NOAA Technical Report NMFS 7

The Feeding Ecology of Some

Zooplankters That are

Important Prey Items

of Larval Fish

Jefferson T. Turner

July 1984

u.s. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service
NOAA TECHNICAL REPORTS NMFS

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St., Washington, DC 20235.
NOAA Technical Report NMFS 7

The Feeding Ecology of Some

Zooplankters That are

Important Prey Items

of Larval Fish

Jefferson T. Turner

July 1984

U.S. DEPARTMENT OF COMMERCE

Malcolm Baldrige. Secretary

National Oceanic and Atmospheric Administration

John V. Byrne. Administrator

National Marine Fisheries Service

William G. Gordon, Assistant Administrator for Fisheries
The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary material

mentioned in this publication. No reference shall be made to NMFS, or

to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or imply that NMFS approves, recommends

or endorses any proprietary product or proprietary material mentioned

herein, or which has as its purpose an intent to cause directly or indirectly

the advertised product to be used or purchased because of this NMFS

publication.
CONTENTS

Introduction .

Acartia , .

Calanus .

Centropages .

Paracalanus .

Pseudocalanus .

Temora .

Cyclopoid copepods .

Oithona .

Corycaeus .. " .

Oncaea .

Planktonic harpacticoid copepods .

Copepod nauplii .

Tintinnids .

Appendicularians .

Marine c1adocerans .

Barnacle nauplii .

Gastropod larvae .

Limacina .

Discussion .

Acknowledgments .

Literature cited .

Appendix tables .

Figures

1

4

5

7

8

8

8

8

8

9

9

9

9

10

10

11

11

11

12

12

13

13

20-28

I.

2.

I.

Percentages of total number of fish larvae, total number of clupeiform larvae, total number of perciform larvae, total

number of pleuronectiform larvae, and total number of gadiform larvae that have been reported to feed upon various

copepod taxa .

Percentages of total number of fish larvae, total number of c1upeiform larvae, total number of perciform larvae, total

number of pleuronectiform larvae, and total number of gadiform larvae that have been reported to feed upon various noncopepod taxa .

Table

Species of fish for which feeding habits of larvae were examined .

iii

4

4

2-3

The Feeding Ecology of Some Zooplankters That are

Important Prey Items of Larval Fishl

JEFFERSON T. TURNER 2

ABSTRACT

Diets of 76 species of nsh larvae from most oceans ofthe world were inventoried on the basis ofInformation

In 40 pubUshed studies. Although certain geographk, slze- and taxon.speclnc pallerns were apparent, certain

zooplankton taxa appeared in the diets oflarvae of a variety of nsh species In numerous 10caUtles. Included were

six genera of calanold copepods (Acartia, Cala"",, Ce"tropage., PartJt:ala"",, P.eudocala"",, remora), three

genera of cyclopold copepods (Corycaell', Oitho"4, O"caea), harpactlcold copepods, copepod naupUl, tintinnids,

c1adocerans of the genera Evad"e and Podoll, barnacle naupUl, gastropod larvae, pteropods of the genus

Limaci"a, and appendicularlans. Literature on feeding habits of these zooplankters reveals that most of the

copepods are omnivorous, feeding upon both phytoplankton and other zooplankton. Some taxa, such as

Cala"",, Paracala"",, P.eudocala"",, and copepod naupUl appear to be primarily herbivorous, while others,

such as Acartia, Ce"tropages, remora, and cyclopolds exhibit broad omnivory or carnlvory. The noncopepod zooplankters are primarily nIter-feeders upon phytoplankton andlor bacterioplankton. Despite the Importance of

zooplankters In larval nsh food webs, speclftc knowledge of the feeding ecology of many taxa Is poor. Further,

much present knowledge comes only from laboratory Investigations that may not accurately portray feeding

habits of zooplankters in nature. Lack of knowledge of the feeding ecology of many abundant zooplankters,

which are also important In larval nsh food webs, precludes realistic understanding of pelagic ecosystem

dynamics.

INTRODUCTION

It has long been thought that fluctuations in year class sizes are

largely dependent upon the success of fish larvae in finding

necessary kinds and amounts of food (Hjort 1914). Understanding

the dynamics of food webs that support fish larvae requires information not only on the kinds and amounts of food required by larval fish, but also of the organisms upon which larvae feed.

Although larvae of numerous fish species, particularly clupeoids,

feed directly on phytoplankton (Blaxter 1974; Lasker 1975, 1978,

1981; Mendiola 1969, 1974), most also feed upon zooplankton, at

least during some stage of larval life. Since larvae of numerous

species of fish may feed selectively upon certain zooplankters,

which in turn may feed selectively on certain phytoplankters,

there is a possiblity that climatic- or anthropogenic-induced

alteration of phytoplankton species assemblages may produce

changes in food chain structures which extend to the top (Greve

and Parsons 1977).

Since many of the thousands of zooplankton taxa present in the

sea may not be directly involved in larval fish food webs, it is important to identify those zooplankters which are the food of larval

fish. The feeding habits of these zooplankters must then be defined before food webs of larval fish can be understood.

This paper reviews existing knowledge of feeding habits of

zooplankters that are important forage items of larval fish. These

zooplankters were identified from 40 published studies inventorying the diets of 76 species of fish larvae from most oceans of the

world. Eleven clupeiform, 38 perciform, 15 pleuronectiform, and

12 gadiform species were represented (Table 1). Care was taken to

'Contribution No. 83-35B, from the Southeast Fisheries Center, Beaufort

Laboratory.

'Southeast Fisheries Center Beaufort Laboratory, National Marine Fisheries Service, NOAA, Beaufort, NC 28516; on leave from: Biology Department, Southeastern

Massachusetts University, North Dartmouth, MA 02747.

use information on feeding of larvae only, not postlarvae.3 All but

a few studies were based on examinations of gut contents of fieldcaught larvae; the few exceptions were based on studies of larvae

feeding upon natural zooplankton assemblages. Collette (1978),

Hureau and Monod (1973), and Robins et a1. (1980) were used to

identify taxonomic synonymies of fish species.

Lists of fish species whose larvae have been reported to eat

various zooplankton taxa are presented in Appendix Tables 1-20.

For zooplankton taxa eaten by larvae of >5% of the total 76 fish

species, the percentages of the total number of fish, as well as

percentages of the total number from each order (Clupeiformes,

Perciformes, Pleuronectiformes, and Gadiformes) that eat various

zooplankters, are presented in Figures 1 and 2.

Certain geographic, size-specific, or taxon-specific patterns are

apparent from these figures. For instance, the cold-water

copepods Calanus and Pseudocalanus are consumed mainly by

cold-water cod larvae (Gadiformes). Conversely, the apparent low

utilization of the copepod genera Corycaeus, Oncaea, and

Microsetella, which are most abundant in tropical waters, is probably a reflection of fewer studies of the feeding of tropical fish

larvae than of temperate fish larvae. Although copepod nauplii are

eaten by larvae of most species, and by larvae of all clupeoid

species examined, they appear to be utilized much more intensively by younger (smaller) larvae. Since many clupeoid larvae are

known to feed directly on phytoplankton, it is no surprise that

most also eat small items such as copepod nauplii and tintinnids.

Although appendicularians were eaten by only a small percentage

of larvae, they were an important food for many temperate flatfish

larvae (Pleuronectiformes). Further, since appendicularians are

dominant zooplankters throughout the year, mainly in oceanic

'it is clear from sizes of fish examined that Lebour's (1918, 1919" 1920) papers

dealt with what we now call larvae. Apparently, larvae in her classification referred

to those which still retained a yolle sac.
Table l.-Specles of nsh for which the feeding habits of larvae were examined.

Species Common name Location' References

Order Clupeiformes (herring-like fishes)

Anchoa mitchelli

Brevoonia patron us

Brevoortia tyrannus

Brevoortia tyrannus

Clupea harengus

Clupea harengus

Clupea harengus

Clupea harengus

Clupea harengus

Clupea harengus

Clupea harengus

Clupea harengus

Clupea harengus

Engraulis anchaita

Engraulis mordax

Engraulis ringens

Harengula pensacolae

Sardina pilchardus

(Clupea pilchardus)'

Sardinops sagax

Sprattus spraltus

(Clupea sprattus)'

bay anchovy

gulf menhaden

Atlantic menhaden

Atlantic menhaden

herring

herring

herring

herring

herring

herring

herring

herring

herring

Argentine a,chovy

northern anchovy

Peru anchovy

scaled sardine

pilchard

California sardine

sprat

FLAE

GOMEX

NCE

DELE

NWECW

NWECW

NWECW

NWECW

NWECW

NWECW

NWECW

GOM

BOF

ACW

CALCW

PCW

FLAE

NWECW

CALCW

NWECW

Detwyler and Houde (1970)

Govoni et al. (1983)

Kjelson et al. (1975)

June and Carlson (1971)

Bainbridge and Forsyth (1971)

Bjl'lrke (1971, 1978)

Blaxter (1965)

Bowers and Williamson (1951)

Gamble et al. (1981)

Lebour (1918,1919,1920,1921)

Ogilvie (1927)

Sherman and Honey (1971)

White (1977, 1980)

Ciechomski (1967)

Arthur (1976)

Mendiola (1974)

Detwyler and Houde (1970)

Lebour (1920,1921)

Arthur (1976)

Lebour (1918,1919,1920,1921)

Order Perciformes (spiny-rayed fishes)'

Agonus caraphractus .rmed bullhead NWECW Lebour (1918,1919,1920)

Ammodytes americanus sand lance NEE Covill (1959)

Ammodytes marinus sand lance NWECW Ryland (1964); Wyatt (1974)

Ammodytes tobianus sand lance NWECW Lebour (1918,1920)

I;tmmodytes lanceolatus)'

Auxis sp. frigate tuna INDO Votani et al. (1981)

Blennius galtorugine blenny NWECW Lebour (1918,1919)

Callionymus lyra dragonet NWECW Lebour (1918,1919,1920)

Coryphoblennius galerita blenny NWECW Lebour (1918,1919)

(Blennius galerita)'

Crystallogobius nilssoni gob; NWECW Lebour (1918,1919,1920)

Ctenolabrus rupestris wrasse NWECW Lebour (1919, 1920)

Cyclogaster montagui sea snail NWECW Lebour (1918,1919)

Diplecogaster bimaculatus clingfish NWECW Lebour (1919, 1920)

(Lepadogaster bimaculatus)'

Eutrigla gumardus gurnard NWECW Lebour (1918,1920)

(Trigla gurnardus)'

Gobius niger goby NWECW Lebour (1919, 1920)

(Gobius paganellus)'

Gobiusculus flavescens goby NWECW Lebour (1918,1919,1920)

(Gobius minutus)'

(Gobius ruthenspa"i)'

Katsuwonus pelamis skipjack tuna INDO Votani et al. (1981)

Labrus.bergylta wrasse NWECW Lebour (1918,1919,1920)

Labrus bimaculatus wrasse NWECW Lebour (1920)

(Labrus mixtus)'

Lagodon rhamboides pinfish NCE Kjelson et al. (1975)

Leiostomus xanthurus spot NCE Kjelson et al. (1975)

Leiostomus xanthurus spot GOMEX Govoni et al. (1983)

Lepadogaster candollei clingfish NWECW Lebour (1918,1919,1920)

Lepadogaster lepadogaster clingfish NWECW Lebour (1918,1919)

(Lepadogasler gouani)'

Micropogonias undulatus Atlantic croaker GOMEX Govoni et al. (1983)

Myoxocephalus aenaeus sculpin NEE Laroche (1982)

Myoxocephalus

oclodecemspinosus sculpin NEE Laroche (1982)

Myoxocephalus scorpius sculpin NEE Laroche (1982)

Pomatoschistus microps goby NWECW Lebour (1920)

(Gobius microps)'

Sebastes marinus redfish IS Bainbridge (1965)

Sebastes marinus redfish GOM Marak (1974)

Scomber scomber mackerel NWECW Lebour (1918,1920)

Taurulus bubalis bullhead NWECW Lebour (1918,1919,1920)

(CoitUS bubalis)'

Thunnus alalunga albacore INDO Votani et al. (1981)

Thunnus albacares yellowfin tuna INDO Votani et al. (1981)

2
Table l.-Continued.

Species

Thunnus moccayi;

Thunnus obesus

Trachinus vipera

Trachurus symmetricus

Trachurus trachurus

(Caranx trachurus)2

Triglops murrayi

Common name

southern bluefin

bigeye tuna

weaver

jack mackerel

horse mackerel

sculpin

Location'

INDO

INDO

NWECW

CALCW

NWECW

NEE

References

Uotani et al. (1981)

Uotani et al. (1981)

Lebour (1918,1919,1920)

Arthur (1976)

Lebour (1918,1919,1920)

Laroche (1982)

Order Pleuronectiformes (flatfish)

Amoglossus sp.

Limanda limanda

(Pleuronectes Iimanda)'

Microchirus variegatus

(Solea variegata)2

Microstomus kilt

(Pleuronectes microcephalus)'

Phrynorhombus regias

(Zeugopterus unimaculatus)'

Phrynorhombus norvegicus

(Scophthalmus norvegicus)2

Platich/hys flessus

(Pleuronectes flessus)2

Pleuronectes platessa

Psetta maxima

(Rhombus maximus)2

Pseudopleuronectes

amer;canus

Scoph/halmus maximus

Scophthalmus rhombus

(Rhombus laevis)'

Solea nasuta

(Solea lascaris)2

Solea solea

(Solea vuIgaris)2

Zeugopterus punctatus

lefteye flounder

dab

thickback sole

lemon dab

turbot

turbot

righteye flounder

plaice

turbot

winter flounder

turbot

brill

lemon sole

common sole

turbot

NWECW

NWECW

NWECW

NWECW

NWECW

NWECW

NWECW

NWECW

NWECW

NEE

NWECW

NWECW

NWECW

NWECW

NWECW

Lebour (1918, 1920)

Last (1978a); Lebour (1918,1919,1920)

Lebour (1918,1919)

Lebour (1918,1919,1920)

Lebour (1918)

Lebour (1918,1920)

Last (l978a); Lebour (1918, 1920)

Last (1978a); Lebour (1920); Ryland

1964; Shelbourne (1953, 1957, 1962);

Wyatt (1974)

Lebour (1920)

Pearcy (1962)

Last (1979)

Lebour (1919,1920)

Lebour (1918,1919,1920)

Last (1978a); Lebour (1918,1919,1920)

Lebour (1918)

cod NWECW

cod GOM

haddock NWECW

haddock GOM

whiting NWECW

hake NWECW

Pacific hake CALCW

blue whiting NWECW

ling NWECW

pollack NWECW

coalfish GOM

bib NWECW

bib NWECW

Ciliata mustela

(Onos mustela)'

Gadus morhua

Gadus morhua

Melanogrammus aeglefinus

Melanogrammus aeglefinus

Merlangius mer/angus

(Gadus merlangus)2

Merluccius merluccius

Merluccius productus

Micromesistius poutassou

Molva molva

Pollachius pollachius

(Gadus pollachius)2

Pollachius virens

Trisopterus luscus

(Gadus luscus)'

Trisopterus minutus

(Gadus minutus)2

rockling

Order Gadiformes (cod-like fishes)

NWECW Lebour (1918,1919,1920)

Last (l978b); Lebour (1918);

Ellertsen et al. (1981)

Marak (1960);

Sysoeva and Degtereva (1965)

Ogilvie (1938)

Marak (1960)

Last (1978b); Lebour (1918, 1919, 1920)

Lebour (1920)

Sumida and Moser (1980)

Conway (1980)

Lebour (1920)

Lebour (1918,1919,1920)

Marak (1960)

Last (1978b); Lebour (1918,1919,1920)

Lebour (1918, 1920)

I ACW, Argentine coastal waters; BOF, Bay of Fundy; CALCW, California coastal waters; DELE, Delaware estuarine waters;

FLAE, Florida estuarine waters; GOM, Gulf of Maine; GOMEX, Gulf of Mexico coastal waters; INDO, Indian Ocean; IS, Irminger

Sea; NCE, North Carolina estuarine waters; NEE, New England estuarine waters; NWECW, northwest European coastal waters;

PCW, Peru coastal waters.

2Previous taxonomic synonymy.

'For the purposes of this review, five species of Order Scorpaenidae (Myoxocephalus aenaeus, M. oc/odecemspinosus. M. scorpius,

Sebas/es marinus, and Triglops murrayi) are included with the perciforms, since their larvae are morphologically and ecologically

"perciform-Iike."

3
% PERCIFORM SPECIES

% GAOlFORM SPECIES

% CLUPEIFORM SPECIES

% PLEURONECTIFORM SPECIES

% GADIFORM SPECIES

% PLEURONECTIFORM SPECIES

% PERCIFORM SPECIES

% CLUPEIFORM SPECIES

• % TOTAL FISH SPECIES

III

D

Figure 2.-Percentages of the total number of nsh larvae (76 species), the total

number of clupelform larvae (11 species), the total number of perclform larvae

(38 species), the total number of pleuronectlform larvae (IS species), and the

total number of gadlform larvae (12 species), listed In Table 1, that have been

reported to feed upon various noncopepod taxa.

ACARTIA

IOO~

n:ldJJI rJNTINNIOS

,':Lw~ EVADNE POOON BARNACLE NAUPLII

,':luo~L o GASTROPOD LARVAE LIMAC/NA APPENOICULARIANS

• "10 TOTAL FISH SPECIES

II ;::W D

COPEf'OO NAJJPLII

;~~Ulom ACARTIA CALANUS CENTROPAGES

,':ww~ o PARACALANUS PSEUOOCALANUS TEMORA

,'~~~L CORYCAEUS O/THONA ONCAEA

,':~L~ o

Figure I.-Percentages of the total number of nsh larvae (76 species), the total

number of clupelform larvae (11 species), the total number of perclform larvae

(38 species), the total number of pleuronectiform larvae (IS species), and the

total number of gadlform larvae (12 species), listed In Table 1, that have been

reported to feed upon various copepod taxa.

waters, it is understandable that they were important prey items

for oceanic tuna larvae (Perciformes).

Aside from such specific patterns, it is clear that the zooplankton taxa shown in Figures 1 and 2 are eaten by larvae of a variety

of fish species in numerous localities. These zooplankters include

adults and/or copepodites of six genera of calanoid copepods

(Acartia, Calanus, Centropages, Paracalanus, Pseudocalanus, and

Temora), adults and/or copepodites of three genera of cyclopoid

copepods (Corycaeus, Oithona, and Oncaea), and adults and/or

copepodites of two genera of planktonic harpacticoid copepods

(Euterpina and Microsetella). Also included are zooplankton often

unidentified, such as harpacticoids, copepod nauplii (some of

which were identified to genus, but most not), tintinnids, the

cladocerans Evadne and Podon, barnacle nauplii, gastropod larvae,

the pteropod Limacina, and appendicularians. However, with the

exception of the appendicularians which were not identified in the

study of Uotani et al. (1981), the other records for appendicularians refer to Oikopleura dioica and Frittilaria borealis.

The remainder of this review will summarize published knowledge of feeding habits of the zooplankton taxa listed above.

£'UTERPINA M/CROSETELLA OTHER HARf'/lCT/COIOS Acartia is a broadly distributed copepod genus (Bradford 1976).

In most estuarine waters of the world one or more Acartia species

are numerically co-dominant, if not the dominant, copepods (see

references cited in Turner 1981).

Acartia is omnivorous. Early gut content studies reveal remains

of diatoms, dinoflagellates, radiolarians, and crustaceans (Lebour

1922; Marshall 1949; Conover 1956). Numerous laboratory

studies have shown that various species of Acartia feed upon

unialgal or mixed cultures of diatoms, dinoflagellates, microflagellates, coccolithophorids, and other phytoplankton (Gauld

and Raymont 1953; Conover 1956, 1960; Anraku and Omori

1963; Anraku 1964a, b; Marshall and Orr 1966; Esaias and Curl

1972; Gaudy 1974; Corkett and Zillioux 1975; Nival and Nival

1976; O'Connors et al. 1976; Piontkovskii and Petipa 1976;

Reeve and Walter 1977; Dagg 1977; Roman 1977; Honjo and

Roman 1978; Tomasini and Mazza 1978; Donaghay and Small

1979a, b; White 1979; Deason 1980a, b; Rosenberg 1980; White

1981; Cahoon 1981; Bartram 1981; Tomas and Deason 1981;

Weiss 1983).

Acartia is also capable of carnivorous feeding. Various species

of Acartia eat their own juveniles (Conover 1956; Roman 1977;

Landry 1978; Lonsdale et al. 1979), nauplii of other copepod

species (Hodgkin and Rippingale 1971; Rippingale and Hodgkin

1975; Lonsdale et al. 1979; Lonsdale 1981), Artemia nauplii4

(Conover 1960; Anraku and Omori 1963; Gaudy 1974),juvenile

chaetognaths (Davis 1977), and tintinnids (Turner and Anderson

in press; Robertson 1983).

4The use of Anemia nauplii as prey in experiments on predation by marine zooplankton is artificial since Anemia is absent from the ocean. References to such

studies will be included, however, since ingestion of Anemia nauplii is the only

evidence of feeding upon animal prey by some marine zooplankters.

4
In addition to eating phytoplankton and zooplankton, Acartia

eats other items. Friedman (1980) presented electron microscopic

evidence that A. tonsa can ingest bacteria as small as 0.5 /-1m in

diameter. It can also ingest particulate macrophyte detritus, but

does not survive well on this diet (Roman 1977). Acartia negligens

collected near coral reefs ingested and assimilated up to 50% of

the organic matter in mucus produced by scleractinian corals

(Richman et al. 1975).

The few attempts to compare the relative importance of plant

and animal food for Acartia have produced mixed results. Anraku

and Omori (1963) found that A. tonsa ate equal amounts of

Artemia nauplii and the diatom Thalassiosirafluviatilis. However,

Gaudy (1974) reported that A. clausi exhibited a marked

preference for phytoplankton over Artemia nauplii. Lonsdale et al.

(1979) demonstrated that the rates of predation by A. tonsa adults

on conspecific nauplii and nauplii of other copepods were not

reduced by the presence of alternative phytoplankton food.

There is also disagreement on the selective feeding of Acartia

on particles. Although many studies have suggested size-selective

grazing upon the largest available cells in algal cultures, recent

studies of the feeding ofAcartia on naturally occurring particulate

assemblages (Hargrave and Geen 1970; Richman et al. 1977;

Poulet 1978; Mayzaud and Poulet 1978; Roman and Rublee 1980;

Ryther and Sanders 1980; Bartram 1981; Turner and Anderson

1983) have produced contradictory results. Richman et al. (1977)

found that A. tonsa and A. clausi preferentially fed upon the

largest available particles, even if these were not the most abundant. However, as these larger particles were grazed down, grazing

pressure shifted to biomass peaks of smaller sized particles. Poulet

(1978) and Mayzaud and Poulet (1978) concluded that A. clausi

was an opportunistic grazer on particles between 1.5 and 150 /-1m

equivalent spherical diameter, and feeding was nonselective with

regard to particle size. As in the study of Richman et al. (1977),

Poulet (1978) reported that grazing A. clausi tracked biomass

peaks. Turner and Anderson (1983) found that in terms of filtration rate,A. hudsonica (= A. clausi) was unselective when feeding

upon combinations of naturally occurring concentrations of a

small dinoflagellate (Heterocapsa triquetra) and a larger

dinoflagellate (Gonyaulax tamarensis), or G. tamarensis and tintinnids. Each type of food was eaten in approximate proportion to its

availability. Ryther and Sanders (1980) examined the feeding of

A. tonsa on natural particulate assemblages in large-volume

continuous-flow enclosures and reported that A. tonsa reduced the

number of centr.ic diatoms (mainly Skeletonema costatum) by 10%

in grazed enclosures, but that small flagellates increased by

35-60%. Presumably this occurred because, as suggested by Conover (1956), Acartia is an inefficient grazer of small flagellates,

and these increased in the grazed enclosures due to reduced

nutrient competition with diatoms.

Other studies suggest that some flagellates are poor food for

Acartia. Tomas and Deason (1981) offered A. hudsonica and A.

tonsa cultures of the diatom Skeletonema costatum and the

flagellates Dunaliella tertiolecta and Olisthodiscus luteus, either

singly or in combination. At intermediate and high concentrations, O. luteus was inhibitory to feeding, both on itself and

on the other phytoplankters. Also, Parrish and Wilson (1978)

found that A. tonsa females produced eggs equally well on diets of

the diatom Thalassiosira pseudonana or the. "large" (100-140

/-1m3 ) flagellate Chroomonas salina, but that when the small

flagellate Isochrysis galbana (40 /-1m3 ) was the only food, egg production ceased, as with starved females, after 3 d.

5

These observations are somewhat at odds with the results of

Acartia culture studies. Flagellates such as Isochrysis galbana,

Dunaliella tertiolecra, and Platymonas succia have been used as

food (usually in combination with diatoms) to rear multiple

generations of A. tonsa, A. clausi, or A. steueri from fertilized egg

to adult (Zillioux and Wilson 1966; Heinle 1969, 1970; Zillioux

1969; Nassogne 1970; Dye 1980a, b" 1981; Tester 1982). However, Klein Breteler (1980) successfully cultured A. clausi only

when dinoflagellates were added as food, not when

microflagellates alone were used. These discrepancies possibly

reflect the suitability of small flagellates as food for Amrtia

nauplii, but additional diatom or dinoflagellate food is needed for

late copepodite and adult stages of Acartia.

Earlier observations of the feeding mechanisms of Acartia

(CoJ;lover 1956; Gauld 1966), supplemented by recent cinematographic studies (Rosenberg 1980), revealed that the feeding

mechanisms of Acartia are complex. Other observations suggest

the possibility of chemosensory detection of food. For instance,

Poulet and Marsot (1978) found that microcapsules enriched with

homogenate of naturally occurring phytoplankton were ingested

preferentially over nonenriched capsules by A. clausi. Lonsdale et

al. (1979) found that predation rates by A. tonsa adults on nauplii

of three other copepod species were significantly higher than

cannibalism of conspecific nauplii. This suggests that Acartia tonsa adults can recognize conspecific nauplii, relative to those of

other species.

The abundance of Acartia in food-rich inshore, particularly

estuarine, environments may relate largely to its feeding ecology.

Dagg (1977) found thatA. tonsa needed fairly continuous food for

survival and egg production. Acartia tonsa starved within 6-10 d

with no or only intermedilite additions of food, whereas other cooccurring copepod species survived for up to 20 d under constant

starvation. Dagg concluded that Acartia is less well equipped than

other copepod species to overcome periods of poor food conditions and subsequently exploit food-rich patches when they occur.

However, in continuous food-rich conditions, such as estuaries,

Acartia can outcompete other species due to its elevated fecundity.

CALANUS

It has long been known that species of the genus Calanus are

omnivorous. Gut contents of North Atlantic C. finmarchicus contain remains of diatoms, dinoflagellates, silicoflagellates,

Phaeocystis, coccolithophorids, radiolarians, tintinnids, molluscan

larvae, and other crustaceans (Dakin 1908; Lebour 1922; Marshall 1924). Marshall (1924) found that the food remains in any

single gut were mixed, that seasonal changes in gut contents

reflected seasonal changes in the microplankton upon which the

copepods fed, and that no particular preference was shown for any

given species of phytoplankton. Guts of Pacific C. ten uicornis, C.

plumchrus, and C. cristatus also contain diatom, coccolithophorid,

silicoflagellate, tintinnid, and foraminiferan remains (Gienrikh

1958; Arashkevich 1969).

Numerous studies have shown that various species of Calanus

eat a variety of cultured phytoplankton diets (Clarke and Gellis

1935; Fuller and Clarke 1936; Fuller 1937; Harvey 1937; Clarke

and Bonnet 1939; Gauld 1951; Marshall and Orr 1952, 1955a, b,

1958, 1961; Conover 1960,1966a,b,c;Mullin 1963; Anraku and

Omori 1963; Anraku 1964a; Corner et al. 1967, 1972; Butler et

al. 1969; Richman and Rogers 1969; PaffenhOfer and Strickland

1970; PaffenhOfer 1971, 1976a, b; Esaias and Curl 1972; Frost

1972, 1975,1977;Gaudy 1974; Dagg 1977; Ikeda 1977; Schnack
1979, 1983; Vidal 1980a, b; Runge 1980; Gifford et a1. 1981; Cox

and Willason 1981; Huntley 1982). Other studies have examined

the filter feeding of Calanus on natural particulate assemblages

(Corner 1961; Cowey and Corner 1963; Adams and Steele 1966;

Butler et a1. 1970; McAllister 1970; Ikeda 1971; Taguchi and Ishii

1972; Gamble 1978; Cowles 1979; Hebert and Poulet 1980; Dagg

et a1. 1980, 1982; Boyd et a1. 1980; Huntley 1981, 1982; Harris

1982). Additionally, C. helgolandicus and C. pacificus (possibly

the same species) have been cultured on phytoplankton diets

(Mullin and Brooks 1970a, b; PaffenhOfer 1970; Vidal 1980a, b).

Many concepts of copepod feeding that have risen to the status

of paradigms are based upon experiments with Calanus. For instance, the concept that copepods preferentially select larger sized

phytoplankton cells began as a result of Harvey's (1937) experiments with C. finmarchicus. Harvey fed the copepods on mixtures

of large and small diatoms, and found that invariably more of the

larger cells were eaten. Richman and Rogers (1969) reported that

C. helgolandicus filtered significantly higher numbers of paired

daughter cells (after cell division, but before final separation) of

the diatom Ditylum brightwelli, than unpaired cells. Since the same

species was offered, and the only difference was in terms of size

rather than nutritional content, Richman and Rogers concluded

that their results demonstrated size-selective feeding. Frost (1977)

found that C. pacificus, presented with different-sized cells of the

same diatom species, invariably ate large cells at a higher rate

than small ones. Frost concluded that this differential filtration of

cells of different sizes was purely mechanical and a function of

variable retention efficiency of the intersetular distances of the

filtering appendages (second maxillae) of the copepods. The

results of Gamble (1978) support the notion of size-selective

feeding on larger particles, in that C. finmarchicus mainly filtered

the largest available cells, which were also the most abundant,

during a spring diatom bloom in the North Sea. Calanus may also

preferentially feed on chain-forming or spine-bearing diatoms

because they occupy a larger volume and are more frequently encountered. Schnack (1979) fed C. helgolandicus on mixtures of

chain-forming diatoms, and found that grazing was most intense

upon species which had long spines. Gifford et a1. (1981) obtained

the same result using C. finmarchicus.

The results of some recent studies of Calanus feeding upon

natural particulate assemblages often do not point to size-selective

feeding. Cowles (1979) found that C. chilensis in the Peru upwelling were size-selective grazers at high food concentrations, but

became less selective as total food concentration declined. Also,

the size spectrum of particles ingested was much broader at low

than at high food concentrations. Cowles concluded that while the

mechanical siev~ mechanism of filtering (Boyd 1976; Frost 1977)

could explain the patterns recorded at high food concentrations, it

could not explain the diet expansion at low food levels. Similarly,

Harris (1982) found that C. pacificus grazed a broad size spectrum

of phytoplankton, similar to that grazed by the much smaller

copepod Pseudocalanus. Harris concluded that both species compete for the same food resource over a wide range of the particlesize spectrum. Hunger of animals also appears to influence size

selection. Runge (1980) found differences in filtration rate versus

cell size relations for starved versus well-fed C. pacificus.

The question of size-selective feeding by Calanus and other

copepods is still unresolved. Inasmuch as laboratory studies using

phytoplankton cultures as food allow reliable quantification of

grazing on distinct size classes of food, they have the disadvantage

of not offering copepods the broad size spectrum and nutritional

6

array encountered in nature. Thus, because experimental conditions are partly artificial, patterns recorded may have questionable

relevance to copepod behavior in nature. Feeding studies using

natural particulate assemblages have the advantage of presenting

animals with a food array found in nature. Unfortunately most

such recent studies have quantified particle number and size by

using electronic particle counters. Harbison and McAlister (1980)

demonstrated numerous artifacts which can result from using

electronic particle counters to quantify natural particle

assemblages. They concluded that many reported patterns of

copepod feeding resulting from such studies are not based upon

experimental evidence.

The study of Huntley (1981) represents one of the few attempts

to define copepod feeding upon natural particulate assemblages in

which particle quantification was performed by visual rather than

electronic means. Huntley found that three species of Calanus

from the Labrador Sea exhibited no selective feeding on various

phytoplankton species (>20 J.lm cell diameter) on the basis of size,

shape, or species of cell. The copepods removed various

phytoplankters in direct proportion to their abundance.

Another copepod feeding paradigm based on experiments with

Calanus is that of the maximum ingestion rate plateau. Frost

(1972) fed monospecific diatom cultures at different cell concentrations to adult female C. pacificus and found that the ingestion

rate increased linearly with increasing food concentration up to a

maximum rate, beyond which the ingestion rate remained constant with increasing levels of food. This maximum ingestion rate

(saturation) was the same, in terms of carbon, for diatoms ranging

in diameter from 11 to 87 J.lm, but as cell size increased the carbon

concentration at which saturated feeding was achieved decreased.

Frost (1972) concluded that C. pacificus can obtain their maximum daily ration at lower carbon concentrations when feeding

on larger cells.

Conover (1978b) suggested that saturated feeding is likely an

artifact of unnaturally high laboratory food concentrations, and

concluded that food concentrations in nature are rarely, if ever,

high enough to induce saturation. This view is supported by experiments of Reeve and Walter (1977) with Acartia tonsa, and

Huntley (1981) with Calanus. Huntley found no evidence of a

saturated ingestion rate for three Calanus species feeding on the

spring diatom bloom in the Labrador Sea. Although Runge (1980)

did record saturated feeding rates by C. pacificus, he found that

there are seasonal changes in the level of food at which saturation

is obtained.

Calanus also exhibits carnivorous feeding. Various species eat

their own eggs (Conover 1960, 1966a), Artemia nauplii (Anraku

and Omori 1963; Gaudy 1974), barnacle nauplii (Corner et a1.

1974, 1976), and the nauplii of other species of copepods (Landry

1980, 1981). Calanus finmarchicus appeared to prefer

phytoplankton over Artemia nauplii in the studies of Anraku and

Omori (1963) and Gaudy (1974), but Landry (1981) found that C.

pacificus can switch from eating diatoms to copepod nauplii,

eating primarily whatever is most abundant.

Other food items eaten by Calanus include fecal pellets and

other forms of detritus (PaffenhOfer and Strickland 1970). Paf~

fenhOfer and Knowles (1979) reared C. helgolandicus from

copepodite stage III to adult on a diet of fecal pellets alone. Boyd

et a1. (1980) found that C. chilensis in the Peru upwelling also

appeared to utilize nonliving detrital particulates as a major component of its diet. Additionally, C. helgolandicus has been shown

to eat "red mud" (microscopic particles of Fe 20 3 residual from

extraction of aluminum from bauxite), but the copepod does not
survive or feed well upon diatoms in the presence of "red mud"

(Paffenhofer 1972).

Although ingestion of nonnutritional particles such as "red

mud" suggests that Calanus cannot discriminate between different

food items, there is also limited evidence to suggest that a choice

may occur. Huntley (1982) found that C. pacificus filtration rates

on a large dinoflagellate during a red tide were minimal compared

with those when a small diatom was used as food. He concluded

that the persistence of the bloom may have been partly due to a

lack of grazing, and that the dinoflagellate was likely avoided by

the copepods.

Although some phytoplankton blooms may not be grazed

because their component species are repulsive to copepods, other

blooms may develop because of a lag in copepod feeding response.

Cox and Willason (1981) found that production of the digestive

enzyme laminarinase by C. pacificus requires presence of the

substrate laminarin (the principal storage product of marine

phytoplankton). Since there was at least a I d lag in the production of the enzyme in the copepods after addition of substrate, Cox

and Willason proposed that such a time lag may be important in

areas of bloom potential, since the lag would allow initial

development of a bloom before it could be destroyed by grazing.

The role of Calanus and other copepods in cropping primary

production has long been of interest (Harvey et al. 1935; Riley

1946; Cushing 1958, 1964; Cushing and Vucetic 1963; Dagg and

Turner 1982) and, as expected, the impact of grazing has varied.

Dagg et al. (1980) found that C. chilensis and two other large

copepods consumed <5% of the daily carbon production in the

Peru upwelling. Dagg et al. (1982) found that three large copepods

(Eucalanus bungii, Calanus (Neocalanus) cristatus and C. (N.)

plumchrus) removed 6-25% of the daily primary production during the spring bloom in the Bering Sea. Dagg and Turner (1982)

found that during the spring bloom in the New York Bight and

Georges Bank areas a copepod community dominated by C. finmarchicus and Pseudocalanus grazed 15-48% of the daily primary

production. In both areas, grazing impact was highest on the outer

shelf (30-48% of primary production) because of large populations of C. finmarchicus. In other regions, grazing appears particularly important. Frost (1980) stated that the lack of seasonal

variation in the phytoplankton biomass of the subarctic North

Pacific, where phytoplankton apparently never reach bloom

concentrations even though conditions favorable to blooming do

occur, is largely due to continuous grazing by subadults of C. (N.)

plumchrus and C. (N.) cristatus. Since. adults of these species do

not feed, reproduction is not dependent upon phytoplankton

pulses, and large numbers of juvenile copepods are present in the

epipelagic waters during the season when phytoplankton blooms

should occur.

Frost et al. (1983) confirmed that Calanus (Neocalanus) cristatus and C. (N.) plumchrus CV copepodites could feed on the low

concentrations of small phytoplankton cells found in the subarctic

Pacific. In addition, these copepodites exhibited predatory feeding

upon copepodites of the small copepod Oithona similis.

In conclusion, it appears that species of Calanus are omnivorous, in that they are capable of carnivorous as well as herbivorous feeding. However, it is likely that this genus is primarily

and broadly herbivorous, resorting to camivory mainly during

periods of phytoplankton paucity (Corner et al. 1974; Digby

1954).

CENTROPAGES

Members of the genus Centropages are broadly omnivorous. Gut

7

contents have included remains of diatoms, dinoflagellates, coccolithophorids, tintinnids, copepods, and molluscan larvae (Lebour

1922; Marshall 1924, 1949; Marshall and Orr 1966). Cultured

phytoplankton diets have been fed to C. hamatus (Ki9irboe et al.

1982; Marshall and Orr 1966; Gauld 1951; Gauld and Raymont

1953; Smith and Hall 1980; Klein Breteler 1980), C. typicus

(Cowles and Strickler 1983; Gaudy 1974; Dagg 1977, 1978;

Tomasini and Mazza 1978; Smith and Hall 1980), and C. furcatus

(in the Atlantic = C. velificatus - see Fleminger and Hulsemann,

1973) (PaffenhOfer and Knowles 1980). Centropages is also

capable of carnivorous feeding (Anraku and Omori 1963; Gaudy

1974; PaffenhOfer and Knowles 1980; Conley 1983).

There is a lack of consensus on the relative importance of plant

and animal food in Centropages diets. Anraku and Omori (1963)

examined the feeding responses of C. hamatus and C. typicus on

diets of Artemia nauplii and diatoms, either singly or in combination. Both species of Centropages ate both types of food, but

animal food (Artemia) was preferred. However, Gaudy (1974)

found that C. typicus fed equally well on Artemia nauplii and

phytoplankton cultures. PaffenhOfer and Knowles (1980) found

that the subtropical species, C. velificatus, ingested a higher

percentage of its body nitrogen as copepod nauplii (laboratoryreared Pseudodiaptomus coronatus) than as unialgal phytoplankton (Rhizosolenia). Conversely, Conley (1983) found that

the temperate species, C. hamatus, ingested a higher proportion of

its body carbon as natural estuarine phytoplankton (predominantly small flagellates) than as cooccurring wild copepod nauplii

(primarily Acartia hudsonica or A. tonsa). Such differences may

reflect the use of different food sources, different experimental

procedures and methods of measurement, or possibly real differences between subtropical continental shelf (c. velificatus) and

temperate estuarine (c. hamatus) species.

There have been few investigations of the grazing of Centropages on natural particulate matter. Dagg and Grill (1980) found

that ingestion rates of C. typicus in the New York Bight were

usually less than maximal, and concluded that ingestion rates were

closely related to food quality because of the narrow range of

available food concentrations. Dagg et al. (1980) calculated that

in the Peru upwelling the population of Centropages brachiatus

consumed an average of 1.5 I.lgC/m2 per d. Since average primary

production rates were> I gClm2 per d, Dagg et al. calculated that

the grazing stress of C. brachiatus and two other large copepods

(Eucalanus inermis and Calanus chilensis) was always <5% of the

daily carbon production. Cowles (1979) examined the feeding

responses of the same three species of copepods in the Peru

upwelling. He found that C. brachiatus was a size-selective grazer

at high food concentration, and exhibited decreased selectivity as

total food level declined.

Boyd et al. (1980) found that in the Peru upwelling C. brachiatus migrated into the food-rich upper 5 m at night, but did not

show a migration pattern when food was scarce. Using gut

fluorescence techniques, Boyd et al. found no indication that C.

brachiatus was grazing phytoplankton in proportion to its abundance, and concluded that this reflected predatory feeding in addition to grazing.

As with Acartia tonsa, Dagg (1977) found that C. typicus is intolerant of starvation and intermittent food. He concluded that C.

typicus is therefore sensitive to small-scale patchiness. The migration data of Boyd et al. (1980) support this proposal, in that C.

brachiatus actively moved into food-rich waters. Their data also

support other observations suggesting that carnivory is a major

component of Centropages ecology.
PARACALANUS

Limited available literature suggests that small « I mm adult

body length) copepods of the genus Paracalanus are primarily

herbivorous. Lebour (1922) found diatom and coccolithophorid

remains in the guts of P. parvus, and several Paracalanus species

have been fed phytoplankton cultures or natural particulate

assemblages (Marshall and Orr 1966; Ikeda 1977; Checkley 1980;

Bartram 1981; Price et aJ. 1983). Bartram (1981) showed that P.

parvus was more efficient at capturing small particles than was

Acartia, and that P. parvus fed at a higher rate on small particles.

Sorokin et aJ. (1970) stated that P. parvus can also filter bacteria.

The calculations of Paffenhofer (1982) indicate that Paracalanus can have a substantial grazing impact on natural systems.

Applying his own grazing rates for Paracalanus sp. from the

southeastern shelf of the United States to the numbers of

Paracalanus denudatus reported by Dagg et aJ. (1980) in the Peru

upwelling, Paffenhiifer calculated that Paracalanus could graze an

average of 33% of the primary production in the Peru upwelling.

PSEUDOCALANUS

Corkett and McLaren (1978:54-80) reviewed feeding studies

published prior to 1977, which showed that Pseudocalanus is

mainly a filter feeder, ingesting a variety of diatom, coccolithophorid, flagellate, and radiolarian foods. However, Marshall

(1949) found crustacean remains in Pseudocalanus guts, and Corkett and McLaren (1978) stated that females will cannibalize their

own nauplii. Pseudocalanus also ingests detrital particles (Poulet

1976). Nonetheless, Corkett and McLaren concluded that

Pseudocalanus is overwhelmingly herbivorous. This conclusion is

supported by culture studies of Paffenhofer and Harris (1976) and

Vidal (1980a). In both cases, Pseudocalanus was cultured from

fertilized eggs to adults on diatom diets.

Recent studies of grazing on natural particulate assemblages

(poulet 1978; Koeller et aJ. 1979; Harris 1982) reveal that

Pseudocalanus ingests a broad spectrum of particle sizes. Poulet

(1973, 1974, 1978) reported seasonal shifts of grazing on the

most abundant particles, regardless of size, and concluded that

Pseudocalanus is an unselective opportunistic grazer. Harris

(1982) found that Pseudocalanus grazed a size spectrum of

phytoplankton similar to that grazed by Calanus, which is 10

times larger in terms of body weight.

Dagg (1977) found that Pseudocalanus can withstand starvation

for at least 16 d, and can produce eggs under discontinous starvation at rates equivalent to those for continuously fed animals.

Dagg concluded that Pseudocalanus can metabolically remove

itself from poor food conditions and wait for food-rich patches.

Vidal (1980a) demonstrated that the growth rate of Pseudocalanus

is relatively insensitive to food concentration. Vidal concluded

that this contributes to survival fitness of Pseudocalal1us in

adverse conditions which frequently occur in its temperate and

boreal habitat.

TEMORA

Early gut content studies (Lebour 1922; Marshall 1949) revealed that Temora is omnivorous, eating diatoms, dinoflagellates, the

haptophycean Phaeocystis, radiolarians, and other crustaceans.

Gauld (1966) stated that Temora is a filter feeder that also feeds as

a grasping predator.

8

Numerous studies show that the temperate species Temora

longicornis eats a variety of cultured phytoplankton diets (Gauld

1951,1953; Gauld and Raymont 1953; Raymont 1959; Conover

1959; Marshall and Orr 1966; Berner 1962; Harris and Paffenhofer 1976a; Cushing 1958; Smith and Hall 1980; Schnack 1983;

Weiss 1983). Studies reaching similar conclusions for the subtropicaJltropical species T. turbinata and T. stylifera include those

of Gaudy (1974), Ikeda (1977), and Paffenhiifer and Knowles

(1978, 1980). Temora longicornis has also been cultured from fertilized egg to adult on various flagellate or diatom diets (Corkett

1967, 1970; Corkett and Zillioux 1975; Harris and Paffenhiifer

1976a; Klein Breteler 1980).

Studies of grazing of Temora on natural particulate assemblages

have had inconsistent results. Hargrave and Geen (1970) found

that T. longicornis ate all sizes of flagellates present in natural

estuarine water, and that feeding rates were highest on larger cells,

even though smaller cells were most abundant. They also reported

that T. longicornis did not eat chain-forming diatoms of the genus

Chaetoceros, or other chain-formers, even though these were the

numerically dominant phytoplankters. This seems peculiar, since

Harris and Paffenhiifer (1976a) have cultured the same copepod

on unialgal diets of chain-forming diatoms, and Gaudy (1974) and

Paffenhofer and Knowles (1978) demonstrated that T. stylifera

and/or T. turbinata ate a variety of chain-forming diatoms.

Poulet (1978) and Mayzaud and Poulet (1978) reported that T.

longicornis feeding on natural particles was an opportunistic

grazer, and nonselective with regard to particle size. Mayzaud and

Poulet (1978) also found that ingestion rate was linear with increasing food concentration, regardless of particle size. In a

similar study with T. longicornis feeding on natural particles,

O'Connors et aJ. (1980) found that different ingestion rate versus

food concentration relations emerged when the food particle

assemblage had differences in species composition and size

distribution. Maximum ingestion rates were higher when T.

longicornis fed upon larger sized chain-forming diatoms than on

naked microflagellates.

Temora is also capable of carnivorous feeding. Various species

eat corpses of other copepods (Lowndes 1935), their own eggs

(Gaudy 1974), Artemia nauplii (Gaudy 1974), and the nauplii of

other copepods (Paffenhofer and Knowles 1980). Although Gaudy

(1974) found that T. stylifera had a higher food intake when

feeding on Artemia nauplii than when feeding on various

phytoplankton cultures, Paffenhiifer and Knowles (1980) found

that T. stylifera ingested more food as phytoplankton than as

nauplii of another copepod.

In addition to phytoplankton and zooplankton foods, T.

longicornis has ingested oil particles following a tanker wreck,

with no apparent ill effect (Conover 1971). Also, Paffenhiifer and

Knowles (1979) found that T. stylifera ate fecal pellets of the

nauplii of another copepod at about the same rate as it ate large

phytoplankton cells (Rhizosolenia) having cell volumes comparable with those of small fecal pellets.

CYCLOPOID COPEPODS

Oithona

There is limited information on the feeding habits of several

species of Oithona, and much of it is contradictory. Taken as a

whole, however, members of the genus apparently are omnivorous.
Lebour (1922) concluded that O. similis is primarily herbivorous after finding diatom remains in its gut. Murphy (1923)

cultured O. nana on pieces of ground kelp, although she stated that

the guts of field-caught specimens contained diatom and copepod

remains. Gauld (1966) and Marshall and Orr (1966) stated that O.

similis feeds little on phytoplankton, and then only on large cells.

Since Gauld (1966) and Marshall and Orr (1966) found that O.

similis eats the nauplii of other copepod species, they concluded

that the species is primarily carnivorous. Timonin (1969, 1971)

reached the same conclusion for several other species of Oithona.

However, Petipa et a1. (1970) considered Oithona copepodites to

be omnivorous and only adults to be exclusively carnivorous.

Oithona nana can eat either Acartia or Calanus nauplii (Lampitt

1978; Lampitt and Gamble 1982), but it can also ingest unialgal

cultures of five phytoplankton species ranging in size from 2.7 to

67.0 !lm in diameter (Lampitt and Gamble 1982). Likewise,

Hargrave and Geen (1970) and Poulet (1978) found that O. similis

fed upon natural particulate assemblages and ingested various

phytoplankton species, but Poulet suggested that O. similis may be

an inefficient grazer. This may partly explain why Dagg and

Turner (1982) calculated that a copepod community dominated

by O. similis was grazing> 100% of the primary production/day

in continental shelf waters. Nonetheless, diets consisting solely of

phytoplankton cultures have been used to rear O. nana (Haq

1965) and O. colcarva (Lonsdale 1981).

Corycaeus

Little is known of the feeding habits of Corycaeus. Lebour

(1922) found remains of diatoms, dinoflagellates, and coccolithophorids in C. anglicus guts, and she classified this species as

primarily herbivorous. Wickstead's (1962) examination of

preserved Indo-Pacific species of Corycaeus led him to conclude

that they were mainly carnivorous. Timonin (1969, 1971) also

classified the genus as carnivorous. Indeed, Gophen and Harris

(1981) found that C. anglicus ate Artemia nauplii, and Johnson

(1969 - cited in Gibson and Grice 1978) classified this species

as a raptorial carnivore with Acartia spp. as primary prey.

Nonetheless, Landry and Zakar (unpub1. data - cited in Landry

1981) found that stomachs of C. anglicus may be full of

phytoplankton during bloom conditions.

Oncaea

The feeding habits of all species of the genus Oncaea are largely

a matter of conjecture. Gienrikh (1958) mentioned that O. conifera guts contain coccoliths and remains of pennate diatoms, but

Wickstead (1962) classified the genus as carnivorous. Bernard

(1963a) and Wickstead (unpub1. data) (both cited by Marshall

1973) hypothesized that Oncaea is actually a temporary plankter,

and that it either feeds in bottom sediments (Bernard) or attaches

to, and feeds upon, gelatinous zooplankton (Wickstead). While

there are little data to support the hypothesis that Oncaea is not

truly holoplanktonic, Alldredge (1972) found that Oncaea mediterranea can feed upon abandoned appendicularian houses and the

microplankton attached to them.

PLANKTONIC HARPACTICOID COPEPODS

Many species of harpacticoids inhabit epibenthic or meiobenthic habitats, and it is likely that benthic harpacticoids are a major

portion of the "unidentified harpacticoid" component of the diets

9

of demersal larvae of many fishes. Some benthic harpacticoids

have been cultured (Omori 1973; Walker 1981) on a variety of

phytoplankton diets. In addition, feeding studies reveal that certain species ingest phytoplankton, protozoans, detritus (Harris

1977; Heinle et a1. 1977; Brown and Sibert 1977; Rieper and

Flotow 1981), and bacteria (Rieper 1978, 1982; Brown and Sibert

1977).

Information on the feeding habits of planktonic harpacticoids is

sparse. Lebour (1922) found phytoplankton and copepod remains

in the guts of Euterpina acuti/rons. This species has also been

cultured on a variety of unialgal or mixed phytoplankton cultures

(Bernard 1963b - cited in Omori 1973; Haq 1972; Nassogne

1970; Neunes and Pongolini 1965; Zurlini et a1. 1978). The only

information located on feeding by Microsetella is that M. rosea

guts contain coccoliths, green remains, and radiolarian spines

(Gienrikh 1958). Bjornberg (1965) found that Macrosetella gracilis requires the blue-green alga Trichodesmium as a substrate.

Roman (1978) confirmed this, and found that M. gracilis ingested

Trichodesmium at rates equivalent to 90-126% of its body

weight/day.

COPEPOD NAUPLII

All copepods, regardless of adult body size, begin life as small

nauplii. Copepods molt through six progressively larger naupliar

stages, followed by six copepodite stages, the sixth being the adult.

Although subadult copepods usually outnumber adults by orders

of magnitude in most marine waters (for example, compare

numbers in Turner 1982), the feeding habits of these juveniles,

particularly nauplii, are poorly known. Intuitively one might expect that copepod nauplii eat smaller phytoplankton cells than

adults, but this is not always the case.

Marshall and Orr (1956) fed Calanus finmarchicus nauplii on

cultures of small flagellates, coccolithophorids, chain-forming

diatoms, and dinoflagellates. Smaller naupliar stages usually did

not eat smaller sized cells than did larger naupliar stages. Rather,

some stages of nauplii ate the same types and sizes of food as

adults. Marshall and Orr found that naupliar stages I and II did not

feed, presumably because their anus had not yet formed. From

naupliar stage III onward, cells of up to 20 !lm in diameter were

eaten, but flagellates as small as 2-4 !lm in diameter were not

eaten.

Nauplii of other copepod species do not always exhibit these

same patterns. Mullin and Brooks (1967) found that Rhincalanus

nasutus nauplii could not eat some species of large diatoms which

were eaten by adults. Mullin and Brooks (1970a) fed unialgal

cultures of Ditylum brightwelli (a large diatom) and Thalassiosira

fluviatilis (a small diatom) to nauplii of R. nasutus and Calanus

helgolandicus. Calanus nauplii were unable to eat the large

diatom, but grew well when eating the small one. The reverse was

true for Rhincalanus nauplii, which are larger than those of

Calanus. Apparently Rhincalanus nauplii are large enough to capture and ingest the large diatom, but their filtering appendages are

too coarse to collect the small diatom. The success of naupliar

feeding is apparently important to the ultimate success of a

copepod. Mullin and Brooks (1970b) found that the body weights

of late-stage copepodites and growth efficiencies were functions

of food concentrations encountered by copepods as nauplii.

Nauplii of Calanus helgolandicus have been cultured on diets of

several dinoflagellate and chain-forming diatom species (Paffenhorer 1970, 1971, 1976a). Ingestion rates increased with both increasing food concentration and particle size. Unialgal diets of

chain-forming diatoms also have been used to rear nauplii of
Pseudocalanus elongatus and Temora longicornis (Paffenhiifer and

Harris 1976; Harris and Paffenhiifer 1976a, b).

Allan et al. (1977) examined the feeding of naupliar stages IlIVI of three estuarine copepods (Eurytemora affinis, Acartia tonsa,

and A. clausi) on natural particulates. In a few cases there was

evidence of size-selective feeding, but most feeding was unselective.

Paffenhiifer and Knowles (1978) offered nauplii of Eucalanus

pileatus, Temora stylifera, and T. turbinata mixtures of two chainforming and one large solitary (Rhizosolenia) diatom species.

Eucalanus pileatus nauplii larger than stage IV, copepodites, and

adults ingested similar percentages of the three phytoplankters.

However, the nauplii of the two Temora species ingested only one

of the chain-formers (Skeletonema costatum), and only nauplii

larger than stage IV ingested the other (Leptocylindrus danicus).

Ingestion of Rhizosolenia by Temora subadults did not commence

until copepodite stage II. Paffenhiifer and Knowles (1978) concluded that the size range of cells eaten by Temora nauplii was

4-20 Jim diameter, whereas that for Eucalanus nauplii was 4.5 to

> 100 Jim.

Fernandez (1979a, b) fed Calanus pacificus nauplii on mixtures

of culture 1 phytoplankton and artificial foods such as plastic

beads, pollen grains, or detritus from senescent algal cultures.

Phytoplankton were selected over artificial foods, but

phytoplankton cells> 45 Jim or < II Jim in longest dimension

were not eaten well, due to the inability of nauplii to manipulate

larger cells or to retain smaller ones. Fernandez found that ingestion rates were dependent upon algal size. Further, dinoflagellates,

regardless of size, were ingested at lower rates than diatoms.

In conclusion, most copepod nauplii appear to eat primarily

phytoplankton, the size and type of which varies with copepod

species and stage of nauplius. Thus far, however, the feeding

habits of nauplii of only about a dozen copepod species have been

examined. Considering the high numbers of nauplii usually present in the sea, the current lack of information on the dynamics of

feeding by copepod nauplii precludes realistic understanding of

pelagic ecosystem dynamics.

TINTINNIDS

Tintinnids are ciliate protozoans which appear to be primarily

herbivorous. Lebour (1922) found that Cittarocyclis serrata ate

eight species of dinoflagellates, one species of diatom, and two

species of tintinnids. Since the tintinnids and dinoflagellates were

present only in summer, and dinoflagellates were the main food

items of the tintinnids, Lebour classified C. serrata as a "peridinian feeder."

Stoecker et al. (1981) also found dinoflagellates to be important

in the diet ofFavell(l ehrenberghii. Successful culture of this tintinnid required dinoflagellate food. Further, in selective feeding experiments, dinoflagellates (II species) were always consumed, but

there was little if any grazing of nondinoflagellates (12 species of

diatoms, microflagellates, and chlorophytes). Selection of dinoflagellates was not on the basis of size alone. One dinoflagellate,

Amphidinium carterae was not eaten; this species has been suggested to produce ectocrines which repel grazers. Blackbourn

(1974), however, found that A. carterae, as well as many nondinoflagellates were eaten by various tintinnid species (see table 2 of

Rassoulzadegan and Etienne 1981 for a summary of Blackbourn's

unpublished work). Stoecker et al. (1981) concluded that F.

ehrenberghii is a specialized dinoflagellate predator which consumes few, if any, of the nondinoflagellates tested.

10

While the conclusion of Stoecker et al. (1981) may apply to F.

ehrenbergii, it clearly does not apply for some other tintinnids.

Gold (1968, 1969a, b, 1970, 1971, 1973) has cultured five other

species of tintinnids on diets of microflagellates. Only one of the

fi ve tintinnids required dinoflagellate food for successful culture.

Heinbokel (1978a) also found that five additional tintinnid

species ingested three species of microflagellates and one species

of coccolithophorid, either singly or in combination. For the three

tintinnid species studied most extensively, one showed increasing

ingestion rates with increasing food concentration, with no

plateau. The other two species exhibited increasing ingestion rates

with increasing food up to a saturation plateau. Since these tintinnids ingested 10-20% of their body weight per day, converted this

ration to tintinnid biomass with> 50% efficiency, and exhibited

doubling times of 12-24 h, Heinbokel (1978a) and Heinbokel and

Beers (1979) concluded that tintinnids are capable of substantial

grazing impact on nanoplankton. Capriulo and Carpenter (1983)

similarly concluded that the tintinnid populations of Long Island

Sound were responsible for ingesting 27% of the annual primary

productivity in the estuary. This community ingestion was equivalent to that calculated for the Long Island Sound copepod

population.

Rassoulzadegan and Etienne (1981) examined the feeding of

Stenosemella ventricosa on natural particulates, and reported this

tintinnid to feed on nanoplankton-sized particles « 27 Jim in

diameter) primarily in the 3-12 Jim size range. Ingestion rate was

a linear function of food concentration, with no saturation over

the range of food concentrations examined. The tintinnid ingested

about 66% of its body weight per day. After feeding for a time on

a c~rtain size range of particles, it adjusted its grazing pressure to

other size categories which had not been previously grazed.

Thus it appears that various species of tintinnids feed mainly on

nanoplankton-sized flagellates and/or dinoflagellates. Part of the

lack of agreement on tintinnid diets probably is due to the large

variation in size of lorica oral diameters of various tintinnid

species. For instance, the oral lorica diameter of Eutintinnus pectinis is 20 Jim (Turner and Anderson 1983), whereas that of

Favella panamensis is > 100 Jim (Turner unpubl. obs.). These differences are important since various tintinnid species have been

shown to ingest particles only < 41-45% of their lorica oral

diameter (Spittler 1973; Heinbokel 1978b). In addition, other

phytoplankters such as the flagellate Olisthodiscus luteus appear to

be unsuitable as food for some tintinnids for reasons related to

toxicity rather than size (Verity and Stoecker 1982).

There has been abundant speculation that tintinnids are a major

trophic link between the nanoplankton upon which they feed, and

larger plankters such as copepods and larval fish which might prey

upon tintinnids (Beers and Stewart 1967; Heinbokel and Beers

1979; Pomeroy 1974; Parsons and Takahashi 1973). Apparently

the first synoptic demonstration of this is the observation of

Turner and Anderson (1983) that Eutintinnus pectinis cells containing the microflagellate Chroomonas amphioxea were ingested

by the copepod Acartia hudsonica.

APPENDICULARIANS

Appendicularians are gelatinous tunicates which are ubiquitous

throughout the ocean. They feed by a ciliary mucoid mechanism

using an external mucus filter. A spherical mucus "house"

expands around the animal, through which water is pumped by

action of the animal's tail, and particles are trapped on incurrent

filters. When the "house" becomes clogged, the animal discards it
and secretes a new one (Alldredge 1977; Alldredge and Madin

1982).

Appendicularians feed primarily on nanoplankton-sized particles (Alldredge 1977, 1981). They can retain particles as small

as 0.1 ~m in diameter, and particles < 2 ~m are the largest component of their gut contents (Alldredge and Madin 1982). PaffenhOfer (1973, 1976c) has cultured Oikopleura dioica and Frittilaria

borealis for multiple generations on diets of the microflagellates

Isochrysis galbana and Monochrysis lutheri. and the small diatom

Thalassiosira pseudonana. Oikopleura dioica also exhibits substantial grazing on bacterioplankton (King 1982; King et al. 1980).

Alldredge (1981) measured in situ filtration rates of two appendicularian species and found that, compared with filtration rates

of other zooplankters, those of appendicularians were high. Multiplying these rates by the number of animals present, Alldredge

found that appendicularians could filter clear up to 38% of each

cubic meter per day, a rate which exceeded particle replacement

by phytoplankton growth.

Appendicularian growth rates are high, and populations of

these animals can "explode" in response to bacterioplankton and

nanophytoplankton blooms (King 1982; PaffenhOfer 1976c;

Alldredge and Madin 1982). Since appendicularians feed on

nanoplankton-sized particles, and are directly consumed by larval

fish (Shelbourne 1953, 1957, 1962; Uotani et al. 1981; Wyatt

1974), they provide a major trophic link between the lower and

upper parts of the pelagic food chain.

MARINE CLADOCERANS

Although most c1adoceran species live in freshwater, three

genera (Podon. Evadne. and Penilia) comprising seven species inhabit the sea. Due to parthenogenic reproduction, c1adoceran

populations can "explode" under favorable conditions and may

become numerically dominant zooplankters at a given place and

time (Della Croce and Venugopal 1973; Gieskes 1971; Onbe

1977).

The existing literature on the feeding habits of marine

c1adocerans is sparse and somewhat contradictory. For instance,

Conover (1978a: 272-273) stated that "Evadne andPodon seem to

feed largely on discrete particles and perhaps detritus... but no

experimental observations have been made." Examination of

feeding appendages prompted Nival and Ravera (1979) to suggest

that Evadne spinifera possesses the ability to catch and hold

animal prey or large algal cells, although no confirming observations were reported. Lebour (1922) found green remains of

Phaeocystis in the guts of Evadne normanni and soft brown remains with no apparent structure in the guts of Podon intermedius.

Bainbridge (1958) also reported unrecognizable remains in the

guts of E. normanni. Pavlova (1967) found no phytoplankton remains in the guts of E. spinifera and E. tergestina. Since some of

the particles were "very much like chitin," Pavlova concluded

that Evadne eats zooplankton and detritus. White (1980) observed

that the gut contents of E. normanni collected during a bloom of

the red tide dinoflagellate Gonyaulax excavata contained "yellowbrown material consisting of chromatophore-like bodies" which

he assumed to be remains of G. excavata. The c1adocerans also

contained intact cells of the large (up to 300 ~m diameter) diatom

Coscinodiscus asteromphalus.

The only other marine c1adoceran that has received any study is

Penilia avirostris. which Pavlova (1959) stated is an indiscriminant filter feeder utilizing only particles <8 ~m in diameter. Gore

(1980) reported, however, that this species ingested glass beads of

up to 50 ~m in diameter but preferred beads < 20 ~m. Several

II

Soviet investigators have stated that P. avirostris feeds on small

phytoplankton and detritus, but primarily on bacteria (Pavlova

1959; Pavlova and Sorokin 1970 - cited in Conover 1978a;

Sorokin et al. 1970; Pavlova et al. 1973; Sorokin 1973,1981;

Petipa et al. 1973). Nonetheless, since copepods as large as Rhincalanus cornutus (body length = 2.7 mm for males, 3.6 mm for

females), Eucalanus attenuatus (body length = 3 mm for males,

4-5 mm for females), and Clausocalanus mastigophorus (body

length = 1.0-1.5 mm for males, 1.2-1.8 mm for females) (lengths

from Rose 1933 and Frost and Fleminger 1968) also have been

stated to ingest 10-75% of their body carbon per day as bacteria

(Sorokin 1981), the claims that c1adocerans eat primarily bacteria

appear deserving of further investigation.

In summary, there is disagreement about, or extreme variability

in, the nature and sizes of food eaten by marine c1adocerans. The

paucity of knowledge on the feeding habits of these animals is

reflected by the fact that in his 278-page review of feeding by

marine animals, Conover (1978a) devoted nine sentences (p. 272,

273, 293, 294) to summarizing existing knowledge on marine

c1adoceran feeding.

BARNACLE NAUPLII

Barnacle nauplii appear primarily herbivorous. Lebour (1922)

found that guts of several species contained remains of green

flagellates, diatoms, and Phaeocystis. Moyse (1963) compared the

suitability as food of unialgal cultures of 17 species of microflagellates, dinoflagellates, and diatoms for nauplii of 5 barnacle

species. The palatability of various phytoplankters to nauplii of

different barnacles varied. Two dinoflagellates, Amphidinium and

Prymesium. were unsuitable food for all nauplii, and Chlorella and

Dunaliella. while eaten, did not allow nauplii to develop completely. Balanus nauplii could be reared on diatoms but not on

flagellates; for Chthamalus nauplii the reverse was true. Eliminius

nauplii grew well on a wide range of phytoplankters. Nauplii of

the oceanic barnacle Lepas grew well on diets offlagellates but not

on diets of diatoms. Moyse (1963) suggested that growth was better because flagellates are usually more abundant than diatoms in

oceanic waters. Conversely, nauplii of the arcticlboreal barnacle

Verruca grew best on diatoms, which usually dominate the

phytoplankton in arctic/boreal waters. Moyse suggested that the

unsuitability of chain-forming diatoms as food for nauplii of some

species was due to their clogging the finely setulated filtering appendages of the nauplii. Clogging did not occur when these

species fed upon small flagellates. Moyse concluded that nauplii

of different barnacle species have different diet requirements.

Other species of barnacles have been reared successfully

through naupliar stages on unialgal or mixed diets of diatoms and

microflagellates (Wisely 1960; Moyse 1960; Barnes and Costlow

1961; Tighe-Ford et al. 1970; Karande 1974a, b). Further,

Costlow and Bookout (1957, 1958) reared two other barnacle

species on diets of Chlamydomonas. but found that completion of

development required addition of starfish (Arbacia) eggs in the

later stages of development. Aside from the observations of

Costlow and Bookout, however, it appears that most barnacle

nauplii are primarily herbivorous although the types of phytoplankton eaten vary with barnacle species.

GASTROPOD LARVAE

Most gastropod larvae examined feed on phytoplankton. Gut

contents contained remains of a variety of diatoms (Lebour 1922),
and Thompson (1959) fed the diatom Phaeodactylum tricornutum

to larvae of numerous gastropod species.

Fretter and Montgomery (1968) examined the feeding of veligers of 19 gastropod species on cultures of 3 dinoflagellate, 7

microflagellate, 4 nanochlorophyte, and 4 diatom species. Some

phytoplankters clearly were unsuitable food. The flagellate

Olisthodiscus was either undigested or, for some veligers, was toxic. Monochrysis was also lethal to veligers of four species.

However, most of the phytoplankters were readily ingested, except for large (> 30 /-1m cell diameter) or chain-forming species,

which clogged the preoral cilia of some veligers.

Mapstone (1970) fed unialgal cultures of the dinoflagellate Exuviaella baltica and two microflagellates to veligers of Nassarius

reticulatus and Crepidula fornicata. Based on activity of veligers

while feeding, the microflagellates were good food for N.

reticulatus veligers but E. baltica was not. The reverse was true for

C. fornicata veligers. Pilkington and Fretter (1970), using the

same phytoplankters as food, examined growth of veligers of the

same species and reached the same conclusions. Of five additional

microflagellate species, four were poor foods, and of these two

were toxic. A diatom was also poor food because the frustules

were too large for convenient ingestion, and they appeared to

stimulate gut pulsations, which Pilkington and Fretter considered

indicative of irritation. Pilkington and Fretter also found that

veligers of both gastropods ingested several strains of bacteria.

The limited available information suggests that gastropod larvae are primarily phytoplankton, and possible bacterioplankton,

feeders.

LIMACINA

Limacina is a genus of pteropod (holoplanktonic molluscs).

Although gymnosomatous or nonshelled pteropods appear to prey

principally, or exclusively, upon thecosomatous or shell-bearing

pteropods (Lalli 1970, 1972; Conover and Lalli 1972, 1974),

thecosomes are ciliary-mucoid filter feeders (Yonge 1926; Morton 1954; Lalli 1970; Gilmer 1972, 1974; Silver and Bruland

1981). The guts of Limacina retroversa, a thecosome, contained

remains of diatoms (Lebour 1922) and Morton (1954) assumed

that it ate dinoflagellates and nanoplankton as well. Paranjape

(1968) found that L. helicina guts contained some remains of

chain-forming diatoms, but that remains of dinoflagellates and

tintinnids were most abundant.

These limited observations suggest that species of Limacina are

primarily herbivorous filter feeders.

DISCUSSION

Several points should emerge from this review: 1) Most zooplankters, particularly copepods, appear to be omnivorous feeders;

2) we still have too little information on the exact mechanisms by

which many zooplankters capture and select (or even if some select) various types of food; and 3) we know next to nothing about

the feeding habits in nature of some very abundant zooplankters.

Since the time of Lebour (1922) it has been known that most

copepods are omnivorous, ingesting plant, animal, and detrital

foods. In the absence of detailed examinations of gut contents and

quantitative comparisons of ingestion of various types of food,

attempts to use mouthpart structures to classify copepods as herbivores or carnivores (Itoh 1970; Arashkevich 1969) appear unwarranted. Indeed, some copepods thought to be primarily carnivorous (pontellids) have been shown to ingest a variety of

12

phytoplankters (Turner 1978; Conley 1983) and some "typical"

herbivores such as Calanus have been shown to feed as carnivores

(Corner et al. 1974, 1976; Landry 1980, 1981). Missing for most

species are not only qualitative descriptions of food items ingested

in nature, but also quantitative comparisons using natural food of

the relative importance of herbivorous and carnivorous feeding.

The studies of Paffenh6fer and Knowles (1980), Landry (1981),

and Conley (1983) represent steps in that direction. Observations

by Landry (1981) that Calanus can switch from eating phytoplankton to eating zooplankton, and vice versa, suggest that

copepod feeding preferences are quite plastic. If this pattern is

generally applicable, the idea that alteration of phytoplankton

species assemblages may result in adverse trophodynamic alteration of zooplankton and ichthyoplankton assemblages appears

simplistic. The true pattern may be that most zooplankters, including larval fish, eat their "favorite" food if it is present and

abundant, and if not, they make do by eating something else.

Many classic concepts of the manner by which copepods gather

food have recently been shown to require revision. The

"textbook" description of copepods using bristled appendages

such as the second maxillae as open rakes to strain particles from

water by filtration (Esterly 1916; Cannon 1928; Lowndes 1935;

Marshall and Orr 1955c; Gauld 1966; Boyd 1976; Nival and

Nival 1976; Frost 1977) has been altered by cinematographic

studies which show that these appendages behave as solid paddles,

capturing whole parcels of water containing food particles which

are brought to the mouth with hardly touching setae or setules

(Alcaraz et al. 1980; Koehl and Strickler 1981; PaffenhOfer et al.

1982; Strickler 1982). There also is increasing evidence that

"selection" of various types of food, or even induction of feeding,

has a chemosensory component (Poulet and Marsot 1978, 1980;

Friedman and Strickler 1975; Cox and Willason 1981). Although

food particle size may broadly dictate what types of food items

can be physically handled by the feeding apparatus of a zooplankter, and although certain sizes of food items appear to be more

efficiently ingested than others, it is becoming clear that selective

feeding, if and when it occurs, is not solely on the basis of particle

size. Indeed, recent experimental (Harris 1982; Huntley 1981)

and gut content (Hayward 1980) studies have revealed that different sized copepods compete for the same broad size array of

food items. In view of these observations, and those of Harbison

and McAlister (1980) on artifacts associated with the use of electronic particle counters to study selective feeding by zooplankters

upon natural particulate assemblages, it may be that we need to

"unlearn" some of what we presently "know" of zooplankton

feeding patterns.

The largest problem with the present state of knowledge of

zooplankton feeding is that we have very little information on the

feeding habits in nature of some ubiquitous taxa that are usually

the most abundant. Particularly, these include cyclopoid

copepods, nauplii of almost all types of copepods, and tintinnids

and other ciliates. The same applies to other taxa such as marine

cladocerans, appendicularians, thaliaceans, and various types of

meroplankters which, although less ubiquitously abundant, exhibit sporadic and localized population increases. Limited information suggests that the trophodynamic impact of such population

pulses can be substantial (Seki 1973; Wiebe et al. 1979).

In conclusion, I return to the thesis presented in the introduction, namely that understanding of fishery dynamics requires

understanding of the dynamics of food webs supporting larval

fish. Sandwiched between a need to understand responses of

planktonic primary producers to climatic, oceanographic, and
anthropogenic-induced variability, and a need to understand

trophic interactions between larval fish and their prey, is a critical

need to understand dynamics and patterns of zooplankton feeding.

ACKNOWLEDGMENTS

I wish to thank D. E. Hoss for the invitation to present a review

of zooplankton feeding habits at a session entitlted "Dynamics of

Larval Fish Food Webs" presented at the 72nd Annual Meeting of

the American Fisheries Society, Hilton Head, S.C., 19-22

September 1982. This invitation provided the stimulus for

preparation of this review. J. J. Govoni straightened out 6.5

decades ofrevisions of fish taxonomy. G. Thayer, P. Tester, and an

anonymous reviewer provided helpful criticism of the manuscript.

Preparation of this review was supported by a contract from the

Ocean Assessments Division, National Ocean Service, NOAA, to

the Southeast Fisheries Center's Beaufort Laboratory, NMFS.

LITERATURE CITED

ADAMS, J. A., and J. H. STEELE.

1966. Shipboard experiments on the feeding of Calanus finmarchicus (Gunnerus).In H. Barnes (editor), Some contemporary studies in marine science,

p. 19-35. Allen and Unwin Ltd., Lond.

ALCARAZ, M., G.-A. PAFFENHOFER, and J. R. STRICKLER.

1980. Catching the algae: A first account of visual observations on filterfeeding calanoids. In W. C. Kerfoot (editor), Evolution and ecology of zooplankton communities, p. 241-248. University Press of New England,

Hanover, N.H.

ALLAN, J. D., S. RICHMAN, D. R. HEINLE, and R. HUFF.

1977. Grazing in juvenile stages of some estuarine calanoid copepods. Mar.

BioI. (Berl.) 43:317 -331.

ALLDREDGE,A.L.

1972. Abandoned larvacean houses: A ufJique food source in the pelagic

environment. Science (Wash., D.C.) 177:885-887.

1977. House morphology and mechanisms of feeding in the Oikopleuridae

(Tunicata, Appendicularia). J. Zooi. (Lond.) 181:175-188.

198 I. The impact of appendicularian grazing on natural food concentrations

in situ. Limnoi. Oceanogr. 26:247-257.

ALLDREDGE, A. L., and L. P. MADIN

1982. Pelagic tunicates: Unique herbivores in the marine plankton. BioScience 32:655-663.

ANRAKU,M.

1964a. Influence of the Cape Cod Canal on the hydrography and on the copepods in Buzzards Bay and Cape Cod Bay, Massachusetts. II. Respiration and

feeding. Limnoi. Oceanogr. 9:195-206.

I 964b. Some technical problems encountered in quantitative studies of grazing and predation by marine planktonic copepods. J. Oceanogr. Soc. Jpn.

20:221-231.

ANRAKU, M., and M. OMORI.

1963. Preliminary survey of the relationship between the feeding habit and

the structure of the mouth-parts of marine copepods. Limnoi. Oceanogr.

8:116-126.

ARASHKEVICH, Yeo G.

1969. The food and feeding of copepods in the northwestern Pacific.

Oceanology 9:695-709.

ARTHUR, D. K.

1976. Food and feeding of larvae of three fishes occurring in the California

Current, Sardinops sagax. Engraulis mordax, and Trachurus

symme/ricus. Fish. Bull., U.S. 74:517-530.

BAINBRIDGE, V.

1958. Some observations on Evadne nordmanni Loven. J. Mar. BioI. Assoc.

U.K. 37:349-370.

1965. A preliminary study of Sebasles larvae in relation to the planktonic

environment of the Irminger Sea. Spec. Pubi. Int. Comm. Northwest Ati.

Fish. 6:303-308.

BAINBRIDGE, V., and D. C. T. FORSYTH.

1971. The feeding of herring larvae in the Clyde. Rapp. P.-V. Reun. Cons.

Int. Explor. Mer 160:104-113.

BARNES, H., and J. D. COSTLOW, Jr.

1961. The larval stages of Balanus balanus (L.) Da Costa. J. Mar. BioI.

Assoc. U.K. 41:59-68.

13

BARTRAM, W. C.

198 I. Experimental development of a model for the feeding of neritic copepods on phytoplankton. J. Plankton Res. 3:25-51.

BEERS, J. R., and G. L. STEWART.

1967. Micro-zooplankton in the euphotic zone at five locations across the

California Current J. Fish. Res. Board Can. 24:2053-2068.

BERNARD, M.

1963a. Adaptation de quelques copepodes pelagiques Mediteraneens adifferents milieux de survie en aquarium. Rapp. P.-V. Reun. Comm. Int. Explor.

Sci. Mer Mediterr. 16:165-176.

1963b. Observations sur la biologie en aquarium de Eu/erpina aculifrons.

copepode pelagique. Rapp. P.-V. Reun. Comm. Int. Explor. Sci. Mer

Mediterr. 17:545.

BERNER, A.

1962. Feeding and respiration in the copepod Temora longicornis (Miiller).

J. Mar. BioI. Assoc. U.K. 42:625-640.

BJ~RKE, H.

1971. The food of herring larvae of Norwegian spring spawners. Rapp.

P.-V. Reun. Cons. Int. Explor. Mer 160:101-103.

1978. Food and feeding of young herring larvae of Norwegian spring spawners. Fiskeridir. Skr. Ser. Havunders. 16:405-422.

BJORNBERG, T. K. S.

1965. Observations on the development and the biology of the Miracidae

Dana (Copepoda:Crustacea). Bull. Mar. Sci. 15:512-520.

BLACKBOURN, D. J.

1974. The feeding biology of tintinnid protozoa and some other inshore

microzooplankton. Ph.D. Thesis, Univ. British Columbia, Vancouver.

BLAXTER, J. H. S.

1965. The feeding of herring larvae and their ecology in relation to feeding.

Calif. Coop. Oceanic Fish. Invest. Rep. 10:79-88.

BLAXTER, J. H. S. (editor).

1974. The early life history of fish. Springer-Verlag, N.Y., 765 p.

BOWERS, A. B., and D. I. WILLIAMSON.

1951. Food of larval and early post-larval stages of autumn-spawned herring

in Manx waters. Annu. Rep. Mar. BioI. Stn. Port Erin 63:17-26.

BOYD, C. M.

1976. Selection of particle sizes by filter-feeding copepods: A plea for reason.

Limnoi. Oceanogr. 21:175-180.

BOYD, C. M., S. L. SMITH, and T. J. COWLES.

1980. Grazing patterns of copepods in the upwelling system off Peru.

Limnoi. Oceanogr. 25:583-596.

BRADFORD, J. M.

1976. Partial revision of the Acania subgenus Acar/iura (Copepoda: Calanoida: Acartiidae). N.Z. J. Mar. Freshw. Res. 10:159-202.

BROWN, T. J., and J. R. SIBERT.

1977. Food of some benthic harpacticoid copepods. J. Fish. Res. Board Can.

34: 1028-1031.

BUTLER, E. I., E. D. S. CORNER, and S. M. MARSHALL.

1969. On the nutrition and metabolism of zooplankton. VI. Feeding efficiency of Calanus in terms of nitrogen and phosphorus. 1. Mar. BioI. Assoc.

U.K. 49:977-1001.

1970. On the nutrition and metabolism of zooplankton. VII. Seasonal survey

of nitrogen and phosphorus excretion by Calanus in the Clyde Sea-area. J.

Mar. BioI. Assoc. U.K. 50:525-560.

CAHOON, L. B.

1981. Reproductive response of Acania /onsa to variations in food ration and

quality. Deep-Sea Res. 28:1215-1221.

CANNON, H. G.

1928. On the feeding mechanism of the copepods, Calanus finmarchicus and

Diaptomus gracilis. Br. J. Exp. BioI. 6:131-144.

CAPRIULO, G. M., and E. 1. CARPENTER.

1983. Abundance, species composition and feeding impact of tintinnid

micro-zooplankton in central Long Island Sound. Mar. Ecoi. Prog. Ser.

10:277-288.

CHECKLEY, D. M., Jr.

1980. The egg production of a marine planktonic copepod in relation to its

food supply: Laboratory studies. Limnoi. Oceanogr. 25:430-446.

CIECHOMSKI, J. D. de.

1967. Investigations of food and feeding habits of larvae and juveniles of the

Argentine anchovy Engraulis anchoita. Calif. Coop. Oceanic Fish. Invest.

Rep. 11 :72-8 I.

CLARKE, G. L., and D. D. BONNET.

1939. The influence of temperature on the survival, growth and respiration of

Calanusfinmarchicus. BioI. Bull. (Woods Hole) 76:371-383.

CLARKE, G. L., and S. S. GELLIS.

1935. The nutrition of copepods in relation to the food-cycle of the sea.
BioI. Bull. (Woods Hole) 68:231-246.

COLLETIE, B. B.

1978. Adaptations and systematics of the mackerels and tunas. In G. D. Sharp

and A. E. Dizon (editors), The physiological ecology of tunas, p. 7-88.

Acad. Press, N.Y.

CONLEY, W. J.

1983. A comparison of herbivory and carnivory for two coastal marine copepods. M.S. Thesis, Southeastern Massachusetts University, North Dartmouth, I 15 p.

CONOVER, R. J.

1956. Oceanography of Long Island Sound, 1952-/954. VI. Biology of Acartia clausi and A. tonsa. Bull. Bingham Oceanogr. Collect. Yale Univ.

15:156-233.

1959. Regional and seasonal variation in the respiratory rate of marine copepods. Limnol. Oceanogr. 4:259-268.

1960. The feeding behavior and respiration of some marine planktonic

Crustacea. BioI. Bull. (Woods Hole) 119:399-415.

1966a. Feeding on large particles by Calanus hyperboreus (Kriiyer). In H.

Barnes (editor), Some contemporary studies in marine science, r. 187 -194.

Allen and Unwin Ltd., Lond.

1966b. Assimilation of organic matter by zooplankton. Limnol. Oceanogr.

II :338-345.

1966c. Factors affecting the assimilation of organic matter by zooplankton

and the question of superfluous feeding. Limnol. Oceanogr. II :346-354.

1971. Some relations between zooplankton and Bunker C oil in Chedabucto

Bay following the wreck of the tanker Arrow. J. Fish. Res. Board Can.

28:1327-1330.

1978a. Transformation of organic matter. In O. Kinne (editor), Marine

ecology, Vol. IV, Dynamics, p. 221-499. John Wiley & Sons, N.Y.

1978b. Feeding interactions in the pelagic zone. Rapp. P.-V. Reun. Cons.

Int. Explor. Mer 173:66-76.

CONOVER, R. J., and C. M. LALLI.

1972. Feeding and growth in Clione limacina (Phipps), a pteropod mollusc.

J. Exp. Mar. BioI. Ecol. 9:279-302.

1974. Feeding and growth in Clione limacina (Phipps), a pteropod mollusc.

II. Assimilation, metab~lism, and growth efficiency. J. Exp. Mar. BioI.

Ecol. 16:131-154.

CONWAY, D. V. P.

1980. The food of larval blue whiting, Micromesistius poutassou (Risso), in the

Rockall area. J. Fish BioI. 16:709-723.

CORKETI, C. J.

1967. Technique for rearing marine calanoid copepods in laboratory conditions. Nature (Lond.) 216:58-59.

1970. Techniques for breeding and rearing marine calanoid copepods.

Helgol. wiss. Meeresunters. 20:318-324.

CORKETI, C. J., and I. A. McLAREN.

1978. The biology of Pseudocalanus. Adv. Mar. BioI. 15:1-231.

CORKETI, C. J., and E. J. ZILLIOUX.

1975. Studies on the effect of temperature on the egg laying of three species

of calanoid copepods in the laboratory (Acartia tonsa, Temora longicomis and

Pseudocalanus elongatus). Bull. Plankton Soc. Jpn. 21:77-85.

CORNER, E. D. S.

1961. On the nutrition and metabolism of zooplankton, I. Preliminary

observations on the feeding of the marine copepod, Calanus helgolandicus

(Claus). J. Mar. BioI. Assoc. U.K. 41 :5-16.

CORNER, E. D. S., C. B. COWEY, and S. M. MARSHALL.

1967. On the nutrition and metabolism of zooplankton. V. Feeding efficiency

of Calanus finmarchicus. J. Mar. BioI. Assoc. U.K. 47:259-270.

CORNER, E. D. S., R: N. HEAD, and C. C. KILVINGTON.

1972. On the nutrition and metabolism of zooplankton. VIII. The grazing of

Biddulphia cells by Calanus helgolandicus. J. Mar. BioI. Assoc. U.K.

52:847-861.

CORNER, E. D. S., R. N. HEAD, C. C. KILVINGTON, and S. M. MARSHALL.

1974. On the nutrition and metabolism of zooplankton. IX. Studies relating to

the nutrition of overwintering Calanus. J. Mar. BioI. Assoc. U.K.

54:319-331.

CORNER, E. D. S., R. N. HEAD, C. C. KILVINGTON, and L. PENNYCUICK.

1976. On the nutrition and metabolism of zooplankton. X. Quantitative

aspects of Calanus helgolandicus feeding as a carnivore. J. Mar. BioI. Assoc.

U.K. 56:345-358.

COSTLOW, J.D., Jr., and C. G. BOOKOUT.

1957. Larval development ofBalanus ebumeus in the laboratory. BioI. Bull.

(Woods Hole) 112:313-324.

1958. Larval development of Balanus amphitrite var. denticulata Broch reared

in the laboratory. BioI. Bull. (Woods Hole) 114:284-295.

14

COVILL, R. W.

1959. Food and feeding habits of larvae and postlarvae of Ammodytes americanus, 1952-1955. Bull. Bingham Oceanogr. Collect. Yale Univ.

17:125-146.

COWEY, C. B., and E. D. S. CORNER.

1963. On the nutrition and metabolism of zooplankton. II. The relationship

between the marine copepod Calanus helgolandicus and particulate material

in Plymouth sea water, in terms of amino acid composition. J. Mar. BioI.

Assoc. U.K. 43:495-5 I I.

COWLES, T. J.

1979. The feeding response of copepods from the Peru upwelling system:

Food size selection. J. Mar. Res. 37:601-622.

COWLES, T. J., and J. R. STRICKLER.

1983. Characterization of feeding activity patterns in the planktonic copepod

Centropages typicus Kroyer under various food conditions. Limnol.

Oceanogr. 28:106-115.

COX, J. L., and S. W. WILLASON.

1981. Laminarinase induction in Calanus pacificus. Mar. BioI. Lett.

2:307-311.

CUSHING, D. H.

1958. The effect of grazing in reducing the primary production: A review.

Rapp. P.-V. Reun. Cons. Int. Explor. Mer 144:149-154.

1964. The work of grazing in the sea. In D. J. Crisp (editor), Grazing in

terrestrial and marine environments, p. 207-225. Blackwell Sci. Publ.,

Oxford.

CUSHING, D. H., and T. VUCETIC.

1963. Studies on a Calanus patch. III. The quantity of food eaten by Calanus

finmarchicus. J. Mar. BioI. Assoc. U.K. 43:349-371.

DAGG, M.

1977. Some effects of patchy food environments on copepods. Limnol.

Oceanogr. 22:99-107.

1978. Estimated, in situ, rates of egg production for the copepod Centropages

rypicus (Krl'lyer) in the New York Bight. J. Exp. Mar. BioI. Ecol.

34: 183-196.

DAGG, M., T. COWLES, T. WHITLEDGE, S. SMITH, S. HOWE, and D. JUDKINS.

1980. Grazing and excretion by zooplankton in the Peru upwelling system

during April 1977. Deep-Sea Res. 27:43-59.

DAGG, M. J., and D. W. GRILL.

1980. Natural feeding rates of Centropages rypicus females in the New York

Bight. Limnol. Oceanogr. 25:597-609.

DAGG, M. J., and J. T. TURNER.

1982. The impact of copepod grazing on the phytoplankton of Georges Bank

and the New York Bight. Can. J. Fish. Aquat. Sci. 39:979-990.

DAGG, M. J., J. VIDAL, T. E. WHITLEDGE, R. L. IVERSON, and J. J. GOERING.

1982. The feeding, respiration, and excretion of zooplankton in the Bering

Sea during a spring bloom. Deep-Sea Res. 29:45-63.

DAKIN, W. J.

1908. Notes on the alimentary canal and food of the Copepoda. Int. Rev.

Gesamten Hydrobiol. Hydrogr. 1:772-782.

DAVIS, C. C.

1977. Sagitta as food for Acartia. Astarte 10: 1-3.

DEASON, E. E.

1980a. Potential effect of phytoplankton colony breakage on the calculation

of zooplankton filtration rates. Mar. BioI. (Berl.) 57:279-286.

1980b. Grazing of Acartia hudsonica (A. clausi) on Skeletonema costatum in

Narragansett Bay (USA): Influence of food concentration and temperature.

Mar. BioI. (Berl.) 60:101-113.

DELLA CROCE, N., and P. VENUGOPAL.

1973. Pen ilia avirostris Dana in the Indian Ocean (Cladocera). Int. Rev.

Gesamten Hydrobiol. 58:713-721.

DETWYLER, R., and E. D. HOUDE.

1970. Food selection by laboratory-reared larvae of the scaled sardine

Harengula pensacolae (Pisces, Clupeidae) and the bay anchovy Anchoa mitchilli (Pisces, Engraulidae). Mar. BioI. (Berl.) 7:214-222.

DIGBY, P. S. B.

1954. The biology of the marine planktonic copepods of Scoresby Sound,

East Greenland. J. Anim. Ecol. 23:298-338.

DONAGHAY, P. L., and L. F. SMALL.

1979a. Long-term food modification by Acartia clausi: A preliminary view.

Mar. BioI. (Berl.) 52:129-136.

1979b. Food selection capabilities of the estuarine copepod Acartia clausi.

Mar. BioI. (Berl.) 52:137-146.

ELLERTSEN, B., P. SOLEMDAL, S. SUNDBY, S. T1LSETH, T. WESTGARD, and

V.I!lIESTAD.

198 I. Feeding and vertical distribution of cod larvae in relation to availability

of prey organisms. Rapp. P.-V. Reun. Cons. Int. Explor. Mer 178:3 I7-319.
ESAIAS, W. E., and H. C. CURL, Jr.

1972. Effect of dinoflagellate bioluminescence on copepod ingestion rates.

Limnol. Oceanogr. 17:901-906.

ESTERLY, C. O.

1916. The feeding habits and food of pelagic copepods and the question of

nutrition by organic substances in solution in the water. Univ. Calif. Publ.

Zool. 16:171-184.

FERNANDEZ, F.

1979a. Nutrition studies in the nauplius larva ofCa/anus pacificus (Copepoda:

Calanoida). Mar. BioI. (Berl.) 53:131-147.

1979b. Particle selection in the nauplius of Calanus pacificus. J. Plankton

Res. 1:313-328.

FLEMINGER, A., and K. HULSEMANN.

1973. Relationship of Indian Ocean epiplanktonic calanoids to the world

oceans. In B. Zeitzschel (editor), The biology of the Indian Ocean, p.

339-347. Springer-Verlag, N.Y.

FRETTER, V., and M.C. MONTGOMERY.

1968. The treatment of food by prosobranch veligers. J. Mar. BioI. Assoc.

U.K. 48:499-520.

FRIEDMAN, M. M.

1980. Comparative morphology and functional significance of copepod

receptors and oral structures. In W. C. Kerfoot (editor), Evolution and ecology

of zooplankton communities, p. 185-197. University Press of New

England, Hanover, N.H.

FRIEDMAN, M. M., and J. R. STRICKLER.

1975. Chemoreceptors and feeding in calanoid copepods (Arthropoda:

Crustacea). Proc. Natl. Acad. Sci. U.SA 72:4185-4188.

FROST, B. W.

1972. Effects of size and concentration of food particles on the feeding

behavior of the marine planktonic copepod Calanus pacificus. Limnol.

Oceanogr. 17:805-815.

1975. A threshold feeding behavior in Ca/anus pacificus. Limnol. Oceanogr.

20:263-266.

1977. Feeding behavior of Ca/anus pacificus in mixtures of food particles.

Limnol. Oceanogr. 22:472-491.

1980. The inadequacy of body size as an indicator of niches in the zooplankton.ln W. C. Kerfoot (editor), Evolution and ecology of zooplankton communities, p. 742-753. University Press of New England, Hanover, N.H.

FROST, B. W., and A. FLEMINGER.

1968. A revision of the genus Clausocalanus (Copepoda: Calanoida) with

remarks on distributional patterns in diagnostic characters. Bull. Scripps

Inst. Oceanogr., Univ. Calif. 12:1-235.

FROST, B. W., M. R. LANDRY, and R. P. HASSETT.

1983. Feeding behavior of large calanoid copepods Neocalanus cristatus and

N. p/umchrus from the subarctic Pacific Ocean. Deep-Sea Res. 30:1-13.

FULLER, J. L.

1937. Feeding rate of Calanus finmarchicus in relation to environmental conditions. BioI. Bull. (Woods Hole) 72:233-246.

FULLER, J. L., and G. L. CLARKE.

1936. Further experiments on the feeding of Calanus finmarchicus. BioI.

Bull. (Woods Hole) 70:308-320.

GAMBLE, J. C.

1978. Copepod grazing during a declining spring phytoplankton bloom in the

northern North Sea. Mar. BioI. (Berl.) 49:303-315.

GAMBLE, J. c., P. MacLACHLAN, N. T. NICOLL, and I. G. BAXTER.

1981. Growth and feeding of Atlantic herring larvae reared in large plastic

enclosures. Rapp. P.-V. Reun. Couns. Int. Explor. Mer 178:121-134.

GAUDY, R.

1974. Feeding four species of pelagic copepods under experimental conditions. Mar. BioI. (Berl.) 25:125-141.

GAULD, D. T.

1951. The grazing rate of planktonic copepods. J. Mar. BioI. Assoc. U.K.

29:695-706.

1953. Diurnal variations in the grazing of planktonic copepods. J. Mar.

BioI. Assoc. U.K. 31:461-474.

1966. The swimming and feeding of planktonic copepods. In H. Barnes

(editor), Some contemporary studies in marine science, p. 313-334. Allen

and Unwin Ltd., Lond.

GAULD, D. T., and J. E. G. RAYMONT.

1953. The respiration of some planktonic copepods. II. The effect of temperalUre. J. Mar. BioI. Assoc. U.K. 31:447-460.

GIBSON, V. R., and G. D. GRICE.

1978. The developmental stages of a species of Corycaeus (Copepoda: Cyclopoida) from Saanich Inlet, British Columbia. Can. J. Zool. 56:66-74.

GIENRIKH, A. K.

1958. On the nutrition of marine copepods in the tropical region. Dokl.

15

Akad. Nauk SSSR 119:229-232 (Engl. transl., Am. Inst. BioI. Sci., Wash.,

D.C.).

GIESKES, W. W. C.

1971. Ecology of the Cladocera of the North Atlantic and the North Sea,

1960-1967. Neth. J. Sea Res. 5:342-376.

GIFFORD, D. J., R. N. BOHRER, and C. M. BOYD.

1981. Spines on diatoms: Do copepods care? Limnol. Oceanogr.

26:1057-1061.

GILMER, R. W.

1972. Free-floating mucus webs: A novel feeding adaptation for the open

ocean. Science (Wash., D.C.) 176:1239-1240.

1974. Some aspects of feeding in thecosomatous pteropod molluscs. J. Exp.

Mar. BioI. Ecol. 15:127-144.

GOLD, K.

1968. Some observations on the biology of Tintinnopsis sp. J. Protozool.

15:193-194.

1969a. The preservation of tintinnids. J. Protozool. 16:126-128.

1969b. Tintinnida: Feeding experiments and lorica development. J. Protozool. 16:507-509.

1970. Cultivation of marine ciliates (Tintinnida) and heterotrophic flagellates. Helgol. wiss. Meeresunters. 20:264-271.

1971. Growth characteristics of the mass-reared tintinnid Tintinnopsis

beroidea. Mar. BioI. (Berl.) 8:105-108.

1973. Methods for growing tintinnida in continuous culture. Am. Zool.

13:203-208.

GOPHEN, M., and R. P. HARRIS.

1981. Visual predation by a marine cyclopoid copepoil, CO,ycaeus anglicus.·

J. Mar. BioI. Assoc. U.K. 61:391-399.

GORE, M. A.

1980. Feeding experiments on Pen ilia avirostris Dana (Cladocera: Crustacea).

J. Exp. Mar. BioI. Ecol. 44:253-260.

GOVONI, J. J., D. E. HOSS, and A. J. CHESTER.

1983. Comparative feeding of three species of larval fishes in the northern

Gulf of Mexico: Brevoortia patronus, Leiostomus xanthurus. and Micropogonias undu/arus. Mar. Ecol. Prog. Ser. 13:189-199.

GREVE, W., and T. R. PARSONS.

1977. Photosynthesis and fish production: Hypothetical effects of climatic

change and pollution. Helgol. wiss. Me~rsunters. 30:666-672.

HAQ, S. M.

1965. The larval development of Oithonina nana. J. Zool. (Lond.) 146:

555-566.

1972. Breeding of Euterpina acutifrons, a harpacticid copepod, with special

reference to dimorphic males. Mar. BioI. (Berl.) 15:221-235.

HARBISON, G. R., and V. L. McALISTER.

1980. Fact and artifact in copepod feeding experiments. Limnol. Oceanogr.

25:971-981.

HARGRAVE, B. T., and G. H. GEEN.

1970. Effects of copepod grazing on two natural phytoplankton populations.

J. Fish. Res. Board Can. 27:1395-1403.

HARRIS, R. P.

1977. Some aspects of the biology of the harpacticoid copepod, Scotlolana

canadensis (Willey), maintained in laboratory cullUre. Chesapeake Sci.

18:245-252.

1982. Comparison of the feeding behaviour of Calanus and Pseudocalanus

in two experimentally manipulated enclosed ecosystems. J. Mar. BioI.

Assoc. U.K. 62:71-91.

HARRIS, R. P., and G.-A. PAFFENHOFER.

1976a. Feeding, growth and reproduction of the marine planktonic copepod

Temora longicornis Muller. J. Mar. BioI. Assoc. U.K. 56:675-690.

1976b. The effect of food concentration on cumulative ingestion and growth

efficiency of two small marine planktonic copepods. J. Mar. BioI. Assoc.

U.K. 56:875-888.

HARVEY, H. W.

1937. Note on selective feeding by Calanus. J. Mar. BioI. Assoc. U.K. 22:

97-100.

HARVEY, H. W., L. H. N. COOPER, M. V. LEBOUR, and F. S. RUSSELL.

1935. Plankton production and its control. J. Mar. BioI. Assoc. U.K. 20:

407-442.

HAYWARD, T. L.

1980. Spatial and temporal feeding patterns of copepods from the North

Pacific central gyre. Mar. BioI. (Berl.) 58:295-309.

HEBERT, R., and S. A. POULET.

1980. Effect of modification of particle size of emulsions of Venezuelan

crude oil on feeding, survival and growth of marine zooplankton. Mar. Environ. Res. 4: 121-134.
HEINBOKEL, J. F.

1978a. Studies on the functional role of tintinnids in the Southern California

Bight. I. Grazing and growth rates in laboratory cultures. Mar. BioI. (Berl.)

47:177-189.

1978b. Studies on the functional role of tintinnids in the Southern California

Bight. II. Grazing rates of field populations. Mar. BioI. (Berl.) 47:191-197.

HEINBOKEL, J. F., and J. R. BEERS.

1979. Studies on the functional role of tintinnids in the Southern California

Bight. III. Grazing impact of natural assemblages. Mar. BioI. (Berl.)

52:23-32.

HEINLE, D. R.

1969. Cuhure of calanoid copepods in synthetic sea water. J. Fish. Res.

Board Can. 26: 150-153.

1970. PopUlation dynamics of exploited cultures of caJanoid copepods.

Helgol. wiss. Meeresunters. 20:360-372.

HEINLE, D. R., R. P. HARRIS, J. F. USTACH, and D. A. FLEMER.

1977. Detritus as food for estuarine copepods. Mar. BioI. (Berl.) 40:

341-353.

HJORT, J.

1914. Fluctuations in the great fisheries of northern Europe viewed in the

light of biological research. Rapp. P.-V. Reun. Cons. Int. Explor. Mer

20:1-228.

HODGKIN, E. P., and R. J. RIPPINGALE.

1971. Interspecies conflict in estuarine copepods. Limnol. Oceanogr.

16:573-576.

HONJO, S., and M. R. ROMAN.

1978. Marine copepod fecal pellets: Production, preservation and sedimentation. J. Mar. Res. 36:45-57.

HUNTLEY, M. E.

1981. Nonselective, non saturated feeding by three calanid copepod species in

the Labrador Sea. Limnol. Oceanogr. 26:831-842.

1982. Yellow water in La Jolla Bay, California, July 1980. II. Su,?ression of

zooplankton grazing. J. Exp. Mar. BioI. Ecol. 63:81-91.

HUREAU, J. C., and Th. MONOD (editors).

1973. Check-list of the fishes of the north-eastern Atlantic and of the Mediterranean, Vol. I. UNESCO, Paris, 683 p.

IKEDA, T.

1971. Preliminary shipboard culture experiments on the feeding and respiration of an oceanic copepod, Calanus eristatus, in the Bering Sea. [In

Engl.) Bull. Plankton Soc. Jpn. 18:5-14.

1977. Feeding rates of planktonic copepods from a tropical sea. J. Exp. Mar.

BioI. Ecol. 29:263-277.

ITOH, K.

1970. A consideration on feeding habits of planktonic copepods in relation to

the structure of their oral parts. [In Jpn., Engl. abstr.] Bull. Plankton Soc.

Jpn.17:1-10.

JOHNSON, G. F.

t969. A contribution [0 the biology ofCorycaeus anglicus, a marine cyclopoid

copepod. M.S. Thesis, University of California, San Diego, 68 p.

JUNE, F. C, and F. T. CARLSON.

197 I. Food of young Atlantic menhaden, Brevoonia tyrannus, in relation to

metamorphosis. Fish. Bull., U.S. 68:493-512.

KARANDE, A. A.

1974a. Balanus variegatus Darwin: The laboratory reared larvae compared

with Balanus amphitrite amphitrite Darwin (Cirripedia). Crustaceana

26:229-232.

I 974b. Larval development of the barnacle Tetraditella karandei reared in

the laboratory. BioI. Bull. (Woods Hole) 146:249-257.

KING, K. R.

1982. The population biology of the larvacean Oikopleura dioiea in enclosed

water columns. In G. D. Grice and M. R. Reeve (editors), Marine mesocosms:

Biological and chemical research in experimental ecosystems, p. 341-35 I.

Springer-Verlag, N.Y.

KING, K. R., 1. T. HOLLIBAUGH, and F. AZAM.

1980. Predator-prey interactions between the larvacean Oikopleura dioica and

bacterioplankton in enclosed water columns. Mar. BioI. (Berl.) 56:49-57.

KIl!lRBOE, T., F. Ml!lHLENBERG, and H. NICOLAJSEN.

1982. Ingestion rate and gut clearance in the planktonic copepod Centropages

hamatus (Lilljeborg) in relation to food concentration and temperature.

Ophelia 21 :18 I -194.

KJELSON, M. A., D. S. PETERS, G. W. THAYER, and G. N. JOHNSO'll.

1975. The general feeding ecology of posllarval fishes in the Newport River

estuary. Fish. Bull., U.S. 73:137-144.

KLEIN BRETELE~, W. C M.

1980. Continuous breeding of marine pelagic copepods in the presence of

16

heterotrophic dinoflagellates. Mar. Ecol. Prog. Ser. 2:229-233.

KOEHL, M. A. R., and J. R. STRJCKLER.

198 I. Copepod feeding currents: Food capture at low Reynolds number.

Limnol. Oceangr. 26:1062-1073.

KOELLER, P. A., J. E. BARWELL-CLARKE, F. WHITNEY, and M. TAKAHASHI.

1979. Winter condition of marine plankton populations in Saanich Inlet,

B. C, Canada. III. Meso-zooplankton. J. Exp. Mar. BioI. Ecol. 37:161-174.

LALLI, C. M.

1970. Structure and function of the buccal apparatus of Clione limaeina

(Phipps) with a review of feeding in gymnosomatous pteropods. J. Exp.

Mar. BioI. Ecol. 4:101-118.

1972. Food and feeding of Paedodione doliiJormis Danforth, a neotenous

gymnosomatous pteropod. BioI. Bull. (Woods Hole) 143:392-402.

LAMPITT, R. S.

1978. Carnivorous feeding by a small marine copepod. Limnol. Oceanogr.

23:1228-1231.

LAMPITT, R. S., and J. C. GAMBLE.

1982. Diet and respiration of the small planktonic marine copepod Oithona

nana. Mar. BioI. (Berl.) 66:185-190.

LANDRY, M. R.

1978. Population dynamics and production of a planktonic marine copepod,

Acania dausii, in a small temperate lagoon on San Juan Island, Washington.

Int. Rev. Gesamten Hydrobiol. 63:77-119.

1980. Detection of prey by Calanus paeifieus: Implications of the first antennae. Limnol. Oceanogr. 25:545-549.

1981. Switching between herbivory and earn ivory by the planktonic marine

copepod Calanus paeifieus. Mar. BioI. (Berl.) 65:77 -82.

LAROCHE, J. L.

1982. Trophic patterns among larvae of five species of sculpins (Family:

Cottidae) in a Maine estuary. Fish. Bull., U.S. 80:827 -840.

LASKER, R.

1975. Field criteria for survival of anchovy larvae: The relation between

inshore chlorophyll maximum layers and successful first feeding. Fish.

Bull., U.S. 73:453-462.

1978. The relation between oceanographic conditions and larval anchovy

food in the California Current: Identification of factors contributing to

recruitment failure. Rapp. P.-V. Reun. Cons. Int. Explor. Mer 173:

212-230.

1981. Factors contributing to variable recruitment of the northern anchovy

(Engraulis mordax) in the California Current: Contrasting years, 1975

through 1978. Rapp. P.-V. Reun. Cons. Int. Explor. Mer 178:375-388.

LASKER, R., H. M. FEDER, G. H. THEILACKER, and R. C MAY.

1970. Feeding, growth, and survival of Engraulis mordax larvae reared in the

laboratory. Mar. BioI. (Berl.) 5:345-353.

LAST, J. M.

1978a. The food of four species of pleuronectiforrn larvae in the eastern

English Channel and southern North Sea. Mar. BioI. (Berl.) 45:359-368.

1978b. The food of three species of gadoid larvae in the eastern English

Channel and southern North Sea. Mar. BioI. (Berl.) 48:377-386.

1979. The food of larval turbot Seophthalmus maximus L. from the west

central North Sea. J. Cons. Int. Explor. Mer 38:308-313.

LEBOUR, M. V.

1918. The food post-larval fish. J. ~ar. BioI. Assoc. U.K. 11 :433-469.

1919. The food of post-larval fish. No. II (1918). J. Mar. BioI. Assoc. U.K.

12:22-47.

1920. The food of young fish. No. III (1919). J. Mar. BioI. Assoc. U.K.

12:261-324.

1921. The food of young c1upeoids. J. Mar. BioI. Assoc. U.K. 12:458-467.

1922. The food of plankton organisms. J. Mar. BioI. Assoc. U.K. 12:

644-677.

LONSDALE, D. J.

1981. Regulatory role of physical factors and predation for two Chesapeake

Bay copepod species. Mar. Ecol. Prog. Ser. 5:34 I -35 I.

LONSDALE, D. J., D. R. HEINLE, and C. SIEGFRIED.

1979. Carnivorous feeding behavior of the adult calanoid copepod Acania

tonsa Dana. J. Exp. Mar. BioI. Ecol. 36:235-248.

LOWNDES, A. G.

1935. The swimming and feeding of certain Calanoid Copepods. Proc. Zool.

Soc. Lond. 1935:687-715.

MAPSTONE, G. M.

1970. Feeding activities of veligers of Nassarius reetieulatus and Crepidula

Jornieara and the use of artificial foods in maintaining cultures of these larvae. Helgol. wiss. Meeresunters. 20:565-575.

MARAK, R. R.

1960. Food habits of larval cod, haddock, and coalfish in the Gulf of Maine

and Georges Bank area. J. Cons. Int. Explor. Mer 25:147-157.
1974. Food and feeding of larval redfish in the Gulf of Maine. fn J. H. S.

Blaxter (editor), The early life history of fish, p. 267-275. Springer-Verlag,

N.Y.

MARSHALL, S. M.

1924. The food of Ca/anus jinmarchicus during 1923. J. Mar. BioI. Assoc.

U.K. 13:473-479.

1949. On the biology of the small copepods in Loch Striven. J. Mar. BioI.

Assoc. U.K. 28:45-122.

1973. Respiration and feeding in copepods. Adv. Mar. BioI. 11:57-120.

MARSHALL, S. M., and A. P. ORR.

1952. On the biology of Ca/anus jinmarchicus. VII. Factors affecting egg

production. J. Mar. BioI. Assoc. U.K. 30:527-548.

1955a. On the biology of Ca/anus jinmarchicus. VlIl. Food uptake, assimilation and excretion in adult and stage V Calanus. J. Mar. BioI. Assoc. U.K.

34:495-529.

1955b. Experimental feeding of the copepod Ca/anus jinmarchicus (Gunner)

on phytoplankton cultures labelled with radioactive carbon (..C). Deep-Sea

Res. 3 (suppl.):11O-114.

1955c. The biology of a marine copepod, Calanus jinmarchicus (Gunnerus).

Oliver & Boyd, Edinburgh, 188 p.

1956. On the biology of Calanus jinmarchicus. IX. Feeding and digestion in

the young stages. J. Mar. BioI. Assoc. U.K. 35:587-603.

1958. On the biology of Ca/anus jinmarchicus. X. Seasonal changes in

oxygen consumption. J. Mar. BioI. Assoc. U.K. 37:459-472.

196I. On the biology of Ca/anus jinmarchicus. XII. The phosphorus cycle:

Excretion, egg production, autolysis. 1. Mar. BioI. Assoc. U.K. 41:463-483.

1966. Respiration and feeding in some small copepods. J. Mar. BioI.

Assoc. U.K. 46:513-530.

MAYZAUD, P., and S. A. POULET.

1978. The imponance of the time factor in the response of zooplankton to

varying concentrations of naturally occurring panicu!ate matter. Limnol.

Oceanogr. 23:1144-1154.

McALLISTER, C. D.

1970. Zooplankton rations, phytoplankton monality and the estimation of

marine production.fn J. H. Steele (editor), Marine food chains, p. 419-457.

Oliver & Boyd, Edinburgh.

MENDIOLA, B. R. de.

1969. The food of the Peruvian anchovy. J. Cons. Int. Explor. Mer 32:

433-434.

1974. Food of the larval anchoveta Engraulis ringens J. fn J. H. S. Blaxter

(editor), The early life history of fish, p. 277-285. Springer-Verlag, N.Y.

MORTON, J. E.

1954. The biology of Limacina retraversa. J. Mar. BioI. Assoc. U.K. 33:

297-312.

MOYSE, J.

1960. Mass rearing of barnacle cyprids. in the laboratory. Nature (Lond.)

185:120.

1963. A comparison of the value of various flagellates and diatoms as food for

barnacle larvae. J. Cons. Int. Explor. Mer 28:175-187.

MULLIN, M. M.

1963. Some factors affecting the feeding of marine copepods of the genus

Ca/anus. Limnol. Oceanogr. 8:239-250.

MULLIN, M. M., and E. R. BROOKS.

1967. Laboratory culture, growth rate, and feeding behavior of a planktonic

marine copepod. Limnol. Oceanogr. 12:657-666.

1970a. Growth and metabolism of two planktonic, marine copepods as

influenced by temperature and type of food.fn J. H. Steele (editor), Marine

food chains, p. 74-95. Oliver & Boyd, Edinburgh.

1970b. The effect of concentration of food on body weight, cumulative ingestion, and rate of growth of the marine copepod Ca/anus he/go/andicus.

Limnol. Oceanogr. 15:748-755.

MURPHY, H. E.

1923. The life cycle of Oithona nana reared experimentally. Univ. Calif.

Publ. Zool. 22:449-454.

NASSOGNE, A.

1970. Influence of food organisms on the development and culture of pelagic

copepods. Helgol. wiss. Meeresunters. 20:333-345.

NEUNES, H. W., and G.-F. paNGOLINI.

1965. Breeding a pelagic copepod, Euterpina acutifrons (Dana), in the laboratory. Nature (Lond.) 208:57 I-573.

NIVAL, P., and S. NIVAL.

1976. Particle retention efficiencies of an herbivorous copepod,Acartia clausi

(adult and copepodite stages): Effects on grazing. Limnol. Oceanogr.

21 :24-38.

NIVAL, S., and S. RAVERA.

1979. Morphological study of the appendages of the marine cladoceran

17

Evadne spinijera by means of the scanning electron microscope. J. Plankton

Res. 1:207-213.

O'CONNORS, H. 8., Jr., D. C. BIGGS, and D. V. NINIVAGGI.

1980. Particle-size-dependant maximum grazing rates for Temora /ongicornis

fed natural panicle assemblages. Mar. BioI. (Berl.) 56:65-70.

O'CONNORS, H. B., L. F. SMALL, and P. L. DONAGHAY.

1976. Panicle-size modification by two size classes of the estuarine copepod

Acartia clausi. Limnol. Oceanogr. 21 :300-308.

OGILVIE, H. S.

1927. Observations on the food of post-larval herring from the Scottish coast.

Rep. Fish. Board Scotl. 1:1-10.

1938. The food of post-larval haddock with reference to the annual fluctuations in the haddock broods. Rapp. P.-V. Reun. Cons. Int. Explor. Mer 107:

57-66.

OMORI, M.

1973. Cultivation of marine copepods. lin Jpn., Engl. abstr.) Bull. Plankton

Soc. Jpn. 20:3-1 I.

ONBE, T.

1977. The biology of marine cladocerans in a warm temperate water. /n

Proceedings of the symposium on warm water zooplankton, p. 383-398.

Nat. Inst. Oceanogr., Goa.

PAFFENHOFER, G.-A.

1970. Cultivation of Ca/anus he/go/andicus under controlled conditions.

Helgol. wiss. Meeresunters. 20:346-359.

197 I. Grazing and ingestion rates of nauplii, copepodids and adults of the

marine planktonic copepod Ca/anus he/go/andicus. Mar. BioI. (Berl.) II:

286-298.

19,72. The effects of suspended 'red mud' on mortality, body weight, and

growth of the marine planktonic copepod, Ca/anus he/go/andicus. Water

Air Soil Pollut. I:314-32 I.

1973. The cultivation of an appendicularian through numerous generations.

Mar. Bioi. (Berl.) 22:183-185.

1976a. Feeding, growth, and food conversion of the marine planktonic copepod Ca/anus he/go/andicus. Limnol. Oceanogr. 21 :39-50.

1976b. Continuous and nocturnal feeding of the marine planktonic copepod

Ca/anus he/go/andicus. Bull. Mar. Sci. 26:49-58.

1976c. On the biology of appendicularia of the southeastern Nonh Sea. fn G.

Persoone and E. Jaspers (editors), Proc. 10th Europ. Symp. Mar. BioI.,

Ostend, Belgium, Sept. 17-23, 1975, Vol. 2, p. 437-455. Universa Press,

Wettern, Belg.

1982. Grazing by copepods in the Peru upwelling. Deep-Sea Res. 29:

145-146.

PAFFENHOFER, G.-A., and R. P. HARRIS.

1976. Feeding, growth and reproduction of the marine planktonic copepod

Pseudoca/anus elongarus Boeck. J. Mar. BioI. Assoc. U.K. 56:327-344.

PAFFENHOFER, G.-A., and S. C. KNOWLES.

1978. Feeding of marine planktonic copepods on mixed phytnplankton.

Mar. BioI. (Berl.) 48:143-152.

1979. Ecological implications of fecal pellet size, production and consumption by copepods. J. Mar. Res. 37:35-49.

1980. Omnivorousness in marine planktonic copepods. J. Plankton Res.

2:355-365.

PAFFENHOFER, G.-A., and J. D. H. STRICKLAND.

1970. A note on the feeding ofCa/anus helgolandicus on detritus. Mar. BioI.

(Berl.) 5:97 -99.

PAFFENHOFER, G.-A., J. R. STRICKER, and M. ALCARAZ.

1982. Suspension-feeding by herbivorous calanoid copepods: A cinematographic study. Mar. BioI. (Berl.) 67:193-199.

PARANJAPE, M. A.

1968. The egg mass and veligers of Limacina he/ieina Phipps. The Veliger

10:322-326.

PARRISH, K. K., and D. F. WILSON.

1978. Fecundity studies on Acartia tonsa (Copepoda: Calanoida) in standardized culture. Mar. BioI. (Berl.) 46:65-81.

PARSONS, T. R., and M. TAKAHASHI.

1973. Biological oceanographic processes. Pergamon, Oxford, 186 p.

PAVLOVA, E. V.

1959. On the feeding of Penilia avirostris. Tr. Sevastopol BioI. Stn. 11:

63-7 I. (Engl. transl., Fish. Res. Board Can. Transl. 967, 18 p.)

1967. Food utilization and energy conversion by Black Sea Cladocera populations. BioI. Morya 4:66-85. (Engl. transl., Fish. Res. Board Can. Transl.

974, 34 p.)

PAVLOVA, E. V., T. S. PETIPA, and Yu. I. SOROKIN.

1973. Bacterioplankton as food for pelagic marine organisms. fn M. E.

Vinogradov (editor), Life activity of pelagic communities in the ocean

tropics, p. 156-165. Israel Program for Scientific Translations, Jerusalem.
PAVLOVA, E:,y., and Y. I. SOROKIN.

1970. Fe.eding on bacteria by the planktonic crustacean Penilia avirostris

Dana in the Black Sea. lIn Russ.] BioI. Morya 19:182-197.

PEARCY, W. G.

1962. Ecology of an estuarine population of winter flounder, Pseudopleuronectes american us (Walbaum). Bull. Bingham Oceanogr. Collect. Yale

Univ. 18:5-78.

PETIPA, T. S., E. V. PAVLOVA, and G. N. MIRONOV.

1970. The food web structure, utilization and transport of energy by trophic

levels in the planktonic communities. In J. H. Steele (editor), Marine food

chains, p. 142-167. Oliver & Boyd, Edinburgh.

PETIPA, T. S., E. V. PAVLOVA, and Yu. I. SOROKIN.

1973. Radiocarbon studies of the feeding of mass plankton forms in the

tropical zone of the Pacific. In M. E. Vinogradov (editor), Life activity of

pelagic communities in the ocean tropics, p. 135-155. Israel Program for

Scientific Translations, Jerusalem.

PILKINGTON, M. c., and V. FRETTER.

1970. Some factors affecting the growth of prosobranch veligers. Helgol.

wiss. Meeresunters. 20:576-593.

PIONTKOVSKII, S. A., and T. S. PETIPA.

1976. Quantitative description of the behavior of copepod Acania clausi

during feeding on algae. Sov. J. Mar. BioI. (Engl. transl. BioI. Morya)

2:40-46.

POMEROY, L. R.

1974. The ocean's food web, a changing paradigm. BioScience 24:499-504.

POULET, S. A.

1973. Grazing of Pseudocalanus minutus on naturally occurring particulate

matter. Limnol. Oceanogr. 18:564-573.

1974. Seasonal grazing of Pseudocalanus minutus on particles. Mar. Bioi.

(Berl.) 25:109-123.

1976. Feeding of Pseudocalanus minutus on living and non-living particles.

Mar. BioI. (Berl.) 34:117-125.

1978. Comparison between five coexisting species of marine copepods feeding on naturally occurring particulate matter. Limnol. Oceanogr. 23:

1126-1143.

POULET, S. A., and P. MARSOT.

1978. Chemosensory grazing by marine calanoid copepods (Arthropoda:

Crustacea). Science (Wash., D.C.) 200:1403-1405.

1980. Chemosensory feeding and food-gathering by omnivorous copepods.

In W. C. Kerfoot (editor), Evolution and ecology of zooplankton communities, p, 198-218. University Press of New England, Hanover, N.H.

PRICE, H. J., G.-A. PAFFENHOFER, and J. R. STRICKLER.

1983. Modes of cell capture in calanoid copepods. Limnol. Oceanogr. 28:

116-123.

RASSOULZADEGAN, F., and M. ETIENNE.

1981. Grazing rate of the tintinnid Stenosemella ventricosa (Clap. & Lachm.)

Jiirg. on the spectrum of the naturally occurring particulate matte: from a

Mediterranean neritic area. Limnol. Oceanogr. 26:258-270.

RA YMONT, J. E. G.

1959. The respiration of some planktonic copepods. III. The oxygen requirements of some American species. Limnol. Oceanogr. 4:479-491.

REEVE, M. R., and M. A. WALTER.

1977. Observations on the existence of lower threshold and upper critical

food concentrations for the copepod Acania tonsa Dana. J. Exp. Mar. BioI.

Ecol. 29:211-221.

RICHMAN, S., D. R. HEINLE, and R. HUFF.

1977. Grazing by adult estuarine calanoid copepods of the Chesapeake Bay.

Mar. BioI. (Berl.) 42:69-84.

RJCHMAN, S., Y. LOYA, and L. B. SLOBODKIN.

1975. The rate of mucus production by corals and its assimilation by the coral

reef copepod Acania negligens. Limnol. Oceanogr. 20:918-923.

RICHMAN, S., and J. N. ROGERS.

1969. The feeding of Calanus helgolandicus on synchronously growing

populations of the marine diatom Dirylum brighrwellii. Limnol. Oceanogr.

14:701-709.

RIEPER, M.

1978. Bacteria as food for marine harpacticoid copepods. Mar. BioI.

(Berl.) 45:337-345.

1982. Feeding preferences of marine harpacticoid copepods for various

species of bacteria. Mar. Ecol. Prog. Ser. 7:303-307.

RJEPER, M., and C. FLOTOW.

198 I. Feeding experiments with bacteria, ciliates and harpacticoid copepods.

Kiel. Meeresforsch. Sonderh. 5:370-375.

RILEY, G. A.

1946. Factors controlling phytoplankton populations on Georges Bank. J.

Mar. Res. 6:54-73.

18

RJPPINGALE, R. 1., and E. P. HODGKIN.

1975. Predation effects on the distribution of a copepod, Aust. J. Mar.

Freshw. Res. 26:81-91.

ROBERTSON, J. R.

1983. Predation by estuarine zooplankton on tintinnid ciliates. Estuarine

Coastal Shelf Sci. 16:27 -36.

ROBINS, C. R., R. M. BAILEY, C. E. BOND, J. R. BROOKER, E. A. LACHNER,

R. N. LEA, and W. B. SCOTT (editors).

1980. A list of common and scientific names of fishes from the United States

and Canada. 4th ed. Am. Fish. Soc. Spec. PubI. 12, 174 p.

ROMAN, M, R,

1977. Feeding of the copepod Acania tonsa on the diatom Nitzschia closterium

and brown algae (Fucus vesiculosus) detritus. Mar. BioI. (Berl.)

42:149-155.

1978. Ingestion of the blue-green alga Trichodesmium by the harpacticoid

copepod, Macrosetella gracilis. Limnol. Oceanogr. 23:1245-1248.

ROMAN, M. R., and P. A. RUBLEE.

1980. Containment effects in copepod grazing experiments: A plea to end the

black box approach. Limnol. Oceanogr. 25:982-990.

ROSE, M.

1933. Copepodes pelagiques. Faune Fr. 26, 374 p.

ROSENBERG, G. G.

1980. Filmed observations of filter feeding in the marine planktonic copepod

Acania clausii. Limnol. Oceanogr. 25:738-742.

RUNGE, J. A.

1980. Effects of hunger and season on the feeding behavior of Calanus pacificus. Limnol. Oceanogr. 25:134-145.

RYLAND, J. S.

1964. The feeding of plaice and sand-eel larvae in the southern North Sea.

1. Mar. BioI. Assoc. U.K. 44:343-364.

RYTHER, J. H., and J. G. SANDERS.

1980. Experimental evidence of zooplankton control of the species composition and size distribution of marine phytoplankton. Mar. Ecol. Prog. Ser.

3:279-283.

SCHNACK, S. B.

1979. Feeding of Calanus helgolandicus on phytoplankton mixtures. Mar.

Ecol. Prog. Ser. 1:41-47.

1983. On the feeding of copepods on Thalassiosira partheneia from the Northwest African upwelling area. Mar. Ecol. Prog. Ser. 11 :49-53.

SEKI, H.

1973. Red tide of Oitwpleura in Saanich Inlet. La Mer (Bull. Soc. FrancoJpn. Oceanogr.) 11:153-158.

SHELBOURNE, J. E.

1953. The feeding habits of plaice post-Iarve in the Southern Bight. J. Mar.

BioI. Assoc. U.K. 32:149-159.

1957. The feeding and condition of plaice larvae in good and bad plankton

patches. J. Mar. BioI. Assoc. U.K. 36:539-552.

1962. A predator-prey size relationship for plaice larvae feeding on Oitwpleura. J. Mar. BioI. Assoc. U.K. 42:243-252.

SHERMAN, K., and K. A. HONEY.

1971. Seasonal variations in the food of larval herring in coastal waters of

central Maine. Rapp. P.-V. Reun. Cons. Int. Explor. Mer 160:121-124.

SILVER, M. W., and K. W. BRULAND.

1981. Differential feeding and fecal pellet composition of salps and pteropods, and the possible origin of the deep-water flora and olive-green "cells."

Mar. BioI. (Berl.) 62:263-273.

SMITH, S. L., and B. K. HALL.

1980. Transfer of radioactive carbon within the copepod Temora longicornis.

Mar. BioI. (Berl.) 55:277-286.

SOROKIN, Yu. I.

1973. Quantitative evaluation of the roleofbacterioplankton in the biological

productivity of tropical Pacific waters. In M. E. Vinogradov (editor), Life activity of pelagic communities in the ocean tropics, p. 98-134. Israel Program for Scientific Translations, Jerusalem.

1981. Microheterotrophic organisms in marine ecosystems. In A. R. longhurst (editor), Analysis of marine ecosystems, p. 293-342. Acad. Press,

N.Y.

SOROKIN, Yu. I., T. S. PETIPA, and Yeo V. PAVLOVA.

1970. Quantitative estimate of marine bacterioplankton as a source of food.

Oceanology 10:253-260.

SPITTLER, P.

1973. Feeding experiments with tintinnids. Oikos 15 (suppl):128-132.

STOECKER, D., R. R. L. GUILLARD, and R. M. KAVEE.

1981. Selective predation by Favella ehrenbergii (Tintinnia) on and among

dinoflagellates. BioI. Bull. (Woods Hole) 160:136-145.

STRICKLER, J. R.

1982. Calanoid copepods, feeding currents, and the role of gravity. Science
(Wash., D.C.) 218:158-160.

SUMIDA, B. Y., and H. G. MOSER.

1980. Food and feeding of Pacific hake larvae, Merluccius producluS, off

southern California and northern Baja California. Calif. Coop. Oceanic

Fish. Invest. Rep. 21:161-166.

SYSOEVA, T. K., and A. A. DEGTEREVA.

1965. The relation between the feeding of cod larvae and pelagic fry and the

distribution and abundance of their principal food organisms. ICNAF Spec.

Publ. 6, p. 411-416. Dartmouth, N.S., Canada.

TAGUCHI, S., and H. ISHII.

1972. Shipboard experiments on respiration, excretion, and grazing of

Calanus crisla/us and C. plumchrus (Copepoda) in the northern North Pacific.

In A. Y. Takenouti (chief editor), Biological oceanography of the northern

North Pacific Ocean, p. 419-431. Idemitsu Shoten, Tokyo.

TESTER, P. A.

1982. The effects of the temperature acclimation of parental generations and

incubation temperature on lability of egg hatching time in the copepod Acarlia tonsa Dana. Ph.D. Thesis, Oregon State University, Corvallis, 53 p.

THOMPSON, T. E.

1959. Feeding in nudibranch larvae. J. Mar. BioI. Assoc. U.K. 38:239-248.

TIGHE-FORD, D. J., M. J. D. POWER, and D. C. VAILE.

1970. Laboratory rearing of barnacle larvae for antifouling research.

Helgol. wiss. Meeresunters. 20:393-405.

TIMONIN, A. G.

1969. The structure of pelagic associations. The quantitative relationship

between different trophic groups of plankton in frontal zones of the tropical

ocean. Oceanology 9:686-695.

1971. The structure of plankton communities of the Indian Ocean. Mar.

BioI. (Berl.) 9:281-289.

TOMAS, C. R., and E. E. DEASON.

1981. The influence of grazing by two Acania species on Olislhodiscus IUleus.

Pubbl. Stn. Zool. Napoli I: Marine Ecology 2:215-223.

TOMASINI, J. A., and J. MAZZA.

1978. Comportement alimentaire de deux copopodes calanoides (Centropages

typicus et Acania clausi) en milieux nutritifs aune seule algue. J. Cons. Int.

Explor. Mer 38:154-179.

TURNER, J. T.

1978. Scanning electron microscope investigations of feeding habits and

mouthpart structures of three species of copepods of the Family Pontellidae.

Bull. Mar. Sci. 28:487-500.

1981. Latitudinal patterns of calanoid and cyclopoid copepod diversity in

estuarine waters of eastern North America. J. Biogeogr. 8:369-382.

1982. The annual cycle of zooplankton in a Long Island estuary. Estuaries

5:261-274.

TURNER, J. T., and D. M. ANDERSON.

1983. Zooplankton grazing during dinoflagellate blooms in a Cape Cod

embayment, with observations of predation upon tintinnids by copepods.

Pubbl. Stn. Zool. Napoli I: Marine Ecology 4:359-374.

UOTANI, I., K. MATSUZAKI, Y. MAKINO, K. NODA, O. INAMURA, and M.

HORJKAWA.

1981. Food habits of larvae of tunas and their related species in the area

northwest of Australia. [In Jpn., Engl. abstr.) Bull. Jpn. Soc. Sci. Fish. 47:

1165-1172.

UYE, S.-1.

1980a. Development of neritic copepods Acania clausi and A. sleueri. I. Some

environmental factors affecting egg development and the nature of resting

eggs. Bull. Plankton Soc. Jpn. 27:1-9.

1980b. Development of neritic copepods Acania clausi and A. sleueri. II. isochronal larval development at various temperatures. Bull. Plankton Soc.

Jpn.27:11-18.

19

1981. Fecundity studies of neritic calanoid copepods Acania clausi Giesbrecht and A. steueri Smirnov: A simple empirical model of daily egg production. J. Exp. Mar. BioI. Ecol. 50:255-271.

VERITY, P. G., and D. STOECKER.

1982. Effects of Olislhodiscus IUleus on the growth and abundance of tintinnids. Mar. BioI. (Berl.) 72:79-87.

VIDAL, J.

1980a. Physioecology of zooplankton. I. Effects of phytoplankton concentration, temperature, and body size on the growth rate of Calanus pacificus and

Pseudocalanus sp. Mar. BioI. (Berl.) 56: 111-134.

1980b. Physioecology of zooplankton. II. Effects of phytoplankton concentration, temperature, and body size on the development and molting rates of

Calanus pacificus and Pseudocalanus sp. Mar. BioI. (Berl.) 56:135-146.

WALKER, L. M.

1981. Reproductive biology and development of a marine harpacticoid copepod reared in the laboratory. J. Crustacean BioI. 1:376-388.

WEISS, T.

1983. Feeding of calanoid copepods in relation to Phaeocystis pouchelii

blooms in the German Wadden Sea area off Sylt. Mar. BioI. (Berl.) 74:

87-94.

WHITE, A. W.

1977. Dinoflagellate toxins as probable cause of an Atlantic herring (Clupea

harengus harengus) kill, and pteropods as apparent vector. J. Fish. Res.

Board Can. 34:2421-2424.

1980. Recurrence of kills of Atlantic herring (Clupea harengus harengus)

caused by dinoflagellate toxins transferred through herbivorous zooplankton.

Can. J. Fish. Aquat. Sci. 37:2262-2265.

1981. Marine zooplankton can accumulate and retain dinoflagellate toxins

and cause fish kills. Limnol. Oceanogr. 26:103-109.

WHITE, H. H.

1979. Effects of dinoflagellate bioluminescence on the ingestion rates of

herbivorous zooplankton. J. Exp. Mar. Bioi. Ecol. 36:217-224.

WICKSTEAD, J. H.

1962. Food and feeding in pelagic copepods. Proc. Zool. Soc. Lond. 139:

545-555.

WIEBE, P. H., L. P. MADlN, L. R. HAURY, G. R. HARBISON, and L. M. PHILBIN.

1979. Diel vertical migration by Salpa aspera and its potential for large-scale

particulate organic matter transport to the deep-sea. Mar. BioI. (Berl.)

53:249-255.

WISELY, B.

1960. Experiments on rearing the barnacle Eliminius modestus Darwin to

settling stage in the laboratory. Aust. J. Mar. Freshw. Res. 11:42-54.

WYATT, T.

1974. The feeding of plaice and sand-eel larvae in the Southern Bight in

relation to the distribution of their food organisms. In J. H. S. Blaxter (editor),

The early life history of fish, p. 245-251. Springer-Verlag, N.Y.

YONGE, C. M.

1926. Ciliary feeding mechanisms in the thecosomatous pteropods. J. Linn.

Soc. Lond., Zool. 36:417-429.

ZILLIOUX, E. J.

1969. A continuous recirculating culture system for planktonic copepods.

Mar. BioI. (Berl.) 4:215-218.

llLLIOUX, E. J., and D. F. WILSON.

1966. Culture of a planktonic calanoid copepod through multiple generations.

Science (Wash., D.C.) 151 :996-998.

ZURLlNI, G., I. FERRARI, and A. NASSOGNE.

1978. Reproduction and growth of EUlerpina aculifrons (Copepoda:Harpacticoida) under experimental conditions. Mar. BioI. (Berl.) 46:59-64.
Reference Fish species Reference

Appendix Table "-Species of nsh whose larvae have been reported to eat copepod naupUi.

I Fish species

CLUPEIFORMES Sebastes marinus Marak (1974)

Anclwa mitchelli Detwyler and Houde (1970) Scomber scomber Lebour (1918,1920)

Brevoortia patronus Govoni et al. (1983) Thunnus alalunga Uotani et al. (1981)

B. tyrannus Kjelson et al. (1975) T albacares Uotani et al. (1981)

Clupea Iwrengus Bainbridge and Forsyth (1971) T. maccoyU Uotani et al. (1981)

Clupea Iwrengus Bjj!lrke (1971,1978) T obesus Uotani et al. (1981)

Clupea Iwrengus Bowers and Williamson (1951) Trachinus vipera Lebour (1918,1919,1920)

Clupea harengus Gamble et al. (1981) Trachurus symmetr;cus Arthur (1976)

Clupea harengus Lebour (1918,1919,1920,1921) T. trachurus Lebour (1920)

Clupea harengus Ogilvie (1927) Triglops murrayi Laroche (1982)

Clupea harengus Sherman and Honey (1971) PLEURONECTIFORMES

Engraulis anchaita Ciechomski (1967)

E. mordax Arthur (1976) Limanda limanda Last (l978a)

E. ringens Mendiola (1974) Limanda limantia Lebour (1920)

Harengula pensacolae Oetwyler and Honde (1970) Microstomus kilt Lebonr (1920)

Sardina pilchardus Lebonr (1920,1921) Phrynorhambus norvegicus Lebonr (1918, 1920)

Sardinops sagax Arthnr (1976) Platichthys jlessus Last (l978a)

Sprat/us sprattUs Lebonr (1918,1921) Pleuronectes platessa Last (1978a)

PERCIFORMES Pleuronectes plolessa Ryland (1964)

Pleuronectes platessa Shelbonme (1953)

Ammodytes americanus Covill (1959) Pleuronectes platessa Wyatt (1974)

A. mar;nus Ryland (1964) Psetta maxima Lebonr (1920)

A. marinus Wyatt (1974) Pseudopleuronectes

A. tobianus Lebour (1918,1919,1920) americanus Pearcy (1962)

Auxis sp. Uotani et al. (1981) Scophthalmus maximus Last (1979)

Callionymus lyra Lebour (1920) S. rhombrus Lebonr (1920)

Crenolabrus rupestris Lebonr (1920) Solea solea Last (1978a)

Eutrigla gumardus Lebonr (1920) GADIFORMES Gobius niger Lebonr (1920)

Gobiusculus flavescens Lebonr (1918, 1920) Ciliata mustela Lebonr (1918, 1920)

Ka/suwonus pelamis Uotani et al. (1981) Gadus morhua Ellertsen et al. (1981); Last (I 978b);

Labrus bergylta Lebonr (1918,1919,1920) Marak (1960); Sysoeva and

L. bimaculatus Lebonr (1920) Degtereva (1965)

Lagodon rhamboides Kje1son et al. (1975) Meianogrammus aeglejinus Marak (1960)

Leiostomus xanthurus Govoni et al. (1983) Melanogrammus aeglejinus Ogilvie (1938)

Leiostomus xanthurus Kje1son et al. (1975) Merlangius merlangus Last (1978b)

Lepadogaster candollei Lebonr (1918,1920) Merlangius mer/angus Lebonr (1918,1919,1920)

L. lepadogaster Lebonr (1918) Merluccius merluccius Lebonr (1920)

Micropogonias undularus Govoni et al. (1983) M. productus Snmida and Moser (1980)

Myoxocephalus aenaeus Laroche (1982) Micromesistius poutassou Conway (1980)

M. octodecemspinosus Laroche (1982) Molva molva Lebonr (1920)

M. scorpius Laroche (1982) Pollachius virens Marak (1960)

Pomatoschistus microps Lebonr (1920) Trisopterus luscus Last (l978b)

20
Appendix Table 2.--Specles of nsh whose larvae have been reported to eat species of Acartia.

Fish species

CLUPEIFORMES

Anchoa mitchetli

Brevoortia patronus

B. IYrannus

B. tyrannus

Clupea harengus

Clupea harengus

Clupea harengus

Clupea harengus

Harengula pensacolae

Sardina pilchardus

PERCIFORMES

Ammodytes americanus

A. tobianus

Catlionymus lyra

Coryphoblennius galerita

Cyclogaster montagui

Gobius niger

Gobiusculus flavescens

Labrus bergylta

Lagodon rhomboides

Leiostomus :canthurus

Leiostomus xanthurus

Micropogonias undulatus

Pomatoschistus microps

Thunnus albacares

Trachurus trachurus

Reference

Delwyler and Houde (1970)

Govoni el al. (1983)

June and Carlson (1971)

Kjelson el al. (1975)

Bowers and Williamson (1951)

Gamble el al. (1981)

Lebour (1920, 1921)

Shennan and Honey (1971)

Delwyler and Houde (1970)

Lebour (1920,1921)

Covill (1959)

Lebour (1918, 1920)

Lebour (1919, 1920)

Lebour (1919)

Lebour (1919)

Lebour (1918,1919,1920)

Lebour (1918,1919,1920)

Lebour (1919, 1920)

Kjelson el al. (1975)

Govoni el al. (1983)

Kjelson el al. (1 975)

Govoni el al. (1983)

Lebour (1918,1919,1920)

Uotani el al. (1981)

Lebour (1920)

Fish species

PLEURONECTIFORMES

Limanda limanda

Microchirus variegatus

Phrynorhombus norvegicus

Platichthys flessus

Scophthalmus maximus

Solea nasuta

Zeugopterus punctatus

GADIFORMES

Ciliata mus/ela

Gadus morhua

Melanogrammus aeglefinus

Merlangius mer/angus

Merlangius merlangus

Micromesistius poutassou

Pollachius pollachius

Trisopterus luscus

Trisopterus luscus

T. minutus

Reference

Lebour (1920)

Lebour (1918)

Lebour (1918,1920)

Lebour (1920)

LaSI (1979)

Lebour (1920)

Lebour (1920)

Lebour (1920)

Sysoeva and Degtereva (1965)

Ogilvie (1938)

Lasl (1978b)

Lebour (1919, 1920)

Conway (1980)

Lebour (1918,1919,1920)

LaSI (1978b)

Lebour (1918,1920)

Lebour (1918,1920)

Appendix Table 3.--Specles of nsh whose larvae have been reported to eat species of Calanu•.

Fish species Reference Fish species Reference

CLUPEIFORMES PLEURONECTIFORMES

Clupea harengus Blaxter (1965) Amoglossus sp. Lebour (1920)

Clupea harengus Bowers and Williamson (1951) Limanda Iimanda Lebour (1920)

Clupea harengus Gamble el al. (1981) Phrynorhombus norvegicus Lebour (1918,1920)

Clupea harengus Ogilvie (1927) Zeugopterus punctatus Lebour (1918)

Engraulis ringens Mendiola (1974) GADIFORMES

PERCIFORMES Gadus morhua Lebour (1918)

Auxis sp. Uotani el al. (1981) Gadus morhua Sysoeva and Deglereva (1965)

Catlionymus lyra Lebour (1918) Melanogrammus aeglefinus Ogilvie (1938)

Crystallogobius nilssoni Lebour (1918) Melanogrammus aeglefinus Marak (1960)

Eutrigla gumardus Lebour (1920) Merlangius merlangus Lebour (1919, 1920)

Sebastes marinus Bainbridge (1965) Mer/ucdus merlucdus Lebour (1920)

Sebastes marinus Marak (1974) M. productus Sumida and Moser (1980)

Thunnus abesus Uolani el al. (198 I ) Micromesistius poutassou Conway (1980)

Trachurus trachurus Lebour (1918,1920) Pollachius pollachius Lebour (1919,1920)

P. virens Marak (1960)

Trisop/erus luscus Lebour (1919,1920)

T. minu/us Lebour (1920)

21
Appendix Table 4.-Specles of fish whose larvae have been reported to eat species of Centropages.

Fish species Reference Fish species

CLUPEIFORMES PLEURONECTIFORMES

Brevoortia ryrannus June and Carlson (1971) Arnoglossus sp.

Brevoonia ryrannus Kjelson et al. (1975) Psetta maxima

Clupea harengus Lebour (1921) Scophthalmus maximus

Clupea harengus Ogilvie (1927) S. rhambrus

Engraulis anchoila Ciechomski (1967) GADIFORMES

PERCIFORMES

Melanogrammus

Ammodyles lobianus Lebour (1920) aeg/efinus

Callionymus lyra Lebour (1918) Melanogrammus

lAgodon rhomboides Kjelson et al. (1975) aeg/efinus

Leiostomus xanthurus Kje1son et al. (1975) Merlangius mer/angus

Lepadogasler Micromesistius

/epadogasler Lebour (1918) poutassou

Sebastes marinus Marak (1974) Pollachius virens

Trachurus rrachurus Lebour (1918,1920)

Reference

Lebour (1920)

Lebour (1918)

Last (1979)

Lebour (1918)

Ogilvie (1938)

Marak (1960)

Lebour (1920)

Conway (1980)

Marak (1960)

Appendix Table 5.-Specles of ftsb whose lanae han been reported to eat species of ParaetJla"u,.

Fish species

CLUPEIFORMES

Brevoortia patronus

C/upea harengus

Clupea harengus

Clupea harengus

Engraulis anchoita

PERCIFORMES

Ammodytes americanus

A. ,obianus

Aruis sp.

Ca//ionymus lyra

Eutrig/a gurnardus

Gobiuscu/us flavescens

lAbrus bergy/ta

L. bimacu/atus

Leiostomus xanthurus

Micropogonias undu/atus

Thunnus a/alunga

T. albacares

T. maccoyii

T.obesus

Trachurus trachurus

Reference

Govoni et al. (1983)

Bowers and Williamson (1951)

Lebour (1920, 1921)

Ogilvie (1927)

Ciechomski (1967)

Covill (1959)

Lebour (1920)

Volani el 01. (1981)

Lebour (1918,1920)

Lebour (1920)

Lebour (1920)

Lebour (1920)

Lebour (1920)

Govoni et 01. (1983)

Govoni el 01. (1983)

Votani el 01. (1981)

Votani el 01. (1981)

Votani el 01. (1981)

UOlani el 01. (1981)

Lebour (1920)

22

Fish species

PLEURONECTIFORMES

Arnog/ossus sp.

Phrynorhombus norvegicus

P. regias

GADIFORMES

Gadus morhua

Me/anogrammus aeg/efinus

Me/anogrammus aeg/efinus

Mer/angius mer/angus

Merlangius mer/angus

Merluccius productus

Mo/va mo/va

Pollachius virens

Trisopterus luscus

Reference

Lebour (1918)

Lebour (1918, 1920)

Lebour (1918)

Last (1978b)

Marak (1960)

Ogilvie (1938)

LaSI (1978b)

Lebour (1918,1920)

Sumida and Moser (1980)

Lebour (1920)

Marak (1960)

Last (1978b)
Appendix Table 6.-Specles of nsh whose larvae have been reported to eat species of Pseudoca/anus.

Fish species Reference Fish species Reference

CLUPEIFORMES PLEURONECTIFORMES

Clupea harengus Bowers and Williamson (1951) Amog/ossus sp. Lebour (1918.1919.1920)

C/upea harengus Gamble et al. (1981) Limanda limanda Last (1978a)

C/upea harengus Lebour (1918. 1919. 1920. 1921) Limanda limanda Lebour (1918. 1920)

C/upea harengus Ogilvie (1927) Microchirus variegatus Lebour (1918)

C/upea harengus Sherman and Honey (1971) Phrynorhambus norvegicus Lebour (1918. (920)

Sardina pu/chardus Lebour (1920.1921) P. regias Lebour (1918)

Sprattus sprattus Lebour (1918.1921) P/euronectes p/atessa Ryland (1964)

PERC1FORMES Psetta maxima Lebour (1920)

Scophtha/mus maximus Last (1979)

Agonus cataphractus Lebour (1920) So/ea nasuta Lebour (1920)

Ammodytes americanus Covill (1959) GADIFORMES A. marinus Ryland (1964)

A. tobianus Lebour (1918.1919.1920) Gadus morhua Last (l978b)

Callionymus lyra Lebour (1918.1919.1920) Gadus morhua Marak (1960)

Eutrig/a gumardus Lebour (1920) Gadus morhua Sysoeva and Degtereva (1965)

Gobius niger Lebour (1919.1920) Me/anogrammus aeg/efinus Marak (1960)

Gobiuscu/us flavescens Lebour (1918.1919.1920) Me/anogrammus aeg/efinus Ogilvie (1938)

Labrus bergy/ta Lebour (1918) Mer/angius mer/angus Last (I 978b)

L. bimacu/atus Lebour (1920) Mer/angius mer/angus Lebour (1918.1919.1920)

Lepadogaster cando/lei Lebour (1919. 1920) Merluccius merluccius Lebour (1920)

L. /epadogaster Lebour (1918) Micromesistius poutassou Conway (1980)

Myoxocepha/us scorpius Laroche (1982) Mo/va mo/va Lebour (1920)

Pomatoschistus microps Lebour (1919. 1920) Pol/achius pol/achius Lebour (1919)

Sebastes marinus Marak (1974) P. virens Marak (1960)

Tauru/us bubalis Lebour (1919. 1920) Trisopterus luscus Last (l978b)

Trachinus vipera Lebour (1918.1919.1920) Trisopterus luscus Lebour (1918.1919.1920)

Trachurus trachurus Lebour (1920) T. minutus Lebour (1918.1920)

Trig/ops murrayi Laroche (1982)

AppendIx Table 7.-Specles of nsh whose larvae have been reported to eat species of Temora.

Fish species Reference Fish species Reference

CLUPEIFORMES Tauru/us bubalis Lebour (1918.1919.1920)

Brevoortia patronus Govoni et al. (1983) Trachinus vipera Lebour(l919. 1920)

B. tyrannus June and Carlson (1971) Trachurus trachurus Lebour (1919.1920)

B. tyrannus Kje1son et al. (1975) Trig/ops murrayi Laroche (1982)

C/upea harengus Gamble et al. (1981) PLEURONECTIFORMES

C/upea harengus Ogilvie (1927)

Sardina pilchardus Lebour (1921) Limanda limanda Last (l978a)

Limanda limanda Lebour (1918.1920)

PERC1FORMES Microchirus var;egatus Lebour (1918)

Agonus cataphractus Lebour (1920) Phrynorhambus norvegicus Lebour (1918. 1920)

Ammodytes americanus Covill (1959) P. regias Lebour (1918)

A. tobianus Lebour (1920) P/atichthys flessus Lebour (1920)

Gallionymus lyra Lebour (1918.1919.1920) Pleuronectes platessa Last (l978a)

Coryphablennius ga/erita Lebous (1918) Pleuronectes platessa Lebour (1918.1920)

Crystal/ogobius nilssoni Lebous (1920) Psetta maxima Lebour (1918)

Cleno/abrus rupestris Lebous (1919. 1920) Scophtha/mus maximus Last (1979)

Cyc/ogaster montagui Lebous (1919) S. rhambrus Lebour (1918.1920)

Dip/ecogaster bimacu/atus Lebour (1920) So/ea nasuta Lebour (1918. 1920)

Eutrig/a gumardus Lebour (1920) S. so/ea Lebour (1918)

Gobius niger Lebous (1919) Zeugopterus punctatus Lebour (1918. 1920)

Gobiusculus flavescens Lebour (1918.1919) GADIFORMES Lobrus bergy/ta Lebour (1918.1919.1920)

L. bimacu/atus Lebour (1920) Ciliata muste/a Lebour (1919)

Logadon rhomboid.. Kjelson et al. (1975) Gadus morhua Lebour (1918)

Leioslomus xanthurus Govoni et al. (1983) Me/anogrammus aeg/efinus Marak (1960)

Le;ostomus xanthurus Kjelson et al. (1975) Me/anogrammus aeg/efinus Ogilvie (1938)

Lepadogaster cando/lei Lebous (1918.1919.1920) Mer/angius mer/angus Last (1978b)

Micropogonias undu/atus Govoni et al. (1983) Mer/angius mer/angus Lebour (1919.1920)

Myoxocepha/us aenaeus Laroche (1982) Pol/achius pol/achius Lebour (1918.1919.1920)

M. octadecemspinosus Laroche (1982) P. v;rens Marak (1960)

M. scorpius Laroche (1982) Trisopterus luscus Lebour (1920)

Pomatoschistus microps Lebous (1920) T. minutus Lebour (1920)

Scomber scomber Lebour (1920)

23
Appendix Table 8.-Specles of nsh whose larvae have been reported to eat species of COryClUUS.

Fish species Reference Fish species Reference

CLUPEIFORMES Trachurus symmetricus Anhur (1976)

Brevoortia patronus Govoni el al. (1983) T. trachurus Lebour (1920)

Clupea harengus Lebour (1918,1920,1921)

Thunnus alalunga Uolani el al. (1981)

T. albacares UOlani el al. (1981) Engraulis anchoita Ciechomski (1967) T. maccoyii Uolani el al. (1981)

PERCIFORMES T. obesus UOlani el al. (1981)

Callionymus lyra Lebour (1918) PLEURONECTIFORMES

Katsuwonus pelamis UOlani el al. (1981) Limanda limanda Lebour (1918) Leiostomus xanthurus Govoni el al. (1983)

Micropogonias undulatus Govoni et al. (1983) Microchirus variegata Lebour (1918)

Trachinus lIipera Lebour (1919) Scophthalmus norvegicus Lebour (1918)

Appendix Table 9.-Specles of nsh whose larvae have been reported to eat species of Oilhona.

Fish species Reference Fish species Reference

CLUPEIFORMES Sebastes marinus Bainbridge (1965)

Sebasles marinus Marak (1974) Brevoortia patronus Govoni el al. (1983) Thunnus alalunga Uolani el al. (1981) B. tyrannus Kjelson el al. (1975) T. moccayii Uolani el al. (1981) Clupea harengus Bowers and Williamson (1951) Trachinus vipera Lebour (1919) Clupea harengus Gamble el al. (1981) Trachurus symmetricus Anhur (1976) Clupea harengus Lebour (1918,1920) T. trachurus Lebour (1920) Clupea harengus Ogilvie (1927)

Clupea harengus Sherman and Honey (1971) PLEURONECTIFORMES

Engraulis anchoila Ciechomski (1967) Phrynorhambus norvegicus Lebour (1920) Sardina pilchardus Lebour (1920, 1921)

GADIFORMES

PERCIFORMES

Ammodytes americanus Covill (1959) Ciliata mustella Lebour (1919)

Gadus morhua Sysoeva and Deglereva (1965) A. tobianus Lebour (1918,1919,1920) Melanogrammus aegle/inus Ogilvie (1938) Labrus bergylta Lebour (1920) Merlangius merlangus Lebour (1920) L. bimaculatus Lebour (1920) Merluccius productus Sumida and Moser (1980) Lagodon rhamboides Kjelson el al. (1975) Micromesistius poutassou Conway (1980) Leio~·tomus xanthurus Govoni el al. (1983) Pollachius pol/achius Lebour (1919,1920) Leioslomus xanthurus Kjelson el al. (1975)

Micropogonias undulatus Govoni el al. (1983)

Pomatoschistus microps Lebour (1920)

Appendix Table IO.-Specles of nsh whose larvae have been reported to eat species of Ollcaea.

Fish species Reference Fish species Reference

CLUPEIFORMES PLEURONECTIFORMES

Brevoortia patron us Govoni el al. (1983) Microchirus variegatus Lebour (1918)

Clupea harengus Lebour (1918) Phrynorhambus norvegicus Lebour (1918)

PERCIFORMES

Solea vulgaris Lebour (1918)

Zeugopterus punctatus Lebour (1918)

Leiostomus xanthurus Govoni el al. (1983) GADIFORMES

Micropogonias undulatus Govoni el al. (1983)

Trachurus symmelricus Anhur (1976) Merlangius merlangus Lebour (1918)

Micromesislius POUlassou Conway (1980)

24
Appendix Table H.-Species of nsh whose larvae have been reported to eat species of EuUrpina.

Fish species Reference Fish species Reference

CLUPEIFORMES Limanda Iimanda Lebour (1918)

Brevoortia potronus Govoni el al. (1983) Microchirus variegatus Lebour (1918)

Microstomus kilt Lebour (1918) C/upea harengus Lebour (1918) Phrynorhombus norvegicus Lebour (1918) Engraulis anchoita Ciechomski (1967) P. regias Lebour (1918)

PERCIFORMES So/ea nasuta Lebour (1918)

S. so/ea Lebour (1918) Callionymus lyra Lebour (1918) Zeugopterus punctatus Lebour (1918) Labrus bergy/ta Lebour (1918)

PLEURONECTIFORMES GADlFORMES

Lebour (1918) Ciliata mustella Lebour (1919) Amog/ossus sp. Pollachius pollachius Lebour (1918)

Trisopterus minutus Lebour (1918)

Appendix Table 12.-Specles of nsh whose larvae have been reported to eat species of Microsetella.

Fish species Reference Fish species Reference

CLUPEIFORMES PLEURONECTIFORMES

Brevoortia potTonus Govoni et al. (1983) Microchirus variegatus Lebour (1918)

C/upea harengus Bjorke (1971, 1978) So/ea so/ea LaSI (1978a)

C/upea harengus Ogilvie (1927) GADlFORMES

PERCIFORMES Micromesistius poutassou Conway (1980)

Leiostomus xanthurus Govoni et al. (1983)

Micropogonias undu/atus Govoni el al. (1983)

Myoxocepha/us aenaeus Laroche (1982)

M. octodecemspinosus Laroche (1982)

M. scorpius Laroche (1982)

Trachurus symmetricus Arthur (1976)

Appendix Table 13.-Spedes of nsh whose larvae have been reported to eat other (mostly unldentlned) harpactlcold copepods.

Fish species Reference Fish species Reference

CLUPEIFORMES Lepodogaster condollei Lebour (1918,1919,1920)

Brevoortia patron us Govoni el al. (1983) L. /epadogaster Lebour (1918)

Micropogonias undu/atus Govoni el al. (1983) B. cyrannus Kjelson el al. 1975) Myoxocepha/us undu/atus Laroche (1982) C/upea harengus Bowers and Williamson (1951) M. octodecemspinosus Laroche (1982) C/upea harengus Gamble el al. (1981) M. scorpius Laroche (1982) C/upea harengus Lebour (1918,1920,1921) Trachurus symmetricus Arthur (1976) C/upea harengus Ogilvie (1927) Trig/ops murrayi Laroche (1982) C/upea harengus Sherman and Honey (1971)

Sardina pilchardus Lebour (1921) PLEURONECTIFORMES

Sprattus sprattus Lebour (1921) Limanda Iimanda Lebour (1920)

PERCIFORMES Microchirus variegatus Lebour (1918)

Agonus cataphractus Lebour (1920) Pseudopleuronectes americanus Pearcy (1962)

Callionymus lyra Lebour (1919) So/ea so/ea Lebour (1918)

Cteno/abrus rupestris Lebour (1920) GADlFORMES

Dip/ecogaster bimacu/atus Lebour (1919) Me/anogrammus aeg/efinus Ogilvie (1938) Gobius niger Lebour (1919)

Gobiuscu/us flavescens Lebour (1919, 1920) Mer/angiu. mer/angus Lebour (1920)

Labrus bergylta Lebour (1919, 1920)

Pollachius pollachius Lebour (1919,1920)

L. bimacu/atus Lebour (1920) Trisopterus luscus Lebour (1920)

Lagodon rhomboides Kjelson et al. (1975)

Leiostomus xanthurus Govoni el al. (1983)

Leiostomus xanthurus Kje1son et al. (1975)

25
Appendix Table 14.-8pecles of Osh whose larvae have been reported to eat tintinnids.

Fish species Reference Fish species Reference

CLUPEIFORMES PLEURONECTIFORMES

Anchoa milchelli Delwyler and Houde (1970) Limanda /imanda Lasl (l978a)

Brevoortia patronus Govoni el al. (1983) Limanda limanda Lebour (1918)

Clupea harengus Bainbridge and Forsyth (1971) Microchirus variegatus Lebour (1918)

C/upea harengus Lebour (1920,1921) Phrynorhombus norvegicus Lebour (1918)

Clupea harengus Sherman and Honey (1971) P/arichlhys flessus LaSI (1978a)

Engraulis mordax Arthur (1976) Pseudopieuronecles

E. ,ingens Mendiola (1974) americanus Pearcy (1962)

Sardinops sagax Arthur (1976) GADIFORMES

Sprallus sprallus Lebour (1921)

PERCIFORMES

Gadus morhua Last (1978b)

Me/anogrammus aeg/efinus Ogilvie (1938)

Leioslomus xanthurus Govoni et al. (1983) Merlangius mer/angus LaSI (1978b)

Micropogonias undulatus Govoni el al. (1983) Micromesistius poutassou Conway (1980)

Trochurus symme/ricus Arthur (1976) Trisopterus luscus LaSI (l978b)

Appendix Table IS.-Specles of Osh whose larvae have been reported to eat species of Evadne.

Fish species

CLUPEIFORMES

C/upea harengus

C/upea harengus

Engraulis anchoita

PERCIFORMES

Auxis sp.

Katsuwonus pelamis

Leiostomus xanthurus

Lepadogasler condollei

Scomber scomber

Thunnus a/a/unga

T. a/bacares

T maccayii

T. obesus

Trachinus vipera

Reference

Lebour (1918, 1921)

White (1980)

Ciechomski (1967)

Uolani el al. (1981)

UOlanielal (1981)

Govoni el al. (1983)

Lebour (1920)

Lebour (1920)

UOlani el al. (1981)

UOlani el al. (1981)

UOlani el al. (1981)

Uotani el al. (1981)

Lebour (1920)

Fish species

PLEURONECTIFORMES

Scophlha/mus maximus

So/ea so/ea

GADIFORMES

Gadus morhua

Me/anogrammus aeg/efinus

Me/anogrammus aeg/efinus

Micromesistius POUlassou

Reference

Lasl (1979)

Lebour (1918)

Sysoeva and Degtereva

(J 965)

Marak (1960)

Ogilvie (1938)

Conway (1980)

Appendix Table 16.-8pecles of Osh whose larvae have been reported to eat species of Podoll.

Fish species Reference Fish species Reference

CLUPEIFORMES PLEURONECTIFORMES

C/upea harengus Ogilvie (1927) Limanda limanda Lebour (1918,1920)

Engrau/is'anchoila Ciechomski (1967) Microchirus variegalus Lebour (1918)

PERCIFORMES Phrynorhombus norvegicus Lebour (1918,1920)

P. regias Lebour (1918)

Ammadyles lobianus Lebour (1919) Psetta maxima Lebour (1920)

B/ennius gallorugine Lebour (1918, J919) Scophlha/mus maximus Last (1979)

Callionymus lyra Lebour (1918, 1920) S. rhombrus Lebour (1918)

Coryphob/ennius ga/erita Lebour (1919) GASIFORMES Cryslallogobius nilssoni Lebour (1918)

Creno/abrus rupeslris Lebour (1920) Me/anogrammus aeg/efinus Ogilvie (1938)

Dip/ec'!gasler bimacu/alus Lebour (1920) Mer/angius mer/angus Lebour (1920)

EUlrig/a gumardus Lebour (1918, 1920) Micromesistius poutassou Conway (1980)

Gobiuscu/us flavescens Lebour (1919) Mo/va mo/va Lebour (1920)

Labrus bergy/la Lebour (1918,1919,1920) Pollachius pollachius Lebour (1920)

L. bimacu/alus Lebour (1920) Trisopterus luscus Lebour (1920)

Lepadogasler candollei Lebour (1919,1920) T. minutus Lebour (1918,1920)

Scomber scomber Lebour (1920)

Trachinus vipera Lebour (1920)

Trachurus trachurus Lebour (1920)

26
Appendix Table 17.-Specles of nsh whose larvae have been reported to eat barnacle naupUi.

Fish species Reference Fish species Reference

CLUPEIFORMES Leiostomus xanthurus Kjelson et al. (1975)

Brevoortia patronus Govoni et al. (1983) Micropogonias undulatus Govoni et al. (1983)

Myoxocephalus aenaeus Laroche (1982) B. tyrannus Kjelson et al. (1975) M. octodecemspinosus Laroche (1982) Clupea harengus Bjj!lrke (1978) M. scorpius Laroche (1982) Clupea harengus Lebour (1918,1920,1921) PomatoschislUS microps Lebour (1920) Clupea harengus Ogilvie (1927) Taurulus bubalis Lebour (1918,1919,1920) Clupea harengus Sherman and Honey (1971) Triglops murrayi Laroche (1982) Harengula pensacolae Delwyler and Houde (1970)

PERCIFORMES PLEURONECTIFORMES

Agonus cataphractus Lebour (1919, 1920) Psetla maxima Lebour (1918)

Callionymus lyra Lebour (1918) Solea solea Lebour (1918)

Diplecogoster bimaculatus Lebour (1919,1920) GADlFORMES

Gobiusculus flavescens Lebour (1919)

Labrus bergylta Lebour (1919, 1920) Merlangius merlangus Lebour (1919, 1920)

Lagodon rhomboides Kjelson et al. (1975)

Appendix Table 18.-Specles of nsh whose larvae have been reported to eat gastropod larvae.

Fish species Reference Fish species Reference

CLUPEIFORMES PLEURONECTIFORMES

Anchaa mitchelli Detwyler and Houde (1970) Limanda limanda Lebour (1920)

Clupea harengus Bowers and Williamson (1951) Pleuronectes platessa Shelbourne (1953)

Clupea harengus Gamble et al. (1981) Scophthalmus maximus Last (1979)

Clupea harengus Lebour (1918,1920,1921) Solea solea Last (1978a)

Clupea harengus Ogilvie (1927) GADlFORMES

Engraulis mordax Lasker el al. (1970)

Sprattus sprattus Lebour (1919,1921) Ciliata mustela Lebour (1920)

PERCIFORMES Melanogrammus aegle/inus Marak (1960)

Melanogrammus aegle/inus Ogilvie (1938)

Ammodytes tobianus Lebour (1920) Micromesistius poutassou Conway (1980)

Diplecogoster bimaculatus Lebour (1919) Pollachius pollachius Lebour (1920)

Gobiusculus jlavescens Lebour (1919, 1920)

Labrus bergylta Lebour (1920)

Lepadogoster lepadogoster Lebour (1919)

PomatoschislUS microps Lebour (1919)

Scomber scomber Lebour (1918)

Taurulus bubalis Lebour (1918,1919)

Trachurus trachurus Lebour (1920)

Appendix Table 19.-Specles of nsh whose larvae have been reported to eat species of Limacina.

Fish species

CLUPEIFORMES

Brevoortia pO/Tonus

Clupea harengus

Engraulis mordax

Sardinops sagax

PERCIFORMES

Leiostomus xanthurus

Micropogonios undulatus

Reference

Govoni et al. (1983)

While (1977)

Arthur (1976)

Arthur (1976)

Govoni el al. (1983)

Govoni el al. (1983)

27

Fish species

GADlFORMES

Melanogrammus aeglejinus

Micromesistius poutassou

Reference

Ogilvie (1938)

Conway (1980)
Appendix Table 20.-Species of nsh whose larvae have been reported to eat appendicularlans.

Fish species Reference I Fish species Reference

PERCIFORMES

Ammodytes marin us

Ammodyles marinus

Auxis sp.

Katsuwonus pelamis

Thunnus alalunga

T. albacares

T. maccayii

T. obesus

Ryland (1964)

Wyatt (1974)

Uotani el al. (1981)

Uotani et al. (1981)

Uotani et al. (1981)

Uotani et al. (1981)

Uotani et al. (1981)

Uotani et al. (1981)

PLEURONECTIFORMES

Limanda limanda

Platichthys flessus

Pleuronectes platessa

Pleuronectes platessa

Pleuronectes platessa

Pleuronectes platessa

Scophthalmus maximus

GADIFORMES

Micromesistius poutassou

28

Last (1978a)

Last (1978a)

Last (1978a)

Ryland (1964)

Shelbourne (1953,

1957,1962)

Wyatt (1974)

Last (1979)

Conway (1980)