AD-A206 973 OREGON STATE UNIV CORVAI.IS SCHOOL Or OCEANOGRAPHY F/e 8/1STUDY OF THE INVERTEBAATES AND FISHES OF SALT MARSWS IN TWO -- ETC(U)JUN 61 0 L HIGL.EY, R L HOLTON DACW?2-77-C-0013UNCLASSIFIED CERC-1-81-5 NL*I*fllll*

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yMR 81-5

7) A Study of the Invertebrates and Fishes( of< Salt Marshes in Two Oregon Estuaries

by

Duane L. Higley and Robert L. Holton

MISCELLANEOUS REPORT NO. 81-5

JUNE 1981

,.,. %.DT!,"

NOV

Approved for public release;distribution unlimited.

Prepared for

* U.S. ARMY, CORPS OF ENGINEERS

COASTAL ENGINEERINGRESEARCH CENTER

Kingman Building

Fort Belvoir, Va. 22060

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IIN(IASS 11 11 11SECURITY CLASSIFICATION OF THIS PAGE (Whe, Date Entered)

REPORT DOCUMENTATION PAGE BERE DMSTIC ORMI REPORT NUMBER GOVT ACCESSION NO. 3. RECIPIENT'S CATALOG NUMBER

4. TTL trr7Jbttf.) 5 TYPE OF REPORT A PERIOD COVERED

A1 q [IVY 01 TIII N\VETl IWA'TLS ANDI- SlIl OF "1 ) i sce elaliCOlls el 4

2jOi "S~r.T MARSI1.S IN TWO ORF(;ON ESTUARIIS; O,, , j i @ . , . . PE,,O RINO ,,. REORT .%,ME *

7. AUTHOR(.) 8 CONTRACT OR GRANT NUMBER(a)

/0 Ilale I'.l]iglev /)

/0 Robert ./Holton ( IACW72-7-c-y/l

9. PERFORMING ORGANIZATION NAME AND ADDRESS 10. PROGRAM ELEMENT, PROJECT. TASK

,chool of Oceanography AREA & WORK UNIT NUMBERS

Oregon State University G31534Corvallis, Oregon 97330 ' /

11. CONTROLLING OFFICE NAME AND ADDRESS ,-

Department of the Army // ,unlCoastal lngineering Research Center 13 NUMBER OF PAGEt'

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4

18. SUPPLEMENTARY NOTES

'S

19, KEY WORDS (C roinue orn reverse side if necessary and Identify by block number)

I ish Netarts Bay, O)regon Si let: Bay, OregonInertebrates Salt ma rshes

20. ABSTRACT (Continue on reverse mide If neceesy and identity by block number) I

This study examines the invertebrate and fish life in the estutIa rine tidalmarshes of Siletz and Netarts Bays, Oregon. -Sweep nets, corers, enclosures, and

c 1 ip-quadrat samplers were used to collect both (ItIlnt i tat ire and non4tiant it at i resamples of invert ebrat es in level ma rsh , pan, t ida I creek, and t ida I flat ha) i -tats located in seven study areas representing variots tvpes of marsh. Fish inthese habitats, as well as in a slough and ii hay channels, were sampled bysei ne and otter trawls. Community taxonomic composition and trophic structure,

(Cont inued , (D AN 3 1473 EDITION OF I NOV 661I OBSOLE TE IINt:LASS 11 11 1) 5

SECURITY CLASSIFICATION OF THIS PAGE (When Dal Entered)

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UNCLASS I Fl IE)SECURITY CLASSIFICATION OF THIS PAGE(Whao Date Entered)

along with fish stomach contents, are presented as relative frequency histogramsand pie charts. Dominant invertebrate taxa in terrestrial collections wereAcarina, Ilomoptera, and I)iptera, and in aquatic collections were Capitellidae(polychaeta), Oligochaeta, Gnorirnosphaerora (Isopoda), and Anisocanrmarua andLOrophiuri (Amphipoda). Three-spine stickleback and young staghorn sculpin wereby far the most common fish species throughout the marsh zone; juvenile sal-monids and other species were captured only in submerged level marshes and in aslough. Trophic structure of terrestrial and aquatic invertebrate communitieswas generally heavily weighted to detritivores and scavengers. The herbivorecomponent increased from low marsh to high marsh and was the dominant trophict.e in the higher vegetation (sweep net collections) of the high marsh. Araneaewas the dominant invertebrate carnivore in the terrestrial communities. Fishconsumed primarily aquatic animals, even those captured in tidal creek and sub-merged level marsh habitats where tidal inundation would be expected to maketerrestrial foods available., The detritus food chain appears more importantthan the grazing food chain in the terrestrial communities, and transfer ofmarsh products to aquatic food chains apparently is predominantly through theexport of detritus rather than by the direct consumption of terrestrial animals.

2 INCLASS I: I IDSCCURITY CLASSIFICATION OF THIS PAGE('WPon Der Entered)

ABSTRACT

This study examines the invertebrate and fish life in the estuarine

tidal marshes of Siletz and Netarts Bays, Oregon. Sweep nets, corers,

enclosures, and clip-quadrat samplers were used to collect both quanti-

tative and nonquantitative samples of invertebrates in level marsh, pan,

tidal creek, and tidal flat habitats located in seven study areas repre-

senting various types of marsh. Fish in these habitats as well as in a

slough and in bay channels were sampled by seine and otter trawls.

Community taxonomic composition and trophic structure, and fish stomach

contents are presented as relative frequency histograms and pie charts.

Dominant invertebrate taxa in terrestrial collections were Acarina,

Homoptera, and Diptera, and in aquatic collections were Capitellidae

(polychaeta), Oligochaeta, Gnorimosphaeroma (Isopoda), and AnisogccIamaus

and Corophiwn (Amphipoda). Threespine stickleback and young staghorn

sculpin were by far the most common fish species throughout the marsh

zone; juvenile salmonids and other species were captured only over

submerged level marshes and in a slough. Trophic structure of terrestrial

and aquatic invertebrate communities was generally heavily weighted to

detrivores and scave:igers. The herbivore component increased from low

marsh to high marsh and was the dominant trophic type in the higher

portions of vegetation (sweep net collections) of the high marsh.

Araneae was the dominant invertebrate carnivore in the terrestrial

communities. Fish consumed primarily aquatic animals, even those captured

in tidal creek and submerged level marsh habitats where tidal inundation

would be expected to make terrestrial foods available. The detritus

food chain appears more important than the grazing food chain in the

-4-

terrestrial communities, and transfer of marsh products to aquatic food

chains apparently is predominantly through the export of detritus

rather than by direct consumption of terrestrial animals. This report

can be used to evaluate the impact of Corps of Engineers projects on

marshlands along the Oregon coast.

r

I -

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A )t ./t

v-.

PREFACE

This report provides baseline and food chain data on the invertebrate

and fish fauna of several marsh habitats located in Siletz and Netarts

Bays, Oregon. The study, sponsored by the U.S. Army Coastal Engineering

Research Center CCERC), evaluates the trophic value of Pacific Coast

salt marshes to provide information for assessing the impact of coastal

engineering projects on these resoureces. Results and conclusions

presented here are those of the authors and are not necessarily accepted

by CERC or the Corps of Engineers.

The following people employed by the School of Oceanography, Oregon

State University, contributed to the production of this report: Kim

Chalopka, Duane Higley, Robert Holton, Kim Jones, John Morgan, Jean

Shaffer, and Francis Stilwell. In addition, several student employees

supported in part by the College Work Study Program worked on the project.

Animal identification and determination of trophic type was aided

by Dr. Norman Anderson, Thomas Dudley, Dr. George Ferguson, Barry Frost,

Dr. John Lattin, Dr. Gerald Krantz, and Gary Peters of the Department of

Entomology, and Dr. Christopher Baynes of the Department of Zoology at

Oregon State University.

R. M. Yancy and A. K. Hurme were the CERC contract monitors for the

report, under the general supervision of E. J. Pullen, Chief, Coastal

Ecology Branch, Research Division.

Comments on this publication are invited.

-4-

CONTENTS

PAGE

I INTRODUCTION ........ ......................

II DESCRIPTION OF STUDY AREAS ..... ...............1. Pacific Northwest Salt Marshes .. ...........2. Siletz and Netarts Bays ............3. Bay Study Areas . . . . . . . . . . . . . . . .

III METHODS . . . . . . . . . . . . . . . . . . . . . . .1. General2. Invertebrate Studies ..... ...............3. Fish Studies ....... ...................

IV RESULTS . . . . . . . . . . . . . . . . . . . . . .1. General . . . . . . . . . . . . . . . . . . . .2. Taxonomic Structure of Invertebrate Communities.3. Trophic Structure of Invertebrate Communities.4. Composition of Fish Communities ........S. Fish Food Habits ....... ................

V DISCUSSION ......... ......................

LITERATURE CITED ....... ....................

APPENDIXA CRITIQUE OF METHODS .................B TAXONOMIC LIST OF INVERTEBRATES ...........C TAXONOMIC LIST OF FISH ...... .................D INVERTEBRATE SAMPLE DATA ...... ...............E FISH SAMPLE DATA ....... ....................F FISH FOOD HABITS DATA ................

TABLES

I Salinity and temperature readings ..............

2 Substrate characteristics of marsh soils .... ........... .

3 Description of sampling gear and methods .... ............

4 Occurrence of fish species in marsh and nonmarsh habitats .

5 Size of fish species in marsh and nonmarsh habitats .....

6 Invertebrates characteristic of terrestrial habitats .......

7 Invertebrates characteristic of aquatic habitats ...........

FIGURES

PAGE

1 Location of study areas in Netarts and Siletz Bays .....

2 Habitats of the salt marsh ecosystem ............ .

3 Taxonomic structure of invertebrate communities ...........

4 Trophic structure of invertebrate communities ...........

5 Fish stomach contents ........ .....................

A STUDY OF THE INVERTEBRATES AND FISHES OF SALT MARSHES

IN TWO OREGON ESTUARIES

by

Duane L. Higley and Robert L. Holton

I. INTRODUCTION

North American salt marsh ecosystems have been intensively studied

because of their high productivity and relatively simple structure.

However this attention has been mainly directed to the Atlantic coast

marshes. Prior studies have investigated community structure and popu-

lation energy flow (Odum and Smalley, 1959; Teal, 1962; Nixon and Oviatt,

1973), nutrient pathways using radionuclide tracers (Marples, 1966), and

faunal distribution (Barnes, 1953; Davis and Gray, 1966). Studies

centered in the Chesapeake Bay region, the Carolina coast; Sapelo Island,

Georgia; and Barataria Bay, Louisiana, have produced the following

information on salt marsh characteristics: a) Primary productivity is

high (about 445 to 2883 grams dry weight per square meter per year,

comparable to the most fertile natural and agricultural systems; b)

little of the marsh production is grazed (<10 percent), most ending up

in detritus food webs of the estuary; and c) the nutritional content of

detrital particles consumed is enhanced by adhering decomposer organisms

(summarized by de la Cruz, 1973). Because of the major importance of

detritus food chains in marsh and other estuarine habitats, recent work

has emphasized determining the rates and outputs of marsh detritus

(Reimold, et al.,'1975), and the structure of the dependent heterotrophic

food chains (Odum and Heald, 1975).

_____ 21

| I I

Floral composition and zonation of salt marshes on the Pacific

Coast have been documented (MacDonald, 1977). The major study of Oregon

salt marsh vegetation is by Jefferson (1974), who characterized and

mapped essentially all of Oregon's coastal marshes except those in the

Columbia River. Her descriptions of species composition, and community

structure, succession and distribution apply to Washington marshes

(MacDonald, 1977). Further description of marsh composition and zonation

is provided by Frenkel, Boss, and Schuller (1978). They studied the

transition zone between intertidal marshes and contiguous upland vegeta-

tion in Oregon and Washington.

Eilers (1979) conducted an intensive study of the salt marshes of

Nehalem Bay, Oregon. He determined plant associations and zonation

relations, and measured primary production and detrital output. Net

primary production vacied between 518 and 1,936 grams per meter square

per year. An excess of 90 percent of the intertidal net production was

transported into the estuary as detritus.

The Environmental Protection Agency (EPA) is presently studying

salt marsh plant productivity in Silet: and Netarts Bays, Oregon. The

EPA study is part of a larger program concerned with defining wetland

boundaries, the reactions of wetlands to perturbation, and the effects

of wetlands on water quality (H. Kibby, Corvallis Environmental Research

Laboratory, EPA, Corvallis, Oregon, personal communication, 1979).

Information on the structure and ecology of the animal communities

of Pacific Coast salt marshes is incomplete. MacDonald (1969) studied

I I I . I I | l I n n

the local, seasonal, and latitudinal variations in molluskan fauna in

level marsh and tidal creek habitats along the Pacific coast from Baja,

California, to Washington. He found Assiminea translucens, a small

prosobranch, to be ubiquitous in level marshes of this region, with

Littorina newcombiana (Prosobranchia) and Phytia myosotis (Pulmonata)

joining Assiminea to form a characteristic Oregonian assemblage. Tidal

creek mollusks were mostly bivalves, a Macoma-Mya assemblage character-

izing the Oregonian Province. The number of species recorded from each

habitat increased from north to south. Level marsh mollusks fed predom-

inantly on algae or plant detritus by rasping; tidal creek forms included

deposit and suspension feeders as well as predators and scavengers.

Whitlatch (1974) observed the growth, production, and seasonal

abundance patterns of Batillaria zonalis, a small introduced prosobranch,

in pans, tidal creeks, mudflats, and SaZicornia level marshes of Tomales

Bay, California. Abundance was greatest in pans and creeks, but recruit-

ment was lacking in the creeks which apparently resulted in the relative

stability of the populations there. Influx was likely due to immigration

from the pans where recruitment was successful.

Two studies have been made of insect populations of San Francisco

Bay marshes. Using a sweep net for collecting, Lane (1909) identified

124 species in Spartina-SaZicornzia marshes. The majority of species

were in the orders Diptera (flies) and [tomoptera (planthoppers); Del-

phacidae (Fomoptera), and Chloropidae, Ephydridae, and Chironomidae (all

Diptera) were the dominant families. Cameron (1972) used a clip-quadrat

method in a similar marsh to study insect trophic diversity and its

relation to resource availability (living and dead plant materials). He

found that herbivore diversity increased with primary production, and

that saprovore diversity increased during periods of detrital input. In

general, trophic deversity showed seasonal patterns relating to physical

factors and (more clearly) to resource availability. Cameron hypothesized

that seasonal increases in diversity occurred as seasonal species joined

persistent species in exploiting expanding resources.

The only major study of trophic relations in a Pacific coast salt

marsh ecosystem is the Coos Bay, Oregon, study sponsored by the National

Science Foundation (Hoffnagle, et al., 1976). Short-term field and

laboratory studies were used to measure net primary production, detrital

production, decomposition rate, nutrition of key species, and the compo-

sition of insect and fish populations of several marsh sites.

In recent years, interest has increased in the role of estuarine

food chains in the growth and survival of seaward migrating juvenile

salmonids along the Pacific Northwest coast. There is evidence that

those juveniles which benefit from favorable estuarine conditions have a

better chance at sea (e.g., Riemers, 1971). These fish seem to adjust

their habitat and feeding strategies to exploit freshwater and marine as

well as estuarine food chains while making the transition to marine life

(Mason, 1974). The fish are found in some marsh habitats, especially

tidal creeks. Dunford (1975) found juvenile chum salmon (Oncorh:inchus

keta) and chinook salmon (0. tshawyjtscha) residing in sloughs and creeks

of the Fraser River estuary marshlands (British Columbia) in the spring

and summer. The salmon consumed a variety of terrestrial, planktonic,

and benthic foods. Dunford identified 13 other fish species in these

habitats.

Juvenile salmonids in nonmarsh habitats may exploit marsh-based

food chains. In the Squamish River estuary (British Columbia), Cliff

and Stockner (1973) discovered heavy feeding by salmon on amphipods

(principally Anisoganarus spp.) which are largely marsh-dependent.

Juvenile chum salmon in the Nanaimo estuary (British Columbia) heavily

exploit harpacticoid copepods and thus use a food chain that depends on

detritus from the marshlands (Healey, 1979).

Although past studies of Pacific coast salt marshes-have been

limited, the data collected suggest similarities of structure and function

between these marshes and the Atlantic coast marshes; e.g., levels of

primary production, contribution to detritus-based food chains, and some

aspects of community composition. Important questions remain regarding

the use of Pacific coast marsh habitats and food chains by various fish

species, especially juveniles; and the trophic structure and function of

these marshes should be determined, especially to evaluate the value of

marshland in relation to human use.

This study characterizes the animal communities and food chains of

marshes in Siletz and Netarts Bays, Oregon. The objectives were to

develop taxonomic lists, to characterize the trophic structure of marsh

invertebrate communities, and to identify the principal fish species

using the marsh and marsh-related habitats. In addition, food habits of

these fish were studied to determine marsh food-chain relations.

II. DESCRIPTION OF STUDY AREAS

1. General.

Salt marshes of the Pacific Northwest are of recent origin and, in

comparison to the Atlantic marshes, are limited in size and distribution.

The steep and rocky coastlines of Washington, Oregon, and California

restrict suitable marsh habitats to a few bays, estuaries, and lagoons.

These marshes generally lack the thick peat layers which reflect long

term accretion (MacDonald, 1969).

In Oregon, interglacial deposits filled river mouths, and post-

Pleistocene drowning produced extensive tidelands in the northern and

central bays. More rapid sediment deposition in the southern bays

matched rises in sea level and thus restricted tideland development.

All of the 27 estuaries in Oregon are presently accumulating sediment.

Fires in the mid-19th century and the Tillamook fire in 1933, augmented

by logging and other detrimental land-use practices, have increased the

erosional sources of bay deposits (Jefferson, 1974).

The climate of the Oregon coast is wet-temperate. Annual precipi-

tation averages about 180 centimeters and temperature about 10 degrees

Celsius. The frost-free season lasts 250 to 300 days, and freezing

weather is infrequent. Pacific winter storms accompanied by gale-force

winds are common, but generally lack the destructive force of tropical

and convective storms common to the Atlantic coast. Winter freshets in

coastal rivers and the diluting effects of the Columbia River discharge

may substantially reduce estuarine salinities. In light of this, Kistritz

(1978) suggests that the term "salt marsh" may often be inappropriate in

describing tidal marshes of the Pacific Northwest.

Mixed diurnal tidal fluctuations result in abrupt changes of immer-

sion and exposure times at about 2.7 meters or mean higher high water

(MHHW), where mean lower low water (MLLW) is the zero datum. Below MHHW

a distinctive salt marsh vegetation characterized by pickleweed (Salicornia

virginica), commonly known as "low marsh," extends down to about mean

lower high water (MLHW). Above MHHW, a "high marsh," characterized by

tufted hair grass (Deschampsia caespitosa), grades into terrestrial

vegetation at about extreme high water (EHW). Jefferson (1974) lists

six vegetation types for Oregon saline-brackish intertidal marshes: (a)

low sand marsh, (b) low silt marsh, (c) sedge marsh, (d) immature high

marsh, (e) mature high marsh, and (f) bulrush and sedge marsh. One to

seven vegetative communities may occur within each vegetation type.

These communities and marshes form complex and somewhat variable relations

with each other and with tidal level which Jefferson treats as succes-

sional. Three successional patterns occur, depending on substrate (sand

versus silt) and freshwater influence. Lyngbeye's sedge (Carex Zyngbyli)

is intermediate in all three patterns, widely distributed, and considered

by Jefferson to typify Oregon salt marshes.

Low marshes typically advance through coalescing colonies of seaside

arrowgrass (Triglochin ma=itima) or rhizomous mats of pickleweed. The

lower edges of the marsh are also commonly lined with three-square

bulrush (Scirpus americanus). Transitions from low marsh to high marsh

.1

may be gradual or abrupt across an eroded bank. Tidal-flat to high

marsh eroded banks may be 1 meter high. Extensive diking, landfills,

and other man-induced effects have significantly changed the marshlands.

Jefferson (1974) states that undiked old, high marsh is nearly nonexistent

in Oregon.

S2. Siletz and Netarts Bays.

Siltez Bay, a spit-protected estuary of about 4.8 square kilometers,

is located on the central Oregon coast (Fig. 1). The bay receives

runoff from the Siletz River and two creeks. The average witner and

summer Siletz River discharge is 45 cubic meters per second and 6 cubic

meters per second, respectively. Logging has caused extensive sedimen-

tation, and diking, roadbuilding, and filling projects have restricted

flushing causing tidelands to increase; therefore, the marshes are

expanding. Salinity varies widely according to discharge and tidal

stage. During winter freshets, the salinity of surface waters is often

less than 5 parts per thousand where the Siletz River enters the bay;

summer surface salinities exceed 20 parts per thousand (Rauw, 1975).

Temperatures generally vary from 7 to 15 degrees Celsius (Rauw, 1975),

but may exceed 18 degrees Celsius in some habitats (Table 1).

Netarts Bay, a shallow, bar-built estuary of about 10.4 square

kilometers, is located on the north-central Oregon coast (Fig. 1). The

bay has a very small watershed, which drains through 13 small creeks,

and is therefore usually completely mixed and marine dominated. Salini-

ties usually exceed 25 parts per thousand. Bay temperatures generally

WASI41NCtCN

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AREAAR4

AREAI BA

CALIFOR-411

AREA 7

TiOALCREEK

AREA 5

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NETARTS BAY -

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STUDY AREAS ' '--.Lwsn as ' R~

Low silt marsh SLT ~:-~I

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3 Sedge marsh N

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4 Immature high marsh

5 Mature high marsh , A

6 Netarts open bay otter trawl p 0 1sites (indicated by A A

7 Netarts low sand marshrL;R'seine site AREA 3

'iIl S

8 Set low sand marsh ,'seine site II

-' SLOUGH9 Suet: open bay otter trawl

sites (indicated byA ) 5.1I

SILETZ BAY

Figure 1. Location of study areas in Netarts and Suet: Bays.

_ I I I-iI

Table 1. Salinity and temperature readings.

Netarts BayArea Habitat Date Salinity Temperature

1 Level marsh 18 Jan. 78 -- 9.0

1 Level marsh 7 Feb. 78 26 9.8

4 Tidal flat 7 April 78 29 17.0

4 Large pan 7 April 78 12 --

I Level marsh 7 April 78 29

6 Bay channel 3 June 78 31 --

I Level marsh 22 July 78 36 27.0

1 Level marsh 17 Oct. 78 33 20.0

5 Tidal creek 17 Oct. 78 33 20.0

5 Marsh channel! 1 Nov. 78 18-30 7.0-11.8

5 Pan 1 Nov. 78 13 --

I Tidal flat 29 Aug. 78 33 21.0

5 Tidal creek 12 April 79 15 11.0

5 Pan 12 April 79 19 11.0

7 Tidal flat 12 April 79 28 11.0

Siletz BayArea Habitat Date 1 Salinity Temperature

(0/o.) (0C)3 Level marsh 18 Jan. 78 -- 9.5

3 Level marsh 6 Feb. 78 28 10.5

3 Level marsh 6 April 78 9 12.5

2 Level marsh 6 April 78 9 12.5

3 Millport sl. 24 June 78 21 --

3 Tidal creek 21 July 78 26 28.0

2 Level marsh 21 July 78 30 25.0

3 Tidal creek 21 July 78 26 23.0

9 Tidal flat 18 Sept. 78 18-20 18.0

3 Level marsh 16 Oct. 78 25 15.5

2 Level marsh 16 Oct. 78 23 16.0

3 Tidal creek 26 April 79 18 17.0

8 Level marsh 26 April 79 27 14.0

3 Pan 26 April 79 15 18.0

reflect ocean temperatures (about 8-15 degrees Celsius); however, tem-

peratures greater than 26 degrees Celsius may occur in the summer over

tidal flats and marshlands (Table 1). Logging on the watershed from

1951 to 1971 caused extensive siltation in the bay, but sediment input

now is apparently low and stable (Kreag, 1979).

High and low marshes fringe the inner shore of the spit, and a

large area of high marsh occupies the southern end of the bay. This

marsh was once diked and used for pasture, but has since reverted to

nearly natural drainage patterns under state ownership.

3. Bay Study Areas.

Nine study areas were established in the two estuaries (Fig. 1).

Areas 1 to 5 were chosen to represent the specific vegetation types

listed by Jefferson (1974), and were sampled most thoroughly. The other

areas are open bay and low marsh habitats each sampled once each for

fish. Elevation data for areas 1, 3, and 4 are based on nearby EPA

itudy sites (H. Kibby, personal communication, 1979).

The study areas were:

a. Area 1, Low Sand Marsh (Netarts Spit). This beach is sandy

(Table 2) and supports a mixed cover of pickleweed and saltgrass (Dis-

tich.is spicata). The lower edge of the marsh is lined with three-

square bulrush. Invertebrate samples were taken in the pickleweed-

saltgrass zone (about 2.4 meter above MLLW), fish samples in the three-

square bulrush zone and the adjacent tidal flat (-2.1 meters above

II

1.

Table 2. Substrate characteristics of marsh soil at level marsh samplingsites. 1

River Netarts Siletz Siletz Netarts NetartsLow Sand Low Silt Sedge Immature High Mature High

Marsh (area 1) (area 2) (area 3) (area 4) (area 5)

Debris 3.3% 10.1% 15.6% 66.1% 23.0%

Sediment 96.7% 89.9% 84.4% 33.9% 77.0%

Sand 92.5% 12.8% 1.1% 67.8% 87.0%

Mud 7.5% 87.2% 98.9% 32.2% 13.0%

SedimentSize Class (mm)

>1.00 0.0% 0.2% 0.2% 0.0% 0.0%

0.500-1.00 0.0% 0.3% 0.3% 0.1% 0.0%

0.250-0.500 71.8% 3.2% 0.3% 2.7% 5.6%

0.125-0.250 19.5% 2.3% 0.3% S7.6% 80.6%

0.063-0.125 1.2% 7.0% 0.1% 7.4% 0.8%

0.063 7.5% 87.2% 98.9% 32.2% 13.0%

Sample cores were processed in the following manner: (a) The whole sample

was wet-sieved on a 2-millimeter screen (> 2 mm = debris, < 2 mm = sediment);(b) the sediment fraction was wet-sieved on a 0.063-millimeter screen (> 0.063mm = sand, < 0.063 mm = mud); (c) the sand fraction was dry-sieved on 1.0-,0.5-, 0.25-, and 0.125-millimeter screens; and (d) all fractions were dry-weighed. The debris fractions included roots, shells, and similar materials.

MLLW). A debris line of dead eelgrass (Zostera marina) frequently forms

at varying levels along this marsh.

5,

b. Area 2, Low Silt Marsh (North of Siletz River). This is an

area along Highway 101 of prograding low marsh. The substrate in the

marsh and the adjoining tidal flat is mud (Table 2). The lower edge of

the marsh is formed in interrupted colonies of seaside arrowgrass invaded

by Lyngbeye's sedge, which is the dominant species at higher elevations.

Aquatic invertebrate samples were taken in this transition zone which is

characterized by frequent flooding, pools of standing water among the

plants, and dense populations of amphipods and isopods. Terrestrial

invertebrate samples were collected higher in the sedge stand. Fish

samples were collected about 100 meters south of these sites in a series

of small tidal creeks that extend from high marsh through the sedge

community and through the bulrush community at the edge of the marsh.

c. Area 3, Sedge Marsh (South of the Siletz River). This marsh

has muddy soil (Table 2) with vegetation dominated by sedge, but floods

less frequently than the low silt sedge marsh. Elevation in the region

of level marsh invertebrate sampling site is about 2.3 meters above

MLLW. A dendritic system of small tidal creeks laces the marsh and

apparently receives some seepage through earthen dikes. A major creek

(maximum 10 meters wide, 0.7 meter deep) dissects the marsh in an east-

west direction. Water in the creek flows in both directions from about

the center of the marsh where the channel is but a shallow depression in

the level marsh. Fish and aquatic invertebrate samples were taken in

various creek, pan, and tidal flat habitats, as well as in Millport

Slough which borders the marsh on the southwest. All of these habitats

have muddy substrates.

d. Area 4, Immature High Marsh (Netarts Spit). This marsh, located

slightly north of the low sand marsh, has an elevation of about 3.2

meters above MLLW and is bordered by an eroded bank. The dominant

vegetation is tufted hairgrass and Pacific silverweed (Potentila pacif-

ica). The soil is peaty with an underlayer of fine sand (Table 2). A

large pan (40 by 10 meters) retains tidal and runoff water during the

winter and spring but dries up by mid-summer.

e. Area 5, High Marsh (South End of Netarts Bay). A branch of

Jackson Creek, which flows directly into the ocean, flows through this

40-hectare marsh. The marsh is dissected by numerous deep tidal creeks

with several openings into the bay. These creeks and the northern edge

of the marsh have steep eroded banks. The marsh soil is peaty with a

sand underlayer. Creek bottom and adjoining tidal flats from brown

sandy mud to black mud. Marsh vegetation is primarily tufted hairgrass

but the composition varies; some areas are dominated by Pacific silver-

weed, pickleweed, rush, and other plants. The creeks are often clogged

with rotting eelgrass. Several pans are scattered throughout the marsh.

Those connected with creeks retain water, while others tend to dry out

in mid-summer.

f. Area 6, Netarts Open Bay. This designates the bay channel and

tidal flat regions in which otter trawls were used to obtain estuarine

fish samples. The channels are mostly shallow, many of them having

eeigrass beds.

g. Area 7, Low Sand Marsh Seine Site (Netarts Bay). This 1-

kilometer section of low sand marsh, located immediately south of area

1, is a narrow strip (about 3 to 20 meters wide) that is mostly vegetated

by pickleweed. Plant cover is variable, and the shoreline is irregular

due to erosion.

h. Area 8, Low Sand Marsh Seine Site (Siletz Bay). This 0.4

kilometer strip of low marsh, located on the southeast edge of the

Siletz spit, has high marsh along eroded banks.

i. Area 9, Siltez Open Bay. This designates tidal flats and

channels which were sampled for estuarine fish using an otter trawl.

Selection of the study areas was partly based on EPA use of Areas

1, 3, and 4 for their productivity studies. The intent was to establish

site specific data on the animal communities of marshes where the EPA

studies were being conducted. The EPA work focused on determining

primary productivity and decomposition rates for selected, nearly mono-

specific vegetation types (pure stands) and determining the availability

of marsh production to detritus-based food chains. The results of this

work are presently being compiled (H. Kibby, personal communication,

1979). Initial conclusions are that primary productivity rates range

from about 500 to 1,800 grams per square meter per year, with Lyngbeye's

sedge having the highest productivity. Biomass of this sedge peaks in

June-July at about 1,200 grams per square meter per year. Seaside

arrowgrass apparently decomposes more rapidly than other ipecies studied,

and is the only species which showed evidence of grazing (probahly by

deer).

The marshlands provide a variety of habitats and subhabitats whose

properties change daily with tidal and seasonal conditions. Animal

populations respond with zonations and marked flucutations which reflect

life cycles, tidal exchange, and migrations to escape inundation. In

this study, it was impossible to fully characterize these fluctuating

populations over the variety of marshes and habitats studied. The

approach was to sample the major habitat types in the marsh ecosystem

(Fig. 2), and to collect comparative samples from other estuarine habitats

such as tidal flats and bay channels. Extensive sampling was conducted

in level marshes, the most widely distributed, and tidal creeks, the

most likely contributors to aquatic food chains of the marsh habitats.

III. METHODS

1. General.

The basic objective of this research was to characterize the inver-

tebrate and fish life of the Siletz Bay and Netarts Bay marshes. Sampling,

which varied with weather and tidal conditions, was conducted at approx-

imately 2-month intervals. Greatest sampling effort was made in the

spring and summer. Most collections were either one-time surveys or

repeated as opportunities arose. The only habitat for which seasonal

data was collected is the submerged level marsh (invertebrate fauna). On

some occasions, two work crews were used to exploit a brief sampling

time frame (e.g., a single high tide). Table 3 lists the various sampling

devices and their uses. Appendix A provides suggestions for gear improve-

ment.

High LowLeve ' Tidal Level

Fiur 2 Hia tso the sal m arshstm(apedfo

Ranwenk 1972)

Table 3. Description of sampling gear and methods.

Device Description Use

Small corer 5.1-cm diameter tube Quantitative infauna sampling;with handles also sediment sampling

Medium corer 10.2-cm diameter tube Quantitative infauna samplingwith handles

Large corer 15.2-cm diameter tube Quantitative infauna samplingwith handles

Small enclosure 27-cm diameter by 30-cm high Quantitative sampling of in-plastic cyliner vertebrates of strand line

Large enclosure l-m-dia;neter by 1-m high Quantitative sampling of in-canvas cylinder with lead- vertebrates in submerged levelline and floats marsh

Aquatic sweep net 0.S-mm mesh nitex Quantitative (with largeenclosure) and nonquantitativesampling of submerged in-vertebrates

Terrestrial sweep net fine mesh muslin Semiquantitative sampling ofinvertebrates on exposedvegetation

Small drift net 0.5-mm mesh nitex net on Nonquantitative sampling of12.S-cm-diameter frame drift organisms in small tidal

creeks

Large drift net 0.5-mm mesh nitex net on Nonquantitative sampling of2S-by 50-cm frame drift organisms in large tidal

creeks

Clip quadrat 2S-by 2S-cm wooden frame Quantitative sampling of in-within which plant material vertebrates on exposed levelwas clipped loose from the marshsoil

3-m seine common-sense seine with Fish collection in small tidal0.6-cm mesh creeks and pans

15-m seine 1.3-cm mesh body and Fish collection in large tidal0.6-cm mesh bag creeks and over low (level)

marshes

52-m seine 2.5-cm mesh body and Fish collection over low1.3-cm mesh bag marshes and adjacent tidal

flats and sloughes

Otter trawl S-m trawl with 3.2-cm Fish collection in baymesh body and O.b-cm mesh channels and mudflats

cod end

2. Invertebrate Studies.

Aquatic invertebrate samples from level marsh, pan, tidal creek,

and adjacent tidal flat habitats were routinely processed and preserved

in the field using a 5 to 10 percent buffered seawater formalin solution.

Occasionally, it was necessary to process samples in the laboratory

after storage in an ice chest for a day. Such treatment had no observable

effect on the stored animals. Except for terrestrial and certain core

samples, all samples were sieved on 0.5 millimeter screens or were

obtained with 0.5 millimeter mesh nets.

After several days storage in formalin solution, the samples were

transferred to a 70 percent isopropanol and stained with rose bengal or

a similar stain to enhance visibility of the animals during sorting.

Samples were sorted under a 3-diopter illuminated lens to broad taxonomic

groups, and later identified. Usually, crustaceans, polychaetes, and

bivalves were identified to genus or species, insects to family, and

other groups to higher taxa (order, class, etc.). When appropriate,

life stage (e.g., adult, larva, pupa) was recorded. Invertebrate clas-

sification follows Barnes (1974) and Borror, DeLong, and Triplehorn

(1976).

The aquatic samples varied widely in quantity of debris and number

of animals collected. To facilitate processing, the samples were separ-

ated by stacked sieves into two size groups (O.S to 2 millimeters, and

>2 millimeters) or split quantitatively with a Folsom plankton splitter.

This process was especially useful for samples collected with the large

enclosure in the fall when detached vegetation was present.

v4V

a. Level Marsh. The principal method for collecting submerged

invertebrates on the level marshes was the large enclosure (Table 3).

It was dropped over a preselected sample point and secured at the soil

by standing on the leadline, which closely conformed to the soil contours.

The 0.5 millimeter mesh aquatic sweep net was then repeatedly swept

within the enclosure until capture rates were very low or zero. The

animals and debris were concentrated and preserved. This method provides

a semi-quantitative measure of the aquatic and terrestrial animals found

near or on submerged vegetation, although in a few cases it was difficult

to remove all of the highly abundant isopods found in the low silt marsh

(Siletz Bay) study site.

Large enclosure studies were designed primarily for the low marshes

although a single sample set was collected on the immature high marsh

during an extremely high winter tide. Samples from the low marshes were

collected on three to four occasions.

Large enclosure sample sites were established where a stand of

selected type of vegetation occurred in a reasonably accessible location.

Each site was a 10-by 10-meter grid divided into 100 sampling areas. On

each sampling day, four randomly preselected areas were sampled. Each

area was sampled only once during the study.

A similar sampling plan was established to study the infauna of

level marshes. A plug of soil and roots 10.2 centimeters in diameter

and up to 25 centimeters deep was removed at selected sampling areas in

a grid (separate from but near to the large enclosure grid). The plug

I - I i I I U I~ I I I -i

was disaggregated by hand under water and then sieved on a 0.5 millimeter

screen. Early results showed that the majority of the animals were near

the surface, so later samples wre only 5 to 10 centimeters deep. It was

also decided that the few animals collected and the relative unlikelihood

of their directly entering aquatic food chains did not warrant the time

and effort required for extensive sampling. Therefore, only one set of

four samples per marsh was collected and completely processed.

Sampling of terrestrial invertebrates of the level marsh was con-

ducted during low tides with the terrestrial sweep nets, clip-quadrat

method, and small enclosure (Table 3). One set of samples was taken at

each marsh. Collections were planned during the warmest and driest

period of the years, but an unusually wet season forced the postponement

of several collecting trips. The collections were finally accomplished

during favorable tides and weather on 29 August 1978 (low sand and

immature high marshes of Netarts Bay), 7 September 1978 (low silt and

sedge marshes of Siletz Bay), and 25 September 1978 (mature high marsh

of Netarts Bay). On these dates, air temperature was 19 to 240 Celsius,

wind 0 to 16 kilometers per hour, and the sky sunny to overcast.

All samples were taken at low tide. The wind was minimal, the air

temperatures were moderate, and the marsh vegetation was slightly damp.

Within each level marsh type, sample sites met the following criteria:

(I) selected vegetation community, (2) uniform vegetational cover, (3)

level ground, (4) easy accessibility, and (5) no evidence of recent

disturbance. A 10-by 10-meter grid at each site was measured and marked

off by corner stakes.

The terrestrial sweep net sampling method (Table 3) was adapted

from Davis and Gray (1966). The net was vigorously swept back and forth

across the upper parts of the vegetation through an horizontal arc of

about 1 meter. Following each sweep, one step was taken and the direction

of the net was reversed. Four samples, each consisting of 20 strokes

(10 in each direction), were obtained, one along each edge of the per-

imeter of the grid.r

After each sample, the contents of the net were placed in a large

ethyl acetate-charged killing jar and later transferred to a wide-mouth

specimen jar. The samples were cooled in an ice chest for processing in

the laboratory where they were then stored in a cold room until the damp

and sometimes succulent plant debris could be removed. The insects were

sorted and stored dry except for soft-bodied species which were preserved

in 70 percent isopropanol.

At each marsh grid, four randomly preselected points were sampled

by the clip-quadrat method (Table 3). The vegetation was first clipped

off 15 centimeters above the ground. The remaining vegetation was then

sliced off at the ground level with a sharp knife and placed in a heavy

plastic bag along with any plant litter that could be gathered at the

base of the plant. Roots were not collected. Insects seen crawling on

the ground inside the quadrat frame were also deposited in the bag. The

bags were inflated and securely fastened to avoid crushing the collected

plants and insects. The inflated bags were packed in an ice chest for

transport to the laboratory. In the laboratory, the plant material was

processed in a Berlese-Jullgren apparatus for 7 days. The insects were

preserved in small specimen jars filled with 70 percent isopropanol.

U-,-

b. Debris Line. Invertebrate life of a 40-by 1-meter (approximate)

debris line on the low sand marsh was sampled using the small enclosure

method (Table 3). Four randomly chosen areas in the line were sampled

by pushing the small enclosure through the debris (principally eelgrass)

and removing the enclosed plants and invertebrates. The samples were

processed in the same manner as the clip-quadrat samples.

All of the terrestrial samples were sorted in a flat container

under a binocular dissecting scope. Terrestrial sweep net samples,

which often contained considerable plant debris, were sorted in a white

enamel pan. Samples processed in the Berlese-Tullgren apparatus were

sorted in a petri dish. Larvae and all animals less than 0.5 millimeter

were not included in the data.

c. Pan. Several samples were taken in pans in immature and mature

high marsh using the aquatic sweep net method (Table 3). Some laboratory

observz.tions of living animals were also made.

d. Tidal Creeks. Tidal creeks were sampled using small corer,

large corer, and aquatic sweep net methods along transects in the mature

high marsh in Netarts Bay (I November 1978) and sedge marsh in Siletz

Bay (24 June 1978). In each bay, the creeks were sampled at equal

intervals as measured along the curves of the creeks, using the small

corer (four samples per station), the large corer (one sample per sta-

tion), and the aquatic sweep one (one sample per station). The small

corer samples were 10 centimeters deep and captured small surface crus-

taceans and worms. Large corer samples penetrated 30 centimeters to

' . . . . .. ...

sample larger and deeper dwelling species such as bivalves. Small corer

samples were screened on a 0.5 millimeter sieve and the large corer

samples on a 2 millimeter sieve.

The mature high marsh transect was 480 meters long and included

five stations spaced at 120 meter intervals). Station 1 was located at

the cre2k mouth, the bottom of which is 28 meters in width and 0.8 meter

below the level marsh. Stations 1, 2, and 3 were located below a dike,

and stations 4 and 5 above the dike in a tributary creek. The creek at

station 5 was 1.1 meter deep and 0.7 meter wide. Aquatic sweep net

samples were taken only at stations 1, 2, and 4.

The sedge transect was 400 meters long with eight stations spaced

at 50-meter intervals. The creek bisects the sedge marsh, and drains in

opposite directions from a shallow center area (station 5). Maximum

creek width was 10 meters and maximum depth was 0.7 meter (station 8).

At station 5, the creek forms an 8-centimeter-wide depression in a

sparsely vegetated, dark muddy area. Because of time constraints,

stations 4 and 7 were not sampled. Two small tidal creeks in the sedge

marsh were sampled by aquatic sweep net on b April 1978. The cieeks are

about 0.5 meter wide and 0.5 meter deep and form part of the dendritic

system that flows into the major creek.

Drift nets (Table 3) were set in the lower regions of the creeks of

the sedge and mature high marshes to collect animals that represent

available fish food. Large drift net samples were collected in a small,

dendritic creek in the sedge marsh on 19 December 1977, and at the

- . . .. , ,... . i .. . . . i .. . . I i . . i.. .

oayward mouth of the large tidal creek on 16 October 1978 and 26 April

1979. A small creek was also sampled on 6 February 1978 using the small

drift net. Large drift net samples in the mature high marsh were col-

lected at a single location in the lower region of a major tidal creek

on 17 October 1978, 1 November 1978, and 12 April 1979. A small drift

net sample was obtained in a small tributary on 12 April 1979.

e. Tidal flats. Infaunal samples were collected by large and

medium corers (Table 3) over 30-by 60-meter grids located on tidal flats

adjoining the low sand (Netarts Bay) and sedge (Siletz Bay) marshes.

The grids were marked at 1 meter intervals producing 1,800 potential

sample areas. Ten of these were randomly selected for each set of

samples. At each area, a 10-centimeter deep medium corer sample and a

30-centimeter deep larger corer sample were collected. Medium corer

samples were screened on 0.5 millimeter sieve and the large corer samples

on a 2-millimeter sieve.

3. Fish Studies.

Fish were collected with sienes and an otter trawl from several

marsh habitats and in the open bay of each estuary. A comparison was

made of the species composition and food habits of the bay fauna and the

marsh fauna.

a. Collection. Major collections of bay species were made by

otter trawl on 2-3 June 1978 in Netarts Bay and on 18 September 1978 in

Siletz Bay. Fish were taken by seine in the sedge marsh (18 September

1978) and mature high marsh (I November 1978 and 12 April 1979), and in

the tidal creeks at sites which previously had been extensively sampled

for aquatic invertebrates. Also, flooded low marshes (areas 7 and 8)

were seined for juvenile salmonids and other species in April 1979. A

total of 20 additional seine samples was collected in pan, level marsh,

creek, mudflat, and slough habitats.

Only part of the catch was generally retained since the primary

objective was to document habitat use according to species and to provide

specimens for stomach content analysis. Thus, where large numbers of

the same species were caught in a single haul or in several hauls, a

subsample of each species representing the size spectrum captured was

retained. Fish were preserved in 10 percent buffered seawater formalin

in the field. The abdominal cavities of all but very small fish were

opened to allow penetration of the preservative. In the laboratory, the

fish were transferred to 70 percent isopropanol for storage. All speci-

mens were identified to species and measured for fork length.

b. Stomach content analysis. The stomach contents of 10 to 12

fish from a sample were analyzed. A total of 237 fish stomachs from 27

samples was analyzed. The fish selected approximated the species compo-

sition and size distribution of the preserved samples.

Stomach content analysis involved removing the stomach and estimating

stomach fullness, digestion state, bolus volume, and volumes and numbers

of the different food items. The analysis was made using a binocular

dissecting microscope and a grided petri dish. Digestion state of the

bolus was rated on a scale of 0 to 9, based on prey recognizability

(i.e., 0 = nothing recognizable; 9 = totally recognizable). The volume

(but not number of items) of unrecognizable materials was recorded as a

separate item. Prey items were identified according to taxonomic groups

used in the invertebrate studies.

RESULTS

1. General.

The structure of the invertebrate and fish communities is first

depicted ona taxonomic basis and then on a trophic basis. In both

cases, the data are presented in the form of relative abundance. For

the trophic interpretation, each invertebrate taxon was assigned to a

trophic type (herbivore, detritovore, carnivore, omnivore, scavenger,

non-feeder, and unknown). Fish trophic relations are based on the

stomach contents data.

Drift net data were omitted from this presentation due to sampling

difficultures (Appendix A) and because the aquatic sweep net collec-

tions in tidal creeks provide very similar information. Appendices B

and C are taxonomic checklists of invertebrates and fish, respectively.

Tabular summaries of the data are provided in Appendix D (invertebrate

collections), E (fish collections), and F (fish stomach contents).

2. Taxonomic Structure of Invertebrate Communities.

Soil infauna, sampled by medium corer, was dominated by oligochaetes

and several dipterous larvae (Fig. 3). Ceratopogonid and chironomid

larvae were especially abundant in the low marshes (sand, silt, and

sedge), while mycetophilid and dolichopodid larvae were most abundant

in the two high marshes, which had a more diverse dipterous fauna. Cer-

tain taxa numerous in the low marshes samples--Acarina, Isopoda, and

the amphipod genera, Anisogamrnarus and Orchestia--are epifaunal forms

which were trapped at the surface by the corer. Another amphipod genus,

Corophium, lives in tubes both in the substrate and on vegetation,

depending on species. The dominant species in the marshes was C.

salmonis, which is an infaunal animal common in muddy estuarine tidal

flats. Its high density in the low silt marsh reflects the fact that

the samples were collected near the edge of a prograding marsh where

it merges with a tidal flat.

The fauna of low vegetaiton (clip-quadrat samples) included high

densities of Acarina in all marshes (Fig. 3). Collembola were

abundant only in the high marshes, while Coleoptera and Homoptera

occurred in both low and high marshes. The isopod, Gnorimosphaeroma

lutea, was abundant only in the low silt marsh. The high marsh fauna

included four families of Collembola, two of Homoptera, and eight of

Coleoptera. Aphididae (Homoptera) and Limnebiidae (Coleoptera) in-

habited some of the low marshes.

The invertebrate fauna of the high vegetation sampled by terrestrial

sweep net was boradly similar for all five marshes in that Acarina,

Homoptera, Diptera, Aranae, and Hymenoptera were abundant in all

marshes (Fig. 3). Hemiptera of the low marshes were predominantly

saldids, and in the high marshes mirids and pentostomids although

these were not abundant. The composition of the Homoptera varied

amoung marshes, although Delphacidae was generally abundant. The dip-

terous fauna tended to be more diverse in the high marshes; the low

number of taxa in the low sand marsh likely relates to the poor vege-

tation cover afforded by pickleweed and salt grass.

The fauna of the low sand marsh debris line was composed chiefly

of Acarina, Collembola, Amphipoda (Orchesti a traskiana), and Aranae

(Fig. 3). This fauna differs in part from the fauna of the low vegeta-

tion and high vegetation habitats of the low sand level marsh, although

Acarina and limnebiid beetles were abundant in all three habitats.

Collembola (mostly isotomids) were abundant in the debris line, but

absent from both high and low vegetation. Debris line dipterans were

mostly sphaerocercids, as contrasted with chironomids and ceratopogonids

found in the low vegetation, and muscids in the high vegetation.

Faunal composition of the submerged level marsh (sampled by the

large enclosure method) was a mixture of aquatic and terrestrial forms

(Fig. 3). Dominant groups were Acarina in the low sand and sedge

marshes, isopods in the low silt marsh, and dipterous larvae in the

immature high marsh. Oligochaetes were moderately abundant in the low

silt and sedge marshes, where they were frequently found inside decay-

ing sedge leaves, a condition which made their quartificatin difficult.

Among coleopterans captured from the submerged vegetation of the low

marshes were limnebiids, staphylinids, and coccinellids. In the im-

mature high marsh, carabids and hydrophilids were collected. It is

interesting that the hydrophilids, which are aquatic, moved into this

high marsh during its rare submergence. These animals probably origin-

ated in nearby pan or eroded bank habitats. Diptera of the submerged

level marfh were primarily larvae psychodids, ceratopogonids, and chiro-

nomids, with some variation among marshes, Homoptera, although not

abundant in the submerged marshes, was represented by three families,

Cicadellidae, Delphacidae, and Aphididae, with the Delphacidae the

most abundant.

Aquatic crustaceans of the submerged level marshes were the amphipods

Corophium spp., Anisogmnarus confervicoZus, and Orchestra traskiana,

the isopod G. Zutea, and the two cumacean genera, Hemieucon and

CumelZa (Fig. 3). Of these, G. lutea and A. confe-vico~us were especially

abundant in the low silt marsh. Dense summer populations of G. Lutea

swarmed in the warm water of shallow depressions between vegetated areas.

On the low sand marsh, large numbers of talitrid amphipods migrated

upshore ahead of advancing tides, seeking shelter in dead eelgrass and

other debris. When this material floated within the large enclosure

sampling grid, amphipod and other animal densities measured very high.

Several pans in the high marshes sampled by aquatic sweep net were

inhabited by a variety of aquatic forms (Fig. 3). The immature high

pan had large numbers of copepods (mostly harpacticpods), the amphi-

pod, A. confervicoZus, and oligochaetes. The mature high pans also

contained amphipods and oligochaetes; corixids, and ephydrid and culicid

larvae were also abundant.

Infauna of tidal creeks in the sedge and mature high marshes were

similar (Fig. 3). Oligochaetes, polychaetes, and amphipods were the

most abundant forms in each creek. Capitellids and anpharetids

dominated the polychaete fauna in both creeks, although spirorbids and

spionids were also abundant in the mature high creek. Amphipods were

mostly Corophium and Anisoaamars conferviculus, but included some

talitrids and Ampithoe in the mature high creek. Macoma baithica, a

small tellinid bivalve, was common in the sedge creek but absent from

the mature high creek.

Animals collected in the tidal creeks by aquatic sweep net were a

mixture of aquatic and terrestrial animals also collected in large en-

closure samples and in creek infauna samples (Fig. 3). Presumably,

terrestrial animals in the creek fell into the water or washed in during

tidal submergence. Diptera of the two creeks were quite different,

being quite diverse in the sedge creek and limited to a few taxa in

the mature high creek. This may reflect the comparatively large amounts

of filamentous algae occurring in the sedge creek at the time of

sampling. The algae appeared to have high densities of dipterous lar-

vae and other taxa capatured by the aquatic sweep net and the corer.

The grapsid crab, Hemigrcapsus oregonensis, was also common in the algae,

although it was not quantitatively sampled.

The infauna of the sedge tidal flat was similar in many respects

to the infauna of the sedge creek infauna (Fig. 3). The tidal flat is

located near the bayward outlet of the creek, and both the creek and

the tidal flat have muddy substrates. The tidal flat infauna was

relatively poor in Diptera, however, having only low densities of

dolichopodid larvae. Other differences included a lower density of a

burrowing cnidarian, and the addition of a sacoglossan gastrod, Aldepia.

The infauna of the sandy tidal flat located below the low sand

marsh (Netarts Bay) differed from the infauna of the sedge tidal flat

in having a relatively greater abundance of polychaetes (principally

ha : Iosco opZcs) and an Eohaustroiius-Paraphoxus amphipod fauna, in con-

trast to the Corophium-dominated fauna of the sedge mudflat. The

decapod shrimp, Uallianassa, and the bivalve, Cryptomya californica,

an inhabitant of Ca"lianassa burrows, was also present in the sandy

tidal flat.

1W

3. Composition of Fish Communities.

Of 26 species of fish captured in seines and trawls, 2 species

(staghorn sculpin, Leptocottus armatus, and the threespine stickle- Iback, Gasterosteus acuZeatus) dominated the catches in both high and

low marshes (Table 4). The two species were common in creeks, pans,

and submerged vegetation at the marsh edge, as well as in non-marsh

habitats. However, staghorn sculpin were not captured in low marsh

pans. Threespine stickleback captured in marsh habitats were juveniles

to adults (12 to 76 millimeters), while staghorn sculpin were juveniles

and young adults (17 to 173 millimeters) (Table 5).

Other species in marsh habitats were juvenile - rfsmelt (Hypomesus

pretiosus) and juvenile chum salmon, captured primarily in low level

marshes (Tables 4 and 5). The young chum salmon were seined along

sparsely vegetated low marshes in both Netarts and Siletz Bays. In

Netarts Bay, these salmon are occasionally abundant in the spring be-

cause of natural reproduction and the release of hatchery-reared

juveniles. Those in Siletz Bay apparently result from a small natural

run.

The most abundant fish species in the slough adjoining the sedge

marsh were the shiner surfperch (Cymatogasta aggregata) and the three-

spine stickleback (Table 4). Nine other species were captured although

in much lower numbers. These species included staghorn sculpin, nor-

thern anchovie (EngrauLis mordax), starry flounder (Patichthyes

stellatus), and juvenile chinook salmon.

The largest variety of fish occurred in the bay channel, where

species found in marsh habitats along with several juvenile marine

species were collected (Tables 4 and 5). The most abundant marine

--

Table 4. Occurrence of fish species in several marsh and nonxnarshhabitats. 1

Hig Marsh I Lo. Marshe ___ OtherTidal Bay

FISH1 Pan Creek Level Pan Creek Slough fa hne

Number of Samples ____ S S 2 a 4 4 11Pacific sandiance tA.nmoayces haapter-sa)Topsmelt (Atharinaps affinia)Speckled sanddab (Cithsorichthysa tig'maeus) XXXXXXX)Staghorn sculpin (L..wtocottusa .-Umaus) xxxxxxx.XXXXXXXX xxxxxxxXXX)XXXX)X xxxx xxx xxxx xxx xxxxx.NBuffalo sculpin (Enochrya bison)Cabe-on (Scorpaenichya xtrnorarius)Prickly sculpin (Cottua aper) I / /Coastal sculpin (Cottusa zauticus) f /Shiner surfperch (Cynroogaster Ia1gregata) ////////XXXXXXX)XXXXXXXWhite surfperch (Pi'anarodon fukxc.~s I//Northern anchovie (Engrau~is ",ordaz) IPacific touctd (Microgadus proximus) /Tubesriout 'AuZorhynchsa I avidus)Threespine stickleback (Gaeterosceua acuZearus) XiXXXXi XX)X~Xa uXxxxxx xxixxx ~XX)XXXX rXXXXXXXI / / / //,Lingcod t'ohiodon elongacue) XXXKXXX)Kelp greenling (Hexagrannos decgrnmrus) XXXXXXXSurf smelt (NPomnerus j'retiosus) / IXXXXXSX / XXXXXX / / / /I/I/Saddleback gunnel (Pholis a narca)////Starr" flounder (P~archth-jo stca tu ) ///i//ixxxxxxx)// ',

English sole (Paropiryo vuaLua) ////xxxxxxx)Sand sole (Psettichthys '7ELanostictus) I / / /Chum salmon (Onorhynchus eta) / XXxxI / /Chinook salmon 'Oworhynocoue tah:j~ta ha) ffI /Steelhead trout (Sainwo gaiz'dnaiii)f/Rockfish spp. (Sebastes app.) I /Snake prick leback iDL.mpunus sagitta)f/Bay pipefish (Syngniathmua leptornycus)f/

'Results are based on seine samples (most habitats) and otter trawl samples(bay channel) collected on several dates in the two bays; XXX=abundant,///=present.

2Low marsh refers to low sand, low silt, and sedge marshes.

Table S. Size (fork length in mm) of fish species collected inseveral marsh and nonmarsh habitats.1

Pactfic sandlance t-yte. heptemaia re wa o ra

Top%.ait (Athertirope affinie) 39:2(32-44)Speckked svddab (Cit.2richthje atiy".aeu) 3 4 5StagltOr s ctlpin (Leptocottua a,.-ast) 56:29(44.76) 49:88(35.82) S8:115(18.67) 44:97(17-124)buffalo 2CaIpta (Eeophrya biwon)Caboton fMcorpoeiihtjys awso)Prickly sculptn (Cotta caper) 16:4(34-41)coastal sctait (Cocttte eaei"sO 37: 1(37)Shinter itterparc (Crenatogastor agggqto 7S:1(7S) 72:1(72)white t. orfptrch (Phanorodais (oreccaNorther" 6nchovte (Emmrolte i~ordawr)Pacific tomicod (Microgmsetwe prczxnsta)?Tlbesni)%it (AstIolhynchoa ftdeidwe)Threptn* stickleback (C~ater'oe,ue ac.tleosus) 41:146(31-62) 39: 216(22-Sa) 41:88(10-60) 22:46(12.33) 30:301(20-76)

r Loegcod (Optliodan flonyotue)sells greenling (Hera7.O..toe deceMetaJSurf -. It tUypon.t Prezioete) 42:1(42) 53:97(40-64)Saddlebiack gunel (PhltLa oi~naca)Starry flouander (Platichthys stelttte)Eglish, tale 1P.UMoplryd a n.s)Sand* sale (Pseethcs netanottueJClou. SCIson (Otterhj,s.a ketaJ 39!1(39) 44:S7(36-66)Ch-ock sain. (C'tt,hynclota taha,wteha~)Steelbead trouat (Salmo gairdneriiJRockfish spp. (Sebast.8 app.)Snake prickleback (Latpenuo aietta)N., p,reftsh 15,'.11 Owe Z.pco-tacloue) _______ ________ _______ _______

______________ OthterSPCIE I6 YT. d c I Da

Slg flat Ch.n1~Pacific sasndlan~e Immotdytes IseoaptaeeTaps.*It (416erias afftoiie)S pecklerd 0.0444k (C ithrtchthys stiqotesei S7:63(2I.1IS)Slatharn sculpin (cpcoettaa aro'.otaa) S7;59(28-173) 90:16(36-193) 7tt(7j3NSuffato sculpin (Enophrsh wn 82:6e4-'14Cabeto. 15 orpoeicAtlhye eaomta) 53:b(346-24)

Prcl Pcapo(Cta ar) 142;1(142) 3(4-6Coa1c11sta ocain 1Ccocttae: oleotwxue)Shtner surfporch (Ceacogaeter agteata) 112:4311(SO0.154) SS:77(1I-119) 64:1(61)whtle sulfperch IPkss""edo. ftaco.e) 76 1:76)hi nr" .r ano..e (Egoo'Li. mrdae) 1 3: 4 73.110)Pactftc Loscosi tMiaroyadae proxe@a) 79:1(79)

Shre~estale (Pet.ctabac oe(.oeeecs e) S4(Z-0 lO:J(60O) IOA:329-1)Chtngo (sodo .1nohoIa etc.u@ cJo 9:462lS 96:3('2-20S~cea trelngft (scrgn grdisearcs me 67:23(59-111locift-It app. su (Seheatee 693(3-7p.9 -2) S;:12)Sdo raackcne (hLa..e aogsa) 942(010) ,:11(2-4122)Starryftounde (S ,aelohe 1. te~te IS2lo.6)_133:.___S_.43 177:17(70.2S

E me sulso' 'amroe baed, on sie sml S6:(most24 habitats )an ot r tawCus apes (baorynch annladtda lt oletdo eerldtsi

the two bays. .

2 Low marsh refers to low sand, low silt, and sedge marshes.3 Mean.4 Sample size.5Range.

species in Netarts Bay was juvenile English sole (Parophrys vetulus),

which invade northwest estuaries in large numbers during the spring.

4. Trophic Structure of Invertebrate Communities.

The trophic structure of the major terrestrial and aquatic marsh

communities is presented in Figure 4. Data from large enclosure and

aquatic sweep net collections have been omitted because these col-

lections include both submerged terrestrial and aquatic species. An

analysis of the trophic structure of such assemblages would be mis-

leading, since they do not represent communites as such.

The major feature of Figure 4 is the predominance of detritovores

and scavengers in most of the communities. Oligochaetes, amphipods

(Ccrop)hi) and Acarina were the principal detritovores of the soil

communities, while Acarina were the most abundant detritovores in low

vegetation, high vegetation, and debris line communities. Herbivore

populations (mostly homopterans) were abundant in the high vegetation

especially in high marshes, where their densities exceeded those of

the detrivores. Scavengers were numerous in the soil marsh (cerato-

pogonid and chironomid larvae), the low vegetation of the low marsh

(isopods, amphipods, limnebiid beetles) and in the debris line (amphi-

pods, limnebiids).

Carnivores generally comprised a small fraction of the animal

life in soil and low vegetation habitats. However, dolichopodid

(Diptera) larvae were abundant in high marsh soils, and also occurred

in low marsh soils. The carnivore populations of low vegetation were

composed primarily of Araneae and staphylinid beetles. High vegetation

carnivores tended to be more numerous, and included several types of

dipterous adults (Dolichopordidae, Ceratopogonidae, and Muscidae) and

Araneae. The debris line carnivores were Araneae and Saldidae (Hemip-

tera) which occurred in moderate abundance.

The trophic structure of infaunal communities of the tidal creeks

and tidal flats was heavily weighted to the detrivore component (Fig.

4). In all creek and tidal flat communities. oligochaetes and capitellid

polychaetes were among the dominant detritovores. Other detritovores

were Haploscoloplos (Polychaeta) and Corophium (Amphipoda). Common

carnivores were the Polychaete Eteone and a small cnidarian polyp.

Although algae covered much of the sedge creek and tidal flat sub-

strate surface at the time of sampling, macrofaunal herbivores were

rare.

5. Fish Food Habits.

Fish stomach contents data are summarized in Figure 5, which

combines data for all sampling sites and dates for each habitat.

Staghorn sculpin, threespine stickleback, and juvenile chum salmon

captured over submerged level marshes consumed a variety of predominantly

aquatic animals, including amphipods (Corophium and AnisogCVnmarus),

harpacticoid copepods, cumaceans (Hemileucon), oligochaetes, and

polychaetes (Fig. 5). The diet is diverse partly because data from

several samples have been combined. Terrestrial prey were not eaten

except by the chum salmon, which ate small amounts of adult insects

and spiders. They also consumed various dipterous larvae and pupae,

especially psychodids, found in marsh habitats. In the chum salmon's

stomach, insect foods often formed a surface layer over a ball of

flatfish larvae, indicating that the salmon fed subtidally and then

9-

fed along the shoreline. The most abundant food organism in the salmon

was Hemileucon, which comprised 39 percent of the stomach content.

Harpacticoids were abundant in the stomachs of staghorn sculpin and

stickleback but not in the chum salmon. Starry flounder mostly ate

decapod larvae, adult CalZianassa, and amphipods. Surf smelt mostly

consumed Hemi leucon.

In marsh pans, staghorn sculpin consumed mostly amphipods, aquatic

isopods, and small fish, while threespine stickleback ate a large

variety of animals, including calaniod and harpacticoid copepods, and

ceratopogonid larvae (Fig. 5). Very little of the diet of the two fish

could be considered terrestrial, although some of the dipterous larvae

live in marsh litter or soils.

Staghorn sculpin and threespine stickleback captrued in tidal

creeks had diets very similar to fish captured in pans (Fig. 5).

Sculpins concentrated on amphipods and isopods, while the stickleback

diet included a total of 40 prey types dominated by harpacticoids and

ceratopogonid larvae.

Several species of fish captured in the slough near the sedge

marsh consumed large quantities of amphipods (Fig. 5). Shiner surf-

perch supplemented this food with the gastropod Alderia and polychaetes.

Ampharetid polychaetes (very likely Hobsonia florida) were eaten by

both the perch and the starry flounder.

Young staghorn sculpin and English sole captured on the tidal

flat below the low sand marsh ate tanaids, amphipods, harpacticoids,

and polychaetes (Fig. 5). These invertebrates are characteristic

forms of tidal flat substrates. There is little indication of use of

marsh foods by the sculpin or sole.

I I I ! 1 U I I I!

Among the dozen fish species examined which were captured in bay

channels, the dominant foods were decapods (especially Cranqon), poly-

chaetes, and a variety of amphipods, fish, and other aquatic animals

(Fig. 5). Terrestrial foods were of minor occurrence.

DISCUSSION

Marsh studies, especially those of vegetation, have concentrated

on level marsh habitats due to their prevalence and importance as pro-

ducers of organic detritus. However, nutrient transfer to aquatic food

chains involves both bay detritus transport and secondary production by

marsh invertebrates in pans, tidal creeks, and adjoining tidal flats.

This study determined community composition, trophic structure, and food

chain relations for fauna in both level marsh and aquatic habitats in

two Oregon estuaries.

Broadly viewed, the study revealed similarities between the terres-

trial invertebrate communities of the Oregon marshes and those studied

elsewhere on the Pacific and Atlantic coasts. The full extent of

this similarity can not be assessed since the level of identification

varied among the studies. The Oregon marsh study did not study season-

ality or identify immature insects collected from exposed vegetation.

However, the data provide a sufficiently accurate picture of community

structure and aquatic food chains for comparison with other marsh

communities. In these comparisons, collection method is discussed in

relation to the portion of the community represented.

The invertebrate fauna of the level marsh, debris line, pan, tidal

creek, and tidal flat habitats are summarized in Tables 6 and 7. The

tables include animals captured by all sampling methods used in each of

these habitats. Taxonomic diversity of the level marsh habits was

highest in the high level marsh, slightly lower in the low level marsh,

and lowest in the debris line (Table 6). However, the habitats share

several taxa. A similar overlap occurred in fauna of aquatic habitats

(Table 7). Composition of the tidal creek infauna is similar to that

of the muddy tidal flat. Taxa from this community also appear in

tidal pans. It is likely that more extensive sampling of pans, es-

pecially in the low marsh, would reveal greater similarities of pan

and creek faunas than indicated here.

The fauna of the marsh soils, dominated by oligochaetes and dip-

terous larvae (Fig. 3), is not diverse partly because samples were

collected dluring the winter and early spring when some insect species

presumably rest in the egg state. The high abundance of oligochaetes

and near absence of polychaetes contrasts with the results of Cammen

(1976) who studied the macroinvertebrates of natural and planted salt

marshes in North Carolina. In the natural marshes and at one bare soil

site, polychaetes dominated (by biomass), while insect larvae and amphi-

pods were dominant in some planted and bare soil sites. Composition of

the marsh and creek polychaete fauna was similar. Among the several

dipterous families Cammen lists, only Dolichopodae was abundant in th.:

Oregon marsh soils. High densities of Ceratopogonidae and Chirononidae

occurred in the Oregon marshes and were sparse or absent from the North

Carolina marshes. Both the North Carolina and Oregon lists are rela-

rively short in comparison to Wall's (1973) list of taxa for Cape Cod

marshes. Thus more extensive collections might show greater similarity

between Atlantic and Pacific coast soil infauna.

4-- -

Table 6. Invertebrates characteristic of terrestrial habitats.1

HABITAT HAB ITAT1?figjF Low High Low

TAXON Level Level Debris TAXON Level Level DebrisMarsh Marsh Line Marsh Marsh Line

Cnidaria Coleopr eraHalaoampa s? p. A Carabidae A A A

Turbellaria A Limnebjidae A A Aematoda A A Staphylinidae

*Palychatta Pselaphidae ACapitellidae A Ptiliidae AHobsonia florida A Heteroceridae A

Oligochaeta A A Coccinellidae A AAranae A A A Corylophidae AAcarina A A A Chrysomelidae ACirripedia Trichoptera

Balanidae A Limnephilidae LCumacea Lepidoptera A A

Cwnela sp. A Pyralidae LIsopoda Dipt era

9 iorimoae'haai'ma Zutea A Tipulidae L A,LLigidium graci~is A Psychodidae A A,LPorcelio saober ACeratopongidae A,L A.L

Amphipoda Chironomidae A,L A,L AAmpithoe sp. A CulicidaeA A

Corophium sp. A Mycetophilidae LAnisogav.-ar-As confer'vicolus A Scatopsidae AOrchestia traskiana A A A Sciaridae A A A

Col lembola Cecidomyiidae AEntomobryidae A Daiichopodidae A,L A.LIsotomidae A A Longchopteridae AOnychiuridae A A Phoridae APoduridae A Sepsidae ASininthuridae A A Sciomyzidae A

Diplura A Sphaeroceridae A A AOrthoptera A Ephydridae A AThysanoptera A A A Chioropidae A AHetniptera Muscidae A A,L

Saldidae A,N A.N Hymenoptera A A ALygaeidae A Chilopoda AMiridae A APentatomidae A A

Homapt eraCercopidae A ACicadeilidae A ADelphacidae A AAphididae A A

A adults, L =larvae, N =nymphs

Table 7. Invertebrates characteristic of aquatic habitats.1

HAB ITAT H fAB ITAT

TAXON Tidal Tidal Flat Tidal Tidal Flat

Lan Creek Sandy Mudd TAXON PA Creek SandyMdd

Cnidaria A A TanaidaceaNemertea A A Pancolus sp. A ANematoda A A A Leptochelia sp. A APolychaeta I sopoda

H~ap 1O8030ZOSO Sp. A Gnorimosohaeroma Zutea A APolyaor-a Sp. A Idotea raecata APaeudopolydora sp. A A A AmaphipodaFldoapio Sp. A A A Ampithoe sp. A AStreb~oapio sp. A A Cororphiwn sp. A A ACapitellidae A A A A Anisogamvarue confervicolus A A A

Neanthea limnicoZa A EohauetoriU3 Sp. AEteane sp. A A A Paraphoxris sp. AArabellidae A Talitridae A AHobaonia fZorida A A A DecapodaSpirorbidae A Callianassa sp. A

Oligochaeta A A A A !iemigrapas cregonensia A A

Gastropoda Colleinbola.Aldgr'ia (?) sp. A A Isotomijdae A

Bivalvia Odonata NCr-dptowya caiifornica A Hiem ipteraM&acona balthica A A Saldidae A,N

Aranae A Corixidae A A

Acarina A HomopceraOstracoda A A Aphididae A ACopepoda Coleoptera

Calanoida A A Hydrophilidae ACyclopoida A A Limnebjidae A

Harpacticoida A A A Staphylinidae ACirripedia Trichoptera

Balanidae A Limnephilidae LCumacea Diptera

CLwmeZ Sp. A A A Tipulidae A,LHeniieuson sp. A A Psychodidae A,L

Ceratopongidae L A,L AChironomidae L A,LCulicidae L ATabanidae LDolichopodidae L A,L LEphydridae L AMusacidae L L

A adult, L larvae, N =nymphs

The low vegetation was inhabited by dense populations of Acarina

and, in high marshes, moderate populations of Collembola (Fig. 3).

Acarina, Homoptera and Diptera were the most abundant invertebrates in

the upper vegetation. Lane (1969) also found that the dominant insect

orders were Homoptera and Diptera in the San Francisco Bay marsh he

studied. He collected by sweep net, aerial net, and blacklight so that

his collections were most similar to the sweep net collections of upper

vegetation made here. Cameron (1972), who also studied a San Francisco

Bay marsh, used a clip-quadrat method which harvested animals from

the total above-ground plant. Thus his methods approximate a combina-

tion of the sweep net and clip-quadrat methods used in Siletz and

Netarts Bays. He found that the orders Diptera, Coleoptera, and Hymen-

optera contributed the most species, but that a pseudococcid homopteran

was the most abundant species throughout the year. In Lane's study,

the dominant homopterans were delphacids and psyllids. Inthe Oregon

marshes, aphidids, delphacids, and cicadellids were variously the most

abundant homopterans, depending on marsh and collection method.

In the Oregon marshes, adult dipterans were almost absent in the

lower vegetation, and both abundant and varied in the upper vegetation,

where ceratopognomids, dolichopodids and muscids were common (Fig. 3).

Dominant dipterans in Lane's (1969) study were Chloropidae, Ephydridae,

and Chironomidae. Cameron (1972) does not provide abundance informa-

tion for Diptera.

On the Atlantic coast, Davis and Grey (1966) collected marsh in-

sects with a sweep net. The dominant orders there were also Homoptera

and Diptera. The most abundant homopterans were cicadellids and

delphacids and the most abundant dipterans were chloropids, dolichopodids,

and ephydrids.

- ---- ... .. . .. i . . .. . - -

Collembolans of the Oregon level marshes were concentrated in the

lower vegetation of high marshes (Fig. 3). The most abundant family,

Isotomidae, also occurred in Lane's (1969) core samples, but were not

abundant in his other samples. Davis and Gray (1966) do not list

Collembola as abundant. In Cameron's (1972) study, a podurid was ex-

tremely abundant in Spartina foZiosa (a low marsh), especially after

high tides. Paviour-Smith (1956) indicates that an isotomid was very

abundant in the high marsh zone of a New Zealand salt meadow which she

sampled using a cylindrical enclosure. She points out that collembolan

densities can be erratic due to rapid summer reproductive cycles and

the animal's habit of floating on incoming tides and then remaining in

dense colonies where the dropping water leaves them.

The coleopterous families Coccinellidae and Chrysomellidae were

collected in the Oregon marshes (Fig. 3), as well as in the Atlantic

coast marsh studied by Davis and Gray (1972), and in San Francisco

marshes (Lane, 1979). Paviour-Smith (19S6) does not list these families.

Several other families (e.g., Carabidae, Staphylinidae, Curculionidae)

are varioulsy mentioned in these studies, but there seems no consistent

pattern to their occurrence. Limnebiidae, abundant in the low sand

marsh of Netarts Bay, is not mentioned in the other studies.

Of four terrestrial families of Hemiptera found in the Oregon

marshes (Fig. 3), Lygaeidae, Miridae, and Pentatomidae, are described

by Davis and Gray (1966) as the most abundant hemipterans in North

Carolina marshes. The remaining Oregon family, Saldidae, is listed by

Lane (1969) along with Miridae, Pentatomidae and two other families

not found in the Oregon marshes as occurring the San Francisco marsh.

The order Hymenoptera was relatively low in abundance in the low

L.." --'.- _ _ _ _ _ _ _ __i IF.. ..... ... ...

marshes and of moderate abundance in the high marshes (Fig. 3). Few

ants (Formicidae) were captured, even in the high marshes. Since the

sampling areas were small, ant colonies could have been missed. The

majority of the hymenopterans collected were wasps and similar flying

forms, which were not further identified. Davis and Gray (1966) stated

that all of the common Hymenoptera in the North Carolina marsh were

ants, while Lane (1969) reported that although an ant species was the

most prevalent soil insect in his study, several wasp species also were

collected.

Thysanoptera were common only in the high marshes (high vegetation)

of the present study (Fig. 3). This order was not important in the

studies of Lane (1969), Cameron (1972), Davis and Gray (1966), or

Paviour-Smith (1956).

Other terrestrial insect orders collected in the Oregon marshes

were Lipidoptera, Diplura, and Orthoptera (Fig. 3). These were all of

low occurrence in the San Francisco marshes (Cameron, 1972; Lane, 1969).

However, Davis and Gray (1966), Teal (1962), and Marples (1966) indicate

that grasshoppers (OrcheZimum) may be common and trophically important

in Atlantic coast marshes. The scarcity of orthopterans in Pacific

coast collections may be both a matter of chance and the animal's

ability to escape collection. However, large populations were never

observed in the Oregon marshes when collections were being made.

The high Acarina populations found in the Oregon marshes (Fig. 3)

cannot be well compired to other marshes because these animals usually

have received little attention elsewhere. However, Paviour-Smith's

(1956) kite diagrams show a strong zonation of mites by family, and

indicate that highest population density occurred in higher marshes.

_ _ _ _ _ _ t

-_ _ ____ _

In contrast, very high densities of mites occurred in Oregon low

marshes.

Araneae populations were relatively low in abundance in the low

vegetation and, excepting the low sand marsh, moderate in abundance in

the upper vegetation (Fig. 3). The present study, like most, has

given little attention to the composition of the Araneae community.

Barnes (1953), however, provides a thorough description of maritime

spider communities in North Carolina.

A striking feature of the Oregon marsh collections is the scarcity

of gastropods, especially in light of MacDonald's (1977) observation

that Assimerea transiucens is ubiquitous across Pacific coast marshes,

and that gastropod densities often reach several thousand per square

meter. Gastropods are common members of level marsh faunas on the

Atlantic coast (Nixon and Oviatt, 1973; Teal, 1962). It seems unlikely

that these animals were common in the areas investigated considering

that several sites were sampled and with varying techniques. Paviour-

Smith (1953) apparently found few or no gastropods in her study.

The fauna of the debris line (Fig. 3) on the low sand marsh is an

interesting blend of taxa found in other habitats. Like other level

marsh habitats, the debris line contained large numbers of Acarina and

low numbers of Araneae. The collembolan family Isotomidae was abundant,

as in the high marsh low vegetation; suggesting that the debris line

of the low sand marsh provides a rich, if unstable, habitat comparable

to the accumulated litter found in high marshes. Other debris line

taxa were the amphipod Orchestia traskrana, found in all the marshes,

Saldidae (Hemiptera), found principally in the low marshes, and Lim-

nebiidae (Coleoptera) found mostly in the low sand marsh. Dipterous

T

adults were not abundant; most were spaerocerids, which occurred in

both high and low marshes.

Several terrestrial taxa were collected from inundated vegetation

during high tide (Fig. 3). Adult Coleoptera, Homoptera, Hemiptera, and

Collembola appeared in many of the submerged marsh samples, and were

especially well represented in the immature high marsh samples, where

several beetle families were collected. Limnebiid beetles were abun-

dant in the submerged low sand marsh as they are during tidal exposure.

Adult Diptera were rare except in the low sand marsh. The data suggest

that more active flying animals (Diptera) are less apt to be covered

than animals less likely to fly (Coleoptera, Homoptera, Collembola,

Hemiptera). Opinions differ as to the ability of terrestrial insects

in salt marshes to escape submergence. This is reviewed by Cameron

(1976) who tested the response of adult insects to submergence by

collecting them from several strata of salt marsh plants during dif-

ferent phases of exposure and submergence. He detected no differences

in these animal communities that would suggest exodus or upward migra-

tion on the plants. He does not provide the taxonomic composition for

his samples, but since he used the clip-quadrant sampling technique,

it seems likely that adult dipterans were not adequately sampled

and that he studied the less active orders of insects such as were

found on the submerged vegetation in the Oregon marshes.

The infauna of pans and tidal creeks includes estuarine animals

(e.g., Polychaeta, Amphipoda, Tanaidacea, Isopoda) and animals of

terrestrial origin (dipterous larvae) (Table 7). Many of the taxa

found in the Oregon tidal creeks also occur in Atlantic coast tidal

creeks or embayments. These include Neanthes, Strebospio, PoZydora,

Hobsonia, Capitellidae, Eteone, Corophiwn, Orchestia, Dolichopodidae,

Ephydridae, and Muscidae (Cammen, 1976; Nixon and Oviatt, 1973). The

polychaete, Hobsonia f'orida, is common on the east coast and is apparently

widespread in northwest estuaries, where it has only recently been

identified (Banse, 1979). The Atlantic coast tidal creeks apparently

are inhabited by a greater variety of decapods, including fiddler crabs

S(Uca), the green crab (Carcinidea nacnas), and the blue crab (Calinectes

saridus) (Nixon and Oviatt, 1973). Only one decapod, Hemigrapsus

oreonensis, was found in the sedge and mature high tidal creeks, al-

though it is possible that such estuarine decapods as Crangon, Callianassa,

and Cancer occur in other Oregon tidal creeks. Molluscan diversity was

also low in the Oregon tidal creeks studied. Only two taxa were abun-

dant, Alderia and Macoma bal.thica. MacDonald (1969) found Macoma

inconspicua (considered here to be synonomous with M. baZthica) and

'!.,z arenaria in a marsh tidal creek of Coos Bay, a southern Oregon

estuary. In Grays Harbour, Washington, he found these species plus

MAcoma nasura and Cryptomya californica. All four species are common

in Northwest estuaries. There was a tendancy for fewer species of

tidal creek molluscs to occur in the Oregonian Province than in the

Californian Province. These tidal creek molluscs are not mentioned in

Cammen (1976) or Nixon and Oviatt (1973), although both Macoma bathica

and Mya arenaria occur in Atlantic coast estuaries.

Few fish species were collected in the marsh habitats. Three-

spine stickleback, staghorn sculpin, and much fewer numbers of prickly

sculpin (Cottus asper), coastal sculpin (C. aleuticus), shiner surfperch,

surfsmelt, and chum salmon were found in the tidal creeks. In tidal

creeks of marshes in the Fraser river estuary, Dunford (1975) collected

juvenile chum and chinook salmon, threespine stickleback, and small

numbers of prickly sculpin. In slough habitats he collected a much

greater variety of fish, including juvenile salmon, starry flounder,

threespine stickleback, prickly sculpin, staghorn sculpin, peamouth

(MyLocheilus caurinus), squawfish (Ptychocheilus oregonensis), and

several species of the minnow family (Cyprinidae). Thus, while the

two studies agree that fish diversity is higher in sloughs than in tidal

creeks, species composition tended toward freshwater species in the

Fraser River sloughs and marine species in the Siletz River slough.

Daiber (1977) working on Delaware marshes, and Slenker and Dean

(1979), working in Sough Carolina marshes, observed high utilization of

Atlantic coast tidal creeks by larval and juvenile fishes. Their

results emphasize the high diel and seasonal variability in catch com-

position. Also, while more species used creeks in the lower more

marine parts of the estuary, variation in use from creek to creek was

high (Daiber, 1977). A total of 22 species and 16 families of larval,

juvenile, and adult fish used the South Carolina creeks. Many of these

are marine species.

Based on Dunford's (1975) study and the Oregon study, the fish

fauna of marsh tidal creeks in northwest estuaries is low in diversity

and does not include large or diverse larval and juvenile populatiois.

Several explanations are possible: (1) The studies did not adequately

represent the fauna studied, with may vary greatly seasonally, daily,

and from creek to creek; (2) The low salinity regime of the estuaries

studied prevented the influx of marine species; and (3) The relatively

simple and spatially restricted nature of Pacific coast marshes has not

encouraged extensive exploitation of the tidal creek habitats by

juveniles of marine species such as has occurred on the Atlantic

coast.

The trophic structure of invertebrate communities in the Oregon

marshes is strongly oriented to the detritus food chain. In the marsh

soil, low vegetation, debris line, tidal creek substrate, and tidal

flat habitats, numbers of detritovores and scavengers far exceeded

the number of herbivores (Fig. 4). Only the upper vegetation sampled

by sweepnet contained a large proportion of herbivores, and this pro-

portion increased from low marsh to high marsh. Herbivores were thus

concentrated on growing plant tissues where their food resources are

greatest, while detritovores and scavengers were abundant in surface

debris and in the soil where their food accumulates. Overall animal

abundance appears to favor detritovores and scavengers and thus the

detritus food chain. This is consistent with the observation that

energy flow in salt marshes is greater through detritus than through

grazing food chains (Teal, 1962), and that marsh plants produce sur-

pluses of organics that are both incorporated into marsh food chains

and exported to other estuarine food chains (Cameron, 1972; Eilers,

1979; Teal, 1962).

As in other studies (Cameron, 1972; Davis and Gray, 1966) spiders

were found to be the dominant invertebrate carnivore in terrestrial

food chains.

Dunford's (1975) study of fish communities in slough and tidal

creek habitats of the Fraser river estuary provides comparative infor-

mation to the Oregon study. Juvenile chum, chinook, and sockeye

(Oncorhunchus keta) salmon which he collected in these habitats con-

sumed mostly aquatic foods. However, there appeared to be more ter-

restrial animals consumed in the tidal creeks than in sloughs, and more

of these animals were consumed in late May than in April. The prin-

cipal prey organisms were Homoptera and Collembola, although other

terrestrial animals were eaten. In some incidences, terrestrial animals

accounted for more than 40% of the prey biomass. The implication is

that the young salmon fed opportunistically on available prey, which

included increasing amounts of terrestrial insects as populations in-

creased during early spring. More insects presumably wash into the

marsh-lined tidal creeks than into sloughs. In other studies of North-

west estuaties, juvenile salmon consumed predominantly benthic amphipods

(Cliff and Stockner, 1973), harpacticoids (Healey, 1979), and a mixture

of amphipods, isopods, dipterous larvae, and copepods (Mason, 1974).

The diurnal variation in juvenile chum and coho (0. kisutch) salmon

foods observed by Mason in a small coastal creek is an excellent illu-

stration of the dietary flexibility exhibited by young salmonids.

Other fish species in Dunford's (1975) study consumed mostly aquatic

foods. The results for the slough habitat were: (1) longfin smelt

(Spirinchus thazeichthys)--mysids; (2) peamouth--cladorera and ostracods;

(3) starry flounder--benthic amphipods and isopods, oligochaetes, poly-

chaetes, and chironomid larvae; (4) prickly sculpin--benthic isopods,

chirononoid and tabanid larvae, and benthic amphipods; (5) staghorn

sculpin--benthic amphipods and isopods, and juvenile salmon; and (6)

threespine stickleback--chironononid larvae, oligochaetes, benthic

amphipods, tabanid larvae, copepods, cladocerans, and terrestrial in-

sects. In the tidal creek, threespine stickleback ate copepods and

amphipods, and prickly sculpin ate mostly benthic isopods and amphipods.

9-.

I |~~I I I I II I ,

In Silet: and Netarts Bays, terrestrial invertebrates were con-

sumed in small amounts by fish collected in marsh habitats, in an

adjoining slough, and in bay channels. Rather, amphipods, isopods,

tanaids, polychaetes, cumaceans, copepods, dipterous larvae and pupae,

and fish were variously dominant food items according to collection

site and species examined. Thus, it appears that energy flows into

the aquatic communities primarily through the detrital pathway, where

it is augmented by inputs from benthic and plantonic primary producers.

This conslusion is consistent with the results of Teal (1962), Odum and

Heald (1975) and similar studies of estuarine food chains.

The information on animal communities and food chain relations

supplied in this report provide a basis for establishing guidelines for

dredging and other activities, either conducted or monitored by the

Corps of Engineers, which may affect Oregon marshlands. Supporting

information is found principally in Jefferson (1974), Eilers (1979),

MacDonald (1969), and EPA studies yet to be published (H. Kibby, per-

sonal communication, 1979).

LITERATURE CITED

BANSE, K., "Ampharetidae (Polychaeta) from British Columbia and Wash--ington," Canadian Journal of Zoology, Ottawa, Vol. 57, No. 8,Aug. 1979, pp. 1543-1552.

BARNES, R.D., "The Ecological Distribution of Spiders in Non-ForestMaritime Communities at Beaufort, North Carolina," Ecological Mono-graphs, Vol. 23, No. 4, Oct. 153, pp. 315-337.

BARNES, R.D., Invertebrate Zoology, 3d ed., W.B. Saunders Co., Phila-delphia, Pa., 1974.

BORROR, D.J., DELONG, D.M., AND TRIPLEHORN, C.A., An Introduction tothe Study of Insects, 4th ed., Holt, Rinehart and Winston, NewYork, 1976, 852 pp.

CAMERON, G.N., "Analysis of Insect Tropic Diversity in Two Salt MarshCommunities," Ecology, Vol. 53, No. 1, 1972 pp. 58-73.

CAMMTEN, L.M., "Abundance and Production of Macroinvertebrates fromNatural and Artificially Established Salt Marshes in North Carolina,"The American Midland Naturalist, Vol. 96, Oct. 1976, pp. 487-493.

CLIFF, D.D., and STOCKNER, J.G., "Primary and Secondary Components ofthe Food-Web of the Outer Squamish River Estuary," ManuscriptReport Series No. 1214, Fisheries Research Board of Canada, PacificEnvironment Institute, West Vancouver, British Columbia, Jan. 1973.

DAVIS, L.V., and GRAY, I.E., "Zonal and Seasonal Distribution of Insectsin North Carolina Salt Marshes," Ecological Monographs, Vol. 36,No. 3, 1966, pp. 275-295.

DE LA CRUZ, A.A., "The Role of Tidal Marshes in the Productivity ofCoastal Waters," The Association of Southeastern Biologists Bulle-tin, Vol. 20, No. 4, Oct. 1973, pp. 147-156.

DUNFORD, W.E., "Space and Food Utilization by Salmonids in Marsh Habitatsof the Frazer River Estuary," M.Sc. Thesis, University of BritishColumbia, British Columbia, Canada, 1975.

EDWARDS, T.D., DUKES, J.C., and AXTELL, R.C., "Soilwashing Apparatus forRecovery of Tabanid Larvae and Other Invertebrates," Journal of theGeorgia Entomological Society, Vol. 9, No. 1, 1974, pp. 32-35.

EILERS, H.P., "Production Ecology in an Oregon Coastal Salt Marsh,"Estuarine and Coastal Marine Science, Vol. 8, May 1979, pp. 399-410.

FRENKEL, R.E., BOSS, T., and SCHULLER, S.R., "Transition Zone VegetationBetween Intertidal Marsh and Upland in Oregon and Washington,"Environmental Protection Agency, Corvallis Environmental ResearchLaboratory, Corvallis, Oreg., Aug. 1978.

HEALEY, M.C., "Detritus and Juvenile Salmon Production in the NanaimoEstuary: I. Production and Feeding Rates of Juvenile Chum Salmon,"JournaZ of the Fisheries Research Board of Canada, Ottawa, Vol. 36,1979, pp. 488-496.

HOFFNAGLE, J., et al., "A Comparative Study of Salt Marshes in the CoosBay Estuary," National Science Foundation Student Originated Study,University of Oregon, Eugene, Oreg., 1976.

JEFFERSON, C.A., "Plant Communities and Succession in Oregon Salt Marshes,"Ph.D. Thesis, Oregon State University, Corvallis, Oreg., 1974.

KISTRITZ, R.U., "An Ecological Evaluation of Frazer Estuary Tidal Marshes:The Role of Detritus and the Cycling of Elements," Technical ReportNo. 15, Westwater Research Center, University of British Columbia,Vancouver, British Columbia, Canada, Oct. 1978.

KLINE, D.L., DUKES, J.C., and AXTELL, R.C., "Salt Marsh Culicoides (Dip-tera: Ceratopogonidae): Comparison of Larval Sampling Methods,"Mosquito News, Vol. 35, No. 2, June 1975, pp. 147-150.

KREAG, R.A., "Natural Resources of Netarts Estuary," Estuary InventoryReport, Vol. 2, No. 1, Oregon Department of Fish and Wildlife,Portland, Oreg., 1979.

LANE, R.S., "The Insect Fauna of a Coastal Salt Marsh," M.Arts Thesis,San Francisco State College, San Francisco, Calif., 1969.

MACDONALD, K.B., "Quantitative Studies of Salt Marsh Mollusc Faunasfrom the North American Pacific Coast," Ecological Monographs,Vol. 39, No. 1, 1969, pp. 33-60.

MACDONALD, K.B., "Plant and Animal Communities of Pacific North AmericanSalt Marshes," Ecosystems of the World 1: Wet Coastal Ecosystems,V.J. Chapman ed., Elsevier Publishing Co., New York, 1977, pp. 167-191.

MARPLES, T.G., "A Radionuclide Tracer Study of Arthropod Food Chains ina Spartina Salt Marsh Ecosystem," Ecology, Vol. 47, No. 2, 1966,

pp. 270-277.

MASON, J.C., "Behavioral Ecology of Chum Salmon Fry (Oncorhynchus keta)in a Small Estuary," Journal of the Fisheries Research Board ofCanada, Ottawa, Vol. 31, No. 1, 1974, pp. 83-92.

NIXON, S.H. and OVIATT, C.A., "Ecology of a New England Salt Marsh,"Ecological Monographs, Vol. 43, No. 4, 1973, pp. 463-498.

ODUM, E.P., and SMALLEY, A.E., "Comparison of Population Energy Flow ofa Herbivorous and a Deposit-Feeding Invertebrate in a Salt MarshEcosystem," Proceedings of the National Academy of Sciences, Vol.45, April, 1959, pp. 617-622.

ODUM, W.E., and HEALD, E.J., "The Detritus-Based Food Web of an EstuarineMangrove Community," Estuarine Research, L.E. Cronin ed., Vol. 1,Academic Press, New York, 1975, pp. 265-286.

RANWELL, D.S., "Ecology of Salt Marshes and Sand Dunes," Chapman andHall, Ltd., London, England, 1972. £

RAUW, C.I., "Seasonal Variations of Tidal Dynamics, Water Quality, andSediments in the Siletz Estuary," M.Sc. Thesis, Oregon State Univer-sity, Corvallis, Oreg., 1975.

REIMERS, P.E., "The Length of Residence of Juvenile Fall Chinook Salmonin Sixes River, Oregon," Ph.D. Thesis, Oregon State University,Corvallis, Oreg., 1971.

REIMOLD, R.J., et al., "Detritus Production in Coastal Georgia SaltMarshes," Estuarine Research, L.E. Cronin ed., Vol. 1, AcademicPress, New York, 1975, pp. 217-228.

SHENKER, J.M,, and DEAN, J.M., "The Utilization of an Intertidal SaltMarsh Creek by Larval and Juvenile Fishes: Abundance, Diversityand Temporal Variation," Estuaries, Vol. 2, No. 3, Sept. 1979, pp.154-163.

TEAL, J.M., "Energy Flow in the Salt Marsh Ecosystem of Georgia," Ecology,Vol. 43, No. 4, 1962, pp. 614-624.

WALL, W.J., JR., "The Intertidal Sand and Salt Marsh Invertebrate FaunaAssociated with the Bloodsucking Diptera of Cape Cod, Massachusetts,"Environmental Entomology, Vol. 2, Aug. 1973, pp. 681-684.

WHITLACH, R.B., "Studies on the Population Ecology of the Salt MarshGastrop Batillaria zonalis," Veliger, Vol. 17, No. 1, July 1974,pp. 47-55.

APf'*, r X A

CRIT1QUf (A: METHODS

Travel among the study areas was time-consuming and the number of

habitats under study was large. These factors combined with weather

and tidal patterns to prevent an adequate study of seasonality. Truly

adequate study of faunal seasonality requires site-intensive study with

summer sampling at one or two week intervals, a schedule beyond the

resources of this study. In retrospect, effort should have been con-

centrated in fewer visits so that the survey aspects could have been

emphasized and thus provide a more evenly distributed data base cover-

ing the various habitats.

Of the sampling methods used, only the corer samples provided

truly quantitative estimates of animal abundance. The enclosure and

clip-quadrat samples were semiquantitative; terrestrial sweep net,

drift net, seine, and aquatic sweep net samples provided estimates of

relative abundance. Because of these varying characteristics, com-

parisons among habitats and samplers have necessarily emphasized rela-

tive rather than absolute abundance. The large enclosure method could

be made more quantitative by using a device which severs the enclosed

vegetation, which could then be rinsed in a dilute formalin solution

to remove attached animal life. This method, as with the one used here,

does not account for organisms such as oligochaetes and insects which

live within living and dead plant tissues and are likely important

factors in detrital and grazing food chains. The enclosure apparently

could be smaller than the 1 m diameter used, since sample counts in

some cases exceeded several thousand for dominant species. However,

this decision should consider the fact that sample counts varied greatly

according to season and site.

Based on the low sample counts obtained for level marsh infauna,

a larger sampler than the 9.8-centimeter diameter corer used would be

desirable, although core depths apparently can be limited to about 5

centimeters. This assumes first that the study of this fauna is war-

rented, and second that an efficient method for separating animals from

the soil is available. The silty soils of Siletz Bay were compacted

and root-bound and thus resistant to simple methods of animal extrac-

tion such as provided by the Berlese funnel. The mostly sandy and

peaty nature of soils at Netarts study sites likely would have allowed

use of the Berlese funnel, although such use would have created dif-

ferences of methodology between the two bays. Other methods tend to be

time-consuming, arduous, or selective for certain taxa, and also may

require special washing racks (Edwards, Dukes, and Axtell, 1974; Kline,

Dukes, and Axtell, 1975).

Measurements of invertebrate drift in tidal channels were non-

quantitative principally because water speeds were too low to operate

the net flow meter (General Oceanics Model 2030). Use of a more sensi-

tive meter or direct measurement of water flow rate appears necessary

if drift is to be quantified. Quantification of fish populations in

tidal creeks apparently can be approached through use of nets described

by Shenker and Dean (1979).

APPENDIX B TAXONOMIC LIST OF INVERTEBRATES

Phylum ProtozoaSubphylum Sarcomastigophora

Class RhizopodeaOrder Foraminifera

Phylum CnidariaClass Anthozoa

Subclass ZoanthariaOrder Actinaria

Halaccmpa ()sp.Phylum Platyhelminthes

Class TurbellariaClass Trematoda

Phylum NemerteaPhylum Nemat-daPhylum Annelida

Class PolychaetaOrder Orbiniida

Family OrbiniidaeHca,7,co Zor Los sp.

Order SpinoidaFamily Spionidae

Pol2ydora sp.Pseudopo~ydora sp.

70~sp1-a sp.Srebospio sp.

Order CapitellidaFamily Capitellidae

Order PhyllodocidaFamily Glyceridae

Gy~zceraz sp.Family Nereidae

Jecvithes 1 mfi-coZ-

Family PhyllodocidaeEteone sp.

Order EunicidaFamily Arabellidae

Order TerebellidaFamily Ampha ret idae

ilobsonia floridaFamily Terebellidae

.Amaeana sp.Order Sabellida

Family SpirorbidaeClass Oligochaeta

Phylum MolluscaClass Gastropoda

Subclass OpisthobranchiaOrder Sacoglossa

A~deria ()sp.Class Bivalvia

Order MyoidaFamily Mlyidae

Cr. ,ptomya ca lifornicaOrder Veneroida

Family Tellenidaetcomna baithica

Phylum ArthropodaSubphylum Chelicerata

Class ArachnidaOrder PseudoscorpionesOrder AranaeOrder Acarina

Subphylum MandibulataClass Crustacea

Subclass BranchiopodaOrder Diplostraca

Suborder CladoceraFamily Polyphemidae

Podon sp.Evadne sp.

Subclass OstracodaSubclass Copepoda

Order CalanoidaOrder CyclopoidaOrder Harpacticoida

Subclass CirripediaOrder Thoracica

Suborder BalanomorphaFamily Balanidae

Subclass MalacostracaSuperorder Peracarida

Order MtysidaceaFamily 1M-ysidae

.eorn'sis 7ercev .-sOrder Cumacea

Family Nannastacidae.w~rela sp.

Family HemileuconidaeiemiZeucol sp.

Order TanaidaceaFamily Tanaidae

PcMCOLUS sp.

Family Paratanaidae

Order Isopoda L~oh .i pSuborder Flabellifera

Family Sphaeromat idaeGnoyriosrphaerona lutea

Suborder ValviferaFamily Idoteidae

I'o tea fewkesiIdotea resecata

Suborder OniscoideaFamily Ligiidae

Ligidiumr graciZisFamily Oniscidae

Porcel17-o sca ,erOrder Amphipoda

Suborder GammarideaFamily Ampithoidae

Arnithoe sp.Family Corophiidae

Corochiwn sp.Family Gammaridae

An7isog=rnrarus confervico c.sFamily Haustoriidae

Eonaustorm.u.s sp.Family Phoxocephalidae

Para,,hoxAs sp.Family Talitridae

orchestia traskianaSuborder Caprel lidea

Family CaprellidaeSuperorder Eucarida

Order DecapodaSuborder Natantia

Family CrangonidaeCracon franciscor'umCrangcn nz,7r-scauaa

Family PandalidaePanda ha danae

Suborder ReptantiaFamiily CallianassidaeFamily PaguridaeFamily Cancridae

Cancer raisterCancer roluctus

Family GrapsidaeHemigrapeus oreconensis

Family MajidaePugettia 17roducta

Class InsectaSubclass Apterygota

Order CollembolaFamily EntomobryidaeFamily IsotomidaeFamily OnychiuridaeFamily PoduridaeFamily Sminthuridae

Order DipluraOrder Odonata

Suborder AmisopteraOrder OrthopteraOrder ThysanopteraOrder Hemiptera

Suborder AmphibicorizaeFamily Saldidae

Suborder GeocorizaeFamily LygaeidaeFamily MiridaeFamily Pentatomidae

Suborder HydrocorizaeFamily Gorixidae

Order flomoptera'Suborder Auchenorrhyncha

Family CercopidaeFamily CicadellidaeFamil:f Delphacidae

Suborder SternorrhynchaFamily Aphididae

Order ColeopteraSuborder Adephaga

Family CarabidaeSuborder Polyrphaga

Family HydrophilidaeFamily LimnebiidaeFamily StaphylinidaeFamily SilphidaeFamily PselaphidaeFamily PtiliidaeFamily feteroceridaeFamily loccinellidae

Family Corylophidae

Family Chrysomelidae

Order TrichopteraFamily Limnephilidae

Order LepidopteraSuborder Frenatae

Family PyralidaeOrder Diptera

Suborder NematoceraFamily TipulidaeFamily PsychodidaeFamily CeratopongidaeFamily ChironomidaeFamily CulicidaeFamily MycetophilidaeFamily ScatopsidaeFamily SciaridaeFamily Cecidomyi idaeFamily StratiomyidaeFamily TabanidaeFamily Dolichopodidae

Suborder CyclorrhaphaFamily LongchopteridaeFamily PhoridaeFamily SyrphidaeFamily SepsidaeFamily SciomyzidaeFamily SphaeroceridaeFamily EphydridaeFamily ChloropidaeFamily Nluscidae

Order HymenopteraSuborder Apocrita

Family FormicidaeClass ChilopodaClass Diplopoda

Phylum EchinodermataClass Stelleroidea

Subclass AsteroideaOrder Forcipulatida

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APPENDIX D

INVERTEBRATE SAMPLE DATA

Abbreviations used for gear in this appendix are:

AN = aquatic sweep net

CQ = clip-quadratLC = large corerLD = large drift netLE = large enclosureMC = medium corerSC = small corerSD = small drift netSE = small enclosureTN = terrestiral sweep net

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F. (3 t A Lam U% AALn C N - .4 m0 Nl

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(A.4 4A v 0' L V A 0D' 00 A 00~ LA -X

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f3 C

0.0.L 0.w4, 4 .4 ( 0 0.0.

M. 0. LU.L AA .*1W

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Table D-17. Density (number per in2 ) of pelagic and epifaunal animalscaptured by LE3 in the submerged level marsh of the Low Sand area,

February 1978.

4 ~ SL SARO L SAND L. SAND I. S4)N!LE 01 IPS, LI L i

LIFE STAG E01ISD

41.4114A SOP ADULT.S 2? 4.3)1 .9

.EIESYA SPP A CATS S .61 1.1NIE4.!,;4

51.942014 SPO £014.13 1.11 .4

A454 SORE .A P ADULTS S 4 1

1.1L040 0O SP 1 4O...O 4A S IP A01M.1 :' 0 42 17161 1. I 1

A-1 EA65 SIP ADULTS 3 4 I. t4 7.4' 6.3.

Al"..5. SUP A.3U..2 & L16 452 97'. 467.1. 736.61

C114I00 OULfTJ : S 3 a.

I 9254

.2. r1SZ.I 91141 so tat,24 2

1Ea~ .01 2.21 1.6)

24...34 SP 41.31.31 .31

.C .4 Et.sT II ISM -545 SUSPP.2 .4t 1.1

JUL5 1 T9 S 31 s 21.It '& 7: 1:

.31 .5,

C041 *J3 2454 4'. 1173 1

7

tAO - 2 -

- - - -~ - 4'4tA r... ** -- -

0 4 - - - 4- - -

A, - - N

-

A- *~ 4 - S a -

4~4

o -s 0 - - 0~tt *4 A, -,Z 4' 0 C - - -

- ~J.h 4

-~ - -'0 -. 0 4...SA ca~,..s I,. -

- 4.1

N A.~: 4'~~ 4 -~A4 44 A, * *-A,*~4A, *- - S

0

C) - - - - - - in a tnt., t, mnu,~n a tot., ntt., avwm, ,,,...at.~ 0, 4

C) 1~.-~ 44~44.J.J .4 5

- a 'S ~ 0 = 4 S'S C 55 ~S 4 S~0 '554 'tOSS ,S44Z~t.t'5'S

.4 4 4 4 4 4 4 .4 40 4 ** .4 4~4 444 4440 8

.4.4.4.CJ440 4

o

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t j .J - , -I , A 9 ' .3 -3 Q. _- 0.CJ A... A CJ 0CA l

of M A .0 - -9 4- -- 01 0 A0 -tf W0-j 22nf(44 0A .1 4 ( A .4 4r .4 'a 4^ le Z4 44 -9.J

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- - a Ill ~ N ON a- C N

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.4 - - *4 N "1 S

- N - '0.440 - C 1.. 4% N.4 CC 44 Q~ 4 SN 0 N

- - 0 .4 N N C 0 - C N'~, C - - .4 - -

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.4 N .4

C ~ ~ 0

C-

0

I'\j -

-~ - C

S.. 5-. 4 .4 N 4.t-4 0. C O.L4 0. -

0. C C 0. .44.4. 44.*,a 4. .4 0 .4 0. '.10. .4 C

4-' 0. 0 0.fl 44014 C04 .4 0. = 4. C *A 4 404.410.

S.. - £ 0. 4.CO 0. 0 VI .~ SlIM 0.O...,.14.*0.C) 0. 4. *044 C 4. '4 4 .4'4 .4 4.0. 4.4041414 .4

.4t40.A~ 0. 0 2 t - ~,O O.~4ON04 ~.0 - .4 14 4 1.44 0 4. CC *44 14 ).3....J..4 -.4 4 C .4014 CO C - .4 14 ~C CC 4.0.00004 -

---------------- 34.00.40 0W 0 ~?£ .4.4 2 £1.3 .4.~g.a%3.Jr.o.ra Cot--.- .4

o c~

C)>- C).0-

Table D-21. Density (number per m 2 of pelagic and epifaunal animalscaptured by LE in the submerged level marsh of the Low Silt area,21 July 1978.

L ~£3 SILT 6SILT L. SILT L. SILT

SAPE027 car 07 GA.

TA34LIFE STAG.E HEA14 (SO I

CNI ZAiAd,.0AAkIA SPP A.)ULTS .3 * S)

POL1C.AETAPW L.NLA FLORIDA A.3ULTS 5 2. 119 31.43 1 50.$)

SL..CCAETA SAP AOULTS 72 a 167 163 44.11I 66.4)

'L.,Ck;A SPA A3UULTS b 1.64 ?.?I

ARA £EAEAR ..NEAE SP AOULTS 146 9 37 '2 4.8.91 36.31

1 6. SA ASULTS 15 3 2. I3 8.34 7.1)

0 STRAA.CSI6A30A SOP A9ULTS 5 L.31 2.23

EMILJCO SP AOULTS to b 6.03 7.31

r.RI0533PP4AEkQP1A LUtEA AaULTS 13620 753L 4.03 7103 651M.21 (.780.23

CQ..CP"IUM SOP AOULTS 306 L. 36 5.3 129.33

'I4.E A PA AOULTS 1 .3( .511JUL7 3PS' 1 2 L.51 2~

, 6 SI31E SPA r z Z 3

STl"r fSNIC SOPA A JUL TS 4. 1 1 3 9.*23 6.1

"CIATELJ~E ~.P ;A LV to 51 45

'J~T3A 1P, AUI.S1CI 4 L. E SP IAuLT 1-6P 131JAEJ: SAP IAULE G1 3 .33 Z..)

341MEOPUTER&

n''IENOT.A P PQAA 52 1.3 2ZZ2

PAT. LT. "ACE

TOT^CG UE P RAL 16? A6 .46 1Q32:2: 0:

_____ -- U EU U --

-~ _____________________-

.1

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-- ..

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S.. S N'O4C.44.04.N4.4.4-4 3.4 - - *0 -4-44.4

4. * , --4-4 .4

U

- -- - .4 .. a - .4 A 4 4 13% '.4 .4 040%34 - 4

a 04 - ~ .lWf 0 S 4 N N 0' 44 ~

C, - -. V~ .0 M N S49 0

4.44. - 4 N .40~- 0 .4 5 .4 -4 4

.400 N N .4 N 4' .44 -- 4 N P - 4.

-4 .4 -- .4-4C)~-4

C) - ... * N .. 4 - - Ca oN a C

~ 4-a

C) 0 -4C4 C, 4, .4 44 44.4 4 .4 .4 31 0 O Li .4 44444.1 44 .4.4a~..aLJ.41.J 4

0

C) '4 -S .5 a-S a a a a a a a .4 aa~~ a o.a----- .4t4

t 4 - S S SO S S S S S S S 4 55.4 5 fl &X42XX*A -0 5 4 50 d S 5 N 44 0 5 050 0b-52.4.g,444o -~ .4 4 4 44 4 4 - 4 4 4 .4 4 *4 4 2.4

4-4

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-4 44* 4 4.0.0. 24 0 44.4

4.4 4. - 54-4 4) .44.S... 0.44 0. 31 0 42 .4 40.0.

0. 0. 0 .4 4' 44.34.40.C) C) ' .4 0. * 4. 0 4.444 0 4.443.40.0.

- 0. 0. 440 0. 0 .4 44 4.54 0.0. *hO4.. 44.0. 0. 4.5 4 0. 0. 0. ~' .4 444 .4 0.44 4. '.34"A

E.a 4, ~ 0 4' 4' 4 5 4444 0...450.S* 4 .44 44 0 4 .4 44 52.5 4. .444 .3452-44

C 4.. 4444 Owa-s.444 .4 - 0 .~ S.......405000 ~. -~ 44444.44j45244)44 S145.,4-5.45..4

4 .44.4 44 .4A 0. .3 4... .. t 440.4 LI4. 4.44 .r .4.5.15 5.4 .. 00 4004454 t4 35 54.4 .5-4 43 44 44 44. 44 .44 45 444.43 12 -Jj 4.444r....44 04 -z u4r4 'Ia 4'a..e4 4.44 d'4 0.444)02 .. 4.424' l.a 04.0 ~4.4 fl 4' 4 03 44 4.jt 40 , 24 44 Fr, .30 4w 4.440 42 0444 .~S.4,c2.Jaa.-.-

.44.5 - 4 2 a 0 4 4 4 0. 0. 0 4. 4 4' 0.ja?4 2 .4 42 a 4 4 U .4 44 54.45.443.4

'A C, 44440.0W 4 2 0. 0 4) 4 4 0 U 42 .4 4 0 2 0

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-~ NJ...N. C NNC - * 9 4.04N*.2 S N C 4.04

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$4 -

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~. S -o a ~s . a'- -. -

Ci S a ,. a - N N - A

-4

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.4 ivY a a a 44444 aaw4.4tiia.,~.4L..hi4a hi a

.44 4 4 44444.44 44.4 3.444 4 4,ii;~. .4 .4 .4 ~ .4 .4.43.4 a 2 -a .44*4.24 S -

- - 0 .4 0 .4 .4 04.4 .4 ~.4.4.j 4 *~t44444 a4 4 4a4 a 44ac a 4 4 4444444 4

C)

N -

0 yhi'4 4.4 4.4 4.4.

N 4 0 4. 40.C) ~5 a. r .4w a .~&4.

C-C) 4. hi & 0 .4 4. N-ta4.41 4.4 4. 4 4.4.404 4. N 4. 4..,, 434.

4. 2 44 hi 4..S4 4. a 4. A4..4.-NI 4.S., C5 4. a 4. 4. .u 4.4..4a.4tm.. a a a 4.-h?4.h44I) $44 - 4. 4. ~ 4.440. 04 - N

444 4. .4424 -i 4 4 4 04N44C) * * a ~ .~ ~ 3 .42-h a aN.4. a4..42N.(.i4.N .t N a 440 44 n.4aN a .tr ~fl Na.44,aS .4.

N 2 ,hi.3 4 444 4iN4 t4.Z4hflaN..4 .

.hia 33X.4ZON 410 a140.h-h4 hi .J4N.4a.h.~Shi -*4 4-N 44 444.a. 34.3 .4C) 4.4044.4* .4.4 .34 044.4.4 .44.tiL.4i44 hiN 4.44.-i- - - - - 04

4 444 44 44 4! 5) .iCt~ .4444444-4 .- t 34 .444 41444.44. 41.-..4 4. aX 04 32 Nt) a.. (3.4N) 0* NtO?'V a ar-ha-

44S5,hira-h *44.4h

2 N 44 22 41 tO .44 4.44.4 4.4 44440 4,4 .44 ... 4. 4.4 C ~44444.h. .444N N .4.-haa*t a a -.4 4. 4 N 4. 4.>.4C.J 44222 4 2 4 - .4 .4 = 4 $3 .4 .4 2

44.N4 .

4 hit) 0 4. 2 04 U N 4 04 - 4 0 04.444 -~

44 44.4~2 4 2AC

'A

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AO-AI06 973 OREGON STATE UNIV CORtVALLIS SCHOOL OF OCEANOGNAPHY F/0 8/1STUOT OF THE INVERTEBATES AND FISHES OF SALT MARSHES IN TWO -- ETCcU)JUN 81 0 L HIGLEY. TR HOL.TON DACW?2-77-C-0013UNCLASSZFZEO CERC-M-81-5 NL

minl~llllIEIIIIIII

EIIIIIIhhhm

Table D-25. Density (number per m 2 of pelagic and epifaunal animalscaptured by LE in the submerged level marsh of the Sedge area, 6April 1978.

sort SIO ( S[OGE SIO1( SEDGE siarl

C41"tO.46A S66 ADULTS 9 1 1s 1 25 94.54 ?.61

A.)101 U6 34. T 523 I71 211 131 345 29%.71 1.7.1

CS4.44 AO'3G44UL.S 1 .31 .1

16'.waSI ADULTS 121% 1182 6,9 41: to? Wi::: Z114.1

-9~1 A A6 430.1 931: 2667 1'464 1234 M41 Is,,.:4 S4.94

~S444LTS 36 441 7 1 7.64 1..61

OO!1?F4AOO IIAUT 3 20 i 33 364 133.64 I4.14t

COP ADULTS LID 65 21! 603 731 349.4 244

ISO4 3.4

.G1.6. Sop66

4o .A S6 4J4JL7SIO~u Ag" 1 1 .IN1 1.361

JEADPODA SIP 4044.12 1 .34 .5s

'4L. I". U07

L A1U 44 E 1 11 .

;0'0 2ER AJ~f 3 1 1 1.4 10CO 1E 34 .7o SI DLS 3

L I ".2. 11 A SRI LA VA 3 6 4 .44

.64466 PPLAVA 1 3.14 0

31-1-A~.5 1.64 I$ a? .3 6 a

J.L3ES- 3413SIP 33~ A6

13...E 366 IF?44 2 31''

A0744. 5613 513 to,5 33 5

CO49i'EU

0 ~ ~ ~ ~ ~ ~ ~ 3 - :C .*4.4q~~ 7s.-- 4.T MM -- s Rea .9w- .- 6,------ ~ _ _ _ __ _ _ _ _ _

C)

C

4)

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4)

4.)

Al 4 ..~ ~.4 ....4A. A 4e4% *4eC~ .4 .. 4.4.A.4r.

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.~ ~ 4% 4* *9'N -

C..

A- .4y .7 4 0 - .40 *.4S4~*

@9 * - q. 4, .4 0a a -. - *

C.40 46 44 9 .40 .40 *A.4 -. 0

eM 04.. 4 .4 9 0 .4... -- a - .4 0. -

0 -

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-4 .4.4

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.4 @4 A S 4 4. * '4a a @4 .4 a

N

- S.. 40.) C) - - en - we even en.........4.4.4.4 en

- 4 W14t4bbbb44 .4 4.4 0

4 4 .4 .444 40 4 4.~en~*.4*4 -

4) ~.s - &.4

~ C) 4 4. a.,

. .4 a ~, a4. 2 - 4. en

i-v ~3 4. 0 *e a. a.44~a *0) 4 et 4.4.4 4. Ma a 4. -~ .44. .4.00) .4 O>fl0 4. 4. 44. 4. '14.2.. 4.

4.4 v.4 - 0. .441 044 - 4.0.4. 4.-enter'v.. t . 4.4 MI 1$ 4 4.4.4.MJefl0~04 C.4 .4 et .44. 444 4~ .4.4 .4 44.0.4.4 0.4.40.4 44

Z ~ 4 4, ~4.4 .4 e44 aSc 4 Oe&Ot...4 4..@4~ Oa.-e 4-. 06.5 *.43.4W -04 -:4) a .4e4 en 2 at 7.44.4 dO .4~ = .42.4 .4.4 92&0Z4440 eO

S 4! .4* 4* 44.4 .40.4 -~ .400. ~-44 0..~ aM)4)4*454 3fl 4~44.4..4C4 4..4~ ~4C) .4 0.4 MO .4.) e,.e..4 -. 441 4.4 tOM, 06-V 0%. M---4. 00.404'- 2.4 '00 t .4* a.4*.4 SvJO ~t4 0? O.4QJ4.4

400eM.. 014.34... .12

- ~4 4 4 0 4. 2 .4 4 0. 4. 44.) 44.44.444 .4 4, .4 .4 .4 0 4, .4 4. -

A 44.44 0.40 4 4 .4 4 .4 0 0 44 0.47.-C 0*

CC4.4 44.4404)0

0.

A.1~

i~i-

~1I

Table D-27. Density (number per m ) of pelagic and epifaunal animalscaptured by LE in the submerged level marsh of the Immature High area,7 February 1978.

'E I,

1835LIFE SIAGE -EA-5505

INVE4YTIkS!

OL4141C.5&ItA 109 ADULTS 6is 2* 13 10 6.1 08.41

GAS t4.PCO

"50.Pool SPA umSPECLF1IO Is2 6.75 10.85

A ":A SP ~ uT: e i 29 .::.41 291.15

IA; Ri1 &O DLS11 1 I S

.I * 6 I A, 'II ID P V t2 1C LSEALA OP A3osLiS 14 it 3 1 10.35 6.5

O t.NLJnA 78G18 L481 IS 8J I 5 5 6.55 1.31

179CTA. 1 1:

'P~~ A.Sf 3 8 U11

CCL tJ.. SIS

AL6$'. 3.5SP jLS o 34 3 46 .75

.. 14"tP4Is.10A 50 888 5 1.5 lZ

t ...QE 10P A*A 15 93 5

V41051,1.. 1

L'0 " sfP aOA 1.4 .2

-. 60P P1.8 LA..sJE .b5 So1

In..I.O so , AASb 1 b .sp?

SISAL li13 308% 63z is.

Table D-28. Number of animals taken by AN (non-quantitative) in alarge pan of the Immature High area, 7 April 1978.

AREA III MZS AMNPLE R ANSITE CL.SAMIPLE coal.

TAAQN LIFE STAGE

INifERT E3IKATES

OLIGO1LHAETAOL GOCHAETA SPP ADULTS 1L5

t ~~~~COP EPODA fC P .UT

CU'IACEACUMELLA SPP ADULTS I

AMPHI POOACONCFPHIUM SPP AJULTSIANLSOGAMl1AUS CONFERVICOLUS AJULTS 350AIPITmOE SPP A13ULTS I

0 0N A TAOOUNATA SPP NYMPHS

HE41IPTERACOeIxI0AE SPP ADULTS L

TRICHlOPTERAL..ZNEPN4Z ..OAE SPP LARVAE 2

OIPTErkAOIPTERA SPP ADULTS I.EPmiYRZ0.AE SPP LAI&VAE I?U.CDAE SPP LARVAE 3CH.RONOMIUAE SPP LAkVAE 1

.;ASTEnGOSTEIDAEGA.4TEi~0STEUS AC ULEATUS ADULTS I.

TOTAL 1080 .

CA I

0

00 0 .N S

4-J V.

$4

el In.C M q .- 4 -d A - -9 -9 .- ..4 4 S P76.4nn i4~

(- CA

a) 0

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0. 0.4 CL TN L 'Iii i I Iii w 4. CLI.4I .

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x-,- ol :1 0 Uoc X.V ,.ja Z

'If 4. -- qa.

a.ca. a avO . a.v3

Q) Mf 4w U%.Dt% N1" .0 0 .. ,

CD LA% a% s (*J* N do ~ M SC .- cm Cary-,a a' 0' N2 4% MrK

4 -s - -- -.T j -o n

o0 C* I"4. uN Na uN00* *

a~) cu

0 -. .4 . N U'c. 0N

c4M (P N M N N o N N

0 . 6% r4 on'S~ . wr v7. JO

(Ao (4 .t 4

t- L

-~~~C 0 o ' C4-o Lo 49 N 49N -

'-I () IAfl(A

0 0 4.4 0 U .4 N .49 .'..

-i .4 4 NLQc

T- (L ( 0 0 L Lm E

-9 -JjOm -4- c -" oA -s IA IAAI IA -t -a IA UIA IIA

La ~ ~ , .J .J o..J.. x3 . 3 . .. 34a X, 0 a:.3 (L 0W Q3 0t -a o.

_jI z- " z '.u 0 _j0 0 0m v 0

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a C% P 19U CP N W .0

- ~ ~ t 0 94 P -% * a

4r -0 -09.j :U (.4"1 0 .N U

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N ~ ~ a a CP N N. NP

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L4 -", - :3.4 -4

La 0 Z0. a i-=0 -40 0mu A0 ~o

0 94 0.9 N441 0~ ZO. 4- 00v 0-- 4 r. -* 0n J" '3.

JXX 44 le 44 4 44O-g w~ Of ,3 o 44- Ix -9 04"I 9.4 0i

uj 0 -C z ,-.J 9.1 ;z X4)4 4Z3

4 . 9 0 4 4 4 4 3 0 0 r1- J 0 9 .~~L.I w u z~ (L 0.~.. ~ UZ J.4 1C) I-- LA w4 04C x. JJ -0

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.4N P.. ?I co Nn 0n M UN I) 0

0 q, -t C*'4 0 4 * ~ C N ~ 0

(4)0 .4 - 4 . Na

W.A 00L.140N L) 0'C 1

~~CC LA 0m A N I~~(74 -0 -00. C.

wu L)L .4 Nl

C4. 0n (n (A O~ V 0V I 0 V) 0 I 0" 4

4~~~- 4 4i _44 4 4 4 4 4

M.L t: 3 3 1 0 am mC)- -99S9 - 9 - a- .

CL~ 0 a -

LA CL. (A _n0 0 4£0 LA..-. b -j -so z. -S0., 0

. 0. 4 U0-h .4 d cc 0 . 1.

4 c Q IWAQ 4O .11 44 .10 .0 4 4. X3 0-0 34 04 Is 8.'.. P.S "a a 09'. a " 4 0 0.

(V I 4 r t 4. '.110 Ul 0 J- 4.4 .. 03. -) 4-vX a- 4 . 04 4.-3 O4 .40 ) -j- 0dl~ 04 Ou. -0( 44"

* 4>r s u w- cK o - u xi 'r t4.'-' (D"1 .L C ' 2 L4 L4N A 0 i .r A" OX1. r14 c0 .

.4 1 4 2 w, -Z U Q4 i A 0 M rooW0 0UM 0 1. .9 0 'A 0 3 Q 4W 0

gA--cn

qT 0 Z

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a' 4-da fn ~ .0' r- N.iU 'aC N4 C2 m Q 4 02

444 00r s

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u ~a r- 4 4 0 N 'N03' 0,.3 L1 7~' . N z' . 2 * a

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CL C) UAv L 0 v a Q

a) Ln (AA 0.. 9 ;'-i - 44 .i a L r V 0

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tn I. 0.C 0~ Ln 0.w 0 U . 0..40. 0 4.4 a 1454r 3 C

4 S of .1J54 44 0 0 4 U , 0

,n x44 41- WS0. 40 UIJ 'M . o4 uJ- OLO

r-LI 0 Si m .It zZ '.3 Cl1 'A0 344 (A1.3.3. m 42L ai w Q z CL0 0 Q , 2 2 -0.1.0.0S (. . 0 U 0 U - .

00

P.. Nv JNI' .0 .04 N C, Ps1 0

u 0 N0c a 0 'J. M .- 0 CYN N4.0

0 o U2 cy IA -0 0 -0 UN N A

0) -) 'A 0i'7 A 0

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U ~ C mA Nl.' 0, N ' 4 N

C) -y

M4- 0 . 0' -0 NJP (P 40 J 14' N .oj tA C N 0. 0 v1 m 40 C'4'0 w4 ' A N. N m

'4 NC) on

:po 1.tC M.0 a.. 0. C N . 1 'M~ .4I F, " N 0' N 0 0 .,'0 r

D. V) C A N A 04 CA A (A4 (AA0'UA

-1 '40 0 O .4 .-4 07. 4

.- *0

a- 0u NI T4 o~'

) - 4A NLC

wC Is A ) OC L

0A 49 -V- 4" ( 1 10 A

U. .4 0- =00 0 0 x 30 .0 0 0 N4. _L3 0 < 0 .4 x" M ~ 0 0 0e 0,'. 0 000If l 4 43 c 4 r 46 'a 4 a4 a I44c C

- 0-

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- 10 M 0 .N N-0 z p

-cr ).

N~C~ .4< 0 Nt.. utP. N 4 - .

4 0 3'U' U'. N 4U

M 4D 40 0 T 40 . 4 .' 4

a 4 . .t-0 UN 9.7

'o. 4-t -VrI

0 ~C Fnu 0 ' OP 0 U' 0 C* . 3.

L5 .4

(A W0L 0 a4 N) W 0 EA 0 0 OW 0

a M 0 In U' =,a 13 x

3~- 0 04 0

0 a. w C0 MU. 0.

0 M 2. 4 1 - V

a-4 - L I W 0 t

.- '-499 C3 1 u- -

W) r

Ac j GA GA- 3c L U'. in 314 LA (AL xA -)L LA U) r4 . .. -r I

-LI .9 z ' Ll t 00 1" 01 -11 0 -, 0 07 OZ M m 4J

LnA

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w It L-0 N

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Z . ;49C 1 44 ~jj .4ffl a, I" tVfl Ne'J .O.%~ J 4 pfj

QCA a.0 N0 N 0'

4 1

-CM c- 10 N 01 0' ' N 40- ' z.0 s0 Wl 9 t r .

=. .4 , .94 10 .90 r N U .99

0e 1

X

(AN 0 (A 0 1^02( u

I j -i N i -j -js -j 0' .4 . - ' -z4 :. ) z9 9:30D D:). 0)z

oN' C, 00 m44. '0 1-3 .-99- .4 4

2~In00

-. 06 m4 L) wN

tn 0' V) a 'LCr - a 0. 0 .5 J I /),0'DO 7

a. 0. Q. -% -, ..C4- . vi .

0 n.. .4.4u

w X

oa O4mro oA w, Ai w U, -- £:)' (5Uf -t- 4 0

ka -~ 0 00 I Z o N

r- r= 0 ", Ax00 00 0 0 4

> z w j 4 r 244 41 44 G4 zi .wh L

w ~ ~ ~ ..4Q vx tz.QIL

z '

0.0.0 0

CA ~ O LAa-'a 10 Mn M n - .tN

-d

4J Ny C,

t4 .-0 C*0

400

u ... 14 'a1 1

4D N

-~ CS~ .. a a CC 0~~ P.. 0 %n

r 1/1'~4 '

0~~~ ~- V. 0 - . U 0

m4 0 A

- 5 b. I-

0 ui -i - j Ji - -i 1-.,

:2 :3 u::flVm ) mt mn V)fl

- 0 Q amm03n 0 3 On0

C) C

'AL4 4 0.0.LCLv Q0

nfil 00 0. 0".L

0. 0. 2, Laa. 00 .0. ~ ~ ~ ~ U 0.00~.- 4 dG

j) t'4In .4 4.3-O J4 al e

ce 4- 1. z4 .4- U).(L o 00 .J 00.4

0. w . 0 z4 CL4~'U .9~

*~~~~ C)r ]04U

tr4 -L )T,~- 04 ..

;44Z50'0~.4 Z 9 40 aN.O4 -444F

- aP..ttv " N P, (r -- j f

4--.i N V..0 N. rm F".4

z C 1'0.00 s ra Q' c2 OPa- .0.4Nn O ~ N ' . P. OP',4 N-

4-J~c 0N P'S na0,4 N% in %'S~l. .4'PNY .40. .0 CY C .

O C 0 -- 0 1 - 41

4-

-C2070 0

U44

5-j

40 =( ~no 0' Ix-x 0

~~- -j J, -A

'EL) 0En

~4,'S~ .5. 5 4S.0Q

- ~ .a - 53M.- "1 4'.54'of

mI a aC4(L L

(L0 V

(LC L (L 0 wCd)n-*L C L f 9V

vi le'4n 4. C3,a

4/4 ..JOl .4 .4 . .4 .3 x.4 Q..j.J. 4FD' 04A - 0 .9 00"CL 80x'L.4 Cuf .40 0w00LO-A 0 0 W0 00000olc 0-544o

01- 4 4 =444 <4 4. 4- 4O 41= 4.44.4-4- X ~ V, g

r~#5IL j

z- 09.m 0 U ) z Iwrt~x A0

W45 .9

4-1

o a

I.,

0 '0

,14 Vw

C)C

I CL

~~aa

f2 O-4 -- 4

j 33.

W4A -t

.1,

41I C.

M- C

z.C3

C-4C

Z,4 -i

->

000n

.- ~ _

- .- L"

0 t jrpr

'A 't MA

Table D-42. Number of animals taken by AN (non-quantitative) in twosmall creeks of the Sedge area, 6 April 1978.r

A4EA SEDGE SEDGESAM.PLER AN AN

tAD LIFE STAGE EuIgI VE TE34ATES

CM.AR P 5 AUL ?S Lao ZT Y 3S 65MENER TE A 3 1 3.5

NEMERTEA SP ADULTS I St4 .51EMATODA SPP AJULtS : ±2 i:::: .51

POL rCMAETA

P1OA ULtS 43 3 5C 3.AD SONIA FLORIDA ADUt 2155 21~±j0OrCMAETAJU S2S 6 4.0 1:.

GLiGUCHAETA SPP ADULTS 291 ?1 5.0 11.4

aL'..A SIPP IARAE I*5 4AL IA Se .JU9ENILES 2 1.0 1:.LAMIRSE

A$..NEAE SPP ADULTS 4 1 2.51 1.S1ACA ;N&A SP ADULTS 26 3 14..5c 11.S3

rOST LOOA

OSI..DDA 5CADULTS 3 1 2.04 1.41

M A, RiMOSAROA SUTE ADULTS 10 3 57.04 '6.S

IO S9 AGULTS 72 21L40 2.41

P.D i'M'AAMUS CONFERVICOLUS A.JUL TS 45 5

SALOZSAE SPOMMIP .54 .5

OIPTE,A0a $TEV OR1 ,PP PUPAE S12

tPMS.C,'QJ2 AE SPP U.AAA I 5 1IC!LMO31QA.E SPP AET .4 12

P I 4 AMj

LC TI OC~f ARPIATUS AJULfS I s 5

TOTAL 891 108e

Table D-43. Number of animals taken by AN (non-quantitative) at fivesampling points in a tidal creek of the Sedge area, 21 July 1978.

A E£SEDGE SEOGE SEDGE $EGE SEDGEt A PilE A AN AN AN Al. AN03 3 03 3

010 02 31 001 0401TAXO% LIFE STAGE MEAN 4S1INVE41 il&TES

562 rA.j2AN.,&IoA SOP ADUJLTS 119 6 6 '.1 36.64 (..7

EnEREA P ADULT S 1.2 .bif NEKAE.OOM2.AT10DA SPP AOYLTS 63 Is 19 6 10 26.21 27.00

POLIl..AETACAPITE..L1337 SPP ADU LTS 1022 5 '0 4 247 263.14 390.14NE-NI0-ES L2HNI;OLA AloI 1.01 2.04P15.10 SPP A .ULTS 5 6 104OaIGNIA FLOR40A AU)ILTS 145 ? 6 30 37 53:.01 67:11

OLISOQ.H.1(T A;L.t.0C'4AETA SOP ADULTS L111 30 86 29 341 320.64 11t.311ASTROPC3A1

AL.EKA 3PP ADULTS 26 1 to 25 12.61 11.31

AA0NEAE SPP ADULTS 2 1 3 1.21 1.20

ACAOZN-...INA SPP ADULTS 6 66 11 10 14.64 16.54

CS16.CJOA SPP AOULTS 1 6 1.02 3.13

CQPO'.CA;i..-..I.OLA 'F' AJULTS I 2 6m.h.lACAC010A SPP AJiUL FS 27 2 7 so 32 Z3.61 17.,:

C A...AAE SPP ADUjLTS 1 .21 .62..UM 54.A;J..CEh Sep ADULTS 2 .651 .5)nE41.EC,A SPP AJuLES 7 2 66 14 56 2644 21.54CU.ELLA SPP ADULTS 31 .64 t.21

;NAMOSPOIAEAO,2 LUTEA AOUILTS 5 7 52 29 295 77.64 110.03

, "" L QC.5 5'CC ADULTS I 2 6co. .- I1j' 50pp AJ.LT3 109 10 36 61 30.02, 3601i;;0A1IZEA SP AUQkt tz -a 0S4 '~l00 C.GNFERVICOLUS A uL r~ 2 £ 1 .71.TAAIAE 'SCCS AU0.1 it 2.4 1:

~..ASUS GREGOkENSIS ADULTS 1 .22 .621140$ CTA

140o. _ASPP LARVAE 1 .24 .60.OEmIPO(&A

,E.,,.FA~ SPP "Y"4PAS 2 .44 .81'£,PTErA SCC A0IOULS I zt4 .64i, LU AEN ULo H! :I.AP.,L4.1OA SPP ADULTS 91 44 1? 9 10 Z7.1 1.50

COLEOPTEPO;JELCP((A SCC LAR44E 6 1 2 12 6.60 6.60ST..P4,o.N2AE $00 A UVTS1JSPT"I IAE $' F ADULTS GZ

LPT1EOO SPP PUPAE 1 .4 .52

j.'fERA ',PP A~viiTS 2 2 3 23 6.04 0.&IEP .1a IC C. A4.LS1.2 64CIC. SPP LA4 AE 6 1.20( 2.4)j 0';.I3P30104( OF LaONVACE 1 2.:0 2.20 K..PCIZE9p A3Jo.TS 2 ,.20 .6CEAfrU0.I6C£i SOP ILA4VAE 30 1 6 1 5 0.601 10.04

AL..0 Is 1 4 .644 ANOM E~ L-4.0S CU2D S', 4A*OJLT 2 64 A" 'A

GATEO) .US A;ULEAtUS AJOJ4TS 1 6 z 41 2.64 2.22

7071. M0. 1se M7 Z*? 126.

~Ii44

o .0 rAdAdamlIa. 0 OS - Add OC.Ad b.~Cl OSIA 0 4 9

- d *W..Ad OW Ad ao Ad Ad - -

I----------------------- -- hi 00.4 0 ~ ~ -

.0 V*.0ft. 9 Pd - S -- .0.4AdC

C)C)

- S.' A. 4 - .4 ftdS'4,09AdAd .0

4~i - -44

- S.. 9AdAd4.Ad.P. 0 .4 - - *NOW 4 .0C) Ad Ad

- A 4 0 Sd '1.0 -4 - IA - Ad

- .4

~-'.4iC)

C C) A4 '4 b45054b4A4,44 (4 '4 (4 '4 '4.4 "WA V4'4'4'4 '4(4 CA Wahi 01 '4 4

hi 4-i.J4~d444 ~4444.i~I(~~4' ~ 0o .~ 2 2ZS~Z23 -~ .4

z z 32'2244'-C 4 '2222334232.403440 03

- 0

~C) - 4 2

2 hi '4 40. .J 4. 0. 2 hi0. Z 0. - 4

0.0. '4 c 0 .44 4 3~ 0. 0. 0. 44 Sd 4 4 (1

.0~ 0. 0. 3.4 0. 0. .4.4 54 4~fl ... 4 0. hiSs 0.00. 0. '4 0. .. 0.U 0.2L40. 0.0. '4 0.50. '4

0. 44d.440 4 0. '4 0.4 4~ 414 4 4554 441 '4 4.0-- '4 3 0. 2 .4... 4.4 '4 . 4 00 ~ hi

~-.S.40.hiv4hi0a4..hi 44.1 hi .0 0.44d - 0Ad.5A.'40...0

4S 2 424 0 240 40 hi 0.44 - -

.4 *'*4Ad0 .0 CS.. 5420 0 .40*4 - C .. 554,004.4 .450 5 44 hi t.4t4 4(44 .44444 40.4 0. 44.. U '444.4 .. ~..4CdL.d.444 '40 AdOS 4442 45 .4. 3P...se.40..,O...3J...S.,5..d

4 4 4AdL4.2254 4- 0O.,4Ad...dOAd,.hiCAd42 4- CAd45 .j .0 t* t~4'4.J4t 4(4 ZI 4.4 44 2~ '40)2 4(444 .540 44 44.44 fl CA4 44 (Y44211ThAd 04 .45) .4 L.a .44hi 0500 .. 044t 0.20 00 aCT... 14 .54

I (C OLd %J4'40.0.0.'4 '-40 00 441 44 .40.4 0.54.4.0 &504b.4Ad'4.404'454Ad 4 4 ~ ~ 0. ~ 0. Ad

0 4 4 2 4 '4 4 hi 0 Ad 0 4O C0.(d0. 040 0. 0 4 0 (1 Ad - = 4 (1 0 40 .1C) ... t.. 4 - 41

44..4 44 4

~4.4

zC)

? I..*1* '.)

C) -~

-~ 4.4

~1~

Table D-45. Number of animals taken by drift net (non-quantitative) intidal creeks of the Sedge area. Site 03 (at the mouth of the largecreek) was sampled by LD for 6-8 hours through an ebb tide on 16October 1978 and on 26 April 1979. Sites 01 and 22 are in smallcreeks and were sampled by SD for one hour on 6 February 1978 (site '01) and for six hours on 26 April 1979 (site 22) during ebb tides.

A10 0 G SEDGE SE G E SEr.E SDGES C 1 03 L 03 so 01.s 2

.AMPLE 010L @102 0001 0101

I A A 0 LIFE STAGE REA8 'iso I

tmIvERTE3 .AT0ES

C".NlAR1A 5CC ADULTS 21 S.It 9.03

.I E..ILTP O S 12 3 3.83 4.9

POL ICoICTAC AP ITEI.LZOAE SP P AqLILS 2 4S5 3 L2.$1 19.61,.E;. 41" S L I ", I'OLA A UL I S I .3 *3"OdS.1A FLORIJA AOULTS 5 3 2.03 2.1t1

0,1...CAETA SPP ADULTS 23 90 57 4-6.01 J8.91

GASTR.PC34J. .TQOPOOA SPP A IULTS 5 1.41 2.23hAL.ER0A SPP ADITS 1 ac .31

, A N.EAE SPP AI3ULTS I .21 .43)

A INA14 SPP ADULTS 40 13 1 1 460.0t 62.31

OSIAI.130A SPP ADULTS 3 1 1.43 1.21COP IP00

C A.AN.0CA SPP ACULTS 2 133 4. 04 5.1HA.3 ACIACO0ZDA SPP AJQLTS 25 35 15.01 15.31

,nhSIS 3IERCEDIS ADULTS S.1.4 .53CUlACE AAE3I1333O SPP AI3ULTS 1d 033 1 213.01 358.31

CLIPE664 SPC AdULTS 1 30 1 3.41 12.71

ISOFO. A..N.AIplSPU0E30NA LUTEA ADULTS 2 13 3.54 5.51

;33.33SeAUULT 15s 30 270 40.361 110.1343AM;.UEASP IIO IULS 1 3 .3A1n, C4£llOlAbLi CONFERVIC04us 0JUL15 1) 5933 20 160.3( M5.61

3O..iPODA SP; IDEA 21 5 16.3 0lOE 43D.A SPC OIICALOPS 1 6: 3 .33

.4 4 D 0E.OCTRA SPP ADULTS 3. 1.04 I.?)

C CLEO 3? 10ACJ,1E6 EA SCC ADULTS 1 .34 .'.l

T11CV34PTERh

Pl , L IDAL SPC LARVAE 1 .3( .1

a 1,,* C P AUPA 23 50.3 100.,

;N& :CGONIAE SP LARIVAE . 31 41

C LOIOIOAE SPP L AI A E4 S 2.63P 5VC..OO..3 O~P L.AV A E5 20ILPULIJAE SPP LARVAE 1 .31 .331

Ft."s S3.C LARVAE 1 21 S.53 9.03

LEI'TI.OTTU.S AR3IATUS ADULTS 3.83 1.33

TOTAL 164 2101 4.04

........

CIO0

c0 C0 ~a0) ~ .. ~ C

SaC)I 0

c rz

CA x0.

to -c-)

a ~.-3s0 .

00 5-. 41-~

73 0

a))

~00-1 a1

Cj 4.aa

Z - W-WUt -

Z %

04

0 UOtl%

0.. In O _NC rr F4 4 4 4 CA5

.40. 4

44 IA .4Ut .9

0 -11U)7

uN N

tAA C! ina 0C U% a9 * .

u- -4 -

w ku

0 CS. a- n. 0 C3 .th 0. . .49 N.

CL C

.4)

IA IA- zA 0 A S 0

c4 LA No N~ w t .4 QUt x LAm0wft C u iXo

- "

C - C-,-400

a *- iii i-

44,. ,,N4. N 0 C

,.• C , ,,.o °- S --SoC,, o. a . S : -. ..

,.13 In-

a N SO .- j S ,N

-4 9 - 40 N , - .o - -O

" ' . . .4- , N ' " . . .. i . .. .i . . ..I

0~ _j

0. mN -n -,

t, C,.4 UN U%

O.j

C 0

V), -U.-

fl N

L/Lr%

C/) UJ a.. -,

Mw) LA J

C3 1. C

14.4 u .

37

"Ut)

00U C1J ao L C

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-j z L.t CA -

a--- 4 ZAJ CA.4

APPENDIX E

FISH SAM'PLE DATA

Abbreviations for gear used in this appendix are:

LS = large seineMS = medium seineOT = otter trawlSS = small seine

Table E-1. Table E-2.

SUmpl., Mssit . Oi

Area I (Low Sand) s..pI., Olol 1 40lSampler MS (¢nnen reus of *4pts

Site 13 - bt .2 Vl tihSample 01

1 Apr-k 1979-

Habitat Low level marsh t -. € .-c ,

Date 17 November 1978 t 27

Fork Length (mm) 32

3 1f30 2 s

32 1 3633 2 3734 1 3 6,35 S936 71|

37 338 639 94s40 56

41 5 49

42 8 0 143 9 1144 6645 1746 S9

75 16

Table E-3. Table E-4.

Saeele rSArea 3 (Sedge)li

f... Sample

g ISite 0123 2Sample 01:97 1 Habitat Low level rarsh 7

Date 6 February 19 "-

Fork Fork Length '57) t,_"A90 I 9

22 I9 2 3 2-3 z 1; 4094

SS 1 43

6061

99 ~ ~ ~ ~ ~ ~ , _-,__________________- !Il

Table E-5. Table E-7.

I S S ". } -I Area 3 (Sedge)

s-, .I.1 S~ll)ISamip Ier SS0s,. 2. X Site IS

SalO 2 .32. 3

(C.O". r~1~~ 3~pI. Smle 01

13Habitat P

.6 PW 0-9

8.11L ~Date 18 Spebr1978 ,L

I8 Fo rk Length (mn.' r-

29 14 3

3it16 238 21 1739 18 349 I 19 1

r 43 I 20 1'4 4 21 1Iss 24

43 2 249so Table E-8.sl 1°1 1

$2S3

543 Area 4 (Immature High)

s5 Sampler MSs7 2 Site 013s IISample 01 j

Habitat Pan A

I Date 7 April 1973

193 IFork Lenctn M'.7 I n

24e, '14 8 1

266 S!5 1

SS606276 1

Table E-6. Table E-9.

Area Z (Low Silt) Area S (Mature High)

Sampler M!S Sa. plIer NISSite 10 Site 15

Sample 01 Sample 17Habitat Pan w -Habitat Pan

Date Is September 1978 -] Date I November 1973

- FOrkL 'Lngth ( ' -" '

Fork Length (mam)

10 1 31 121 33

22 1 34 125 3 35 11

26 4 36 427 S 37 1

28 3 38 2s

29 4 39 1830 1 40 25

32 2 41 1033 1 42 8

43 4

44 145 2

46 148 1

a-I II

Table E-1O. Table E-12.

Area S (Mature High) Area 2 (Low Silt) ISamper NS Sampler MSSite 1 . Site 02Sample 02 Sample 02Habitat Pan Haia Smal tida cree

Date 12 April 1979 Date 18 September 1978 -:I4 .:-. .4!

Fork Length (mm) .Fork Length mm)44 1 20 145 1 21 248 1 22 2

So 1 23 SSl 1 1 24 152 3 1 2S 1453 3 26 14S4 1 1 2755 4 3 28Sb 2 29 1257 3 1 30 a58 131 S59 2 32 660 1 33 362 1 34 1 S63 1 3S I65 1 36 1 1

•__- 3 8 1' 391

40 1411

44 176 1

Table E-11.

,'..2 . I Table E-13.S~i. 02s-.,,' 'L,1 Tia -r~

oa. . 19... , - Area 3 (Sedge)= 2 ; Sampler SS

SSites 01, 02i," [tSamples Ol, Ol2!• "2 |(Combined results of 2samples) ,Z} Habitat Small tidal creek .o

j4 Date 6 April 1978

27:s Fork Length ,mn)

S 223

ea I l

p-25I34 38is 4631 471 3 48

1o~ 5;.

4.47

11

51

so

-411 I I

- - I

r.fl s a, -

- !2H

. . . .. " ... . i .. . . i " "!I II

111

-L

A 4

Table E-17. Table E-]8.

A".9 3 S9*

1Z 6 ii OS27" 31 Pil I

Sit I2 IIt

No 1-": 92i -! t mt2

2S136 1016037 * 6i'

4 1 is 605I3 66 I644 2 7 II

460 60 1

37 I 7

59 72

SO 637

60 4616S

4 66 S1

676 0 4I 6 11

40 11

04 772

78 IsI 8

60 1 7

20

04 i 90lOS

so 2

114 2 I

997 16 II

73 9 19942 I194

60 1 1105020

1097

12, 1119

131 1 ts1I1

1 94?

2 4

I's_______;1__

Table E-19. Table E-21.

s4 .r 3 (S.0Area I (Low Sand)

sc. o Sampler MSS.~.r* 02Site Il

Dat 26 aple 01S .:~, pH bitat Tidal flat (sand).

F.r L.2th' -- 1 7 Date 7 February 197836 I

6411 [ Fork Length (mm)4S 3647

4142 I

34

$s Table E-22.6o42

44 Area I (Low Sand)6s 2 Sampler MS

Site 01I 6-

Sample 0269 Habitat Tidal flat (sandy)

73 3 Date 3 June 197875B I Fork Length (,."l '. -.:

lo 33

309 35

3235

113 45

s 46 1US 4so

1151

13S3

336 36

Table E-20. Table E-23.

Area 2 (Low Silt)Area 3 (Sedge) Sampler MSSampler MS Site 11 t!Site 23 Sample 01Sample 01 Habitat Tidal flat (muddy)

Habitat Slough Cshallow region) Date 18 September 1978

Date 26 April 1979Fork Length uI7

Fork Length mmi 11 1

28 1 2.13S 1 22 136 1 23 L

39 1 24 440 2S 6

41 26 1042 2 27 443 3 28 8

44 3 29 8I4S 3 3046 3 31 447 32 148 1 33 249 1 34 350 1 35 IS 2 37 1S2 2953 41 I54 1 52 1So 60

60 1 67 36873

!! I I I II Ii , _

i

Ii

Table E-24.

Area 9 (Silet Trawl)Sampler OTSite 11Habitat Tidal flat (muddy) 'i

Date 18 September 1978 - - --

Fork Length~ timm) W-F X

65 170 175 177 179 1 180 283 214 1 1Is I1871

8 8193195s 1197 199 2100102 1103 11061113

11 52 1116

117119 1

124128130 1133 1135141 1144 2S O 1150 1165 1167 1182 118S Ii

193 12011218 12431

t

-UJ~~J -- - - -- - - -

- - -, - - - - - - -Z ;

diai

, , l '"S

Oa41 S "

,.EE

- --,

"" J d I '

,PU - -

I]

-' ,

Table E-26.

Area 9 Si let: Tra'd

Sample r UTSites 12. 1s. 16Sample 01 (each site)(Combined results of 3 samples.Habitat Say' channel ). 0o an ~ ~~Date 18 September 1978 0 - a 00a

32 1

53 1

62 166 16870 179 1821

9091959798 199103107 1Ill I

118 1120 1123 1124 1 1

126 21341

135 1138 1144 1146159 1171 1181 1193 1324 13844!5

APPENDIX F

FISH FOOD HABITS DATA

Stomach contents of fish captured in marsh and bay channel habitats.Each food habits table is referenced to the appropriate table in AppendixE which provides species and length-frequency data for the sample. Meanprey volumes are shown for all fish examined in a sample (excluding fishwith empty stomachs). Means shown as ".0" represent values <.05%.

Fish species codes are interpreted in the following table:

Family Scientific Name Common Name

0301 Ammodytidae ,t.nodytes hexapterue Pacific Sandlance0401 Atherinidae Atherinore affinia Topsmelt0901 Bothidae Citharihthys atig-naeus Speckled Sanddab1601 Cottidae Lptocottus armatua Staghorn Sculpin1,02 Cottidae Enophr's lison Buffalo Sculpin1603 Cottidae Scorpaenichthys marnoratus Cabezon1604 Cottidae Cottua asrer Prickly Sculpin1605 Cottidac Jttue aleuticut Coastal Sculpin2201 Embiotocidae Cmatogaeter 7:?reaata Shiner Surfperch2;02 Embiotocidae PAwerodn )uraatus lfhite Surfperch2301 Engraulidadae Engraula mordax Northern Anchovy2401 Gadidae .V!cr," ius rroxzius Pacific Tomcod.501 Gasterosteidae Auorh,nc;vi -avidus Tubesnout2502 Gasterosteidae ;iateroate:,3 juieatus Threespine SticklebackZ901 1 exairammidae edh- n e'arus Lznecod2902 Ilexagranaidae Hexaqrrn s s decagrtmnua Kelp Greenling3401 Osmeridae HYreesua protionua Surf Smelt3901 Pholidae Pholia ornat7 Saddleback Gunnel4001 Pleuronectidae F&yhr/ia A.-Itua Starry Flounder•1002 Pleiironectidae Partphrys t'eru~ua English Sole4003 Pleuronectidae Psett'ehtha: melaoatictus Sand Sole4401 Salmonidae nclorhuwhe ke ! Chun Salmon4402 Salmonidae Ocorh nhue teshawstacha Chinook Salmon4403 Salmonidae o iranerit Steelhead Trout4801 Scorpaenidae Sebastes srp Rockfish sppS301 Stichaetidae L.penua sagita Snake PricklebackS401 Syngnathidae Syngnatha ieptornycaa Bay Pipefish

Abbreviations for gear used in this appendix are:

LS = large seineMS = medium seineOT = otter trawlSS = small trawl

Table F-I. (Reference Table E-1)

fK : . 3 3.83.~ Iu.i. z9 9.1 "Id336

L.A4 J 0 ".- Is. 29 .

PREY NU Ol 03.. I %6.39 6 l I 9.10 VOL X. NON@9 VOL Z 0.196 VOL X '.1.0 VOL. X

J-e. 112 MD9 61.9 A0 53.9 NO0 19.0 NO 00.1 NJ 19.0 ho 62.2

P-&4.L1L . SIP £23.83 2 .4

ILI.2.2.'L..E SIP 003.03

A' - SOP OOu6TS

..j.TAC..0 SOP VOU0% I .

'.40TCI0SOP OSL...i 1.16 20.9 11, 13.5 Z61 19.0 &.0 1.. S1 11.'. 22. L.6

I Ito I P.(A $10 LARVAE 1 1. .5 29. 0 2..2 1.0 2 .

09, . SP AQ3~..32 .

207aNAO6 SP 4O.S .1 a 4. o 911 . i..

a ".ODLVIJ6( A I 1.2.0Is21.I,.1

A(09~2010 '.P 1;962 ,. S.P 13.2.s 1 4.1 . 2. 97 3 .

Z.000.lstA SIp I.;0. it V1.'.1.

Table F-1. (Concluded)

S-IF9 SAV

Sp,; FP.i Z542 2502 2162 2562 2;4iSP.Cil: , 7 1 9 1 0 ti

L:~ a9t .n1.8 5.3 139 6. 15

AC IA I I

0: ?14 51

aI.Gc;A~f IPP X047 3366 95.6 X7.044.1Ik~ dL NP440

.3 rj a AE A4

zhJsfbCiA I'P ADULtS .1

2S59.-11'A ISP DQu..8S 6 .2 2 .3 27 .. 7 .?

nAtacTSo SP# L9..27 1.0 38 1.1 46 is.i as 2.7 V.S

:I.9P1, '

;Ppa CE&4 a j

2J4#LA SP44 AuLlS is 0.8 2 1.8 .

644,,A16,fa SOP 0004.71 18 F.5 2 .2 8 5.8 91 01.6 4.2

£..1. AP ILu.T t..4

4)1T.OL 1,6 Aaua.To .

3z , ta app LARVAE s .0 .4

.'l Fa I p LAA,&C z2

!4;~07P.rfl:fLOA( 169 LARVAL I L.S 7 .i a .2 36 6. .1GC"68LW0401 P LAWtVAC 1 1.3

Table F-2. (ReferencL. Table E-2)

4 4 . 3i18

i js,0 (' *J1 3 8 1186 3.6 65. 801oi. S f7 6 6 4 3

PRE I69 MUNI . 13 X u.188 VL X "U411 VOL X6~ 6 40 XO NUMB4 40L1 1, 8 9 31 0

osr(.f:t3 NO 05.6 N0 6.5 %05 6%.6 NO 65.6 "a 54.9 ho 32.4

lv4E"ITOOA

149850710 199 aoUU,5t04.

4T88fM1 11480018 JuL11 1 94.1

7601&6U--1E

.40CT 3. SIP a-tn.7$ 19 240 2 1.6

.ji'06& &Pr AOU FS

4540. .uh SIP L3ULtS $ 68.6 1 7.0

;PP D4 : 1 2.

J16.O4;I 399.NLF 18611 . .6 2.

178&.01. 1C*t0 uM560 6 1s. 13.6.

.. p),SP U' E

SA uLS1 453 &. 0 2:6 .

Table F-2. (Continued)

1S1:F.1,83 1

l LtI 5 169 1 69 01 60 ~ A6:' 9

31. STATE & 3 4

PRE I M6ING V34. 1 %U%6 VOL Z HUA& VO. 1 WHO1 VOL I NUMS VOL X hNgM VOL X

1hF'11. C 7.9 go 48.6 No 100.0 go 36... No 16.9 No 42.3

$014C610061Q SF9 ADU&TfS

FO46T FFusa

OAt4IT.(. SF9 ADULTs

I ZO403 A CIA

*J44 1910

L*4', NSu6 A509I Issau 4. LrS.

al.'p.. 59JIQWIS 1 2.3.

OLFT1A 3F FJP6ILZ 1 6.6

50642.,SP ~

tts

Table F-2. (Continued)

140.[ 916£ 4 1. 1 0 19. No 1S4 17 19 0 k

1-4 .3 .. 1 1 304

073 ~.3 4 1 I 4 19 1.0.2 9L 3*3939 .4 33 .9.3 S

3R.1 :LrE 4 0

13E13 1 43.3, 43.r oroloS

,IAV 339C3& SP 439.7364

41.34704 111.

ta41FLC , A~ OL

T" A 1 OA14 SIP £001.732 91 0 . .

Av..6043 1 311

U . I i CEvL & 199 O W

394 1.6A0 O ULIS1r' 13 3 .A Z2 ..

A11. SOP.4

.1376. A 7 U13,

413P114SPU.

Sf140 M9104 099 ~ t ASP401

Table F-2. (Continued)

SaP.ES, b.1 16&301 .:

81 24 28 24 22

Srll' F.LL 4 a89 94. 43 18893OSV8 A08 M-3 88.8 1,a.4 91.1 6.60 4.8 27'.4

PREY PLANS V36. I NV48 VOL X 14WAO M. NU644 VOL X rNUAS fOL Z NVd VOL X 2 A

No8ECFi Z1 2.8 00 61.11 00 42.9 ND 128.8 0 PA 8$.7 N 4.3.1 A.1

444? Ipp t

M8.,~. r8 8u.S .2.

P. . T SPNIOL ADL1,12 1.6

:1. AC8.8

28i I S8 28 $62 35 1:5

.A AV8

." S,' Aou.As 8 .3 9 4.8 .

40 ).883 5.2 2 S.7 3:1

Pkt16..12.. COFE8 I v 2.( 2oij 88. :. .6 .'4.

L AUOS PP 2a .2

31'prf*i $2.6

2802184 ik $G30l LAN842 8 8.

Table F-2. (Concluded)

itl! I I

PRE AWA4V,91 $ 4d691x I

IN L0 iN.163.jIJ "lull Ni*. va .

Ali SO44A RUt rOIRIOV .L

IRE 'PT~ V3

RA. NM 59 I M A

Table F-3. (Reference Table E-5)

SP E8 C * :400 MES al 1 1 60 8 2 5PIKN Ll. 46 sin so 4.8 06 S2V. .,.,I VO 9V &.E a" .8.. 1.6 1,: 1.68 1DIG STATE p a 2 2

PatE f RUN6V.S 069 WA VOL X 0*M VOL X RUNS0 VOL I RUNS 4OL I "UPS VOL Z

W805(C:F113 go L1.8 10 &1.4 RD 15.8 RD '13.6 No 08.7

8'19848(tSA so9 ADULTS5

ISTAACOIA09 6~

"64980t160136 SPIP ADULTS

CUR80(2V(MILIU:04 59 ADULTS 8 .1 6 9.8

04T80PTEX&00801EADU9hOLTS * 1.4

3IPTE6A 199 PUPAL 20 ?.a Z2 1.4

*Or~o .0 3 p, 9496A

FORCIO 1181 ADULT *8.1 16.

S9(009(8 &be 16 -. 14lt Cs KRN F7 1 96 &a

60.6 :O$ w MR3 1!, .1 %.a .aot18 STATE 1 5 3

:.IT 0.680 VU28 04* VOLt IRUNS VOL I 84* 6 1~ REAMVol. %

1645P(CZVZ40 40 58.41 go 86.4 Re 786 MIT 100.0 5.

INVv(07160471

AOIAAG81

a4 1020 39 a .16 3.6 1 .40.

SI PICR*34(6 599 &0184. 2.2

;4qh?191*16JAA SF9P LARVAI a1

psVc800L.A. SI, PUPAE L .1 .

F6MI"1 SIP 0091E6*.

'00026419 OIS.

P1CCTP90 6AVI .

Table F-4. (Reference Table E-5)

A-L** s SEZO

4461 hoot .261 601 430t

S t0 l. fL x 3 6

144~ 05 0.8 143 3.0 No 2.0 %a 61.7 NO 9.9 .0

C' LVQ pp IaLT I. j

01"J S70 LOULrS So 64.3 't 4.0 164

f42 00 300 PL A 30.0 o 3. 1.:5

CALLI.MSA 3P Ag Is a96.

I, Pua - - - - -

Table F-5. (Reference Table E-5)

1 u LL 'I 322

W sV I 11N VO a IO U* 4 I &WS06X AR O

#4 ~ ~ 3UI 1.4 MM4.2412 2

~~.Stat*. ~ L

,FlACASP £3ULIS 1 .

40.AAIeSPAuI1 .VAPQ.S13 SP 9 8(JuLrs

;38I1,-I.LA see £ou~ts . 1 .

£&4I.1811 S0' LouLTI 0 .

.4) 3, .to t s. A

;o:3PI4 109 6.u9t MO3.8 1 8 % 26.8 of .20

~3~.oll ,V8 IP LAMEVit 1 .

swr..uLG* S ooLt PUAt 13 4.6 32 318 19. 1 .0 to 8.M .

orv.Io',,fa ( 9 5.420 AV~t 1.4 24 2. &.8 .

Olt KI,.AtS' Sp AJokmL 44.7.

*I.0 110410CH MIE LAMVAI 14 1.43 .8 2 2.

a- -. O.8 SImaps8 . . 1 18 3 .

Table F-5. (Concluded)

SF121 1 440 1 4461 0441

SPECIMENL 4 t 9 11.

1 2LUS VOL, 41--I 12.9 17.6 16.0 6.4.2f SITTE a 6 5

VOL X

1ff;rtLlrtEj go 1W. no i.e No 9.2 No tos. 428.6

INVEOSISB.ILS

£4,J II £6. 'tPALiC13 SOP AOuTI

COSI P .E A

COPIJTT. AU DLS.Jk £C14 SPFL A401 1U T 1 .6

.IAATTCD SO P ANILE ± . .2

;3 UL.L'4U0.F A

CJLEL(.MVUII 3FFf b4 .2

INLTkA S 1AAULS

:3L~tE, A SF1 . AIULSi 5.

011l.0 1VA IIL:A 2 .

PIK ..OPL1SE soFF LAIVIEL.1; OUD LAE FF AVC9 262 . . .6

SY1CwOO£TIA 5FF AsSI1L44&S 6 2.0 41 3.; 1.9

4IM&EUFAA""IENUPTESA 5FF ADU4.T f

,C OAt 1FF ADU06S 2.6

JOSIENVP E LAREAE 6 6. ? 6. 29.6

Table F-6. (Reference Table E-6)

lIES#1i49 E; I .?

79 49. IV21 Z1 9 2. 22

01; STATE 3 A

MUNIS d)61 X3 W". 409 . YO Mum8 VGA. x 9v9 40L9 IJ 920L X0 k" 9400. 4

4VpcP. 0 9026 6.9 No 916Ho5.3 43 30.1 No IS.8

0 ,ZIIC.ALTA IP8Vr .

0L~ftQHAA SP AVULrS 1 419 0 7. ..

ACAIF.VAA C ,IN&9 SPP houk.Ts 1I .

341:.C23011 11 AOUITS 1 .4 Z .2

CIICLA

,I.JI ISOP 32.1 .6

.VIopIIV~APVARMA .27(8 88U471 2 17.6

,...oSV P ADVLIs 2 15.8

P1IZ4 5 P2598 1 1.9 1 L.9'usL.V IS'tVAL 1 2.2

III'1 15 98 2 1.4

:14ATlsP.4391 88E SOP "849 C 11.41 12 1 9. 0 29 . 2 37.9 27 17.8PS, CPII.IAE SP P 89028 7 1.1.52L4.E SP LAIVII L .4 1 .2

2922 2962 2581 i982

74.4 I27 16 27 26

511 FULL I &9 7s 7902421 VII Z9.* . 1.1 9.3 24.4222 SATE a 6 5

p4EY 6046 V3.. X XQPW VIOL x VNlVO #6. 1 9204 84 z "CADMVOL.

I9ISPCLFtO 40 98.? VI 87.3 NO 69.o N 960 4.7 $9.6

2.sVE4TEaqATES

112VVISPp AOU.9S 5 F.1 L S.1 .3 1.9

GASTiDIPO1.0A 00k 28 SP AOVD.tS &G1.9 1 3.9 9.8

888619 SP ADULTS 4 .6 .2 .1

"HAIS:4A?113 IS" £Iu.TS St S.? 0 .1 S? ~..1 . 3.5 9.8

.EN 28V11148051 A S 242.

1105088.llAZ31P089098418 8124286.9

ANO-15002

'0IIPOIO SP 83842

3151868I O PiA SOO OU041 .

4,S21 S o 46888254 V21158 SPP LA4A18 13

a.1?.P..,9I81ISP 480814 .9 2.7 21 4. S8 11.J 21.3

1 pl.I08 a6 48'0 A98 3 11.3 A..

Table F-7. (Reference Table E-8)

ailfs 3063 356 2542 2542 2562 2sor511V1 2 3 6 4$YL&8nk 62 56 2 9M

"f3 F,4, z 28 Is IS1 $1s...J VOL ,I'll, 17.6 21.9 10.6 4.6 1.: 3.all STATE. 6 9 7?

PREY *356 90. 1 "long VOL x RUNS VOL Moo84 VOL x 4448 VOL I RUNaa VOL 1

.j 3t~ I. 0 5.3 53 69.6 NO 6.8 no 62.1 A,) .a #4 0.6

so 'r L 31.9tTILCAC1.AA "1P ADULTS M N T tt 14.9

CP41,00

AM553&INR4US CakFZRVICDLU A&ILI& 1. L.8 5 2.7 3 1.1 1 5. 4 1.7

2' PT FaAPT5(6 C 5 Q*A6 1o AA 1.6

'A 4L1G54R76

R..1 VL j 12.5

PRET SUNS V)6 0 MEANVOL 0

UNSPECIFIED 40 30.6 35.2

0.1 1OCHAkth3.Z.I 'aL05A SP AJ.4.1 .2

.6*AGATZC.jj3* sop sOU~ro .

ASiPHPOI* 3AG M'IS 50 3.0" '61.504A tns*1fl CCfFERVZC*U sJLaYs 2 1. 12.

l7Lmas I LARVAE .2

Table F-S. (Reference Table E-9)

tillS I a IS -i?:~z 20 52 sz 2

S~l~1554 12 3 58

l2L3S 135 44-l3 S. .4 22.4 8.8 9.6 t1aa1 1 a a a

Pi1 "04130 52 3 UNl8G 3 *6 VOL. X hill8 800. 1 MUOVO UNGS VOL X hIrMh sitL X

4SICZISO No Hg. 44 91.6 NO 91.6 No 9g.i No 46.6 hi 99.4

'Wt~idt II Is

"I.Its UP 8STS 2 l.a

4DAT:0934 SPP A83Tst 1t .s ii 3~

3 17(5. :11 flLMbS

CZWC0 ATPL4OAE Sip LAJt.. "QAO .4t. SP LAAE 1 .4 S 2.5 3 -49.0

net7t as# 8Z42 25ot ago2

C' 1 4 17" I2 Th S4A

314 1IAE J 3 2 1& 0

1344P(CZI..0 140 ?$.a No 43.8 No 44.1 No0 iit.O 84.1

* 0111 SIP A it.13 5 -.5 :3

C." jA SPP kut.I 1 154 ..

.r~a&TA&co~ in AOU..TS I54.

;"pFrF.j .OPS 83u.AS I 43.8 4.3

eD'4sG:A , I 0-I~ 1 4P4 71.8 Z.5

CWSGOI8 SPLDI a 7.5 3 .3 9.1

Table F-9. (Reference Table E-10)

SAO$. a 2

i16I1S- b1 2902 i681 1641 &bat 29?.1 2 3 a.

r, . 613 55, ?6 56403, F.,.. "t 68 4636163.41$ 42664L ii.: 6.6 .% 6.4 .1it s144 E76 1 8 9

98789 4SJ". 836 I 642 261 VOL 2 AWN8 40 . 1 h" Q 6Z48A 406 4 6686. t40L

NO2 10 60 0)) No n4.e Ito %.S N.) 0.4 M, v6.0

4oa l I t ij

A-:J)..4 606.7

2.1,31.~jT 049 1:'.

I vo3'c26co 2Ja,694 .271 £01.7 1 66.4

LIP, I*3

j1 6.flEO6 494 826211 9.9 D 7* 4 .? :,*

. 4162041.11" E46 .

'024,

7t~x Po 'I Is Z4

14 I'. 44br S.8. 39

S3Vj 40644 14 1 .4, . .ZROMNIS' SULL 1. I28 0

42q 277 2kk I GS 8 7

Table F-10. (Reference Table E-11

441S Lot 51..?~a

0 11*0 N 1. L No 9562 NO ?44 4 4 . ho 65.

~.,2u... t ... a. .OLT

01 . - "I 62 6 ? k. a1104 104 0 6. 5

&J.C~IA

ALA '4 A3U Y 4 F :

40404.4COL SA9 4201S 4 4. 3. 46 3 G

LAEAOJ SV* a - , ,

;.EQ11tt uAE S.' 306102 1 L.6 . .

4WP4L.OAL [" 44

:r51.'.IF $PP 4 2 3is5.

2.1 601

:L itLP11zA 16L6o6

1601 160 .63 61 52 21

50 61.3 50 6.4 5 2.6 2s 1.5 9t 23.31 1 342

.. C 1, "0 Sop AC IA A&

4CAV.PIA IPP I Is.3

h.5420 2ofRI0% 266 2.

*~~~As.334 SpP1..5

4444115iE1.,(4L 40'll I L4A 031

Co LAA-OL A~

Table F-lb. (Concluded)

A Z-.J5 VL.L -- 1 5. 125.031 701 7 5TAT

401. X

JL!2h2MD 5. NO 26.5 37.6

AU s7 33.5 1

2SINQACO.A SPP 5014t1 2 .

.JL'.41.2 87£3JUT$ 1. ? 2 .

.'3 0f3, M&EMA 2.5TE AM02.55 3.0

'ft~j34A%"nQs5 CONFEnVIL u aQUfS 262.5 1 21.61 1?1.'

31I0VM14 P5UAE I 2*.& 21.3

CE'ls LP PUU. 15 .

1.: so.a~as sn 28952 .P.VC4OO&E SOLARVAL .

TLP6al0at 188 L.456I 1 2.3 .

Table F-il. (Reference Table E-12)

SD~C. C 2 2 jA. zon 25 22 27 72 3.

;;GSTATE 0 S 7

WINI V04 3. 5 4040 VOL I 0044 100. X SuS VOL 404 NOiOL £ noflJ 4;

"aFfC F0 7.1 NO 69.5 NO 7.0 No 5t.7 No 7M2 No J3

Pj I.tL.. A 0tA SPp A00.75 5 .0

CL1 . C V 0174J.i .'~f S ' A3U 11t5 2 6.'.

1010 S? £001.55 3 2.4

i40. PP U36kfS

SOPLIN .4 .3

34434SIP A2U31762. 5 2.0 2 .0

.. "t. f41.71w4)4 2,? 4301.73 14 11.2 61 I3.. . 79 11.5 32 12.2 63 3.7

.. P.4SF? LARVAE1 3 0.9

i6 .1.P 4OA E &uot f1

4 p ~ Ik~~ ; 0.7 Av-T

7'. :761a46 S4y 4401.-5 1 1.3 .

t.lt3lo 1O RA46 .

AP.-II3.E 5PP 63035

'ta12501 SP4 PuA &E 1 0. .?. 1 .

3;L:C"0#G;2500E S.P p .'a

C'112062.23.1 SI9 1..3.25. 4.

7100.1361 3op L4.46 2 u.9 1 1..

Table F-li. (Continued)

aac.. L x 6Lt '

AAN.tS 2 ? .sS.:2

7"o a0 90. 140 663 N .. ioi.6 . 4 00 &.

11 T61 'I 12 02 6

aTt' A.& 1r~ 7 1 70&.757

St O ;179i 1. . 56 195 2.

0'PAT OD SOP 60.5f 6.3 60a 604 9 .3 30 3.6 # 58 6 .

P.4E4A SOP LARuL4

I . ; G. T Sn J3 LJ -i l0a

A .

*fl 40PeRCA SUP J £UV,1 346 0730. .

AA £45154S . 6.

4 JS66451 SIP AA JL 1 .179 3.1 1 3.

£144064 SOP &A011 2 .

410IIA t ISP *4.1$ 1.2 9 .

114 O I 2OL

SOPalLc-oaSUP LAS.A

*VQI43.VEACR L115 6.0. 1 4 36. 3. 1?.3?

*4' 0.A E P A.&.S 62 1.7?. 39 .

~~Zi1......... SW JxS6 3.

Table F-li. (Concluded)

13 1..1air; 36 1

t4S W': :4 M A-1 1 4. 9.

X.; STAtE 0 A

Ito 0 1.7 40 43.6 NO 21.7 41.3

Z2~7 74. A4

3431.01.4

A4 . "kI 41A-LT .

A41 1 ' PP4U.

A Sfl6 :944LI S 3.1.

.46$1~63*AuS612.6. 2 25.3 . 5.9

44f093.. No9 U305 .0 01. 14..

GJ.1L10 r9 AloS L.64:4

l%SECTI

1311 f 9 PWPAC6 .1

I41";At1 b* *I At4C0 . 1911: ; at ' A.dk(:100.1c341 111 L&v* .a a S.? C3.

tlPuLIQ.t SOP L0.6466 4 .3 3.

Table F-12. (Reference Table E-.13)

ht14 'I'L 1:0....61.F, ll 1",11

oiVl v It43

go E2.8 g "A1 146 4.1 no 1461 too 461 N 42.

SPCI"E 1 203 Z t.6

J'l.71(1FPAA3.Q TA T I0 3.5 40 4.4 4 49.1 MO 46.8 0 6.40 4.

*;..5 Sp AOUL tS 1 1.6

C~kVIEL 11.

,A6FAt fO SP £0" 1 I's 21 .

£ i (,;.j (N PUEVAA1.7( -- U,.3 1s3.8 91 2 4.

1f '1UC 3.T

P-E :WHO s'PL IaQ4 V L Z E k

1.34~N 2,3,3 CPA LAV( 6.3 1.4 9 .

s'r::. 2ic

;E .2 MM 6.4 4 .

Table F-13. (Reference Table E-14)

aift. ?5&.

WP:&s 52 2582 2502 Z5C2 Z502 23.Z

~6 '4 '4 31 4 i 39- 3.SC. !'J~,.3 too7 6 65 73

3L J .... I NN'4 .6 32.6 10.6 219 J5.9 £.

WOO4 HaA . X 98*64 #G, X 4.44 4L i NUMB OL Z 90U44 .0j. Z k6miK 420 ZS

. 1 F' 2 , 5.1 No £2.0 h63 65.4 No 43.2 ha 71.7 i 0.

£JUrs 2 .3

; 4"'.ZAe .35P *28*201 1 3.8

-3-78 4j.6t.

A .P3 2 .3 9 2.4 2 .

IT A 6 SAP ADULTS a7 3.5 41 5.4

~.t.cros.,u~r 2 .6 1 1S"4&?~23 5661 A 3I * .8 49 3.0 So 6.? U9S4i~o 2 29.2

G. ".-CRMALoEtAU T

A4*1 05 0411OL A0*20

PP lj3

100,

I PtlrJ-

*62 aJ~1MA4 C252%2CII A2.

..i:. at 5?IA

-US0'flO. .,A31

97SPA 1?AR#915 . 6 .

~t9..10.22ezQ.C s#P 5.28*226 65.8 1S 2.3 1 .0 3 5 10 92 36 17ILr l26.26£842* 2.

'aaa

A -. ,, -. . .'.' .d .C... . .... . .

p ,4C4 - ,. N .S

C, e .- C.."..; ,. .

- "5 -* - 4 -d.4

r Nx

4 .

Cd 44 - P4

.

Table F-14. (Reference Table E-14)

I E 3.10L RO

Ira. FULL II 0*

" L! vUN Wha.o I3 u sfd VOL X MUNS so.. X "Ca,VOL1

bl18100*

113OU

(*4*fOPCU3WtOC SIP LAA4 a 0.8 %J

Table F-i5. (Reference Table E-15)

S81 981 pik .

1104 FILL 1. 8 1 64 1. a

3l1. STATE p 3 6 S

4000 NUS 31 X MISUP VOL. Z %ai4@ 8. X NM8 806X110 VOL8 X MR VOL I0.

jhEsIIFIL No 35.8 gao 67.2 No 28.4 90 58.1 No8 11.6 NG 30.4

11A9.14E1N8 IMP A0ULIS 1 .

f.TO.,APaqA5Ir.C 5FF ADULTS 9

3aOCAESIA

111,1300, 1

ICA ST$I DA SOO

ADULTS

ART 61008811 .FI IM. J.JIMIE 10- .

.8 r." ADULT$S

CA.hAIAA svF &GULFIS if 1.1 - 1S L

'.IPI 4,13 "9 83..s It It.5 aI s.F 19.4 12 27.7 39 3.6 1a L4.8

Licrf SF10AULT

:OAPtaa0 S PF 1.8188 £ ;1 1.

11 OPL 40 a P0 :"I

.. of 1 ADULTS 1.6 81.

Table F-15. (Concluded)[

slit;. IAt got

01f 4 7 " 4 , 8

RE Y 1& 03L 49 RU&VLI NUN VO .UN O I MA

J152. CFL S 0-o I. 584.? n 648 1 4 Z.

ADLULOLT$* A. 1. 1 2 4.

Al STTE SO7DUT 4 4. r.

STA1tuE8 TAILAUTS1 4. .

041,ASt. AFDLULTS 4 4.7 .

ACA LV NA. CT O Ag r Z0

,OATA 5FFIQ AO ,6r 42 44as 4 6 ..1 2.37:; L)

84)484(8 111 01544. .

; , t,'.TA E P LAS aLT a 1. L 1.8

A1TOPIA SPP 40UeTS 1 0.4 .

I4414POA :wEaEA

: , F F A U L T .5 4 . 1:

:.P...OL. S P Asuao S a.2

C.A3ELA l F F ADUL.TS 14..412.a O', 310.1 S0 to'4aL .

VAAq AE SAP A C31 .

6L O-GXOe A S P kAJUVC8L. 1.3;o~..u 1F AGULT 5 4.5 .toot 7.LAVA

"11411191,8$S CNRVC LA " 8 1 3 6. .7

SIP TI4CT &F OALO A .2.

Table F-16. (Reference ;1 V-)

L~tE- .0

226SZ1 228L 2202. 2201. 2201 22012 3

S44 F6. Z I a I7

31; r0TE 1000 £

Ni 3. 4 4446~ VOL Z Nwva VOL X h.$Ev1 VOL X "LOW %31. X 11015 M. X

1No i4 I. 05 go 7.7 02 03.A.

Ie fI ) I U S

k3.14.13L.'4S6 SIP Aaurfs

6Si "."A SIP AgIJ~s

&1 4AAa* 0. E see A0ULrs

IIIA13 3 .

.,1.aPAO" SP AaO 4lS 3 .1

w , LO ICIO&S3 SP 0DWLIS

Cj4OCE.4 A 1.1 1& 1.74ILEU.S0 SIP £SuLIS

.,.3..AE01tQMA LVTIA ADULTS

S4.~P0IP ADUJLTS$0JVV0SLCS3£ s.i

^.0~.j* SOP AUlS 0IJ.

A ; 10APA0Ul c~tb.M Y ~U6Ts~j C2ULTS

400Q 1.0120.000 AoU6TS

I 0..*'UA SIP 0L35

5 1.

Aw1T SP ADULTS

C14600G600110AE SIP LA ht

PIS"

S S6~ WA01troC. UNPECIFICO

Table F- 16. (Continued)

;q-41 E E

1S"Esf alai ala1 241 Z02 1 &61 &6!1

17a. SUL.. x 1: -3 so I0.IILJS "~ i MR,3 4. 16.4 4.4 OS. 6 7S. C0011120 S3 3 4 94 a

Pq : v M74 VOL. 4 4.JWI VOL. X 240.. NU 44 4 06.UM 0VL 14A440L X M1148 00.. X

JNPS42;ON 4F.0 410 4.0.6 Nol 66.4 40o Z3.5 No Z.0

#4ACOAGUr

*C..f6-.A Svp ACU..TS

A.,E: .1.1 0 AQU .TS if S7.2

.s023 , 9 4A2.7 1 .0A 1P .1W 1 .7

.PCTIC.ZJ& SF9 DOL.7 S 1 1 . 3 R.0

4021 0 19 IJuL70 S2 Uf .0 1 7

23 Z3ISP-' UU.TS

Aopm9.OA SPP 2326.7 no0 7.21040" 599E ,2441E 9.1

M, 9 £51,.. M 14 L8.0 ma $6.1 7 12.134110.4,44j C0'SFRV501 A06."T7 2 1.2

J; . 9034ASOM ICP

57401 4301 &.IIS

Z 11 $P 6.274I

49.31,. LAA9 £34 97.

5110801 GE

2.64 IjUtfPC*CN .SCXVZS 9B.0

LAWAj'MI

Table F-16. (Continued)

IStS tilt 3401 4402 ..4 -3

1ltf N I1M3I 14.t 14 ;.7i: So nt tt So& 105 .

110.4 SiLt X 00 1 74 75OJt41 aQmNn*3 44t., I.: 07.0 7v2.5 434 23.3

Pill nvm flA . Z NUNS VOL. a Nn. VOL I MNN OOL 4 J 4 ZII -14114 V6 Z

.u.,PLCIP0LD go 15.6 940 100.4 NoD 100.0 NO 95.1 ma3 04.2 No3 3.0

0C10.1

-Ft'.4C5411A SF9

ALJLOA ADULTS

44A4080 SF &Et IS 3A A0-1

IAOJ SP ADULTS

A., a f ATSPAU S

,S1404XJa., SAF ADULTS4

40 1O"OL.. A Awl

KOITSIJAC 4FFADULTS 2 .

1049 0 ., 59 A ~ ~ 3n 3 1 . 0 1.

i 11MU. A9 A(OAI.NoFN

1J..Ct S9 4.13 I 0..Ao .. 1

040140 11 Lk04 -

1,10. 0.8JU SNJ L.

;CA ..A opO F ADUJL2

,,',,, fOOA.3 2.

;C 0rP 4t0(8 SAOF AAn 0ftihi(

,isI- -GI-60,

Table F-16. (Continued)

S310 C.1 .14..4G& 91 3!

I P( I Iz 2z3;

6~. I 108 44 a 12 &0? 3StZ :'J.L Z I's 50 41 3607SOL.174OL -M-~3 34.0: Z44. 188.4 6.d! 4.4 10.4

S11t7 A4 4 6

.91, NUNS V3i. % 3634 VOL X NUNS8 VOL. 1 4UKS VOL. X wOod 921. I L~ 464 3L X 1 04

No0YCI 9.*0 33 4.8 NO 4.3. No 34.3 No 33.4 NO 15.0 08.4

P73440144t00 SOP 83uIf1.

,L,1ADUpoL~uTS 0 .0 .1

A E .o4Uz.E SOP &3U03I 6 53.1 0.3

.CqA 3FF AOBL f 0 1.0

1&ADULTS. Z.9

48081 f0 SOP LA3h .1

I401~~~l.0 tubCA £ou IS

o.IJ. 09 42v7 No 43.8 1.9pp8., 59 .uvEN0136

A3Z1n F OtT . 3 1.:4 3 0.4 3 13.4 7.7&:oassmaconfERvzCOtU o.j £ 14 40. 140 .9 5.0 1 12z .7. 9 ?3-6 ..

SP '.01 Cr0

I'Stcf I , .r A.51I Srt SP ."A,1 .0

A 1.1106 SOP LA9t41A

S.fc SJF0RC 4 0pi00. 38*.

Table F-16. (Concluded)

"STf. 06 21 01 001 ma za

P, LAO dUMV 01 MN LX UaVLX % & 1OL Z0 6? VA 4 X114 J

goDC1104 3t.0 43 100.0 Na 100.0 N0 ?6.5 No 75.3

00.GY.nAiT SI D £00.75 NO 14.3 1 4.6

3.70*203.Is(r4431d WO ADULTS 1 .3

nIlPA fZCOSO4 000 ADULTS0 1 .2

1A1~~*SOP LARVAE 3 .4

"IS01I A44 OPp ADULTS 1 0.6

C 4NA " A

n1ENZ;hOm SOP ADULTS5 3 6.0 1 .

;.4001 50041*1.0 11 40.6G 9 4.4 9 1.1SSOANASCOS.t4VICOtu ,QLT 2 0.

;IPIERA*S C.00.061 £00-w. 1 5.5

SPIESIE3 2001 22c1

MR .51, NM 1 la1273. 5,. 01 6b.1

.1 1047 3 0

DOE? NUNS NDL X 0 U46 VOL I IAN0

i41PECIFOLO 00 65.6 640 72.4 7.0

*2?"AI IS AUL6TS 1 .5 a 6.5 4.6

OS?" AL04

f14.4SDL~f .0

ID,.OTLS3 SOD 3JfS 1 .1 .

.:10,1 100w he14 .4

4t4:4:a SOD 60u,.TS 4 1.4 1.7

4404 10 :3:oko&p.,n.0 432.7 156 00. 2 16.1 1.2~A 'I1;"AnAaJI CUNFEVLCIU D JL0 a.0I

':

01

;a

0-44

Table F- 17. (Reference to Table E1-22)

F. ~ *~ As 35 55 9a33

a SC anS3 so55633i.S1f 6 . 5 L95 . 6

N,0? 41, 2. 1 tUld YVu. A 'W44 ';L x NO86 620o 1, MUddo 40 0 u.08 'S'

JvPZ'Zl8 .4 69 26.2 ma 61. b0 20.2 NO3 L1.4 53 35.8

PAC..±Ti SPP ADULT2S

A AS:ADU..1S 23 5 NO 51.? 6 6.8

ETC. .0IT ..P SLu( 1 32.5

3, C tr2 SAP 80u'T1 3 L.6 5 3.5 8 .5 2 .8

5 t "...3) SAP A£~f

'PCTS:&:3 SAJaUL;$ 1 .6 .1 8 t .5 14 36.3

faa.8.aSAAOUurs 65 58.5 74 66.6 t 35.5 t5 65.G 1& 36.2 317 61.8

atV.aSOPJDCLC 2 6.8 a.. 7 2.;;4cE .. t SP A3 V6 r

2.51. SP 03OU.. 2 .

F.~~7 2S. 4 g ± 585.2 .0. t**31465.1 4 11.5 3%.3 1.31.. STaT $ 7 s

P8(y548 tIL I uuAd VOL Z 4a4* #I-. X 6a8 VOL Z NUMa 406. 0 KEAN

JSEI'FZ3 No ±r.o 4a 2%.3 N3 sj.7 No 34.7 NO3 Z.t.. O.

1. . sp 0 0 ! 826

A": SOP 53D20F1 6 .

0;.:A5 SPA2.1T

r . _.8A SAP aZU6j.&A25& 2 .7 a .6916.6 .

2.281. 1A.Au 75.

" 8 ' 8 1 5 f 3 3 8 AP u~ r S2 . u . 3.

"Mr5..-h

Table F-18. (Reference Table E-2S)

AA&P.tAS L

spCS29 I'l 2 3 4 5 9F.. tNI. Z.. 863 9 3 A1 L30.S 1. 9 90

4..SP(CF~t i4 .0 N. 1&.S N0 1.0 '.0 3.0

i0AtZQAE SOP A3U.TS I 1.

ACO AAAACO U

ZaICpL9SP 1.t, t

ar'..P1J SP JU73S .

fLA2". SOP Aj..t.

.IE-AE UPP Ajo.3S 75 53.8

:OANp.3N F.Ai.10Auf 1 u..f

L%jA&55 3UAE 959.15 I IS.S

It NIZNTZFLEO UNSPC1; 1 1.6 i.

1940z 5301, 2981 iLt :0 n

I~S L5 24 &5 44*.

312 STATE .1 4. 2 2

pu" UNS V3L X MUfl VOL X NUNS VOL 2 NSARI VOL X NUN@ VOL Z NuNA VOL. X A'S

No 2;.; N;0 3.8 No 10G.J NO Z9.0 .No a.

30.1

St * Uft3 . Rsea 7..

: ;AP: P A')~t 46IS2SOP?: P AjJI.TO 3. S

JLUAPtab 1.0 A.V

---EL,& A. -1 13..4-'5

- 4 S 84 *A .4 .4 -

.4*.4 - - - - - -

* - - - a - - --

* ~ , 4 .4 a SO

a C - - - --, S - '4-

a --

- .4 .4 a S S* - a

a 4. 0

- - C

- a - 0

a a , -

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S- - *4 C

a a

.4 S*4isaC * - aS

.4 0 0 --

1~' - --

LI) .4 Sc-i

I '4

C) S

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C.)U

- -, 8' -- ~ .4tui.~t4nkia0ais..44'4 -C) a J .JJ.JJ.J *. - a

.44.44.4 .4 C -- C .4 0 CC -4 .4 a CaCao C 4.4C CO a aQ aaaaaaaaa aS~c .40 a a

C) d" .4 a

4. - 0 5. 4.4 4 - a

.4 '4 4. 4.4 4. 4. 4.t.t ~.4 .. a A a a .4I - 4. 4.', - 4. *'4a &a, U a *

).L. a 3 .. a 4. a a a 444 ~ at aitt C-------------0 .5.4 .4.4

C) Ca ii. Ca ~.40 .. a 40.4 C COa.4 aa.4*2.3 a5t44 aa 04 eCU a .40 .4.4 54.14.4.4 aaaai a'a a

- Lv-.,.. 4CC .44 *44 CS. .4C.i 0.. 4.a4.~..a O4~.... t.,..a U a4. .4 .45 4.44. 3... a.J U.. .. 4.84..a 4.4Z45 44,.. .4

*3 tav4IaSa&.ia4.aa~ .4t4 .. ista .4 4.4.4 2' 24 04 4. C alt. .44.40CC 4.-..4, 4 ac5 ~240 *..4 4. ,aa.4taA.4 Ciii Cisa*,attisai) 0.4iiJi aCt a -5- V * .4 3% '8 .3 - - a a .4 - a

1~

Table F-20. (Reference Table E-26)

S.Ec us. .go1 1481 220t 146C1 .301 .8

74 ~ ~ ~ 1 aSC AM83 71 7. : 7013 auA 1 a0 a8 15 637.

5G S "VOL Is. 167 21.4 I.: 26.6 72.9 &d.441L STarTL I j 1 z s 3

PIYNUNS V)6. X Is i5 VOL X NUNS VOL 2 Mu4i 401. Z NUNS dOL 46.0d VOL X

jC'PCCFI.3 mo 14.3 no 45.5 No I9. No2 99.6 N3 62.21 40 66.5

~E'A430Ah SPtO P 07 JUL12

Is..A5P ppADULTS 4 4.2

JI A ADUioLTS 2 65.6

.f~31..CA SPUP L0u.TS1 .

~S W" SPP Ajw6.1 L 1.8 15 32.5 35 16.4 .

A%,$3.ANVJS C(AFIRVOLU A965,1 L. 1.6

A 7234 37* ..1AV40 2 24.4

1?LE: * 1s3 2600 4121 .001SAL't. a 9 1 G

.. a lit~MsS 1 322 $47

fL., vo1. ..4'3 12S.0 24.6 5,6

P84(1 NUNS 43. X 4a06 OOL C 04.NS 42, X h03N VOL Z OVAN

maP:1'i *0 00,0 C, 97.5 nO Z.a $q.'

*3.7~t 3.. .

2.FK 11OL U V . S

A~n~pUPa 3

:l00'z.. LI 6C.L114 2. 41

Table F-21. (Reference Table E-26)

Ai"P.LI 1

12 3 44

1. S1404. n4*3I S5.3 28. 3.7. 2.37.631w STTC 5 ? 6

IRE5 I48 NU .. 4) AM&~ 4OL X MoiG 40 ZO Nb0M6 tO., X NMAn tOt X 48 L 42Q1

No1EC1. MD 1.5 40 11.5 No N4. oM 7.0 411 4.0 No L.3

;salTA.:34 F8 43.S 1 3:8

3184.vla SP. ADULTS 1 L.8 1 2.0

ljnL.4 DPPAo.ts

;szn~ 1~ ~AlOji 1 3.4

£74.1 C"" 1..0AW, 1 .5 1 &.2A184i. Dios S2~S1 .1.112.AAatuS4071t10'u AD TS 12 74. 1 4.0

t8I~lA& 18$ADUTS 1 Z.8 1 3.8

4.40 444G&1

£731 'i A .1S ??7

J43.41 .. 3r' 454.81. 9.

1C~~I T.TI

No~CS~ $0 ".a's 17.4 "a 21.0 No 1.2 11.4

* .. ER4TE3441

E'L SO AO 7 1' 1.9 £ 9 .

"I3[.fA 0 ADULY I I

3.~iA SP80473 2 7. 1 80.0 8s 75..0 41.4

C _rt8CE A p ^

44~t 39 2MS, 1L.52 8 .0

*-A- 4-'

0 u0 0. w1 -0I , va I1 v I.. e I1 a

a 3 '0 . .4 8.- >1 4110 0U

.- - .14 00- 0 0-r.- r- :- N wa0.-000 .-Aa 41)4 w .-4 41O00 W - I Oa 4114 q, v

.004141.4. 49 0011

41-411 00 00 w1 *'0k41

j4 .. b a 01 Z 1..1 0I ~ ~ ~ ~ ~ W l-U4 411k-*I.U 4

0-l *.-00 0 1 14 I I .t0 . 1 00 0 41 ) 4 800 4 4 0 0 . 10-4

41~ .41 0 w 000- 44 . 0 0 00w-4 4 04 .0 41 1 M. 0 1 .w 1 . 44C 10

4' > 0, 11 w 1 8 I., I > 041 ~ ~ .0 0 40 41- . ~

a ~ 0 v- a1.N441. a0 a a4 4J.441 do ...C 10..-4 a4 4 k4 41 00 c 0. -.- I41aa . 42- 4f 2 1

.00 w1.4.0 0 0 . 4 .0'4

r . 0 44~~ 0 1 00 0 wI .40 ..- wa - 01 0w

0 11 41 0 441 9 -.4 Um 00 1, 1 1. 1 00.1. a 041 1.C v 1. w

a> * 00.0. a4 1 0 . .0.. a1 v 00.4) 04 Z144 04

k 4 0 041 0 .4 44I 0 04 M .14 4

00 00 0W

0 IN 0

41 V 0 10 401 .0 C V

rI wa M 1 k, C, ,. 01a0 ' . a.- 41 w 10.--. 4 41 a

w w w00 u a w Oww0kO4 M 4 0 41 -I.414 w 11 ;:7 .40.4w 41.1 kOj w ~ - 00 w 0 1-4 O a al- 000a l

OD4 8' 410 v4k 0 010 81144 440 .0 -- w .- 44 .. a4 ' 0.-- 0.1 -0.'

;10I0'I 1 9 1 w0- 4100 01.000k&- 2 1 1 w 0-2

w0 1,. *4

g. 1

.041,r.0

0 kl J I 4144 w 0 z .l4.

.0041 1a -U .00 1400

1 04 0 ' o -. 4 41 .40 00 1 1 0 '0 4 41C41,00 7k. . 1 00 101 Vkk 00k-4 0f0 %0 v 10,I kI 114- W- M4 1 0 U41 41...0 0- c 44 .0 4

.)0 0'.0 4 .0 -I, -. ac .. 00 0

'0 004411 04 C .w w1.C . 00 Dw-I c~ c 40-

00 044 1 01 0

41.40 400 411 41414k. 0 k N11.I4l 4 ce w 1.. r

414 0 4 4 10 41 0 I

41 41. 0.4 aU-.0 00 01 4.0--4 w4)444 00 .00 0

5 0 w u4 0)I1 -. .W .410 4 Do'0

210, .0 " 104 02141.0

0 0, M .. uu .40.k0 k>.. ... 0 .W 0 0 - E0 0 0 k>.1..'

>W04 k-41. 001 0 w0 w .404 kto14

' 0 1 .. 0 k41 ~ ~ ~ ~ ~ ~ ~ ~ 4 k io1-0* 1 01 ~ ~ * 4k~l

0044 11 k21 0004 A00U'.0 k 104W. 4 1 0 1U 0.4w1WOO12 04

,a14 .001 4 > 0 41 00.a - .0 k .4.0 00 1 k0 a-- I k. c 0 0.4 9) 0IM 0u4 41 I's4 a 410 16 004 .0k1" CO 1* 4.0c W 1. 4- we leW1 l0~ r 4 0 .0 1 1 w1 1 0. k w1 . . 4 1 a. f

.0 0014.44 414 14 4141 .4 CM414 0

11.

DAT

DI 4