonetal. 1982, Brodeur, R. D., H. V. Lorz, and W. G. Pearcy. 1987.
Mahnken et al. 1982). Thomas (1985) found a Food habits and diet variations of pelagic nekton off
correlation between coho salmon hatchery production Oregon and Washington, 1979-1984. NOAA Tech.
and a decline in central California Dungeness crab Rep. NMFS 57, 32 p.
(Cancer magister) abundance, probably related to
coho salmon feeding on crab megalopae. El Niio also Burner, C. J. 1951. Characteristics of spawning nests
affects coho salmon abundance (Hayes and Henry of Columbia River salmon. Fish. Bull., U.S. 61(52):97-
1985). Finally, Japanese high-seas fishing fleets take 110.
unknown numbers of coho salmon andthesquid gillnet
fisheries may also take coho salmon incidentally. Chapman, D. W. 1966. Food and space as regulators
of salmonid populations in streams. Am. Nat. 100:345-
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144

145

Oncorhynchus mykiss
Adults

10 cm

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

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

146

Steelheadcontinued

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

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

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

147

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

148

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

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

149

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

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

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

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

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

150

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

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

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

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

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

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

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

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

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

151

Oncorhynchus nerka
Adults

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

152

Sockeye salmon continued

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

153

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

154

Sockeye salmon continued
Eggers 1978). During their downstream migration, pollutants, etc.) which can be a result of poor forest
smolts may feed heavily on gammarid amphipods practices, industrial waste, mining and refining effluents,
(Muir and Emmett 1988). In estuaries, euphausiids, agriculture practices,and urban development. Physical
fish larvae, juvenile shrimp, insects, amphipods, and disturbance of the redd (by erosion, subsequent
mysids are eaten (Levy and Yesaki 1982, Simenstad et spawners, ice scour) and predation can also diminish
al. 1982). In the ocean, juvenile sockeye salmon feed freshwaterproduction (Foerster 1968, Hart 1973). River
on euphausiids, hyperiid amphipods, copepods, obstructions such as dams (manmade and natural,
decapod larvae, pteropods, juvenile and larval fish, such as Hell's Gate and the Fraser River rock slide of
squid, and other invertebrates. The primary prey 1913) can affect upstream and downstream migrations
consumed depends on the location, time of day, and (Foerster 1968). Columbia River sockeye salmon runs
fish's age (Andrievskaya 1957, Allen and Aron 1958, have diminished primarily as a result of dams and
Ito 1964, LeBrasseur 1966, Foerster 1968, Pearcy et irrigation diversions of spawning rivers (Mullan 1986).
al. 1984). In lakes and in the ocean, juvenile sockeye The abundance of food relative to parr numbers in
salmon appear to feed primarily at dusk or at night reservoirs and lakes also affects production; when
(Doble and Eggers 1978, Pearcy et al. 1984). Parr may sockeye parr densities are high, food may limit their
not feed during the winter in lakes (Doble and Eggers growth which in turn can reduce smolt size and marine
1978). Juveniles (ocean- and lake-dwelling) feed near survival (Foerster 1954,1968, Kyle et al. 1988). Nutrient
the surface, except in lakes when surface temperatures fertilization of lakes has been attempted to increase
are high (Foerster 1968). lake primary production and zooplankton standing
crop and thus juvenile sockeye salmon growth and
Biological Interactions survival (LeBrasseur et al. 1978, Hyatt and Stockner
Predation: Primary fish predators of fry and parr in 1985). Predators and competition can reduce
fresh water are coho salmon (0. kisutch), cutthroat populations in reservoirs (Foerster 1968). Ocean
trout (0. clark,), char (Salvelinus spp.), rainbow trout conditions may also reduce production as a result of
(0. mykiss), Dolly Varden (Salvelinus malma), lake density-dependent mortality (Peterman 1980). The
trout (Salvelinus namaycush), lake whitefish Japanese high seas fishery (located west of long.
(Coregonus clupeaformis), mountain whitefish 174�W) intercepts many North American sockeye
(Prosopium williamsonm), northern squawfish salmon (Fredin et al. 1977). This fishery took over 46
(Ptychocheilus oregonensis), burbot (Lota Iota), and million North American sockeye over a 20 year period.
sculpins (Foerster 1968, Fresh 1984). Gulls, common This catch, togetherwith the accidental mortalities and
loon (Gavia immer), red-necked grebe (Podiceps lost additional weight gain before North American
grisegena), common merganser (Mergus merganser), harvest, represents a substantial loss to U.S. fishermen
belted kingfisher (Megaceryle alcyon), terns, and large (Ricker 1976, Fredin et al. 1977). Hatchery releases of
predatory birds [osprey (Pandion haliaetus) and bald sockeye salmon are used to maintain this species'
eagle (Haliaeetus leucocephalus)] are important avian abundance in some areas (Wahle and Smith 1979).
predators (Fresh 1984). Marine predators include
lamprey (Lampetra spp.), spiny dogfish (Squalus References
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
et al. 1979, Fiscus 1980). Bears and other mammals wildlife. Alaska Dept. Fish Game, Juneau, AK, 429 p.
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
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155

Sockeye salmon continued
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Forrester, C. R. 1981. Statistical yearbook 1978.
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Doble, B. D., and D. M. Eggers. 1978. Diel feeding
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in the Columbia River, 1984. Reg. Riv. Res. Man. 2:1-
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Sci. 43(8):1643-1655. salmon caught in gill nets during a 24-hour period in the
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158

159

Oncorhynchus tshawytscha
Adults

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

160

Chinook salmon continued

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

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

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

161

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

162

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

163

Chinook salmon continued
Factors Influencina Ponulations: High mortality occurs Allen, M.J., R. J. Wolotira, Jr., T. M. Sample, S. F. Noel,
during the early freshwater life stages (eggs, fry, parr). and C. R. Iten. (in press). Salmonids: life history
This mortality is caused by redd destruction, siltation descriptions and brief harvest summaries for salmonid
and destruction of spawning grounds, extremely high species of the northeast Pacific Ocean and eastern
or low watertemperatures, low dissolved oxygen, loss Bering Sea. Tech. Memo., NOAA, NMFS, Northwest
of cover, disease, and predation (Reiser and Bjornn Alaska Fish. Cent., Seattle, WA.
1979). Besides the above factors, man-made changes
such as river flow reductions, the creation of dams and Ames, J. 1983. Salmon stock interactions in Puget
reservoirs, pollution, and logging practices, have Sound: a preliminary look. In M. A. Miller (editor),
affected population abundances (Raymond 1979, Southeast Alaska coho salmon research and
Netboy 1980, Stevens and Miller 1983). Estuaries management review and planning workshop, May 18-
appearto play a vital role in chinook salmon life history 19,1982, p. 84-95. Alaska Dept. Fish Game, Juneau,
(MacDonald et al. 1988). In the ocean, this species is AK, 109 p.
affected by disease, predation, food availability, and
oceanographicconditions. Overfishing has not allowed Beach, R. J., A. C. Geiger, S. J. Jeffries, and S. D.
optimal spawning escapement and has reduced the Tracy. 1981. Marine mammal- fishery interactions on
age and size structure ofsome populations (Fraidenburg the Columbia River and adjacent waters, 1981. Second
and Lincoln 1985). Also, the high-seas gill net fishery Annual Rep. to NOAA, NMFS, Northwest and Alaska
for squid is taking an unknown number of chinook Fish. Cent., Seattle, WA. Wash. Dept. Game, Olympia,
salmon. The release of millions of juvenile chinook WA, 186 p.
salmon by public and private hatcheries has helped
maintain some runs (Phinney 1986), and the United Beauchamp, D. A., M. F. Shepard, and G. B. Pauley.
States-Canada Salmon Interception Treatyshould allow 1983. Species profiles: life histories and environmental
more escapement in the future. The survival of hatchery requirements of coastal fishes and invertebrates (Pacific
smolts to maturity is influenced by time of release, size Northwest) - chinook salmon. U.S. Fish Wildl. Serv.,
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adults typically spawn in deeper water and use larger in central Columbia River. Fish. Bull., U.S. 71:387-400.
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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.
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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.

168

169

Hypomesus pretiosus
Adult

5cm

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

170

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

171

Surf smelt continued

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

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

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

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

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

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

172

173

Spirinchus thaleichthys
Adult

2cm

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

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

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

174

Longfin smelt continued

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

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

175

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

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

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

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

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

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

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

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

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

176

177

Thaleichthys pacificus
Adult

5cm

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

178

Eulachon continued

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

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

179

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

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

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

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

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

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

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

180

181

Microgadus proximus
Adult

5cm

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

182

Pacific tomcod continued

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

183

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

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

184

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

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

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

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

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

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

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

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

185

Atherinops affinis
Adult

5cm

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

186

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

187

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

188

Topsmelt continued

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

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

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

189

Atherinopsis californiensis
Adult

10cm

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

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

190

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

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

191

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

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

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

192

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

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

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

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

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.

193

Gasterosteus aculeatus
Adult

2cm

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

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

194

Threespine stickleback continued

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

195

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

196

Threespine stickleback continued

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

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

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

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

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

197

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

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

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

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

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

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

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

198

199

Morone saxatilis
Adult

10 cm

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

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

200

Striped bass continued

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

201

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

202

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

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

203

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

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

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

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

Fay, C. W., R. J. Neves, and G. B. Pardue. 1983. Miller,D.J.,andR. N. Lea. 1972. Guidetothecoastal
Species profiles: life histories and environmental marinefishesof California. Calif. Fish Game, Fish Bull.
requirements of coastal fishes and invertebrates (Mid- 157, 235 p.
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and invertebrates in west coast estuaries, Volume I:
Ganssle, D. 1966. Fishes and decapods of the San data summaries. ELMR Rep. No. 4. Strategic
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Ecological studies of the Sacramento-San Joaquin 240 p.
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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.,
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Radovich, J. 1963. Effectofoceantemperatureonthe environmentally impacted estuaries. Mar. Poll. Bull.
seaward movements of striped bass, Roccus saxatifis, 19(9):466-477.
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Chesapeake and San Francisco Bays: Two

205

Striped bass continued
Turner, J. L. and T. C. Farley. 1971. Effects of
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102.

206

207

Paralabrax clathratus
Adult

10cm

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

208

Kelp bass continued

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

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

209

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

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

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

210

Kelp bass continued

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

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

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

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

211

Paralabrax nebulifer
Adult

10cm

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

212

Barred sand bass continued

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

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

213

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

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

214

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

215

Atractoscion nobilis
Adult

25 cm

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

216

White seabass continued

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

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

217

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

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

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

218

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

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

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

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

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

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

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

219

Genyonemus lineatus
Adult

5cm

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

220

White croaker continued

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

221

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

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

222

White croaker continued

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

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

223

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

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

224

225

Cymatogaster aggregata
Adult

2cm

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

226

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

227

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

223

Shinerperch continued

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

Clemens, W. A., and G. V. Wilby. 1961. Fishes of the Horn, M. H. 1974. Fishes. InA summary of knowledge
Pacific coast of Canada. Fish. Res. Board Can., Bull. of the southern California coastal zone and offshore
No. 68, 443 p. areas, Chapter 11. South. Calif. Ocean Stud. Consort.,
Fullerton, CA, 124 p.
Earnest, R. D., and P. E. Benville, Jr. 1971. Correlation
of DDT and lipid levels for certain San Francisco Bay Jeffries, S. J., S. D. Treacy, and A. C. Geiger. 1984.
fish. Pest. Monitor. J. 5(3):235-241. Marine mammals of the Columbia River estuary.
Columbia River Estuary Data Dev. Prog., CREST,
Earnest, R. D., and P. E. Benville, Jr. 1972. Acute Astoria, OR, 62 p. plus appendices.
toxicity of four organochlorine insecticides totwo species
of surf perch. Calif. Fish Game 58(2):127-132. Morrow, J. E. 1980. The freshwater fishes of Alaska.
Alaska Northw. Publ. Co., Anchorage, AK, 248 p.
Eigenmann, C. L. 1892. Cymatogaster aggregata
Gibbons; a contribution to the ontogeny of viviparous Moyle, P. B. 1976. Inland fishes of California. Univ.
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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
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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.
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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.
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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,
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2479:1-10.

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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
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disposal in the oceans; minimizing impact, maximizing
benefits, p. 194-208. Westview Press, Boulder, CO.

Stober, Q. J. 1973. Summary and overview of
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proposed Kiket Island nuclear power site, p. 441 -448.
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guide to the early life histories. Tech. Rep. 9.

230

231

Ammodytes hexapterus
Adult

5cm

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

232

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

233

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

234

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

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

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

235

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

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

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

236

237

Clevelandia ios
Adult

2cm

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

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

238

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

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

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

239

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

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

240

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

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

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

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

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

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

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

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

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

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

241

Ophiodon elongatus

25 cm

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

242

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

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

243

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

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

244

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

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

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

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

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

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

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

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

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

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

245

Leptocottus armatus
Adult

5cm

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

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

246

Pacific staghorn sculpin continued

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

247

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

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

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

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

248

249

Paralichthys californicus
Adult

10cm

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

250

California halibut continued

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

251

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

252

California halibut continued
scale changes in the marine environment, overfishing, Barnett, A.M., A. E. Hahn, P. D. Sertic, and W. Watson.
alterations and destruction of estuarine habitat, or a 1984. Distribution of ichthyoplankton off San Onofre,
shift in population centers (Plummer et al. 1983). California, and methods for sampling shallow coastal
Pollution, (e.g., watersoluble fractions of crude oil) can waters. Fish. Bull., U.S. 82(1):97-111.
reduce hatching success, reduce size of larvae at
hatching, produce morphological and anatomical Barry, J. P., and G. M. Cailliet. 1981. The utilization of
abnormalities, and reduce feeding and growth rates shallow marsh habitats by commercially important
(MBC Applied EnvironmentalSciences 1987). Initiation fishes in Elkhorn Slough, California. Cal.-Nev. Wildl.
of feeding by larvae appears critical for larval survival Trans. 1981:38-47.
(Gadomski and Petersen 1988). Natural production
has recently been augmented by hatchery production Cailliet, G. M., B. Antrim, D. Ambrose, S. Pace, and M.
(Crooke and Taucher 1988). Substantial genetic Stevenson. 1977. Species composition, abundance
variation between two populations of California halibut and ecological studies of fishes, larval fishes, and
in the southern California Bight, suggests that the zooplankton in Elkhorn Slough. In Ecologic and
natural population is subdivided (Hedgecock and Bartley hydrographic studies of Elkhorn Slough, Moss Landing
1988). Wide fluctuations in young-of-the-year Harbor and nearshore coastal waters, p. 216-386.
recruitmentexist, but noexact causehasbeenidentified Moss Landing Marine Lab., Moss Landing, CA.
(Allen 1988). Southern California estuaries and
protected shallow water habitats play a critical role in California Department of Fish and Game. 1987. 1987
the life history of this species. California sport fishing regulations. Calif. Dept. Fish
Game, Sacramento, CA, 12 p.
References
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previous studies and the status of the sport fishery.
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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
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Tomales Bay area. Mariscos Publ., Hampton Bays,
NY, 143 p.

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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.
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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
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254

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

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

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

255

Hypsopsetta guttulata
Adult

5 cm

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

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

256

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

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

257

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

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

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

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

258

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

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

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

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

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

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

259

Pleuronectes vetulus
Juvenile

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

260

English sole continued

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

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

261

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

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

262

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

Becker, D. S. 1988. Relationships between sediment Hogue, E. W., and A. G. Carey, Jr. 1982. Feeding
character and sex segregation in English sole, ecology of 0-age flatfishes at a nursery ground on the
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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.
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Hulberg, L. W., and J. S. Oliver. 1979. Prey availability
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Data Devel. Prog., CREST, Astoria, OR, 113 p. plus OR, 40 p. (ORESU-T-81-001).
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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.
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Bull. 56:1-50. Ketchen, K. S. 1947. Studies on lemon sole
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263

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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.,
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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
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Olson, R. E., and I. Pratt. 1973. Parasites as indicators
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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
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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.
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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
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1982. Age and growth of a pleuronectid, Parophrys Can. J. Fish. Aquat. Sci. 44:48-53.
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E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list
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98. Thornburgh, and L. J. Bledsoe. 1979. Food web
relationships of northern Puget Sound and the Strait of

264

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Juan de Fuca. U.S. Interagency (NOAA, EPA) Energy/
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Sopher, T. R. 1974. A trawl survey of the fishes of
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Toole, C. L. 1980. Intertidal recruitment and feeding in
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Van Cleve, R., and S. Z. El-Sayed. 1969. Age, growth,
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the Pacific coast of the United States. Ph.D. Thesis,
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Westrheim, S. J. 1955. Size composition, growth, and
seasonal abundance of juvenile English sole (Parophrys
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Yoklavich, M. 1982. Growth, food consumption, and
conversion efficiency of juvenile English sole (Parophrys
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Grant, Univ. Wash., Seattle, WA.

265

Platichthys stellatus
Adult ,

10cm

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

266

Starry flounder continued

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

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

267

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

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

Factors Influencina Populations: Contaminants can Frey, H. W. 1971. California's living marine resources
impairreproductivesuccess (Whipple etal 1978, Spies and their utilization. Calif. Dept. Fish Game,
et al. 1985) and may cause fin erosion disease and Sacramento, CA, 148 p.
lethal skin tumors (Wellings et al. 1976, Campana
1983). Endoparasitic flukes and monogenetic Garrison, K. J., and B. S. Miller. 1982. Review of the
trematodes have been found on the gills (Bane and earlylifehistoryofPugetSoundfishes. Fish. Res. Inst.,
Bane 1971). Population sizes are probably greatly Univ. Wash., Seattle, WA, 729 p. (FRI-UW-8216).
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.

268

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
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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,
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Monaco, M. E., R. L. Emmett, S. A. Hinton, and D. M. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker,
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269

Starry flounder continued
Spies, R. B., D. W. Rice, Jr., P. A. Montagna, R. R. ecological impacts of oil spills, p. 757-806. Amer. Inst.
Ireland, J. S. Felton, S. K. Healy, and P. R. Lewis. Biol. Sci., Keystone, CO.
1985. Pollutant body burdens and reproduction in
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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
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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.

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

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

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

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

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

270

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272

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

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

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

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

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

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

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

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

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

ANTHROPOGENIC-Refers to the effects of human activities.

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

AREAL-Refers to a measure of area.

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

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

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

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

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

BIGHT-An inward bend or bow in the coastline.

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

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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 3�C winter isotherms. In neritic
waters of western North America, it extends from Point Conception, California, to the southern Bering Sea, Alaska.

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

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

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

CALCAREOUS-Composed of calcium or calcium carbonate.

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

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

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

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

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

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

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

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

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

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

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

CONGENER-Referring to members of the same genus.

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

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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.

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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).

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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.

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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.

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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

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Glossary continued

predator population and negative (decreasing) for the prey population. See PARASITISM, SYMBIOTIC,
CARNIVORE, and TROPHIC LEVEL.

PRODUCTION-Gross primary production is the amount of light energy converted to chemical energy in the form
of organic compounds by autotrophs like algae. The amount left after respiration is net primary production and
is usually expressed as biomass or calories/unit area/unit time. Net production for herbivores and carnivores is
based on the same concept, except that chemical energy from food, not light, is used and partially stored for life
processes. Efficiency of energy transfers between trophic levels ranges from 10-65% (depending on the organism
and trophic level). Organisms at high trophic levels have only a fraction of the energy available to them that was
stored in plant biomass. After respiration loss, net production goes into growth and reproduction, and some is
passed to the next trophic level. See FOOD WEB and TROPHIC LEVEL.

PROKARYOTIC-Organisms that have nuclear bodies, but lack chromosomes, nucleoli, and nuclear mem-
branes.

PROTANDRY-A type of hermaphroditism in which an individual initially develops as a male, then reverses to
function as a female. Common for some species of shrimp.

PROTISTAN-Pertaining to the eukaryotic unicellular organisms of the kingdom Protista, including such groups
as algae, fungi, and protozoans.

PROTOZOA-A varied group of either free-living or parasitic unicellular flagellate and amoeboid organisms.

PYCNOCLINE--A zone of marked water density gradient that is usually associated with depth.

RACE-An intraspecific group or subpopulation characterized by a distinctive combination of physiological,
biological, geographical, or ecological traits. In salmonids, a race is determined by when it returns to its natal
stream.

RADULA-A toothed belt or tongue in the buccal cavity of most molluscs that is used to scrape food particles from
a surface, or modified otherwise to serve a variety of feeding habits.

RANGE-(1) The geographic range is the entire area where a species is known to occur or to have occurred
(historical range). The range of a species may be continuous, or it may have unoccupied gaps between
populations (discontinuous distribution). (2) Some populations, orthe entire species, may have different seasonal
ranges. These may be overlapping, orthey may bewidely separated with intervening areas that are at most briefly
occupied during passage on relatively narrow migration routes. (3) Home range refers to the local area that an
individual or group uses for a long period or life. See DISTRIBUTION and TERRITORY.

RECRUITMENT-The addition of new members to a population or stock through successful reproduction and
immigration.

RED TIDE-A reddish coloration of sea waters caused by a large bloom of red flagellates. The accumulation of
metabolic by-products from these organisms is toxic to fish and many other marine species. The accumulation
of these metabolites in shellfish makes shellfish toxic to humans.

REDD-A gravel nest dug by spawning female salmon and trout. After eggs are released and fertilized by the
male, the female covers them with gravel by sweeping movements of the tail.

REPRODUCTIVE POTENTIAL-The total number of offspring possible for a female of a given species to produce
if she lives to the maximum reproductive age. This is found by multiplying the number of possible reproductive
periods by the average number of eggs or offspring produced by females of each age class. This potential is
seldom realized, but this and the age of first reproduction, or generation time, determine the maximum rate of
population increase under ideal conditions.

282

Glossary continued

RESIDUALISM-Occurswhen juvenile salmon smolts do not migrate to sea but revert backto parr, usually loosing
their ability to osmoregulate in seawater.

RHEOTAXIS-A response movement by an animal toward or away from stimulation by a water current.

RIVERINE-Pertaining to a river or formed by a river or stream.

ROE-The egg-laden ovary of fish, or the egg mass of certain crustaceans.

RUN-A group of migrating fish (e.g., a salmon run).

SALT WEDGE-A wedge-shaped layer of salt water that intrudes upstream beneath a low-density freshwater lens
that has "thinned" while flowing seaward.

SAN DIEGO PROVINCE-A zoogeographical designation for faunal distributions that, based on minimum
temperature requirements, extends from Point Conception, California, to Magdalena Bay, Baja California Sur.

SCAVENGER-Any animal that feeds on dead animals and remains of animals killed by predators. See
DECOMPOSER and DETRITIVORE.

SEAMOUNT-An undersea mountain rising more than 3000 feet (914 m) from the sea floor, but having a summit
at least 1000 feet (305 m) below sea level (in contrast to an island).

SEDENTARY-Refers to animals that are attached to a substrate or confined to a very restricted area (or those
that do not move or move very little). See SESSILE.

SEMELPAROUS-Animals that have a single reproductive period during their lifespan.

SESSILE-Refers to an organisms that is permanently attached to the substrate. See SEDENTARY.

SETTLEMENT-The act of or state of making a permanent residency. Often refers to the period when fish and
invertebrate larvae change from a planktonic to a benthic existence.

SHOAL-(1I) A sand bar in a body of water that is exposed at low tide. (2) An area of shallow water. (3) A group
of fish (school). (4) As a verb, to collect in a crowd or school.

SIPHONS-The "necks" or tubes of clams and other bivalves that carry water containing food and oxygen into
the gills, and then expels water containing waste products (exhalent siphon).

SLOUGH--Ashallow inlet orbackwaterwhose bottom may be exposed at lowtide. Sloughs often borderestuaries
and typically have a stream passing through them.

SMOLT-A juvenile salmon or anadromoustrout that is in the process of migrating to the ocean and physiologically
adapting to seawater. Smolts are usually very silvery and have very faint parr marks. See PARR.

SPAT-Juvenile bivalve molluscs which have settled from the water column to the substrate to begin a benthic
existence.

SPAWN-The release of eggs and sperm during mating. Also, the bearing of offspring by species with internal
fertilization. See PARTURITION.

SPECIES-(1) A fundamental taxonomic group ranking after a genus. (2) A group of organisms recognized as
distinct from other groups, whose members can interbreed and produce fertile offspring. See POPULATION,
SUBPOPULATION, and SUBSPECIES.

283

Glossary continued

SPERMATOPHORE-A capsule orgelatinous packet (extruded by a male) containing sperm and used to transfer
sperm to females. Spermatophores are produced by certain invertebrates and some primitive vertebrates.

SPIROCHETE-A spiral-shaped, non-flagellated bacterium of the order Spirochaetales. This group can be free-
living or parasitic. Some members cause diseases.

SPIT-A long, narrow sand bar or peninsula extending into a body of water which is at least partly connected to
the shore. See SHOAL.

SPOROCYST-A simple larval stage of parasitic trematode worms. Contact with the host causes a metamorpho-
sis from an earlier stage to this stage.

STENOHALINE-Pertaining to organisms that are restricted to a narrow range of salinities, in contrast to
EURYHALINE.

STIPE-A thickened, stalk-like structure in kelps that bears other structures, such as blades. Also, the basal
portion of the thallus or plant body of alga.

STOCK-A related group or subpopulation. See POPULATION and SUBPOPULATION.

SUBADULTS-Maturing individuals that are not yet sexually mature.

SUBLITTORAL-The benthic zone along a coast, or lake that extends from mean low tide to depths of about
200 m.

SUBPOPULATION-A breeding unit (deme) of a larger population. These units may differ little genetically and
taxonomically. See SUBSPECIES. Subpopulations may intergrade with some interbreeding, orthey may occupy
a common seasonal range prior to the mating season. The units may have different reproduction times and be
separated spatially or temporally. See RACE, STOCK, and POPULATION.

SUBSPECIES-A taxonomic class assigned to populations and/or subpopulations when interbreeding (gene
flow) between populations is limited, and there are significant differences in some combination of characteristics
between subspecies (e.g., appearance, anatomy, ecology, physiology, and behavior). While successful
interbreeding can occurwhen the groups are in contact, undernatural conditions reproductive isolation is complete
and the groups are considered distinct. Classification of such groups is based on the comparative study and
judgement of phylogenists. A second epithet for each subspecies is added to the binomial for the species (e.g.,
Oncorhynchus clarki clarki). See SPECIES, POPULATION, and SUBPOPULATION.

SUBTIDAL-See SUBLITTORAL.

SUPRALITTORAL-The splash zone of land (adjacent to the sea) that is above the mean high tide level.

SUSPENSION FEEDER-An animal that feeds directly or by filtration on minute organisms and organic debris
that is suspended in the water column.

SYMBIOSIS-The relationship between two interacting organisms that is positive, negative, or neutral in its
effects on each species. See COMPETITION, MUTUALISM, PARASITISM, and PREDATION.

TAXONOMY-A system of describing, naming, and classifying animals and plants into related groups based on
common features (e.g., structure, embryology, and biochemistry).

TEMPERATE REGION-Oceanic waters between the 13 and 200C winter isotherms. The temperate region of
the neritic zone on the Pacific coast of North America extends from Point Conception, California, to Magdalena
Bay, Baja California Sur.

284

Glossary continued

TEMPORAL-Pertaining to time. Used to describe organism activities, developmental stages, and distributions
as they relate to daily, seasonal, or geologic time periods.

TERRITORY-An area occupied and used by an individual, pair, or larger social group, and from which other
individuals orgroups of the species are excluded, often with the aid of auditory, olfactory, and visual signals, threat
displays, and outright combat.

TEST-A rigid calcareous exoskeleton produced by some echinoderms in the class Echinoidea (e.g., sea urchins
and sand dollars).

THERMOCLINE-A relatively narrow boundary layer of water where temperature decreases rapidly with depth.
Little water or solute exchange occurs across the thermocline, which is maintained by solar heating of the upper
water layers.

TREMATODA-A class of parasitic flatworms of the phylum Platyhelminthes. Trematodes have one or more
muscular, external suckers and are also known as flukes.

TRIPLOIDY-The occurrence of threetimes the haploid numberof chromosomes. When genetically engineered,
randomly occurring traits may be selected for commercial applications. For example, the Pacific oyster
experiences a degradation in flesh quality associated with spawning. Non-reproducing triploid cultures avoid this
seasonal problem.

TROCHOPHORE-A molluscan larval stage (except in Cephalopoda) following gastrulation (embryonic stage
characterized bythe development of a simple gut). It is commonly ciliated, biconically shaped, and free-swimming;
it establishes an evolutionary link between annelids and molluscs, since both groups display a similar life stage.

TROPHIC LEVEL-The feeding level in an ecosystem food chain characterized by organisms that occupy a
similar functional position. At the first level are autotrophs or producers (e.g., kelps and diatoms); at the second
level are herbivores (e.g., copepods and snails); at the third level and above are carnivores (e.g., salmon and
seals). Omnivores feed atthe second and third levels. Decomposers and detritivores may feed at all trophic levels.
See FOOD WEB and PRODUCTION.

TROPICAL REGION-Oceanic waters between the 20�C winter isotherms in the southern and northern
hemispheres. Tropical neritic waters along the west coasts of North and South America extend from the southern
tip of Baja California, Mexico, to about lat. 50S along the coast of Peru.

TURBELLARIA-A class of mostly aquatic, non-parasitic flatworms that are leaf-shaped and covered with cilia.

UPWELLING-The process whereby prevailing seasonal winds create surface currents that allow nutrient rich
cold water from the ocean depths to move into the euphotic or epipelagic zone. This process breaks down the
thermocline and increases primary productivity, and ultimately fish abundance.

VELICONCHA-A bivalve larval stage. A veliconcha has two larval shells and moves by using its velum.

VELIGER-A ciliated larval stage common in molluscs. This stage forms afterthe trochophore larva and has some
adult features, such as a shell and foot.

VELUM-The ciliated swimming organ of a larval mollusc.

VIVIPAROUS-Refers to animals that produce live offspring; eggs are retained and fertilized in the female (as
compared to OVIPAROUS).

WATER COLUMN-The water mass between the surface and the bottom.

285

Glossary continued

YEAR-CLASS-Refers to animals of a species population hatched or born in the same year at about the same
time; also known as a cohort. Strong year-classes result when there is high larval and juvenile survival; the reverse
is true forweak year-classes. The effects of strong and weak year-classes on population size and structure may
persist for years in species with long lives. Variation in year-class strength often affects fisheries. See
DISTRIBUTION and STOCK.

ZOEA-An early larval stage of various marine crabs and shrimp; zoea have many appendages and long dorsal
and anterior spines.

ZOOPLANKTON-Animal members of the plankton. Most range in size from microscopic to about 2.54 cm in
length. They reside primarily in the epipelagic zone and feed on phytoplankton and each other. Although they
have only a limited ability to swim against currents, many undertake diel migrations. Taxa include protozoa,
jellyfish, comb jellies, arrowworms, lowerchordates, copepods, water fleas, krill, and the larvae of many fish and
invertebrates that are not planktonic as adults.

286

Appendices

Appendix 1: Summary table example: Spatial distribution and relative abundance

Appendix 2: Summary table example: Temporal distribution

Appendix 3: Summary table example: Data reliability

Appendix 4: Presence/absence of 47 species in west coast estuaries

Appendix 5: Life history tables: Life history characteristics of 47 west coast species
Table 5Ak Biogeography
Table 5B. Habitat Associations
Table 50. Biological Attributes and Economic Value
Table 5D. Reproduction

Appendix 6: Definitions of terms used in life history tables

287

West Coast Estuaries

Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay
Species/Life Stage T M S T M S T M S T M S T M S T M S T M S T M S
Blue mussel A 0 a 00 00 0
Mytilis S � a 00 0 0 0
edulis J O * O 00 00 0
L a* * i 1S 0 0 00 0 1i
E I � tWJ 0 0 00 0 ( O
Pacific oyster A 0 0 a a S
Crassostrea S
gigas J * U 0 � *
L
E
Horseneck gaper A 0 1 1 1 0 0 0 0 0
Tresus S 0 0 0 0 0
capax J O0 � 0 C 0 0 0
L 00 �1 00 0 0 0�
E 0 0 0 0 0
Pacific gaper A O � 0 0 0
Tresus S � 0
nuttallii J O� O 0 0
L � 0I 00
E �� 01 0
California jackknife A
clam S
Tagelus J
californianus L
E
Pacific littleneck A le - a 00 00 a 0o
clam S � �� � 0 0 00 a 00
Protothaca � � �� �a 00 00 � 0
staminea L a 0 00 a C
L I* - I I - O0 O0 � OO
E U U a 00 00 a 0C
T M ST M ST M ST M ST M S T M S T M ST M S

Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay

West Coast Estuaries

Relative Abundance Salinity Zone Life Stage/Activity

* Highly Abundant T - Tidal Fresh A - Adults
i Abundant M - Mixing S - Spawning
O Common S - Seawater J -Juveniles
Blank Not Present, Rare, or L - Larvae
No Data Available E - Eggs

288

West Coast Estuaries
Puget Sound Hood Canal Skagit Bay
Month JFMAMJJASOND JFMAMJJASOND JFMAMJJASOND
Species/Life Stage
Blue mussel A l
Mytilis
edulis J ............ .'........................ji.. ..............

E i i

Pacific oyster A
Crassostrea S
gigas J
L
E
Horseneck gaper A .iii iiiiiiiii ....ii ....... ......"j.f ......
Tresus S : I :I.
~capax I I i: I I
E
Pacific gaper A !!ii iii i ii iiiii ii ii i.iiii iiiiiiii !!!!!!!iiiiiii iiii ii
Tresus S "
nuttallii J . iiiiiiiiiiiiiiiiiiiiiiiii iiiiiiii . .iii .iiiiiiiiiiiiiiiiiii. I
L 1 5_iiiiiiiiiii!i:i iiiii!iiiiml liiiiiiiiiiiiiii iiii i I I

California jackknife A
clam
Tagelus J
californianus L
E
Pacific littleneck A
clam S
Protothaca J
staminea L
E
JFMAMJJASOND JFMAMJJASOND JFMAMJJASOND
Puget Sound Hood Canal Skagit Bay
West Coast Estuaries

Relative Abundance Life Stage/Activity
Highly Abundant A - Adults
~iiiiiii~l Abundant S - Spawning
J - Juveniles
[I | Common L - Larvae
E - Eggs
Blank Not present, Rare, or
No Data Available

289

h.. * * * 0 3*T

West Coast Estuaries

Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay
Species/Life Stage
Blue mussel A 0 NEl
MONilS S *IliU El 171E
edulis i U1 n o UU
L U j li l lE

Pacific oyster A 0 0 0 U U U
Crassostrea S 0
gigas 'J U
L U U U U U 0
E 0 N 0
Horseneck gaper A 0 U 9 U N 0
Tresus S E
capax i' UUUU
L r- 1-U1lI]UElE
E
Pacific gaper A * UU
Tresus S l1E UUU
nuttallii 'J U
L El 13 El a U U

California jackknife A a 0 U U
clam S * U UUU
Tagelus J UUUU UUU
californianus L UUU UUUU

Pacific littleneck A *UUE
clam S I! F I! IN] 13 13 U 0 0
Prolothaca J U a U U El U U U
staminea L El 13 9~ 17- El U 13 U
E El El nol E El a El U

Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay

West Coast Estuaries

Reliability Life Stage/Activity

N Highly Certain A - Adults
FRI Moderately Certain ~S - Spawning
iii Moderately Certain j~~~ - Juveniles
El Reasonable Inference L - Larvae
E - Eggs

290

Note: Due to post-publication revisions of the presence/absence information in VolumeI
(Table 5, pp. 185-197), data in this appendix has been updated and supersedes that presented in volume I

Index to Appendix 4: Page location of presencelabsence table for each species and estuary

Estuary

Common and Scientific Name .9
Blue mussel (Mytilu eduli)
Pacific oyster (Crassosfrea gigas)
Horseneack gaper (Tresus capax)
Pacific gaper (Tresus nutralliO
California jackknife clam (Tagalus californianus)
Pacific littleneck clam (Protothaca starminea)
Manila clam (Venerupisjaponica)
Softshell (AMya aranaria)
Geoduck (Panopea abrupra)
Bay shrimp (Crangon franis corum) 292 293 294 295
Dungeness crab (Cancer magjste,)
Leopard shark (Thakis semifasciata)
Green sturgeon (Acipenser madirosrins)
White sturgeon (Acipenser transmonf anus)
AmerIcan shad (Alasa sapidissima)
Pacific herring (Clupea pallast)
Deepbody anchovy (Anchaa compressa)
Slough anchovy (Anchoa delicatissima)
Northern anchovy (Engraufis mordax)
Cunthroat trout (Oncortryncthus clarki)
Pink salmon (Oncorhynchus gorbuscha)
Chum salmon (Oncorhynchus kaea)
Coho salmon (Oncorhynchus kisutch)
Steelhead (Oncorhynchus mykiss)
Sockeye salmon (Qncortrynchus nerka) 296 297 298 299
Chinook salmon (Oncortrynchus Ishawytscha)
Surf smell (l-lypomesus prefiosus)
Longtin smelt (Spirinchus thaleichthys)
Eulachon (Thaleichthys pacificus)
Pacific tomrcod (Microgadus proximus)
Topsmelt (Athhannops at finis)
Jacksmell (Atherinopsis califomniensis)
Threespine stickleback (Gasterosteus aculeaf us)
Striped bass (Momone saxatlfis)
Kelp bass (Paralabrax clathratus)
Barred sand bass (Paralabrax nebuliler)
White seabass (At tact oscion nobffis)
White croaker (Genyonemus fineatus)
Shiner parch (Cyrmalogastrer ggregala) (303233
Pacific sand lance (Amnmodytes haxaplerus)30313233
Arrow goby (Clevelandia ios)
LUngcod (Ophiodon elongatus)
Pacific staghorn sculpin (Leptocorus arrnatus)
California halibut (Paralichthys califomricus)
Diamond turbot (Hypsopserta gutfulata)
English sole (Pleuronectes vefulus)
Starry flounder (Platidrihys stellatus)

291

Appendix 4 continued

Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay
Species T M S T M S T M S T M S T M S T M S T M S T M S
Blue mussel A 44 : 1 V 4 / i I 4 I
Mytilus J '14 I '14 44 1 ' 1 4 3 4
edulis L 1 4' 4 4 74 4 4 '4 3 4
Pacific oyster A 4 4 4 4 4 4 / 4 4 /
Crassostrea J 4 ' 1 4 4 '4 ' ' I
gigas L
Horseneck gaper A 4 4 J J '4 4 4J /1 1 V
Tresus J 4 4 : 44 4 4 '] '4 4 4 J
ca:ax LJ J 4
Pacific gaper A / 4 '4 4 '/ 4 '4 '4
Tresus J '4 4 3 4 3 '
nuttallii L 4 4 3 4 4 4
California jackknife clam A
Tagelus J
californianus L
Pacific littleneck clam A 44 4 '4 '4 44 4 4 4/
Protothaca J ' 4 41 3J 3 ' 4 J 3 3 I
staminea L 4 3 3 3 3 J 3 4 4 4
Manila clam A 44 1 / / I 4 44 4 44J 4
Venerupis J 4J 3 34 3 i J ' 44 3 l
iaponica L 44 44 J V 44J 4 4 3
Softshell A '4 44 4 '4 V 4/ ' 4 4
Mya J ' 3 4 3 ' 3
arenaria L 4j 4 '3 4 4 I V 4 4 '4 J
Geoduck A 4 / :
Panopea J 4J ' 4 4 4
abrupta L 4/ '4 4 4
Bay shrimp A '4 4 4 V 4 V 4 44
Crangon J * V 4 4 V 4 4 4 4 ' 4 4 4 4
franciscorum L ' / J 44 I J 4 4 4 44 4 4 4 4
Dungeness crab A 4 V 44 -4 -, V J 4 4J 44 '4 4 4
Cancer J '1 44 ' 44 4 I J 4 4 4 4 i J
magister L ' 4 J 3 J ' 4 4 ' V 4
Leopard shark A
Triakis J
semifasciata P
Green sturgeon A 4 4 4 4 4/ 4J
Acipenser J J 3
medirostris L
White sturgeon A 4 4 4 4 x/ J . . t/ q ' �J /
Acipenser J 4
transmontanus L
American shad A 4 V4 - 'J / V V ", " , V 44 '4 44 4 4 44
Alosa J 4J 4 J -
sapidissima L
Pacific herring A V 44 44 4 44 4
Clupea J 4 4 V V '4 V 4 4 V 4
pallasi L 44 3 44 4 4 '4 '
Deepbody anchovy A
Anchoa J
compressa L
Slough anchovy A
Anchoa J
delicatissima L
Northern anchovy A " "J 44 4 4 '4 4J ' 44 4 4
Engraulis J 3 I ' I 3 3 4 4 3
mordax L 4 4 44 1 4 44 4
T MS T MS T MS TM ST MS TM ST MS T MS
Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay
Legend:
T = Tidal fresh zone A = Adults 3 = Species / lifestage is present
M = Mixing zone J = Juveniles Blank = Species / lifestage is not present
S = Seawater zone L = Larvae
P = Parturition

292

Appendix 4 continued

Netarts Siletz Yaquina Alsea Siuslaw Umpqua Coos Rogue
Bay River Bay River River River Bay River
Species T M S T M S T M S T M S T M S T M S T M S T M S
Blue mussel A -4 '4 '4 '4 '4 -' '
Mytilus J '4 '4 '4 ' '4 '4 '4 '
edulis L 4
Pacific oyster A '4 '4 '4
Crassostrea J '4 ' '4 '4 '4
.0igas L
Horseneck gaper A ' ' '4 ' ' 4
Tresus J '4 ' ' ' '4'4
capax L ' ' ' ' '4 '4'4
Pacific gaper A '
Tresus J '
nuttalliN L '
California jackknife clam A
Tagelus J
californianus L
Pacific littleneckclam A '4 ' ' '4 ' ''
Protothaca J '4 ' '4 ' '
staminea L
Manila clam A ' ' '
Venerupis J ' ' '4
japonica L ' '4
Softshell A '4 '4 ' '4 ' ' '4'4
Mya J '' 4'' 44 ' 4''
arenaria L '4 '4 '4 '4 ' ' '4'4
Geoduck A '
Panopea J
abrupta L '4
Bayshrimp A '4 '4 - 4 ' 4' '4 '4'4 ' 4'
Crangon J '4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4
franciscorum L '4'4 '4'4 4' '4 ' ' '4'4
Dungeness crab A '4 ' ' ' ' '4 ' '4'4
Cancer j '4 ' ' '4 '4 '4 '4 '4'4
maoister L '4 ' '4 ' '
Leopard shark A
Ttiakis J
semifasciata P
Green sturgeon A ' '4 '4 '4 '4'4'4'4'4
Acipenser J ' '4 '4 '4 '4'4'4'4'4
medirostris L
White sturgeon A ' 4 ' ' '4'4'4'4'4
Acipenser J ' ' ' 4 '4'4'4'4'4
transmontanus L
American shad A '4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4'4
Alosa J '4'4'4'4'4'4'4'4'4'4'4'J'4'4'4'4'4'4
sapidissima L '
Pacific herring A '4 '4 ' '4 '4 '4 '4
Clupea J ''4 '' '44 '' '4 '' '44 44
pall1asi L ''4'' '44 '''4 44
Deepbody anchovy A
Anchoa i
compressa L
Slough anchovy A
Anchoa J
delicatissima L
Northern anchovy A ' ' ' ' '4 '4 '4 '4,4
Engraulis J ' 4 '4 '44 '4 '4'4
mordax L '4 4 q 4 4
T MS TM ST MS T MS T MS T MS T MS TM S
Netarts Siletz Yaquina Alsea Siuslaw Umpqua Coos Rogue
Bay River Bay River River River Bay River
Legend: A Adults
T = Tidal fresh zone J Juveniles '4= Species I lifestage is present
M = Mixing zone L Larvae Blank = Species / lifestage is not present
S = Seawater zone
P =2Parturition

293

Appendix 4 continued

Klamath Humboldt Eel Tamales CentralSan South San Elkhorn Morro
River Bay River Bay Francisou Bayt Francisco Bay Slough Bay
Species TM ST M STMST M ST M S*M S S S*
Blue mussel A'1. 4' 4. .44.44
Mytilus J 4 4 4
edulis L444' 44444
Pacific oyster A . 4. 4' 4.
Crassostrea J4444444

Horseneck gaper A'44
Tresus i
capoax L44
Pacific gaper A4 4..44'44
Tresus j 444
nuttalIl L 4
California jackknife clam A'1..44
Tagelus 4444
californianus L4444
Pacific littleneck clam A .4 '4 -4 .4 . 4
Protothaca J 4
staminea L 4
Manila clam A.4..4 ' 4 4. 4
Venerypis J444 44 444
ia,~onica L444 44 44
Softshell A 4. 4. 4. .44.44
Mya J 4 4 4
arenaria L 4 4 4
Geoduck A .
Panopea J44
abrupta L44
Bay shrimp A -4 . 4. 4. 4
Crangon J44 44 44 4
franciscorum L44 444 44 44
Dungeness crab A 44.4 44 4. 4
Cancer j 4 4 4 4 4
mapister L444 44 444
Leopard shark A 4. 4 . 4. 4.
Triakis j 44 44 4
semnifasciata P 444 4444
Green sturgeon A...44 44444 44
Acipenser i4 4 4 4 444 4
medirostris L4
White sturgeon A.44.44 '44
Acipenser J44444444 44
transmontanus L4
American shad A . 4. 4. 4. /.
Alosa j4444444 4
sapidissima L
Pacific herring A.4 ..4 . 44. 4.44
Clupea J 4 4 44 4 4
vallasi L44 44 44 44
Deepbody anchovy A
Anchoa J
compressa L
Slough anchovy A
Anchoa J
delicatissima L
Northern anchtovy A 4 4 4 4 4 44.
Engraulis J 4 4 44 4 4
mordlax L 4' 4 4
T MST M ST MS T MS T MS M MS S *5**
Klamath Humboldt Eel Tomales Coentral San South San Elkhorn Morro
River By River Bay Francisco Bayl Franciso Bay Slown a

Legend: 1Includes San Pablo and Suisun Says.
T =Tidal fresh zone A = Adults 4=Species I lifestage is present
M =Mixing zone J = Juveniles Blank = Species I lifestage is not present
S =Seawater zone L = Larvae
=Salinity zone is not present P = Parturition

294

Appendix 4 continued

Santa San Pedro Alamitos Anaheim Newport Mission San Diego Tijuana
Monica Bay Bay Bay Bay Bay Bay Bay Estuary
Species * * S * * S * * S * * S * * S * * S * * S * * S
Blue mussel A : : /1 J '4 ' : 4
Mytilus J / : :: :4 4 4 '
edulis L 4 4 4J 4 4 4 '
Pacific oyster A
Crassostrea J
gigas L
Horseneck gaper A
Tresus J
capax L
Pacific gaper A '4 V ' '4 '4 i '4 '/
Tresus J 4 4 4 'J 4 4 4 J
nuttailli L 4 J 41 4 4
California jackknife clam A V : :: . ' V '4 '/
Tagelus J 4 4 : 4 :: 4 :
californianus L 4 J 4 :l 4 lJ lJ
Pacific littleneck clam A 4 / V ', '4 ' q ' V
Protothaca J 4 4 1 4 44
staminea L 4 4 J 4' 4 4 4
Manila clam A
Venerupis J
japonica L
Softshell A
Mya J
arenaria L
Geoduck A '4'
Panopea J: :
abrupta L
Bay shrimp A
Crangon J
franciscorum L
Dungeness crab A
Cancer J
magister L
Leopard shark A ' ' V '4 V 'J
Triakis J 4 4 4 '
semifasciata P 4 l
Green sturgeon A
Acipenser J
medirostris L
White sturgeon A
Acipenser J
transmontanus L
American shad A
Alosa J
sapidissima L
Pacific herring A 4/
Clupea J 4
pallasi L
Deepbody anchovy A '4 ' '4 ' '4 '4 ',
Anchoa J '
compressa L 4
Slough anchovy A '4 '4 ' '4 '4 '
Anchoa J 4 4 J 4 4
delicatissima L l 4
Northern anchovy A ' ', 4 ' ' '
Engraulis J 4 '4 4 4 4 'J
mordax L 1 J 4 4 4 4 44
* S * * S * * S * * S * * S * * S * * S
Santa San Pedro Alamitos Anaheim Newport Mission San Diego Tijuana
Monica Bay Bay Bay Bay Bay Bay Bay Estuary
Legend:
T = Tidal fresh zone A = Adults 4 = Species / lifestage is present
M = Mixing zone J = Juveniles Blank = Species / lifestage is not present
S = Seawater zone L = Larvae
= Salinity zone is not present P = Parturition

295

Appendix 4 continued

Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay
Species T MST M ST M STM ST M STM ST M STM S
Cutthroat trout A 44 4 44 4 4 4 4 4 4 4
Oncorhynchus i -444 4 4 4 4 44 4 4 4
clarki L
Pink salmon A 4 4 4 4 4 4 4
Oncorhynchus J44444 4444
gorbuscha L4
Chum salmon A 4 -4 4 -4 4 -4 4 4 V 4 4 -4 4i 4 -4 4 4 4 4 4 -4 4 q -4 -4
Oncorhynchus, J 4 4 4 4 4 44 4 4 4 4 4
keta L
Coho salmon A 4444444444444 444444444 44
Oncorhynchus J i
kisutch L
Steelhead - fall A
Oncorhynchus J
mykiss (F) L
Steelhead - half pounder A
Oncorhynchus j
mykiss (H) L
Steelhead -summer A44 4444 4 4 -4 4
Oncorhynchus J4 4 4 4 4 4 4
mykiss (B) L
Steelhead -winter A 444 4444 44444 44 444 444 4 444
Oncorhynchus J i
mykiss (W) L
Sockeye salmon A 4 444 4 44 44
Oncorhynchus J i
nerka L
Chinook salmon -fall A 44 444444 4444 444 44 444 444 4
OncorhynchusJ44 44 4 4 44 4 ' 44 ''
tshawytscha (F) L
Chinook salmon - late fall A
Oncorhynchus J
tshawytscha (LF) L
Chinook salmon - winter A
Oncorhynchus J
tshawvtscha (W) L
Chinook salmon -spring A 4 4 4444 444444444444
Oncorhynchus, j4 4 4 4 4 444 4
Ishawyfscha (Sp) L
Chinook salmon - summer A44
Oncorhynchus i 4
Ishawytscha (Su) L
Surf smelt A 44 4 44 4 44 4 44 4
Hypomesus i 4 4 4 4 4 4 4 4
pretiosus , L 44 4 4 44 4 44 44
Longfin smelt A 4 4 4 4 4
Spirinchus , i
thaleichthys L 4 4 4 4 4 4
Eulachon A 44 4 4 4 4
Thaleichthys J
pacificus L4444 4444 4
Pacific tomood A 44 4444 444 444
Microgadus J 4 4 4 4 4 4 4 '
proximus L 44 4 44 4 44 4 44 4
Topsmelt A 4444 4
Atherinops J 44 4
affinis L
T M ST MS TMS T MS T MS T MS T MS TM S
Puget Hood Skagit Grays Willapa Columbia Nehalem Tillamook
Sound Canal Bay Harbor Bay River Bay Bay
Legend:
T =Tidal fresh zone A =Adults 4=Species / lifestage is present
M Mixing zone J =Juveniles Blank = Species / lifestage is not present
S Seawater zone L =Larvae

296

Appendix 4 continued

Netarts Siletz Yaquina.- Alsea Siuslaw Umpqua coos Rogue
Say River Say River River River Bay River
Species T M STM ST M STM ST MS TM ST MS T MS
Cutthroat trout A '4 '4~ '4 '4 '4 '4 '4 '4 '4 '4 '4 '-4 '4 '~ '4 '~ '4 '4 '4 '4 '4 '4 '4 '4
Oncorhynchus J '4 4 '4444444444444 '4 44 '4444
clarki L
Pink salmon A4'4''4''44'44
Oncorhynchus J
gorbuscha L
Chum salmon A'4'44''444''' 4444'
Oncorhynchus J''i 4444 - - j ' 4'4 -4'q4' 4' 4 44
keta L
Coho salmon A '4 '4'4' 4 '4 '4 '4 '4 '4 '4 4 4 '4 '4 '4 '4 '4 '4 4 '4 '4 '4'
Oncorhynchus J 4' 4' 4' 4' 4' 4' 4' 4' 44' 4'
kisutch L
Steelhead - fall A
Oncorhyrnchus J
mykisqs (F) L
Steelhead - half pounder A
Oncorhynchus J 4''
mykiss (H) L
Steelhead -summer A ''' 44 444'''
Oncorhynchus i 44'4 444'4
mykiss (8) L
Steelhead -winter A'4''4''4''44''4''4'4'4''4''44'44'44
Oncorhynchus J 4' 44' 44' 4'4' 4' 4' 4' 4' 4'
mykiss (14/ L
Sockeye salmon A
Oncorhynchus J
nerka L
Chinook salmon -fall A ' 4' 4' 4' 4' 44' 4' 4' 4' 4' 4'
Oncorhynchus J 4' 4' 4' 4' 4' 4'4' 4' 4' 4' 4' 4'
tshawytscha (F) L
Chinook salmon - late fall A
Oncorhynchus J
tshawytscha (LF) L
Chinook salmon - winter A
Oncorhynchus J
tshawytscha (14' L
Chinook salmon -spring A q V -V -4 -q q 4 44' 4' '4 4'
Oncorhynchus i 4 4 4 4 4'''''''''
tshawytscha (SP) L
Chinook salmon - summer A
Oncorhynchus J
tshanwvtscha (Su) L
Surf smelt A '44 '44 '4 '4 4 4' 44'
Hyporresus J '4'4 '4 '4 4 '4 44 44 ''4 44
pretiosus L 4 4' 4'4' 4'
Longfin smelt A ' 444' 4' 4'
Spirinchus i 44' 4' 4'
thaleichthys L ' 4
Eulachon A '''4''''
Thaleichthys J
pacilicus L4 4
Pacific tomood A '4 '4 '4 4 4 44 4
Microgadus J 4 4 44 '4 '' 44 '' 4
proximus L'4''4 ''4 '4 '44
Topsmelt A '' 4 44 '4' 4
Atherinops J 4 4 44 '4' 4
affinis L '4'4 444'4
T M STM ST MS TM ST M STM ST M STM S
Netarts Siletz Yaquina Alsea Siuslaw Umpqua Coos Rogue
Bay River Bay River River River Bay River
Legend:
T =Tidal fresh zone A = Adults '=Species / lifestage is present
M =Mixing zone J = Juveniles Blank = Species / lifestage is not present
S Seawater zone L = Larvae

297

Appendix 4 continued

Klamath Humboldt Eel TaMaleS Central San South San Elkhomn Morro
River s ay River' Bay Francisou Say' Francisco Bay Slough Bay
Species T MST M ST M S T M ST M S M S*S *5
Cutthroat trout A 4 4q 4 4 4
Oncorhynchus j 44 4
clarki L
Pink salmon A444
Oncorhynchus J
gorbuscha L
Churnsalmon A 44- 1
Oncorhyrnchus 444
keta L
Coho salmon A444444444444
Oncorhynchus J i
kisutch L
Steeihead -fall A444
Oncorhynchus J 4.
rnvkiss (F) L
Steelhead - half pounder A
Oricorhynchus J 4 44
mykiss (H) L
Steelhead -summer A 4 4 4
Oncorhyrichus J i
mykiss (S) L
Steellhead -winter A 4 i 4144444
Oncorhynchus J i ~
mykiss 1W) L
Sockeye salmon A
OncorhynchusJ
nerka L
Chinook salmon -fall A4444444444444
Oncorhynchus j 4 4 4 4 4
tshawytscha (F)
Chinook salmon - late fall A1 44
Oncorhynchuvs 44
tshawytscha (LF)
Chinook salmon - winter A 44
OncorhynchuVS J4 44
tshawytscha (W) 1L
Chinook salmon -spring A 4 -444444
Oncorhynehus J 4 4 V
tshawiflscha fSp) L
Chinook salmon - summer A
Onco~rhynchus J
Ishawytscha (Su) L
Surf smelt A 4 44 44
Hypomeasus J 4 4 44 4
pretiosus L 44444
Longfin smelt A 4 4 4 44 4 4
Spininchus J i
thaleichthys L4 4 44444
Eulachon Al
Thaleichthys J
Pacificurs L444
Pacific tomcod A444 44 44
Microgadus , J 444 4 4
proximus L
Topsmelt A 14 1

Atherinops j 4 4 4
T M TMS T MS T S T M1S M MS * S
Klamath HmbolIdt Eel Tmls CnrlSn SuhSn Eko or
River Bay River S ay Frlanciac Bay1YFrancisco Bay Slough Say

Legend: 1Includes San Pablo and Suisun Bays.
T Tidal fresh zone A =Adults 4 = Species / lifestage is present
M =Mixing zone J =Juveniles Blank = Species I lifestage is not present
S =Seawater zone L =Larvae
=Salinity zone is not present

298

Appendix 4 continued

Santa San Pedro Alamitos Anaheim Newport Mission San Diego Tijuana
Monica Bay Bay Bay Bay Bay Bay S ay Estuary
Species S 5 * S S 5 * 5 S S
Cutthroat trout A
Oncorhynchus J
clark.i L
Pink salmon A
Oncorhynchus J
gorbuscha L
chum salmon A
Oncorhynchus J
kera L
Coho salmon A
Oncorhynchus i
kisutch L
Steelhead - fall A
Oncorhynchus J
mvkiss (F) L
Steelhead - half pounder A
Oncorhynchus J
mykiss (H) L
Steelhead - summer A
Oncorhynchus J
mykiss (S) L
Steelhead - winter A
Oncorhynchus J
mykiss (W) L
Sockeye salmon A
Oncorhynchus J
nerka L
Chinook salmon - fall A '
Oncorhynchus J
tshaw-ytscha (F) L
Chinook salmon - late fall A
Oncorhynchus U
tshawytscha (LF) L
Chinook salmon - winter A
Oncorhynchus J
tshawytscha (Wi L
Chinook salmon - spring A
Oncorh~ynchus J
tshawytscha (SP) L
Chinook salmon - summer A
Oncorhynchus J
tshaw-ytscha (Su) L
Surf smelt A
Hypomesus J
pretiosus L
Longfin smelt A
Spirinchus J
thaleichthys L
Eulachon A
Thaleichthys J
pacificus L
Pacific tomcod A
Micro gadus J
pro ximus L
Topsmelt A'4''4''44'44
Atherinops J '
affinis L ' 4444'
S * S * S 5 * S
Santa San Pedro Alamitos Anaheim Newport Mission San Diego Tijuana
Monica Bay Bay Bay Bay Bay Bay Bay Estuary
Legend:
T =Tidal fresh zone A =Adults 4=Species / lifestage is present
M =Mixing zone J Juveniles Blank = Species / lifestage is not present
S =Seawater zone L =Larvae
=Salinity zone is not present

299

Appendix 4 continued

Puget Hood Skagit Grays Willapa Columbia Nehalem Tillarnook
Sound Canal Bay Harbor Bay River Bay Bay
Species T MST M S TM STM ST M ST M STM STM S
Jacksmelt A
Atherinopsis J
calitomiensis L
Threespine stickleback A ' 4' 4' 4' 4' 4' 4' 4' 4' 4' 4' 4'
Gasterosteus J' 4' 4' 4' 4' 4' 4' 4' 4' 4' 4' 4'
aculeatus L'44 '' 44 '' 44 '' 44 ''
Striped bass A ' .'
Morone
saxati/is L
Kelp bass A
Paralabrax i
clathratus L
Barred sand bass A
ParalabraxJ
nebuliler L
White seabass A '
Atracloscion J
nobilis L
White croaker A
Genyonemus J
lineatus L
Shiner perch A '44 '' '44 '' '44 ', '4 44
Cymatogaster J 4' 4' 4' 4' 4' 4' 4' 4' 4' 4' 4' 4'
aggregate p '4''4''4'''4 44
Pacific sand lance A '44 '.4 ', 4' '4 . ', 44
Ammodytes j 4 44 '' 4W 'W 'W '' 4
hexapterus L 'W 'W '' 44 '' 4 4'
Arrow goby A '44 '' '44 '4 ''44
Clevelandia j 44 '' 4' 4 44'
ios L '' 44 '' 44 ''
Ungcod A '4 '4 '4
Ophiodlon J 44 '' 4' 4 ' 4 '4'
eloaqatus L '4' '4'. ''4 4'4 4 4
Pacific staghorn sculpin A '44 '' 44 'W ''''' 4,4 ',
Leptocottus 4' 4' 4' 4' 4'4' 4' 4' 4' 4' '4'44'44
armalus L '44 '' 44' 4 '' 4 '
California halibut A
Paralichthys J
califomicus L
Diamond turbot A
Hypsopsefla J
_quttulata L
English sole A ' 4'
Pleuronectes J '4 '4 '4 W' '4 ''4 44 'W
vetulus L 'W '' 4 4 44 '4' 4
Starry flounder A ', 44 '' 44 'W ', 44 ''
Platichthys J 4 ' 4 ' 4 ' 4 ' 4 ' 4 ' 4 ' 4 ' 4 ' 4 ' 4 ' 4 '
stellatus L '4 '44 '' W' '44 '''4 44
T MS8 T M S T M S T M S T M S T M S T M S T M S
Puget Hood Skagit Grays Willapa Columbia Nehalern Tillamook
Sound Canal Bay Harbor Bay River Bay Bay

Legend:
T =Tidal fresh zone A =Adults '=Species / lifestage is present
M =Mixing zone J =Juveniles Blank = Species/I lifestage is not present
S =Seawater zone L =Larvae
P =Parturition

300

Appendix 4 continue,

Netarts Siletz Yaquina Alsea Siuslaw Umpqua Coos Rogue
Bay River S ay River River River Bay River
Species T M STM ST M STM STM ST M ST M STM S
Jacksmelt A 4'4
Atherinopsis J V
californiensis L
Threespine stickleback A ' 4V VV ' 4' 4V V '
Gasterosteus i
aculealus L V V V V V V V V
Striped bass A V'4' '4 4' 4
MoronejV V V
saxatilis LVVV
Kelp bass A
Paralabrax J
clathratus L
Barred sand bass A
Paralabrax J
nebulifer L
White seabass AVV
Atractoscion J
nobilis L
White croaker A
Genyonemus J
fineatus, L
Shiner perch A V '44 V4 V4V VVV V4
cymatogaster i V V V V V V V V V
aggregate P VV V VV V VV V VV V
Pacific sand lance AVVV'44VV
Ammodytes i
hexapterus LVVVV VV
Arrow goby AVV'4V4V V V
Clevelandia J i
ios L VV
Lingcod AV
Ophiodon i f
eloniqatus LV
Pacific staghorn sculpin A V 44 V V V V 'V V
Leptocottus JV V V V VV V VV V V V V V V V V V V V V V V
armatus LVVVVVVVV
California halibut A
Parafichthys J
cafifornicus L
Diamond turbot A
Hypsopsetta J
guttulata L
English sole A
Pleuronectes i VV VV VV VV VV V V
vetulus LVVVVV
Starry flounder A -4VV VVV VVV
Platichthys i V V
stellatus L VV V VV
T M S T M S T M S T M S T M S T M S T M S T M S
Netarts Siletz Yaquina Alsea Siuslaw Umpqua Coos Rogue
Bay River Bay River River R iver Bay River

Legend: A =Adults
T Tidal fresh zone J =Juveniles V=Species / lifestage is present
M =Mixing zone L =Larvae Blank = Species I lifestage is not present
S =Seawater zone P =Parturition

301

Appendix 4 continued

Klamath Humboldt Eel Tomrales Contra] San South San Elkhorn Morro
River s ay River Bay Francisco Bay' Francisco Say Slough Bay
Species T M ST MST M STM ST M S*M S*S *
Jacksmelt A444 44 4444
Atherinopsis i
californiensis L444 44 4444
Threespine stickleback A 44 4 4 4 4 4 4
Gasterosteus J44 4 4 4 4 4 4
aculeatus L44 44 444444
Stnped bass A4
Morone J
saxatilis L 4
Kelp bass A
Paralabrax J
clathralus L
Barred sand bass A
Paralabrax
nebulifer L
White seabass A44 44
Atra ctos cion J i
nobilis L
White croaker A444 44
Genyonemus i
lineatus L4 4444
Shiner perch A 44 44 -4 44 44 444
Cymatogaster i 4 4 44q 444 4 44 4
a!aqreqata p 4 4 4 4 4
Pacific sand lance A444444
Ammodytes J i
hexapterus L44
Arrow goby A444 44 4444
Clevelandia j444 44 4444
iOs L 4 4 4
Lingood A -4
Ophiodon J444 44444
elongatus L444
Pacific staghorn sculpin A 4 4 4 4 4
Leptocottus J 4 4 444 4 4
armatus L4444 44 444
California halibut A44444
Paralichthys i
calitornicus L44444
Diamond turbot A 44 4
Hypsopsetta i 4 4
piuttulata L 44
English sole A
Pleuroftectes J 4 4 4 44 4 4 4
vetulus L4444 4 44
Starry flounder A4444 44 444
Platichthys J 4 4 4 4 4 4 4
stellatus L 4 4 4 4
T MS TM T MS T Mr' .Tn.MS * S * 5* *
Klamath Humboldt Eel Toae etrlSn Suh Sa.Ekor or
River Bay River Bay Francisco Bay1 Francisco Bay Slough Bay
1Includes San Pablo and Suisun Bays.

Legend: A = Adults
T =Tidal fresh zone J = Juveniles 4=Species / lifestage is present
M =Mixing zone L = Larvae Blank = Species / lifestage is not present
S =Seawater zone P = Parturition
=Salinity zone not present

302

Appendix 4 continued

Santa San Pedro Alamitos Anaheim Newport Mission San Diego Tijuana
Monica Bay Bay Bay Bay Bay Bay Bay Estuary
Species * * S * * S * * S * S * S * * S * * S * * S
Jacksmelt A H' ' : 4
Atherinopsis J 41 :: . :
califomiensis L : '4 :
Threespine stickleback A
Gasterosteus J
aculeatus L
Striped bass A '4 :: '4
Morone J
saxatilis L
Kelp bass A ' 4 4 '4 ' / '4 '
Paralabrax J ' ' 4 '4 '4 ' '
clathratus L '4
Barred sand bass A '4 : ' / il ' i
Paralabrax J : '4 'J '4 . l l '4
nebulifer L ' 'J 4
White seabass A ' '4 I
Atractoscion J '
nobilis L
White croaker A V V '4 ' ' ' ''
Genyonemus J ' l l '4 4 'j '4
lineatus L '4 l 4 4 4 4 '
Shiner perch A '4 '4 I' '4 '4
Cymatogaster J '4 ' '4 '4 '4 'j
aq.qreqata P 4 ' 4 4 '
Pacific sand lance A
Ammodytes J
hexapterus L
Arrow goby A '4 '4 '4 ' '4 '
Clevelandia J '4 ' '4 ' ' '4
ios L ' l l 4 4 ' '
Lingcod A : ::
Ophiodon J ' '4
elongatus L
Pacific staghorn sculpin A '4 V 4 '4 4 '4 '
Leptocottus J ' ' ' ' l '4 '4 V
armatus L l '4 '4 'I
California halibut A 4V 4 ' 4 '4J '
Paralichthys J 4 4 '4 ' '4 J J 'j
califomicus L ' 4
Diamond turbot A ' '4 '4 '4 ' '4 ' '
Hypsopsetta J '4J ' '4 '4 '
Quttulata L 'J ' ' 4 '4 l
English sole A '4
Pleuronectes J 4 l
vetulus L '4 'J
Starry flounder A
Platichthys J
stellatus L
* * S * * S * * S * * S * * S * * S * * S * * S
Santa San Pedro Alamitos Anaheim Newport Mission San Diego Tijuana
Monica Bay Bay Bay Bay Bay Bay Bay Estuary

Legend: A = Adults
T = Tidal fresh zone J = Juveniles ' = Species / lifestage is present
M = Mixing zone L = Larvae Blank = Species / lifestage is not present
S = Seawater zone
= Salinity zone is not present

303

304

Index to Appendix tables 5A-5D: Page location of Biogeography, Habitat Associations, Biological Attributes
and Economic Value, and Reproduction tables for each species.

Common and Scientific Name Cmmon and Scientific Name
Blue mussel (My/i/us edufi6) Blue mussel (M'y/i/us edulis)
Pacific oyster (Crassos/rea gigas) Pacific oyster (Crassostrea gigas)
Horseneck gaper (Tresus capax) Horseneck gaper (Tresus capax)
Pacific gaper (Tresus nuttaffl~ 306 1 1 Pacific gaper (Tresus nflu/a//i
California jackknife clam (Tagelus cafifornianus) California jackknife clam (Tagelus ca/iforniarnus)
Pacific littleneck clanm (Proto/haca stamninea) Pacific littleneck clam (Protiofhaca s/aminea)
Manila clam (Venerupis japonica) Manila clam (Venerupisjaponica)
Softahell (Mya arenania) Softshell (Mya arenaria)
Geoduck (Panaopea abru p/a) Geoduck (Panopea abrupla)
Bay shrimp (Crangon franciscorum) Bay shrimp (Crangon franciscorum)
Dungeness crab (Cancer magistei) Dungeness crab (Cancer nragistei)
Leopard shark (Triakis semdiasciata) 307 313 319 Leopard shark (Triakis semifasciata)
Green sturgeon (Acipenser medirostris) Green sturgeon (Acipenser medirostris) 324
White sturgeon (Acipenser transmontanus) White sturgeon (Acipenser /ransmontanus)
American shad (Alasa sapidissima) American shad (Alosa sapidissirfa)
Pacific herring (C/upea pa//asi) Pacific herring (C/upea pal/asi)
Deepbody anchovy (Anchoa compressa) Deepbody anchovy (Anchoa compressa)
Slough anchovy (Anchoa deficalissimna) Slough anchovy (Anchoa de/icatssima)
Northern anchovy (Engraulis mnorday) Northern anchovy (Engrau//s mwrdax)
Cutthroat trout (Oncorhynchus c/ark,) 308 314 320 Cutthroat trout (Oncorhynchus clarki)
Pink salmon (Oncorfhynchus gorbuscha) Pink salmon (Oncorhynchus gorbuscha)
Chum salmon (Oncorhynchus keta) Chum salmon (Oncorhynchus keta)
Coho salmon (Oncorhynchus kis utch) Coho salmon (Oncorhynchus k/s utch)
Steelhead (Oncorhynchus mykiss) Steelhead (Oncorhynchus mykiss)
Sockeye salmon (Oncorhynchus, nerka) Sockeye salmon (Oncorhynchus nerka)
Chinook salmon (Oncorhynchus tshawy/scha) Chinook salmon (Oncorhynchus tshawy/scha)
Surf smelt (Hypamesaus pretiosus) Surf smelt (Hyporresus pretiosus)
Longfin smelt (Spirinchus tha/eichthys) 0 1 2 Longfin smelt (Spirinchus fthaeichthys)
Eulachon (Tha/eicifhys pacificus) 391521Eulachon (Tha/eichthys pacificus)
Pacific tomcod (Microgadus proximus) Pacific tomrcod (Micro gadus proximus)
Topsmelt (Athernnops affinis) Topamnelt (Athermnops affinis)
Jacksmelt (A/herinoposis ca/ifomniensis) Jackamelt (Afherinopsis ca/ifomniensis)
Threespine atickleback (Gas/erG steus aculeatus) Threespine stickleback (Gasterosteus acu/eatua)
Striped bass (Mfororne saxa/ilis) Striped bass (Morone sairafi/is)
Kelp bass (Para/abrax c/athratus) Kelp bass (Paralabrax c/athra/us)
Barred sand bass (Par/alabras nebu/ifer) 30 16 22 Barred sand bass (Para/abrax nebu/ife,)32
White seabass (Atractoscion nobi/is) 31 1632White seabass (Atractoscion nobi/is)32
White croaker (Gern'onemus lineatus) White croaker (Genyonemus lines/us)
Shiner perch (Cyrnatogaster aggregata) Shiner perch (Cymnatogaster aggregata)
Pacific sand lance (Ammodytes hexapterus) Pacific sand lance (Ammodytes hexapterus)
Arrow goby (C/eve/and/a ios) Arrow goby (C/eve/andia ibs)
Lingcod (Ophiodon e/onga/us) Lingcod (Ophiodon elongatus)
Pacific staghom sculpin (Leptocottus arma/us) Pacific staghorn sculpin (Leptoco//us armatus)
California halibut (Para/ich/hys ca/ifornicus) 311 317 323 California halibut (Para/ich/hys ca/ifom/icus)
Diamond turbot (Hypsopset/a gu/tu/ata) Diamond turbot (Hypsopsetta gut/u/a/a)
English sole (Pleuronectes ve/u/us) English sole (P/euronectes ye/u/us)
Starry flounder (P/a tich/hys s/el/a/us) Starry flounder (P/a tichthys ste//a/us)

305

L**. * **. I *-.

BIOGEOGRAPHY
IMarine Estuarine IRiverine
Li f sIg/ctv Salinity Range EsuryTp Stratifi-
A - Adults ISAB I Venice System caio
S -Spawning adults - ~ -'~

L -Larvae ~4
E -Eggs 0

J q0 0 0000 0 0 0
LC. 0 0 0 0 .* Q. 0 0 0~- .0 . 0L

Pacific oyster A 55 000 SSSS@S0S@ A
Crassostrea S 55 SS S SS S
gigas J 0* *. S0S J
L 0S* S @ O @ L
E 550SS S O E
Horseneck gaper A 55 55 5. 5 5 5 A
Tresus capax 55 5 **S* *** S
JO 55 ** .S SS J
LO So *S e e e L
ES 55 0OSSO E
Pacific gaper A S 55 *S S S S A
Tresus nuttalIN 55 SS SS S S S S
J 55 55 So . . .
LS OS SO S S S L
ES 55 SS S S S E
California jackknife A SSSSSSSSSA
clam 5 5OSS* S
Tagelus J SSS S55J
californianus L 55 0S S 55 00 L
E 55 0SO 55 00 E
Pacific littleneck AS 55 *SSS SSS A
clam 55 55* * * * * S
Protothaca J 55 55@ 5 5 5 5 J
staminea LO 55* S S S S L
ES 55 *S S SS E
Manila clam A 55 0 SSSSSSSS A
Venerupis 5 5*sS S * S
japonica J 55* *iS S
L 5 O S S SSL
E O SS S eeE
Softshell A 5 O SSO A
Mya arenaria S 5 * SS * S
J 50 SSSSS4S00 00 00 J
L 50 SSSSSSSSSS0 0 L
E 5SO O S S SE

306

Appendix 5A continued

BIOGEOGRAPHY
A - Adults I ~~~~Marine IEstuarine IRiverine
S - Spawning adults I aiiyRne Estuary Type I tatioi-
M - Mating ISAB IVenice System IIcto
J -Juveniles ' e0'
L -Larvae
E -Eggs C
P -Parturition Q-

Geoduck AS S00 0 0 00 * OS A
Panopea abrupta S 00 0 0@0 0 0 *000 S
i 0 0 0O 0 0 050 0 00 J
LOS 0 0 0 0 @ 0 0 00 L
ES 0 00 00 00 000 E
Bay shrimp A 0 0 0 0 @ 0 0 0 A
Crangon S 0 00S00 e o
franciscorum J 0 0 00 *@O@@@Se@@ee j 0
L 0 0 0 *0 0 0 0 0 00 L
E S 0 0 0 0 0*00 0000 E
Dungeness crab ASS 0 0 0 0 0 0 0 0 A
Cancer magister M 0 0 0 0 0M

L 0 000 0 0 0 0L
E O 0 0 E
Leopard shark AS 0 00 0 0 0 A
Triakis M 500 0 0 0M
semifasciata JO 0 00 0 0 0
P 0 0 0 0 0 P
Green sturgeon AS 0 0SO O OS O O A
Acipenser S * S
medirostris J 0 0 @0 0 000 05 50 J
L 0 0 L
E S E
White sturgeon A 0O 00 0 0 0 O O O OA
Acipenser 5 005
transmontanus J 0 0 0 00 00 00000 0 0 0 00 0J
L 0 0 0 OS L
E 0 E
American shad A 0 0 000 000000000 0 0SOSOOS *A
Alosa S 0 0 0 0 005
sapidissima J 0 0 0 00*0 0000 0 OSOSOOSJ *
L 0 0 0 OS L
E 0 0 0 OS E
Pacific herring A 0 0 0 0 0 0 00 00 000 A
C/upea 0 0 0 0 0050 0 00 00 0S
pallasi JOS00 S O @0 0 0 0 0 J
0 *0 0 00 SO S @S O L
E 0 0 0 050&060 000 05 E

307

Appendix 5A continued

Life -stage/activity IBIOGEOGRAPHY
A - Adults IMarine IEstuarine IRiverine
S- Spawning adults I Salinity Range 'Estuary Type Stratifi-
J - Juveniles ISAS Venice System c aion

E -Eggs

Deepbody A 0 0 0 0 0 A
anchovy S 0 0 S
Anchoa J 0 0 0 0 0 J
compressa L 0 0 0 L
E 0 0 E
Slough anchovy A 0 0 S S0 A
Anchoa S S 0 5
delicatissimna J 0 0 5 00J
L 00 0L
E 05 00 E
Northern A @ 0 05 *SSOSSOSOO A
anchovy S 0 0 00 0500 S
Engraulis J OS @5 0 0 0S@SS@O J
mordax L 0 @ 0*S0 0 0 0 L
E 0 0@ @5 0E
Cutthroat trout A 0 *@OS@OSO@OS0 @0 0 00 A
Oncorhynchus S S 0 00 S
clarki J 0 0 0 0 0 0 00 0 050 00 J
L 0 0 L
E 0 0 E
Pink salmon A 0 @00 *0 0 *0 0 050 0 S SOO A
Oncorhynchus S 0 0 0 0 0SSS
gorbuscha J 0 0 00 0 0 0 0 00 S *0 0 00 *0
L0 0 0 S S L
E 0S 0 S S E
Chum salmon A S 0 0 0 0 0 60 0 0 00 S OOSA
Oncorhynchus 5 0 0 0 S
keta J *0 0 0 0 00 0 0 0 *6 0 00 0 J
L0 0 0 0 OS L
E0 0 0 S O E
Coho salmon A S *OS00 0 0 050 0 0 0A
Oncorhynchus S 0 0 0000 0 005
kisutchJ** * * * * * * * * **
L 0 000 S L
E 0 5 I5 5*O E
Steelhead A 0 0**0000S SO SS 0S 0 0 0 0 0A
Oncorhynchus S 0 I I 0 00*0 0 005
mykiss J * ** * *D '* *********
L 0 00 *@ L
E 5 5 *O E

308

Appendix 5A continued

Life stagelacti~~ity BIOGEOGRAPHY
Lif- sAdulatsvt Marine Estuarine Riverine
S - Spawning adults I Salinity Range E s ur Typ Stratifi-
J - Juveniles SAB I Venice System EstaryTp tion
L - Larvae ~ ~ ~
E -Eggs '

Oncor y n ch u S 0 0
L 6 0 00
E-0 0C. 0 * 0 0 *. E
ChinookC. sa0nA0 0 0 0 .0 0. 0 00 0 0 0 00. A -
Oncorhynchus 5 * * 00 0
nerkysca J ** * ** * * 0 *S@SSSJ 000 000
L 0 0 S000 L
E 0 *0@E
Chinoosmelto A 0 0 *S00 0 0 0 0 0 0 0 0A
Oncorhyncus 5 0 0 @000 0 00s

L 0 000 * L
E 0 00 *0 E
Lourfinsmelt AO 0 @0 **S@@00 0 * 0 SOS A
Hypomescus SO0 SO @0 0 *000 0S
prthliosuhs J 0 00 0 0 0 @ 0 J
L 00 00 00 L
EO 0 0 00 E
ELaongfnml A 0 * 00 00 00 00 00 00 0 00 A
Spirinchuhs S 0 0 0 S
thaceichths J 0 00 *@iOO @
L *O@SS@@@O@OO@@@O0 0 0 L
E 0 0 0 0 0 E
Euacihconco A SO *@ OO SO 0 0 SO A
ThMichthyds 5 0 S * **
pacximius J S 0 J
L ***** 000 L
E 0 0 0 0 E
Tpacmfltomo A * 000 05 A
Micognadus 5 0 0
proxinus J 0 00 00 0 00 J
L 0000 0 00 L
E 0 E
Jacksmelt AO @ 0 50 50 0 A
Atherinopsi 5 0 0 * S00 0 5
calffminis J 00 OS0 0 0 0 0
L @50 0 0 0 0 L
E 500 0 0 0 E

309

Appendix 5A continued

I ~~~~BIOGEOGRAPHY
Life stage/activety Marine IEstuarine IRiverine
A - Adults I Salinity Range I Staii
S - Spawning adults SBIVncSytmIEstuary Type cattion
M -Mating SBIVnc ytmIcto
J -Juveniles - '~
L -Larvae ~ ~ o~
E -Eggs
P -Parturition z �

Threspie, 0 00 0 0 0 0 00 A~

stickleback S 55 55 e 0 0 SSOS* S
Gasterosteus J 550 0 *SS S * * S S S J
aculeatus L 550 05 0 SSSSSSS0 00 L
E 55 * 5 0 S0 0 0 S 00 E
Striped bass A S SO S S O 05 S A
Morone S 0 0 0 0O S S
saxails J 0O S J
L 55 5 5 5 @ ~ SL
E 55 S S5S S SE
Kelp bass ASS A
Paralabrax S S S S
clathratus 55 S 0 i
L 055 55L
E 0 E
Barred sand bass A S 0 A
Paralabrax 5 0 S
nebulifer 55 0 0 0 0 5 J
L S L
E 0 E
White seabass A S 0 A
Atractoscion S 6 S
nobilis 55 0 SO 0 J
LO 0 0 L
E 0 E
White croaker AS 55 SO S 0 55 000 A
Genyonemus 5 0 0 55 0 S
lineatus JO 55 55 5 5 5
LO 05 05 5 S L
E S 5 E
Shiner perch AS 55 50 5 55 A
Cymatogaster M 5* SS SS SM
aggregata J SO 0S S S S S S S J
P D S *S S SS S P 00 0I
Pacific sand lance AS *S * 00 0SOSSSSS0 00 A
Ammodytes 5550 5 55 @0 0 0 5
hexapterus J 55 55 SS * S * J
L S S SS 0 0S S L
ES 5 55000 E

310

Appendix 5A continued

BIOGEOGRAPHY
A - Adults I Marine Estuarine Riverine
S- Spawning adults Salinity Range E ca
J - Juveniles Venice System stuaryTypI Stiti
L-Larvae �
E-Eggs

a P

a ~ $ 0`~u0 C) b i~

Arrow goby A 0 S *O@@SSOOOSO A
Clevelandia S 0 O @ 0 s 0 0 0OSSO a S
los J 0 0 OSSOOSSOOSO J
L S OS SOOSSOOSSO L
E O OSOOSSOOSS E
Lingcod A 5 55 00000 005 A
Ophiodon S 0 0 s 05 0 5 5
elongatus J S SO OSOSOSSOOS J
L 0 0 05005500 L
E 0 00 0 0 E
Pacific staghorn A 0 0 *SOOOOOSOO A
sculpin 5 0 005050 0 S
Leptocottus J S *o 0 o .ooo..... J
armatus L 0 00 @55000505 L
E S SOSOOSOS E
California halibut A 0 5 5 0 0 0 0 5 A
Paralichthys S S S
californicus 00i 05 0000 0 5 J
LOS OS *oo S OS L
E * S E
Diamond turbot A 0 0 0 5 * A
Hypsopsetta S 0 0 0 0 S
gutfulata J 0 00 00000 0 0 J
L 0 0 0 0 0 00 L
E 0 0 0 0 E
English sole A 0 0 0 0 0 A
Pleuronectes S 0 s
vetulus 00i 0 0 * 0 0 0 00 J
LOS OS 0505000 0 L
E 0 5 0 0 000 E
Starry flounder A 0 5 5050 0050 A
Platichthys S 0 0 0 s
stellatus J 0 *SOOOSOOOO ees o J
L 5 00 OOSOOOOO0o L
E 0 0 5 E

311

IAff 0 .-o:lm�

I ~~~~~~~HABITAT ASSOCIATIONS
Habitats Substrate preference Domain
I Benthic I elagic I EstuarineI

Life staoe/a~~~~~~~~~~~~~~~~~~jy& ~~~~Littorall Sublittoral I13athyall

S -Spawning adults
J -Juveniles
L -Larvaea
E- -Eggs aa

Blue mussel A 0 C 00000
Mytikis edulis S *SO 0 COOC 0*S
J CC@ 0. 0 ..0
L OS C * C C L
E C@ C C C CE
Pacific oyster A a *C00 :0 * C A
Crassostra S 0000S
gigas J 0 0C0
L 0 COOC0L
E * 0 S S C S E
Horseneck gaper A *@ 0 9 CC0 0O S S A
Tresus capax S SC 0 *55*500
J Ce S @5 5**0
L * 0 0 0 * S L
OC S 0 * S C E
Pacific gaper A 00 0 *S 0C * C A
Tresus nuttafli S @0 S S5 0
J 0 0 0 OS 00 0 46J
L @ 00 * 0 0 O S L
E @0 0 0 0 * S E
California jackknife A * * 0@ 0@*C A
clam S SC 9 000 S* S
Tagelus J CS Ce OS 000
califomianus L 0 0 0 * L
E ge * S O E
Pacific littleneck A *g0 * * * 0* * S A
clam S em 0 see *e0 60S
Protothaca J 0 0 S *@* 00 CO C * J
staminea L *@0 0 0 C C CO L
E *@0 q 0 * * @ E
Manila clam A 0 @0 *5 CC O A
Venerupis S 0 00 *0 0
japonica J 0 CC0 0 00
L 0 0 0 0 S OL
E 0 C 0 CCE
Softshell A 0 CC 0 CC * C A
Mya arenania S Ce 0 000 O C S
J ~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 0C0 @0 . .J
L @6~~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~ 0 * C L
E OS0 t C C * C E

312

Appendix 5B continued

HABITAT ASSOCIATIONS
Habitats Substrate preference IDomain
I Benthic I1Pelagic I Estuarine
Life stacielactivety ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~~~ Littoral ISublittoral lBathyall
A -Adults
S -Spawning adults
M -Mating
J -Juveniles
L -Larvae
- Eggs
P -Parturition

Geoduck A 0 SO 60 50 000 0S A
Panopea abrupla S 0 @ 0 S @0 0 0 0o 0s

L @0 0 0 5 * O L

Say shrimp A 06 00 9 e A
Crangon S @6 000 a 000 S
franciscorum J @ 0 @0 00 0 0 060
L @ 0 0 O 0 0S 0 * 0 L
E @ 0 0 @00 0 0 E
Dungeness crab A @ 0 0 00 000500 @550 A
Cancer magister M 0 0 0066 M

L 0 00 060 0 000L
E @ 0 0 E

Leopard shark A 0 05 0 0 0 0 0@0000 0 0 A
Triakis M ** 0 M
semifasciata J @ 0 0 0 0 0 0560 0555 000
P P
Green sturgeon A 0 @ 0000 0@0 . 0 0 05 00500 *0 0A
Acipenser S 0 0 S
medimostris J 00 0 0 00 00 0 00 0 0 0S*0
L 0 0 * oL

White sturgeon A @ 0 0 60 6 @00 * 0 A
Acipenser S 0 S0
transmontanus J @ 0 0 00 0 0 00 0 0 0 0 0 0 0 0
L @5@ 000 0 L
E 0@02
American shad A S 0 0 0000 0 A
Alosa S 55 0 @0 S
sapidissirna J 0 00 00 0 S0 0 0
L @5 0 0 L
E 06 00 @5 E
Pacific herrnng A @ 0 0 6 00 00 A
Clupea S 06 60 00
pallasi J @0 5 0 0 0 0 0 0
L @60 0 0 0 * S L
2 00 55 @0~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ *E

313

Appendix 58 continued

HABITAT ASSOCIATIONS
Habitats Substrate preference I Domain
I Benthic I Pelagic Estuarine
[LItlorall Sublittoral I1athyall
Life stage/agjjy&t
A- Adults
S - Spawning adults
J-Juveniles -_X
L -Larvae
2- Eggs aa

Deepbody A * 0 6 0 0 0 0 A
anchovy S 0 0 0 S
Anchoa J 0 0 0 0 *00 J
compressa L 0 S S 000 L
2 0 0 @0 E
Slough anchovy A 0 0 0 0 0@ 0 A
Anchoa S 0 0 S
delicatissima J 0 i 0 S @ U
L a 0 000 L
E 00 2
Northern A 00 00 0 @0000 A
anchovy S 0 @ 0 00 S
Engraulis J 00 00 0 @0000 J
mordax L *0 0 00000 L
E 0 00 0 @0 E
Cutthroat trout A 0 0 0 00 0 A
Oncorhynchus 5 0 00
clarki f 0 000 0 0000J
L 0 00 L
E E -( ::::
Pink salmon A 0 0 0 0 * 000 A
Oncorhynchus S 0 000 S
gorbuscha J 0 00 0 aOOOOOJ
L @0 *0 L
E 00 E
Chum salmon A 0 00 0 00 00 A
Oncorhynchus S 00 00 S
keta J 0 00 0 00 0000J
L @0 @0 L
E @0 @ 2
Coho salmon A 00 0 0 0 A
Oncorhynchus 5 0 00 S
kisutch J 00 0 0 0 000000 J
L 0 00 L
E 0 00 E
Steelhead A @0 0 0 0 0 0 0 0 A
Oncorhynchus S 0 0 S
mykiss i 00 00 0 0. 006600 J
L 0 00 L
E 0 00 E

314

Appendix 5B continued

HABITAT ASSOCIATIONS

Habitats Substrate preference I Domain
I Benthic I Pelagic I Estuarine I
ILittoral I Sublittoral IBathyall

A -Adults
S - Spawning adults 44
J -Juveniles e
L -Larvae
- Eggs

Sockeye salmon A

Oncorhynchus S 0 C S
nerka JC CC 0 0 iee
LC 0 0 L
2CC CC E
Chinook salmon AC @0 C C C C C C CA
Oncorhynchus S C C S
tshawytscha J 0@ a@ e0 0 0 0 i
L C C L
2 E C
Surf Smelt A 0C O 0 *CC*CC0 0A
Hypomesus S C CC C
pretiosus J 0 0 a
L C C C CCL
2- E 0C
Longfin smelt A 0C CC C C 4 6 A
Spirinchus S 0C C C S
thaleichthys J C C C 0 C C C C J
L 0C C C * 0 L
E 0C E
Eulachon A CC6 0 C 00 0C A
Thaleichthys S C C
pacificus J C 0 J
L CC C C @ L
2- E
Pacific tomcod A C 400 4C CCC A
Microgadus SC S
proximus J C OCC 0C 0C J
L C CCC C L
- IF2
Topsmelt A CC 000 CS C C C C A
Atherinops S C C C C C C C S
affinis J C CC C C C CJ
L CC 0 0 C C C CL
E CS C C C CE
Jacksmelt A C 1 SI C C C C C CA
Atherinopsis S 000 0C CC C C S
califomniensis J a CI CC C C 0 4CC
L CC0 CSL
2 C C C

315

Appendix 5B continued

I ~~~~~~~HABITAT ASSOCIATIONS
Habitats ~~Substrate preference Domain
I Benthic IPelagic IEstuarine
~~~~~~~~~~~~~~~~~~~~~~~~~~~~ILifesaeatlyLtorall Sublittoral IBathyall
A -Adults
S - Spawning adults
M -Mating
J -Juveniles
L -Larvae
- Eggs
P -Parturition e a-o u

Threespine A 0 0 0 * 0 @ A
stickleback S@ O 0 0 @0 OS S
Gasterosteus i 0 0 0 0 *0@O
aculeatus LOO0 e S 00 L
20000 00 @ 0 r-
Striped bass A *e * 0 00 * SOS A
Marone S OS S S
saxatifis J **0 0 0 0 *S*OJ

L @0 O@0 0 L

Kelp bass A *0 @0 0 0 0 00 SA
Paralabrax S 0 0 @ S
cla thra tus J ***O 0 05 * ** J
L 0 00L

Barred sand bass A O* O *0 0 S *0 @ 0 A
Paralabrax S 0 S
nebulifer J 0 * 0 0 *0 0e 0 000 J
L Se * 0 0 L
2 0 E
White seabass A 0 0 0 * 0 A
Atracloscion S S
nobifis J 00 00 *@ Se 0 000 J
L 0 0 * * 0 L
2n a r-S
White croaker A e@ ** * 0 00 0 0 O SA
Genyonemus S S 0 S
lineatus J SO0 0 0 0 0 0 * S J
L @ 0 000 @ 0 L
2 00
Shiner perch A @0 0 * 0 0 0 *0 0 A
Cymatogaster M @0 0 0 0 0 0 0 M
aggregata J 0 00 0 05 0 0 *@i
P 00 0 0 0 0 e eP
Pacific sand lance A @0 0 0 S4S A
Ammodtytes S 0 0S05 0 0 S
hexapterus J 000 0 0 .000 40 0 J
L 0 0 0 00 0L
2 S E S

316

Appendix 5B continued

I ~~~~~~~HABITAT ASSOCIATIONS
Habitats Substrate preference Domnain
Benthic I Pelagic IEstuarine
ILittorall Sublittorall I1athyall

A -Adults ~-
- Spawning adults
J -Juveniles CD~'
L -Larvaea 2 Z-s a
- Eggs

Arrow goby A I @0 @0 @000
Clevelandia S @0 @ 00 @00 * e s
jos J 0 @0 @0 000
L I 0 0 0 0 0L
E @0 @0 @0 00 2
Ungcod A 0060 @ 0 0 0 00000 A
Ophiodon S 0 0 00* 0 00 S
elongatus J 0 0 0 0 4 0 0 0 000 0 J
L 0 0L
2 0 0 0
Pacific staghorn A 0 0 0 0 *00 0 0 0@0 00 0 40 0 A
sculpin S 0 0 0 40 0 S
Leptocottus J 000 0 0 00 0 00 0 0 000
armatus L **4 * 00 00 L
E @0 0 0 0 E
Califomnia halibut A 0 @0 S5 00 0 A
Paralichthys S 0 0 0 S
califomnicus J SO 0 0040 0 0 0 0 0 0 *
L @0 @ 0 0 0 0 L
2 0 0 F-
Diamond turbot A @0 S 000 0 O SA
Hypsopsetta S 0 0S
gutut~lata J @0 0 0@ 0 O
L 00 S00 0 L
E 6 00
English sole A 0 0@ 0 0 A
Pleu~ronectes S 0 0 0 0S
vetuilus J 0 .0 000 0 00 0 0 S*0 0J
L @0 @ @00 L
2- E000
Starry flounder A @0 @00 00 00 A
Pla ach thys S 0 50S
stellatus J 000 0 0 0 00000
L @0 a 000 L
E 0 0

317

Life sage/atiylI BIOLOGICAL ATTRIBUTES IEconlomic
A - Adults jFeeding type Spatial strategy Longevity Value
S - Spawning adults
J -Juveniles c ~ bt
L -Larvae

E~~~~~~~~~~~~ - Eg

Blue mussel A 0 0 A
Mytilus edlulis S **S
J @5 5@ 0 J
L 0 50L
E @0 0E
Pacific oyster A 06 0 0 0 0 A
Crassostrea S 0 S
gigas J 0 0 0 0J
L 0 ~L
E 0 0E
Horseneck gaper A 0 A
Tresus capax 5 0
J 0@0
L 000L
E 0 0E
Pacific gaper A 0 0 0 0 A
Tresus nuttailli S 0
J S@ 0 J
L 0 00 L
E E
California jackknife AS 0 0 0 0 S S A
clam 5 S
Tagelus J 50 0 S J
californianus L 0 * L
E S SE
Pacific littleneck A 0 0 0 0 S A
clam S @6
Protothaca J 0 0 0 0 J
stamninea L 5 0 L
E. 0 E
Manila clam A SS 0 S S S A
Venerupis 5 0 S
japonica J 0 0 0 0 J
L SSSL
E 5 5E
Softshell A 5O * 0 0 A
Mya arenaria 5 S
J 05 0 5 ~~~~~~~~~~~~~~~J
L @500L
E S SE

318

Appendix 5C continued

BIOLOGICAL ATTRIBUTES IEconomic
AL- Adults Feeding type jSpatialstrategy Longevity Value
S - Spawning adults
M- Mating
J-Juveniles - '- 0~
L -Larvae
E-Eggs
P- Parturition

Geoduck A @0 00 0 0 0 A
Panopea abrupta S 0 0 5
J 00 i0 J
L 0 00 0 L
E 000 E
Bay shrimp A 0 0 0 0 A
Crangon S 0 0 5
franciscorurm J 0 0 0 J
LO 0 L
E 00 E
Dungeness crab A 0 00 0 0 0 A
Cancer magister M 0 M
J0 o J
LO 0 00 0 L
E 0 0 E
Leopard shark A 0 0 0 00 0 A
Triakis M 00 M
semifasciata J 0 0 0 J
P 00 P
Green sturgeon A 0 00 0 0 0 A
Acipenser S 0 5
medirostris J0 0 0 0 J
L 0 0 L
E 0 0 E
White sturgeon A 0 0 0 0 0 0 A
Acipenser 5 0 S
transmontanus J 0 0 6 0 00 0 J
L 0 0 L
E 0 0 E
American shad A 0 0 0 0 0 A
Alosa S 0 5
sapidissima J 0 000 0 J
L 0 0 L
E 0 E
Pacific herring A 0 0 0 0 0 0 0 A
Clupea S 00 0 5
pallasi J 0 00 00 0 0 0 J
L 0 0 0 L
E 00 0 0 E

319

Appendix 5C continued

Life stage/activity I BIOLOGICAL ATTRIBUTES IEconomic
A-Adults Feeding type j Spatial strategy Longevity Value
S - Spawning adults
J-Juveniles
L- Larvae

8~ e
E - Eggs0

Deepbody A 0 0 0 0 A
anchovy S 0 S
Anchoa J 0 0 S 0 J
compressa L 0 S 0 L
E S E
Slough anchovy A 0 0 0 A
Anchoa S S
delicatissima J 0 0 0 J
L 0 0 0 L
E 0 0 E
Northern A SS0 0 A
anchovy S 00 S
Engraulis J 0 5 0 0 0 S 0 J
mordax L S 00 L
E @0 5 E
Cutthroat AO @00 00 A
trout 5 S
Oncorhynchus Ji 0 0 0* 0 * 0 J
clarki L 5 0 L
E S 0 E
Pink salmon A 0 S S S S 0 A
Oncorhynchus S S S
gorbuscha Ji 55500 0 J
L S S L
E 0 E
Chum salmon A 0 00 5 0 0 A
Oncorhynchus S 0 5
keta JO 00055 00 J
L 0 5 L
E 5 0 E
Coho salmon A 5 5 0 S A
Oncorhynchus S S S
kisutch JO 0 0 J
L S L
E S S E
Steelhead A 0 0 0 0 A
Oncorhynchus S 0 S
mykiss JO 0000 0 J
L 0 0 L
E 5 0 E

320

Appendix 5C continued

Life stage/activity I BIOLOGICAL ATTRIBUTES IEconomic
A - Adults Feeding type ISpadal strategy Longevity Value
S - Spawning adults
J-Juveniles
L-Larvae
E ~ ~ F- Eggs

Sockeye salmon A0 000 0 0 0 0 A
Oncorhynchus S 0 5
nerka Ji 0 00000 @ J
L 0 0 L
E S E
Chinook salmon A 0 0 0 0 0 0 0 A
Oncorhynchus S 0 S
tshawytscha J * * 0 ' 0 0 0 0 .0 0 i
L S 0 L
E 0 0 E
Surf smelt A 0 0 0 0 0 0 0 A
Hypomesus S 0 0
pretiosus JO S 0 @0 0 J
L 0 000 L
E 00 S E
Longfin smelt A 0 0 0 00 0 0 A
Spirinchus S 0@0 S
thaleichthys JO 0 0 000 0
LO 0 0 0 L
E 0 0 E
Eulachon AS 0 0 @00 0 0 0 A
Thaleichthys S 0 S
pacificus JO 0 0 0 0 J
LO 0 @00 0 L
E S @0 E
Pacific tomcod A 0 0 0 0 5 A
Microgadus S 0 S
proximus J 0 0 a 0 J
LO0 0 L
E S E
Topsmelt A 0 0 00 00 0 S A
Atherinops S * S
affinis J 000 00 J
L 00 00 0 L
E 00 00 E
Jacksmelt A 0 0 00 0 0 0 A
Atherinopsis 5 0@ S
californiensis J 0 0 0 0 0 0
L @0 00 00 L
E 00 0 E

321

Appendix 5C continued

Life stage/activity j BIOLOGICAL ATTRIBUTES IEconomic
A- Adults Feeding type Spatial strategy Longevity Value
S - Spawning adults
M- Mating
J -Juveniles
L - Larvae
E-Eggs
P- Parturition

Threespine A * 0 0 0@ 0 A
stickleback S S 0
Gasterosteus J 0 0 0 J
aculeatus L 0 0 0 0 L
E 00 0 0 E
Striped bass A 0 0 0 0 A
Morone S 0 S
saxatilis JO 00 00 J
LO 00 0 L
E E
Kelp bass A S 0 0 A
Paralabrax S 0 S
clathratus JO 00 0 J
LO 0 0 L
E 000 E
Barred sand bass A * 4 1 0 0 A
Paralabrax S 0 S
nebulifer J 00 J
L 0 L
E 0 0 E
White seabass A 0 0 0 0 0 0 A
Atractoscion S 0 S
nobills JO 00 0 J
L 0 0 0 L
E 0 E
White croaker A 0 0 0 0 0 0 A
Genyonemus S 0 S
lineatus J * 0 0 J
LO 0 00 0 L
E 0 0 E
Shinerperch A 0 0 0 0 0 A
Cymatogaster M 0 M
aggregata Ji 0 00 J
P 0 0 P
Pacificsand lance A 0 0 0 000 0 0 A
Ammodytes S 0 0 s
hexapterus J0 0 0 00 0 J
LO 00 0 L
E 00 00 E

322

Appendix 5C continued

Life stagie/activitv I BIOLOGICAL ATTRIBUTES IEconomic
A - Adults Feeding type Spatial strategy Longevity Value
S - Spawning adults
J -Juveniles
L- Larvae B
E-Eggs 4

Arrow goby A 4 0 0 A
Clevelandia S 0 S
ios JO 0 0 J
L 0 0 0 L
E 0 0 E
Lingcod A S 0 S 0 A
Ophiodon S S S
elongatus JO 0 0 0 J
LO 0 0 L
E S E
Pacific staghorn A 00 0 A
sculpin S 0 S
Leptocottus J 0 0 0 J
armatus L0 0 @0 0 L
E 00 E
California halibut A 00 0 A
Paralichthys S S
californicus JS 0 0 S J
LO 0 0 S L
E S E
Diamond turbot A 0 0@ A
Hypsopsetta S 0 S
guttulata Ji 0 0 0
L 0 0 L
E 0 0 E
English sole A S 0 0 A
Pleuronectes 5 0 S
vetulus JO 0 0 0 J
LO 0 0 S L
E 0 E
Starry flounder A 0 0 0 0 0 0 0 A
Platichthys S 0 S
stellatus J 0 0 J
L 0 0 L
E 0 0 F

323

I.,- * **~~ I *.I. ..0 .

REPRODUCTION
Fertilization! I Spawning Spawning TPeriodicity Domain
Egg Developmentl type behavior mpora ceue

Blue mussel
Mytilus edulis
Pacific oyster * * * * *
Crassostrea gigas
Horseneck gaper * * * * * *
Tresus capax
Pacific gaper * * * **
Tresus nuttaffli
California jackknife clam * * * * * a 0 * * * *
Tagelus califomianus
Pacific littleneck clam
Pratothaca staminea
Manila clam
Venerupisjaponica
Softshell * a
Mya arenaria
Geoduck
Panopea abrupta
Bay shrimp 0
Crangon franciscorumr
Dungeness crab 0 0 0 0 41111 0 0 6 0
Cancer magister
Leopard shark
Triakis semifasciata
Green sturgeon * * * * * * *
Acipenser medirostris
White sturgeon * * * * * * * *
Acipenser transmontanus
American shad
Alosa sapidissima 0
Pacific herring
Clupea pallasi .
Deepbody anchovy * * * * * * *
Anchoa compressa
Slough anchovy * * * * * * *
Anchoa delicatissima
Northern anchovy 0 0 0 0 0 0 0 :400 0 0 0 0 0
Engraulis mordax
Cutthroat trout * * a * a
Oncorhynchus clark,
Pink salmon
Oncorhynchus gorbuscha 0 0 a
Chum salmon
Oncorhynchus keta
Coho salmon 0 0 0 0 4
Oncorhynchus kisutch
Steelhead * * * *
Oncorhynchus mykiss
Sockeye salmon
Oncorhynchusnerka 5 0 *
Chinook salmon
Oncorhynchus tshawytscha

324

Appendix 5D continued

I ~~~~~~REPRODUCTION

Egg Development tySpewin behavnior Temporal Schedule Periodicity Domain

Hypomesu pretious 0 0 0a 6:4 a0 0 0.

Longfin smelt 0 0
Spirinchus thaleichthys 00
Eulachon
Thaleichthys pacificus C5 065
Pacific tomcod * * 06 0 ...
Microgadus proximus
Topsmielt @
Atherinops affinis C Se .
Jacksmelt 0 0 0 CS0

Threespine stickleback 00000000
Gasterosteus aculeatus 550 5CC0565
Striped bass * *
Morone saxatilis
Kelp bass 0a80
Paralabrax clathralus 0 S SS5
Barred sand bass 000
Paralabrax nebulifer 0 00 60 @ 6
White seabass 00
Atractoscion nobilis S 500 0
White croaker * * * * * * * * * * a
Genyonemus lineatus
Shiner perch
Cymnatogasteraggregata S S 5S C
Pacific sand lance*****
Ammodytes hexapterus
Arrow goby 0 0411
Clevelandia ios0 se . . .S
Lingood * * ** a**000
Ophiodon elongatuis
Pacific staghorn sculpin * * * * * * * *
Leptocattus armatus
California halibut
Paralichthys califomnicus 0 0 0 50 0 0S 0
Diamond turbot 000000
Hypsopsetta guttulata 0 S0 0
English sole * * * a * * * *
Pleuronectes vetulus
Starry flounder*****
Platichthys stellatus 0 S

325

326

BIOGEOGRAPHY
Marine- Distribution of life stages in seawater areas.
*Beach- Exposed shore areas receiving ocean waves and wash.
*Continental shelf- Waters over the gradually-sloping continental seabed from shore to a depth of
about 200 m.
*Continental slope- Waters over the steeply-sloping seabed from continental shelf to 1000 m.
*Oceanic- The open ocean waters beyond the continental shelf. Defined here as the ocean beyond
the continental slope.

Estuarine- Distribution of life stages in estuarine areas.
Salinity Ranae: SAB (Strateaic Assessment Branch classification)
* Tidal fresh- Salinities of 0.0-0.5%oo.
*Mixing- Salinities of 0.5-25.0?/0.
*Seawater- Salinities >25%0.

Salinity Ranae: Venice classification
*Limnetic- Salinities of 0.0-0.5%0.
* Oligohaline-Salinities of 0.5-5.0%/o.
*Mesohaline- Salinities of 5-18%o.
*Polyhaline- Salinities of 18-30%o.
*Euhaline- Salinities >30%0.

Estuary type
*Drowned river- Estuaries resulting from valleys being inundated by rising sea levels (e.g., Grays
Harbor and Columbia River estuary).
*Bar-built- Estuaries resulting from the building of barrier islands or spits (e.g., Netarts Bay and
Humboldt Bay).
*Fjord- Glacier-formed estuaries with deeply-carved, steep-sided channels (e.g., Puget Sound and
Hood Canal).
* Tectonic- Estuaries formed by faulting orsinking of the earth's crust (e.g., Tomales Bay and South San
Francisco Bay).

Stratification
*Highly-Very little mixing between surface and bottom layers, resulting in marked differences between
surface and bottom salinities.
*Moderately- Moderate mixing between surface and bottom layers primarily due to tidal-induced
turbulence. Surface salinities are usually lower than bottom salinities.
*Homogeneous- High mixing of surface and bottom layers resulting in equivalent salinities.

Riverine- Distribution of life stages in freshwater areas.
*Coastalplain- River portions in the relatively flat land along a coast.
-Piedmont- River portions at the base of mountains.
* Upland- River portions in mountainous areas.

HABITAT ASSOCIATIONS

Habitats- General habitat of life stages.
*Lake- Freshwater non-flowing areas with riverine connections to the sea.
*River/stream- Areas with flowing fresh water.
*Estuarine- Embayment with tidal fresh, mixing, and seawater zones.
*Bay- Semi-enclosed water body that has predominantly seawater salinities.
*Coastal (high energy)- Nearshore areas subject to significant wave or current action.
*Coastal (low energy)- Nearshore areas subject to only minor wave or current action.
*Offshore- Offshore areas beyond the coastal high or low energy areas.

327

Appendix 6 continued

Substrate preference- Size of substrate that life stages reside on or in.
*Mud/clay/silt- Fine substrates <0.0625 mm in diameter.
*Sand/granule- Substrates 0.0625-4.0 mm in diameter.
*Pebble- Substrates 4-64 mm in diameter.
PCobble- Substrates 64-256 mm in diameter.
-Boulder- Large substrate >256 mm in diameter.
�Rocky outcrop (bedrock)- Exposed solid rock.
*Estuarine vegetation- Aquatic plants within an estuary.
�Marine vegetation- Aquatic plants in marine waters.
'None- No reported preference.

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

Benthic-Sublittoral
�Inner shelf (10-50 m)- On the bottom of the continental shelf at depths of 10-50 m.
�Middle shelf (50-100 m)- On the bottom of the continental shelf at depths of 50-100 m.
*Outer shelf (100-200 m)- On the bottom to the edge of the continental shelf at depths of 100-200 m.

Benthic-Bathval
*Mesobenthal (200-500 m)- On the bottom of the continental slope at depths of 200-500 m.
*Bathyobenthal (>500 m)- On the bottom of and beyond the continental slope at depths >500 m.

Pelaiic
-Neritic- Residing within the water column from the shore to the edge of the continental shelf.
Oceanic- Residing within the water column beyond the edge of the continental shelf.

Estuarine
'Mainstem channel- The deep, drowned river channel of an estuary
-Subsidiary channel- Small tributary channels emptying into the mainstem channel of an estuary.
-Channel edge- Rim of an estuarine channel where the bottom slopes upward and meets shallow flats.
'Intertidal flat- Shallow, often almost level estuarine areas alternately covered and left bare by tidal waters.

BIOLOGICAL ATTRIBUTES AND ECONOMIC VALUE
Feeding type- Trophic role of life stages.
'Carnivore- A flesh-eating organism.
'Herbivore- A plant-eating organism.
'Omnivore- An organism that eats both plants and animals.
'Planktivore- A plankton-eating organism.
�Detritivore- A detritus-eating organism.

Spatial strategy- Use of habitats by life stages.
-Coastal migrant- An organism which migrates within nearshore waters of the continental shelf.
�Ocean migrant- An organism which migrates in ocean waters beyond the continental shelf.
-Freshwater resident- An organism which resides primarily in freshwater habitats.
�Estuarine resident- An organism which resides primarily in estuarine habitats (salinities 20.5 and <25%0).
-Marine resident- An organism which resides primarily in seawater habitats (salinities >25%o).

Longevity- Average lifespan of a particular life stage ( day to >20 years).

Economic Value- Monetary worth (direct and indirect) from harvesting a species.
�Recreational- Harvested by sport anglers.
'Commercial- Harvested by professional fishermen for sale in markets.

328

Appendix 6 continued
REPRODUCTION
Fertilization/Egg Development- Method of egg fertilization and development.
*External- Egg fertilization occurs after eggs and sperm are shed into the water.
-Internal- Egg fertilization occurs when a male inseminates an egg within a female.
-Oviparous- Eggs are laid and fertilized externally.
�Ovoviviparous- Eggs are fertilized and incubated internally, and usually released as larvae. Little or
no maternal nourishment is provided.
* Viviparous- Eggs are fertilized, incubated, and develop internally until birth. Maternal nourishment is
provided.

Mating Type- Mate selection strategy.
*Monogamous- A single male and a single female pair for a prolonged and exclusive relationship.
*Polygamous- A male mates with numerous females or vice-versa.
*Broadcast spawner- Numerous males and females release gametes during mass spawning.

Spawning- Spawning mode.
-Anadromous- Species spends most of its life at sea but migrates to fresh water to spawn.
*Iteroparous- Species reproduces repeatedly during a lifetime.
*Semelparous- Species reproduces only once during a lifetime.
-Batch spawn- Species spawns (releases gametes) several times during a reproductive period.

Parental Care- Type of egg protection.
*Protected- Eggs are protected by parent(s); eggs are buoyant or attached to substrates, but not buried.
*Nests- Eggs develop in the shelter of a nest.

Temporal Schedule- Period when spawning typically occurs.
*Early spring- From mid-March through April.
*Late spring- From May to mid-June.
*Early summer- From mid-June through July.
-Late summer- From August to mid-September.
-Early fall- From mid-September through October.
-Late fall- From November to mid-December.
*Early winter- From mid-December through January.
*Late winter- From February to mid-March.

Periodicity- Frequency of spawning events.
-Annual spawning- Spawning once each year, usually during a restricted season.
-2 or more per year- Spawning more than once each year (more than one spawning season).
*2 or more years- Spawning events separated by at least two years.
*Undescribed- Spawning frequency not documented.

Domain- Location of spawning.
-Oceans- Spawning occurs primarily in open marine waters.
-Estuaries- Spawning occurs primarily in estuarine waters (to head of tide).
-Rivers- Spawning occurs primarily in fresh water, above head of tide.

329

The Strategic Environmental Assessments (SEA) Division of NOAA's Office of Ocean Resources Conservation
and Assessment (ORCA) was created in response to'the need for comprehensive information on the effects of
human activities on the Nation's coastal ocean. The SEA Division performs assessments of the estuarine and
coastal environments and of the resources of the U.S. Exclusive Economic Zone (EEZ).

In June 1985, the National Oceanic and Atmospheric Administration (NOAA) began a program to develop a
comprehensive information base on the life history, relative abundance and distribution of selected fishes and
invertebrates in estuaries throughout the Nation (Monaco 1986). This program, the Estuarine Living Marine
Resources (ELMR) program, is conducted jointly by the Strategic Environmental Assessments Division of the
Office of Ocean Resources Conservation and Assessment and laboratories of the National Marine Fisheries
Service (NMFS). Currently, the Point Adams (Hammond), OR; Galveston, TX; Beaufort, NC; and Oxford, MD
laboratories are compiling information for the contiguous West Coast, Gulf of Mexico, Southeast, and Northeast
regions. Also, the Virginia Institute of Marine Science is compiling data for the Chesapeake Bay area.

Three salinity zones, as defined in Volume 1 of NOAA's National Estuarine Inventory Data Atlas (NOAA 1985),
provided the spatial framework for organizing information on species distribution and abundance within each
estuary. These salinity zones are tidal fresh (0.0 to 0.5%o), mixing (0.5 to 25.0%o), and seawater (25.0%o and
greater). The primary data developed for each species for each salinity zone include spatial and temporal
distributions and relative abundance by life stage (i.e., adult, spawning or mating adults, juvenile, larva, and egg).
In addition, a detailed estuarine life history summary is written for each species.

Additional information on this or other programs of NOAA's Strategic Environmental Assessments Division is
available from:
Strategic Environmental Assessments Division
Office of Ocean Resources Conservation and Assessment
National Oceanic and Atmospheric Administration
6001 Executive Blvd., Rm. 220
Rockville, Maryland 20852
FTS/Comm. (301) 443-0453/8921

Reports available from NOAA's Estuarine Living Marine Resources program include:

Monaco, M. E., T. E. Czapla, D. M. Nelson, and M. E. Pattilo. 1989. Distribution and abundance of fishes and invertebrates
in Texas estuaries. ELMR Rep. No. 3. Strategic Assessment Branch, NOS/NOAA, Rockville, MD, 107 p.

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

Bulger, A. J., B. P. Hayden, M. E. Monaco, D. M. Nelson, and M. G. McCormick-Ray. 1990. Aproposed estuarine classification:
analysis of species salinity ranges. ELMR Rep. No. 5. Strategic Assessment Branch, NOS/NOAA, Rockville, MD, 28 p.

Williams, C. W., D. M. Nelson, M. E. Monaco, L. C. Clements, S. L. Stone, L. R. Settle, C. lancu, and E. A. Irlandi. 1990.
Distribution and abundance of fishes and invertebrates in eastern Gulf of Mexico estuaries. ELMR Rep. No. 6. Strategic
Assessment Branch, NOS/NOAA, Rockville, MD, 105 p.

Czapla, T. E., M. E. Pattillo, D. M. Nelson, and M. E. Monaco. 1991. Distribution and abundance of fishes and invertebrates
in central Gulf of Mexico estuaries. ELMR Rep. No. 7. NOAA/NOS Strategic Environmental Assessments Division, Rockville,
MD, 82p.

Emmett, R. L., S. L. Stone, S. A. Hinton, and M. E. Monaco. 1991. Distribution and abundance of fishes and invertebrates
in west coast estuaries, Volume II: Species life history summaries. ELMR Rep. No. 8. NOAA/NOS Strategic Environmental
Assessments Division, Rockville, MD, 329 p.

D. M. Nelson, E. A. Irlandi, L. R. Settle, L. C. Coston-Clements, and M. E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in southeast estuaries. ELMR Rep. No. 9. NOAA/NOS Strategic Environmental Assessments Division,
Rockville, MD, 167 p.