National Oceanic and Atmospheric Administration
                     Environmental Research Laboratories
                     Seattle WA 981 15
EPA
United States
Environmental Protection
Agency
Office of Environmental Engineering
and Technology
Washington DC 20460
EPA 600 7 79-259
September 1979
            Research and Development
            Food Web
            Relationships of
            Northern Puget
            Sound and the
            Strait of Juan de Fuca
            A Synthesis of the
            Available Knowledge

            Interagency
            Energy/Environment
            R&D Program
            Report

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                RESEARCH REPORTING  SERIES

Research reports of the Office of Research and Development, U.S. Environmental
Protection Agency, have been grouped into nine series. These nine broad cate-
gories were established to facilitate further development and application of en-
vironmental technology. Elimination of traditional grouping was consciously
planned to foster technology transfer and a maximum interface in related fields.
The nine series are:

      1.  Environmental Health  Effects Research
     2.  Environmental Protection Technology
     3.  Ecological Research
     4.  Environmental Monitoring
     5.  Socioeconomic Environmental Studies
     6.  Scientific and Technical Assessment Reports (STAR)
     7.  Interagency Energy-Environment Research and Development
     8.  "Special" Reports
     9.  Miscellaneous Reports

This report has been assigned to the INTERAGENCY ENERGY-ENVIRONMENT
RESEARCH AND DEVELOPMENT series. Reports in this series result from the
effort funded  under the 17-agency Federal Energy/Environment Research and
Development Program. These studies relate to EPA's mission to protect the public
health and welfare from adverse effects of pollutants associated with energy sys-
tems. The goal of the Program is to assure the rapid development of domestic
energy supplies in an environmentally-compatible manner by providing the nec-
essary environmental  data and control technology. Investigations include analy-
ses of the transport of energy-related pollutants and their health and ecological
effects;  assessments  of, and development of, control technologies for energy
systems; and  integrated assessments of a wide range of energy-related environ-
mental issues.
This document is available to the public through the National Technical Informa-
tion Service, Springfield, Virginia 22161.

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                         FOOD WEB RELATIONSHIPS OF
           NORTHERN PUGET SOUND AND THE STRAIT OF JUAN DE FUCA

                   A Synthesis of the Available Knowledge
                                    by

          Charles A. Simenstad, Bruce S. Miller, Carl F. Nyblade,
                 Kathleen Thornburgh, and Lewis J. Bledsoe

                       Fisheries Research Institute
                           College of Fisheries
                         University of Washington
                        Seattle, Washington  98195
Prepared for the MESA (Marine Ecosystems Analysis) Puget Sound Project,
               Seattle, Washington, in partial fulfillment
                                    of

                 EPA Interagency Agreement No. D6-E693-EN
                        Program Element No. EHE625-A
                         This study was conducted
                          as part of the Federal
                      Interagency Energy/Environment
                     Research and Development Program
                               Prepared for
            OFFICE OF ENVIRONMENTAL ENGINEERING AND TECHNOLOGY
                    OFFICE OF RESEARCH AND DEVELOPMENT
                   U.S. ENVIRONMENTAL PROTECTION AGENCY
                         WASHINGTON, D.C.
                              September 1979

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     This work is the result of research sponsored by the Environmental
Protection Agency and administered by the Environmental Research Laboratories
of the National Oceanic and Atmospheric Administration.

     The Environmental Research Laboratories do not approve,  recommend,
or endorse any proprietary product or proprietary material mentioned in
this publication.  No reference shall be made to the Environmental Research
Laboratories or to this publication furnished by the Environmental Research
Laboratories in any advertising or sales promotion which would indicate or
imply that the Environmental Research Laboratories approve, recommend, or
endorse any proprietary product or proprietary material mentioned herein, or
which has as its purpose an intent to cause directly or indirectly the
advertised product to be used or purchased because of this Environmental
Research Laboratories publication.
                                     ii

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FOREWORD

     Substantially increased petroleum transfer and refining activities
are anticipated in the northern  Puget Sound and Strait of Juan de Fuca areas.
These activities will likely increase the chances of chronic and/or acute
oil inputs into the marine environment.   These areas are currently stressed
to only a limited degree by petroleum.  The study reported here was undertaken
to identify biologic means by which petroleum constituents may be transferred
from lower to higher trophic level populations and to identify those populations
and pre-predator links that are of critical importance to maintenance of major
biological communities.   Interruption of these critical links by loss of
important prey groups could drastically  change the composition and/or productivity
of higher trophic level populations.  The study was conducted by scientists at
the Fisheries Research Institute, University of Washington and involved primarily
a compilation of existing data.
                                   iii

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                            TABLE OF CONTENTS

I.    Introduction                                                        1
II.   Conclusions                                                         3
III.  Recommendations                                                     6
IV.   Materials and Methods                                               8
      IV-A.  Community Organization                                       8
      IV-B.  Documentation and Quantification  of  Food  Web Linkages       9
      IV-C.  Stomach Analyses                                            11
      IV-D.  Index of Relative Importance  (IRI)                          11
      IV-E.  Trophic Diversity                                           13
      IV-F.  Sources of Food Web and Community Data                      13
      IV-G.  Definitions                                                 14
      IV-H.  Place Names and Locations and Associated  Habitats           16

V.    Results and Discussion                                             18
      V-A.   Food Web Structure of Northern Puget  Sound
             and the Strait of Juan de Fuca                              18
      V-B.   Prey Assemblages of Major Importance  to
             Upper Trophic Levels                                        35
      V-C.   Relative Importance of Autotrophic Versus
             Heterotrophic Energy Bases to Nearshore Food Webs          39
      V-D.   Ecological Impact of Introduction or  Incorporation of
             Petroleum Hydrocarbons into Food  Weo  Structures            41
      V-E.   Comparison of Food Web Structures at  Existing or
             Potential Oil Terminal Sites and  an Evaluation of
             Their Relative Importance                                  45
      V-F.   Utilization of Empirical Food Web Data for
             Modeling Energy Flow in Marine Ecosystems                  48

      References                                                        49

Appendix A:  Algae and Invertebrates                                    55
Appendix B:  Fishes                                                     79
Appendix C:  Sea Birds and Shore Birds                                 214
Appendix D:  Marine Mammals                                            259
      D-l.   Cetaceans                                                 259
      D-2.   Pinnipeds                                                 278
Appendix E:  Seasonal Distribution and Abundance of Food Web Nodes
             and Number and Relative Importance of Food Web Linkages   291
Appendix F:  Mechanisms of Petroleum Hydrocarbon Influence
             on Food Web Structure                                     306
      F-l.   Effects Upon and Within Marine Organisms                  306
      F-2.   Indirect Subletnal Effects                                308
      F-3.   Uptake and Effects of Petroleum Hydrocarbons
             Transferred Via Food Web Linkages                         309
      F-4.   Ecological Effects of Petroleum Hydrocarbons              311
Appendix G:  Bioenergetic Model of Westcott Bay                        322
                                    iv

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                               LIST OF TABLES

   1.  Example computation of IRI values and percentages of total
       IRI from data illustrated in Fig. 1                                 12

   2.  Distribution and abundance of food web nodes for food webs
       characterizing nearshore habitats of north Puget Sound and
       the Strait of Juan de Fuca.                                         19

   3.  Number and relative importance of food web linkages to trophic
       levels characterizing nearshore habitats.                           20

   4.  General listing of species or functional groups which are of
       major importance to the upper trophic levels of the nearshore
       food webs.                                                          36

 B-l.  Representative nearshore fish assemblages and functional feeding
       groups of component species.                                        80

 B-2.  Prey composition of spiny dogfish reported by Jones and Geen
       (1977) (n=14,796).                                                  84

 B-3.  Prey composition for juvenile kelp greenling in neritic
       waters of Strait of Georgia (Barraclough 1967a).                  124

 B-4.  Prey composition (frequency of occurrence) of lingcod in
       southern California kelp beds (Quast 1968).                        128

 B-5.  Prey composition (frequency of occurrence) of Pacific staghorn
       sculpin in Anaheim Bay, California (Tasto 1975).                  154

 B-6.  Prey composition (numerical and gravimetric composition) of
       Pacific staghorn sculpin in Everett Bay, Washington (Conley 1977) 155

 B-7-  Prey composition (numerical composition) of tidepool sculpin
       at Port Renfrew, Vancouver Island, B.C.  (Nakamura 1971).          157

 B-8.  Prey composition (numerical composition) of fluffy sculpin at
       Port Renfrew, Vancouver Island, B.C. (Nakamura 1971).             161

 B-9.  Prey composition (freq. occurr. and grav. comp.) for general
       prey categories consumed by juvenile, subadult, and adult
       cabezon in central California (Connell 1953).                     162

B-10.  Prey composition (freq. occurr.) for specific prey categories
       consumed by juvenile, subadult, and adult cabezon in central
       California (Connell 1953).                                        163

B-ll.  Prey composition (freq. occurr.) of cabezon from southern
       California kelp beds (Quast 1968).                                165

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B-12.  Prey composition  (grav. comp.) of shiner perch  in  spring,
       summer, fall, and winter  in Anaheim  Bay, California
       (Weller 1975).                                                     176

B-13.  Prey composition  (freq. occurr.)  of  shiner perch in upper
       Newport Bay, California  (Bane  and Robinson 1970).                  178

B-14.  Prey composition  (freq. occurr.)  of  striped seaperch in
       Yaquina Bay, Oregon  (Gnose 1967).                                  180

B-15.  Prey composition  (freq. occurr.  and  grav.  comp.) of striped
       seaperch at  Santa Cruz and Santa Barbara (Alevizon 1975).         181

B-16.  Prey composition  (freq. occurr.)  of  pile perch 100-199 mm and
       200-299 mm long in southern California kelp beds (Quast 1968).     182

B-17.  Prey composition  (freq. occurr.)  of  high cockscomb at San
       Simeon, California  (Barton 1974).                                  186

B-18.  Prey composition  (freq. occurr.)  of  high cockscomb at Second
       Narrows, Burrard  Inlet, B.C.  (Pepper 1965).                        187

B-19.  Prey composition  (freq. occurr.)  of  black  prickleback at  San
       Simeon, California (Barton 1973).                                  189

B-20.  Prey composition  (freq. occurr.) of  rock prickleback  at San
       Simeon, California (Barton 1974).                                  192

 C-l.  Relative abundance of  marine and shore birds known  to  the
       northern Puget Sound and  Strait of Juan de Fuca area.              215

 C-2.  Functional feeding groups  and representative prey taxa of
       58 marine and shore birds  common to northern Puget  Sound and
       the Strait of Juan de  Fuca.                                        218

 C-3.  Percent composition of numbers of prey delivered to rhinocerus
       auklet nestlings on Protection Island (Manuwal 1977).              249

 D-l.  Cetaceans occurring in Washington State and British Columbia.       260

 D-2.  Functional feeding groups  and representative prey taxa of marine
       mammals known or suspected to occur in north Puget Sound and the
       Strait of Juan de Fuca.                                           262

 E-l.  Distribution and abundance of food web nodes of neritic habitats.  292

 E-2.  Number and relative importance of food web linkages to trophic
       levels of neritic habitats.                                       293

 E-3.  Distribution and abundance of food web nodes of rocky
       sublittoral habitats.                                             294
                                      VI

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 E-4.   Number and relative importance of food web linkages to trophic
       levels of rocky sublittoral habitats.                             295

 E-5.   Distribution and abundance of food web nodes of rocky and
       cobble littoral habitats.                                         296

 E-6.   Number and relative importance of food web linkages to trophic
       levels of rocky and cobble littoral habitats.                     297

 E-7.   Distribution and abundance of food web nodes of gravel-cobble
       shallow sublittoral habitats                                      298

 E-8.   Number and relative importance of food web linkages to trophic
       levels of gravel-cobble shallow sublittoral habitats.             300

 E-9.   Distribution and abundance of food web nodes of sand/eelgrass
       shallow sublittoral habitats.                                     302

E-10.   Number and relative importance of food web linkages to trophic
       levels of sand/eelgrass shallow sublittoral habitats.             303

E-ll.   Distribution and abundance of food web nodes of mud/eelgrass
       shallow sublittoral habitats.                                     304

E-12.   Number and relative importance of food web linkages to trophic
       levels of mud/eelgrass shallow sublittoral habitats.              305

 G-l.   Standing stock and primary production in coastal detritus-
       based system.                                                     324

 G-2.   Estimated primary energy sources (wet weights) in Westcott
       Bay, San Juan Island.                                             325

 G-3.   Seasonal growth rates and conversion efficiencies, Westcott
       Bay.                                                              329

 G-4.   Estimated annual production  (wet weights) of secondary
       consumers in Westcott Bay.                                        330

 G-5.   Seasonal changes in standing crop of secondary consumers at
       Westcott Bay.                                                     331
                                     vn

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                               LIST OF FIGURES

   1.   Example IRI (Index of Relative Importance) diagram.                12

   2.   Locations of DOE and MESA sampling sites from which nearshore
       community and food web data were obtained.                         17

   3.   Composite food web characteristic of neritic habitats.             21

   4.   Composite food web characteristic of sublittoral rocky/kelp bed
       habitats.                                                          24

   5.   Composite food web characteristic of rocky littoral
       habitats.                                                          27

   6.   Composite food web characteristic of cobble littoral habitats.     28

   7.   Composite food web characteristic of exposed gravel-cobble,
       shallow sublittoral habitats.                                      30

   8.   Composite food web characteristic of protected sand/eelgrass,
       shallow sublittoral habitats.                                      32

   9.   Composite food web characteristic of protected mud/eelgrass,
       shallow sublittoral habitats.                                      34

B-l to B-77.  IRI prey spectra.                                      86-209

 G-l.   Bioenergetic food web model of epibenthic  community of
       Westcott Bay.                                                     323

 G—2.   Estimated flows to detritus, grazed  and dissolved organic
       matter compartments from primary production  compartments.          327
                                    viii

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                            Acknowledgements

     We thank everyone who provided information, guidance, and criticism
for the prolonged synthesis process.

     Catherine Terry, Steve Ralph, Tom Crawford, Craig Staude, and
Theresa Clocksin spent considerable time digging into the obscure
literature on fish, bird, gammarid amphipod, and marine mammal food
habits, or helped diagram countless food webs.  Dan Moriarity and Susan
Oliver of Graysmarsh Wildlife Refuge, and Bob Jefferies and Richard
Parker of the Washington Department of Game's Skagit Wildlife Laboratory
provided data sets on the seasonal occurrence, distribution, and
abundance of shorebirds and waterfowl in northern Puget Sound and the
Strait of Juan de Fuca.  Dr. David Manuwal, University of Washington,
also provided expert criticism of the seabird and shorebord discussions.

     Bob Everitt and Cliff Fiscus of the NMFS Marine Mammal Lab at Sand
Point, Seattle, provided expert criticism of earlier drafts of the report
and a number of literature sources on marine mammals.  Ken Balcomb and
Rich Osborne of the Orca Survey, and Mike Bigg of Pacific Biological
Station, Nanaimo, B.C., supplied valuable preliminary, unpublished data
on the abundance and distribution of orcas in the region.  Jeff Cross's
data and knowledge of the littoral and kelp bed fish communities in the
Strait of Juan de Fuca contributed precise information on these little-
documented habitats.  Craig Wingert provided invaluable data and assistance
with the numerical classification of nearshore fish assemblages.  Kim
Marshall provided the exceptionally fine renditions of the composite food
web structures.

     To  the  biologists, technicians, and students who suffered the
elements to haul beach seines and townets around northern Puget Sound and
the Strait of Juan de Fuca, or spent hours on end turning fish stomachs
inside out, we are indeed indebted.
                                    IX

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                            I.  INTRODUCTION

     A principal concern about the long-term effects of  oil  spills  in the
marine environment is the fate and effects of petroleum  hydrocarbons
transferred through food web pathways and of the  disruption,  through toxic
effects, of important food web linkages, causing  significant  alterations
in the overall community structure of the biota.   In 1977  MESA contracted the
University of Washington's Fisheries Research Institute  to document the
structure of the marine food webs of northern Puget Sound  and the Strait
of Juan de Fuca using data from the literature, from unpublished  sources,
and from the ongoing MESA studies in the Strait of Juan  de Fuca.   This
report is intended to synthesize all the information available on the
food organisms, feeding behavior, and trophic  (predator-prey) relationships
of marine organisms and to discuss the  possible effects  of an oil spill on
these ecological relationships.

     Long before completion of the Alaska pipeline, it became apparent that
a  sizable volume of crude oil from Alaska:s North Slope  would eventually
have to reach  the high-demand areas of  the continental United States,
especially  the midwestern region and the northern tier states.  At  that
time, only  Long Beach,  California, and  Puget Sound, Washington, had deep-
water ports  capable of  handling the deep-draft supertankers  (125,000 DWT)
that would  transport Alaskan crude oil.  Notification of eventual
diminution  of  crude oil shipment from Canada to the existing  refineries
at Cherry Point in north Puget Sound further intensified political
pressure  to establish a western terminus in north Puget  Sound.

     Political pressure began mounting  in 1976 to prevent  establishment
of an oil port in northern Puget Sound.  Opponents to such a  port cited
the navigational hazards of traversing  the narrow channels through  the
San Juan  Islands and the risk of oil spill damage to the region's natural
resources,  and lobbied  for location of  a facility west of  Port Angeles
along the less hazardous Strait of Juan de Fuca.   Political pressure
culminated  in  a Washington state law, the Tug Escort Law,  prohibiting
tankers greater than 125,000 DWT from operating in north Puget Sound east
of Port Angeles, and requiring tug escorts of tankers larger  than
50,000  DWT.  This law was eventually modified by  the U.S.  Supreme Court
 (ARCO vs. Ray) on 6 March 1978.  Since  then, however, the  U.S. Coast
Guard has declared temporary navigation rules prohibiting  tankers greater
than 125,000 DWT from entering waters east of Port Angeles.   By that
time, the State Legislature had enacted a law prohibiting  construction of
an oil  port and pipeline terminus east  of Port Angeles,  which was vetoed
by the  governor.  The controversy, however, moved to the U.S.  Senate where
Washington  Senator Warren Magnuson won  approval of a similarly constructed
amendment  (Public Law 95-136) to the Marine Mammal Act which  was  enacted
on 17 October  1977.

     Facing an almost complete lack of  economic and environmental
information about the ncrf:-~ Puget Sound region, in 1974  Washington  State
initiated the north Puget Sound baseline study through the Department of
Ecology which was designed to evaluate  the oceanographic,  biological,  and

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economic resources of the region (Gardner 1978).  The State also implemented
studies of potential offshore and inshore oil transshipment systems and
sites by the Washington Oceanographic Institute in 1974 (Ocean. Comm. Wa.
1974, 1975).

     Recognizing the lack of knowledge of the biological communities and
oceanographic conditions of the Strait of Juan de Fuca, the National
Oceanic and Atmospheric Administration's Marine Ecosystem Analysis (MESA)
Program initiated baseline studies in 1975 which sought to document the
character and dynamics of the biological communities, the oceanographic
conditions, and the existing pollutant levels, as well as to model probable
oil spill trajectories in the eastern region of the strait.

     Although subtidal marine environments have been shown to be susceptible
to the effects of petroleum hydrocarbons (North, et al., 1964; Blumer,
et al., 1971; Kolpack, et al., 1971; Sanborn 1977), it is the nearshore
(littoral and shallow sublittoral) and surface water (neritic) habitats
that are the most available to pollutant introductions and effects.
Petroleum pollution has generally had the most dramatic impact on these
environments especially in estuaries (Clark and Finley 1977).  Thus, the
MESA biological studies, like the earlier Department of Ecology baseline
studies, focused on the biological communities of the nearshore environ-
ment.  The results of the first three years' studies were reported in
Simenstad, et al. (1977), Cross, et al. (1978), Nyblade (1978, 1979),
Webber (1979), and Everitt, et al. (1979).

     This report resulted from the need to synthesize existing knowledge
of the structure of food webs in nearshore marine habitats of northern
Puget Sound and the Strait of Juan de Fuca, in order to identify the
potential transfer processes of petroleum hydrocarbons through the marine
ecosystem of the region.  The objectives of this investigation were to:
(1) identify the food web structures of biological communities of neritic,
shallow sublittoral, and littoral habitats; (2) document seasonal, site,
and regional variability in food web structure; (3) identify important
predator-prey linkages that could be disrupted by a pollutant, and the
potential consequences of disruption to the community; (4) identify the
main prey organism groups utilized by economically or ecologically
important predators; and (5) identify food chains having the greatest
potential for transferring pollutants to higher trophic levels.

     In addition to nearshore habitats, we focused on sites of existing
or proposed oil terminals—specifically, Cherry Point and Fidalgo Bay
(March Point) in northern Puget Sound, Burrows Bay at the eastern end of
the Strait of Juan de Fuca, and Port Angeles.

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                            II.  CONCLUSIONS

     In general, existing information on the structure of  food webs  in
nearshore marine habitats of north Puget Sound and the Strait of  Juan de
Fuca shows that food web complexity or "connectivity" increases with
decreasing exposure, decreasing sediment particle size,  and  increasing
deposition of algal and vegetative detritus.  Greater food web diversity
implies greater energy flow, although the  efficiency of  energy transfer
may be less.  Under some circumstances, the more diverse the food web,
the less liable the overall community structure is to change dramatically
as a result of removal or alteration of linkages.

     Except in neritic food webs, detritivores are the major prey organisms
leading to higher trophic levels in the region's nearshore ecosystem.
Direct herbivory by suspension feeding and grazing on macroalgae  is  less
important.  The most important detritivores are epibenthic organisms such
as gammarid amphipods, harpacticoid copepods, flabelliferan  isopods,
tanaids, mysids, and polychaete annelids,  which are  the  principal prey of
nearshore carnivores, fishes and shorebirds and seabirds.  Neritic
phytoplankton-zooplankton-planktivorous fish food webs effect more
rapid, direct transfer of organic matter among trophic levels, whereas
detritus-based  food webs are typically more complex and  effect slower
transfer among  trophic levels.  Additionally, heterotrophic  processing of
autotrophically produced algal and vegetative carbon is  a  critical
mechanism limiting  the productivity and transfer rates of  detritus-based
food webs.

     The effects of petroleum  hydrocarbons would appear  to be more
pronounced  and  long-term on food webs based on detritus  processing than
on food webs  based  on herbivory.  The incorporation of hydrocarbons  into
 fine,  unconsolidated sediments and detritus pools of contained embayments
 can result  in prolonged recycling of persistent hydrocarbon  components
 through  the detritus-decomposer-detritivore food web, continually providing
 contaminated  epibenthic prey to the upper  trophic levels,  including  species
utilized by man.

      Except for the neritic habitats, the  majority of nearshore habitats
 in north Puget  Sound and the Strait of Juan de Fuca have food web structures
which  could be  altered for years by the introduction of  petroleum hydro-
 carbons.   The contained embayments, eelgrass beds, and saltmarshes of
north  Puget Sound,  however, have the potential to suffer longer from
perturbations than  the more exposed environs of the Strait of Juan de
 Fuca.   The  sand/eelgrass and mud/eelgrass  habitats in the  eastern Strait
of Juan  de  Fuca are also sensitive to petroleum effects.   Of the  four
areas  of  existing or planned oil terminal  sites, March Point near Anacor-
tes may have  the greatest sensitivity to ecological disruption by oil
spill  because of the diverse communities and complex food  webs character-
izing  the mud/eelgrass habitats which predominate there.   Exposed cobble
habitats at the other three sites have less complex food webs, and there
is less chance  of incorporation of unweathered oil into  the  more  consolid-
ated sediments.  In these latter three areas, Cherry Point has the most

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diverse food web, followed by Port Angeles and Burrows Bay.  The food web
structure of the Port Angeles vicinity, however, showed the highest
connectivity (average number of linkages per node) of any of the areas,
including Fidalgo Bay.

     In almost all instances, food web structures were most diverse—
greatest number of nodes as well as linkages—during spring and summer.
During these seasons diversity of food web was most pronounced in the
neritic and some of the exposed cobble-gravel habitats where a multitude
of marine organisms at several trophic levels spend their early life
history before settling into benthic habitats or moving into the shallow
sublittoral region.  The larvae and juveniles of many economically
important species, such as Pacific salmon, Pacific herring, and Dungeness
crab, are also particularly prominent (and susceptible to toxic pollutants)
during these seasons.  Juveniles of many species, especially fishes,
increase the diversity of food web structures characterizing the more
protected habitats in spring and summer, and seabirds and shorebirds are
responsible for maintenance of similarly diverse food webs in fall and
winter.  Because of the importance of juvenile recruitment for sustaining
the adult populations and the greater vulnerability and sensitivity of
juvenile forms to toxic petroleum hydrocarbons, food webs in spring and
early summer have the greatest potential for disruption by spilled
petroleum and its incorporation into the nearshore environments of the
region.

     A number of taxa or assemblages of organisms were identified as
being critical to upper trophic levels, either because they provide food
resources for important consumer organisms, or because they convert or
transfer organic matter to trophic levels where it is available to higher
level consumers, e.g., detritus processors.  The two most obvious groups
were calanoid copepods and gammarid amphipods.  Calanoids are the key
herbivores in the neritic food webs and the principal prey of important
consumers such as Pacific herring, Pacific sand lance, and juvenile Pacific
salmon, which in turn are the main secondary consumers utilized by higher
level carnivores in that habitat.  In the shallow sublittoral zones of the
nearshore environment, gammarid amphipods head the list of detritivorous
crustaceans which are the main food of nearshore consumers.  Their
conversion of detrital carbon into food biomass for nearshore fishes and
shorebirds provides the principal structure for almost all of the near-
shore food webs.  Other detritivorous crustaceans such as harpacticoid
copepods, flabelliferan isopods, cumaceans, mysids, and shrimps also
contribute significantly to important upper-level consumers.

     As a great number of the nearshore food webs are apparently supported
by detritivores, the sources of detrital carbon are also of ultimate
consequence in sustaining a productive biotic community.  Although there
have been no studies to estimate the relative contributions to the total
annual detritus by kelps and other macroalgae, microalgae, eelgrass,
saltmarsh plants, and riverine inputs, it would appear that eelgrass and
kelps are the main sources in north Puget Sound and the Strait of Juan de
Fuca.  Riverine inputs, principally from the Fraser and Skagit rivers,

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may be more significant in the neritic habitats in north Puget Sound.
The physical and biological processes of breaking down and conditioning
these large organic particles into particles small enough to be used by
the small detritivorous crustaceans have not yet been examined.  They
appear to be different in different habitats.  Exposed habitats such as
rocky  and cobble littoral and gravel-cobble shallow sublittoral
habitats may act as giant grinders, physically reducing  the kelp plants
to smaller particles.  Contained  sand/eelgrass and mud/eelgrass embayment
habitats may act as detritus traps, where  a pool of eelgrass  and other
organic particles decomposes primarily  through microbial activity.   In
both cases the critical step of conversion to usable biomass  involves
microbial colonization and reduction  (making more  surface area available
for colonization) of detritus particles.

     While a number of specific food  webs  or prey-predator linkages were
identified as prominent or important  in the region's nearshore marine
communities, several stand out because  they form the trophic  base of
economically important species.   The  phytoplankton - calanoid copepod
food web in neritic habitats forms the  resource base for  juvenile pink
and chinook salmon and Pacific herring,  all of which as adults support
high-value fisheries.  Calanoid copepods are also used by secondary
consumers such as fish and crab larvae which are in turn consumed by
juvenile coho salmon.  Thus, for  the  critical period of their juvenile
residence in Puget Sound during migration  into the Northeast Pacific
Ocean, three of the five species  of Pacific salmon of this region  are
dependent upon this short, simple, but substantial neritic food web.

     Pacific herring and the other major neritic secondary consumer,
Pacific sand lance, subsequently  form the principal food  organisms of many
recreationally or commercially important fish species  (lingcod, rockfishes)
of the rocky/kelp bed habitats,  of resident Puget  Sound  salmon ("black-
mouth," chinook), and aesthetically valued carnivores  such as  orca,  Dall
porpoise, and alcid seabirds.  In the shallow sublittoral habitats,  the
detritus-detritivore, gammarid amphipod, harpacticoid  copepod  food web
supports juvenile chum salmon and a number of  juvenile flatfish  (English
sole, rock sole) which are commercially exploited as adults.   The impor-
tance of the shallow sublittoral habitats,  especially  eelgrass, as
nurseries and juvenile rearing environments for  fish and  invertebrates
economically important in the Puget Sound region cannot be overstated.

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                         III.  RECOMMENDATIONS

     Many lacunae in our data and in our understanding of nearshore
community and food web structure were discovered during this analysis of
the northern Puget Sound and Strait of Juan de Fuca region; some of them
were glaring.  For example, it is most unfortunate that there are no
quantitative data on the species composition and food web relationships
of the fish communities of the extensive rocky/kelp bed habitat in this
region.  Having no data comparable to those obtained during the DOE
studies in northern Puget Sound severely inhibited our ability to
evaluate the structure of the food webs characterizing the outer strait.
The only available data were from Barkley Sound along the northwest
coast of Vancouver Island and cannot be considered representative of the
outer strait.  Many of the fishes composing the communities  of this
habitat are important recreational or commercial species (lingcod, rock-
fish, greenling).  SCUBA-diver transect studies such as conducted by
Miller, et al. (1977), and Moulton (1977) are definitely needed.

     The role of pelagic plankton as the base of the neritic food webs is
obvious; the structure and dynamics of nearshore pelagic zooplankton
communities are not obvious.  Existing documentation of these communities
in the region, including the MESA studies (Chester, et al., 1977), are
oceanographic examinations of mid-channel, deep-water stations.  Comparison
of fish assemblages at such stations with nearshore assemblages suggests
that the nearshore environs, and especially contained embayments, harbor
much higher densities of neritic fishes and decapod larvae.  There is an
obvious need to determine the relationship between offshore pelagic
zooplankton and those populations found in the nearshore neritic environ-
ment, whether the latter are an advected component of the former or a
unique community characteristic of nearshore habitats.

     The lack of data on the prey organisms of marine mammals and seabirds
and shorebirds inhabiting north Puget Sound and the Strait of Juan de Fuca
is apparent from reading Appendices C and D.  Considering their important
roles as secondary and tertiary carnivores and their ecological and
recreational importance, quantitative documentation of prey composition
and consumption rates are necessary before the magnitude of their predation
upon lower trophic levels and their dependence on specific food web
linkages can be properly assessed.

     The importance of detritus in nearshore food webs is apparent, but
studies of the decomposition process and the interaction between pollutants
and organic detritus particles are completely lacking.  We need a much
better understanding of the mechanisms, rates, and rate-limiting factors
regulating the microbial, chemical, and physical processing that make
detritus accessible to detritivores and to incorporation into the food web.
And although it has been established that detritus particles often adsorb
hydrocarbons, the conditions dictating the processing and incorporation
of the hydrocarbons by microflora and the transfer to detritivores are
unknown.  This crucial process must be examined before we can hope to
understand the pathway of pollutants through detritus-based food webs.

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     The question still remains of whether a more diverse or connected food
web is necessarily more stable—i.e., less prone to be severely altered by
removal of a portion of its nodes and linkages—than a simpler food web.
Efforts to answer this question have been only theoretical or through
simulation modeling.  As yet, no one has experimentally perturbed a
documented food web in the laboratory or in the field by introducing a
toxic substance and following the acute and sublethal effects through time.
Similarly, although there have been a few laboratory experiments to
examine the transfer of petroleum hydrocarbons between trophic levels,
there is a need for more detailed, multi-trophic level experiments which
include documentation of sublethal effects on the growth, behavior, and
reproduction of secondary and tertiary consumers.

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                       IV.  MATERIALS AND METHODS

     Synthesis of known food web relationships was made with information
gathered through a combination of analytical, laboratory, and field sampling
tasks, including (1) comprehensive quantitative analysis of existing raw
data residing in NOAA/MESA and our own data bases; (2) review of published
and unpublished literature and inclusion of appropriate data in the data
base for analysis; (3) analysis of hitherto unprocessed fish stomach
specimens from the north Puget Sound region; (3) further taxonomic and
size analysis of representative prey retained from previous processing of
fish stomach contents; (4) quarterly sampling of nearshore demersal and
neritic fishes along the eastern end of the Strait of Juan de Fuca
(Burrows Bay to west Whidbey Island), for the purpose of collecting stomach
samples representative of that area's nearshore fish communities; and
(5) interviews with experts knowledgeable in the food habits of the region's
marine invertebrates and fishes, seabirds and shorebirds, and marine
mammals.

     The overall objective was to formulate conceptual food web models
which (1) documented the major species or taxa involved in carbon flow
through the region's nearshore ecosystems;  (2) illustrated regional,
habitat, and seasonal variations in food web structure and energy flow, and
(3) provided some semiquantitative evaluation of the quality (frequency of
occurrence, etc.) and quantity (proportion of prey biomass transferred) of
the food web linkages between prey and predators.  The food web models were
then summarized as to their complexity (i.e., number of species or taxa
nodes; and number of primary, secondary, tertiary, and incidental linkages
between nodes) according to trophic compartments (i.e., detritus processors,
herbivores, planktivores, benthivores, and omnivores) and compared according
to the objectives described at the end of Introduction.

     Two steps were required in formulating the food web structures:
(1) Definition of the principal species—from planktonic and macrophytic
algae to marine mammals—composing the region's biotic communities, and
(2) synthesis of predator-prey data on these organisms.

                      IV-A.  Community Organization

     Definition of the component species varied according to the diversity
of organisms and the extent of quantitative data on their seasonal abundance
and distribution.  Because of the scattered and often meager survey data for
marine birds and mammals, only subjective and often conjectural definitions
of their community composition were possible.  On the other hand, the data
base for the region's marine benthic invertebrates has become voluminous,
and unfortunately  has not been subjected to any detailed analyses such as
recurrent group analysis (Fager 1957) or numerical classification (Clifford
and Stephenson 1975; Smith 1976).  In these three cases, therefore, all raw
data and relative indices of frequency of occurrence and abundance have
been treated subjectively to provide the community descriptions.

     The extensive, uniform data base for nearshore fishes provided by the
DOE studies in north Puget Sound (Miller, et al., 1977) and the NOAA/MESA
investigations in the Strait of Juan de Fuca (Simenstad, et al., 1977;

                                    8

-------
Cross, et al., 1978), however, was adequate for such analysis.  Distribu-
tional analysis of many of these data was already in process by Wingert
(unpubl. Ph.D. Thesis, Univ. Washington), who has applied hierarchical
numerical classification techniques to the beach seine and  townet data.
This technique was further applied to the SCUBA transect data  gathered in
the rocky/kelp bed habitat by Moulton (1977) as a part of the  DOE studies
and to the littoral fish collections provided by the NOAA/MESA program
(Cross, et al., 1978).  While this scheme cannot be validly applied  to the
combined data sets as a whole, it can differentiate species groups or
spatial patterns within the data subsets described by the different
collection techniques.  Thus, distinct species groups have  been identified
within the neritic environment sampled by the townet, shallow  sublittoral
habitats sampled by beach seine and SCUBA transect methods, and the  littoral
zone as sampled by using fish-specific narcotics.

     The classification analysis utilized was of the agglomerative-polythetic
variety,* applied to  numerical fish catch data by species.  Although the
addition of  life history stage designations was desirable,  it  was felt that
the  food habits data  could not be meaningfully divided into that many subsets.
Before  calculation of inter-entity distances,* the data were scaled  using  a
square-root  transformation.   Both normal or site classification and  inverse
classification were used, normal analysis for indication of species-site
associations and inverse analysis for interspecific associations.  The data
matrix  was  further standardized using a species mean standardization,

                                   .,  nst
                              ,,. . , 1  r
                              Xlj /—  I
                                  nz k=t

 (where  nz = number of non-zero elements in row i)for normal analysis and a
 species maximum standardization, Xij/Xmax (where Xmax = largest element  in
 row i)  for  inverse analysis.  A flexible fusion strategy (adjusted mean  of
 Dhi and Dhj :  Dhk = aDhi + aDhj + BDij where a = l/2(l-g) and  6 was  set  at
 -0.25)  (Lance and Williams 1967) was utilized for the process  of clustering
 entities and groups of  entities together.

      Wingert's classification analyses were performed using the Ecological
 Analysis Package  (EAP,  R.W.  Smith, Allan Hancock Foundation) system  which
 was installed at  the  University of  Southern California's IBM computer.
 The analyses of Moulton1s  (1977) SCUBA transect data and the MESA tidepool
 fish collections  (Cross,  et  al., 1978) were made utilizing  program CLUSTER
 an interactive clustering  program developed at Oregon State University and'
 adapted to  the University  of Washington's CDC 6400 computer.

       IV-B.  Documentation and  Quantification of Food

      As the quality  of  data available  for different  trophic levels is usuall
 unequal, an analysis of the structure  and functional feeding relationships


      *See Clifford  and  Stephenson  (1975)  and  Smith  (1976) for  explanation
 of these approaches  to  numerical  classification.

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linking identifiable communities of organisms was considered more
appropriate than, say, a bioenergetic analysis, even though the ultimate
documentation of food web dynamics requires quantification of the transfer
rates, efficiencies, and partitioning of carbon from primary producers
through tertiary carnivores.

     The process of energy acquisition by marine organisms can be
characterized by three quantitative parameters which reflect both the
trophic contribution by the prey organism to the predator and the energy
expended by the predator to capture and consume the prey.  These parameters
are (1) the frequency of occurrence of each prey organism or functional prey
group in the predator's diet, and (2) the percentage of the total number and
(3) the percentage of the total ingested biomass contributed by each prey.
Together they are important indicators of both the trophic (energy gained)
and the behavioral (energy expended) processes which characterize predation.
The simultaneous measurement of these three variables provider the most practical
measurement of importance of food web linkages.  As is described later in
the section on modeling energy flow in marine ecosystems, all three
variables are necessary to predictably quantify transfer rates between food
web nodes.

     It is characteristically uncommon, however, to find quantitative food
habits or predation data in the literature which include all three
variables.  Often it is impossible to obtain all three, especially when
it is desirable or necessary to obtain the data by observation or without
killing the predator.  Even if it is possible to obtain the stomach contents
intact, the food items may be unidentifiable because of digestion, or
uncountable (e.g., algae consumed by grazers), or of questionable trophic
importance (e.g., rocks, bivalve shells, algae, matchsticks).  Any biologist
who has examined the stomach contents of marine predators will attest to the
difficulty of explaining the significance of many food items, and how and
why the predators consume them.

     The optimal approach is a bioenergetic one including measurements of
the caloric content of each food group, the assimilation efficiencies
associated with each, and the energy expended to obtain them.  Applications
of the bioenergetic approach on the scale of a food web are rare, however.

     When all three variables characterizing food web linkages were
available, as in the DOE- and MESA-supported fish stomach analysis, they
were combined into an Index of Relative Importance (I.R.I.) after Pinkas,
et al. (1971), and Cailliet (1977).  This index arbitrarily attaches equal
weighting to the three variables and is expressed as the area occupied by
each prey group plotted on a three-axis graph  (see following section
describing IRI in detail).

     Previous species accounts of marine mammal food habits and feeding
behavior are usually not detailed enough to generate quantitative representa-
tions of a species' trophic link to its prey species.  In many cases the
probable prey spectrum required inference from data sources originating
outside the area of interest, such as the outer coasts of Washington and
British Columbia and the Gulf of Alaska, but there is always an implied
error associated with the different prey assemblages characterizing these

                                    10

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regions.  In these instances, Puget Sound organisms which are  functionally
equivalent to the recorded prey in another  region  are  considered  probable
prey organisms.

                          IV-C.  Stomach  Analyses

     Whole specimens or  intact  stomach samples of  economically important
fishes retained from the west Whidbey  Island - Burrows Bay  collections were
examined according to a  systematic, standard procedure (Terry 1977) which
identifies the numerical and gravimetric composition of prey organisms, the
stage of digestion of the contents, and  the degree of stomach fullness.  In
the laboratory, the stomach samples were removed  from the preservative (10%
buffered formalin), or   from the  preserved  whole  fish, and  soaked in cold
water for at least two or three hours  before examination.   The stomach was
then identified according to information on the label and then processed.
Processing involved taking a total  (damp) weight  to nearest 0.1 g, and
removing the contents from the  stomach and  weighing the empty stomach to
obtain  the total  stomach contents weight by subtraction.   Subjective
numerical evaluations of the stomach  condition or  degree  of fullness—
scaled  from 1  (empty) to 7 (distended)—and stage  of  digestion—scaled from
1  (all  digested)  to 5  (no digestion)—were  made at this time.   The stomach
contents were  then sorted and  identified as far as was practicable, and the
sorted  organisms  were counted  and a total  (damp) weight of  each taxon
obtained  to nearest 0.001 g.   If  a sorted taxon was represented by too many
individuals to count, the number  was  estimated using  a random grid-counting
procedure.

                 IV-D.   Index of Relative Importance (IRI)

      When possible,  the  relative  importance of food web linkages  has been
 represented as the percentage  of  the  total  IRI contributing to the total
 prey spectrum  of  a predator.   Though  this was possible for  most of the
 nearshore fish data,  insufficient data prevented an  IRI assessment of the
 other biological  groups.  In these cases, the percentage  of total biomass
 was considered the most  important measure of trophic  importance;  the
 percentage of  total  prey abundance was considered  second  in the absence of
 biomass data;  frequency  of occurrence  data  were considered  only in the
 absence of the other  two measures.  If two  measures were  given they were
 both considered.

      The three-axis  IRI  graph  (Fig. 1) illustrates frequency of occurrence
 (the proportion of  stomachs  containing a specific  prey organism)  plotted
 sequentially  on the  horizontal  axis.   Percentage of  total abundance (number
 of prey)  is  plotted  above the  horizontal axis.  Percentage  of  total weight
 of prey is plotted below the horizontal  axis.   All prey groups, including
 those which  had to be assigned  to a broad taxonomic  level (family,  order,
 class), have  been arranged from left  to  right by decreasing frequency of'
 occurrence.   Prey taxa  in differing stages  of digestion (e.g.,  partly
 digested shrimp,  "Natantia unidentified," as opposed  to family, "Pandalidae "
 or species,  "Pandalus borealis")  were  graphed separately.

      The IRI  value was  computed as follows:
                                     11

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         INDEX OF RELflTIVE  IMPORTflNCE (I.R.I.)  OlflORFIfl
UJ
U
oo
cr
CO
O
Q_
H
o
CJ
o
Q_
5

5

CD

Z
O
to
O
Q_
z:
o
O
     lOOr
      80
      60
      40
      20
      20
      40
      60
      80
     100
                 20       40       60       80      100      120



                            PERCENT FREQUENCY  OF OCCURRENCE
                                                                     140
160
     Fig.  1.   Example of  Index of Relative Importance (IRI)  diagram.
     Table 1.
               Example  computation of IRI values and percentages

               of  total  IRI from data illustrated in Fig.  1.
Prey
category
1
2
3
4
5
6
7
8
9
% Freq. of
occurrence
44.44
33.33
11.11
11.11
11.11
11.11
11.11
11.11
11.11
7. Numerical
composition
33.33
16.67
3.33
6.67
13.33
16.67
3.33
3.33
3.33
% Gravimetric
composition
0.21
37.71
0.17
1.43
1.37
12.07
0.06
32.14
14.85
Prey IRI
1490.8
1812.5
38.9
90.0
163.4
319.3
37.7
394.1
202.0
% Total
IRI
32.78
39.85
0.86
1.98
3.59
7.02
0.83
8.66
4.44
                                     12

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        IRI  =  % Frequency of
                  occurrence.
       [% Numerical     +  % Gravimetric
         composition.        composition.I
and is equivalent to the area encompassed by the bar for each prey
category i composing the IRI diagrams.  In order to compare the IRI values
between prey spectra with different sample sizes, the overall importance
of general prey taxa (e.g., all shrimp added together, including "Natantia
unidentified" and those identified to family and species) has been discussed
as a percentage of the summed total IRI values for individual spectra.
Table 1 is an example of the IRI values and percentages of total IRI
generated from the data diagrammed in Fig. 1.  The advantage of the IRI
value is that the more representative prey are not dominated by numerically
rare but high biomass prey (e.g., preyg, Fig. 1), by infrequently occurring
but abundant or high biomass (when eaten) taxa, or by numerically abundant
or frequently occurring taxa which contribute little in the way of biomass
(e.g. , preyj, Fig. 1).

                        IV-E.  Trophic Diversity

     Three quantitative indices of the numerical and biomass composition
of predator diets are used to describe trophic diversity (see Pielou 1975) :

      (1)  Percent dominance index:

                          % Dominance  =  Z(p.)2

where p.'s are ratios of the number or biomass of prey i to the total prey
abundance or biomass.

      (2)  Shannon-Wiener diversity index:

                                   s
H
where p.'s are the same as above.
                                      (p  In  p )
                                               X
      (3)  Evenness index:

                          e  =  H/lnS

where H = mean H, S= number of species, and lns= Hmax.

              IV-F.  Sources of Food Web and Community Data

     The amount of quantitative data on nearshore marine community structure
in north Puget Sound and the Strait of Juan de Fuca has been greatly
increased by DOE and NOAA/MESA baseline studies.  Quantitative food web data
are generally restricted to fishes, however.  Trophic data for the remaining
taxa usually are for other temperate ocean regions or for related spe'cies.

                                    13

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The  following  sources have  provided  the  data  base  for  the  region.
Algae
 (Qualitative)
 Invertebrates
 (Semi-
 quantitative)

 Fishes
 (Quantitative)
Seabirds and
shorebirds
(Semi-
 quantitative)
Marine mammals
(Qualitative)
   Data existing for
 north Puget Sound and
Strait of Juan de Fuca*

DOE and MESA baseline studies
(Nyblade 1977,1978; Webber
1977,1979); other Puget
Sound studies

DOE and MESA baseline studies
(Nyblade 1977,1978;
Webber 1977,1979)

DOE and MESA baseline studies
(Miller, et al. , 1977;
Simenstad, et al., 1977;
Cross, et al., 1978; Moulton
1977); other Puget Sound
studies (Simenstad and Kinney
1978; Fresh, et al., 1978;
Fresh 1979)

Natl. Wildl. Fed. study
(Manuwal 1977); other studies
(Richardson 1961; Wilson 1977;
Hartwick 1973; Salo 1975)
MESA baseline studies
(Everitt, et al., in
press; Bigg 1969;
Manzer and Cowan 1956;
Pike and MacAskie 1969;
Scheffer and Sperry 1931;
Scheffer and Slipp 1948)

           IV-G.  Definitions
Unpublished  data  and
sources  of expertise**

T. Mumford,  DNR
C. Nyblade,  UW
R. Thorn, UW
C. Nyblade, UW
B. Webber, WWU
B. Miller, UW
C. Simenstad, UW
J. Cross, UW
S. Borton, Seattle
  Aquarium
D. Manuwal, UW
S. Spiech, UW
B. Jeffries, Wn. Dept. Game
R. Parker, Wn. Dept. Game
T. Wahl, Bellingham

S. Rice, NMFS-MML
C. Fiscus, NMFS-MML
B. Everitt, NMFS-MML
T. Newby, NMFS-MML
K. Balcomb, Orca Survey
R. Osborne, Orca Survey
M. Bigg, Dept. Environ. Can.
     The following definitions are of terms and abbreviations used in this
report.
     *See pertinent appendices for references associated with each biotic
group.
    **DNR = Washington Department of Natural Resources
      UW = University of Washington
      WWU = Western Washington University
      NMFS-MML = National Marine Fisheries Service, Marine Mammal Laboratory
                                    14

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 Assemblage:  A restricted group of taxa or organisms which are found
      together.

 Autotrophic:  Self-nourishing; denoting those organisms capable of
     constructing organic matter from inorganic matter.

 Benthivore:  Organism which feeds on benthic organisms.

 Carnivore:  Organism which feeds on other organisms.

 Community:  The aggregation of organisms, plants and animals, within  a
      specified area which are interrelated in some manner.

 Consumers:  Heterotrophic organisms, chiefly animals, which  ingest  other
      organisms or particulate organic matter.

 Demersal:  Living on or near the bottom.

 Deposit feeder:  Organism, typically benthic, which is either somewhat
      selective or almost completely unselective in feeding;  includes
      organisms which sweep the surface or use ciliary tracts along
      extensile tentacles.

Detritivore:  Organism which utilizes detritus and/or its associated
     microflora for food.

Detritus:  Finely divided sinkable material of organic or inorganic origin
     which is suspended in the water.

 DOE:  Washington State Department of Ecology.

Entrapment carnivore:  Organism which, by using tentacles or mucous webs,
     entraps other organisms.

Epibenthic:  Associated primarily with the surface of the bottom but also
     with the water column directly above the bottom.

 Facultative feeder:  An organism which is not constrained to feeding on
      one general type of plant or animal but may feed on organisms from
      several trophic levels.

 Filter feeder:  Carnivore which feeds by engulfing large numbers of prey
      organisms as they swim through the water.   The filtering apparatus
      (such as gill rakers) retains the prey but lets the water pass out
      of the mouth.

 Food  web:  The network of organisms,  each of which provides  food
      or more organisms in the same or higher trophic level.          °ne

 Food  web linkage:  The trophic connection between food web nodes

 Food  web node:  Species, taxon, or functional feeding  group  constitut- •
      a unique prey or predator compartment in a food web.              ng

 Habitat:  The total of environmental conditions of a specific place th
      is occupied by an organism, a population,  or a community         at
                                    15

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Herbivore:  Organism which feeds on plant material.

Heterotrophic:   Dependent on organic matter for food.

IRI:  Index of Relative Importance (see Section IV-D).

Littoral zone:   The zone between the high and low water marks—
     i.e., the intertidal zone.

MESA:  NOAA's Marine Ecosystem Analysis Program.

Neritic zone:  Shallow surface water zone extending from the high-tide
     mark to the edge of the continental shelf.  Neuston nets, surface
     trawls, and townets were assumed to sample neritic organisms.

Obligate feeder:  Organism constrained by morphology or behavior to feeding
     on one general type of plant or animal.

Omnivore:  Organism which feeds on both plant and animal matter.

Pelagic:  Inhabiting the water column.

Planktivore:  Organism which feeds on suspended microorganisms.

Raptoral carnivore:  Organism which pursues and individually captures
     its prey.

Sublittoral zone:  The benthic zone extending from mean low water (the
     seaward limit of the littoral zone) to 200 m, or the edge of the
     continental shelf usually defined as being 200 m deep.  Beach seines
     were assumed to sample the shallow sublittoral, just below the littoral
     zone.

Suspension feeder:  Typically a benthic organism which processes the water
     flowing over the substrate, feeding on diatoms and other microscopic
     organisms and suspended detritus.

Trophic level:  A group of organisms in a food web that secures food in
     the same general manner.
        IV-H.  Place Names and Locations and Associated Habitats

     Locations of DOE and MESA sampling sites, and their representative
habitat types, which are cited in this report are indicated in Fig. 2.
                                    16

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 Habitat Abbreviations
R/K
C
G
CL
RL
S-C
S
S/E
- Rocky, kelp bed
- Cobble
- Gravel
- Cobble littoral
- Rocky littoral
Sand-cobble
Sand
Sand/eelgrass M/E - Mud/eelgras
^ \^~*f
^
$
•o
 NEAH
        EAGLE COVE (S/E;

          SOUTH

KYDAKA BEACH(S)
         PILLAR POINT(R/K)
           /  TWIN RIVERS(G)
                    OBSERVATORY POINT(RL)
                                           .OWS ISLAND (R/K),
                                         WEST BEACH(G)—
                     MORSE CREEK(S-C)
                             DUNGENESS SPIT(G)
                                JAMESTOWN/PORT
                                   WILLIAMS(M/E)
                                        BECKETT POINT (S/E;
•DRAYTON HARBOR(M/E)
-BIRCH BAY (S/E)
CHERRY POINT(C)
        LUMMI BAY(M/E)
     -VILLAGE POINT (G)
     BARNES ISLAND(R/K)
        -WESTCOTT BAY (M/E)
      POINT GEORGE(R/K)
     EAST GUEMES ISLAND(S/E)
      'ADILLA BAY (M/E)
    OUTH GUEMES ISLAND(G)
      TDALGO BAY(MAR H POINT)(M/E)
         BURROWS BAY(S)
       •ALLAN ISLAND (R/K)


         NORTH BEACH(CL)
Fig.  2.   Locations  of  DOE  and  MESA sampling  sites from which
           nearshore  community and  food  web data were  obtained.
                                              17

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                        V.   Results  and  Discussion

 V-A.   Food  Web  Structure  of Northern Puget  Sound  and  the  Strait of Juan
       de  Fuca

      Synopses of  the  principal  nearshore  communities  and  the  prey spectra
 of  the prominent  species  are in Appendix  A  for  algae  and  invertebrates,
 Appendix  B  for  fishes,  Appendix C for seabirds  and  shorebirds,  and Appendix
 D for marine  mammals.   The  results  of the stomach analyses  conducted
 specifically  for  this  report are in Appendix  B, and the results of the
 nearshore fish  sampling along the western shoreline of Whidbey  Island  and
 at  Burrows  Bay  have been  presented  in Cross,  et al. (1978), and Miller,
 et  al.  (in  press).

      Composite  food webs  have been  constructed  for  seven  representative
 nearshore habitats characterizing north Puget Sound and the Strait of
 Juan  de Fuca—neritic,  rocky/kelp bed sublittoral,  rocky  littoral,  cobble
 littoral, and shallow sublittoral zones of  gravel-cobble, sand-gravel/
 eelgrass, and mud/eelgrass  habitats.  These composite webs are  illustrated
 as  being  much more complex  than they actually are at  any  one  time because
 both  energy sources and consumers change  seasonally and vary  according to
 differences in  the location's environmental character  (e.g.,  wave exposure,
 sediment  sources, freshwater influences).   Accordingly, the food  webs
 were  constructed  by season  and  individual location where  data on  community
 structure existed.  These structures have been summarized in  tables  describ-
 ing the distribution and  abundance  of food web nodes  among functional
 trophic groups  and the  number and relative  importance of  food web linkages
 between trophic levels  (Appendix E).  The following discussion  is based on,
 and constrained by, these simple representations of very  complex,  dynamic
 "structures."

 V-A - 1  Neritic Food Webs

      Neritic  food webs, i.e., those  of the  surface waters and water  column
 in  the  nearshore  region,  are the only webs based principally  on autotrophic
 production  (Fig.  3) and are  the least complex (least number of  linkages
 per node, 1.76) although  the cobble  littoral  food web exhibits  a  slightly
 less  diverse  structure  (Tables  2 and 3).  Phytoplankton (chrysophytes,
 diatoms,  dinoflagellates, microflagellates) produce organic carbon which
 is  grazed by  pelagic zooplankton, principally through suspension  feeding.
 These  small animals may in  turn be utilized by larger zooplankton such as
 primary carnivores.  Although plankton studies in the Strait  of Juan de
 Fuca  and north  Puget Sound have been restricted to offshore waters,
 Chester,  et al.  (1977), suggested that diatoms  (Skeletonema costatum,
 Thalassiosira. sp., Chaetoceros  sp.)  are the principal components  of  the
 plankton blooms and various  microflagellate species form  the  dominant
 non-bloom component of  the community.  Among  the herbivores,  small
 calanoid copepods (Pseudocalanus sp., Acartia longiremis, Microcalanus
 sp., Oncaea borealis) and the cyclopoid copepod Oithona similis^ numerically
 dominate the  surface water zooplankton community.  Larger calanoids  Calanus
 plumchrus and (\ marshallae  migrate  into the surface layers from  deeper
water at night  (Parsons, et  al., 1969; Chester, et al., 1977).  Larger

                                     18

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Table 2.  Distribution and abundance of food web nodes for food webs  characterizing
          nearshore habitats of north Puget Sound and the Strait of Juan  de  Fuca.
Herbivores


Detritus
Habitat processor
Neritic 2
Rocky
sublittoral
w/kelp beds 6
Rocky
littoral 7
Cobble
littoral 5

C
O CO
>-i CO >H l-i
n) -u co cu
tH C C TJ
3 tfl 4) CU
u f-i ex cu
CO CX. CO 4-1
CO 3
5


2

3

3

cu
TO CO
60 U
Microal,
graze



1

3

2

cu
TO co
ao u
Macroal;
graze



1

2

1


Mixed
algae/
detritus Omnivores
1


2 3

2

2 1
Planktivores

CO
cu
rH t-l
Raptora
carnivo
12


14

10

4

4-1 CO
C CU CO
CU J-) M
go cu
d. > H T3
n) -H cu cu
1-1 C 4-1 CU
4J U rH 4-1
C « -H
PJ CJ P*-i
1 3


1



1
Benthivores

CO O CO
J-i -H }-i
•U CU CO CU
•H T3 C T3
CO CU CU CU
0 CU (X CU
O. 4-1 CO 4-1
CU 3
O C/3



1

1

1

CU
Carnivo



11

13

12
Piscivores


0
•H
60
cfl
rH
CU
Pi
6


5





rH
Demersa



1

5

3
rH 60
'£ C v Total
•P -H 0) ,, ,
CO 60 CU It Of
CU -rl VW
M M i nodes
HOC „
cu o N
H !Z n
1 7 38


48

46

35
Gravel-cobble
shallow
sublittoral

Sand/eelgrass
shallow
sublittoral

Mud/eelgrass
shallow
sublittoral
                              10
2   1
10
                  1 11
                                                   13
1    11
                            41
                                           40
46

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Table 3.  Number and relative importance of food web linkages to trophic levels character-
          izing nearshore habitats of northern Puget Sound and the Strait of Juan de Fuca.
          1° = primary, 2° = secondary, 3° = tertiary, Incid. = incidental trophic
          linkages.

Neritic
Rocky sub-
littoral
w/kelp beds
Rocky littoral
Cobble
littoral
Gravel-cobble
shallow
O sublittoral
Sand/eelgrass
shallow
sublittoral
Mud/eelgrass
shallow
sublittoral
Phy toplankton M.icroalgae
2 5 21443
5 4 12 11 2316652
7 51 21 122 214 16 53
5 4 1 241 213
6 3 4952 486
4 1 4 5 13 7 2233
4 3 5 4 25 432244
13
11
13
11
17
37
18
9
31
9
16
15
13
32
Tertiary
22 4 16 14 5 23 25
223 7 12 6 26 26
25 2 21 8 45 27
15 9 12 5 30 27
12 19 10 26 36
12 8 52 22
2 2 16 11 47 40
Tolal t
linkages
Nl
67
104
101
74
91
94
114
x No.
link. if.es
per noJc
1.76
2.17
2.20
2.11
2.22
2.35
2.48

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                                                    NERITIC FOOD WEB
ISJ
                                                              //     / M  /I
                                                              X         ,..„
      Fig.  3.   Composite food web  characteristic of neritic  habitats in northern Puget  Sound and the Strait
               of  Juan de Fuca.

-------
grazers include euphausiids (Euphausia pacifica, Thysanoessa longipes,
T_. spinifera),  larvaceans (Oikopleura dioica), and larval and juvenile
stages of other crustaceans.

     Carnivorous zooplankton includes cnidarian (Hydromedusae, scyphozoans),
ctenophores  (Beroe sp.,  Pleurobranchia sp.)> hyperiid amphipods (Parathemisto
pacifica, Hyperoche medusarum), larvae of large benthic crustaceans
(brachyuran crabs), and chaetognaths (Sagitta elegans, j^. lyra).

     The principal secondary consumers are neritic schooling fishes such
as juvenile Pacific herring, Pacific sand lance, northern anchovy, longfin
smelt, and surf smelt.  Some of these species are present in the community
for only a part of their life history, as in the case of herring which
occupy the neritic waters of the region for their first year before entering
the North Pacific Ocean.  Some species such as Pacific sand lance are present
in the community from larvae to adults.  Other species are even more
transient though ecologically important during their short residency—for
example, juvenile salmonids which occupy neritic waters for two weeks to
six months during their migrations out of Puget Sound and the Strait of
Georgia.  Several species, including coho salmon, may spend their whole
life cycle in the region's inland waters, changing trophic levels several
times.

     Almost all the marine birds and mammals occupy positions of tertiary
carnivores.  Only the gray whale, which may enter the region during its
oceanic migration, can be considered a secondary consumer by its
utilization of zooplankton  (euphausiids and crab larvae).  The high
production of calanoid copepods, which are eaten by neritic fishes, is
undoubtedly responsible for maintaining large numbers of tertiary consumers
and their diversity.  For example, the various alcid seabirds seem to have
evolved their reproductive period and nesting location in conjunction with
the peak occurrence of larval and juvenile neritic fishes.  The Protection
Island colonies appear to be very dependent on the fluxes of neritic fishes
occupying the region and juvenile salmonids migrating through it.

     The diversity in nodes and linkages of the various neritic food webs
also reflects the importance of spawning regions of the adult fishes.  In
general the regions around Cherry Point, Anacortes, and the San Juan
Islands, which are in proximity to spawning areas of Pacific herring, tended
to exhibit more complex food webs than western Whidbey Island and the Strait
of Juan de Fuca (Appendix Tables E-l, E-2).  This also reflects the role
of contained embayments in providing nursery and rearing area for the
larvae of neritic fishes, regardless of whether or not these embayments
were significant spawning areas  (Fresh 1979).  Thus, north Puget Sound
with its abundant embayments  (Birch Bay, Padilla Bay, Fidalgo Bay, Westcott
Bay, Lopez Sound) tends to have more highly developed neritic food webs
than the eastern Strait of Juan de Fuca, which has only two embayments
(Discovery Bay and Sequim Bay), and the western strait which has no
embayments.

     There are several neritic fishes  (e.g., surf smelt and longfin smelt)
which utilize sand-gravel beaches for spawning.  The abundance of this
habitat along the Strait of Juan de Fuca obviously contributes to the

                                    22

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neritic food webs of that region more than in north Puget Sound.

     Seasonal variations in the neritic food webs are probably the most
pronounced of any of the habitats examined (Appendix Tables E-l, E-2).
This would appear to be the result of the extreme seasonality of the
primary production cycle in the temperate waters at these latitudes.
Accompanying this effect is the reproductive cycle of most of the nerz
fishes, which produce larvae at the time of maximum availability or
appropriate food organisms.  Despite the decline in many lower trop ic
level organisms, the overall diversity (number of nodes) of the fal  a
winter food webs is maintained by seabirds wintering in the protected
inland waters of the region.

V-A - 2  Rocky Sublittoral Food Webs

     Rocky sublittoral habitats of the region and their associated kelp
bed  (Nereocystis, Macrocystis) community showed a mixture of neritic and
sublittoral food webs (Fig. 4) and thus the highest number of food web
nodes of any habitat and the second highest number of linkages  (Tables 2,
3).  This in part reflects the steep gradient, well flushed character of
this habitat, which makes neritic organisms available for use by kelp bed
carnivores.  This is evident in the number of secondary carnivores which
occupy the kelp bed habitat, for protection from predation or some other
non-food-oriented purpose, but which typically feed upon neritic organisms.
Black, quillback, and yellowtail rockfish, juvenile gadids and rockfish,
Heermann's gulls, and Brandt's cormorants prey on food resources character-
istic of the neritic communities.  A completely different food web is
organized around the production of macrophytic algae and the accumulation
of detritus.  These carbon sources are in turn utilized by epibenthic
zooplankton and benthic organisms which are prey for epibenthic- and
benthic-fceding carnivores.  Although the bottom habitat is not as capable
of supporting large numbers of infaunal species as the soft-sediment
environments, epibenthic shrimps, crabs, mysids, gammarid amphipods,
isopods,  and copepods occupy the microhabitat provided by the kelp
holdfasts and the macroalgae understory.  Important shrimp species include
Spirontocaris prionata,  Crangon stylirostris, £. franciscorum, and Hepta-
carpus stimpsoni.  Crabs include Cancer gracilis, C^. oregonensis, Oregonia
gracilis, Scyra acutifrons, Hyas lyratus, Pagurus beringanus, P^. dalli,
P^. hirsutiusculus,  Petrolistes eriomerus, Loxorhynchus erispatus, and
Pachycheles sp.   Mysids include Holmesiella anomala, Neomysis awatschensis,
and Archaeomysis grebnitski.  Gammarids include Eusiroides sp., Hyale
frequens, Parapleustes pugettensis, Erichtonius brasiliensis, Photis sp.,
Amphithoe simulans, and _A. lacertosa.  Isopods include Exosphaeroma
amplicauda, _E.  media, Gnorimosphaeroma oregonense, and Dynamenella sheareri
(Nyblade 1977,  1978; Leaman 1976).

     Secondary carnivores which utilize these epibenthic organisms are all
demersal or bottom-oriented fishes, including kelp greenling, copper
rockfish, cabezon,  longfin sculpin, striped seaperch, lingcod, scalyhead
sculpin,  red Irish lord, and blackeye goby.  Although the data base for  the
prey composition of tertiary carnivores, marine mammals, is inadequate,
it appears from existing information that harbor seals prey on smaller
demersal fishes, northern sea lions on the neritic-feeding rockfish, and

                                    23

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                                                   SUBLITTORAL ROCKY/KELP BED FOOD WEB
N3
         l^l^H PRIMARY 175-100% OF TOTAL I R I )
         ^—^— SECONDARY 150-T4% Of TOTAL I R I )
         	TERTIAUT 12S- 49% OF TOTAL I R I I
                      VI   SEA
                    / '*\  """"
                |	1  ^
                 yiCROPHYTIC
        Fig   4    Composite food web  characteristic  of sublittoral rocky/kelp bed  habitats in northern Puget
                   Sound  and the  Strait of Juan de  Fuca.

-------
orcas on the large demersal fishes and harbor seals.

     Comparison of the rocky/kelp bed communities and food webs  in
different regions of north Puget Sound and the Strait of Juan  de Fuca
difficult because of the lack of a uniform data base.  While the DOb   ^
baseline studies produced comparable data for three areas in north  Pug
Sound (Miller, et al., 1977; Moulton 1977), no such data exist for  the
Strait of Juan de Fuca.  Only Leaman's (1976) studies of the kelp bed  ^^
communities in Barkley Sound, outer Vancouver Island, contain  quanti a
data on species composition, abundance, and food habits, although onlyt°
season was represented.  Examination of the food web structures of  nor
Puget Sound (Appendix Tables E-3, E-A) indicated that there may not be
significant differences between the different regions sampled  and that
seasonal variation is comparatively low.  Moulton (1977) indicated that
of the dominant species in the community, quillback rockfish,  copper
rockfish, and longfin sculpin tended to show strong seasonal fluctuations.
The meager data from the strait indicate some significant differences in
species composition, seasonality, and prey resources.  Both Leaman s  (.I?/ )
data and unpublished records from repeated SCUBA dives  (J. Cross, FRI, UW)
in the region suggest that both seaperch and rockfish may be more prevalent
and that several of these species (striped seaperch, canary rockfish, and
black rockfish) show strong seasonality.  In addition, Leaman's  stomach
analysis data indicated that caprellid amphipods may provide more  important
prey resources for demersal fishes, especially red Irish  lord,  kelp
greenling, and scalyhead sculpin, than documented for north Puget  Sound.
Assuming a predominantly detritivorous feeding mode  for  caprellids, these
data suggest that detritus production may be higher  on  the exposed coast,
perhaps because of the extensive Macrocystis kelp beds  which  occur there.
(Macrocystis extends into the Strait of Juan de  Fuca only as  far as
Crescent Bay.)  Although such differences may be very  real, Barkley Sound
is not necessarily representative of the  exposed coast  and the Strait of
Juan de Fuca, since high temperatures, high  salinities,  and low tidal mixing
occur in Barkley Sound in summer  (T. Mumford, Jr.,  Wn.  Dept.  Nat.  Res.).
Nonetheless, the importance of Macrocystis as a  perennial habitat  and
source of detritus should be considered  in evaluating  the structures  of
the rocky/kelp bed community and  food  web in the Strait of Juan de Fuca.

     The importance of the rocky/kelp  bed  habitat,  and  the lack of community
or food web data, in the Strait of  Juan de Fuca  should  not be ignored in
the future.  This habitat probably  forms  the  largest proportion of shoreline
west of Port Angeles, hence playing a  major  role in  the production of
macroalgae and detritus at the base of  the food  webs in all habitats  of
this region.

V-A- 3  Rocky Littoral Food Webs

     The invertebrate fauna of the  littoral portion  of  the rocky/kelp bed
habitat has been extensively studied throughout  north Puget Sound  (Kozloff
1973).  The fish fauna has been only peripherally examined in north Puget
Sound (Friday Harbor Laboratories,  class reports), though sampled extensively
in the Strait of Juan de Fuca as part of the MESA studies  (Cross, et al.,
1978).  The information that does exist for the  San Juan Islands (J.  Cross,
FRI) suggests that the community composition is similar to that documented
for the strait, although there is unquantified evidence that the rocky
                                    25

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littoral fish fauna in north Puget Sound is not as diverse or as abundant
as along the Strait of Juan de Fuca.  The eastern shoreline of north Puget
Sound has very little rocky littoral.  The food webs summarized in Fig. 5,
Tables 2 and 3, and Appendix Tables E-5 and E-6 represent the two regions
combined.

     Detritivores and grazers form the basis of the food web leading to
the secondary carnivores in the rocky littoral.  Detritivorous gammarid
amphipods are especially important, accounting for over half of the food
web linkages to the secondary trophic level (Fig. 5).   Although many of
the amphipods may enter the littoral system with each tidal exchange,
resident populations are probably sustained in the littoral habitats
because of the protection and food supply provided by the extensive
macroalgal community typical of this region (Carefoot 1977).  Detritus
production is sustained both by senescence of annual macroalgae and by the
action of herbivores such as chitons, limpets, sea urchins, and snails
which by their grazing release macroalgae from the substrate.  The seastar
Pisastej: ochraceus plays the "keystone" role in maintaining a diverse
algal community by preferentially feeding on the space-dominating mussel
Mytilus sp. on the outer exposed coast (Paine 1974).  Gastropods such as
Thais sp. and the seastars Leptasterias sp. fill that role inside the
Strait of Juan de Fuca.  As pointed out by Paine's exclusion experiments,
removal of a keystone species results in dramatic shifts in community
dominance and diversity.  Theoretically, removal of these carnivores would
decrease macroalgae production in the rocky littoral and ultimately reduce
the supply of detritus for this habitat and adjacent ones.

     At the base of the food web seasonal fluctuations are quite prominent,
especially those associated with the annual die-off of macroalgae and the
massive recruitment of barnacles and mussels.  Very little seasonal
variation is evident among the principal food web relationships at the
higher trophic levels, however (Appendix Tables E-5, E-6).  Any variation
can be attributed to the occurrence of less common fishes such as saddle-
back sculpin, fluffy sculpin, and sharpnose sculpin, and migratory shore-
birds such as surfbird, whimbrel, and black turnstone.

V-A. 4  Cobble Littoral Food Webs

     Compared with the rocky littoral, the cobble littoral is generally
less diverse and less complex in its community and food web structure
(Fig. 6; Tables 2,3), especially at the secondary carnivore  (fishes) level.
The principal components are typically the same species or functional
groups, but food web nodes and linkages are three-quarters as numerous
as in the cobble littoral.  It is important to point out here that although
the food web structure is less diverse or connected, the extensive
beneath-rock habitat, combined with the rock benthic epifauna, may make
this habitat one of the most productive on the basis of standing stock of
the community and the extent of this habitat in the region.

     Some striking differences in food web structure, as compared to the
rocky littoral, include the increased importance of idoteid isopods
(Idotea urotoma, Pentidotea montereyensis), the decreased importance of
shorebirds, and the presence of the pigeon guillemot, a diving piscivore.
                                    26

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                                                      ROCKY LITTORAL FOOD WEB
           RELATIVE IMPORTANCE
               0«
           FOOD wee LINKAGES

      Mi^w efl uiO, (75-100% Of TOTAL I R I I
      -   SECONDARY I50-T4% OF TOTAL I R I ]
      	T£RTiA»T US- 49% OF TOTAL I R I )
                                       HEKHIW 6ULL,
                                      CALIFORNIA 6ULL,
                                       WESTERN QULL,
                                     SLAUCOUS-WINGCD CULL
     NUDI8RANCHS -*,  ^
    I	—J    ;
            i ZOOXANTHELLAE
Fig.  5.   Composite food web  characteristic of rocky  littoral  habitats in northern Puget  Sound and the
           Strait  of Juan de Fuca.

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                                                    COBBLE LITTORAL FOOD WEB
S3
00
      Fig. 6.   Composite food web characteristic of  cobble  littoral habitats in northern Puget Sound and the
                Strait  of Juan de Fuca.

-------
Detritus grazers still form the basis of the food  web  leading  to  the
secondary carnivores.  As in the rocky littoral, seasonal variation 1
slight (Appendix Tables E-5, E-6) .

V-A - 5  Gravel-Cobble Shallow Sublittoral Food Webs
                                                         .   the overall
     Perhaps the most widespread and most common habitat in     ra^ habitats,
study area, after the rocky littoral and rocky/kelp bed su   ,  e to sediment
is the gravel-cobble beach.  Variations in this habitat are     gence  of
size and exposure to wave action, which largely restrict     hitat tend  to
macroalgae or eelgrass.  In general, the food webs in  tn^s     ^ut ^e most
be the least diverse  (in number of nodes) of all the food we t>  ,    fcg
connected  (^ No. of linkages/node = 2.22) of all the exposed
(Fig. 7, Tables 2,3).
     Even though this habitat does not typically support
macroalgal communities, detritus is apparently transported and
in the unconsolidated sediments which act as a physical grinder,
up kelp and algal fragments into smaller particles easily utilize   y
detritivores.  Detritus-grazing epibenthic crustaceans such as gammar
amphipods, cumaceans, and harpacticoid copepods, and mixed algae-de ri
grazers such as mysids and f labellif eran and valviferan isopods were the
most important elements of the lower trophic levels which supported the
majority of the secondary consumers.  Key species  include the  amphipods
Ischyrocerus anguipes, Paraphoxus   sp. , Melita desdichata, Paramoera mohri,
Pontogeneia ivanovi , Hyale rubra  f requens ;  the cumaceans Cumell% sp . ;  the
mysid Archaeomysis  grebnitzki; the  f labellif eran isopods Gnorimosphaeroma
oregonensis, Exosphaeroma amplicauda,  and Dynamene 1 1 a shear eri;  and  the
valviferan isopods  Idotea wosnesenski,  Synidotea sp. , and  Idotea monterey-
ensis.  Gammarid amphipods supported  7  of  the  10 epibenthic  planktivores
and accounted for 16 of  the  46 linkages to  secondary  carnivores.

     There are a number  of variations  in food  web structure  among the  ten
gravel-cobble habitat  locations  examined (Appendix Tables  E-7, E-8).   In
general, the more protected  the  location,  the  more complex the community
and food web structure.  Relatively protected  locations like Legoe Bay,
Deadman Bay, and south Guemes Island  appeared  to have the  highest number
of food web nodes and  linkages.   At the other  extreme, exposed sites like
Alexander's Beach,  Dungeness  Spit,  and Kydaka  Beach indicated  the least
diversity.  This is not  always the  case, however.   Relatively  exposed
locations such as Cherry Point and  South Beach exhibited more  diverse  food
webs than might be  expected.  At  Cherry Point,  large  boulders  and sparse
kelp beds (Nereocystis)  immediately adjacent to  the beach probably acted
to diversify the habitat and  decrease  its exposure to wave action.  Similarly,
kelp beds were present offshore of  the  gravel-cobble  beach at  West Beach  on
the west coast of Whidbey Island.   South Beach,  on the southwestern shore
of San Juan Island, exposed to the  Strait of Juan  de  Fuca, is  a high-energy
beach with no such protection.  The relatively high diversity  of  the
community is therefore difficult to explain in the same manner.   Some of
the additional species may be attributed to the expansive sandflat habitat
immediately offshore,  but the complex food web documented in summer was
still unexpected.


                                    29

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                                       EXPOSED GRAVEL-COBBLE SHALLOW SUBLITTORAL FOOD WEB
             	TERTIARY 123-49% OF TOTAL IR I)
             	— INCIDENTAL (IO-Z4%OF TOTAL I R I
                                     I SILVERSPOTTEOI   A £ _--   \ ,     I   CNGLISM
                                   llf\  SCULPIN  •«-  i "fl    x^	I/    V    SOLE
                                  ,     F^	x"  > 'I   /I  "^  I M   IJ''V'
                                        I «H,,E,PO,TED ' V  I  ^ »•   !J""°""  M\ ,^ W
Fig.  7.   Composite food web  characteristic of   exposed gravel-cobble,  shallow sublittoral habitats  in
           northern Puget Sound and  the  Strait of Juan de Fuca.

-------
     One obvious difference in food web structure among the various
gravel-cobble locations was the increased importance of mysids at,  e
locations along the Strait of Juan de Fuca.  These shrimp-like eP"
crustaceans  (predominantly Archaeomysis sp.) appeared to supplant  ^ ^
role of gammarid amphipods as prey for many epibenthic caynivoreSult of
region.  It  cannot be established, however, whether this is a res
increased exposure at sites in the western portion of the ^ral  .
greater influence of oceanic environment.  Cumaceans (CumeTLa. sp.
tended to be more important in the strait.

     Seasonal variations are not consistent (Appendix Tables E~7^ ~um
In many locations there is a greater food web diversity evident i
and fall, especially at some of the north Puget Sound locations.  Locat
in the strait, however, maintain approximately the same number ot too  w
nodes throughout the year, even though there is usually an increase in tne
total number of food web linkages in summer.

V-A- 6  Sand/Eelgrass Food Webs

     Sand/eelgrass habitats generally represent more protected versions of
the gravel-cobble habitat.  They are shallow, semi-enclosed  embayments
which have low- to moderate-energy beaches, allowing sand  and mixed  fine
gravel to accumulate and stabilize.  Only one the  sites  typified a moderate-
to high-energy location—Eagle Cove on the western shore of  San  Juan Island.
The principal characteristic is the presence of "beds" of  the  seagrass
Zostera marina.

     The influence of the more stable substrate is illustrated  in the
community and food web structure  (Fig. 8).  Benthic infauna and epibenthic
fauna which  inhabit the top layers of the  bottom  sediments are more
prevalent, as are carnivores which prey  on  benthic forms.   In addition,
the increased abundance of small epibenthic crustaceans  (harpacticoid
copepods, cumaceans, tanaids) provides the  necessary prey  resources  for
juvenile demersal fishes which can seek  protection from predation among
the eelgrass beds and in the shallow waters.

     Eelgrass is responsible in many ways  for the  complexity of  the  food
web, which has the second highest average number of linkages per node, even
though the total number of nodes is only average  (Tables 2,3).   As a
structural habitat, eelgrass increases the  substrate available  for the
growth of epiphytic algae and associated fauna, reduces wave and current
action, traps sediments and detritus, maintains high dissolved oxygen
concentrations through photosynthetic activity, and by shading at low
tide minimizes fluctuating temperatures  that would be induced by direct
sunlight (Kikuchi and Peres 1977).  More important, eelgrass and  its
associated epiflora provide great quantities of detrital carbon  to the
nearshore system through autumn die-back and atrophy of the  emergent
growth.

     Detritus, which is presumably eelgrass-derived to a great extent,
provides energy directly to detritivores and indirectly to  primary
carnivores preying on benthic meiofauna.   Important detritivorous
crustaceans include harpacticoid copepods;  the gammarid  amphipods

                                    31

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                                    PROTECTED SAND-GRAVEL/EELGRASS SHALLOW SUBLITTORAL FOOD WEB
U)
      Fig.  8.  Composite  food web characteristic  of  protected sand/eelgrass,  shallow sublittoral habitats in
               northern Puget Sound and the  Strait  of Juan de Fuca.

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 Anisogammarus  confervicolus, Pontogeneia sp., Ischyrocerus anguipes,
 Paraphoxus  sp., Photis brevipes, Aoroides sp., Atylus tridens, and
 Podoceropsis inaequistylus;  the tanaid Leptochelia dubia; the leptostracan
 Nebalia  pugettensis;  the cumacean Cumella vulgaris; and pagurid crabs
 Pagurus  sp.  Crustaceans which are able to utilize both detritus and
 epiphytic macroalgae  include the flabelliferan isopods nnorimosphaeroma
 oregonense  and Exosphaeroma  sp., and the valviferan isopod Synidotea^
 nodulosa.   Primary  carnivores upon benthic meiofauna include crangonid
 shrimp,  crabs, tanaids, and  polychaete annelids.

      Of  the secondary carnivores which exploit the diverse epibenthic and
 benthic  fauna, juvenile and  adult flatfish are one of the most important
 groups,  including juvenile English sole and  sand sole, and adult rock sole.
 Other species  compose an assemblage which is apparently characteristic of
 eelgrass beds, including shiner perch, bay pipefish, penpoint gunnel, and
 tube-snout.  It is  in this habitat, as well  as in the mud/eelgrass habitat,
 that  benthic-feeding  shorebirds are prevalent—greater yellowlegs, sander-
 ling,  least sandpiper, and western sandpiper.  As the only true herbivores
 upon  the eelgrass,  Canada geese, American coot, and black brant commonly
 inhabit  the sand/eelgrass habitats during their seasonal stay in the region.

      In  general, the  sand/eelgrass communities and food webs at the different
 locations have the  same structure (Appendix  Tables E-9, E-10).  Birch Bay
 appears  to  be  slightly more  diverse in structure than the other three sites,
 mainly because of a greater  number of benthic carnivores.  Pacific sanddab,
 whitespotted greenling, and  buffalo sculpin were common in this habitat
 only  at  Birch Bay.  Other important differences were the absence of
 herbivorous waterfowl in the western Strait  of Juan de Fuca  (where the
 eelgrass beds are reduced and sparser than in north Puget Sound and  the
 eastern  strait) and the presence of redtail  surfperch only in the western
 Strait of Juan de Fuca.  There is no definite pattern in the seasonal
variability of the number of food web nodes  or linkages.

 V-A - 7  Mud/Eelgrass Food Webs

     Undoubtedly,  the most complex and highly connected habitat is the
mud/eelgrass habitat and the saltmarsh environment often associated with
it (Fig.  9).  Overall, the highest number of food web nodes and linkages
and the highest average number of food web linkages per node were found
for this habitat (Tables 2,  3).  Most of the increase, as compared with
the sand/eelgrass habitat, originates at the carnivore level where there
are twice as many incidental linkages leading to the secondary carnivores.
The principal reason for the increase is the presence of the benthic-feeding
shorebirds  (sanderling, longbilled dowitcher, shortbilled dowitcher,
greater yellowlegs).  At the tertiary consumer level, the great blue heron
preys on a number of demersal fishes.

     The principal species involved in the food web transfer are basically
the same as in the sand/eelgrass habitat.  It would appear that, rather
than an  increase in species  richness, the actual production is higher,
resulting in higher densities of each taxon.  If there is any increase in
species or  functional groups in the mud/eelgrass habitat it is probably
to be found in small epibenthic zooplankton and meiofauna.   Nyblade's (1977,

                                    33

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                                       PROTECTED MUD/EELGRASS SHALLOW SUBLITTORAL FOOD WEB
OJ
•P-

^^

PHtMA
SECO*
TERTl
1NCID
OF
It (75-100% OF TOTAL 1 H 1 )
OARY (50-74% OF TOTAL 1 R I )
ftRY 125- 49% OF TOTAL 1 H 1 )
ENTAL 
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1978) documentation of the littoral and shallow sublittoral benthic
communities shows that although the species richness may be slightly
higher in the lower intertidal levels of the mud/eelgrass habitat, the
densities and biomass of organisms usually are appreciably higher  than in
the sand/eelgrass habitat.  Neither the DOE nor the MESA sampling  design
quantified organisms such as epibenthic zooplankton or meiofauna;
therefore, this difference may be more pronounced  than indicated.  MESA-
sponsored epibenthic plankton sampling during August 1978 along the  strait
(Simenstad and Kinney, in prep.) also substantiates the fact  that  epibenthic
crustaceans such as harpacticoid copepods less than 1 mm in size are  much
denser in the mud/eelgrass habitat at Jamestown and Point Williams than in
the  sand/eelgrass habitat at Beckett Point  (crustaceans in DOE and MESA
benthos  studies sieved down to 1 mm; the epibenthic plankton  pumping
filtered to 0.209 mm).  Furthermore, densities were almost five times
higher within the eelgrass bed than in the base sediment.

      One possible reason  for the high benthic diversity and production, in
 addition to the sediment  particle size and  its ability to entrain  organic
matter,  is the input of organic detritus from the  saltmarshes which  usually
 occur in the  estuarine end of the contained embayments in this region.
 This vascular-plant vegetation perennially  produces high biomasses of
 organic  material which tends to accumulate  on the  mudflat and partly
 decompose there, and then is transported into the  estuary with the spring
 runoff and spring  tides.   A lag of 14 months between peak carbon uptake by
 the saltmarsh vegetation  and eventual flushing into the estuary has  been
 documented  for  saltmarsh  ecosystems on the  Atlantic coast  (Hopkinson  and
 Day 1977).

      Perhaps  because  the  habitat is protected, there are no distinct
 indications  of  seasonal change in food web  structure or diversity.
 Differences  in  food web structure between the three mud/eelgrass locations
 are minimal  except  for the relative unimportance of flabelliferan  isopods
 at the Strait of Juan de  Fuca location  (Jamestown  - Port Williams) as
 compared to  their  prominance in the food webs in north Puget  Sound.

 V-B.  Prey Assemblages of Major Importance  to Upper Trophic Levels

      Several  functional groups or taxa of organisms stand out as important
 energy sources  or  prey resources for upper  trophic levels  (Table 4).   The
 criteria determining  importance are that the organisms  (1) provide the
 majority of  the  energy sources for consumer organisms at some time,
 (2) provide  important conversion or transfer of organic matter to  trophic
 levels where  it  is  available to higher level consumers, or  (3) hold
 "keystone" roles in  structuring the composition of the community and  the
 directions and  rates of food web energy  flow.

      The simple  but  extensive transfer of phytoplankton biomass to
 planktivorous neritic fishes by calanoid copepods  is a prime  example.
 Calanoids rapidly  convert autotrophic carbon into  high density plankton
 particles which  are  utilized by dense larval and juvenile neritic  fishes,
 especially herring  and Pacific sand lance.  These  carnivores  are in  turn
 the principal prey  of the alcid seabirds and several marine mammals.
 This calanoid chain, unlike the detritus-based chain, is characterized

                                     35

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Table 4.  General listing of species or functional groups which are of
          major importance to the upper trophic levels of the  nearshore
          food webs of northern Puget Sound and the  Strait  of  Juan de Fuca.
          See text for criteria determining importance.
Primary
energy
sources
Decomposers   Herbivores
Diatoms:

Chaetoceros sp.
Ske~ietonema GO station
Thalassiosir>a sp.

Macroalgae:

Nereocystis sp.
Laminaria sp.

Vascular plants:

Zostera marina
Fungi
Bacteria
Flagellates
Calanoid copepods:

Acartia longremis
Calanus mavshallae
C. plwnchrus
Epilabidocera amphitrites
Microcalanus sp.
Oncaea borealis
Pavacalanus sp.
Pseudocalanus sp.

Cyclopoid copepods:

Oithona similis

Euphausiids :

Euphausia paaifica
Thysanoessa longipes
T. spinifera

Sea urchins :

Stvongylocentrotus sp.

Polychaete annelids:
                                               s sp.
                                     Lwnbrineris sp.

                                     Molluscs :

                                     Lacuna sp.
                                     Acmaea sp .
                                     Transenella sp.
                                     Katar-ina  tunioata
                                     Mopalia  sp.
                                     Littorina sp.
                                     Mytilus  sp.
                                     Balanus  sp.
                                     36

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Table 4, cont'd
Detritivores
  Primary
carnivores
                                                       Secondary  and
                                                         tertiary
                                                        carnivores
Harpacticoid copepods:     Seastars:
                                                       Fishes:
Amphiasooides cinctus
Dactylopodia sp.
EC tinos omidae
Harpaetieus uniremis
Bimtemarmia jadensis
Tisbe sp.
 Zaus sp.

 Cammarid amphipods:

 Amphithoe simulans
 A. lacertosa
 Aoroides columbiae
 Atylus tridens
 Anisogaomarus sp.
 Eohaustorius sp.
 Erichtonius brasiliensis
 Eusiroides sp.
 Hydle rubra
 Ischyrocerus sp.
 Callvpiella pratti
 Melita desdiohata
 Paramoera mokri.
 Panxphoxus sp.
 Parapleustes pugettensis
 Photis brevipes
 Pontogeneia sp.

 Flabelliferan isopods:

 Dynamenella sheareri
 Exosphaeroma amplicauda
 Gnorimosphaeroma
   oregonense

 Idoteid isopods:

 Idotea wosnesenski
 I. Montereyensis
 I. upotama
 Sirnidctea sp.
Pisaster sp.
Leptasterias

Fishes:
               Ophiodan slangatus*
sp.
Armodytes hexaptems
Anoplarchus purpwescens
Clicpea harengus pdllasi.
CymatogasteT aggregata
Gobiesox maeandricus
Hypomesus pretiosus
Leptocottus armatus
Oligocottus maciilosus
Oncorhynchus keta
0.  kisutch
0.  tshauytscha
Parophrys vetulus
Psettichthys melanostictus
Spirinchus thaleichthys
 Molluscs:

 Octopus dofleini
 Thais  sp.

 Polychaete annelids:

 Dorvillea sp.
 Glycera sp.
 Hemipodus sp.
 Micpopodarke sp.
 Nephtys sp.
 Ophiodromus sp.
 Phyllodocidae
 Ppotodorvillea sp.

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Table 4, cont'd
Detritivores, cont'd
My sids:

Aoanthomysis sp.
Archaeomysis grebnitzki
Holmesiella anomala
Neomysis mercedis

Cumaceans:

Cumella sp.

Tanaids:

Leptochelia dubia

Leptostracans:

Nebalia pugettensis

Shrimp:

Crangon sp.
Hippolyte  sp.
Lebbeus sp.
Sp-Lvontocavis  sp.
Heptacarpus sp.

Polychaete annelids:

Capitellidae
Cirratulidae
Paraonidae
Spionidae
                                     38

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by high turnover rates, high energy demands  on the part  of  the consumers,
and a relatively contracted time  scale—in spring and early summer when
the spring phytoplankton hlooms develop,  the larval fish enter the neritic
environment, and the alcids are scouring  the neritic fish communities for
food for their nestlings.

     Overall, the most consequential  organisms to nearshore food webs were
gammarid amphipods, which dominated the detritivore group in every
environment except the neritic.   Although the taxonomic  documentation of
this diverse group of crustaceans is  very incomplete, we have distinguished
a number of genera which appear to be critical participants in the food
web.

     Other  important detritivorous crustaceans are harpacticoid copepods,
 flabelliferan and idoteid isopods, cumaceans, mysids, tanaids, leptostracans,
 and shrimps.

      Of the carnivores, besides Pacific herring and Pacific sand lance
 already mentioned,  shiner perch,  Pacific  staghorn sculpin,  tidepool sculpin,
 surf smelt, longfin smelt,  salmon (principally juvenile  chum, coho, and
 chinook),  juvenile  sand  sole,  and northern clingfish were identified as
 contributing a significant  proportion of  the primary and secondary food web
 linkages to tertiary carnivores.   Octopus and lingcod are important predators
 on sublittoral fish and  invertebrate  communities, and themselves are
 utilized by marine  mammals  and man.   In the shallow sublittoral habitats,
 several seastars,  gastropods,  and polychaete annelids by preferentially
 feeding on certain components  of  the  lower trophic levels determine the
 course of competitive interactions,  regulate the community structure, and
 influence the pathways of trophic energy  flow.

 V-C.  Relative Importance of Autotrophic  Versus Heterotrophic Energy Bases
       to Nearshore Food  Webs

      The importance of detritus as a carbon source for consumer organisms
 is readily apparent from our documentation of the nearshore food webs of
 north Puget Sound and the Strait  of Juan de Fuca.  Only in the neritic
 food webs do autotrophic (phytoplankton)  sources provide more numerous and
 more important linkages to consumers than detritus, and even in this case
 many herbivores are functionally capable of exploiting both phytoplankton
 and detritus particles.   Even in the neritic food web, the established
 paradigm of the dominant role of phytoplankton production is becoming
  increasingly challenged in favor of microbial utilization of dissolved
 organic matter and non-living organic particles  (Pomeroy 1974).  Although
 much of the macroalgae and vascular plant vegetation in the nearshore
  environs are directly grazed by herbivores such as sea urchins, isopods,
 and brachyuran crabs, the majority of the first level consumers are
  detritivores.  Thus, while some autotrophically produced biomass is
  directly transferred to higher trophic levels, the majority appears to
  reach maturity and detach and decompose in the nearshore region, eventually
  providing a pool of suspended and dissolved organic matter available for
 heterotrophic conversion to decomposer biomass.  The apparent unimportance
  of food web transfer by herbivory may be partly due to the control of
  herbivores by predators.  Much of the reason may also lie in the fact that

                                      39

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most marine invertebrates cannot digest structural polysaccharides because
they lack adequate digestive enzymes (Kristensen 1971).  In the case of
eelgrass, the only significant grazers appear to be sea urchins (uncommon
in the eelgrass beds of this region) and herbivorous waterfowl such as the
black brant (Kikuchi and Peres 1977).  Even in the case of benthic
macroalgae, herbivores may cycle more biomass into the detritus "pool"
by detaching large pieces of algae from the substrate  in the act of
grazing the holdfasts.

     Just as live macroalgae cannot be used by grazing invertebrates,
subsequent detritus particles also are not directly convertible.
Structural and nutritional decomposition of the particles by microflora
such as marine bacteria appears to be a critical process in conditioning
the detritus before utilization by detritivorous fauna.  As has been
recently established, the associated microflora may be the actual food
source of the detritivores (Mann 1972; McRoy 1970; Brown and Sibert 1977;
Mclntyre 1969; Seki 1966).

     Other recent investigations into the trophic structure of food webs
characterizing estuaries  and coasts have also pointed to the importance
of the transfer of primary production into detritus and the linkages
between the detrital carbon, the associated microflora, the detritus-
stripping fauna, and the fishes which feed on them (Williams, et al.,
1968; Odum 1970; Qasim and Sankaranrayanan 1972; Shubnikov 1977; Simenstad,
et al., 1977).

     Thus, the dynamics and mechanisms of the processes which determine
(1) the transport of detritus sources into the estuarine and nearshore
environment,  (2) the physical and chemical conditioning of the detritus,
and (3) the development of microflora on the detritus particles will
dictate how detritus-based food webs can be altered by pollutant effects
or how pollutants may be transferred to higher trophic levels through
detritus particles.  In general, these processes are poorly understood
and seldom quantified.
                                    40

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V-D.  Ecological Impact of Introduction or Incorporation of Petroleum
    Hydrocarbons Into Nearshore Food Web Structures of North Puget
                 Sound and the Strait of Juan de Fuca

     Petroleum hydrocarbons in the nearshore habitats of north Puget Sound
and the Strait of Juan de Fuca could disrupt both the structure and the
energy flow dynamics of their marine food webs.  The resulting ecological
effects would vary according to the quantity and type of petroleum spilled
and the conditions of weathering and biodegradation before and after the
petroleum impacts the habitat.  The potential for extensive and long-
lasting alterations to the food web would in turn depend on which nearshore
habitats are impacted and the season in which the incident occurred.

     A brief introduction to the documented toxic effects of petroleum
hydrocarbons to  temperate marine organisms is in Appendix F.  The existing
knowledge of petroleum effects on marine organisms and ecosystems, and
documentation  of the observed effects of accidental oil spills, has been
utilized  to assemble the  following description of the most probable and
serious ecological effects which could be imposed on the nearshore food
webs of the region.

     Disruption or alteration of a food web could occur through several
mechanisms:   (1) Selective elimination of species or functional groups
which provide  important food web linkages to higher trophic levels,
 (2) disruption of key processes which control the conversion of non-
utilizable carbon into biomass available for consumption by lower trophic
level organisms, (3) selective elimination or reduction in populations of
"keystone" predators  (Paine 1969) or "foundation" species (Dayton 1972)
which control  or dominate competitive interactions between components of
lower trophic  levels, and  (4) sublethal effects on metabolic activity,,
behavior, growth, and reproduction causing key species to lose their
competitive advantage or  reducing their effectiveness as predators or
herbivores.  Also to be considered is effects via inter-food web relation-
ships, where disruptions  in one community may eventually result in
disruption of  the food web in another community. This is particularly evident
where a species occurs in one habitat as a juvenile and in another as an
adult.  Examination of the region's food web structures indicates there is
viable potential for alteration through several mechanisms.

     One  of  the most obvious cases is that of the short, simple food web
 involving neritic phytoplankton, herbivorous calanoid copepods, and
planktivorous  fishes, specifically Pacific herring and Pacific sand lance.
Discussions  in Appendix F point out that levels of prey abundance and size
are critical determinants of fish survival during the transition from larval
 to juvenile  stages.  It is also at this stage that these raptoral
planktivores are probably least able to switch to alternative prey organisms
without significant cost  in survival.  Specifically, the populations of
 small surface-dwelling calanoids  (Pseudocalanus sp., Microcalanus sp.,
Oithona sp.),  which are the group most vulnerable to acute toxic effects

                                      41

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from spilled petroleum in neritic habitats, provide the high density prey
aggregations which larval Pacific herring, Pacific sand lance, surf smelt,
and longfin smelt depend on.  This highly structured predator-prey linkage
may specifically include the early life history  (nauplius, copepodite)
stages of these small calanoids, which are most  densely aggregated in  the
upper surface  (0 m to -10 m) and which may form  the limited size fractions
(less than 0.500 mm) the larval fish first feed  on.  High densities of
adult Pseudocalanus sp., Microcalanus sp., and Oithona sp. extend down to
approximately  -25 m (Mark Ohman, UW Dept. Ocean.), so it is improbable
that the total population of adults would be severely depleted by the  toxic
effects of soluble petroleum components diffusing from the surface.  The
extent of depletion would depend on the extent of surface mixing of the
water column.  However, we do not know whether the larval fishes, still
relatively planktonic at that stage, are functionally entrained in the top
surface layers and vulnerable to toxic hydrocarbons or subject to reduction
in the only food resources available to them.  In any case, considering
the potential  vulnerability of these larval fishes and their unique prey
resources and  the contribution the neritic fishes make to upper trophic
levels, the potential for significant and long-term food web disruption
is extremely high during March through June when the larvae occupy the
region's surface waters.

     The later juvenile stages of these neritic  fishes, as well as juvenile
salmonids (coho, chinook, and pink salmon) which continue to occupy neritic
habitats through late fall and early winter, do  not appear to be as
vulnerable to  the effects of spilled petroleum hydrocarbons.  By this  stage
their diet, though still selective by prey size, has become more diverse.
In most cases, large calanoid copepods (Calanus  sp.) which ascend from
deep water populations constitute the majority of their diet.  Thus,
although these preferred prey may be locally depleted, there is still  the
diel migration of prey from unaffected populations in deep water to re-
supply the surface waters.  Only two prey taxa would be severely affected
by surface-oriented pollutants such as oil—drift terrestrial insects,
prominent in the diet of juvenile chinook salmon, and the neustonic
calanoid copepod Epilabidocera amphitrites, which are preferentially con-
sumed by all juvenile salmonids during daylight  foraging.  The capability
of temporarily switching to other appropriate prey—even  though they may
not provide optimal energy  (caloric) content for the energy required to
capture them—would probably enable these fishes to obtain alternate prey
without adverse effects.

     The previous impacts have involved acute toxic effects responsible
for immediate elimination or reduction in predator-prey linkages.
Sublethal effects may also be manifested  in the neritic food web and
could contribute to long-term effects  through several generations  of
prey and predator populations.   Through the consumption of petroleum
hydrocarbon-contaminated copepods,  neritic fishes could accumulate
enough hydrocarbons or their metabolic byproducts to affect behavior or


                                    42

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reproduction.   Thus, recruitment of ecologically and commercially
important species could be adversely affected through the lifetime of  the
yearclass* and its progeny.  This transfer of contaminants does not stop
at the neritic fishes, of course.  As noted earlier, they support major
trophic linkages to alcid seabirds and marine mammals that could
subsequently suffer adverse sublethal effects similar to those described
for the fishes.

     In the case of the nearshore littoral and shallow sublittoral food
webs, we envision the potential for one of the most deleterious impacts
from spilled petroleum reaching nearshore habitats.  Considering the
obvious importance of detritus to all the region's nearshore food webs,
any disruption or contamination of the detritus pool, its replenishment
from macroalgae and eelgrass or the physical-biological process of
decomposition and colonization by microorganisms implies extensive, long-
term alterations of the structure of the food web or its productivity.
While growth of sulfide-generating bacteria may be enhanced by the
adsorption of petroleum by detritus particles, the deleterious sublethal
effects on detritivorous zooplankton and meiofauna (harpacticoid copepods,
gammarid amphipods, cumaceans, shrimps, tanaids, leptostracans, polychaete
annelids, oligochaetes) may more than compensate for any possible benefit
due  to  increases in microbial activity.  Although the populations of the
detritivores may not  be directly (acutely) affected as a result of their
stripping of hydrocarbons off the detritus particles, incorporation of
hydrocarbons into  their tissues would probably result in some decrease in
reproductive potential.  Reduction in the density of epibenthic detritivores
could in  turn  reduce  the importance of the linkages leading to many of the
carnivores which feed upon these numerous epibenthic organisms.  During
certain seasons  (especially late spring to early summer) the carnivore
compartment includes  juvenile stages of many ecologically or commercially
important species  (i.e., English sole, chum salmon, staghorn sculpin,
tidepool  sculpin)  which utilize  the shallow sublittoral zone as a nursery
area, partly because  the epibenthic organisms form appropriate prey items.

      The  persistence  of  this impact will be determined in part by the
availability of  the petroleum hydrocarbon to weathering and biodegradation.
This has  been  shown to be  particularly associated with the degree of
unconsolidated sediments characterizing the habitat where the petroleum
has  accumulated.   The most extreme  case documented to date is evidenced
by the  prolonged  incorporation of hydrocarbons into soft sediments
generally associated  with  estuaries and marshlands.  This has been well
 illustrated by the history of the West Falmouth spill of No. 2 fuel oil
       *Yearclass denotes  those  fish of a species which were spawned
 during one year and are usually identified  through an analysis of  the
 population's age structure.
                                      43

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in 1969.  Long-term documentation of the effects has indicated that
petroleum degradation in the soft-bottom saltmarsh habitats occurs slowly,
especially below the surface of the bottom sediment (Blumer and Sass 1972).
Almost seven years after this relatively small spill was stranded in the
area 's marshlands there was evidence of aromatic hydrocarbons present in
the marsh sediments at concentrations above background (Teal, et al., 1978).
While the lighter aromatics were responsible for immediate and toxic effects
upon benthic organisms, such as the trophically important fiddler crab Uca
pugnax, long-term reductions in crab density, sex ratio changes, reduced
juvenile recruitment, high overwinter mortalities, hydrocarbon assimilation
into tissues and behavioral abnormalities appear to have resulted from
extended contact to sublethal concentrations entrained within the sediments
(Kreb and Burns 1977).  Thus, even though more resistant fauna were able
to recolonize the surface, subsurface sediments still showed oil below
2.5 cm was fresh and presumably  toxic after 24 months as the oil at the
surface was after 10 months.

     Thus, detritivorous benthic meiofauna, which either graze the detritus
particles in the interstitial water or filter the water completely, may
thus be continually subjected to hydrocarbons leaching from the lower
sediments (Roesijadi et al. 1978).  In addition to the infaunal species,
the many epibenthic organisms residing in the top sediment layer (often
referred to as the flocculent layer),      by their periodic migration
into the water column where they are susceptible to predation, act as
transporters of still active hydrocarbons between the sediments and the
water-column biota.

     A third result which might be predicted from petroleum hydrocarbon
pollution in the nearshore environs involves the selective mortality
of the ecologically dominant species listed in Section V-C.  These species
by their selective predation on competitively dominant species at lower
trophic levels or by their own competitive dominance play deterministic
roles in structuring much of their nearshore community.  In the first
instance, elimination of predators such as seastars and gastropods, which
control herbivore and sessile bivalve and barnacle populations, would
probably result in the successive dominance by more efficient occupiers
of space, thus reducing or eliminating macrobenthic algae.  This effect
could be profound, not only to the food web structure of those affected
habitats but to any food web which is based on the detritus generated by
the periodic die-off of these macroalgae.  Those habitats in which
macroalgae detritus collects and decomposes, even though the pollutant
may not be impacting them, could have their food webs indirectly altered
by the cessation or reduction of detritus input.

     Alternately, in areas where herbivores are dominant such as in rocky
sublittoral habitats occupied by sea urchins, removal or significant
reduction in these herbivore populations would drive the benthic community
more toward one dominated by macroalgae, with a likely decrease in
                                     44

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diversity of benthic epifauna and a potential increase in detritivorous
zooplankton such as mysids and gammarid amphipods.

     When considering the potential for disruption of established food
web structures by introduction of pollutants, questions of the food web  s
"stability" arise, its inherent susceptibility to collapse to a different,
usually less diverse, structure under a perturbation (removal of linkages
or nodes), and whether or not the food web is actually "unstructured,"
i.e., carnivores randomly utilize energy from the mean composition of  the
food web above the primary production level  (Isaacs 1972, 1973).  The
concept that a community is proportionally more stable with increasing
diversity of energy pathways  (increasing connectance) has become an
established paradigm  (MacArthur 1955; Watt 1964) although more recent
laboratory and mathematical modeling exercises have suggested that randomly
connected systems  tend to become less stable as the connectance increases
 (Gardner and Ashby 1970: May  1972, 1973).  De Angelis (1975), however,
has  illustrated  a  number of plausible cases  in which a similar food web
model  does  show  increased stability with increased connectance.  While
 this argument  still smolders  among theoretical ecologists, there are
 inadequate  empirical  data, either laboratory or field, which have  tested
 this hypothesis  by perturbing a  diverse spectrum of food webs.  The most
 appropriate example is  that  of Paine's  (1966) manipulation of rocky  shore
 communities of Washington's  exposed coast.   In absence of such verification,
 the existing information suggests  that  elimination of certain community
 dominants and their associated food web linkages  can result  in dramatic
 alteration (decreased diversity  and  connectance)  of the  food web structure.
 As a result,  relatively complex  food webs  should  not necessarily be  con-
 sidered any more stable than less  diverse  food webs if pollutants  can
 selectively impact populations of  these dominant  species  or  groups.

 V-E.  Comparison of Food Web Structures at Existing or Potential Oil
      Terminal Sites and an Evaluation of  their  Relative  Importance

      Four areas in northern Puget Sound and the  Strait of Juan de  Fuca
 have been considered for location of major oil  transshipment  facilities:
 Cherry Point, March Point (Anacortes),  Burrows  Bay, and  Port  Angeles  (Ediz
 Hook).   Discussion of the habitats and food webs of  these  areas will
 contribute information useful in evaluating these sites  for  such  facilities.

      The neritic  food webs at the four areas are fairly  similar  in structure,
 However, food webs of the Anacortes area (Padilla Bay,  Fidalgo Bay)  tend to
 be more diverse in summer and less diverse in fall and winter than food
 webs of the other  areas (Appendix Table E-l), mainly  because of  additional
 detritivores  (valviferan isopods, cumaceans^ and juvenile salmon in summer.
 Cherry Point had  the greatest connectance (X number of linkages  per node)
 during  the spring, Port Angeles had the greatest in summer and fall,  and
 Anacortes  the greatest  in winter.  In terms of numerical abundance,'however
 the Cherry Point  area  (Cherry Point, Lummi Bay, Birch Bay)  consistently


                                      45

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exhibited the highest density and standing crop of neritic fishes during
the DOE studies (Miller, et al., 1977; Fresh 1979) while the MESA
collections in the Port Angeles vicinity were comparatively lower than in
any other area (Simenstad, et al., 1977; Cross, et al., 1978).  The high
catches were principally due to high densities of postlarval Pacific herring
which result from the sizable herring stock spawning adjacent to Cherry
Point (Trumble, et al., 1977), although threespine stickleback and surf
smelt also contributed to the high catches in this area.  The Port Angeles
area is also qualitatively different from the other three in several other
ways, i.e., longfin smelt functionally replace surf smelt in the neritic
fish assemblage and the tufted puffin is prominent.

     The littoral habitats at Cherry Point, Burrows Bay, and Port Angeles
are generally exposed, gravel-cobble beaches which grade into sand and
sparse eelgrass in the shallow sublittoral zone.  Food webs at all
locations are based mainly on detritus and detritivorous crustaceans.
Overall, Port Angeles shows the highest number of food web nodes in the
spring and summer and Cherry Point appears to have the most diverse food
web in the fall and winter (Appendix Table E-7) • In terms of food web
connectance, however, the Port Angeles area appears to have the highest
average number of linkages per food web node during all seasons.  The
Cherry Point food web is based on gammarid amphipods which support a
variety of epibenthic carnivorous fishes, while the food webs at Burrows
Bay and Port Angeles are much more broadly based on detritivorous
crustaceans (gammarid amphipods, cumaceans, harpacticoid copepods, pagurid
crabs, hippolytid shrimp), suspension feeders (bivalves), and mixed food
processors (mysids).  In Burrows Bay and Port Angeles mysids become more
important than gammarid amphipods in the diets of several prominent epi-
benthic carnivores that occur in all areas (e.g., Pacific staghorn sculpin,
silverspotted sculpin).

     The mud/eelgrass habitat which dominates the shallow sublittoral zone
of March Point (Anacortes) has a much more diversified food web, possessing
half again as many nodes as the food webs of the other areas (Appendix
Table E-ll).  The average number of linkages per node is not appreciably
different (Appendix Table E-12).  Much of the increased complexity is due
to the numerous seabirds and shorebirds which occupy the mudflats throughout
the year.  The tremendous abundance of waterfowl (black brant, snow goose,
Canada goose) present from fall through spring feeding on eelgrass and
saltmarsh plants, also increase the complexity.  Benthic meiofauna and epi-
benthic detritivorous crustaceans and their predators effect the transfer
of detrital carbon to upper trophic levels.  Secondary consumers include
one of the most diverse assemblages of nearshore fish of any habitat
sampled during the DOE and MESA studies.

     One of the main differences between this food web structure and the
others is the extensive linkages between eelgrass and herbivorous waterfowl
and the large annual contribution of eelgrass to the total detritus pool.
                                      46

-------
     As regards vulnerability to ecological disruption from pollution,
March Point shows the greatest sensitivity because of the diverse
communities and productive food webs of its predominantly mud/eelgrass
habitats.  The unstructured sediments typical of this habitat would
probably entrain toxic petroleum components for a long time and they would
continue to leach into the epibenthic food web through the detritus-
decomposer linkage.  Loss of eelgrass would seriously limit the foraging
of wintering waterfowl and dramatically alter the accumulation of detritus,
the principal energetic source of the area's food web.

     Cherry Point is the second most vulnerable area because of its
extensive neritic food web.  No other area in north Puget Sound or the
Strait of Juan de Fuca produces as great a standing stock of post-larval
neritic  fishes, especially the commercially important Pacific herring.
These fish are apparently restricted to nearshore habitats during the
most sensitive stage of their life history, which means they would be
vulnerable to both acute and sublethal effects of petroleum hydrocarbons.
Overall, the north Puget Sound region is more vulnerable to long-term
adverse  effects of petroleum pollution because of the high proportion of
contained embayments and eelgrass and saltmarsh habitats characterizing
the nearshore region.

     In  the Strait of Juan de Fuca, the Port Angeles area exhibited the
most highly connected food webs, which suggests that it would be less
vulnerable to  instability caused by removal of or reduction in food web
linkages.  The food web of the immediate area is more complex and
productive than  the food web identified at Burrows Bay.  Also, increased
exposure and more consolidated sediments than March Point would allow more
weathering and degradation of spilled petroleum.  Fewer toxic compounds,
and in smaller amounts, would become incorporated in the sediments. The
habitats adjacent to Port Angeles increase the area's overall vulnerability,
however.  The  exposed gravel beach at Dungeness Spit and the mud/eelgrass
and saltmarsh habitats behind the spit and at Discovery and Sequim bays to
the east, are  areas that would potentially suffer long-term effects from
spilled  oil, equal to those postulated for March Point.  Dungeness Spit
also provided  the highest shallow sublittoral catches of juvenile salmon
of the sites along the Strait of Juan de Fuca.  These commercially
important  species may aggregate along the spit during their outmigration,
which would expose them to any pollutant transported east from Port Angeles.

     Burrows Bay characteristically had the least diverse and least
productive food web of the four areas.  The high energy of this embayment
would probably cause considerable weathering of spilled petroleum. However,
 the MESA nearshore fish collections at Alexander's Beach also documented
consistently high catches of juvenile chum salmon, suggesting that juvenile
 salmon migrating out of Skagit Bay through Deception Pass may aggregate
 in the bay.
                                     47

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V-F.  Utilization of Empirical Food Web Data for Modeling Energy
                        Flow in Marine Ecosystems

     Although analysis of the structure of food webs provides measurable
insights into the organization and importance of predator-prey linkages
and inter-trophic-level transfers, only a comprehensive bioenergetic
approach will ultimately produce clues to the dynamics of the system.
Only the bioenergetic approach can generate predictions of the effects
of perturbations on components of trophic levels.  Bioenergentic analysis,
however, requires relatively precise estimation of population density and
standing crop levels of component species or functional groups and
quanti fication   of energy flow through predator-prey linkages in order
to determine trophic, ecological, and population efficiency rates between
and within trophic levels.  Few natural systems have been empirically
analyzed to this degree.

     Although the limited data available for north Puget Sound and Strait
of Juan de Fuca nearshore and neritic food webs do not allow so comprehensive
an analysis, a preliminary analysis was performed on a component food
web identifiable at one site in north Puget Sound—the mud/eelgrass, con-
tained embayment habitat of Westcott Bay on San Juan Island.   Low exposure,
a relatively closed system, and DOE and other data (Thornburgh 1978) on
top consumers and their prey made feasible the assembling of a simple
energetic model for this site.  The bay is used by several species of
secondary consumers as a nursery area  and the species assemblage might
be considered typical of other mud/eelgrass sites in Puget Sound.

     The results of this exercise (Appendix G), although not the product
of a systematic experiment specifically designed and conducted to answer
such questions, have illustrated significant gaps in our understanding
of food web flow, ecological and assimilation efficiencies,  food consump-
tion and population integrity.  Appearance of a 40-50X error factor between
the annual consumption rate estimated from growth of juvenile English sole
and estimated from their calculated daily consumption indicates that our
basic  knowledge of gastric evacuationand assimilation efficiencies is
inadequate to meet the accuracy demanded by a bioenergetic model.
Similarly, the standard techniques for estimating the standing stock of
secondary consumers may also be inappropriate.  What the exercise does
show is that energy flow between prominent food web compartments (nodes)
is quite sensitive to feeding behavior, including frequency of feeding on
a  specific prey, actual prey biomass consumed, periodicity and intervals
in feeding, gastric evacuation rates, and the bioenergetic cost involved
in the feeding process itself.  Some of these functional relationships
can be accurately estimated from well-designed laboratory experiments, but
much will depend upon more precise quantification   of the feeding of
fishes and other high-level consumers in the environment.
                                    48

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APPENDIX A:  ALGAE AND INVERTEBRATES
                  55

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Taxon: Genera
Regional
  # of
species   Trophic level
                            References
Chlorophyta
  Volvocales: Clamydomonas,  Dunaliella,  Thalassomonas
  Ulotrichales: Blidingia, Enteromorpha,  Entocladia,
    Internofet-ia, Monostvoma, Percuvsia,  Phaeophila,
    Ulothrix, Ulva
  Schizogoniales: Pras-iola,  Rosenvingiella,  Schizogoniwn
  Cladophorales: Chaetomorpha,  Cladophora,  Lola,
    Nhizoolaniwn, Spongomorpha,  Urospora
  Chlorococeales: Chloroohytvium,  Codiolwn,  Gomontia
  Codiales: Bryopsis, Coditon, Derbesia,  Halicystis

Bacillariophyta: Pennales

Phaeophyta
    Agarwn, Alar-La, Analipus, Carpomitra,  Coilodesme,
    Colpomenia, Compsonema,  Costaria,  Cymathere,  Cystoseira,
    Desmarestia, Diatyoneuropsis,  Dictyoneurum,  Dictyo-
    siphon, Diatyota, Ectocarpus,  Egvegia,  Eisenia,
    Elaohista, Eudesme, Feldmania, Fucus,  Giffordia,
    Haplogloia, Hecatonema,  Hedophyllum,  Laminaria,
    Leathesia, Leptonematella,  Lessoniopsi-s, Maerocystis,
    Melanosiphon, Myrionema, Nereoaystis,  Pelvetiopsis,
    Petalonia, Petrodevma, Phaeostrophion,  Pleurophyaus,
    Postelsia, Pterygophora, Punataria,  Pylaiella, Ralfsia,
    Sargasswn, Saundersella, Scytosiphon,  Soranthera,
    Sphaoelaria, Sphaerotriohia, Spongonema, Stietyosiphon,
    Streblonema, Syvingoderma
   27
    3

   23
    5
    7
  102
          Primary producers
Primary producers

Primary producers
29


29
29

29
29
29

18

36

-------
Taxon: Genera	

Rhodophyta
    AaTfodhaetiwn, Aglaothamnion, Ahnfeltia, Amplisiphonia,
    Antithamnion, Antithamnionella, Arthroaardia,
    Asterocolax, Audouinella, Bangia, Besa, Bonnemaisonia,
    Bossiella,  Botryoaladia, Botryoglosswn, Branchio-
    glosswn,  Calliarthron, Callithamnion, Callocolax,
    Callophyllis, Cauloeanthus, Geranium, Chondrus,
    Choreooolax, Clathromorp'hwn, Coeloseira, Conohocelis,
    Constantinea, CoTa.11i.na, Cruoria, Cruoriopsis, Cryp-
    tonemia,  Cryptopleuva, Cryptosiphonia, Cwnagloia,
    Cumathamnion, Delesseria, Dermoeorynus, Dilsea,
    Endoaladia,  Erythroaladia, Erythrodermis, Erythro-
    glosswn,  Erythrophyllum, Erythrotviaia, Euthora,
    Farlouia, Fauchea,  Fauaheooolax, Fryeella, Gastvo-
    cloniion,  Geli-diwn,  Gigartina,  Gloiopeltis,
    Gloiosiphonia,  Gonimophyllum,  Goniotrichopsis,
    Goniotr-idhum,  Graoilaria, Gracilariophila,
    Gracilariopsis,  Grateloupia, Griffiths-La,
    Gymnogongrus,  Halosaecion, Halymenia, Harveyella,
    Hepposiphonia,  Heterosiphonia,  Hildenbrandia,
    Hollenbergia, Holmesia,  Hydrolithon,  Hymenema, Ividaea,
    Janezewskia,  Kallymenia,  Kylinia, Laurencia, Lepto-
    fauchea, Lithophyllum,  Lithothamniwn, Lithothrix,
    Lomentaria,  Lophosiphonia, Melobesia, Membranoptera,
   Mesophyllwn,  Microaladia, Myriogramma, Nemalion,
   Neoagardhiella,  Neodilsea, Neopolyporolithon,
   Neoptilota,  Nieburgia,  Nitophyllum,  Odonthalia,
    Opwitiella,  Petrocelis,  Peyssonelia,  Phyoodrys, Pikea,
    Platysiphonia, Platyfhamnion, Pleonosporiwn, Plocamium,
   Plocamiooolax, Polyneura,  Polyneuvopsis, Polysiphonia,
   Porphyra,  Porphyrella,  Porphyropsis,  Pvionitis, Pseudo-
   gloiophloea, Pseudolithophyllum,  Pterochondria, Ptero-
   oladia, Pterosip'honia,  Ptilota,  Ptilothamniopsis,
   Pugetia, Ehodoohovton,  Khodoglosswn,  Rhodomela,
312
Primary producers

-------
Taxon: Genera
Regional
  // of
species   Trophic level
References

    Khodophysema, Khodoptilum, Rhodymenia, Rhodymeniooolax,
    Savcodiotheoa, Scagel-ia, Scagelonema, Schizomenia,
    Serraticavdia, Smithora3 Stenogramme , Tenarea,
    Thuretellopsis, Tiffaniella, Trailliella, Twmerella,
    Weeks-la, Whidbeyella, Yamadaea

Spermatophyta
    Phy 1 lospadix, Zostera

Porifera
  Calcarea: Leucosolenia, Soypha
  Hexactinellida: Aphrocall-istes
  Demospongiae : Cliona, Ealichondfia, Haliclona, Micpoaiona
    Mycale, Myx-illa, Ophlitaspongia, Ploaamia, Suberites

Cnidaria
          Primary producers                16
          Suspension feeders on phyto-     3
    2+    plankton, bacteria, and small   19
    4     protozoans                      19

   39+                                    19
  Hydrozoa
    Hydroida: Abietinaria, Aequorea, Bougainvilla, Campanulina, 65+
      Clava, Coryne, Gavve-ia, Gonionemus, Hydraatin-ia,
      Phialidium, Plwnular-ia, Probosc-idaotyla, Sertularella,
      Sertularia, Tubularia
    Trachylina
    Hydrocorallina: Stylantheca
    Chondrophora: Velella
    Siphonophora
  Scyphozoa
    Discomedusae: Awfeli-a, Cyanea
    Stauromedusae: Haliclystus

  Anthozoa
    Stolonifera: Clavularia
    Alcyonacea:  Gersemia
    Gorgonacea:  Paragorgia
          Hydroids suspension feeders      3
          on zooplankton; Medusae carni-  19,13,
          vores on zooplankton            18
          Proboscidaotyla a commensal/
          parasite on sabellid
          polychaetes
    3                                     19
    1                                     19
    1                                     19
    5                                     19
          Primary/secondary carnivores      3
    7     on zooplankton and fish         19
    2     Primary carnivores on caprellid 19,18
          amphipods
          Primary/secondary carnivores on   3
    1     zooplankton, fish, and benthic  19
    1     invertebrates                   19
    1                                     19

-------
Taxon: Genera
Regional
  # of
species   Trophic level
                           References
    Pennatulacea: Ptilosarous                                   3

    Ceriantharia: Pachycerianthus                               1
    Zoanthiniaria: Epizoanthus                                  1
    Scleractinia: BaTanophyllia                                 2
    Actinaria: Anthopleura, Cribrinopsis, Diadianene,  Epiaatis,  20
      Halaampa, Metridium, Peachia, Stomphia,  Teal-la
Ctenophora: Beroe, Bolinopsis, Pleurobrachia
Platyhelminthes
  Cestoda
  Trematoda
  Turbellaria
    Acoelida
    Polycladida:  Kdburakia,  Notoplana
    Neorhabdocodida
    Proseriata:  Itaspiella
    Tricladida:  Procerodes

  Orthonectida
    34-
   20
    8+
    4+
          Primary carnivores on
          zooplankton
          Diadumene and Metvidiwn zoo-
          planktivores; Anthopleuva
          "farm" zooxanthellae, carni-
          vores on benthic crustaceans
          and dislodged mussels
          Primary carnivores on zoo-
          plankton; Beroe secondary
          carnivores on other
          ctenophores
Entoparasites
Ecto- and ento-parasites
Herbivores on diatoms; carni-
vores on small crustaceans
Herbivores on diatoms; carni-
vores on protozoans and
copepods
Carnivores on small crusta-
ceans, gastropods, worms, and
ascidians
Some commensals in sipunculans,
crinoids, echinoids, and
holothuroids
          Parasites in ophiuroids and
          polychaetes
                                 19

                                 19
                                 19
                                 19
                                 19,18,
                                 30,15
 3
 3
18

19,15,
 3

19,18,
15

19
                                19
                                19

                                19

-------
   Taxon: Genera
Regional
  # of
species   Trophic level
References
     Dicyemida                                                     4
     Nemertea: Amphiporus, CerebTatulus, Empleotonema, Lineus,    26+
       Ualaoobdella, Micrura, Paranemertes, Tubulanus
     Rotifera                                                      ?



     Gastrotricha                                                  7+

     Kinorhyncha                                                   7

g    Acanthocephala                                                ?



     Nematomorpha                                                  ?

     Nematoda                                                     83+



     Entoprocta: Barentsia                                         8
   Mollusca
     Amphineura
       Acanthochitonidae: Cryptoohiton                             1
       Callistoplacidae                                            2
       Chaetopleuridae                                             1
       Ishnochitonidae: Cyanoplax,  Ishnoohiton,  Tonioella         10
          Parasites in cephalopods        19
          Primary carnivores on poly-     19,18,
          chaetes, small molluscs, and    28
          crustaceans; Papanemertes up
          to 80% on Platynere-Ls;
          Malacobdella commensal  with
          Siliqua
          Suspension feeders on phyto-    19,3
          plankton; primary carnivores
          on protozoans,  rotifers, and
          other small metazoans
          Consumers of bacteria,  diatoms  19,3
          and small protozoans
          Detritivores;  herbivores on     19,3
          microalgae
          Parasites on isopods and        32,3
          amphipods as juveniles;
          marine mammals,  birds,  and
          fish as adults
          Parasites on decapod crusta-    32,3
          ceans
          Parasites; herbivores on        19,35,
          microalgae;  detritivores;        3
          primary carnivores on small
          metazoans
          Suspension feeders on phyto-    19,3
          plankton and small zooplankton
          Herbivores on microalgae
          Toniaella herbivores on
          microalgae and encrusting
          corallines
   18,3,15
     19,4
     19,4
     19,4
     19,4,
     18

-------
Taxon:  Genera
                                                            Regional
                                                              # of
                                                            species   Trophic level
                           References
    Lepidopleuridae
    Mopaliidae:  Ka.tha.rina,  Mopalia
      Plaaiphorella
Gastropoda
  Prosobranchia
    Acmaeidae: Acmaea,  Collisella,  Notoaomaea

    Lepetidae: Cryptobranchia
    Scissurellidae
    Haliotidae: Haliotis
    Fissurellidae: Diodofa, Puncturella
    Trochidae: Tegula,  Calliostoma, Margarites, Lirularia
   Turbinidae: ^straea., Homalopoma
   Lacunidae : Lacuna
   Littorinidae: Littorina
   Rissoidae: Alvinia, Barleei-a
   Assimineidae
   Vitrinellidae
   Caecidae
   Turritellidae
   Vermetidae

   Cerithiidae :  B-Lttiim,  Cevithiopsis
   Potamididae:  Batillaria
   Hipponicidae
   Calyptraeidae :  Calyptraea,  Crepidula, Crepipatella.

   Trichotropididae:  Tviohotropis

  Naticidae:  Natiaa,  Polinioes
                                                               2
                                                              15+
                                                              13

                                                               1
                                                               1
                                                               1
                                                               9
                                                              14
                                                               4
                                                               2
                                                               3
                                                               6
                                                               1
                                                               1
                                                               2
                                                               1
                                                               3

                                                               7+
                                                               1
                                                               2
                                                               7
Placiphorella carnivores on
amphipods and shrimp as well
as herbivores on microalgae
Herbivores on microalgae
and encrusting macroalgae
Herbivores on microalgae
Herbivores on microalgae
Herbivores on microalgae
Herbivores on microalgae
Herbivores on microalgae;
Calliostoma carnivores  on
hydroids
Herbivores on microalgae
Herbivores on microalgae
Herbivores on microalgae
Herbivores on microalgae

Commensals with chitons
Suspension feeders on  phyto-
plankton
Carnivores
Suspension feeders on phyto-
plankton
Suspension feeders on phyto-
plankton
Carnivores on thin-shelled
bivalves, e.g., Macoma
19,4
19,4,15
19,4,
32,18
19,4,32
19,4,32
19,4,32
19,4,32
32,19,
4,32,18

19,4,32
19,4,18
19,4,18
19,4
19,4
19,4,32
19,4
19,4
19,4,32

19,4,32
19,4
19,4
19,4,
32,15
19,4

19,4,
18,15

-------
   Taxon:  Genera
Regional
  # of
species   Trophic level
                           References
ON
       Lamellariidae
       Velutinidae
       Cymatiidae: Fusitriton
       Epitoniidae:  Epitonium,  Cpalia

       Eulimidae:  Balds
       Thyonicolidae
       Muricidae:  Cevatostoma,  Ocenebra,  Urosalpinx,
         Borcotrophon
       Thaididae:  Thais (Nucella)

       Buccinidae:  Buecinum
       Neptuneidae:  Bevingius,  Colus,  Neptunea,  Searlesia
       Columbellidae:  Amphissa,  Mitrella

       Nassariidae:  Nassarius
       Volutidae
       Mitridae
       Marginellidae:  Granulina
       Olividae: Olive I la

       Cancellariidae: Actmete
       Turridae: Oenopota,  Ophiodermella
    Opisthobranchia
       Bullidae: Volvulella
       Acteocinidae: Cylichna
       Aty i d ae:  Eami.no ea
       Philinidae
       Gastropteridae: Gastropteron
    5
    3
    9
    5
    1
    1
    1
    2

    2
   11

    1
    4
    1
    2
    1
Carnivores on ascidians
Carnivores on ascidians
Carnivores on StTPongy'Locent-
rotus, bivalves, chitons,
asteroids
Ectoparasites on anemones

Ectoparasites
Entoparasites in holothuroids
Carnivores on barnacles,
bivalves
Carnivores on small mussels
and barnacles
Carnivores
Carnivores on bivalves;
Searlesia carnivores and
scavengers on barnacles,
limpets, and chitons.
Scavengers on dead animal
tissue
Carnivores
Carnivores
Carnivores
Carnivores
Scavengers on dead animal
tissue
Carnivores
Carnivores on polychaetes
                                        19,4,32
                                        19,4,32
                                        19,4,10
19,4,
32,15
19,4,32
19,4
19,4,18

19,4,33

19,4
19,4,21
19,4,18

19,4
19,4
19,4
19,4
19,4,18

19,4
19,4,31

19,4
19,4
19,4
19,4
19,4

-------
                                                         Regional
                                                           # of
                                                         species   Trophic level
                                  References
  Aglajidae:  Aglaja
  Pyramidellidae:  Odostomia, Turbonilla
  Aplysiidae:  Phyllaplysia
  Clionidae: Clione
  Elysiidae: Elysia
  Stiligeridae: Hermaea
  Pleurobranchidae: Berthella
  Corambidae:  Corcwibe

  Dorididae: Anisodovis, Arshidoris, Cadlina, Diaulula,
   Diseodoris, Rostanga
  Onchidoridae: Acanthodoris, Onchidoris
 Polyceridae: Polyoera,  Triopha, Lai la
 Arminidae:  Armina
 Dendronotidae:  Dendronotus

 Dotonidae:  Doto
 Durauceliidae:  Duvaucelia (Tritoni-a)
Firabriidae: Melibe

Aeolidiidae: Aeolidia,  Hermissenda
Cuthonidae: Cratena, Cuthona
Dironidae: Dirona

Flabellinidae: Eubranchus
Zephyrinidae: Antiope'lla.
Onchldiidae: Onohidetla
21
 1
 2
 3
 4
 1
 2
 6
 1
 8

 3
 4
 1

 2
 4
 2

 7
 2
 1
Carnivores on polychaetes,
ostracods, amphipods,  and
small bivalves
Ectoparasites
Carnivores on bryozoans
(e.g., Membranipora)
Carnivores on sponges

Aaanthodoris carnivores  on
bryozoans, Onchidoris  on
barnacles
Carnivores on bryozoans
Carnivores on Pt-ilosarcus
Carnivores on hydroids,
D. irus on Paehycerianthus

Carnivores on alcyonaceans
and pennatulaceans
(Ptilosaraus)
Carnivores on small
crustaceans  (e.g., Caprella)
Carnivores: Aeoli,dia on
anemones, Hermissenda  on sea
pens, hydroids, and colonial
tunicates

Carnivores on bryozoans,
ascidians, and small snails
Carnivores on hydroids

Herbivores on microalgae
                                                                                                  19,4
  19,4,32
  19,4
  19,4
  19,4
  19,4
  19,4
  19,4,
  18,15
  19,4

  19,4,18
  19,4,18
19,4,18,5
  19,4,27

  19,4
  19,4,5
  19,4,18

  19,4,18
  19,4
  19,4,18

  19,4,18
  19,4
  19,4

-------
Taxon: Genera
Pulmonata
Ellobiidae: Phytia, Assiminea
Siphonariidae: Siphonaria
Bivalvia
Nuculidae

Nuculanidae: Nuculana, Yoldia

Malletiidae

Glycymeridae
Philobryidae
Mytilidae: Cvenella, Modiolus, Musculus, Mytilus
Pectinidae: Chlamys, Hinnites, Pecten
Anomiidae : Pododesmus
Limidae
Ostreidae: Cvassostrea, Ostrea
Lucinidae
Ungulinidae
Thyasiridae
Kelliidae: Kellia, Mysella
Leptonidae: Lasaea
Carditidae: Cardita, Glans
Astartidae
Cardiidae: Clinoaardiwn
Mactridae: Mactra, Spisula, Tresus
Solenidae: Siliqua* solen
Tellinidae: Macona, Tellina

Psammobiidae: Gari

Semelidae: Semele
Kelliellidae : Turtonia
Veneridae: Transennella, Gemma, Tapes (Venerupis),
Regional
// of
species

2
1

1

7

2

1
1
8
8
1
1
3
2
1
4
5
1
5
3
5
4
2
18

1

1
1
9
Trophic level References

Herbivores on microalgae
Herbivores on microalgae

Detritivores , deposit
feeders
Detritivores, deposit
feeders
Suspension feeders on phyto-
plankton
it
ii
ii
ii
ii
M
ii
ii
ii
ii
ii
ii
11
ii
ii
11
"
Detritivores, deposit
feeders
Suspension feeders on phyto-
plankton
ii
M
ii

19,4
19,4

32,19,
4,3
32,19,
4,3
19,4,3

19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3
19,4,3

19,4,3

19,4,3
19,4,3
19,4,3
Psephidia, Himilaria

-------
                                                            Regional
                                                              // of
                                                            species   Trophic level
                                  References
     Petricolidae

     Cooperellidae
     Myidae:  Cryptomya, Mya, Hiatella, Panopea
     Pholadidae:  Netastoma, Penitella, Xylophaga, Zirfaea
     Teredinidae: Bankia, Teredo
     Pandoridae
     Lyonsiidae:  Lyonsia, Mytilimeria
     Thraciidae
     Cuspidaridae: Cardiomya
   Scaphopoda: Dentalium, Cadulus
   Cephalopoda
     Sepiolidae: Eossia
     Loliginidae: Lol~igo

     Gonatidae: Gonatus

     Onychoteutidae:  Moroteuthis
     Octopodidae: Octopus

Annelida
  Polychaeta
    Errantia
      Amphinomidae
      Aphroditidae

      Arabellidae: Arabella, Drilonereis
      Chrysopetalidae:  Paleanotus
      Dorvilleidae:  Dorvillea,  Protodorvillea
      Eunicidae:  Eunice
      Euphrosinidae
1
9
6
2
5
4
4
2
6

1
1
1
3
1
5
1
8
2
      Suspension feeders on phyto-
      plankton
Detritivores
                                19,4,3

                                19,4,3
                                19,4,3
                                19,4,3
                                19,4,3
                                19,4,3
                                19,4,3
                                19,4,3
                                19,4,3
                                19,4,3
Carnivores on shrimp and  fish
Carnivores on small fishes
and shrimp
Carnivores on small fishes
and shrimp
                                19,4,7
                                19,4,
                                18
                                19,4,
                                18
                                19,4
Carnivores on crabs, bivalves,  19,4,
and gastropods                  18
                               2,12
Carnivores on polychaetes,     2,12
e.g., terebellids and sabellids
Juveniles parasites  in poly-   2,12
chaetes and echiurans; adults
carnivores
Carnivores                     2,12
Carnivores                     2,12
Carnivores on annelids,        2,12
ostracods, copepods, and
bivalves
                                      2,12

-------
Taxon: Genera
Glyceridae: Glyoeva, Hemipodus
Goniadidae: Glycinde, Goniada
Hesionidae: Gyptis, Kefersteinia., Mieropodarke ,
Oph-iodrorms
Lumbrineridae: Lwnbrineris
Regional
# of
species
8
6
4
13
Trophic level
Carnivores on polychaetes
Carnivores
Carnivores on annelids ,
cumaceans , and tanaids
Herbivores on micro- and
References
2,12
2,12
2,12
2,12
  Nephtyidae: Nephtys
13
  Nereidae: Cheilonereis, Micvoneveis, Nereis, Platynereis 17
  Onuphidae: Diopatra, Onuphis                              7
  Phyllodocidae: Eteone, Eulalia, Hesionura, Phyllodoce    27
  Pilargidae: Pilavgis, Sigambra                            3
  Pisionidae: Pisione                                       1
  Polynoidae: Arctoroe, Eunoe, Halosydna, Earmofhoe,
    Lepidasthen-ia, Tenon-la                                 29

  Polyodontidae: Peisidice                                  1
  Sigalionidae: Pholoe, Thalenessa                          4

  Sphaerodoridae: Sphaerodorops-is, Sphaerodorwn             3
  Syllidae: Autolytus, Eusyllis, Exogone, Odontosyllis,    37
    Pionosyllis, Sphaerosyllis, Streptosyllis, Syllides,
    Syllis, Trypanosyllis
Sedentaria
  Acrocirridae                                              1
  Ampharetidae: Ampharete, Asabellides                     18
  Apistobranchidae: Apistobranehus                          2
  Arenicolidae: Abarenioola, Arenicola, Branchiomaldane     4
macro-algae; carnivores;
detritivores
Carnivores on small benthic    2,12
animals
Herbivores on macroalgae       2,12
Herbivores on macroalgae;      2,12
scavenger on animal and plant
tissue
Carnivores on polychaetes and  2,12
other benthic animals;
scavengers
Carnivores or omnivores        2,12
Detritivores                   2,12

Carnivores; Harmothoe on       2,12
amphipods
                               2,12
Carnivores on small            2,12
invertebrates
Detritivores                   2,12
Autolytus, Syllis, Trypano-    2,12
syllis, carnivores; other
genera detritivores

Detritivores                   17,12
Detritivores                   17,12
Detritivores                   17,12
Detritivores                   17,12

-------



Taxon : Genera
Capitellidae: Barantolla, Capitello, Deaamastus,
MediomastuSj Notomastus
Chaetopteridae: Mesochaetopterus , Phyllochaetopterus,
Spioehaetopterus
Cirratulidae: Caulleriella, Chaetozone, Cirratulus,
Cirri formia, Dodeeaeeria, Tharyx
Cossuridae
Ctenodriidae
Disomidae: Trochochaeta
Flabelligeridae : Pherusa
Mage Ion idae: Magelona
Maldanidae: Axiothella, Euclymene, Maldane, Nioomaohe
Notoproctus
Opheliidae: Armotrypane, Armandia, Ophelia, Travisia
Orbiniidae: Naineris, Ovbinia, Protoariaia, Sooloplos
Oweniidae: Myviochele, Owenia
Paraonidae: Aricidea, Paraonella
Parergodrilidae : Stygocapitella
Pectinariidae: Pectinaria
Questidae: Questa
Sabellariidae: Idanthyrsus , Sabellaria

Sabellidae: Chone, Euchone, Eudistylia, Fabricia,
Jasmineira, Manayunkia, Oriopsis, Potamilla, Sabella
Sabe 1 las tarte, Schizobranohia
Scalebregmidae : Scalibregma
Serpulidae: Pseudoahitinopoma, Serpula, Spirorbis

Spionidae: Laonioe, Malaeoceros , Polydora, Pv-Lonospio,
Pygospio, Scolelepis, Spio, Spiophanes
Sternaspidae: Sternaspis
Terebellidae: Eupolurmia. Neoamphitrite, Nicoleas

Regional
// of
species
13

4
17

1
2
1
6
2
22

10
10
2
11
1
3
1
2

31
i

3
22

34

1
31
— . 	 — 	 	


Trophic level
Detritivores

Suspension feeders
detritivores
Detritivores


Detritivores

Detritivores
Detritivores
Detritivores

Detritivores
Detritivores
Detritivores
Detritivores

Detritivores

Suspension feeders
plankton
Suspension feeders
plankton and small

Detritivores
Suspension feeders
plankton


References
17,12

and 17,12
17,12

17,12
17,12
17,12
17,12
17,12
17,12

17,12
17,12
17,12
17,12
17,12
17,12
17,12
on phyto- 17,12

on phyto- 17,12
zooplankton

17,12
on phyto- 17,12

Detritivores; carnivores on 17,1*
zooplankton
Detritivores
Detritivores

17,12
17,12
  Pista,  Poly cirrus, Proalea, Soionella, Thelepus
Trichobranchidae
17,12

-------
Taxon: Genera
Regional
  # of
species   Trophic level
                                                                                                  References
  Archiannelida
      Polygordidae: Polygordius
      Protodrilidae: Protodrilus, Protodriloid.es
      Nerillidae: Nerilla
      Saccocirridae: Saccocirrus
      Dinophilidae: Trilobodrilus3 Diurodrilus, Dinophilus
  Oligochaeta
      Enchytraeidae: Enchytraeus, Lianbricillus
      Naididae: Nais, Paranais

      Tubificidae: Pelosoolex, Monopylephorus, Lirmodrilus
  Hirudinea
Priapulida: Priapulus caudatus

Sipuncula: Themiste, Pascolosoma, Golfingia

Echiura: Echiurus, Arhychite, Nellobia

Tardigrada: Echiniscoides, Batillipes
Pycnogonida: Aahelia, Halosoma, Nymphon, Phoxichilidiwn,
  Pyanogonwn
Arachnida
  Pseudoscorpionida: Halobisium
  Acari, Halacaridae
Crustacea
  Cladocera: Podon, Evadne
  Ostracoda
    1
    2
    3
    1
    3
    1

    4

    3

    2


   20
    5
   42
Detritivores                   19,12
Detritivores                   19,12
Detritivores                   19,12
Detritivores                   19,12
Detritivores                   19,12

Detritivores                   32
Detritivores; carnivores on    32
small mesozoans
Detritivores                   32
Carnivores, parasites on       32
fishes, shrimp

Carnivores on polychaetes      19,32,
and priapulids                 3
Detritivores                   19,18,
                               32
Detritivores                   19,32

Herbivores (suctorial) on
macroalgae

Juveniles parasites on cnid-   19,32
arians; adults carnivores on
hydroids, anemones, mussels
          Carnivores on mites            32,19
          Herbivores (suctorial) on      19,32
          algae; carnivores
Carnivores on copepods          19,3
Detritivores                    19,3

-------
Regional
  # of
species
                                                                    Trophic levej^
                                                                                                References
Copepoda
  Calanoida:  Calanus,  Centropages,  Acartia,  Eucalanus,        75
    Farac-alanus,  Pseudocalanus,  Microcalanus,
    Sjaphocalanus,  Metridia
  Harpacticoida:  Harpacticus,  Zaus,  Tisbe,  Oxyjulis,          98+
    Porcellidium,  Amonardia,  Diosaccus, Anrphiascopsis,
    A'nphiascus,  Daotylopodia,  Parathalestris,  Diarthrodes
  Cyclopoida:  Oncaea,  Corycaeus,  Clausidium, Oithona          23+
   Monstrilloida                                               A
   Caligoida                                                   ?
 Branchiura: Argulus                                           1
 Cirripedia
   Thoracica: Pollicipes, Lepas, Chthamalus, Balanus          21

   Acrothoracica                                               1

   Rhizocephala                                                8

Leptostraca: Nebalia                                          2

Mysidacea: Mysis, Archaeomysis, Holmesiella, Neomysis,       30
  Acanthomysis
Cumacca: Lampvops, Diastylopsis, Leptostylis, Diastylis,     31
  Eudorella,  Leptocuma, Cumella
Tanaidacea:  Fancolus,  Anatanais, Leptochelia                  5

Isopoda
  Epicaridea
    Bopyridae:  Hemiarthrus, Pseudione, Argeia, Phyllodurus    6

    Cryptoniscidae:  Cryptothir                                1
    Entoniscidae: Portunion                                   1
  Valvifera
    Idoteidae:  Edotea,  Idotea, Synidotea                     16
  Anthuridea
    Anthuridae:  Colanthura, Paranthura                        2
          Suspension feeders on
          phytoplankton

          Detritivores
                                                                                                    19,3,9
                                                                                                    19,3,9,
                                                                                                    20
          Suspension feeders on          19,9
          phytoplankton; commensals with
          benthic invertebrates
          Parasites in polychaetes       19
          Ectoparasites on fish          19
          Ectoparasites on fish          19,3,32

          Suspension feeders on phyto-   19,3
          plankton and small zooplankton
          Suspension feeders on phyto-   19,3
          plankton
          Parasites on decapod           19,3
          crustaceans
          Suspension feeders on detritus 19,3
          and microalgae
          Suspension feeders on          19,3
          microalgae and detritus
          Detritivores                   19,3

          Detritivores; carnivores on    19,3
          small benthic animals
          Parasites on shrimp and hermit 19
          crabs
          Parasites in barnacles         19
          Parasites in Hemigrapsus       19
          Detritivores
          Scavengers on macroalgae       19,18
          Detritivores
                                         19

-------
Taxon: Genera
Regional
  # of
species   Trophic level
                           References
    Flabellifera
      Aegidae: Rocinela,  Aega
      Cirolanidae: Cirolana
      Cymothoidae: Lironeca
      Limnoriidae: Lirnnoria

      Sphaeromatidae:  Dynamenella,  Exosphaeroma,
        Gnorimosphaeroma
    Microcerberidea
      Microcerberidae: Micro cert ems
    Asellota
      Jaeropsidae: Jaeropsis
      Janiridae: Janiropsis, Janiralata
      Munnidae:  Munna, Munnogonium
    Oniscoidea
      Armidillidiidae: Armadillidiwn
      Ligiidae:  Ligia
      Oniscidae: Alloniscus} Porcellia
      Scyphacidae: ArmadilloniscuSj  Detonella
  Amphipoda
    Hyperidea

    Gammaridea
      Acanthonotozomatidae:  Panoploea
      Ampeliscidae: Ampelisca,  Byblis
      Amphilochidae: Amphilochus
      Ampithoidae: Ampithoe, Cymadusa
      Aoridae: Aor>oides
      Argissidae:  Argissa
      Atylidae:  Atylus
      Calliopiidae: Calliopiella,  Oligochinus
      Corophiidae: Covoph-ium,  Ericthonius
      Dexaminidae: Guemea,  Polycheria
      Eusiridae: Accedomoera,  Paramoeva, Pontogeneia
    5
    3
    2
    2
   10

    1

    3
    7
    6

    1
    1
    3
    2
    1
    6
    2
    6
    1
    1
    3
    2
    7
    2
   11
Detritivores; parasites
Ectoparasites on fishes
Scavengers on animal tissue

Boring "consumers" of kelp
and wood

Scavengers on macroalgae
Detritivores

Detritivores
Detritivores

Scavengers on macroalgae
Carnivores on planktonic
cnidarians

Piercing/sucking mouthparts
Detritivores
Detritivores
Herbivores on macroalgae
Detritivores
Detritivores
Detritivores
Detritivores
Detritivores
Detritivores
Detritivores; carnivores
19,3
19,32
19
19,18
19,18

19

19
19
19

19
19,18
19
19

19,3
34
34
34
34
34
34
34
34
34
34
34

-------
Regional
# of
Taxon: Genera

Gammaridae : Anisogarmarus , Ceradocus, Maera,
Megaluropus , Melita
Haustoriidae: Eohaustorius
Hyalidae: Allorchestes , Hyale,


Para I torches tes
species
9

2
8
Trophic level References
Detritivores; Anisogarmarus
herbivores
Subsurface detritivores
Detritivores; herbivores on
34

34
34
     Isaeidae: Gammaropsis, Kermystheus,  Photis,
       Podoceropsis, Protomedeia
     Ischyroceridae: Isahyroeevus,  Jassa,  Microjassa,
       Microjassalitotes

     Leucothoidae:  Leucothoe

     Lysianassidae: Anonyx, Hippomedon, Lepidepecreum,
       Orehomene, Pachynus
     Nainidae:  Naina
 9

 5


 1

10

 1
     Oedicerotidae: Monoculodes,  Synchelidium,  Westwoodilla     7
     Phoxocephalidae:  Heterophoxus, Metaphoxus, Paraphoxus,    17
     Pleustidae: Parapleustes,  Pleusirus,  Pleustes,  Pleusymtes 6
     Podoceridae: Dulichia, Podocerus                           2
     Stenothoidae: Metopa, Stenothoides                         2
     Synopiidae: Tiron                                          1
     Talitridae: Orohestia, Orchestoidea                        3
  Caprellidea: Caprella, Cerops, Deutella, Metacaprella,
    Tritella
Euphausiacea: Euphausia, Thysanoessa

Decapoda
    Penaeidae:  Gennadas
    Sergestidae: Sevgestes
    Pasiphaeidae: Pasiphaea
    Oplophoridae: Hymenodora, Notostomus
26
13
 1
 1
 1
 2
macroalgae
Suspension feeders on
detritus; detritivores
Suspension feeders on
detritus; herbivores;
detritivores
Detritivore (commensal  in
sponges)
Detritivores; carnivores
34

34


34

34

34
Detritivore  (burrowing  in
Alaria)
Detritivores                    34
Subsurface detritivores         34
Detritivores                    34
Suspension feeders on detritus  34
Detritivores                    34
Detritivores                    34
Detritivores; scavengers on     34,18
macroalgae
Herbivores on microalgae;       19,32,
detritivores; carnivores on     18,8
harpacticoid copepods, amphi-
pods, and ostracods
Suspension feeders on phyto-    19,3,1
plankton

Carnivores on pelagic animals   19,3
Carnivores on pelagic animals   19,3
Carnivores
Carnivores                      19

-------
    Taxon: Genera
Regional
  # of
species
Trophic level
References
-vl
ho
          Pandalidae: Pandalopsis, Pandalus                        10
          Crangonidae: Argis, Paracvangon, Sclerocrangon, Crangon  18
          Alpheidae: Betaeus                                        2
          Hippolytidae: Hippolyte, Lebbeus, Eualus, Spirontocaris ,
            Heptacavpus
          Axiidae: Aociopsis
          Callianassidae: Callianassa, Upogebia
          Hippidae: Emerita
          Galatheidae: Munida, Munidopsis
          Porcellanidae: Paahycheles, Petrolisthes

          Lithodidae: Plaoetron, Hapalogaster, Oedignathns,
            Cvyptolithodes, Phyllolithodes, Lopholithodes
          Diogenidae: Pagwistes
          Paguridae:  Elassoohirus, D-iscorsopaguicus, Labidoehirus
            Orthopagurus,  Pagurus
          Parapaguridae: Parapagurus
          Majidae:  Chionoecetes, Choril-ia,  Eyas, Mimulus,
            Oregonia, Pugettia,  Scyva
          Cancridae:  Cancer,  Telmessus

          Xanthidae:  Lophopanopeus
          Pinnotheridae: Fabia,  Pinnixa,  Pinnotheres,  Scleroplax

          Grapsidae:  Hemigrapsus, Planes

    Insecta
      Collembola: Anurida
      Thysanura: Neomachilis
      Diptera
          Tipulidae:  Limonia
          Chironomidae:  Paraclwiio
   32
    1
    3

    1
    2
    4

   10
    2
   25

    1
   10
    1
   10
    1
    1
Carnivores
Carnivores
Carnivores

Carnivores
                                         19
                                         19
                                         19
Detritivores by suspension
feeding
Suspension feeders             19
Carnivores                     19
Suspension feeders on detritus 19
and plankton
Carnivores, Lopholithodes on   19
asteroids; Hapalagoster and
Oedignathus suspension feeders
on detritus and plankton
Detritivores                   19
Detritivores                   19
    19
    19
    19,18
Detritivores
Herbivores on macroalgae and
eelgrass
Carnivores; C. magister on
bivalves

Parasites of bivalves,
polychaetes
Scavengers on animal matter
Scavengers on macroalgae
          Larvae macroalgae herbivores
          Larvae macroalgae herbivores
    19
    19,8

    19,18

    19
    19,18

    19
    32
    32

    32
    32

-------
  Taxon: Genera
                                                              Regional
                                                                # of
                                                              species
          Dolichopodidae: Aphrosylus                                ?
          Helcomyzidae
        Coleoptera
          Staphylinidae: Diaulota,  Emplenota,  Liparocephalus,        ?
            Pontomalota
          Salpingidae: Aegialites                                   '

    Phoronida: Phoronis, Phoronopsis                                3

    Bryozoa (Ectoprocta)
      Ctenostomata:  Alcyonidium,  Bowerbankia,  Buskia,  Clavopora,     7+
        Flustrella,  Nolella,  Triticella
      Cyclostomata:  Bicrisia,  Crisia,  Crisidia,  Diaperoecia,        19+
        Diplosolen,  Discocytis, Disporella,  Filicrisia,
        Filifascigera,  Heteropora,  Idmonea,  Lichenipora,
£{       Oncousoecia, Plagioecia,  Proboscina, Stomatopora,
        Tubulipora
      Cheilostomata: Aetea, Alderina,  Arthropoma,  Bugula,  Caberea,  55+
        Caulibugula, Caulorarrrplius,  Cellaria,  Cheilopora,
        Codonellina, Colletosia,  Conopewn, Copidozown,
        Corynoporella,  Costazia,  Cribrilina, Dendrobeania,
       Doryporella, Electra, Ellisina, Eurystomella,Fenestratulina,
       Hincksina, Hippodiplosia,  Hippomonavella,  Hippothoa,
       Holoporella, Lagenipora, Lyrula, Mentoranipora, Miaropora,
       Mieroporella, Microporina, Mucronella, Myriozown,  Para-
       smittina,  Phidolopora, 'Porella, Puellina,  Reginella,
       Rhamphostomella, Schizomavella, Sohizoporella, Serupo-
       aellaria,  Smittina,  Stephanosella, Tegella, Termino-
       flustra,  Tricellaria,  Trypostega, Umbonula

  Brachiopoda:  Hemithyris,  Laqueus, Terebratalia, Terebratulina    4
                                                                        Carnivores on chironomids
                                                                        Scavengers on macroalgae

                                                                        Carnivores on dipteran
                                                                        larvae
                                                                        Suspension feeders on
                                                                        phytoplankton

                                                                        Suspension feeders on
                                                                        phytoplankton
                                                                        Suspension feeders on
                                                                        phytoplankton
                                                                        Suspension feeders on
                                                                        phytoplankton
                            References

                                32
                                32

                                32

                                32

                                19,3


                                19,3

                                19,3




                                19,3
                                                                        Suspension feeders on
                                                                        phytoplankton
                                19,3
Echinodermata
  Crinoidea: Florometra
Detritivores
                                                                                                         19,3

-------
Taxon: Genera
Regional
  # of
species   Trophic level
                           References
  Ophiuroidea: Amphiodia, Amphiphol-is , Dianrphiodia, Govgono-   25
    cephaluSj OphiophoUs> Ophiura, Unioplus
  Asteroidea: Cvossaster, Dermasteriast EvasteriaSj Henvicia,  19
    Hippasteria, Leptasterias, Luidia, Mediaster, Orth-
    asteifias, Pisaster, Pyenopodia, Solaster
  Echinoidea: Dendvastev, Strongyloaentrotus
  Holothuroidea: Cucwnavia, Eupentacta, Leptosyna.pta,
    Molpadia, Pavastiehopus, Psolus, Thyone
Chaetognatha: Eukrohnia, Sagitta
Hemichordata: Saecoglossus

Chordata-Urochordata
  Larvacea: Fritillaria, Oikopleura
   13
Detritivores                    19,3

Crossaster carnivores on        19,5,
Ptilosaraus and opistho-        18,26,
branchs; Devmastevias on        23,24,
Ptilosarcus, actinians,         25,22
sponge and holothurians;
Hippasteria on Ptilosavcus;
Leptastevias on barnacles,
limpets, Lacuna, Littorina,
chitons, holothurians;
Mediaster on Ptilosarous, algae,
detritivore; Pisaster on
Mytilus, barnacles, limpets,
chitons; Pyanopodia on large
bivalves, sea urchins; Solaster
dawsoni on asteroids; Solaster
endeca and S. stimpsoni on
holothurians
Dendraster detritivores;        19,3
Strongylocentvotus  herbivores
on macroalgae
Leptosynapta and Parastiohopus  19,3,
detritivores; rest suspension   11
feeders on detritus and
phytoplankton

Carnivores on zooplankton and   19,3
small fish

Detritivores                    19,3
          Suspension feeders on           19,3
          phytoplankton

-------
Taxon: Genera
                     P    ^
                                          , Ascidia, Boltenia,
                                        Cnemidocarpa,  Corella,
                      D^p^oSoma,  Distaplia, Haloeynthia,
                                           ,  Pycnlclavella,
            Salpidae

            Doliolidae
                                                                  Regional
                                                                    // of
                                                                  species   Trophic level
                                                               42      Suspension feeders on
                                                                      phytoplankton
                                                                     Suspension  feeders  on
                                                                     phytoplankton
                                                                     Suspension  feeders  on
                                                                     phytoplankton
                                                                                                 References
19,3
19,3

19,3
-vl
Ln

-------
                                 REFERENCES
 1.   Banner,  A.H.   1950.   A taxonomic study of the Mysidacea and Euphausiacea
        (Crustacea)  of the northeastern Pacific.   Part III.   Euphausiacea.
        Trans.  Royal Canad. Inst.  28:1-63.

 2.   Banse,  K.,  and K.D.  Hobson.   1974.  Benthic  errantiate  polychaetes of
        British Columbia  and Washington.   Bull.  185,  Fish.  Res.  Board Can.
        Ill  pp.

 3.   Barnes,  R.D.   1963.   Invertebrate zoology.   W.B. Saunders Co.,
        Philadelphia.   632 pp.

 4.   Bernard,  F.R.   1970.   A distributional checklist of the marine molluscs
        of British  Columbia; based on faunistic  surveys since 1950.
        Syesis  3:75-94.

 5.   Birkeland,  C.E.  1970.  Consequences of differing reproductive and
        feeding strategies for  the dynamics and  structure of an  association
        based on the single prey  species, Ptilosarus  garneyi (Gray).
        Ph.D.  Thesis,  Univ. Washington, Seattle.   99  pp.

 6.   Bloom,  S.A.  1974.   Resource  partitioning among  the doridacean nudibranch
        molluscs of the San Juan  archipelago,  Washington—a  guild hypothesis.
        Ph.D.  Thesis,  Univ. Washington, Seattle.   157 pp.

 7.   Brocco,  S.L.   1971.   Aspects  of the  biology  of the sepiolid squid Rossia
        pacifica Berry.   M.Sc.  Thesis, Univ. Victoria, British Columbia.   151  pp.

 8.   Caine,  E.   1978.   Univ. Washington,  Friday Harbor Laboratories.


 9.   Davis,  C.C.  1949.   The pelagic Copepoda of  the  Northeastern Pacific
        Ocean.   Univ.  Wash. Publ.  Biol. 14:1-117.

10.   Eaton,  C.M.  1971.   The reproductive and feeding biology of the prosobranch
        gastropod Fusitriton oregonensis  (Redfield) (Family  Cymatiidae).
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11.   Engstrom,  N.A.   1974.  Population dynamics  and prey-predator relations of
        a dendrochirote holothurian, Cucumaria lubrica, and  sea  stars in  the
        genus Solaster.   Ph.D.  Thesis, Univ. Washington, Seattle.  144 pp.

12.   Fauchald,  K.,  and P.A. Jumars.  1979.   The  diet  of worms:  A study of
        polychaete  feeding guilds.  MS.

13.   Fraser,  C.McL.   1937.  Hydroids of the Pacific coast of Canada and the
        United States.  Univ. Toronto Press, Toronto.  207  pp.
                                      76

-------
14.   Conor,  J-J-   1964.   Structure and function of the digestive system of
        Aglaja diomedea  Bergh,  a marine snail (Opisthobranchia, Cephalaspidea).
        Ph.D.  Thesis,  Univ.  Washington, Seattle.  254 pp.

15.   Hedgepeth, J.W.   1968.   Between Pacific tides.  4th ed.  Stanford Univ.
        Press, Stanford.  614 pp.

16.   Hitchcock, C.L.,  and A. Cronquist.  1973.  Flora of the Pacific
        Northwest.  Univ. Washington Press, Seattle.  730 pp.

17.  Hobson, K.D., and K. Banse.  MS.  Benthic sedentariate polychaetes of
        British Columbia and Washington.

18.  Kozloff,  E.N.  1973.  Seashore life of Puget Sound, the Strait of Georgia,
        and the San Juan Archipelago.  Univ. Washington Press,  Seattle.   282  pp.

19.  Kozloff,  E.N.  1974.  Keys to the marine invertebrates of  Puget  Sound,
        the San Juan Archipelago, and adjacent regions.  Univ.  Washington
        Press, Seattle.  226 pp.

 20.  Lang,  K.  1965.  Copepoda Harpacticoidea from the California  Pacific
        coast.  Kungl.  Sverska Vetenskapakad. Handl. ser. 4, 10(2):1-560.

 21.  Londa,  S.  1968.   Student  report, Friday Harbor Laboratories, Univ.
        Washington, Friday  Harbor.

 22.  Lubchenko, J.A.  1971.  Resource  partitioning between  two  intertidal
        predaceous asteroids,  Pisaster ochraceus  Brandt and Leptasterias
        hexactis  (Stimpson).   M.Sc.  Thesis, Univ. Washington,  Seattle.   36 pp.

 23.  Mauzey,  K.P. 1966.  Feeding behavior and  reproductive cycles in
        Pisaster  ochraceus.  Biol.  Bull.  131:127-144.

 24.  Mauzey,  K.P., C. Birkeland,  and P.K.  Dayton.  1968.  Feeding  behavior of
        asteroids and escape responses  of  their prey in  the Puget  Sound
        region.   Ecology 49:603-619.

 25.  Menge,  B.H.  1970.   The population ecology and community  role of the
        predaceous asteroid, Leptasterias  hexactis  (Stimpson).   Ph.D. Thesis,
        Univ.  Washington, Seattle.   213 pp.

 26.  Paine,  R.T.  1974.   Intertidal  community structure.  Oecologia  15:93-120.

 27.  Robilliard,  G.A.   1971.   Natural  history, niche exploitation, and
        coexistence in  the  genus  Dendronotus  (Mollusca:  Opisthobranchia).
        Ph.D.  Thesis, Univ. Washington, Seattle.  170 pp.

 28.  Roe,  P.   1971.   Life history and  predator-prey interactions of  the
        nemertean Paranemertes peregrina  Coe.   Ph.D. Thesis, Univ. Washington
         Seattle.  129 pp.                                                      '
                                      77

-------
29.  Scagel, R.F.  1966.  Marine algae of British Columbia and northern
        Washington, Part I:  Chlorophyceae (green algae).  Bull. 207,
        Biol. Ser. No. 74, National Mus. of Canada, Ottawa.  257 pp.

30.  Sebens, K.P.  1977.  Habitat suitability, reproductive ecology, and
        the plasticity of body size in two sea anemone populations
        (Anthopleura elegantissima and A. xanthogrammica).   Ph.D. Thesis,
        Univ. Washington, Seattle.  258 pp.

31.  Shimek, R.L.  1977.  Resources utilization and natural history of some
        northeastern Pacific Turridae.  Ph.D. Thesis, Univ. Washington,
        Seattle.  216 pp.

32.  Smith, R.I., and J.T. Carlton, eds.  1975.  Light's manual:  Intertidal
        invertebrates of the central California coast.  Univ. California
        Press, Berkeley.  716 pp.

33.  Spight, T.M.  1972.  Patterns of change in adjacent populations of an
        intertidal snail, Thais lamellosa.  Ph.D. Thesis, Univ. Washington,
        Seattle.  308 pp.

34.  Staude, C.  1978.  Univ. Washington, Friday Harbor Laboratories.


35.  Wieser, W.  1959.  Free-living nematodes and other small invertebrates
        of Puget Sound beaches.  Univ. Wash.  Publ. Biol. 19:1-179.

36.  Widdowson, T.B.  1973.  The marine algae of British Columbia and northern
        Washington:  Revised list and keys.  Part I:  Phaeophyceae (brown
        algae).  Syesis 6:81-96.

37.  Widdowson, T.B.  1974.  The marine algae of British Columbia and northern
        Washington:  Revised list and keys.  Part II:  Rhodophyceae.
        Syesis 7:143-186.
                                      78

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                             APPENDIX B:   FISHES


      Numerical classification of the nearshore fish collection data from
northern Puget Sound and the Strait of Juan de Fuca indicated that 64 species
composed the principal components of the region's nearshore fish communities
(Table B-l).  Sixteen assemblages were distinguished in this analysis,
including two neritic, eleven shallow sublittoral, two littoral, and one
(rocky) sublittoral assemblage.  Two species, copper rockfish and tidepool
sculpin.  occurred in more than one assemblage.

      Identification of functional feeding groups based on the DOE and MESA
food habits data based and pertinent literature, summarized in the following
species accounts section, indicated that facultative epibenthic planktivores
and benthivores, 26 and 16 species, respectively, predominated (Table B-l).*
Obligate epibenthic planktivores and facultative pelagic planktivores were
secondary  in importance.

      In the following species accounts of food habits, unless cited otherwise,
references  to data collected in northern Puget Sound resulted from the DOE
baseline studies (Miller, et al., 1977; Moulton, 1977) and in the Strait of
Juan de F-aca, under the auspices of NO\A's Puget Sound MESA program
(Simenstad  et al. 1977; Cross, et al., 1978).

                              Species Accounts

Spiny Dogfish

      Spiny dogfish examined during DOE studies in northern Puget Sound had a
high percentage  (71%) of the stomach contents digested and unidentifiable.
Epibenthic  gammarid amphipods  and shrimp and benthic gastropods dominated the
diet numerically, while Pacific sand lance (Ammodytes hexapterus) contributed
95% of the total consumed (identifiable) biomass.  The one spiny dogfish from
the eastern region  sampled by  WWSG contained four gammarid amphipods, one
polychaete annelid, and one  turbellarian.

      The  most comprehensive compilation of food habits data for spiny dogfish
in northern Puget Sound was  that reported by Jones and Geei (1977) for British
Columbia waters, principally the Strait of Georgia.  Analysis of almost 15,000
stomachs over  30 years indicated that Pacific herring  (Clupea harengus pallasi)
euphausiids, unidentified eggs, and caridean crustaceans were the principal
prey organisms for  all life history stages combined (Table B-2).
        See Methods  section  in body of report for definition of  terms describing
 feeding types.
                                      79

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Table  B-l.   Representative nearshore fish assemblages  of  northern  Puget
               Sound and the Strait of Juan de  Fuca  and  functional feeding
               groups of component species.   See methods  section  for  assem-
               blage determinations and definition of feeding  groups;  F =
               facultative  feeder, 0  = obligate feeder.
         Species
   Zone  Assemblage
  Component
   Species
                  Functional Feeding Group

  Pelagic    Epibenthic  Epibenthic  Meiobenthic
Planktivore  Planktivore Benthivore  Benthivore  Omnivore
Neritic
                     Pacific herring,
                     Clupea harengus
                     pa I lasi *
                     Threespine  stickle-
                     back, Gasterosteus
                     aculeatus
                     Pacific sand lance,
                     Aimodytes hexapterus
                     Pink salmon, Oncor-
                     hynchus gorbuscha*
                     Coho salmon, 0.
                     kisutch*
                     Chinook salmon, 0.
                     tshawytscha*
                     Tadpole sculpin,
                     Psychrolutes paradoxsus
                     Surf smelt, Hypomesus
                     pretiosus
                     Longfin smelt, Spirin-
                     chus thaleichthys
                     Northern anchovy,
                     Engraulis mordax
                     Soft sculpin
                     Gilbertidia sigalutes
                       F?

                       0?
Shallow     3
Sublittoral
Rock sole, Lepidopsetta
bilineata
C-0 sole, Pleuronichthys
coenosus
English sole,  Parophrys
vetulus*
Shiner perch,  Cymatogaster
aggregate
Pile perch, Rhacochilus
vacaa
                                            80

-------
Table B-l  (continued).
   Zone
          Species
         Assemblage
Component
 Species
                    Functional Feeding Group

  Pelagic    Epibenthic   Epibenthic  Meiobenthic
Planktivore  Plauktivore  Benthivore  Ber.thivore  Omnivore
                       Pacific tomcod,
                       Microgadus proximus'
                       Sturgeon poacher,
                       Agonus acipenserinus
                       Roughback sculpin,
                       Chitonotis pmjetensis
                                               f

                                               F
                       Speckled sanddab,
                       Cithariehthy3
                       stigmaeus
                       Pacific sanddab,
                       C. sordidus

                       Staghorn sculpin,
                       Leptocottus armatus
                       Starry flounder,
                       Platichthys staltatus
                       Snake prickleback,
                       Lwirpenus sagitt^
                       Tidepool sculpin,
                       Oliaocottus maculosus
                        Sand  sole,  Psettich-
                        thys  melanostictus*
                        Redtail surfperch,
                        Amphisticus frenatus
                        Chum salmon,  Oncor-
                        hynchus keta*
                        Penpoint  gunnel,
                        Ai'G,K*'rithys  flavidus
                        Cresent gunnel,
                        Pholis laeta
                        Silverspotted  sculpin,
                        Blepsias  cirrhosus
                        Whitespotted greenling,
                        Hexagrammos  stelleri
                        Tubenose  poacher,
                        Pallasina barbata
                                                   81

-------
Table B-l  (continued).
          Species
   Zone  Assemblage
Component
 Species
                    Functional Feeding Group

  Pelagic    Epibenthic   Epibenthic  Meiobenthic
Planktivore  Planktivore  Benthivore  Benthivore  Omnivore
           10         Padded sculpin,
                      Artedius  fenestralis
                      Buffalo sculpin,
                      Enophrys bison
                      Great sculpin,
                      Myoxocephalus
                      po lyacan thoceplTalus
                      Sharpnose sculpin,
                      Clinocottus acutiaeps
            ]1         Tube-snout, Aulor-
                      hynchus flavidus
                      Bay pipefish, Syngn-
                      athus griseolineatus
                      Cabezon, Scorpaen-
                      ichthys marmoratus'
            12         Copper rockfish,
                      Sebastes caurinus*
                      Smoothhead sculpin,
                      Artedins lateralis
                      Manacled sculpin,
                      Synahirus gilli
                                  F

                                  0

                                  0?
            13         Walleye pollock, .
                      Theragra chalcograjvma
                      Spiny lumpsucker,
                      Eiarricrotremus orb-is
                      Spiny dogfish,
                      Squalus acanthias
                      Ratfish, Hydrolagus
                      colliei
 Littoral    14         High cockscomb,
                      Anoplarchus purpurencens
                      Northern clingfish,
                      Gobiesox maeandricus
                      Rosylip sculpin,
                      Ascelichthys rhodorus
                      Ringtail snailfish,
                      Liparis rutteri
                                  F

                                  F?
                                                 82

-------
Table B-l  (continued).
          Species
   Zone  Assemblage
Component
 Species
                    Functional Feeding Grou£

  Pelagic    Epibenthic   Epibenthic  Meiobenthic
Planktivore  Planktivore  Benthivore  Benthivore  Omnivore
             15        Black prickleback,
                       Xiphister atropurpwceus
                       Tidepool sculpin,
                       Oligocottus maculosus
                       Mosshead sculpin,
                       Clinocottus globiceps
                       Saddleback  sculpin,
                       Oliqocottus rimensis
                        Fluffy sculpin,
                        Oligocottus snyderi
                        Rock prickleback,
                        Xiphister mucosus
                        Calico sculpin,
                        Clinocottus embryum
                        Tidepool snailfish
                        Liparis florae
                                  F

                                  F



                                  0

                                  F



                                  F

                                  F
                               F?
   Rocky/     16        Yellowtail rockfish,
   Kelp                 Sebastes flavidus
   Bed                  Black rockfish,
                        S. melanops
                        Copper rockfish,
                        S. caurinus
                        Puget Sound rockfish,
                        5. emphaeus
                        Kelp greenling,
                        Hexagramnos decagrammes
                        Lingcod, Ophiodan
                        elongatus
                        Longfin sculpin,
                        Jordania zonope
                        Striped seaperch,
                        Embiotoca lateralis
                                               F

                                               0
   Total (w/o duplication)
                                           8 F/3 0
                                                        26 F/10 0    16 F/2 0
                                                                                  1 F
                                                                                             1 F
                   *Predominantly juveniles.
                                                   83

-------
Table B-2.   Prey composition  of  spiny dogfish reported by  Jones and
               Geen (1977)  (n =  14,796).
                                                                     (SPINY OOGF19H
                       *  Of  Pt'L&TIvF IMPORTANCE (I.w.I.)  TABLE
                        w ruF  IHFNT.  PFK  13.  STATION  TOTAL
                                        FPFO    NIIM.    GRAV.  PPFY      PERCENT
                                        OCCUR   CO^P.   COMP.  I.R.I.   TOTAL  IRI
     rLi'°F"-  HAPfMf,i|-  DAILASI          16.33   P1.B4
     rip,iAII=;TAC>-_n                      15.68   1 ' . 6 3
     IINl'IFNTIF Il'fi  rr,<,                 1 0. «fl    9.6=1
     °LrOCYFHA Tt-fAi) IOF A               9.«9    R.03

     THALEICHTHYS PACIFICUS                 '.?3    5.53
     "F:;uiM i'ir- "-(•:'i|iCT"<;              ?.^>P    6.79
     PLEUROBRANCHIA  SP.                    ?.3^    ?.40
     I 0| I '--M  c;,.                         ?. 14    1.71
     OCTOPUS  DOFLEINI                     ?. 1 <•    3.05
     HYDROLAGUS COLLIEI                    1.65    ?.^B
     P|  FM^i'Mri. T I l«i-                     1 .64    5.89
     DO| vrnAFT a             "'           1.63     .?9
     /IMMnnYTFs HFxOPTFivim              1.60    1.6ft
     FMPIOTOCIQftF                       1.5R     .31

     AMrMJoo^A                          1.47     .60
     SCAPHOPODA                       '     -q7    l-l7
                                           .69    1.35
             TAxA  'vIT^ fu>FO. OCCui-1. LF<=S  THAN 5 AND  NUMERICAL  ANO  GRAVIMETRIC
                  IN  POT*I I F^S  THAN i A«F  FxCLuneo FPOM  THF TAHLF.  ANO PLOT
        ( *HT 'JOT  FI-OM  CAI UiLATIOr,' OF DIVERSITY INOICFS)

           PFr-CFNT  L>OM UlAMCF.  INOFX              .11     .00                .00
           SHiMMrvg-w'" INFD OIVFP^ITY            3.46     .00                .00
           FvrvF<;c;  INHFX                        .PI     .on                .00
                                               84

-------
Ratfish

      A nocturnal predator  In  the nearshore demersal fish assemblage in
exposed gravel-cobble habitats in the San  Juan  Islands, the ratfish illustrated
one of the most diverse  diets  examined.  Eight  ratfish fed on an  array of
brachyuran crabs  (Cancer magister,  C^. oregonensis,  Pugettia gracilis, and
Telmessus cheiragonus),  valviferan  (Synidotea sp.)  and flabelliferan isopod
gammarid amphipods  (Paraphoxus spinosa,  Esiroides  sp., Pontogeneia sp.,  Photis
californica,  Photis sp., Lyssiansidae sp., Aorides sp.),  gastropods, other
diverse peracaridan crustaceans (Hippolyte clarki, Heptacarpus stimpsoni,
Parapaguridae,  and  Paguristes  sp.), hyperiid amphipods,  oniscoiden isopods,
polychaetes,  bivalves,  fish, tanaidaceans, and amphineurans.   Composition  of
the prey by weight  was  dominated by unidentifiable fishes (54%) and brachyuran
crabs  (13%).

Pacific Herring

       Juvenile Pacific  herring constitute the predominant species in North
Puget  Sound's neritic fish assemblage (Fresh 1979).  The specimens examined
indicated  a  low mean fullness factor, high stages of digestion, and the fifth
highest  percentage of empty stomachs: this was perhaps due to high digestive
and  gastric  evacuation rates and because these fish, which may be predominantly
 diurnal-feeding  fish, were  generally collected  at  night.

       Calanoid copepods dominated the overall diet composition by frequency of
 occurrence,  percentage abundance, and percentage biomass in fish collected in
 northern Puget Sound (Fig.  B-l).  Harpacticoid and cyclopoid copepods,
 hyperiid amphipods, barnacle nauplius and cypris stages,  and crab zoea were
 less important while camaceans, gammarid amphipods, and  mysids contributed
 significant percentages of the total biomass of prey organisms.  In addition
 to the prey indicated in Fig.  B-l,  crab megalops,  euphausiids,  crustacean eggs,
 fish larvae, and diverse crustacean larvae contributed to the total diet.

       Juvenile Pacific herring were also captured during beach seine collections
 along the eastern  shoreline, primarily along the northeast and south sides of
 Guemes Island, and at the Cherry Point and Birch Bay sites.   These fish fed
 principally on shallow sublittoral epibenthic organisms  such as harpacticoid
 copepods which composed 82 percent of the total IRI; shrimp larvae and  other
 pelagic organisms, accounted for only 13.5 percent (Fig.  B-2).   Other  epibenthic
 organisms—gammarid amphipods, oniscoidean and valviferan isopods, shrimp  and
 bivalves—made only incidental contributions to the total IRI.

       The diet composition from several sites and sampling periods indicated
 major differences  in food habits between habitats and  seasons.   A series of
 samples obtained from three of the northeastern Puget  Sound sites—Birch Bay.
 Cherry Point, and  Padilla Bay—indicated that different  planktonic organisms
 were  the principal dietary components at the different sites,  i.e.,  calanoid
 copepods in  Padilla Bay, mysids in Birch Bay, and barnacle (cypris)  nauplii
 at Cherry Point.                                                       F
                                       85

-------
                    OD
                    -z.
                    O
                           INDEX  OF RELRTIVE IMFORTPNCE (I.R.I.) OIRGRRM
                           FROM FILE IDENT. N PCSD. STRTION RLSTR
                                     PREORTOR 8747010201
                                  (PRCIFIC HERRING    1
                                 CLUPER HRRENCUS PRLLRSI
                                RDJUSTED SRMPLE SIZE   115
                        lOOr
                         80
                         60
                         40
                         20
                    2    20
                         40
                         60
                         80
                        100
                                                                           ••-  (J to J
                                   20
                                           40
                                                    60
                                                            80
                                                                    100
                                                                             120
                                                                                     140
                                          CUMULRTIVE FREQUENCY OF OCCURRENCE
FREO
P»EY 1TEU nccnp
CALAHOIDA ?q.of)
HARPACTKOIOA 17.00
AMP*lIpr)r)a_MYPFPI IL)f A 	 - - If,. 00
ClIMflCEA 14.00
CTcoIDFTI/r 	 13.00
n^TOACnnt 10.00
OFT ADOpA-«uArMYiiPA 10.00
f-ttMM4BIDEA 9.00
«YTin«crA •"• 	 9.00
FIIDHAIJM ACEA ?.00
" n^TFIfwTHYES l.nn"
MUM.
COMP.
f>7.63
.78
.54
2.9S
3.2?
1.73
.53
1.74
9.P2
.04
. 0*i
COMP.
40.50
.04
.4R
7.02
.45
1 .80
.85
5.91
22.24
1.45
10.80
PPEY
I.O.I.
3135. B
13.9
17.0
140.0
11.0
35.3
13. B
86.1
288.5
3.0
"" 10.9
PERCENT
TOTAL IRI
83.50
.30
.50
3.70
.30
.90
.30
2.30
7.70
.10
.30
             Dory TAXA WITH FDEO. DCCMW. LE1:^ THAN S AND NU^E^ICAL  ANO  GDAV
             COMPOS I Tln-g NOTH LEcs TH«N 1  A3F EXCLUDED FOOM  THE  TAflUE AND  °LOT
           	(BllT-MOT FRfli' CALCULATION OF  nlUEPSITY INOICES)   	  -- --
niMlrjJN'CE INCirx  —
iyF INEP DTVEP5ITY
                                                    .47-   .23"
                                                   1.45   2.20
                                                    .42    .63
                                                  1 .01
                                                    .29
Fig.  B-l.   IRI prey spectrum  of  juvenile Pacific herring in neritic waters
              of  northern Puget  Sound.
                                                86

-------
                         INDEX OF RELRTIVE IMPORTflNCE (I.R.I.) OlRGRftM
                         FROM FILE IDENT. MM  BS, STRTION  HLL
                                 PREOflTOR   8747010201 - CLUPEH HRRENGUS PflLLfiSI
                               (POCIFIC HERRINO    )  HOJUSTEO SRMPLE SIZE = 81
                      100


                       80


                       60


                       40


                       20
                  §    20
                       40
                       60
                  w    80
                      100
                                                                           -Q.
                                 20
                            40       60       80      100

                          CUnULflTIVE FREQUENCY OF OCCURRENCE
                                                                         120
                                                                                  140
                  PREY ITFM
                          FPEO   NU".    GRAV.  PREY     PERCENT
                         _OCCUR  CO«P.   COMP.
	 — - - 	
CALANOIDA
GAPMARIDEA
CRUSTACEA
HARPACTIC01DA
CRUSTACEA "
ROCK
PEMAEIOAE
PERACARIOA-ISOPOOA-ONISCOIDEA

5i.no
27.00
27.00
1ft. 00
7.00
4.00
1.00
1.00

^4.22
3.49
30. *2
_ 3.09
.35
5.19
.45
.42


3.75
76.46
.00
1.95
3.91
2.74
1 •K.I.., 	 Tl

195.5
723.1
1304.3
2.5
28. ft
4.4
3.2
OI»L IWI

3.70
25.00 _
.00
.50
~ .00
        	PREY TAXA WITH FREQ. OCCUR. LESS THAN 5 »NO NUMERICAL AND GRAVIMETRIC
           COMPOSITION BOTH LESS THAN  1 ARE EXCLUDED FOO«« TM£~ TABLE win PLOT"
           (BUT NOT FRO" CALCULATION OF DIVERSITY INDICES)
              PERCENT DOMINANCE INDEX
              SHANNON-VEINEP DIVERSITY
              EVENNESS  INOE*
                                   .39     .62
                                  1.64    1.20
                                   .55     .43
 .41
1.57
Fig.  B-2.
IRI prey spectrum of juvenile Pacific herring in shallow
sublittoral waters along the eastern shoreline of northern
Puget Sound.
                                              87

-------
      Pacific herring were also the most commonly encountered neritic fish
throughout the Strait of Juan de Fuca occurring at all townet sites as post-
larvae and juveniles through midwinter.  Although fish larvae (primarily
Pacific herring and Pacific sand lance) accounted for over 80% of the prey
biomass, calanoid copepods were the most common prey both in occurrence and
numerical composition and thus formed over 96% of the total IRI (Fig. B-3).

      Extensive data on prey composition of larval and post-larval Pacific
herring caught in the Strait of Georgia's surface waters (Barraclough 1967a,
b & c; Barraclough & Fulton 1967, 1968; Robinson et al. 1968 a & b;
Barraclough et al. 1968; Robinson 1969)  indicated that Calanus plumchrus
was the predominant calanoid copepod consumed; Pseudocalanus minutus,
Acartia sp., the euphausiid, Thysanoessa longipes, and the hyperiid amphipod,
Parathemisto pacifica were less important zooplankters and larval fish
(Thaleichthys pacificus, Leuroglossus stilbius, Merluccius productus) appeared
often in the diet but seldom in abundance.  Pacific herring from the nearshore
environs of the east and south coasts of Kodiak Island also have been shown to
almost exclusively (>99% by numbers and biomass) feed upon calanoid copepods
 (Harris and Hartt 1977).

 Northern Anchovy

       Northern anchovies were commonly captured in spring and early summer in
the northeastern study areas (Padilla Bay, Birch Bay, and Cherry Point) during
the DOE baseline studies).  As with herring,  the state of the anchovy stomach
samples indicated a high rate of digestion with a significant proportion of
unidentifiable material in the total stomach  contents sample  (65.4 percent)

      The identifiable organisms (Fig. B-4) indicated rather unselective
planktonic food habits.  Fish (juvenile rockfish) and fish larvae (Clupeidae),
harpacticoid and calanoid copepods, crab and  barnacle larvae, and insects, in
descending order, composed the prey organisms with the highest IRI values.

Pink  Salmon  (Juveniles)

      Pink salmon spawn cyclically in odd years in this region.  Thus, the
juveniles are present in abundance only during even-numbered years.  Juvenile
pink  salmon were present in the largest numbers in the neritic waters of
northern Puget Sound from June-August and were most evident in the sand/eel--
grass habitats.  In this region they preyed on a diverse assortment of epi-
benthic and neritic plankton, with calanoid copepods providing the highest
percentage of the total IRI, and harpacticoid copepods, gammarid amphipods,
barnacle larvae, and cumaceans contributing lower, but fairly equal
proportions  (Fig. B-5).

      Larvaceans and calanoid copepods were the only important preh organisms
composing the overall prey spectrum of juvenile pink salmon caught in the
Strait of Juan de Fuca  (Fig. B-6).

      Amphipods  (Orchomenella sp., Calliopius sp., Parathemisto pacifica),
calanoid copepods (Calanus pacificus, Eucalanus bungi bungi, Acartia sp.,
Pseudocalanus minutus), larvaceans and a diverse array of insects  (principally
families of Diptera) composed the majority of the prey consumed by juvenile
pink  salmon collected in the Strait of Georgia  (Barraclough 1967a, b &  c;
                                      88

-------
                        INDEX OF RELRTIVE ItlPORTfiNCE 11.R.I.) OIHGRRM
                        FROM FILE IOENT.  76-78. STRTION RLSTfl
                                      8747010201   CLUPEfl HflRENGUS PflLLRSI
                                                     PRCIFIC HERRING
                                          RDJUSTED SflMPLE SIZE   477

UJ
2
cc
D
1
ID
CC
>-
OO
Z
o
k-
UD
CD
0-
CD
(_)
1 —
-
CD
•z.
O
to
o
if
o
o

t—
LJ
Q_





P»FY
CALA-OIOA
"Y^IDACEA
HA3PACTICOIOA
TF| FP«;TE1
CLI °F IO»E
AMMOOYT?DAE
100
80



60



40



20

0
20


4


6




8



10






























1 	 1
1






„
•H 11
O rtl
0) fl) O iH QJ -C
-Q 4) ^1 ft) (3 -H
•H u *j *j -a w
O « U U3 -H ^
C T3 •> 
-------
               CAI
                               INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) DIRGRflM
                               FROM FILE IDENT. N PGSD. STflTION flLSTfl

                                           PREDflTOR  8747020101 - ENGRHULIS MORDHX
                                      (NORTHERN flNCHOVY    )   flDJUSTEO SflMPLE SIZE
                                                     31
                            IDOr
                             80
                             60
                             40
                            20
                             20
40
                             60
                             80
                            100
                        POEY  ITE"
                       en
                       •z
                       o
£

u
nl
01
O
J%

CJ
-H
V)

2
u
tn
O
o.
ra
X
E u

U f—
"a
3
W
(T
3

U
re
e

U
*

c
a>
c

2
U)
O
re
U

«
                                      20
                                               40
                                                       60
                                                                80
                                                                        100
                                                                                 120
                                             CUMULFITIVE FREQUENCY OF OCCURRENCE
                                               FOFC
                                               OCfiiP
                         M J M .
                         COMP.
GPAV.
coup.
POC Y
J.O.I.
 PEPCENT
TOTAL IRI
               EiiP"Aii<; I tCK A
1 ft . 00
1 3. no
n.no
13.00
13.00
1 3.00
l?.no
ft. 00
ft. 00
ft.no
ft.no "
ft . no
ft. 00
3.00
'.oo
3.31
17.00
13.0?
?.43
1.10
15. «9
1.10
3.09
4.4?
1 .3?
.88
.44
33.11
.??
.S3
.00
5.ft4
1 .03
?.4ft
??.ft7
.8?
1 .34
• ?1
1.13
.in
1 .03
1 1 .79
?4.6?
?0.51
6.15
53.0
?94.">
1 8?. ft
17.9
309.0
'17.?
?9.-*
19.8
33. 1
8.5
11. S
73.4
63.7
6?.?
?!.]
4.30
?3.80
14.90
1 .40
'5.oo
17.50
?.40
1 .60
?.70
.70
.90
5.90
5.10
5.00
1.70
               rnc 4oioc

                 PPFY  TA»A  v, I rn  F^FO.  OfCli'3.  LF<;=;  THtig  5  ANP NU"E3IC»l  ANO
                 ( OMPO^ITlnM  ROT"  LE^c  TH4M  ]  iSF  ExCLIIDFD  FPOM  THF  TABLF ANO  PLOT
                 (M'lT  'IfT rcn« CaiCUL'TION OF  DIVERSITY
                                                        .19
                                                               .IB
                                                              2.PI
                                                               .74'
                                                  3.37
                                                   .Hft
                                                                                         140
Fig. B-4.   IRI prey  spectrum of northern  anchovy  in  neritic  waters  of
              northern  Puget Sound.
                                                90

-------

                         INDEX OF RELflTIVE IMPORTflNCE (I.R.I.) OlflGRfln
                         FROM FILE IDENT. N PGSD. STflTION flLSTfl

                                  PREDRTOR 8755010201 - ONCORHYNCHUS OORBUSCHfi
                                (PINK SRLMON        )   flDJUSTED SHMPLE SIZE = 38
                      100
                       80
                       60
                       40
                       20
                  £    20
                        40
                       10
                                                                          .
                                                                         4  3,5
                                   50
                                            100
                                                      150
                                                                200
                                                                          250
                                                                                    300
                                        CUnULflTIVE FREQUENCY OF OCCURRENCE
PBEY
ITFM
mto
OCCUR
' MUM/
CO«P.
GRAV.
COMP.
POEY
I.Q.I.
PERCENT
TOTAL IR1
          CAL A
                D A-RR ACHY U1* A
          COPFPIHA     ' ~ ^
        — OJPTFPA •--
          f 6POFLL IDEA
*?. no
4?. 00
<.n. oo ~
37.00
3;>.oo~
?4.00
n.oo
i«(.no
«.on~
«.oo
s.oo —
s.oo
IP. 65
17.7?
10. M
23.33
10.71
.61
7.9?
3.«3
?.?!
.5fl
— .05
.OR
3. IP 	 3.?1
?9.96
^.??
?1.0?
«.07
1.1?
1.68
10.77
5.31
1 1.05
3.59
.00
.40
3.79
?OM.6
1005.S
l?f>5.?
1013."
379.?
55. n
336.4
166.3
106.?
33.4
.3
?.4
15.0
31. «0
15.70
19.70
  .80
  .90
  .90
  .?0
  .60
  .70
  .50
  .10
  .10
  • ?n
               15.
               5.
                 .
               5.
               2.
               1.
            copy T«T4 glTM FOEO.  OCClR. LS';^ THAN 5 AND NUMERICAL ANH ROAVIMETRIC
            rOMPnsITIHM POTH LESS THAN  ]  »SE TXCLDDEO FOOM THE TABLE ANO PLOT
           - I HOT-MO T->QriM C«i.CULATInM OF  DIVERSITY INDICES)
                       OOMIN4NCF-
               SH4NNON-WFI^EB DIVERSITY
.17
.87
.80
  .20
  .6«,
  .7?
Fig.  B-5.   IRI prey  spectrum of  juvenile  pink salmon in neritic waters of
              northern  Puget Sound.
                                               91

-------
                   a
                   g
                   o
                        lOOr
              80
                         60
                         40
                           INDEX OF RELFITIVE IMPORTflNCE (I.R.I.) OlflORflM
                           FROM FILE IDENT.  76-78. STflTION flLSTfi

                                         8755010201 - ONCORHYNCHUS GORBUSCHfl
                                                         PINK SRLMON
                                             flDJUSTED SflMPLE  SIZE =   39
                   en
                   o
                   fc
                   o
                         20
                         20
                         60
                         80
                        100
                                                                 *J  U U .
                                                        u  w  t-t
                                                                     O H
                                                                     U O
                                                                     03 -C
                                  50
                                           100
                                                   150
                                                           200
                                                                    250
                                                                            300
                                                                                     350
                  OREY  ITEM
                                         CUMULRTIVE FREQUENCY OF OCCURRENCE
                                        ~FPEO   NIJMT   GRAV^  PREY     ofRCENT
                                        OCCIJP  COMP.  COMP.  I.P.I.  TOT4L IRI
C4LAMOID*
LARV«CEA-COPEL»TA
HYPEPIIDAF
GAWMARIDAE
CUWACEA
FUCARlOA-OECAPnDA
IN5ECTA
HARPACTICnlOA
WY=IOACF.A
CP|."5TACFA
CTFNOPHORA
OSTF1CHTHYS
HECApnn*
CICPIPFOIA-THORACICA
NATAMTIA
O'lTRACOOA
UOCCHOROATA
6<3.23
5T.H5
PO.^1
?O.S1
17.95
17. 95
17.95
17.95
1?.P?
in. ?f>
10. ?6
10.2ft
7.^.9
7.69
7.»!9
5.13
5.13
?3.B6
67. 6S
1.10
1.01
.63
.39
.19
1.70
.35
.3"
.?*
.11
.?5
.03
.03
.0?
1.51
53.78
33.55
1.42
1.94
.33
.9]
.50
1.13
.80
.56
?.94
.27
.16
.01
.02
.11
.59
=.-<75. n
S451.0
53.4
60.5
17.2
?3.T
12.5
50.7
14.8
9.7
3?.B
3."
3.1
.3
.4
.7
10. fl
4P.32
49.00
.48
.54
.15
.21
'11
.46
.13
.09
.29
.03
.03
.00
.00
.01
.10
PREY TA»A WITH FPFO.  nccuR.  LESS  THAN  5  ANO  NUMERICAL  ANO GRAVIMETRIC
COMPOSITION ROTH LESS THAN ]  ARE _FxCLUOED  FnOMJTHE _J_ABLF_ AW PLOT
              CALCULATION OF  OIvTRSlTY' INDICES)

                                       .52    .40      	       .47
                                      1.46^1.82              1.24
                                       .33    .41               .21
              PFRCENT  n.TMlNANCF  tNOFX
             "SffANNON-WF IwrR" D I VTRSTTY"
              EVENNESS INDEX
Fig.  B-6.   IRI prey spectrum of  juvenile pink  salmon  in neritic waters  of
              the Strait  of  Juan de Fuca.
                                               92

-------
Barraclough & Fulton 1967; Robinson  et  al.  1968 a & b;  Barraclough and Fulton
1968; Barraclough et al. 1968; Robinson 1969).

      Harpacticoid copepods dominated the prey spectra of juvenile pink salmon
captured in nearshore habitats at  Kodiak Island while calanoid copepods,
barnacle larvae, crustacean nauplii,  harpacticoid copepods and fish eggs were
prevalent  in the diet of those fish  captured in pelagic habitats (Harris and
Hartt 1977).

Chum Salmon (Juveniles)

      Juvenile chum salmon occurred throughout  the  north Puget Sound DOE study
 area from May-August,  principally  in the neritic fish assemblages where they
 were most common in collections  from Lummi Bay, Burrows Island, and South
 Beach.

       Chum juveniles had a less  diverse overall prey spectrum than  the pink
 juveniles.  Calanoid copepods completely dominated the  total IRI (80 percent),
 with hyperiid  amphipods being second in importance (12 percent).  Harpacticoid
 copepods, gammarid amphipods, cumaceans, and euphausiids were other less
 important prey items.  This prey spectrum indicates that juvenile chums had a
 more pelagic feeding behavior than juvenile pinks, although epibenthic
 organisms also were important prey in the chum diet, especially when they
 frequented shallow sublittoral habitats (this occurred generally when the
 juvenile are < 55 mm  in length).   Thus, when the combined prey compositions
 of  juvenile chum salmon caught in beach seine collections (Fig. B-7) were
 compared with those collected by townet (Fig. B-8), it was apparent that the
  (earlier, smaller) chums frequenting the shallow sublittoral environment fed
 predominantly upon epibenthic organisms—harpacticoid copepods, gammarid
 amphipods, and oniscoidean isopods—while neritic  (later, larger) chums
 utilized  pelagic organisms—calanoid copepods and hyperiid amphipods.

        Juvenile chum salmon were captured in high numbers during spring beach
  seine  collections along the Strait of Juan de Fuca, especially at Kydaka
 Beach, Beckett Point, and Alexander's Beach, and also were in townet
  collections at Beckett Point.  Main prey of chum fry included epibenthic
 organisms  (harpacticoid copepods and gammarid amphipods) and pelagic organisms
  (calanoid copepods and fish larvae)  (Fig. B-9).

       This transition in feeding behavior from epibenthic to neritic organisms
 appears to be representative  of  most regions of Puget Sound,  including Hood
 Canal  (Feller and Kaczynski 1975;  Simenstad and Kinney 1978),  Nisqually Reach
 (Feller and Kaczynski 1975;  Fresh, et al.,  1978), east central Puget Sound
 (Feller 1977),  and in the Strait of  Georgia (Healy, et al.,  1976).   Detailed
 analysis of prey consumed  in  neritic waters (Barraclough 1967a,b,c;  Barra-
 clough and Fulton 1967,  1968:  Robinson, et al., 1968a,b; Barraclough,  et al.,
 1968; Robinson 1969;  Simenstad and Kinney,  unpubl.  data) suggests that juvenile
 chums predominantly utilize large  calanoids (e.g.,  Calanus pacificus,  (?.
 plumchrus), which are the  principal  components of diel-migrating deep-water
 community, rather than the smaller but  more abundant calanoids (i.e.,  Pseudo-
 calanus minutus),  which characterize the plankton community in the region's
 surface waters.


                                      93

-------
                         INDEX OF RELRTIVE IMPORTANCE (I.R.I.) DlflGRflM
                         FROM FILE IDENT. N PGSD. STHTION flLSTfl

                                    PREDflTOR  8755010202 - ONCORHYNCHUS KETfl
                                (CHUM SflLMON       )   flDJUSTED SRMPLE SIZE
35
PCT. COMPOSITION BY WEIGHT PCT. COMPOSITION BY RBUNOflNCE
o no o> 4* ro ro ** a) o> o
_o oooooooooo




•o «
U rH
ID 0}
Q. ^3
u ra
tO T-1
ffi
V
>. re
j: u
4J OJ
JI CO
U ,£
•H O
01 >•.
tn o
0 cu



a
•o
a)
a
D 4t X
to u s*, t n
V (0 £ rt (0 13
•a -^ *-> a -a o
•H (0 wljriflOOD.
>d (U 3UUCOD.O
to u (ai-tuTaT-ivj
E E D.«->tnc/ia.c/)
(0 3 3WC>,ert
o u wo»-<:£
-------
                        INDEX OF RELATIVE IMPORTHNCE (I.R.I.) DIHGRRM


                        FROM FILE IDENT. N PGSD. STfiTION flLSTfl


                                   PREOflTOR  8755010202   ONCORHYNCHUS KETfl


                               (CHUM SflLMON        )   flDJUSTED SflMPLE SIZE =  32
                 (J

                 i
                 o
                      100
80
                      60
























UJ

O
£ 40
1


S 20
>—
o
a. „
0
1 20
£
>-
CD
2 4°

t—
§ 6
fe
S

^ s
u


10

























CJ
m A
01 U
XCQ
£ 1
4J R)
U O
0) fl)
*J U
 J:
o. o «
X U TH
1 OQ O «
^ -O nl Bt
O D- O CJ ^ V *-"
C -H O. rt ,
P E 41
CJ U P*
'
250

>i--t.r) NUM. OWAV. pot- Y P^WLLNT
uQFt TTFM ncCIJB CO^P. Cn"P. I.O.I. TOTAL IB!
f H 4-ir-
i'-nMjp
rcr «D If
"FI-4C,;
cor /
r - 11.40
PA-Bu*(-«yi|B4- 17. OH l.«7 1.?7 190.? P.OO
Ti""n-i i< F-CO. 1,'fnP. L^1^^ THAN S AMI NUMERICAL AND GDAVIMETP

,/*'.'('_ SC ]•!"• < .f^O ,S9 .11
	 - —

_









Ir


Fig.  B-8.   IRI prey spectrum of  juvenile  chum salmon in  neritic waters  of

             northern Puget  Sound.
                                             95

-------
                     CO
                     CD
                     Q_
                          lOOr
                             INDEX OF RELflTIVE IMPORTHNCE (I.R.I.)  OlflGRflM

                             FROM FILE IDENT.  76-78. STflTION ALSTA

                                             8755010202 - ONCORHYNCHUS  KETfi

                                                            CHUM SRLM.ON

                                               RDJUSTED SfiMPLE SIZE =  63
                          60
                          40
                          20
                     S    20
                           40
                          60
                          80
                          100
                                      50
                                               100
                                                         150
                                                                   200
                                                                             250
                                                                                       300
                                           CUMULflTIVE FREQUENCY OF OCCURRENCE
PPEY ITEM
HAPPACTICOIOA
CALAMOIOA
GAKM4RIDEA
GAWWAPIDAE
Cl|««CEA
OSTRACODA
KATANTIA
CIPTFRA
TFLEOSTFII
FBFO
OCCLI"
b't.ttl
frl .90
«?.^6
3T.11
?0.^3
1?.70
1?.70
11.11
?.7
l.?3
1.71
.22
1.23
1.54
<..!?
GRAV.
COMP.
25.83
15.55
5.87
5.06
3.33
.74
8.91
10.6S
21.76
PP£Y
I.R.I.
M9P.4
19«3.«
421.6
149. A
104. n
12.?
128. »
iss.n
?05.4
PERCENT
TOTAL IRI
66.16
?1.17
4.50
1.60
1.11
.13
1.37
1.69
2.19
           J3RFY TAXA WITH FPEO.  OCCllR_.  LES^  THAN  5  AND  NUMERICAL  AND GRAVIMETRIC

           "COMPOSITION BOTH LE":?" THAN^l  ARF" FXCLUOEO  FROM  THE  TABLE ANO PLOT

            (BUT NOT FRO" r»irUL»HON OF  OlVERSITY INDITES)
               PEPCENT DOMINANCE  INDEX
               SHANMON-W^INFR DIVERSITY
               EVENNESS INDEX
 .4P   ~  ".16

1.67    2.92

 .38     .66
1.57

 .36
Fig. B-9.   IRI prey  spectrum of  juvenile chum  salmon  from Strait of  Juan de
              Fuca.
                                                96

-------
      Juvenile chum salmon from nearshore habitats of Kodiak Island had fed
principally upon harpacticoid copepods although fish larvae, gammarid
amphipods, and mysids comprised much of the prey spectrum in terms of biomass
(Harris and Hartt 1977).

Co ho Salmon (Juveniles)^

      During the DOE Baseline Studies juvenile coho salmon were  found throughout
the neritic waters of north Puget Sound from April to October.   Townet
collections at Birch Bay, Eagle Cove, Shannon Point, Cherry Point, and  Padilla
Bay typically produced the highest catches.  The overall prey  IRI  spectrum
(Fig. B-10) showed that juvenile coho in that region fed upon  both epibenthic
and pelagic organisms, but apparently equally so on those which  were available
within a  certain size range.  The pelagic organisms included drift insects
(the most commonly taken item), crab zoea and megalops, hyperiid amphipods,
and fish.  Epibenthic prey included only crustaceans—gammarid amphipods,
shrimp  (Crangonidae), flabelliferan isopods, and ostracods.

       In  terms  of  the total  IRI, the highest contributors were fish  larvae
 (23 percent),  insects (22 percent) peracaridian crustaceans  (15  percent),
oniscoidean  isopods  (14 percent), gammarid amphipods (9 percent),  and crab
larvae  (7 percent).   Identifiable amphipods included Eusiroides  sp.  (the most
common),  Atylus sp.,  Allorchestes sp., Eohaustorius sp., Calliopius  laeviusculus,
Talitroidea  sp.,  Paraphoxus  (Trichophoxus) spp., Pontogeneia spp., and
 unidentified Hyperiidae  species.  The mysid was Hornsiella anomala, the  isopods
were  predominantly Gnorimosphaeroma oregonense, and the calanoid copepod was
 Epilabidocera amphitrites. The  identifiable fish were all larval or  juvenile
 herring.

       Although never in  high numbers,  coho salmon  juveniles were frequently
 encountered in spring and summer beach seine collections at  almost all  sites
 along the Strait of Juan de  Fuca.  Gammarid amphipods  (over  90%  of the  total
 IRI)  and  fish larvae (including Pacific herring and Pacific  sand lance) were
 the most  important prey;  cumaceans, polychaetes, sphaeromatic  isopods
 (Gnorimosphaeroma oregonensis), insects, and mysids were of  secondary
 importance (Fig.  B-ll).   The relative importance of epibenthic crustaceans such
 as gammarid  amphipods, cumaceans, harpacticoid copepods and sphaeromatid isopods,
 of neritic zooplankton  such  as  crab larvae, hyperiid amphipods and euphausiids,
 and of fish larvae is sustained in other areas of  Puget Sound.  Brachyuran
 crab larvae and euphausiids  predominated in the prey spectrum  for  juvenile
 coho in Hood Canal (Simenstad & Kinney 1978); those collected  over the  1978
 outmigration period (late March - mid-June) in Nisqually Reach showed  a
 transition in predominant prey  from  gammarid amphipods, mysids and harpacticoid
 copepods  early in that  period toward  euphausiids and crustacean  larvae  in May
 and June  (Fresh et al.  1979).   Although not exceedingly common in  the Pacific
 Biological Station's 1966-68 neritic  collections in  the Strait of  Georgia,
 juvenile  coho preyed exclusively upon large zooplankters  (3.5-10 mm),
 principally the calanoid copepod Calanus plumchrus,  the hyperiid amphipod,
 Parathemisto pacifica,  and euphausiids (Euphausia  pacific.  Thysanoessa
 raschii and T\ spinifera).  Fish larvae,  including  Pacific  herring, eulachon
 (Thaleichthys pacificus), and Pacific sand lance,  became prevalent prey after
 June (Barraclough 1967  a-c;  Barraclough  &  Fulton 1967  and  1968;  Robinson et
 al.  1968  a & b; Barraclough et  al.  1968; Robinson  1969).
                                       97

-------
                             INDEX OF RELRTIVE IflPORTRNCE (I.R.I.) DlflGRRM
                             FROM FILE IDENT. N PGSD. STflTION RLSTR

                                        PREDRTOR  8755010203 - ONCORHYNCHUS KISUTCH
                                    (COHO SHLMON        )   RDJUSTED SRMPLE SIZE = 140
                     o
                     •z.
                     cc
                     a.
                     §
                     o
                          100
80
                           60
                           40
ZO
                     §    20
                           40
                           60
                     t-    80
                     
-------
                               INDEX OF RELflTIVE IMPORTflNCE (I.R.I.I DlflGRRM
                               FROM FILE IOENT.  76-78. STflTION  ALSTA

                                              8755010203 - ONCORHYNCHUS KISUTCH
                                                              COHO SflLriON
                                                 flOJUSTED SflMPLE SIZE    16
                                        50
                                                  100
                                                            150
                                                                      200
                                                                                250
                                                                                          300
                                              CUMULflTIVE FREQUENCY OF OCCURRENCE

GAUMARIDEA
CUXACEA
POLYCMAETA
"YMDACEA
GAMMARIOAE
?i:£"
TFLEOSTEI
UNIDENTIFIED
CHLOROPHYTA
CAPRFLLIOAE
HACPACTICOIOA
AMMOOYT1UAE
COI.ErPTERA
FREO Nil'
> OCCUR CO
62.50 4?
25.00 5
i::75 I
1».75
1?.50 4
12.50 8
1?.SO
1?.50
5:11
6.25
".
MP.
.83
.45
.83
.61
.65
.89

.40
!?0
.20
GRAV.
COMP.
?6.02
.87
3.68
.42
1.23
.45
.44
48.81
.17
.00
.02
.00
1.52
.00
ORF.Y
I.P.I.
4303.1
158.1
87. S
798.5
19.1
73.4
116.7
10.6
615.?
?4.B
2.5
1.4
1.3
10.8
1.3
PERCENT
TOTAL IPI
67.80
1.92
1.38
12.58
.30
1.16
1.84
.17
9.69
.39
.04
.02
.02
.17
.0?
               PRFY TAXA tilTH FREO. OCCUR. LES<; TMAN~trTt*fTT4llMEeTCAL~Aii40" GP~*VP4ET«»TC	
               COMPOSITION BOTH LE<;? THAN i ARE EXCLUDED FROM THE TABLE  ANO PLOT
               (RUT NOT FROM CAI.CUL»TIOM OF DtVEPSITY INDICES)
                  PERCENT OflMlN«NCF INDEX
                  SHANNON-MFINEO DIVERSITY
                           1NOE>
                                                      .59
 .33
^.06^
 ".51
  .49
 1.69
~
Fig.  B-ll.   IRI prey  spectrum of juvenile  coho salmon in the Strait of Juan
               de Fuca.
                                                 99

-------
Sockeye Salmon (Juveniles)

     Except for a sizable catch at Birch Bay in May 1976, juvenile sockeye
were infrequently encountered during the DOE Baseline Studies in the neritic
waters of north Puget Sound, usually only in the northeastern study sites
during May through September.

     Juvenile sockeye salmon were dependent upon euphausiids, shrimp and fish
larvae, and general  (unidentifiable) eucaridan crustaceans (typically epi-
benthic organisms),  and to a lesser degree upon pelagic and surface prey
items such as calanoid copepods, barnacle nauplii, and hyperiid amphipods
(Fig. B-12).

     No juvenile sockeye were collected during the MESA collections in the
Strait of Juan de Fuca (Cross et al. 1978).

     Juvenile sockeye salmon captured in the neritic waters of the Strait
of Georgia (Barraclough 1967 a-c; Barraclough & Fulton 1967 and 1968;
Robinson et al. 1967 a & b; Barraclough & Fulton 1967 and 1968; Robinson
et al. 1967 a & b,;  Barraclough et al. 1968; Robinson 1969) had
characteristically consumed either drift insects (especially after June),
including many dipterans and hymenopterans, or pelagic zooplankters,
including calanoid copepods (Calanus plumchrus, Paracalanus parvus,
Pseudocalanus minutus, Fucalanus bungi bungi), euphausiids (Thysanoessa
raschii), hyperiid amphipds (Parathemisto pacifica), chaetognaths  (Sagitta
elegans) and larvaens.

Chinook Salmon (Juveniles)

     Juvenile chinook salmon were ranked among the  10 most common  neritic
fishes in north Puget Sound and were the most common juvenile  salmonid in the
region's neritic waters (Fresh 1979).  All the stomach samples originated
from the eastern study sites from May through September with the largest
samples from Padilla Bay, Birch Bay, and Burrows Island  in July and August.

     The generalized prey spectrum from the DOE collections  indicated both
epibenthic and pelagic feeding beahvior with an emphasis  on  the latter
 (Fig. B-13).  Overall, the most important prey taxa were  crab  megalops,
insects, juvenile and larval fish  (Pacific herring  surf  smelt, Hypomesus
pretiosus) and gammarid amphipods.

     Juvenile chinook salmon were caught in beach  seine  collections  in
southern North Sound (Guemes Island sites, Padilla  and Fidalgo bays)  during
July and August.  Their diet was almost entirely  taken up by crab  larvae
 (megalops) which constituted 89 percent of  the total  IRI for that  species.
Insects, gammarid amphipods, and polychaetes formed the  majority of  the
remaining  incidental prey organisms.

     In the Strait of Juan de Fuca collections chinook  salmon  juveniles
and  a  few maturing residents were  common to both  beach  seine and  townet
collections from May and August, especially at Morse Creek and Beckett Point.
Dipteran  insects,  shrimp  larvae, and gammarid amphipods  predominated in the

                                     100

-------
                          INDEX OF RELRTIVE IMPORTANCE (I.R.I.) DIRGRfitt
                          FROM FILE IDENT. N PGSO. STRTION flLSTfl

                                     PREDRTOR 8755010205 - ONCORHYNCHUS NERKR
                                 (SOCKEYE SflLMON      )   RDJUSTED SfltlPLE SIZE -  39
                       100
                        80
                        60
                        40
                        20
                   §    20
                        40
                        60
                        80
                        100
                                                                       tt -a
                                                                      •o o> ra
                                                                       o a u
                                      50
                                                 100
                                                             150
                                                                         200
                                                                                     250
                                         CUMULflTIVE FREQUENCY OF OCCURRENCE
                   PPFy
       NUM.
OCCUR  COMP.  COMP.
                                                               PREY
                                                               I.P.I.
              PERCENT
              TOTAL  IPI
           FUPHAUMACtA
                       PFi5I IDFA
           EllCAOlDA
           PLEnCYEMATA-CADIOFA
?p.no 	 1.59
2S.OO 51.71
26.00 11.75"
23.00
21 .00"
15.00
13.00~
11.00
1 1 .00
10.00
~ P . n o
8.00
3.00
1.31
~.75~
1.73
.9ft
1 .60
.16
3.9ft
2.77
.16
1 .14
~3.2T
17.37
3.^1
4.40
3.08
" 1.32
25.63
7.89
5.29
.08'
.15
.57
                      139.2
                     71B8.P.
                      120.1
                      108.?
                       72.?
                       29.9
                      354.0
                       PP.ft
                       92.5

                        2.5
                        5.1
                 3.10
                48.60
                28.40
                 2.70
                 2.40
                 1.60
                  .70
                 7.90
                 2.00
                 2.10
               ""  .50
                  .10
                  .10
             CPFY TA^A UITH  FPEO.  OCCUR. LFSS THAN 5 AND NUMERICAL AND GRAVIMETRIC
         	ro«POMTlON BOTH  LE<;S THAN  ] ARE EXCLUDED FOOM THF TABLE AND PLOT
             fOTT NOT FROM CALCULATION OF DIVERSITY INDICES)
                        OOMlNAfJCF  INOEX
                SH4MI.JON-WFINFP DIVERSITY
                EVENNESS INOF»
          .37
         1.94
          .52
2.81
 .7ft
 .33
2.17
 .59
Fig.  B-12.   IRI  prey spectrum of juvenile  sockeye  salmon  in  the  neritic
               waters  of northern  Puget Sound.
                                              101

-------
                         INDEX OF RELRTIVE IHPORTRNCE (I.R.I.) DIRORfltt
                         FROM FILE IDENT. N PCSO. STRTION RLSTfl
                                  PREDRTOR  8755010206
                                (CHINOOK SRLMON      )
                               ONCORHYNCHUS TSHHWYTSCHR
                               RDJUSTED  SRHPLE SIZE = 131
                  en
                  >-
                  00
                  -z.
                  a
                      100
                       80
60
                       40
                       20
                       20
                  CD
                  -z.
                  o
40
                       60
                      100
                                                                            -
                                                                       o c a. a)  ID -
                                                                       nj u ra J-*  jr
                                                                       E E O CL 0. r
                                  so
                                            too
                                                      150
                                                                200
                                                                          250
                                                                                    300
                                        CUMULflTIVE FREQUENCY OF OCCURRENCE
	 — 	 — • FREf)
PREY ITEM OCCUR

GAWM4RIDEA
OSTnCHTHYES
UROCHOOim*
ruuictT1'
HYWF'JOPTFKA
U 1 U 1 1. u 1
FlICHAIISIACEA
rA^'olDA
EYPOMES^TPO'ETII

JPA '-• 67.00
43.00
41.00
21.00
NUM.
COMP.
~26.09
11.36
15.21
1.65
20.00 " 18.06
1^.00 3.?2
IS. 00
in.no
«.on
JPA 8.00
7. no
7.00
6.00
4.00
osus i.oo
4.49
4.14
3.97
2.88
1.80
.66
.90
.78
".12
.0?
T3RAV.
COMP.

15.43
4.57
	 3.91
48.77
" .09"
.39
"5.35
.67
" .64
1.15
1.54
.27
.22
' "S/30
5.63
PREY PERCENT
I.P.I. TOTAL 1RI

685. n
779. B
1058. P
363.0
65. n
48. 1
" 41.5
32.?
16.6
15.4
8.?
6.0
33.7
5.7

45.70
11.30
12.80
17.40
6.00
1.10
2.40
.80
.70
.50
.30
.30
.10
.10
.60
.10
          DPFY TAXA «1TH FQEO. OCCUR. LESS THAN 5 AND NUMERICAL  AND  GRAVIMETRIC
          -Tn«POSITlnN~fTITH T.ESS THAN~l ARE EXCLHOED FROM  THE  TABLE AND  PLOT
           (SUT NOT FROM CALCULATION OF DIVERSITY  INDICES)
              PERCENT  DOMINANCE  INDEX
                       E~IMFR
                       TNOEx
Fig. B-13.   IRI  prey spectrum of juvenile  chinook salmon in northern
               Puget Sound.
                                              102

-------
most diverse prey spectrum of all  the  juvenile salmonids  (Fig.  B-14).
Osstracods, post-larval fishes  (including Pacific sand  lance  and rockfish,
Scorpaenidae), polychaetes (including  Syllidae),  mysids,  and  calanoid  copepods
were of secondary importance.

     Juvenile chinook from Hood  Canal  and southern Puget Sound had remarkably
similar prey spectra (Simenstad  and  Kinney 1978;  Fresh  et al. 1979) to those
from northern Puget Sound and the  Strait of Juan  de Fuca; insects
(principally dipterans), crab and  shrimp larvae,  shrimp (principally crangonids),
and fish larvae also formed  the  most prevalent prey organisms in these regions.

Surf Smelt

     Surf smelt were similar to  Pacific  herring in their distribution through
northern Puget Sound.   Eastern  townet  sites,  especially Birch Bay and
Padilla Bay, sampled during  the  DOE  studies produced large samples.  The
 total  sample was approximately two-thirds juveniles and  one-third adults.

     The prey  spectrum  from  this region  (Fig.  B-15) include both pelagic and
 epibenthic  organisms as important  prey.   According to the total IRI,  epibenthic
 flabelliferan  isopods were the most  important prey organisms followed by
 cumaceans,  larvaceans,  and calanoid  copepods.   One specimen of Lophopanopeus
 bellus was  also  found in a stomach  but  was not included in the IRI graph
 because  it did not  represent a  significant portion of the stomach contents.

      Surf  smelt  were also  caught in  the  shallow sublittoral zone all along
 the eastern shoreline of northern  Puget  Sound during beach seine collections;
 the highest catches obtained for stomach samples  were from Cherry Point in
 December.   Of  the diverse  prey  organisms consumed, 26.7 percent of the total
 IRI were larvaceans;  27.0  percent, caprellids; 9.9 percent, gammarid amphipods;
 11.8 percent,  calanoid  copepods; 7.7 percent,  penaeid shrimp; 5.0 percent,
 harpacticoid copepods  (Fig.  B-16).  Thus, close to 60 percent of the prey
 from these collections  were  epibenthic organisms.

      All life history  stages of surf smelt were commonly caught throughout
 the Strait of Juan de  Fuca during the  MESA studies, but catches were highest
 at Twin Rivers,  Morse  Creek, Alexander's Beach, and West Beach.

      Although not among the most abundant species caught in the shallow
 sublittoral zone (approx.  10th one year, not caught the next), surf smelt
 usually ranked in the top  five species sampled in the neritic waters by the
 townet (Cross et al.  1978).

      In accordance with their predominantly neritic distribution, calanoid
 copepods provided the most trophic input (80.6% of total IRI) to the overall
 surf smelt prey spectrum (Fig.  B-17).   Harpacticoid copepods (12.7%) and
 polychaete annelids (3.8%)  were second in importance.  Fish, although
 infrequently consumed,  accounted for 27.5% of the total prey biomass.
                                     103

-------
                      100
                         INDEX OF RELflTIVE IHPORTflNCE (I.R.I.) OlflGRfltl
                         FROM FILE  IDENT.   76-78. STflTION ALSTA

                                      8755010206 - ONCORHYNCHUS TSHflWYTSCHfl
                                                      CHINOOK SfiLMON
                                           flOJUSTED SflttPLE SIZE    67
                                                                      n  —t o c.^ u &
                                                                      c  n TJ -H a M «
                                                                      I-H  CJ M B H O U
                      100
                                  50
                                            100
                                                      150
                                                               200
                                                                         250
                                                                                   300
                                        CUMULflTIVE FREQUENCY OF OCCURRENCE
PREY ITFM
OIPTFRA
GAWMARJDEA
fc ATANT I A
POLYCHAETA
T«TRACOOA
wyclDACF A
INCECTA
CALANOIDA
IDCTEIOAE
HIPPOLYTIDAE
TELEOSTEI
OSTEICHTHY5
CAPPELLIOEA
^Yl L IDAE

40.1
166.7
43.1
.0
9.4
20.4
17.1
PERCENT
TOTAL IRI
52.55
15.58
17.22
3.83
1.40
.24
1.95
.09
.67
2.78
.7?
.01
.16
.34
. •2R .
         PPEY TAXA MlTH FPEQ. OCCUR.  LESS THAN 5 AND NUMERICAL  AND GRAVIMETRIC
         COMPOSITION BOTH LESS THAN 1 ARf EXCLUPEO FonM THE  TARLE_ANn PLOT	
         ("BUT" NOT' FRO» CAICULATION OF DIVERSITY INDICES)
            PERCENT DOMINANCE INDEX	
            SHANNON-WF1NFP DIVERSITY
            EVENNESS INDEX
 .30    .10
?.50   3.64
 .48    .69
 .13
2.21
Fig.  B-14.   IRI  prey spectrum  of juvenile chinook salmon in the Strait of
               Juan de Fuca.
                                               104

-------
                           INDEX OF RELflTIVE  IMPORTRNCE (I.R.I.) DIRGRRM

                           FROM FILE IDENT. N PGSO. STRTION RLSTfl


                                      PREDflTOR  8755030101 - HYPOMESUS PRETIOSUS

                                  (SURF SMELT          )    HOJUSTED SRMPLE SIZE =  56
(T


>-
(D
                        100
                         80
                         60
J-    40


s
a.


o    '0


h-
CJ

       0



I—


§    20
LU
2

>-
(D



I    4°
»—



1    60

IT
O
O


,-1    80
o
0.




     100
                                                                                  O.T3
                                                                                  o o
                                                                                  e a.
                                  20
                                        40
                                               60
                                                      80
                                                            100
                                                                   120
                                                                          140
                                                                                 160
                                                                                        180
                                           CUnULHTIVE FREQUENCY OF OCCURRENCE
f RfO MUM.
PWEY ITF.M OCCIIP COMP.

CAI.AN01DA

r IPSIPFQI A
LAUVACt A
TPIJSTACEA
COPFPOOA- -
HAOOACTICOIOA

I OOA-MYPESI If
"STFICHTHYE*;
CTFNOPHOPA
T ^ ^nrtr\


?7;oo" 19.
?5.00 13.
"?5700 17.
?o.on 2.
is.no 30.
13.no ?.
7.00 1.

>FA 7. on i.
5.10 	 .
5.00
5.PO
• " 1 •
7.10
62
50
*V
07

09
19
19
c n
32
GRAV.
COMP.
13.55
3.01
59.0?—
.27
S.fl?"
.10

.41
	 .85 —
.64
.00 ~

1.79
PPF.Y PEPCtNT
I. 5. I. TOTAL IPI
415.?
1913.0
53.8
32. ?
60. »>
8.?
8.n
10.5
6.0
.9
18.?
19.0

21 .80
10.10
46.60 	 ~
1.30
16.00
1.50
.20
.20 '
.30
.?0
• 10
.00
.40
• 50
            PPFY-TATA ^ITH FDFO. OrcilS. LF.5<;-THAN  5  AND NUMEPICAC AND GRAVIMETRIr
            CoUooSITIO,, POTH LE<;? TH4(J  ,  iQF  fxc            THE


           - (BUT- NOT F90M CALCULATION OF  PlVEPSITY IMnICES)
               PF.PCENT
                                                    .19
                                                           .37

                                                          l.BO

                                                           .47
                                                         .30
Fig. B-15.    IRI  prey spectrum  of  surf smelt in northern Puget Sound.
                                                105

-------
                         INDEX OF RELRTIVE IMPORTRNCE  (I.R.I.) OIRORRM
                         FROM FILE I DENT. HM  BS. STRTION  RLL
                                    PREDRTOR  8755030101 - HYPOMESUS PRETIOSUS
                                (SURF SMELT         )   RD JUSTED SRMPLE SIZE  = 42
                       100
                       80
                       "
                       40
                       20
                  S   20
s
fe
S
                        so
                       80
                       100
                                                           JJ^
                               20    40     60     80     100    120    140

                                       CUMULATIVE FREQUENCY OF OCCURRENCE
                                                                          160
                                                              180
PREY ITEM
CAL4NOIOA
HARPACTICOIDA
GAMMAR1DEA
lADVACfA
CAPRELLIDE*
OSTEICHTHYES
O«;TEICHTHYES
CRUSTACEA
ROCK
CU«ACEA
PENAEIDAE
NE"ATOOA
OSTRACOOA PLATYCOPA
CIRRIPEOIA
FRFO
OCCIJR.
29.00
24.00
2A.OO
19.00
1?.00
10.00
10.00
7.00
7.00
S.OO
S.OO
s.oo
s.oo
2.00
NUM.
COMP.
6.52
4.90
3.03
?3.67
?3.92
.75
6.06
6.67
8.34
.09
2.56
.70
.14
10.82
GRAV.
COMP.
3.31
.1?
6.S6
9.9S
30.25
2.56
2.56
.35
1.45
.00
34.32
3.20
.00
4.07 "
PREY PERCENT
I.R.I. TOTAL IPI
2R5.1
120. S
237.4
642.6
650. n
33.1
86.2
49.1
6B.5
.4
184.4
19. S
.7
2~9 . fl
ll.flO
5.00
9.90
?6.70
27.00
1.40
3.60
2.00
2.80
• 00
7.70
1.00
.00
1.20
            PREY TAXA WITH FPEQ. OCCUR.  LESS THAN 5 AND NUMERICAL ANO GRAVIMETRIC
            COMPOSITION ROTH LESS THAN 1  ARE EXCLUDED FoO" THE  TABLE ANO PLOT
            (BUT NOT FROM CALCULATION OF  DIVERSITY INDICES)
              PERCfNT DOMINANCE INDEX
              SHANNON-WEINER DIVERSITY
              EVENNESS INDEX
                              .15
                             3.02
                              .79
 .23
3.56
 .71
2.87
 .80
Fig.  B-16.   IRI prey spectrum of surf  smelt  along eastern shoreline of
              northern Puget Sound.
                                           106

-------
                    d
                    §
                        100
                         80
                         60
                           INDEX OF RELflTIVE  IMPORTflNCE  (I.R.I.) DIflORfltl
                           FROM FILE IOENT.   76-78. STRTION  ALSTA

                                           8755030101 - HYPOMESUS PRETIOSUS
                                                           SURF SMELT
                                              fiDJUSTED SRMPLE  SIZE = 188
                    £    40
                    §
                    fc
                    5    20
                    2    20
                    £
                    £
                         40
                         60
                         80
                        100
                                                                  i-t tC T3 u Ifl
                                                                  J= *J d ^
                                                                   d. o "J 0)
                                                                  < eu u H
                                                                               (0   4J C-
                                10
                                      20
                                            30
                                                  40
                                                         50
                                                               60
                                                                     70
                                                                           80
                                                                                 90
                                                                                       100
                                          CUMULflTIVE FREQUENCY OF OCCURRENCE
PPEY
ITFM
FPEO
OCCUP
NUM.
COMP.
r.RAv.
COMP.
PPEY
I.P.I.
OEPCENT
TOT4L IRI
        HAePACTlCOinA
        POLYCHifTA
                                        43.62   51.34   23.08  3246.1
        we ir«CEA
        NATAKiTIA
        «Pt-OODITIDAF
        POTAMOGFTOSiCElE
        TELEnSTF.I
        COTTIDAF
n.io
10.11
 8.51
                                         S.R5
  .53
  .53
  .53
  .53
  .53
11. 80

  .31
  .42
  .11
~ .08
  .01
  .01
  .01
  .01
  .01
 5.24
 2.47
 4.11
  .19
 4.46
 1.88
 3.23
15.30
 9.1?
 1 .44
 9.7?
17.68
 145.1
  36.4
   4.0
  28.6
	8.5
   8.8
   8.1
   4.9
    . p
   5.2
   9.4
                                                          80.60
                                                          12.65
                                                           3.60
                                                            .95
                                                            .10
                                                            .71
                                                            .21
                                                            .22
                                                            • 20
                                                            .12
                                                            .02
                                                            .13
                                                            .23
          PPFy T«»A WITH FPEC.  OCCllP.  LE^-; TH«M S AND NUMERICAL ANO GRAVIMETRIC
           nnPO<;lTI1\ POTH LESS THAN 1  APE EXCLUDED FPOM THE TABLE AND PLOT
          (PUT MOT FBO» CAI. CULAtlnN OF  DIvEPSITY INDICES)	    	
PEOCENT
SHANNON-
EVENNESS
                           ,CE  IK'DF.X
                            DlvEPSITY
         .39    .11
        1.70   3.2»
         .34    .65
                           .67
                          1.06
                           .21
Fig.  B-17.   IRI prey  spectrum of surf  smelt  in  Strait  of  Juan de Fuca.
                                               107

-------
           n smelt were not a common component of the DOE collections  in
towne      8et Sound'  Three adult longfin smelt caught during a September
cala   .^ollection at Cherry Point had consumed principally crab larvae,
and »a™  c°Pepods, and mysids, with supplemental contributions by hyperiid
        1?r^-amPJip0ds-  This pre^ composition suggested a basically
        reeding beahvior .
     In the Strait of Juan de Fuca, however, longfin smelt of all life-
history stages were caught frequently, and were especially abundant in
August and October townet collections at Twin Rivers and Pillar Point and
in January beach seine collections at West Beach.  Epibenthic crustaceans
predominated the overall prey spectrum of longfin smelt (Fig. B-18) .   Gammarid
amphipods accounted for 61.6% of the  total IRI; mysids (Archaeomysis
grebnitzki and Neomysis sp.), 24.4%;  and cumaceans, 5.1%.  Pelagic prey
organisms were not important.

Northern Clingfish

     Northern clingfish were common members of  the  intertidal  fish
assemblages documented during the MESA studies  in  the  Strait of  Juan de Fuca;
they were found to be especially abundant at North  Beach,  Morse  Creek,
Observatory Point, Twin Rivers, and Slip Point.  Epibenthic  and  benthic
crustaceans and benthic molluscs were the most  important  prey  organisms
(Fig. B-19) .  Gammarid amphipods and  isopods  (Gnorimosphaeroma oregonensis,
Exosphaeroma amplicauda, Dynamenella  sheareri,  Idotea  urotoma, and
Pentitotea montereyensis) made up  68.1% of  the  total  IRI,  and  limpets
(Collisella pelata, C. digitalis,  (3.  strigatella,  Notoacmea  scutum,  W.
persona, and 11. fenestrata) , 24%.

     The prey spectrum of northern  clingfish  in the littoral environs of
northern California (Johnson 1970)was very  similar to  that of the Strait of
Juan de Fuca; gammarid amphipods  (specifically  Amphithoe sp.)- flabelliferan
isopods, and benthic gastropods  (Acumaea  sp., Nassarius sp.) were the
principal prey organisms in the spectrum  (Fig.  B-20) .   Northern clingfish
collected in the rocky/kelp bed habitat on  theouter coast of Vancouver Island
appeared to have a relatively similar prey  spectrum including isopods (26.7%
by weight), unidentified crustaceans  (26.7%)  and chitons  (20.0%)  (Leaman
 1976).

 Pacific Tomcod  (juvenile)

     During the DOE Baseline Studies  in northern Puget Sound, juvenile Pacific
 tomcod were often caught in large  numbers  during the beach seine  collections
 along  the eastern shoreline, especially at  Shannon Point  in July  and  Birch
 Bay in  December.  Penaeid  shrimp  (28.9) percent of total  IRI),  gammarid
 amphipods  (38.0) percent of  total  IRI)  and  calanoid copepods  (19.6  percent
 of total IRI)  composed  the majority of  the  prey organisms from  these samples
 (Fig. B-21) .   This  composition  suggests a predominantly epibenthic  planktivores
 feeding beahvior but  with  some  neritic  feeding, also, perhaps at  night.


                                    108

-------
                        100
                           INDEX OF RELflTIVE  IMPORTfllCE (I.R.I.) OIRGRflM
                           FROM FILE IDENT.   76-78. STflTION HLSTfl
                                         8755030402 - SPIRINCHUS THflLEICHTHYS
                                                         LONOFIN SMELT
                                              flOJUSTED SflflPLE SIZE =  93
                    0
                    1
                    a:
                    v
                    m
                    s
                    fc
                    o
                    o
                    Q_
                    §
                    o
                                          CUnULRTIVE FREQUENCY OF OCCURRENCE
                  P»Fv  ITF"
                                         FPEO
                                         OCCUR
MUM.
COMP.
                                                       COMP.
PPEY
I.R.I.
 PERCEN'T
TOTAL IP!
        _MY«:inArEA
         CU«ACF A
         GAUMARIOAE
         SPt-AEPOMATlnAF
         CAL AMOIDA
         POLYTHAETA
25.81 ?7.17 32.21 1532.3 54
2?.58_
|fl .'8
9.A8
-7-5 3_
4.10
4.10
3.?3
?• 15
?. 15
?. 15 	
1.08
1 .08
1..08
I .08
9.58 .20.96 _
4.20
13.65
.98 	
30.64
.4ft
.52
6.7ft
.59
- .36. .
?.43
1.64
_ .07
.07
3.70
ft. 41
.56 	
9.40
1.97
3.34
4.14
1.48
2.47 	
2.82
.52
ft89.6 	 24
144.4 5
194.1 ft
. 11_. 6
172.? ft
10.5
12.5
?3.4
4.5
. 5.9 	
5. ft
2.3
6.90 	 7.5
2.1ft
2.4
.31
.44
.12
.88
.4]
.10
.37
.44
.83
.16
,21
.20
.08
• 27_
.08
         EliCAQIDA-DECAPOOA
         N4TANTIA
         njirtc^THYS ______
         CIHOIPF.DIA
         EUPHAUMACEA
         LUxRolNER
         K'FOFIOAE

        _  POFY  Ti>4 WITH FPET.  OCCUR.  LE«;t;  THAM  s AND WUMEPICAL AND GRAVIMETRIC
           COMPOSITION  POTH  LE«;C  THAN  1  APE  EXCLUDED FR~0M THE TABLELAND PLOT
           (RUT  NOT FRO"  CALCULATION PF  DIVERSITY INDICES)
              PE"CF'JT  DOMlNAMCF  INDFX
              cH^^"gO"-wFIN';o DIVERSITY
              EVENNESS INOE»
  .?0    .17
 ?.74   3.13
  .63    .7?
             .37
            1.96
             .45
Fig.  B-18.   IRI prey  spectrum of  longfin smelt in the Strait  of Juan de Fuca.

                                                109

-------
                                  INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) DIfiCRfin
                                  FROM FILE IOENT.  76-78. STflHON flLSTfl
                                                8784010101 - 008IESOX  MEHNDRICUS
                                                             N. aiNCFISH
100
80
60
40

20



•20
-40
-60
fln
100

MUJUOILU SHnrLt OltL = CVQ
m
V
00
i
u
a .
.
V Oi -0 C. £
u ^ o. u -o
o i- z: o u at <
	 1 — r-tj-Tkn
| | 1 — 1 U^
u \\
\\
,
•0 « M 0 4 I
-a 4 oa-QUT^-aw
-i 
O < O*-
-------
                          INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) OlfiGRfitl
                          FROM FILE IOENT. JOHNSN. STfiTION  1970
                                   PREDHTOR 8784010101  GOBIESOX MERNORICUS
                               (N.  CLINOFISH      )   BOJUSTED SfiMPLE SIZE =  78
UJ
(_)
1 80
! so
m
§
~ 40
1

2

§ -20
5
1 -60 • i
1 : 1 j
* on w - -80 • -« * o *
S i S S
31 C 3 i
3
U «
*^ 3
^ u
• V « •
|r-i
*^ ^ i
3655
— i — . rr~i ^T_
— i — (_j Lr|-i:[iti
S
•
U
i 5.1
• 1 -H £ W
!« o a.
n a. » u t- • « K
2 5 " S •? S 851? SS
: s : s g-ssss-ss
I • ue. MO.««U-H£
3« ^ o £ qcvatx^
Z <-*0. KXUHOrM
0 50 100 150 200 250 300 350
CUflULflTIVE FREQUENCY OF OCCURRENCE
	 _ FO£0 NU*. GPAV. PPEY PERCENT
P"E»"lTF" " * OCCUP CO»P. COMP. I.P.I. TOTAL IPI

•MOMITMOF so. ss.no i°.«-n
_LITTnPI..l so. }?'.t)0 io!so
•C-AFA SP. ------ 3l^nn 10>fr0
f-AM-APIOEA 2fl.no 11. «0
HACSAPMS 5R. ?n.OO 4.90
IOOTFA SP. 19.no 3.40"
OOCPXT3A 17.00 .00
_pM|70°nOFA-rno«MlNlFEPI9A I7.no 1?.?0
^AooaCTicnlla u.nn *.30
"I"1!^'5 »,!no U70
•jsTPicno* ,,.no - 3.«o
KATHAK1-4A TllWlf«T» s]no ]SO
CncAI.L INATE4 Von .On
Tat s=. i.no ~:?n
	 P°FY T'»a WITH F°FO. icri.o. I.F<;<; TMJN s »Nr
rn-fco<;iTInM OATH i.tss TMCM | «OF ExCLuOEP F
(RllT 'JO' roni.' C»l CULATI"" OF OlVFPSJTY [NO
SH«..MO»,-^r|..Ft, OIvEc^nr 3.34
Evfv-tss I«.n£, .R<1
2S.no ?4Si.o 4S.oo
9.00 «,7S.S 1?.40
l.nn 3A«.n A. BO
12.00 700.* '12.90
2.00 186.4 7.10
.no o«.n 1.80
4.00 140. is 2.AO
9.00 153.0 ?.iO
_.0o ?n7. s.sn
.00 g«. " " J.fto
6.00 4?. .90
.00 10. .20
.00 22. .40
3.00 72. .40
2.00 14. .10
•i.OO 77. .SO
1.00 3. .00
*.00 19. .40
3.00 3.? .00
MJ"F()ICAt AND G°»VIMETPIC
"O'" THE TARLF »«m PLOT ~
CFS)
3.03 2.71
.
-------
                           INDEX OF RELRTIVE IMPORTHNCE (I.R.I.) DIflGRRM


                           FROM FILE IDENT. WW  BS. STRTION  flLL


                                      PREDflTOR  8791030601  - MICROOflDUS PROXIMUS

                                  (PflCIFIC TOMCOD      )   fiOJUSTED SflMPLE SIZE = 46
                        lOOr
                         80
                    :    so
                    CD

                    §

                    "    40
                         20
                    §    20
                    2    4°
                    I—



                    1    60






                    S    80

                    a.




                        100
                                                                   « r
                                                                   t- i
                                                                   to ra
                                                       V   y-.   ,-n
                                                               u. s: o a.
                                20
                                      40
                                           60
                                                 80
                                                      100   120    140   160   180    200
                                         CUMULflTIVE FREQUENCY OF OCCURRENCE
PREY ITEM
GAHHARIDFA
CALANOIDA
PEMAEIDAE
ClIWACEA
VALVIFERA
FLARELLIFFRA
M Y^ I DAE
OSTEICHTHYES
PAGURIOAE
OECAPODA-RRACHYURA
PFRACARIDA-MICROCEPREPIDEA
FREQ NUM.
OCCUR COMP.
70.00 19
41.00 29
33.00 12
IS. 00 5
13.00 1
11.00 3
7.00
?.oo
?.oo
2.00
?.00 2
41
.PI
.94
.63
.64
.05
.94
.47

.47
.P2
COMP._
7.67
.52
f 3.74
.72
2.3?
?.89
.2?
8.12
1.11
2.31
P. 89
PREY PERCENT
I.R.I. TOTAL IRI
l?97.n 44.
1?43.S 16.
2530.4 34.
95.? 1.
5] .S ,
65.3
«. 1 .
17.? .
2.7
5.6 .
23.4
70
90
50
30
70
90
00
20
00
00
30 ~~~
            PREY TA»A WITH  FREO. OCCllP. LE";1; THAN 5 AND NUMERICAL  ANH OOAVIMETRIC

            COMPOSITION BOTH  LESS THAN 1  ARE" F XCLIIDF n FOO" THE TABLE ANO PLOT

            (HUT NOT FROM CALCULATION OF  DIVERSITY  INDITES)
               PEBCENT"OnMlNANCF INDEX

               SHANNON-WFINER DIVERSITY

               EVENNESS INDFX
                                   .27    .43


                                   .2?   l.BR

                                   .64    .54
 .35

1.7?

 .57
Fig.  B-21.
IRI prey spectrum of  juvenile  Pacific  tomcod from shallow

sublittoral  habitats  along the eastern shoreline of  northern

Puget Sound.
                                            112

-------
     Pacific tomcod,  mainly juveniles (65%),  were often caught in abundance
in both beach seine and townet collections along the Strait of Juan de Fuca.
Beckett Point, Jamestown and Point Williams,  Morse Creek, and West Beach
contributed the most specimens.   The overall  prey spectrum (Fig. B-22) is
composed almost exclusively of epibenthic crustaceans, including gammarid
amphipods (84.3% of total IRI),  mysids (4.0%  ,  including Archaeomysis
grebnitzki), hippolytid shrimp (3.5%, including Heptacarpus brevirostris).
harpacticoid copepods,  cumaceans,  and unidentified shrimp.

     Juvenile tomcod collected in Nisqually Reach (Fresh et al. 1979) had
fed predominantly upon epibenthic crustaceans,  gammarid amphipods and
hippolytid and crangonid shrimp.

Walleye Pollock (juvenile)

     Most of the juvenile walleye pollock collected in northern Puget Sound
originated from winter beach seine collections  at Birch Bay and Cherry Point.
Epibenthic or benthic organisms were the principal prey organisms; gammarid
amphipods constituted 69.6 percent of the total IRI, valviferan isopods
contributed 7.8 percent, while hyperiid amphipods, shrimp, and calanoid
copepods made up lower contributions (Fig. B-23).

     In the Strait of Juan de Fuca, juvenile  walleye pollock occurred mainly
in fall and winter beach seine collections at Beckett Point and Dungeness
Spit and in townet collections at Jamestown and Point Williams.  Calanoid
copepods, because of their numerical predominance, constituted the most
important item in the IRI prey spectrum (67.1%  of the total IRI) (Fig. B-24).
Gammarid amphipods (15.8%), hippolytid shrimp (including Heptacarpus
brevirostris, 8.8%), mysids (including Archaeomysis grebnitzki, 3.1%), and
cumaceans (1.1%) were the other prey of significance.

    Post-larval pollock collected in late spring in the neritic waters of
the Strait of Georgia (Barraclough 1967a-c);  Barraclough and Fulton 1967,
1968; Robinson et al. 1967 a, b: Barraclough  et al. 1968; Robinson 1969)
indicated that calanoid copepods and crustacean larvae were the prevalent
organisms consumed by the pelagic juveniles.

Threespine Stickleback

     Threespine sticklebacks were the second  most frequently encountered
neritic species in northern Puget Sound.  The more important prey organ-
isms were both epibenthic—harpacticoid copepods (67.0% of the total IRI)
and polychaetes (3.7%)—and pelagic—calanoid (14.2%) and euphausiids
(13.3%) (Fig. B-25).

     Threespine stickleback collected by beach  seine in the southern North
Sound sites at Guemes Island and Padilla Bay  had also consumed epibenthic
crustaceans; 76.4 percent of the total IRI was  gammarid amphipods, 14.8 per-
cent harpacticioid copepods, 4.6 percent crab larvae and 1.3 percent cumaceans
(Fig. B-26).
                                   113

-------
             lOO
               INDEX OF RELRTIVE IMPORTRNCE (I.R.I.) DlfiGRRM
               FROM FILE IDENT.  76-78. STfiTION RLSTR
                               8791030601 - MICROOflDUS PROXIMUS
                                            PflCIFIC TOMCOO
                                 RDJUSTED SRMPLE  SIZE r 272
                              CUMULflTIVE FREQUENCY OF OCCURRENCE
                             FDEO   NUM.    GPAV.   PREY     PEPCENT
PREY ITEM
GAMMARIDEA
"YSIDACF.A
CIJMACEA
SPHAEROMATinAE
GAUMARIDAF
HIPPOLYTIDAE
POLYCHAETA
NATANTIA
HABPHCTICOIOA
CALANOIDA
_TRf NGONlDAE 	 	 	 	
FtlOHAUSIACEA
PAsOALIDAf
OSTFICHTHYS
CLIIPEIDAE
Alll OPHVJCHinAE
OCCUR (
55.15 '
?3.53_
19.85
17.65
17. ?«
13.60
1P.13
11.76
7.15
*.62
	 6.62 	
4.04
?.9ii
1.84
.17
.17
:OMP.
56.40
6.4)
?.78
i .e.R
1.P5
1 .66
1 .44
2.17
9.78
7.95
.55
1.84
.24
.06
.01
.01
COMP.
35.79
*.09 ...
.84
1.16
1 .55
13.86
3.6?
3.14
.09
.20
5.65
1.01
13.3?
8.34
2.19
1.21
I.P.I.
^083.9
246.9
71.9
53.6
93.3
?13.H
6) .It
62.4
7?. f,
54. n
41.1
11.5
19.9
15.5
.8
. i.
TOTAL IRI
P?.84
4.02
1.17"
.87
1.5?
3.48
1.00
__... 1.02
1.18
.88
.67
.19
.65
.25
.01
.01
PREY. TAXA KITH FREQ. OCCUR. LESS THAN 5 AND NUMERICAL  AMI  GRAVIMETRIC
COMPOSITION POTM  LESS THAN 1 APE EXCLUDED FPOM THF  TAPLE AND PLOT
(RuT NOT FPOM  CAI CULATION OF DIVERSITY INDITES)
Fig.  B-22.
   PEPCENT  DOMINANCE  INDEX
   SHANNON-WMNEP  DIVERSITY
  .EVENNESS INDEX  . .  ....
                                                  .34
                                                 2.50
                                                  .47
 .18
3.?0
 .60
 .69
1.24
 .23
    IRI  prey  spectrum of  juvenile Pacific tomcod in  the  Strait  of
    Juan de Fuca.
                                   114

-------
CD
s
g
fe
s
                           INDEX OF RELRTIVE  IMPORTRNCE (I.R.I.) DIRORRM
                           FROM FILE IDENT. WW  BS. STflTION flLL

                                     PREOflTOR 8791030701 - THERRGRR CHflLCOORRMHfi
                                  (HRLLEYE POLLOCK     )    ROJUSTEO SRHPLE  SIZE * 20
                        100
                         80
                         60
                         40
                         20
                    2    20



                    i    40
                    I—
                    I    60


                    ^    30
                    u

                        100
                                  20
                                           40
                                                   60
                                                           80
                                                                   100
                                                                           120
                                                                                    140
                                         CUdULflTIVE FREQUENCY OF OCCURRENCE
	 P°EY ITEM
FREQ
OCCUR
NUM.
COMP.
GPAV
COMP
PREY
. 	 I.P.I. _
PERCENT
TOTAL IRI
           VALVIFEPA
           AMPHIPOOA-HYPERIIDFA
           CA|
           CRUSTACEA
          .POLYCHAETA _
           EUPMAUSIACEA
           FL'BELLIFERA
                      ?n.no   7.94
                      10.00 __ 3.18
                      in.00   11.90
                       ^.00    .79
                       5.00 _  .79
                       ^.00   1.59
                       5.00   2.38
                                   10.95  ]084~.7
13.93
 3.98
17.91
  .00
 1.00
 3.48
171.1
158.8
 93.5
.  *.0_
 12.9
 98.9
A9.90
17.80
 3.40
 ?.80
 ?.60
 1.50
  .00
  .20
 1 .60
             POFY TAXA WITH FREQ. OCCUR.  LF^^ THSN 5 AND NUMEOICAL  ANfl HP4VIMETRIC
             COMPOSITION BOTH LESS THAN  1  ARE EXCLnnEH FonM THF  TAPLF ANin PLOT
                  i"T FBOM_CA| CUL4 T ION OF
                PERCENT DOMINANCE INDEX
                SH4NNON-WFINER DIVERSITY
                EVENNESS  INDEl
                              .28    .28
                             2.20   2.26
                              .70    .75
                  .52
                 1.46
                  .49
Fig.  B-23.   IRI prey  spectrum of juvenile walleye pollock in  shallow  sub-
               littoral  habitats along  the  eastern  shore of  northern Puget  Sound.
                                             115

-------
                      >-
                      CO
                      s
                      5
                      §
                      CJ

                      I—
                      o
                           lOOr
                              INDEX OF RELRTIVE IMPORTRNCE  (I.R.I.) DIRCRRM
                              FROM FILE  IDENT.  76-78. STRTION RLSTR
                                            8791030701 - THERRORR CHRLCOORRMMR
                                                           WRLLEYE POLLOCK
                                                RDJUSTED SRMPLE SIZE =  74
                                                                            200
                                                                                        250
                    PPFv ITEM
                                            CUMULRTIVE FREQUENCY OF OCCURRENCE
                                                  NUM
                                           FREO
                                           OCCUR
                     GRAV.
                     COMP.
P9EY
I.B.I.
 PERCENT
TOTAL IRI
           GAMUARIDAE
           CALAN01DA
           GAMM4P10E*
           CilMACEA
          . MY.<;IDACFA _____ ___
__  	 31.08 _79
       21 .*>?
           NATANTIA
           EIIPHAUSIACEA ______
           OSTPACOOA
           HAPPACTICOIDA
          -ElJCAPIDA-DECAPODA-SpAChYRIYNCH
           PSC'OALIDAE
           POLYCHAFTA
                     .
           CRANGONIOAE
        14.8ft
        13.SI
        10.81
        8.11

        6.76.
        S.4]

        ?!70
58
71
b2
82
82
90
49
77
20
38
15
20
15
05
0?
8.92
21.07
6.61
1.57
_ 5.75
P6.84
1.90
3.48
.26
.08
. 15
10.70
1.30
7.24
1 .35
388.6
3132. 3 _
348.1
51 .*.
- 143.? 	 .
412.3
32.3
- 45.9
3.7
9.9
a.o- 	
S8.9
7.9
_. 19.7
1.9
8.32
67.06
7.47
1 .10
-3.07
8.83
.69
-- .98
.08
.21
,04
1.26
.17
.4?
.04
             RRFY TAxA_nITM FDfQ. OCCUR.  LF^S_THAM «i  AND  NUMERICAL  AND  G" A V I METR I C_
             COMPOSITION HOTH LESS THAN 1  ABE EXCLUDED  FROM THE  TABLE  AND PLOT
             (BUT NOT FRO" CALCULATION OF  DIVERSITY INDICES)
                PERCENT DOMINANCE INDEX
                SHANNON-WEINE" DIVERSITY
                EVENNESS INDE«	
                 .6^     .15
                1.31   3.22
                 .27     .67
            .47
           1.81
            .38
Fig. B-24.   IRI prey  spectrum of juvenile  walleye pollock in Strait of Juan
               de Fuca.
                                              116

-------
          Along the Strait  of  Juan  de  Fuca,  adult  threespine stickleback
were most common in Beckett Point,  Jamestown,  and  Point Williams beach
seine collections.  Unlike  northern Puget Sound, it was not often
encountered in the townet collections.  Threespine stickleback appeared
to be feeding throughout the nearshore water column, as pelagic calanoid
copepods and epibenthic harpacticoid copepods were qually important (Fig.
B-27).  Limited sample sizes,  however, do not allow us to determine whether
this  catholic feeding beahvior is due to diel changes, site differences or
collection methods.   Secondary prey organisms were mostly epibenthic forms,
including gammarid amphipods and mysids.

Bay Pipefish

           Several adult  bay pipefish from beach seine collections at Birch
 Bay were large  enough to  permit analysis of their  stomach contents; 86.8
 percent of  the  total IRI were isopods, the remainder, gammarid amphipods.
 Tube-snout

           Tube-snouts were frequently captured in the mud/eelgrass  and  sand/
 eelgrass habitats and pocket gravel beaches in northern Puget  Sound.  Common
 identifiable organisms included gammarid amphipods  (90.0  percent of total
 IRI);  only  polychaete annelids  (6.3 percent), and crab larvae  (1.8  percent)
 were secondary prey.  Tube-snouts  from beach seine  collections along the
  eastern shoreline,  Birch  Bay and northeast Guemes Island, tended to have more
  pelagic organisms  in  their diet.   Pelagic calanoid  copepods  composed 73.5
  percent of the total IRI,  while  harpacticioid copepods  (32.1  percent), gammarid
  amphipods (23.9 percent), and mysids  (7.6 percent)  made up the principal
  epibenthic  prey composition.

            Along the Strait  of  Juan de Fuca, beach  seine and townet  collections
  at Beckett Point and  Morse  Creek produced numerous  tube-snouts.  As in the
  case of threespine stickleback, both calanoid  and  harpacticoid copepods were
  the principal prey species  of  tube-snouts  feeding  in nearshore habitats
  (Fig.  B-28) .  Shrimp larvae,  though constituting 28.9% of the  total prey
  biomass, were not abundant prey items.

             Tube-snout from the Nereocystis  kelp beds at Barkley Sound examined
  by Leaman  (1976) had consumed primarily barnacle larvae  and caridean
  crustaceans.

  Kelp Greenling

             Kelp  greenling were the most commonly observed  fish in the rocky/
   kelp bed habitats  of northern Puget Sound  and often characterized  the protected
   gravel beach  enviornments such as those at Deadman Bay and Legoe Bay   The
   prey spectrum for  kelp greenling  (Fig. B-29)  was one of   the most diversified
   so   rank^T d°C™ented-  *npMpods, principally Eusiroides sp.,  Amphithoides
   sp., ranked as the most  important (46.1 percent of total IRI)  prey, followed -
   by crabs,  Cancer  magister, Pugettia ^racilis ,  Oregonia gracilis  Telmessus
   chexra.  and  unidentified           -
                                      117

-------
                            INDEX OF RELRTIVE IMPORTflNCE (I.R.I.) DIHGRflM
                            FROM FILE IDENT. N POSD. STHTION flLSTfl
                                      PREDftTOR  8818010101
                                    (THREESPINE STICKLEBK)
 CflSTEROSTEUS flCULEflTUS
RDJUSTED SflMPLE SIZE    14
                     m
                     g
                     8
                     Q_
                         100
                          80
                          60
                          40
                          20
                     -    20
                          40
                     °    60
                     §
                     (_>
                     i-l    30
                          100
                                                                               I  3 C
                                  20
                                         40
                                                60
                                                      80
                                                             100
                                                                   120
                                                                          140
                                                                                 160
                                                                                       180
                                           CUMULflTIVE FREQUENCY OF OCCURRENCE
                     PREY ITFM
                                            OCCUR
                                                   NUM.
                                                   COMP.
CPAV.
COMP.
              PREY
              I.R.I.
 PERCENT
TOTAL IRI
           -HAopACTlcoirrA
            POLYCHAFTA
            CORF.PODA	
            CALANOI'lA

            FUPHAUMACEA
            AMRHIPonA-rlYPEWI IDEA
            LARVACFA	
            FLARFLLIFFRA
~~ 50.00
43.00
14.00
14.00
14.71(1
7.00
~~ ~~ 7. on
7.00
7.00~~
7.00
ACFA 	 7.00""
55.97
.65
33.77-
.47
" .19
.65
.47'
1 .03
.09 -
.09
2.15
.00 ?798.S
2.90
P. 17- '
.00
.00
7H.84
.-.1"
.41
.00
4. 15
.4]
152.6
594.7 —
6.6
2.7
556.4
6.2
10.1
.6
29.7
17.Q
67.00
3.70
14.20
.20
.10
13.30
.10
.20
.00
.70
.40'
              PPFY TMXA  WITH FDFO.  nccMB.  irc.s. THAN'S AND NUMERICAL" ANT GRAVIMETRIC
              cn>'pn<;iT ln\ POTH L^S THAN i  AOE FKCLHPF.O FROM THE TABLE AND PLOT
              (R'lT NOT  FRO" CA| CUL'TION OF  nivFR^TTY
                 PF5CENT DOMUJANCF INOFX  "
                 SHAN'JON-WF INf P RIVE>"^ITY
                          INPtt
— .43
 1.19
  .40
 1.00
                  1 .49
                   .45
Fig. B-25.    IRI  prey spectrum  of  threespine  stickleback  from northern  Puget
                Sound.
                                               118

-------
                    s
to
g
I—
§
fc
s
                           INDEX OF RELflTIVE IMPORTflNCE (I.R.I.) DIflORRM
                           FROM FILE IDENT. WW  BS. STflTION  flLL
                                     PREDflTOR  8818010101 - GflSTEROSTEUS flCULEflTUS
                                  (THREESPINE STICKLEBK)   RDJUSTED SRMPLE SIZE   15
                         100
                         80
                         60
                          40
                         20
                    §   20
                          40
                         60
                          80
                         100
                    PREY ITEM
                                20
                                      40
                                            60
                                                  80
                                                       100   120    140    160   180   200
                                          CUMULflTIVE FREQUENCY OF OCCURRENCE
                                          FREQ
                                          OCCUR
                             NU".
                             CO«P.
             GRAV.
             COMP.
              PREY
              I.R.I.
               PERCENT
              TOTAL  IRI
                       _ __
           " HiBPACTICOIDA
            OEC»POD»-BPACHYU"A
            CU-ATEA     ____
            OLIGOCHAETA
            CPL'STACFA
            CIRRIPEDIA
            LARVACEA
33.00
27.00
27.no
n.no
 7.00
 7.00
 7/00
 7.no
 27.74
"34.35
  4.84

   .48
 14.6«
	.16
   .16
   .65
1.75
8.77
1.75
7.02
 .88
 .00
3.51
 .00
6)49.3
1191.3
 167.5
 108.3
  97.^5
 108.9
_  1.1
  ?5.7
                                                    76.40
                                                    14.80"
                                                     4.60
                                                     1.30
                                                     1.20
                                                     1.40
                                                   	.00
                                                      .30~
                                                      .10
             PRFY TA«A 1.ITH FPEQ.  OCCllH. L^S^ THAN 5 AND  NUMERICAL ANn GOAVIMETRIC
             COMPOSITION OQTH LESS THAN 1 ARE EXCLUDED FROM  THE  TARLF ANO PLOT
             _[B|lJ_NOT_FHnu_CALCULAT^ON OF DIVERSITY INDICES)
                PERCENT DOMINANCE  INDEX
                SHANN"N-WI\FO DIVERSITY
                EVENNESS INOFi
                               .22
                              1 .90
                               .60
                .42
               1.42
                .51
                          .61
                         1.19
                          .40
Fig.  B-26.   IRI prey  spectrum  of  threespine stickleback from shallow sub-
               littoral  habitats  along the eastern shoreline  of northern  Puget
               Sound.
                                             119

-------
                      o
                      o
                      
-------
                      fc
                          100
                           80
                           60
                           40
                           20
                           20
                           40
                           60
                           80
                           100
                             INDEX OF RELHTIVE IttPORTflNCE (I.R.I.) DlflORflM
                             FROM FILE IDENT.  76-78.  STfiTION flLSTfl
                                            8818020101 - flULORHYNCHUS FLflVIOUS
                                                             TUBE-SNOUT
                                               flOJUSTED SflMPLE SIZE    82
                                      20
                                                40
                                                          60
                                                                    80
                                                                              100
                                                                                        120
                                            OMILRTIVE FREQUENCY OF OCCURRENCE
PREY
ITEM
FPFO
OCCUR
NUM.
COMP.
GRAV.
COMP.
PREY
I.O.I.
PERCENT
TOTAL IRI
3S.37 4?. 7R
34.15 28.59
in. 98 ?.34
o.76 8.RS
7.3? - _ .8?
30.9?
6.11
8.64
?8.89
3.47
9.73
7.84
2.98
7606.5
1184.1. _
120.5
368.?
31 .-5
81.9
— . 4.1 —
58.53
26.60
2.7)
8.27
-7]
1.09
1 .84
- .09 	 	 	 _
           CALANOIDA
           HAPPACTICOIOA	

           MATANTIA
           GAMMARIOEA	
           OSTFJCHTHYS
           I APVACEA
           -H1PPOLYTIDAE	  -

             ODFY TAXA uITH FPEO. OCCUR. LESS THAN S ANO NIJHERICA|  AND GRAVIMETRIC
           	CO.MPHSITIIN POTH LESS THAN i_ ARE EXCLUDED FDOM THE TABLE *ND_OLO.I	
             (BUT NOT FPQM CALCULATION OF DIVERSITY INDICES)
                PERCENT QQMINANCE .INDEX 	
                SHANNON-wFlMfp DIVERSITY
.29    .21
.ll   ?.65
.SS    .70
 .42
1 .65
 .43
Fig. B-28.   IRI prey  spectrum of  tube-snout  in  Strait  of  Juan de Fuca.
                                               121

-------
                                INDEX OF RELflTIVE IMPORTflNCE (I.R.I.) OlfiGRRM

                                FROT1 FILE IDENT. N PGSD. STHTION BLSTfl

                                         PREOflTOR  8827010101  - HEXflGRHtlMOS OECflGRflftflUS

                                       (KELP GREENLINC     )   RDJUSTED SflMPLE SIZE   31
                              100
                              80
                               60
                               40
                               20
                         1    20
                               40
                               60
                               80
                              100
                                                             £
a.  e  f. —
o  x  u -*
u  •£  H ^

3  1  SC
                                                                                 •o a
•O M -O X — u u
O -Q -4 p^ k. O U
o. o i- o 3 -< n

u
7
2
6
n



33.
4.
1.
S.
.
	 3.
.
B
.
1.

90
40
80
no
00
20
80
30
00
40
10
30
80
9n
60
10
10
00
10
70
• JO
GRAVIMETRIC 	
Ann PLOT

2.

01
Fig.  B-29.   IRI  prey spectrum of  kelp  greenling  in  northern  Puget  Sound.
                                                    122

-------
isopods (34 percent) fishes  (1.9 percent),  polychaetes (3.7 percent),
gastropods  (3.5 percent), and  tanaids  (1.2  percent).   Although algae
(Rhodophyta  33.9 percent), constituted a measurable percentage of the total
biomass,  these were  considered incidental items,  byproducts of the feeding
of  kelp greenling  on the predominantly benthic prey items.

      The  only food data for  kelp  greenling in the Strait of Juan de Fuca,
 that from Leaman's (1976)  collections  at Barkley Sound, Vancouver Island,
 indicated that caprellid (Caprella gracilion, £.  laeviscula, C^. ferrea, C_.
 incisa,  and £.  equilibra)  and gammarid amphipods (Parapleustes pugettensis,
 Erichtonius brasiliensis.  Photis  sp.,  Podocerus sp., Metaphoxis sp.) were the
 most important prey; fish (Artedius lateralis. Synchirus gilli), crabs
  (Pachycheles sp., Cancer gracilis, C.  oregonensis, Pagurus sp. , Oregonia
  gracilis, Loxorhynchus crispatus, Petrolistes eriomerus) and shrimp
  (Heptacarpus sp.) were of secondary importance.  The diet of greenling between
  101-200 mm in length was dominated by isopods while that between 201-700 mm
  was  prevalently crabs.

      Juvenile kelp  greenling were reported by Barraclough and his co-workers
  to be one  of the  most  common  fishes in the neritic waters of the Strait of
  Georgia  in May-June (Barraclough 1967 a, b, & c; Robinson et al. 1968 a & b).
  Calanoid copepods (Calanus plumchrus, Pseudocalanus minutus), cladocerans
  (Podon sp.)  hyperiid amphipods (Parathemisto pacifica) and larval fish
  (eulachon) were important prey taxa at this stage (Table B-3).

  Whitespotted Greenling

       In  northern  Puget Sound  adult whitespotted greenling was collected most
  often at pocket gravel beaches such as at Deadman Bay during beach seining.
  It was also  collected  in the  rocky/kelp bed habitat, but in much smaller
  numbers  than the  kelp  greenling.

      The  prey spectrum of the whitespotted greenling was very similar to  the
  kelp greenling (Fig. B-30).   Gammarid amphipods, especially Eusiroides sp.,
  Amphithoe  sp., constituted the most important food item, contributing 55.7
  percent  of the total IRI.  Shrimp (Heptacarpus stimpsoni), with 22.4 percent,
  and  various brachyuran crabs  (Cancer oregonensis, Pugettia gracilis), with
  11.4 percent of the total IRI, were secondary; fish and polychaete annelids
  provided less than  5 percent each.  Incidental algae, however, was not as
  significant  in the  overall diet composition ofthe whitespotted greenling  as
  the kelp greenling.  Whitespotted greenling were often collected during
 beach seine sampling at northeastern Guemes Island, Birch Bay, Cherry Point
 and Legoe Bay along the eastern shoreline.  Despite the different areas and
 habitats, prey composition of this sample was very similar to  those from  San
 Juan Island collections, with gammarid amphipods responsible for 65.9 percent
 of the total IRI; penaeid and callianassid shrimp, 21.6 percent; brachyuran
 crabs, 10.0 percent, and polychaetes, 1.3 percent (Fig. B-31).

      Along the Strait  of Juan de Fuca, whitespotted greenling, most of which
 were juveniles, were included in beach seine collections at  Beckett
 Point and Point Williams.  The overall prey spectrum of whitespotted greenling


                                      123

-------
Table  B-3.  Prey composition  table for  juvenile kelp greenling in  neritic
             waters of  Strait  of  Georgia,  documented by Barraclough (1967a)
 PREDATOR ^8927010101  - HEXAGRAMMOS  DECAOPAMMUS               (KELP GREENLING
           	INDEX OF~RELATIVE  IMPORTANCE  (I.P.I.) TABLE
                 FROM FILE IDENT.  REF   1,  STATION  ALSTA

FREO "" NUM. GRAV. PPEY PERCE
PREY ITEM OCCUR COMP. COMP. I.R.I. TOTAL
:NT
IRI
  -CALANOIDA ----  -------------- 98.90   97.50
   AMPHIPQDA                        S.40    1.00
  "OSTEICHTHYES --------- -------- 3.30  ---- .'30
   EUPHAUSIACEA                     ?.20     .10
  "CRUSTACEA          ~ -------- ?.?0    1.00
   THERAGRA CHALCOGRAMMA            1.10     .10
  ~OIKOPLEURA^SP.                    1 .10     MO
               WTTH^FREO.~~OCCuRT~LE:<;s"THAN  S "ANO~NUMERICAL
     COMPOSITION ROTH LESS  THAN  1  ARE  EXCLUDED FPQM THE TABLE AND PLOT
     
-------
                           INDEX OF RELRTIVE IMPORTRNCE M.R.I.) DIRCRfitl
                           FROM FILE IDENT. N PGSO. STRTION RLSTfi
                                    PREDRTOR 8827010104 - HEXflGRRdMOS STELLERI
                                 (HHITESPOT GREENLING 1   HOJUSTED SflflPLE SIZE =  13
100
B
1 80
! 60
m
•z.
o
~ 40
8
8:
8 2°

t—
<_>
o> „
0
I
COMPOSITION BY HEIO
8 £ S
>- 80
inn

1 —

mi—
1 «

I
ti
t « 0
S 5 .
S £• 6
V % S
« a 4 i- . W « « J=
« <• q £ X •-> 1 C.
•v -a e *j£«*«
•H 5 4i ,C G. * T3 O
^ a- & ^£-5S.5
E £ O «IOX«h>
E & » Wf-i^tuo
0 5 2 OUbOU




Trf B^
P
Lr
0)
•
V
k. a
>. • -b u x-r* w w 
acectnoniwoc u «

                                  SO      100      ISO      ZOO      250

                                        CUMULflTIVE FREQUENCY OF OCCURRENCE
                                                                         300
                                                                                 350

PRCT ITEM
~Gl»M»RIntA~
*HPH1POO<
PLtOCYFM»T»-CAR10E»
pf-D«c«Pln»-SPEL»fO&P»P>-»CE*
O^TFICnrMYt5
CHI.OnOPMTT*
POLTCKAET* "~
OEC»nOD»-BP*CHYljB«
OMnP|MOSPM«EPOM« OBEOONENSIS
PH«frPHYt»
o«r>"PlD*E
P»Kn«LI04E
MY<;IO»CE«
p"COnpMYT»
rL«1F.LLIFE»»
OE<»«P.EStI»CE»E
MOOD
PEK»EID»E
DEOPnO*
V«LVIFEP».
FBEQ
OCCUS
3M.OO
3A.OO
38.00
31.00
31.00
33.00
Z3.00
15.00
15.00
e.oo
A. 00
x.no
B.OO
ft. 00
A. 00
e.oo
8.00
8.00
A. 00
«.oo
NUM.
COMP.
27.91
11.63
<..65
16.7*
3.3*
3.72
2.79
?0.93
2.3)
.93
.93
.'7
.47
.*7
.47
.*7
.47
1.40
.47
.47
GP*V.
COMP.
1 T74 "
5.67
16.54
8.04
4.69
1.26
6.26
1.42
.23
.07
22.46
16.37
.00
.03
.00
.10
.04
7.8*
1.20
1.01
PREY
I.R.I.
"f 126.7 ~
657.4
805.2
768.2
372.9
114. s
208.1
315.2
38.4
8.0
187.1
134.7
3.8
4.0
3.8
4.6
4.1
74.)
13.4
11.4
PERCENT
TOTAL IRI
T3~.10
13.50
16.50
15*80
7.60
2.30
4.30
6.90
.80
.00
3.80
2.80
.00
.00
.00
.00
.00
1.50
".30
.20
             PBFY TAXA yITM rpfQ.  OCCljK. LESS  THAN 5 AND NUMERICAL AND GRAVIMETRIC
             COMPOSITION BOTH LESS THAN 1 ARE  EnCLUOEO FPOM THE TABLE AND PLOT
             <«IUT NOT FRO" CALCULATION OF DIVERSITY INDICES!
                PEoCENT Or-lNtNCE  IMCEX
                S^iv.^'J-wF I M P DIVERSITY
 .17
3.0'
 .71
 .13
3.19
 .7'
 .14
3-11
Fig.  B-30.    IRI  prey  spectrum of  whitespotted  greenling  in northern  Puget
                 Sound.
                                                   125

-------
                           INDEX OF RELflTIVE IMPORTANCE (I.R.I.) DlflGRflfl
                           FROM FILE IOENT. WH  BS. STflTION  RLL

                                      PREDRTOR  8827010104   HEXflGRflMMOS STELLERI
                                  (HHITESPOT GREENLINO )   flDJUSTED SflMPLE SIZE =  13
                         100
                         80
                    :    eo
                    oa
                    o
                    £    40
                    if
                    S    20
                    2    20
                    £
                    >-
                    (O
                    z    40
                    g    60
                         80
                         100
                                20
                                      40
                                            60
                                                  80
                                                       100   120   140    160    180   200
                   PREY ITEM
                                          CUMULRTIVE FREQUENCY OF OCCURRENCE
FREO   NUM.    GRAV.  PREY     PERCENT
OCCUR  COMP.   COUP.  I.P.I.  TOTAL IRI
          _GAMMARir>FA^	6P.no   75.70
           PENAE'IOAE                       38.00    6.21
           EUCARIDA-OECAPDDA-RRAC^YRHYNCH  23.00    3.95
           OECAPODA-BRACHYI)RA_	     15.00    1.13
           POLYCHAETA                  "   15.00    1.69
           TANAIOACEA                       8.00    2.82
           DECAPOOA-BRACHYUPA	«.00    2.26
           EUPHAUSIACEA   ~                  fl.OO     .56
           VALVIFERA                        S.OO     .56
          _TARDIGRADA                    	 8.00   _ .56
           PERACARIOA-MICROCERBERIDEA       B.no     .56
           PLEOCYEMATA-CAPIDEA              «.oo     .56
           CALL1ANASSIOAF                   8.00     .56
16.83
23.12
15.60
P. 83
3.69
.22
6.26
.35
1.10
.06
?.20
1 .69
15.97
3B02.'
1116.5
669.6
59.6
77.7
26.3
6B.O
7.3
13.3
5.0
22.1
Ifl.n
132.?
                              65.90
                              19.30
                               7.80
                               1 .00
                               1.30
                                .60
                             	1.20
                                .10"
                                .20
                                .00
                               " .60
                                •30
                               2.30
             PRFY TAXA WITH FREO. OCCUR.  LESS  THAN 5 AND NUMERICAL AND GRAVIMETRIC
           _ COMPOSITION ROTH LESS THAN 1  ARE  EXCLUDED FROH_THE TARLE ANH PLOT
             (HUT NOT FROM CALCULATION OF~DIVFRSITY INDICES)"           " ~
               PERCENT DOMINANCE INDEX
               SHANNON-HEINEP DIVERSITY
               EVENNESS INDEX
         .58	  .13
        1.63   2.65
         .39     .72
 .60
1 .61
 .65
Fig.  B-31.   IRI spectrum of  whitespotted greenling  in  shallow  sublittoral
               habitats along  the  eastern  shoreline in northern Puget Sound.
                                             126

-------
was one of the most  diverse encountered.  Gammarid amphipods were the most
important prey  (60.9%  of  total  IRI)  but  tanaids, polychaete annelids,
hippolytid shrimp (Heptacarpus  sp.), crangonid shrimp, bivalves and bivalve
siphons, and majid and pagurid  crabs all composed  more than 1% of the total
IRI  (Fig. B-32).

Lingcod

      The lingcod was commonly observed  in rocky/kelp bed habitats of northern
Puget Sound  (57 percent  frequency of occurrence during SCUBA transect
observations) and consituted the major top-level carnivore  in the fish assemblage
characterizing that habitat.  Stomach contents examined from eight adults and
six juveniles indicated  that lingcod were primarily piscivorous, with 36.1
percent of  the total  IRI being  fish.  Although the fish were usually digested
beyond recognition, rockfish (Scorpaenidae)  were  identified.  The remaining
 secondary food items  were benthic gastropods,  siphonophores, ascidians,
polychaetes, and  incidental algae.   Except  for the fish, which may or may not
be bottom-oriented, all  the prey items  were benthic.

      Lingcod sampled  during Quast's (1968)  detailed examination of  southern
 California's kelp bed  fish communities  had  consumed predominantly fishes
 (Perciformes, Clupeiformes) while algae and cephalopods were less representative
 prey (Table B-4) .

 Copper Rockfish

      Copper rockfish  were commonly  caught in beach seine collections in
 northern Puget Sound  during July and August  at Deadman Bay (gravel  habitat)
 and were frequently sighted along all SCUBA transects in the rocky/kelp bed
 habitat.

      The composite  prey  spectrum (Fig.  B-33)  suggests that copper rockfish
 were facultative  epibenthic feeders, having  consumed both  benthic and pelagic
 organisms.   General percaridan  crustaceans  were the more important  prey;
 these  included gammarid  amphipods (40.3 percent of total IRI), mysids  (6.0
 percent), shrimp  (3.0  percent)  brachyuran crabs (Cancer gracilis, Petrolistes
 eriomerus and Scyra acutifrons, 1.9 percent),  flabelliferan (4.9 percent)
 isopods, and cumaceans (1.4 percent).   Fish  (Pacific sand  lance and juvenile
 rockfish) accounted for  17.6 percent of the  total IRI.

      Juvenile copper  rockfish sampled by beach seine along the eastern shore-
 line (Legoe Bay)  had  a relatively similar diet composition based on epibenthic
 and pelagic prey.   Shrimp (Crangonidae,  Pandalidae, and Penaeidae)  and gammarid
 amphipods predominated,  with 36.1 oercent and  31.8 percent, respectively, of
 the total IRI.  crab larvae (15.9 percent) and  fish (threespine stickleback,
  J--J-/ percent)  formed  secondary  diet components.

  to
                                       127

-------
Table  B-4.  Prey composition (frequency of occurrence)  of lingcod in southern
             California kelp beds documented by Quast  (1968).
 PREDATOR  «p?7nin?ni  -  O°HIODON  KLONGATUS
                                            (LINGCOD
               innF.x OF RELATIVE  IMPORTANCE  (I.P.I.) TABLE
                      FILE  IPFNT.  REF 59. STATION SLSTA
            PRF.Y ITEM
                  FPFO   NUM.   GRAV.
                  OCCUR  COMP.  COMP.
                                                       PREY
                                                       I.R. I
                                                            PERCENT
                                                            TOTAL  IRI
TELt'OSTEI
UNin^NTIFI
THEUTHIDIDA
ALG4F
                                   1?."0
                                   11.00
          TAXA WITH FPFO.  OCCtlK.  LPSS  ThAN S AND NUMERICAL ANO GRAVIMETRIC
     COMPOSITION «OTH LESS THAN 1  ARF  FxCLUDFH FROM THF. TARLE ANO PLOT
     (RUT MOT FROM CALCULATION OF  DIVERSITY  INDICES)
        PERCENT DOMINANCF INDEX
        SH4NMON-WFINFC DIVERSITY
        EVFNNESS
                           .00
                           .00
                           .00
                                               .00
                                               .00
                                               .00
.00
.00
.00
                                         128

-------
                            INDEX OF RELHTIVE IMPORTHNCE (I.R.I.) DIBORRfl
                            FROM FILE IDENT.  76-78. STHTION RLSTfl

                                        8827010104 - HEXflORHnnOS STELLER1
                                                  HHITESPOT OREENLINO
PCT. COMPOSITION BY WEIGHT PCT. COMPOSITION BY RBUNDHNCE
SssftS^KSSSS
flDJUSTED SfltIP



«
• N « -5 «
u u n ArFA ?B.'6 11.32 .17 1?4-7 8.03
GAMMAPinAE 26.09 l?.9fl .33 347.0 8.58
HIP°nLYTlD»E 23.91 2.91 6.09 215.6 5.33
PHI YCHAFTA 10.57 4.81 9.81 »»*-«. 7. no
PtvALVIA 19.57 2.93 1.86 93.7 2.32
tPANKONlDAE 17.19 2.04 7.19 160.5 3.97
NATANTIA 15.?? 1.65 2.34 60. 0 1 .5(1
PAf-IIPIOAE 15.?? .89 |
GASTROPODA 13.04 .76
UNIOFNTIFIEr) 11.04 8.5? 1
.87 42.0 1.04
.25 13.3 .33
.94 136. S 3.38
DErAPnOA-HPACHYIlPA *.52 .39 2.94 21.7 .54
"AjTOAE 6.52 1.02 11.16 79.4 1.96
<-ADOFILIIJFA 6.s? |.27 .13 9.1 .23
FlirAP|r>A-r>S.CAPr>OA 4.15 1.15
CPI STACFA 4. 115 ?-f,7
_£.A.ir>aLinAr ,.17 _,s <;
.66 7.9 .19
.58 14.1 .35
.84 ?I.O .54
ATFLFCYCLIOAE ?,17 .25 15.06 33.1 .82
tA-^rptrjAF ?-17 -13 1-n 2>7 .07
-nSlEiCHTH^S 	 ^,7 .13 1.58 3.7 .09
"Tim»E z.,7 .3, 3.,H ?;; .;,
EMPioTocinir ,-l7 >n ,6>,6 3T>, _,z
	 »;01r_LlEA£ 	 ?,17 ,11 1.9? 4.s .11
"5Ir>T?¥I "'TH F"E0' nCC|ls- LES?^5127:' LE-iS^ LI»N_ L_ »Be._F.»ci.nneri F^OM THE TARLE AND BLOT
iSin NOT Ffco« CALCULATION OF OIvE»SITY INDICES) 	 	 	
EV-NNES5 ,ND,., ^^ 3;^ Z.«
	 ' 	 . 	
Fig.  B-32.   IRI p^y spectrum of whitespotted greenling  in shallow
              sublittoral  habxtats along Strait of Juan de Fuca.
                                            129

-------
                               1NOEX OF RELflTIVE  IMPORTANCE tI.R.I.] DIRGRflM

                               FROM FILE IDENT. N PCSO. SlflTION RLSTfl

                                           PREDRTOR  8826010108   SEBflSTES CRURINUS

                                      (COPPER ROCKFISH     ]   RDJUSTED SRMPLE SIZE    52
    100




g    80





I    6°

§
~    40

tn


3    20




°~     0


I—

I    20


>-
m


1    4°


1    60




^    80
a.


    100
                                                             >i  i-
                                                             J=   OJ "O U •-
                                                                   •o   •.- moiiootrc
                                                                       .    ,
                                                                   CE  QU^Ci
                                                                   nj 3    (_> o a. a.
                                          50         100         ISO         200


                                              CUMULRTIVE FREQUENCY OF OCCURRENCE
                                                               Y.—PPEY
                        P»ET ITEM
                                             -FUEO	NU"~
                                              OCCUR  COWP.   COMP.  I.R.I.   TOTAL  1RI
                                                                           PtBCtNT
                                                                                       250

PEPACARIDA-SPELAEOGRAPHACEA

DFCABOOA-flRACHVlIRA
PAK'DALinAF
CAPRFLLIDFA
BOLYCHAFTA
fiACTPOPOOA
- CA^FW ')P»C1LIK
<;CroPAFN|OAE
AMTMIIOIOAF
PETRnl.ISTHKS EBIOMFRUS
COMOOSITIOM ROTH LE5? THAN 1
	 TB1IT NOT FROM CALCULATION tlF~
5HAMNOij-wF"ii'r'E°NOI«£BMTY
EVt'JNLSS INUtY

31.00 17.
31.00 9 .
30.00 3.
lh.no ID.
l?.oo
l?.no 3.
10.00
in. no
lo.no
8.00
6.00 3.
A. no
4. no
3.00 1.
3.00 .
?.nn
:F«;«-rHnfr-5-
AUC FKCLllDE
"TJIVEB^ITY^
3.
•
74 	 2.~45 ~14
45 3.49 6
73 .51 3
41 34.00 5
SO .46 1
89 1.58
S4 4.04
8» 5.55
76 4.30
63 .0*
51 15.01 1
OS 1.44
38 .45
51 16!ll
S3 .11
1"< 	 5.17 	
35 3.38
13 3.34
ANTT-NDMEBICAL
0 FOQM THE TA
NUICtS)
16 . !«•
88 3.30
6S . fS

49.1
IS.n
39. f.
91. n
64.1
49. >,
7.1
ll.o
?4.3
43.9
16.4
9.7
66.^
13.5
3.3
10. «.
5.1
6~.9
Aijrr
1LE «




40.30
18.10
6.00
15.30
4.90
.80
3. SO
1.80
1.40
.30
.30
3.50
i.ao
.50
.30
1.90
.40
.10
.30
.10
."
(,OAVIMtTBlC
i«jn PLOT

3.77
.63
Fig.  B-33.   Spectrum of  copper  rockfish  from  northern  Puget  Sound.
                                                  130

-------
      Prince and Gotshaul (1978)  documented the prey spectrum of 241
copper rockfish captured around an artificial reef in South Humbolt Bay,
California.  In terms of frequency of occurrence and percent volume, juvenile
Cancer magister dominated the diet (40% of total IRI) while gammarid amphipods
composed 31% of the total number of prey items.  Crangonid shrimp, caprellid
amphipods and northern anchovy provided secondary food items.

Puget Sound Rockfish

     Puget Sound rockfish were documented only during SCUBA transect
observations in the rocky/kelp bed habitats along San Juan channel (Pt. George,
Shaw Is.)-  The overall prey composition indicated a relatively unspecialized
planktonic feeding behavior.  Calanoid copepods (57.3 percent of total IRI),
siphonophores (17.8 percent), and crab larvae (16.0 percent) constituted the
more important prey; hyperiid amphipods (5.1 percent), and crabs (1.6 percent)
were secondary in importance.

Yellowtail Rockfish

     Although juvenile yellowtail rockfish were often caught in the beach
seine sampling at Deadman Bay in northern Puget Sound, the majority originated
from rocky/kelp bed habitats around San Juan Island and Burrows Island.

     Prey composition, similar to those of Puget Sound rockfish, emphasized
pelagic organisms.  Calanoid copepods accounted for the highest proportion
(34.4 percent) of the total IRI while mysids (20.1 percent, Neomysis
 awatchensis)*, fishes (17.5 percent including Pacific sand lance),  crab
larvae (10.6 percent), chaetognaths (8.4 percent), hyperiid amphipods (2.1
percent), and fish larvae (1.2 percent) composed the other food items.
Gammarid amphipods (including Pontogeneia sp.,  and Eusiroides sp.,  Atylus sp. ,
Ischyrocerus sp., and Photis californicus)  were not very important.


Black Rockfish

     Adult black rockfish constituted over 15 percent of the total  fish
enumerated along the SCUBA transects in northern Puget Sound.  Prey organisms
were predominantly pelagic organisms.   Hyperiid amphipods (79.9 percent of
total IRI) were most important, followed by fishes (13.9 percent;  including
Pacific sand lance and tadpole sculpins),  crab larvae (1.1 percent),  and crabs
(1.0 percent; Cancer sp.).  Incidental items such as  rocks accounted for 1.7
percent of the total IRI.  Gammarid amphipods (predominantly Eusiroides sp.,
Pontogeneia spp., and Atylus sp., but also Ischyrocerus sp., Hyale  sp.,
Paraphoxus spinosa (?), Photis californica, Photis sp.,  Amphithoe  lacertosa,
and Amphithoe sp.) were not significant in the total IRI.

     Juvenile black rockfish from shallow sublittoral gravel habitats at
Deadman Bay and Guemes Island had an entirely different diet composition which
emphasized epibenthic prey such as shrimp,  harpacticoid copepods,  and gammarid
amphipods.                                                            6

     *
      Holmquist (1973) suggests that N. awatchensis should be considered as N.
mercedis m the Northeast Pacific.  We have, however,  left it as N.   awatchenlis
to minimize confusion.                                          —   	
                                    131

-------
Scalyhead Sculpin

     Scalyhead sculpins were the second most common cottid observed in the
rocky/kelp bed SCUBA observations in the vicinity of San Juan Island.  The
few specimens which were procured by slurp gun or spearing indicated a
diverse array of organisms—pelagic, epibenthic, and benthic—were included
in its diet.  Harpacticoid copepods were the most important prey.  Chaeto-
gnaths (pelagic arrow worms), calanoid copepods, crabs (Petrolisthes
eriomerus), crab larvae, mysids, shrimps, gammarid and hyperiid amphipods,
fishes, euphausiids, and caprellid amphipods were of secondary importance.

     Caprellid amphipods (Caprella mendax, £. laeviscula, C^. equilibra,
£. natalensis, £. incisa, ^. guacilior, C. ferrez, Metacaprella kennerlyi)
and tammarid amphipods  (Photis californica, P_. bifurcata, Parapleustes
pugettensis, Podocerus  sp., Erichtonius brasiliensis, Lembos sp.) and shrimp
(Heptacarpus sp.) were  listed as prey of scalyhead sculpins collected in
Nereocystis kelp beds on the outer coast of Vancouver Island (Leaman 1976).

Smoothhead Sculpin

     Smoothhead sculpins were not common in beach seine collections in either
northern Puget Sound or along the Strait of Juan de Fuca.  They appeared in
almost all the tidepool collections along the strait and were especially
common at Observatory Point and Slip Point.  Gammarid amphipods and hippolytid
shrimp together formed  84.0% of the total IRI.  Fish (including Pholis sp.),
sphaeromatid isopods (including Gnorimosphaeroma oregonensis, Exosphaeroma
amplicauda, and  Dynamenella sheareri), polychaete annelids, and pagurid
(hermit) crabs (including Pagurus beringanus) were of secondary importance
(Fig. B-34).

     Leaman's  (1976) collections of smoothhead sculpins in the Nereocystis
kelp beds  in Barkley Sound, outer Vancouver Island, indicated that amphipods
(caprellids, Caprella equilibra, C^. incisa and gammarids, Photis bifurcata,
P_.  californica) composed the highest proportion of the prey, followed by
shrimp  (Betaeus setosus, Lebbeus lagunae), crabs  (Pachycheles sp., Podocerus
sp.) and fish  (northern clingfish, longfin sculpin).

Rosylip  Sculpin

     While not found abundantly during  the DOE studies in northern Puget Sound,
rosylip  sculpins were ubiquitously distributed among the intertidal collection
sites along the Strait  of Juan de Fuca.   In that  region gammarid amphipods
 (67.8% of  total IRI) and sphaeromatid  isopods  (including Gnorimosphaeroma
oregonensis, Exosphaeroma amplicauda,  and Dynamenella sheareri;  20.8%) composed
the majority of the IRI prey spectrum  (Fig. B-35).  Idoteid  isopods  (including
Idotea wosnesenski), polychaete annelids, crustacean larvae, and mysids
 (including Archaeomysis grebnitzki) composed most of the remaining  important
prey organisms.
                                     132

-------
                           INDEX OF RELRTIVE IMPORTUNES (I.R.I.) OIRORflH
                           FROM FILE IDENT.  76-78. STRTION RLSIfi

                                         8831020403 - RRTEOIUS LflTERflLIS
                                                    SMOOIHHERO SCULPIN
                                           RDJUSTED SWLE  SIZE =  77
                                  20
                                          40
                                        CUMULATIVE FREQUENCY OF OCCURRENCE
PBEY ITF"
(••Au»ABIOE>
HTC°ni YTTDAF
«PHAFPO"I»TIOAE
TELECISTE1
PAOIRlOAE
NATANT1 A
r.AcTonpnnA
CPUST ACEA
POT AMOGFTONACF AE <'
P"1LIOir>AE
UNinFNTlFlED
k-ACPACTICOini
PA* OAL I naF
DH"ricPHYTA
FOEO
OCCUR
46.75
14.29
10.19
7.79
5.19
2. SO
?.*a
1.10
1.10
1.30
NUM.
11.11
	 3__03 	
2.53
2.53
	 U52_
1.01
1.5?
	 L.0_l_
1.01
3.01
	 L.01_
1.01
1.01
	 l.ni
1.01
COMP.
10.69
J?j91_
19I37
_S.70_
7.66
!o8
.0?
	 ^94
.04
.25
_LO_,80_
.25
.00
	 L.64
.02
PREY PERCENT
I.R.I. TOTAL IRI
_?_?5.1 	
193.1
2*3.?
	 68.0 	
66.?
36.4
A,n
2.8
4.0
	 5U 	
2.7
8.S
	 30.7 	
3.1
1.1
1.3
62.37
21.59
*.51
5.6"
1 .so
1.54
.85
-19
.07
.09
.1?
.06
.20
.7?
.08
• 03
.11
.03
              PPFY TAi*  *1TH FOFC.
                        ~
                          _FOF_O. irCi_io_._LE5<:_TMAN 5 ANn NUMF.OICAL J^iO GRAVIMETRIC
                          '" l'
-------
o
o
                            INDEX OF RELflTIVE IMPORTfiNCE U.R.I.) OIHCRBM
                            FROM FILE IOENT.  76-78. STflTION PLSTfl
                                         8831020501  RSCELICHTHYS RHOOORUS
                                                      ROSYLIP SCULP IN
                                            flDJUSTED SflMPLE SIZE = 276
                                  20     40
                                               60
                                                      80
                                                            100
                                                                   120
                                                                          140
                    PUEY  ITEM
                                         CUHULBTIVE FREQUENCY OF OCCURRENCE
                                        FREO   NUM.   GRAV.  PREY     PERCENT
                                        OCCUR  COMP.  COMP.  I.R.I.  TOTAL  IRI
                                                                                160
GAMMABIDEA
<;pHAFJ>OMATIDAE
lOOTEIOAE
POLYCHAETA
Mve;!OArFA
GAMMARIDAE
PAOIIOIDAE
OXRHYNCHA
UNIDENTIFIED '
HIPROl.YTIOAF.
CRUSTACEA
COTTlDAE

61.59
31.5?
9.4?
9.4?
«..BB
3.43
4.15
•>. an
2.54
7.54
1.81
1.45
1.45
1.09
?9.66
9.87
1.01
2.54
7.71
6.20
.37
!2S
.21
.17
4?. 37
.11
.08
14.76
17.33
17.76
10.07
1.91
5.21
2.89
1.53
1.7?
3.87
1.55
.15
5.23
2.47
1.29
1.31
857.7
176.9
U8.R
4-;. a
67.0
14.2
4.4
S.n
7.4
3.1
61.6
2.»
1.1
	 .5 —
66.32
20.78
4.29
2.88
l -l \
1.50
.34
.30
• 11
.1?
.18
.08
1.49
.19
.07
.03
.01
              PPFY TA«> >_m(_F-PF,_0._OCCMR.. LESS TH«N_5 _»N[> NUMERICAL ANO__GPAy I METRIC
              COMPOSITION HOTM LESS >MAN 1  AUE EXCLUDED FPO" THE TARLE AND PLOT
              (HUT MOT FROM CALCULATION OF  DIVERSITY INDICES!
                 PEPCENT OnulNSNCF INOE«
                 SHANNOM-ME^'ER DIVERSITY
 .?»
2.41
 .44
 .11
3.75
 .66
 .49
1.61
Fig.  B-35.   IRI prey spectrum of  rosylip sculpin in littoral habitats
                along Strait  of  Juan  de  Fuca.
                                               134

-------
     Caprellid (Caprella incisa,  £. equillbra) and gatnmarid amphipods
(Parapleustes pugettensis,  Advoides sp.) were the principal prey of rosylip
sculpins in the Nereocystis kelp bed habitat in Barkley Sound, outer Vancouver
Island (Leaman 1976).

Padded Sculpin

     Principal prey of padded sculpins from northern Puget Sound were
gammarid amphipods (89.1 percent of total IRI) and several other epibenthic
crustaceans—the flabelliferan isopod (Gnorimosphaeroma oregonense) (6.5
percent) and tanaids (3.0 percent).  In these collections from the eastern
shoreline (i.e., Legoe Bay) gammarid amphipods also were the most prevalent
prey  (88.7 percent of total IRI), followed by unidentified Caridean crustaceans
(shrimp; 5.7 percent), and sphaeromatid isopods (3.7 percent) (Fig. B-36).

      In the Strait of Juan de Fuca padded sculpin were common at Twin  Rivers
and Beckett Point, and were especially abundant in winter.  Epibenthic
crustaceans—gammarid (including Corophiidae) amphipods, hippolytid shrimp
(Heptacarpus kincaidi, H. tenuissimus). crangonid shrimp, sphaeromatid
isopods  (Gnorimosphaeroma oregonensis and Exosphaeroma amplicauda), and
idoteid isopods  (Synidotea sp. and Idotea wosnesenski)—were more abundant
than  benthic prey organisms such as polychaetes (Fig.B-37).

Silverspotted  Sculpin

      In northern Puget  Sound silverspotted sculpin commonly appeared in  the
pocket gravel  beach habitat  (Deadman Bay) beach seine collections  from July
through October.  The spectrum of prey  identified from these specimens was
oriented  toward  epibenthic crustaceans, specifically gammarid amphipods
 (40.1 percent  of total  IRI; including Amphithoe sp.). flabelliferan isopods
 (39.6 percent)  and shrimp  (9.6 percent; Heptacarpus stimpsoni) (Fig. B-38).

      Silverspotted sculpin specimens from cobble and gravel habitat sites at
Cherry Point,  Shannon Point, and Legoe Bay  appeared  to be even more specialized
in their  diet.   Gammarid amphipods were 32.7 percent of  the total  IRI, shrimp
7.2 percent.

      Of  the  sites sampled along  the Strait of Juan de Fuca, beach  seine
collections  at Twin Rivers, Morse Creek, and Jamestown generally provided the
most  silverspotted sculpin.  The prey spectrum (Fig. B-39) was almost  evenly
divided  between mysids  and gammarid amphipods.

Roughback Sculpin

      Roughback sculpin  were collected in the shallow sublittoral habitat only
in the Strait  of Juan de Fuca, in winter beach seine collections at Beckett
Point.  Shrimp,  including hippolytids such as Heptacarpus  tenuissimus,
crangonids such as Sclerocrangon alata  and Crangon sp.,  and unidentified
pandalids,  composed 83.5% of the total  IRI.  Gammarid amphipods and polychaete
 annelids  were  of minor  importance  (Fig. B-40).
                                      135

-------
                      g
                      £
                     I
                     §
                     CO
                     o
                     fe
                     8
                             INDEX OF RELRTIVE  inPORTRNCE (I.R.I.) OIRGRRM
                             FROM FILE IDENT. UW  BS. STRTION RLL

                                       PREOflTOR  8831020401 - flRTEOIUS FENESTRRLIS
                                    (PRDOEO SCULP IN     )   HDJUSTED SRMPLE SIZE = 15
                          100
                80
                           60
                           40
                20
                     °    20
                           40
                SO
                           80
                          100
                                  20
                                        40
                                             60
                                                   80
                                                         100
                                                               120
                                                                     140
                                                                           160    180
                                                                                      200
                                           CUMULHTIVE FREQUENCY OF OCCURRENCE
PREY
ITEM
FREO
OCCUR
NUM.
COMP.
GRAV.
COMP.
PREY
I.R.I.
PERCENT
TOTAL IRI
 GAMMAP.IDEA	SO.OjO	47.92
 "SPHAEROMATID»E                  33.00    2.92
 PLEOCYEMATA-CARIDEA             20.00    1.67
 _HARPACTICOJ_OA	      	13.00_   3.33
 TAK'AIDACEA         .  -  -     -   n.oo   " .83
 POLYCHAETA                       7.00     .42
 EDCARIDA-DECAPQOA-BPACHYRHYNCH
 BIVALVIA     "   "                7.
 PROSOBRANCHIA                    7.
i-RRACHYRHYNCH	7.00	.42	
                  00
                  00
42
83
42
_LSO£ODA	  	7 . 0_0_	 1 .25__
12.14
3.23
13.69
.13
.16
.03
.33
.26
.33
1.32
4804.8
202.9
307.?
45.0
12.Q
3.2
5.J3
7.6
5.3
18. Q.
                                                                          PP. 7Q
                                                                            .70
                                                                            .70
                                                                            .80
                                                                            .20
                                                                            .10
                                                                            .10
                                                                            .10
                                                                            .10
                                                                            OO
              PREY TAXA WITH FREO.  OCCUR. LESS THAN 5 AND NUMERICAL AND GRAVIMETRIC
             _COMPOSITION ROTH  LESS  THAN  1 ARF EXCLUDED FoQM THE TABLE_ANO PLOT ___
              (BUT NOT FROM CALCULATION OF DIVERSITY INDICES)
                 PERCENT DOMINANCE  INDEX_
                 "SHANNON-WFINEP  DIVERSITY
                 EVENNESS INDEX
                                          .23     .03
                                         1.21""  1.11
                                          .37     .33
                                                .79
                                                .70
                                                .21
Fig.  B-36.   IRI prey spectrum of padded  sculpin in  shallow sublittoral  habitats
               along  eastern shoreline  of northern Puget  Sound.
                                             136

-------
                     o
                     0-
                     §
                         100
                          80
                          60
                          40
20
                            INDEX OF RELFtTIVE ItlPORTRNCE (I.R.I.) OlflGRRM
                            FROM FILE IDENT.  76-78. STRTION flLSTfl

                                           8831020401 - flRTEDIUS FENESTRflLIS
                                                         PflDOED SCULPIN
                                              flOJUSTEO SflMPLE  SIZE = 143
                     2    20
                          40
                     to
                     o
                     fe
                     o
                     o
60
                          80
                         100
                                                                          oj nj eu -o m a)
                                                                          m 01 o IH n , «
                                                                        SE  IH S H
                                  20
                                          40
                                                 60
                                                         80
                                                                100
                                                                       120
                                                                               140
                                                                                      160
                                          CUMULATIVE FREQUENCY OF OCCURRENCE
                                          FPEO   NUM.   GPAV.   PPEY     PERCENT
                                          OCCKR  COMP.  COMP.   I.P.I.  TOTAL  IRI
p^^A
f, AMMAPJOAF
ii'HOFKJTjritn
>> ftTAK'TI A
I OP TF I n AE
MYSinACEA
TAKiAjnarEA
CANCpTnAE
PArnoIDAE
_F IIC An I nA-nFCAonoA^-QPArpYPHYNC
CriPOPHi PJAE
CTITHAF IPAF
IB.lfl
IS. 39
11. R9
4.79
5.S9

"..'0
?.90
'».... ?.io .
'.10
.70
vl3
5.7S
1 .09
3.17
1 .79
.40
.99
.40
1.19
.in
S.06 ]
16.53
2.23
10.62
1 ft 1
l.'lS
.1?
L5.23 	
3.96
4.10
.09
6.79
ms.fi 43
463. fl 15
315.4 10
.11B.3 10
100.' 3
143. n 4
1C -,

B.O
.59.0 1
12.?
9.4
9.4
3.S
1.9
4.R
1.73
.19
.33
.47
.09
.71

.72

.96 	
.40
• 31
.31
.1?
.06
.16
            POFY Ttxi *TTH FOEO.  iccnp. LESS THAN •; ANO NUMEPICAI  ANn GPAUIMFTOIT
         _.CnMP«|T.ins POTH LE,,  TH.N  , APE E.CLi.Otn FPOM THE TAR*E ANn ^LOT
            (RuT NOT FPOM CA|CULATION OF nivEOSITY INDICES)""       	    	
                        ifjnF»
                              CIVEOSITY~
3.37
 .66
                                  ^
                               3.51
                                .69
    -
?.64
 .51
Fig.  B-37.   IRI  Prey spectrum of padded sculpin  in shallow  sublittoral
               habitats along  Strait  of Juan  de  Fuca.
                                              137

-------
                           INDEX OF RELRTIVE IttPORTRNCE (I.R.I.) DIHGRRM

                           FROM FILE IDENT. N PGSD. STRTION RLSTfl

                                      PREDRTOR  8831020602  BLEPSIRS CIRRHOSUS
                                  (SILVERSPOTTED SCULP )   RDJUSTED SRMPLE SIZE =  20
UJ
(_>

Q

1
d
                        100
                         80
                         60
                         40
                         20
                    2    20
                    CD
                         40
                    |    60
                    o
                    o

                    ,-i    so
                        100
                                                                               u C 0.
                                                                               U) fll (0
                                                                               OHO
                                    20
                                             40
                                                       60
                                                                 80
                                                                          100
                                                                                    120
                                         CUflULRTIVE FREQUENCY OF OCCURRENCE
PPEY I
Fl.BFttlFE*.—
CAWMAR IDEA
PLFOCYEWATA-CAO
pr U&CARIDA-SPEL

OSTRACODA
TAK'AIDACEA 	
CAPPELLIDFA
...... - FPEQ —
TE" OCCUR
.._ 	 lc.>no
35.00
'IOE4 	 --- 15.00 —
AFor,pAoHACEA l^.no
=•.00
5.00
	 .... _ _ . S.OO
5.00
NUM.
COMP.
43 ?4
40.54
4.05
7.43
"? • 03
1.35
.68
.68
GRAV.
COMD.
8.44
P2.88
?5.30
4P.45
.57"
.15
.00
.19
PPEY
I.R.I.
1BOB fl
??19.7
440.?
748.?
13. n
7.5
3.4
4.3
PFRCENT
TOTAL IRI
34.50
4?. 30
8.40
14.30
• ?0
.10
.10
.10
             pppv  TA«A  WITH FDEO. nccu". LE<;<; THAN 5 ANO NUMERICAL  ANO
             CluPt^ITlnN  «nTM-cE<;s THAN I ARE EXCLUDED TPOM fHF.  TABLE ANO PLOT
             OUT  MOT FROM CALCULATION OF DIVERSITY INDICES)
                PERCENT  DOMlNflNCF INDEX
                <;HnMNO'J-WF INFP DIVERSITY"
                EVENNESS INOE«
                               .36
                              1 .«i
                               .60
 .30
1 .89
 .67
 .33
1.80
 .60
Fig.  B-38.   IRI prey spectrum  in silver-spotted sculpins  in shallow  sub-
               littoral waters of  northern Puget  Sound.
                                              138

-------
                     100
                        INDEX OF RELRTIVE IMPORTRNCE  (I.R.I.) DIRGRRM
                        FROM FILE IOENT.  76-78. STflTION RLSTR
                                        8831020602  - BLEPSIRS CIRRHOSUS
                                                   SILVERSPOTTED SCULP
                                          RDJUSTED SRMPLE SIZE = 132
                  DOEY ITEM
                                         CUMULflTIVE FREQUENCY OF OCCURRENCE
                                        FPEO   NU«.    &OAV.  POEY     PFPCENT
                                        OCCiJO  COMP.   COMP.  I.R.I.  TOTAL IRI
uycnACEA 64. 19
r.af'io inFA 	 	 _ 54.55 _
r.a>"'6«I riiF 25.76
1 nnTF IDAE 6.R2
»fcTA>:TLA. 	 -- 	 	 6.06--
HIPPOLVTlnAF 6.06
CP^N^.ONinAE .76
CYCLOPTES1DAE. 	 . . _. 	 . .76-
41.11
4Q.65
11.35
1 .00
- 1 .64
.90
.11
.05 _
36.76
28.21
12.81
8.02
2.46.
3.61
1.85
- 1.85
5144. 1
1756.4
622.4
61 .5
24.8
27.1
1.5
1.4
51.32
3H.93 	 	 	
6.45
.64
.26 __.
• 2»
.02
.01
             ooff T»«A 
-------
      §
      o
                          lOOr
                          80
                          60
                           40
                          20
                             INDEX OF RELATIVE IMPORTANCE (I.R.I.) DlfiORflM
                             FROM FILE IDENT.  76-78.  STRTION RLSTfl
                                           8831024001 - CHITONOTIS PUCETENSIS
                                                        ROUGHBHCK SCULPIN
                                              RDJUSTED SRMPLE SIZE =  39
                     §    -20
                           40
                     , U
                                                                   JH *H
                                                                   a w
                                                                   o o
                                                                   o a.
                                   20
                                          40
                                                  60
                                                         80
                                                                100
                                                                       120
                                                                               140
                                                                                      160
                    PPEY  ITEM
                                           CUMULflTlVE FREQUENCY OF OCCURRENCE
                                          FPEO
                                          nccim
                                  NUM.
                                  COMP.
                                                        COMP.
      POEY
      I.P.I.
 PERCENT
TOTAL IPI
           HIDPOLYTIDAE
           M4TANTJA
                  "
           TAKA10ACEA
           MALACOSTPACA  PHYLLOCAPIDA
             .
           HAPPACTICOIDA ______
35.^0
?3.ne
17.QS
i5;. is
1^.13
7.«,q
7.f>9
5.13
^.13
S.13
=-.13
«.13
?.<;6
2.56
15.07
in.?7
5.'.S
8.2?
?3. . 14
1.23
9.13
.43
4.0<(
fc.19
.00
.05
5.62
.32
.00
.00
1600.0
?66.5
890.6
1<^.<>
509.2
19.1
5?. 14
28.5
7.0
7.1
35.9
8.7
3H.6
24.6
                                                          44.01
                                                           7.33
                                                          24.50
                                                           4.00
                                                          14.01
                                                            .53
                                                           1 .44
                                                         ..  .78
                                                            .19
                                                            .?0
                                                         ._  .90
                                                            .24
                                                           1 .06
                                                            .68
             PPFY TAXA dlTH  FPEO. ICCllR. LFS^ THAN 5 AND NUMERICAL  ANP  GPAVIMETPIC
            _£OuonsITION  BOTH  LE5« THAN  I AHF E»CLIlO£0 FROM THE TAHLF  Awn  PLOT
             (P'JT MOT  FPOM CALCULATION OF DIVERSITY INDICES)
                PERCENT  DOMINANCE  INDEX
                SHANNO'J-WF INFO DIVERSITY
                EVENNESS INDE»
                                    .13
                                   3.21
 .30
2.20
 .56
    .28
   2.31
    .59
Fig. B-40.
IRI prey  spectrum of  roughback sculpin in shallow sublittoral
waters  of  the  Strait  of Juan  de Fuca.
                                              140

-------
Sharpnose Sculpin

     Sharpnose sculpin were found  to be common members of the tidepool fish
assemblages along the Strait  of  Juan de Fuca,  especially at Slip Point, Morse
Creek, and North Beach, but only at  Point Williams were they collected by
the beach seine.  Gammarid amphipods and sphaeromatid isopods (Gnorimosphaeroma
oregonensis, Exosphaeroma amplicauda, and Dynamenella sheareri) made up
94% of the total IRI  (Fig. B-41).

Calico Sculpin

      The  stomachs of  several  calico   sculpins collected at Fidalgo Island
 (Anacortes region)  contained  23  gammarid amphipods and nine harpacticoid
 copepods.

      Along  the  Strait of  Juan de Fuca, however, calico sculpin  appeared often
 in tidepool  collections,  mostly  at Observatory Point and Slip Point.   As with
 the sharpnose sculpin,  gammarid  amphipods and sphaeromatid isopods made up
 the majority of the IRI prey  spectrum for the calico sculpin: however,
 barnacles (principally cirri) were also a numerous (40% of total prey abundance)
 component in the diet (Fig.  B-42).

 Mosshead Sculpin

      Mosshead sculpins were not included in the collections  in  northern Puget
 Sound, due, perhaps, to the lack of  sampling in rocky  littoral  habitats as
 they were often included in tidepool collections along the Strait  of Juan de
 Fuca.  They were particularly abundant at Slip Point and Observatory Point.
 The  diet was more diverse and quite  different from the other two Clinocottus
 species.  Barnacles were the predominant prey organism while gammarid amphipods
 and  sphaeromatid isopods did not contribute significantly  to the diet (Fig.  B-43).
 Algae  (including Urospora mirabilis, Porphyra sp. , and Iridaea sp.)  composed
  38.7% of the total IRI, followed by  harpacticoid copepods,  sabellid  annelids,
 nemertean worms, and ostracods.

  Buffalo Sculpin

       Buffalo sculpins  caught in northern Puget Sound  had in  their  stomachs
  numerous pieces of algae  (ulvoid  type),  constituting  61.8  percent  of the total
  IRI, accompanied by two  amphipods  (25.0  percent),  and  one  partly digested fish
  (13.2 percent).  The sample  size  was too small, however,  to  determine whether
  the consumption of algae is  representative  of  the  food habits.   Buffalo
  sculpins originating from the Cherry Point  region  also indicated a high
  contribution by algae.  Nonalgae  prey taxa  included  gammarid amphipods
   crab! ?sTf    ^tal ??' ±nSeCtS (11'2  Pe««*),  polychaetes (9.2 percent),
   and orlanic dehrf  O^   3nC^ (4'7 Perce*t), pychnogonids  (4.1 percent),  sticks
   and organic debris  (2.5 percent), and flabelliferan  isopods  (2.4 percent'
   primarily  Exosphaeroma amplicauda).                              percent,

        Among the beach  seine  collection^ alnr,™ «-K~  o_
   juvenile  buffalo  sculpins were most common^ Twin *£"»  *Tl *t ^
   Tidepool  collections  at Observatory Point and N^rtb^I  ^ *?     k6" P°int'
   specimens.  Gammarid  amphipods, algae (IncSint JV        ° pr°vided a few
   Ulva fenestrata,  Porphyra sp.,    8
   annelids were identified  as the
                                        141

-------
           m
           z
           o
           I—
           CO
           o
           o_
           c
           o
           o
                  INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) DIRGRfiM
                  FROM FILE IOENT.  76-78. STflTION fiLSTfi
                                8831020701
               lOOr
                80
                60
                40
                20
           "    20
                           40
                           60
                80
               100
         P3EY  ITFM
                                                       CLINOCOTTUS flCUTICEPS
                                                         SHPRPNOSE SCULPIN
                                               flOJUSTED SfltlPLE SIZE   124
                         20
                                  40
                                          60
                                                  80
                                                          100
                                                                  120
                                                                           140
                                CUMULRTIVE FREQUENCY OF OCCURRENCE
                               ncciip
                                      NUM.
                                      COMP.
                                             COMP.
                                                               PPEY
                                                               1 .0.1
                     PFRCENT
                    TOTAL IRI
                               15.65  3?.P6  33.^9   36"?.
OIPTFBA
HAB3ACTICOIOA
1DCTF.1QAE ..
UNIOFNT IFIEH
MY«:IDACFA_
                                           10.4*)
                                 .81
17.14
 3.76
 9.6?
                                      -1.1?
                                       1.51
-17.87. .
   .94
   .23
.  1.35
  3.11
   .56
                                               .01
                                       3.87  ?7.86
               53.0
              103.3
               12.9
               92.3
                7.9
              ._ 3.7.
                1.?
               33.7
                                                                         67.08
                                                                         21- IS .
                                                                           .96
                                                                          1.8*)
                                                              1
                                                                            6*)
                                                                            14
                                                                            07.
                                                                            02
                                                                            61
  PPFY TAXA WITH  FOFO. OCCllW. LESS THAN 5 AND NUMERICAL  AND GPAvIMETPIT
  COMPOSITION  <>OTH  LESS THAN 1 APE EXCLUDER FPQM THE  TABLE AN1 PLOT
  (RUT MOT  FRO«.CALCULATION OF OIVESSITY INOITES)_	
     PEPCFNT  DOMINANCE  INOEX
     SHANNOM-KF1NKP DIVFPS1TY
     EVEMNtSS INDEX
 .)»
?.9?
    "
                                                           .29
                                                         2.14
                                                           .49
                         .52
                         1.27
                         .29
Fig.  B-41.
   IRI prey  spectrum of  sharpnose sculpin in shallow sublittoral
   and littoral  habitats along  the  Strait of Juan de Fuca.
                                  142

-------
                         100
                         80
                         60
                         40
                         20
                         20
                         40
                         60
                         80
                         100
                            INDEX OF RELRTIVE IMPORTRNCE (I.R.I.) DlflORflM
                            FROM FILE IDENT.  76-78. STRTION fllSTfl
                                           8831020702 - CLINOCOTTUS EflBRYUM
                                                         CRLICO SCULP IN
                                             ROJUSTEO SRMPLE SIZE -  20
                                   u  tj « «
                                   ffl  O *J 14
                                   u  m u v
                                20
                                      40
                                            60
                                                  80
                                                        100    120    140    160    180   200
                   PSJFY ITFM
CUnULRTIVE FREQUENCY OF OCCURRENCE
FPEO   NUM.    GPAV.   PPFY     PF9CENT
OrCuo  CO"".   COMP.   I.R.I.   TOTAL  I&I
cpHAFOOVftTlOAF
Ml)' >l!DAF
wACPACTICr-IOA
JUPTEPl 	 . . _
7o.no
5n.oo
3n.io
in.no
IP. 00
s.on
s. no
5. no
26.19
40.48
1.79
— . 1.19_
4.7*.
1.79
1.7.9
66.52
21 .54
4. S3
.54
.05
.97
.. ...91.
•>419.8
?1 18.7
1350.3
23.?
	 50.9 _
24.1
13. •»
i3.a__
IS4.29
20.99
13.39
.23
.__._. 50 	
.24
.14
.14
            POFY T».A bITH FPfO.  OCCltP.  LFSS  THAN  5  AND NUMERICAL ANO GOAVIMETPIC
           _rriMPOs!TION BOTH (.ESS TM»N 1  AHF  E«CLlinEO Fop" THE TABLE ANO PLOT
             (PijT NOT TRIM CALCULATION Or  OIvEPSlTY INDICES)                 	
               PFOCFNT
                         INEO Dl"vER«SITY
                                                  ?.19
               ^
               1.50
                .45
                                                                         _
                                                                         1.39
                                                                          • 4?
Fig.  B-42.   IRI prey spectrum  of calico sculpin in littoral  habitats along
               Strait  of  Juan  de  Fuca.
                                             143

-------
                      s
                      J—
                      8
                      5
                      5
                           tOOr
                              INDEX OF RELRTIVE IMPORTRNCE (I.R.I.) DIflGRRM
                              FROM FILE IDENT.  76-78, STRTION RLSTR
                                            8831020703 - CLINOCOTTUS GLOBICEPS
                                                         MOSSHEflD SCULPIN
                                               RDJUSTED SflMPLE  SIZE =  77
                                                                                      160
                    P=!EY ITEM
 CUMULflTIVE FREQUENCY OF OCCURRENCE
•"pfo   NUM".   'GPAV." "PREY     PERCENT
OCCUR  COHP.   COMP.  I.R.I.  TOTAL IRI
CIBPIPfDIA
..UJLaTRlCHALES 	
UNIDENTIFIED
ME«EPTFA
-GAMMApiDEA 	 	
pMnrjnRHYTA
H»OPACTICOIOA
CHLOOOPHyT*
«ARFLLIOAE
rPHAEPO"ATIOAF
.C.STOAC.onA . 	
ACRO^lPrtrtNlACFAE
POI VCMA<"T«
PAVGIACEAE
GACTRDPODA
C-ir:AOTlN4C E»
31.17
.23.J8
10. 4«
1 1 .ft9
10.39
in.. 19
9.09
9.09
S.19
S.19
, -3.90
?.f>0
?.«)0
?.ftO
?.isn
?.«,o
fcl .89
_.6.22
4.3?
1.35
1.22.
?.70
9.71
1.76
11.7ft
.54
10.54
1.76
.?7
.?7
1.08
3.78
3.68
30.04
?3.??
?.8S
.78
4.75
.26
13.63
1.95
.?4
1 .9«;
1.35
1.7?
3.00
.68
8.54
1420.1
847. 6 	
*36.7
49.1
20.7 	
77.4
90. »
139.9 _
71.?
4.1
48.7
8.1
5.?
8.5
4.6
3?.0
4?. 17
.25.17 __
15.93
1.46
-. .62. _
?.30
2.70
. 4.15 _
?.ll
.1?
l.*5 . _
.24
* 15
.25
.14
• 95
             PREY TAXA WITH FPEO.  OCCUR. LE^S THAN 5 AND NUMERICAL  AND
             COMPOSITION ROTH LESS THAN  1 AHF EXCLUDED FPOM THE  TARLE AND PLOT
           	(J?_MT_ NOT FRO" CA| CULATION OF DIVERSITY INDICFSI          _
                PTOCENT
        2.9?	3.05	
         .67    .69
                                                                           .27
                                                                           •'•O.
                                                                           .55
Fig.  B-43.   IRI  prey^spectrun, of  mosshead sculpin in littoral habitats along
               the  Strait of Juan de Fuca.                                              &
                                              144

-------
 spectrum  (Fig. B-44) .  The high  incidence  (31.5% of  total  IRI)  of  algae  a
 suggests  that they may constitute more  than an  incidentally  consumed  rou

     Adult buffalo sculpin collected  in a  gravel-cobble  habitat in Nisqually
 Reach, southern Puget Sound  (Fresh  et al.  1979), had  consumed algae (also
 principally ulvoid types) almost exclusively, such  that 93% of the total IKi
 was algae.  Most of  the prey animals  were  gammarid amphipods and polychaete
 annelids.  Johnson's  (1968)  in-depth  analysis of the food  habits of buffalo
 sculpin in Humboldt  Bay,  California,  indicated  that  caprellid (Caprella sp.)
 and gammarid amphipods (inc.  Amphithoe  sp.) were the predominant food
 organisms, composing  66.3% of the total IRI  spectrum (Fig. B-45) .  Again,
 algae (Ulva lobata) were  a major component (16.4%  of total IRI) in the diet.

 Red Irish Lord

     Red Irish lord were  characteristic of the  demersal fish assemblage in
 the gravel-cobble pocket  beach habitats in northern  Puget Sound and were often
 found in the rocky/kelp bed  habitat.  They appeared  to be an almost completely
bottom-oriented carnivore, preying  on flabelliferan  isopods (42.8 percent of
 total IRI), brachyuran crabs (39.9  percent;  Cancer magister, C^. oregonensis.
 C. productus. Pugettia gracilis, Mimulus sp.),  fish  (13.4 percent), and
 shrimp (1.3 percent).

     Although they were not  frequently  collected  along the Strait of Juan
 de Fuca during the MESA studies, Leaman (1976)  identified them as common
 members of the Nereocystis kelp  bed assemblage  in  Barkley Sound, outer
 Vancouver Island, his documentation of  their  prey  organisms indicated
 approximately equal  gravimetric contributions  by crabs (Pachycheles rudis,
 Loxorhynchus crispatus, Cancer oregonensis) .  gammarid amphipods (Erichtonius
 brasiliensis) and caprellid  amphipods (Caprella equilibra. C. incisa  C~!
 natalensis, C^. laeviscula, jC . mendax , £. ferrea) .          ~        ' ~

 Longfin  Sculpin

      The longfin sculpin  was the most frequently observed  cottid and  third
 most common  species  over  the combined SCUBA trains*^ ~u               cnira
 frequency of  occurrence)  in  the  rocky/kelp bed  haMtat T^10"8 ^ PerCent
 Sound.   The  prey spectrum was very diverse  with Tn   J    *     er° Puget
 included in  the  overall  sample.  Harpacticoid COD    J   °f  rare Prey ite*»s
 prey, contributing  55.4  percent  of  the  total  TUT     i  WSre  the  most  lmP°rtant
 crabs (8.7 percent), gammarid amphipods (5 7  J,   po|ychaetes  (23.9 percent),
 and crab larvae, (1.5 percent) were of  second*™-    ' shrimP  d-5 percent),
 benthic organisms were  taken more frequently  hi ^?°rtance-  Although epi-
 greatest contribution to  the total  prey biomass        organisms made  the

      Longfin sculpins also appear tn
                                 r  n  o
 habitats along the Strait of Juan  de F^^ef fT^" °f the ro<*y/kelp bed
 data) and m the Nereocystis kelp  bed habitat  i     ' ^±V' Wash- > unpub
 Vancouver Island (Leaman 1976).  In the 1»^     ^ the Outer coast of
 (Parapleustes pugettensis. Photis  calif ornlrf Z*K±°*' 8ammarid
 appeared to be their most ^^u^l~~?-> ^ichtoniu
 crustaceans and polychaete annelids       SuPPlemented by
                                      145

-------
                            100
                              INDEX OF RELflTIVE IMPORTANCE (I.R.I.I OIBGRflM
                              FROM FILE IDENT.  76-78. STPTION RLSTR
                                             8831021001 - ENOPHRYS BISON
                                                       BUFFflLO SCULP IN
                                              flDJUSTED SHMPLE SIZE =  116
                        S   20
                            100
                                   20
                                         40
                                               60
                                                    80    100   120   140    160
CUMULRTIVE FREQUENCY
PPFY 1TFM
POI YCHAFTA
IINIOFNTIFIEO
III UATFAF
TANAIOACFA
GAMMAPinAE
HIPPOLYTIPAF.
PHnnoPHYTA
BIVALVIA
CHLOROPHYTA
CANCRIOAE
OSTEICHTHYS
PACURIHAE
ATFLECYCLIDAE
COTTID4E
CALLIANASSlOAF
HA»!NOEIOAE
CLIIPEIOAE
PHYI.LOOnCIOAE
FPFO
OCCIIP
4,. .18
15.52
10.14
10.34
8.62
6.90
6.03
5.17
4.11
4.11
3.45
1.7?
1.7?
1.72
.BIS
."6
NUM.
COMP.
G"AV.
COMP.
25.06 1.11
~ 6!27 2.17
S.90 7.01
	 I.** 	 35.26.
6.87 .40
3.98 .03
	 Z.65 	 U.1%.
2.41 .21
.84 .88
1.45 .92
1.20
4.34
.36
6.27
.60
.84
.60
.1?
.12
.47
9.50
.67
1.01
1.J7
5.60
3.33
2.33
1.S2
3.27
OF OCCURRENCE
PPF.Y
I.R.I.
	 41S.5 .
131.0
	 S0?.8_
75.?
34.5
	 9_S.J_
15.8
10.?
7.?
47.7
9.1
17.9
2.B
10.1
3.0
1.5
2.9
PERCENT
TOTAL IRI
40.25
15.54
4.86
7.03
18.79
2.79
1.28
3.54
.59
.33
.38
.27
1.77
.34
.67
.10
.13
.37
.11
.09
• 06
.11
               _PPE.T_IAXA KITMJPEJ. OCCUg._J.ESS_tHAN 5 _ANO_ NUMERICAL
                COMPOSITION POTH LESS THAN 1 APE FxCLIIHEO FPOM THE TAHLE  ANO PLOT
                (BUT NOT FROM CALCULATION OF DIVERSITY
                   PERCENT OOMINANC";  [HDF»
                   SHANNON-WE1NEP 01VEPSITY
  .11
 3.85
_.71
.19
.3fr
.62
   .23
  ?.73
	.50
Fig. B-44.    IRI  prey spectrum of  buffalo  sculpin in shallow  sublittoral
                habitats along Strait of Juan de  Fuca.
                                              146

-------
                     INDEX OF RELRTIVE IMPORTHNCE (I.R.I.) OIRORRM
                     FROM FILE IOENT. JOHNSN. STHT10N  1968
                                 PREORTOR 8831021001  - ENOPHRYS BISON
                            (BUFFRLO SCULPIN    )  ROJUSTED SHMPLE SIZE
                   100 r

•
1

~4
2
TJ O.
« a
• — a
EJ 5 1
SiSfl
• &•
",S|
"M
o 5
sld
                     100
                                           Too        iso        200
                                    CUI1ULHTIVE FREQUENCY OF OCCURRENCE
                                     FPEO   NUM.   G««V.  POET    PERCENT
                                     nrdlP  COMP.  CQMP.  I.P.I.  TOTtL  1P1

ROCK
Gau">Rinf a
• AMFHITHOE SP.
NASSARIUS SP.
OHFl T* SP. 	
	 EULALIA AVICULISETA
M4CO"* SP.
II^TOFNT IF 1EO
	 ELAlMOfUS St.
10CTFA SP.
rp»i«ir,OM SP.
— pr>L»e*»E"i4
FLtBFLLIfERA
CLYCERA SP.
— NECII<; SP.
	 (TiMPn^TmKr>
LESS THAN 5 AKIO NUMERICAL Akin G°l
*RE F.ltLliOED TooM THE TABLE »nn
nivERSITT INOirfsi
.P2 .16
2.40 3.14
.65 .8?
33.40
10.30
24.10
16.40
3.80
.80
.30
.20
.40
.00
.30
.50
.10
.10
.00
.20
WIMET9IC
PLOT
.2?
2-51
.64
Fig.  B-45.
IRI  prey spectrum of  buffalo  sculpin  generated from  data
included in  Johnson's (1968)  analysis of  their food  habits
in Humboldt  Bay,  California.
                                            147

-------
Pacific Staghorn Sculpin

     Staghorn sculpins were probably the most ubiquitous cottid in the shallow
sublittoral region of northern Puget Sound.  Considering the sample size from
this region, the overall prey spectrum (Fig. B-46) is not very diverse.
Emphasis is on benthic organisms, with flabelliferan isopods (32.2 percent
of total IRI), and bivalve siphons (29.6 percent) being equally important;
polychaetes follow with 11.5 percent.  Crabs account for 8.9 percent of the
total IRI; fish, 51.7 percent: crab larvae, 4.3 percent; tanaids, 3.3 percent:
gammarid amphipods (including Atylus sp., Allorchestes sp., Paraphoxus
spinosa, and Euhaustorius sp., Paraphoxus spinosa, and Euhaustorius sp.), 1.3
percent; and bivalves, 0.5 percent.  Although not as frequently preyed upon,
fish (including juveniles and larvae of Pacific herring and juvenile striped
seaperch, Embiotoca lateralis) and oxyrhynchan crabs actually composed the
majority of the biomass ingested.  Included in the decapod and general
peracaridan crustacean categories were Crangon franciscorum, Idotea resecata,
and Cancer magister.

     Also, the most common nearshore demersal species collected along the
eastern shoreline, they occurred dominantly in collections at the Fidalgo Bay,
Drayton Harbor, and Padilla Bay mud/eelgrass sites and the Brich Bay sand/
eelgrass site.  Overall prey composition from the eastern sites is dramatically
different than those in the western area (Fig. B-47).  Gammarid amphipods
are much more important, providing 69.8 percent of the total IRI.  Other prey
taxa are rather equally represented by crabs (8.8 percent, Hemigrapsus
oregonensis, E. nudus, Cancer magister, Pinnixa sp., and Paguridae), shrimp
(8.2 percent; Crangonidae, Callianassidae, including Upogebia pugettensis,
and Penaeidae), isopods (5.7 percent, Exosphaeroma amplicauda, E_. media) ,
fish (3.2 percent; juvenile Staghorn sculpins, shiner perch, and rockfishes),
and polychaete annelids (3.2 percent).

     Along the Strait of Juan de Fuca, Pacific Staghorn sculpin were one of
the few nearshore demersal species which occurred commonly in the beach seine
collections at all sites; however, collections at Jamestown and Beckett Point
provided more specimens than the other sites.  The diverse prey spectrum
(Fig. B-48) included both benthic and epibenthic organisms and, unlike the
northern Puget Sound spectrum, was dominated by fish (46% of total IRx, including
buffalo sculpin, Enophrys bison, shiner perch, Cymatogaster aggregata, Pacific
sand lance, tube-snout; Aulorhynchus flavidus, juvenile salmon, Oncorhynchus sp.,
other Pacific staghorn sculpin, Leptocottus armatus, and unidentified flatfish,
Pleuronectidae), true shrimp  (17.5% of total IRI, including Heptacarpus taylori,
Pandalus danae, Crangon alaskensis, and C_. stylirostris) , mysids (11.7% of total
IRI, including Neomysis awatschensis), polychaete annelids, and crabs (6.2% of
total IRI, including Pugettia richi, Telmessus cheiragonus, Cancer magister,
and Hemigrapsis oregonensis).

     Jones (1962) extensive studies of Leptocottus populations in Tomales Bay,
California, included documentation of an overall diet oriented almost exclusively
toward benthic shrimp (Crangon sp. and Upogebia pugettensisa, 92.3% of total
IRI combined); fish (northern anchovy, Engraulis mordax and shore crabs
(Hemigrapsus oregonensis) were secondary prey organisms  (Fig. B-49).  Life
history studies of staghorn sculpins in Anaheim Bay (Tasto 1975) similarly
illustrated both the variety of food organisms consumed and the importance of
decapod crustaceans, specifically Pinnixa sp., Hemigrapsus oregonensis and
                                      148

-------
                         INDEX OF RELRTIVE  IMPORTANCE (I.R.I.) DIHGRflM

                         FROh FILE IDENT. N PGSD. STflTlON flLSTfl

                                     PREDHTOR  8831021801 - LEPTOCOTTUS flRMRTUS
                                 (PRC- STRGHORN SCULPN)  flOJUSTED SflHPLE SIZE =  51
  y
  g
  I
                   o

                   *—
                   o
                      100
                       80
                       60
                       40
                        20
                  2    20
                        40
                                                           AJ  O U
                                                           £ »w O
                                                           O -H 4J
                                                           •H  01 O
                                                           o)  C.-H
                                                           w  3 .fl
                                                           « T-I e
                                                           O U W
                                                                     S.-S
                                                                  01
                                                                  O
                                                      »-i U  > 4J
                                                      QJ 0) -H 10
                                                      PL. Q PQ O I
                                      50          100         150          200


                                          CUMULflTIVE FREQUENCY OF OCCURRENCE
                                                                        250
                                                  MUM;	GRAVi— v°Ef	PFPCENT
                                           OCCUR  COMP.  COMP.  I.R.I.  TOTAL  IRl
POLYCHAET6 	 	 — — .
f . AUCI 1 T F F Q A
r (_ B 7*r l_L 1 * t w *»
G AMM AO 1 OF A " " " — — -- —
PF CAPOOA — flOACHVllPft

p 1 V fi L v T ft - 	
TAKftlDACFA.
O^C&°OOft - 	 _
pt DAPARinfi — CDTi ArnrnAnL.A/-r>
»'c.H'*i,'«**iuiivti^r LAtOoRAP^flCEA
- DtC»DOOA-RRACMTI|0« 	 -- --
BIVALVI A
— O^TEICMTHYE^ 	 	
OXYRHYNCHA
CL'fPEA HAPENGM*^ DA[tA<:l
O^TF ICHTHvFc
TLIIOE IFOO"E<;-CLUPEnlDFI
FMPlnTOCA LiTFoALK
47.00
?4.00 3
??.oo
?'.no
?n.oo " ?
IB. oo
16.00
10.00
«.00~
i.no
— «. . o o — -
fi.no
?.oo
?."0
2. no
?.no
4. 06 --
13.4?
1.7? —
«.l?
3.5T
?.50
1.39 -
.69
?.«5 --
1 .64
.35~
.43
.09
.09
.09
.17
6.06
3.14
~ .19 —
3.12
6.8?
?.53
6.14
.53
.73
.14
14.31
18.??
4.06
1 .50
1.2?
?7.54
475.6^"
S77.4
41. P
315.9
«06";^"
90.5
120.3
12.2
29. f,
14.?
98.0
111.9
8. 1
3.?
2.6
S5.4
" 15.60 	
28.70
1 .40 	
1 0.30
?6 .40
3.00
3.90 " 	
.40
.90 	
.50
2.90 	 	
3.70
.30 	
.10
.10 	
1 .80
             PPFY 'All 
-------
                             INDEX OF RELRTIVE IMPORTANCE (I.R.I.) DIRGRRM
                             FROM FILE IDENT. HH  BS. STRTION  fiLL

                                        PREDfiTOR  8831021801   LEPTOCOTTUS flRtlflTUS
                                    CPfiC.  STRGHORN SCULPN)   fiDJUSTED SRHPLE SIZE  - 91
                           100
                      3
                           80
                           60
§
" 40
tn
o
a.
§ 20
\—
o
°~ o

t—
s
2 20
§ 40

^~"
I 60




hi 80
o
Q_


inn













•

ffl
01

(U
1

u







— 1 — I — I — I""1 n r*"*
	 ^ ^ — ~* — 1 1 [ I
^u u
, :
.
-o i-
-p- T3 3
5 S « 1


*J 4J ocj

                                  20    40    60    80    100   120    140    160    180   200

                                            CUMULflTIVE FREQUENCY OF OCCURRENCE
                      PPEY  ITEM
                                            FREO   NUM.
                                            OCCUR  COMP.
      r,RAV.   PPEY     PERCENT
      COMP.   I.P.I.  TOTAL IRI
            _GAMM4RJOE4_ 	 	
             POLVCHAETA
             PENAEIDAE
             GRAPSIOAE		
             ISOPODA "  "
             OSTEICHTHYES
             PLEOCYEMATA-CARIOEA  	 	
             FUCAPIDA-DECAPODA-BRACHYRHYNCH
             CALLIANASSIOAE
             SPHAERQMATIDAE_
             EMPIOTOCIOAE"
             COTTIDAE
             VALVITERA       	  _
             DECAPODA-8RACMYURA
             CANCRIDAE
4 ? . 0 0
2?.00
18.00
lfc.00
1A.OO
1P.OO
9. no
B.OO
6.00
•5.00
fc.no
4.00
3.00
?.oo
1.00
sn.n?
3.8ft
4.13
1.33
11. 9B
1.70
1.31
l.?0
.93
1 .ftn
.93
.53
1 .fiO
4.13
.13
1.97
4.24
7.10
18.00
.2«
5.57
3.38
3.78
1?.7?
.09
13.81
7.3?
.30
2.13
17.49
??37.7
178.?
?0?.l
?98.«,
175.S
87.?
42.4
39. p
S4.6
«.4
59.0
31.4
5.7
12. S
17.6
                      64.5n
                       5.10
                       5.8n
                       ».60
                       5.10
                       ?.SO
                      _l.?n
                       1.10
                       1.60
                         .20
                       1.70
                         .90
                      _  •?"_
                         .40
                         .50
               PRPY  TAX*  WITH FpEO. OCCUR. LE55 THAN'5  AND NUMEOICAL AND GRAVIMETRIC
               COMPOSITION ROTH LESS THAN 1 ARE EXCLUDED FPOM THE TABLE ANn PLOT
               (BUT  NOT FROM CALCULATION OF DIVERSITY INDICES)
                  PERCENT DOMINANCE INDEX
                  SHANNON-WEINER DIVERSITY
                  EVENNESS
 .?7    .1?
?.??   3.26
 .57 — .83
 .43
2.03
 .5?'
Fig.  B-47-   IRI  prey spectrum of staghorn sculpins  from  shallow  sublittoral
               habitats along  eastern  shoreline of  northern Puget Sound.
                                               150

-------
                              INDEX OF RELflTIVE IMPORTANCE (I.R.I.) OIHORHM
                              FROM FILE IDENT.  76-78. STfiTION HLSTfl
                                            8831021801   LEPTOCOTTUS flRflflTUS
                                                      PflC. STHOHORN SCULPN
                                               flOJUSTED SfWPLE  SIZE = 294
                            100
                                   20
                                        40
                                              60
                                                        100    120   140   160   180   ZOO
                                            CUnULflTIVE FREQUENCY OF OCCURRENCE 	
                                           FOEO   NU".   CP»V.  PPEY     PERCENT
                                           OCCIIH  COMP.  COMP.  I.R.I.  TOT»L  1RI
UHIOENT.FIED
CPAVGONIOAf.
Pni YCHAFTA
TFLEOSTFI
NATANTIA
H1DOOLYT IDAr
C»>roln«E
TAKA 1 DACF A
SP^AFROMATIHAF.
PA* OAL IOAE
FllCAOinA-OECAPnOA-flPACHYPMYNCH
OSTEICHTHYS
FIICAOIOA-nEClPOOA
COTTIDAF
ATFLECYTLIDAE
NFMATOOA
MYM1AE
CLUOEIOAE
20.41
20.07
15.65
11.61
11.9(1
10.20
9.86
0.50
7.«2
7.14
5.7C
5.7H
3.74
3.40
3.40
1.06
2.72
2.72
2.18
1.16
1.16
1.36
.34
2.33
_3.85_
1.24
9.06
.90
.74
.72
.49
.50
2.30
.S4
.29
.98
68.44
.!•*
.12
.10
.12
.n»
2j2l
1.76
.04
.16
.05
.01
4.46
.93
7.22
1.19
_2*«5
16.68
1.58
1.73
7.69
.03
.38
5.60
1.80
10.23
2.84
4.29
4.86
8.05
2.71
.05
2.63
1.04
1.30
1.12
152.4
97.6
169.7
160.4
207.3
23. S
69. A
18.?
34.0
16. 0
J94.4
10.3
15. n
15.?
22.?
7.6
5.4
3!6
?.7
1.8
10.73
6.87
11.95
11.29
1.21
14.60
1.6S
1.54
4.9Q
1.21
.46
2.40
1.13
20.73
.7?
1.06
J.07
1.57
.54
.38
.37
.26
.19
.13
• 03
                 PRFY TAIA HUM FPEQ. OCClH.  LESS THAN 5 AND NUMERICAL AND GRAVIMETRIC
                 COMPOSITION «OT« LESS THAN I  ARF EXCLUDED FOOM  THE TABLE ANO PLOT
                 j9aLNOJLEflO"__£A.LCULA-I-lOM_OF_alvEP.SlT»_lNOICESJ	
1MINANCF INOEH            .48
rI"FO OIVFDSITY	_2.|4
                                                          .07
                                                         4.4Q
 •11
j.63
Fig.  B-48.   IRI  prey spectrum of  staghorn  sculpin  in  shallow
                sublittoral habitat of Strait  of  Juan  de  Fuca.
                                               151

-------
                           INDEX OF RELflTIVE IMPORTflNCE (I.R.I.) DlflGRRM
                           FROM FILE IDENT.  JONES. STflTION  1962

                                      PREORTOR  8831021801   LEPTOCOTTUS RRMHTUS
                                  (PRC. STflGHORN SCULPN)   PDJUSTED SRMPLE SIZE
87
10U
1 80
CD
•z.
ID
GO
^ 60
00
•z.
o
E 40
Q_
S 20
\—
C_)
0
t—
§ 20
UJ
3
>-
CD
40
o
i—
£ 60
1
^ 30
<_>
Q_
100







Crangon sp .








Upogebia pugettensi:
) 50 100








,1,1 T"Dn
IP

,raulis mordax
nigrapsus oregonensis
dentif ied
eostei
lianassa longimana
cer magi s t er
jjiea harengus pallasi
cliis sp.
C Oi C Oi .'ANCE  INDEX
                              OI
-------
Callianassa sp. (Table B-5).   Fish, mostly.the arrow goby, Clevelandia ios,
also occurred frequently.

     Leptocottus from Everett Bay, central Puget Sound, examined by
Conley (1977) had consumed mostly Corophium sp. and other gammarid amphipods,
based on numbers, while other Leptdcbttus armatus,and mud shrimp, Callianassa
sp., composed the majority of the total prey biomass (Table B-6).

Great Sculpin

     Great sculpins were retained only in collections from northern Puget
Sound.  One juvenile great sculpin from a beach seine collection at Deadman
Bay had two unidentifiable decapods in its stomach.  Another from a beach
seine collection at Guemes Island contained 17 gammarid amphipods, 17 benthic
gastropods (Littorina scutulata) and a piece of alga.

Tidepool Sculpin

     Tidepool sculpin were numerically the predominant fish in the intertidal
collections along the Strait of Juan de Fuca.  They were especially abundant
at four sites—Slip Point, Observatory Point, Twin Rivers, and North Beach.
Gammarid amphipods and sphaeromatid isopod species (Gnorimo sphaeroma oregonensis
and Exosphaeroma amplicauda)  dominated the prey spectrum (Fig. B-50),
combining for 80.3% of the total IRI.  Harpacticoid copepods (8.2% of total
IRI), polychaete annelids (4.9%), and barnacles (4.3%) constituted the more
important secondary prey items.  Sphaeromatid isopods were quite important
at the sites in the eastern half of the Strait but were only fourth or fifth
in importance at the three western sites: there barnacles and harpacticoid
copepods or hermit crabs replaced sphaeromatid isopods.

     Nakamura's (1971) analysis of the food habits of tidepool sculpin at Port
Renfrew, on the west coast of Vancouver Island, B.C., showed harpacticoid
copepods and gammarid amphipods to numerically predominate; insects, polychaete
annelids and isopods were of secondary importance (Table B-7).

Saddleback Sculpin

     The saddleback sculpin was reported only from tidepool collections at
Slip Point and Observatory Point in the Strait of Juan de Fuca.  They had
fed primarily upon gammarid amphipods (Fig.  B-51); the second most important
prey for the saddleback sculpin was harpacticoid copepods.

Fluffy Sculpin

     Fluffy sculpins were common in collections from Slip Point, Observatory
Point, and Neah Bay along the Strait of Juan de Fuca.  Among the three
Oligocottus species, gammarid amphipods contributed more to the prey spectrum
of the fluffy sculpin; accordingly, sphaeromatid isopods were the least
important in the diet of this species (Fig.  B-52).  The three isopod species—
Gnorimosphaeroma oregonensis, Exosphaeroma amplicauda, and Dynamenella sheareri—
were equally represented.  Harpacticoid copepods, polychaetes, and idoteid
isopods were secondary food organisms.

                                      153

-------
Table  B-5.   Prey composition (frequency of occurence)  of Pacific staghorn
              sculpin  in Anaheim Bay,  California documented by Tasto  (1975).
    PREDATOR  8831021801 - LEPTOCOTTUS ARMATUS
                                                                 (PAC. STAGHORN SCULPN)
                  INDEX OF RELATIVE IMPORTANCE  (I.R.I.) TABLE
                    FROM FILE IDENT. REF 67. STATION SLSTA
               PREY ITEM
FREO   NUM.
OCCUR  COMP.
                                               GRAV.   PREY
                                               COMP.   I.R.I.
                                                                    PERCENT
                                                                   TOTAL IRI
 PINNIXA
 MEMIGRAPSUS OREGONENSIS
 CALLIANASSA SP.
 CLEVELAND IA IDS
 OECAPODA
 COROPHIUM ACHERUSICUM
 TELEOSTEI
 AMPHITHOE  SP.
 FUNDULUS SP.
 HYALE PLUMULOSA
 OECAPODA
 OIPTERA
 POLYCHAETA
 GAMMARIDEA
 OXYUROSTYLIS SMITHI
 CRANGON SP.
 EULALIA SP.
 HYALE SP.
"TAGELUS SP.             ""
 GONIADA SP.
 LEPTOCOTTUS ARMATUS
 ALLORCHESTES SP.
 CANCER ANTENNARIUS
 POTAMIDIDAE
 ATHERINOPS AFFINIS       "
 HYALE RUBRA
 CAPRELLA EOUILIBRA
 BIVALVIA
 PLEURONECTIDAE
 CLAUSIDAE      _
                                      ai.f.0
                                      20.70
                                      17.40
                                      10.30
                                       8.90
                                       6.10
                                       3.80
                                       ?.40
                                       1 .90
                                       1.40
                                       1.40
                                       1.40
                                        .90
                                        .90
                                        .90
                                        .90
                                        .90
                                        .90
                                        .90
                                        .90
                                        .50
                                        .50
                                        .50
                                        .50
                                        .50
                                        .50
                                        .50
                                        .50
                                        .50
                                        .50
        PREY TAXA WITH FREO. OCCl/R, LESS THAN 5 AND NUMERICAL AND GRAVIMETRIC
        COMPOSITION BOTH LESS THAN 1 ARE EXCLUDED FROM THE TABLE AND PLOT
        (BUT NOT FROM CALCULATION OF DIVERSITY INDICES)
PERCENT DOMINANCE INDEX
SHANNON-WEINER DIVERSITY
EVENNESS INDEX
         .00
         .00
         .00
                                                 .00
                                                 .00
                                                 .00
                                                                       .00
                                                                       .00
                                                                       .00
                                          154

-------
Table B-6.   Prey composition (numerical and gravimetric composition) of
             Pacific staghorn sculpin in Everett  Bay, Washington, documented
             by Conley  (1977).
                     -  L^OTOCOT TUS AP'«ATIJS
                                                          (PAC. STAGHODN
  PI.
  r-c
  TF I
            vnF> nF
             FD1:< FILE IOFNT
                                          (I.P.I.) TABLE
                                    f, -> ,  STATION SLSTA
FPEO NUM.
OOF_C IT^" OCCllP COMP.
V* .10
ii'/arp 5 .,„
• ?0
o.Mnn, C3. >40
i_4T fOr-iy ^ = T^LL^TIJS 10
j*~lr\nf~r- • i '
^TOArrM
.'CTOfOTTtiC^r-'i-r^ ' • f- /
I'JV-'llJtlKit,. HJ|j^ _-
e *t if o T i^c. A - *
L**^l)rfl i * *»
:,rnc^i 14>A°
•I I 'i«,l-^ ^D -10
-a^^-^D. tn
;^^^Q4 »nu
lc;r|c -1 J,;.. CD. *7"
r T f-'po ^ Tp-. , ,- A/-J.I c *T.,^ 79.30
^'-i('-i>a(jl[_f.*Tlje; .
r Aoc.oi-pt: ar -,VI ,ca -- 	 	 *
•^OPVTF<; HrxapfFons *^°
T n MC CD. ' «?0
.10
C-RAV.
COMp.
.10
.80
.50
.10
.10
45.00
.40
.60
?3.00
.10
2.90
1.00
3.00
.80
.10
PPFY PEPCENT
I.O.I. TOTAL IPI












" "


= = FV TA>A  vITH KPFO. OCCUC. LES"; THAN  S  'NO K'HME°ICAL ANO
ro.-onsT TIT.  -^OTH L^S^ THAN 1  AP^ FxCLU.DED
("MT MOT FPOM C«l CPL4TIO-J OF  DI»'
                                                 THE TABLE AND PLOT
                    DIVERSITY
                                        1.08
                                               1 .
                                                .4?
                                                             .00
                                                             .00
                                                             .00
                                      155

-------
                              INDEX OF RELflTIVE IMPORTANCE (I.R.I.) DIflGRfln
                              FROM FILE IDENT.  76-78. STflTION flLSTfl

                                            8831022401   OLIGOCOI7US nflCULOSUS
                                                        TIOEPOOL SCULPIN
                                               flOJUSTED SflnPLE SIZE = 512
                            60
                            60
                            100
                                                     80
                                                           100
                                                                 120
                                                                       140
                                                                             160
                                                                                   160
                                            CUnULRTIVE FREQUENCY OF OCCURRENCE
PPFY ITF«
OAMUAPIQE&
H*oP4CTICOl()a
ciP»i°Ef>iA
pnt vCNAiiT*
OfPTFRA
^rpTEA^
IIMIOFNTIFIED
CRIKTACE',

COTTIDAE
FPfO
OCCUR
53.93
14.77
1 ft. 0?
11.67
7.11
s. o«
1.71
2.73
ll95
1.17
.98
.19
.19
.39
NUM.
CO*P.
77.07
_1J.9«!
P4.30
1.77
1.57
1.09
.54
.19
1.45
5.79
.05
.11
.05
GPAV.
COMP.
73.46
.61
1.38
14.7?
.26
.59
.37
.30
1.0"
3.00
7.00
2.00
1.74
3.84
1.35
1.76
1.57
PRFY
I.P.I. '
7674.7
1755.9
199.1
740.0
15.9
7.4
5.2
4.4
\f ."
8.0
11.7
6.1
1.5
.9
.7
PERCENT
fOTAL IP1
54.6?
75.65
8-15
4.3?
4.90
.3?
.24
.15
.11
.09
.17
.38
.18
.78
.11
.01
.0?
.01
.01
                PPRY T«xA  «TTH FCTQ.  OCCUR. I F5S _THAN 5 «Nn_MIIMEP IC«L_»NO Q.0.4" IME TBiT,_
                cn»pn<;iiinN '>OT» i.tss THAN i  AOE ExCLiiOEN  run" THE  TIRLE ANO PLOT    "~
                (BUT NOT FRO" TA| CIIL'TIOM OF  niVFP^ITY TNOITFS)
                  PF.PCENT  DOMINtNTE  IMOFX
                  S^AKiNON-v'F INFO DIVERSITY
                  EVENNESS iwntj	
 .38
1.90
Fig.  B-50.   IRI  prey  spectrum  of  tidepool  sculpin  in  littoral  habitats
                along  the Strait of  Juan  de  Fuca.
                                                156

-------
Table B-7.  Prey composition (numerical composition)  of tidepool sculpin
             at Port Renfrew, Vancouver Island, B.C.,  documented by Nakamura
              (1971).
PREDATOR  8831032401 - OLIGOCOTTUS MACULOSUS
                                                           (TIOEPOOL SCuLPIN
              INDEX  or  RELATIVE  IMPORTANCE   TABLE
                FROM FILE  IDENT.  REF 68* STATION SLSTA
           PREY ITEM
                                  FREO   NUM.   GRAV.  PREY     PERCENT
                                  OCCUR  COMP.  COMP.  I.R.I.  TOTAL IRI
  OSTRACOOA
  COPEPODA
  MOLLUSCA
  GAMMARIDEA
  DECAPOD*
  INSECTA
  OSTEICMTHYES
  ISOPOOA
  ANNELIDA
  CAPRELLIDEA
                                           .70
                                        61.40
                                           .40
                                        23.60
                                          1.00
                                          4.30
                                           .40
                                          3.50
                                          4.30
                                           .10
    PPEY TAXA WITH FREO.  OCCUR.  LESS THAN 5 AND NUMERICAL AND GRAVIMETRIC
    COMPOSITION BOTH LESS THAN  i ARE EXCLUDED FROM THE TABLE AND PLOT
     (BUT NOT FROM CALCULATION OF DIVERSITY INDICES)
PERCENT DOMINANCE  INDEX
SHANNON-WEINEP DIVERSITY
EVENNESS INDEX
 .44
1.67
 .so
                                                 .00
                                                 .00
                                                 .00
                                                                  .00
                                                                  .00
                                                                  .00
                                       157

-------
                       a
                       i
                       a
                       s
                       o
                            lOOr
80
                            60
                               INDEX OF RELRTIVE  IMPORTRNCE (I.R.I.) DIRGRRM
                               FROM FILE  IDENT.   76-78.  STRTION flLSTfl

                                             8831022402 - OLIGOCOTTUS RIMENSIS
                                                          SfiDDLEBRCK SCULPIN
                                                 RDJUSTEO SRMPLE SIZE =  75
                             40
20
                       2    20
                             40
                            60
                            80
                            100
                                      20
                                              40
                                                       60
                                                               80
                                                                       100
                                                                                120
                                                                                        140
                      PREY  !TCM
                                             CUMULRTIVE FREQUENCY OF OCCURRENCE
                                            FREO   NUM.   GRAV.   PREY     PERCENT
                                            OCCUR  COMP.   COMP.   I.R.I.   TOTAL  IRI
GAUMARIOEA
HARPACTICOIOA
SPWAFRIV'AT II1AE
POLYCHAFTA
CIOOIPEDIA 	 	 .
FLABFLLIFFRA
ISOPODA /
CHL030PHYT-A/ _ _. -. 	
ARGHL IHEA
NATANTIA
COTTIDAF
56.00
	 3?. 00
??.67
4. 00
	 4.00.
?.67
1.33
1.33
1.33
1.33
^ 1.33
3fl 14
31.27
14.43
1.03
2.75
1 .03
.34
... .34
4.12
.34
.34
55.04
1.03
16.57
1 .44
.. .98
1 .44
1.92
7.68
.02
3.64
	 6.96_
"18.5
1033.7
702.7
9.9
1<(.9 	
6.6
3.0
10.7
5.=;
4.0
9.7
74.16
14.69
9.99
.14
.21
.09
.04
	 .15 	
.08
.06
.14
               DPFY  TAXA  WITH FREO. OCCl'R. LES«; THAN S ANH NUMERICAL  AND  GRAVIMETRIC
               _Cn-«POSITION_woTH LESS THAN  ]_ARF ExCLMOFO FPOM_THE TABLE_ANO  PL.OT	
               (RUT  NOT FROM CA|_CULATIO" IF DIVERSITY INDICES)
                  PFRCFNT _OOMlMANiCE_ INOEX	
                 "SHANNON-WFINEP DIVERSITY
                  EVENNESS
_
 ?.50
  .54
                               ?.35
                                .51
1.1 fl
 .25
Fig.  B-51.   IRI prey spectrum  of  saddleback  sculpin  in littoral  habitats along
               the Strait  of  Juan de Fuca.
                                                158

-------
                             INDEX OF RELRTIVE ItlPORTRNCE (I.R.I.) DIROfWI
                             FROM FILE IDENT.  76-78. STRTION RLSTR
                          lOOr
                                            8831022403 - OLIGOCOTTUS SNYDERI
                                                          FLUFFY SCULPIN
                                               flDJUSTED SfittPLE SIZE - '86
                      8
                      a.
                                           CUMULRTIVE FREQUENCY OF OCCURRENCE
FPEO
PREY ITEM
GAMMSPTDEA
e;pt- AF_ROM AT I DAE
HARPACT ICOIOA
POLVCHAETA
..TANA1DACE-A 	 	 	
IPOTEIPSE
CIRRIPEDIA
P<;TPACODA
NE«FPTEA
CAPPELLIDEA
..S'PELUPAE
OCCUR
5«
?*•
17
1?
	 12
fl
5
^
3
?
1
.14
,74

.79
.79__
.14
.«!
.65
.49
.33
. 16
NUM.
CO
47
in

3
_3
2
3
1

1
4
MP.
.59
.39
.01
.46
.93-
.54
.00
.15
.69
.15
.62
GRAV.
COMP.
58
1?

15
1
7


1
1

.10
.51
.76
.64
.34_
.14
.14
.01
.00
.00
.75
PPEY
I.R.
6143
612
3?7
?44
	 67
78
18
5
5
5
. 	 6.
PERCENT
I. TOTAL IPI
.7

.5
.3
.-< 	
.ft
.3
.5
.9
.0
.?
81
R
4
3

1





.64
•14
.35
.25
.89 	
.05
.24
.07
• 08
.07
.08
             PREY TAXA WITH FPEO.  OCCUR. UFSS THAN 5 AND NUMERICAL AND GRAVIMETRIC
             rOMPO'UTION HOTH..LtS.S.THAN 1 ARE EXCLUDED FPOM_THE TABLE..AND. PLQT_
(BUT NOT FROM CALCULATION OF
PFPCENT DOMINANCE IMDF.X
SHANNON-WFINEP DIVERSITY
EVENNESS INDEX
DIVERSITY INDICES)
.23
2.M
.60
.38
1.99
.46
.68
1.09
.25
Fig.  B-52.   IRI  prey spectrum of  fluffy  sculpins  from littoral habitats in
               Strait of Juan de Fuca.
                                              159

-------
     In Nakamura's (1971) comparison of the food habits of Oligocottus
maculosus and (). snyderi at Port Renfrew, Vancouver Island, B.C., (). snyderi
was shown to feed much more upon gammarid amphipods than 0^ maculosus, while
polychaete annelids and isopods were of secondary importance  (Table B-8).
Accordingly, harpacticoid copepods, which composed over 60% of the total
number of prey consumed by CK maculosus, composed less than 5% of the prey
consumed by (). snyderi.

Cabezon

     Juvenile cabezon caught by beach seine at a cobble beach habitat (South
Beach) in northern Puget Sound had full or nearly full stomachs.
Oniscoidean isopods were the most important prey (55.6 percent of total IRI).
Various epibenthic decapods (41.0 percent) including a shrimp (Heptacarpus
stimpsoni), a crab (Cancer oregonensis), and amphipods (mostly Eusiroides sp«,
also Atylus sp. and Allorchestes sp.), and a parapagurid hermit crab (3.4
percent) were also included in the prey spectrum.

     Cabezon from northeastern Guemes Island and Legoe Bay collections along
the eastern shoreline had consumed Dungeness crab, Cancer magister, shrimp
parts, and a rock.

     O'Connel (1953) described the food habits of cabezon in California
throughout their life history.  Based on both frequency of occurrence and
gravimetric composition, crustaceans (primarily shrimp, Spirontocaris sp.
and Hippolytidae, crabs, Cancer sp., Pugettia sp. and Grapsidae, and gammarid
amphipods) and fish (Cottidae, Gibbonsia sp. and Blennidae) were important
prey of juveniles (Tables 9a and lOa); crustaceans (Cancer sp.,  Pugettia sp.,
Scyra sp., Grapsidae) and molluscs (Haliotis sp., Acmaeidae) were important in
the diet of subadult (Tables 9b and lOb); and crustaceans (Cancer sp.,  Majidae,
Pugettia sp., Phyllolithodes sp., Scyra sp., Hemigrapsus sp., Pasurus sp.
Cryptolithodes sp., Idotea sp.), molluscs (Haliotis sp., Acmaeidae, Cephalopoda,
Mimulus sp., P'olyplacophora. Bivalvia) and fish (Sebastes sp., Citharichthys  sp.,
Cottidae) were all important in the diets of adult cabezon (Tables 9c and lOc).
O'Connel (1953) also illustrated seasonal differences where, based on gravimetric
composition, crustaceans became more important than fish in the spring diet
spectra for juvenile cabezon; molluscs were important to subadult cabezon just
during spring; and fish and molluscs increased in proportional contribution
during winter and spring in the diet of adult cabezon.  In Ouast's (1968)
analysis of food habits of kelp bed fishes, decapod crustaceans and ceaphalo-
pods (Theuthidida), predominated the diet of cabezon (b ased on frequency of
occurrence) while fish (Perciformes), unidentified  algae, other crustaceans
and gastropods also occurred frequently (Table B-ll).

Tubenose Poacher

     Beach seine collections in the Strait of Juan de Fuca (Twin Rivers and
Morse Creek) provided tubenose poacher stomach samples which  illustrated
a very specialized diet oriented almost exclusively toward mysids (Fig. B-53) .
                                      160

-------
  Table B-8.   Prey composition  (numerical composition) of  fluffy sculpin at
               Port Renfrew, Vancouver Island,  B.C., documented by Nakamura(197l).
PREDATOR  8631022*03  - OLIGOCOTTUS SNYDERI
                                                           (FLUFFY SCULPIN
              INDEX  OF  RELATIVE IMPORTANCE d.R.I.I  TABLE
                FROM FILE  IDENT. REF 68, STATION SLSTA
              • «»••••••••*•*•«**•**»•**<>•»*»<
           PREY ITEM
FREQ   NUM.
OCCUR  COMP,
GRAV.  PREY
COMP.  I.R.I.
                                                              PERCENT
                                                              TOTAL IRI
  OSTRACOOA
  COPEPOOA
  MOLLUSCA
  GAMMARIOEA
  OECAPODA
  INSECTA
  ISOPODA
  ANNELIDA
  CAPRELLIDEA
        ~ .20
        3.60
         .60
       70.00
         .60
         .60
       11.80
       12.20
         .SO
    PREY TAXA WITH FPEO. OCCUR. LESS THAN 5 AND  NUMERICAL AND GRAVIMETRIC
    COMPOSITION BOTH LESS THAN 1 ARE EXCLUDED FROM THE TABLE AND PLOT
    (BUT NOT FROM CALCULATION OF DIVERSITY INDICES)
       PERCENT DOMINANCE INDEX
       SHANNON-WEINER DIVERSITY
       EVENNESS INDEX
          .52
          .46
          .46
   .00
   .00
   .00
.00
• 00
.00
                                      161

-------
 Table B-9.   Prey composition  (frequency of occurrence and gravimentric
              composition) for  general  prey categories consumed by  juv-
              enile  (a), subadult (b),  and adult  (c) cabezon in central
              California documented by  Connell(1953).
PREDATOR  883t-;3101  -  SCORPAENICHTHYS MARMORATuS
                          (CABEZON
              INDEX  or  RELATIVE  IMPORTANCE 
-------
Table B-10.   Prey composition  (frequency of  occurrence)  for specific prey
               categories consumed by juvenile (a), subadult  (b),  and adult
               (c) cabezon in central California documented by  Connell(1953),
 •REOATOR  8831023101 - SCORPAENICHTHYS MARHORATMS
                         (CABEZON
              INDEX OF RELATIVE IMPORTANCE (I.R.I.)  TABLE
                FROM FILE IOENT. REF 69. STATION SLSTA
              »•••••*•*••••••••••••••»•••»•«**••••*•••••••
   a. juvenile
           PREY  ITEM
FREO
OCCUR
                                         NUM.
                                         COMP.
GRAV.
COMP.
PREY
I.R.I.
 PERCENT
TOTAL IRI
  SPIRONTOCARIS SP.
  CANCER  SP.
  OSTEICHTHYES
  GAMMARIOEA
  COTTIDAE
  GIBBONSIA SP.
  PUGETTIA  SP.
  BLENNIDAE
  GRAPSIOAE
  HIPPOLYTIDAE
  COPEPOOA
42.86
17.14
14.29
14.29
 5.71
 5.71
 ?.86
 2.86
 2.86
 2.86
 2.86
     PREY  TAXA  WITH  FREO. OCCUR. LESS THAN 5 AND NUMERICAL AND GRAVIMETRIC
     COMPOSITION  BOTH LESS  THAN  1 ARE EXCLUDED FROM THE TABLE AND PLOT
     (BUT  NOT FROM CALCULATION OF DIVERSITY INDICES)
        PERCENT  DOMINANCE  INDEX
        SHANNON-WEINER DIVERSITY
        EVENNESS INDEX
         .00
         .00
         .00
  .00
  .00
  .00
   b. subadult
            PREY ITEM
FREO   NUM.   GRAV.  PREY
OCCUR  COMP.  COMP.  I.R.I.
            .00
            .00
            .00

         PERCENT
        TOTAL IRI
   CANCER SP.
   PUGETTIA SP.
   OSTEICHTHYES
   SCYRA SP.
   HALIOTIS SP.
   GRAPSIDAE
   ACMAEIDAE
le.ie
18.18
 9.09
 9.09
 9.09
 9.09
 9.09
     PREY TAXA WITH FREO. OCCUR. LESS THAN 5 AND NUMERICAL AND GRAVIMETRIC
     COMPOSITION BOTH LESS  THAN  1 ARE EXCLUDED FROM THE TABLE ANO PLOT
     (BUT NOT FROM  CALCULATION OF DIVERSITY INDICES)
        PERCENT DOMINANCE  INDEX
        SHANNON-WEINEP  DIVERSITY
        EVENNESS INDEX
         .00
         .00
         .00
  .00
  .00
  .00
            .00
            .00
            .00
                                        163

-------
Table B-10  (continued).
. adult
PREY
ITEM
FREO
OCCUR
NUM.
COMP.
6RAV.
COMP.
PREY
I.R.I.
PERCENT
TOTAL IRI
 CANCER  SP.
 MAJIDAE
 PUGETTIA SP.
 HALIOTIS SP.
 ACMAEIOAE
 PHYLLOLITHODES SP.  _
 SCYRA SP.
 SEBASTES SP.
 CEPHALOPODA
 UNIDENTIFIED
 MIMULUS  SP.
 OSTEICHTHYES
 POLYPLACOPMORA
 BIVALVIA
 PAGURUS SP.
 CITHARICHTHYS SP.
 HEMI6RAPSUS SP.
 COTTIOAE
 CRYTOLITHOOES SP.
 IDOTEA  SP.
45.65
45.65
43.46
16.30
16.30
 9.78
 8.70
 5.43
 5.43
 4.35
 4.35
 4.35
 4.35
 2.17
  .09
 1.09
 1.09
  .09
 1.09
 1.09
   PREY TAXA WITH FREO. OCCUR. LESS THAN 5 AND NUMERICAL AND GRAVIMETRIC
   COMPOSITION BOTH LESS THAN 1 ARE EXCLUDED FROM THE TABLE AND PLOT
   (BUT NOT FROM CALCULATION OF DIVERSITY INDICES)
      PERCENT DOMINANCE INDEX
      SHANNON-WEINEP DIVERSITY
      EVENNESS INDEX
         .00
         .00
         .00
.00
.00
.00
.00
.00
.00
                                         164

-------
Table  B-ll.  Prey composition (frequency of occurrence)  of
              southern California kelp beds documented by Quast(19oH>
          8B31023101 - SCORPAtNICHTHYS MAPMORATMS
                                                           (CARE70N
              IN1E*
                     F  PELATIVE  IMPORTANCE (I.P.I.)  TARLF
                     FILE  IOENT. REF S<3 . STATIO" 5LSTA
                                FPEO
                                OCCUR
                                        NUM.
                                        COMP.
GRAV.
COMP.
                                                     I.R.I,
 PERCENT
TOTAL IRI
               ALGAF
THEUTHIDIDA                       69.00
                                as.oo
                                25.00

                                20.00

                                 7.00
                                 7.00

  PRFY TAXA JTTH FPEO. OCCUR. LESS THAN 5 AND NUMERICAL  AND GRAVIMETRIC
  COMPOSITION "OTH LE = S THAN 1  ARE EXCLUDED 'FooM THE  TARLE AND PLOT
  (PUT NOT F90M CALCULATION OF  DIVERSITY INDICES)         "  	"
  COIJSTACE4
  PLFnCYfVA
 "GASTROPODA
               DOMINANCF  [NOFX
       SHA^j^JO^^-w(r IMF.D
                INDEX
                                         .00
                                         .00
                                         .00
  .00
  .00
  .00
    .00
    .00
    .00
                                        165

-------
                      (X
                      >-
                      CO
                      z
                      o
                           100
                           80
60
                           40
                           20
                      2    20
                           40
                             INDEX OF RELRTIVE IMPORTRNCE (I.R.I.)  DIRGRRM
                             FROM FILE IDENT.  76-78. STRTION RLSTR

                                             8831081101 - PRLLRSINR BRRBRTR
                                                         TUBENOSE PORCHER
                                               ROJUSTED SRMPLE SIZE   35

                           60
                           80
                           100
                                      20
                                                40
                                                          60
                                                                   80
                                                                             100
                                                                                       120
                     PREY ITEM
                                            CUMULRTIVE FREQUENCY OF OCCURRENCE
                                            FREQ   NUM.   C-RAV.  PRFY     PERCENT
                                            OCCUR  COUP.  COMP.  I.R.I.   TOT4L  IPI
PAMMARIDEA
CHWACEA
C-AMMABIOAE
MATANTIA
17.14
S.71
?.7)
- 5.7L.
94.07
1.34
.57
_3.63
«8.SO 13S84.6
?.2\ ISO. 9
1.1? e.f.
1.H3 13.7
6.04_ ._55.3-
98.99
.44
.10
_. .40
              OBEY TAXA »ITH fPEO.  nCCljR.  LES1?  THAN «» AND NUMERICAL ANH GOAylMETPIC
              -COMPOSITION BOTH LESS THAN  1. APE  EXCLUDED EROM THE TABLE AND .PLOT.	
              (RilT NOT rpox CA|CUL*TIO'J Of"  DIVERSITY INDICES)
                __
                 SHAMNON-UFINEP DIVERSITY
                          INDEX
                         .89	,79_
                         .41    .70
                         .11    .30
.•99
 .10
 • 04
Fig.  B-53.   IRI prey spectra of tubenose poacher from  shallow  sublittoral
               habitats in Strait  of  Juan de  Fuca.
                                               166

-------
Tadpole Sculpin
                                                  n rtions from both northern
     Although they appeared in neritic (townet) co^e"      originated only
Puget Sound and the Strait of Juan de Fuca, stomach conte        dg and mysids
from collections in the Strait of Juan de Fuca.  Ga»"*"a   epibenthic feeding
were the prominent food organisms (Fig. B-54)  suggesting an ep
behavior.

Pacific  Spiny Lumpsucker

     Beach seine collections at Cherry Point and Legoe Bay provided the  few
Pacific  spiny lumpsucker specimens  for analysis of stomach Contents.   Gammarid
amphipods supplied 56.1 percent of  the total IRI, hyperiid amphipods, -">•*
percent.  Caprellid amphipods (3.0 percent), valviferan isopods IJ.l percent;,
and cumaceans (0.6 percent) were incidental prey.

 Sturgeon Poacher

      Sturgeon poachers in  the  north Puget  Sound  collections originated from
 collections at Birch Bay.   The total  IRI was  rather  evenly distributed among
 cumaceans, gammarid  amphipods,  shrimp (Crangonidae and Penaeidae), and
 harpacticoid copepods.   Polychaetes  and tanaids were also found in the
 stomachs.

      Along  the  Strait of Juan de Fuca, winter beach seine collections at
 Beckett  Point and West Beach furnished the greatest number of sturgeon
 poachers  for  stomach analysis.  The overall prey spectrum (Fig. B-55) was
 divided  among cumaceans (36.3 of total IRI), gammarid amphipods (29.6/.) ,
 and harpacticoid copepods (22.9%)  as primary prey organisms,  and "angonid
 shrimp  (including Crangon alaskensis and  C. stylirostris) as secondary  prey.

 Ribbon  Snailfish

       Beach seine and tidepool collections at Jamestown and Twin Rivers  along
  the  Straits of  Juan de Fuca produced stomach samples of ribbon snailtisn,
 wherein over 94% of the total IRI was contributed by gammarid amphipods.
  Sphaeromatid isopods (Gnorimosphaeroma oregonensis and Exosphaeroma amplicauaa;
  and  mysids provided the remaining 6% (Fig. B-56).

  Tidepool Snailfish

       Some tidepool snailfish from a beach seine  collection at Birch Bay  had
  consumed principally gammarid amphipods (41.4 percent of total IRI), polychaetes
  (39.8 percent)  and valviferan isopods (15.4 percent), with shrimp (Penaeidae)
  providing a small contribution (3.4 percent).
   intertidal  th^  Strait'  tidepool  snailfish  were  the  most common snailfish in the
   Point  and  si'   p*""8'  and were regularly collected  at Morse Creek,  Observatory
                        '   AS i?  the C3Se  °f  the ribb°n  snailfish,  gammarid
                        woT    ° I**** ^^ IRI'   Id°te±d "opods^ncluding
                                                    montereyensis)  and  harpacticoid
                                         167

-------
                               INDEX OF RELflTIVE  IHPORTHNCE (I.R.I.) OlflGRRM
                               FROM FILE IDENT.   76-78. STflTION PLSTfl

                                        PREDHTOR  8831070101 - PSYCHROLUTES PRRflDOXUS
                                      (TflDPOLE SCULPIN    1   flOJUSTED SHMPLE SIZE =  12
                            100
                             80
                             60
2
O
£
§
fe
s
£
1
s
§
8
Q-
§

t—
u
Q-




40


20
0
20

40
60



80


100
























— 1



V
•0 tJ
s . 5 £
•° S e S
•S « 61!
g 5 S g
g ^ nt -o
I III
1 1 1 1 1 1
0 20 40 60 80 100 12
                                             CUMULflTIVE FREQUENCY OF OCCURRENCE

OS.fi TTFM

MYCJHACEA
PPHAEPOMATIOAE
I'WI DFNT IF ICO
FREO
occu"

SO. 00
33.33
16.67
P. 33
NU".
COMP.

37. SO
50.00
8.33
4.17
GRAV.
COMP.

58.47
37.04
3.37
1.12
PPF.Y
I.R.I.

™l:;
44.]
PERCENT
TOTAL IRI

60.44
36. S4
2*46
• 56
                PPFY TA^»_wTTH_FPEO^_OCCl!P. LFSS_THAN 5 AND NUMERICAL ANO GRAVIMETRIC
                ch'Mposifioiii ROTH LESS" THAN  i" ARE EXCLUDED FQOM THE TABLE AND PLOT
                ('BUT NOT FROM CAl.CUL'TION OF DIVERSITY INDICES)
                   PEOCFNT  DOMINANCE  INDEX
                   SMANNON-WEIMFO  DIVERSITY
                   EVENMFSS INOE«
 .40     ,4R
1.5?    1.22
 .76     .61
 • 50
1.14
 .57
Fig. B-54.   IRI prey spectrum  of  tadpole  sculpins  in  neritic  waters  along
               Strait  of  Juan de  Fuca.
                                                168

-------
                                  INDEX OF RELRTIVE IMPORTHNCE (I.R.I.) DIRORRM

                                  FROM FILE  IDENT.  76-78. STflTION RLSTR

                                                    8831080802 - ROONUS RCIPENSERINUS

                                                                   STURGEON  PORCHER

                               1QO                      RDJUSTEO SfittPLE SIZE =  82
                                80
                                 60
                                 40
                           8     20
                                100
                                                                              CJ  4J  -o  «  efl n)     T-(
                                                                           «  U  Qi  IH  E    a.
                                                                           «  «a  o  M  a  x rn e -H ••-»
                                                                           Z  H  PL.  u  w E C 3 CQ 3C
                                              50
                                                          100
150
                                                                                200
                                                                                           250
                                                                                                      300
                                                     CUMULATIVE FREQUENCY OF OCCURRENCE
oREr ITEM
f, 4«M4RIDE4
CU"ACEA 	 	 	
H4CD4CT ICOID4
GAMMAPinAE
MATAMT1A 	
T4^ AICiACEA
DHL YrnA£T4
ro»Nc,oN]B4f
<;P»-AFOO^AT IDAE
WYC In A TEA
^*> TDACQOA
H'llnENTIFIED
RTV4L VI A
_HI_P£nLYT IOAT

FREO NUM.
OCCUR COMP.
51.22 17.
	 49.7B_..30.
3?. 93 43.
??
ft. in
ft. 10
33
4S_.
31
68
92 	
53
62
6B . .
49
30
09
33 ~
15
21
GRAV.
COMP.
17. IS
20.19
4.13
6.27
PREY PERCENT
I.R.I. TOTAL IRI
1765.9
2471.6 	
1562.?
?51.3
.,8.68 	 1B7.4 	
1.74
3.10
29.58
.41
i.as
• Oft
~.93
.19
_?l64
43.X
50.0
	 406.0 	
11.0
?3.9
I y
	 	 - 1 . C. 	 _
9.?
2.0
. L7.4
?5.91
36.27 	
?2.93
3.69
_.2.75 —
.64
.73
. 5.96-_
.16
.35
. n ?
	 • V C. 	
.13
.03
	 .26 	
                               OTHPf?;«.°TC"h'* ^^  ™ftN  S *Nn  NUMERICAL AND  GRAVIMETRIC

                                C», rt), AT!HM  or ^E  fXCLL)OED FOPM THE  TABLE ANO PLOT
                                LAICULATJON  OF OIVFRSITY INDICES)  ~      '  	      	
Fig.  B-55
             •
PFDCE'JT OnMlMA'.'Cr

SHANHON-WFIMFP

EVEi-wF. SS  INOE.
                                                             .31

                                                            ?.09

                                                             .44
 •I7

3.0?

 .64
                    .26

                  ?.29
                                                     169

-------
                               INOEX OF  RELflTIVE IMPORTflNCE (I.R.I.) DIflGRRM
                               FROM FILE IDENT.  76-78. STflTION RLSTfi
                                                8831090806
                            lOOr
           LIPRRIS CrCLOPUS
           RIBBON SNRILFISH
flDJUSTEO SRMPLE  SIZE    39
                            80 •
V 60
CD
ION BY HEIGHT PCT. COMPOSITION
-p. N) N> *.
O O O O O
§ 60
H










\ 1 1

11
n
•u
•H a>
(U «> 4-t PI
                            80
                                 -O 
-------
                       INDEX OF RELRTIVE IMPORTRNCE (I.R.I.) DIRORRH
                       FROM FILE IDENT.  76-78. STRTION RLSTR

                                         8831090810 - LIPRRIS FLORHE
                                                   TIOEPOOL SNRILFISH

y
CQ
CC
>_
CO
s
1—
§
0
k-
a
1—

LJ
3
£
2
O
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**-




PREY
Oft WM ABIDE A
_I On T t 1 0 ftF
^P^AfftOMAj I n A £"
HADP6CTICOIDA
C AOPri I T nr- A
-w r C-L_l_iUt- *^ 	
H]DP(^l_YT ID6F
IMCFCTA
L'NIDENTIFIEO
NATAMT 14

	 °OFV TOIA WI
CnMDQMTiorg
I^UT MOT FPO
100
80

60


40

2C
f
L
20

40

60



80


100
RD JUSTED SRMPLE SIZE = 87


.


•


















	 "^LTtij





.
(0 CO
TS -a a) i
•H TH eg to 1
HE TABLE AND PLOT
    	-EvE'.'jEss'
                        NCF  IMOEX
                         DIVERSITY
 .49    .4,7
1.65   1.79
 • 3S.._ ,38_
.89
.43
.09.
B-57
                                    de^ucf  Sna±lfish  in littoral habitats
                                         171

-------
Shiner Perch

     Shiner perch were relatively common in the nearshore beach seine catches
in northern Puget Sound, especially at Deadman and Westcott bays during summer;
they were also caught in the townet at Birch Bay, Cherry Point, and Burrows
Island.  Prey composition was relatively equally divided between a number of
epibenthic organisms (Fig. B-58), gammarid amphipods (38.6 percent of total
IRI), cumaceans (29.2 percent), and polychaetes (22.2 percent) with caprellid
amphipods making a lesser congribution (8.0 percent).

     Shiner perch also ranked high among the most abundant species in the
collections along the eastern shoreline, being especially common at the
Cherry Point (cobble habitat), Padilla Bay (mud/eelgrass), and Legoe Bay
(gravel) sites.  Compared to the San Juan Islands samples, the prey composition
from these collections was considerably less diverse (H* = 2.61 for abundance
and H' = 2.60 for biomass, versus H1 = 1.53 for abundance and H' = 1.26 for
biomass) and was dominated by gammarid amphipods (95.6 percent of total IRI)
with only minor contributions by calanoid copepods and isopods (Fig. B-59).

      Shiner perch was one of the principal schooling nearshore fishes
characterizing the eastern MESA sampling sites along the Strait of Juan de
Fuca,  especially at  Beckett Point and Jamestown.  The diverse IRI prey spec-
trum was composed of gammarid amphipods  (47% of total IRI), cumaceans, harpac-
ticoid  copepods, tanaids, sphaeromatid isopods  (including Gnorimosphaeroma
oregonensis and Exosphaeroma amplicauda), algae, and calanoid copepods (Fig.
B-60).
     DeMartini (1969) listed bivalve molluscs, gammarid amphipods, tanaids,
cumaceans, polychaete annelids and ostracods as the principal prey organisms of
shiner perch.

     Weller's (1975) analysis of shiner perch food habits in Anaheim Bay,
California, indicated that unidentified eggs dominated the prey biomass during
all seasons (Table B-12a-d) although topsmelt (Atherinops affinis) were
somewhat important in the spring, insects and mussels (Mytilus edulis)
appeared in the summer and fall diets and mussels, polychaetes, gastropods and
topsmelt occurred during the winter.

     The studies of Bane and Robinson (1970) in upper Newport Bay, California
indicated only Potoamogetonaceae plants (includes eelgrass, Zostera marina),
crustaceans and rocks as the principal prey of shiner perch; this is based,
however, only on frequency of occurrence data (Table B-13).

Striped Seaperch

     Striped seaperch from northern Puget Sound (Deadman Bay, Guemes Island
and Cherry Point) had a prey composition composed almost entirely of epibenthic
or benthic crustaceans—gammarid amphipods, valviferan and flabelliferan
isopods, crabs and shrimp, while in the Strait of Juan de Fuca (Beckett Point,
Twin Rivers, and Morse Creek) over 90% of the striped seaperch prey spectrum
was made up of gammarid amphipods, supplemented by sphaeromatid (Gnorimosphaeroma
oregonensis) and idoteid (Synidotea nodulosa) isopods.


                                     172

-------
                             100
                                INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) DIHGRRM


                                FROM FILE  IDENT. N POSO. STRTION RLSTH



                                           PREORTOR   8835600201 - CYMHTOGflSTER RGGREGflTfl

                                        (SHINER PERCH        )   flDJUSTEO SflMPLE SIZE -  31
                              80 r-
                              60
" 40
-











=i- 4p-

                        o

                        fe
                        o
                        o
                              40 h
                              60 r
                              80 h
                             100L
                                                                   QJ « efl     eg CD

                                                                   w-i -H V     *O I

                                                                   1-1 -o u  « « o «
                                                                   ^ 0) « T) -H D. "B

                                                                   i-t a-a  o > o o
                                                                   01 -H ft  O.f-1 VJ O.
                                                                   £ b m  ai  « O
                                       20
                                               40
                                                      60
                                                              80
                                                                     100
                                                                            120
                                                                                    140
                                                                                           160
                                                                                                  180
                                                 CUMULflTIVE FREQUENCY OF OCCURRENCE
°RFY ITF"
r-A"'J«p IO(£A 	 	 	
POI vfMir TA
rt|^*^fa — 	 	
TAnnFLL 1 DE A
^t "P^LLTTrc A 	

CloolPFOIA
TA^AJOACE A " — — — - - — 	 	
P I V A L V 1 A - 	 	 .
GASTROPODA
DFCAPnoA-ypacnYUPA 	 	 	

"Cp^;? -'i;;/°r«: "EC,,*. LE
FRFO
OCCUR
55.00
17.00
?*.oo
?1.00
6.00"
6.00
- 6.00" '
1.00
1.00"
1.00
1 • 00 -
:«<; THAN
Nl.1'1 .
COHP.
38.82
2.17
16.30
4.31
" 2.90
1 .60
'• .43
10.31
1.11 —
1 .2?
.05-
5 AND
GRAV.
COMP.
?9.10
34.10
16.81
12.09
.95
.03
.03
.48
"4.35
.02
1.2T
NUMER
PREY
I. a. i.
18585.6
1 160.6
1180.9
177.?
?3.1
9.0
2.«
12.4
— 16.4
3.7
— ~ 3.9 '
IC»L ANn
PEBLtNT
TOTAL IRl
96.10 	
5.40
6.40 "" ~
1.70
.10
.00
.00
.20
" .10 	
.00
	 .00"
GRAVIMETRIC
                                           AN  '  *RE EKCLUOEO  FoQM THE  TABLE ANO CLOT

                                                -
Pig.  B-58.
."•IT HOT  rpo» CALCULATION  OF-D1VERSITY INDITES.	




   ^"CFNT-OOMlNA.jCF INDFX	-,„—   „

   '•"A'.NO'J-WFINFP  DIVFC^ITv              •3°     "?5
   tvEM.j£<;c  TNf1F.  UIVtP^ITr             ??1


                                           .6?     .65




IRI prey
                                                                                       .75"

                                                                                       .80

                                                                                       .29
                                                     173

-------
a
i
                       £
                       g
                              INDEX OF RELRTIVE INPORTRNCE (I.R.I.) OlflORfiH
                              FROM FILE IDENT. HH  BS.  STRTION flLL
                                       PREORTOR  8835600201  - CYMflTOORSTER flOOREGflTfl
                                     (SHINER PERCH       I   flDJUSTED SflMPLE SIZE = SI
                           100
                            80
                            60
                       e    40
                       8
                       fc
                       3    20
                       2    20
                            40
                            60
                            80
                           100
                                    10
                                          20
                                                30
                                                       40
                                                             SO
                                                                   60
                                                   70
                                                                                80
                                                                                       90
                                            CUMULflTIVE FREQUENCY OF OCCURRENCE
PREY ITEM
fiAMMARIOEA
CALANOIDA
ISOPOOA
CRUSTACEA
ANTHOZOA
PACURIDAE
FREO
OCCUR
53.00
14.00
12.00
4.00
?.oo
2.00
NUM.
CO"P.
ft7 fl  PLOT
              _(Ry_T  NOT_FROM_CAl CULAjriON_OF_gIj/ERSJTY  INDICES)	
                  PERCENT DOMINANCE INDEX
                  SHANNON-WEINER. DIVERSITY
                  EVENNESS 'INDEX
                                                     .52
                                     .63
                             l.?5_   1.11
                              .<•<»     .43
.86
.45
.22
Fig.  B-59.   IRI  prey spectrum of  shiner perch  from shallow sublittoral  habitats
               along the eastern shoreline of northern Puget  Sound.
                                              174

-------
                          INDEX OF RELHTIVE IMPORTRNCE (I.R.I.) D1RORRM
                          FROM FILE IDENT.  76-78. STRTION RLSTR

                                        8835600201 - CYHRTOORSTER ROGREGRTR
                                                         SHINER PERCH
                        100
                         80
                         60
                         40
                         20
                    2    20
                         40
                    8
                    a.
60
                         80
                        100
                                             RDJUSTED SRttPLE SIZE = 168
                                                                                Q
                                        •o
                                        •H
                                        o
                                   20
                                            40
                                                    60
                                                             80
                                                                     100
                                                                              120
                                                                                      140
                                          CUMULflTIVE FREQUENCY OF OCCURRENCE
                                         FREQ    NUM.    GRAV.   PPEY      PERCENT

r,Au
PPF.Y ITFM
M40IOEA
OCCUR
38
.A9
ClJ"ACEA 1°.45
SPh
TAN
4rpOMAj 10AF
SIDACEA
HAODACTICOIHA
irn
"01
GA"
y Y<=
1 11. O
TAL
K. A T
r A i
1 01
TFIDAE
YCHAFTA
•••4DJ04E
TDACFA
TD I CHA[_ES
4 • 1 0 I 0 A
AMTIA
L I AN Ac S I Pi AF
11
1 1
1 1
7
ft
=;
4
4
	 3
1

.10
.90
.11
.74
.55
.Tft
.7ft
.17
.57
.19
.60
COMP.
10.49
77.31
3.61
14.09
71.70
.45
.32
5.9ft
1.35
.97
10.P5.
.05
.03
COMP.
24
9
10
7
2
2
3
7
2
17
1
1
7
.43
.50
.07
.45
.99
.18
.75
.88
.77
.19
.24
.14
.46
I.R.
1151
e.79
179
256
101
2.0
26
74
19
75
	 41
1
4
I. TOTAL IRI
.1 44
.3 ~t
.1
.4
.9 	 	
.1
.ft
.2
.ft
.7
. 0 	
.4
.5
•2
5
8
9


2

2
.1


.51
• 38 _
.90
.45
.95 ..
.67
.88
.44
.65
.49
.35_ _
.05
.15
                   A wITM FREO. TCr,,
                         C4|
                                       LESS THAN! ; AND NUMERICAL  AND  GRAVIMETRIC
                                       1 ARF EXCLUDED FopM  THE  TABLE AND  PLOT
                          .18
                        7.85
                          .58
                                                          .13
                                                         3.34
 .27
2.42
 .49
Fig. B  60.   IRI prey  spectrum of  shiner  perch  in  the  Strait  of  Juan de Fuca.
                                             175

-------
 Table B-12
PREDATOR
           Prey composition  (gravimetric  composition) of  shiner perch
           during spring  (a),  summer (b),  fall  (c), and winter  (d) in
           Anaheim Bay, California,  documented  by Weller(1975).
                     - CY 'ATO'jASTFP
                                                             (SHINFR PERCH
              IMOE* "F PF.LATIVF  IMPORTANCE  (I.P.I.)  TABLE
                FRO" FII_F  IDENT.  PEF  51.  STATIOM  SL5TA
                 ITEM
                              FPFO   NUM.    GPAV.  POFY
                              OCCUR  COMP.   COMP.  I.R.I
                                                                 PERCENT
                                                                TOTAL IRl
  FMTPROMOQPH4
  Ch| OaOPHYTO
  TEGULA SP.
  r-4CT
  MLVA
  CADOF
  OL'VCLLA S°.
   ATHERINOPS AFFINIS
     OCYEwaTa-CA0
     i| lit;  FD'jl, IS
                                             1.70

                                             1.10



                                            16.90

                                             5.00

                                             1.30

                                             1.10

                                             3.70
     TAXA  WITH FOF.Q.  OCCUR. LES1^ THSN S AND NUMflCAL ANO
        lnr. MOTH LE<=<: T^AN 1  ARF EXCLUDED FOP" THE TABLE
(»I|T  ^|OT  F9OM C A|_ C.IJL AT I OM OF  OI'/ERSITY INDICES)
                                       .00
                                       .00
                                       .00
  C JOTEPA
               -WFINFO  DIVERSITY
                                                                  PLOT
                 .00
                 .00
                 .00
                                   FOFO    NUM.    G"AV.   POEY     PERCENT
                                   OCC'IR   COMP.   COMP.   I.R.I.   TOTAL IRI
  r'apoFLL IDEA
  TEGULA AP .
  COPFPOD4
  4MPHIPO04
  ui.yA so.
                                                  6.70
                                                  5.30
    PPFY T/i»a WITH FDFO. OCCll=. LESS  THftN  R  ANO  NUMERICAL  ANO  GPAVluETRlC
    COMPOS I T tn.xi ^tOTH LESS TH4N  1  APF  PxCLliOEt'  FPOM  THE  TABLE  ANQ PLOT
    (-?IIT MOT FQOV CAI CULATION OF  DIVERSITY
       PERCENT OOMIMA'JCF  INDEX
                FlNFo DIVERSITY
                TNOtX
                                       .00
                                       .00
                                       .00
.as
.54
.00
.00
.00
                                          176

-------
Table B-12 (continued).
                                           G4Tfl
                                                                         PERCH
                    OF =>EL&TIVF Iv = ORTA-iCF  (I.O.I.) TAPl.F
                   M riLc  I^'NT. RrF  CT. c,TATlON SLSTA
           c i. if * I T F •
                                   F0£0   NUM.    GPAV.   PREY      PERCENT
                                   OCCUR  COMC.   COMP.   I.P.I.   TOTAL IPl
   1 <; I r- a r F A
  PLFOTYK •JiTa-CA0I
  r>i I rjrMT IF I n Fr,r
 "
 VT ! i MS FIMIL I ^
                                                    1.70

                                                    ? • 30

                                                    1 •hi
    CDCY TA«a  «ITH FPEO. OCCl"3.  LFS^ THIN  5 AND NUMERICAL AND  GRAVIMETPlC
    CO"PO=;ITIO'j  BOTH LE"5?  TH&N  1 ARF E'CLUPEO F^O"  THE TABLF AND PLOT
         NOT FPOM CALCULATION OF DIVERSITY
^ •-• A f i* '0* J - V ^ JK'PP DIV^D<-ITY .00
,. ,r1n FPFr) NUM.
P°EY ITFu OCCUR CO'
-------
Table  B-13.  Prey composition (frequency of  occurrence)  of shiner  perch
              in  upper Newport Bay,  California,  documented by Bane  and
              Robinson(1970).
                                                         (SHIMEP PEPCH
            INDEX  OF  RELATIVE IMPORTANCE (I.D.I.)  TAP.LF
              rpnM  fiLf  IOENT. PEE Si., STATIOM SL5TA
         PPFy
                               FOFQ   NUM.
                               OCCUR  COMP.
                                             GRAV.
                                             COMP.
     POFY
     I.P.I,
 PFPCENT
TOTAL IPI
POC
CHI. O°OOHYTA
fi * T E I C H T H V E S
     IDA
                               6=5.00
                               s^.on
                               si.no
       TAVA  WITH  FRFO. OCCIJP. LFS^ THAM 5 AiMf) NUMEPICAI.  ANH  GRAVIMETRIC
              ROTH  LF.<;S  THAN  i APE EXCLUDED FRO^ THE  TAHLE ANH PLOT
           FPOU ("/II_CUI_ATIOM OF DIVERSITY INDICES)
     PERCENT OOMlMANCF
              EINFR  DIVERSITY
              INOKX
                                        .00
                                        .00
                                        .00
.00
.00
.00
    .00
    .00
    .00
                                       178

-------
    DeMartini's (1969) analysis  of  the  comparative  feeding mechanism
morphology among the  surfperches indicated that bivalve molluscs, acmaeid
limpets, the shrimp,  Callianassa californiensis, isopods, gammarid amphipods,
gastropods (especially Thais  sp.)  and caridean crustaceans were important
prey of striped seaperch.

    Gnose's  (1967)  detailed study of striped seaperch in Yaquina Bay, Oregon,
indicated  that the  gammarid amphipods Amphithoe sp. and Anisogammarus sp.
comprised  the most  frequently eacCn prey item (Table B-14); mussels, barnacles,
chironomid insects  and  isopods (Idotea  sp.) were of secondary importance.
Alevizon  (1975) did an  extensive comparative study of striped seaperch feeding
 ecology in southern California.   He found that, on the basis of prey biomass,
 gammarids, polychaete annelids,  caridean crustaceans, caprellid amphipods and
 isopods were equally important at Santa Cruz Island (Table B-15a) while just
 gammarid and caprellid amphipods and caridean crustaceans  were of importance
 to fish collected along the Santa Barbara shoreline (Table B-15b).

 Pile Perch

     Pile perch collected in the vicinity of Guemes Island  and Cherry Point had
 consumed mostly valviferan isopods  (73.1 percent of the  total IRI), bivalves
  (10.5  percent), crabs (9.7 percent), and gammarid amphipods  (4.5  percent).
 Those  collected along the  Strait of Juan de Fuca, however, had consumed
  gastropods  (32.1%  of the total  IRI), pagurid crabs (Pagurus  beringanus, P_.
  granosimanus, P_. hirsutiusculus; 22.6%), gammarid amphipods,  (13.6%), brachyuran
  crabs, (11.5%),  sphaeromatid  isopods,  (2.6%), valviferan isopods  (5.9%) and
  tanaids,  (2.8%).

      Pagurid crabs, fissurellid  and acmaeid  limpets, the  bivalves  Pododesmus  sp.,
  Mytilus  sp. and  Hinnites sp.  and  the gastropod  Thais  sp.  were  listed  as
  principal prey of  pile  perch by DeMartini  (1969).

      Wares (1968)  has provided an extensive  analysis of  prey  composition,  using
  frequency of occurrence and  estimated  percent volume,  of pile  perch in  three
  regions of  Yaquina Bay, Oregon.   In general, barnacles  (Balanus  sp.), mussels
  (Mytilus sp.),  crabs (Cancer magister,  jC.  productus,  and C^.  oregonensis)   Other
  bivalves (Clinocardium sp.,  Prototheca sp.)  and shrimp (Upogebia sp.) were the
  prevalent food  organisms.   When examining variations  in prey composition by
  seasons, crabs,  particularly Cancer magister,  appeared predominantly in the
  spring while barnacles  and mussels comprised  the greatest proportion of  the
  diet during the  rest of the  year.   In  comparing the  three regions of the bay
  (upper, mid and  lower)  there were no consistent trends although  decapods
  appeared to be  most important in the diet of  pile  perch occupying the lower
  bay and bivalves other than mussels appeared  predominantly in fish from the
  upper bay.   In  general, however,  there was  no  change  in feeding  ecology,  as
  all prey were epibenthic or benthic organisms  which were "picked" from the
  bottom.

      Quast's (1968) documentation of the food habits  of pile perch in southern
  California kelp beds indicated  that decapod crustaceans, bivalve molluscs
  and ophiuroids  were the principal prey of all size classes while only
  gastropods entered significantly into  the prey spectra of fish 200-299 mm in
  length (Table B-16a & b).
                                      179

-------
  Table B-14.   Prey  composition (frequency  of occurrence)  of striped  seaperch
                 in Yaquina Bay,  Oregon, documented  by Gnose(1967).
PREDATOR
                                 LATE"ALIS
                                                             (STRIPED
                     F RELATIVE  IMPOPTANCE  (I.o.I.)  TARLE
                FPOM FILE IOENT. REF S7. STATION SLSTA
           POEY ITEM
                                  FPEO   NUM.   GRAV.  PREY
                                  OCCUR  COMP.  COMP.  I.R.I,
                                                 PERCENT
                                                TOTAL  IRI
  4MPHJTHOE S°.
  ANISOGAMMARUS SP.
  MYTH. us s°.
  RAI.AMIIS SP.
  OIPTFRA-CHI=>ONOMIPAE
         SP.
SP.
  MEMBRANIPORA SP.
  IIPOIFBIA SP.
  POL YCHAFTA
  lIMTOFNTIFIEO
  i ITTORIMA SP
  Cl.!»inCAOOIIIM
  PSTOACODA
  COLEOPTERA
 "AMPHISSA SP.
  ACMAFIOAE
  BUCCINUM SP.
 ~T~ACUMA "SP.
  SILIQUA AP.
                   ?o.so
                    9.40
                    7.70
                                    1.AO
                                    3.40
                                    T.40
                    ?.60

                    1 .70
                    1 .70

                     .90
                     .90
    CRFY TAXA WITH FOEQ. OCCll^. LESS THAN 5 ANfl NUMERICAL  ANO
    COMPOSITION POTH LESS THAN 1 ARF ExCLUOEO FPOM  THE  TARLE AND  PLOT
    (P-IlT NOT FROM CAICULATION OF DIVERSITY  INOICFS)
       PERCENT nnMlMlNCF INOE»
       SHANNON-WFINFP DIVERSITY
                            .00
                            .00
                            .no
.00
.00
.00
.00
.on
.00
                                          180

-------
Table B-15.   Prey composition (frequency of occurrence  and gravimetric
               compsoition)  of striped seaperch  at Santa  Cruz  (a) and  Santa
               Barbara (b),  California, documented by Alevizon(1975).
                                                           (STPIPF.O SEAPEPCH
           OF*  nF  PELATIVF
           FQ  STATION SLSTA
PPFY TTF«
r-ACTPODIQA
FCI-IMOIOEA
nFCApnoA-PLFOCYE"ATC
GALATHEIDAE
nFCAPr)r)A-PLFr>rvEMATA
ArwAFIDAE
Ppl_YDLAl~nPHOC4
PI V/ALVI A
PQTF K^T-oFc;
ppi-lnppinEA
FPFO MI«. G"AV. PPEY
OCCHP CriMp. COMP. I.P.I.
7^.00
?1 .<3(i
19. «0
1P.SO
1?.SO
1P.SO
Q.'iO
3. in
3. 10
•». 10
PERCENT
TOTAL IPI


                                            17.00
                                            ?1 .00
                                            10.00
                                            I'..00
                                            7.00
                                            IP.00
    DDF i  Till  -I-TH FOFQ. OCCi:». LF9q THAN S ANn NUMERICAL ANO GP»VI ME TRIC
    ri"0l<;i T ION HO™ Lt^S THAN ] AP>- F»CLHOFO Foo" THE TABLE AMP PLOT
    (SiiT  NOT FPOM CALdJi.ATIOM OF OWEP^ITY IN.DICES)
       <;"4NNOM-JF IKFP DlvFPSITY
       FVr .•j"'-IS  INOf»
                                       .no
                                       .oc
                                       .00
2.35
 .91
.00
.00
.00
 b.
                                 FPFO   NUM.
                                 OCCMP  CO"P.
                                            GPAV.   PPEY      PEPCENT
                                            COMP.   I.P.I.   TOTAL IPI
  PI Vi| VIA
  IIMHFNT IF IEP
 _PPhJliOOl,',F A
  C.A"MAoir,F4
  PLFOCYf'l,T 'iOT
                    f^. OCfli". L'lSe; THAN = AND *iUMEDIC*L "NO
                    LECS Tt-aN  1 4 u F. ^xTLUDEO FPOM  THE TABLE ANO  PLOT
                    riji.4TIOf OF DIvfPSITV I'.OirES)
                     DIVF.PSITY
                                       .00
                                       .no
                                       .00
                                                  .93
                   .00
                   .00
                   .00
                                          181

-------
 Table B-16.   Prey  composition (frequency of  occurrence) of  pile perch
                100-199mm (a)  and 200-299mm long (b)  in southern California
                kelp  beds documented by Quast(1968).
                     -  RHACOCHILUS  VACCA
                                                            (PILE PERCH
              INDEX  nF  RELATIVE  IMPORTANCE  (I.P.I.)  TAfiLE
                FPriM FIl.E  [DENT.  REF  ^9.  STATION'
a.
           PREY  ITE"
FPEQ   NUM.   GPAV.  PREY
OCCUR  COMP.  COMP.  I.R.I,
              PERCENT
             TOTAL  IRI
  DECAPODA-PLEOC'-'E'-'ATA
  BIv/ALVIA
  PLEOCYE'iATi-CAPIOfa
  RPYO?OA(ECTOPOOCTA)
 _IOrTEA REStCATA
 "GASTROPODA
               ALGAE
.b-
68.00
6«.on
19.00
10.00
  .00
  .00
  .50
  .sn
  .so
    PPFY TAXA WITH FPEQ.  OCCllW.  LES^  THAN  S  AND  NUMERICAL  ANO  GRAVIMETRIC
    COMPOSITION ROTH LF«;S THAM  1  ARE  EXCLUDED  FPOM  THE  TAflLE AND  PLOT
    (BUT NOT  FROM  CALCULATION of  DIVERSITY INDICES)
       PERCENT  DOMINANCE
       SHANNON-WEIMER DIVERSITY
       EVEN'MESS INDEX
         .00
         .00
         .00
.00
.00
.00
.00
.00
.00
PQF.Y ITP>
FPEO
OCCUR
NUM.
COMP.
GRAV.
COMP.
PPEY
I.R.I.
PERCENT
TOTAL IRI
  DECAPODA-PLEOCvEMATA
  PIVALVIA
  GASTROPODA
 "OPPIIIROIDEA
  8PY070A (ECTOPROCTA)
  UNIDENTIFIED ALGAE
  GAMMARIDEA
  PLEOCYEMATA-CAPIDFA
  IOOTEA
  CAPOIIOAE
  PAGURIOAF
47.00
44. 00
PS.OO
14.00
 6.00
 S.OO
'5.00
 5.00
  .50
  DONAX SP.

    PDFY TAXA vdTH FPFO.  OCCl'O.  LESQ  THAN  S  AND  NUMERICAL  AND  GPAvIMETRlC
              N  POTH LESS THAN  i  APE  EXCLUDF.D  FPOM  T^E  TABLE AND  PLOT
         NOT FPOM  CALCULATION OF  DIVERSITY INDICES)
       PERCENT  DOMINANCE.  INDEX
       SHANNON-WFIMED  DIVERSITY
       EVENNESS INDFX
         .00
         .00
         .00
.00
.00
.00
.00
.00
.00
                                         182

-------
Redtail Surfperch

     Redtail surfperch occurred abundantly only along the western  Strait of
Juan de Fuca (Twin Rivers and Hydaka Beach).  Over 70% of the  prey spectrum
was gammarid amphipods.  Mysids (including Neomysis awatschensis),
sphaeromatid isopods (including Gnorimosphaeroma oregonensis and Exosphaeroma
sp.), flabelliferan isopods, idoteid isopods  (including  Idotea resecata and I.
wosnesenski), and polychaetes were secondary  prey  (Fig.  B-61).

Snake Prickleback

     Stomach samples from snake prickleback from northern Puget Sound
characterized this species as principally a benthic feeder.  Bivalves
composed 48.7 percent of the total IRI, tanaids and polychaetes both accounted
for  21.2 percent, and gammarids amphipods contributed 7.3 percent.

     Oligochaetes accounted for 84.4 percent  of the total IRI; gammarid
amphipods,  11.0  percent: and polychaetes, 4.6 percent,  in stomachs from fish
collected  in  the mud/eelgrass habitat  (Fidalgo Bay) near Anacortes.

     Gammarid amphipods  and harpacticoid copepods  were  the major prey, based
 on numbers, and  polychaete annelids provided  significant contributions to the
 prey biomass  of  snake  prickleback collected in nearshore habitats  of Kodiak
 Island, Alaska  (Harris and Hartt 1977).

 High Cockscomb

      The most ubiquitous prickleback  in littoral habitats along the Strait,
 A. purpurescens  was  a  predominant member of the intertidal assemblage at
 Slip Point, Observatory  Point,  Twin Rivers, and Morse Creek.   Nemertean worms,
 gammarid amphipods,  and  polychaete annelids predominated in  the overall prey
 spectrum (Fig.  B-62).

      Based on frequency  of  occurrence, gammarid amphipods, polychaete annelids
 and gastropod molluscs predominated in the diet of high cockscomb examined from
 San Simeon, California by Barton  (1974)  (Table B-17).

      Based on frequency  of  occurrence, Peppar (1965)  assessed  the diet of
 high cockscomb  collected at  Second Narrows, Burrard Inlet, British Columbia.
 He found algae,  polychaete  annelids,  gammarid amphipods and  flatworms to be
 the most commonly occurring  food  items (Table B-18).

 Black Prickleback

      Almost three-quarters  of  the  total prey  IRI  (Fig.  B-63)  of black
 pricklebacks collected during  the MESA studies in  the Strait were gammarid
 amphipods.  Sabellarid and  polychaete worms and several algae were also
 important.

      Anomuran crabs,  gammarid  amphipods and the alga, Ulva  sp. were the
 major constituents,  based on frequency of occurrence, of the diet of black
 prickleback at  San Simeon,  California (Barton 1973; Table B-19).

                                      183

-------
                         100 r
                          80
                          60
                          40
                            INDEX OF RELRTIVE IMPORTANCE (I.R.I.) DlflGRRfl
                            FROM FILE IDENT.  76-78, STflTION PLSTfl

                                          8835600701   flMPHISTICUS RMODOTERUS
                                                        REDTfllL SURFPERCH
                                              flOJUSTED SflMPLE SIZE   142
                          20
                          20
                          40
                          60
                          80
                         100
                                                                              oc o» ffl
                                                                              C O T3
                                                                              ra >, c
                            I-H  , -H > QJ 4)
                            it  —in ^H en u
                            —'  O > nl C VI
                            U.  0. Z > « O
                                       50
                                                  100
                                                              150
                                                                          200
                                                                                      250
                        JTFM
CUhULflTIVE FREQUENCY OF OCCURRENCE
              &PAV,  PREY      PERCENT
              COUP.  I.P.I.   TOTAL  IRI
                                          OCCUR
                                                 NUM.
                                                 CO«(P.
                                          <•]. ?5  ?5.43  ?0.f>0

                                          ?
9.7
?5.3
'>.'•
_ 1.0
3.7
NUMERICAL AMD
                       rjow T ^!a^J
                      -«F [Mf p
                                 INOr
                .1?
               3.M
                .7<4
                                                                          ?.3S
                                                                           .51
Fig. B-61.   IRI  prey spectrum  of redtail  surfperch  in  shallow sublittoral
               habitats along  the Strait  of  Juan de Fuca.
                                             184

-------
                        INDEX OF RELRTIVE IMPORTHNCE 11.R.I.) OlfiORWI
                        FROM FILE IOENT. 76-78. STRHON flLSTH

                                   8842120402 - HNOPLHRCHUS PURPURESCENS
                                                 HIGH COCKSCOMB
                                       BOJUSTED SflMPLE SIZE = 273
PCT. COMPOSITION BY HEIGHT PCT. COMPOSITION BY flBUNOSNCE
SaasSoSSSSS







; 3
! 1 1
0 20 40 60 80
CUMULRTIVE FREQUENCY
FBEO NUM. GPAV.
PaFY 1TE" OCCUR COMP. COHP.
HFUEBTEA 34.07 ll.BT 25.50
_r.AM-iBl-)EA 3?.«.0 16. ?» 6.72
POLYCMA^TA 27.71 6.97 20.86
HAOPACT1COIOA 8.79 4.19 ,0«
_LiHIPFNTIFIED fc.9ft 1.99 7.IQ
PHnnoPHYTA «,.?3 1.90 ?.02
CM|_nonPHYT4 4.7>> 1.85 3.0.1
(!4«tDnPnDA 4.7A l.)4 4.3»,
TEOEPEILIOAE . 7.70 .71 5.8?
SAFrLLinAF J.R3 1.77 .81
CarsTACFA no 1i.77 ^3
Hni:AC«-4F .73 ,14 i.t,
rpA>ir,iriin4E .73 .n, 3.23
^r'n10*' A '?3 •'* z*:ji
nE«»A9ESTUCF.tF .37 . 4 ,.7;,
M»T»"TI« .37 .07 ,.03
POFY TAI4 WITH FOEQ. nCCllB. L1"1;1; TMAN 5 ANO NUMfo
	 raupocJjJriM BgTn_)_F5;<; JMJN, 1 ABF Fji^L'JUEJ) FPOM TM
SHAk4»jON*WF |r>l
EVENNESS INO
AL<~ULATM» ot" nivFasltv INDITES)
JARCF 1NDFJ .Ifj m 13
to OlvEBSITY 3.M 3.«j
t* .6? .66
«
U II 41
« 4 O
e v -o a
1:15. ss
SSSHSSll
* 3 >, ffl v. E "i
JC
I l t_{ 	 J^J |U ~"W
|
a .3. • i

5 S S J!S6333SS
100 120 140 160
OF OCCURRENCE
PBFY PERCENT
I.P.I. TOTAL IRI
1272.9 43.11
749. s 25.38
631.9 21.4(1
37. * 1.27
25.0 .85
22.9 .7R
23.' .79
76.7 .«9
27.7 .77
4.5 .15
14.1 .49
8.4 .28
36.0 1.2?
1.? .04
2.5 .08
I.A .04
.5 .07
.7 .02
.4 .01
1CAL ANO G°>VI*
-------
Table  B-17.  Prey composition  (frequency  of occurrence) of high cockscomb
              at San Simeon, California, documented  by Barton(1974).
                                                         
-------
Table B-18.   Prey composition  (frequency of  occurrence) of  high cockscomb
              at Second Narrows,  Burrard Inlet,  B.C., documented by Pepper
              (1965).
        r-*u?\?n'tn?. - S
                                                        (HIGH  COCKSCOMH
                 OF o^LATIVF  IMDOPTAMCF.  (I.P.I.) TABLE
                  FILE I OF NT.  &EF  «>0. STATION SLSTA
                                     MIM.
                                            G"AV.
                                            COMP.
                                                I.P.I.
                                                         PfPCENT
                                                        TOTAL  IRl
             A|
                               17.^0
                               1S.*0
 '.•F.PEIDAF
                                ?. 10
PDFY  TArA WITH FOF.O.  OCCUR. LE^^ TH4N  S  ANQ NUMERICAL AND GPAVIMETPlC
           BOTH LESS TH«N 1 4CF EXCLHOFO FOOM THE TARLF AND PLOT
           " CALCULATION OF DIVERSITY INDICES)
        MOT
      SM4MMON-WF
                    DIVERSITY
                                     .00
                                     .on
                                     .00
.00
.00
.00
.00
.00
.00
                                       187

-------
                           INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) OlflORWt
                           FROM FILE IOENT.  76-78. STRHON HLSTH

                                      8842121401   XIPMISTER RTROPURPUREUS
                                                  BLRCK PRICKLEBHCK
PCT. COMPOSITION BY WEIGHT PCT. COMPOSITION BY RBUNOflNCE
ococn.ik.ro N) *• en CD o
ooooooooooo



V






a •)
Rhodophy
Polychae
) 20 40 60 80 100
CUMULflTIVE FREQUENCY OF
FRE-)
PRF.Y ITEM OCCitP
r>AUMAR|DEA 67.XD
«APELL»PinAF 19.70
RHrOOPHYTA 14.39
POLYCHAETA 11.16
HAPOACTICOIOA «.11
IJNIOFMTIFIED fl.33
PAr.T-IACEAE 7.59
CHLOP1PHYTA 5.10
l.'LOTRICHiLES 3.79
nF_CM/\H£5T. 1 AT£AF JL.JL9
NE«FRTF.4 3.79
•5FLLOTA, ?.?7
SAPELLIOAF 1.5?
MOLLUSCS 1.5?
NATAMTIA 1.5?
AWr-MR4 " K5?
CH|.nPOPHYCEAE-7Yr,MFMATAUFS .76
STICHAFIOAE
PHOLIOIOAE
.76
.76
MJ-.
COMP.
47.01
16.3?
6.74
-SjJL-
1.94
4.6R
_a.i7^
3. ao
.57
_1J4
1.01
1.01
1.01
.57
1.14
.46
.?3
.34
.73
!s7
.11
.11
.11
• n v
q fl 4t -O ig IQ
T4 T3 « -H ^ tO -0
•S'u-Sco-j.cSc.S
UM^_-:Z=.
	 . ) __- ' ^^* 1
LJ^^
01
v n
m v ti
•a T» M u
O « W U -0 ^4

1 Il^slll
120 140 160 180
OCCURRENCE
GPAV. PREY PERCENT
COMP. I.R.I. TOTAL 1RI
9.04 1709. p 73.39
.66 134.6 7.65
10.60
10. ao
	 .SLl 	
5.10
17.31
_a.o7 	
3.10
1.10
7.0« 	
.70
.05
.00
1.83
.04
1.31
a. 93
2.41
4.67
1.69
1.03
l.SH
5.74
a. 06
a49.5 5.70
145.0 3.33
34.4 .79
58.7 1.34
166. «. J.8I
33.4 .76
6.1 .15
6.5 .15
a. 5 .06
7.1 .05
5.5 .17
l.n .04
7.7 .06
4.0. .11
4.7 .10
7.4 .17
7.9 .07
1.7 .03
1.5 .03
4.4 .10
1.6 .04
PREV T»xA WITH FDEO. OCCl'P. LF-5S THAN 5 AND MJMEPICAL. Ann GRAVIMETRIC
COMPOSITION POTH LESS THAN 1 ARE EXCLUDED FROM THE T»8LE ANO PLOT
(O.I1T NOT FROM CA1 CAII ATIO'! OF DIVERSITY INDITES)
PERCENT On**TK|ANCF INPE1*
.77
.OH
4.74
.55
1.67
Fig.  B-63.   IRI  prey  spectrum  of black  prickleback  in littoral habitats
               along the Strait of Juan de Fuca.

                                           188

-------
Table B-19.   Prey  composition  (frequency of occurrence)  of black prickleback
               at  San Simeon, California, documented by Barton(1973).
                                                           "(BLACK  PPICKLER4CK   )
             I'lflF' OF DEL4TIVF I'.'POPTfMCF  (I.R.I.)  TARLF
               FRIIM FIlE IH^NT. 9FF SS. 9TATIOM SLSTA
          PPFY ITFw
FPFO
OCC'IP
                                        MIJM.
                                        CO^P.
     GP.AV.
     COMP.
            I.P.I.
 PFPCENT
TOTAL IRI
 HLVA SP.
 R I r- 1 P T T »! A
_SMITHORA SP.
                                  P. 00
                                  <, . o n
  PETROLISTHES SP.

    CPFY  TivA  WITH  FPFO.  OTi|3.  LFSS  THAN  S  AND  NUMERICAL AMD GRAVIMETRIC
    <-T>PnciITl'Vi  ROTH  LE<^<^ T~,AN  1  A = F  FxCLUDEO  FoQM  THE TARLF. ANP PLOT
    (HUT  NOT  F30M  CAlCijLATlON Of  OIvtPSlTY INDITES)
             T  OOWI'I\NCF  I'lOfx
       <;HAMNI)N-WFIMFP  CIVEPMTY
      "
.00    .00
.00 _  .00
.00  " .00
                                  .00
                                  .00
                                  .00
                                          189

-------
Rock Prickleback

     Although not as prevalent in intertidal collections as the black
prickleback, the rock prickleback occurred at most of the intertidal sites
along the Strait of Juan de Fuca and were common at Slip Point and Twin
Rivers.  Numerically, the IRI prey spectrum of rock pricklebacks  (Fig. B-64)
was more diverse than that of black pricklebacks.  Gammarid amphipods were
the predominant prey organism but composed less than half of the  total IRI.
Instead, algae made a greater contribution; Rhodophyta, Chlorophyta,
Phaephyta, Ultrichales, Bangiales, and Gigartinacea combining for 44.8% of
the total IRI.

     Algae, principally Smithora sp., Ulva sp. and Plocamium sp., were also
the pricnipal diet constituents, based on frequency of occurrence, of rock
prickleback collected at San Simeon, California (Barton 1974); gammarid
amphipods were the only prey animals (Table B-20).

Ribbon Prickleback

     A ribbon prickleback from beach seine collection in northern Puget Sound
had 14 gammarid amphipods in its stomach.

Penpoint Gunnel

     Penpoint gunnel from gravel pocket beaches in northern Puget Sound had
consumed oniscoidean isopods and gammarid amphipods (43.8% and 43.4% of the
total IRI, respectively), valviferan isopods (9.3%).  shrimp (1.2%),  and
several other epibenthic crustacean taxa.

     Penpoint gunnels also appeared frequently in both beach seine and tidepool
collections along the Strait of Juan de Fuca and were most numerous at Twin
Rivers and Beckett Point.  Epibenthic crustaceans—gammarid amphipods,
harpacticoid copepods, mysids, and valviferan isopods (Idotea sp.)—were the
most important prey in the spectrum (Fig. B-65), composing 88% of the total
IRI.  Benthic polychaete and nemertean worms composed 7.4%.

Crescent Gunnel

     As with the penpoint gunnel, crescent gunnel in northern Puget Sound
preyed upon epibenthic and benthic organisms.  Gammarid amphipods were the
principal prey item, totaling 78.4 percent of total IRI.  Harpacticoid copepods
were less important, accounting for 10.2 percent.  Tanaids made up 4.7
percent; polychaetes, 2.5 percent; valviferan isopods, 1.1 percent; and a
variety of epibenthic crustaceans contributed less than 1.0 percent of the
total IRI.  The diet composition of crescent gunnels from the eastern
shoreline was quite similar to those in FRl's San Juan Island collections.
Gammarid amphipods predominated (85.4 percent of total IRI), while polychaetes
(8.8 percent), crab larvae (2.7 percent), and hyperiid and caprellid amphipods
(each at 1.1 percent) were the less important prey.
                                     190

-------
                      100
                         INDEX OF RELRTIVE IflPORTflNCE (I.R.I.] DIRGRflM
                         FROM FILE IDENT.  76-78. STHTION RLSTR
                                         8842121402   XIPHISTER flUCOSUS
                                                      ROCK PRICKLEBRCK
                                            RDJUSTED SflMPLE SIZE    87
60
£
§
£ 40
8
§ 20
(_)


§ 20
UJ
3
&
40
O
1 60
^ 80
o
Q-
POFY TTFf
CP^ATROMAT j n &r
CHpnriPHYC^ A P - -4. fl *• I T-
WIDENTIFIED"
TMcrrra
n
j:
(.
c
5
11
i r-h-i \
— -KTT
	 	 ' — — T MJ

0}
01
,H
rt
00
c

, s: aixiai^Exci
•H ^ O O(XtO£OX'H4
M a. -^ m o jr a. ^ D. 4J
n o P . -H >-. a o at o t-
£ O O -HC^Hr-Hrt^O&O
(8 ,C ^H £3(5^0X10-^1^^
0 20 40 60 80 100 120
CUMULRTIVE FREQUENCY OF OCCURRENCE
FPFO Nil". GRAV. PPET PERCENT*
ncC'io COUP. COMP. I.P.I. TOTAL IRI
36.7% ?fl.?l 3.09 1151.3 43.01
	 	 IP. 10 10.^3 9.7B _ 379.) 14.16
]7.?<, S.06 4.19 ?11.? 7.B9
]3.«-4 5.5i, 4.e>0 ]?8.? 4.79
	 ._ 10.34 11. 5.1. _ 1?.60 __,?50.2 	 9.35__
9.?0 If.O" 7.«3 164.6 6.15
fl."5 2.0? 15.2? 13S.7 5.1«
	 f-.9Q 3.53_ 7.69 _ 77.3.-.. 2.B9_
6.9(1 ?.02 .5? 17. c .65
£LE<; "=.75 ^.04 3.B5 51.1 1-91
'.30 1.01 .00 2.3 .09
1.15 .25 4.54 5.5 .21
	 1.15 .25 1.01 1.4 .05
1.15 1.01 1.56 2. .09
l.lc .25 9.56 " 11.3" .42
13
ra o
^ *J D.
U O
-, IK •**
£6
\l
«
RU
\
o» \
10 01
01 10
U 01
3 <0 U
H C (0
3 O ^
H oi j:
-i oo a.
J O O
H E rH
S. « tH
- 4J X
S££
40 160
	
	
        _  PPFY  TAx«  .ITH FDEQ. OCCUR. LF-;"; THAN 5 AND NI)«EPICAL  AND  OPAvI»ETR!C
           COwPn-ITlON HTTH LE^"; THJ-J J  APF >•_ xCL'lOt'D F'of)" THE  TABLE  ANn PLOT
           (BUT  Nl^T FCA.. rt| ClJ'_AT I^V OF  0
                      nnM!»jn;iCE" INOfx
                     -WF T^lK c? DIVERSITY
 .13
3.55
 .73
 .09
3.74
 .77
                                                                        2.79
Fig.  B-64.   IRI  prey spectrum of rock prickleback  in  littoral habitats  along
               Strait  of Juan  de Fuca.
                                            191

-------
  Table B-20,  Prey composition (frequency of  occurrence)  of rock prickleback
                at  San Simeon,  California, documented by  Barton(197A).
PPFOATOP
                                'iijcosus
                          (ROCK PRICKLF8AC"
                    OF  RELATIVE  IMPORTANCE  d.p.i.) TABLE
                FPOM FIlE  IOENT.  OFF S5. STATIOM 5LSTA
           PQFY I
                                 FREQ
                                        NUM.
                                        COMP.
              GPAV.
              COMP.
     PPEY
     I.P.I.
 PERCENT
TOTAL IRI
  SMITHORA SP.
  PLOCAMIUM SP.
l?.no
 4.00
 4.00
 4 . il 1
 4.00
  RHODOGLOSSUM SP.
 _CLADOPHORA SP.
  PHVLLOSP60IX  S
    DPFY TAxA  ^ITH  FPFO.  OCCllrt. LF^S THAN S ANO NUMERICAL AMO
    COMPOSITION  BOTH  I ESS THAN  I ARF EXCLUDED FPOM THF TARLE ANO PLOT
    (RUT NOT F30« CA| CUL*TION OF DIVERSITY INDICES)
       PERCENT  OnMInAMCP  INDEX
       SH4NMOM-WFIMFp  DTVFPSITY
                INOF*
         .on
         .no
         .no
.on
.00
.00
    .00
    .00
    .00
                                          192

-------
                   CO
                   •z.
                   o
                   8
                   Q_
                          INDEX OF RELRTIVE IMPORTRNCE (I.R.I.) OIBGRRH
                          FROM FILE IDENT.  76-78. STflTION RLSTfi
                                        8842130101   RPOOICHTHYS FLRVIDUS
                                                      PENPOINT GUNNEL
                       100                  ROJUSTED SHMPLE SIZE   88
                        80
                        60
                        40
                   Q.
                   g
                   (_)
                        20
                        20
                        40
                        60
                        80
                       100
                                                                     :rflk-i-uj
I
                                                                            eooait.
                                 20
                                        40
                                               60
                                                       80
                                                              100
                                                                     120
                                                                            140
                                                                                   160
                                         CUMULRTIVE FREQUENCY OF OCCURRENCE
                                       FPEO   NU»'.   GCAV.  PPEY     PERCENT
                                       OCCUP  COUP.  COMP.  I.R.I.   TOTAL  IPI
H4CPACT1C01DA.. .__ 	 	
MYCIOACEA
Wf.l. V If EPA
comrpouAT IHAF
K 4TAMT I A
POLYCHAETA
CPfY TAX' WITH foEO. OCCllrf.
	 rn"on<; ! T Io>, ROTH Lf.5 THAN 1
(&'JT '.TIT CBOM CALCULATION o>"
	 PfPCFNT nOMlMflhjCF INOEX
SM AfyMO'J-WF I MTP h I V^RS 1 T~Y~
51.14 ?S.96 13.22 'n03.7
13.64^28.75-. .23— -395.7
I'.SO 10.96 19.54 381.3
1?.SO 14.33 21.3« 446.4
1?.50 	 2.02_ie.92 	 761.7
ti.OQ ?.8» .9"? 35.1
7.95 ?.?) .73 ?3.4
	 7.95 -.2.02- 3.55_ 44.3
6.S2 S.19 4.75 67.1
4.55 .87 ?.71 16.?
.._ 4.55 .87 3.78 ?1.1
4.55 .67 ?.31 13.^
3.41 .87 3.44 14.7
?.?7 	 .19 3.96_ 	 9.4
LF<;<; THAN 5 AMP NUMERICAL AMO
ARF FxCLIIOEO FDpM THE TARLE
DIVERSITY INOICFS)
.19 .15
?.9? 3.14"
.64 .68
53.60
	 10.57 	
10*20
11-94
	 7.00 	
.94
.63
	 1.13 	
1.81
.43
.56
• 36
.39
	 .25. 	
GPAvIMETPIC
ANn DLOT
.33
?.26~
.49
Fig.  B-65.   IRI  prey  spectrum of  penpoint  gunnel  in Strait of  Juan de Fuca.
                                           193

-------
      In the Strait of Juan de Fuca, crescent gunnels were common in inter-
tidal collections at Twin Rivers, Morse Creek, and North Beach and beach
seine collections at Twin Rivers which all contained crescent gunnel.  Except
for dipteran insects, all the major prey organisms were epibenthic crustaceans.
Gammarid amphipods composed over 80% of the IRIj isopods (including sphaeromatid,
idoteid, and valviferan species), 7.4%, munnid crabs, 3.4%- and harpacticoid
copepods, 2.7% (Fig. B-66).

Saddleback Gunnel

     Saddleback gunnels occurred in the same habitats in northern Puget Sound
and had much of the same prey composition as the more common crescent gunnel.
Amphipods were also the principal prey (49.7 percent of the total IRI) but
oniscoidean isopods (not found in 7_. laeta stomachs) ranked a close second
(32.9 percent) in importance.  Polychaete annelids (7.4 percent), harpacticoid
copepods (5.6 percent), cumaceans (2.0 percent), and valviferan isopods (2.0
percent) were also in these stomachs.

     In the Strait of Juan de Fuca, gammarid amphipods were important to the
diet (75% of the total IRI); secondary prey were sabellid worms, mysids, and
juvenile hippolytid shrimp (Fig. B-67).

Pacific Sand Lance

     Over both northern Puget Sound and the Strait of Juan de Fuca, Pacific
sand lance were similar in occurrence and distribution to juvenile Pacific
herring.  In the former region, beach seine catches of Pacific sand lance
were most frequent and numerous at Eagle Cove, and townet catches, at Point
George and Westcott Bay.  Along the Strait, both beach seine and townet
collections at Dungeness Spit and Kydaka Beach yielded high numbers.

     Pacific sand lance were basically pelagic feeders with an even more
specialized prey spectrum than juvenile Pacific herring.  In northern Puget
Sound calanoid copepods composed 88.5 percent of total IRI and gammarid
amphipods 9.0 percent.  The prey spectrum from the Strait of Juan de Fuca
was even more oriented toward calanoid copepods (Fig. B-68).

     Over 75% of the prey biomass of Pacific sand lance collected in nearshore
habitats at Kodiak Island, Alaska, consisted of calanoid copepods, supplemented
by crustacean zoea and nauplii and larvaceans (Harris and Hartt 1977).

     Calanoid copepods are, in fact, the almost universal prey organism of all
other sand lance species (Ammodytes americanus, A_. dubius, A_. marinus. A_.
tobianus and A^ personatus) occurring in north temperate waters of the Atlantic
and Pacific Oceans (Meyer et al. 1979; Reay 1970; Scott 1973; Sekiguchi 1977;
Senta 1965).

Speckled Sanddab

     Speckled sanddabs from the Strait of Juan de Fuca (Beckett Point) con-
tained principally polychaetes and gammarid amphipods.  Other less commonly
utilized prey included bivalves and their siphons, calanoid copepods, shrimp,

                                     194

-------
                          INDEX OF RELRTIVE IMPORTHNCE (I.R.I.I  Q1RGRRH
                          FROM FILE IOENT.  76-78. STRT10N  RLSTR

                                            8842130205   PHOLIS LRETR
                                                      CRESCENT GUNNEL
                                            HOJUSTEO SHMPLE SIZE = 97
                        100
                                20
40   60    80   100   120   140   160

   CUMULRTIVE  FREQUENCY OF OCCURRENCE
                                                                            180
                   PREY ITEM
                                       ~FREONUM.   GR»V.  PPEY     PERCENT
                                        OCCUR  COMP.  COUP.  I.R.I.  TOTAL  IRI
SPHAFRO'<^ _•; AND NUMERICAL Axn Go Ay T"E TB 1 C
             COMPOSITION POTM Lf.sc THAN 1 ABF f«CLnl)Eri FRO" THE  TAULF "iNO^OLOT
                              .
             (BUT HOT FPOM  CALCULATION OF DIvFH«;ITY  INDICES)
                PERCENT DOMINANCE INOEI
                SHA»,NON-'
-------
                                INDEX OF RELflTIVE IMPORTANCE (I.R.I.) DlflORflM

                                FROM FILE IDENT.  76-78. STRTION flLSTfl
                                                 8842130206,  PHOLIS ORNfiTfl

                                                           SflODLEBflCK GUNNEL
                        a
                             lOOr
                             80
                             60
8
a.

§
u
00

z
o
                             40
                             20
                             20
                             +0
                             60
                             80
                            100
                                                                  (U
                                                              LI _ )  dj
                                                              X\  TJ
                                                        t-> O


                                                        IS
                                                        t-> O t) -H O >, O

                                                        o) D. 
-------
                       s
                       o
                       in
                       o
                       a.
                       £
                            100
                             80
                             60
                             40
             20
                             20
                            •40
             SO
                             80
                               INDEX OF RELRTIVE IHPORTRNCE (I.R.I.) OIRORHH
                               FROM FILE IDENT.  76-78. STflTlON flLSTR

                                              8845010101 - RMtlODYTES HEXRPTERUS
                                                           PRCIFIC SRNO URNCE
                                                 flDJUSTED SRhPLE SIZE =  43
                            100
                                            10     15     20     25
                                                                      30
                                                                             35
                                                                                    40
                                                                                           45
                      PPEY ITEM
                                              CUMULfiTIVE FREQUENCY OF OCCURRENCE

                                             F4EO   NUM.   GRAV.   PBEY      PERCENT
                                             OCCUR  COMP.  COMP.   I.P.I.   TOTAL  IRI
             CALANOIDA
                                             30.53  90.21
                                              2.33	.02
                                                           T7.06
                                                             97.3?
                                                    145.9	?.63.
               P°FY TAXA WITH FBFO. OCCUR. LE";1; THAN ^ ANO NUMERICAL  AND  GOAVIMETP.IC
              _CBMPQ£lllfl?j JOIH LE«,S JLMAN 1 . ARE. EXCLUOEO FROM. THE. TABLE. AND_PLO.T_	
               (PUT NOT FSOM CALCULATION OF OIVEPSITY INDICES)
                  PFOCFNT. OOMlKi
                  SHf^^lN^N-wF INF
                                 DIVERSITY            .09     .99               .18
                                                      .03     .3S               .Oft
Fig. B-68.
IRI prey  spectrum of  Pacific sand  lance  in Strait of Juan de  Fuca.
                                               197

-------
amomuran crabs, one isopod, and algae.

Pacific Sanddab

     Winter beach seine collections at Dungeness Spit and Beckett Point
produced Pacific sanddabs.  Gammarid amphipods were common prey to three,
contributing 68.4 percent of the prey and 3.8 percent of the prey biomass.
Mysids (Neomysis sp.), shrimp (61.2 percent total biomass), and polychaetes
were the other prey items.

Rock Sole

     Adult rock sole were caught, though not in abundance, in the cobble
and sand/eelgrass habitats of southwestern San Juan Island.  The prey
spectrum from northern Puget Sound was extremely broad.  Prey items, in
descending order of importance, were flabelliferan isopods, gammarid amphipods,
bivalve siphons, polychaetes, cumaceans, bivalves, brachyuran crabs, and fish.
Rock sole from collections at two Guemes Island sites and Cherry Point along
the eastern shoreline had consumed principally gammarid amphipods (88.9 percent
of total IRI); crabs (4.4 percent), bivalves (3.0 percent), and polychaetes
(2.4 percent) were only supplemental organisms.

     Rock sole in the Strait of Juan de Fuca were benthic feeders, preying
principally on polychaete annelids (75% of the total IRI) (Fig. B-69).   Epi-
benthic gammarid amphipods (12.6%) and tanaids supplemented the diet.

     On the basis of biomass, fish (Pacific sand lance) were more prominent
in the diet of rock sole collected in the nearshore environs of Kodiak  Island,
Alaska, although gammarid amphipods, polychaete annelids and bivalves were
important numerically (Harris and Hartt 1977).

English Sole (Juveniles)

     Juvenile English sole were the most frequently caught species of the
nearshore demersal assemblages in northern Puget Sound, and were most
prevalent at Westcott Bay (mud/eelgrass) and Eagle Cove (sand/eelgrass).
Overall, cumaceans dominated the prey spectrum with 74.8 percent of the total
IRI (Fig. B-70).  Gammarid amphipods (11.7 percent), polychaete annelids
(8.8 percent), tanaids (1.1 percent), crabs (1.0 percent), and bivalves (0.3
percent) were of secondary importance.  Juvenile English sole were also common
in sand/eelgrass and mud/eelgrass habitats along the eastern shoreline.  All
the prey taxa were similar; diet composition in this region was dominated
by gammarid amphipods (87.7 percent of total IRI), with cumaceans (8.4 percent),
polychaetes (2.0 percent), and bivalves (1.4 percent) providing lower inputs
(Fig. B-71).

     Juvenile English sole were the most widely and evenly distributed species
at the eight beach seine sites sampled by the MESA program in the Strait of
Juan de Fuca.  The overall prey spectrum (Fig- B-72) was equally divided among
benthic glycerid and gonaid polychaetes, bivalves (including Clinocardium
nuttalli), epibenthic gammarid amphipods, cumaceans, harpacticoids, tanaids,
and mysids (including Archaeomysis grebnitzki).

                                    198

-------
                              INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) OIHDRRn
                              FROM FILE IDENT.  76-78. STRTION RLSTH

                                          8857040801 - LEPIOOPSETTH BIL1NEHTR
                                                          ROCK SOLE
                                              fiOJUSTEQ SRMPLE SIZE   66
COMPOS
BY HEIGH
                                            100
                                                    150
                                                           200
                                                                   250
                                                                          300
                                                                                  3SO
                                           CUMULHTIVE FREQUENCY OF OCCURRENCE
PPEY ITEM
POLYCMAETA
TANAIOACFA
C.AuvARIO'E
PIVALVIA
UNIDENTIFIED
EUCAPIDA-DECAPOUA
CAPPFLLIPAE
GASTDOPOOA
NE"ATOOA
NATAHT1 A
CADOI IDAF '
ATFI FCYCL IOAF
CANCPJDAF
PIIIC'II.ANIC'E
PHPL 10IOAF.
iMunDYTIDAE
ANTHIIPIDAE
FHEO
OCCUB
84. «5
54.55
54.55
50.00
Pfl.79
15.15
9.09
9jfl9
6.06
4.55
4.55
.5?
.52
:i .52
.5?
.52
.52
Nil". GBAV.
COMP. COMP.
56.91 29.36
6.6A .20
17.94 2.06
5.11 7.37
4.41 .41
i.eo 1.01
.46 .94
.21 .01
l.?S .00
.73 2.15
.23 17.49
.12 1.77
.06 8.31
.12 1.67
.06 11.51
.12 7.13
.17 1.4?
.12 1.30
PBEY
I.B.I.
7119.5
375.0
1091.2
624.1
138. B
42.6
12.7
7.1
1.5
5.1
10. »
80.6
2.9
13. t
2.7
17.5
3.1
11.1
2.4
7.1
PERCENT
rOTAL IRI
74.84
3.83
6.39
1>42
.44
.13
.on
.02
.06
.11
.8?
.03
.11
.03
.18
.03
.11
.02
.02
                POFY TA»A »1TM FOFB._QCCL|P. LESS THAN S A^ll JUUMEPlCJLL__A»(D_0a^v 1 METRIC
                COMPOSITION BOTH LESS  T1«N 1  APF FXCLIIOED FPO» TM£ TABLE  AND PLOT
                («I|T MOT FROM CALCULATION OF  DIVERSITY INDICES)
PFOCENT OOHlMtNCF IMOF«
SHANNON-UFINFP DIVERSITY
FuFNNFSS INDE«
.37
2.21
.15
3.36
.51
1.39
Fig.  B-69.   IRI  prey  spectrum of rock sole  in  shallow sublittoral
                habitats  along  Strait  of  Juan de Fuca.
                                              199

-------
                             INDEX OF RELflTIVE IttPORTHNCE (I.R.I.) OIRGRflfl
                             FROM FILE IDENT. N PGSO. STflTION flLSTfl
                                         PREDflTOR   8857041301   PRROPHRYS  VETULUS
                                    (ENGLISH SOLE       )   PDJUSTED SflHPLE SIZE -  46
                          100
                           80
                           60
                           40
                           20
                     H     20
                           40
                     s
                     u
                          60
                          80
                          100
                                     50
                                               100
                                                         150
                                                                  200
                                                                            250
                                                                                      300
                                           CUMULflTIVE FREQUENCY OF OCCURRENCE
PPEY
ITFM
rpfrj
OCCliP
NUM.
COUP.
G»AV.
COMP.
PPEY
I.O.I.
PtPClNT
TOTAL IP!
             r- 4u
             POI YCHAF.TA
             TAKA1DACEA
            HAOP4CTICOIOA
             OXYRHYNCHA
f.7.00
63.00
•i^.OO
35.00
I3;00
13.00
11.00
9.00
?;no
- 49. BO
?7.ei
	 8.74
?.40
I^I^i —
.41
	 4; 17 —
1 .44
	 .9?
33.07
7.98
7.0?
.15
— .66
.00
.04
?.61
37.05
~?37«~
  5.3
 46.3
 36.5
 75.9
                         6?.00
                         ?S.?0
                          9.70
                          1.00
                            .30
                            .10
                         ~  .50
                            .40
                         —  .10
              -pOFY~TAr« -WlTH-TDfO; OCCIIRT Lfet^" THAfJ"?  AND" NUMEPICAC ~*ND 'GP*V IMETR 1C
              COMPOSITION BOTH LESS THAN 1  APE  EXCLUDED  EOO"  THE TABLE AND PLOT
               (RUT NOT rpf)M CA( CULATION OT  HIVEPSITY  INDICES)	
                 PERCENT
                                DIVERSITY
------ .34'-   ,2f>
   1.90   1.8?
---- .60-   .61
                                                                           1.44
                                                                            .45"
Fig. B-70.   IRI prey  spectrum of  juvenile  English sole in shallow sublittoral
               habitats  of northern  Puget Sound.
                                              200

-------
                           INDEX OF RELflTIVE IHPORTfiNCE (I.R.I.) OIRGRRM
                           FROM FILE IOENT. UH  BS. STRTION  flLL

                                       PREOflTOR  8857041301 - PRROPHRYS VETULUS
                                   (ENGLISH SOLE       )   MJUSTEO SflHPLE SIZE = 120
                         100

                    8
                    |    80


                    I    -
                    §
                    E    40

                    |    20
                          20
                     £
                     £
                     §    40
                     »-•
                     I—
                          60
                         100
                                                                             =a-
                                                                          •o 1-1 ex M o
                                       50         100         150         200

                                           CUMULATIVE FREQUENCY OF OCCURRENCE
                                                                                     250
                     PREY ITEM
 FPEO   NUM.   GPAV.  PPEY
_OCCUR_ .COHP.  CQMP. __ l.R..J_.
                                                                        PEPCENT
                                                                       IPJAL .IP_I_
GAMMARIDEA
ClJMACEA
POLYCHAETA
RIVALVIA
ISOPODA
PLEOCYEMATA-CARIOFA
Jf_ANAIDACEA
CNIOARIA
CIRPIPEDIA
AMPHARETIOAE
NE-ATOOA
92.00
42.00
2S.OO
17.00
p. no
».no
a. oo
A. 00
ft. 00
A. 00
fl.OO
73.06
17.96
1.63
1.2?
2.45
.41
.41
.41
.41
1.63
.41
60.33
9.92
6.26
9.92
T.31
1.65
.00
1.65
1.65
3.31
.00
12271.9
1171.0
247.2
189. 4
46.1
16. S
3.3
2.1
2.1
39.5
3.3
87.70
8.40
1.80
1.40
.30
.10
.00
.00
.00
.30
.00
              PREY_TA«A WITH FPEO. OCCll». LESS_THAN_5_ANn NUMERICAL, AMn_GPAVIMETRlC_
              COMPOSITION BOTH LESS  THAN 1 ARE EXCLUDED FROM THE  TABLE  AND PLOT
              (BUT NOT FROM CALCULATION OF DIVERSITY INDICES)
                 PEPCFNT DOMINANCE  INDEX
                 SHANNON-WFINEP  DIVERSITY
                 EVENNESS INDEX
          .57
                "73<»
         1.3*   2.0?
          .39    .64
.78
.72
.26
Fig.  B  71.   IRI  prey  spectrum of juvenile  English  sole  in  shallow  sublittoral
               habitats  along eastern shoreline of northern Puget  Sound.
                                              201

-------
                               INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) DIflGRflM
                               FROM FILE IDENT.  76-78. STflTION flLSTfl

                                               8857041301  - PflROPHRYS VETULUS
                                                             ENGLISH SOLE
100
LJ
o
i so
1
• «
03
§
£ 40

8
~

8 20

*-:
LJ
fl
U
(_
r
2 20
UJ
3
CD
2 4°
»—
| BO
8

.-1 80
u
a.

inn
iooc
nuuuon_u onnri_i_ oitc. = ooo





o>
0) (0
0) (0 ^
CO ••-< 4-« O
flj C >N >,
4-1 O <-H U
1-. OC O 0)
E > (Q E E ci-tn--i^-i*-t
ft. u cj: OPQHSZXXZ
i i i i
) 50 100 150 200 2E
                           ITFM
                                             CUflULflTIVE FREQUENCY OF OCCURRENCE
                                            FPFO   NUM.    GPAV.  DPEY     PEPCENT
                                            OCCUR  COMP.   COMP.  I.P.I.  TOTAL IPI
Pfl YfHAFTA
CI.PMACEA 	 	
r-AUMJPlflf A
GAVMAPTQ4E;
PI»A| v! ^
TAVAIDACEA
HAPPACTICOIOA
MY«in*<"EA 	
Hni.OTHiioniDFA
HOLfiTHiiPnlOFA AsPinnCHionTACf A
h'AT'tMTIA
NFUFPTEA
TQANGOfJIDAE
HIPUOLYTJDAE
ATFLFCYCLIDAE
44 S4
4ft. 71 	
39.89
1°.95
]O.40_.
19.40
1ft. 94
in. 38
3.^1
3.83
3 . ?8
?.4ft
1 .09
.27 _
.'7
13.08
PO.S3
17.08
ft. 46
2.45.
6.10
?ft.79
1 . *9_
.59
.47
1.09
.13
.03
.02
.0?
28.68
13.51
17.03
9.44
_3.60
2.05
1.00
6.4ft
3.31
3.37
.39
1.57
1.39
1.44
1.07
IflftO.n
1385.7
1 360. ft
317.1
117.4
158.?
470.8
87.7
14.9
14.7
4.1
4.?
1.5
	 .4
.3
32.02
23.86
?3.4?
5.4ft
_2>02
2.7?
8.10
-- 1.51. . .._
.26
.25
.08
.07
.03
	 .01
.01
              J?°.EY_ TA^A WITH  F.PEO. OCCUR. LE5S_JHA_N .5  AND_ NUMERICAL .AN
               roMpntjiTioM '•OTH  LE^S THAN i ARE EXCLUDED  FROM  THE TABLE AND PLOT
               (R|lT MOT FPD" C«l CUI.ATIOtJ OF DIVERSITY  INDITFS)
                  PERCENT OOHlNAr4CE  INDE*
                                 DIVERSITY
 .17    .15
2.99   3.4?
 .54 __   .61.
       .23
      2.47
	..44
Fig.  B-72.   IRI  prey  spectrum of  juvenile English sole in  shallow  sublittoral
               habitats  along Strait of  Juan de  Fuca.
                                              202

-------
Starry Flounder

     Adult starry flounder were  frequently captured in beach seine collections
in northern Puget Sound  (Eagle Cove  and South Beach)  from July through
November though never in large numbers.  The most frequently consumed prey
organisms were flabelliferan  isopods which accounted for 58.9 percent of
the total IRI.  Fish  (18.2 percent)  were second in importance, followed by
gammarid amphipods  (8.2  percent),  epicaridan isopods (4.5 percent),
polychaetes  (3.7 percent), gastropods (3.3 percent),  and turbellarians (1.2
percent).  The amphipods were primarily Atylus sp. but also Eusiroides sp.
and Amphithoe so.   All prey,  except  perhaps the fish, were epibenthic or
benthic organisms.

      Juvenile starry  flounder also appeared in beach seine collections along
 the  eastern  shoreline, principally in sand/eelgrass (Birch Bay, East Guemcs
 Island) and  mud/eelgrass (Padilla  Bay,  Drayton Harbor) habitats in August
 through December.   While isopods (primarily valviferan) were still important
 (30.2 percent of  total IRI)  in  these samples, gammarid amphipods (33.4 percent),
 barnacles  (16.7 percent),  and oligochaetes (11.8 percent) were much more
 prevalent  in the  diets of  starry flounders on the eastern shore than at San
 Juan Island  (Fig.  B-73).

      Although not as  numerous as English sole, starry flounder occurred
 along the Strait  of Juan de  Fuca in  all but the Dungeness Spit beach seine
 collections; juveniles  and adults  occurred in approximately equal proportions.
 The overall prey  spectrum (Fig.  B-74) was quite similar to that of the rock
 sole (Fig.  B-69):  polychaete annelids,  gammarid amphipods, and tanaids
 supplied the greatest proportions of the total IRI.

 C-0 Sole

      C-0 sole were caught during beach seine collections at South Beach and
 Deadman Bay in northern Puget Sound.  Their principal prey items were
 flabelliferan isopods (45.8  percent  of total IRI), fish (21.4 percent),
 polychaetes (14.3 percent),  amphipods (9.2 percent), and turbellarians (4.4
 percent).   One C-0 sole from a  beach seine collection at southern Guemes
 Island in July 1974 had 12 bivalves  (98.0 percent of total biomass) and 10
 pieces of algae (Rhodophyta) in its  stomach.

      C-0 sole collected from the Strait of Juan de Fuca almost exclusively
 originated from the protected,  sand-eelgrass habitat at Beckett Point.
 Polychaete annelids,  with over  80% of the total IRI, were the most important
 prey taxa and bivalves contributed almost 13% (Fig. B-75).

 Sand Sole
      In sand sole from the eastern shoreline of northern Puget Sound  (Birch
 Bay), gammarid amphipods were the most important prey taxa, with 82.1 percent
 of  the total IRI (Fig. B-76).  Polychaetes supplied 8.3 percent and epibenthic
 organisms—tanaids (3.8 percent), cumaceans (3.4 percent), and valviferan
 isopods (1.3 percent)—accounted for the remaining proportion.
                                      203

-------
                             INDEX OF RELRTIVE IhPORTRNCE (I.R.I.) OIRGRHM
                             FROM FILE IOENT. HH  BS. STflTION  HLL

                                       PREDHTOR  8857041401 - PLHTICHTHYS STELLRTUS
                                    (STfiRRY FLOUNDER     )    ROJUSTED SRNPLE SIZE = 22
                          100
                      a
                      i    so

                      I    60
                      i
                      r    40
                      I
                      5    20
                      §    20
                      8
                           4°
                           60
                           80
                          100
              j: •-* -o -H
              U O O -H
              O CUV
                                   20
                                          40
                                                 60
                                                         80
                                                                100
                                                                       120
                                                                              140
                                                                                     160
                                           CUMULRTIVE FREQUENCY OF  OCCURRENCE
PSEY ITEM
GAMHARIDEA
ISOPOOA
TANAIDACEA
POLYCHAETA
BIVALVIA
OLIGOCHAETA
CAIANOIDA
NEt'ATOOA
FLA8ELLIFERA
CIRPIPEOIA
VALVIFEPA
PLEOCYEMATA-CAPIDEA
OSTEICHTHYES
PENAEIOAE
FPEO
OCCUR
3?. no
??.no
1<>.00
K.OO
10.00
B.OO
s.no
•5.00
s.oo
5.00
5.00
s.oo
s.oo
5.00
NUM.
COMP.
23.92
?7.?H
<..98
1.31
.<)6
i.on
3.65
GRAV.
COMP.
2.07
.61
.31
.3P
1 .99
11.03
.00
.04
1.91
76.60
.00
1.39
.38
3.06
PREY
I.R.I.
B31.7
752. n
74.1
23.9
29. *
293. «S
1.6
20.1
21.2
416.2
1 .6
10.2
6.Q
33.5
PERCENT
TOTAL IRI
33. 01)
29.90
2.90
1.00
1.20
11.70
.00
.80
.80
16.50
.00
.40
.30
1.30
              _PRFY TAXA_WITH FPEO.  OCCU".. LESS THAN. 5_ANO NUMEP 1C AL .AND GPAVIMETPlC._
              COMPOSITION BOTH LESS THAN  1 ARE EXCLUDED FoOM  THE TABLE AND PLOT
              (RUT NOT FROM CALCULATION OF OIvFPSITY INDICES)
                "PERCENT DOMINANCE  INDEX
                 SHANNON-WFINER DIVERSITY
                 EVENNESS INDEX
 .19    .60
2.76   1.36
 .73    .38
 .24
2.3S
 .66
Fig.  B-73.   IRI  prey spectrum of  starry  flounder  in shallow sublittoral
               habitats along eastern shoreline  of northern Puget  Sound.
                                              204

-------
                               100
                                 INDEX OF RELRTIVE inPORTflNCE (I.R.I.I  DIBGRBH
                                 FROI1 FILE IOENT.  76-78. STBTION BLS'B
                                              8957041401 - PLHTICHTMtS STELLBRIS
                                                           STBRRY FLOUNOER
                                                 BOJUSTEO SBHPLE SIZE = 117
                           1
                           ~    40
§
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1
COMPOS I
5

20
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40
60
80
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£ 3 till; 33fl3Z

                                             40    60

                                               OMJLBT1VE FREQUENCIf OF OCCURRENCE
                             iKinri nr on»Tiyr  IMPBRTANCF il.P.I.I  T»8i F	
                             USING FlLElOa  76-79.   »LST« FOR PLOT
                                              FRFO   NUM.   GRAV.  PREY     PERCENT
                                              OCCUR  CO«P.  COMP.  I.R.I.  TOT»L  1RI
GlWKOlnFt J9

r.Iv'dR"olFl01E
• unOYTIOit
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26
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114
04
04
63.16 3344.
.70 761.
.16 374.
2.97 69.
.91 170.
.20 13.
1.13 82.
.01 IS.
.19 29.
14.54 65.
3.?9 9.
2.45 7.
1.54 1.
1.56 1.
1.16 1 .
1 . 35 1 .
6§.06
7.63
1.4?
1.64
.11
.59
1.32
.14
• 01
.01
• 02
.0?
                     P-P-£Y_IAX4_klIH_rj:.F_0. -OCCUR. LESS JM1X 5_»SJ) NU»E"1C»L_»N!1 C-»»VI«ET9IC
                     ClwpnsITlnN PIU.I LC^1;  TH»N |  »RF E»CUinED Fon» THE HBLF «NO PLOT
                     OUT NOT FPO" C»I.CIJL*TInM OF  01VERSIT* INOIrFS)
                       PEOCC'IT

                      _£VfN'lt!S
Fig.  B-74.    IRI prey  spectrum of  starry  flounder  in  shallow
                  sublittoral habitats  along  Strait  of  Juan  de Fuca.
                                                    205

-------
                           100
                              INDEX OF RELflTIVE IMPORTRNCE (I.R.I.) DlflGRflM
                              FROM FILE IDENT.  76-78. STflTION flLSTfl
                                            8857041601  - PLEURONICHTHYS COENOSUS
                                                              C-0 SOLE
                                                flOJUSTEO SflMPLE SIZE =  19
1
1
DO
Z
o
"
1
§
I—
0-

1
UJ
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76
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7.44
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7.19
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1.97
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.22
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1.31
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.22
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1.73
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JR49.? 	 12
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1 10.9
12.2
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14.6
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?• 1
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l. __________
                "ERCENT nn"lNANCE INPFX
                SHA'JNO'J-WEI^EP DIVERSITY.	
                EVENNESS  INOF»
 .48    .47
1.81 _ 1.53
 .49    .41
 .69
_.96_
 .26
Fig. B-75.   IRI  prey spectrum  of C-0  sole in shallow sublittoral  habitats
               along Strait of Juan de Fuca.
                                              206

-------
                    I


                    to
                    z
                    o

                    8
                    fc
                    S
                           INDEX OF RELRTIVE INPORTHNCE (I.R.I.) OIRGRRM
                           FROM FILE IOENT. WW  BS. STRTION  flLL
                                   PREDflTOR 8857041701 - PSETTICHTHYS MELRNOSTICTUS
                                  (SRND SOLE         )   ROJUSTED SRHPLE SIZE = 16
    100


     80


     60


     40


     20
                    2    20


                    CD

                    S
                    I—

                    P    60
^    80
K

    100
                                                                             «   -H C O
                                                                             u   > o c.
                                                                             U   fH 00-H
                                                                             to   id C JZ
                                      50
                                                 100
                                                             150
                                                                         200
                                                                                    250
                                         CUMULRTIVE FREQUENCY OF OCCURRENCE
                    PREY ITFM
                      FPEO
                     _C)CCUR
NUM.
COMP.
GRAV.
COMP.
PREY
I.R.I.
 PERCENT
TOTAL IRI
GAMMARIDEA
POLYCHAETA
VALVJFERA
TASAIDACEA
CUMACEA
BIVALVIA
CRANGONIOAE
AMPHIPOD^A-HYPERI IDFA
94.00
56.00
31.00
25.00
19.00
13.00
6. no
6.00
62.90
3.77
5.51
1*.81
8.99
1.45
.?9
.?9
                                    53.27 J09?0.0_
                                    15.89  1111.0
                                      •00   170.R
                                     3.74   513.7
         . 5*
     _ 5.61
         .00
                                            103.9
                                             35.4
                                              1.7
                                                                         8?.1Q_
                                                                        "  B.*0
                                                                          1.30
                                                                          3.90
                                                                         "3.VO"
                                                                           .HO
                                                                           .30
                                                                           .00
                  TAXA WITH FREO. OCCUR. LESS THAN 5 AND NUMERICAL  AND  GRAVIMETRIC
             COMPOSITION BOTH  LESS  THAN  1 ARE EXCLUDED FPOM THE  TABLE  AND  PLOT  ~ "
             (HUT NOT FRO- CALCULATION OF DIVERSITY INDICES)
                PERCENT DOMINANCE  INDEX "
                SHANNON-WfINER  DIVERSITY
                EVENNESS INDEX
                               .44    .34
                              1.71   1.98
                               .57    .77
                          .68
                         1.04
                          .37
Fig.  B-76.   IRI  prey  spectrum of  sand  sole in shallow sublittoral habitats
               along eastern shoreline of northern Puget Sound.
                                             207

-------
     In the Strait of Juan de Fuca.sand sole were almost as abundant as
English sole but originated primarily from the four western beach seine
sites.  The overall prey spectrum (Fig. B-77) showed a radical difference
from northern Puget Sound and from the other flatfish by the predominance
of epibenthic mysids (primarily Neomysis awatschensis but also including
N_. rayi and Archaeomysis grebnitzki, accounting for 88.9% of the total IRI)
and the absence of polychaetes.  Other epibenthic crustaceans, such as
gammarid amphipods, cumaceans, and shrimp (including Crangon stylirostris
and Pandalus danae), were the secondary prey.  Fish (including juvenile
Pacific herring, Pacific sand lance, tidepool snailfish, Liparis florae,
and sand sole, Psettichthys melanostictus) accounted for 31.5% of the total
prey biomass but were not common or abundant enough to provide a high (1.5%)
proportion of the total IRI.
                                     208

-------
                            INDEX OF RELflTIVE  IflPORTflNCE (I.R.I.) DIRGRflM
                            FROM FILE IDENT.   76-78. STRTION RLSTfi
                                         8857041701 - PSETTICHTHYS HELHNOSTICTUS
                                                            SRND SOLE
                                               flOJUSTED SRMPLE SIZE = 308
                    PPF.Y ITFM
                                           CyrjULflTIVE FREQUENCY^ OF OCCURRENCE
                                           F&EO   MUM.  " GRAV.   PREY     PERCENT
                                           OCCIIP  CO«P.   COMP.   I.P.I.   TOTAL  IRI
r,AWMARIOEA
CM'MCEA
OSTFICHTHYS
MNIOFNTIF IED
k'ATANTI A
CLI'PEIDAE
. AMworiYTIPAE
PI_FllPO"'FCTlnAF
67. ?1
3?. 14
9.09
A. 17
3.
-------
                                  REFERENCES
Alevision, W. S.  1975.  Comparative feeding ecology of a kelp bed embiotocid
      (Embiotoca_ lateralis).  Copeia, 1975(4):609-615.

Bane, G. and M. Robinson.  1970.  Studies on the shiner perch, Cymatogaster
      aggregata  Gibbons, in upper Newport Bay, California.  Wasmann J. Biol.
      28(2):259-268.

Barraclough, W. E.  1967a.  Data record.  Number, size and food of larval and
      juvenile fish caught with an Isaacs-Kidd trawl in the surface waters of
      the Strait of Georgia, April 25-29, 1966.  Fish Res. Bd. Canada. M. S.
      Report Series 926, 79 pp.

Barraclough, W. E.  1967b.  Data record.  Number,  size composition
      and food of larval and juvenile fish caught with a two-boat surface
      trawl in the Strait of Georgia, June 6-8, 1966.  Fish Res. Bd. Canada.
      M.S. Report Series 928, 58 pp.

Barraclough, W. E. & J. D. Fulton.  1967. Data record.  Number, size
      composition and food of larval and juvenile fish caught with a two-boat
      surface trawl in the Strait of Georgia, July 4-8, 1966.  Fish Res. Bd.
      Canada.  M. S. Report Series 940, 82 pp.

Barraclough, W. E.  1967c.  Data record.  Number, size and food of larval and
      juvenile fish caught with a two-boat surface trawl in the Strait of
      Georgia, April 25-29, 1966.  Fish Res. Bd.  Canada, M. S. Report Series
      922, 54 pp.

Barraclough, W. E. & J. D. Fulton.  1968.  Data record.  Food of larval and
      juvenile fish caught with a surface trawl in Saanich Inlet during June
      and July 1966.  Fish Res. Bd. Canada, M. S. Report Series 1003, 78 pp.

Barraclough, W. E., D. G. Robinson & J. D. Fulton.  1968.   Data record.
      Number, size composition, weight, and food of larval and juvenile fish
      caught with a two-boat surface trawl in Saanich Inlet, April 23-July 21,
      1968.  Fish Res. Bd. Canada, M. S. Report Series 1004, 305 pp.

Barton, M. G.  1974.  Studies on the intertidal vertical distribution, food
      habits, and movements of five species of eel (Pisces:  Stichaeidae and
      Pholidae) at San Simeon, California.  M. A. Thesis,  Calif. State Univ.,
      Fullerton.  100 pp.

 Conley, R. L.  1977.  Distribution, relative abundance, and feeding habits
      of marine and juvenile anadromous fishes of Everett Bay, Washington.
      M. S. Thesis, Univ. of Washington, 57 p.
                                      210

-------
Cross, J. N., K. L. Fresh, B. S. Miller, C, A,  Simenstad,  S.  N.  Steinfort
     and J. C. Fegley.  1978.  Nearshore fish and  macroinvertebrate
     assemblages along the Strait of Juan  de  Fuca  including food habits of
     the common nearshore fish:  report of two years  of  sampling.  Annual
     Report to NOAA, MESA Puget Sound Office.   Fish.  Res.  Inst., Coll.  Fish.
     Univ. Washington, Seattle, FRI-UW-7718.   Also NOAA Tech. Memo. ERL MESA-32.

De Martini, E. E.  1969.  A correlative study of the  ecology and comparative
     feeding mechanism morphology of the embiotocidae (surf fishes) as evidence
     of  the family's adaptive radiation into  available ecological niches.
     Wasmann J. Biol. 27(2):177-247.

Feller,  R. J. and U. W. Kaczynski.  1975.   Size-selective  predation by juvenile
     chum  salmon (Oncorhynchus keta) on epibenthic prey  in Puget Sound. J.
     Fish. Res. Bd. Canada 32(8):1419-1429.

Feller,  R. J.  1977.  Life history and production  of  meiobenthic harpacticoid
      copepods in Puget Sound.  Ph.D. Thesis,  Dept.  Ocean.  Univ.  Washington,
      Seattle.  249 pp.

Fresh,  K.  L., D. Rabin, C. A. Simenstad, E. 0.  Salo,  K.  Garrison and L.
      Matheson.  1978.  Fish ecology studies in the Nisqually Reach area of
      southern Puget Sound, Washington.  Ann.  Prog.  Rept.,  March 1977 - June
      1978, to Weyerhaeuser Co., Fish. Res.  Inst.,  Coll. Fish., Seattle, FRI-UW-
      7812.  151 pp.

Fresh,  K.  L.  1979.   Distribution and abundance of fishes  occurring in the
      nearshore  surface waters of northern  Puget Sound, Washington.  M.S. Thesis,
      Coll. Fisheries, Univ. Washington, Seattle, 120  pp.

Gnose,  C.  E.  1967.   The  ecology of th striped seaperch, Embiotoca lateralis,
      in Yaquina Bay,  Oregon, M. S. Thesis,  Oregon  State  University, 51 pp.

Harris,  C. K. and  A.  C. Hartt.  1977-  Assessment  of  pelagic and nearshore
      fish in three bays on the east and south coasts  of  Kodiak Island, Alaska.
      Final Rept. to OCSEAP/BLM  (proposal RU 485, A-7, A-8, A-9 & A-ll).  Fish.
      Res.  Inst., Univ. Washington. FRI-UW-7719.  190 p.

Healey,  M. C. ,  R.  J.  LeBrasseur, J. R. Sibert,  W.  E.  Barraclough and J. C.
      Mason.   1976.  Ecology of young salmon in Georgia Strait,  pp. 201-207.
      In  G.  R.  Gundstrom  (Ed.) Proc. 1976.  N.E. Pacific pink and chum
      salmon  workshop, Alaska Dept. Fish &  Game, Juneau.

Holmquist, C.   1973.  Taxonomy, distribution  and ecology of three species of three
      species, Neomysis  intermedia  (Zerniavsky), N.  awatchensis (Brandt) and
      N.  mercedis  (Holmes)  (crustacea, mysidacea).   Zool. J. B. Syst. Bd. 100,
      5:197-222.

 Johnson, C.  L.  1968.   Food of  the buffalo sculpin, Enophrys bison. J. Fish.
      Res.  Bd.  Canada  25(4):807-812.
                                      211

-------
Johnson, C.L.  1970.  Notes on the intertidal life history of the northern
     clingfish Gobiesox maeandricus (Girard).  Amer. Midi. Nat. 83(2) -.625-627.

Jones, A.C.  1962.  The biology of euryhaline fish (Leptocottus armatus
     Girard).  Univ. California Publ. Zool. 67(4):321-367.

Jones, B.C., and G.H. Geen.  1977-  Food and feeding of spiny dogfish  (Squalus
     acanthias) in British Columbia waters.  J. Fish. Res. Board Can.  34(11):
     2067-2078.

Leaman, B.M.  1976.  The ecology of fishes in British Columbia kelp beds;
     Barkley Sound Nereocystis beds.  Kept, to Map Resources Branch, British
     Columbia Dept. Recreation and Conservation.  Inst. Anim. Resource Ecology,
     Univ. British Columbia, 80 pp.

Meyer, T.L., R.A. Cooper, and R.W. Langton.  1979.  Relative abundance, behavior,
     and food habits of the American sand lance, Ammodytes americanus, from
     the Gulf of Maine.  Fish. Bull. 77(1):243-253.

Miller, B.S., C.A. Simenstad, K.L. Fresh, F.C. Funk, W.A. Karp, S.T. Borton,
     and L.L. Moulton.  1977.  Puget Sound baseline program; nearshore fish
     survey.  Final Rept. to Wash. State Dept. Ecology, June 1974 - June 1977-
     Fish. Res. Inst., Coll. Fish., Univ. Washington, Seattle.  FRI-UW-7710,
     220 pp.

Moulton, L.L.  1977.  An ecological analysis of fishes inhabiting the rocky
     nearshore regions of northern Puget Sound, Washington.  Ph.D. Thesis,
     Univ. Washington, Seattle, 181 pp.

Nakamura, R.  1971.  Food of two cohabitating tide-pool Cottidae.   J. Fish.
     Res. Board Can. 28(6):928-932.

O'Connell, C.P.  1953.  The life history of the cabezon Scorpaenichthys
     marmoratus (Ayres).  Calif. Dept. Fish and Game 93:1-76.

Quast, J.C.  1968.  Observations on the food of kelp-bed fishes.  In W.J.
     North and C.L. Hubbs, eds., Utilization of kelp-bed resources in southern
     California.  Calif. Fish and Game, Fish. Bull. 139, 264 pp.

Reay, R.J.  1970.  Synopsis of biological data on North Atlantic sand eels of
     the genus Ammodytes, A. tobianus, A. dubius, A. americanus and A. marinus,
     from the Gulf of Maine.  Fish. Bull. 77(1):243-253.

Robinson, D.G., W.E. Barraclough, and J.D. Fulton.  1968a.  Data record:
     Number, size composition, weight and food of larval and juvenile fish
     caught with a two-boat surface trawl in the Strait of Georgia, May 1-4,
     1967.  Fish. Res. Board Can., M.S. Rept. Ser. 964, 105 pp.

Robinson, D.G., W.E. Barraclough and J.D. Fulton.  1968b.  Data record:
     Number, size composition, weight and food of larval and juvenile fish
     caught with a two-boat surface trawl in the Strait of Georgia, June 5-9,
     1967.  Fish. Res. Board Can., M.S. Rept. Ser. 972, 109 pp.


                                     212

-------
Robinson,  D.  G.   1969.   Data record.   Number, size composition, weight
     and food of larval and juvenile fish caught with a two-boat surface
     trawl in the Strait of Georgia.   June 4-6, 1967.  Fish. Res, Bd. Canada,
     M.  S. Report Series 1012.   71 pp.

Scott, J.  S.   1973.   Food and inferred feeding behavior of northern sand lance
     (Ammodytes  dubius). J. Fish.  Res. Bd. Canada 30:451-454.

Sekiguchi, H.  1977.   Further observation on the feeding habits of planktivorous
     fish sand-eel in Ise Bay.   Bull.  J.  Soc. Sci. Fish. 43:417-422.

Senta, T.   1965.  Nocturnal behavior of sand-eels, Ammodytes personatus
     (Girard) Bull.  J.  Soc. Sci. Fish. 31:506-510.

Simenstad, C. A., B.  S. Miller,  J. N.  Cross, K. L. Fresh, S. N. Steinfort and
     J.  C. Fegley.  1977.  Nearshore fish and macroinvertebrate assemblages
     along the Strait of Juan de Fuca including food habits of nearshore fish.
     Fish. Res.  Inst. Rept. to MESA Puget Sound Project.  NOAA Tech. Memo.
     ERL-MESA-20, 144 p.

Simenstad, C. A. and W. J. Kinney.  1978.  Trophic relationships of out-
     migrating chum salmon in Hood Canal, Washington, 1977-  Final report
     to Wash. Dept. of Fish, Fish.  Res. Inst. Coll. Fish., Univ. Washington,
     Seattle.  FRI-UW-7810, 75 pp.

Tasto, R.  N.   1975.   Aspects of the biology of the Pacific staghorn sculpin,
     Leptocottus armatus Girard, in Anaheim Bay. pp.  123-135.  In: E. D. Lane
     and C. W. Hill (Eds.).  The marine resources of Anaheim Bay. Calif. Dept.
     F\.sh. and Game.   Fish. Bull.  165, 195 pp.

Wares, P.  G.   1968.   Biology of the pile perch (Rhacochilus vacca).  M. S.
     Thesis,  Oregon State Univ., Corvallis.  93 pp.

Weller,  D. B.  1975.   The life-history of the shiner seaperch, Cymatogaster
     aggregata Gibbons, in Anaheim Bay, California, pp.  107-115 In:  E. D.
     Lane and C. W.  Hill (Eds.)  Marine resources of Anaheim Bay, Calif. Dept.
     Fish & Game. Fish. Bull. 165, 195 pp.
                                     213

-------
                  APPENDIX C:  SEA BIRDS AND SHORE BIRDS

     Eighty-one species of marine or estuarine-associated birds are reported
or could be safely assumed to consistently occur in north Puget Sound and
the Strait of Juan de Fuca (Jewett, et al., 1953; Robbins, et al., 1966;
Alcorn 1971; Salo 1975; Manuwal 1977).  Ten species occur in abundance and
represent sizable breeding or wintering populations; an additional 54 species
could be considered common, many of which are seasonal migrants and residents.
The remaining species are either not commonly encountered—e.g., they migrate
through the region offshore—or numerically unimportant (Table C-l).

     The following species accounts discuss food habits and feeding behavior
of all species for which such data could be found.  For many species,
especially the shore birds, there were few, or very unspecific, data avail-
able for this region.  In those cases, information for that species occurring
outside the Pacific Northwest was utilized, or in the instances of no such
data, the diet was inferred from congeneric species.  Several review
references, notably Jewett, et al. (1953), Eaton (1975), and Salo (1975),
were extremely useful in locating data on the general food habits of the
less-known birds.

     The functional feeding groups of the 55 abundant and common marine and
shore birds associated with north Puget Sound and Strait of Juan de Fuca
habitats are summarized in Table C-2.  Of these species, 19 (35%) utilize
fishes as their principal prey resources, as either obligate or facultative
piscivores; 18 (33%) feed  mainly on benthic invertebrates in the littoral or
shallow sublittoral zones; 7 (13%) are omnivorous, eating both plants and
invertebrates; 5 (9%) are planktivorous; 4 (7%) are herbivorous; and 2 (3%)
are either parasitic, or like the bald eagle, facultative avivores preying on
both sea birds and fish.

     Sixteen (29%) of the bird species feed in the shallow sublittoral zone,
which includes much of the sand to cobble shoreline and embayments in this
region; 14 (25%) feed in the salt marsh and mudflat environments character-
istic of large estuaries or contained embayments; 11 (20%) feed in neritic
waters; 4 (7%) obtain their prey on relatively exposed sand-gravel beaches;
3 (5%) utilize the fauna of the rocky littoral environment or the adjacent
kelp bed fauna; and 4 (7%) gulls have to be essentially universal to almost
all of these habitats.

                             Species Accounts

Arctic Loon

     Arctic loons occur in north Puget Sound between October and May,
primarily as migrants and winter residents (Jewett, et al., 1953; Alcorn 1971;

                                    214

-------
Table C-l. Relative abundance of marine  and  shore  birds  known to the
           northern Puget Sound and  Strait of  Juan de Fuca area.
           A = abundant, C = common,  NC  = not  common, R  = rare;
           asterisk denotes seasonal occurrence.
Scientific name
                                     Common name
                         Abundance
Order Gaviiformes

    Gavia avctioa
    G. irnner
    G. stellata

Order Podicipediformes

   Family Podicipedidae
    Aechniophorus  occidentalis
    Podiaeps  auritus
    P. grisegena
    P. nigrioollis

 Order Procellariiformes

   Family Procellariidae
     Filmarus  glacialis
     Puff inns  griseus
   Family Hydrobatidae
     Oceanodroma leucorhoa

 Order Pelecaniformes

   Family Phalacrocoracidae
     Phalacroaorax penicillatus
     P.  auritus
     P.  pelagieus

 Order Anseriformes

   Family Anatidae
     Branta aanadensis
     B.  nigvicans
     Anser albifrons
     Chen hyperborea
     Anas platyrhynchos
     A.  aeuta
     A.  olypeata
     A.  carolinensis
     Mareea americana
     Aythya valisineria
     A.  marila
     A.  af'finis
Arctic loon                   C*
common loon                   C
red-throated loon             C*
grebes
western grebe                A
horned grebe                 C*
red-necked grebe             C*
eared grebe                  C*
fulmars
fulmar                        NC
sooty shearwater              NC
storm petrel
Leach's petrel                NC
cormorants
Brandt's cormorant            C*
double-crested cormorant      C
pelagic cormorant             C
waterfowl
Canada goose                  C*
black brant                   A*
white-fronted  goose           NC*
snow goose                    C*
mallard                       A*
pintail                       C*
northern shoveler             C*
green-winged teal             NC*
American widgeon             C*
canvasback                    NC*
greater scaup                 NC*
lesser scaup                  A*
                                    215

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Table  C-l. Relative abundance of marine and shore birds known  to the
           northern Puget Sound and Strait of Juan de Fuca area.
           A = abundant, C = common, NC = not common, R = rare;
           asterisk denotes seasonal occurrence, continued.
Scientific name
Common name
Abundance
    Buaephala olangula
    B. islandiea
    B. albeola
    Clangula hyemlis
    Histrionicus histrionicus
    Melanitta deylandi
    M. perspicillata
    M. nigra
  Family Anatidae
    Mergus merganser1
    M. sernaton

Order Falconiformes

  Family Pandionidae
    Pond-ion halinetus
  Family Accipitridae
    Aalineetus leuooaephalus

Order Ciconiiformes

  Family Ardeidae
    Ardea herodias

Order Gruiformes

  Family Rallidae
    Rallus limicola
    Fulica americana

Order Charadriiformes
common goldeneye             C*
Barrow's goldeneye           C*
bufflehead                   C*
oldsquaw                     C*
harlequin duck               NC*
white-winged scoter          C*
surf scoter                  A
black scoter                 NC*
mergansers
common merganser             C
red-breasted merganser       C*
ospreys
osprey                       NC
kites, hawks, eagles
bald eagle                   C
herons, bitterns
great blue heron
rails, gallinules, coots
Virginia rail                R
American coot                C
  Family Haematopodidae
    Haematopus baohmani
  Family Charadriidae

    Charadrius semipalmatus
  Family Scolopacidae
    Numenius phaeopus
    Actitis macularia
    HeteYOScelus inccmwn
    Limnodromus griseus
    L.  soolopaoeus
    Aphriza virgata
oystercatchers
black oystercatcher          C
plovers, surfbirds,
  turnstones
semipalmated plover          NC*
sandpipers
whimbrel                     C
spotted sandpiper            A*
wandering tattler            NC*
short-billed dowitcher       C
long-billed dowitcher        C
surfbird                     C*
                                   216

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Table C-l. Relative abundance of marine  and  shore  birds  known to the
           northern Puget  Sound and  Strait of  Juan de Fuca area.
           A = abundant, C = common,  NC  = not  common, R  = rare;
           asterisk denotes seasonal occurrence, continued.
Scientific name
                                     Common name
                         Abundance
    Arenaria interpres
    A. melanocephala
    Tringa melanoleuoa
    Calidris ptilocnemis
    C. alpina
    C. aanutus
    C. minutilla
    Ereunetes mauri
    Crocethia alba
   Family Phalaropodidae
    Phalaropus  tricolor
    Lobipes  lobatus
ruddy turnstone
black turnstone
greater yellowlegs
rock sandpiper
dunlin
knot
least sandpiper
western sandpiper
sanderling
phalaropes
Wilson's phalarope
northern phalarope
NC*
C*
C*
NC*
C*
C
C
A
C*

NC*
NC*
   Family Stercorariidae
     Stevoovarius parasiticus
     S. pomarinus
   Family Laridae
     Larus glauceseens
     L. occidentalis
     L. nurgentatus
     L. californicus
     L. delawarensis
     L. oanus
     L. heermanni
     L. Philadelphia
     Xema sabini
     Rissa tridactyla
     Sterna hirundo
     S. paradisnea
     Hydroprogne aaspia
   Family Alcidae
     Uria aalge
     Cepphus oolumba
     Lunda airrhata
     Cerorhinea monocerata
     Ptyahoramphus aleutica
jaegers, skuas
parasitic jaeger
pomarine jaeger
gulls
glaucous-winged gull
western gull
herring gull
California gull
ring-billed gull
mew  gull
Heermann's gull
Bonaparte's gull
Sabine's gull
blacklegged kittiwake
common tern
Arctic tern
Caspian tern
alcids
common murre
pigeon guillemot
tufted puffin
rhinoceros auklet
Cassin's auklet
C*
NC*

A
C*
C
C*
C*
C*
C*
A
NC
C*
C*
C*
C*

C*
C
C
A
R
     Synthliboramphus antigiam
     Brachyramphus mamoratum
 ancient murrelet
 marbled murrelet
C*
C
                                    217

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    Table C-2. Functional  feeding groups and representative prey taxa of 58 marine and shore birds
               common to northern Puget Sound and the Strait of Juan de Fuca.
    Habitat
Trophic position
Predator species
Prey taxa
    Offshore neritic
Obligate piscivore
Common murre
Black-legged kittiwake
Common tern
Rhinoceros auklet
Western grebe
10
H
oo
Facultative piscivore
Tufted puffin
Marbled murrelet
Ancient murrelet
                             Obligate planktivore
                             Cassins  auklet
Northern anchovy
Eulachon
Pacific herring
Pacific sand lance
Juv. rockfish
Juv. Pacific salmon
Surf smelt
Night smelt
Walleye pollock
Threespine stickleback

Pacific sand lance
Pacific herring
Surf smelt
Northern anchovy
Rockfish
Shiner perch
Juv. rockfish
Sea urchins
  (Strongylocentrotus sp.)
Bivalve molluscs
  (Mytilus sp.)
Euphausiids

Calanoid copepods
  (Calanus sp.)
Hyperiid amphipods
  (Parathemisto sp.)
Euphausiids

-------
 Table C-2.  Functional feeding groups  and  representative prey taxa of 55 marine and shore birds
            common to northern Puget Sound and  the Strait of Juan de Fuca, continued.
Habitat
                          Trophic position
 Predator species
Prev  taxa
                         Facultative planktivore     Mew gull
                                                     Bonaparte's gull
                         Parasite

Nearshore kelp beds      Facultative avivore
                        Obligate piscivore
                        Facultative piscivore
 Parasitic jaeger

 Bald eagle
Brandt's cormorant
Heermann's gull
Arctic tern
Euphausiids
Hyperiid amphipods
Pacific herring  (larv.?)
Pacific sand  lance
   (larv.?)

Foods of gulls and  terns

Gulls
Pigeon guillemots
Cormorants
Puffins
Pacific herring
Pacific salmon
Dolly Varden
Cutthroat trout
Flatfishes
Sculpins
Sea urchins
Crabs (Pugettia producta)

Redtail surfperch
Kelp greenling
Black rockfish
Cabezon
Pacific sand lance

Pacific herring
Pacific sand lance

-------
    Table  C-2. Functional  feeding  groups  and representative prey taxa of 55 marine and shore birds
                common  to northern  Puget Sound and the Strait of Juan de Fuca, continued.
    Habitat
Trophic position
Predator species
Prey taxa
     Inshore  rocky  littoral    Obligate benthivore
                            Black oystercatcher
                            Whimbrel
                            Black turnstone
ho
o
     Inshore  sand-gravel
      beaches
Obligate benthivore
Spotted sandpiper
Surfbird
Least sandpiper
Sanderling
    Nearshore                 Obligate  piscivore
       shallow  sublittoral
                            Double-crested cormorant
                            Red-necked grebe
                            Common merganser
Limpets
  Collisella digitalis
  Notoacmea scutum
  Aamea mitra
Chitons
  (Katharina tunioata)
Bivalve molluscs
  Mytilus edulis
  M. oalifornianus
Barnacles
  (Pollioipes polymerus)
Polychaete annelids
  (Nereis sp.)

Polychaete annelids
  Nereis sp.
  Streblospio benedicti
  Eteone longa
Amphipods
  Anigogarmarus conferrico lus
  Corophium sp.
  Orchestoidea pugettensis
Bivalve molluscs
Univalve molluscs

Penpoint gunnel
Crescent gunnel
Pacific sand lance
Shiner perch
Snake prickleback
Staghorn sculpin
Pacific herring
Juv. Pacific salmon
Northern anchovy

-------
    Table C-2. Functional feeding groups  and  representative prey  taxa of 55 marine and shore birds
               common to northern Puget Sound and  the  Strait of Juan  de Fuca,  continued.
    Habitat
Trophic position
Predator species
Prey taxa
                             Facultative  piscivore
                            Arctic loon
                            Common loon
                            Red-throated loon
                            Pelagic cormorant
                            Pigeon guillemot
                            Red-breasted merganser
                            Caspian tern
ro
ro
                            Obligate planktivore
                            Facultative benthivore
                           Eared  grebe
                           Lesser  scaup
                           Common  goldeneye
                           Bufflehead
                           Oldsquaw
                           Surf  scoter
                            Crescent  gunnel
                            Pacific sand  lance
                            Penpoint  gunnel
                            Staghorn  sculpin
                            Northern  clingfish
                            Snake prickleback
                            Pacific herring
                            Surf smelt
                            Black prickleback
                            Threespine prickleback
                            Juv. flatfish
                              (Pleuronectidae)
                            Snake prickleback
                            Shrimp
                              Spirontocopis
                               brevirostris
                              Crago alaskensis
                              Pandalus sp.
                            Crabs
                              (Hemigrapsis sp.)

                            Mysids
                            Amphipods

                            Bivalve molluscs
                              Tapes japonica
                              Ostrea  lurida
                              Mytilus edulis
                              Mya arenaria
                              Maooma  sp.
                            Crustaceans
                            Fish
                            Pacific herring eggs
                            Eelgrass  (.Zoetez-a  sp.)

-------
    Table C-2. Functional feeding groups and representative prey taxa of 55 marine and shore birds
               common to northern Puget Sound and the Strait of Juan de Fuca>  continued.
    Habitat
Trophic position
Predator species
Prey taxa
    Inshore, saltmarsh
      and mudflats
Obligate herbivore
Canada goose
Black brant
Snow goose
American coot
                             Omnivore,
                               Facultative herbivore
ro
to
                            Mallard
                            Pintail
                            Northern shoveler
                            American widgeon
                             Omnivore
                            Dunlin
                            Knot
                            Western sandpiper
Eelgrass (Zostera sp.)
Saltmarsh plants
  Saliaornia sp.
  Distichlis spiaata
  Cuscuta salina
  Trigloahia maritimwn
  Canex lyngbyei

Eelgrass (Zostera sp.)
Saltmarsh plants, seeds
  Saliaornia sp.
  Trigloahia maritinum
  Canex lyngbyei
Amphipods
  (An is ogcanmarus
   eonfervioolus)
Insect larvae
  (Aphrosylus sp.)

Saltmarsh plants, seeds
  Spevgularia marina
  Triglochia maritinwn
  Deschampsia oespitosa
Amphipods
  Covophiim sp.
  Anisogarmarius conferviaolus
  Orahestoidea pugettensis
  Amphitoe sp.
Polychaete annelids
  Nereis sp.
  Streblospio benediati
  Amphioteis mucronata
  Eteone lonqa

-------
     Table C-2.  Functional  feeding groups and representative prey  taxa  of 55 marine and shore birds
                common to northern Puget Sound and the Strait of Juan de Fuca,  continued.
     Habitat
Trophic position
Predator species
Prey taxa
NJ
NJ
                             Obligate piscivore
                             Obligate benthivore
                            Great blue heron
                            Shcrt-billed dowitcher
                           Long-billed  dowitcher
                             Facultative benthivore
                            Greater yellowlegs
                            Oligochaetes
                            Bivalve molluscs
                              Maaoma inaonspiaua
                              Mya arenaria
                            Tanaids
                              (Pancolus californiensis)
                            Nematodes

                            Staghorn sculpin
                            Starry flounder
                            Shiner perch
                            Penpoint gunnel

                            Polychaete annelids
                              (Nereis sp.)
                            Univalve molluscs
                              Nassariidae
                              Littorina sp.
                            Bivalve  molluscs
                              (Mytilus edulis)
                            Crabs
                            Shrimp
                            Isopods
                              (Exosphaevoma oregonensis)
                            Amphipods
                              Corophiufn spinicorne
                              Eahinogammarus  oohotensis

                            Molluscs
                            Crustaceans
                            Fish

-------
      Table C-2.  Functional feeding groups and representative prey taxa of 55 marine and shore birds
                 common to northern Puget Sound and the Strait of Juan de Fuca,  continued.
      Habitat
Trophic position
Predator species
Prey taxa
      Universal
Facultative benthivore
Glaucous-winged gull
Western gull
California gull
Ring-billed gull
SJ
Chitons
Starfish
Sea cucumbers
Sea urchins
  (Strongyloaentrotus sp.)
Crabs
  Cancer sp.
  Hemigrapsus sp.
Bivalve molluscs
  Mytilus edulis
  Clinocardium nuttalli
Polychaete annelids
  (Nereis sp.)
Pacific herring
Northern anchovy
Surf smelt
Pacific herring eggs
Cormorant fledglings
Murre fledglings

-------
Salo 1975).  They are quite common in the San Juan Islands, often in large
aggregations (up to 1,000) during migration (Salo 1975; Heilbrum, et al.,
1977).  When on marine waters arctic loons are assumed to eat fish, crusta-
ceans, and molluscs (Martin, et al., 1951; Salo 1975).

Common Loon

     The only year-round resident loon, common loons tend to be more abundant
in marine habitats during winter, though  seldom in as  large concentrations as
arctic loons (Jewett, et al., 1953; Alcorn 1971;  Salo  1975).  Wintering
common loons appear to feed primarily on  fish  (Martin, et al., 1951; Jewett,
et al., 1953).  Salo  (1975) reported a  common  loon preying on small 10-13 cm
flounders off Sandy Point, Whidbey  Island.

Red-Throated Loon

     Red-throated  loons,  the  least  common of  the  three loon  species, are
migrants and winter residents in north  Puget  Sound,  and  occur in aggregations
of up to 100 birds.   They appear to be  fairly abundant in  the San Juan
Islands  (Jewett, et al.,  1953;  Alcorn 1971;  Salo  1975).  Jewett,  et al.
(1953), and Martin, et al.  (1951),  reported  the  generalized  diet of red-
throated loons as  including fish, crustaceans, molluscs, fish eggs, leeches,
insects, and some  aquatic  vegetation.

Western Grebe

     Overall, western grebes  were the most abundant  of the species observed
in Manuwal's (1977) seven  transect  surveys in the San Juan Islands conducted
from ferries between  November 1973  and  April  1974.   They appear  primarily as
wintering populations, although some non-breeding birds  may  reside in  Puget
Sound through the  summer  (Jewett, et al., 1973; Salo 1975).  Fish are  the
primary prey organisms of western grebes  (Munro 1941a; Jewett, et al., 1953;
Phillips and Carter 1957;  Salo  1975).   Species which have  been identified as
prey include Pacific  herring  (Clupea harengus  pallasi),  especially during
spawning periods,  staghorn sculpin  (Leptocottus armatus) and other sculpins,
shiner perch (Cymatogaster aggregata),  and smelt  (Osmeridae).  Shrimp
(Pandalus goniurus) were also reported  as prey.

Horned Grebe

     Breeding horned  grebes are common  migrants on Puget Sound in winter,
typically from September through May, and a few non-breeding birds occur in
summer  (Munro 1941a;  Jewett,  et al., 1953; Bakus  1965; Salo 1975).  In
Manuwal's  (1977) ferry transect surveys through the  San  Juan Islands,  they
were observed to be numerous only once, in January 1974.  Fish (Pacific sand
lance, Ammodytes hexapterus, Pacific herring, and staghorn sculpin) and small
shrimp  (Spirontocaris brevirostris. Crago alaskensis. and Pandalus  sp.) and
crabs  (Hemigrapsis sp.) appear  to be most important  to the horned  grebe
feeding on marine waters  (Munro 1941a; Guiguet 1971b).
                                    225

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Red-Necked Grebe

     Of the species of grebes frequenting Puget Sound, the red-necked grebe
is probably the least abundant.  It follows the same pattern of occurrence,
however, residing as a winter migrant between November and May  (Munro 1971a;
Jewett, et al., 1958; Salo 1975).  It occurred in abundance during two of
Manuwal's (1977) San Juan Island ferry transect surveys, in January and April
1974.  Spawning Pacific herring, Pacific sand lance, pilchard (Sardinops
caerulea?), threespine stickleback (Gasterosteus aculeatus), blenny (Pholidae
and Stichaeidae?), and sculpin  (Cottidae) have been reported as food organisms
(Munro 1941a; Guiguet 1971b; Salo 1975).

Eared Grebe

     Like the other grebes, eared grebes are mainly winter migrants, and are
similar to horned grebes in distribution and abundance (Jewett, et al., 1953;
Salo 1975; Heilbrum, et al., 1977).  They did not appear in significant
numbers, however, during Manuwal's (1977) ferry transect surveys through the
San Juan archipelago.  Unlike the other grebes, eared grebes feeding in marine
habitats eat planktonic crustaceans—mysids and amphipods (Munro 1941a).

Fulmar

     Jewett, et al.  (1953), and Sanger (1965, 1970) provide indications and
data that fulmars are generally abundant in nearshore areas along the
Washington coast in fall (September-October), although it appears that their
incidence inside the Strait of Juan de Fuca is rare.  Martin (1942) suggested
that fulmars were apparently more abundant off the northwest coast of
Vancouver Island in winter.

     Sanger and Baird (1977) listed fish (70% frequency of occurrence), squid
(70%), and crustaceans (20%) as the principal prey of fulmars collected in
Alaskan waters between 1969 and 1976.  Fulmars collected in April from along
the Bering Sea icepack had consumed only squid.

Sooty Shearwater

     Sooty shearwaters appear in the region's offshore waters between March
and October  (Martin 1942) but seldom occur in nearshore waters  (Dr. Dave
Manuwal, Univ. Washington, pers. comm.).  Anchovies (Engraulidae) contributed
80% of the diet by volume of sooty shearwaters off Oregon, and  other fishes
and squid provided the difference, as reported by Wiens and Scott  (1975) who
calculated their total annual energy demand to equal 1.653 k cal m~2.
Feeding off northwest Vancouver Island was observed to include  Pacific sand
lance and euphausiids (Martin 1942).  Sanger and Baird's (1977) account of
the major food items of sooty shearwaters collected in Alaskan  waters between
1967 and 1976 indicated that fish occurred in 61% and squid in  46% of the
stomachs examined.

Leach's Petrel

     Martin  (1942) described the Leach's petrel as the second most frequently
seen bird in offshore areas from February through June, though  never in high
                                    226

-------
abundance.  Sanger  (1970), however, reported this species as the least
abundant and least  frequently seen during  his surveys.  According to
Manuwal's (1977) surveys, 3,655 pairs of Leach's petrels were breeding on the
outer coast of Washington, centered at Carroll Island; their occurrence^
inside the Strait of Juan de Fuca is rare, however.  In Wiens and Scott's
(1975) energy modeling  study, hydrozoa and euphausiids were listed as the
principal dietary components by percent volume for Leach's petrel.   The model
output predicted a  total annual energy demand of 0.106 k  cal m~2  for this
species, the lowest estimate of the four  species modeled.

Brandt's Cormorant

     The majority of Brandt's  cormorants  appear  to migrate into the  north
Puget Sound region  from California breeding colonies between October and
April, peak abundance occurring between November and February (Wahl  1977).
Breeding colonies of 20 to  50  mating  pairs, however, do occur on the outer
coast at Point  Grenville,  Grenville Arch, Willoughby Rock, and Quillayute
Needles  (Manuwal  1977); these  birds may constitute many of those feeding in
the western Strait  of Juan de  Fuca.   The 1976 Audubon Christmas bird count
(Heilbrum, et al.,  1977)  reported 479 sightings  at Victoria,  B.C.,  438 at
Fender Islands, B.C.,  12 in the San Juan Islands,  110 at Bellingham, and 15
in the Sequim-Dungeness area.

     Gabrielson and Jewett's  (1940)  account of Brandt's cormorant in Oregon
listed "trash fish" as the  primary food.   Scott's (1973) data indicated that,
on the basis of percentage  of  stomach contents volume, anchovy, rockfish
(Scorpaenidae), sculpin,  and other fishes contribute equally to the Brandt's
cormorant diet  off  Oregon in  summer;  in winter,  rockfish, sculpin,  and other
fishes make up  the  diet.   Using  these data,  Wiens and Scott (1975)  determined
that the  total  annual energy demand  for this species was 2.308 k cal m~2,
second highest  of the four  species that they modeled.

     Hubbs's, et  al. (1970), account  of the diverse prey spectrum of Brandt's
cormorants feeding  in southern California indicated that they typically
foraged  in Macrocystis kelp beds  where they captured midwater fishes (black-
smith, Chromis  punctipinnis, senorita,  Oxyjulis  californica,  white seaperch,
Phanerodon atripes, kelp perch, Brachyistius frenatus, and vermillion  rock-
fish,  Sebastes  miniatus).   Over sand  bottoms,  the diving birds utilized
various  species of  flatfish (speckled sanddab,  Citharichthys stigmaeus,
fantail  sole, Xystreurys liolepis,  and curlfin turbot, Pleuronichthys
decurrens) and  roughback sculpin,  Chitonotus pugetensis.

     Martini  (1966) suggested  that the fish otoliths in the pellets regurgi-
tated  by western  gulls actually originated from  fish captured by Brandt's
cormorants.  These  fish included  Pacific  hake (Merluccius productus),  Pacific
 sanddab  (Citharichthys sordidus),  pink seapearch (Zalembius rosaceus),  short-
belly  rockfish  (Sebastes jordani),  sculpin (Icelinus sp.), and blackeye  goby
 (Coryphopterus  nicholsi)  which may be taken at depths of 20 to 50 m or more
 (Hubbs,  et  al.,  1970).

      Sculpin  (including cabezon,  Scorpaenichthys marmoratus).  juvenile rock-
 fish,  and greenling (Hexagrammidae) were  the principal prey taxa consumed  by

                                     227

-------
Brandt's cormorants off Oregon, except in mixed-species flocks when they  fed
on northern anchovy and "smelt" (Scott 1973); surfperches  (Embiotocidae)  were
also observed in their beaks.

     These data would tend to indicate that Brandt's cormorants feeding in
north Puget Sound and the Strait of Juan de Fuca would also prey upon
rocky/kelp bed fishes, which probably would include redtail surfperch
(Amphistichus rhodoterus), kelp greenling (Hexagrammos decagrammus), white-
spotted greenling (H. stelleri), several rockfish species, Pacific herring,
and Pacific sand lance.

Double-Crested Cormorant

     Manuwal (1977) estimated 390 nesting pairs on Washington's outer coast
and 64 pairs in the San Juan Islands and the Strait of Juan de Fuca; the  most
numerous populations were located at Bird Rocks and Viti Rocks.

     Robertson's (1974) report of food organisms regurgitated by double-
crested cormorant chicks in British Columbia included 11 fishes of which
gunnels (Apodichthys flavidus and Pholis laeta) were the most important
(46.6% total prey abundance, 51.6% total prey weight), followed by Pacific
sand lance (20.5%, 4.6%), shiner perch (15.5%, 20.5%), snake prickleback
(Lumpenus sagitta, 11.5%, 10.2%), staghorn sculpin (2.7%, 5.9%), Pacific
herring (1.4%, 2.7%), threespine stickleback (0.4%, 0.1%), juvenile salmon
(Oncorhynchus sp., 0.2%, 0.9%), and northern anchovy (Engraulis mordax, 0.2%,
0.1%).

     Unidentified crustaceans occurred in the one stomach sample listed by
Sanger and Baird (1977).  Scattergood's (1950) analysis of the contents from
35 stomachs collected in Maine included demersal fishes, primarily cunner
(Tautogolabrus adspersus, 35.2% of total abundance of identifiable prey),
shorthorn sculpin (Myoxocephalus scorpius) and longhorn sculpin (M. octodeci-
mospinosus, 11.1% each), redfish (Sebastes marinus, 9.3%), Atlantic herring
(Clupea harengus, 7.4%), winter flounder (Pseudopleuronectes americanus,  7.4%),
rock gunnel (Pholis gunnellus, 7.4%), and alewife (Alosa pseudoharengus,  5.6%).
Rock gunnel, winter flounder, cunner, and pollock (Pollachius virens) were
the predominant fish regurgitated by chicks at the White Islands, Maine,
rookery in early July, whereas in early August silversides (Meniclia notata)
suddenly became predominant, followed by cunner, redfish, and rock gunnel.

Pelagic Cormorant

     Pelagic cormorants are the most abundant of the nesting cormorants in
the region.  Manuwal  (1977) listed 995 pairs in the population breeding along
the outer coast and 395 pairs in the inland waters of the  San Juan Islands
and the Strait of Juan de Fuca.  Included in the latter estimate are large
colonies at Protection Island, Viti Rocks, Williamson Rocks, Bare Island, and
Colville Island.

     Pelagic cormorants' prey reported by Robertson  (1974) included shrimp
(19.4% prey abundance, 6.8% prey weight), and six species  of fish:  Crescent
gunnel (Pholis laeta, 34.9%, 37.1%), Pacific sand lance  (31.1%, 18.9%),

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penpoint gunnel (Apodichthys flavidus. 4.9%, 11.1%), staghorn sculpin  (2.9%,
13.2%), northern clingfish (Gobiesox maeandricus, 2.9%, 5.6%), and snake
prickleback (3.9%, 7.3%).  It was also suggested that prey size was larger
for pelagic cormorants than for double-crested cormorants.

     Sculpins (Myoxocephalus sp.) were the only identifiable prey organisms
taken by pelagic cormorants collected off Yaquina Head, Oregon, although
these birds were also observed eating northern anchovy and "smelt" while
feeding in large mixed-species flocks, and benthic  crustaceans while  feeding
singly.  Palmer (1962) listed demersal fishes  (sculpin, greenling, pholids,
poacher, Agonidae, and flatfish) and midwater  fishes  (Pacific herring and
Pacific tomcod, Microgadus proximus) as  the  principal food organisms  found in
31 stomachs collected in  Alaska.

Canada Goose

     Canada geese  are both winter migrants  and residents  along  the Washington
coast, specifically  during March through April and  September  to October.
Although migrating flocks are composed of thousands of birds, wintering flocks
are small, less than 100  individuals  (Salo  1975).   The 1976 Audubon Christmas
bird count recorded  as many as 515  Canada geese at  Vancouver, B.C., 87 at
Victoria, B.C., 54 at Nanaimo, B.C., and a  few individuals at  Sequim-Dungeness,
Bellingham, and Everett,  Washington (Heilbrum,  et al., 1977).  Martin, et al.
(1951), cites the  diet of wintering Canada  geese in this  region to include
primarily reeds and  the vegetative  parts of  barley, bulrush,  and  hardstem,
and secondarily wheat, wild barley, and  bromegrass.  When Canada  geese fre-
quent marine shoreline habitats, however, eelgrasses  (Zostera sp.) and salt-
marsh plants  (e.g.,  Salicornia sp., Distichlis  spicata, Cuscuta salina, and
Triglochin maritimum) may also enter their diet.

Black Brant

     Black brant  frequent the north Puget Sound region throughout the winter
and spring, both  as  migrants and local winter  residents,  but they appear to
be most abundant  in  March and April (Jewett, et al.,  1953; Einarsen 1965;
Salo 1975).  The  principal wintering sites include  Padilla Bay, Samish Bay,
Discovery Bay, and Sequim Bay.  In  April, as many as  18,000 birds have been
recorded in Samish Bay, 55,000 in Padilla Bay, and  6,000  in Discovery Bay
 (Salo  1975; D. Moriarity  and S. Oliver,  Graysmarsh  Wildl. Ref., pers.  comm.;
R.R. Parker, Wash. Dept.  Game, unpubl. data).  The  1976 Audubon Christmas
bird count  (Heilbrum, et  al., 1977) included black  brant  sightings all over
the north Puget Sound and Strait of Juan de Fuca region but in abundance
only at  Sequim-Dungeness.  Eelgrass is the main component of the black
brant's  diet, augmented slightly by ulvoid algae (Einarsen 1955; Salo 1975).

White-Fronted Goose

     White-fronted geese  occur as migrants during the  fall migration
especially  in September,  along the  northern Olympic Peninsula in areas such
as Dungeness Bay  (Salo 1975).  The  only  diet information  is from the  Califor-
nia wintering grounds where rice, vegetative parts  and seeds of grasses
sedges,  and wild  millet,  and barley were the most important foods (Martin,


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et al., 1951).  Hence, they resemble Canada geese in their foraging and may
utilize similar food sources—e.g., saltmarsh plants and eelgrasses—when
feeding in estuarine habitats of this region.

Snow Goose

     Like black brant, snow geese frequent the estuaries of northern Puget
Sound as migrants and as local winter residents.  Thus, their peak abundance
occurs twice, in November and again in April (Jeffrey 1950; Jewett, et al.,
1953; Alcorn 1971; Salo 1975).  Contrary to the brant distribution, snow
geese tend to congregate on the east side of Puget Sound, especially in
Padilla Bay, Skagit Bay, and Port Susan, reaching maximum densities of 27,000
to 30,000 geese.  They are reported to maintain local movements between these
areas and the Fraser River delta, British Columbia (D. Manuwal, pers. comm.).
Marsh plants, especially the roots and bulbs, are their principal food items
when foraging in the estuary  (Jeffrey 1950; Jewett, et al., 1953; Salo 1975).

Mallard

     The most common of the game ducks taken recreationally in Washington,
the mallard occurs mostly in freshwater habitats, but also frequents
estuarine habitats, principally between November and April (Munro 1943;
Jewett, et al., 1953; Alcorn 1971).  The largest concentrations in north
Puget Sound appear to be at Dungeness Bay (4,000-5,000 individuals), and the
region encompassing Padilla Bay, Samish Bay, Skagit Bay, and Port Susan
(19,500-44,000) (Salo 1975).  Mallards collected from Grays Harbor on
Washington's outer coast were feeding mainly on seeds of saltmarsh plants
(including Triglochin maritimum) and incidentally on polychaete annelid
worms (Smith and Mudd 1976).

Pintail

     Pintails occur concurrently with American widgeons, principally in the
upper tideflat regions of estuaries, as a fall and spring migrant, and
probably do not represent a significant breeding population (Munro 1944;
Jewett, et al., 1953; Alcorn 1971; Salo 1975).  Dungeness Bay and the eastern
shore between Port Susan and Boundary Bay tend to exhibit the highest concen-
trations of pintails (Salo 1975).  Skagit Bay undoubtedly has the greatest
numbers, up to 52,300 birds in October.  Padilla Bay is also an important
feeding area, with 13,000 to 15,000 birds in October-November (R.C. Parker,
Washington Dept. Game, unpub. data).  Smith and Mudd (1976) provided an
indication of a diverse diet spectrum for pintails foraging in estuarine
habitats at Grays Harbor, including eelgrass, seeds of the saltmarsh grasses
Carex lyngbyei and Triglochia maritimum, amphipods (Anisogammarus confervi-
colus), and insect larvae (Aphrosylus sp.)-

Northern Shoveler

     Northern shovelers occur in flocks of 20 to 500 birds on sheltered
estuaries and embayments between September and May (Kortright 1942; Jewett,
et al., 1953; Salo 1975).  They were sighted throughout the region during the
1976 Audubon Christmas bird count (Heilbrum, et al., 1977) but Victoria,

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B.C. (98 birds), and Bellingham, Washington  (58 birds), had  the  greatest
counts.  Kortright  (1942), Martin, et al.  (1951), and  Salo  (1975)  described
the diet of northern shoveler as 65% plant matter—pondweed,  sedge,  and grass
seeds—and 34% invertebrates, 19% of which were gastropod molluscs.

Green-Winged Teal

     A common game  bird in Washington's  freshwater habitats, green-winged
teal are also common in the upper estuarine  habitats of the State.  They are
particularly abundant in  the mud/eelgrass  estuarine  habitats along the
eastern shoreline of Puget Sound, including  Skagit  Bay (max. of approximately
25,000 in October), Port  Susan  (max. of  13,700 in December), Padilla Bay
(max. of 2,500 in November), and Samish  Bay  (max. of 5,100 in October) (R.C.
Parker, Washington  Dept.  Game,  unpub. data).  Smith  and Mudd (1976)  listed
invertebrates—amphipods  (Anisogammarus  confervicolus) and insect larvae
(including Sanderia sp. and Manayunkia  sp.)—as the  major prey items of
green-winged teal collected from mudflat habitats in upper Grays Harbor.

American Widgeon

     American widgeons  utilize  the  upper tideflat habitats of most of the
major  estuaries of  coastal Washington and  Puget Sound in winter.  At that
time (October to April) they usually are the most common duck on the region's
marine waters  (Salo 1975).  Port  Susan,  Skagit Bay,  Samish Bay, and Padilla
Bay tend  to  support the largest concentrations, peak abundances of 32,000 to
39,000 occurring  in October-November (R.C.  Parker,  Wn. Dept. Game, unpub.
data).   October-November  was also  the period of maximum abundance (3,500)
observed  at  Sequim  Bay, although  counts  usually stayed above 1,000 until
after  March  (D. Moriarity and  S.  Oliver, Graysmarsh Wildlife Refuge, pers.
 comm.).   The 1976 Audubon Christmas bird count (Heilbrum, et al., 1977)
 reported the greatest  number of sightings  in north  Puget Sound at Victoria
 (8,201)  and  Vancouver  (3,978)  in  British Columbia,  and Bellingham, Washington
 (1,014).   Widgeons  examined  from  four  sites  in upper Grays Harbor had been
 feeding principally upon  eelgrass  and dwarf  European eelgrass, "L_. nolti
 (Smith and Mudd 1976).                                          ~

 Canvasback

      Northeast Puget Sound bays and estuaries, including Drayton Harbor,
 Birch Bay, Padilla Bay, and  Samish Bay,  are the principal areas of canvasback
 concentrations, where peak abundances  of up to 1,000 are reached in November
 (Jewett, et al.,  1953;  Salo  1975;  R.C.  Parker, Wn.  Dept. Game, unpub. data).
 The eastern Strait of Juan de  Fuca may  also harbor  some winter residents,
 especially at Dungeness Bay and Victoria,  B.C. (Salo 1975; Heilbrum, et  al.,
 1977).  Jewett, et al.  (1953),  indicate that eelgrass and marine polychaete
 annelids and crustaceans  constitute the canvasback's diet.

 Greater Scaup

      The Strait of Georgia and northern Puget Sound harbor concentrations  of
 winter migrant and resident  greater scaup from December to February  (Munro
 1941b; Jewett, et  al.,  1953;  Salo 1975).  Relatively high abundances were

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enumerated at Vancouver, B.C. (1,067), and Bellingham, Washington  (1,055),
during the 1976 Audubon Christmas bird count  (Heilbrum, et al., 1977).
Padilla Bay, where peak abundances occurred of approximately 2,700 birds,
appears to harbor the highest concentrations  in north Puget Sound  (R.C.
Parker, Wn. Dept. Game, unpub. data), although Munro (1941b) reported as
many as 8,000 in one flock.  Manuwal  (1977) listed greater scaup as the
third most abundant diving species observed during his seven ferry transect
surveys through the San Juan archipelago.

     Littoral and shallow sublittoral molluscs and crustaceans appear to
constitute the diet of greater scaup foraging in marine environments  (Salo
1975), although these birds are also reported to congregate near spawning
Pacific herring in March and April (Munro 1941b).  Martin and Uhler (1939)
listed the native oyster, Ostrea lurida, along the Washington coast as one of
the principal food items of greater scaup.

Lesser Scaup

     Salo  (1975), using information from Munro (1941b), Jewett, et al. (1953),
Bakus  (1965), and Alcorn (1971), suggests that lesser scaup are distributed
mainly in  freshwater habitats during their winter residency in northern Puget
Sound; several sources, however, had documented lesser scaup occurring
commonly on the region's marine waters.  The  1976 Audubon Christmas bird
count  (Heilbrum, et al., 1977), in fact, included high counts at both
Vancouver  (452) and Victoria (1,012), B.C.  Although preference for aquatic
and marsh  plants was shown by Munro (1941b) and Martin, et al. (1951), the
diet of lesser scaup in marine waters may, like greater scaup, be oriented
toward invertebrates such as molluscs and crustaceans (Rogers and Korschgen
1966).

Common Goldeneye

     Common goldeneye was considered a common migrant and winter resident by
Munro  (1939), Jewett, et al. (1953), and Salo (1975).  These birds typically
occur in the channels and small bays in flocks of 20 to 30 individuals, often
in association with other diving ducks.  Drayton Harbor and Birch Bay in
northeast  Puget Sound appear to harbor the greatest wintering abundance of
any of the sites surveyed by the Washington Department of Game in north Puget
Sound and  the Strait of Juan de Fuca  (R.C. Parker, Wash. Dept. Game, unpubl.
data).  Kortright's (1942) analysis of 395 common goldeneye stomachs indi-
cated that crustaceans (32%) and insects (28%) were the major prey taxa,
followed by molluscs (10%) and plant  (pondweed 9%) remains.

Barrow's Goldeneye

     Generally thought less abundant than the common goldeneye, Barrow's
goldeneye  are similarly distributed as migrants and winter residents through-
out the north Puget Sound region (Munro 1939; Alcorn 1971; Salo 1975).
Populations wintering in the Strait of Georgia may be more numerous, as
evidenced by the 1976 Audubon Christmas bird  count (Heilbrum, et al., 1977),
wherein over 2,000 individuals were counted at Nanaimo, B.C., and 1,269 at
Vancouver, B.C.  Insects (36%), and especially molluscs (19%) and crustaceans

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(18%), are important food taxa for Barrow's goldeneye occupying marine
habitats (Kortright 1942; Salo 1975).

Bufflehead

     Bufflehead appear to be widely  distributed,  abundant  migrant  and winter
residents, characterizing shallow, sheltered  marine habitats (Munro 1942;
Jewett, et al., 1953; Erskine  1971;  Salo  1975).   They  appear to  be especially
concentrated in northeastern Puget Sound,  where  flocks of  3,800  to 12,800
birds are reported, and  Dungeness Bay,  where  peak abundances of  4,600 birds
are recorded in December (Salo 1975).   Abundances reported at Sequim Bay
peaked at over 300 during October-November and March but remained between
135 and 150 during the rest of the winter (D. Moriarity and S. Oliver,
Graysmarsh Wildlife Refuge, pers. comm.).   The maximum counts in the 1976
Audubon Christmas bird count were at Victoria (1,198), Bellingham (656), and
Dungeness Bay  (581)  (Heilbrum, et al.,  1977).  In marine habitats, crusta-
ceans, molluscs, and  to  a lesser extent,  small fish appear to be the most
important food items  (Munro 1942;  Salo  1975); herring eggs laid  in shallow
sublittoral waters may also be consumed when available in the spring.

Oldsquaw

      Also a migrant and  winter resident,  oldsquaw in Washington State occur
rather uniformly though  sparsely throughout north Puget Sound and the Strait
of  Juan de Fuca  (Kortright  1942; Jewett,  et al., 1953; Larrison and Sonnen-
berg   1968; Alcorn  1971; Salo  1975).  Kortright  (1942) listed crustaceans
 (48%), molluscs  (16%),  insects (11%),  and fish (10%) as the primary prey taxa
 found in  227  stomachs.   Like many of the other diving ducks, oldsquaw have
been  observed  foraging  on  the  eggs  of Pacific herring in the spring (Jewett,
 et  al.,  1953).

 Harlequin Duck

      Harlequins  appear  to be  one of  the few diving ducks which are year-long
 residents in the State,  although many of those inhabiting Puget  Sound in
 winter may include migrants (Jewett, et al.,  1953; Alcorn 1971;  Salo 1975).
 They occur in congregations of up to 200 birds along rocky shores of the San
 Juan Islands,  Dungeness  Spit  and Bay,  and along  the northern shore of the
 Olympic Peninsula.   Crustaceans (57%)  and molluscs (25%) composed the diet of
 64 harlequins examined  by Kortright  (1942, cited in Salo 1975).

 White-Winged Scoter

      A common and often the most abundant sea duck in winter in Washington's
 inland and coastal waters,  white-winged scoters  have been estimated to number
 up to 42,000 birds (Jewett,  et al.,  1953;  Alcorn 1971; Salo 1975).  Manuwal
 (1977) reported this species  as the  second most  abundant over all seven ferry
 transect surveys conducted between  November 1973 and April 1974  in the San
 Juan Islands.  They tend to be more  abundant  in  the eastern Strait of Juan de
 Fuca; approximately 400 are observed during October-November in Sequim Bay
 and they are also common at Dungeness Spit (D. Moriarity and S.  Oliver
 Graysmarsh Wildlife Refuge,  pers.  comm.;  1977 Olympic Pen. Audubon Soc!
 Christmas Count Newsletter,  unpub.).

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     One specimen collected in Grays Harbor had eelgrass in its esophagus
(Smith and Mudd 1976).  Cottam (1939) described the stomach contents of  819
white-winged scoters collected in the Pacific Northwest.  They contained 75%
bivalve molluscs (Manila clams, Tapes japonica, Olympia oysters, blue mussels,
and scallops), 13% crustaceans, and incidentally fish and insects.  In Puget
Sound they have been observed foraging over shallow-water commercial clam
and oyster beds and on herring eggs (Jewett, et al., 1953; Salo 1975).

Surf Scoter

     Surf scoters are generally similar to white-winged scoters in occurrence,
distribution, and abundance (Jewett, et al., 1953; Salo 1975).  According  to
the 1976 Audubon Christmas bird count, in fact, they were more numerous  at
Nanaimo (1,943), Vancouver (2,160), Victoria (361), Bellingham (417), and
Everett (58)  (Heilbrum, et al., 1977).  Salo (1975) included reports of
2,400 to 3,000 birds at Dungeness Spit between September and November, 40,000
at Destruction Island in October, and 1,200 in the San Juan Islands in
December.  Manuwal (1977) designated surf scoters as the sixth most numerous
diving species observed along his ferry transect surveys, winter 1973 to
spring 1974.

     The prey of surf scoters examined by Cottam (1939) was primarily
molluscs (61%, 29% of which were blue mussels), crustaceans (10%), and
insects (10%).  The stomachs of 21 surf scoters from Dabob Bay, Washington,
contained primarily Manila clams, but blue mussels and soft-shell clams, Mya
arenaria, and Macoma sp. clams also occurred (Glude 1967).

Black Scoter

     Black scoters appear to be less abundant than the other two scoter
species, and tend to be distributed more along the exposed coast (Salo 1975).
Black scoters were sighted during the 1976 Audubon Christmas bird count  in
the Strait of Georgia—e.g., Nanaimo (280) and Vancouver (342) (Heilbrum,
et al., 1977).  October and May appear to be the peak fall and spring
migration periods (Kortright 1942).  Cottam's (1939) analysis of the contents
of 124 black scoter stomachs indicated that 65% was molluscs (27% blue mussel,
7% razor clam, Siligua patula, 6% oysters, Ostrea lurida?, and 5% littleneck
clams), 17% crustaceans, and 3% insects.  Glude (1967) identified Manila
clams, blue mussels, and barnacles (Balanus sp.) in the stomachs of scoters
feeding in Dabob Bay, Washington.

Common Merganser

     Though common mergansers reside in terrestrial aquatic habitats through
the spring and summer, they congregate in marine and estuarine habitats
during fall and winter (Jewett, et al., 1953; Salo 1975).  Nanaimo, Fender
Islands, Victoria, and Vancouver, B.C., all had between 100 and 200 sightings
during the 1976 Audubon Christmas bird count (Heilbrum, et al., 1977) and
300 have been sighted at Dungeness Spit (Salo 1975).  Yet, little quantitative
diet information exists for the common merganser.  It is assumed to dive for
and consume small fishes, apparently including juvenile salmon (Salo 1975).
Small demersal fishes (cottids, pholids, stichaeids, pleuronectids) and

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schooling neritic fishes (Pacific herring, Pacific sand lance) may  also  be
consumed.

Red-Breasted Merganser

     The red-breasted merganser  is more of a migrant  and  winter  resident
than the common merganser.   It occupies north  Puget Sound waters between
September and April and reaches  maximum abundance in  November (Salo 1975).
As many as 125 birds were  seen forming one premigration flock at Dungeness
Spit in April and 141 birds  were observed there in December 1977 (1977
Olympic Pen. Audubon Soc.  Christmas  Count Newsletter, unpub.).  Fish (sculpin
and Pacific herring) and crustaceans are  important prey items for red-breasted
mergansers foraging in marine habitats, and  they have been observed to
concentrate on spawning Pacific  herring during February and March (Munro and
Clemens 1939; Kortright 1942; Salo  1975).

Osprey

     Ospreys do not appear to frequent marine habitats of Washington, except
in  the  San Juan Islands where they  occur  from April to October  (Retfalvi
1963; Bakus 1965).  Fishes are  generally  considered to be the basic prey of
ospreys  (Bent 1937; Salo 1975).

Bald Eagle

     Although they  do  not  actually  occupy the marine waters, bald eagles at
times  forage  in  the nearshore environment,  and thus periodically appear as
upper  trophic level consumers in those food  webs.  Of the 221 bald eagle
nests  reported  in a 1975  survey of  western  Washington (Grubb, et al., 1975),
47% (103 nests) were  located in the San  Juan Islands and 26%  (57 nests) were
 located in the  northern section of  the Olympic Peninsula; between 60% and 64%
 of these nests  were considered successful.   Despite increased human activity
 in the vicinity of  nests  on San Juan Island  reported by Retfalvi (1965),
 Newman, et al.  (1977),  reported a 100% increase in eagle nests between
 1962-63 and 1975.

      Although they commonly scavenge for  food, bald eagles raid seabird
 colonies, taking gulls, guillemots, cormorants, and puffins  (Campbell 1969;
 Salo 1975), and capture benthic invertebrates such as crabs  (Pugettia
 producta), sea urchins (Strongylocentrotus  sp.), and abalone  (Haliotis
 kamschatkana),  and fish,  including salmon (Oncorhynchus sp.), Pacific herring,
 Dolly Varden (Salvelinus malma), cutthroat  trout  (Salmo clarki), Pacific
 halibut  (Hippoglossus stenolepis). and sculpin  (Hawbecker 1958; Campbell
 1969; Ofelt 1975; Salo 1975).

 Great Blue Heron

      Commonly observed singly or in small groups in shallow waters of tide
 flats and estuaries, great blue herons are distributed throughout north Puget
 Sound and eastern Strait of  Juan de Fuca (Bakus  1965; Salo 1975).  In general,
 great blue heron prey on aquatic and estuarine animals, including  fish,
 amphibians, and crustaceans, but also insects and small terrestrial mammals

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(Martin, et al., 1951; Jewett, et al., 1953; Lowe 1954; Meyerriecks 1962).
Those studied by Krebs (1974) on the Fraser River delta near Vancouver, B.C.,
had consumed either staghorn sculpin or starry flounder (Platichthys
stellatus), these two species accounting for 67% of all prey.  Other
incidental prey were shiner perch, penpoint gunnel, and shrimp.  Although
great blue heron typically forage in shallow waters, one was observed by
Godin (1977) to effectively capture 5-9 cm long shiner perch while standing
on a platform floating over 10 m of water.

Virginia Rail

     These rather rare shorebirds are found on Pacific Northwest estuaries,
tideflats, and sand beaches in winter (Robbins, et al., 1966).  Heilbrum,
et al.  (1977), included reports of one Virginia rail each at Vancouver and
Victoria, B.C., during the December 1976 Audubon count.  Little information
is available on their diet; it is assumed that, as in the diets of other
shorebirds, shallow-water crustaceans may be important.

American Coot

     Coots are common resident birds in Washington and British Columbia which
winter in sheltered marine habitats (Jewett, et al., 1953; Alcorn 1971; Salo
1975).  Large aggregations have been reported throughout north Puget Sound
and the Strait of Juan de Fuca, especially in the semi-urban areas near
Vancouver and Victoria, B.C., and Bellingham, Washington (Salo 1975; Heilbrum,
et al., 1977).  The stomach of one American coot collected in Grays Harbor by
Smith and Mudd (1976) contained eelgrass and a small amount of algae.

Black Oystercatcher

     Black oystercatchers are common residents of the coastal regions of
western Washington, usually frequenting the rocky exposed  shoreline and
nesting on the islands and islets along the Pacific coast  of the Olympic
Peninsula and in the San Juan and Gulf islands (Salo 1975).  Eighty to 100
pairs were estimated to nest on the coastal islets of the  Washington Islands
National Wildlife Refuge, 80 pairs in the San Juan Islands, and approximately
90 pairs in the Gulf Islands.  Ninety-two to 100 breeding  pairs were estimated
for the outer coast and 25 to 34 for the San Juan Islands  and the Strait of
Juan de Fuca in Manuwal's (1977) survey.  Principal nesting locations included
Long, Protection, and Smith islands.

     Marine gastropods and bivalves, e.g., limpets, mussels, and chitons, of
rocky littoral habitats constitute the common prey of black oystercatchers
(Jewett, et al., 1953).  Webster (1941) estimated that their diet in south-
eastern Alaska was composed of 30% ribbed limpets, Collisella digitalis; 20%
blue mussels (Mytilus edulis; 15% California mussels, M. californianus; 15%
gooseneck barnacles, Pollicipes polymerus; 13% shield limpets, Notoacmea
scutum; 5% black chitons, Katharina tunicata; 1% duncecap  limpets, Acmaea
mitra; and polychaete annelids, Nereis sp.  Hartwick (1973) indicated that
while mussels formed the greatest part by weight of the oystercatcher's diet,
crabs were of importance when chicks were present.
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Semipalmated Plover

     Salo (1975) considered the semipalmated plover a common migrant  in
western Washington, frequenting sandy beaches and tideflats during  spring
(April-May) and fall  (September-October).  Large flocks of up to  1,000 birds
have been reported along the  eastern end of the Strait of Juan  de Fuca and  in
the San Juan Islands  during these  periods.  Wintering birds, however, may be
uncommon.  Heilbrum,  et al.  (1977), reported only two birds during  the 1976
Audubon Christmas bird count  in the San Juan Islands and  the Strait of Juan
de Fuca.  Bent  (1929) listed  littoral  invertebrates (molluscs  such  as
Littorina sp. and Mytilus  sp. and  crustaceans  such  as Gammarus  sp., Limnoria
sp., and Orchestria sp.) and  insects as the principal food  organisms of
semipalmated plovers.

Whimbrel

     Whimbrels  apparently  are common only during  their  fall (August) and
spring  (May) migration periods, when 100  to  170 birds have  been counted  in
the Washington  Islands National Wildlife  Refuge and the San Juan Islands
(Salo 1975).  Bent  (1929)  and Jewett,  et  al.  (1953),  described the  diet  of
wintering whimbrels as including  polychaete  annelids, molluscs, and
crustaceans.

Spotted  Sandpiper

     Although some spotted  sandpipers winter  in  Washington, most are summer
residents which breed in the  region  (Jewett, et al.,  1953;  Alcorn 1971;  Salo
1975).   When present  in marine habitats,  principally  sand and  gravel beaches
and mudflats, spotted sandpipers  eat mostly benthic invertebrates,  including
polychaete  annelids,  gammarid amphipods,  and molluscs.

Wandering Tattler

     Jewett, et al.  (1953),  and Salo  (1975) consider  the  wandering  tattlers
as  locally  common migrants, most  often observed in  rocky  shoreline  habitats
of  the  exposed  coast, the  Strait  of Juan  de Fuca, and the San  Juan  Islands.
Their diet  according  to  Bent  (1929) includes small  molluscs, crustaceans,
polychaetes, and  insects.

 Short-Billed Dowitcher

     Although migrants to  Washington,  short-billed  dowitchers  are commonly
 observed along  the  coast and  through the  Strait of  Juan de  Fuca to  the San
 Juan Islands.   The peak  spring migration  is in April  and  May and the peak of
 the fall migration  is in September and October (Larrison  and Sonnenberg  1968;
 Salo 1975). They feed  in  small flocks of  10 to 50  birds  in sand and mudflat
 habitats,  extracting  marine annelids,  flatworms,  small  molluscs, fish eggs,
 and occasional  plant  matter from  the shallow waters (Salo 1975).

      Sperry's   (1940)  analysis of  the stomach contents of  191 short-billed
 dowitchers, collected mostly from the  Atlantic coast  of the United  States
 indicated that  insects  (29.1%, including  18% dipteran larvae,  primarily   '

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Stratiomyiidae, Empldidae, and Tabanidae, and 8.9% Coleoptera, including
aquatic Hydrophilidae and Dytiscidae), marine annelids  (27.4%, including
Nereis sp.), and marine molluscs  (20.9%, including Melampus sp. and
nassariid snails among the gastropods; and Gemma gemma, Ostrea virginica,
Mytilus edulis, and Venus mercenaria among the bivalves) were the principal
components of their diet.  Crustaceans, including crabs (Uca sp.), shrimp
(Crago sp.), isopods  (Cynthura carinata). ostracods, and amphipods, and
horseshoe crab (Limulus polyphemus) eggs were also found to a lesser degree
in the stomach contents of short-billed dowitchers.

Long-Billed Dowitcher

     Although considered less common than short-billed dowitchers, migrant
long-billed dowitchers have been observed in great numbers in the coastal
embayments of Washington and are assumed to occur frequently in the San Juan
Islands (Salo 1975).  Small flocks may overwinter in the region, as evidenced
by the 1976 Audubon Christmas bird count (Heilbrum, et al., 1977) which
included 17 individuals observed at Victoria, B.C.


     Jewett, et al. (1953), and Salo (1975) assumed that long-billed
dowitchers feed upon coastal marine invertebrates—e.g., annelids, small
crustaceans, and molluscs—and some plant matter.  The majority of the 107
stomach samples examined by Sperry (1940) were collected in inland areas,
but marine annelids (Nereis sp.), crustaceans (amphipods, including Echino-
gammarus ochotensis and Corophium spinicorne, and the isopod Exosphaeroma
oregonensis), and molluscs (gastropods, including Littorina sitchana) were
included as prey items.  Smith and Mudd (1976) collected only a few specimens
in Grays Harbor.  Examination of the esophagus of one specimen indicated
amphipods (Corophium sp.) and polychaetes were important prey but further
examination of the gizzard (which biases toward hard-food organisms) suggested
that clams (Macoma inconspicua and Mya arenaria) were also important.

Surfbird

     Surfbirds migrate into western Washington in August and reside on rocky
shores and gravel beaches until April  (Jewett, et al., 1953; Salo 1975).
They are most common along the exposed coast and rarer around Puget Sound.
The 1976 Audubon Christmas bird count  (Heilbrum, et al., 1977) in this region
recorded surfbirds in abundance only at Nanaimo, B.C.  Crustaceans and small
molluscs of the rocky littoral zones are the prey of surfbirds (Bent 1929).

Ruddy Turnstone

     Ruddy turnstones, though fairly common migrants along the exposed coast,
are uncommon inside the Strait of Juan de Fuca and rarely over-winter in this
region (Salo 1975).  Their diet resembles that of the sandpipers, i.e.,
polychaetes, molluscs, and crustaceans of beaches and mudflats (Bent 1929;
Jewett, et al., 1953; Salo 1975).
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Black Turnstone

     Black turnstone is much more common than its congener ruddy  turnstone
and is locally abundant in northern Puget Sound and the Strait  of Juan de
Fuca as a migrant and winter resident  (Jewett, et al., 1953;  Salo 1975).
Heilbrum's, et al.  (1977), account of  the 1976 Audubon Christmas  bird count
listed black turnstones at almost all  stations in this region,  but the
highest counts were at Victoria  (212)  and Vancouver  (147), B.C.  Molluscs
(limpets), barnacles, and other  crustaceans  common to the rocky littoral zone
are the basic prey  items of black turnstones (Bent 1929;  Jewett,  et al.,
1953).

Greater Yellowlegs

     Migrating greater yellowlegs occur in  the region in April-May and July-
August, and are most  common  in northern Puget Sound  (Jewett,  et al., 1953;
Salo 1975; Heilbrum,  et al.,  1977).   The highest count  of overwintering birds
during the 1976 Audubon Christmas bird count was 34  at  Victoria,  B.C.  Jewett,
et al.  (1953), described their food  as including molluscs, crustaceans, and
small fish.

Rock Sandpiper

     Larrison and Sonnenberg  (1968)  described rock sandpipers as uncommon to
rare spring and fall  migrants, although there may be some casual winter
residency  (Alcorn 1971).  Although  little is known of their diet, small
crustaceans, molluscs, and  insects  are assumed to be important (Eaton 1975).

Dunlin

     Dunlins are  often one  of  the most common shorebirds encountered on the
coast of Washington,  especially  on  the region's extensive mudflat habitat.
Dunlin  dominated  the  shorebirds  (almost 2,800 birds  counted)  at Dungeness
 Spit during  the 1977  Audubon Christmas bird count (1977 Olympic Pen. Audubon
 Soc. Christmas  Count  Newsletter, unpub.).

     Couch (1966) reported  that  wintering dunlins in northern Puget Sound
 consumed mostly amphipods,  especially Anisogammarus  confervicolus at low tide
 and Orchestoidea  pugettensis at  high tide.   Benthic  invertebrates, mainly
 amphipods  (69.7%  of total number of identifiable prey,  including Corophium
 sp.,  52.5%;  Anisogammarus  confervicolus, 16.5%;  and Amphitoe sp., 0.7%),
 polychaete annelids (8.5%  of identifiable prey,  including Nereis sp.,
 Streblospio  benedicti,  Amphicteis mucronata, and Eteone longa), oligochaete
 annelids (3.2%),  molluscs  (1.9%, including Macoma inconspicua and Mya
 arenaria), tanaids (0.7%,  Pancolus  californiensis),  and unidentified
 nematodes (1.5%), were  the principal prey of dunlins wintering  in Grays
 Harbor (Smith and Mudd 1976).   It was also determined that during high tide,
 dunlins supplemented the invertebrate-based diet with seeds of the saltmarsh
 plants Spergularia marina and Triglochia maritinum.   Few differences were
 noted in diet composition among  five sampling sites.
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     Recher's (1966) analysis of the contents of 46 stomachs of dunlin
collected in California documented the nereid polychaete Nereis diversicolor
as the principal prey (70% frequency of occurrence), followed by ostracods
and amphipods.  Studies by Bengtson and Svensson (1968) in southern Sweden,
and Ehlert (1964, cited in Kawaji and Shiraishi 1979) in the Baltic Sea
(Helgoland) illustrated generally the same preference for Itf. diversicolor.
The data provided by Madon (1935, cited in Wolf 1969) indicated that in
addition to ^J. diversicolor, molluscs, crustaceans, insects, and vegetable
matter were also important in the stomachs of dunlin along the Atlantic
coast of France.  The Wild Bird Society of Japan (1975, cited in Kawaji and
Shiraishi 1979) and Kawaji and Shiraishi (1979) found that the polychaete
annelid Neanthes japonica; the gastropod molluscs Fluviocingula nipponica,
Salinator takii, and Ellobium chinense; and the bivalve mollusc Musculus
senhousia supplied the principal food items in nearshore regions of Japan.
Davidson (1971) showed a similar diet composition for dunlin collected in
Morecambe Bay, Lancashire, England.

     The only deviation from the above documentation of benthic polychaetes
and molluscs as principal prey was that of Holmes's (1966) description of
dunlin food habits during its breeding season in arctic Alaska, when it
utilized larval and adult dipteran insects.

Knot
     Both the American knot  (Calidris canutus rufus) and the Pacific knot
 (C.c^ rogersi) are found in Puget Sound infrequently (Alcorn 1971; Jewett,
 et al., 1953; Salo 1975).  When observed they are often in large flocks on
 tidal flats and along the sand beaches of exposed coasts.  The stomachs of
 219 American knots collected from the east coast of the United States  (Sperry
 1940) contained, by volume, 59% molluscs (mainly Nassariidae, Littorina sp.,
 and Melampus sp. among the univalves, and Donax sp., Gemma gemma, Mytilus
 edulis, and Modiolus demissus among the bivalves); 14.8% insects (including
 12.7% Diptera, i.e., Stratiomyiidae, Ephydridae, Tabanidae, Empididae, and
 Tipulidae; 1.2% Coleoptera, i.e., Hydrophilidae; and 0.9% other insects, i.e.,
 Formicidae, Pentatomidae, and Cydnidae); 8.9% crustaceans (including crab
 eggs); 2.2% fish, marine annelids, and other incidental animals; and 15.2%
 plants (primarily seeds of widgeongrass and other estuarine plants).

 Least Sandpiper

     Least sandpipers, common migrants and occasional winter residents in
 this region, are quite abundant along the outer coast but also occur
 frequently at Dungeness Spit and in the San Juan Islands (Jewett, et al.,
 1953; Alcorn 1971; Salo 1975).  April and May mark the spring migration,
 August-September, the fall migration.  Winter residents may be uncommon in
 northern Puget Sound as few were reported during the 1976 Audubon Christmas
 bird count (Heilbrum, et al., 1977).  The principal food of least sandpipers
 feeding on beaches and mudflats is the amphipod Anisogammarus confervicolus
 (Couch 1966).  Sanger (1970) indicated that elevated mudflats were preferred
 foraging areas at high tide, whereas sandier mudflats were utilized during
 low tide.
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Western Sandpiper

     Western sandpipers are common spring and fall migrants  in north  Puget
Sound, occurring in April and May and again  in July  through  December.   Small
flocks are especially common in the  San Juans and at Dungeness Spit  (Jewett,
et al., 1953; Alcorn 1971; Salo 1975; Heilbrum,  et al.,  1977).

     Smith and Mudd  (1976) listed a  wide variety of  prey items of western
sandpipers wintering in Grays Harbor.  Polychaete annelids (24.4%, including
Streblospio benedicti and other unidentified spionids  and Eteone longa),
nematodes  (13.3%), amphipods  (8.9%,  Euhaustorius washingtonius), adult and
larval insects  (6.7%), and tanaids  (2.2%,  Pancolus  californiensis) composed
the animal prey  items, whereas  saltmarsh  plant  seeds (40%, primarily Spergu-
laria marina but also including Triglochia maritimum and Deschampsia
cespitosa) were  the  main  diet when  the birds fed at high tide.   Western
sandpipers collected from the  San Juan Islands  and  other Pacific Northwest
sites had  eaten  mostly amphipods,  including Anisogammarus confervicolus and
Corophium  sp.  (Couch 1966).

Sanderling

      Sanderlings are common migrants and  winter residents which form large
 (e.g., 5,000 birds)  flocks on  the outer  coast but  smaller   (e.g., 50) flocks
in north Puget  Sound (Jewett,  et  al.,  1953;  Alcorn 1971; Salo 1975).   They
were  especially  numerous  at Bellingham and Dungeness Spit during the 1976
Audubon Christmas  bird count  (Heilbrum,  et al.,  1977).  Amphipods (Anisogam-
marus  confervicolus,  Orchestoidea pugettensis)  and  bivalve molluscs (razor
clams  on the outer coast)  are  their  usual prey  organisms (Cough 1966).

Wilson's Phalarope

      Wilson's  phalarope  is a  casual  summer migrant  and resident  which
migrates  to  the  area in  May and leaves in August (Jewett, et al., 1953;
 Alcorn 1971;  Salo  1975).   Jewett,  et al.  (1953), listed insects  and their
 larvae as  the  major food  of  sanderlings,  although  small crustaceans and
 plants may also  be consumed  (Bent 1927).

 Northern  Phalarope

      A much more common  migrant  than Wilson's phalarope, the northern phala-
 rope occurs  in high numbers  (10,000-17,000 birds)  in northern Puget Sound
 in May and August  as well as  offshore (Jewett,  et  al., 1953; Salo 1975).
 While insects  appear to  form much of the  diet (80%), small crustaceans (9%),
 polychaete annelids, small molluscs  and  fish, and  plant material also
 occurred in their  diet  (Bent  1927).

 Parasitic Jaeger

      Parasitic jaegers  occur  in Washington's marine habitats frequently
 during their spring (April-May)  and  fall  (July-December) migration through
 the region (Jewett,  et  al.,  1953; Alcorn  1971;  Salo 1975).  Since they
 commonly obtain food by  harassing gulls  and terns  into dropping  or disgorging

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their food, parasitic jaegers are ultimately linked to the food resources of
gulls and terns (Salo 1975).

Pomarine Jaeger

     Little information is available on the distribution or relative
abundance of pomarine jaegers in the region.  Guiget  (1971a) states that they
appear offshore of Vancouver Island, B.C. in small numbers in mid-July
increasing through September; occurrence in inland waters is apparently
scarce.  Although they capture small birds and rodents, when in marine
environments they appear to gain most of their food by robbing gulls, terns,
and other marine birds.

Glaucous-Winged Gull

     Glaucous-winged gulls have been described as the most commonly observed
gull in nearshore areas along the Washington coast (Sanger 1965) but are
encountered in abundance offshore only during November and January (Sanger
1970).  Manuwal's (1977) survey of the breeding seabirds in Washington State
included an estimate of 6,234 breeding pairs in the San Juan Islands and the
Strait of Juan de Fuca; the largest populations included colonies on Colville
and Protection islands.  They were the most prominent gull species surveyed
at Dungeness Spit in December 1977, totaling almost 600 birds (Olympic Pen.
Audubon Soc. 1977 Christmas Count Newsletter, unpub.).

     Outram (1958) listed glaucous-winged gulls as one of the major causes
of herring egg mortality; the stomachs of 12 gulls that he examined contained
an average of 13,800 eggs.  Fish, primarily capelin (Mallotus villosus)
occurred in the stomachs of 33% of the glaucous-winged gulls collected in
Alaskan waters by Sanger and Biard (1977).  Pollock averaging 20 cm in length
were found in the stomachs of gulls associated with the Bering Sea pack ice
in April (Divoky, et al., 1977).  James-Veitch and Booth (1954) listed chitons,
sea cucumbers, sea urchins, crabs (Cancer sp., Hemigrapsus sp.), blue mussels,
and polychaete worms (Nereis sp.) as prey of foraging glaucous-winged gulls.

Western Gull

     Western gulls are the most common residents of Washington's outer
beaches and non-breeding gulls are common through the Strait of Juan de Fuca
and northern Puget Sound (Jewett, et al., 1953; Alcorn 1971; Salo 1975).
Hunt and Hunt (1976) examined the food items of western gull chicks on Santa
Barbara Island, California.  They found that 89% of the food was composed
mostly of schooling fishes, and included northern anchovy (45% frequency
occurrence), jack mackerel  (Trachurus symmetricus, 5.2%), Pacific saury
(Cololabis saira, 2.8%), midshipman (Porichthys sp.,  1.9%), and squid.

Herring Gull

     The common, ubiquitous herring gull occurs in every habitat of northern
Puget Sound and the Strait of Juan de Fuca and is especially well adapted to
urban coastal areas (Jewett, et al., 1953; Salo 1975).  Verbeek (1977)
established that in the Cumbria, England, area, immature herring gulls relied

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more on stealing than on foraging, until they developed feeding  skills,
whereupon starfish became the principal food item.  Herring  gull predation
upon spawned herring eggs in British  Columbia was reported by Outram (1958),
who documented an average of 8,500 eggs per stomach for the  nine stomachs
examined.

California  Gull

     California  gulls  occur in western Washington in abundance during their
fall and spring  migrations  and small numbers overwinter in Puget Sound
 (Jewett,  et al., 1953;  Alcorn 1971;  Salo 1975).  Though observed frequently,
they were not  very abundant during the 1976 Audubon Christmas bird count
 (Heilbrum,  et  al.,  1977).   As many as 1,000 have been recorded at Dungeness
Spit in August (Salo 1975).  Though the California gull apparently is as
omnivorous  as  the other gulls (Jewett, et al., 1953; Salo 1975), it also
preys  specifically on juvenile sea or shore birds (Chura 1962).

Ring-Billed Gull

     Ring-billed gulls are common winter residents in western Washington and
non-breeding birds may be found throughout the summer (Alcorn 1971; Salo
 1975;  Eaton 1975).   Like the other large gulls, ring-billed  gulls consume
 almost anything available and edible, including small fish,  refuse, insects,
and small rodents and birds (Guiguet 1971a; Salo 1975; Eaton 1975).

Mew Gull

     Mew gulls are common winter residents along the Pacific Northwest coast
 from October to April.   Despite its common occurrence in this region, few
 data on its food habits exist.  In Alaskan waters they feed  mainly on
 unidentified crustaceans (83% frequency occurrence) and much less on fishes
 (17%)  (Sanger and Baird 1977), but are documented as feeding primarily on
 small  fishes when wintering in north Euget Sound and the Strait  of Juan de
 Fuca (Guiguet 1971a).

 Heermann's  Gull

     As a common summer migrant, Heermann's gulls are most abundant in Puget
 Sound  from August to October (Alcorn 1971; Guiguet 1971a; Salo  1975).  School-
 ing neritic fishes, such as Pacific herring, or those of kelp beds are the
 most common food organisms of Heermann's gulls, but they are also known to
 parasitize Bonaparte's gulls and cormorants (Guiguet 1971a;  Salo 1975).

 Bonaparte's Gull

      Bonaparte's gull occurs in Washington's marine waters primarily as a
 migrant and winter resident, reaching peak abundance in October  (Salo 1975).
 Counts in the San Juan Islands National Wildlife Refuge included approximately
 7,000 in April and up to 15,000 in August.  Wahl (1977) reported that after
 high abundances in October and November,  only a few hundred  were seen at
 Active Pass, B.C., until April and May when the northward-migrating eulls
 passed through.


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     Little detailed information exists on the diet of Bonaparte's  gull.
Jewett, et al. (1953). recorded that at times insects form the majority of
its prey.  Salo (1975) suggested that "small fish and other marine  life"
generally composed the diet.  Guiguet (1971a) reported that small Pacific
herring and Pacific sand lance, forced to the surface by feeding salmon,
were fed upon by Bonaparte's gulls.

Sabine's Gull

     Sanger (1965) reported one or two Sabine's gulls within  five miles of
the Washington and Oregon coast during September and October, but it  is
doubtful that they frequent the inshore waters of the Strait  of Juan  de Fuca
or northern Puget Sound  (D. Manuwal, pers. comm.).  Guiguet (1971a) reported
that Sabine's gulls feed upon crustaceans, small fishes, and  marine annelid
worms.

Black-Legged Kittiwake

     Kittiwakes winter well offshore the coasts of Washington and British
Columbia but may occasionally be encountered inshore, especially during
storm periods (Gabrielson and Jewett 1940; Jewett, et al., 1953; Sanger 1970).
Sanger and Baird (1977) examined the stomachs of 21 of 76 specimens collected
in Alaskan waters between 1969 and 1976.  Fish (Pacific sand  lance, 43%
freq. occur., and Pacific herring, 14%) occurred in 76% of these stomachs,
crustaceans (the hyperiid amphipod  Parathemisto libellula, 29%, and  the
euphausiid Thysanoessa raschi, 14%) in 19%, and squid in 5%.

     Both fish and invertebrates entered the diet of black-legged kittiwakes
collected in the Cape Thompson vicinity of Alaska (Springer and Roseneau
1977).  Fishes included polar cod  (Boreogadus saida, 33% freq. occur.),
saffron cod (Eleginus gracilis, 33%), flatfish (7%), and ninespine  stickle-
back (Pungitius pungitius, 7%); and invertebrates included gastropods
(Trochidae, 13%), crabs  (13%), nereid polychaetes (7%), shrimp (Pandalus
goniurus, P_. montagui, Pandalus sp., and Eualus gaimardi, 7%  each), isopods
(Saduria entomon, 7%), and insects (7%).

     Fish (principally cod and capelin) and euphausiids (Thysanoessa  inermis,
T_. longipes, T_. raschi, and T_. spinifera) were the main prey  items  of kitti-
wakes collected in the Pribilof Islands (Hunt 1977).  Birds feeding along the
southern edge of the Bering Sea icepack in March and April had consumed
capelin and walleye pollock (Theragra chalcogramma)  (Divoky,  et al.,  1977).
The diet of black-legged kittiwakes of the Fame Islands in Great Britain
was almost completely composed of  fish (98% prey abundance, 99% biomass),
primarily sand lance  (Ammodytidae), herring  (Clupeidae), and  cod  (Gadidae).

Common Tern

     Whereas common terns migrate  rapidly along the Pacific coast in  the
spring, the fall migration during  August to October brings them to  the coastal
and Strait of Juan de Fuca areas  (Jewett, et al., 1953; Alcorn 1971;  Salo
1975).  Salo  (1975) reported 250 common terns observed at Dungeness Spit  in
August.  Common terns feed mainly  on small neritic fishes  (Salo 1975) which

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in this region would include Pacific herring, Pacific  sand  lance,  smelt
(Osmeridae), and juvenile salmon.

Arctic Tern

     Arctic terns are common migrants  along the Pacific Northwest coast and
in north Puget Sound  (Guiguet  1971a).   A small  breeding colony (7-12 pairs)
has been reported to occur on  Jetty Island near Everett, Washington
(D. Manuwal, pers. comm.).  Crustaceans (unidentified) occurred in 100% of
the stomachs of birds collected  by Sanger and Baird (1977)  from Alaskan
waters between 1969 and  1976.  Pacific sand lance and herring composed the
majority of the prey abundance (87%) and biomass (85%) of arctic terns in the
Fame  Islands of Great Britain (Pearson 1968).

Caspian Tern

     Although at the  approximate northern extremity of their West Coast
distribution, Caspian terns are  common, both as fall migrants and as breeding
birds  (approximately  2,500),  on  the outer Washington coast (Robbins, et al.,
 1966;  Eaton 1975).   Smith and Mudd (1976) reported the species composition
 of 31  fish removed  from  a Grays  Harbor ternery.  Shiner perch composed over
 half of  the fish collected; juvenile chum salmon (Oncorhynchus keta) and
 Pacific  staghorn  sculpin were also common; white seaperch  (Phanerodon furca-
 tus).  whitebait  smelt (Allosmerus elongatus), snake prickleback, cutthroat
 trout, and longnose dace (Rhinichthys  cataractae) were also found.  Bent
 (1921) also suggested shrimp,  mussels, and eggs and young of other birds as
 prey items.

 Common Murre

      Common murres breed in large concentrations in crevices or on steep
 cliffs, on the large stacks and islands of the outer coast, especially around
 LaPush, Washington (Cody 1973),  and are very abundant in the Strait of Juan
 de Fuca from August through October.  Manuwal  (1977) documented 11,950 pairs
 of common murres nesting on the outer coast of Washington.   The largest
 breeding populations were located in the Point Grenville and Willoughby Rock
 vicinities.

      The best documentation of common murre prey spectrum is in Scott's
  (1973) study at Yaquina Head, Oregon.   His results substantiated earlier
 reports by Belopol'skii (1957) and Bedard  (1969) that common murres feed
 mainly on midwater fishes.   Scott's study indicated that eulachon (Thaleich-
 thys pacificus) and northern anchovy were the most important prey of the
 regon.  Juvenile rockfish became a major constituent of the diet of fledg-
 lings after they had left the breeding rocks.  In one year, however,
 epibenthic crustaceans  (euphausiids, mysids) constituted 86.2% of the total
 prey volume, a result that Scott attributed to the lack of midwater schooling
 fishes in the area at that time.  Using Scott's data, Wiens and Scott  (1975)
 calculated the total animal energy demand of common murres off Oregon to be
 3.488 k cal nr2,  the highest of the four species they examined.
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     Steele and Drury (1977), Hunt  (1977), and Sanger, et al.  (1977), also
documented the importance of midwater fishes in the prey spectrum of common
murres in Alaskan waters.  The principal prey species were capelin, Pacific
sand lance, walleye pollock, a prickleback (Lumpenus fabrici), and several
other unidentified cod and smelt.   Fish, primarily Pacific sand lance,
occurred in 77% of the stomachs examined from Alaskan waters by Sanger and
Baird (1977); crustaceans (unidentified) occurred in only 15%.

     Common murres in the Cape Thompson vicinity of Alaska appeared to
utilize primarily polar cod  (42% freq. occur.), saffron cod  (33%), Pacific
sand lance (17%), and unidentified  sculpins  (17%) (Springer and Roseneau
1977).  Murres collected over the Bering Sea pack ice in March by Divoky,
et al. (1977), had consumed  7.5-18  cm long pollock and 10-14 cm long capelin,
but the hyperiid amphipod Parathemisto sp. occurred the most frequently in
the stomach, though providing less  than one percent of the total prey
volume; euphausiids appeared more often (57% freq. occur., 20.4% volume) in
April.

     Tuck's  (1960) summary analysis of the diet of North Atlantic murres
also confirmed that they eat midwater schooling fishes, specifically gadids
(cod and haddock), sand lance, herring, and capelin, but also included such
diverse benthic fishes as sculpin and flatfish.  Although they were not a
significant proportion of the total prey volume, a variety of invertebrates
(shrimp and cephalopod molluscs) also occurred in the diet.  Common murres
in the Pembrokeshire region  of Great Britain were observed to feed their
chicks sand lance and small  herring (Clupea sprattus) in the 50-175 mm size
range (Harris 1970).

Pigeon Guillemot

     Pigeon guillemot is ubiquitous throughout the region, breeding in small
colonies on high cliffs both offshore and inshore (Cody 1973).  Manuwal (1977)
estimated 161 breeding pairs along  the outer Washington coast and 194 pairs
in the San Juan Islands and  the Strait of Juan de Fuca; prominent colonies
in the latter region include Castle, Protection, Skipjack, and Smith islands.
They were tenth in total abundance  of the diving birds observed during the
San Juan Island ferry transect surveys.

     In Cody's (1973) ecological analysis of Washington's alcid communities,
the pigeon guillemot's diet  is distinguished from the other alcids1 diets by
its dependence on  shallow sublittoral fishes, specifically "blennies"
(Pholidae and Stichaeidae?), and clingfish (Gobiesocidae).  Cody's analysis
was supportive of earlier, more general reports, including Drent  (1965) who
reported that over 70% of the diet  of pigeon guillemot on Mandarte Island,
B.C., was composed of blennies, flatfish, and sculpins, and Thoresen and
Booth (1958), who listed Pacific sand lance, surf smelt (Hypomesus pretiosus),
black prickleback (Xiphister atropurpureus), snake prickleback, and small
flatfish as the principal prey fed  to nestlings.  Fourteen of 16 prey items
consumed by pigeon guillemots in Yaquina Bay, Oregon, were flatfish; the
other two were blennies (Scott 1973).  The same study also mentioned one
flatfish, eight planktonic crustaceans, and two crabs as the prey found in
two adults collected offshore.  Both fish and crustaceans  (euphausiids,

                                    246

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Thysanoessa inermis) were represented in the stomachs of pigeon  guillemots
collected in Alaskan waters between 1969 and 1976 by Sanger  and  Baird  (1977).

Tufted Puffin

     The only puffin which occurs  in north Puget Sound  and the Strait  of Juan
de Fuca and which  plays  any  significant role in the food web is the tufted
puffin  (Lunda cirrhata).   It is particularly abundant  along Washington's
northwest coast  where  it nests in colonies on the larger stacks and islands
such as Carroll  Island,  Alexander Island,  and Cake Rock (Cody 1973).  One
population  estimate for  the  outer coast region was 7,343 nesting pairs
(Manuwal 1977).   Tufted  puffin do not,  however, nest in abundance inside the
Strait  of Juan de Fuca,  as Protection Island which has the largest breeding
colony  in the region has only 25 to 30 nesting pairs (Manuwal 1977).

     Cody's (1973) comparative study of the alcids indicated that primary
prey delivered  to nestlings  by adult birds were, in decreasing order of
importance,  Pacific sand lance, northern anchovy, rockfish, and smelt.
Sealy  (1973)  also listed Pacific sand lance as the major prey in June
delivered by nesting adults.  Manuwal  (1977), however, did not identify
Pacific sand lance as an important prey, rather smelt  (Hypomesus?), Pacific
herring, and northern anchovy.  Manuwal (1977) and Bent (1929) also suggested
that feeding adults have a broader prey spectrum that  includes crustaceans,
mussels, sea urchins,  surf smelt, Pacific sardine  (Sardinops sagax), Pacific
herring, seaperch (Embiotocidae), Pacific sand lance,  northern anchovy, and
rockfish.

     Amaral (1977) and Manuwal and Boersma (1977) reported  94.5% of the prey
delivered  (89.5% freq. occur.) to tufted puffin chicks on the Barren Islands,
Alaska, was capelin;  prowfish (Zaprora silensus) and squid  also occurred but
were insignificant.  Sanger  and Baird  (1977), however, noted that fish and
squid  occurred  approximately equally in their collections of tufted puffin
in Alaskan  waters between 1969 and 1976; Pacific sand  lance was the only
fish  species identified  from the stomach contents.  Sanger, et al.  (1977),
listed capelin,  Pacific  sand lance, walleye pollock, and invertebrates as
the principal prey of tufted puffin in the Kodiak Island area.  Cod and
capelin composed all of  the  prey of tufted puffin sampled by Hunt  (1977) on
 St.  Paul  Island  in the Pribilofs in 1976,  but a year earlier the diet had
been volumetrically dominated by unidentified crabs.

 Rhinoceros  Auklet

      More  than  11,000 breeding pairs of rhinocerus auklets were estimated
 along  the  outer  coast, principally at Destruction Island, while 9,800 pairs
 were estimated  for the San Juan Island and Strait of Juan de Fuca region,
 these  breeding  mainly at Protection Island (9,200) and Smith Island (600)
 (Manuwal 1977).   As one  of the most abundant seabirds  occurring in the Strait
 of Juan de Fuca and the  outer northwest Washington coast, rhinoceros auklets
 of this region have been studied extensively over the  past few years,
 especially their diet (Richardson 1961; Cody 1973; Leschner 1976; Manuwal
 iy//;  Wilson 1977).  In  most cases, these  data were collected for breeding
                   f°°d f°r nestlin8s'  mainly on Destruction and Protection

                                     247

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     Richardson's (1961) documentation of prey delivered to nestlings on
Protection Island listed 4-6 inch long Pacific sand lance as the main prey
item, the only other fish being a surf smelt.  The bill loads averaged 6.4
Pacific sand lance (ranging from 1 to 13 fish) and included extremely fresh
fish, suggesting feeding grounds within 10 miles of Protection Island.

     Annual variation was most obvious in the data obtained by Leschner
(1976) from Destruction Island.  Northern anchovy accounted for 56.0%,
Pacific herring, 20.8%, and surf smelt, 15.3%, of the prey delivered to
nestlings in 1974.  In 1975, night smelt (Spirinchus starksi) and Pacific
sand lance predominated (31.9% and 31.7%, respectively), followed by northern
anchovy (26.8%).  Species composition also varied considerably over the
nesting period.

     The comprehensive study by Wilson (1977) indicated that Pacific sand
lance and Pacific herring were the most important prey of rhinoceros auklets
nesting on Protection Island in 1975 and 1976, composing 90.6% of the total
weight of 1,198 prey items over that time.  Juveniles of all four species of
Pacific salmon, adult northern anchovy and surf smelt, juvenile walleye
pollock, and threespine stickleback occurred to a lesser extent.  Prey
species composition reported by Wilson varied less during the nesting period
between 29 June and 29 August 1976 than the prey composition reported by
Leschner (1976).  In Wilson's study, juvenile salmonids declined slightly
and Pacific herring increased slightly over that period.  Peak utilization
of Pacific sand lance occurred at the end of July, coinciding with the peak
abundance of chicks being fed on the island.  Wilson also documented an
average daily consumption rate for nestling rhinoceros auklets of 54-55 g
Pacific herring per day.

     Manuwal's  (1977) studies provide the most extensive data on rhinoceros
auklet prey items and quantities.  Although there was some annual variation
between the two years sampled, prey delivered to nestlings on Protection
Island were mostly Pacific sand lance and herring in terms of both numbers
and biomass, augmented by juvenile salmon and several other neritic fishes
(Table C-3).  This prey spectrum was quite dissimilar to the spectrum at
Destruction Island on the outer coast, where northern anchovy and night smelt
were more important, and where annual variation in prey composition was more
pronounced.  Prey composition based on weight also shifted slightly over the
nesting period  (nine weeks) at Protection Island.  Juvenile salmon were more
important in the first four weeks, whereas Pacific herring gradually
increased in importance through the nesting period.  The average number of
fish delivered to chicks on Protection Island was similar during the two
years (5.62 in 1975, 5.65 in 1976) but the average weights of the loads were
not similar (32.28 g in 1975, 29.52 in 1976) and the patterns over time were
also dissimilar.

     Outside the north Puget Sound and Strait of Juan de Fuca area,
euphausiids appeared to be more important in the diet of adult rhinoceros
auklets and their nestlings (Grinnel 1899; Linton 1908; Kozlova 1957; Komaki
1967).  Pacific sand lance, however, dominated the stomach contents of four
rhinoceros auklets collected in Alaskan waters (Sanger and Baird 1977).
                                    248

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Table C-3.  Percent composition of numbers of prey delivered to rhinoceros
            auklet nestlings on Protection Island (from Munuwal 1977).
1975
Prey
Pacific sand lance
Pacific herring
Chinook salmon, juv.
Pink salmon, juv.
Chum salmon, juv.
Coho salmon, juv.
Northern anchovy
Surf smelt
Walleye pollock
Threespine stickleback
Squid (unidentified)
% Number
82.78
14.44
0.56
0
1.11
0
0.56
0
0
0.56
0
% Weight
70.60
26.02
1.55
0
0.87
0
0.77
0
0
0.19
0
1976
% Number
74.36
20.53
1.08
2.26
0
0.10
0.79
0.39
0.39
0
0.10
% Weight
63.76
25.72
3.01
3.50
0
0.19
2.09
1.43
0.13
0
0.17
                                    249

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Cassin's Auklet

     Although common offshore, Cassin's auklets are not frequently
encountered nearshore, but they have been reported to breed on Carroll and
Alexander islands and on Cake Rock on the outer coast  (Cody 1973).  According
to Cody, Cassin's auklet is the only alcid in Washington that forages on
plankton at long distances.  The plankton is regurgitated as a soup to the
nestlings.  Thoreson  (1964) and Manuwal (1974) documented euphausiids
(Thysanoessa spinifera), amphipods (Phromema sp.), and squid as  the main
components of their diet off the coast of California.  Sanger and Baird
(1977) listed only unidentified crustaceans in the stomach contents of five
Cassin's auklets collected in Alaskan waters.

Ancient Murrelet

     Although neither Salo (1975) nor Manuwal  (1977) includes ancient
murrelets in his survey, they appear, although infrequently, in  the northern
reaches of Puget Sound during migration from their breeding sites in Alaska
and British Columbia and many may overwinter in the region (Sealy 1976).
The 1976 Audubon Christmas bird count recorded 377 at Victoria,  B.C., and
incidental sightings at Pender Islands and Vancouver, B.C., and  Bellingham,
Washington (Heilbrum, et al., 1977).

     Sealy's (1972) detailed comparison of the feeding ecology of the marbled
murrelet and the ancient murrelet during the breeding season in  the Queen
Charlotte Islands, B.C., illustrated that adult ancient murrelets prey
specifically on euphausiids (Euphausia pacifica, Thysanoessa spinifera),
which composed 92.4% of the total prey abundance, while fish (Pacific sand
lance and shiner perch) made up only 7.2%.  Subadult birds, however, fed
more upon fish, primarily Pacific sand lance 30-60 mm in length, which
accounted for 45.7% of the total prey abundance vs. 55.3% for invertebrates.
Juvenile ancient murrelets were essentially piscivorous (98.3%).  The
euphausiid IS. pacifica prevailed in the diet in March and April  before it was
supplanted by T_. spinifera; Pacific sand lance and shiner perch  occurred
during the period between mid-June and mid-July.  Sealy (1973) also noted
that ancient murrelets often fed in conjunction with black-legged kittiwakes
and glaucous-winged gulls, which were feeding on similar prey organisms at
the surface.  The results of Sanger and Baird's (1977) examination of ancient
murrelets in Alaskan waters were generally similar, the euphausiid Thysano-
essa inermis being the principal prey of adult birds.

Marbled Murrelet

     Marbled murrelets are an abundant summer bird along the Pacific North-
west coast (Drent and Guiguet 1961) and are presumed to nest inland on large
rivers (Hoh and Quileute) of the Olympic Peninsula (Cody 1973).

     Compared with the prey spectrum of adult ancient murrelets, that of
adult and subadult marbled murrelets is much more oriented toward fish.
Sealy (1972) documented that small (less than 60 mm in length) Pacific sand
lance provided 61.8% of the total prey abundance of prey organisms for
marbled murrelets in the Queen Charlotte Islands, B.C.; shiner perch,

                                    250

-------
rockfish, osraerids (smelt, capelin, eulachon), and prickleback were other
important fishes.  Euphausiids  (Thysanoessa spinifera)  larger than 12 mm
formed the majority of the invertebrates consumed, which  overall contributed
only 26.8% of the total prey abundance, and occurred only early  (mid-April to
late June) in the breeding season.  Shiner perch and osmerids appeared in the
diet primarily in the latter half of  the breeding season  (early  July to mid-
August).  Sealy  (1973) also found marbled murrelets feeding upon T\ spinifera
in conjunction with glaucous-winged gulls.

     Manuwal and Boersma  (1977) also  listed fish  (unidentified)  as the main
constituent  (67% abundance) of  the food of marbled murrelets collected in
the Gulf of Alaska and the southeastern Bering Sea.  Both fish  (67% freq.
occur.) and unidentified  crustaceans  (33%) occurred in marbled murrelets
collected in Alaska' south-central waters  (Sanger and  Baird 1977).
                                    251

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    Alaskan Continental  Shelf.  Annual Rept. Prin. Invest, for year ending
    March  1977.

 Sanger,  G.A.,  and  P.A.  Baird.  1977.  The  trophic relationships of marine
    birds  in the Gulf of  Alaska and  the southern Bering Sea.  NOAA-OCSEAP,
    Environmental assessment  of the  Alaskan Continental Shelf.  Vol. IV.
    Receptors—birds.   Annual Rept.  Prin.  Invest, for year ending March 1977.

 Scattergood, L.W.   1950.   Observations on  the food habits of the double-
    crested cormorant,  Phalacrocorax auritus auritus.   The Auk 67(4):506-508.

 Scott, J.M.  1973.   Resource allocation in four synoptic  species of marine
    diving birds.  Ph.D.  Thesis,  Oregon State Univ., Corvallis.

 Sealy, S.G.  1972.   Adaptive differences in breeding biology in the marine
    bird family Alcidae.   Ph.D. Thesis,  Univ.  Michigan.

 Sealy, S.G.  1973.   Interspecific  feeding  assemblages of  marine birds off
    British Columbia.  The  Auk 90:796-802.

 Sealy, S.G.  1976.   Biology  of nesting ancient murrelets.  The Condor
    78(3):294-306.

 Smith, J.L., and D.R.  Mudd.   1976.   Impact of dredging on the avian fauna in
    Grays Harbor.  Appendix H in  U.S. Army  Corps of Engineers, Maintenance
    dredging and the environment  of  Grays Harbor, Washington.  Seattle, WA.

 Sperry, C.C.  1940.   Food  habits of a group of shorebirds:  Woodcock, snipe,
    knot,  and dowitcher.   Bur.  Biol. Survey, Wildlife Res. Bull. 1, 37 pp.
                                     257

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Springer,  A.M.,  and D.G. Roseneau.  1977.  A comparative seacliff inventory
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   Environmental assessment of the Alaskan Continental Shelf.  Vol. V.
   Receptors—birds.  Annual Rept. Prin. Invest, for year ending March 1977.

Steele,  B.B.,  and W.H. Drury.  1977-  Birds of coastal habitats on the south
   shore of the Seward Peninsula, Alaska.  Pages 7-119 in NOAA-OCSEAP,
   Environmental assessment of the Alaskan Continental Shelf.  Vol. III.
   Receptors—birds.  Annual Rept. Prin. Invest, for year ending March 1977.

Thoresen,  A.C.,  and E.S. Booth.  1958.  Breeding activities of the pigeon
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Thoresen,  A.C.  1964.  Breeding behavior of the Cassin's auklet.
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Verbeek, N.A.W.  1977.  Comparative feeding behavior of immature and adult
   herring gulls.  The Wilson Bull. 89(3):415-421.

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Webster, J.D.  1941.  Feeding habits of the black oystercatcher.
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Wiens, J.A., and J.M. Scott.  1975.  Model estimation of energy flow in
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                                    258

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                         APPENDIX D.  MARINE MAMMALS
                               D-l.  Cetaceans

     Scheffer and Slipp (1948) recorded 20 species of cetaceans  (whales,
dolphins, and porpoises) in the marine waters of Washington  State; Pike and
MacAskie (1968) listed 21 from British Columbia  (Table Dl).*  These
comprehensive accounts, however, were based principally upon records  of
the coastal whaling stations** and of strandings, rather  than upon
systematic survey observations with estimates of abundance.   As  a result
they do not necessarily reflect whale populations of today,  many of which
(sperm and humpback whales) have been seriously depleted  by  overharvesting;
only gray whales have recovered to their estimated original  abundance.
In addition, only a few of the cited species were ever common to the
enclosed waters of Puget Sound, the Strait of Georgia, and the inner  Strait
of Juan de Fuca, though many were encountered during aboriginal  whaling
activities by  the Makah Indians, who were located on the  western end  of the
Strait of Juan de Fuca.  Of the modern occurrences of cetaceans  in  the
region's inland waters, only two cetaceans—orca (killer  whale)  and harbor
porpoise—can  still be considered abundant, and  four—Pacific white-sided
dolphin, Ball  porpoise, gray whale, and Minke whale—are  still relatively
common.  The following species accounts consider all cetaceans reported in or
adjacent to  the Strait of Juan de Fuca and northern Puget Sound, but  the
preceding food web section discusses only the six species which  now occur in
the  region.

     All the cetaceans documented for the region have been categorized
according to six functional feeding groups (Table D-2).   Of  the  common or
abundant species, gray and Minke whales were determined to be obligate
planktivores and the Pacific white-sided dolphin and the  Ball porpoise,
 facultative  piscivores  in pelagic habitats; in nearshore  habitats,  the Pacific
harbor porpoise has been classified as an obligate piscivore while  the orca
or killer whale sits at the peak of the marine food web as a  facultative
carnivore.
      *Nomenclature has been standardized according  to  Rice  (1977).

     **Statistics  came mostly  from one station  operating at  Bay City,
 Washington,  from  1911 to 1925 and another at Coal Harbour on Vancouver Island,
 British Columbia, from 1948 through 1959 and 1962 through 1967.
                                     259

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  Table D-l.  Cetaceans occurring in Washington State and British Columbia.  A = abundant, C = common,
              NC = not common, R = rare, c = coastal only, i = inshore Puget Sound and Strait of
              Juan de Fuca.


                                                              Washington State  British Columbia
                                                                (Scheffer and      (Pike and     Reported
Order Cetacean—whales and dolphins                              Slipp 1948)     MacAskie 1969)   habitat


Suborder Odontoceti—toothed whales and dolphins
   Family Ziphiidae—beaked whales
      Berard-ius bai.vdi.-L Stejneger, Baird's beaked whale              NC                C            i
      Mesoplodon stejnegeri True,
        Stejneger beaked whale                                        R                Re
      M. oavlhubbsi Moore, Hubbs' beaked whale                                         R            c
      Ziphius cavirostris Cuvier, Cuvier's beaked whale                                R            i
   Family Physeteridae—sperm whales
      Physeter oatodon Linnaeus, sperm whale                          C                C            c
      Kogia breviaeps Blainville, pygmy sperm whale                                   NC            i
   Family Delphinidae—ocean dolphins
      Stenella sp., spotted dolphin                                                    R            i
      Delphinus delphis Linnaeus, Pacific common dolphin              R                R            i
      Lissodelphis bmvalis Peale,
         northern right-whale dolphin                                 R                Re
      Lagenorhynchus obliguidens Gill,
         Pacific white-sided dolphin                                  C                C            i
      Orcinus oroa Linnaeus, killer whale or orca                     A                A            i
      Grampus gr-iseus Cuvier, gray grampus or
         Risso's dolphin                                                               R            i
      Pseudorca OTassidens Owen, false killer whale                   R                             i
      Globiccphala macvovh'ijnc'ha, shortfin pilot whale                NC               NC            i
      Thocoena phoaoena Linnaeus,
         Pacific harbor porpoise                                      A                A            i
      Phojoenoides dalli True, Dall porpoise                          C                C            i
   Family Monodontidae—Arctic dolphins
      Delphinapterus leucas, beluga whale                             R                             c

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       Table D-l.   Cetaceans occurring in Washington State and British Columbia.   A = abundant, C - common,
                    NC = not common, R = rare, c = coastal only, i = inshore Puget Sound and Strait ot
                    Juan de Fuca - cont'd
    Order Cetacean—whales and dolphins
                                                               Washington  State   British Columbia
                                                                 (Scheffer and       (Pike and     Reported
                                                                  Slipp  1948)      MacAskie 1969)   habitat
NJ
Suborder Mysticeti—whalebone whales
   Family Eschrichtiidae—gray whales
      Eschrichtius robustus Lilljeborg,  gray whale
   Family Balaenopteridae—furrow-throated whales
      Balaenoptera physalus Linnaeus, fin or finback whale
      B.  borealis Lesson, sei whale
      B.  acutovostrata Lacepede,
         little piked whale, minke whale
      B.  muscutus Linnaeus, blue  whale
      Megapteva novaeangliae Borowski, humpback whale
   Family Balenidae—smooth-throated whales
      Balaena glacialis Muller, northern or
         black right whale
 C
 C

NC

NC


 R
 C
 C

NC
NC
NC
1
c

i
c
i

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     Table D-2.  Functional feeding groups and representative prey taxa of marine mammals known or suspected
                 to occur in north Puget Sound and the Strait of Juan de Fuca.  Species common or abundant  in
                 the region are underlined.  Species involving questionable sightings or extremely rare
                 species are in parentheses.
     Habitat
Feeding group
Predator species
Representative prey taxa
     Pelagic
Obligate planktivore
ON
                 Facultative
                 (macro-)planktivore
                 Obligate piscivore
Gray whale*
Finback whale
Sei whale
Minke whale
Blue whale
North Pacific right
  whale
                        Humpback whale
                        (Shortfin pilot whale?)
                        Gray grampus dolphin?
                        False killer whale
                        Northern fur seal
                        Northern elephant seal
                        (Beluga whale)
Calanoid copepods (Calanus sp.)
Euphausiids (Euphausia paeifioa, Thysano-
  essa sp.)
Crab zoea (Pachyaheles Tudis, Cancer sp.)
Squid (Ommastrephes sp.)
Pacific saury (Coloabis saira)
Northern anchovy (Engraulis mordax)
Walleye pollock (Theragra chalcogTcama)
Pacific sand lance (Ammodytes hexaptevus)

Squid (Gonatidae, Histioteuthidae,
  Ommastrephidae, Onychoteuthidae)
Rockfish (Sebastes sp.)
Lingcod (Ophiodon elongatus)
Skate (Rajiidae)
Lancetfish (Alepisaw?us sp.)
Shark (Squaliformes)
Surf smelt(Hypomesus pretiosus)
Atka mackerel (Pleurogrammus
  monoptery gins')
Pacific sardine (Sardinops sagax)
Euphausiids (E.  pacifica, Thysanoessa sp.)

Capelin (Mallotus villosus)
Pacific herring (Clupea harengus pallasi)
Pacific sardine (5.  sagax*)
Salmon (Oncorhynchus sp.)
Northern anchovy (E.  mordax)
Walleye pollock (T.  ohalcograrma)
Eulachon (Thaleichthys pacifiGus)
Pacific sand lance (A.  hexapterus)

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 Table D-2,  cont'd
 Habitat
Feeding group
                                    Predator species
Representative prey taxa
             Facultative piscivore
Nearshore   Obligate piscivore
                        Baird's beaked whale
                        Stejneger beaked whale
                        and  other Mesoplodcni sp.
                        (Cuvier's beaked whale)
                        (Spotted dolphin,
                        Stenella sp.)
                        (Pacific common dolphin)
                        Pacific white-sided
                        dolphin
                        Northern right-whale
                        dolphin?
                        Pall porpoise
                        Pygmy sperm whale


                        Harbor porpoise
                        Northern sea lion
                        California sea lion
                        Pacific harbor seal
Squid  (Loligo opalescens, Gonatus  sp.,
  Ormastrephes sp., Onyehoteuthis  sp.)

Lanternfish  (Myctophidae, Diaphus  sp.)
Pacific hake  (Merlucaius produotus)
Northern anchovy  (E. mordax)
Pacific saury (C. saira)
Squid  (Gonatus sp., Onychoteuthis  sp.,
  L. opalescens}
Shrimp (Bentheogennema borealis, Pasiphaea
  pacifioa, Pandalus sp.)
Pacific herring  (C. harengus pallasi)
Capelin (M. villosus)
Juv. rockfish (Sebastes sp.)
Salmon (Oncorhynchus sp.)
Octopus (Octopus  so.)

Pacific herring  (C. harengus pallasi)
Pacific sand lance  04. ~hexa.ptems)
Walleye pollock  (T. chatcogramma)
Salmon (Oncorhyna'hus sp.)
Starry flounder  (Platiehthys stetlatus)
Pacific tomcod (Miorogadus pacificus)
Rockfish (Sebastes  sp.)
Skate  (Rajiidae)
Pacific cod  (Gadus maopocephalus)
Pacific hake (M.  productus)
Spiny dogfish (Squalus aoanthias')
Plainfin midshipman (Porichthys notatus)
Greenling  (Hexagrammidae)
Shiner perch (Cymatogaster aggregata)

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    Table  D-2,  cont'd
Habitat     Feeding group
                                         Predator species
Representative prey taxa
                Facultative  carnivore   Orca (killer whale)
S3
Pacific staghorn sculpin  (Leptooottus
  armatus)
Eelpout (Zoarcidae)
Blackfin sculpin (Malacocottus ki-ncaidi)
Shrimp
Crabs (Cancer sp.)
Octopus (Octopus sp.)

California sea lion  (Zalophus
  californianus)
Northern  sea lion (Eumetopias jubatus)
Harbor seal (Phoca vitulina)
Elephant seal (Mirounga californianus)
Harbor porpoise (Phoeoena phocoena)
Ball porpoise (Phocoenoides dalli)
Minke whale (Balaenoptera acutorostrata)
Nursing calves of humpback  (Megaptera
  novaeangliae'), finback  (Balaenoptera
  physalus), and gray whales (Eschrichti-us
  robustus)
Lingcod (0. elongatus)
Salmon (Oncorhynchus sp.)
Steelhead trout (Salmo gairdneri)
Pacific halibut (Hippoglossus stenolepis)
Pacific herring (C. harengus pallasi)!
         *0n  feeding  grounds  outside the north Puget  Sound  -  Strait  of  Juan de  Fuca  region,  gray whales feed
    on benthic and  epibenthic organisms.
         ?—denotes subjective assignment of  predator species to  feeding  group  in  absence  of food habits data.

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                              Species Accounts

Baird's Beaked Whale

     Looking much like a small sperm whale (10-15 m long at maturity),  Baird's
beaked whale appears to occur principally along the coast, peak occurrences
being in the fall.  Six whales were reported in early October 1976,  75  km
offshore of Westport, Washington, by Wahl (1977).  Of thirteen stomachs
examined from the Coal Harbour station between 1950 and 1958, three  were
empty.  Squid and rockfish (Sebastes spp.) bones appeared in seven stomachs
and skate egg cases were found in the stomach contents of two.

Stejneger Beaked Whale and Other Mesoplodon sp.

     Another group of beaked whales includes three species of Mesoplodon,
of which M. stejnegeri appears to be the prominent species.  They appear
typically along the coast, although an early record by Scammon (1874)
indicated a large number may have occurred in Port Townsend Bay (Scheffer and
Slipp 1948).  There is no recorded account of stomach contents of any of these
species.  We assume they would feed similarly to the other beaked whales, i.e.,
basically upon squid and fishes.

Cuvier's Beaked Whale

     Although no  confirmed records of Cuvier's beaked whale originate from
Washington, there have been a number of records from the west coast  of  Van-
cover Island and  the Queen Charlotte Island area.  Little stomach contents
information is presently available.  Recently, a Cuvier's beaked whale, 4.3 m
in length, washed ashore on the north side of Nizki Island in the western
Aleutians, and the stomach was examined and reported to contain squid remains
 (R.M. Mayer, Fish. Res. Inst., personal communication, 1979).

Sperm Whale

     Although now seldom reported off the coast of Washington, the largest
of the  toothed whales (15-20 m at maturity) were third in abundance  in  the
catch at Bay City, Washington, stations, 1913-1915 and 1918-1919 (Scheffer
and Slipp 1948), and composed almost 80% of the whales taken at Queen Charlotte
Islands' whaling stations between 1933 and 1943, and 32% of those taken at
Coal Harbour between 1948 and 1959 (Pike and MacAskie 1968).  Sperm  whales
therefore may have been a predominant species before coastal whaling off
Washington and southern British Columbia and present high seas whaling  reduced
their numbers.  They appeared from late spring through early fall in large
harem schools in the open ocean up to 200 miles from the west coast  of  British
Columbia (Pike and MacAskie 1968).   This concentration was apparently a
feeding aggregation, as most stomachs contained food organisms, including

                                     265

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squid (Moroteuthis robusta and Gonatus sp.), hagfish  (Eptatretus  stouti),  and
rockfish.   Octopi, codfish (Gadidae), and a lamprey (Petromyzontidae)
constituted incidental prey items (Robbins, et al., 1937).

     Small to moderate sized cephalopod molluscs  (squid and octopus) appeared
to be the prevalent prey items of sperm whales on most localities including
the western Gulf of Alaska (Thompson 1940; Kawakami 1976), the Bering Sea
(Okutani and Nemoto 1964), California (Fiscus and Rice 1974, Japan  (Okutani,
et al., 1976), the Kurile Islands (Bethesheva and Akimushkin 1955), New
Zealand (Gaskin and Cawthorn 1967), Spain (Clarke and MacLeod 1974), and
Northwest Africa (Clarke 1962).  Taxonomic composition of the squids varied
according to the geographic region, but the family Gonatidae composed more
than half of the squid prey reported for the North Pacific.  Okutani, et al.
(1976), identified 15 species belonging to 9 families where species of the
families Histioteuthidae, Ommastrephidae, and Onychoteuthidae numerically
predominated.  The so-called giant squid, Architeuthis sp. and Moroteuthis
robusta,  did not occur frequently but contributed high biomass proportions
when they did occur.

     Fish appeared less frequently than squid in almost all circumstances
but often contributed considerably to the total prey biomass.  Gaskin and
Cawthorn (1967) documented a 1.7:1 ratio of squid to fish by weight.  All of
the 14 sperm whale stomachs examined at Port Hobron in 1937 and 8 of the 13
from Akutan in 1938 had cephalopod remains.  Fish  (lingcod, Ophiodon
elongatus), rockfish, skates (family Rajiidae), and their egg cases) and
hermit crabs (Paguridae) were secondary prey items (Thompson 1940).

     Backus (1966) described the contents of one sperm whale from the Azores
as containing "10 or 12 photophore-bearing cephalopods, each about 1 m in
total length, 2 lancetfish (Alepisaurus sp.), each about 1 m long, and the
partial remains of a large shark" (probably Cetorhinus maximus).

Spotted Dolphins

     No specimen records nor stomach contents of spotted dolphins (Stenella
styx,  S^. euphrosyne, and S^. caeruleoalba) have been examined from the
northern Washington or southeastern British Columbia coasts.  Wahl (1977)
reported one sighting of J5. caeruleoalba, 72 km northwest of Westport in late
September 1976.

     Miyazaki, et al. (1973), provided detailed stomach contents analyses of
27 specimens of S^. caeruleoalba from Sagami Bay, Japan.  An average of 53.6
+ 43.6 (+ 1 s.d.) prey organisms comprising 7.6 + 3.5 species was contained
in each stomach, weighing an average of 1,074 + 418 g.  Fish numerically
dominated the stomach contents at 59% of the total prey abundance.  Lantern-
fishes were the prevalent group, especially Diaphus sp., accounting for 64%
of all identified fishes.  The estimated body lengths of consumed fishes
ranged from 60 to 300 mm.  Shrimp, especially Bentheogennema borealis, were
also common prey components, followed by squid.  All prey species were pelagic
or epipelagic.  Since the coastal environments of central Japan and northern
Washington and southern British Columbia are similar, the food habits of this
genus of dolphin may also be similar.


                                    266

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Pacific Common Dolphin

     One specimen of this species, relatively  rare in the region, recovered
at Victoria, B.C., was not examined  for  stomach  contents.  Fiscus and Niggol
(1965) reported the stomach contents of  four animals from central California.
Fish—Myctophidae, northern anchovy  (Engraulis mordax),  Pacific saury (Colo-
lab is saira)—and squid  (Gonatus sp., Onychoteuthis sp.,  Loligo opalescens)
were the predominant prey items.

Northern Right-Whale Dolphin

     One right whale found stranded  on a southwestern Washington coastal
beach had one squid beak in its stomach  (Scheffer and Slipp 1948).   About 20
individuals were observed by Wahl  (1977)  in association with gray grampus
dolphins in October 1974, 85 km west of  Westport.

Pacific White-Sided Dolphin

     Pike and MacAskie (1969) listed 32  sightings of Pacific white-sided
dolphins from British Columbia and northern Washington which suggested an
inshore movement in winter and an offshore movement in summer.   The northern-
most observation made by Fiscus and  Niggol  (1965)  was off Point Grenville;
they reported few Pacific white-sided dolphins inside the 100-fm and few
outside the 1,000-fm curve.  Wahl  (1977)  included one report of 25-30 Pacific
white-sided dolphins observed 72 km  northwest  of Westport in late September
1976.  Extremely large pods of 1,000-2,000 animals have been reported in the
Gulf of Alaska  (Pike 1959; Fiscus, et al., 1977);  55 other sightings have
been reported between 1958 and 1975  (Fiscus, et  al. ,  1977).

     Few stomach samples have been examined in the northeastern Pacific
region.  Five from California coastal waters reported in  Fiscus and Niggol
 (1977) had  consumed fish (northern anchovy, 45-95%,  and  Pacific hake, Mer-
luccius productus, 5-10%) and squid  (Gonatus sp.).   Another specimen from
California was  reported  to have fed  upon squid (Scheffer  1950b) while
another from that area had consumed  Pacific sardine (Sardinops  sagax)
 (Higgins 1919).	—

     Lanternfishes were  the main fish (77% of total stomach contents volume)
of the prey consumed by  13 Pacific white-sided dolphins  examined by Wilke,
et al. (1953);  anchovy,  Engraulis japonica, and  chub mackerel,  Scomber
Japonicus, were the other prey fishes.   Squid  beaks and  eye lenses, probably
from Watasenia  scintillans. composed only 7% of  the stomach contents volume.

Orca  (Killer Whale)

     Undoubtedly the most ubiquitous toothed whale in north Puget Sound  and
the Strait of Juan de Fuca is the orca.   As of 1977,  approximately  115 whales
occurred in this area, of which 65-70 could be considered residents (compos-
ing four distinct pods designated J, K,  L 8, and L 10)  and the  remainder
transients  (composing seven pods, A5, E,  G, M, N,  0,  and  Q)  (Bigg , et al.,
™ ™. !eCent Si8btl'n8s su8gest that the resident  pods may now comprise
75-80 individuals (Balcomb 1978;  K. Balcomb and  R.  Osborne,  pers. comm.).
                                    267

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There was no distinct pattern to their distribution and movements  (M.S.
Bigg, Pac. Biol. Sta.,  pers. comm.), which appeared to be related  directly
to feeding or the search for food (Balcomb and Goebel 1976).  Only J pod
appeared to be completely residential in Puget Sound, the Strait of Georgia,
and eastern Strait of Juan de Fuca, having a range of about 210 nautical
miles.  The other three pods apparently resided in the region 8-9  months of
the year, entering and leaving through the Strait of Juan de Fuca; there is
no information on their movement along the outer coast.  The transient pods
occurred infrequently,  moving into and out of the northern Strait  of Georgia
from the north, and have never been seen inside Puget Sound proper.  Several
aggregations of 3 to 17 animals were sighted, 25-100 km off Westport by
Wahl  (1977) between May and September.

     Scheffer and Slipp (1948) summarized orca food habits and feeding beha-
vior known to that time.  A diverse array of marine mammals, sea birds,
fishes, and cephalopod molluscs were the principal prey taxa described.
Smaller individuals and nursing calves of humpback (Megaptera novaeangliae),
finback (Balaenoptera physalus), and gray (Eschrichtius robustus)  whales,
harbor porpoise (Phocoena phocoena), hair seal, black brant (Branta nigri-
cans), greenling (Hexagrammidae), lingcod, salmon (Oncorhynchus sp.), and
squid were the specific organisms itemized.

     Nishiwaki and Handa (1958) were the first to evaluate orca food habits
on a large scale.  Their analysis of the stomach contents of 364 orcas from
the coast of Japan indicated that the whales fed mostly on fishes  and
cephalopods, but also on large numbers of cetaceans and pinnipeds.

     Detailed information on orca food habits was assembled by Rice (1968).
He summarized the stomach contents of ten orcas collected in offshore waters
of the eastern North Pacific between Kodiak Island, Alaska, and San Miguel
Island, California.  Eight of the ten stomachs examined contained  marine
mammal remains, specifically California sea lion (Zalophus californlanus),
northern sea lion (Eumetopias jubatus), elephant seal (Mirounga anguisiros-
tris), harbor porpoise, Dall porpoise (Phocoenoides dalli), and minke whale
(Balaeroptera acutorostrata), which made up over 75% of the total  prey.
Three stomachs contained fish, including Pacific halibut (Hippoglossus
stenolepis), opah (Lampris regius), and carcharinid sharks.  Only  one speci-
men contained cephalopod remains.  Martinez and Klinghammer (1969) generally
itemized orca prey organisms, which included 10 species of cetaceans, 8
pinnipeds, 11 teleost fishes, 6 other vertebrates, and 3 elasmobranch fishes.
One specimen from California examined by Fiscus and Niggol (1965)  was pre-
sumed to have eaten one California sea lion, one cetacean, and as  many as
four elephant seals.

     Other recorded prey organisms of orcas have included northern fur seal
(Callorhinus ursinus) in the western North Pacific (Bychkov 1967); bearded
seal  (Evignathus barbatus) and walrus (Odobenus rosmarus) in the Bering  Sea
(Zenkovich 1938); eagle ray (Myliobatis sp.) off Brazil  (Costello  1977);
leatherback sea turtle  (Dermochelys coriacea) in the Lesser Antilles  (Cald-
well and Caldwell 1969); and minke whale along the northwest coast of
Vancouver Island, B.C.   (Hancock 1965).

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     Recent observations of orca feeding activity and collections  of  prey
remains made by Pacific Biological Station biologists off southeastern
Vancouver Island identified chinook (Oncorhynchus tshawytscha),  coho  (0.
kisutch)  and sockeye salmon (0. nerka), and anadromous  (steelhead)  trout
(Salmo gairdneri) as important food items in the region  of  concern (M. Bigg
and G. Ellis, unpub. data).

     Existing reports of orca food habits imply that pinnipeds and small
cetaceansconstitute the preferred prey  organisms and the staple element of
the orca diet in most regions of the world. Rice  (1968)  has pointed out,
however  that, "in Puget Sound, where the killer whale population is probably
denser  than  anywhere else  in the world,the marine mammal population (mostly
harbor  porpoise, Minke whale, Pacific harbor seal,  and a few northern sea
lions)  does  not  appear large enough to  provide a major proportion of the
killer  whale's diet.  It is widely believed that runs of spawning salmon  are
a major food there  during  the summer, but no data are available to support
 this  assumption."   As supported by Bigg's recent evidence,  salmon and
 abundant schooling  forage  fishes such as Pacific herring (Clupea harengus
 pallasi) and Pacific sand  lance  (Ammodytes hexapterus) may  be the most stable
 trophic contribution to  the orca's diet in northern Puget Sound and the Strait
 of Juan de Fuca, supplemented by such pinnipeds and small whales as they  may
 encounter.

      Feeding behavior may  also  vary from pod  to pod.  Balcomb and Goebel
  (1976)  identified L pod  as being more  aggressive  in its  food habits than
 others; i.e., it was  the only  one  observed  to  take  marine mammals.  The L pod
  typically frequented the waters on the  west  side  of Vancouver Island and  the
 Olympic Peninsula,  venturing  periodically  into inland waters through  the
  Strait of Juan de Fuca.

 Gray Grampus or Risso's  Dolphin

      Stroud (1968)  documents  one of the few  records of  gray grampus dolphin
  in Washington.   His description of a young male  specimen which washed ashore
 at Makkaw Bay (northwest,  exposed  coast of Olympic  Peninsula) included
  contents of the stomach,  which,  though  the animal  had been  dead about a
 month, included 44 (55  g)  identifiable  squids.   Seven  taxa  were represented;
  Gonatus fabric!, two other gonatids,  and Chiroteuthis verangi were the most
  prominent forms but Onychoteuthis  banksi and  Octopodoteuthis sicula were also
  present.  Several verified records and  several credible  sightings of  gray
  grampus dolphin exist  for British  Columbia which  is apparently the northern
  limit of its range (Guiguet and Pike  1965;  Pike  and MacAskie 1968); no food
  habits were determined,  however.   Wahl  (1977)  reported  three sightings, one
  in October 1974 and two  in September  1976,  of gray grampus in groups  of 40 to
  60, often mixed with other species such as  Pacific striped dolphin.

  False  Killer Whale

      Scheffer and Slipp (1948) provided the only record of  a  false killer whale
  in Washington and British Columbia, an animal stranded  near Olympia  (south
  Puget  Sound) in 1937.   Its stomach contained some  salmon remains.
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Shortfin Pilot Whale

     Although sightings of Pacific pilot whales have occurred frequently,
especially in 1958 when there appeared to be a northward shift in distribution,
there were but a few verified specimens in the Washington and British Columbia
region (Pike and MacAskie 1968).  They do not appear to move into the region's
coastal waters before mid-April (Fiscus and Niggol 1965).  Sighting data
recently compiled by the Moclips Cetological Society's Orca Survey suggest
that five to ten groups may frequent Puget Sound each year, principally in
spring and summer (Rich Osborne, pers. comm.).  No diet information was
reported.

Harbor Porpoise

     Considered the most frequently seen cetacean in Washington State by
Scheffer and Slipp (1948) . harbor porpoise range the breadth of Puget Sound
and the Strait of Georgia, through the Strait of Juan de Fuca, and well
offshore Washington and British Columbia.  They were frequently sighted by
Wahl (1977) close inshore of Westport and in Grays Harbor channel, usually
in groups of one to five animals.  They have been captured in nets set on the
bottom as deep as 44 fm.  When seen in Puget Sound they were often in groups
of two to five, occasionally ten to twelve.  The 1977 Orca Survey, conducted
by the Moclips Cetological Society (Balcomb 1978) also reported seven
sightings of harbor porpoise in the San Juan Islands; compilation of their
total data, however, suggests that between 30 and 60 are seen each year in
northern Puget Sound (Rich Osborne, pers. comm.).

     Scheffer and Slipp (1948) concluded from the contents of four stomachs
that, "the favorite foods of the harbor porpoise are probably fishes under
a foot in length; of slender form and soft flesh; lacking stiff spines and
armor; including types that commonly run in schools near, but not on, the
bottom.  Such speedy swimmers as the salmon and trout; bottom dwellers;
heavily armed species like the rock-cod and sculpin, and invertebrates with
the exception of the squid, are probably not important in the diet of the
porpoise."

     Five Pacific herring were found in the stomach of one female harbor
porpoise examined at Port Townsend, Washington, in May 1950 (Wilke and Kenyon
1952).

Pall Porpoise

     Despite the high abundance of Ball porpoise in Washington and British
Columbia waters, second only to the harbor porpoise, few described specimens
existed (Scheffer and Slipp 1948; Pike and MacAskie 1968).  Ball porpoise
appeared to be most abundant offshore and common in the Strait of Juan de
Fuca, though seldom seen in either the Strait of Georgia or Puget Sound
proper.  Fiscus and Niggol (1977) observed Ball porpoise six times during
April 1959, all within 50 km of shore.  Sightings of Ball porpoise were made
on 24 of the 34 cruises between July and October described by Wahl (1977),
but sightings occurred on only 2 of 12 cruises between mid-April and mid-May.
The majority of these observations were made more than 50 km offshore and

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were of groups of 3 to 6 porpoise.   Dall  porpoise generally travel in
schools of 5 to 14 (usually 6) but  are  encountered inshore in groups of 30 to
100 in spring and fall.  Eight sightings  of Dall porpoise were reported in
the 1977 Orca Survey  (Balcomb 1978),  principally along western Whidbey Island
between Rosario Strait and Possession Point.  More recent assessment of the
survey data indicates that three  to  four  groups, comprising 40 to 60
individuals, of Dall porpoise frequent Puget Sound (Rich Osborne, pers. comm.).

     Pacific herring were the sole  prey found in the stomachs of four Dall
porpoise taken in Queen Charlotte Sound,  B.C., by Cowan (1944).  The stomachs
of  the Dall porpoise  collected in 1937  between Sauk Inlet and Port Angeles
along the  Strait of Juan de Fuca  contained fish and squid beaks and unidenti-
fied "eyes" (Scheffer and Slipp 1948).  Of five specimens collected offshore
Washington and British Columbia between 1957 and 1959, the stomachs of two
were empty, two had "mostly squids," and  one had "mostly herring." Stomach
contents of nine Dall porpoise collected off the California and Washington
coasts between 1964 and 1972  (Stroud et al. 1964-72) contained 4 species of
pelagic fish, one flounder and six  species of squid. Fiscus and Niggol (1965)
examined the stomach  contents of  five specimens collected off northern
California; one stomach was empty,  the  other four contained only remains of
squid.

      The stomach contents of  25 Dall porpoise from Monterey Bay were examined
by Loeb  (1972).  She  found  that hake (Merluccius productus), juvenile rockfish,
and the  squid Loligo  opalescens were the  prevalent prey between May and
December,  and northern anchovy and  Pacific herring became important in winter.
During  the period between October and April other fish species became
 important: Myctophids  (including predominantly Tarletonbeania crenularis),
night smelt  (Spirinchus  starksi), Pacific sanddab (Citharichthys sordidus),
 spotted  cusk-eel  (Otophidium  taylori),  pompano (Petrilus simillimus), juvenile
 sablefish  (Anoplopoma fimbria), California smoothtongue (Bathylagus stilbius),
 pinpoint  lampfish  (Lampanyctus regalis),  snailfish (Liparis sp.), eelpouts
 (Zoarcidae),  grenadiers  (Macrouridae),  and eels.  Pelagic cephalopods found
 in the  stomachs  included  the  squids Abriolepsis felis,  Gonatus sp., and
 Onychoteuthis boreali-japonicus,  and the  octopus Octopus bimaculatus.

      Capelin  (Mallotus villosus)  were the only prey found in the stomachs
 of two  porpoise  from  the  Gulf of  Alaska examined by Scheffer (1953).  Squid
 predominated  in  the  stomach contents of 148 Dall porpoise from the Bering
 Sea and  North Pacific examined by Mizue,  et al. (1966); fish, including
 sockeye  salmon,  and  shrimp were secondary components of the diet.

      Ninety-eight percent of  the  total  volume of the stomach contents of four
 Dall porpoises collected  off  Japan  by Wilke et al. (1953) was squid,
 predominantly Ommastrephes  sloani pacificus; a gadid fish, Laemonema
morosum,  composed the remainder of  the  contents.  Lanternfishes (predominantly
 Notoscopelus  sp. and  Tarletonbeania taylori as well as Diaphus sp.,
 Lampanyctus  sp., Myctophum  sp. and  other  unidentified myctophid species) and
 a fish  species of the family  Sudidae (Paralepis sp.) were the principal prey
 of seven  Dall porpoise described  by Wilke and Nicholson (1958); squids
 (including Watasenia  scintillans  and Ommastrephes sloani pacificus), though
 frequently fed upon,  did  not  provide a  high percentage of the stomach contents

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volume.

Beluga Whale

      Scheffer and Slipp (1948) included a 1940 report of a white or beluga
whale observed in south-central Puget Sound.  This was an extremely rare
occurrence for a whale which was previously not seen south of Cook Inlet,
Alaska.

      Belugas apparently have a broad prey spectrum.  Kleinberg, et al. (1964),
reported prey including arctic cod  (Boregadus saida), capelin, salmon,
flatfish (Pleuronectidae), herring, and crustaceans.  Johnson, et al.  (1966),
examined the stomachs of two belugas taken in the Cape Thompson region of
Alaska and described the contents as arctic  cod and at  least  three species
of shrimp.

Pygmy Sperm Whale

      Very few specimens of this species were reported from the Pacific
coast of North America, though one  from the central coast of Washington was
examined in 1942 (Scheffer and Slipp 1948).  Although it had been dead for
two weeks,  its stomach still contained evidence of 15 squid, at least 10 fish
(including Pacific sandfish, Trichodon trichodon), crab, and shrimp (including
Pasiphaea pacifica, Pandalus borealis?, and Pandalopsis dispar?).

Gray Whale

      Gray whales pass close inshore through Washington and British Columbia
coastal waters during their migration between their calving grounds along
southern California and northern Mexico and the summer feeding grounds in the
Bering Sea.  They usually appear during their northward migration between
February and May and in December and January during their southward migration
(Pike and MacAskie 1968), traveling individually or in groups of two, three,
or four.  Wahl's (1977) four sightings all occurred in May and were
typically close inshore.  It was primarily during the northward migration in
spring that the Indian whalers of the Pacific Northwest concentrated on the
gray whale, highly susceptible to their harvest because of its small size,
moderate speed, and tendency to pass close inshore (Scammon 1974; Scheffer
and Slipp 1948).  They generally cross the mouth of the Strait of Juan de
Fuca in a line between Cape Flattery and Pachena Point, Vancouver Island
(Hatler and Darling 1974; Hart 1977), or Carmanah and Cape Beale (Pike 1962).
The incidence of gray whales venturing inside the Strait of Juan de Fuca is
not high.  One was reported at Point Defiance, in south-central Puget Sound,
in about 1938, and recently a young calf was discovered  floating dead  in
Possession Sound (Seattle P-I, June 21, 1978), and one beached near Neah Bay
in August 1978.  Included in Balcomb's (1978) 1977 Orca Survey were eight
sightings of gray whales, two in the Port Angeles vicinity, two around
Bremerton,  and four in southern Puget Sound.  In general, 20 individuals may
be assumed to frequent Washington's inland waters annually  (Rich Osborne,
pers. comm.).

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      Scammon (1874). Pike (1962), and Rice and Wolman  (1967,  1971)
maintain that in general gray whales do not feed  significantly along the
southern section of their migration route; only one  of  the  84  stomachs
examined by Rice and Wolman (1967) contained prey organisms, zoea of the crab
Pachycheles rudis. Wilke and Fiscus (1961) . Hewell and  Huey (1930),  Rau and
Schevill (1974, citing Ken Balcomb, pers.  comm.), and Hart  (1977),  however,
have provided indications of feeding activity or  significant stomach contents
for some gray whales during the migration  along the  Pacific Northwest coast.
Planktonic euphausiids (Euphausia pacifica) and nektonic  fishes (rainbow
smelt, Osmerus mordax?, northern anchovy)were the prevalent prey in these
instances.  The stomach of the dead calf from Possession  Sound was reported
to contain crab zoea (Bob Everitt, NOAA, pers. comm.).

      On their usual feeding grounds in the Bering and  Chukchi seas (Pike
1962), benthic and epibenthic organisms are the prevalent food, specifically
gammarid amphipods but also other epibenthic crustaceans  such  as mysids,
cumaceans, and isopods (Pike 1962; Rice and Wolman 1967,  1971; Tomilin 1957;
Zimushko and Lenskaya 1970).  Wilke and Fiscus  (1961) described the feeding
whales  making large muddy blotches in  the  water as they came to the surface
to blow, implying  that, "in feeding along  the bottom the whales gathered mud
along with food and were expelling it  through their  baleen  as  they rose to
the  surface."  Tomilin (1957) supposed that the whales  fed  by  scooping and
plowing their mouths into the bottom,  but  Ray and Schevill  (1974),  from
observations of a  captive juvenile gray whale, documented a bottom-sweeping
 feeding sequence wherein "the whale rolls  over far enough so that the cheek
 is about parallel  with the bottom, and the lip is opened  as the tongue,
pressing against  the palate, pushes the gular region away so that it expands,
producing an inflow which brings  in the epibenthic food.  Then the tongue
relaxes and  the gular musculature tightens, reducing the  size  of the mouth
cavity  and expelling water; the food is trapped in the  baleen  fringes.  We
do not  know  exactly what happens next: perhaps a  slight renewed suction of
water  removes  the  food from baleen fringes, and swallowing  presumably follows."
This  behavior,  if  real, would explain  the  asymmetric distribution of
barnacles along the head, producing predominantly "right-sided" (feeding on
 left  side) animals (Kasuya and Rice 1970).

Finback (Fin) Whale

       Historically,  fin whales were the first or  second most abundant whale
 of  the  Pacific Northwest coast.  Pike  and  MacAskie  (1969) indicated that fin
whales  were  the most abundant of  the baleen whales found  along the coast of
 British Columbia;  they were the second most important species  in the commercial
 catch  there  and off  the Washington coast  (Scheffer and  Slipp 1948).   Several
 reports exist of  fin whales sighted or captured in Puget  Sound and the Strait
 of  Georgia.  Although many of the adult whales appear to  be migrants traveling
 between summer  feeding grounds in the  North Pacific  and winter breeding grounds
 to  the  south, young  animals may reside off the Washington and  British Columbia
 coast  during the  summer  (Pike and MacAskie 1968).

       While the little that  is  known of fin whale feeding  in  our region is
 to be found principally in  Pike  (1950),  reports of their food  organisms in
 the North Pacific include  the  expansive work by Nemuto (1959)  , and  Andrews

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(1909), Bethesheva  (1954,  1955),  Mizue (1951),  Nemoto and Kasuya (1965), and
 Thompson  (1940).   Nemoto  (1959)  and Tomilin (1954) showed that fin whales
 have  the  second greatest  number  of baleen plates and the third smallest baleen
 fringe diameters of  all the baleen whales in the North Pacific.  Fin whales
 also  have comparatively coarse baleen fringes, similar to those in blue whales.
 They  found  that euphausiids and  calanoid copepods served as the primary food
 items and pelagic  schooling fishes (primarily  capelin, juvenile walleye pollock,
 Theragra  chalcogramma, and  Pacific  herring) provided secondary contributions.
 Thysanoessa inermis  and _T.  longipes were the principal euphausiid species, and
 Calanus^ cristatus  and ^C.  plumchrus the principal copepod species occurring in
 the samples examined by Nemoto  (1959) .  Copepods predominated in the stomachs
 of whales taken offshore  whereas euphausiids predominated in specimens
 captured  along  the coast  (Nemoto and Kasuya 1965).  Euphausiids CT. inermis)
 were  the  dominant  prey in 48 of  the 50 stomachs with food examined from
 Akutan, Alaska, in 1937 and 1938;  walleye pollock (Theragra chalcogramma)
 and calanoid copepods  (Calanus cristatus) predominated in the other two
 stomachs  (Thompson 1940).   Fin whales were also feeding on euphausiids
 (Euphausia  pacifica, Thysanoessa spinifera) off the coast of California in
 1937; only  one  of  the  14  stomach samples had copepods as the prevalent prey
 (Thompson 1940).

 Sei Whale

      For  many years  the sei whale was largely  ignored by the commercial
 whalers off the Pacific Northwest coast because of its small size and poor
 yield of  oil, but  it became more important with the decline of the fin whales.
 Of British  Columbia's  commercial catch from 1962 through 1966, 57% was
 composed  of sei whales  (Pike and MacAskie 1969).  At the Bay City whaling
 station,  however,  only 21 sei whales were taken between 1911 and 1925
 (Scheffer and Slipp  1948).   There are no confirmed reports of sei whales
 appearing in the inland waters of Puget Sound.


      Amongst the  baleen whales of the North Pacific, sei whales have an
 intermediate number  of baleen plates but have one of the lowest diameters of
 baleen fringes,  second only to young right whales  (Tomilin 1954; Nemoto 1959).
 The  fine  filtering capability of the sei's baleen morphology  is reflected in
 the  predominance  of  small prey organisms, almost exclusively  calanoid
 copepods.  Nemoto  (1959), Nemoto and Kasuya (1965), and Kawamura  (1973)
 illustrated that  calanoid copepods,  including Calanus cristatus, C\ plum-
 chrus,  C. pacificus, and  Encalanus bungi bungi, were the primary food
 species,  £. cristatus  being more dominant in offshore waters  than  the others.
 Other prey  organisms occurring less  frequently in the stomach contents of sei
 whales  were euphausiids  (predominantly Euphausia pacifica), saury, chub
 mackerel, Japanese sardine, Sardinops melanostica, and squid, Ommastrephes
 sloani  pacificus  (Nemoto  1959;  Kawamura 1973).  Fish appeared prominently in
 the diets of whales  collected south of 40°N.
      The  stomach  of  one  sei whale examined at Port Hubron, Alaska, in 1937
 contained copepods,  Calanus cristatus,  and a few amphipods, whereas 12 taken
 in coastal California  had  eaten only euphausiids (Thompson 1940).

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Minke (Little Piked) Whale

     Because minke whales, the smallest of  the  baleen whales occurring along
the Pacific Northwest coast, were not  favored by  the commercial whalers, the
historical records did not reflect  their  actual abundance.   Wahl (1977)
included one report of a minke 179  km  offshore  Westport in September 1976.
Collins (1892) and Waterman  (1920)  reported that  the native whalers off Cape
Flattery and the Strait of Juan de  Fuca did take  minkes.   Scammon (1874)
reported the first specimen  taken in Puget  Sound, a 27-foot female washed
ashore in Admiralty Inlet.   Scheffer and  Slipp  (1948) included four other
verified reports of minke whales in various locations off Puget Sound,
including Whidbey Island, McAllister Creek  (Nisqually Reach), Snohomish River,
and Deception Pass, and one  at Waadah  Island off  Neah Bay.   Pike and MacAskie
 (1969) described five records of minke whales along the British Columbia coast,
but none from inside waters.  Only  two records  of minke whales occurred  in the
commercial whale statistics  from British  Columbia (Pike and MacAskie 1969),
none  from Washington  (Scheffer and  Slipp  1948), and the one stomach of these
which was examined was empty.  Balcomb's  (1978) 1977 Orca Survey also included
 21 sightings of minke whales, all occurring in  northern Puget Sound and
 Admiralty Inlet.  One was sighted in southern Possession Sound west of Everett.
 Recent assessment of  the  survey's sighting  data suggests that between 5 and  20
 minke whales occur  annually  in  the  region's inland waters (Rich Osborne,
 Moclips  Cetological  Society, Orca Survey, pers. comm.).

      Having  an  average of 280 baleen plates (per  side, range 260-300),
 minke whales are  in the  group of right and  gray whales, but they have baleen
 fringe  diameters  closer  to  sei whales  and are classified as a swallowing
 (gulping)  type  (Nemoto 1959).  Accordingly, their diet elsewhere in the North
 Pacific  is  quite  catholic for a baleen whale,  including euphausiids (Euphausia
 pacif ica, Thysanoessa inermis, T_.  longipes,  and T_.  raschi);  fish (Pacific sand
 lance,  walleye  pollock,  and  saury)jcopepods  (Calanus finmarchicus); and
 squid (Nemoto  1959;  Omura and Sakiura  1956).

 Blue Whale

      Prior  to  the protective regulations  imposed  by the International
 Whaling Commission  (IWC)  in 1965, blue whales were one of the most prized
 catches  of  the  commercial whalers operating off the Pacific Northwest coast.
 Because  of  their  large size, speed, and  tendency  to pass well offshore, they
 were not overly abundant  in the catches;  only  13  were taken during the
 operation of the  Bay City whaling station (Scheffer and Slipp 1948) and 320
 in British  Columbia waters  (Pike and MacAskie  1969).  There were no reports of
 blue whales  naturally occurring within the  region's inland waters.

      Blue whale baleen is one of the coarsest  filtering apparatuses amongst
 the baleen whales,  having the highest  number of baleen plates,  the largest
 diameter baleen fringes,  and the second  lowest  number of baleen fringes
 (Nemoto  1959).   In  the North Pacific  their  prey is almost exclusively the
 larger  euphausiids  (Euphausia pacifica,  Thysanoessa inermis, and T\ longipes),
 and only incidentally copepods  (Mizue  1951; Nemoto 1959).  Thompson's  (1940)
 analysis of  blue  whale stomachs brought  to  Akutan and Port Hobron, Alaska,
 whaling stations  substantiated their dependence  upon euphausiids (primarily

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^T. inermis) in that region and those reported  from  California  in  that  source
were identical.

Humpback Whale

     Humpbacks once made up the majority of whales  taken  commercially  off  the
coast of Washington and British Columbia, but  under heavy exploitation there
and in California and Alaska  they declined rapidly  until  the early  1960s
when most of the whaling operations were forced  to  terminate (Scheffer and
Slipp 1948; Pike and MacAskie 1969).  Humpbacks  were placed under IWC
protection in 1966 and 1967-  They appeared to represent  migrants which
veered inshore during their migration to Alaska  in  July and August.  The
wandering humpbacks often ventured into the inland  waters of Puget  Sound and
the Strait of Georgia and the Strait of Juan de  Fuca;  Scheffer and  Slipp
(1948) described three such occurrences in south and central Puget  Sound.
Recent compilation of the Orca Survey's (Moclips Cetological Society)  sighting
data illustrates that about two whales venture into Puget Sound annually
(Rich Osborne, pers. comm.).

     Unfortunately, despite the frequency of occurrence of humpback whales,
few stomachs were examined in the region.  Classified  as  a "swallowing
(gulping) type" like blue whales by Nemoto (1959),  humpbacks have an inter-
mediate number of baleen plates, mid-range baleen fringe  diameters, and a
number of baleen fringes similar to the blue,  fin,  and minke whales.   Pike
(1950) described the prevalence of euphausiids (Thysanoessa spinifera,
Euphausia pacifica) in humpback stomachs in British Columbia, and Hewell and
Huey (1930) listed the euphausiid  Euphausia pacifica  and Pacific sardines.
In the North Pacific, euphausiids (primarily  Thysanoessa  inermis) were the
dominant prey organisms, but  schooling pelagic and  epipelagic  fishes
(especially Atka mackerel, PIeurogrammys monopterygius. capelin, Pacific
sand lance, and walleye pollock, Theragra chalcogramma) were also prominent.
Thompson (1940) described the stomach contents of humpbacks taken in the
western Gulf of Alaska as principally euphausiids,  Thysanoessa inermis and
T_. gregaria, and surf smelt,  Hypomesus pretiosus.

North Pacific Right Whale

     Records of right whales  along the coast of  Washington originated
primarily from unverified reports of native catches (Waterman  1920), but
four were taken during the commercial whaling  off British Columbia  (Pike and
MacAskie 1969).  Scammon (1874) suggested that they were  once  found off
Oregon,  often in high numbers.  Fiscus and Niggol  (1965) recorded  three
right whales off northwest Washington in April 1959.   Gilmore  (1956) suggested
that the coastal waters of California, Oregon, Washington, and British
Columbia were wintering grounds for right whales during former periods of
abundance.  Although no records exist of right whales  occurring in  the region's
inland waters, a recent sighting placed three  whales off  Cape  Flattery
(Rice and Fiscus 1968).

     Neither food habits nor feeding behavior  have  been documented  for this
region.   Right whales have a low number of baleen plates  and the lowest
diameter and highest number of baleen fringes  of  all the  baleen whales, and

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are considered a unique skimming type of feeder (Nemoto 1959).  As a result,
the smaller planktonic crustaceans such as the copepods Calanus plumchrus and
C^. finmarchicus form the bulk of their diet while larger euphausiids were
incidental.
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                              D-2.  Pinnipeds

     Marine mammals from two pinniped families are represented in Puget
Sound—Otariidae,  the eared seals, and Phocidae, the earless seals.  Northern
fur seal,  northern or Steller's sea lion, and California sea lion are Otarii-
dae; Pacific harbor seal, Phoca vitulina richardsi, and northern elephant
seal are Phocidae.  Only northern sea lions and Pacific harbor seals are now
prevalent in the Strait of Juan de Fuca and northern Puget Sound.  The follow-
ing species accounts document the trophic relationships of all pinnipeds
reported from the region; however, the preceding food web discussion considers
only the two prevalent species.

                             Species Accounts

Northern Fur Seal

     Transient fur seals, migrating between their  summer breeding grounds
in  the Bering Sea and their winter feeding grounds off California, occur along
the Washington and British Columbia coasts in March, April, and May.  Wahl
(1977) reported sightings on 14 of 47 trips offshore Westport, Washington,
mostly in May.  Inshore winter residents include mostly 1- to 3-year-old
seals, the older females moving farther south (Spalding 1964).  Commercial
sealers took large numbers offshore until the North Pacific Fur Seal Treaty
of  1911, and native Indians were  reported to have  captured nearly 50,000 fur
seals off British Columbia between 1912 and 1940  (Pike and MacAskie 1969).
Apparently, many fur seals venture into the region's inland waters; Manzer and
Cowan  (1956) listed sightings in  British Columbia's coastal waters through
1955.

     Stomach contents analyses on northern fur  seals in the Pacific Northwest
region were performed as early as the 1930s when  Clemens and Wilby (1933)
and  Clemens, et al. (1936), examined 593 fur seal  stomachs from off the coast
of  British Columbia.  They found  that 84% of the  volume of stomach contents
was  Pacific herring.

     Kenyon's (1950a) synopsis of early food habits data on fur seals taken
in  coastal waters between Washington and southeastern Alaska in spring and
early  summer prior to 1950 indicated that Pacific  herring composed almost
three-quarters of the total prey  composition by weight; other fish, including
eulachon, Pacific salmon, Pacific sardine (pilchard) and Pacific sand lance
were prey of secondary importance.

     Detailed collp.ctions and examinations initiated as a result of the 1957
Interim Convention on Conservation of the North Pacific Fur Seals included
2,113  fur seal stomachs procured  from 1958 to 1961 in British Columbia's
coastal waters (Spalding 1964).   Clupeid fishes (Pacific herring, Pacific
sardine, and northern anchovy) and squid (Loligo  opalescens, Gonatus magister)
formed  the basis  of the  fur  seal  diet.  Whereas the adult  seals migrating  off-
shore  the British Columbia coast  fed principally  upon  clupeids,  the  diet  of
young  seals residing in  protected coastal waters  from  January  through  mid-April
shifted from squid, ratfish, Hydrolagus  colliei,  and sablefish  to  only the squid
Gonatus magister.

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The few young animals left in the region in summer had  consumed  Pacific
herring, Pacific hake, Pacific cod, Gadus macrocephalus,  and  various squid
species; it was at this time that species of Pacific  salmon entered the diet
to any significance, 10% of the total food intake.  Age specific food data
also indicated that larger prey (Pacific salmon, Pacific  cod, Pacific hake,
rockfish, and shad, Alosa sp.) became increasingly important  with increasing
age of the predator.

     The stomachs of fur seals taken off the coast of Washington contained,
in order of decreasing volume, walleye pollock, eulachon  (Thaleichthys
pacificus), American shad (Alosa sapidissima), rockfish,  Pacific herring, and
northern anchovy    (Schultz and Rufn 1936; Wilke and  Kenyon 1954).  Northern
anchovy, rockfish, capelin, and several species of Pacific salmon
(Oncorhynchus kisutch, £. tshawytscha) combined to form 92.3% of the total
food volume in 190 fur seals collected off the coast  of Washington in 1969
(Fiscus and Kajimura 1971).  Seals taken at the entrance  to the  Strait of
Juan de Fuca, however, appeared to have fed principally upon  Pacific herring.

     Wilke and Kenyon's  (1952) collections of 148 fur seals from southeastern
Alaska  in 1950 and  1951  showed that wintering female  fur  seals preferred
Pacific herring  (99.5%) over walleye pollock (0.5%) and squid (Loligo sp.,
trace)  in that region.

     Capelin, eulachon, and Pacific sand lance appear to  assume  greater
  importance as the migrating fur seals move into the  western  Gulf of Alaska
and eastern Aleutian  Islands (Taylor, et al., 1955; Wilke and Kenyon 1957),
and are supplemented by walleye pollock once the seals  enter  their summer
residency in  the  Bering  Sea (Lucas 1899; Taylor, et al.,  1955; Wilke and
Kenyon  1954).  Pacific sand fish (Trichodon trichodon)  composed   94.2% of the
total prey volume and 99% of the total prey numbers from  stomach contents
of  27  fur seals  collected at St. Paul Island in the Pribilof  Islands,  and
sturgeon poacher,  Agonus acepenserinus, composed most of  the  rest (Kenyon
1956).

     Stomachs of  445  fur seals captured in the pelagic  waters off northern
Japan were reported  to contain predominantly lanternfishes, 55.3% by volume,
and squid  (Ommastreph.es  sloani pacificus, Watasenia scintillans), 43.5%
 (Wilke  1951).  Stomachs  of 559 fur seals collected in November in the Sea
of  Okhotsk contained Asian greenling  (Pleurogrammus azonus),  Pacific sand
lance,  smelts  (Osmeridae), walleye pollock, and Pacific salmon (Far Seas
Fisheries Research  Laboratory 1979).  Data collected  by USSR  (VINRO/TINRO
1977)  indicated  that  squid (Gonatus magister, Onychoteuthis banksi). and
salmon  were important prey organisms of fur seals feeding off the southeast
coast of Hokkaido in  1975, and squid  (Onychoteuthis banksi)   and lantern-
fish of fur seals feeding near the eastern shore of Honshu in 1976.

     In general,  fur  seals feed upon epipelagic and neritic schooling fishes
and squid, usually  consuming them whole underwater.   Larger prey such as
lingcod  (Ophiodon elongatus), Pacific salmon, and rockfish are often brought
to  the  surface where  they are vigorously shaken apart (Spalding 1964).  In
almost  every  region,  feeding appears to occur principally at  night when many
prey species have migrated closer to the surface  (Wilke 1951; Fiscus and

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Kajimura 1971; Wilke and Kenyon 1954; Fiscus, et  al., 1963;  Spalding  196A).
Estimates of daily food consumption varied from 5%  to 7%  of  the  total body
weight with an observed maximum of 10%  (Spalding  1964).

     Because of the composition and abundance of  neritic  schooling  fishes
in the Strait of Juan de Fuca, it is probable that  fur  seals feeding  in  that
region would be utilizing Pacific herring, Pacific  sand lance, surf smelt,
longfin smelt (Spirinchus thaleichthys), immature Pacific salmon, and northern
anchovy.  Some demersal fishes such as  Pacific tomcod (Microgadus proximus)
and Pacific cod would probably also enter the diet  incidentally.

Northern or Steller's Sea Lion

     Northern sea lions reside and breed along the exposed coast  of  the Pacific
Northwest, principally on the Scott Islands and Cape St.  James in British
Columbia.  A 1961 census in British Columbia indicated  approximately  1,500
pups and 4,500 adults residing in the province's  waters (Spalding 1964).
Bigg's (1973) report of California sea  lion abundances  on southern  Vancouver
Island included data on northern sea lions occupying the  same haulout  areas;
more than 953 individuals were observed at peak abundance, occurring  in
December.

     Wahl (1977) reported them far offshore the coast of  Washington and
they are commonly dispersed into the inland waters  of Puget  Sound and  the
straits of Georgia and Juan de Fuca when not occupying  the rookeries  in
fall and winter.  Preliminary results from NOAA-MESA-sponsored aerial
surveys in northern Puget Sound, conducted between  November  1977 and  June
1978, indicated a maximum abundance at  Race Rocks (southern  tip  of  Vancouver
Island) of approximately 260 individuals.  Abundances declined rapidly after
April when the majority of the animals  departed Puget Sound  (Bob Everitt,
NOAA, unpubl. data).

     The stomach contents of northern sea lions collected off the British
Columbia coast by the Fisheries Research Board of Canada  included a diverse
spectrum of large prey, principally octopus, rockfish,  walleye pollock,
Pacific cod, Pacific hake, and spiny dogfish (Squalus acanthias) (Spalding
1964).  Rockfish were the most common prey throughout the year,  whereas
Pacific herring was a staple prey in winter, walleye pollock in  summer,  and
rockfish, Pacific salmon, walleye pollock, and Pacific  hake  in fall.

     Northern sea lions widely collected along the  Gulf of Alaska had  fed
principally upon walleye pollock (55% by volume), salmon  (13.3%), starry
flounder (Platichthys stellatus), octopus, skate, and Pacific tomcod;
Pacific salmon, however, occurred in only two of  the 15 stomachs examined
(Imler and Sarber 1947).  Mathisen, et  al. (1962).  described the principal
dietary components of northern sea lions from the Shumagin Islands  in the
northwestern Gulf of Alaska as including nearshore  fishes (mainly greenlings,
 Hexagrammidae,  and rockfish) and invertebrates  (bivalves,  squid,  and
octopus), occurring in 42% and 85% of the stomachs,respectively.  Bulls
tended to consume fish more often than  did cows and yearlings.   There were
no obvious changes in diet during or after the pupping  season.   Stomach
contents of two sea lions from St. Paul Island in the Pribilofs  were

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described in Wilke and Kenyon (1952) as being dominated by either Pacific
sand lance or Pacific halibut with additional contributions by Pacific  cod,
walleye pollock, and starry flounder.

      In general, northern sea lions feed predominantly upon large  fish
occurring throughout the water column as well as small schooling fishes when
abundant in pelagic waters.  Like the fur seal, sea lions prefer to bring
large fish to the surface where the prey is torn apart into edible  pieces  by
violent shaking (Spalding 1964).  They also appear to be nocturnal  feeders,
exhibiting a decline in mean stomach contents volume from a maximum at  dawn
to empty 13.5 hours after sunrise.  Daily food consumption has been estimated
to range from 2% to 4% of the total body weight (Scheffer 1958).

California Sea Lion

      Although at the northern extremity of their reported range, California
sea lions have been sighted and collected along the west coast of Vancouver
Island, British Columbia (Cowan and Guiguet 1965; Guiget 1953), and thus
periodically may enter the Strait of Juan de Fuca to feed.  Bigg's  (1973)
censuses along  southern Vancouver Island in 1971 and 1972 provided  indications
of a population numbering at least 473 individuals occupying eight  haulout
areas.  Peak abundances were recorded in February.  Bigg (1973) also  reported
a haulout area  in the Strait of Georgia on the southeastern side of Dodd
Narrows.

      The NOAA-MESA surveys indicated a maximum of approximately 76 California
sea lions occupying the Race Rocks haulout area between December 1977 and
late April 1978.  Like the northern sea lion, their numbers decline rapidly
thereafter as they depart Puget Sound for their breeding areas off  California
and Mexico  (Bob Everitt, NOAA, unpubl. data).

      Investigations of the food habits of California sea lions in  southern
California have established squid (Loligo sp.) and small fish  (northern
anchovy and Pacific hake) as their main prey items in that region (Bonnot
1928; Scheffer  and Neff 1948; Fiscus and Baines 1966; Peterson and  Bartholomew
1967).  Mate (1973), however, found no squid in the stomachs of California
sea lions collected on the Oregon coast in fall and winter.

      Mate  (1973) also documented a generally nocturnal feeding behavior for
the Oregon sea  lion population that he closely observed, showing 88%  of the
stomachs containing food in morning, 21% at midday, and 7% by evening.
"Feeding trips" were estimated to average 4.2 days.

      Although  there are no data existing for the diet of California  sea lions
occurring in Washington and British Columbia waters, it could be assumed that
their diet would be based principally upon the schooling epipelagic fishes
(Pacific herring, northern anchovy, Pacific sand lance) and some of the more
midwater and demersal forms (Pacific hake, Pacific cod, and walleye pollock)
of the region.
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Pacific Harbor Seal

      The most abundant and ubiquitous pinniped occurring in  the protected
waters of Washington and British Columbia, Pacific harbor seals are most  common
in estuaries and river deltas but are seen in shallow sublittoral waters  along
the entire coastline and even ascend large rivers to reside in lakes  (Cowan
and Guiguet 1965; Pike and MacAskie 1969).  Hart  (1977) reported one  sighting
80 km offshore Westport, Washington.

      No quantitative estimates of harbor seal populations have been  made for
the region as a whole, although Pike and MacAskie (1969) estimated
approximately one seal per mile of shoreline—e.g., 17,000 seals—for coastal
British Columbia.

      Aerial survey counts recently conducted by NOAA under the auspices of
the MESA program have provided the first detailed inventory of harbor seals
in northern Puget Sound and along the Strait of Juan de Fuca.  The total
number of seals enumerated during these surveys varied between 643, taken in
December 1977, to 1,618 (excluding 143 pups), taken in August 1978 (Bob
Everitt, NOAA, unpubl. data).  Of the 17 sampling areas subdividing the
region, the following eight had the highest abundances, in decreasing order:
San Juan Island, Smith and Minor islands, Patos Island to Lummi Island,
Rosario and Haro straits, Bellingham to Padilla Bay (all in north Puget Sound);
Protection Island, Dungeness to Sequim Bay, Becher Bay to Discovery Island
(along the Strait of Juan de Fuca).

      Scheffer and Sperry (1931) were the first to quantitatively analyze
harbor seal stomach contents from Puget Sound.  The 100 stomachs examined
contained, by volume, 93.6% fish (Pacific tomcod, flounder (Pleuronectidae),
Pacific herring, Pacific hake, sculpin (Cottidae), codfishes  (Gadidae),
walleye pollock, and shiner perch, Cymatogaster aggregata), 5.8% molluscs
(squid and octopus), and 0.6% crustaceans (shrimp).  Scheffer (1928)  had
recorded the stomach contents of 22 adult harbor seals from southern  Puget
Sound (Nisqually Flats) to be Pacific herring, Pacific tomcod, shiner perch,
sculpin, shrimp, crab, squid, octopus, skate, starfish, and flounder; only
two stomachs contained remains of Pacific salmon.

      The most recent and comprehensive data on harbor seal food habits in
Puget Sound have been assembled by Dr. John Calambokidis (undergraduate
study project, Evergreen State College, 1978), based upon the composition
and abundance of fish otoliths (ear bones) in the feces deposited at  haulout
areas.  Pacific hake (73% of total numbers), plainfin midshipman (Porichthys
notatus^ 7%), and staghorn sculpin (Leptocottus armatus, 6%) predominated in the
overall diet combined for all regions, but there were distinct differences in
the dominant prey species for the different regions of Puget Sound.   Pacific
hake and plainfin midshipman were most prevalent in the Hood Canal region;
staghorn sculpin and Pacific hake predominated in south Puget Sound;  and
blackbelly eelpout (Lycodopsis pacifica), an unidentified eelpout (Lycodes sp.)
and Pacific herring were most important in the region of concern, north Puget
Sound (based on collections at Smith Island).  Calambokidis' data also
indicate some seasonal changes in the diet composition.  Pacific hake clearly
dominated harbor seal fish consumption in April and July through September 1977,

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plainfish midshipman predominated in early May. blackfin  sculpin (Malacocottus^
kincaidi) in early June, and walleye pollock in late June.

     In British Columbia, Fisher (1952) summarized  the  stomach contents of
50 harbor seals taken from the Fraser River, the Queen  Charlotte Islands,  and
the Skeena River between August 1945 and November 1946.   Rockfish and octopus
were the most commonly occurring prey, but pink salmon  (Oncorhynchus gorbuscha) .
Pacific herring, and other unidentified fishes composed most  of the total
stomach contents volume.  Fisher (1952) also indicated  that  there was
considerable temporal and spatial variability in the seals' diets and that
they appear to move along the Pacific coast in response to fluctuations in
abundance of preferred food organisms.  Spalding's  (1964) comprehensive
comparison of harbor seal feeding habits with those of  fur seals and sea
lions along the coast of British Columbia showed a  diet quite similar to the
sea lions, based upon salmon, octopus, squid, clupeids, and rockfish.  The
contribution by Pacific salmon, however, was biased by  a  high number of
collections made in the vicinity of salmon spawning streams  in fall.  Although
seasonal  sampling of harbor seals was inconsistent, it  appeared that eulachon
may be most important in winter; octopus, Pacific herring, rockfish, and
salmon in summer; and salmon in fall.

     Harbor seals in Alaskan waters have shown a similar  feeding preference
for fish, including walleye pollock and Pacific herring in Prince William
Sound and eulachon at Copper River delta (Imler and Sarber 1947; Pitcher 1977),
and hexagrammids  (Hexagrammos lagocephalus , Pleurogrammus monopterygius)
and walleye pollock in  the Aleutian Islands (Wilke  1957;  Kenyon 1965).
Walleye  pollock  (83% by volume) and Pacific herring (10%) were the predominant
food organisms of harbor seals sampled in the Sea of Okhotsk  off Hokkaido
 (Wilke 1954).  Octopus  and squid commonly appeared  in the diet in all areas,
though never as  a high  percentage of  the prey volume.

     Harbor seal pups and juveniles have been reported  to feed specifically
upon shrimp (Havinga 1933; Fisher 1952; Bigg 1973)  and  small  fishes (Pitcher
1977) at a general increase in prey size with increasing  predator size
 (Spalding 1964).  Unlike fur seals and sea lions, harbor  seals do not appear
to be nocturnal  feeders  (Spalding 1964) .  Daily food consumption by harbor
seals has been estimated at 3.7% (Pitcher 1977), 5% (Havinga  1933), and 6%
 (Scheffer 1958)  of the  total body weight.

Northern Elephant Seal

     Although the nearest breeding site of the northern elephant seal is
1,000 miles to the south, off southern California and Mexico, they have been
reported into the Gulf  of Alaska, including a specimen  found  at Prince of
Wales Island in  southeastern Alaska (Willet 1943) and several reports at
Middleton Island and Baranof Island (Fiscus et al.  1977).  Wahl (1977)
reported three sightings between 50 and 65 km offshore  Westport, Washington,
and numerous sightings  and specimens were reported  from British Columbia,
especially in Hecate Strait (Pike and MacAskie 1969) .   The stomach of one
specimen recovered from the west coast of Vancouver Island contained hagfish.
The only feeding behavior information from the Puget Sound region is of an
                                                  N° P°lnt in n°rth-Central
                                     283

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Mathisen, 0. A., R. T. Baade &  R. J. Lopp.  1962.  Breeding habits, growth
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Miyazaki, N.,  T. Kusaka  & M. Nishiwaki.   1973.   Food  of Stenella caeruleoalloa.
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Mizue, K.  1951.  Food of whales  in  the adjacent waters of  Japan.   Sci.  Rep.
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      porpoise  in the Bering Sea and North Pacific  Ocean.  Bull. Fac.  Fish.
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Nemoto,  T.  1959.   Food  of  baleen whales  with reference to  whale movements.
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Nemoto,  T. & T. Kasuya.   1965.  Foods  of  baleen  whales in the  Gulf of Alaska
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Nishiwaki, M.  & C.  Handa.  1958.  Killer  whales  caught in the  coastal waters
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                                      287

-------
Okutani, T.  & T. Nemoto.  1964.  Squids as the food of sperm whales  in  the
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Pike, G. C.   1962.  Migration and feeding of the gray whale (Eschrichtius
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                                     288

-------
Scammon,  C.  M.   1874.   The marine mammals of the north-western coast of
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     348.

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                                     289

-------
Tomilin,  A.  G.   1957.  Cetacea.  Mammals of the USSR and adjacent  countries.
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Wilke, F. and K. W. Kenyon.  1954.  Migration and food of the northern fur
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Wilke, F. and K. W. Kenyon.  1957.  The food of fur seals in the eastern
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Wilke, F., T. Taniwaki and N. Kuroda.  1953.  Phocoenoides  and Lagenorhynchus
     in Japan,  with notes on hunting.  J. Mammal., 34(4) :488-497.

Wilke, F.  1954.  Seals of northern Hokkaido. J. Mamm. 35(2):218-224.

Wilke, F.  1957.  Food of sea otters and harbor seals at Amchitka  I.  J. Wildl.
     Mgmt. 21(2):241-242.

Wilke, F. and A. J. Nicholson.  1958.  Food of porpoises in waters  of Japan.
     J. Mammal 39(3):441-443.

Wilke, F., and C. H. Fiscus.  1961.  Gray whale observations.  J.  Mamm. 42:
     108-109.

Willet, G.  1943.  Elephant seals in southeastern Alaska.   J. Mammal. 24:500.

Zenkovich, B. A.  1938.  On the grampus or killer whale, Grampus orcaLin.
     Priroda 4:109-112 [in Russian].

Zimushko, V. U. and S. A. Lenskaya.  1970.  Feeding of the  gray whale
     (Eschrichtius gibbesus) at foraging grounds.  EkologL>a Akad.  Nauk SSSR
     l(3):26-35 [in Russian].  Engl. transl., Consultants Bureau,  Plenum
     Publ. Corp., 1971, Ekologiya 1(2):205-212.

VlNRO (Ail-Union Research Institute of Marine Fisheries  and Oceanography/
     TINRO (Pacific Research Institute of Fisheries and  Oceanography)  [USSR]
     1977 The USSR fur seal investigations conducted in  1976.  Moscow.
                                     290

-------
                           APPENDIX E

SEASONAL DISTRIBUTION AND  ABUNDANCE OF  FOOD WEB NODES AND NUMBER
  AND RELATIVE IMPORTANCE  OF  FOOD  WEB LINKAGES CHARACTERIZING
             NEARSHORE HABITATS  OF NORTH PUGET SOUND
                 AND THE STRAIT  OF JUAN DE FUCA
                               291

-------
Appendix Table E-l.
Distribution and abundance of food web nodes characterizing neritic habitats
of north Puget Sound and the Strait of Juan de Fuca.

Cherry Point



Anacortes



W. Whidbey I.
Burrows Bay


San Juan Is.



Port Angeles




Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter

3
3
1
1
3
4

1
1
1
1
1
3
2
1
1
1
1
1
1
Ik-rbivorcs
5
2
1
2
3
3
1
1
3
2
1
2
3
1
1
2
3
2
1
1
Hlxcd
1
1
1
1 1
1
1 1
1
1
1



111
111

1
1
1 1
1
1
Pl.inklivorcs
10
8 1
3
5
11
11 1
2
2
5
7
2
4
8
7 1
5 1
3
6
6
3
3

1

1
1
1


1
1

1
2
1

1
2
1

1
1
lionthivores PisCtvorcs
1
1
1
3
1
1
1
2
1
3
2
2
1
3
2
3
1
4
2
3
Terrestrial Non-

1 6
3
1

1 5
3
1

5
3


6
3
1

1 6
3
1
ToMl
nndi'K
21
22
11
14
20
28
8
9
12
18
10
10
19
23
14
13
13
22
12
11

-------
                 Appendix Table  E-2.   Number  and  relative  Importance of  food web  linkages to  trophic levels characterizing neritic habitats
                                      of  north  Puget  Sound  and  the  Strait of Juan de  Fuca.   (1° - primary, 2° - secondary, 3°  -  tertiary,

                                      Incid - incidental trophic  linkages,  Nn - number of food web nodes indicated on Table E-l.)
ro
VO
10


Cherry Point



Anacortes



W. Whidbey I.
Burrows Bay


San Juan I .



Port Angeles





Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
p,-
P-S«
3
3
1
1
3
4

1
1
1
1
1
3
2
1
1
1
1
1
1

'•""' r.r."er» rli"
3* Incid 1* 2* 3" Incld 1* 2
5
2 1
1
2
3
3 1
1
1
3
2
1
2
3 1
1 1
1
3
3
2
1
1

• 3' Incld !• 2' 3' Incld 1* 2'
2
2 1
2
5
2
2 1
2
2
2



2
4 1
1
2
2
4
2
2
rlnviry
3' Incld
2
2


2
2


2
2


A
2
2

2




1"
2
3
4
4
3
2
1
2
3
4
2
4
4
41
5
2
4
3
1
2

2' 3' Incld
37 8
25 4
2
2 2
36 9
5 5 7
1 2
3 2
14 1
3 3
2 1
2 1
8 6
2 6
1
13 2
14 1
4 5
31 3
21 2

1' 2' J'
2

1
111
2


1 2
2
1 2
3
1 1
2
1 2
3
1 3
2
112
3
2 1

Incld

8
3
5

8
3
1

8
3

1
10
5
3

12
3
2

!•
10
9
6
8
9
11
2
5
7
8
4
8
11
10
8
7
8
7
3
4

2'
3
2

3
3
5


1


1



1
1
1
3
4

3' Incld
13 8
10 12
3 5
6 7
10 9
9 17
3 5
7 3
8 1
7 11
5 4
2 1
16 2
10 16
5 6
8 5
10 1
10 17
6 6
4 4
Totnl 1
"l
34
33
14
24
31
42
10
15
17
26
13
12
29
36
19
21
20
34
18
16
M No.
llnk.igtf
(Nj/Nn)
1.62
1.50
1.27
1.71
1.55
1.50
1.25
1.67
1.42
1.44
1.30
1.20
1.53
1.57
1.36
1.62
1.54
1.55
1.50
1.45

-------
              Appendix fable E-3.   Distribution and abundance of food web nodes characterizing rocky sublittoral habitats
                                   of north Puget Sound and the Strait of Juan de Fuca.
S3

RCT.IOTI (location)
Cherry Ft.
(Barnes I.)


W. Whidbey I.
Burrows Bay
(Allan I.)

San Juan Is.
(Pt. George)


Strait of
Juan de Fuca
Composite

Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter




6
6
6
6
6
6
6
5
6
6
6
6

4
6
llcrblvorei
211
211
211
211
211
211
211
211
211
211
211
211

212
2 1 1
HUcd
1
1

1

1




1
1

1
2
Planktlvorcs
36 1
35 1
32 1
31 1
38 11
38 11
25 11
23 11
37 11
26 11
33 11
32 1

46 11
3 14 1
Ili'tlthivor
1
1
1
1
1
1
1
1
1
1
1
1

1
1
•cs
9
9
8
9
7
9
7
7
8
9
9
10

9
11
Piscivorcs
3
1 3
1 4
4
3
1 3
1 3
1 2
1 2
1 5
1 4
3

1 5
5 1

34
34
30
30
34
38
31
27
34
36
34
31

38
48

-------
Appendix Table E-4.  Number and relative Importance of food web linkages to trophic levels characterizing rocky sublittoral
                     habitats of north Puget Sound and the Strait of Juan de Fuca.   (1* • primary, 2° • secondary,
                     3° - tertiary, Incid - incidental trophic linkages, Nn - number of food web nodes indicated on Table E-3.)

Cherry Point
(Barnes I.)


W. Whidbey I.
Burrows Bay
(Allan I.)

San Juan Is.
(ft. George)


Strait of
Juan de Fuca

Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter



1' 2* 3' Incld
5
5
5
5
5
5
5
4
5
5
5
5

3

1' 2" 3' In
4
4
4
4
4
4
4
4
4
4
4
4

4

cli) 1' 2' 3' InclJ !•
1
1
1
1
1
1
1
1
1
1
1
1

1 1

2'
2
2
1
2
1
2
1
1
1
1
2
2

2

>• I:
9
8
8
9
8
9
6
6
8
6
9
9

10
od
neld
2
2
2
2
2
2
2
2
2
2
2
2

4

3 3
3 3
211
2 1
2 6
4 6
3 3
3 3
215
4 5
3 3
2 1

3 3

6
6
6
6
6
6
6
6
6
6
6
6

6

229
229
118
129
219
2 1 12
4 8
1 8
129
3 1 10
229
317

1 3 15

20
18
19
19
20
18
16
16
20
20
17
17

20


1
1


1
1


1
1


1

2
2
2
2
3
2
2
3
2
2
2
1

1

3
3
2
3

2
1

2
2
2
4

4

3 15
2 16
2 14
3 13
3 14
3 17
3 14
1 13
1 13
3 18
3 16
2 15

6 13

6 24
6 23
5 19
6 22
5 23
5 29
7 18
4 17
6 24
4 23
6 23
4 21

6 32

31
28
29
30
31
29
27
25
29
31
28
27

36
Tot.ll »
"l
76
73
67
71
73
80
66
59
72
76
73
67

87
x No.
llnkmci
(Nj/Nn)
2.24
2.15
2.23
2.37
2.15
2.11
2.13
2.19
2.12
2.11
2.15
2.16

2.29

-------
Appendix Table E-5.  Distribution and abundance of food web nodes characterizing rocky and
                     cobble littoral habitats of north Puget Sound and the Strait of Juan de Fuca.








(SO
VO
ON

Rocky
Strait of Juan
de Fuca and
San Juan Is.

Cobble
Strait of Juan
de Fuca and
San Juan Is.



Spring
Summer
Autumn
Winter

Spring
Summer
Autumn
Winter


6
7
6
7

4
6
5
6


3
3
3
3

3
3
3
3
Herbivore";

2
2
2
2

2
1
2
2
Hixvd

2 2
2 3
2 2
2 3

2 1
121
121
121


10
9
9
8

6
5
5
6
Plankctvorcs






1
1
1
1
Ronthivores

1
1
1
1

1
1
1
1


14
13
12
14

10
10
11
10
Plscivores

1
1
1
1

2
2
2
2


41
41
38
41

32
33
34
35
Composite
                                                                                                                  12
                                                                                                                                                  35

-------
N)
IO
                      Appendix Table E-6.  Number and relative importance of food web linkages to trophic levels characterizing rocky and cobble
                                           littoral habitats of north Puget Sound and the Strait of Juan de Fuca.  (1* « primary, 2° • secondary,
                                           3* • tertiary, Incid - incidental trophic linkages, Nn - number of food web nodes indicated on Table E-5.)

Rocky
Strait of Juan
de Fuca and
San Juan Is.

Cobble
Strait of Juan
de Fuca and
San Juan Is .



Spring
Summer
Autumn
Winter

Spring
Summer
Autumn
Winter


6
7
6
7

3
4
4
5
n,v, „,,,„.

5 1
5 1
5 1
5 1

4
3
4
4


2 1
2 1
2 1
2 1


1
1
1
"r f d


1 22
1 24
1 22
1 33

2 4
2 4
2 4
2 4
114
214
114
114

1
1
1
2


16
16
16
16

3
3
3



3 4
2 4
1 7
2 4

2 3
2 2
1
1 3


21
19
16
16

9
6
17
9


1
6
1
7

15
15
13
17



11
10
11
10

11
12
12
11
2 17
1 18
2 15
2 17

8 9
7 10
9 9
9 13


9
9
12
10

6
5
4
5

38
37
33
33

24
22
33
24


19
23
19
25

26
25
25
26
Totnl 1
"l
83
87
79
85

65
84
71
68
x No.

2.02
2.12
2.08
2.07

2.03
2.54
2.08
1.94

-------
            Appendix Table E-7.
Distribution and abundance of food web nodes characterizing gravel-cobble shallow sublittoral

habitats of north Fuget Sound and the Strait of Juan de Fuca.
ro
vo
oo

Cherry Point



(Legoe Bay)



Anacortes
(Cuemes S.)


W. Whidbey I.
Burrows Bay
(Alexander's
Beach)
(West Beach)



San Juan Is.
(Deadman Bay)


(South Beach)




Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter

5
4
5
4
4
5
5
3
5
5
5
4
4
5
5
5
5
4
5
4
4
5
5
4
4
4
4
4
Ht-rbtworos


1 1
2
1
2 1
2 1
1
2
1 1
2 1
2




1



1
1
1
1

1 1
1
1
HlKcd
1
1
3
1 2
3
3
2

1
2
3
2
1
1
1


1
2
2
2
2
3
2
1
3
1
1

4
2
4
3
5
5
5
3
3
4
4
4
2
2
3
3
3
3
4
2
4
5
3
4
4
5
5
4
Planktlvorcs
4
5
8
4
6
7
12 1
6
3
4
10 1
3 1
1
1
5
1
3
4
3
1
13 1
16 1
8 1
12
6
12 1
6 1
4 1



1
1


1
1
1

1
1

1
1
1


1
1

1
1
1
1
1
1
1

3
2
8
9
6
6
8
3
6
6
13
9
5
4
10
4
3
9
10
6
4
11
16
7
6
11
9
8
risclvores
1
1
2
2
1
1
2
2
1
1
2
2
1
1
2
2
2
1
1
2
1
1
2
2
1
1
1
2

1 19
15
1 34
27
1 27
1 31
1 40
1 20
22
24
42
28
14
15
27
16
17
22
26
1 19
30
43
40
33
23
40
29
26

-------
Appendix Table E-7, cont'd

Port Angeles
(Dungeness
Spit)

(Morse Creek)



(Kydaka Beach)




Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
IU rblvor.'s
5
4
3
5
6 11
6 1
4
4
4
4
3
3
MUtd
1
1
1
1
2
2
2
1
2
1
1
2

2
1
3
2
2
3
2
2
2
2
2
2

1
2
2
1
3
3
1

1

1
1

6
6
4
5
7
6
6
7
4
7
4
5
TUclvorcs
1
1
2
2
1
1
2
2

1
2
2

16
15
15
16
22
22
17
16
13
15
13
15

-------
                Appendix Table E-8.
                                     Number and relative  importance of  food web  linkages  to  trophic  levels  characterizing gravel-cobble
                                     shallow sublittoral  habitats of north Fuget Sound  and the  Strait  of  Juan de Fuca.   (1°  • primary,
                                     2° = secondary,  3° '• tertiary, Incid = incidental  trophic  linkages,  Nn - number of food web
                                      nodes  indicated on Table E-7.)
CO
O
O

Cherry Point



(Legoe Bay)



Anacortes
(Guemes, S.)


W. Wliidbey I.
Burrows Bay
(Alexander's
Beach)
(West Beach)



San Juan Is.
(Deadman Bay)



Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter

4 1
4
4
it 1
4 1
5 1
4 1
4
4 1
4 1
4 1
3 1
4
5
4
5
5
4
5
4
4
5
5
4
Fhytorlonkco. H,c,o,l,
1 1

12 12
4 12
12 1
13 1
14 12
3 12
13 1
1 3
13 1
12 1

1 1
1 1
1 1
1

1 1
1 1
1
2 1
2 1
2 1

1

3
2
2
2
1

1
2
3
2




1
1
2
1
2
3
2
2

6
4
7
6
4
7
9
3
5
7
8
7
4
4
4
4
3
5
8
5
8
8
8
7

7
3
6
4
6
6
6 2
61 2
4
5
52 2
52 2
4 1
4 1
4 1
5
4
4 1
5 1
4
4 2 >'• 2
42 2
42 2
52 2
•y
4
4
4
4
4
4
4
<4
4
4
5
5
5
4
4
4
4
4
4
4
4
4
4
4

3 3
5 2
7'5
3 5
7 4
6 5
5 7
3 2
2 4
5 3
10 6
4 4
3
1 1
6 4
2 1
2 2
5 4
4 5
2 1
9 2
9 7
10 5
10 3

3

7
7
3
7
13
6
5
6
15
9
,7
5
7
5
5
8
9
6
10
15
14
9

4
3
7
7
5
6
5
1
6
4
5
3

2
4
1
2
3
7
3
6
15
11
4

2 8
2 9
6 13
4 13
3 13
1 14
7 13
2 13
2 9
2 12
7 19
4 11
1 5
1 7
5 12
2 8
3 8
5 10
4 10
3 7
2 16
5 18
7 19
9 18

5 9 18
2 4 10
8 14 25
8 13 20
7 9 20
8 14 19
9 22 25
2 11 12
6 10 18
6 13 16
10 25 24
7 18 19
3 11 10
1 9 12
4 11 18
1 9 13
3 8 13
5 13 16
7 17 21
2 11 15
4 18 16
10 23 29
7 24 20
5 18 23
Total 1
"l
40
25
60
54
49
55
69
38
43
47
78
55
29
29
45
31
32
44
55
35
54
80
70
64
X No,
llnk.igti
(Nj/Nn)
2.11
1.67
1.76
2.00
1.81
1.77
1.73
1.90
1.95
1.96
1.86
1.96
2.07
1.93
1.67
1.94
1.88
2.00
2.12
1.84
1.80
1.86
1.75
1.94

-------
                   Appendix Table E-8,  cont'd
Ul
O

(South Beach)



Port Angeles
(Dungeness
Spit)

(Morse Creek)



(Kydaka Beach)




Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
fir.ncr< gr.ncrs pl.int £
1' 2' 3* Incid 1' 2* 3' Incld 1* 2* 3
4 11
5 31
4 31
4 21
5
4
3
5
6 2
6 1
4
4
4
4
3
3

' Incid I' 2
2
4
2
2


1

1
2
1

1

1
1

• 3'
6
10
8
6
4
2
5
4
5
6
5
4
5
4
3
5

Incid
4
3
3
4
4
3
4
4
4
4
4
4
4
4
4
4
Prim
I 1' 2' 3'
1 2
1 2
1 2
1 1
1
1
1
1 1
1
1
1 1

1
1
1

Incid
4
4
4
4
5
4
4
4
5
4
4
5
4
4
4
4

!• 2'
4 4
7 7
7 2
4 2
1 2
4 3
2 2
1 2
1 4
2 1
2
3
2 1
1,4
1 2
3 1

3* ]
9
19
7
6
3
2
2
5
8
7
9
6
5
3
2
3

Incld !• 2'
7
17
10
7
4
2
2
1
10
8
3
4
2
6
2
2

1 3* Incn
4
6
5
5
2
2
4
5
3
1
5
5

2
3
4

i i'
9
16
15
11
7
8
6
6
10
10
5
5
6
6
5
7
Suf
• 2'
6
11
4
4
3
3
3
2
6
3
3
4
2
4
5
2
itot.i
J' I
15
31
17
14
7
4
7
9
13
13
14
10
10
7
3
8
i
ncld
20
25
23
21
13
11
14
14
22
17
16
18
10
16
13
14
Tot.il 1
llnk.nsoi
"l
50
83
59
50
30
26
30
31
51
43
38
37
28
43
26
31
lln
per
IN1
2.
2.
2.
1.
1.
1.
2.
1.
2.
1.
2.
2.
2.
2.
2.
2.
No.
k.in
-------
Appendix Table E-9.
Distribution and abundance of food web nodes characterizing sand/eelgrass shallow sublittoral
habitats of north Fuget Sound and the Strait of Juan de Fuca.
Herbivores Mixed
Rer.lon (location)
Cherry Point
(Birch Bay)


San Juan Is.
(Eagle Cove)


j5 Strait of Juan
de Fuca
(Beckett Ft.)

(Twin Rivers)



Season
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
urocessor planes feeders
5 2
4
421
431
4 2
4
4 2
3 3
3 1
4
3 2
331
3 1
3 1
4 2
3 2
fcr.i7.crs r.r.izcrs detriti
1 4
1 4
1 2
2
2
3
3
2
2
1 2
2
2
4
4
3
3
is Oinnlv<
4
4
4
4
2
3
4
2
4
3
4
6
3
5
4
4
Pl.-mkLivorcs Benthlvoi
res Plscivorcs
ores carnivores carnivores feeders feeders feeders Carnivores PelaKlc Demersal
5
6
5
3
2
5
7
1
2
7
4
6
6
9
6
5
12
13
13
9
10
10
10
8
10
10
7
11
9
9
9
8

origin fcndlnK nodrri
33
33
32
26
22
25
30
19
22
27
22
32
26
31
28
25

-------
                  Appendix Table E-10.
U>
O
U>
Number and relative importance of food ueb linkages to trophic levels characterizing sand/eelgrass
shallow sublittoral habitats of north Puget and the Strait of Juan de Fuca.  (1° - primary, 2' -
secondary, 3* - tertiary, Incid ™ incidental trophic linkages f Nn « number of food web nodes

Indicated on Table  E-9.)

B..CIOII (loc.itlon)
Cherry Point
(Birch Bay)


San Juan Is.
(Eagle Cove)


Strait of
Juan de Fuca
(Beckett Ft.)

(Twin Rivers)




Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Bi.ircrs
5
4
4
4
4
4
4
3
3
4
3
3
3
3
4
3

1
1
2
1





1

1

1
2


2

2
3
2

2
3
1

2
3
1


2

6
6
It
4
3
3
3
3
2
2
2
2
4
4
3
3

10
12
8
7
6
9
11
5
10
10
9
14
10
14
13
11

6
6
6
7
6
6
6
6
6
7
7
6
6
7
7
7

2
1 2
1 2
2
2
2
2
2
2
1 2
2
2
2
2
2
2

2 4
2 5
2 4'
2 1
2 1
2 3
2 2
2
2 5
3 5
2 1
2 3
2 5
2 6
2 4
2 2

4
4
4
3
3
4
7
3
1
5
1
4
4
7
5
6

16
16
19
15
15
13
16
11
12
11
17
17
12
12
13
11

11
11
8
4
9
12
11
2
9
10
5
11
10
10
9
5

12
11
13
9
7
7
8
6
9
10
6
10
9
10
10
5

10
10
8
7
6
7
10
6
3
8
3
6
8
11
8
9

28
30
29
24
23
24
29
18
24
23
28
33
24
28
28
24

19
19
16
13
17
20
19
10
17
20
14
19
18
19
18
14
Tot.il 1
"l
69
70
66
53
53
58
66
46
53
71
51
68
59
67
64
52
K NO.
lui.ir,"
(Nj/Nn)
2.09
2.12
2.06
2.04
2.41
2.32
2.20
2.42
2.41
2.63
2.32
2.13
2.27
2.16
2.29
2.08

-------
Appendix Table E-ll.
                                   Distribution and abundance of food web nodes characterizing mud/eelgrass

                                   shallow subllttoral habitats of north Puget Sound and the Strait of Juan de Fuca.
O
•C-
Herbivore* Mixed
Rrelon {locution)
Anacortes
(Fidalgo Bay)


San Juan Is.
(Westcott Bay)


Port Angeles
(Jamestown-
Graysmarsh)

Season processor
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
Spring
Summer
Autumn
Winter
4
4
It
4
4
4
4
4
5
2
3
3
plants
2

2
4



1
3

2
4
feeders
1
1
2
2
1
3
3
2
1
2
1
1
Erasers ernzcrs detritus Omnlvores
1 1
1 1
1 1
112
3
2
1 1
1
112
1
2
1
5
6,
8
5
5
8
8
3
3
4
6
4
rlanktivores Bcnthivorcs Plsclvores
carnivores carnivores feeders feeders feeders Carnivores Pelaelc Demersal origin
8
5
3
4
9
10
9
6
6
4
5
2
5
8
11
9
8
1 11
1 10
6
1 16
5
6
6
4
1
1
1
1
1
1
1
1
1
1
1

feeding. nodcH
31
27
33
33
31
40
38
24
40
19
26
22

-------
Appendix Table E-12.
Number and relative Importance of food web linkages to trophic levels characterizing mud/eelgrass shallow
•ubllttoral habitats of north Puget Sound and the Strait of Juan de Fuca.   (1* - primary, 2* - secondary,
3* - tertiary, Incld - Incidental trophic linkages, Nn - number of food web nodes Indicated on Table E-ll.)

teflon (location)
Anacortes

San Juan Is.


Port Angeles
(Jamestown-
Graysmarsh)

Fhytoplankton Nacronli;**
Spring
Autumn
Spring
Autumn
Winter
Spring
Summer
Autumn
Winter
4
4
4
4
4
5
2
3
3
2
3
1
3
2
2
2
1
1
2 26
2

1
1
4

2
4
2 21
4 12
2 21
1 9
2 10
1 11
2 18
1 11
4
4
5
5
4
6
4
4
4
1 2
1 2
2 2
2 2
2 2
2 2
2
1 2
2
lucid
2
2
2
2
2
2
2
2
2

6 3
13
2 6
1 7
5
5 4
2 4
1 5
3

8 18
9
8
12
4
6
5
4
7
9
12
22
8
31
2
8
4

2 2 14 6 18 26
2 2 10 6 34 17
2 1 7 12 24 20
2 1 9 11 37 31
2 1 7 8 17 15
2 2 16 8 20 41
2 1 6 5 20 9
2 1 7 8 26 15
2 8 4 22 10
Total (
"l
64
67
63
87
47
85
40
62
44
• No.
llnk«|*«
(Mj/Hn)
2.06
2.03
2.03
2.29
1.96
2.13
2.11
2.38
2.00

-------
                               APPENDIX F

  MECHANISMS OF PETROLEUM HYDROCARBON INFLUENCE UPON FOOD WEB STRUCTURE

              F-l.  Effects Upon and Within Marine Organisms

     The literature describing the biological effects of petroleum
hydrocarbons on marine organisms is extensive.  Recent symposia and
syntheses have summarized the state of the knowledge (Baker 1978; Am. Inst.
Biol. Sci. 1976; Wolfe 1977; Malins 1977; Mclntyre and Whittle 1977; Fish.
Res. Board Can. 1978) and the reader is referred to these for a thorough
discussion of petroleum in marine ecosystems.  Documentation of the acutely
toxic levels and fractions of petroleum hydrocarbons has provided accurate
indications of short-term effects on marine biota (Craddock 1977).  In the
long run, however, sublethal or chronic effects may be more important to
food web structures than acute effects, in that subtle changes in behavior,
growth, natural mortality, reproduction, and general physiological condition
may alter or eliminate critical predator-prey linkages (Blumer 1970; Sprague
1971; Tarzwell 1971).  A brief introduction to     biological fate and
effects in selected marine organisms is necessary for our understanding of
the potential incorporation and transfer of petroleum hydrocarbons in food
webs of north Puget Sound and the Strait of Juan de Fuca.

F-l-a.   Toxic Components

     Of the two basic categories of the water-soluble fractions of petrol-
eum oils, the volatile and the non-volatile, the volatile components have
generally been shown the more toxic, and thus have received the most
attention (Moore and Dwyer 1974; Donahue, et al., 1977; Lee, et al., 1973:
Struhsaker 1977; Morrow, et al., 1975).  Although more persistent and more
likely to enter the food web, non-volatile aromatics and their biological
effects have not been as well studied (Emery 1970; Winters, et al., 1976).
For example, Lee and Nicol  (1978) pointed out that although the volatile
aromatics were measurably more toxic to the marine amphipod Elasmopus
pectenicrus, the concentrations which induced acute mortality were higher
than would naturally occur in the water-soluble fraction.  The non-volatile
aromatics, although much less toxic, had a greater potential of biological
interaction with the marine organism because of their persistence at high
concentrations in the environment.

F-1-b.   Uptake, Retention, and Depuration

     Uptake and retention of petroleum hydrocarbons appear to vary accord-
ing to the type as well as the form of hydrocarbon.  In general, aromatic
hydrocarbons of high molecular weight will be retained in an organism's
tissues longer than hydrocarbons of low molecular weight, total release
of hydrocarbons requiring from 2 to 60 days  (Neff, et al., 1976).
Concentrations of aromatics in tissues are assumed to reach an equilibrium
with external concentrations, and hydrocarbons of lighter molecular weight
tend to reach this equilibrium at a faster rate than heavier compounds.
Retention of petroleum hydrocarbons in animal tissues appears to be a

                                   306

-------
passive process, primarily through the partitioning  of  the hydrocarbons
between the water and tissue lipids  (Neely,  et  al.,  1974;  Neff,  et al.,
1976).  Active biological processes  such as  ventilation, blood circulation,
excretion, and hydrocarbon metabolism determine the  actual uptake and
release.

     According to Stone  (1975) and Neff, et  al.  (1976).  the incorporation
of hydrocarbons with tissue lipids is probably  by  hydrophobic interaction,
which means that the hydrocarbons remain in  an  exchangeable form.   Exposure
to petroleum hydrocarbons for a long period  of  time,  however, allows a
small fraction to enter a stable state in  the tissue from  which  it is not
readily exchangeable (Stegeman and Teal 1973; DiSalvo,  et  al., 1975; Neff,
et al., 1976).  When the organism is eventually re-exposed to hydrocarbon-
free water, hydrocarbon is released  from this stable fraction more slowly
than hydrocarbon is bound by hydrophobic reaction.

     Experiments on the uptake and fate of petroleum hydrocarbons (radio-
labelled  paraffinic and aromatic hydrocarbons)  by  the blue crab  Callinectes
sapidus indicated that up to 10% of  the hydrocarbons can be taken up from
either water or food organisms  (Lee, et al., 1976).   All the assimilated
hydrocarbons were metabolized or eliminated  through  fecal  matter.   More
than half of the metabolic activity  involving the  labelled hydrocarbons
occurred  in the hepatopancreas, which was  the only site of radioactivity
25 days after exposure.  In fish, however, the  liver and gall bladder
appear  to be the primary organs involved in  the metabolism of hydrocarbons
ingested  or assimilated  (Lee, et al., 1972).

      Both feeding mode and intrinsic capabilities  appear to determine the
extent  of hydrocarbon uptake by benthic invertebrates.   Roesijadi, et al.
 (1978), conducted experiments with deposit-feeding bivalves (Macoma
inquinata) and  sipunculids (Phascolosoma agassizi) and  suspension-feeding
bivalves  (Protothaca staminea) exposed to  aliphatic  and diaromatic
hydrocarbons.   In general the deposit feeders accumulated  the hydrocarbons
to a  greater extent than did the suspension  feeders.  Roesijadi, et al.
 (1978), also indicated that compounds of higher molecular  weight will be
relatively persistent in the tissue  of exposed  bivalves.

      Even though depuration or release of  assimilated petroleum is usually
rapid,  it is most often  incomplete.  Small quantities of hydrocarbons are
often retained  long enough to be transferred to a  higher trophic level.
Lee  (1975) found that detectable residues  of labelled benzo(a)pyrene were
evident in Calanus helgolandicus after 9 days,  in  £.  plumchrus after
16 days,  and in C^. hyperboreus after 4 weeks.   Molluscs appear to be unable
to metabolize hydrocarbons  (Lee, et  al., 1972;  Carlson  1972) but able to
eliminate accumulated hydrocarbons by depuration (Lee,  et  al., 1972;
 Stegeman  and Teal 1973;  Neff and Anderson  1975).

 F-l-c.    Detection and Avoidance

      Many references suggest that fish have  the ability to avoid petroleum
hydrocarbon-saturated waters  (Boesch, et al.,  1974;  North, et al., 1965),
 especially offshore waters.  The ability of  many fish to detect petroleum
hydrocarbons at low levels has been  illustrated in several cases but there

                                    307

-------
are a number of contradicting reports  (Patten  1977).   Syazuki  (1964)  showed
that goby (Chaenogobius heptacanthus), crescent perch  (Therapon  jarbua),
and striped mullet (Mugil cephalus) exhibited  threshold responses  at
between 0.7 ppm for crude oil and 48 ppm for "Mobil" oil.  Larvae  of
Atlantic cod (Gadus morhua), Atlantic herring  (Clupea  harengus harengus).
and plaice (Pleuronectes platessa), however, showed no ability to  avoid
water contaminated by three different types of crude oil  (Kuhnhold 1970).
Rice (1973) showed that the ability of juvenile pink salmon  (Oncorhynchus
gorbuscha) to avoid water-soluble fractions of Prudhoe Bay crude  oil in
a flowthrough system varied according to the fishes' adaptation  to sea
water, their age, and possibly the water temperature;  in  sea water the
avoidance level ranged from 1.6 ppm to 16.0 ppm.

                    F-2.   Indirect Sublethal  Effects

F-2-a.    Effects on Reproduction, Growth, and Metabolism

     Incorporation of sublethal levels of hydrocarbons into tissues has
also been shown to alter reproductive success  of crustacean populations
by decreasing egg production, increasing mortality rates  for egg and
larval stages, and altering basic metabolic processes  (Johnson 1977).

     Donahue, et al. (1977), examined the physiological response of mud
crab (Rhithropanopeus harrisi) larvae continuously exposed to naphthalene
or phenanthrene under different regimes of temperature and salinity and
found decreased survival to metamorphosis, increased duration of larval
development, increased respiratory rates, and  increased sensitivity to
acute salinity stress.  They concluded that sublethal  hydrocarbon  stress
acted to divert assimilated energy from growth to maintenance processes.

     Water-soluble fractions of No. 2 fuel oil were lethal to the  amphipod
Elasmopus pectenicrus at 4 ppm for short exposure periods, but growth and
fecundity were inhibited at lower concentrations of 0.6 and 0.2  ppm,
respectively (Lee, et al., 1977).  When adult  isopods  (Sphaeroma quadri-
dentatum) were exposed to low levels  of oil for one month, the juvenile
stages suffered significantly higher  mortalities, more than 70%  in five
weeks (Lee and Nicol 1977).  In oil concentrations of  0.3 to 0.4 ppm, the
amphipod Gammarus oceanicus produced  significantly fewer  larvae  than  the
controls in experiments conducted by  Linden  (1976).  Ustach  (1977) found
that the soluble fraction from 200 ul crude oil per liter sea water
significantly reduced egg production  by ^he harpacticoid  copepod Nitocra
affinis, as well as one-half and one-fourth dilutions; the mean  length of
life and the mean number of broods were not significantly affected.

     Decreasing fecundity and rate of  egg laying have  also been  shown to
result in birds which had consumed petroleum  (Hartung  1963;  Grau,  et  al.,
1977); this may be a function of both direct deposition  in the ovary  by
transport through the intestinal tract and liver and by  changes  in the  ion
composition of the yolk as a result of osmoregulatory  inhibition (Crocker,
et al., 1974, 1975; Grau, et al., 1977).

     Oritsland  (1975) found that fresh Norman  Wells  crude oil  did  not
significantly change the insulating values of  ringed  seal fur  in air  and

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at varying wind speeds.  It was suggested, however,  that  the  solar heating
of the seal's skin during haulout would be increased.

     Enzymatic and histological evidence of kidney damage,  possibly due  to
an unsuccessful attempt to concentrate or excrete the  oil or  its  metabolites
through the urinary system, was evidenced in  Smith and Geraci's  (1975) oil
immersion studies with ringed seals, Phoca hispida.  They suggested that
25 ml to 75 ml ingested crude  oil would probably represent the upper
limit of what an animal might ingest without  irreversible damage.   Thus,
ingestion of oil by immersed seals may not be significant except  through
the consumption of live contaminated prey.  However, an accumulation of
over 75 ml of petroleum hydrocarbons by this  route of  entry would require
significant levels of the toxicant in the prey.

     Geraci and Smith  (1977) suggested that more important  consequences
of an oil pollution incident may be eye damage, prolonged moulting, and
the influence of oil on stress of the pinnipeds, rather than  accumulation
of petroleum hydrocarbons in tissues.  Starvation would probably  result.
Pollution incidents would tend to be more detrimental  to  older  seals and
seals in poor nutritional condition.

F-2-b.    Tainting of Tissue by Accumulated Petroleum  Hydrocarbons

     Long-term  incorporation of hydrocarbons  by predators also  has the
potential to limit their utilization as food  by man.   Even  though most
marine  organisms are capable of eventually depurating  or  metabolizing
assimilated hydrocarbons, those of commercial value  which may smell and
 taste  in a manner reminiscent of petroleum, will be  unmarketable  for a
 long  time   (Blumer, et al., 1970; Bourcat and Mallet 1965;  Cahnmann and
 Kuratsune 1957; Mackie, et al., 1972; Nitta,  et al., 1965;  Shipton, et al.,
 1970;  Vale, et  al., 1970).

 F-3.     Uptake  and Effects of Petroleum Hydrocarbons Transferred  Via Food
                              Web Linkages

 F-3-a.      Food Web Biomagnification

      While  there has been much discussion of  the potential  for biomagnif-
 ication of  petroleum hydrocarbons with transfer  through the food  web, no
 definitive  studies have illustrated such a phenomenon.  In  fact,  although
 most   marine organisms are capable of accumulating high concentrations
 (200-300 times  in crustaceans and fish) directly  from  sea water,  there is
 no evidence that petroleum burdens in tissues actually increase at higher
 trophic levels  (Varanasi and Malins 1977).

 F-3-b.      Transmittal via Food Web Linkages

      Trophic  transfer,  i.e., from prey to predator,  of petroleum is well
 documented,  however, and there is good evidence  of  sublethal effects
 resulting  from the  incorporation or metabolism of  the  hydrocarbons by the
 consumer.
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     Corner, et al. (1976), found that at least for zooplankton, uptake
of hydrocarbons via food particles was a more significant and longer-lasting
route than via direct uptake from the water solution.  Not only was  the
total accumulation of hydrocarbons (naphthalene) higher from the food web
transfer but depuration was considerably slower such that a third of the
radiolabelled hydrocarbons remained in calanoid copepods ten days after
exposure to contaminated food particles.  Conover  (1971) found that
copepods (Temora longicornis, Calanus finmarchicus) ingested suspended
bunker C oil particles from the ARROW spill in Chedabucto Bay and accounted
for up to 10% of the oil in the water column.  Ingestion of the oil had no
apparent effect on the copepods, as they passed it with their feces which
were shown to contain up to 7% oil.  This mechanism was considered to be
one of the most important pathways of the pollutant between the pelagic and
the benthic systems—as much as 20% of the total suspended particulate oil
was estimated to precipitate to the bottom as zooplankton feces.  Parker,
et al. (1971), calculated that the calanoid copepod Calanus finmarchicus
could graze up to 1.5 x W~^ g of oil per day.  Thus, assuming a density
of 2,000 per m3 over a 1-km2 area to 10 m depth, as much as three tons of
oil could be encapsulated and precipitated as feces per day!

     Such high rates of uptake illustrate the rapidity with which
suspended petroleum can be biologically converted into a form available
for ingestion by higher trophic level organisms, through ingestion of
either the contaminated copepods by pelagic carnivores or of their feces
by benthic detritivores.

     The predators of the hydrocarbon-burdened zooplankton, unlike the
zooplankton, will often incorporate the hydrocarbon into their tissues
rather than excrete it.  Post yolk-sac larvae of Pacific herring (Clupea
harengus pallasi) that were fed rotifers (Brachionus plicatilis) contain-
ing high accumulated levels of labelled benzene rapidly incorporated the
monoaromatic hydrocarbon into their tissues (Struhsaker 1977).  Assimila-
tion of the hydrocarbons or metabolism products usually takes place in
high-lipid tissues, especially the reproductive organs.  In this case,
effects upon the organism's reproductive efficiency result.  Struhsaker's
(1977) exposure of female Pacific herring to the aromatic hydrocarbon
benzene caused a pronounced reduction in survival of the ovarian eggs and
resultant embryos and larvae through yolk absorption; the later life
history stages suffered the highest mortality, 43%.  Again, the total
effect of incorporation of the hydrocarbons into the food web would not be
manifested until the next generation of the carnivore populations, which
in some cases might not occur for two to three years.

     Ogata, et al. (1977), documented the accumulation of paraffins,
organic sulfur compounds, and aromatic hydrocarbons by eels reared in a
crude oil suspension.  Paraffinic hydrocarbons were detected in salmon
(Oncorhynchus keta), mullet (Mugil cephalus), and black sea bream (Mylio
macrocephalus) collected from waters polluted by a crude oil spill outside
Niigata Harbor in the Sea of Japan (Motohiro and Inoue 1973).  Indications
of higher n-paraffin concentrations in the digestive organs suggested that
the hydrocarbon contamination in this case was acquired principally  through
feeding.  Brown trout (Salmo trutta) collected 11  days after a diesel fuel
spill were found to still contain n-paraffin  (C-IO~CIQ) hydrocarbon compounds

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(Mackie, et al., 1972).  These results showed that many  of  the aliphatic
saturated hydrocarbons and possibly some of the aromatics characteristic
of the spilled diesel oil were also present in the flesh of the trout.

           F-4.   Ecological Effects of Petroleum Hydrocarbons

F-4-a.    Vulnerability of Lower Trophic Levels

     Studies of the toxicity of petroleum hydrocarbon components to marine
biota indicate that in general the lower the trophic level,  the more
sensitive the organism.  For example, marine crustaceans are more vulnerable
to petroleum hydrocarbons than are marine fish and marine mammals.  This
suggests that although the organisms at the upper end of the food web,
which are most often utilized by man, have greater tolerance of the effects
of petroleum hydrocarbons in the marine environment, the productivity and
stability of lower trophic levels can be altered to the  point  where basic
food limitations can be effected upon the higher consumers.  This process
is potentially one of the longest lasting and most deleterious effects of
petroleum pollution, and it has to be the most difficult to  detect,
considering the present knowledge of food web dynamics and natural
variability in food resources and the consequential effects  on the
consumers.

F-4-b.   Disruption of Autotrophic Production

     It is not unreasonable to assume that some effects  are  going to be
manifested at the very base of the trophic pyramid, the  photosynthetic
generation of carbon by phytoplankton.  The presence of  oil, undergoing
the processes of weathering and degrading on the surface film,  can
radically alter the utilization of organic and inorganic nutrients
(nitrogen, phosphorus) by the primary producers (Williams 1967;  Feldman
1973).  No. 2 and No. 3 fuel oils and Venezuelan crude oil generally
inhibited phytoplankton growth at concentrations greater than  30-50 pg/1
in radiocarbon-uptake experiments by Gordon and Prouse (1973).   Stimulation
of photosynthesis as reported by Mironov and Lanskaya (1969) and Strand,
et al.  (1971), was apparent only for low concentrations  of the crude oil
during spring.  (Gordon and Prouse indicated that the highest  oil
concentration they had ever measured in sea water, 25 cm below a 2-day
crude oil slick, was 800 ug/1.)

F-4-c.    Effects on Herbivores

     The existence of water-soluble petroleum hydrocarbons in  the water
column can also affect the ability of suspension-feeding herbivores to
utilize phytoplankton.  One effect is through behavioral inhibition of
the herbivores' feeding process.  A decrease in copepod  filtering rate
was surmised from a decrease in fecal pellet production  by copepods
exposed to an oil droplet concentration of 10 pom for 20 hours (Spooner
and Corkett 1974).

     Reduction of food resources of zooplankton has the  effect of reducing
overall production by influencing both the occurrence of adults and their
sex ratio  (Conover 1965; Omori 1970).  Even subtle alterations in the

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community structure of the primary producers available  as  food  for  the
zooplankton have the potential to inhibit growth and reproduction.   This
is a prominent process in natural systems which has been evolutionarily
accommodated by the various life history strategies exhibited by  the
zooplankton.  The question arises, however, of the effects of changes in
the available phytoplankton beyond magnitudes and time  frames to  which
the zooplankton have adapted.

     Parsons, et al. (1967), provided evidence that although zooplankton
may consume a variety of foods, only a few food items may provide a
satisfactory diet for growth and reproduction.  Specifically, they found
that the diatoms Chaetoceros debilis and C\ socialis were poor  food  sources
for three zooplankton organisms in the Strait of Georgia.  Consequently.
the growth rate of late stage Calanus plumchrus populations (an important
prey of juvenile salmon) under the Fraser River plume, declined when the
only food supply available consisted of small flagellates in late April
(Parsons, et al., 1969).

F-4-d.    Disruption of Heterotrophic Production

     In the nearshore ecosystem, where the decomposition of detritus by
marine microflora is an important process in the annual production of
food resources available for epibenthic zooplankton and benthic meiofauna,
petroleum hydrocarbons could influence the rates or timing of this
conditioning process.  There is little evidence to indicate that bacteria
would be negatively affected by the introduction of petroleum.   Hodson,
et al. (1977), reported that although bacteria were the first organisms
in the CEPEX enclosures to be affected by introduced oil, their rapid
generation time, the diversity of strains,  and their ability to mutate
allowed a rapid recovery of heterotrophic activity.   There is some question,
however,  of the possible effects of hydrocarbons on the behavior of the
bacteria.  Walsh and Mitchell (1973) have indicated that the chemotaxic
behavior of the bacteria—i.e., the ability to detect and move to food
sources—was significantly inhibited by various petroleum products and
components.  Such an effect, though seemingly critical, may be completely
compensated for by the increased growth of sulfide-generating bacteria in
response to the petroleum hydrocarbons as suitable organic matter (Colwell
and Walker 1977; Karrick 1977).  Westlake,  et al. (1J78), have illustrated
that such microbial populations exist to some degree at 22 sites in north
Puget Sound and along the Strait of Juan de Fuca and would logically be
capable of expanding and utilizing petroleum spilled in the region.  It
remains to be verified, however, whether these oil-degrading bacteria
would naturally utilize these detrital sources or would be acceptable to
bacteria-stripping zooplankton and meiofauna.

F-4-e.    Alteration of Food Resources of Consumer Organisms

     The effect of food limitation upon consumers such as planktivorous
fish may be even more pronounced than for zooplankton as the potential for
adaptation and capability for rapid population responses tend to be more
limited.   For instance, the abundance and size composition of zooplankton
has been suggested as critical determinants of fish survival during  the
transition from larval to juvenile stages (Thayer, et al., 1974).  The

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early life histories of many economically and ecologically  important  fish
species of the region (Pacific salmon, herring, sand lance,  greenling,
and smelt) include larval and juvenile stages which reside  in  the  region's
surface waters for weeks or months.  At this time, their growth  and
ultimately their survival are in part dependent upon their  achieving
an adequate ration of prey organisms.

     Blaxter (1965) provides evidence from Soviet investigations that
reasonable survival of fish larvae requires concentrations  of  prey
around 20,000 per m3 and that lower survival rates would be suspected at
prey concentrations less than 5,500 per m3-  LeBrasseur, et al.  (1969),
calculated that the waters of the Fraser River plume in the southern
Strait of Georgia possessed approximately 10,000 copepods  (assuming all
Microcalanus sp.) per m3 in early March when considerable numbers  of
larval fish are in the area.  Plankton sampling in the Strait  of Juan de
Fuca (Chester, et al., 1977) suggests significantly lower abundance in
that region, but they sampled no stations in the nearshore  environs where
zooplankton populations may be larger.

     Just as important as total abundance, the size composition  of the
zooplankton also determines the extent of exploitation by piscivorous
fish.  LeBrasseur, et al. (1969), illustrated that larval fish occupying
the  region of the Fraser River plume in the Strait of Georgia  grazed
mainly on zooplankton in the 500-y size range while juvenile fish  utilized
700-y  to  2,000-y plankters.  This is not a static pattern but  changes with
the  changes in morphology and bioenergetic demands and feeding capabilities
which  occur as the fish grow.  For instance, Pacific sand lance  (Ammodytes
hexapterus) less than 20 mm in length feed specifically on  zooplankton
 (nauplii  and copepod eggs) less than 500 y, while Pacific sand lance
greater than 40 mm prey on zooplankton in the 500-y to 1,000-p range
 (LeBrasseur, et al., 1969).

     These findings imply that since only a small portion of the available
prey resource is usually exploited by predators at a particular  time, any
deleterious influence on this fraction, despite the maintenance  of the
unexploited fraction, may actually limit the production of  the predator
population.  The principal example in the case of the neritic  food webs
of north  Puget Sound and the Strait of Juan de Fuca is the  surface layer
zooplankton community which is made up primarily of small calanoid
copepods  (Pseudocalanus sp. and Microcalanus sp.).  It is this community
which  is  heavily utilized by larval and juvenile fish during late  winter
and  spring and which would appear to be the most susceptible to  water-borne
petroleum hydrocarbons.  Deepwater zooplankton, composed primarily of
large  calanoids, Calanus sp., would be less affected, and through  diel
vertical  migrations such zooplankton would still be available  to the  fish.
This implies, however, the necessity of the fish to switch  from  one prey
resource, which appears to provide an optimal opportunity to gain  an
adequate  daily ration, to one which may not be adequate.  Although prey
switching may be feasible (Murdoch, et al. , 1975; Oaten and Murdoch 1975;
Reed 1969) because of the availability of sufficient numbers of  alternative
prey and  plastic feeding behavior of the predator, the alternative prey
resources may not be optimal from the standpoint of their size,  avoidance
capabilities, or nutritional value.  The inability to secure an  adequate

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ration usually results in a sacrifice in growth, reproduction,  or  ultimately
survival.

     In the example just cited, the small calanoid copepods Microcalanus
sp.  and Pseudocalanus sp. appear to be unsuitable as a sustained food
source for juvenile salmon (pink and chum) because of their size and
behavior,  whereas the larger copepods such as Calanus sp. can provide
adequate rations at lower concentrations (LeBrasseur, et al., 1969).
Koeller and Parsons' (1977) studies of juvenile chum salmon
(Oncorhynchus keta) also illustrated that the young salmon had  greater
difficulty obtaining their required ration from small copepods  (0.07-
0.12 mg—such as Pseudocalanus minutus, Paracalanus parvus, Corycaeus
anglicus)  than from large copepods (0.4-2.0 mg—such as Calanus plumchrus,
Centropages abdominalis), even though the smaller prey were ten times
more numerous than the larger.  Apparently, energy acquired from feeding
on the more numerous smaller copepods was utilized for the extensive
feeding activity and basic metabolism, whereas energy obtained  from eating
a few large copepods allowed energy in excess of basic metabolism and
feeding activity to be diverted into growth.  Thus, alteration of the
available  prey community by a pollutant introduction may have a more
critical effect than the direct effect upon the predator, especially in
the long run.

     More  important than basic somatic growth, reproduction in fishes is
highly dependent on food supply.  Although there is some variability in
the species' strategies under the stress of inadequate ration, most
investigators have found reproduction parameters—numbers of spawnings,
percentage of population reproducing, total egg production, percentage of
ovarian oocytes with yolk, post-spawning survival—are directly correlated
with ration level (Bagenal 1969; Scott 1962; Tyler and Dunn 1976; Wootton
1977).  Thus, if a pollutant alters the food supply of a significant
portion of the reproductive population in one year, the production of
that year  class will be affected for several years thereafter.

F-4-f.    Effects of Removal or Reduction of Keystone Species

     While disruption of the food web structure by impacts at the lower
trophic levels is likely, just as likely is disruption of the community
structure  by selective mortality or severe sublethal effects on keystone
species at higher trophic levels.  Seastars (Pisaster sp, Leptasterias sp.),
sea urchins (Strongylocentrotus sp.), predatory gastropods (Thais sp.),
and large  infaunal bivalves (Tapes sp., Mya sp.) typically have very low
recruitment ratess and so removal or dramatic reduction of these species
would probably result in a sustained modification of the nearshore
community  structure for years or decades.  Such community disruption has
the potential to dramatically affect production of macrophytic algae, a
major component of the detritus-based food webs.
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                                    317

-------
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                                   321

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                               APPENDIX G
BIOENERGETIC MODEL OF THE EPIBENTHIC  FOOD WEB IN THE MUD/EELGRASS SHALLOW
                   SUBLITTORAL HABITAT  AT WESTCOTT BAY


     A simplified bioenergetic model  was constructed for  one  nearshore
site in northern Puget Sound  for which  we had the most reliable  quantita-
tive data—the relatively enclosed, mud/eelgrass embayment  habitat at
Westcott Bay on the northwestern coast  of San Juan Island.  The  epi-
benthic food web was emphasized because we had community  and  trophic data
on the epibenthic and demersal fish and invertebrate communities (Miller,
et al., 1977; Nyblade 1978) and fairly  meager,  unquantitative data on
food webs of infauna communities.  We recognize that such an  approach has
little applicability to the Westcott  Bay system as a whole, and  much less
to other mud/eelgrass, contained embayment habitats  in north  Puget Sound.

     Food web data have been  summarized in a  very simplified  diagram
(Appendix Fig. G-l) where population  abundance (mean + 1  s.d.),  biomass
(mean + 1 s.d.), and frequency of occurrence  over time have been estimated
for the nodes of the web.  Transfer rates between nodes include  the number
of prey consumed (mean + 1 s.d.), the biomass consumed (mean  + 1 s.d.),
and the frequency of consumption.  In addition,  growth rates  were
estimated for secondary consumers in  order to calculate node  production.

Primary Production

     Many herbivores are present in tropical  marine  waters but not in
temperate-water habitats such as Westcott Bay.   In a Nova Scotian  kelp
community, Mann (1973) found  that herbivores  consume less than 10% of the
primary production and 90% entered detritus food chains.  Thayer,  et al.
(1975), found detritus deposition rates in eelgrass  beds  to be 12-22 mm
per year and this was either  utilized as food in the system or exported.
Harrison and Mann (1975) showed that  eelgrass detritus provides  a  better
food source than dead intact  leaves because there is a rise in the
percentage of nitrogen during bacterial decomposition.  The assimilation
efficiency of detritus by macrofauna  in eelgrass beds has been estimated
at 46%-48% (Adams and Angelovic 1970).   As has been shown in  this  report,
it is  evident that detritus input from  eelgrass and macroalgae forms the
base of the nearshore food webs in coastal marine ecosystems. The
relative importance of pelagic primary  production has not been accurately
assessed in nearshore ecosystems, however.

     Standing stock and production of macroalgae and rooted vegetation
have been estimated in many areas  (Appendix Table G-l).  Four categories
of primary energy sources were identified in  Westcott Bay:  Plankton,
rooted macrophytes (eelgrass), epiphytic and  epibenthic algae, and
macroalgae.  McRoy (1977) estimates that epiphyte dry weight  equals that
of leaf dry weight in eelgrass beds.  Based on literature estimates,
standing crop and production were estimated for the primary  energy sources
in Westcott Bay (Appendix Table G-2).  The production of  epiphytic algae,
a principal energy source for grazers,  was estimated with less  confidence.
                                    322

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Staghorn sculpin
     .2 1
       t gio.O
      33
                                         Phytoplankton
Starry flounder
  4.715.2.
 2>7.0t 49.
English sole
                                     Polychaete
Snake prickleback
   40.2. t- es.(<,
  142.S tZ3t>.
 Shiner  perch
 Penpoint  gunnel
 Surf smelt
Appendix  Fig.  G-l.
     Westcott  Bay.
Bioenergetic food web model of epibenthic  community of
Fish population and consumption data were  obtained in
     a  900-m2  sampling area.   Node and linkage values are, from top  to
     bottom, number  m"1,  grams m"1, and % frequency of occurrence.
                                    323

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Appendix Table G-l.  Standing stock and primary production in coastal
     detritus-based system.  Numbers have been converted to g/m2 wet
     weight.
System                            Standing stock      Production

Eelgrass, Alaska                                „           „
(McRoy 1970)                      585-13,590 g/m    243 g/m /day

Eelgrass, Denmark
August                            3,987  g/m         9,900 g/m /yr
(Sand-Jensen  1975)

Brown algae,  Nova  Scotia
(Mann 1973)                       2,001  g/m         13,000 g/m /yr

Eelgrass, North Carolina
Detritus                          46,000 g/m                 2
Primary production                                 3,400 g/m /yr
(Thayer, et al., 1975)

Eelgrass, Scotland
Plankton                                            2,085 g/m2/yr
Substrate-primary  production                       88-197 g/m2/yr
(Mclntyre and Eleftheriou 1968)
                                   324

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Appendix Table G-2.   Estimated primary energy sources (wet weights)
                     in Westcott Bay, San Juan Island.  Values are
                     annual means.
                            Standing crop          Annual production
                                   2                        2
Source                          g/m                      g/m
Phytoplankton                     14                      560

Rooted macrophytes             2,500                 1,740-9,900

Epiphytic and
 epibenthic algae              2,000                   20,000

Macroalgae                       200                    2,000
                                   325

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     Each of these four primary production compartments contributes energy
to grazers, detritus, and dissolved organic matter.  These flows were
estimated for Westcott Bay as contributions to annual production  (Appendix
Fig.  G-2).  In the absence of any quantitative data the detritus in
eelgrass beds was considered simply as steady state with no net erosion
or accumulation necessary to maintain the bed.  This provides a first-
order estimate of detritus available to fish predators through
detritivorous prey, but may grossly underestimate the total amount of
organic carbon (such as dissolved organic carbon) which is contributed
to the food web by eelgrass.

Primary Consumers

     The primary consumers form the important energy link between primary
production and higher trophic level carnivores.  Brook (1977) in a
Florida eelgrass bed found that polychaete and epibenthic crustacean
populations limit the fish predator population.  Most high trophic level
carnivores in eelgrass beds are at least partly benthic predators.
Mclntyre and Eleftheriou (1968) estimated the benthic meiofauna standing
crop in a flatfish nursery to be 1.25 g C m~2 dry weight and the annual
production was approximately twice the standing crop.

     The primary consumer level may be divided into two classes based on
feeding mode—suspension feeders and deposit feeders.  A loss of organic
matter to the system at this level may be caused by maintenance,
inefficient ingestion processes, production of pelagic gametes, and
conversion of organic matter into unusable elements (Levington 1972).
Selectivity in feeding by size and energy content of particles was shown
for a deposit-feeding bivalve (Hylleberg and Gallucci 1975).   Brinkhurst,
et al. (1972), found the same two criteria in particle selection for a
tube-building oligochaete and they estimated respiration and growth to
be 84.5 kcal/g annually.

     Feller (1977) estimated production and standing crop of benthic
harpacticoid copepods to be 1 g C m~2 yr"1 and 106 m~2, respectively.
The standing crop estimate is a summer maximum and it is much lower in
the winter.  Cohorts took six months to mature and the instantaneous
daily population increase was -0.3 to 2.2.

     Dagg (1975) estimated carbon and nitrogen budgets for an amphipod
which feeds on both diatoms and copepods.  The younger individuals are
primary consumers and they switch to a more carnivorous diet with age.
The assimilation efficiencies of both carbon and nitrogen ingestion were
90%.  Dagg found this higher than assimilation efficiencies for other
carnivorous aquatic invertebrates:  Polychaetes, 82-89%, euphausiids, 84%.
Ingestion rates increased when the amphipod switched from diatoms to
copepods.  The daily ingestion rates on diatoms were 4-60 g C and 0.7-12
g N and copepod ingestion rates were 120-1,000 g C and 40-190  g N.

     Population information for the primary consumers  in Westcott Bay
was derived from subtidal core samples and epibenthic pumping.
                                    326

-------
             Primary
            Producers
planktoji     t-
560 g/irr/yr  j\
         8c/
         ,0
                 UOM
                           \
     75";
   8%
              I
rooted        I
  macrophyles j
(6000)g/m?/yrj
                 DOM
           epiphytic and j
             epibenthic  .'
           macro
           2000

                 DOM
                  2',
                                       ft irnry
                                      f.onv.nw-rr

                                                (inv,rt-)

                                                    /\
                                                                     ro,,-.,
                                              7000 n/n.' ,/yr
                                          \
                                            ^  detritus
                                                                ..
                                           •1	
                                                             .•^^
                                                         .) '   V •
Appendix Fig. G-2.  Estimated flows  to  detritus, grazed and dissolved
     organic matter (DOM) compartments  from primary production
     compartments.
                                    327

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Secondary Consumers

     More detailed information is available for population at the
secondary consumer level than at any other.  Beach seine and stomach
analysis data from Westcott Bay were collected from July 1974 to
September 1975 (Miller, et al., 1976).  Seasonal growth rates and
conversion efficiencies are estimated for the dominant species, staghorn
sculpin and juvenile flatfish (Appendix Table G-3).  The growth rate
varies from 0.01 to 0.86 g day"1 and the maximum is in juvenile flatfish
in early summer decreasing as they grow during the summer.  Based on a
200-day growing season and growth rate of 0.08 g day"1, the annual
production of secondary consumers in Westcott Bay was estimated
(Appendix Table G-4).

Estimate of Daily Rations of Fishes

     Detailed stomach content analysis was available for most secondary
consumer species but the number of stomachfuls ingested per day was
unknown.  Some demersal and schooling neritic fish are believed to
forage at dawn and dusk (Hobson 1975; Cooney 1967).  If the stomach is
filled during each of these activity periods, then an ingestion rate
may be calculated based on two stomachfuls per day, although as we shall
see this may be an underestimate.  This assumption has been used for
estimating ingestion rates in this study.

     Three modes of feeding are exhibited by the secondary consumers in
Westcott Bay.  The first group are the pelagic feeders, composed of
Pacific herring and Pacific sand  lance.  This feeding is driven by
neritic production.  This  group is very abundant numerically and is
subject to heavy predation by birds  and larger  fish.  The second group,
to which perch belong,  are epifaunal  feeders.  This  feeding mode is
driven by eelgrass production with a  time  lag for settlement and growth
of epifauna.  Bottom  feeders  are  the  third  and  largest  category,
comprising starry  flounder,  English  sole,  gunnel, prickleback,  and  surf
smelt.  The  top  predator,  staghorn sculpin,  switches opportunistically
between benthic  feeding and  feeding  on pelagic  fish.

      Seasonal variability in growth  rate  and conversion efficiency  is
 quite high.   As  may be expected in a nursery area,  seasonal changes in
biomass  are  also high at this trophic level.  Monthly estimates of
 secondary consumer standing crop for Westcott Bay are shown in Appendix
 Table G-5.   From core samples and stomach analysis, there is also
 evidence for seasonal switching of prey by English sole which follows
 changing prey availability (Thornburgh 1978), although high variability
 may obscure the trends.  The same predator's diet changes significantly
 between the two sites.  With this very high seasonal variability in
 growth rate, conversion efficiency, standing crop, and diet, it is
 clear that initial food web models should be limited to season, or
 separately iterated over each season.

      Anomalies in the Westcott Bay food web are apparent,  even at  the
 highest trophic level where the  data were  assumed  to be  the most precise.
 To assess the accuracy of estimates of annual  production  of  secondary
 consumers, yearly food intake of English  sole was  compared with  its
                                     328

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Appendix Table G-3.
Seasonal growth rates and  conversion  efficiencies,
Westcott Bay.
 Staghorn  sculpin
 English  sole
 Starry  flounder
Apr-May
July-Aug
Aug-Sept
Sept-Oct

Mar-Apr
Apr-Jun
July-Sept
Sept-Oct

July-Aug
Aug-Sept
Sept-Oct
                                 Daily
                                weight
                               change (g)
0.02
0.13
0.90
0.05

0.01
0.07
0.07
0.13

0.03
0.05
0.20
                            Daily
                         intake (g)
                            Conversion
                            efficiency
                          (weight change)
                            intake (%)
0.36 ± 0.36
1.34 ± 2.56
3.42 ± 9.12
0.06 ± 0.04
0.24 ± 0.20
0.78 ± 3.42
                                              0.24  ±  0.40
 6
10
 3
17
29
 9
                    21
                                   329

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Appendix Table G-4.  Estimated annual  production (wet  weights)  of
                     secondary consumers  in Westcott Bay,  San Juan
                     Island.
Species
     Annual
               t-
production, g/m
Staghorn sculpin

Starry flounder

English sole

Snake prickleback

Shiner perch

Penpoint gunnel

Surf smelt

   Total
      2.10

      0.09

      2.10

      0.90

      1.60

      0.02

      1.60

      8.41
                                     330

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Appendix Table G-5.  Seasonal changes in standing crop  of  secondary
     consumers at Westcott Bay, given in grains per  square  meter.
July 1974
August
September
October
November
December
January 1975
February
March
April
May
June
July
August
September
4.09
1.11
4.16
3.80
1.71
0.91
0.24
0.13
0.04
0.13
0.20

0.87
0.94
6.60
                                   331

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production.  For an annual production of  2.1 g m  2  and  10%  growth
efficiency, English sole would have to consume 21 g m~2 yr"1.  Annual
consumption rates estimated from daily consumption  are  0.35-0.50
g m~2 yr"1.  This error factor of 40-50 shows that  more information  is
still necessary even at the level of higher consumers.  Possible sources
of error are:  (1) Daily intake rate of two stomachfuls is  too low for
rapidly growing juveniles.  (2) Ten percent growth  efficiency is too
high.  (3) Mean growth rate is too high—it may be  nearer the low end
of the range during most of the growing season.   (4)  Intake estimates
from partly digested stomach samples are  too low.   (5) Population biomass
estimates are in error.  These feeding and growth rates could be more
accurately measured by laboratory studies.

     If such great discrepancies exist between feeding and growth rates
at high trophic level, even greater errors may be expected for the
primary consumers where less quantitative data were available.  Metabolic
studies have been conducted for several species of  amphipods (Chang and
Parsons 1975; Halcrow and Boyd 1967).  Information  on particle selection
and assimilation efficiency is also known for species of various taxa.
However, this knowledge is quite patchy.  A concentrated effort on the
primary consumers of an area must be undertaken to  estimate standing
crop, feeding and growth rates, and production.

     An important subsystem of Westcott Bay was overlooked initially in
efforts to simplify the system.  This food web consists of the large
macroinvertebrates such as starfish, bivalves, crabs, and anemones
that are typical of most eelgrass areas.  These macroinvertebrates are
suspension feeders, macroalgae grazers, and carnivores, and therefore
represent unaccounted for nodes and linkages in the food web.  A
comprehensive bioenergetics analysis of any habitat as a whole will
have to incorporate every component system despite  its complexity.

     It is assumed that after identifying and quantifying the principal
energy pathways in a food web, the effects of perturbations can be better
assessed.  Three possible outcomes of a perturbation  are possible for a
population:   (1) Population numbers decrease,  (2) population numbers
remain constant but the mean weight per individual  decreases,  (3) the
population dies or leaves the area.  Knowing the  mobility of the population
and  the ability of individuals to switch  to different prey, as well  as  the
basic information on growth and  feeding rates  and production, would  be
necessary to predict the outcome.
                                    332

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                               REFERENCES

Adams, S.M.,  and J.W.  Angelovic.  1970.  Assimilation of detritus  and its
   associated bacteria by three species of estuarine animals.   Ches.  Sci.
   11(4)=249-254.

Brinkhurst,  R.O., K.E. Chua, and N.K. Kaushik.  1972.   Interspecific
   interactions and selective feeding by butificid oligochaetes.   Limnol.
   Oceanogr.  17:122-133.

Brook, I.M.   1977.  Trophic relationships in a seagrass community  (Thalassia
   testudinum), in Card Sound, Fla.  Fish diets in relation  to  macrobenthic
   and cryptic faunal abundance.  Trans. Am. Fish. Soc. 106:219-229.

Chang, B.D.,  and T.R. Parsons.  1975.  Metabolic studies on  the amphipod
   Anisogammarus pugettensis in relation to its trophic position in the
   food web of young salmonids.  J. Fish. Res. Board Can.  32:243-247.

Cooney, R.T.   1967.  Diel differences in trawl catches  of  some  demersal
   fishes.  M.S. Thesis, Univ. Washington, Seattle.

Dagg, M.H.  1975.  Complete carbon and nitrogen budgets for  the carnivorous
   amphipod,  Calliopus laevisculus.  Ph.D. Thesis, Univ. Washington,  Seattle,
   175 pp.

Feller, R.J.   1977.  Life history and production of meiobenthic harpacticoid
   copepods in Puget Sound.  Ph.D. Thesis, Univ. Washington,  Seattle,  249  pp.

Halcrow, K.,  and C.M. Boyd.  1967.  The oxygen consumption and  swimming
   activity of the amphipod Gammarus oceanicus at different  temperatures.
   Comp. Biochem. Physiol.  23:233-242.

Harrison, P.G.,  and K.H. Mann.  1975.  Detritus formation  from  eelgrass
   Zostera marina, the relative effects of fragmentation,  leaching, and
   decay.  Limnol. Oceanogr. 20:924-933.

Hobson, E.S.  1975.  Feeding patterns among tropical reef  fishes.
   Am. Sci. 63:382-392.

Levinton, J.   1972.  Stability and trophic structure in deposit-feeding
   and suspens'.or -feeding communit \m.  Air,. ?-Tat. 106:472.

Mann, K.H.  1973.  Seaweeds:  Their productivity and strategy for  growth.
   Sci. 182:975.

Mclntyre, A.D.,  and A. Eleftheriou.  1968.  The bottom  fauna of a
   flatfish nursery ground.  J. Mar. Biol. 48:113-142.

McRoy, C.P.  1970.  Standing stocks and other features  of  eelgrass popula-
   tions on the  coast of Alaska.  J. Fish. Res. Board Can. 27:1811-1921.
                                    333

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Miller, B.S., C.A. Simenstad, K.L. Fresh, F.L. Funk, W.A. Karp, S.F.
   Borton, and L.L. Moulton.  1977.  Puget Sound baseline program; near-
   shore fish survey.  Final Kept., July 1974 - June 1977.  Fish. Res.
   Inst., Coll. Fish., Univ. Washington, Seattle.  220 pp.

Sand-Jensen, K.  1975.  Biomass, net production and growth dynamics in an
   eelgrass population in Vellerupvig, Denmark.  Ophelia 14:185-201.

Thayer, G.W., S.M. Adams, and M.W. LaCroix.  1975.  Structural and
   functional aspects of a recently established Zostera marina community.
   Est. Res. 1:518-540.
                                    334

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4. TITLE AND SUBTITLE
 FOOD WEB RELATIONSHIPS OF NORTHERN PUGET SOUND AND THE
 STRAIT OF JUAN DE FUCA -- A Synthesis of the Available
 Knowledge                         	
                                   TECHNICAL REPORT DATA
                                  u Instructions on the reverse before completing)
1  REPORT NO
                                                           3. RECIPIENT'S ACCESSION NO.
                                          5. REPORT DATE
                                             September 1979
                                          6. PERFORMING ORGANIZATION CODE
7 AUTHOR.S)  Charles A> simenstad,  Bruce S.  Miller, Carl  F.
 Nyblade, Kathleen Thornburgh,  &  Lewis J.  Bledsoe
 Fisheries Research  Incrit.nt.p!  llniv  nf Washington	
                                          8. PERFORMING ORGANIZATION REPORT NO

                                              NOAA Report No.
 r I i>nci ica—rvcacoi in—iiia <- i L u L.IT ,—mi i v .—
9 PERFORMING ORGANIZATION NAME AND ADDRESS
 MESA Puget Sound Project
 National Oceanic and Atmospheric  Administration
 Tower Bldg.,Room 120
 7600 Sand Point Way NE, Seattle,  WA   98115
                                          10. PROGRAM ELEMENT NO.

                                             EHE 625-A
12. S°ONSORING AGENCY NAME AND ADDRESS
 Office of Energy, Minerals  and  Industry
 Office of Research and Development
 U.S. Environmental Protection Agency
 Washington, D.C.  20460
                                          11. CONTRACT/GRANT NO.
                                             IA6 No. D6-E693-EN
                                          13. TYPE OF REPORT AND PERIOD COVERED

                                           Final  Report  (3-77 thru  12/78)
                                          14. SPONSORING AGENCY CODE
                                                EPA-600/17
15 SUPPLEMENTARY NOTES
16 ABSTRACT                                                                  ~~~~~	

 Substantially increased  petroleum transfer and refining  activities are anticipated
 in the northern Puget Sound  and  Strait of Juan de Fuca areas.   These activities will
 likely increase the chances  of chronic and/or acute oil  inputs into the marine
 environment.  These areas  are currently stressed to only a  limited degree by
 petroleum.  The study reported here was undertaken to  identify biologic means by
 which petroleum constituents may be transferred from lower  to  higher trophic level
 populations and to identify  those populations and pre-predator links that are of
 critical importance to maintenance of major biological communities.  Interruption
 of these critical links  by loss  of important prey groups could drastically change
 the composition and/or productivity of higher trophic  level  populations.  The study
 was conducted by scientists  at the Fisheries Research  Institute, University of
 Washington, and involved primarily a compilation of existing data.
	 KEY WORDS AND DOCUMENT ANALYSIS

 DESCRIPTORS	I^IOENTIFIERS/OPEN ENDED TERMS
 Ecology
 Environments
                                                                         c. COSATl I icId/Group
 Release to Public
                            'is 'Si'-.;. PIT- CLASS 'Tins Repcrtl   j 21. NO. OF PAGES
                              Unclassified            '     334
                                             ' 20 -3 t C _ t= \ T v C LASS / This pugg;
                                               Unclassified
       2220-;
                                                       |22. PRICE

                             EDITION ,5 OfiSOLETt

                                          335

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