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Dissertations, Theses, and Masters Projects Theses, Dissertations, & Master Projects
1975
Food Habits of Some Demersal Fishes of the Continental Slope and Rise
George R. Sedberry College of William and Mary - Virginia Institute of Marine Science
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Recommended Citation Sedberry, George R., "Food Habits of Some Demersal Fishes of the Continental Slope and Rise" (1975). Dissertations, Theses, and Masters Projects. Paper 1539617460. https://dx.doi.org/doi:10.25773/v5-mj9w-zx56
This Thesis is brought to you for free and open access by the Theses, Dissertations, & Master Projects at W&M ScholarWorks. It has been accepted for inclusion in Dissertations, Theses, and Masters Projects by an authorized administrator of W&M ScholarWorks. For more information, please contact [email protected]. FOOD HABITS OF SOME DEMERSAL FISHES OF THE
CONTINENTAL SLOPE AND RISE
A Thesis
Presented to
The Faculty of the School of Marine Science
The College of William and Mary in Virginia
In Partial Fulfillment
of the Requirements for the Degree of
Master of Arts
/ >-*t tf?« ^ * .•'♦vrunjA !n*>TfT±ej:'i \ v of i '■ , .*AP;r:.r J by '---- George R. Sedberry, III
1975 APPROVAL SHEET
This thesis is submitted in partial fulfillment
of the requirements for the degree of
Master of Arts
George R. S&dberry,
Approved, August 1975
G- „Jonn A. fMsick
Paul A. HaefnerC Jr / 9 ,
Gfeorge Cx Grant
Donald F. Boesch
. Merriner
ii TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS...... iv
LIST OF T A B L E S ...... vi
LIST OF FIGURES...... vli
ABSTRACT...... viii
INTRODUCTION ...... 2 METHODS AND MATERIALS...... 5
RESULTS AND DISCUSSION ...... 10
CONCLUSIONS...... 52
APPENDIX...... 58
LITERATURE CITED ...... 61
ili ACKNOWLEDGMENTS
I thank Dr. John A. Musick, chairman of my thesis committee, for suggesting this problem, and for his suggestions and criticisms. I also thank members of my committee, Drs. P. A. Haefner, Jr., G. C. Grant, D. F.
Boesch, and J. V. Merriner, for their criticisms and review of the manuscript.
For their help in identifying food items, I thank Dr.
Clyde F. E. Roper of the U. S. National Museum of Natural
History, who identified some of the cephalopod beaks; Dr. E.
L. Bousefield of the National Museum of Canada, who identified many of the amphipods and gave me information on their distribution; and John E. Olney and Dr. G. C. Grant of the Virginia Institute of Marine Science (VIMS), who identified many of the copepods. Dr. P. A. Haefner, Jr. introduced me to the literature on crustaceans and gave me access to his library, and Elizabeth G. Lewis of VIMS identified some of the decapods.
Ship time and financial support were provided by
National Science Foundation (NSF) Grants GD-32560, CG-00005
(Cruises E-l-73 and E-2-74 respectively), NSF grant GA-37561
(Cruises CI-73-10 and GI-74-04 respectively), and Sea Grant
04-4-1-158-31 (Cruise D-2-74).
For sharing their knowledge and suggestions with me, I thank my fellow graduate students, especially Douglas
Markle and Labbish Chao. Special thanks are extended to
Charles Wenner, who spent many hours cutting out stomachs and recording data for me, and who also gave me many helpful suggestions.
A special thanks is extended to my wife, Joan, for typing an earlier draft of the manuscript, for her material and moral support, and for her patience and encouragement throughout the duration of this study.
v LIST OF TABLES
Table Page
1. List of fishes examined...... 5
2. Percent frequency occurrence, percent numerical dominance, percent volume displacement, and index of relative importance of food items in the stomachs of Synaphobranchus kaupi...... 13
3. Percent frequency occurrence, percent numerical dominance, percent volume displacement, and index of relative importance of food items in the stomachs of Halosauropsis macrochir...... 22
4. Percent frequency occurrence, percent numerical dominance, percent volume displacement, and index of relative importance of food items in the stomachs of Aldrovandia spp...... 29 5. Percent frequency occurrence, percent numerical dominance, percent volume displacement, and index of relative importance of food items in the stomachs of Phycis chesteri . 35
6 . Percent frequency occurrence, percent numerical dominance, percent volume displacement, and index of relative importance of food items in the stomachs of Lycodes atlanticus...... 43
7. Percent frequency occurrence, percent numerical dominance, percent volume displacement, and index of relative importance of food items in the stomachs of Coryphaenoides armatus...... 48
vi LIST OF FIGURES
age
Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance for higher taxonomic groups of food in the diet of Synaphobranchus kaupi...... 15 Percent frequency occurrence of pelagic fishes in stomachs of Synaphobranchus kaupi in relation to time of day of collection . 19 Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance for higher taxonomic groups of food in the diet of Halosauropsis macrochir...... 24 Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance for higher taxonomic groups of food in the diet of Aldrovandia spp...... 28 Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance for higher taxonomic groups of food in the diet of Phycis chesteri...... 38 Percent frequency occurrence of pelagic fishes and euphausiids in the stomachs of Phycis chesteri in relation to time of day of collection...... 41 Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance of higher taxonomic groups of food in the diet of Lycodes atlanticus...... 45 Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance of higher taxonomic groups of food in the diet of Coryphaenoides armatus ...... 50
vii ABSTRACT
Stomach contents of 742 fishes comprising 14 species were examined from the continental slope and rise of the
Middle Atlantic Bight. Analysis of the stomachs show two main types of feeders among benthic deep-sea fishes: those that feed primarily on pelagic food items, and those that feed by rooting in the sediments, ingesting infauna1 and epifaunal invertebrates. The diets of both pelagic and benthic feeders are found to be unspecialized. This suggests the benthic feeders may play a role in maintaining the high species diversity found in benthic invertebrates, as Dayton and Hessler (1972) have hypothesized.
v• m • • FOOD HABITS OF SOME DEMERSAL FISHES OF THE
CONTINENTAL SLOPE AND RISE INTRODUCTION
Little is known of the food of demersal fishes of the
continental slope and rise and of the abyss (Marshall, 1965
Bright, 1970; Grassle and Sanders, 1973). Thus, on other
than a theoretical basis, relatively few studies have dealt
with trophic structure in the deep sea. Marshall (1954) reported the food of some deep living invertebrates and
fishes and reviewed the two main theories of food provision
to the deep sea: the rain of dead plankton (Agassiz, 1888)
and vertically migrating food chains (Vinogradov, 1953 as
cited by Vinogradov, 1962). Vinogradov (1962), on the
basis of feeding patterns among the deep-sea zooplankton,
proposed that overlapping vertical migrations were the most likely mechanism of food transport from the productive
surface waters down to great depths. Menzies (1962) again
reviewed the theories of food sources and proposed the rain
of dead plankton as the most important food source on the
basis of his observations of isopods and the literature.
Studies related to the mechanism of food transport and the
food of deep-sea invertebrates have been reported by Krogh
(1934), Sokolova (1957,1959), Barnard (1962), Isaacs (1969)
Sanders and Hessler (1969), Harding (1973), Dayton and
Hessler (1972), Grassle and Sanders (1973), and others
(see Zenkevich and Birstein, 1956 for earlier studies). 3
Food of deep-sea pelagic fishes has received some attention in recent years (Marshall, 1954; Haedrich, 1964;
Haedrich and Nielson, 1966; Duka, 1969; Collard, 1970;
Childress and Meek, 1973; and others; see Merrett and Roe,
1974 for review), but less information is available for benthic and benthopelagic fishes. Stomach contents of some species have been reported cursorily in taxonomic and other works (Bigelow and Schroeder, 1953; Marshall, 1954;
Cohen, 1958; Nielson, 1964; Marshall, 1965; Bright, 1968;
Robins, 1968; Marshall and Iwamoto, 1973; McDowell, 1973).
Bright (1970) examined the stomachs of many species of deep-sea fish from the Gulf of Mexico and constructed a food web based on data from 81 small specimens. Clarke and
Merrett (1972) discussed the significance of pelagic food in the stomachs of deep living benthic fishes they examined.
Haedrich and Henderson (1974) and Pearcy and Ambler (1974) studied the food habits of macrourids of the genus
Coryphaenoides.
Dayton and Hessler (1972) predicted that deep-sea benthic fishes should be extreme food generalists, preying on populations of smaller deposit feeders, and thus maintaining a high diversity of deposit feeders by reducing the probability of competitive exclusion. Sanders (1968) and Grassle and Sanders (1973) speculated that deep-sea benthic carnivores are not generalists in their food habits, but would be expected to have specialized feeding habits.
They admitted that their statements were speculative 4 because of a lack of data. The purpose of this report is to quantitatively describe the food habits of several species of fishes from the continental slope and rise, from these data to provide a clearer picture of trophic structure in these habitats, and to evaluate the role of these fishes in maintaining the high diversity (Sanders, 1968; Sanders and
Hessler, 1969) of the deep-sea benthic fauna. METHODS AND MATERIALS
Collection of specimens
In selecting species for examination, primary consideration was given to those species that are dominant on the continental slope and rise. Some dominant species, such as macrourids, were avoided because they usually regurgitate their food due to expansion of gas in the swimbladder. For this reason, only those fishes which appeared to have their stomachs intact were selected.
A total of 742 fishes comprising 14 species was examined (Table 1). Most specimens were collected from the
Norfolk Canyon area (bounded by latitudes 36°58'N and
37°10fN, and extending from 75m to beyond 2000m) and an open slope area to the south of Norfolk Canyon (bounded by latitudes 36°34'N and 46°46rN and encompassing the same depth as the canyon stations). Fishes from these areas were collected on cruise CI-73-10 of the R/V Columbus Iselin in June, 1973, and cruise GI-74-04 of the R/V James M.
Gilliss in November, 1974. Additional study materials were collected off the North Carolina and Virginia coast, including the Norfolk Canyon area, on the R/V Eastward, cruise E-2-74, (April, 1974), and cruise E-l-73 (April,
1973). Several specimens were collected in the Hudson
Canyon area on the R/V Delaware 11, cruise D-2-74, in May, 5 TABLE 1
List of Fishes Examined
SPECIES NUMBER EXAMINED
Order Squaliformes
CentroscyIlium fabricii (Squalidae) 4 Centroscymnus coelolepis (Squalidae) 5
Order Chimaeriformes Harriota raleighana (Pvhinochimaeridae) 10
Order Anquilliformes Nessorhamphus ingolfianus (Nessorhamphidae) 6 Synaphobranchus kaupi ( S~ynaphob ranch idae) 279 Simenchelys parasiticus (Simenchelyidae) 23 Derichthys serpentinus (Derichthyidae) 17
Order Notacarithiformes Halosauropsis macrochir (Halosauridae) 83 Aldrovandia spp. (Halosauridae) 39
Order Aulopiformes Bathysaurus agassizi (Bathysauridae) 13
Order Gadiformes Antimora rostrata (Moridae) 7 Phycis chesteri (Gadidae) 217 Lycodes atlanticus (Zoarcidae) 34 Coryphaenoides armatus (Macrouridae) 3
6 7
1974 (see appendix for station data). Fishes collected on E-2-74 and E-l-73 were caught using a 30 ft (9.1m) lined semiballoon trawl with the following stretch mesh dimensions: 3.81cm stretch mesh in the wings, 3.81cm stretch mesh in the body, 3.81cm stretch mesh in the cod end and 1.27cm stretch mesh in the cod end liner. All other fishes were collected using a 45 ft (13.7m) lined semiballoon trawl with 4.45cm stretch mesh in the wings,
3.81cm stretch mesh in the body, 3.96cm stretch mesh in the cod end and 1.27cm stretch mesh in the cod end liner.
Trawls were towed on the bottom for 30 minutes at depths less than 2000m and for 60 minutes at deeper stations.
Stomachs from selected fishes were either excised on board from fresh specimens or from preserved whole specimens.
Stomachs to be examined were labeled, fixed in ten percent seawater formalin in separate vials or cheesecloth bags, and later transferred to either 40% isopropanol or 70% ethanol for storage.
Stomach content analysis
In the laboratory, stomachs were cut open, their contents were identified, and food items were counted and sorted by taxa. Food items were identified to the lowest possible taxon. Whole or relatively undigested items were usually identifiable at least to genus. Frequently food items were digested to the point that only fragments remained. Some fish remains (myctophids) could be identified by comparing otoliths that occurred in stomachs 8
with the otoliths from whole fishes found in other stomachs
of the same predator species. Fish remains were tallied as
one fish for the numerical percentage analysis.
Cephalopod remains (beaks) were identified by Dr. Clyde
F. E. Roper of the United States National Museum and his
identification series were used as reference material.
Other beaks were identified using Clarke’s (1962) key, or
by comparison with beaks taken from whole squids caught in the trawl. Cephalopod remains in a stomach were counted as
one cephalopod unless there was more than one beak present.
Other fragments of animals such as crustacean parts,
polychaete setae, or ophiuroid fragments were counted as
one animal. Frequently, numerical abundance could be
estimated by counting pairs of eyes or antenna! scales
(crustaceans), discs (ophiuroids), and other parts.
Several methods were used to measure the contribution
of different food items to the total diet, since some methods of food habits analysis are biased, either in favor
of a few bulkier items or very abundant small items (Hynes,
1950; Pinkas et al, 1971; Windell, 1971). First, the number
of stomachs in which a food item occurred is noted and
expressed as a percentage of the total number of stomachs
examined for a given fish species (frequency of occurrence method). Second, the number of individuals of each type of
food was counted and expressed as a percentage of the total number of food items from all stomachs for a given fish
species (numerical dominance method). Thirdly, the volume 9 displacement of food items was measured and expressed as a percentage of the total volume of food from all stomachs examined of a particular species (volume displacement method). Volume displacement was measured by placing the food item in a calibrated vial or graduated cylinder and filling to the calibration mark with a self-leveling buret according to the method of McEachran (1973). In the case of small food items, displacement was estimated by measuring the displacement of several of them together and then placing the items on a grid to estimate the percentage that each item contributed to the total volume (Windell, 1971).
From these three measurements an index of relative importance, IRI (Pinkas, et al, 1971), was calculated as follows:
(N+V ) F=IRI
where N = numerical percentage
V = volumetric percentage
F = frequency of occurrence percentage
IRI = index of relative importance
This index has been useful in evaluating the relative importance of different food items found in fish stomachs
(Pinkas, et al, 1971; McEachran, 1973). RESULTS AND DISCUSSION
In this section, the results of the stomach analysis and a discussion will be presented for each species. A general discussion will follow.
CentroscyIlium fabricii (CI-73-10, station 97; D-2-74, station 3B . 4 specimens examined)
Three of the four specimens of CentroscyIlium fabricii examined had food in their stomachs: one contained unidentified fish fragments, one contained a myctophid
(Ceratoscopelus maderensis), and the third contained crustacean fragments. The relatively intact condition of the myctophid indicates that it was probably consumed alive rather tLan scavenged from the bottom, although it may have been consumed in the trawl. Bigelow and Schroeder (1953) reported cephalopods, pelagic crustaceans, and medusae in the stomach of CentroscyIlium fabricii taken from Greenland waters. Clarke and Merrett (1972) found decapod remains in one longlined specimen. As the only known food of this shark consists of cephalopods, myctophids, pelagic crustaceans, and medusae, it apparently depends on pelagic food. Whether this food is consumed in the water column or scavenged from the bottom is not known.
10 11
Centroscymnus coelolepis (CI-73-10 stations 61, 63, 98. 5 specimens examined)
Four Centroscymnus coelolepis were collected from wire lobster traps set in deep water (952-1171m), and one was caught by trawl. Of the four which were caught in traps, two had empty stomachs and the others contained only well digested flesh, possibly the bait from the traps. The trawl specimen had eaten a squid, Mastigoteuthis.
Bigelow and Schroeder (1948) reported an argentine
(Argentina silas) from one stomach they examined, suggesting that Centroscymnus coelolepis has a fish diet. Clarke and
Merrett (1972) found pelagic cephalopod remains, cetacean remains, and fish fragments in specimens that they caught on bottom longlines. The capture of deep-sea benthic organisms in baited traps is indicative of a scavenging habit (Shulenberger and Hessler, 1974). Capture of
Centroscymnus coelolepis in baited traps and on baited longlines, and the presence of cetacean remains indicates that they probably scavenge much of their food from sinking dead carcasses.
Harriota raleighana (CI-73-10 station 97; D-2-74 stations 2, 3B; E-l-73 station 46. 10 specimens examined)
Harriota raleighana stomachs contained small amounts of sediment and well-digested remains (’’shrimp paste”) . In addition, one specimen contained 1 small gastropod and pelecypod shell fragments, another contained 9 small gastropods and 3 amphipods, and a third had 6 small gastropods in its gut. Cestodarian parasites (Gyrocotyle 12 sp.) were present in the anterior part of the spiral valves of several specimens: three fish had only one, and three had two of the parasites. None of the Gyrocotyle were attached to the intestine wall, but rather they appeared to be free moving in the spiral valve. Harriota raleighana feeds in the sediment, probably using its elongated snout as a sensory probe to find small food organisms.
Nessorhamphus ingolfianus (CI-73-10, stations 48, 52, 82, 83, 84, 97. 6 specimens examined)
Nessorhamphus ingolfianus feeds mainly on pelagic euphausiids and sergestid shrimps. Crustacean remains were found in the 5 stomachs containing food. One stomach contained a large Sergestes arcticus. Three stomachs contained euphausiiu remains, probably Stylocheiron sp., and three contained only crustacean fragments. Beebe
(1935b) found euphausiid and Sergestes remains in six stomachs he examined.
Nessorhamphus ingolfianus is apparently a pelagic animal, though it may spend some time close to, or on, the bottom, as it is often caught in bottom trawls.
Synaphobranchus kaupi (CI-73-10, stations 49, 81, 82, 83, 85, 90, 91, 95, 97. 279 specimens examined)
Food items found in the stomachs of Synaphobranchus kaupi included three families of cephalopods, three orders of crustaceans, and two families of fishes. (Table 2 and
Fig. 1). Of the 279 stomachs examined 102 (36.6%) were empty. Table 2 Percent frequency occurrence (F), percent numerical
dominance (N), percent volume displacement (V),
and index of relative importance (IRI) of
food items in the stomachs of
Synaphobranchus kaupi
Taxon Item F N V IRI
Polychaeta unidentified fragments 0.7 1.5 0.3 1 Mollusca Cephalopoda Enoploteuthis sp. 1.12.2 1.4 2 Enoploteuthidae 0.7 1.5 0.9 2 Histioteuthis sp. 1.8 3.7 1.5 7 Illex illecebrosus 0.4 0=7 25.4 10 Ommastrephidae 0.7 1.5 0.7 2 Cephalopod eggs (?) 0.4 1.5 0.6 1 unid. Cephalopoda 1.8 3.7 2.0 10 total 678 14779 32.7 324
unid. shell fragments 0.4 0.7 0.3 trace Total Mollusca 7.2 15.7 33.0 349
Crus tacea Tanaidacea Neotanaidae 0.4 0.7 0.1 trace
Euphausiacea Stylocheiron elongatum 0.7 1.5 0.4 1 Stylocheiron maximum 0.7 1.5 0.6 1 Stylocheiron sp. 0.4 0.7 0.1 trace Thysanopoda micropthalma 0.7 1.5 1.3 2 Thysanopoda acutifrons 0.4 0.7 0.5 trace Thysanopoda sp. 0.4 0.7 0.8 1 Euphausiidae 5.7 14.9 3.1 103 total 9.0 TITS' 678 256
13 Table 2. Continued
Taxon Item F N V IRI
Decapoda Sergestes arcticus 4.7 9.7 8.7 86 Sergestes sp. 0.4 0.7 0.2 trace Penaeidea 0.4 0.7 0.5 trace Natantia 0.4 0.7 0. 7 1 Pentacheles sculptus 0.7 1.5 0.9 2 Pentacheles sp. 0.4 0.7 0.4 trace Brachyuran megalopae 0.4 0.7 trace trace total 7.1 14.9 11.5 187 unid. Crustacea fragments 25.4 --- Total Crustacea 35.4 37.3 18.3 1968
Pisces Ceratoscopelus 4.3 9.7 12.2 94 maderens is Myctophidae 2.2 4.5 10.2 32 Nemichthys scolopaceus 0 .4 0.7 13.7 6 unid. teleost fragments 12.5 26.9 12.1 488 Total Pisces 19.4 41.8 48.3 1748
Miscellaneous Fecal pellets 1.4 3.7 0.1 5 Sediment 1.4 -- -
Unidentified 14.7 remains
Percent containing food 63 .4
14 15
Figure 1. Percent frequency occurrence, percent number,
percent volume displacement, and index of
relative importance (IRI) for higher taxonomic
groups of food in the diet of Synaphobranchus
kaupi. PERCENT VOLUME PERCENT NUMBER - 0 4 - 0 3 20 - 0 3 - 0 4 |9.4 - 0 5 0 2 - 0 5 0 1 IOH O
- -
E NT FREQUENCY T EN C PER yahbacu kaupi Synaphobranchus 6.8 B 9.0 7.1 D . DECAPODA D. PHAUSIACEA EU C. PISCES A. . CEPHALOPODA B. 1748 IRI 4 2 3 6 5 2 187 16
Crustaceans occurred most frequently (35.4% of all
stomachs examined); euphausiids being more frequent (9.0%>)
than decapods (7.1%). Unidentified crustacean fragments were found in 25.47, of the stomachs examined. Fishes had a high frequency of occurrence (19.47,), and cephalopods were present in many stomachs (6.87,). Sediment was found
in 1.47, of the stomachs examined.
Fishes were the most abundant food item by numerical dominance (41.87,), followed by crustaceans (37.3%).
Euphausiids (21.6%) were numerically more important than decapods (14.97,). Cephalopods (14.97,) were as abundant as
decapods. Polychaetes and shelled mollusks were of minor numerical importance.
Fishes were the most important item volumetricaily in
the diet of Synaphobranchus kaupi. Fishes comprised 48.3% of the volume of all food items, and averaged 75.27, of the volume of food in stomachs in which they occurred.
Myctophids (including Ceratoscopelus maderensis) were the most important (22.4%), followed by one entire specimen of
Nemichthys scolopaceus (13.77,) and unidentified fish remains consisting mostly of bones (12.17,).
Cephalopods ranked second in volumetric importance
(32.77,), mostly due to the occurrence of an entire large
specimen of Illex illecebrosus (25.47,). Other unidentified ommastrephids, enoploteuthids (including Enoploteuthis),
Histioteuthis, and damaged beaks and gragments of flesh made up the rest of the cephalopod remains. 17
Crustaceans ranked third in volumetric importance
(18.37,). Decapods, especially Sergestes arcticus and other natantians, made up the bulk of the identifiable crustacean food (11.57o). Euphausiids were also important (6.87,).
Mollusk shell fragments and polychaete fragments were of minor importance volumetrically.
The index of relative importance shows fishes
(IRI=1748) are the most important food of Synaphobranchus kaupi. Cephalopods (IRI=324), euphausiids (IRI=256), and decapods (IRX=187) are also important foods. Polychaetes and shelled mollusks are insignificant in the diet of
S . kaupi.
Synaphobranchus kaupi feeds mainly on pelagic food items. Pelagic organisms comprised 89.27, of the total number and 97.07, of the total volume of food. Robins
(1968) speculated that synaphobranchid eels probably scavenge dead and dying bathypelagic fishes from the bottom. This feeding mode could account for the presence of pelagic fishes (such as Ceratoscopelus maderensis and
Nemichthys scolopaceus), pelagic squids, decapods, and euphausiids in the stomach of S_^ kaupi. However the depths at which S_^_ kaupi were taken is also the lower limit of the daily vertical migrations undertaken by many of these pelagic animals (Vinogradov, 1962; Voss, 1967; Gibbs, et al,
1971). Vertical migrations in slope waters would bring these prey species in close proximity to the bottom, enabling S^_ kaupi which lives on or just above the bottom 18
(Robins, 1968; Heezen and Hollister, 1971), to take advantage of this available food supply. Percent frequency of occurrence of pelagic fishes (mostly myctophids) in the stomach, in relation to time of day at which Synaphobranchus kaupi were captured (Fig. 2), shows more pelagic fishes consumed during midday and midafternoon than at night. The prey species are nearest to the bottom during midday and midafternoon. The nearly intact condition of many organisms found in Synaphobranchus kaupi stomachs also indicates that they probably are consumed alive, rather than being scavenged from the bottom.
Simenchelys parasiticus (CI-73-10 stations 48, 51, 52, 53, 57, 83, 91, 96, 97; D-2-74 station 3B; Gl-74-04 station 72. 23 specimens examined)
Simenchelys parasiticus generally contained varying amounts of well digested food. Seven stomachs (30.4%) were completely empty and two of the remainder contained only small amounts of sediment. Usually the entire gut would be filled and distended with digested material, but in several stomachs only traces of it were present. The consistancy of this material ranged from hard (potato-like) to soft (gelatin-like). Nothing that could be identified as a particular animal was found, however in one stomach pieces of what appeared to be blood vessels were observed.
Simenchelys paras iticus has been reported to be partially parasitic, burrowing into the flesh of living halibut and other fishes (Goode and Bean, 1896; Bigelow and 19
Figure 2. Percent frequency occurrence of pelagic fishes
in stomachs of Synaphobranchus kaupi in
relation to time of day of collection. N is
the sample size of kaupi and n is the
number of stomachs with fishes. n=8 50- >- o z LU 40- ZD LU O o LJ 2 a: LU 30- u_ cn i- cc
20 - LU.1 o° n=!4 O o CC n = 6 LU 10 CL
TIME 0201 0634 1434 1544 2112 2206 N 97 58 29 16 55 19 20
Schroeder, 1953). Gut contents of metamorphic and juvenile forms from the Pacific Ocean included epibenthic copepods and amphipods, while stomachs from eleven adults examined in the same study contained only portions of flesh (Solomon-
Raju and Rosenblatt, 1971).
The material found in the stomachs of S_^ parasiticus is probably flesh scavenged from dead or dying fishes.
The capture of S_^_ parasiticus in baited traps (Solomon-Raju and Rosenblatt, 1971) indicates that they are scavengers.
None of the specimens captured in this study were attached to other fishes. Those reported by Goode and Bean may have been attached to dead or dying fishes.
Derichthys serpentinus (CI-73-10, stations 48, 49, 50, 52, 53, 56, 81, 83, 91. 17 specimens examined)
Derichthys serpentinus stomachs contained crustaceans in varying states of digestion and Sergestes arcticus was the dominant food item found. (35.3% frequency of occurence and 87.4% of the total food volume). Six specimens (35.3%) had empty guts. One euphausiid, Stylocheiron sp., was the only other identifiable food item found. Four stomachs
(23.5%) contained unidentifiable crustacean fragments.
Beebe (1935a) found shrimps, one of which he recognized as
Sergestes sp., in five stomachs he examined.
Derichthys serpentinus is captured in midwater trawls
(Castle, 1970) and probably forages in the water column for food. It appears to be a specialized feeder, feeding almost entirely on Serges tes arcticus. Two of the four D_^_ 21 serpentinus stomachs which were collected at night contained only traces of crustacean remains and the other two were empty. Those fishes collected during the day in which food was present in the stomach, had freshly ingested shrimps in their guts, indicating some feeding periodicity by D. serpentinus.
Halosauropsis macrochir (CI-73-10 stations 90, 95, 96; E-2-74 stations 6, 27, 33, 34, 39, 41; D-2-74 stations 4B, 6B, Alvin 2; GI-74-04 stations 84, 86. 83 specimens examined)
Stomachs of Halosauropsis macrochir contained food items including sponge remains, polychaetes, echiurids, eight orders of crustaceans, and two classes of echinoderms
(Table 3 and Fig. 3). Sixteen (19.3%) stomachs were completely empty. Sediment was found in 48.2% of all stomachs examined.
Crustaceans occurred in 55.4% of all stomachs examined.
Small crustaceans were the most frequently occurring food item. Small isopods, similar in appearance and presumably one species, were found in 18.1% of the guts examined.
Small amphipods (13.3%,), decapods (9.6%,) and mysids (8.4%) were also frequent. Other small crustaceans (copepods,
3.6%; cumaceans, 2.4%,; tanaids, 1.2%) occurred less frequently. Echinoderms were occasionally found; opiuroids occurred in 3.6%. and holothurians in 2.4%, of guts examined.
Polychaete fragments and shelled mollusk fragments were found in several stomachs (5.9% and 4.8% respectively).
Echiurids were found in two stomachs (2.4%). Table 3
Percent frequency occurrence (F), percent numerical dominance
(N), percent volume displacement (V), and index of relative
importance (IRI) of food items in the stomachs of
Halosauropsis macrochir.
Taxon Item F N V IRI
Porifera sponge spicules 2.4 2.0 0.4 6
Polychaeta unidentified fragments 5.9 5.0 1.0 35
Mollusca gastropod shell fragments 1.2 5.0 0.1 6 pelecypod shell fragments 2.4 2.0 2.7 11 unid. shell fragments 1.2 - - - Total Mollusca 4.8 7.0 2.8 47
Echiurida Ikedaidae 1.2 1.0 30.5 38 Unidentified Echiurida 1.2 1.0 4.1 6 Total Echiurida 2.4 2.0 34.6 88 Crustacea Copepoda Calanoida 3.6 3.0 0.2 12
Cumacea Diastylidae 1.2 1.0 0.1 1 unid. Cumacea 1.2 1.0 0.1 1 total 2.4 2.0 0.2 5
Tanaidacea Neotanaidae 1.2 1.0 0.1 1
Isopoda unidentified 18.1 31.0 1.6 590
Amphipoda Monoculodes sp. 1.2 1.0 0.1 1 Liljeborgia sp. 1.2 2.0 0.1 3 Harpinia sp. 1.2 1.0 0.1 1 Ampeliscidae (Ampelisca sp.?) 1.2 2.0 0.1 3 Lysianassidae (Paracallisoma sp.?) 1.2 2.0 0.1 3
22 Table 3. Continued
Taxon Item F N V IRI
Pardaliscidae (Princaxelia sp.?) 2.4 2.0 0.3 6 Pardaliscidae 1.2 1.0 0.1 1 Stegocephalidae 1.2 1.0 0.1 1 unidentified 3.6 3.0 3.3 23 total 13.3 13.0 ZT3 237
Mysidacea Michthyops parva 1.2 1.0 0.1 1 unidentified 7.2 9.0 0.7 70 total 8.4 10.0 0.8 91
Euphausiacea Thysanopoda (?) sp. 1.2 5.0 0.3 6 Bentheuphausia amblyops 1.2 1.0 0.1 1 unidentified 1.2 2.0 0.3 3 total 3 7 5 8.0 0.7 31
Decapoda Hymenopaneus laevis 1.2 1.0 3.8 6 Benthesicymus bartletti 1.2 1.0 5.8 8 Sergestes rabustus 1.2 1.0 3.6 6 Sergestes sp. 2.4 2.0 1.5 5 Ac an t he phy r a sp. 1.2 1.0 12.5 16 Caridea (Acanthephyra sp.?) 1.2 1.0 18.4 23 Pentacheles sculptus 1.2 1.0 6.0 8 total 9 7 5 870 517 5 572
unid. crustacea fragment 27.7 - Total Crustacea 55.4 78.0 59.5 7618
Echinodermata Holothuroidea unidentified fragments 2.4 2.0 1.2 8
Ophiuroidea unidentified fragments 3.6 3.0 0.4 12 Total Echinodermata 6.0 6.0 1.6 46
Pisces ctenoid scales 1.2 1.0 0.1 1
Miscellaneous Sediment 48.2 - Unidentified
23 24
Figure 3. Percent frequency occurrence, percent number,
percent volume displacement, and index of
relative importance (IRI) for higher
taxonomic groups of food in the diet of
Halosauropsis macrochir. o CVJ Is- — 0 0 in —
O
I 13.3
CD CD *» i---- 1----- O o o o O O O O o o vr ro cvj — CJ ro i n CD yaawriN ±N30d3d 3 W m O A ±N3DH3d 25 Numerically, crustaceans were by far the most abundant food item (78.07,). Isopods comprised 31.0% of the total individual food items. Small amphipods, mysids, decapods and euphausiids were relatively abundant (15.0%, 10.0%, 8.0%, and 8.07, respectively). Shelled mollusks made up 7.0% of food items. Polychaetes (5.0%), ophiuroids (3.07o), copepods (3.07o), echiurids, cumaceans, sponges, holothurians (2.07o for each) and tanaids (1.0%) were less abundant. Crustaceans were the most volumetrically important food (59.5%). The larger decapods (51.6%) made up most of this volume, whereas the smaller crustaceans, though abundant, contributed little to volume. Echiurids contributed 34.67o to the volume of food. Amphipods represented 4.37. of the stomach contents, followed by shelled mollusks (2.87,), echinoderms (1.6%), isopods (1.6%) and polychaete fragments (1.0%). Sediment comprised most of the volume of gut contents (39.7%). Stomachs and intestines were frequently packed with sediment, with food items imbedded in the sediment. Crustaceans had the highest index of relative importance for Halosauropsis macrochir (IRI=7618). Isopods (IRI=590) and decapods (IRI=572) were the two most important identifiable crustaceans. Amphipods (IRI=257) and mysids (IRI=91) were also important. Echiurids, due to their greater volume, had a relatively high IRI (88). Echinoderms (IRI=46), polychetes (IRI=35) and euphausiids (IRI=31) were less important. Copepods, pelecypods, gastropods, 26 poriferans, cumaceans, and tanaids were of minor importance as food. McDowell (1973) speculated that Halosauropsis macrochir roams the bottom, gulping down infaunal and epifaunal organisms along with considerable amounts of non-nutritive sediment. Harrisson (1966) and McDowell (1973) point out that Halosauropsis macrochir has the largest mouth of any halosaur and that the musculature and bone structure are adapted for taking large gulps. Deep-sea photographs of halosaurids show them hovering just over the bottom with the head down, apparently searching and rooting in the bottom for food (Marshall and Bourne, 1964). The inferior mouth, long snout, and long tapering tail and anal fin adapt these fishes for this type of feeding behavior (Marshall, 1934; Marshall and Bourne, 1964). McDowell (1973) believes the snout of halosaurs to be too weak to "root in the oozes” and turn up food organisms as Marshall and Bourne (1964) describe. He believes that the snout is used primarily as a sensory probe. The results of this study suggest that Halosauropsis macrochir is strictly a benthic feeder. The high incidence and volume of sediment in the stomach and the exclusive occurrence of benthic food organisms indicates a totally benthic feeding mode. Most food organisms, such as cumaceans, the amphipods Monoculodes sp., Liljeborgia sp. and Harpinia sp. (E.L. Bousefield, personal communication), polychaetes, echiurids, tanaids, and ophiuroids are 27 infaunal. Some epifaunal predation or scavenging may take place as evidenced by the presence of large epifaunal decapods in some stomachs. One stomach contained a caridean in an advanced state of digestion, indicating that it may have been scavenged. Most shrimps, however, were relatively intact. Many specimens (27.77>) contained what McDowell (1973) has called ’’shrimp paste". He believed this to be partially decomposed detritus. This "shrimp paste" may represent scavenged dead crustaceans. Aldrovandia spp. (CI-73-10 stations 57, 83, 90, 91, 95, 97; E-2-74 stations 27, 28; D-2-74 station 2. 39 specimens examined) Several species of halosaurs of the genus Aldrovandia were examined. Because of the relatively few specimens examined, and the similarity in morphology and distribution of these fishes, all species were combined for the IRI calculation (Fig. 4). Table 4 shows the results of this analysis and notes the occurrence of the various food items in the different species of Aldrovandia. A total of 39 specimens were examined; 18 Aldrovandia phalacra, 11 affinis, 2 A^_ gracilis, 7 specimens of an undescribed species, and 1 unidentified specimen. Of these specimens, 11 (28.2%) had empty stomachs. Food items consisted of polychaetes, shelled mollusks, small crustaceans (mostly amphipods) and brittle stars. Sediment occurred in 20.5% of the stomachs examined. Crustaceans were the most frequently occurring food (41.0%). Unidentified crustacean fragments occurred in 28 Figure 4. Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance (IRI) for higher taxonomic groups of food in the diet of Aldrovandia spp. 3I!N tmOA ± Dd3d 3D ±N A O ftim 3 d 3 d 0 3 N ± U30IA!nN Aldrovandia spp. cr O <3- D U L LU O O CD < Q_ X (D O >- CL o 0 0 O < O ro -1 ---- o o d n ro in cd oo ro . a o LU CL -J LU LU - O >- < CL Q- o o o O — <\J O o O < Q 1 "T Q CD < c h- O O O < < D O CD CC CL LU Q n LU U_ 0 1 CDT" > CO CL X ZD cc o l CD _ ro o LU < < O Is-' Is- O L n i — lc\J . .. .. CJ o 12.8 - 17.9 ro o O m O s ' Is- Is- PERCENT FREQUENCY Table 4 Percent frequency occurrence (F), percent numerical dominance (N), percent —1 r— nO •r-l rP TJ •p nO Ip p P r •H •P > o cp cu P Pi co cd CU o u cu B CU d cd p d p CU d cd o UO CU X > CU tx o p B cd d o cu o «s d cd CO a a O * UTpUUAOJtpXV s *d AOU d tV p J O A B U p T U *dS 'sTpTDUJL^ * Y ■ejLOBjTS TTJ *V STUTJJF 29 • d q M > P^! M F i—i p y p O) B 00 o v m o ^ r 1—1 » CNI r». cp nO ^ i— r^i nO 0 ucd a cu d d . • • • • n «p «p IP s O JPc cd cd P cu d CO CJ 0 pH O 1 cu Ov 1 o CO Oin CO 00 rP i— 1 cd cu d 0 d d O ) - • VO vO v a i— r-> i m d r o0 no no CO — p p P 0 P p cd d CO O p cd CO o d d cu p. a cd d cd O o O cd Cu CO • • i i vO 04 cp nO r—1 1 •P p — cd §3 p CO cu d d CO d d B 1 I I • • 1 n i VO 00 VO - o a> rP o CNI «n i— rP tP rP S H X X P cd CO d 0 O cd O • i O p cd no a) p U 0 CO vo O cn oi ro nO nO •r"I O p X p cd o P O co o cd § O p 0 CO Table 4. Continued u o a V X p a O A U U p T F s d * v x p r : o A e u p T u s d * SJUTJJB *V —l r O V o c i i— i i— l - r D v o o n i o o n 1-1 i o r o » - r o c o X •H •r4 •rl XJ o u i v c X o 4J 0) U cd d o P TO p CU cd o p cu p o • • • • • o t < X •rH X rH O 30 X cd cd cd a • t-4 X - > r r—l I O C N I ■U 1 — 4-» X cd X o • t < ^ o o N | l - r O V p v n i h n i t i i— t i— i i— r-O O O - r o c o c o c *H j cn < d •H xi 1 — CJ X X cd TO Cu CU TO P o cd e P P • • • • • • • • • • • h 3 6 •H •1—1 i n c X cd TO X u P cd d p. • h * r n i 2 6 X •rH X •i-lX o 4-4 m cd u cu cu o o CU 7 3 X . ^ x •l—l •1—1 •l—l 4J CU 1 i— d -u d cu d d 1 3 9 X p I o> P < —i i— o o CTi l - r N C T O C O C 4-» 1 — X X X cd o • • • • O V p i '* O o o O C CNJ •H x> PQ •H £ TO TO TO X o o u cd CU >- cu cd e TO o cu • 3 3 vo U-l CNJ n i X •rH u ■u ■u X X cd cd cd X X X cu X a g *-l X X o TO d p P i i i • O C m ± < CNJ —1 1— o - ^ r I— I X H CJ 4~> 4-1 cd X cd cd TO o 0) O B •• • • • 5 7 • P V O v Q V O O C o O C O C O C —i n i i i— i n rP N CNJ CNJ •rH •rH r-H •H o •rH X o — u cd o P W p p p cu o P e e • • i in i i i— i vo vo X M-l •rH •rH •H X 4-4 X X cd u P cu CU d d O v I— I X r—l •rH n i —I T— rCl •H H o 4-4 X u cd O cd P o cu P • X 3 n i CNJ O •H CO x 4-> X X cd X X cu o CU CU TO P TO d cu o d 1 I 1 X U-4 5 *H •H X •H •H < CsJ P V 4J cd Vl CU d (U cu TO 6 d X X X X t 1 1 1 - •H pH •H M-4 X 00 4-1 ■u }-l cu cu cd o o o d 00 o o d d d 31 25.6% of the guts examined. Amphipods had a high frequency of occurrence (17.9%). Copepods (7.77,), ostracods (2.6%) and mysids (2.6%) occurred less frequently. Pelecypod mollusks were a frequent food item (12.87,). Polychaetes (7.77,), gastropods (5.17,), and ophiuroids (5.1%,) were also consumed. Small crustaceans were also the most important food group by number (57.17,), with amphipods representing the most abundant item (39.37,). Shelled mollusks (25.07,), especially pelecypods (17.97,), were also relatively numerous. Polychaetes (10.77,), copepods (10.77,), and ophiuroids (7.1%) were less abundant. Ostracods and mysids were of minor numerical importance (less than 47,) . Polychaetes (47.57,) dominated the volume of food items. Crustaceans (29.4%), especially amphipods (16.9%), and shelled mollusks (19.6%), especially pelecypods (13.67,), were also volumetrically important foods. Other small crustaceans and ophiuroids in the diet contributed little to food volume. Sediment comprised 29.57, of the total volume of gut contents. Crustaceans (IRI=3547), especially amphipods (IRI=1006) were the most important food items found. Shelled mollusks (IRI=687), especially pelecypods (IRI=403), and polychaetes (IRI=448), are also important prey of Aldrovandia spp. Calanoid copepods (IRI=108), gastropods (IRI=66), and ophiuroids (IRI=55) were of lesser importance and mysids (IRI=33) and ostracods (IRI=10) were of minor importance. 32 McDowell (1973) considered tiny crustaceans, mainly amphipods and copepods, as the main food items for fishes of the genus Aldrovandia, with small mollusks occurring only rarely. The results of the present study agree that crustaceans are an important food, but disagree in that pelecypod mollusks were also shown to be important food for some species. Aldrovandia phalacra stomachs which were examined in this study contained copepods, infaunal amphipods, crustacean fragments, sediment, and '’shrimp paste” . In addition to these foods, McDowell (1973) reported foraminifera, which may have been ingested along with sediment. Aldrovandia affinis had the greatest diversity in its diet of any Aldrovandia, feeding on polychaetes, mollusks, ostracods, mysids, and ophiuroids, in addition to copepods, amphipods, sediment, and shrimp paste. McDowell reported A^ affinis as the only species he found feeding on pelecypods and that it included polychaetes and small crustaceans in its diet. There is an apparent resource partitioning among the Aldrovandia species, with A^_ affinis feeding more on polychaetes and mollusks and the other Aldrovandia species feeding more on crustaceans. However, the small sample size in this study render any statements relative to competition and niche separation as speculative. One Aldrovandia gracilis stomach was empty and the second contained an infaunal amphipod (Harpinia sp.), crustacean fragments, a broken pelecypod valve and a small 33 brittle star. McDowell reported only crustacean and polychaete fragments in specimens he examined. The undescribed species of Aldrovandia contained only polychaete fragments and shrimp paste in its gut. The unidentified specimen contained mollusk fragments, crustacean fragments, sediment, and shrimp paste. Aldrovandia, like closely related Halosauropsis macrochir, feeds mainly on and in the bottom, probably hovering over the bottom, probing the sediments with the snout and gulping down infaunal organisms, detritus, and sediment simultaneously. The smaller mouth of Aldrovandia however, limits its food to smaller items. Bathysaurus agassizi (CI-73-10 stations 95, 97; E-2-74 stations 6, 27; D-2-74 station Alvin 2; GI-74-04 stations 72, 95. 13 specimens examined) Three (23.17,) of the stomachs of Bathysaurus agassizi contained fish remains, one (7.77,) contained a decapod crustacean, one contained sediment, and one contained only well digested flesh. Seven (53.87,) specimens had completely empty guts. Fish remains comprised the greatest volume of stomach contents (83.37o) , followed by the decapod (9.47,) , digested flesh (7.27o) and a trace of sediment. Bathysaurus agassizi is morphologically adapted for life on the bottom (Mead, 1966) and the stomach contents of the specimens examined here indicate feeding on or very near the bottom. A partially digested specimen of the benthic fish Halosauropsis macrochir (gnathoproctal length approximately 180mm), which had been swallowed tail first, 34 was the only identifiable fish remains. Glyphocrangon longirostris, the crustacean found in one stomach, is an epibenthic decapod and was also swallowed tail first. Observations of Bathysaurus agassizi from the D.S.R.V. Alvin show this fish sitting on the bottom, occasionally darting up, apparently in pursuit of food (J.A. Musick, personal communication). Antimora rostrata (E-2-74 stations 6, 33; GI-74-04 station 86. 7 specimens examined) Antimora rostrata is a dominant fish on the lower continental slope and rise (Musick, 1975). When captured, it almost always comes up with its stomach protruding through its mouth because of expansion of gas in the swimbladder. Only seven specimens which appeared not to have regurgitated their food were examined. Of these, three had empty stomachs. One had eaten a large penaeidean shrimp (Pleisiopeneus armatus) and another had consumed a smaller penaeidean (Funchalia villosa?) and a hyperiid amphipod. A third specimen had a brachyuran cheliped in its stomach, and another had cephalopod remains in its gut. Antimora rostrata appears to feed on pelagic food (cephalopod, hyperiid amphipods, and possibly Funchalia villosa (?) ) as well as benthic decapods (Pieisiopeneus armatus, branchyrans). It does not appear to forage in the bottom ooze for food. A^_ rostrata has a large mouth and is capable of taking large prey. 35 Phycis chesteri (CI-73-10 stations 49, 51, 53, 74, 75, 76, 81, 8T, 84, 85V 217 specimens examined) Food items of Phycis ches teri included polychaetes, shelled mollusks, cephalopods, eight orders of crustaceans, and several families of fishes. (Table 5 and Fig. 5). Ninety-seven of the stomachs examined (40.1%,) were empty. Crustaceans were highest in frequency of occurrence (44.77o), and were represented by crustacean fragments, decapods, amphipods, mysids, and euphausiids (19.47,, 12.4%, 8.3%, 5.1%,, and 4.1% respectively). Fishes were also a frequent food item (9.77o). Polychaetes and copepods occurred less frequently (3.7%, and 2.3%, respectively). Sediment was present in 5.5% of stomachs examined. Numerically, crustaceans comprised 87.4%, of all food items. Euphausiids were the most numerically abundant food item (63.6%,). This was due primarily to the occurrence of 259 small Meganyctiphanes norvegica in one stomach. Decapods and fishes were of equal abundance (8.3%), and amphipods were nearly as important (6.9%). Copepods, mysids, polychaetes, and cephalopods were less abundant (4.5%,, 2.9%,, 1.9%, and 1.0% respectively). Shelled mollusks, cumaceans, ostracods, and isopods were of minor importance. Although fishes ranked third in frequency and numerical abundance they were the most important item volumetrically (40.8%>). Myctophids made up most of the volume (29.0%). Cephalopods were relatively unimportant in frequency of occurrence and numerical dominance, but ranked second in percent volume displacement (28.5%,). Total crustacean Table 5 Percent frequency occurrence (F), percent numerical dominance (N), percent volume displacement (V), and index of relative importance (IRI) of food items in the stomachs of Phycis chesteri. Taxon Item F N V IRI Polychaeta unidentified fragments 3.7 1.9 1.4 12 Mollusca Gas tropoda Turridae 0.5 0.2 trace trace gastropod shell fragments 0.5 0.2 trace trace total 0.9 .5 trace 1 Pelecypoda Abra (lioica?) 0.5 0.2 0.1 trace Cephalopoda Illex illecebrosus 0.9 0.5 26.1 25 unid. squid 0.9 0.5 2.4 2 total 1.8 1.0 28.5 53 Total Mollusca 3.2 1.7 28.6 97 Crustacea Ostracoda unid. ostracod 0.5 0.2 trace trace Copepoda Pareuchaeta norvegica 2.3 4.5 0.3 11 Cumacea Diastylis sp. 0.5 0.2 trace trace Campylaspis sp. 0.5 0.5 trace trace total 0.9 0.7 trace 1 Isopoda unidentified 0.5 0.2 0.2 trace Amphipoda Ceradocus (?) sp. 1.4 1.0 1.5 3 Neohela monstrosa 2.3 2.1 1.4 8 Ampelisca sp. (nr, declivitatis) 0.5 0.2 trace trace Hippomedon (?) sp. 0.5 0.2 trace trace Lysianassidae 0.5 0.5 trace trace Eusiridae (nr. Rhachotropis) 0.5 1.2 trace 1 Phoxocephalidae (new genus near Ha 0.5 0.2 trace trace 36 Table 5. Continued Taxon Item F N V IRI Stegocephalidae (?) 0.5 0.2 trace trace unidentified 2.3 1.2 0.6 4 total 8.3 6.9 3.6 87 Mysidacea Pseudomma roseum 1.8 1.2 0.1 5 Boreomysis tridens 0.5 0.2 0.2 trace B . arctica 0.5 0.2 0.2 trace unidentified Mysidae 2.3 1.2 0.2 3 total 5.1 2.9 0.7 18 Euphausiacea Bentheuphausia amblyops 0.5 0.2 trace trace Meganyctiphanes norvegica 1.8 62.4 3.8 119 Stylocheiron elongatum 0.5 0.2 0.2 trace unid. Euphausiid 1.4 0.7 0.3 1 total 4.1 63.6 4.2 278 Decapoda Sergestes arcticus 8.8 6.2 8.9 132 Serges tes sp. 1.8 1.0 1.0 4 Penaeidea 0.9 0.5 3.3 4 Sclcrocrangon agassizi 0.5 0.2 2.1 1 Pentacheles sculptus 0.5 0.2 1.9 1 Geryon quinquedens 0.5 0.2 7.8 4 total 12.4 8.3 20.2 353 unid. crustacea fragment 19.4 - - Total Crustacea 44.7 87.4 29.2 5212 Pisces Aldrovandia rostrata (?) 0.5 0.2 7.8 4 Cyclothone sp. 0.5 0.2 trace trace Sternoptyx diaphana 0.5 0.2 0.6 trace Ceratoscopelus maderens is 3.2 3.1 20.6 76 Myctophidae 2.3 1.9 8.4 24 Lycenchelys verrilli(?) 0.9 1.2 1.2 2 unidentified remains 2.8 1.4 2.1 10 Total Pisces 9.7 8.3 40.8 475 Miscellaneous fecal pellets 0.5 0.7 trace trace sediment 5.5 - Unidentified ’’shrimp paste” 4.1 - 37 38 Figure 5. Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance (IRI) for higher taxonomic groups of food in the diet of Phycis chesteri. o r- Phycis chesteri 9£ £ 1 o CD CO O UJ . a CO * f 8 l d 3 d 0 3 N ! flN 38W U < ro <£ ro O UJ Q m CL O CO O o in UJ < o CO X < 3 OJ 3 J< UJ o s GO Is- GO Q. o o o < Is- X jUL uj Q . 0 OlM o (III < UJ o X o CL o 00 ro <£ -J o 0L in ro — UJ o o < o CL X o CL . — < 0Q c 111-— L z c x I L t q l | , . , T n , .. , , vt-‘ 5.1 MmOA A O m lM 3 to GO q o CVJ — 3.7 ~~ 12.4 o ro d 3 d 0 3 N ! - - - - - 8 -J O oS N- o PERCENT FREQUENCY 39 percent volume (29.2%,) ranked higher than cephalopods, however no individual higher taxon within the Crustacea was more important than the Cephalopoda. Decapods (20.2%,), euphausiids (4.2%), and amphipods (3.6%) made up the bulk of crustacean food. Ostracods, copepods, cumaceans, mysids, polychaetes and shelled mollusks were of minor volumetric importance. The index of relative importance indicates that, in general, crustaceans are the most important food for Phycis chesteri (IRI=5212). However, no individual taxon within the Crustacea was as important as fishes (IRI=475). Decapods (IRI=353), euphausiids (IRI-278), and amphipods (IRI=87) are the most important crustacean foods. Cephalopods (IRI=53) are a less important food item. Mysids (IRI=18), polychaetes (IRI=12), copepods (IRI=11), and gastropods, pelecypods, ostracod.s, cumaceans, and isopods (IRI less than 1 for each) were of lesser importance. The diet of Phycis chesteri, as presented, is very similar to that of closely related Urophycis chuss, U . tenius, an<^ IL_ ^egius (Bigelow and Schroeder, 1953) . P_;_ chesteri feeds extensively on both pelagic and benthic food items. Pelagic food items comprised 84.9% of the total number and 76.6%, of the total volume, and included squids (Illex illecebrosus), calanoid copepods (Pareuchaeta norvegica), amphipods (Ampelisca, Eusirids), euphausiids (Meganyctiphanes norvegica and Stylocheiron elongaturn) , decapods (Sergestes arcticus) and fishes (Cyclothone sp. , Stemoptyx diaphana, 40 myctophids). Again, as in S_;_ kaupi, pelagic fishes in the stomachs of P_^_ chesteri increased in frequency of occurrence during the day (Fig. 6). Euphausiids were also more frequently consumed during the day (Fig. 6), when they would be at their lower limit of vertical migration (Mauchline and Fisher, 1969). Benthic food items included polychaetes, gastropods, pelecypods, cumaceans, amphipods, decapods and fishes. Also, the presence of sediment in several stomachs indicates that I\_ chesteri feeds on the bottom. Most of the amphipods are believed to be either epifaunal (Ceradocus (?) sp. and Neohela monstrosa) or infaunal (Hippomedon (?), Phoxocephalidae) (E.L. Bousefield, personal communication). Phycis ches teri probably feeds on and just above the bottom. This fish appears to be an active predator on pelagic and benthic crustaceans and fishes. The capture of P . chesteri in baited lobster traps during cruise CI-73-10 indicates that they may also scavenge dead organisms. Lycodes atlanticus (CI-73-10 stations 85, 90, 95, 97; E-2-74 stations 28, 39; E-l-73 station 46. 34 specimens examined) Food items of Lycodes atlanticus included sponge remains, polychaetes, shelled mollusks, pycnogonids, three orders of crustaceans, and brittle stars. None of the stomachs examined were completely empty, however some contained only sediment or nshrimp paste". Sediment occurred in 82.4% of examined stomachs (Table 6 and Fig. 7). Ophiuroids had the highest frequency of occurrence, occurring in 44.1%> of guts examined. Crustaceans occurred in 41 Figure 6. Percent frequency occurrence of pelagic fishes and euphausiids in the stomachs of Phycis chesteri in relation to time of day of collection. N equals the sample size of P . chesteri and n is the number of stomachs with fishes or euphausiids. CO Q CO 3 < X GL 3 UJ U_ 2 0 - o > y UJ ^ U J o c r 1 0 - u j c r 0 ^ 3 Ll O o ° ^ O TIME 0321 0736 1000 1607 2015 2112 2206 N 5 15 83 21 54 12 22 co UJ X CO u_ 20 - >- o o X UJ UJ o 3 UJ n=9 o c r UJ 10- c r c r u . 3 o o o TIME 0321 0736 1000 1607 2015 2112 2206 N 5 15 83 21 54 12 22 42 32.4% of all guts examined, amphipods occurring most frequently (26.5%), followed by ostracods (2.9%) and isopods (2.9%,). Shelled mollusks were frequent food items; pelecypods occurred in 29.4%, gastropods in 17.6%, scaphopods in 2.9%, and unidentified fragments in 8.8%, of stomachs examined. Polychaetes (14.7%,), sponge fragments (8.8%) and pycnogonids (5.9%) were also present. Pelecypods were the most important food by number (42.7%,). Crustaceans (15.5%>), mainly amphipods (12.6%), and ophiuroids (15.5%) and gastropods (14.6%) were also numerically important. Sponge fragments, polychaetes, pycnogonids, ostracods, and isopods were low in numerical dominance. Volumetrically, ophiuroids were the most important food (58.2%,). Several specimens of Lycodes atlanticus had their guts packed entirely with ophiuroids. Shelled mollusks (27.7%,) were also volumetrically important foods, with pelecypods comprising the greatest volume (16.9%,), followed by gastropods (9.3%) and scaphopods (1.5%). Crustacean food items (3.8%,) consisted of small amphipods (3.4%,), isopods (.3%,), and ostracods (.1%), and hence were not important foods volumetrically. Sponge fragments (4.9%,) and polychaetes (4.4%,) were as important volumetrically as crustaceans. The two small pycnogonids contributed little to volume (1.0%). Sediment represented 65.6%, of the total volume of stomach contents. Ophiuroids had the highest IRI (3250) of any higher Table 6 Percent frequency occurrence (F) , percent numerical dominance (N), percent volume displacement (V), and index of relative importance (IRI) of food items in the stomachs of Lycodes atlanticus. Taxon Item F N V IRI Porifera sponge spicules 8.8 2.9 4.9 69 Polychaeta Eteone longa (?) 2.9 1.9 2.4 12 Drilonereis longe (?) 2.9 1.0 0.2 4 unidentified fragments 8.8 2.9 1.7 40 Total Polychaeta 14.7 5.8 4. 4 150 Mollusca Gastropoda Scaphander punctostriatus; 14.7' 7.8 5.1 190 Atlanta peroni 2.9 1.9 0.3 6 Natica clausa 2.9 1.0 2.4 10 unidentified fragments 5.9 3.9 1.5 10 total 17.6 14.6 9.3 421 Pelecypoda Limatula sp. 5.9 12.6 1.5 83 Yoldia iris 2.9 3.9 2.4 18 Hyalopecten sp. 11.8 8.7 2.9 137 Lyonsie11a sp. 2.9 1.0 3.9 14 Malletia polita 2.9 8.7 4.4 38 unidentified fragments 17.6 7.8 5.3 231 total 29.4 42.7 16.9 1752 Scaphopoda Caludus (=Platyschides) 2.9 1.0 1.5 7 grandis unid. mollusk fragment 8.8 - — Total Mollusca 47.1 58.3 27.7 4051 Py cn og on i da Nymphon macrum 2.9 1.0 0.5 4 unidentified fragments 2.9 1.0 0.5 4 Total Pycnogonida 5.9 1.9 1.0 17 Crus tacea Ostracoda unidentified ostracod 2.9 1.0 0.1 3 Table 6. Continued Taxon Item F N V IRI Isopoda unidentified isopod 2.9 1.9 0.3 6 Amphipoda Harpinia sp. 17.6 7.8 2.0 172 Orcnomenella (?) sp. 2.9 1.0 0.3 4 Lembos sp. 5.9 2.9 0.7 21 unidentified 2.9 1.0 0.4 4 total 26.5 12.6 3.4 424 unidentified fragments 8.8 __ Total Crustacea 32.4" 15.5 3.8 625 Ophiuroidea Ophiacantha bidentata 2.9 1.0 2.0 9 Oph i omu s ium lyman i 2.9 1.9 0.5 7 unidentified fragments 38.2 12.6 55.7 2609 Total Ophiuroidea 44.1 15.5 58.2 3250 Miscellaneous Sediment 82.4 - - - Unidentified remains 8.8 -- - Percent containing food 100.0 44 45 Figure 7. Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Lycodes atlanticus. d38WflN ±N30d3d Lycodes atlanticus O IO CD o ro o O cvj O o — LU o d§ cvi cd m 1^-. cd oo o CO 3WfnOA ±N30d3d 3WfnOA - O O CJ CVJ CVJ lO LO r r CVJ cr> <3- D i CO I I liJ Q O CD < CL O m UJ — ° £ ° CVJ L LU CL O ZD C D < w < < o o § o 9 o — oJ O \ N ^ d- d ^ N C\J o — ro " > q _j ro o ^ Q_ o □_ m < CO CO 1— DC O o CL O O O — CO lO N cn O o >- o o LU O ^ cr < L L CL CL CL £ < LU c o cr j lO o _ Q o O ^ < CO o CO o sj- j- v PERCENT FREQUENCY 46 taxon of food for Lycodes atlanticus. Pelecypods (IRI-1752), small crustaceans (IRI=625), mainly amphipods (IRI=424), and gastropods (IRI=421) and polychaetes (IRI=150) were also relatively important food sources. Poriferans, scaphopods, pycnogonids, ostracods, and isopods were relatively unimportant foods. Although there is nothing in the literature on the food of Lycodes atlanticus, the food items found here are very similar to those reported by Andriashev (1964) for other species of Lycodes. L . atlanticus is primarily a benthic feeder, preying on infaunal and epifaunal organisms, especially ophiuroids, pelecypod mollusks and amphipods. Infaunal organisms included polychaetes, pelecypods, scaphopods, amphipods (Harpinia, Orchomenclla), and possibly the sponges and brittle stars. Epifaunal food items included sponge fragments, gastropods, pycnogonids, isopods, amphipods (Lembos), and ophiuroids. No pelagic food items were found in L^_ atlanticus. This zoarcid feeds primarily on infaunal organisms, apparently indiscriminately gulping down large amounts of sediment along with its food (one specimen had ingested a piece of coal). It is doubtful that they lie in wait for prey and pounce on it suddenly, as Marshall (1971) has speculated. Coryphaenoides (Nematonurus) armatus D-2-74 stations 6A, Alvin 2. 5 specimens examined) Stomach contents of Coryphaenoides armatus were mainly pelagic food items, including three families of cephalopods 47 and five orders of crustaceans. All stomachs examined contained food (Table 7 and Fig. 8). Amphipods were the most frequently occurring food item. Hyperiid amphipods occurred in all stomachs examined and gammaridean amphipods occurred in two stomachs. Unidentified crustacean fragments occurred in four stomachs, and decapods were found in three stomachs. Cephalopods, copepods, isopods, and euphausiids occurred with equal frequency (two stomachs). Amphipods were the most important food item by numerical dominance (50.0%). Decapods were also abundant (23.17o), followed by copepods (9.67o), cephalopods (5.87>), euphausiids (5.87o), and isopods (3.8%). Decapods made up the bulk of the volume of food (73.1%). Cephalopods (10.9%) and amphipods (8.67o) were less important. Euphausiids, isopods, and copepods were of minor volumetric importance. Crustaceans are the most important source of food for Coryphaenoides armatus. Amphipods (IRI=5860) and decapods (IRI=5772) are the most important identifiable food items. Cephalopods (IRI=668), copepods (IRI=409), and euphausiids (IRI=388) were less important as food. Isopods were of minor importance (IRI=260). Coryphaenoides armatus feeds mainly on pelagic crustaceans, especially amphipods and decapods. Pelagic cephalopods are also eaten. Haedrich and Henderson (1974) and Pearcy and Ambler (1974) found similar results. The presence of sediment, isopods, and gammaridean amphipods in Table 7 Percent frequency occurrence (F), percent numerical dominance (N), percent volume displacement (V), and index of relative importance (IRI) of food items in the stomachs of Coryphaenoides armatus. Taxon Item F N V IRI Mollusca Cephalopoda Histioteuthis sp. 20.0 1.9 0.2 42 Gonatidae 20.0 1.9 1.6 128 Thysanoteuthidae 20.0 1.9 6.2 162 total 40.0 5.8 10.9 668 Crustacea Copcpoda Euchirella rostrata 20.0 1.9 0.2 42 Calanoida 20.0 7.7 0.4 162 total 40.0 9.6 0.6 409 Isopoda Valvifera 40.0 3.8 2.7 260 Amphipoda Hyperiidae 100.0 46.2 7.8 5400 Gammaridea 40.0 3.8 0.8 184 total 100.0 50.0 8.6 5860 Euphaus iacea Nematoscelis microps 20.0 1.9 0.4 46 Thysanopoda acutifrons 20.0 1.9 3.3 104 unidentified fragments 20.0 1.9 0.2 42 total 40.0 5.8 3.9 388 Decapoda Paguridae larvae 20.0 1.9 0.2 42 Caridea 20.0 1.9 71.4 1466 Brachyura megalopae 40.0 19.2 1.4 824 total 60.0 23.1 73.1 5772 unid. malacostracan larvae 20.0 1.9 0.2 42 unid. crustacea fragments 80.0 Total Crustacea 100.0 94.2 89.5 18370 48 Table 7. Continued Taxon______Item F N V IRI Miscellaneous sediment 60.0 cellophane 20.0 Unidentified remains 20.0 Percent containing food 100.0 49 50 Figure 8. Percent frequency occurrence, percent number, percent volume displacement, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Coryphaenoides armatus. PERCENT VOLUME PERCENT NUMBE QZ 60- 50- 40- 30- 0 2 70 40 20 30 50 10 10 0 - - PERCENT FREQUENCY PERCENT oyheods armatus Coryphaenoides 100 A 60 B 40 40)40 D 40 C. F. ISOPODA D. COPEPODA B. E. EUPHAUSIACEA A. C PAOOA668 CEPHALOPODA DECAPODA AMPHIPODA 5772 5860 IRI 409 388 260 51 some stomachs suggests that some benthic feeding may occur. Pearcy and Ambler (1974) have reviewed the possible mechanisms by which the pelagic food is consumed by this supposedly benthic fish. They are probably correct in concluding that armatus preys on active pelagic animals and will opportunistically scavenge dead items from the bottom. This macrourid may move well away from the sea floor (Okamura, 1970), enabling it to forage in the water column. CONCLUSIONS Menzies (1962) discussed possible sources of food for deep-sea organisms and ranked these sources in the following order of importance: (1) rain of dead plankton (Agassiz, 1888); (2) turbidity currents (Heezen, Ewing, and Menzies, 1955); (3) living vertical migrations (Vinogradov, 1958); (4) floating benthic marine plants; (5) floating terrestrial plants (Agassiz, 1892; Brunn, 1957). He wrote that far at sea the rain of plankton might be followed in importance by living vertical migrations, but that near the continental margins the major sources of food, in order of importance, were turbidity currents, the rain of dead plankton, and floating terrestrial and marine plants. He also suggested that turbidity currents bring rich organic sediment and detritus from nearshore estuarine and terrestrial areas to deep oceanic waters. Submarine canyons are believed to be high incidence areas for turbidity currents (Heezen, Ewing, and Menzies, 1955). Even though much of the present study was carried out in Norfolk Canyon, nothing of direct shallow water origin was observed in any of the fish stomachs examined. Thus turbidity currents are probably not an important direct course of food for fishes. Other workers have reported small amounts of neritic algae and tracheophytes from deep-living fishes (Haedrich and 52 53 Henderson, 1974; Pearcy and Ambler, 1974), but there is no way to determine the route by which this material entered the deep sea. Sanders and Hessler (1969) have suggested that turbidity currents are of minor importance in food transport to the deep sea. Such currents may even reduce food supply to fishes by destroying the habitat and burying deposit feeders under thick layers of sediments (Heezen, Ewing, and Menzies, 1955), and by selecting for large macrobenthic organisms (Rowe, 1972), most of which are too large to be ingested by benthic fishes. Consequently, turbidity currents are probably of neutral or even negative value to fishes which are benthic feeders. Turbidity currents may indirectly play an important role in the nutrition of fishes which feed on pelagic prey by resuspending detritus particles and increasing the food supply for planktonic animals near the bottom, thus increasing the density and availability of pelagic animals to serve as food for benthic fishes. Phycis chesteri, Synaphobranchus kaupi, Antimora rostrata, and Coryphaenoides armatus, dominant species from the upper, middle, and lower slope and rise respectively, feed heavily on pelagic euphausiids and decapods,which in turn feed on suspended detritus particles. In addition, Vinogradov's (1953) vertically migrating food chain appears to be the most important food source for many dominant deep-sea fishes. Many of the euphausiids, decapods, myctophid fishes and cephalopods which are important in the diet of P^ chesteri, S . kaupi, A. rostrata, 54 and armatus are also species which undergo extensive daily vertical migrations. On the continental slope, these vertical migrations bring mesopelagic animals down to the bottom, where they are available as prey for the dominant fishes, Synaphobranchus kaupi and Phycis chesteri. Other dominant benthic slope fishes, such as macrourids, also feed heavily on vertically migrating animals (Podrazhanskaya, 1967; Okamura, 1970; Marshall and Iwamoto, 1973). Vertical migrations may be an important source of food for abyssal fishes, such as Antimora rostrata and Coryphaenoides armatus, which had pelagic food items in their guts. Menzies (1962) is probably correct in asserting that the rain of dead plankton Is the most important food source in the oceanic realm, although these small refractory particles are of little use to fishes directly. Filter and deposit ‘feeding animals were important food items for the deeper living fishes examined in this study (Halosauropsis macrochir, Lycodes atlanticus, and in smaller Coryphaenoides armatus [Haedrich and Henderson, 1974]) and in stomach contents of fishes reported by other workers (Nielson, 1964; Bright, 1968, 1970). Rains of small particles support populations of deposit and filter feeders, which in turn serve as food for the benthic deep-sea fishes, (Halosauridae, Zoarcidae, some Macrouridae, chimaeroids). Indiscriminate rooting in the ooze and ingestion of sediment and benthic invertebrates, that many fishes exhibit, may play a role in maintaining the high diversity 55 (Sanders, 1968; Sanders and Hessler, 1969) in deep-sea communities of deposit feeding invertebrates. Disturbance caused by uprooting and ingestion of infaunal and epifaunal animals by species such as Halosauropsis macrochir, Lycodes atlanticus, and Aldrovandia spp., and some macrourids (Marshall and Bourne, 1964) may regulate the population sizes of these prey organisms, thus reducing the probability of competitive exclusion (Dayton and Hessler, 1972). In support of this hypothesis, and as an alternative to the stability-time hypothesis (Sanders, 1968; Sanders and Hessler, 1969), Dayton and Hessler (1972) have suggested that deep sea predators such as fishes should be non- selective ,fcroppersM in feeding, ingesting living particles exclusively or In combination with dead or inorganic material. Other workers (Ivlev, 1961; Schoener, 1971) have also theorized that in areas of low food availability such as the deep sea, predators should be generalized feeders. Cropping benthic feeders examined in the present study exhibit little selectivity in their feeding. They consume any item which is small enough for them to eat, although no comparison of stomach contents with the benthic fauna can be made. Species with larger mouths (Halosauropsis macrochir, Lycodes atlanticus) consume some larger prey than species with smaller mouths (Aldrovandia spp.), but there is much overlap in food items. Amphipods were important food items for Halosauropsis macrochir, Aldrovandia spp. Lycodes atlanticus, and Coryphaenoides armatus, with several genera 56 shared as common food. The wide variety of food items consumed by these croppers indicates a generalized diet. The stomach contents of Halosauropsis macrochir included seven phyla, many classes and orders within these phyla, and at least 30 species. Lycodes atlanticus had consumed five phyla of invertebrates and at least 21 species. Aldrovandia spp. fed on four phyla with representatives of at least 10 species. Stomach analyses on other species of deep living fishes have shown similar results (Bright, 1968, 1970; Haedrich and Henderson, 1974; Pearcy and Ambler, 1974). Benthic fishes which fed mostly on pelagic food items (Synaphobranchus kaupi, Phycis chesteri, Antimora rostrata, Coryphaenoides armatus) also had generalized diets. Since they feed little or not at all on deposit feeders, they would not be important in reducing competitive exclusion at that trophic level. However they are important in transferring energy from pelagic to benthic ecosystems. Also, they may be important in dispersing large isolated falls of dead food (Issacs, 1969; Clarke and Merrett, 1972) by consuming them above or on the bottom and spreading them out as feces available to deposit feeders (Dayton and Hessler, 1972). The "shrimp paste" found in many stomachs and reported by other workers (McDowell, 1973) may represent scavenged larger parcels of flesh which are in the process of being dispersed over the bottom. Scavengers such as the deep-sea shark Centroscymnus coelolepis, and the eel Simenchelys parasiticus, probably function at this trophic 57 level. Several species of fishes had consumed large amounts of sediment. Many specimens of Halosauropsis macrochir, Lycodes atlanticus, Harriota raleighana, and some Aldrovandia had a greater volume of sediment (filling the entire gut cavity) than food in their stomachs. How much nutrition is derived from this sediment is not known. McDowell (1973) believed the sediment he found in halosaurids to be "non-nutritional" but gave no supportive evidence for this conclusion. Whether deep-sea fish can obtain nutritional value by ingesting sediment can only be determined by future research. PPENDIX Appendix Cruise, station number, locations, and mean depths of stations from which stomachs were collected Cruise Station Latitude Longitude Depth number______number______(N)______(W)______(m) 48 36°34.75 f 74° 38.9' 751 49 36°41.51 74°37.4' 743 50 36 °44.251 74°35.41 786 51 36°38.5 ' 74°39.4' 681 52 36°33.51 74°42.751 274 53 36°35.41 74°42.01 753 56 36°35.6' 74°37.5' 1190 57 36°43.61 74°32.8' 1194 61 37°02.71 74°34.45' 1171 63 37 °02.5’ 74°35.5’ 952 74 37 °06.2' 74°40.31 249 75 37 °04.6’ 74°36.6’ 403 76 37°09.01 74°34.41 270 81 36°56.25 f 74°36.6 f 729 82 37 °04.4' 74°32.01 716 83 37°03.5’ 74°33.31 986 84 37°03.21 74°34.1’ 636 58 Appendix. Continued Station Latitude Longitude Depth number______(N)______(W) (m) 85 37 °03.91 74 ° 31.5' 776 90 36°54.65’ 74°27.5' 1488 91 37°03.0' 74°20.6' 1719 95 37°01.61 74°22.5 1 1591 96 36°45.41 74°17.5' 1678 97 36°50.21 74°32.5' 1350 6 35°23.2’ 74°43.4' 2105 0 O 27 -vj 00 74°24.4' 1670 28 37°07.6' 74°19.4' 1617 33 37°03.8' 74°07.3' 2165 34 36 °58.1' 74°11.0' 2200 39 36°09.2' 74°18.91 2060 41 35°17.51 74°43.8' 2195 2 39°09.01 72°19.5' 952 3 39°04.0' 72° 32.01 1142 3A 39°02.01 72°29.0' 1352 3B 39°02.0' 72 °25.0' 1207 4B 38°50.0' 72°34.5' 2196 4C 38°43.0' 72°30.0' 2379 6A 38°28.01 71°55.0' 2745 6B 38°38.O' 72°06.0' 2379 Alvin 2 39°10.0' 71°50.5' 2288 59 Appendix. Continued Cruise Station Latitude Longitude Depth number number (N) (W) (m)__ GI-74-04 72 36°36.3' 74°24.8' 1730 84 36°36.O' 74°24.4' 1763 86 36°41.6' 73°47.0' 2642 95 37°05.0' 74°12.5' 2100 98 36°59.71 74°33.5' 750 E-l-73 46 35°49.1' 74°33.0' 1900 60 LITERATURE CITED Agassiz, A. 1888. 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