OCS Study 

MMS 87-0060 

Northern Gulf of Mexico 

Continental Slope Study 

Annual Report 

Year 3 

Volume II : Technical Report 

\ " ~i " ~7" ?~1t :: 

.S 

LORICFERA X1126 

U.S. Department of the Interior 

Minerals Management Service 

Gulf of Mexico OCS Regional Office 
COVER: 

In 1983, Reinhardt gristensen of the University of Copenhagen, Denmark 

described a new phylum of animals, Loricifera, based upon specimens 

collected in 25 to 30-m depths near Roscoff, France ; from 480-m deep 

waters near the Azores and from 15-m depths off Fort Pierce, Florida. 

This new phylum depicted on the cover was found to be well represented 

(506 specimens) across the entire continental slope of the northern Gulf 

of Mexico, from Florida to Texas, at depths from 298 to 2959 m" 
OCS Study 

MMS 87-0060 

Northern Gulf of Mexico 

Continental Slope Study 

Annual Report 

Year 3 

Volume II : Technical Report 

Contributing Editors 

Benny J . Gallaway 

Larry R. Martin 

Randall L. Howard 

Prepared under MMS Contract 14-12-0001-30212 

Published by 

U.S. Department of the Interior 

Minerals Management Service New Orleans 

Gulf of Mexico OCS Regional Office February 1988 
111 

DISCLAIMER 

This report was prepared under contract between the Minerals Management 

Service (MMS) and LGL Ecological Research Associates, Inc. and Texas A&M 

University. This report has been technically reviewed by the MMS and 

approved for publication. Approval does not signify that contents 

necessarily reflect the views and policies of the Service, nor does 

mention of trade names or commercial products constitute endorsement or 

recommendation for use. It is, however, exempt from review and compliance 

with MMS editorial standards. 

REPORT AVAILABILITY 

Extra copies of the report may be obtained from the Public Information 

Unit (Mail Stop OPS-3-4) at the following address: 

U .S . Department of the Interior 

Minerals Management Service 

Gulf of Mexico OCS Regional Office 

1201 Elmwood Park Boulevard 

New Orleans, Louisiana 70123-2391 

Attention: Public Information Unit (OPS-3-4) 

Telephone Number : (504) 736-2519 

CITATION 

This study should be cited as : 

Gallawaq, Benny J ., Larry R . Martin and Randall L . Howard 

(Eds .) . 19$8 " Northern Gulf of Mexico Continental Slope 

Study, Annual Report : Year 3 . Volume II : Technical 

Narrative. Annual report submitted to the Minerals Management 

Service, New Orleans, LA. Contract No . 14-12-0001-3212 . OCS 

Study/MMS 87-0060 . 586 pp. 
1V 

ACKNOWLEDGEMENTS 

Many persons (over 100) deserve acknowledgements for the completion 

of this phase of the study including the crews of the research vessels ; 

the overworked, underpaid field sampling and laboratory technicians and 

their supervisors ; the taxonomic consultants ; the data management staff at 

LGL; the Contracting Officer's Technical Representative ; and members of 

the Scientific Advisory Committee (SAC) . All of these have been listed by 

name in previous reports and we again gratefully thank them for their 

efforts . 

Two people deserve special thanks for these volumes. First we 

acknowledge Ms . Jean ErWin who typed and handled all phases of the 

production of not only this set of final tomes, but also their preceding 

drafts . In fact, she should probably be given credit as the rightful 

editor of this year's report, but following common practice among 

scientists and their staff, others will be given the formal credit. The 

project participants all know who did most of the editorial work . Thank 

you Jean. Second, we especially acknowledge the scientific illustrations 

of the fish and decapod crustaceans prepared by Mrs . Jan Fechhelm . We 

believe these to be especially good and accurate, and suspect that they 

will reappear again and again in the formal literature--without credit--as 

do most illustrations of this type. Thank you Jan, we are proud to be the 

original source of your excellent illustrations. 

Several persons contributed substantially to the writing of this 

report, although we must emphasize that this project was truly a team 

effort with much interaction and discussion of ideas, results and 

implications of the findings . However, below we recognize 

(alphabetically) those who "put words on paper" . 

Joshua S . Baker G . Fain Hubbard 

Gregory S . Boland Mahlon C . Kennicutt, II 

James M . Brooks George S . LeWbel 

Charlie R . Chandler Lynn M. Maritzen 

Bennq J . Gallaway Linda H. Pequegnat 

Randall L . Howard Willis H . Pequegnat 

Ian Rosman 
V 

TABLE OF CONTENTS 

Page 

1.0 INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 

1.1 PROGRAM OBJECTIVES AND CONTENT OF THIS REPORT . . . . . . . . . . . . . 3 

1.2 PROGRAM DESIGN AND INTEGRATION . . . . . . . . . . . . . . . . . . . . . . . . . S 

1.2.1 PROGRAM SAMPLING DESIGN . . . . . . . . . . . . . . . . . . . . . . . . . . 5 

1.2 .2 FINAL REPORT PLAN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

1.3 C*i TIDE TO AND NATURE OF THIS REPORT . . . . . . . . . . . . . . . . . . . . . . . 19 

2.0 STUDY AREA AND METHODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

2.1 STUDY AREA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 

2.2 CRUISES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 

23 METHODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28 

2.3 .1 FIELD METHODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

2.3 .2 LABORATORY METHODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

23.3 DATA MANAGEMENT AND STATISTICAL ANALYSES . . . . . . . . . . 66 

3.0 PHYSICAL AND CHEMICAL RESULTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 

3.1 HYDROGRAPHY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 

3.1 .1 TEMPERATURE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .79 

3.1 .2 SALINITY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 

3.13 DISSOLVED OXYGEN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 

3.1 .4 NUTRIENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .84 

3.1.5 PARTICULATE ORGANIC CARBON . . . . . . . . . . . . . . . . . . . . . . . 86 

3.1 .6 DISSOLVED ORGANIC CARBON . . . . . . . . . . . . . . . . . . . . . . . . . 86 

3.2 SEDIMENT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .86 

3.2 .1 SEDIMENT CLASSIFICATION . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 

3.2 .2 SEDIMENT HYDROCARBONS . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 

33 CARBON ISOTOPES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .115 

3.3 .1 SEDIMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .115 

3.3 .2 ORGANISMS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119 

3.4 ORGANISM HYDROCARBONS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 

4.0 BIOLOGICAL OCEANOGRAPHY OF THE NORTHERN GULF OF MEXICO 

CONTINENTAL SLOPE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .137 

4.1 MEIOFAUNA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .141 

4.1 .1 COLLECTION RESULTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 

4.1 .2 SEASONAL COMPARISONS . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143 

4.1 .3 COMPARISON OF MEIOFAUNA WITH MACROFAUNA BY DEPTH . . . 143 

4.1 .4 CHANGES IN DENSITY WITH DEPTH . . . . . . . . . . . . . . . . . . . . . . 146 
V1 

Page 

4.1.5 VARIATION IN COMPOSITION WITH DEPTH . . . . . . . . . . . . . . . . . 151 

4.1.6 DISCUSSION OF COMMUNITY ASPECTS . . . . . . . . . . . . . . . . . . . . 158 

4.2 MACROFAUNA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .167 

4.2 .1 INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167 

4.2 .2 COMMUNITY STRUCTURE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168 

4.2 .3 MACROFAUNA ABUNDANCE PATTERNS . . . . . . . . . . . . . . . . . . 170 

4.2 .4 MACROFAUNA DIVERSITY PATTERNS . . . . . . . . . . . . . . . . . . . . 174 

4.2 .5 MAJOR GROUP ACCOUNTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 

43 MEGAFALJNA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .229 

43.1 INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 

4.3.2 COMMUNITY STRUCTURE AND DIVERSITY . . . . . . . . . . . . . . . . 229 

4.3.3 MAJOR INVERTEBRATE GROUP DESCRIPTIONS . . . . . . . . . . . . . 265 

4.3.4 DECAPOD SPECIES ACCOUNTS . . . . . . . . . . . . . . . . . . . . . . . . . 294 

43.5 FISH SPECIES ACCOUNTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336 

5.0 BENTHIC PHOTOGRAPHY RESULTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401 

5.1 LEBENSSPUREN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .401 

5.1 .1 STATION SUMMARIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402 

5.2 FAUNA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .432 

5.2 .1 NUMERICAL ABUNDANCE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433 

5.2 .2 ABUNDANCE BY DEPTH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441 

5.2 .3 ABUNDANCE BY TRANSEC'I' . . . . . . . . . . . . . . . . . . . . . . . . . . . 443 

5.2 .4 DENSITY DIFFERENCES WITHIN TRANSECTS . . . . . . . . . . . . . . . . 444 

5.2.5 SEASONAL DIFFERENCES IN NUMBER OF TAXA . . . . . . . . . . . . . 444 

5.2.6 SIZE MEASUREMENTS FROM BENTHIC PHOTOGRAPHS . . . . . . . . 446 

5.2.7 SELECTED SPECIES ACCOUNTS . . . . . . . . . . . . . . . . . . . . . . . . . 447 

5.2.8 COMPARISONS OF BENTHIC PHOTOGRAPHY AND 

TRAWL DENSITIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 454 

5.3 OUAI_.ITATIVE RESULTS OF CRUISE I PHOTOGRAPHY . . . . . . . . . . . . . . 465 

5.3.1 STATION Cl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 466 

5.3 .2 STATION C2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 466 

5.3 .3 STATION C3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 467 

5.3 .4 STATION C4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 468 

5.3 .5 STATION CS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 470 

6.0 REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .473 

7.0 ATTACHMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 483 
V11 

LIST OF FIGURES 

Figure Page 

1-1 Analysis design diagram for region, season and year comparisons based upon samples 

taken at selected depths, using density as the example response variable . . . . . . . . . . . . . . 7 

1-2 Two contrasting views of the expected faunal changes that might be observed on the 

continental slope of the northern Gulf of Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 

1-3 Analysis design diagram for zonation studies using density as the example response 

variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 

1-4 Analysis design diagram for samples taken in the Eastern Gulf in spring 1985 to 

examine latitudinal versus depth variation in samples taken at selected depths . . . . . . . . . . 12 

1-5 Analysis design diagram for samples taken between the Central and Western 

transects in summer 1985 to examine longitudinal versus depth variation in samples 

taken at selected depths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 

2-1 Stations sampled on each cruise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 

2-2 Box core and configuration of in 5iw subsampling tubes in the box corer . . . . . . . . . . . . . . 33 

2-3 Schematic diagram of photographic system used for benthic photography . . . . . . . . . . . . . 36 

2-4 Sediment hydrocarbons analytical scheme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 

2-5 Hydrocarbon analysis scheme for organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 

2-6 Schematic representation of digitizing apparatus used for processing benthic 

photographs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 

2-7 Proportion of macrofauna organisms in disturbed and undisturbed box core samples . . . . . . 60 

2-8 Example gas chromatograms of possible contaminants collected from sampling vessels . . . . . 67 

2-9 Data flow sequence for Continental Slope Study . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 

3-1 Hydrographic profiles, all cruises combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .80 

3-2 Variation in temperature at selected depths measured at three stations across the 

Gulf during Cruise II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .81 

3-3 Variation in salinity at selected depths measured at three stations across the Gulf 

during Cruise II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .83 

3-4 Variation in dissolved oxygen at selected depths measured at three stations across the 

Gulf during Cruise II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .85 

3-5 Variation in nitrate at selected depths measured at three stations across the Gulf 

during Cruise 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .87 
V111 

Figure Page 

3-6 Variation in phosphate at selected depths measured at three stations across the Gulf 

during Cruise II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .88 

3-7 Variation in particulate organic carbon at selected depths measured at three stations 

across the Gulf during Cruise II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .89 

3-8 Sediment map . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .91 

3-9 Molecular level alkane distributions for sediments from the central transect during 

Cruise I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .95 

3-10 Variation in extractable organic matter and the aliphatic unresolved complex mixture 

along transects in the eastern, western, and central Gulf of Mexico continental slope . . . . . .98 

3-11 Variation in hydrocarbon source parameters along transects in the eastern, western, 

and central Gulf of Mexico continental slope . . . . . . . . . . . . . . . . . . . . . . . . . . . . .99 

3-12 Variation in planktonic, terrestrial, and petroleum hydrocarbons along three transects 

presented as a cumulative concentration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 

3-13 The relationship between alkane petroleum indicators and a terrestrial indicator . . . . . . . . 102 

3-14 The relationship between the aliphatic unresolved complex mixture and the 

planktonic and terrestrial alkane indicator . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 

3-15 Variations in extractable organic matter and the aliphatic unresolved complex 

mixture during three samplings of the central transect . . . . . . . . . . . . . . . . . . . . . . . 105 

3-16 Variations in hydrocarbon source parameters during three samplings of the central 

transect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .106 

3-17 Variation in plankton, terrestrial and petroleum hydrocarbons during three samplings 

of the central transect presented as a cumulative concentration . . . . . . . . . . . . . . . . . . 107 

3-18 Variation in the carbon preference index as a function of depth along the Central 

Transect during Cruise III and a representative fused silica gas chromatogram of the 

aliphatic hydrocarbons from Station C5, Cruise III . . . . . . . . . . . . . . . . . . . . . . . . . 108 

3-19 The relationship between a terrestrial hydrocarbon indicator and the clay content of 

the sediment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 

3-20 The relationship between a terrestrial hydrocarbon indicator and the clay and sand 

content of the sediment during Cruise II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 

3-21 The relationship between two petroleum hydrocarbon indicators and the clay content 

of the sediment during Cruise II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 

3-22 Summary of the carbon isotopic composition of sedimentary organic matter from all 

five cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118 

3-23 Summary of the carbon isotopic composition of organism tissues by organism type . . . . . . . 120 
1X 

Figure Page 

3-24 Summary of the carbon isotopic composition of organism tissues by transect and 

organism type for all five cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 

3-25 Summary of the total alkane concentration in organism tissues for Cruises I, II and III . . . . . 123 

3-26 Selected examples of alkane distributions in fish tissues . . . . . . . . . . . . . . . . . . . . . . 126 

3-27 Selected examples of alkane distributions in crab and shrimp tissues . . . . . . . . . . . . . . . 127 

3-28 Selected examples of alkane distributions in eel tissues . . . . . . . . . . . . . . . . . . . . . . . 128 

3-29 Selected examples of terrestrial hydrocarbons detected in crab and fish tissues . . . . . . . . . 130 

3-30 Selected examples of terrestrial hydrocarbons detected in shrimp and fish tissues . . . . . . . 131 

3-31 Summary of total alkane concentrations from Cruises I, II and III as a function of 

tissue type of all organisms analyzed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133 

3-32 A suggested scheme for the sources and sinks of hydrocarbons on the Gulf of Mexico 

slope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .134 

4-1 Comparison of median numbers of permanent and nauplii meiofauna arranged by 

season collected . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 

4-2 Relative occurrence of meiofauna and macrofauna with depth at all sampling stations . . . . . 145 

4-3 Mean densities of all meiofaunal groups collected at Central Transect stations . . . . . . . . . 147 

4-4 Mean densities of all meiofaunal groups collected at Eastern Transect stations . . . . . . . . . 148 

4-5 Mean densities of all meiofaunal groups collected at Western-Central Transect 

stations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .149 

4-6 Mean densities of all meiofaunal groups collected at Cruise II, Western Transect 

stations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .150 

4-7 Densities and approximate wet weights for the six most abundant meiofaunal groups 

collected during Cruise I, Central Transect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 

4-8 Densities and approximate wet weights for the six most abundant meiofaunal groups 

collected during Cruise II, Western, Central, and Eastern Transects . . . . . . . . . . . . . . . 154 

4-9 Densities and approximate wet weights for the six most abundant meiofaunal groups 

collected during Cruise III, Central Transect . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156 

4-10 Densities and approximate wet weights for the six most abundant meiofaunal groups 

collected during Cruise IV, Eastern Transect . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 

4-11 Densities and approximate wet weights for the six most abundant meiofaunal groups 

collected during Cruise V, Western-Central Transect . . . . . . . . . . . . . . . . . . . . . . . 159 

4-12 Comparison of macrofauna and meiofauna densities for four major infaunal groups . . . . . . 161 
X 

Figure Page 

4-13 Abundance of Nematoda and Harpacticoida plotted against percent sand and clay . . . . . . 164 

4-14 Plot of number of macrofaunal species versus number of individuals showing the great 

number of species that were collected few times compared to the fewer species that 

occurred in large numbers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171 

4-15 Comparative levels of macrofaunal densities by region, season, year and selected 

depth interval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 

4-16 Comparative levels of macrofaunal density by depth on the Central Transect, Fall 1984 . . . . 175 

4-17 Comparative levels of macrofaunal density along selected isobaths in the Eastern Gulf 

of Mexico during Spring 1985 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176 

4-18 Comparative levels of macrofaunal density along isobaths between the Central and 

Western Gulf of Mexico sampling transects, Summer 1985 . . . . . . . . . . . . . . . . . . . . 177 

4-19 Comparative levels of macrofaunal diversity by region, season, year and selected 

depth interval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178 

4-20 Comparative levels of macrofaunal diversity by depth on the Central Transect, Fall 

1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .179 

4-21 Comparative levels of macrofaunal diversity along selected isobaths in the Eastern 

Gulf of Mexico during Spring 1985 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . 181 

4-22 Comparative levels of macrofaunal diversity along isobaths between the Central and 

Western Gulf of Mexico sampling transects, Summer 1985 . . . . . . . . . . . . . . . . . . . . 182 

4-23 Station densities for Porifera at all transects on all five cruises . . . . . . . . . . . . . . . . . . 184 

4-24 Station densities for Nemertea at all transects on all five cruises . . . . . . . . . . . . . . . . . 190 

4-25 Station densities for Nematoda at all transects on all five cruises . . . . . . . . . . . . . . . . . 193 

4-26 Polychaete densities at all stations, all transects, and all five cruises . . . . . . . . . . . . . . . 194 

4-27 Polychaete feeding types showing number of families, taxa, and individuals in each 

feeding type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198 

4-28 Numbers of polychaete species at each station along the Central Transect . . . . . . . . . . . . 201 

4-29 Species diversity and number of species of megafauna invertebrates for the Eastern 

Transect, Cruises I-V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235 

4-30 Species diversity and number of species of megafauna invertebrates for the Central 

Transect, Cruises I-V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236 

4-31 Species diversity and number of species of megafauna invertebrates for the Western 

Transect, Cruises I-V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237 

4-32 Species diversity and number of species of fishes for the Eastern Transect, Cruises I-V . . . . 238 
X1 

Figure Page 

4-33 Species diversity and number of species of fishes for the Central Transect, Cruises I-V . . . . 239 

4-34 Species diversity and number of species of fishes for the Western Transect, Cruises 

I-V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .240 

4-35 The range of depth covered by trawling isobathymetrically as nearly as possible along 

three "horizontal transects" of increasing depth in the Eastern sampling area . . . . . . . . . . 242 

4-36 The range of depth covered at individual stations by trawling isobathymetrically as 

nearly as possible along three "horizontal transects" of increasing depth in the 

West-Central sampling area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243 

4-37 All invertebrates taken by trawl from the 15 stations of Cruise IV on the Eastern 

Transect clustered by NESS at 30 individuals . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 

4-38 All fishes taken by trawl from the 15 stations of Cruise IV on the Eastern Transect 

clustered by Ness at 30 individuals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246 

4-39 All invertebrates taken by trawl from the 12 stations of Cruise V clustered by NESS at 

30 individuals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247 

4-40 All fishes taken by trawl from the 12 stations of Cruise V clustered by NESS at 30 

individuals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248 

4-41 All invertebrates from Cruises III, IV and V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250 

4-42 Numbers of individuals of four common fishes collected from the 15 isobathymetric 

stations of Cruise IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251 

4-43 Results of isobathymetric trawling during Cruise IV on the Eastern Transect . . . . . . . . . . 252 

4-44 Numbers of individuals of two common brachyuran crab species collected from the 15 

isobathymetric stations of Cruise IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253 

4-45 The depth and station distribution patterns of a gorgonian coral and chirostylid 

decapod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .255 

4-46 The belief that the eel preys upon the polychelid is reinforced by the concordance in 

their distribution among 15 stations of Cruise IV on the Eastern Transect . . . . . . . . . . . . 256 

4-47 Species recruitment curve for fish species taken at all stations on the Central Transect . . . . 259 

4-48 Chi-squared values for fish species taken on the Central Transect . . . . . . . . . . . . . . . . 259 

4-49 Species recruitment curve for fish species trawled on the Western Transect . . . . . . . . . . . 260 

4-50 Chi-squared values for fish species taken on the Western Transect . . . . . . . . . . . . . . . . 260 

4-51 Species recruitment curve for fish species trawled on the Eastern Transect . . . . . . . . . . . 261 

4-52 Chi-squared values for fish species taken on the Eastern Transect . . . . . . . . . . . . . . . . 261 
X11 

Figure Page 

4-53 Species recruitment curve for invertebrate species trawled from the Central Transect . . . . . 262 

4-54 Chi-squared values for invertebrate species trawled from the Central Transect . . . . . . . . . 262 

4-55 Species recruitment curve for all invertebrates trawled on the Western Transect . . . . . . . . 263 

4-56 Chi-squared analysis of invertebrate species trawled on the Western Transact . . . . . . . . . 263 

4-57 Species recruitment curve for all invertebrates trawled on the Eastern Transect . . . . . . . . 264 

4-58 Chi-squared analysis of invertebrate species trawled on the Eastern Transect . . . . . . . . . . 264 

4-59 Average number of individuals of selected decapods at collecting stations in the depth 

intervals shown on abscissa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272 

4-60 Average density of three sibling species of the genus enone naeus . . . . . . . . . . . . . . 275 

4-61 Average number of individuals per station of species in the genus Plesionika . . . . . . . . . . 278 

4-62 Average density of four species of the genus Gl,Xphocrangon and of Paranandalus 

willisi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .279 

4-63 Relative abundance of the major groups of decapods . . . . . . . . . . . . . . . . . . . . . . . 286 

4-64 Total individuals collected of the five echinoderm classes . . . . . . . . . . . . . . . . . . . . . 288 

4-65 Locations where B1ax ~hla were collected . . . . . . . . . . . . . . . . . . . . . . . . . . 295 

4-66 Frequency histogram and size-weight relationship for carapace widths of $]~ 

I=hla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .297 

4-67 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace widths of $atwlax ~ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297 

4-68 Frequency histograms for carapace widths of $]ax a from individual cruises . . . . 298 

4-69 Locations where Benthesicvmus barteletti were collected . . . . . . . . . . . . . . . . . . . . . 300 

4-70 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Benthesicymus tti . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300 

4-71 Frequency histogram and size-weight relationship for carapace lengths of 

Benthesic; mus es . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302 

4-72 Frequency histograms for carapace lengths of Benthesicvmus barteletti from 

individual cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302 

4-73 Locations where Benthochascon ti were collected . . . . . . . . . . . . . . . . . . . . . . 304 

4-74 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace widths of Benthochascon ti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304 
X111 

Figure Page 

4-75 Frequency histogram and size-weight relationship for carapace widths of 

Benthochascon gcbmitg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305 

4-76 Frequency histograms for carapace widths of Benthochason schmitti from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .305 

4-77 Locations where G,y,phocrangon a were collected . . . . . . . . . . . . . . . . . . . . . 308 

4-78 Frequency histogram and size-weight relationship for carapace lengths of 

Glvnhocrangon aculeata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308 

4-79 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Jynhocrangon a . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310 

4-80 Frequency histograms for carapace lengths of G ynhocranQOn aculeata from 

individual cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310 

4-81 Locations where Munida valida were collected . . . . . . . . . . . . . . . . . . . . . . . . . . . 312 

4-82 Frequency histogram and size-weight relationship for carapace lengths of Munid.a 

valida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .312 

4-83 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Munida ya]ida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313 

4-84 Frequency histograms for carapace lengths of Munida valida from individual cruises . . . . . 313 

4-85 Locations where Munidopsis robusta were collected . . . . . . . . . . . . . . . . . . . . . . . . 316 

4-86 Frequency histograms and size-weight relationship for carapace lengths of 

Munidopsis robusta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316 

4-87 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Munidopsis robusta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317 

4-88 Frequency histograms for carapace lengths of unidopsis robusta from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .317 

4-89 Locations where Nematocarcinus rotundus were collected . . . . . . . . . . . . . . . . . . . . 319 

4-90 Frequency histograms and size-weight relationship for carapace lengths of 

Nematocarcinus rotundus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319 

4-91 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Nematocarcinus rotundus . . . . . . . . . . . . . . . . . . . . . . . . . . . 321 

4-92 Frequency histograms for carapace lengths of Nematocarcinus rotundus from 

individual cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321 

4-93 Locations where Penaeopsis serrata were collected . . . . . . . . . . . . . . . . . . . . . . . . 323 

4-94 Frequency histogram and size-weight relationship for carapace lengths of Penaeo 

serr3ta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .323 
x iv 

Figure Page 

4-95 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Penaeopsis serrata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324 

4-96 Frequency histograms for carapace lengths of Penaeopsis serrata from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .324 

4-97 Locations where Plesionika holthyjsi were collected . . . . . . . . . . . . . . . . . . . . . . . . 326 

4-98 Frequency histogram and size-weight relationship for carapace lengths of Plesionika 

holthuisi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .326 

4-99 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Plesionika si . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 327 

4-100 Frequency histograms for carapace lengths of Plesionika holthuisj from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .327 

4-101 Locations where Rochinia cL3SSa were collected . . . . . . . . . . . . . . . . . . . . . . . . . . 329 

4-102 Frequency histogram and size-weight relationship for carapace widths of Rochinia 

crassg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .329 

4-103 Frequency histograms for carapace widths of Rochinia &IaSSa from individual cruises . . . . . 330 

4-104 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace widths of Rochinia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 330 

4-105 Locations where Stereomastis scu ,pta were collected . . . . . . . . . . . . . . . . . . . . . . . 333 

4-106 Frequency histogram and size-weight relationship for carapace lengths of 

Stereomastis scu to . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333 

4-107 Median size, 25th and 75th percentiles, and approximate confidence intervals on 

carapace lengths of Stereomastis a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334 

4-108 Frequency histograms for carapace lengths of Stereomastis sculnt a from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .334 

4-109 Locations where Dibranchus atlanticus were collected . . . . . . . . . . . . . . . . . . . . . . . 337 

4-110 Frequency histogram and size-weight relationship for total lengths of Dibranchus 

atlauticus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .338 

4-111 Frequency histograms for total lengths of Dibranchus as from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .340 

4-112 Stomach contents found in Dibranchus us plotted by volume and weight and 

by percent frequency of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341 

4-113 Locations where Nezumia jS were collected . . . . . . . . . . . . . . . . . . . . . . . . . 343 

4-114 Frequency histogram and size-weight relationship for total lengths of Nezumia 

aegua1iS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .344 
XV 

Figure Page 

4-115 Frequency histograms for total lengths of Nezumia aeqya]Lq from individual cruises . . . . . . 345 

4-116 Stomach contents found in Nezumia jq plotted by volume and weight and by 

percent frequency of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347 

4-117 Locations where Bembrons gobioides were collected . . . . . . . . . . . . . . . . . . . . . . . 349 

4-118 Frequency histogram and size-weight relationship for total lengths of Bemrons 

gobioides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .350 

4-119 Frequency histograms for total lengths of Bembrops gobioides from individual cruises . . . . 351 

4-120 Stomach contents found in BembiQgs gobioides plotted by volume and weight and by 

percent frequency of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 353 

4-121 Locations where Coelorinchus caribbaeus were collected . . . . . . . . . . . . . . . . . . . . . 355 

4-122 Frequency histogram and size-weight relationship for total lengths of Coelorinchus 

caribbaeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 356 

4-123 Frequency histograms for total lengths of GoelorinchLS caribbaeus from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .357 

4-124 Stomach contents found in Coelorinchus caribbaeus plotted by volume and weight 

and by percent frequency of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359 

4-125 Locations where Poecilosetta bganj were collected . . . . . . . . . . . . . . . . . . . . . . . . . 361 

4-126 Frequency histogram and size-weight relationship for total lengths of Poecilosetta 

12jDaIIi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .362 

4-127 Frequency histograms for total lengths of Poecilosetta ku~ from individual cruises . . . . . . 363 

4-128 Stomach contents found in Poecilosetta dpi plotted by volume and weight and by 

percent frequency of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365 

4-129 Locations where Gadomus longifilis were collected . . . . . . . . . . . . . . . . . . . . . . . . 367 

4-130 Frequency histogram and size-weight relationship for total lengths of *adomus 

longifilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .368 

4-131 Frequency histograms for total lengths of Gadomus longifilis from individual cruises . . . . . 369 

4-132 Stomach contents found in Gadomus longifilis plotted by volume and weight and by 

percent of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 

4-133 Locations where Chaun ~ pi= were collected . . . . . . . . . . . . . . . . . . . . . . . . . . 373 

4-134 Frequency histogram and size-weight relationship for total lengths of Chaun ~ Pjau . . . . . 374 

4-135 Frequency histograms for total lengths of Chaunax pi= from individual cruises . . . . . . . 375 

4-136 Locations where Coelorinchus coelor ynchus were collected . . . . . . . . . . . . . . . . . . . 377 
XV 1 

Figure Page 

4-137 Frequency histogram and size-weight relationship for total lengths of Coelorinchus 

coelor ynchus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 

4-138 Frequency histograms for total lengths of Coelorinchus coelor ynchus from 

individual cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 

4-139 Locations where Chloropthalmus iA were collected . . . . . . . . . . . . . . . . . . . . . 381 

4-140 Frequency histogram and size-weight relationship for total lengths of Chloronthalmus 

agassiZi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .382 

4-141 Frequency histograms for total lengths of .hloropthalmLS jzj from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .383 

4-142 Stomach contents found in hloropths apj plotted by volume and weight 

and by percent frequency of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384 

4-143 Locations where Synaphobranchus oregoni were collected . . . . . . . . . . . . . . . . . . . . 386 

4-144 Frequency histogram and size-weight relationship for total lengths of 

Synaphobranchus oregoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 387 

4-145 Frequency histograms for total lengths of Synaphobranchus oregoni from individual 

cruises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .388 

4-146 Stomach contents found in Synaphobranchus oregoni plotted by volume and weight 

and by percent frequency of occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 390 

4-147 Locations where Urophvsis cirratus were collected . . . . . . . . . . . . . . . . . . . . . . . . . 392 

4-148 Frequency histogram and size-weight relationship for total lengths of Uronhvsis 

cirratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .393 

4-149 Frequency histograms for total lengths of Uro hnT1 cirratug from individual cruises . . . . . . 394 

4-150 Locations where Peristedion avae were collected . . . . . . . . . . . . . . . . . . . . . . . . . 396 

4-151 Frequency histogram and size-weight relationship for total lengths of Peristedion 

gLayae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .397 

4-152 Frequency histograms for total lengths of Peristedion grayae from individual cruises . . . . . . 398 

5-1 Total lebensspuren densities for Cruise II stations . . . . . . . . . . . . . . . . . . . . . . . . . 410 

5-2 Major lebensspuren category densities by station, Cruise II . . . . . . . . . . . . . . . . . . . . 412 

5-3 Cruise II station densities for separate major lebensspuren categories . . . . . . . . . . . . . . 413 

5-4 Total lebensspuren densities for Cruise III stations . . . . . . . . . . . . . . . . . . . . . . . . 428 

5-5 Total lebensspuren densities for Cruise II and III stations . . . . . . . . . . . . . . . . . . . . . 429 

5-6 Major lebensspuren category densities by station, Cruise III . . . . . . . . . . . . . . . . . . . . 430 
XV11 

Figure Page 

5-7 Cruise III density by depth for separate major lebensspuren categories . . . . . . . . . . . . . 431 

5-8 Total densities of major groups for all stations, Cruises II-V . . . . . . . . . . . . , . . . . . . . 436 

5-9 Total biota density from benthic photography by station, Cruises II-V . . . . . . . . . . . . . . 442 

5-10 Distribution by depth and length frequency distribution of peniagQne sp., Cruises II 

and III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .449 

5-11 Distribution by depth and length frequency distribution of $,embrops gobioides, 

Cruises II-V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451 

5-12 Distribution by depth and length frequency distribution of Chloronthalmus Wasei .i, 

Cruises II-V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452 

5-13 Distribution by depth and length frequency distribution of $yna,phobranchus sp ., 

Cruises II-V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 453 

5-14 Station distribution of density for Munida valida, Cruise IV . . . . . . . . . . . . . . . . . . . . 455 

5-15 Megafauna density comparisons of trawl and benthic photography: Cruise III . . . . . . . . . 459 

5-16 Megafauna density comparisons of trawl and benthic photography: Cruise IV . . . . . . . . . 460 

5-17 Megafauna density comparisons of trawl and benthic photography: Cruise V . . . . . . . . . . 461 

x ix 

LIST OF TABLES 

Table Page 

2-1 Summary of the precision of replicate gravametric analyses of digestion fractionation 

and its comparison to Soxhlet extraction/fractionation . . . . . . . . . . . . . . . . . . . . . . . .61 

2-2 Summary of MMS QC sediment digestion/extraction replicate gas chromatographic 

analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .62 

2-3 Summary of replicate digestion gas chromatographic analyses of QC whitefish tissue . . . . . .63 

2-4 Comparison of the precision of digestion/extraction gravametric analysis between two 

different analysts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .64 

2-5 Summary of accuracy experiments for selected n-alkanes . . . . . . . . . . . . . . . . . . . . . .65 

3-1 The ranges and averages for selected hydrocarbon parameters in Gulf of Mexico 

continental slope sediments . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . .93 

3-2 Summary of Gulf of Mexico sediment hydrocarbon analyses . . . . . . . . . . . . . . . . . . . . .94 

3-3 Suggested molecular-level indicators of specific hydrocarbon sources to sediments on 

the Gulf of Mexico continental slope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .96 

3-4 Variability in hydrocarbon parameters along isobaths-eastern Gulf of Mexico . . . . . . . . . 110 

3-5 Variability in hydrocarbon parameters along isobaths-west/central Gulf of Mexico . . . . . . . 111 

3-6 Comparison of sediment hydrocarbon parameters at two different topographic 

settings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 

3-7 Summary of the incidence of detectable hydrocarbons in organisms muscle tissue . . . . . . . 124 

3-8 Species in which no tissue hydrocarbons were detected in organism in the present 

study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 

3-9 Species in which a sediment hydrocarbon signature was detected in organism tissues . . . . . 129 

4-1 Overall abundance and estimated biomass of the meiofaunal collection . . . . . . . . . . . . . 142 

4-2 Relative abundance of major macrofaunal groups . . . . . . . . . . . . . . . . . . . . . . . . . 169 

4-3 Rank order of dominant macrofaunal species . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172 

4-4 A list of the 50 most abundant polychaetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200 

4-5 Densities (individuals per hectare) of megafaunal invertebrates and demersal and 

benthopelagic fishes at six depth intervals on three major transects . . . . . . . . . . . . . . . . 231 

4-6 Total densities of the most abundant megafaunal invertebrate groups for sampled 

depth intervals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232 

4-7 Densities of the most abundant species of decapoda that account for the density data 

seen in Table 46 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233 
xx 

Table Page 

4-8 Summary of starts and stops of fishes trawled on the isobathymetric series of 15 

stations on the Eastern Transect during Cruise IV . . . . . . . . . . . . . . . . . . . . . . . . . 257 

4-9 Megafaunal arthropods collected by trawl during Cruises I, II, III, IV and V of the 

LGLStudy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269 

4-10 Comparison of the five most abundant penaeid species from the LGL and TerEco 

studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273 

4-11 Comparison of the five most abundant caridean species from the LGL and TerEco 

studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276 

4-12 The most abundant anomurans on the slope of the northern Gulf of Mexico as 

determined by LGL and TerEco projects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 

4-13 The most abundant macruran decapods on the slope of the northern Gulf of Mexico, 

as noted by the LGL and TerEco projects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284 

4-14 The five most abundant brachyuran crabs collected by the LGL and TerEco projects . . . . . 285 

4-15 Total numbers of the five classes of megafaunal echinoderms . . . . . . . . . . . . . . . . . . . 287 

4-16 Abundance of the five most abundant asteroid species collected from the three major 

transects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 

4-17 The six most abundant ophiuroid species collected from the three major transects . . . . . . . 290 

4-18 Total numbers of the five most abundant sea-urchins trawled from the major transects . . . . 291 

4-19 The four most abundant holothuroid species collected from the three major transects . . . . . 292 

4-20 Complete inventory of crinoids collected in the project . . . . . . . . . . . . . . . . . . . . . . 293 

5-1 Total area surveyed by benthic photography, Cruises II-V . . . . . . . . . . . . . . . . . . . . . 401 

5-2 Design table for Cruise II benthic photography lebensspuren . . . . . . . . . . . . . . . . . . . 403 

5-3 Lebensspuren major catagory densities Cruise II stations . . . . . . . . . . . . . . . . . . . . . 404 

5-4 Design table for Cruise III benthic photography lebensspuren . . . . . . . . . . . . . . . . . . 405 

5-5 Lebensspuren major category densities Cruise III stations . . . . . . . . . . . . . . . . . . . . . 406 

5-6 Distance between Central Transect station photo transects between Cruises II and III . . . . . 419 

5-7 Total grouped densities, Cruises II-V . . . . . . . . . . . . . , . . . . . . . . . . . . . , . . . . 435 

5-8 Benthic faunal density, Cruise II Western . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437 

5-9 Benthic faunal density, Cruise III Central . . . . . . . . . . . . . . . . . . . . . , . . . . . . . , 438 

5-10 Benthic faunal density, Cruise IV Eastern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439 
XX1 

Table Page 

5-11 Benthic faunal density, Cruise V West-Central . . . . . . . . . . . . . . . . . . . . . . . . . . . 440 

5-12 Total biota densities for all stations sampled on multiple cruises . . . . . . . . . . . . . . . . . 445 

5-13 Numbers of unique taxa for all stations and transect totals, Cruises II-V . . . . . . . . . . . . . 445 

5-14 Sample of length measurements from benthic photography . . . . . . . . . . . . . . . . . . . . 448 

5-15 Total megafauna density from trawling and benthic photography: Cruise III . . . . . . . . . . 456 

5-16 Total megafauna density from trawling and benthic photography: Cruise IV . . . . . . . . . . 457 

5-17 Total megafauna density from trawling and benthic photography Cruise V . . . . . . . . . . . 458 
1 .0 INTRODUCTION 
1 .0 INTRODUCTION 

3 

Since 1983, LGL Ecological Research Associates, Inc . (LGL) in 

conjunction with Texas ABM University (TAMU) has been conducting a study 

of the continental slope of the northern Gulf of Mexico for the Gulf of 

Mexico Regional Office of the Minerals Management Service (MMS) . The 

overall objective of the program is to develop a basic knowledge of the 

deep Gulf fauna, their environment, and ecological processes in advance of 

extensive petroleum development. 

Prior to initiation of this study, MMS had funded TerEco Corporation 

Inc. to synthesize all available environmental information for the Gulf 

slope. The resulting report (Pequegnat 1983) described and interpreted 

information through 1982, and, is large part, served as the basis for 

formulating the objectives of the present program . The Pequegnat (1983) 

document thus provided the background or baseline data against which the 

information gathered by this study can be compared and evaluated . 

1 .1 PROGRAM OBJECTIVES AND CONTENT OF THIS REPORT 

The specific objectives of the Gulf of Mexico Continental Slope Study 

are: 

(1) To determine the abundance, structure, and distribution of 

animal communities in the deep-sea in the Gulf of Mexico . 

(2) To determine the hydrographie structure of the water 

column and bottom conditions at selected sites within the 

study area . 

(3) To determine and compare sedimentary characteristics at 

selected sites within the study area. 

(4) To relate differences in biological communities to 

hqdrographic, sedimentary, and geographic variables . 

(5) To assess seasonal changes in deep-sea biological 

communities in terms of abundance, structure, animal size, 

and reproductive state . 
4 

(6) To measure present levels of hydrocarbon contamination in 

the deep-sea sediments and selected animals prior to, and 

in anticipation of, petroleum resource development beyond 

the shelf-slope break . 

(7) To assemble together and synthesize appropriate published 

and unpublished data with the results of this study, 

summarizing on a seasonal and spatial basis all 

biological, habitat, and environmental observations and 

parameters . 

(8) To compare the biological and non-biological 

characteristics of the deep Gulf of Mexico with that of 

other temperate and subtropical deep-sea regions . 

(9) To assess the need for, and determine the type of studies 

to be conducted in future program efforts. 

TWO additional objectives were mainly met during the first two years 

of the program : 

(10) To conduct an effective quality assurance and quality 

control program which insures that all data acquired are 

accurate and repeatable within standards normally required 

for each type of observation, measurement, or 

determination. 

(11) To critically review, interpret, and analyze all 

observations and data acquired to redefine as necessary 

the research program in such a way as to avoid or minimize 

redundancy and to optimize the efficiency of all field, 

laboratory, and data management operations for future 

deep-sea studies sponsored by M MS in the Gulf of Mexico. 

Activities during Years 1 and 2 Were dedicated to field sampling and 

laboratory sample analyses. Year 3 Chas been dedicated to finishing the 

sample analyses, and compiling the data in usable and interactive format . 
5 

The acquisition of data for meeting the stated objectives occurred 

over the course of five cruises, all conducted during the first two years 

of the program. Because of the small size and taxonomic complexity of the 

macrofauna, all of contract Years 1-3 (1983-1986), as well as a time 

extension into 1987, were required to complete sample analysis . This 

report represents the first time that all data have been available for 

analysis . 

The primary goals of the Year 3 Annual Report were to (1) provide a 

detailed description of all the field collection, laboratory analyses and 

data management methods that have been used (this volume), and (2) provide 

a comprehensive summary, in hard copy, of all the data that were collected 

and that have been submitted to NODC in specified tape report (Volume III 

Appendices) . In the Year 4 Final Report, emphasis will be placed upon 

statistical analyses, interpretation and reporting of the findings, and 

only summary descriptions of the methods and data detailed herein will be 

included in that report, mainly by reference . 

In this report we present results of some initial exploratory 

analyses, we purposefully provide little in the way of interpretative 

comment. The Year 3 Annual Report has, as an analogy, the results section 

of a standard scientific paper. Following the same analogy, discussion of 

the results in terms of program objectives will be the focus of the Final 

Report for Year 4. 

1 .2 PROGRAM DESIGN AND INTEGRATION 

As discussed above, this report is a descriptive treatise, that may 

appear to the reader to be a curious collection of disjointed results . 

The purpose of this section is to clarify how the results will be 

integrated for the Final Report planned for Year 4 . 

1 .2 .1 PROGRAM SAMPLING DESIGN 

The program sampling plan was structured to first (based upon 

sampling conducted on Cruises I, II, III (in part) and IV (in part) 

compare environmental and biological attributes of the slope, by depth, 
6 

among planning regions (Eastern, Central and Western Gulf, for site maps 

see Panels A, B and C of Fig. 2-1, Section 2) between seasons (fall versus 

spring), and between years (1983-84, 198-85) by season (fall, spring, 

respectively) (Fig. 1-1) . The depth selections were not random or evenly 

spaced down the slope but were rather the approximate mid-points of 

previously-defined (Pequegnat 1983) biological depth assemblages or 

"zones" (Fig. 1-2), namely (1) the Shelf-Slope Transition Zone (150 to 450 

m) ; (2) Horizon A of the Archibenthal Zone (475-740 m) ; (3) Horizon B of 

the Archibenthic Zone (775-950 m) ; (4) the Upper Abyssal Zone (975-2250 

m) ; and (5) the Mesoabyssal Zone, Horizon C (2275-2700 m) . The purpose 

here was not to either prove or disprove the concept of zonation versus a 

continuation of change with depth, but rather to evaluate the predictive 

value of the Pequegnat (1987) zonation scheme . 

The same fine stations on the Central Transact were sampled in fall 

1983 . spring 1984 and fall 1984. On the spring 1984 cruise, the same 

depth intervals sampled on the Central Transact were sampled on both an 

Eastern and Western Gulf transact, and in spring 1985 the stations 

comprising the Eastern Transact were resampled. The design thus allows a 

sequence of specific contrasts, proceeding from comparisons by region, to 

season within region, to year within season to depth patterns within 

region, season and year. Our basic depth comparison strategy will be to 

first contrast the Shelf/Slope Transition station to the deeper slope 

stations. The next basic division will be to contrast variables for 

stations located in depths shallower than 1000 m to those located deeper 

than 1000 m. This depth, or thereabouts, has long been viewed as a mayor 

break in the slope environment as it is here that light from the surface 

can no longer be distinguished and that temperature becomes uniformly cold 

(< 4°C) . The last contrasts will be made between the two depths 

corresponding to Pequegnat's (1983) defined assemblages within the shallow 

(< 1000 m) and deep (> 1000 m) categories. In contrast, according to 

Camay et al. (1983), we should find only (1) a distinct shelf fauna above 

1000 m, (2) a distinct abyssal fauna below 2000 m, and (3) in between, an 

indistinct slope fauna that is partially obscured by immigration from the 

two larger areas shallower and deeper (Fig. 1-2) . 
MEIOFAUNA DENSITY - Number/10 cm2 

REGION - SEASON ~YEAR COMPARISONS 

Station-Mean Depth (m ) 

I _W5 

2 -620 

3 -850 

4 -1400 

5 -2602 

1000191 

u 

0 

500 

v 

z 

Figure 1-l . Analysis design diagram for region, season and year comparisons based upon samples taken at 

selected depths, using density as the example response variable . Each tick mark represents 

the sample locations in time and space . 

v 

l J t J / 2 .r I J , ^" '+w / t .! 4 'J ~'STAT/AV iVLl~i'R 

ArE3'T CENTA.IL E"IST ~AWG/ON 
00 

(Pequeqnat, 1983) ( Carney et al ., 1983) 

0 ~~~~~~~ ~~~~~~ ~y\ \~~~\\ \ \\\~\\ ~\\\\ \\ \\\\\ \\ ~\\\\\ ~~\\\~ \~~ ;

PHOTI~ Z\ONE.~~~~,~~~~~ 

C/ SHELF-SLOPE TRANSITION 

C6 

ARCHIBENTHAL DISTINCT 

;~.; . C2 SHELF HORROR A 

. .. 

,... 

:~ ~ ,;,'°~s . . : .:~ C3 ARCHIBENTHAL 

s~ HORIZON 9 B

1000 

. . 

Cz------- - ---- - - - - ---- 

~ C 9 

~ C4 INDISTINCT 

FAUNA 

UPPER ABYSSAL 

-r) ~t .t ~ r ,-~ . 

r y. r 

2000 . a ..t .~ . . . .n_. . ..r.. .. . . .. . . . _ . .. ..t4 . . _. . .: .i . . . . ::it" .~`.~1-?':!:~x:. :42 . . . . . _n. . . .. . . . :~::1Fi'~~.5"~ . f 

,t :. . ..Y...~r. . .\.:Y.r. .,,T" l

., ~ r7I . . :: .Y," :. A. .. . . .,.. .. .~. .. :; .,. . .. .,< :: ~ : ::~; : ' ;,;:.:. s'. . ~ . r " . . ; Ir ESOABYSSAL 

. .'1 . . . . .~t . . : ~ .e.~ .U . . . .'. 

.. .. . . 

.r. .y~ .. l C5 

" HO I ZON C ISTINCT . .: .. n : . . . , ~ "4"tis ; .~ .a. . .: : .r. . . " .: LOPE 

. f . .'Rs: :..h. " . . ~s . . : MESOABYSSAL . .. ..: .I .1. . . . . HORIZON D 

3000 

0 50 100 150 200 

DISTANCE OFFSHORE (nmi) 

Figure 1-2 . Two contrasting views of the expected faunal changes that might be observed on the continental 

slope of the northern Gulf of Mexico . C1-C12 represent stations sampled in this program. during 

Fall 1985 . 
9 

The sampling design for Cruise III was focused towards evaluating and 

testing selected zonation hypotheses (e .g., Carney et al . 1983 versus 

Pequegnat 1983, Fig . 1-2) . Twelve stations were sampled on the Central 

Transect at depths of 356, 492, 633, 881, 1017, 1191, 128, 1465, 2100, 

2518 and 2915 " A site map is provided in Panel C of Figure 2-1 , Section 

2 ; an analysis design diagram is shown by Figure 1-3 . 

The first contrast will be to test the hypotheses of a real break at 

about 1000 m by comparing data taken at the first four (356 to 881 m) 

depths to data taken at the deeper depths (1017 to 2945 m) . Within the 

shallow group of stations, we can then compare the hypothesized ShelfSlope Transition Zone to the Archibenthal Zone and ; lastly, between the 

two depths within each zone . Given the relatively even spacing among 

these more shallow stations, this sequence of contrasts should delineate 

between the dichotomous views of sharp faunal breaks (zones) versus a more 

conservative view of the nature of changes with depth . 

The next contrast will be to compare variables at depths equating to 

Pequegnat's (1983) Upper Abyssal Zone versus those equating to his 

Mesoabyssal Zone ; and for the latter, between the two hypothesized 

horizons. These should not be different according to the Carney et al . 

(1983) concept (Fig . 1-2) . 

At this point we will be left with the stations between 1000 and 2000 

m. The first step will be to contrast the "'2000-m deep station which, 

although within the Upper Abyssal Zone, was widely separated from the 

other stations (-_ sampled depths) . We will next compare the two shallower 

sites within the zone (881, 1017 m) to the deeper sites (1128, 1465 m), 

and then make comparisons between the individual depths within the two 

sets. The shallower set was more widely spread than the deeper set (see 

Figs . 1-2 and 1-3), thereby providing a test of the hypothesis that 

differences are mainly a function of distance between sites when placed 

along a depth gradient . 

The sampling regimes for Cruises IV (Eastern Gulf) and V (Western 

Gulf) had similar objectives but for different regions. The overall goals 

were to sample along given isobaths to determine the degree of 

longitudinal or latitudinal variation as compared to depth variation, and 
r 

O 

MEIOFAUNA DENSITY - Number/10 cm2 

CENTRAL REGION - FALL "'A" 

Figure 1-3 . Analysis design diagram for zonation studies using density as the example response variable . 

See also Figure 1-2 for relationship of sampling stations to hypothesized faunal zones . 
1I 

to determine some of the sources of the observed variation by means of 

specific contrasts (e.g ., sandy bottom areas versus silt and clay bottoms, 

petroleum seep areas versus non-seep areas, topographic attributes, ete.) . 

In the Eastern Gulf, the selected depths (Fig. 1-4, see also Fig . 2-

1, Panel D, Section 2 for site map) were "350 m (four stations), 625 m 

(six stations), 850 m (five stations), and 2900 m (one station) . Once 

more, the first step will be to compare the Shelf-Slope Transition Zone 

(350-m deep stations) to "true" slope habitats. Next, the 2900-m deep 

station (Mesoabyssal, Horizon D) Will be contrasted to those of the midslope Archibenthal Zone, Horizons A (625 m) and B (855 m) . These 

contrasts will be followed by a comparison of Horizons A and B within the 

Archibenthal Zone. 

Within the Shelf-Slope Transition stations, one (E1A) was located on 

a silty-clay bottom whereas the others (E1B, E1 and E1C) were all sited, 

by design, on sand-silt-clay bottoms. This provides the basis for the 

initial contrast among these stations--the effects of sediment grain size. 

Next, the centrally-located station (E1) will be compared to the flanking 

stations E1B and EM And finally, the two distal stations will be 

contrasted . This sequence of tests will enable an evaluation of the 

effects of distance on observed biological differences along an isobath. 

The placement of stations along the 625-m depth contours were all on 

the same sediment type, but three stations (2B, 2 and 2C) Were tightly 

grouped as a core, with one station (2A) widely spaced from these to the 

northwest and two stations (2D and 2E) separated to the southeast. This 

provides the basis for the first contrast within this group of stations. 

Within the core group of stations, E2C Was farthest removed from the other 

two and data from this station will be contrasted to data for E2 and E2B, 

which mill then be compared to each other. Lastly, for this sequence of 

comparisons, the stations at the opposite ends of the transeet will be 

compared, namely information for station E2A versus E2D and E2E combined, 

and then the latter versus one another . The results of these analyses 

should yield information enabling one to evaluate whether variation along 

an isobath is equivalent to variation observed on the vertical depth 

scale . 
N 

MEIOFAUNA DENSITY - Number/10 cm? 

EASTERN REGION -SPRING 1985 

71000 

N 

O 

500 

r ~m 

z 

. a' 

IA 

A'. 1s~0' 

3A v 

ao X: X. 

" 3C 

ao ''2 . C " I C 

38 PE 

11!°~~ 

*!.'W 94W IAO 'WOO `GIG .?G17 4AW .lGIO 

OEPTilr lmJ 

Figure 1-4. Analysis design diagram for samples taken in the Eastern Gulf in spring 1985 to examine 

latitudinal versus depth variation in samples taken at selected depths . Density is used 

as the example response variable . 
13 

The station array for the 825-m deep stations enables an evaluation 

of the effects of latitude along an isobath. Here, stations E3B (28°02') 

and E3 (28°09" ) were the most southerly, and variables from these areas 

will be contrasted with information from stations EN (28°15'), E3D 

(28°21') and E3A (28°29') treated as a group. Data from E3B and E3 will 

then be contrasted to each other followed by a contrast of data for E3A to 

EN and E3D. Lastly, information for EN and E3D will be contrasted as a 

measure of fine-scale latitudinal differences. 

The isobathic sampling in the Western Gulf was conducted between the 

previously sampled Central and Western Transacts, and stations included 

were given a WC (West-Central) prefix (see Fig . 2-1, Panel D, Section 2 

for site map) . An analysis design diagram is provided by Figure 1-5. Of 

the 12 stations sampled, two (HC1 and WC5) were in the hypothesized ShelfSlope Transition Zone (150-450 m) ; five (WC2, WC4, WC6, WC7 and WC8) were 

in Horizon A of the Arehibenthal Zone (475-740 m) ; three (WC3, WC9 and 

wC10) Were in Horizon B of the Archibenthal Zone (775 to 950 m) ; and two 

(WC11 and WC12) were in Horizon B of the Upper Abyssal Zone (975-2250 m), 

following Pequegnat (1983)" 

Following precedent, the first comparison will contrast the 

hypothesized Shelf-Slope Transition Zone to the stations deeper on the 

slope ; and within the Shelf-Slope Transition Zone, a contrast of WC1 to 

WC5 enables an evaluation of the effects of sediment type (silty-clay 

versus sandy-clay) . As before, we will then proceed to contrast the 

deepest stations (WC11, NC12) to the shallower stations on the slope 

proper (i .e ., not including the transition zone) . The contrast of WC11 

(topographic low surrounded by shallower water) to WC12 (topographic high 

surrounded by deeper water) provides a direct examination of the effects 

of depth her se . 

The neat set of contrasts will be to address whether significant 

differences are apparent between the two hypothesized horizons within the 

Archibenthal Zone. Within Horizon A, a contrast of data for Stations wC6 

and WC7 to the others in this zone (WC2, WC4 and WC8) provides an 

evaluation of areas with suspected petroleum seeps to areas believed to be 

devoid of any known petroleum seeps . The two seep areas x111 then be 
r 

A 

ME IOFAUNA DENSITY - Number/10 cm2 

WESTERN to CENTRAL REGION - SUMMER 1985 

1000 

u 

0 

7500 

. 

~

. ti .. ti.;. S I" 

`i`: ti`: `t~ A00 z 

:" 1Y::ti": "titi 

6t90 

AP 1 

V 

0 

/p00 

7- .c7Gi ~ loor 

93°00' 9P °Olp' 9/°00' 

~as~rvoEFigure 1-5 . Analysis design diagram for samples taken between the Central and Western transects in 

summer 1985 to examine longitudinal versus depth variation in samples taken at selected 

depths . Density is used as the example response variable . 
15 

compared to each other; Station WC2 will be compared to WC4 and WC8; and 

WC4 will be compared to WC8 . The latter contrasts will provide an 

estimate of longitudinal variation within the same depth range and 

sediment type. Within Horizon B, a contrast of WC3 to WC9 and WC10, and 

contrast of WC9 to WC10 will also provide an estimate of longitudi nal 

variation within the same depth range and sediment type . 

1 .2 .2 FINAL REPORT PLAN 

In the Final Report, environmental (hydrography, sediment 

characteristics, hydrocarbon chemistry) and some biological data will be 

integrated to shoal how the slope habitat differs among regions, depths, 

seasons and years. Differences will be determined by inspection as well 

as by more quantitative means such as Principal Components Analyses (PCA) . 

In our project design, we have up to 40 environmental or habitat variables 

that were measured as potential factors affecting biota. PCA enables one 

to transform a large original set of variables into a smaller sat of 

combinations that account for most of the variance of the larger, original 

set. The purpose is thus to explain as much of the total variation in the 

data as possible, with as few of these factors as possible. 

The outputs of PCA enables one to group entities (in our case, 

stations by depth, seasons and year) in terms of their physical/ chemical 

attributes . This, in effect, provides an environmental classification 

against which we can compare results of various biological classifications 

of the same stations. The question being addressed by this approach is : 

Does the distribution and abundance patterns of biota on the slope 

correspond to environmental differences? 

The biological analyses will first focus on each of the mayor 

taxonomic groups associated with soft bottom habitats, namely the 

meiofauna, macrofauna and megafauna, the latter of which were sampled 

using trawls as well as benthic photography. 

The meiofauna section of the Final Report will describe the 

composition of the meiofauna by mayor group and numerical abundance 

patterns by season, region and depth. Differences will be evaluated by 
16 

inspection because of the lack of any detailed taxonomic resolution. The 

abundance of meiofauna will be correlated to physical factors as well as 

to maerofauna abundance and biomass. Historically such contrasts have been 

used to depict overall community structure and a reflection of how this 

structure changes over depth (e .g ., Thiel 1983). Based upon the above 

information, our findings will be compared to other regions, with any 

unique or unusual attributes of the Gulf meiofauna so identified. 

The project meiofauna data set was the most complete of any project 

data set at the time this report was being prepared and the analyses 

presented in Section 4 .1 are reasonably detailed . Nevertheless, the 

results presented herein are preliminary only, and do not represent the 

style and content which will be used in the Year 4 Final Report. 

For each of the maerofauna and megafauna designations, the Final 

Report will first present an overview paper describing the species 

composition of the overall group and relative abundance patterns among 

species. Such descriptions will include species area and/or rarefaction 

curves as an index to assemblage structure. Seasonal and spatial (region 

and depth) patterns of diversity (Shannon-Wiener Index) will be described 

using Analysis of Variance (ANOVA) to detect significant differences and 

the contrast scheme detailed above (orthogonal contrasts) will be used to 

define the nature of the diversity differences by region, season and 

depth . 

A number of diversity indices might be used, each having certain 

attributes and problems. All are influenced at least to some degree by 

sample size and all must be considered representative of the samples 

versus whole assemblages given known problems with sampling efficiencies 

of standard sampling devices (e .g ., trawls and box corers) operated at 

great depths over different types of substrates. We have selected the 

Shannon-Wiener Index because (1) it has been demonstrated to be reasonably 

independent of sample size and, within limits, is normally distributed 

(Bowman et al . 1971) and, most importantly, (2) it has been previously 

used to define species diversity of the Continental Slope collections made 

in the northern Gulf of Mexico (Pequegnat 1983), allowing for direct 

comparisons . 
17 

The next step in the overview sections will be to apply cluster 

analysis techniques to biologically classify stations by region, season 

and depth for comparison, by inspection, to the classification scheme for 

the same stations yielded from PCA applied to physical/ chemical 

attributes. Our cluster analysis approach will follow Grassle and Smith 

(1976) using a Normalized Expected Species Shared (NESS) as the similarity 

measure . 

The findings of the described analyses will be compared to historical 

analyses of similar nature for other regions ; and, in this context, any 

unusual attributes particular to the Gulf of Mexico will be identified . 

Also as part of the overview sections, we will identity the ecologically 

important or numerically dominant component groups within each of the 

macrofaunal and megafaunal designations . These groups kill serve as 

subjects for a series of sections dealing with that group per se. 

The same "com munity-type" analyses described above will be applied to 

the data for each mayor component group of the macro- and megafauna. In 

addition to these, we will also subject the species abundance data for 

depths and longitude to a chi-square analyses following Baekus et al . 

(1965) and Gage (1986), designed to detect apparent faunal boundaries. By 

inspection and/or correlation, the findings will be related to distance 

between sampling sites. 

In these sections of the Final Report, we will identify the most 

abundant species within the component groups and compare abundance 

patterns of these over time and space using ANOVA and orthogonal contrasts 

as defined above. An appropriate transformation will be applied to the 

data, if warranted, prior to the analyses. Likewise, correlation analyses 

will be conducted to determine the apparent associations of species 

abundance to physical/ chemical attributes of the environment using data 

provided by the hydrography and sediment investigations . These 

discussions will also include a description of the present levels of 

hydrocarbons in animal tissues, as provided by the hydrocarbon chemistry 

studies . 
18 

Life-history accounts will be provided for numerically dominant or 

otherwise considered important species of megafauna. These will include 

discussion of food habits, size distribution, apparent growth patterns and 

length-weight relationships . For dominant macrofaunal species groups 

(e .g., the polychaetes) we will attempt to classify the populations by 

feeding type. All of this information is for the purpose of developing an 

overall conceptual model of the Continental Slope ecosystem. Some of the 

species accounts and feeding type information has already been developed 

and is included in this progress report as Attachments. The reader should 

bear in mind, however, that these accounts are not necessarily complete, 

and that no attempt is made herein to either interpret or integrate any of 

the information into a system context. 

The Year 4 Final Report will also contain a section dealing with 

megafauna based upon our benthic photography surveys. While these results 

generally lack the taxonomic resolution required for many analyses, they 

have enabled an evaluation of the overall megafaunal densities estimated 

based upon trailing . Also, at least one very abundant species was 

photographed regularly that was never taken by trawling . These topics 

will be addressed in the Final Report, with the results, along with 

habitat observations, making mayor contribution to the system conceptual 

model . Many of the basic findings from the photography studies are 

presented in this report, but the reader is again forewarned, that little 

interpretation and no integration of the findings is attempted in this 

report . 

As already mentioned above, none of the results of our chemosynthetic 

community investigations are addressed in this report, but a complete 

synthesis of information on these communities will appear in the Final 

Report, including results of project-specific studies . 

The concluding section of the Year 4 Final Report will present a 

conceptual model of the Continental Slope ecosystem of the northern Gulf 

of Mexico, based upon an integration of all the program findings . In this 

section we will define the various types of assemblages that are 
19 

represented and identify the energy sources and flows within and among 

assemblages. We will also identify areas of mayor uncertainties about the 

system and how these might be addressed by future studies. 

1 .3 GUIDE TO AND NATURE OF THIS REPORT 

In the following sections of this report we first describe the study 

area and detail our study methods (Section 2 .0) . We next describe the 

results of the physical/ chemical oceanographic studies on a geographic and 

seasonal basis (Section 3 .0) of this report . It should be noted that the 

hydrocarbon part of Section 3.0 is a reprint of the same material that was 

presented in the final Year 2 Annual Report . It is reprinted (as 

previously approved for the Year 2 Annual Report) only for the sake of 

completeness. In Section 4 .0, we provide our updated biological 

oceanographic results, and in Section 5 .0, the updated results of the 

benthic photography studies are presented . References are provided as 

Section 6 .0 and various attachments are included as Section 7 .0 . An 

Executive Summary is provided under separate cover (Volume I) . Complete 

data listings are also provided under separate cover as the Appendices 

(Volume III) . 

During Year 3, submersible studies were conducted at selected sites 

where ehemosynthetic seep communities were believed to be present. The 

results of these program studies will be provided as a separate report 

which, along with a synthesis of all other available information 

concerning Gulf of Mexico seep communities, will be incorporated in the 

Year 4 Final Report . 

Some final introductory comments are in order here. Various sections 

of this report have been prepared by one or more of 12 individuals 

representing two institutions. Because of the preliminary and progress 

report nature of this volume, a mayor effort has not been given to 

standardization of data presentation, graphics and tables or consistency 

of text among sections. By program design, emphasis to this date has been 

placed upon quality assurance of the sample analyses and data compilation 
20 

as opposed to data analysis, interpretation and the preparation of this 

report . With these observations in mind, we ask the reader's indulgence . 

However, despite the inconsistencies in style and content, we believe the 

discerning reader will find this report informative . 

2.0 STUDY AREA AND METHODS 
23 

2 .0 STUDY AREA AND METHODS 

The Request for Proposal for this study limited the program to waters 

north of 25°N having depths between 200 and 2600 m . It was further 

specified that stations were to be located in depths likely to "delineate 

faunal zonation or areas of transition" and that sampling stations were to 

be located in each of the Minerals Management Service (M MS) Western, 

Eastern, and Central Gulf of Mexico Lease Planning Areas . Guidance was 

also provided to all potential contractors as to the general level of 

research effort being anticipated by MMS, the general categories of 

samples to be collected and the nature of the kinds of laboratory analyses 

which MMS believed appropriate. 

The Statement of Work for the first two years of sampling defined the 

allocation of work as one cruise to the Central Lease Planning Area during 

fall-Winter of 1983, sampling of all three Lease Planning Areas during 

spring-summer of 1984, and intensive sampling of the Central Lease 

Planning Area during fall-winter of 1984. These requirements provide a 

context for the study area and methods descriptions provided below. 

2 .1 STUDY AREA 

As required, the sampling strategy included surveying stations in 

each of the three MMS Gulf of Mexico Lease Planning Areas. The stations 

were initially located on down-slope transacts, although these transacts 

were later expanded laterally by additional stations . The locations of 

the transacts and stations are shown in Figure 2-1, and a summary of the 

rationale for their selection is outlined below. A few characteristics of 

each lease planning area, and the approximate positions of the original 

three transacts that formed the nucleus of the sampling plan are : 

1 . Central Lease Planning Area - This transact extended 

across the slope in the vicinity of the Mississippi 

Trough, from approximately 28020'N, 89°40'W to 26°40'N, 

89020'W . The area is characterized by extremely active 

sediment movement, relatively high terrigenous inputs, an 
24 

Figure 2-l. Stations sampled on each cruise . 
SZ 

(P,IuoD) T-Z aznSz3 
26 

absence of topographic features, and is occasionally 

bathed by the Loop Current. 

2 . Western Lease Planning Area - This transact extended 

across the slope dust south of the Flower Garden Banks, 

Prom 27°25'N, 93°40'W to 25°50 " N, 93030'W. The area is 

characterized by the relatively sluggish circulation of 

the western Gulf, a number of pronounced topographic 

features, moderate to low declivity compared to the 

Mississippi Trough Transact . A fair amount of data is 

available from proximal areas. 

3 . Eastern Lease Planning Area - This transact crossed the 

Florida Escarpment from 2?°k0'N, 85°15'W to 27°30'N, 

85°40'W . The area is characterized by high declivity 

(especially on the lower slope), a low rate of terrigenous 

input and sedimentation, and by moderate to strong 

currents along the face of the slope . 

Station locations within each faunal zone were also influenced by 

water mass distribution and characteristics . The shallowest station in 

each transact was located towards the deeper end of the Shelf/Slope 

Transition Faunal Zone, below the zone of Gulf Common Water in Tropical 

Atlantic Central Water. Going down-slope, the next two stations were 

located in the Antarctic Intermediate Water mass, and the two deepest 

stations were in the Gulf Deep Water. Variation in water mass properties 

would be expected to be minimal at the deeper stations with the exception 

of events related to the passage of cold- and warm-core rings from the 

Loop Current . 

2 .2 CRUISES 

On Cruise I, fine stations at five different depths in the Central 

Gulf were sampled on the Central Transact during fall 1983 (Fig . 2-1A) . 

The primary purpose of Cruise I Was to provide a first look at the study 

area within previously-defined faunal zones over a wide depth range (300 

to 2100 m), and provide data which could be used to refine future sampling 

and modify project hypotheses if necessary. 
27 

Cruise II, conducted during spring/sum mer 1984, extended the 

geographic coverage to the western and eastern regions of the Gulf, and 

re-sampled the stations occupied on Cruise I (Fig . 2-1B) . Based on 

results of Cruise II, comparisons could be made among geographic areas and 

depths, as well as between seasons for stations on the Central Transect . 

During Cruise III, the five original Central Transact stations were 

sampled again in fall 1984 , along with seven additional stations (Fig. 2-

1C) . The seven new stations were located on the Central Transact at 

different depths from the first five stations. The locations for the 

additional stations were mainly in suspected transition areas and were 

based upon the advice of the Scientific Advisory Committee. Sampling the 

five original Central Transact stations allowed annual comparisons between 

the fall cruises of 1983 and 1984. 

During the spring/summer 1985 Cruise IV, 16 stations were sampled 

near the Eastern Transact, including those previously occupied on the 

Eastern Transact on Cruise II (Fig. 2-1D) . The new stations were grouped 

by depth at approximately 350 m (4 stations), 625 m (6 stations), 850 m (5 

stations), and 2900 m (1 station) . The intention was to examine depthrelated differences in the context of observed variability along depth 

contours. Annual variability could also be studied by comparing the data 

from Stations E1, E2, E3, and E5 on Cruise IV to data from Cruise II, when 

the same stations were visited a year earlier. 

Station locations for Cruise V (Fig . 2-1D), also conducted during 

spring/summer 1985, were chosen on the basis of several criteria . The 

area between the Western and Central Transacts was subject to ongoing and 

expected future oil and gas exploration and development activities. Many 

sites were selected along depth contours in order to offer wide geographic 

coverage of the area and to document within-depth variability in sediment 

and biological characteristics. Hydrocarbon seeps had also been reported 

in the area, and the associated biota had not been well documented. Two 

suspected areas of hydrocarbon seeps (Stations WC6 and WC7) were thus 

chosen to compare with probable "control" (non-seep) areas at comparable 

depths (Stations WC8 and WC2). 

The sampling strategy described above permitted project scientists to 

make the following basic contrasts : 
28 

Contrast 

Seasonal variation by depth 

Geographic variation by depth 

Annual variation by depth 

Zonation patterns 

Variation within depths, 

Western Gulf 

Variation within depths, 

Eastern Gulf 

Data Source 

Central Transact, Cruises I & II 

Western, Central, and Eastern 

Transects, Cruise II 

Central Transact, Cruises I & III 

Central Transact, Cruise III, plus 

data from all other cruises 

West-Central Transact, Cruise V 

Eastern Transact, Cruise IV 

2 .3 METHODS 

2 .3 .1 FIELD METHODS 

2 .3 .1 .1 Hydrographic Measurements 

Continuous and discrete measurements of hydrographic parameters were 

obtained throughout the water column (surface to bottom) at five stations 

on each cruise. A Neil-Brown Mark III CTD/Rosette/Transmissometer System 

was used to obtain continuous data and discrete water samples. Continuous 

conductivity (salinity), temperature, depth, and transmission records were 

provided by the Neil-Brown CTD. A 12-bottle rosette attached to the CTD 

was used to collect at least 12 discrete water samples at each station . 

Bottles were spaced throughout the water column in order to delineate the 

mayor mater masses at each site. The CTD/Rosette/ Transmissometer system 

was deployed with a pinger so that the cast could be safely lowered to 

within a few meters of the bottom . This was done in order to discern 

whether there were bottom nepheloid layers at each site. 
29 

Continuous Measurements 

The shipboard Neil-Brown CTD system consisted of a demodulator, 

digital display and digital-to-analog converters. Digital output of each 

scan (every 32 ms) was transmitted via armored cable to the shipboard unit 

in "TELETYPE" format using frequency-shift-keyed modulation designed to 

transmit up to 127 bytes (8 bits) per scan. 

The housing with the conductivity, temperature, and pressure sensors 

was pressure tested to 10,000 psi for one hour . The range, accuracy and 

resolution of the individual sensors were: 

SENSOR TYPE RANGE ACCURACY RESOLUTION 

Pressure 0-320 decibar 

0-650 decibar 

0-1600 decibar 

0-3200 decibar 

0-6500 decibar 

Temperature -32 to +32°C 

Conductivity 1 to 65 mmhos 

0 .1% of FS 

(standard) 

0 .5% 

(optional) 

0 .005°C 

(-3 to +32°C) 

0 .005 mmhos 

0 .0015x FS 

(all ranges) 

0 .0005°C 

0 .001 mmhos 

Transmission profiles were provided by a Sea Tech Inc . 

transmissometer interfaced to the CTD system. This instrument has a 25 em 

light path with a light emitting diode with a wavelength of 660 nm as a 

light source . This instrument, with proper calibration, provides data 

with an error less than 0 .5% transmission . It has a depth capability of 

ca . 2500 m. 

The data from the CTD/transmissometer were stored both as a hard copy 

from an x-Y recorder and on magnetic tape. An HP-1000 computer was used 

aboard the R/Y re for data storage . 
30 

Discrete Measurements 

Discrete measurements of temperature, salinity, dissolved oxygen, 

nutrients, and particulate organic carbon (POC) were obtained by 

collecting samples from PVC Niskin bottles mounted on a General Oceanics 

Rosette sampler. Subsamples for dissolved oxygen were drawn first. All 

the discrete measurements were performed at sea. Measurements of 

temperature, salinity, and POC Were all performed in duplicate . Ten 

percent of the oxygen and nutrient samples were also duplicated to 

establish sampling and analytical precision, and to assure data 

reliability . 

Thermometry. Deep-sea reversing thermometers were attached to Niskin 

bottles mounted on the Rosette. These precisely calibrated thermometers 

were from Texas ABM University's Department of Oceanography collection. 

Most have long histories of calibration to ± 0 .005°C. The thermometers 

were equilibrated at depth for at least 5 min before tripping . All 

thermometers were equilibrated read in duplicate by separate observers. 

Salinity. Samples for salinity were collected in 500-ml airtight 

citrate bottles that were rinsed three times with sample water before 

collection. Samples taken from salinity were analyzed using either a 

Plessey Environmental Systems Model 6230N Laboratory Salinometer or a 

Guildline Model 8400 Autosal Laboratory Salinometer. The Plessey system 

utilized an inductively-coupled conductivity sensor to establish a 

conductivity ratio between an unknown sample and a standard at 

approximately 35 ppt salinity. A dual-element platinum thermometer and 

its associated circuitry sensed the temperature of the sample and applied 

the appropriate compensation. The Plessey system had a range of 0 to 551 

ppt, with an accuracy of ± 0 .003 ppt, and temperature compensation of ± 

0.0007 ppt/°C. The Autosal system measured conductivity directly and had 

better accuracy and precision than did the Plessey system . 
31 

en. Oxygen samples were always the first samples drawn from a 

cast, and were drawn as soon as possible. The samples were taken using a 

length of Tygon tubing with the tip of the tube near the bottom on the 

flask so that it could be filled slowly without agitation. The flask was 

rinsed and air bubbles removed from the tubing with a small amount of 

sample before the flask was filled . The flask was overflowed one full 

volume and the stopper inserted to avoid trapping air bubbles . 

The technique used for analysis of oxygen was the modified Winkler 

technique of Carpenter (1965) . Oxygen samples were analyzed at sea. At 

least 10% of the oxygen samples were taken and analyzed in duplicate . A 

summary of the technique follows: 

As soon as possible after collection, the samples are 

"pickled" by the addition of a divalent manganese solution, 

followed by strong alkali . The precipitated manganous 

hydroxide is dispersed evenly throughout the seawater sample, 

which completely fills a stoppered oxygen flask. Any dissolved 

oxygen rapidly oxidizes an equivalent amount of divalent 

manganese to basic hydroxides of higher valeney states. When 

the solution is acidified in the presence of iodide, the 

oxidized manganese again reverts to the divalent state, and an 

amount of iodine directly proportional to the original 

dissolved oxygen content of the water is liberated. The iodine 

is titrated with standardized sodium thiosulfate per Strickland 

and Parsons (1972) . 

Nutrients. Water samples for nutrient analysis (phosphate, nitrate, 

nitrite, silicate) Were drawn into "Whirl-Pak" sampling bags. If samples 

were not analyzed immediately, they were frozen until analysis later 

during the cruise. Generally, analyses were performed immediately in the 

field following the methods outlined by Strickland and Parsons (1972) . 

Specific methods for each of the nutrients are also given by Technicon 

Instruments Corporation of Andsley, New York, Industrial Methods Bulletins 

100-70W, 98-70W, 161-71 aB, and 155-71W. 
32 

2 .3 .1 .2 Box Core Sampling 

Box core samples were taken at each station to obtain material for 

macroinfauna and meiofauna identification; sediment grain size; carbonate; 

total organic carbon; carbon isotopes ; and hydrocarbons . Six replicate 

samples were taken at each station, except on the Western and Eastern 

Transect stations during Cruise II, when only three replicates were taken 

per station. The replicates were then subdivided to provide material for 

the various types of analyses . 

Box corers were deployed in yoked pairs, using a TAMU-modified 

version of the Gray-0'Aara modification of the J&0 box corer. Under ideal 

conditions, only one cast was required to collect two replicates. 

The box corer (Fig . 2-2) measured 24 .5 x 24 .5 x 44 cm . It was fitted 

with a hinged door to prevent washout of samples, and had up to 135 kg of 

ballast. The door was open until the device had penetrated the substrate, 

whereupon the haws and the door closed . The amount of ballast was 

adjusted to ensure adequate substrate penetration. 

The box corer contained six metal coring tubes, 43 .5 cm long and 3 .5 

cm in internal diameter. During Cruise I, these tubes were mounted in 

three pairs on a wire rack in the center of the box . This design was 

improved on successive cruises by mounting all six tubes against one wall 

of the box and securing them behind an aluminum partition that extended 

the full depth of the box. As each box corer came onboard, the overlying 

water was carefully siphoned into the macrofauna container, and the 

remaining subsamples processed according to their intended uses. 

Sediments for Physical and Chemical Analyses 

Undisturbed, uncontaminated sediment samples for analysis of 

hydrocarbons, grain size, carbonate, and total organic carbon were 

subsampled from the box core immediately after decantation of overlying 

water. The subsample for hydrocarbon, carbonate and TOC analyses were 

stored frozen in a glass far, while the samples for grain size analysis 

were placed in whirl-pak bags and refrigerated . All samples were 

appropriately labeled . 
Box Core Configuration -Cruise I 

: : :0 

Meiofeune and Sediment Sample Core3 

( Area-9.6 cm2 / core) 

Macrotauna Sample 

(I1rra=0 .0569 m2) 

Box Core Configuration - Cruises II- Y 

00000e Meiofeune end Sediment Sample Core*' 

( Area-9 .6 cm2 / core ) 

Mscrofauna Sample 

(Area=0.0475 m2) 

Figure 2-2 . Box core and configuration of in situ subsampling tubes in the box corer . 

w 

w 
34 

Meiofauna 

Four of the tubes were used for meiofauna samples . The top five 

centimeters of sediment in the meiofaunal tubes was extruded using a 

plunger and placed directly into a glass or plastic sample far. An 

isotonic solution of magnesium sulfate was immediately added to narcotize 

meiofaunal organisms . After the sample had been in a cool place out of 

the sun for about 30 min, it was preserved by adding neutral buffered 

formalin with rose bengal until the contents reached a concentration of 5%-

formalin. The far was gently shaken to achieve a uniform mixture of the 

preservative. Preserved samples were stored at ambient temperature. 

Macrofauna 

The remainder of the box core contents was removed to a depth of 20 

em and washed carefully on a 300 micrometer sieve with a gentle stream of 

cartridge filtered sea water. Material retained on the screen was placed 

in suitable containers, labeled, and narcotized in an isotonic magnesium 

sulfate solution. Following narcotization, organisms were preserved in a 

solution of 10% neutral buffered formalin in sea water, to which rosebengal stain was added. These samples were stored in a cool place as soon 

as possible following collection. 

2 .3 .1 .3 Trawl Sampling (MeAafauna ) 

Megafauna sampling was performed with a standard 9-m, semi-balloon 

otter trawl with 60-em steel doors, 3 .8-em stretch mesh, and 1 .3-cm cod 

end mesh. Target trawling times were one hour at stations shallower than 

1300 m, and two or more hours at deeper stations . The amount of time on 

the bottom was arbitrarily measured as the time from winch brake 

application until the winch was started again for trawl retrieval. At a 

towing speed of one to three knots, a ratio of 3 .5 :1 between amount of 

wire out and the depth produced good samples. 

The contents of each retrieved trawl was dumped into metal tubs . 

Fishes and invertebrates intended for hydrocarbon assays were quickly 

removed, photographed, and frozen. The remaining organisms were usually 
35 

rough-sorted into three categories (fish, decapods, and "other") . They 

were then narcotized with isotonic magnesium sulfate if necessary, and 

preserved in 10% neutral buffered Pormalin in sea water . 

2 .3 .1 .4 Benthie Photography 

Benthic photography samples were obtained with the use of a 8enthos 

Model 372 deep-sea camera fitted with a 28-mm lens (angle of view 35° x 

48 .5°), and equipped with a 200 watt-second Benthos strobe . On each visit 

to every station, the camera took 800 exposures of Kodak Ektaehrome 

Professional 5936 film, ISO 200 . Photographs were taken every eight 

seconds. A summary of the number of photos taken per transact summed over 

all cruises follows : 

CRUISFJ TRANSECT N0. OF STATIONS N0 . OF PHOTOGRAPHS 

1/Central 5 4,000 

II/Western 5 4,000 

II/Central 5 4,000 

II/Eastern 5 4,000 

III/Central 12 9,600 

IV/Eastern 16 12,800 

Y/West Central 12 9,600 

Total : 60 48,000 

The photographic gear was mounted inside a protective framework 

similar to that of hoods Hole Oceanographic Institution's Mini-Angus (Fig. 

2-3) . The frame and its mounted components are called "BUCS" (Benthie 

Underwater Camera stem) for convenience in this report. BUCS had a clock 

and altimeter that recorded the time and altitude above the bottom in the 

corner of each photograph. The altimeter had a resolution of ± 0 .1 m. 
36 

Pinger\ 

41 -- Camera 

45° 

Strobe 

' ~ 

-- 

Altimeter 

--~ Altitude 

Frame Length 1 .85 m 2 m 

Weight i n Air 240 kg Optimum 

Photograph Di mensions 

1_3X 1 .8m=2.33qm 

at 2 m altitude 

Figure 2-3 . Schematic diagram of photographic system used for benthic 

photography . 
37 

BIICS Was suspended from the vessel by a hydro wire, and allowed to 

drift near the bottom along transacts 1500 to 5000 m long. Suspending 

BIICS in this way prevented skipping and bouncing on the bottom, thereby 

minimizing disturbance and reducing the chances of attracting or 

frightening animals away. 

The plane of view was parallel to the bottom . Altitude was 

maintained by adjusting the vessel's winch in response to an acoustic 

signal transmitted by a 12 kHz bottom-finding pinger on BUCS. The signal 

was portrayed continuously on a strip chart recorder. Optimum camera 

altitude was approximately 2 m above the bottom, which produced shots that 

included 2 .27 m2 of the bottom. However, acceptable shots were obtained at 

altitudes from 0 .7 m (0 .27 m2 area) to 4 .0 m (9.09 m2 area) . At higher 

altitudes, larger areas were included in each shot, but image resolution 

was poorer; the converse was true at lower altitudes. 

2 .3 .1 .5 Quality Assurance and Quality Control (QA/QC) 

Box Core Sampling 

A concerted effort was made to obtain the highest quality samples . 

This effort included 1) using experienced and competent field biologists 

for sampling; 2) employing discrete criteria for quantifying the quality 

of each sample before it was accepted for on-board processing; 3) closely 

supervising sample handling, washing, narcotizing, and preliminary 

sorting; and 4) maintaining a sample tracking procedure to ensure proper 

chain-of-custody procedures. 

In the field, the following criteria were used to fudge whether a box 

core was acceptable: 

1 . The doors should be fully and properly closed ; 

2 . The sediment in the box should be covered with clear 

water, because cloudy water suggests mixing of water and 

sediment during retrieval ; and 

3 . The sediment should be level within the box, because 

sloping sediment suggests non-vertical sediment 

penetration . 
38 

Coring continued at each station until the requisite number of replicates 

met these criteria. For further information, please refer to Section 

2 .3 .2 .5 Quality Assurance and Quality Control (Laboratory) . 

Benthic Photography 

All BOCS equipment was fully tested prior to each cruise . Fresh 

batteries were installed to poker the 12 kHz bottom-finding pinger and 

camera data chamber. Nickel-cadmium batteries used in the high-intensity 

flash were drained and recharged to full capacity before every cruise, in 

order to prevent premature poker loss during camera transacts . 

After each photographic transact was completed, a test strip of 

exposed film was developed while on station to confirm that no mechanical 

difficulties affecting the photography had occurred. The test strip was 

taken from the end of the 800-exposure roll, to determine whether or not 

the camera had exposed frames from the beginning to the end of the roll. 

If the test strip included an excessive number of poor shots, or if the 

photography had to be aborted while underway, the photographic transact 

was repeated at that station. That proved necessary on only two 

occasions, out of 60 transacts completed. 

2 .3 .2 LABORATORY METHODS 

Laboratory activities included both physical/ahem ical and hydrocarbon 

determinations for sediments and biota ; carbon isotope analyses for 

sediments and biota; sorting, identifying, enumerating and weighing, and 

measuring biota as well as analyzing their gut contents ; and the analyses 

of photographs for biota and lebensspuren. 

2 .3 .2 .1 Sediment Samples from Box Cores 

Grain Size 

Sediment grain size followed the laboratory procedure of Folk (1974) . 

Samples were homogenized and treated with an aliquot of 30% hydrogen 

peroxide (H202) to oxidize organic matter, than washed with distilled 
39 

Water to remove soluble salts . Sodium hexametaphosphate was added to 

deflocculate each sample . The samples were then wet-sieved using a 62.5 

micron (4 .0 0) sieve to separate the gravel and sand from the silt-clay 

fraction. 

The total gravel and sand fraction was oven dried (40°C) weighed, and 

sieved at half-phi intervals (-1 .5, -1 .0, -0 .5, 0.0, 0 .5, 1 .0, 1 .5, 2.0, 

2.5, 3"0, 3 .5, and 4.0) . Each collected fraction was examined for 

aggregates, disaggregated if necessary, and reweighed by fraction to three 

significant figures. 

The silt-clay fraction was analyzed for particle size distribution by 

the pipette (settling rate) method at 4 .5, 5 .0, 5 .5, 6 .0, 7 .0, 8.0, 9 .0, 

and 10.0 phi intervals. 

Organic Carbon 

Organic carbon determinations were made using a Leco WR-12 Total 

Carbon System . Sediment subsamples (0 .2-0 .5 g) Were weighed into 

disposable 5 ml polystyrene beakers and treated with concentrated HC1 to 

remove inorganic carbon (carbonate) . Acid was added dropWise until no 

degassing was observed. The treated samples were then dried at 50°C in a 

recirculating oven for 24-36 hours to remove excess acid and moisture . 

After drying, the sample was quantitatively transferred to a sintered 

crucible. Iron accelerator and tin coated copper catalyst were added and 

analyzed by total combustion on the Leco instrument. Organic carbon was 

converted to C02 and analyzed with a non-dispersive infrared 

spectrophotometer. Blanks and standards mere run on a daily basis. All 

samples were analyzed in duplicate and averaged . Periodically samples 

were combusted at >800°C in a high vacuum, Craig-type combustion system as 

a check on the combustion efficiency of the Leco system. 

Carbonate Carbon 

Carbonate carbon was determined for the same freeze-dried, 

homogenized sediment samples that were used for organic carbon and 

hydrocarbon determinations. Carbonate carbon in Cruise I samples was 

determined by difference between total carbon and carbonate-free (organic) 
40 

carbon, using the Leco WR-12 Total Carbon System. For samples from Cruise 

II, carbonate carbon was determined directly by acidification in a carrier 

stream, followed by infrared detection. 

Carbon Isotope Analyses 

Carbon isotope analyses were performed on sediments and selected 

organisms to determine their food source . Stable carbon isotopes have 

been shown to be useful in delineating the flow of carbon through 

ecosystems since there is considerable evidence for minimal carbon 

isotopic fractionation along marine food chains (Parker 1964, Degens et 

al. 1968, DeNiro and Epstein 1978) . Plants preferentially assimilate 12C 

over 13C during photosynthesis, and the degree of 13C fractionation in 

plants is dependent on the biochemical pathway used for carbon fixation. 

Photosynthetically derived carbon from marine algae generally have carbon 

isotopic values ranging from -19 to -21 ppt . Carbon from terrestrial 

sources is generally at least 7 ppt lighter (more negative) due to the 

uptake of C02 as opposed to bicarbonate in the sea. However, there are 

other pathways that can contribute to variations in the organic carbon 

isotopic content of terrestrial plants. 

Organisms that feed on photosynthetically derived carbon from marine 

algae have carbon isotopic values in the same range as the plankton (-19 

to -21 ppt) . However, tissue from mussels recovered at the Pacific vents 

have delta 13C values near -33 ppt (Rau 1981, Rau and Hedges 1979, 

Williams et al. 1981) . The vent communities of the Pacific are based on 

chemautotrophic bacteria that gain energy from the oxidation of hydrogen 

sulfide (Earl et al. 1980) . In turn, the associated filter feeding 

organisms feeding on these isotopicallp light bacteria have similar 

isotopic values. 

Stable carbon isotopes (delta 13C values) were determined on freezedried sediment organic carbon and tissue samples. The stable carbon 

isotopic C02 composition derived from combustion of the organic matter was 

determined on a Nuclide Corporation six inch, 60° sector, isotope ratio 

mass spectrometer. The carbon isotope values are reported as per mil 

deviations from the Pee Dee Belemnite (PDB) standard : 
41 

delta 13C . L(13C/12C) sample - (13C/12C) std/(13C/12C) std] x 1000 

High !Molecular Weight Hydrocarbons 

This study involved the measurement of high molecular weight (HMW) 

hydrocarbons in megaPauna (fishes and invertebrates) and sediments in 

samples collected on the Gulf of Mexico slope . Sediment samples were 

screened for aromatic hydrocarbon contamination using total scanning 

fluorescence, but primary detection and quantification of petroleum 

contamination was based on high resolution capillary gas chromatography 

and GC/MS/DS analysis . The purpose of the HWM hydrocarbon analyses were: 

1 . To determine the suite of HMW hydrocarbons present and 

their concentrations; 

2 . To determine probable sources of the HMW hydrocarbons as 

either thermogenic (from natural seepage or anthropogenic 

sources) or biogenic ; 

3 . To determine the relationship between HMW hydrocarbons and 

trophie levels ; 

4 . To establish the extent of contamination with respect to 

distance from shore and/or offshore oil/gas production; 

5 . To determine the relationship, if any, between hydrocarbon 

chemistry, water depth, mayor current systems, and 

sediment physical characteristics; and 

6 . To compare the findings to known values for shallow water 

habitats in the Gulf of Mexico and subtropical U .S. Atlantic 

waters . 

Protocols for analyzing both the sediment and benthic organisms were 

very similar (Figs . 2-4 and 2-5)" To avoid repetition, HMW hydrocarbon 

methods for sediments are described below in detail, noting the 

differences in analytical techniques between methods for sediments and 

organisms . 

Sediment Hydrocarbon Analyses. A three-tiered sediment hydrocarbon 

analysis program was implemented. First, extracts of surficial sediments 
42 

SEDIMENTS 

FROZEN SAMPLE 

FREEZE DRYING 

DRY WEIGHT 

KOH/ETHANOL SAPONIFICATION 

SEPARATORY FUNNEL EXTRACTION 

SULFUR? 

rEs 

NO 

SOLVENT REMOVAL ON 

ROTARY EXAPORATOR 

REMOVAL ON COPPER 

COLUMN CHROMATOGRAPHY 

HEXANE BENZENE 

SATURATE FRACTION I I AROMATIC FRACTION 

CAHN WT DETERMINATION CAHN WT DETERMINATION 

GAS CHROMATOGRAPHY 

RESOLVED PEAKS u UNSOLVED COMPLEX 

I MIXTURE 

IGC INTEGRATIONI 

KOVAT ASSIGNMENT C~.T .CULATE 

QUANTIFICATION 

i= 

RATIO INFORMATION 

DATA ANALYSIS ANA SYNTHESIS 

Figure 2-4. Sediment hydrocarbons analytical scheme . 
43 

ORGANISMS 

HYDROCARBON LABORATORY PROCESSING 

THAWED SAMPLE 

POOLING OF ORGANISMS 

AND HOMOGENIZATION 

FREEZE DFiYING 

DRY WEIGHT 

KOH/ETHANOL DIGESTION 

AND ( " H10) SAPONIFICATION 

SEPAfiATORY FUNNEL EXTRACTION 

SOLVENT REMOVAL ON 

ROTARY EVAPORATOR 

NON-SAPONIFABIE LIPID 

COLUMN CHROMATOGRAPHY 

HEXANE BENZENE 

SATURATE FRACTION AROMATIC FRACTION 

CAMN WT DE TERMINA fION CAHN WT DETERMINATION 

-t t 

GAS CHROMATOGRAPHY 

RESOLVED PEAKS UNSOLVED COMPLEX 

MIXTURE 

GC INTEGRATION 

KOVAT ASSIGNMENT CALCU~AT 

QUANTIFICATION 

RATIO INFORMATION 

DATA ANALYSIS AND SYNTHESES 

Figure 2-5 . Hydrocarbon analysis scheme for organisms, . 
44 

from each box core were analyzed by total scanning fluorescence to 

determine the presence or absence of aromatic hydrocarbons. Second, 

detailed saturate and aromatic capillary gas chromatography was performed 

on individual and/or pooled samples from each station. Third, selected 

samples were analyzed by gas chromatography/ mass spectrometry (GC/ MS) For 

compound identification and structure confirmation. 

Sediment samples were obtained from box cores in a clean environment 

on board ship . A clean steel tube was used to subsample the upper 8-cm of 

the sediment. Samples were stored in bars that were solvent-washed and 

combusted at 450°C. The bars were sealed with tef lon-lined caps, labeled, 

and stored frozen. 

The establishment and maintenance of adequate procedural blanks is 

imperative in trace level hydrocarbon analysis. A quality control and 

quality assurance program was strictly followed . Preeleaning of all 

equipment included extensive washings with Micro cleaning solution and 

rinsing with distilled water, acetone, and methylene chloride . All 

solvents were triple glass-distilled, nanograde purity (Burdick and 

Jackson, Inc .) or its equivalent . Final rinses were evaluated by 

gravimetry, gas chromatography and gas chromatography/ mass spectrometry. 

Large volumes (one liter) of solvent were routinely evaporated and tested 

in a similar manner. When possible all equipment (i.e., glassware) was 

combusted at X50°C overnight, after the cleaning procedure is completed. 

Blanks were maintained at negligible levels for all parameters monitored. 

Total scanning fluorescence--determination of trace levels of many 

organic species was possible using fluorescent measurements. Fluorometry 

is inherently selective, and generally at least ten times more sensitive 

than other absorption methods. Fluorescence methods are particularly 

useful for the detection and measurement of complex organic compounds 

containing one or more aromatic functional groups. All oils contain 

significant amounts of aromatics with one to four (or more) aromatic rings 

and their alkqlated analogs. The aromatic composition of an oil provides 

a distinctive "fingerprint", and this fingerprint when used in conjunction 

with other analyses can provide significant typing information. 

Fixed- wavelength and synchronous scanning fluorescence suffer from 

non-selectivity, and are generally ineffective in structural elucidation 
45 

information for the neat and more detailed characterization of the 

polynuclear aromatic hydrocarbons. 

Fixed-Wavelength fluorescence measurements were used to select a 

sample dilution where quenching was insignificant. Next, total scanning 

fluorescence measurements were performed on each sample, using a PerkinElmer Model 3500 or 3600 Data Station equipped with dual 5-1/4" floppy 

disk drives, controlling an automated 650-u0 W-spectrofluorometer. Three 

complete systems were in operation. The extracts were scanned to acquire 

data on three-dimensional fluorescence spectra of emission wavelength, 

excitation wavelength, and intensity. Using custom software, data 

acquisiton time was shortened to 30 min, with a wavelength resolution of 5 

nm for the emission spectrum. 

The emission spectrum was obtained by scanning the emission 

monoehromator at a fixed excitation wavelength. The excitation spectrum 

was acquired by monitoring a fixed emission wavelength while scanning to 

excitation monochromator . The total fluorescence excitation-emission 

wavelength array was filled for each sample by sequential stepping of the 

excitation monochromator over the wavelength rang of interest, and 

scanning with the emission monochromator . 

The data were stored in active memory. Standard wavelength intervals 

were 200-600 nm for both excitation and emission . Intensity data were 

acquired during each emission scan at preset wavelength intervals . 

Programming flexibility permitted acquisition speeds of from 60 to 450 

nm/min on each monoehromator. Up to 2500 discrete intensity values were 

acquired per spectrum, with a 1 nm resolution at slower scan speeds . 

After acquisition, the data were permanently stored on 5-1/4" diskette. 

The data were then transformed via a three-dimensional conversion routine, 

and a spectral plot generated with a standard graphics package on a CRT. 

The output was hard-copied on a graphics printer/plotter. 

Gas chromatography/mass spectrometry--Sediment samples for fused 

silica capillary chromatography were freeze-dried before extraction to 

minimize sale entrainment into the extracts. Sample handling was kept to 

a minimum to avoid contamination of the samples. After freeze-drying, a 

sediment dry weight was obtained. The freeze-dried sample (50 g dry 

weight) was placed in a round bottom flask (500 m1), with standard taper 
46 

(particularly for mixtures) . Despite the ability to select both the 

excitation and emission wavelengths, the conventional luminescence methods 

have limited applicability since most spectra of complex mixtures often 

cannot be satisfactorily resolved . To overcome these problems, Texas A&M 

University developed a software package and methodology for threedimensional total scanning (excitation-emission spectra) of fluorescence. 

The method is in routine use in the AbM laboratory, as well as those of a 

number of oil companies to analyze the extracts of surficial sediments, 

source rocks, and oils . This fluorescent method can differentiate gasprone provinces with condensate aromatic signatures from oil-prone areas 

with oil signatures, and has been used by Brooks et al. (1983), Rennicutt 

et al . (1983), and Rennicutt and Brooks (1983) to estimate and type 

hydrocarbon concentrations in sediments and oils. 

Approximately 20 g of sediment was required for total scanning 

fluorescence. A subsample was lyophilized, ground to a uniform size with 

a mortal and pestle, and Soxhlet-extracted for 12 hours in a 100'X .hexane 

solvent system. All glassware and aluminum thimble were precleaned with 

Micro cleaning solution, washed with nanograde solvents, and combusted at 

500°C for at least four hours. The extracts were concentrated in a Buchli 

Rotovapor R to a volume of about fine ml. Care was exercised at all time 

to ensure that the extract was not brought to complete dryness, in order 

not to volatilize the lighter sample components. The volume of the 

extract was raised to seven ml with hexane, and the extract was stored at 

4°C in the dark until further analysis . A total system blank was run 

routinely for every set of samples, processed and checked by both 

fluorescence and gas chromatography to ensure acceptable blank levels. 

The first level of fluorescence was similar to that of Hargrave and 

Philips (1975) " Analyses were performed with a Perkin-Elmer 650-40 

spectrofluorometer. Emission at 370 nm (excitation 265 nm) Was used as 

the primary parameter to determine the presence or absence of petroleumderived aromatic hydrocarbons. Fluorescence is this spectral region is 

primarily produced by aromatic hydrocarbons with three or more rings and 

provides the greatest sensitivity for the method. Samples which exhibit 

significant fluorescence at the original dilution can be further diluted 

until a proportional decrease in the emission spectrum indicates that 

quenching is insignificant. This level of characterization also provided 
47 

ground glass neck, and mixed with 95% ethanol (150 ml), 50 ml of hexane, 

several glass beads or boiling chips, and KOH (10 g) . The mixture was 

refluxed at 80°C for four hours. 

Following the gOH/ethanol reflux, the digested material was 

transferred to a Teflon-stoppered separator funnel (one liter) using 

distilled H2O (80 ml) and two portions (50 ml each) of hexane . The 

mixture was shaken by hand for five minutes to allow equilibration, and 

the solvent and aqueous phases were allowed to separate . Additional 

hexane was added if the two phases did not separate. The two phases were 

drained into separate flasks and the aqueous phase was returned to the 

separatory funnel using a hexane (50 ml) wash. The extraction and 

separation was repeated three times . 

The combined hexane extracts are washed (minimum three times) With 

aliquots (500 ml each) of distilled H2O to remove solids and residual 

alcohol. A saturated salt solution is often used to break the emulsion. 

To remove any residual water from the hexane extract, anhydrous Na2S04 (2-

3 g) is added . Copper turnings are added to the flask and the extract is 

refluxed for one hour to remove sulfur. The extracts are then rotoevaporated to near dryness and transferred to clean vials with methylene 

chloride. At all times care is exercised to ensure that the extract does 

not go to complete dryness to prevent loss of the more volatile 

components . 

The extracts were fractioned into saturate and aromatics/esters 

fractions on alumina/silica gel columns. The silica gel was activated at 

150°C for 16 hours and the alumina was activated at 350°C for 12 hours . 

The alumina and silica gel were then deactivated with 5% : water. 

Deactivated packings were prepared immediately prior to their use. Ten 

grams of alumina and 10 g silica gel were hexane slurried individually 

over a plug of glass wool. The columns were cleaned with 100 ml of hexane 

which was then discarded . 

The sample extract was dissolved in approximately one ml of hexane 

and applied to the surface of the column. A hexane (100 ml) and a 

benzene :hexane (100 ml, 50:50) fraction was then eluted . Optimum liquid 

chromatographic conditions and recoveries were tested using authentic 

standards . After collection each fraction was roto-evaporated, 

transferred to vials, and dried. The hexane fraction weight was obtained 
48 

by dissolving the sample in 100 microliters of methylene chloride from 

which a 20 mieroliter aliquot was withdrawn and applied to a pre-weighed 

filter pad. The solvent evaporated and the aliquot was weighed to a tenth 

of a microgram. 

The benzene:hexane fraction was further purified using a Sephadex LH20 column (25-100 u mesh). The Sephadex was slurried in the eluting 

solvent (cyclohexane :methanol:dichloromethane ; 6:x :3), allowed to swell 

overnight, and slurry-packed in a glass, teflon-stoppered column. Each 

column was calibrated with authentic aromatic standards to determine the 

fraction to be collected. The column was pre-rinsed with 200 ml of the 

eluting solvent, then the sample, dissolved in the eluting solvent, was 

applied to the top of the column. The first 40 ml of the eluant was 

discarded while the next 100 ml was collected. This fraction was rotoevaporated, transferred to a vial, and weighed as for the hexane fraction. 

Each sample was spiked with a known amount of several internal 

standards to correct for variability in recoveries and extraction 

efficiencies . Several compounds of similar structure but not naturally 

occurring were added for both aliphatic (i.e., 1-ehlorooctadecane, 

polyolef ins, or branched alkanes) and aromatic (i.e., hexamethylbenzene, 

branched aromatics, etc .) analyses. The concentrations of the internal 

standards were in the same range expected for naturally occurring 

compounds. The compounds chosen must be sufficiently resolved from all 

sample components. Authentic standards have shown that this analytical 

procedure provides the desired results. 

The aliphatic and aromatic fractions from the columns were quantified 

by fused silica capillary gas chromatography. Hewlett-Packard gas 

chromatographs (Model 5880) were utilized in a spitless capillary mode. 

Fused silica capillary columns coated with a bonded phase (BPI/QC2; SGE, 

Ltd .) Were used to attain separation of the extract components. Baseline 

separation of n-Cl7 and pristane and n-C1 8 and phytane was maintained at 

all times to insure proper resolution. The columns were 50 m long with an 

inside diameter of 0 .25 mm. 

Helium gas was added as a makeup gas between the capillary column and 

the flame ionization detector, in order to obtain the maximum sensitivity 

of the detector . The injection port was operated at 300°C and the 

detector at 350°C. Typical instrumental parameters are shown below : 
49 

PARAMETER SETTING 

Initial temperature 80°C 

Initial hold 0 min 

Rate 6°C/min 

Final temperature 300°C 

Final hold 20 min 

Infection port temperature 300°C 

FID temperature 350°C 

Chart speed 0 .50 cm/see 

Gas chromatograms were quantified with authentic standards. A 

combination of commercially available quantitative standards and standards 

prepared in the ABM laboratory were used. Sample peaks were identified by 

comparison of retention times with standards. All peaks were assigned a 

Kovats Index based on the retention times of a 20-component hydrocarbon 

standard containing normal alkanes from C13 to C34" The alkanes, by 

definition, were assigned a Kovats Index equal to 100 times the number of 

carbon atoms they contain (i .e., n-C14 K" I" = 100) . The standard was 

then used to calculate Rovats Indices for other compounds based on 

interpolation between normal alkanes. All significant peaks in a sample 

were assigned a Kovats Index . govats Indices compensated for daily 

variations in operating conditions and allowed the direct comparison of 

data from different runs. 

The Hewlett-Packard gas chromatographs were linked with an HP 1000 

data system used for laboratory automation and manipulation of data. 

Existing analysis packages had the capacity to calculate response factors, 

which were a measure of the response of the detector versus the area of 

the peals. From the daily quantitative standard the gas chrom atograph was 

calibrated in an external standard mode. The retention time and amount of 

each standard peak was used to calculate response factors for the standard 

compounds, and this information was stored on tape and/or hard disk. The 

output included the response, the retention time, the Rovats Index and the 

calculated concentration. The response factor was also used to determine 

the concentrations of peaks that do not correspond to components in the 
50 

standard. Every attempt was made to obtain authentic standards for all 

significant sample components . The unresolved complex mixture 

concentrations were calculated based on average n-alkane response over the 

volatility range covered . 

At least 10$ .of all samples were analyzed by gas chromatography/ mass 

spectrometry (GC/ MS) to confirm the identity of the sample components and 

to identify when possible any unknown compounds . The gas 

chromatography/ mass spectrometry was conducted with a Hewlett Packard 5996 

GC/MS system coupled with a Hewlett Packard 1000 data system . Gas 

chromatographic columns and conditions were identical to the quantitative 

GC analyses . A splitless infection technique was used . The total column 

effluent was routed directly into the ion source of the mass spectrometer. 

Standard n-alkanes were run daily to confirm Kovats Indices. Typical 

operating conditions for the mass spectrometer are listed below : 

FACTOR CONDITION 

1 . Source temperature 300°C 

2 . Detector gain 

3 . Source conditions 

a. Drawout lens 10 volts 

b . Repeller 22 volts 

c . Ion focus lens 30 volts 

d . Electron energy 70 eV 

e . X-ray 3 

f. Electron focus 0 

g . Scanning rate 215 amu/sec 

h. Electron emission 160 uA 

i . Target 160 uA 

4 . Calibration with perfluorotributylamine (PFTBA) 

The mass spectrometer was repetitively scanned from m/z 33 to 400 

every 2 .1 sec . Ionization was accomplished using 70 eY electrons. The 

ion source temperature was maintained at 3000C. All substantial peaks had 

their fragmentation patterns hard copied and all data stored on tape for 
51 

future reference. Spectral interpretations were aided by computer library 

searches (in-house and NIH/EPA/CIS), the eight-peak index, reference 

texts, and the periodical literature. 

2 .3 .2 .2 Biological Samples from Box Cores 

Meiofauna 

Meiofauaal samples were gently rinsed through a 300 micron sieve to 

remove larger organisms, and then through a 63 micron sieve. The material 

on the 63 micron sieve was than placed carefully--small amounts at a 

time--into a sorting dish partially filled with water. Individuals were 

sorted by mayor taxon under a dissecting scope, using an Irwin loop to 

transfer specimens to vials containing 70$ ethanol . The vials were 

uniquely labeled according to collection date, location, replicate number 

etc., taxon, and the number of individuals contained in the vial. Biomass 

was estimated based upon published literature values for the size ranges 

of organisms in the samples (Faubel 1982, Route et al. 197u)" 

Macrofauna 

Maerofaunal samples were gently rinsed with water to remove 

preservative, placed in a Petri dish--small amounts at a time--and 

examined under a dissecting microscope. Specimens were removed and sorted 

by mayor taxonomic group into labeled vials containing 70¢ ethanol. Wetweight biomes was estimated for each taxonomic group based upon values 

reported in various published literature (Faubel 1982, Rowe et al . 197) 

All mayor taxonomic groups except Nematoda, Harpacticoida, 

Aplacophora, Seyphozoa (strobilas), Priapulida, and Aearina were given 

either to in-house or consulting taxonomic specialists for identificaion 

to the species level, if possible. The following is a list of taxonomic 

groups and the specialists responsible for identification of specimens 

within each group : 
52 

TAXONOMIC GROUP SPECIALIST(S) 

Porifera Shirley Pomponi (Consultant), Daniel La Chance 

(Consultant) 

Actiniaria Daphne Dunn Fautin (California Academy of Sciences) 

Alcyonaria Jennifer LoWrq (Consultant), Steven A . Viada (LGL) 

Scleraetinia Stephen Cairns (Smithsonian Institution), Steven A. 

Viada 

Hydrozoa Dale Calder (Royal Ontario Museum) 

Nemertea Judy Wern (Texas A&H University) 

Oligochaeta Christer Erseus (University of Goteborg) 

Polychaeta G. Fain Hubbard (LGL), Paul Wolf & Linda Sierte 

(Barry A. Vittor & Assoc, Inc.) 

Bivalvia Frank Rokop (Scripps Institute of Oceanography), 

Thomas R. Waller (Smithsonian Institution) 

Cephalopoda Michael Sweeney (Smithsonian Institution) 

Gastropoda Philippe Bouchet (Museum National d " Historie 

Naturelle) 

Scaphopoda J .N . Rraeuter (Baltimore Gas & Electric) 

Pycnogonida C . Allan Child (Smithsonian Institution) 

Pogonophora Meredith Jones (Smithsonian Institution) 

Amphipoda Larry MeRinney (Texas Parks & Wildlife) 

Cirripedia Henry Spivey (Florida State University) 

Cumacea Norman Jones (University of Liverpool) 

Isopoda George Wilson (Consultant) 

Ostracoda Louis Bornicker (Smithsonian Institution) 

Stamatopoda David Camp (Florida Dept . of Natural Resources) 

Tanaidacea Richard Heard (Gulf Coast Research Lab) 

Decapoda Linda Pequegnat (Consultant), Willis Pequegnat 

(Consultant), Patsy MeLaughlin (Florida International 

University) 

Asteroidea David Pawson (Smithsonian Institution) 

Crinoidea Charles Messing (University of Miami) 

Echinoidea David PaWSOn (Smithsonian Institution) 

Holothuroidea Robert Carney (Louisiana State University) 
53 

Ophiuroidea Gordon Hendler (Los Angeles County Museum of Natural 

History 

Brachiopoda G. Arthur Cooper (Smithsonian Institution) 

Brqozoa A .J .J. Leuterman (Dresser Industries) 

Sipuneulida Mary Rice (Smithsonian Institution) 

Tunicata Claude & Franeoise Monniot (Museum National 

d'Historie Naturelle) 

Fishes Charlie Chandler (LGL), Eddie Matheson Texas A&M 

University), John McEaehran (Texas ABM University) 

2 .3 .2 .3 Trawl Samples ( Megafauna) 

Identification and Enumeration 

Megafauna from the trawl samples were removed from the storage 

containers, rinsed to remove formal in, sorted, identified, and counted to 

major taxonomic group. Specimens were examined by the taxonomists listed 

above (2 .3 .2 .2 Biological Samples from Box Cores , Maerofauna), who 

identified most to the species level. 

Size and Reproductive Condition 

Fish and selected decapod species were weighed to the nearest 0 .1 g 

(drained wet weight) and measured using the standard method for the 

taxonomic group in question. Where possible, sex and state of maturity 

were determined externally or internally if the specimen was examined for 

gut contents. 

Gut Contents 

Guts of selected representatives of common fishes were dissected and 

examined for assessment of food habits. Stomach contents were examined 

for the following parameters: 
54 

1 . Percentage fullness, by volume ; 

2 . Percentage composition by food item group, by volume ; 

3 . Wet weight of each food item group ; and 

4 . Number of individuals in each food item group . 

High Molecular Weight Hydrocarbons 

As mentioned in Section 2 .3 .2.1 ( Sediment Samples from Box Cores), 

the organism hydrocarbon analytical scheme was vary similar to the one 

used for sediments (Fig. 2-5) . No fluorescence screening was performed. 

Since organisms do not generally contain large amounts of sulfur, 

desulfurization with copper was not necessaary . Three tissue types 

(liver, gonad, and muscle) Were analyzed in fish specimens. Only muscle 

tissue was analyzed in other benthic fauna (shrimp, crabs, etc .) . 

Organisms are frozen at -20°C on board ship. Dissection was performed in 

a shore-based, clean laboratory . 111 utensils were pre-cleaned using 

procedures described in the sediment section. The target sample weight 

was 15 g wet weight. The method of digestion of tissues was identical to 

that used for sediment. The methods used in column separation, gas 

chromatography (GC), and gas chromatography/ mass spectrometry (GC/MS) were 

also identical to those used in the sediment analytical scheme. 

Carbon Isotopes 

Carbon isotope methods are described in Section 2 .3 .2 .1 . 

2 .3 .2 .1 Benthic Photography 

A procedure for detailed evaluation of benthic photographs was 

developed specifically for this project. Benthic photography samples 

obtained from photographic transacts were processed on a digitizing pad 

driven by a microcomputer. The sizes of objects seen in the photographs 

were calculated from their distance from the camera (i.e. camera altitude, 

recorded in the corner of each shot) and the acceptance angles of the 

camera lens. Knowledge of the scale of the photographs made it possible 

to calculate the area shown in each photograph, and to measure the sizes 
55 

of various features and biota. Each photo thus served as a quantitative 

quadrat sample of the survey site (Grassle et al. 1975) . 

The acceptance angles of the lenses were 35° (length) and X8 .5° 

(Width) . The dimensions of one half of a quadrat can be calculated 

trigonometrically as : 

1/2 quadrat length - altitude x tan 17,50 

1/2 quadrat width -_ altitude u tan 24 .25° 

The area included in a photograph is twice the product of these two 

numbers . 

Processed film was projected through a modified bulk film projector 

and a front-silvered mirror mounted at a u5° ale (Fig. 2-6). The mirror 

reflected the photographic image onto a Houston Instruments Hi-Pad DT-11YA 

digitizing pad connected to an Apple IIe microcomputer. The software for 

the computer was developed by LGL 

A subsample of either 100 frames (Cruise II) or 200~frames (Cruises 

III-IV) Was selected from an entire roll from each station, using a 

systematic sampling technique described by Coehran (1977) . From each 

frame., the digitizer operator first recorded the following on 

microdiskette : 

1 . Cruise number ; 

2 . Date (day, month, year) ; 

3 . Station number; 

4 . Time of day of exposure (hour, minute, second) ; 

5 . Bedform type (deviations from a flat bed) ; 

6 . Sediment color ; 

7 . Bottom tape ; 

8 . Camera altitude ; and 

9 . Bottom depth . 

The digitizer's cursor was then used to count and measure the 

subjects in the photograph. The operator had the ability to select any of 

three means to measure objects seen, depending on his judgement of the 

best representation of the object. He could determine whether the object 
56 

film projector 

45° mirror 

digitizing palette 

cursor commands . O ^^ ' ' R, - .- ,. 

0 

O 

cursor 

~ I 

Figure 2-6 . Schematic representation of digitizing apparatus 

used for processing benthic photographs . 
57 

was most appropriate to measure as a point, a straight line, or a closed 

figure. Each procedure utilized a different software routine, which could 

be activated with a cursor command. 

Data records from the digitizer were transmitted from the Apple He 

to a database. The custom software used the scale of the photographs to 

convert the digitized images and counts into estimates of lengths, areas, 

and densities. Types of data records or categories resulting from 

photographs included the following: 

1 . Numbers of benthic invertebrates, identified to the lowest 

possible taxon, encoded as points or lines representing 

appropriate lengths or widths for each taxon; 

2 . Numbers of fish, identified to the lowest possible tagoa, 

encoded as points or lines representing total length; 

3 . Man-made artifacts such as cans, bottles, plastic, etc ., 

encoded as points, lines, or areas ; 

Terrigenous or near-shore materials such as sea grass 

blades, encoded as points, lines, or areas; 

5 . Consolidated materials such as hard rocks, siltstone, 

etc., generally encoded as a closed figure or area; and 

6 . Lebensspuren or traces in sediment left by living 

organisms, encoded as lines or areas if possible, but in 

some instances as points (and then converted to areas 

represented by the minimum resolution of the digitizer) . 

2 .3 .2 .5 Quality Assurance and Quality Control ( QA/QC) 

Biological Samples 

The biologists who were involved in the sorting and identification to 

mayor taxonomic groups were periodically checked and instructed to 

maintain the high level of efficiency needed for this project . Each 

sorter was given an indoctrination and training session to familiarize 

that person with the project's goal and objectives, and to teach him/her 

the details of the specific techniques he/she would be using for sample 
58 

processing. Most sorters were biologists with bachelor's or master's 

degrees . 

Macrofauna sorters were initially given close supervision and 

training while processing their first sets of samples. These "training" 

samples were rechecked by an experienced supervisor before a new sorter 

was allowed to proceed further. Additional instruction and training was 

given if necessary . If the sorter seemed unable to perform 

satisfactorily, he was not allowed to process any more samples. Once the 

laboratory supervisor was satisfied with the sorter's performance, he was 

placed in the routine schedule for sample Workup . 

After the training period, each sorter was periodically tested for 

sorting efficiency, proficiency in taxonomic identification, and ability 

to follow sample handling protocols such as chain-of-custody procedures 

and proper labeling techniques. The usual standard was to accept a 

sorter's past work if random lq-selected samples were sorted with at least 

90% accuracy, i .e. at least 90% of the individuals in the samples were 

correctly identified. Counts of organisms in each vial were checked, and 

the sediment residue was also examined. If the work was fudged 

inadequate, all samples previously handled by the sorter were re-worked. 

Continued poor performance resulted in the sorter's dismissal or transfer 

to other duties. 

Upon arrival in the lab, samples were inventoried and organized prior 

to sorting. Samples were subsequently checked out individually on a 

"Sample Check-out Sheet" located in the storage area. The sheet required 

each sorter to record his initials, the sample number, station, replicate, 

and date checked out for sorting. After sorting, the date of completion 

was recorded on the Sample Check-out Sheet, and on a "Sample Disposition 

Status Sheet" 

Sorted taxonomic groups were separated and inventoried by taxon, and 

sent to the appropriate taxonomic specialist(s) along kith copies of the 

Sample Disposition Status Sheet, station data sheets, and work 

authorization forms. Samples were sent by Certified U.S. Mail, Return 

Receipt Requested . Taxonomic specialists were instructed to send 

identified material directly to the Smithsonian Institution, U.S. National 

Museum of Natural History, along with the proper labeling information. 
59 

Mis-sorted material was returned to LGL, and re-directed to the 

appropriate specialist. 

As part of an ongoing effort to document the quality of samples 

collected, a study was conducted on box core macrofaunal samples, using 

the basic assumption that a "good" sample should have the same proportions 

of mayor taxa in both the box and the meiofaunal tubes from the same 

replicate. Meiofaunal tubes were more protected by the box core lid from 

washout, and were sieved under controlled laboratory conditions as opposed 

to field conditions. Paired samples (box and tubes) were thus analyzed 

for relative proportions of mayor taxa . 

Following Hessler and Jumars (197) and Jumars (1980), a Chi-squared 

test for proportions was used to evaluate significant departures from 

expectation. Afield-assigned quality designation was indicated by 1, 2, 

or 3, representing low to high quality based on observations when the 

corers came on deck . Only seven of the 30 replicates tested were 

demonstrated to have lower quality (Fig . 2-7) " In general, borderline 

samples were characteristically better than anticipated, based on field 

observations . 

Hydrocarbons 

Replicate analyses were performed on both a standard sediment and a 

standard fish muscle tissue sample. Results for alkane analysis are 

reported in Tables 2-1 through 2-5" Replicate gravimetric analyses were 

precise within 20 to 30% (± 1 sigma) at the ppm level (Table 2-1) . 

Replicate analysis of individual n-alkanes varied from -5 to 50% (± 1 

sigma) at the ppb concentrations in sediments depending on molecular 

weight (Tables 2-2 and 2-3) " Comparison of two analysts processing the 

same sediment sample is illustrated in Table 2-4 (gravimetric analysis 

only) . Recovery of surrogate standards was generally in the 80-90x range, 

with this percentage reaching a constant value at n-C1 9 ( i .e ., the lower 

molecular weight alkanes are lost in roto-evaporation, Table 2-5) " 

Each set of eight analyses contained six samples, one reagent blank, 

and one reagent blank plus the complete quantitative n-alkane and aromatic 

standard. All samples were spiked with an aliphatic and an aromatic 
60 

D 

D 

5 

M 

H 

e . 

O 

a 

E+ 

O 

H 

k 

e. 

. Box Cares 

IN 

Meiof auna Tubes 

Figure 2-7 . Proportion of macrofauna organisms in disturbed and 

undisturbed box core samples . 

hlydNeU vAutd, Ilu'ietieeika Oriruoia Iso'era Tmiidxa linlvia Sipiowlids 

MycAxt& IkWteli Ilvrutlalka O1kamia ImpdI TaNiJMI liv8riI SiFAMlila 

Taxa 
Table 2-1. Summary of the precision of replicate gravametric analyses of digestion/fractionation 

(MMS Analytical Scheme) and its comparison to Soxhlet extraction/fractionation 

(concentrations in ppm, ug/gm) . 

Sample Repl . ~~ 

Digestion 

Extraction 

Soxhlet 

Extraction 

Q.C . Miss . Delta Sediment 

Aliphatic Fractions 1 9 .1 

(PPM) 2 8 .9 Ave .= 10 .3 (9 .2)i, 25 .2 Ave .=31 .8 

3 12 .0 S .D .= 3 .3 (1 .4)* 33 .0 S.D .= 5 .3 

4 17 .8%; C .V .= 32 .1%(14 .9)* 38 .1 C.V .=16 .8 

5 8 .6 30 .8 

6 7 .6 --- 

7 9 .4 --- 

8 8 .6 --- 

Aromatic Fraction 

(PPM) 1 9 .1 5 .2 Ave .= 8 .8 

2 6.3 Ave .= 6 .7 7 .7 S .D .= 3 .4 

3 8.4 S .D .= 1 .5 13 .4 C .V .=39 .0 

4 6.3 C .V .= 22 .2 8 .9 

5 5 .7 --- 

6 4 .5 --- 

7 7 .4 --- 

8 6 .2 --- 

Whitefish O .C . Tissue 

Alivhatic Fraction 

(PPM) 1 20 .1 Ave .=29 .4 (24 .8)* 45 .7 Ave.=48 .0 

2 27 .7 S .D .=10 .7 ( 3 .5)* 43 .1 S .D .= 8 .8 

3 27 .3 C .V .=36 .3°,6(14 .2%) 41 .0 C .V .=18 .4% 

4 24 .2 63 .2 

5 47 .7* 46 .8 

Aromatic Fraction 

(PPM) 1 20 .6 66 .2 Ave.=54 .8 

2 14 .0 Ave .= 20 .1 38 .7 S .D .=14 .3 

3 19 .4 S .D .= 4 .5 60 .2 C .V .=26 .Ox 

4 19 .6 C .V .= 22 .5°,6 40 .3 

5 26 .7 68 .4 

* Calculated excluding the one aaomalously high concentration . 
62 

Table 2-2 . Summary of MMS QC sediment digestion/extraction replicate 

gas chromatographic analyses (concentrations in ppb) . 

Carbon/ 1 2 3 

Replicate 

4 5 6 Ave . S .D .*%" C .V .*** 

14 - - 3 .7 3 .0 1 .9 - 2 .9 0 .9 31 .3 

15 - 1 .9 3 .4 2 .4 1 .6 - 2 .3 0 .8 34 .3 

16 3 .7 7 .3 19 .0 11 .9 12 .5 7 .0 10 .2 5 .4 53 .1 

17 11 .9 17 .4 42 .3^ 21 .8 18 .0 16 .0 17 .0 3 .6 21 .0 

Pristane 17 .0 23 .9 34 .0 25 .7 26 .3 21 .4 26 .4 7 .3 27 .5 

18 26 .5 30 .5 71 .9^ 40 .7 34 .8 35 .6 33 .6 5.4 16 .0 

Phytane 28 .4 36 .5 51 .1 40 .8 38 .7 37 .8 40 .6 5 .4 13 .2 

19 34 .1 36 .6 83 .0% 48 .8 39 .0 41 .9 40 .1 5 .7 14 .1 

20 32 .5 33 .2 74.4* 38 .8 33 .9 37 .0 35 .1 2 .7 7 .7 

21 28 .2 28 .0 64 .7 30 .7 28 .4 31 .5 29 .4 1 .6 5 .5 

22 28 .4 27 .3 58 .6^ 29 .3 27 .1 30 .3 28 .4 1 .3 4 .8 

23 46 .4 49 .0 74 .8 45 .9 45 .8 49 .2 47 .3 1 .7 3 .6 

24 38 .4 36 .7 56 .2* 32 .6 32 .7 34 .6 35 .0 2 .5 7 .2 

25 143 .1^ 108 .4 111 .4 105 .2 100 .9 96 .9 104 .6 5 .8 5 .5 

26 56 .9 53 .4 68 .5* 48 .4 51 .7 51 .9 52 .5 3 .1 5 .9 

27 168 .1 199 .2 203 .4 172 .2 186 .0 185 .0 185 .7 14 .1 7 .6 

28 73 .3 60 .7 70 .1 53 .6 55 .4 56 .4 61 .6 8.2 13 .4 

29 353 .1 330 .3 310 .2 296 .9 294 .3 295 .2 313 .3 23 .8 7 .6 

30 51 .8^ 40 .8 43 .7 32 .0 35 .5 30 .1 36 .4 5 .6 15 .8 

31 195 .3 194 .7 189 .5 175 .8 175 .5 186 .2 186 .2 9 .9 5 .3 

32 6 .2 7 .2 6 .6 6 .1 6 .9 6 .5 6 .6 0 .4 6 .3 

Total Resolved 

Alkanes 1343 .3 1323 .3 1640 .5-` 1262 .6 1256 .9 1250 .5 1287 .2 42 .8 3 .3 

* values excluded were more than 3 a from the average . 

'^^ S .D . = Standard deviation 

C .V . = Coefficient of variation (S .D ./x x 100) 
63 

Table 2-3 . Summary of replicate digestion gas chromatographic analyses 

of QC whitefish tissue (concentrations in ppb) . 

Carbon # 

Replicate # 

1 2 `3 4 5 Ave* S .D .** C.V.*** 

16 -- -- -- -- -- 

17 18 .7 60 .8 54 .2 40 .0 40 .0 48 .7 10 .5 21 .5 

Pristane 124 .4 208 .5 186 .4 167 .6 156 .7 168 .7 31 .6 18 .8 

18 164 .1 193 .4 237 .4 264 .2 204 .9 200 .8 26 .3 13 .1 

Phytane 200 .9 198 .5 250 .1 226 .6 227 .7 220 .8 21 .4 9 .7 

19 193 .8 384 .3 517 .0 463 .2 482 .5 461 .7 56 .2 12 .2 

20 426 .4 362 .3 491 .7 444 .2 470 .7 439 .1 49 .6 11 .3 

21 191 .3 154 .7 223 .9 196 .9 221 .3 197 .6 28 .0 . 14 .2 

22 103 .6 88 .6 115 .6 115 .3 121 .3 108 .9 13 .0 12 .0 

23 39 .7 26 .5 58 .6 48 .4 65 .1 47 .7 13 .7 28 .7 

Total Resolved 

Alkanes 1462 .9 1677 .6 2134 .9 1906 .4 1990 .2 1834 .4 265 .7 14 .5 

Ave . = Average Concentration 

JHC S .D . = Standard Deviation 

~`*^` C .V . = Coefficient of Variation (S .D ./x x 100) 
64 

Table 2-4 . Comparison of the precision of digestion/extraction gravametric 

analysis between two different analysts (concentration in ppm, 

Mg/gm) . 

Sample Repl . # 

Analyst 

#1 

Analyst 

#2 

Q.C . Miss . Delta Sediment 

Aliphatic Fraction 1 9 .1 6 .3 

2 8 .9 12 .6 

3 12 .0 16 .7 

4 17 .8 7 .1 

S 8 .6 13 .7 

6 7 .6 13 .3 

7 9 .4 -- 

8 8 .6 -- 

Ave .=10 .3 Ave.=11 .6 

S .D .= 3 .3 S.D .= 4.1 

C .V .=32 .1% C.V .=35 .0% 

Aromatic Fraction 1 9 .1 10 .5 

2 6 .3 6 .1 

3 8 .4 7 .4 

4 6 .3 11 .3 

S 5 .7 8 .3 

6 4 .5 7 .3 

7 7 .4 -- 

8 6 .2 -- 

Ave .= 6 .7 Ave .= 8 .5 

S .D .= 1 .5 S .D .= 2.0 

C .V .=22 .2% C.V .=26 .6% 
65 

Table 2-5 . Summary of accuracy experiments for selected n-alkanes 

(reported as percent recovery of individual compounds) . 

Replicate 

% Recovery 

Carbon# --l 2 3 4 Ave . 

13 2.6 0 .0 0.0 0 .0 0 .7 

14 9 .0 1 .6 1 .4 2 .3 3 .6 

15 16 .6 2 .3 6.0 39 .7 8 .7 

16 24 .6 6 .9 13 .2 21 .0 16 .5 

17 34 .5 23 .3 23 .0 32 .1 28 .2 

Pristane 42 .7 27 .7 29 .4 36 .1 34 .0 

18 65 .4 71 .0 52 .7 59 .5 62 .2 

Phytane 51 .2 51 .7 41 .4 45 .5 47 .5 

20 88 .3 94 .9 76 .6 76 .0 84 .0 

21 83 .6 91 .7 75 .0 72 .5 82 .8 

22 84 .0 95 .0 77 .1 73 .6 82 .4 

23 85 .6 99 .9 78 .3 76 .0 85 .0 

24 83 .2 100 .2 77 .6 75 .0 84 .0 

25 82 .5 114.7 78 .9 76 .8 88 .2 
66 

internal standard. All gas chromatographic calculations were corrected 

for internal standard recoveries but gravimetric data were not. 

Diesel oil, lube oil, and bilge water were collected from the 

vessels used in sample collection. Examples are shown in Figure 2-8. 

2 .3 .3 DATA MANAGEMENT AND STATISTICAL ANALYSES 

The sequence of data management and analysis procedures used by LGL 

is shown in Fig . 2-9 " Most of the project data comes to the data 

management group on coding forms which were designed by the data manager 

and key project personnel at the outset of the program before any data 

were collected. Field and laboratory data were coded onto these data 

forms by laboratory personnel and then processed by the procedure 

described below. Data received on magnetic tape from TAMU was transferred 

either to the Amdahl 470V/6 or to the UNIX, depending on where the 

analysis took place. 

2 .3 .3 .1 Data Entry, Verification, and Validation 

Introduction 

The process of data entry, verification and validation relies heavily 

on standard practices and experience. Various analytical determinations, 

simulations and actual implementation determine an appropriate "best" 

method for a given system and set of hardware. Since every data file is 

inherently unique, there is really no one best method . The following 

descriptions and comparisons/contrasts detail the standard methods used at 

LGL. These methods are the result of a six-year data management study 

during which various types and combinations of the methods for entry, 

verification and validation were tried, each with its own successes and 

failures . 

Data Entry 

The encoding of the data was based on the batch system. The encoding 

involved copying the field or lab information to a suitably-devised coding 
67 

574 .12 

4.1e.29 

338.45 

220.81 

102.77 

-15.05 

INJECTED i/1000 

n 

Z 

Z 

t'a 

M 

0 .00 8.75 13 .50 20 .25 27.00 33.75 40 .50 47 .25 54.ou 

Fuel Oil 

183. u 

00 

148.09 

!12.71 

77 .38 

42 .09 

M 

Z 

8.88 1 ~ ~ ~ ~ -

0 .00 8.75 13 .60 20 .25 27 .01 33 .78 40 .51 47 .27 64 .02 

Lube Oil 

M 

Figure 2-8 . Example gas chromatograms of possible contaminants 

collected from sampling vessels . 
68 

Coding sheets and anal yaia for mats are devised 

Date recieved from the field 

and laboratory 

More date?Jl --~ Data are coded in lab 

Data keypunched onto UNIX system 

Date listed end keypunch 

verification performed 

Erroro corrected I .-YES r-Error3? 

NO 

Date listed and verified by 

Project invatigatora 

Errors? 

Data tran3fered to machine on which 

it will be analyzed 

Anelysia of date 

Data sent to contractor at 

project's conclusion 

Figure 2-9. Data flow sequence for Continental Slope Study . 
69 

form, and manual verification that the coded form matched the original 

form. Usually this was performed by two groups, one to encode and one to 

verify content . Once a batch was complete, it was given to the Data 

Manager for incorporation into the database via the data management cycle 

of entry, keypunch verification, Project Investigator 

validation/verification and computer validation. Once received by the 

Data Manager, a Summary Report Form (SRF) was started (and attached 

permanently to the batch) which detailed information such as : 

1 . Study name, data set identification, batch ID and encoder 

ID ; 

2 . Date of receipt of coding forms by the Data Manager ; 

3 . Date of keypunch entry and ID of keypuncher ; 

4 . Total records entered for batch ; 

5 . Date of keypunch verification 

a . whether or not errors were found, 

b . date of correction of error, 

e . date of verification of errors, and 

d . back to Step 5 until no more errors ; 

6 . Date of Project Investigator validation verification 

a . Whether or not errors were found, 

b . date of correction of error, 

e . date of verification of errors, and 

d . back to Step 6 until no more errors ; 

7 . Date of integration into database ; 

8 . Date of archival ; and 

9 . Additional notes/comments pertaining to batch or data set 

as a whole . 

Once the Data Manager initiated the batch via this form, it was ready for 

entry by keypunch personnel using entry software on the LGL UNIX System V 

Operating System. A batch was entered by only one person in order to 

maintain strict control and traceability of the data. Once entered, the 

keypuncher dated and initialed the SRF, which was given to the Data 

Manager along with the batch coding forms. At this point, the Data 
70 

Manager verified dates, initials and locations of the batch, and produced 

a hard copy printout for verification. 

Keypuncher Verification 

Keypunch verification was performed by the keypuncher under the 

direction of the Data Manager. This method entailed the verification of 

the batch in a character-by-character, and line-by-line, comparison 

process of the hardcopy printout against the coding form. The coding form 

was used as the standard rather than the actual lab forms, since the 

process of keypunch verification is designed only for catching 

typographical errors, not validation errors which were caught using the 

original data forms. Since the batch was entered using field entry 

software, values which were constant for a specific field or fields needed 

to be checked only once since the field was static and therefore not 

subject to change. 

Any errors found were noted on the hardcopy printout, dated, and 

initialed by the keypuncher who performed the verification. Also, the SRF 

was updated, indicating whether or not errors were found . The batch, 

printout and SRF were then turned over to the Data Manager, who performed 

any and all corrections, noting the date of correction on the printout 

where the errors were found . Once all noted errors had been corrected, 

the SRF Was again updated to indicate this, a new hardeopy printout was 

obtained, and the Data Manager gave the batch, printout and SRF to another 

keypuncher who performed a verification of corrections. This cycle was 

continued until there mere no more typographical errors found in the data, 

at which point the Data Manager printed a final hardcopy of the batch for 

Project Investigator validation verification. 

Project Investigator Validation/Verification 

Project Investigator validation/ verification, performed by the 

Project Investigators under the direction of the Data Manager, was similar 

in most respects to the method of keypunch verification. However, Project 

Investigator validation, unlike keypunch validation, involved comparing 

the printout with the original forma This allowed any coding errors or 
71 

inconsistencies to be detected. Any errors found were noted on the 

hardcopy printout, dated, and initialed by the Project Investigator who 

performs the validation/verification. Also, the SRF was updated, 

indicating whether or not errors were found. The batch, printout and SRF 

were then turned over to the Data Manager, who performed any and all 

corrections. Once all noted errors have been corrected, the SRF was again 

updated to indicate this, a new hardcopy printout was made, and the Data 

Manager gave the batch, printout and SRF back to the Project Investigator 

who performed a verification of corrections . This cycle was continued 

until there were no more validation errors found in the data, at which 

point the Data Manager printed a final hardeopy of the batch to be used in 

computer validation. 

Computer Validation 

This method involved the compilation of a variables' options and 

constant list for a batch compiled by the Project Investigator under the 

direction of the Data Manager . This list details all possible, 

checkable", options and ranges which were valid for the batch variables. 

It does not list such things as actual lengths, weights, and other finite 

variables ; however, the ranges provided indicated a problem (e .g ., if a 

copepod is shown to bane a length of 25 mm when the valid range is from 1 

mm to 2 mm) . The actual validation is performed via a validation 

algorithm, based on a general scheme of validation for any study (dates, 

cruises, stations, replicates, ete.) in addition to options designed for 

the batch and data set specifically. Due to its design, modification for 

specific options was easily performed interactively during program 

execution. The Data Manager created a file containing the variables' 

options and constants, and verified the information to match the original 

list. Then the computer validation program was run on the batch, at which 

time any batch specific options were indicated by the Data Manager to the 

program. The final output of the program was a file and hardcopy listing 

of any errors. This output included : 

1 . The location in the batch of the error ; 

2 . The error ; 
72 

3 . Why the error was detected ; 

4 . How the error was detected ; 

5 . Whether the error was "constant" (perhaps the variables' 

options and constant list needs to be updated) ; and 

6 . The total number of errors found in the batch . 

The Data Manager and the Project Investigator then used the information to 

trace the error(s) . Once traced, the error was noted on the printout of 

the data, and the SRF was updated to indicate the batch progress at this 

point. The Data Manager performed any corrections, produced a new 

hardeopy printout, and re-ran the computer validation program. This cycle 

was continued until no further "computer-catchable" errors were found, at 

which time the SRF Was updated to indicate that the batch was ready for 

incorporation into the data base . This method of data verification 

resulted in error rates of <0 .5x" 

A cope of the data files is then transferred to magnetic tape which 

has the following format : unlabeled, 9-track, 1600 bpi, 6160 fined block 

format (blocking factor 77), 80 logical record size (blank-filled), 

EBCIDIC, tape density of 3 . This tape is provided to NOD C. Data are 

fully traceable through the system, with summary reports available at all 

times to indicate project status. Magnetic tapes containing the data 

files along with all relevant documentation and accompanying letters of 

transmittal will all have been provided at the time of draft final report 

submission. 

After complete data verification the data set was ready for 

summarization and statistical treatment. Most data analysis was performed 

on Apple microcomputers using commercially available or custom designed 

software . 

2 .3 .3 .2 Statistical Analyses 

All of the data analyses presented in this report are of a 

preliminary and exploratory nature, and the methods used do not 

necessarily represent those that will be used in the final report. 

Because of their preliminary nature, little justification is warranted, at 
73 

present, concerning their appropriateness and use. This will be provided 

in the final report. 

Clustering 

The clustering presented in this report follows the method of NESS 

presented in Grassle and Smith (1976) . This was performed with no taxa 

deleted following the description in the reference. The data used t'or 

these cluster analyses were the megafaunal counts and densities from trawl 

sampling. As noted in the description of the method, the parameter (m) 

specified for this analysis is the expected number of species shared . 

After several trial runs using various values of m, an m of 30 was 

selected to reflect similarities and dissimilarities in the dominant taxa. 

Diversity and Evenness Calculations 

All diversity calculations presented in this report were the ShannonWeiner index of diversity computed as follows : 

_E~=1 ~~) loge 

N N 

where 

S -_ number of species, 

Ni -_ number of individuals in the collection of species 3, 

N 

S N 

J 

, the total number of individuals in the entire 

= E ._1 

~-

collection. 

The evenness calculation was made using the following: 

V' = H"/loge(s) 

These formulas and further descriptions of these summary statistics 

may be found in Pielou (1969) . 
74 

The diversity and evenness values were calculated for species level 

data only. Ultimately we plan to make statistical comparisons of the 

diversity and evenness values. Following Bowman et al . (1971) the 

estimate of diversity has an approximately normal distribution. That is, 

the diversity from a single collection with sufficient number of 

individuals and species (see Bowman et al . 1971 for specific limits) is 

approximately normal with stable variance (i .e ., with variance not 

depending on the underlying mean diversity) . 

Given normalcy and stable variance, Analysis of Variance on the 

calculated diversity and hence evenness values will be justified . The 

diversity will be calculated for each of the replicated macrofauna 

collections, and all replicates will be used in the ANOVAS. 

As an ad-hoc test for single collection diversity (trawl collections 

of megafauna) the approximate sample standard errors are used in this 

report to construct 95% confidence intervals around the diversity 

estimates . The form of the standard error estimate is, following 

Hutcheson 1970: 

Var (H") _ [E~_1 ~ loge () - (,j=1 -1 loge (N )) 

N N N 

Density Calculations 

All densities appearing in the report derived from box core sampling 

are calculated as count divided by area and multiplied by the appropriate 

factor to give density in square meters or centimeters. The area sampled 

for each of the density calculations appears in the appropriate design 

table (see Appendices) . 

Density calculations for the trawl sampling was somewhat more 

complicated. The calculations involved first calculating bottom time of 

the net and multiplying this value times the ships speed to determine the 

distance the net was towed . This value was then multiplied times the 

width of the trawl to obtain the area sampled . The counts were divided by 
75 

this area and multiplied by the appropriate factor to obtain number per 

hectare . 

Length Weight Regressions 

To determine the relationship between length or weight in the species 

accounts of megafauna, the usual log-log transformation was made. This 

was made after a significant deviation from linearity was observed in the 

raw length weight regression residual plots. 

Bathymetric and Longitudinal Zonation Analysis 

To evaluate whether bathymetric and longitudinal zones were suggested 

by the megafaunal species distributions, the analysis of species 

homogeneity found in Gage (1986) and Backus et al . (1965) was used . The 

approach used in the application of these procedures was to detect areas 

of species heterogeneity between stations. The application of the methods 

outlined in the above papers is straightforward . First, the empirical 

distribution of number of species starts and stops is generated. Next, 

the observed patterns of species starts and stops is compared to the 

homogeneous pattern using a Chi-squared test . Significant Chi-squared 

values for a start/stop point indicate a region of significant 

heterogeneity or discontinuity. For a more detailed description of this 

analysis see Backus et al . (1965) " 

In Section 4 .3 the trawl collections of megafauna are described with 

the results summarized by depth interval. The project plan was for trawl 

collections to have been taken within defined bathymetric limits 

corresponding to Pequegnat's (1983) faunal assembages as defined 

previously (see Section 1 .2) . With the trawl collections, maintaining the 

sampling within the defined bathymetria limits proved difficult because 

(1) the hauls were long, (2) the bathymetry maps used to plan the tows 

were not necessarily accurate nor precise on the spatial scale being 

sampled, and (3) other considerations (e .g., safety) sometimes dictated 

the ships course while underway . 
76 

When the depth ranges covered by each trawl-tow over all 

cruise/station combinations were plotted it was found that, overall, the 

samples were best summarized by six depth intervals : 

Median Depth Overall Depth 

Within Interval Interval 

Sampled (m) Sampled (m) 

375 328- 423 

608 465- 750 

876 751-1000 

1275 1050-1500 

1800 1550-2050 

278 2100-2855 

The differences between median depths, from shallow to deep, were 233 

268, 399 525 and 678 m. The depth difference between sampled depths 

increased over the depth range and this factor may account for some of the 

observed differences ; i.e., some apparent faunal breaks may be more 

related to a greater depth-Wise "distance" between some stations as 

opposed to others, than to real faunal breaks . 

3 .0 PHYSICAL AND CHEMICAL RESULTS 
79 

3 .0 PHYSICAL AND CHEMICAL RESULTS 

3 .1 HYDROGRAPHY 

Tabulated results of hydrographic profiles taken at five stations on 

each cruise are shown in Appendix B1 of Volume III of this report series. 

Depth related changes in physical and chemical constituents followed 

expected trends, with some variation by cruise and station location within 

the Gulf. Surface to bottom profiles of temperature, salinity, dissolved 

oxygen, phosphate, nitrate, and silicate values for each cruise are 

provided in Attachment A1 .1 . 

3 .1 .1 TEMPERATURE 

From the surface (upper 25 m) to approximately 1300-1400 m depths 

temperature dropped from near 200 (19.01-25 .08°C) to about u.22-4 .350C at 

1300-100 m depth (Fig. 3-1) . Below 1400 m temperatures remained constant 

within this low range. Between 300 and 1400 m, temperature variations by 

depth within transects was not great. On the central transect, Stations 

C1, C2, C3, C4 and C5 were each occupied three times (Cruise I-III) . The 

maximum variations over the three cruises at representative depths of 300, 

600, 900, and 1400 m were 2 .59°, 0 .78°, 0 .29°, and 0 .10°, respectively. 

Similarly on Cruise IV in the eastern Gulf, profiles were measured at 

stations with bottom depths of 380, 665, 885, 915, and 2920 m . 

Temperature variations by depth along the transect were low ; ranges were 

1 .93° (18 " 35-20 .28°C) at 100 m ; 1 .72° (9 .31-11 .03°C) at 400 m ; 0 .68° 

(7 .05-7 .73°C) at 600 m and 0 .16° (5 .10-5 .26°C) at 900 m. 

On a large scale, temperature differences from the western to the 

eastern Gulf also showed only minor changes (Fig. 3-2) . From the western 

to the eastern Gulf the most temperature variation by depth was only 

0 .49°C at 275-m depth. 

3 .1 .2 SALINITY 

Salinity profiles exhibited the characteristic pattern for the Gulf 

of Mexico (see Attachment A1 .1) . Surface salinity ranged from about 35 to 
80 

Temporetur't (C) 

10 20 30 

WATER 

TROPICAL ~ x' 

ATLANTIC 

CENTRAL 50 T 

ANTARCTIC ; . 

lNTERMEDIAT 

1000 

E 

r 1500 

.r 

a 

O 

2000 

GULF DEEP 

WATER 

2500 

3000 

Phosphate (0) 

0 1 2 3 

0 

500 

1000 ., 

E 

v 

r 

1500 

2000 

2500 

3000 

Cruises 1- Y 

Salinity (ppt) Dissolved Oxygen WA) 

34 35 36 37 2 3 4 5 6 

. . .~.~ 0 SAL MAX 

.

~r 02 MIN 

." 

T ' N03/P`04 MAX 

8AL MIN 

. SILICATE MAX 

1000 1000 ' 

1500 S 1500 

2000 2000 

2500 2500 

3000 3400 . 

Nitrate OM) 

10 20 30 40 

Silks CUM) 

10 20 30 40 

SAL MAX 

02 MIN 

N03/POq MAX 

SAL MIN 

SILICATE MAX 

Figure 3-1. Hydrographic profiles, all cruises combined . 
81 

.. 

U 

O 

v 

41 

H 

v 

a 

H 

14 .0 

13 .5 

13 .0 

12 .5 

12 .0 

11 .5 

11 .0 

10 .5 

10 .0 

9.5 

9.0 

8 .5 

8 .0 

7 .5 

7 .0 

6 .5 

6.0 

5.5 

5.0 

4.5 

4.0 

m ~- .0 

93' 19 .2' 89' 45 .5' 86' 34 .6' 

Longitude 

275 m Tamp 

+ 400 m Temp 

f 775 m Tamp 

-a- 1100 m Temp 

r 1425 m Temp 

Figure 3-2. Variation in temperature at selected depths measured at 3 

stations (W4, C4, E4) across the Gulf during Cruise II . 
82 

about 36 .5 °/oo and increased with depth to about 100-200 m where the 

salinity maxima, characteristic of the core of the Gulf Common Water, is 

found (Fig . 3-1) . Below the salinity maxima, salinity decreased to a 

depth of about 600-800 m where the sali-.ity minimum is located . The 

salinity minimum is used as an indicator of the core of the Antarctic 

Intermediate Water Mass. 

Using the same depths that were used above to compare seasonal 

temperature differences by depth along the central transect stations 

during Cruises I-III, salinity showed the following variability for depths 

between 300 and 1400 m : 

Depth Extreme Range 

300 m 35 .16-35 .70 0/00 0 .54 0/00 

600 m 3 .90-3 .94 0 .04 

900 m 34 .89-34-95 0 .06 

100 m 34 .94-34 " 98 0 .04 

Thus, less than 0 .1 °/oo salinity variation was observed for any depth on 

the Central Transect over three cruises . 

Longitudinal variations in salinity based on measurements taken 

during Cruise II at similar depths from stations in the western, central 

and eastern Gulf are shown in Figure 3-3 . At 275 and 400 m salinity was 

slightly higher in the eastern Gulf, with the central Gulf station (CU) 

having the lowest salinity at these depths. (Extreme difference of 0.07 

°/oo.) At 775 m and below (1100-1400 m) there was a slight but general 

decrease in salinity from west to east (less than 0 .04 °/oo difference) . 

These differences were less than the observed difference along the central 

transact at similar depths over three cruises for December 1983 to 

November 1981 . 

3 .1 .3 DISSOLVED OXYGEN 

Dissolved oxygen concentration (Fig . 3-1) ranged from about 2 .5 to 

5 .5 ml/1 (-_ -3 .6-7 .8 mg/1). These concentrations are not considered to be 

limiting to aerobic fauna found in the ocean. Dissolved oxygen 

concentrations were higher in the surface waters (<50 m) decreasing to the 
83 

35.6 

35 .5 

35.4 

35.3 

.. ~ 275 m 

c + 400 m 

a 35 2 - 

775 m 

. 

r ~ 1100 m 

c 1 t 1425 m 

35.1 

35.0 -

34 .9-

34.8-

9319 89 45 8635 

Longitude 

Figure 3-3. Variation in salinity at selected depths measured at 3 

stations (W4, C4, E4) across the Gulf during Cruise II . 
84 

oxygen minimum layer (about 400 m depth), which is used to identify the 

Tropical Atlantic Central Water Mass . In the eastern Gulf, a secondary 

oxygen minimum lager at 150 to 300 m is sometimes found, associated 

with the Eastern Gulf Loop Current (Nowlin and MeLellan 1967) " This 

phenomenon was apparently present during Cruise II, on the eastern 

transect station E4, where two oxygen minima were observed (one at 174 and 

one at 400 m. 

Below the oxygen minimum, D.O, concentrations increased to a depth of 

approximately 1500 m . From 1500 m to 2200 m dissolved oxygen 

concentrations remain relatively constant at approximately 5 ml/1 . 

Station WC12 sampled on Cruise V produced some anomalous D.O, results 

although temperature and salinity profiles were normal (see circled values 

on Fig . 3-1 ; see Fig. A1 .1-7 in Attachment A1 .1) . No explanation other 

than sample handling or analytical error have been postulated for these 

results, although there is no reason to expect these. 

Across the Gulf, dissolved oxygen levels varied less than 0.2 ml /l at 

a given depth except for measurements made at 1400 m (Fig . 3-4) . The 1400 

m depth is an area of increasing D.O. levels and small-scale differences 

in depth of measurement would have more impact than measurements made at 

other depths. 

3 .1 .4 NUTRIENTS 

Nitrate, phosphate and silicate concentrations are used as indicators 

of Antarctic Intermediate and Gulf Deep Water Masses. The nitrate and 

phosphate maxima, which usually coincide along with the salinity minimum, 

delineate the core of Antarctic Intermediate Water while the silicate 

maximum marks the upper boundary of Gulf Deep Water (a mixture of North 

Atlantic Deep Water and Carribbean Midwater Masses) . 

Nitrate levels increased from near zero at the surface to "'30-33 UM 

at 500-700 m depths (Fig . 3-1) . Below 500-700 m there was a gradual 

decrease to about 20 pM (Fig. 3-1) . Phosphate levels followed a similar 

pattern with maximum values of 2 .0-2 .6 uM (Fig . 3-1) . Both nitrate and 

phosphate levels usually exhibit marked increases at about 100-200 m, 

probably a result of decreased photosynthetic activity. 
85 

5.0 

4.5-

.. 

0 4.0 

°° ~ 275 m 

o + 400 m 

f 775 m 

v .o- 1 loom 

a 3.5 f 1425 m 

.r., 

A 

3 .0 -

2 .5 

93 19 8945 86 35 

longitude 

Figure 3-4 . Variation in dissolved oxygen at selected depths measured 

at 3 stations (W4, C4, E4) across the Gulf during Cruise II . 
86 

On Cruise II, there was a general west to east increase in nitrate 

concentrations below 275 m with maximum values observed on the Central 

Transect (Fig. 3-5) " Phosphate concentrations by depth were usually lower 

in the east with maximum values in the west or central transect depending 

on depth (Fig. 3-6) . 

Silicate concentrations showed less variability at a given depth than 

either phosphate or nitrate (Fig . 3-1) " Silicate levels increased from 

near zero values at the surface to maximum values of about 25 to 30 uM at 

800-1000 m. From these depths to the bottom, values decreased on the 

order of 1-5 p M down to " 3000 m . Values were less variable with 

increased depth. 

3 .1 .5 PARTICULATE ORGANIC CARBON (POC) 

POC values ranged from 1 to 83~pgC/1 (Appendix Table B1) . Many of 

the higher values were observed at depths of less than 100 m with a 

tendency for lower levels to be associated with increased depth. However, 

levels in the 30-45 ugC/1 range were observed across 500-2500 m depths. A 

general decrease from west to east was observed in POC values during 

Cruise II (Fig . 3-7) " 

3 .1 .6 DISSOLVED ORGANIC CARBON (DOC) 

Dissolved organic carbon concentrations, measured on Cruises II, III, 

and V, decreased with depth. In the top 100 m, values usually were 

greater than 1 .0 mgC/1--at times reaching 2.85 mgC/1. Below 100 m, levels 

were usually from 0.5 to 0 .9 mgC/1 although the maximum value observed was 

2 .85 at 299 m (Cruise III, Station C3) . 

3 .2 SEDIMENTS 

3 .2 .1 SEDIMENT CLASSIFICATION 

Sediments were classified into categories using the graphical 

sediment triangle representing percentages of sand, silt and clay in the 

sample. Forty-five stations were sampled during a total of five cruises. 
87 

34-

32 -

30 -

F 28 -

4w e 

26 

f 

24 

22 

d 

20 

93 19 8945 

Longitude 

8635 

f 

t 

275 m 

400 m 

775 m 

1100m 

1425 m 

Figure 3-5 . Variation in nitrate at selected depths measured at 3 

stations (F14, C4, E4) across the Gulf during Cruise II . 
88 

2.0 

1 .5 

0 

t t .o 

0.5 

OA 

-& 275 m P04 

f 400 m P04 

t rrs m Pa+ 

too m Poa 

1400 m Pa 

9319 8945 0635 

Longitude 

Figure 3-6. Variation in phosphate at selected depths measured at 3 

stations (W4, C4, E4) across the Gulf during Cruise II . 
89 

20- 

18 

16 - 

14 - 

.. 

U 12- 

.. 10 - 

U 

O 

a 8 - 

6 - 

4 - 

2- 

0 - 

93 19 8945 

Longitude 

8635 

275 m 

-f- 400 m 

f 775 m 

-o- 1100 m 

-t 1425 m 

Figure 3-7 . Variation in particulate organic carbon at selected depths 

measured at 3 stations (W4, C4, E4) across the Gulf during 

Cruise II . 
90 

Fine of these stations (C1-C5) Were occupied three times and four (E1-E3, 

E5) Were occupied twice . This resulted in a total of 59 station 

measurements with replicated sediment measurements for which the data are 

summarized in Attachment A1 .2 . 

As summarized in Figure 3-8, the most common (over 55% of the 

stations) sediment type was silty-clay found in all areas of the Gulf. 

(For detail, see Figs. A1 .2-1 through A1 .2-5 in Attachment A1) There were 

small amounts of variation within this type depending on the areas of the 

Gulf sampled . In the eastern Gulf this type had slightly higher 

percentages of sand than in the western or central areas (Attachment A1, 

Figs . A1 .2-6 through A1 .2-12) . Along the Central Transect there were 

slightly higher percentages of silt than clay at the deeper stations C5 

and C12 . 

The second most common sediment type was clay, represented by nine 

stations in the western and central Gulf at relatively shallow stations 

(<1226 m) . Stations with clay sediments were relatively uniform--i .e ., 

with little variation in the sand-silt-clay proportions . 

Sandy clay was observed in the western Gulf at Stations W1, W2, W5 on 

Cruise II ; at WC5 on Cruise Y ; and in the eastern Gulf at Station E5 on 

Cruise II. At Station E5 on Cruise IV, the sediment was predominantly 

clay with approximately equal mixtures of sand and silt, compared to the 

Western Transact station which had sand in higher proportions than silt 

(see Attachment A1, Figs. A1 .2-6 through A1 .2-12) . 

The sediment type "sand-silt-clay" has approximately equal 

proportions of each sediment size fraction. Sand-silt-clay sediment was 

found in the eastern Gulf at Stations E1, E1b, E1e and E2d (at these 

shallow stations clay was the most abundant of the three parameters and 

sand was the smallest fraction) and at E4 where clay was the largest 

fraction but the sand proportion was higher than silt. 

Station WC-6 (Cruise V) was difficult to characterize because of the 

variation among the six replicates. Silty clay is probably the best 

classification based on average values . 
e92 

082 

00£ 

" daui juamTpag " g-C azn2i3 
92 

3 .2 .2 SEDIMENT HYDROCARBONS 

Sediments on the Gulf of Mexico continental slope contain a mixture 

of terrestrial, petrogenic and planktonic-sourced hydrocarbons. The 

molecular-level alkane distribution was similar in all samples, but the 

quantitative importance of the three mayor sources varied with location, 

time of sampling, and depth. Hydrocarbon concentrations were relatively 

uniform across the slope, given the large geographical area studied. 

Concentrations (pg/gm dry weight of sediment) ranged from 4 .0 to 94 .2 

for extractable organic matter, from 0 .1 to 5 .2 for the aliphatic 

hydrocarbons, and from 0.7 to 81 .4 for the aliphatic unresolved-complexmixture (Table 3-1) . A comparison with previous studies presented in 

Table 3-2 shows that these concentrations are generally lower than 

reported values for Gulf of Mexico sediments . However, the baseline values 

in Table 3-2 are primarily for coastal and shelf sediments. 

Individual hydrocarbon compounds were detected at concentrations of 

<0.01 to >0.5 jug/g. In general, the qualitative molecular-level alkane 

distribution was similar at all sites sampled. The dominant normal alkane 

between n-C15 and n-C22 was variable, whereas between n-C23 and n-C32 the 

dominant n-alkanes were consistently n-C29 or n-C31 " The alkane 

distribution in samples from the Central Transect during Cruise I are 

typical of all locations sampled (Fig . 3-9) " Detailed sediment 

hydrocarbon data for each station are found in Appendices B-4, B-5, and 

B-6 . 

3 .2 .2 .1 Hydrocarbon Sources--The Approach 

Molecular-level and bulk parameters can be used to estimate the 

relative importance of hydrocarbon sources at a given location. These 

parameters are based on the premise that a hydrocarbon source has a unique 

"fingerprint", i .e., a recognizable suite of compounds. In nature, 

however, few unique end-members occur. Several diagnostic indicators were 

monitored to better understand the dynamics of hydrocabons in Gulf of 

Mexico slope sediments (see Table 3-3) " 
93 

Table 3-1 . The ranges and averages for selected hydrocarbon parameters in 

Gulf of Mexico continental slope sediments (ug/g dry weight 

of sediment) . 

ruise 

Location 

(Transect) 

Extractable 

Organic Matter 

(ppm) 

Aliphatic 

Hydrocarbons 

(ppm) 

Aliphatic 

UnresolvedComplex-Mixture 

(ppm) 

I Central 28 .4 1 .6 23 .3 

(13 .9 - 61 .3) (1 .3 - 2.0) (19 .3 - 29 .8) 

II Central 21 .7 1 .7 8 .9 

(18 .0 - 25 .2) (1 .6 - 1 .8) ( 6 .0 - 14 .0) 

Western 26 .0 1 .1 11 .1 

(14 .0 - 55 .2) (0 .8 - 1 .3) ( 5 .2 - 11 .4) 

Eastern 8 .6 0 .7 5 .4 

( 7 .6 - 10 .9) (0 .5 - 1 .0) ( 3 .2 - 7 .3) 

III Central 18 .1 1 .4 9 .7 

( 4 .0 - 44 .4) (0 .6 - 4 .6) ( 4 .4 - 17 .4) 

IV West/Central 30 .0 0 .9 16 .8 

(17 .7 - 94 .2) (0 .4 - 5 .2) ( 4 .2 - 81 .4) 

V Eastern 7 .2 0 .2 2 .0 

4 .7 - 13 .4) (0 .1 - 0 .4) ( 0 .7 - 5 .0) 

Ranges in values . 
94 

Table 3-2 . Summary of Gulf of Mexico sediment hydrocarbon analyses . 

Location 

Texas/Louisiana - Coastal 

Texas/Louisiana - Coastal 

Gulf of Mexico - Coastal 

Florida ( Bay) - Sandy Sediments 

Muddy Sediments 

N.E . Coast - Sandy Sediments 

STOCSf - Coastal 

3TOC3 - Coastal 

(Before, during and after 

drilling activities) 

Concentrations in nit xI Predominant 

Total 9C Saturated HC Source Reference 

20-190 B(P) Smith, Jr . 1952 

Low Concentratioasl B Stevens et al . 1956 

Hiogenic Wazea2 B Bray and Evens 1961 

eve .=q .a2 eve .=2 .o2 s Palaces et al . 1972 

Ave .=86 .02 Ave.-30 .02 B 

0 .2-10.02 0.1-3 .82 B(P) Palaces et al . 1976 

Ave.-6 .25 Ave.=1 .b 

Very low levela2 B(P) Palaces et al . 1976 

0 .22-5 .602 (0-1-0-5)2 B(P) Parker et al . 1976 

ev8 .=1 .1u eoe .=o .2 

Texas/Louisiana - Coastal Banks 0 .02-0 .802 B Parker 1978 

MIIFI.I**- lisarahore Florida (00 a) lve.=1 .902 0 .29-1 .602 B Boehm 7979 

Ave .=0 .86 

- »0 a Florida Ave-=1 .392 0 .29-1 .892 

Ave-=0-83 

Mississippi/Alabama Shelf Ave.-1 .612 0 .28-2 .892 H(P) 

lve .el .1 

Freeport, Texas - Coastal 0.9-452 0 .1-2 .42 H Slowey 1980 

Ave-_39 .T Ave .=0 .71 

Texas/Louisiana - Coastal 5.71-87 .22 3.13-X9 .52 B(P) 8ulton et al . 1981 

Ava ._39 .7 Ave ._21 .4 

Texas Shelf Ave .=1 .32 B Lytle and Lytle 19T9 

Florida Coastal (<60 s) Ave-z3.12 B hearing et al . 19T6 

Y of Mississippi R. Coastal (<60 d) eve .-1t .72 B(P) Gearing et al . 1976 

Method - graviaetry (expressed as a percentage of the total organic extract recovered from the silica 

gel) . 

2 Method - GC, GUMS. 

0 South Texas Outer Continental Shelf 

"Wiaaisaippi Alabama Florida 

B Biogenio . 

P Petrogsnio . 
500 

250 

0 

500 

250 

Q 

fl. 

500 

Z 

O 

F250 

H 

Z 

W 

U o 

z 

O 

U 

500 

250 

500 

250 

0 

0 

0 

15 17 19 21 23 25 27 29 31 

CARBON NUMBER 

95 

Figure 3-9 . Molecular level alkane distributions for sediments from the 

central transect during Cruise I (ng/g dry wet of sediment,ppb ) . 
96 

Table 3-3 " Suggested molecular-level indicators of specific hydrocarbon 

sources to sediments on the Gulf of Mexico continental slope . 

Indicator Compound Source Abbreviation 

n-C15, 17, 19 and Pristane Planktonic/Petroleum PL-1 

n-C16, 18, 20 and Phytane Petroleum/(Planktonic?) PE-Lo 

n-C25, 27, 29, 3 1 Land/(Petroleum) TERR 

n-C24, 261 289 30 Petroleum/(Biogenic) PE-Hi 

Unresolved-Complex-Mixture Petroleum/(Biogenic?) UCM 

Certain assumptions need to be understood in order to properly 

evaluate these distributions as indicators of hydrocarbon sources. 

Plankton generally produce a simple mixture of hydrocarbons, including 

n-C15, n_C17, n-C19 and pristane, so the presence of these compounds can 

indicate a planktonic hydrocarbon source (PL-1). Petroleum also contains 

these compounds, but usually also contains comparable amounts of the 

alkanes n-C16, n-C18, n-C20, and phytane as well . Thus a low molecular 

weight petroleum indicator (PE-Lo) can be used to assess the petroleum 

component of the planktonic indicator. To make this calculation we assume 

that the contribution of petroleum to each indicator is equal ; therefore 

the planktonic component can be inferred as the diffferenee between PL-1 

and PE-Lo . Straight chain biowaxes with n-C25, n-C27, n-C29, and n- C31 

alkanes have been used extensively as an indicator of terrestrial or landderived input. As such, the sum of these four normal alkanes can be used 

to indicate the terrestrial (TERR) hydrocarbon component . As with the 

planktonic indicator, these normal alkanes can also have a source in 

petroleum. Again, in general, petroleum also contains a near equal amount 

of the even alkanes n-C24, n-C26, n-C28, and n-C30 (PE-Hi) . As in the 

planktonic indicator, the terrestrial component can be estimated by 

subtracting the PE-Hi from the TERR concentration. Plants themselves can 

also contain significant amounts of indigenous even carbon alkanes; thus 

this type of indicator provides a measure of maximum petroleum 

contribution and a minimum terrestrial contribution over this molecular 

weight range . In summary the planktonic input is estimated as [(PL-1)- 
97 

(PE-Lo)], the terrestrial input as fTERR)-(PE-Hi)], and the petroleum 

input as [(PE-Lo)+(PE-Hi)] . These parameters are used to assess 

hydrocarbon inputs and not bulk organic matter since hydrocarbons 

represent only a small fraction of the total organic carbon present in a 

sediment . 

These parameters, with the above mentioned limitations, can be used 

to study the hydrocarbon dynamics on the continental slope as a function 

of water depth, location, and time of sampling . Also included is the 

evaluation of parameters such as the unresolved-complex-mixture (UCM), an 

indicator of petroleum input ; the carbon preference index (CPI), an 

indicator of the relative amounts of odd and even normal alkanes ; and bulk 

sediment characteristics. 

3 .2 .2 .2 Areal Distribution 

Cruise II assessed the distribution of sediment hydrocarbons on 

transacts from the central, western, and eastern Gulf of Mexico 

continental slope. Extractable organic matter (EOM) is a composite of 

biogenic and petroleum related material. Extractable organic matter 

concentrations were generally lowest on the Eastern Transact and nearly 

equal on the Western and Central Transacts, with the exception of Station 

W1 (Fig . 3-10) " The aliphatic unresolved-complex-mixture (UCM), a 

petroleum indicator, was similar over all three transacts though slightly 

elevated in Western Transact sediments (see Table 3-1) . The elevated EOM 

at Station X11 was due to an increased IICM i.e., petroleum component. 

The amount of terrestrial hydrocarbons decreased from the Central to 

the Western to the Eastern Transact (Figs . 3-11 and 3-12) . Terrestrial 

hydrocarbon concentrations, as indicated by the sum of n-C25, n-C27, nC29, and n-C31 (TERR) concentrations, were relatively uniform with water 

depth on the Central and Western Transacts; whereas terrestrial 

hydrocarbons increased with water depth on the Eastern Transact. The 

influence of the land- and/or river-derived material was readily apparent 

in all three regions and accounts for a majority of the GC-resolvable 

alkanes . 

In general, planktonic inputs accounted for less than 10x, of the GCresolvable alkanes . Sediment biogenic hydrocarbons on the slope were 
98 

so 

so 

ao 

30 

20 

10 

0 

Eastern Transect 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2 .6 3 .0 

Western Transect 

so 

Q so 

a 

z ao 

O 

F-

< 30 

FW 20 

Z 

O io 

U 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2_6 3 .0 

0 

Central Transect 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

WATER DEPTH (km) 

- .. .. . . " EOM -Aliphatic UCM 

so 

so 

ao 

30 

20 

10 

0 

Figure 3-10 . Variation in extractable organic matter (EOM) and the aliphatic unresolved complex mixture (UCM) along transects on 

the eastern, western, and central Gulf of Mexico continental 

slope . 
99 

Eastern Transect 

0.5 

E 

a 

a 

1 .0 

Z 

O 

0.5 

H 

Z 

W 

Z o 

1 .0 

. . . . . . . . . . . . ';'YV~w~Vwir+~... . ....rmswnnn~ennnnenn4 .lA.Lt:iiwr~ 

ti+ 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

Western Transect 

~,---------------------

v 

., 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 U 

Central Transect 

1 .0 

0.5 

O 'i T -I 1 

' T* 

r~' I I ~l 

0.2 0.6 1 .0 1 .4 1 .8 2 .2 2.6 3.0 

WATER DEPTH (km) 

-PL-1 "---Terr -.-PE-Lo -. . . . .. . -PE-Hi 

Figure 3-11 . Variation in hydrocarbon source parameters along transects 

in the eastern, western, and central Gulf of Mexico continental 

slope (for definition of source parameters see Table 3-3) . 
100 

1 .4 

Central Transect 

1 .0 

- 

- 

--- - - - - - - - - - - - 

0.5 

O+_ 

0.2 

1 .4 

Q 

fl. 

v 

Z 1 .0 

H 

Q 

H 

Z 0.5 

W 

Z 

O 

U 

0 

0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

Eastern Transect 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3 .0 

Western Transect 

1 .4 

1 .0 

4 

------ - ------------- 

0 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

WATER DEPTH (km) 

"---Terrestrial " ~~~~~~~~~ " Petroleum Planktonic 

Figure 3-12 . Variation in planktonic, terrestrial, and petroleum hydrocarbons 

along three transects presented as a cumulative concentration 

(i .e ., the first concentration plotted is terrestrially sourced 

hydrocarbons, second is terrestrial + petroleum, and third is 

terrestrial + petroleum + planktonic) . 

0.5 
101 

dominated by the more resistant terrestrial components and the degree of 

dominance was a function of proximity to the Mississippi River and delta. 

The steepness of the slope and the prevailing currents affect the observed 

distributions as well. Planktonie inputs were low and often difficult 

to discern on the Central and Western Transacts (Figs . 3-11 and 3-12) . 

The planktonic input was generally higher at the shallower stations of 

these two transacts. The low planktonic hydrocarbon concentrations on the 

Central and Western Transacts may be due to a high sedimentation rate 

and/or dilution with riverine material . On the Eastern Transact the 

planktonic input was discernible and relatively constant with depth. 

Petroleum inputs, measured both by alkane parameters and the 

unresolved-complex-mixture, were detected at all sites (Figs . 311 and 3-

12) . In general, petroleum input was greatest on the Central Transact, 

with lesser amounts at the Western and Eastern Transacts . The maximum 

estimate of petroleum-sourced hydrocarbons indicates relatively low 

concentrations at all locations . To determine if the petroleum 

hydrocarbons detected were due to transported terrestrial particles (most 

likely by mass movement and turbiditq flows after initial deposition near 

the river's mouth) or to upward migration from deeper reservoired 

petroleum, the petroleum indicators were compared with the terrestrial and 

planktonic indicators (Fig . 3-13). These comparisons suggest a dual 

source for the sediment petroleum hydrocarbons. Low molecular weight 

hydrocarbons (PE-Lo) correlate with a terrestrial input on the Eastern 

Transact, but not on the Central and Western Transacts. The higher 

molecular weight petroleum indicator (PE-Hi) also correlates with the 

terrestrial indicator (TERR) on the East but not on the other transacts. 

An estimate of the amount of petroleum hydrocarbons is provided by the 

UCM. The IICM Was generally independent of the planktonic or terrestrial 

input (Fig. 3-14) Which suggests an additional source of hydrocarbons 

(such as upward migration from deeper reservoirs) on the Central and 

Western Transacts. An anomaly was seen at Station W1, which contains 

petroleum hydrocarbon concentrations that were significantly elevated in 

comparison with other Western Transact stations. 

This attempt to correlate petroleum and terrestrial sediment inputs 

assumes that the ratio of petroleum to terrestrial hydrocarbons 

transported to a location is constant with time, which may or may not be 
102 

280 

220 

a 

d 

160 

0 

J 

100 

Eastern 

40 -K 

0.2 0.4 0.6 0.8 1 .0 

250 

.a 

Q. 200 

d 

W 150 

0_ 

100 

\w1 

w 

0.2 0.4 0.6 0.8 1 .0 

TERR (ppm) 

" Central " Eastern " Western 

Figure 3-13 . The relationship between alkane petroleum indicators and a 

terrestrial indicator (for definition of the indicators see 

Table 3-3) . 
103 

30 

20 

10 

L1 

Q 

~~W1 

0.2 

U 

U 

H 

Q 

Z 

30 

20 

W1 

10 

0.4 0.6 0.8 1 .0 

TERR (ppm) 

50 80 110 140 170 200 230 260 

PL-1 (ppb) 

" Central " Eastern " Western 

Figure 3-14 . The relationship between the aliphatic unresolved complex 

mixture (a petroleum indicator) and the planktonic and 

terrestrial alkane indicator (see Table 3-3) . 
104 

true. However, extensive natural hydrocarbon seepage has been documented 

on the Gulf of Mexico continental slope, further supporting natural 

seepage as a mayor petroleum hydrocarbon input to Gulf of Mexico 

continental slope sediments. It is also evident that some fraction of the 

petroleum hydrocarbons are transported to the slope by river/land-derived 

particles most likely by secondary movement such as slumps and/or 

turbidity flows. 

3 .2 .2 .3 Temporal Variations 

Cruises I (November 1983), II April 1984), and III (November 1984) 

sampled the Central Transact in an attempt to document variability between 

samplings . The distribution of EOM and aliphatic UCM during these three 

sampling cruises is shown in Figure 3-15 . On an average, the aliphatic 

UCM, a petrogenic indicator, was highest on Cruise I (Fig. 3-15, Table 3-

1) . UCM concentrations during Cruise II and at the shallower stations 

(<1500 m) of Cruise III, were similar. During Cruise III the UCM was 

markedly higher at stations deeper than 1500 m than had been observed on 

Cruises I and II. Molecular-level indicators were similar along the 

Central Transact during Cruises I and II (Figs. 3-16 and 3-17), but 

substantial differences were also observed on Cruise III at depths greater 

than 1500 m. 

Significant variability with depth was observed during Cruise III 

sampling. Shallower stations (<1500 m) during Cruise III were lower in 

hydrocarbons than the two previous samplings, which suggests dilution by 

inorganic material. Compared with Cruises I and II, terrestrial 

hydrocarbons were significantly reduced over the entire transact . The 

deepest stations (>1500 m) on Cruise III had elevated levels of petroleum 

hydrocarbons. This is substantiated by the hydrocarbon-source parameters 

previously discussed (Figs. 3-16 and 3-17) . Examination of carbon 

preference index (PI) distributions and gas chromatograms suggest the 

presence of relatively fresh petroleum hydrocarbons, probably from oil 

seepage, at the deepest stations (Fig . 3-18) . Station C7 also had a low 

CPI, suggesting anomalouslq high petroleum hydrocarbons . These 

differences between samplings most likely represent the patchiness of 

hydrocarbon distributions rather than a temporal change such as an influx 
105 

Cruise I 

lo 

60 

50 

40 

30 

20 

10 

0 

t 

". . . . . . . 

0.2 0.6 

70 

60 Q 

v 

50 

40 

H 

Q 30 

20 

10 U 

Z 

O ° 

Cruise 11 

V 0.2 

70 

so 

50 

40 

30 

20 

1d 

Cruise III 

.e 

Station number 

1 6 2 3 7 8 94 115 12 

R 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

WATER DEPTH (k m) 

.. . - EOM -- Aliphatic UCM 

Figure 3-15 . Variation in extractable organic matter (EOM) and the aliphatic unresolved complex mixture (UCM) during three samplings of the central transect . 
106 

2.0 

0 

Central Transect II 

1 .0 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

E Q 2.0 

a 

.. 

z 

O 

F-Q 1 .0 

H 

Z 

W 

U 

Z o 

Central Transect I 

0.2 o .s 

U 

2.0 

1 .0 1 .4 1 .8 2.2 

Central Transect III 

2.6 3.0 

1 .0 

i i i i ~. 

i ~. 

Station number 

1 6 2 3 7 8 94 

o 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

WATER DEPTH (km) 

---PL-1 "--"Terr "---..-PE-Lo ". . .. . . . .. PE-Hi 

Figure 3-16 . Variations in hydrocarbon source parameters during three 

samplings of the central transect (for definition of source 

parameters see Table 3-3) . 
107 

Cruise I 

d 

2.0 

1 .0 ------------- 

0 I rT- t r 1 . 1 r r-r j 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2.6 3.0 

E Cruise 11 

d 

-------------------- 

0.2 o.s 1.0 

20 

Z 

Q 

1 .0 

FZ 

W 

U 

Z 

O o 

Cruise III 

2.0 

1 .0 11 5 X121 

0.2 0.6 1 .0 1 .4 1 .8 2.2 2 .6 3.0 

WATER DEPTH (km) 

----Terrigenous "..... . . -Petroleum -Planktonic 

0 

Figure 3-17 . Variation in plankton, terrestrial and petroleum hydrocarbons 

during three samplings of the central transect presented as a 

cumulative concentration (for an explanation see Figure 3-12) . 
108 

Central Transect 

X 

4.0 

U 

C 

3.0 

aD 

^L 

Ll 

2.0 

L 

12 

Cruise I 

. . . .. .r~~''~ '~ . . . . . . . . . . Cruise II 

--- Cruise III 

~ ~ .r .. . . . . . . . . . . . . . .. . . . . . . . . . . . . 

. 

` I ~~\ r 

f 

0.2 0.6 1 .0 1 .4 1 .8 2.2 

Water Depth (krr,) 

U 

2 .6 3.0 

Cruise III ,r, 

Station C-5 U crv 

2405 m z z °' N 

U 

Z 

N 

U 

r- z 

U 

n Z CO M U) U 

UU 

Z Z 

0 3.65 7.30 10.95 14.60 18.26 21 .91 25.56 29.21 

RT in Minutes 

Figure 3-18 . Variation in the carbon preference index (CPI) as a function 

of depth along the Central Transect during Cruise III and 

a representative fused silica gas chromatogram of the 

aliphatic hydrocarbons from Station C5, Cruise III (CPI = 

[En-C23,25,27,29,311 / [En-C24,26,28,305321)- 

295 

236 

C 

d 

177 

L 

O 118 U 

N 

50 

0 
109 

of hydrocarbons . Station C7 was later documented to have oil seepcommunity organisms in residence . 

3 .2 .2 .4 Variability Along Isobaths 

Cruise V in the eastern Gulf of Mexico occupied stations along three 

isobaths to assess lateral variation in the measured parameters . 

Hydrocarbon parameters are summarized in Table 3-4. Bulk and molecularlevel hydrocarbon parameters were low during this sampling and represent 

some of the lowest values measured during this study (Table 3-1) . 

Therefore, the variability observed along this transact is most likely a 

maximum . Cruise V sampling sites were also chosen to contrast sediment 

texture which will also contributes to the high degree of variability of 

hydrocarbon parameters observed . (Detailed sediment texture data for 

Cruises I-Y may be found in Appendix B-2 .) The aliphatic IICM and total 

EOM varied by factors of 1 .? to 7 .6 at a given depth. Molecular-level 

indicators (i.e., individual component sums) varied by a factor of 2 .0 to 

7 .6 along a given isobath. This data suggests that, at these low 

concentrations, hydrocarbons are as variable along isobaths as they are 

with water depth. This again emphasizes the patchy nature of hydrocarbon 

distributions. Bulk sediment parameters varied by as much as a factor of 

3 along an isobath illustrating the variations observed in sediment 

texture . 

Samples along isobaths in the central and western Gulf were also 

taken. Stations from Cruises I, II, III and IV at 350 m are compared in 

Table 3-5 . The variability in hydrocarbon parameters reflecting 

terrestrial input show the greatest variation, i .e., as much as 40 fold. 

The planktonic indicators are also highly variable, most likely due to 

dilution with terrestrially soureed material . Bulk parameters such as 

clay content varied by a factor of 2 and sand content varied from 0 .5 to 

36 .6% at these six locations . These variations reflect the substantial 

influence of river/land derived material. Three samples from Cruise IV 

along the 550- and 750-m isobath were relatively uniform. The lateral 

extent covered at these isobaths was small relative to the 350-m isobath 

sampling. Bulk parameters were also uniform along these two isobaths. 
110 

Table 3-4 . Variability in hydrocarbon parameters along isobaths - eastern 

Gulf of Mexico. 

Variable Ranges 

Depth (m) 342-383 619-634 819-859 

Parameter n=4 n-_6 n=5 

Total EOM 5 .8 - 13 .E 4 " 7 - 9 .9 4 .9 - 8 .2 

(9 .7)l (6 .8) ( 5 .8) 

Aliphatic 0 .7 - 5 .0 0 .5 - 3 .8 O " 7 - 3 .1 

(PPm) ( 3 .0) ( 1 .8) ( 1 .7) 

PL-12 11 .5 - 94 .1 8 .1 - 59 .1 6 .9 - 44 .8 

(PPb) (54 .3) (29 .9) ( 23 .4) 

TERR2 36 .0 - 74 " 0 55 .9 - 119 .8 23 .E - 147 .5 

(PPb) (55 .9) (78 .6) (121 .6) 

PE-Lo2 13 .3 - 100 .7 13 .5 - 39 .8 11 .0 - 27 .3 

(PPb) (56 .2) (21 .1) ( 18 .8) 

PE-H12 14 .4 - 30 .3 20 .4 - 53 .1 17 .5 - 77 .6 

(PPb) (22 .3) (33 .1) ( 48 .6) 

Terrigenous3 

(PPb) 

Petroleum3 

(PPb) 

Planktonic3 

(PPb) 

21 .6 - x+9 .8 

(33 .6) 

33 .5 - 118 .3 

(78 .5) 

0 .0 - 2 .8 

( 0 .7) 

24 .0 - 66 .7 

(45 .4) 

34 .6 - 67 .9 

(54 .3) 

0 .0 - 20 .1 

9 .5) 

5 .9 - 99 .5 

( 72 .8) 

28 .5 - 103 .5 

( 67 .4) 

0 .0 - 17 .5 

7 .3) 

Average . 

2PL-1 = En-C15, 17, 19 and Pristane ; TERR = En-C25, 27, 29, 31 ; 

PE-Lo = 

6+ 18v 20 and Pt~ytane ; PE-Hi = En-C24, 26, 2g, 3p 

Terr genous -_ (TERR) - (PE-Hi) ; Petroleum ~E-Lo + PE-Hi) ; Planktonic = 

( PL-1) - (PE-Lo) . 
111 

Table 3-5 . Variability in hydrocarbon parameters along isobaths -

west/central Gulf of Mexico . 

Depth 

Parameter 

298-371 

n=6 

547-550 

n-_3 

748-759 

n=3 

Total EOM 15 .9 - 61 .3 17 .E - 23 .9 17 .0 - 57 .9 

(PPm) (3u .5) (20 .6) (3~~9) 

Aliphatic UCM 6 .o - 31 .4 6 .9 - 7 .9 5 .6 - 11 .9 

(PPm) ( 15 .6) ( 7 .6) ( 8 .4) 

PL-1 36 .3 - 17 .9 47 .4 - 65 .3 50 .0 - 68 .0 

(PPb) (121 .8) (56 .x) (59 .9) 

TERR 93 .4 - 1082 .8 109 .8 - 273.1 169 .7 - 180 .5 

(PPb) (546 .7) (201 .2) (176 .3) 

PE-Lo 36 .2 - 151 .5 39 .1 - 52 .5 43 " 4 - 48 " 9 

(PPb) (100 .3) ( X3 .7) ( 45 .2) 

PE-Hi 70 .9 - 279 .7 80 .6 -121 .9 48 .0 - 96 .6 

(PPb) (172 .2) ( 99 " x) ( 64 .8) 

Terrigenous 22 .5 - 860 .6 29 .2 - 252 .2 83 .9 - 130 .8 

(PPb) (37u-5) (101 .7) (111 .5) 

Petroleum 123 .3 - 388 " 119 .7 - 161 .5 91 .4 - 140 .0 

(PPb) (127 .4) (143 .2) (110 .1) 

Planktonic 0 .1 - 57 .7 4 .0 - 26 .2 6 .6 - 19 .1 

9PPb) ( 21 .5) ( 12 .7) ( 1u .7) 

lCruises I, II and III, Stations E1 and W1 ; Cruise IV, Stations WC1 and 

WC2 . 

2Stations WC2, WC4, WC8 . 

3Stations WC3, WC9, WC10 . 
112 

Detailed sediment hydrocarbon data for each station sampled during 

Cruises I-n may be found in Appendices B-4, B-5, and B-6 . 

3 .2 .2 .5 Topographic Features 

One set of paired stations (WC-11 and WC-12) were taken to compare 

topographic differences . In this pairing both stations had equivalent 

depths (1200 m), WC-11 was a topographic convexity, whereas WC-12 was a 

topographic concavity. The sediment sample at the convexity was elevated 

in petroleum hydrocarbons (Table 3-6) . This difference cannot simply be 

ascribed to topographic differences and more likely suggests that seepage 

has occurred at Station WC-11 . More detailed studies will need to be 

performed to understand the relationship between hydrocarbons and 

topographic expressions. 

3 .2 .2 .6 Relationship to Bulk Sediment Parameters 

In general, the highest aliphatic hydrocarbon concentrations were 

associated with the more clay sediments rich in organic carbons . To 

understand more fully sediment ological relationships it is most effective 

to consider the three primary hydrocarbon sources individually since their 

distribution may be controlled by different factors. The data must also 

be considered in the context of the hypothesis the sampling was designed 

to test, i .e ., areal, temporal and water depth dependence . The 

terrigenous or land-derived component sometimes correlated with grain size 

distribution changes within a given sampling period, however, when the 

data were considered as a complete set, these trends were not readily 

apparent (Fig . 3-19) " This is most likely due to the ratio of clay to 

terrigenous organic matter changing with time and location. Detailed 

sediment data from Cruises I-V are found in Appendix B-2 . 

Among the samples, the Cruise II Central and Cruise IV 

western/Central samples correlated least with grain size. This may be due 

to a substantial petroleum input to the TERR indicator that is independent 

of the river. It is also probable that the seaward distance the material 

is transported and the composition of the transported material vary with 

time. The largest range in clay content was observed during Cruise II 
113 

Table 3-6 . Comparison of sediment hydrocarbon parameters at two different 

topographic settings (for definitions of hydrocarbon 

parameters see Tables 3-3 and 3-q) . 

Depth (m) 

WC-11 

Topo-Hi 

1226 

WC-12 

Topo-Low 

1236 

Total EOM 18 .9 17 .1 

(PPm) 

Aliphatic UCM 81 .E 4 .2 

(P Pm) 

PL-1 266 .6 67 .5 

(PPb) 

TERR 3068 .3 182 .6 

(PPb) 

Petro-Lo 248.2 45 .7 

(PPb) 

Petro-Hi 852 .7 44 " 

(PPb) 

Terrestrial 2233 .7 137 .7 

(PPb) 

Planktonic 18 " 4 21 " 

(PPb) 

Petroleum 1100 .9 90 .6 

(PPb) 
114 

90 

80 

} 70 

Q 

v 60 

50 

40 

30 

0 

0 C1 

O " ~ " -

0 -I 4 t 

p 

W C 3 

O ~ 

~ ~ 

~% 

p C5 O 

o 11111 600 

:5 

o O E2 M 

o 0 

E1 

0 0.2 0.4 0.6 0.8 1 .0 1 .2 

TERR (ppm) 

*Western/Central Cruise IV "Western/Eastern/Central Cruise II 

OCruise I Central # Cruise III Central O Eastern Cruise V 

Figure 3-19 . The relationship between a terrestrial hydrocarbon indicator 

and the clay content of the sediment . 
115 

when the western, Central and Eastern Transacts were sampled (Fig . 3-20) . 

In this case a positive correlation with clay content and a negative 

correlation with sand content is readily apparent ; however, within a given 

transact the correlation is less striking . The correlation of the 

terrigenous indicator with grain size supports the premise that these nalkanes represent riverine or land-derived material . In general the 

relative importance of riverine material between geographical areas can be 

estimated though variability within a given area, i.e ., along a transact, 

can be substantial. 

The petroleum indicators were generally independent of grain size, 

though as previously mentioned some component of the petroleum is related 

to river-associated particles (Fig. 3~-21) . This river influence can be 

seen most clearly on the Eastern Transact Where the PE-Lo indicator 

generally increases with increasing clay content . In contrast the 

aliphatic UCM is apparently independent of clay content (Fig. 3-21), which 

confirms a dual source for petroleum hydrocarbons. Phytoplankton-derived 

hydrocarbons did not correlate with grain size. 

3 .2 .2 .7 Aromatic Hydrocarbons 

Sediment aromatic hydrocarbons were below the detection limit (~5 

ppb) at all locations sampled. The presence of aromatic hydrocarbons at 

low concentrations was inferred by total scanning fluorescence analyses, 

supporting the conclusion that a low level petroleum input is present at 

all locations sampled. 

3 .3 CARBON ISOTOPES 

3 .3 .1 SEDIMENTS 

The carbon isotopic composition of sedimentary organic matter for all 

fine cruises is summarized in Figure 3-22 . Isotopic data confirms the 

previously inferred influence of river-borne terregenous material on the 

Gulf of Mexico continental slope. Though there are numerous complicating 

factors, in general a more negative carbon isotopic composition suggests 

greater land influence. Terrestrially sourced organic material, delta 13C 
116 

80 

0 '~~ 70 %WOO 

z 

UJ 60 

Z 

6 50 

J 40 

U 

30 

c1 

W~ 4 

E C3 ca 

w2 

E5 C5 

W5 

i w1 

E3 

E1 

0.2 0.4 0.6 0.8 

TERR (ppm) 

40 

1 .0 

F- 

.30 

Z 

W 

Z 20 

0 

U 

10 

Q 

El W1 

W5 

t "3 ~ 

E4 0 

E5 

W2 

ca 

W3 W4 

"

o 

~ 

C5 C3 

C1 

0.2 0.4 0.6 0.8 1 .0 

TERR (ppm) 

Figure 3-20 . The relationship between a terrestrial hydrocarbon 

indicator and the clay and sand content of the sediment 

during Cruise II . 
117 

80 

70 

60 

50 

40 

v 

Z 30 

c1 

j4 W 3 

a 

w2 

E5C5 

W5 

E2 

E1 

ca 

i~~ 

z O 

U 

Q 

J 

V s0 

70 

60 

50 

40 

30 

6 10 14 18 22 28 3C 

PE-Lo (ppb) 

c1 

W3 C2 

a 

ca w2 

C5 E5 

W5 

i wi 

E3 

E1 

40 80 120 160 200 240 280 

ALIPHATIC UCM (ppm) 

Figure 3-21 . The relationship between two petroleum hydrocarbon 

indicators and the clay content of the sediment during 

Cruise II . 
118 

Cruise I Central Cruise II Eastern 

z - 

1 

0 

2 

1 

0 

Cruise III Central Cruise IV West/Central 

-25 .5 -21.5 -17 .5 -25.5 -21 .5 -17.5 

to to to to to to 

-26.0 -22 .0 -18.0 -26 .0 -22.0 -18.0. 

b13 C Cy 0o~ 

a 

U 

Z 

W 

../ 

W 

LL o 

Cruise II Western Cruise V Eastern 

2 

i 

0 

3 

0 

s 

3 

1 

0 

Figure 3-22 . Summary of the carbon isotopic composition of sedimentary 

organic matter from all five cruises (613C in 0 /oo, vs . 

Pee Dee Belemite) . 
119 

varies from approximately -25 to -280 0/00, wile planktonic-derived 

carbon delta 13C varies from approximately -16 to -21 000. As can be 

seen the average delta 13C of sedimentary organic material is most 

positive at the eastern stations, most negative at the Central and Western 

sampling sites. This trend suggests an increased influence of planktonic 

material at the eastern sites. In general the average delta 13C suggests 

a substantial planktonie contribution to the bulk organic matter, whereas 

the hydrocarbons are generally dominated by terrestrial sources. 

3 .3 .2 ORGANISMS 

Stable carbon isotopes have been widely used in marine research to 

trace the flow of carbon in ecosystems and the biogeochemical systems 

(Parker 1964, Sackett 1964, Calder and Parker 1968, Haines 1976, DeNiro 

and Epstein 1978, Haines and Montague 1979, Fry and Parker 1979, Fry 1981, 

Gearing et al . 1980 . Several assumptions are inherent in their use 

including (1) a relatively constant isotopic composition of organic carbon 

produced by each plant source, (2) an unchanging isotopic ratio in plant 

carbon as it decomposes and is broken down into detritus, and (3) little 

or no isotopic fractionation between consumer and its carbon supply. 

Gearing et al . (1981) have recently provided an excellent review of the 

carbon isotopic literature relating to these assumptions. 

The first assumption is generally applicable within a few parts per 

thousand for marine plankton . isotopic variability in marine 

phytoplankton has been correlated with species composition, temperature, 

water masses, latitude, and delta 13C of the inorganic carbon that is 

fixed . Little isotopic fraction is generally observed in the aerobic 

decomposition of organic matter in seawater. They observed an overall 

pattern of increasing carbon isotope ratios with trophic levels, 

progressing from diatoms (-20 .3 °/oo), to zooplankton (-19 .8 0/00), 

meiofauna (-19 .5 °/°°), non-carnivorous maerofauna (-18 .6 0/00), and 

benthic predators (-16.6 0/oo) . Similar patterns have been documented in 

other studies and support an isotopic shift of a few parts per thousand to 

more positive values with increasing trophic level . The organism tissue 

isotopic data for all fine cruises is summarized in Figures 3-23 and 3-2u. 
120 

Other Decapods 

U 

Z o 

W 

LU 18 

LL 15 

12 

9 

8 

3 

n-18 

S 

r I °. 

I 

Other Fish 

n"58 

Synaphobranchus or^ egoni 

5 

4 

3 

2 

1 

0 

n=27 

S " J 

I 

I 

I S 

f 

Y f 

<-22 .0 -20 .0 -18 .0 -16.0 to to to 

-19 .5 -17 .5 -15.5 

b' 3 C (9'00 

s 

s 

3 

n=34 

I 

I 

I ti 

I 

I 

I 

Geryon quinquedens 

5 

4 

3 

z 

Figure 3-23 . Summary of the carbon isotopic composition of organism 

tissues by organism type (S13C in O/oo vs . PPB) . 
Cruise II Central 

n=9 ) ;1 

~ 1!', I 

2 

I n 

I 

I 

I 

Cruise II Western 

2 

i 

Cruise II Eastern 

Cruise III Central 

Cruise IV West/Central 

6 

3 

Cruise V Eastern 

N 
122 

Detailed carbon isotope data for organisms for each station sampled may be 

found in Appendix B-3 " 

3 .4 ORGANISM HYDROCARBONS 

Hydrocarbon levels in Gulf of Mexico slope organisms are highly 

variable. Detailed hydrocarbon data for organisms sampled from Cruises IV are found in Appendices B-7, B-8, and B-9. Total resolved alkanes in 

samples from Cruises I, II and III varied from non-detectable to >8000 ppb 

(Figure 3-25) . Due to the variety of organisms and organ types analyzed, 

trends in variations as a function of location, water depth, and time of 

sampling are difficult to discern. The species collected and analyzed 

were highly variable thus making direct comparisons difficult if not 

impossible . No trends in hydrocarbon distributions were evident as a 

function of any taxonomic group of organisms evaluated (3-25) " Of the 

organisms analyzed, decapods had the lowest incidence of hydrocarbon 

occurrence in muscle tissue (Table 3-7) . More than ~75% of the fish 

muscle tissues analyzed contained detectable hydrocarbons. In the 

sampling provided by this study there were no detectable hydrocarbons in 

several species, primarily shrimp (Table 3-8) " The sampling is often 

limited to a single individual and therefore cannot properly represent the 

entire population. 

Tissue hydrocarbons were generally dominated by pristine, n-C17, nC15 and n-C19 (Figs. 3-26 to 3-28)" These hydrocarbons are predominantly 

planktonic in origin . In general n-C16, n-C18 and phytane were also 

present . These may have a petroleum source but due to the low 

concentrations observed their source is not clear. It is also possible 

that this range of n-alkanes has a bacterial source . A few organisms also 

contained measurable amounts of higher molecular weight petroleum and 

plant bioWax hydrocarbons (Table 3-9, Figs. 3-29 and 3-34) " These 

organisms were all collected at the Central Transect except for one shrimp 

at Station E3 during Cruise II that contained primarily petroleum 

hydrocarbons . The presence of the plant biowaxes suggests that these 

organisms have acquired the hydrocarbon signature of the sediments. These 

hydrocarbons presumably reside in sediments and not in the water column at 

these water depths. In two organisms the plant bioWaxes were detected in 
123 

(All Transacts) 

Guise I, II and III 

n=13 

J 

Q 

D 

> > 

O 

Z 

O 

W 

Z 

Cruise i Central 

4-

2- n=27 

0- 

8-

6 

a 

0 

Cruise II Eastern 

Cruise 111 central Cruise II Western 

4 

1 

1 

Guise II Central 

3 

2 

1 

0 

0 . 250 750 1250 1750 2250 2750 >3000 0 250 750 1250 1750 2250 2750>3000 

to to to to to to to to to to to to 

500 1000 1500 2000 2500 3000 500 1000 1500 2000 2500 3000 

TOTAL ALKANES (ppb) 

Geryon quinquedens 

Other Decapods 

Synaphobranchus oregoni 

D Other Fish 

Figure 3-25 . Summary of the total alkane concentration in organism 

tissues for Cruises I, II and III . 
124 

Table 3-7 . Summary of the incidence of detectable hydrocarbons in 

organises muscle tissue (Cruises I-V) . 

No. of No . of 

Type of No. of Individuals Individuals 

Organisms Species Analyzed Hydrocarbon Free 

Fishes 28 80 21 (26 .3%)* 

Eels 20) 17 3 (17 .6) 

Crabs 4 19 11 (57 .9%) 

Shrimp 18 40 25 (62 .50 

*Percent of individuals analyzed . 
125 

Table 3-8 . Species in which no tissue hydrocarbons were detected in 

organism in the present study . 

No . of 

Individuals Type of 

Cruise Station Analyzed Species Organism 

5 E3D 3 Munida valida Shrimp 

E2 E 

E2B 

u WCS 1 Hymenopenaeus robustus Shrimp 

1 C2 1 Acanthephyra armata Shrimp 

1 C3 1 Benthesicumus bartletti Shrimp 

3 C6 1 Nephropsis aculeata Shrimp 

3 C1 1 Parapenaeus longirostris Shrimp 

3 C2 1 Plesionika holthuisi Shrimp 

4 WC6 1 Merluccius albidus Fish 

5 E2D 1 Metanephrops binghami Shrimp 

1 C4 1 Cataetya sp . Fish 

5 E1 1 Pocilopsetta beam Shrimp 

4 WC5 1 Polycheles typhlops Lobster 
126 

80 

Bathypterois quadrifilis 

Muscle 

Cruise II E4 60 

40 

20 

0 

.O 

Q 240 

200 

O 

~ iso 

CC 120 

H 

Z so 

w 

V z ao 

O 

U o 

Urophycis cirratus 

Muscle 

Cruise III C I 

40 

30 

20 

10 

0 

Urophycis floridanus 

Muscle 

Cruise I C 1 

15 17 Pr 18 Ph19 21 23 25 27 29 31 

CARBON NUMBER 

Figure 3-26 . Selected examples of alkane distributions in fish tissues 

(Pr = pristane, Ph = phytane) . 
127 

220 

180 

140 

100 

60 

20 

Geryon quinquedens 

Muscle 

Cruise II E 1 

.fl 

Q. 

p, 400 

v 

Z 

300 

H 

200 

FZ 

W 100 

U 

Z 

O o 

200( 

150( 

1001 

501 

Stereomastis sculpta 

Muscle 

Cruise II C3 

Nematocarcinus rotundus 

Muscle 

Cruise il W3 

15 17 F'f 1 S F'h 1 y 21 23 25 27 29 31 

CARBON NUMBER 

Figure 3-27 . Selected examples of alkane distributions in crab and 

shrimp tissues (Pr = pristane, Ph = phytane) . 
128 

2,500 

2,000 

1,500 

1,000 

500 

O 

Synaphobrancus oregoni 

Liver 

Cruise II C3 

Q- 12,000 

d 

10,000 

Z 

8,000 

H 

6,000 

F - Z 

W 

V 2,000. 

Z 

O 

U 

Synaphobrancus brevidorsalis 

Gonad 

Cruise II E3 

600 

Synaphobrancus ore9oni 

Gonad 

Cruise II W3 

400 

200 

0 

15 17 Pr 18 Ph19 21 23 25 27 29 31 

CARBON NUMBER 

Figure 3-28 . Selected examples of alkane distributions in eel tissues 

(Pr = pristane, Ph = phytane) . 
-129 

Table 3-9 " Species in which a sediment hydrocarbon signature was 

detected in organism tissues . 

Cruise Station Species Organ Type 

III C6 Urophycis cirratus Gonad Fish 

III C6 Urophycis cirratus Liver Fish 

III C10 Geryon quinquedens Gonad Crab 

III CS Geryon quinquedens Muscle Crab 

III C10 Geryon quinquedens Muscle Crab 

III C7 Geryon quinquedens Gonad Crab 

III C1 Steindachneria 

argentia Gonad Fish 

III C8 Coryphaenoides 

mexicanus Muscle Fish 

III E3 Nematocarcinus 

rotundus Muscle Shrimp* 

III C7 Heterocarpus orb Gonad Shrimp 

III C7 Heterocarpus oryx Muscle Shrimmp 

*Petroleum signature . 
130 

Ger on quinguedens 

Gonad 

Cruise III C10 

so 

60 

Coryphaenoides mexicanus 

Muscle 

Cruise III Ci3 

40 

20 

0 

15 17 Pr 18 Ph19 21 23 25 27 29 31 

CARBON NUMBER 

1200 

1000 

800 

600 

400 

200 

0 

Ger on guinquedens 

Muscle 

Cruise III C10 

280 

Q. 

240 

Z 200 

f- 160 

Q 

120 

H 

80 

40 

U o 

Figure 3-29 . Selected examples of terrestrial hydrocarbons detected 

in crab and fish tissues (Pr = pristane, Ph = phytane) . 
131 

800 

600 

400 

200 

0 

1000 

d 

Q 

v 

800 

Z 

600 

Q 

400 

Z 

W 

U 200 

z 

0 

U o 

soo 

600 

0 

Nematocarcinus rotundus 

Muscle 

Cruise II E3 

400 

200 

15 17 Pr 18 Ph 19 21 23 25 27 29 31 

CARBON NUMBER 

Figure 3-30 . Selected examples of terrestrial hydrocarbons detected 

in shrimp and fish tissues (Pr = pristane, Ph = phytane) . 
132 

the liver and/or gonads as well as the muscle tissue. The presence of the 

sediment hydrocarbons in tissues suggest that these organisms either 

ingested sediments and/or other organisms that had previously ingested 

sediments . 

On average the highest alkane concentrations were detected in liver 

and gonad tissues, though there are high hydrocarbon levels present in 

some muscle tissues as well (Fig . 3y31) . A suggested scheme of 

hydrocarbon sources and sinks on the Gulf of Mexico continental slope is 

shown in Figure 3-32 " 
133 

45 

40 

35 

v 30 

W 25 

W 20 

15 

10 

5 

0 500 1000 1500 2000 2500 3000 

TOTAL ALKANE CONCENTRATION (ppb) 

Muscle D Liver " Gonads 

Figure 3-31 . Summary of total alkane concentrations from Cruises I, II 

and III as a function of tissue type of all organisms 

analyzed . 
Vc -16 to -200/. 

. . . . . . 

. . 

Plankton Pelagic Tar 

n-15,17,19 

Pristane 

6"C -16 t0-20 °/oo 

7 

Pelagic Organisms 

9 

n-15 .17,19 

Pristane 

m 

-16 to-19°/~0 

. . . : : : : : DeMer9al Fish 

Runoff 

Biowaxes 

i 

Petroleum 

Biowaxes ' 

6 13c -28 to -38°/. 

-16 to-20°/00 63"s -2 to +5°/. 

-20 to -22o

/oo 

Benthic o~genl ams 

. : . . . .. 

. :`. .Seep Organisms . .,~ 

. . . . . . . . . 

Sediments Biowaxes ~ ' MICROBES 

`, Petroleum` 

---- -------- ----------- ~ 

------------- 

: :Oil : 'CH. . ~HzS 

------------------- 

----------- ' Seepage -45 to -60°ioo 

6"c -27°i. 

6'4s -4°/00 

Planktonic Biogenic Major Biological Pathways H.C.'s 

Terrestrial Biogenic ------ Major Sediment Inputs H.C.'s 

Petroleum Unknown or Minor Pathways 

w 

41 

Figure 3-32 . A suggested scheme for the sources and sinks of hydrocarbons on the Gulf of Mexico slope 

(note : the size of the various boxes do not represent quantitative importance) . 

4 .0 BIOLOGICAL OCEANOGRAPHY OF THE NORTHERN GULF 

OF MEXICO CONTINENTAL SLOPE 
137 

4 .0 BIOLOGICAL OCEANOGRAPHY OF THE NORTHERN GULF 

OF MEXICO CONTINENTAL SLOPE 

The faunal groups discussed in this study include the meiofauna 

(0 .063-0 .3 mm), collected by box corer, the macrofauna (greater than 0.3 

mm in size) also collected by box corer, and the megafauna (both 

invertebrates and demersal fishes), which were collected by trawling. Of 

these groups, only the meiofauna and certain macrofaunal groups, such as 

Nematoda, Harpacticoida, Rinorhyncha, and a few others, were not 

identified to the species level . 

The results of the five cruises discussed in this report suggest that 

it is now possible to predict with a reasonable degree of certainty the 

basic composition of the faunal communities on the northern Gulf slope 

between the 300 and 2500-m isobaths lying between 85° and 94° west 

longitude . Furthermore, it is also evident that there is a reasonable 

degree of agreement between the faunal findings discussed in a previous 

MMS study of the Gulf (Pequegnat 1983) and the present MMS study conducted 

by LGL. This is not to say that significant new discoveries have not been 

made, for they have . For instance, the discovery of the "seep 

communities" in the north-central Gulf goes well beyond the recognition of 

a unique kind of community in that it shows that matter and energy 

reductions can be a mayor factor limiting community developments in the 

deep sea. Also, new species of organisms have been uncovered during every 

cruise . The numbers of new species in such macrofaunal groups as the 

Tanaidacea, Cumacea, and Ectoprocta, to mention but a few, were not 

anticipated. Very likely the same would be true of the meiofauna were we 

to study it intensively . More surprising, perhaps, is the fact that a 

rather large number of new species and first records for species were 

revealed by a careful study of the megafaunal decapods alone. For 

instance, two new species of galatheids (in the genera Munidopsis and 

Phylladiorhynchus ) are being described (L.H. and W .E. Pequegnat), two new 

species of brachyuran crabs in the genus Cymonomus are being described by 

D. Felder, a new pagurid in the genus Parapaxurus is being described by R. 

Lemaitre, and two new species of carideans, viz., Bythocarls sp . A, and 

Palaemonella sp. A are being described by R. Heard. 
138 

The number of species of decapods found for the first time in the 

Gulf or that are considered extremely rare by specialists is too large to 

detail here, but the tally is an impressive one . A complete list of the 

species identified from Cruises I-V from all sampling efforts can be found 

in Volume III of this report series . 

4 .1 MEIOFAUNA 
141 

4 .1 MEIOFAIINA 

4 .1 .1 COLLECTION RESULTS 

Meiofauna samples were taken from the 2 cm (i .d.) coring tubes afixed 

to the sides of the box cores. (See Section 2 .3 .1 .2 for a description of 

box core methods and materials .) The usual sample design was to collect 

duplicate tubes (for a total of 19 .4 em2) from each of six replicate box 

cores at each station. The counts in these tubes were combined and 

considered one replicate sample per box core. Of the planned 60 sampling 

stations, 59 were successfully completed . During Cruise II, three or four 

replicates were collected at the Eastern and Western Transact stations ; 

six replicates were collected on all of the remaining cruises. Appendix 

Table C1-1 shows the sampling design for the entire meiofauna collection 

obtained during the program. 

Meiofauna were defined as metazoan organisms retained on a 63 um 

screen ; counts of protozoan Foraminifera, though noted, have not been 

included in this discussion. This is consistent with the definition 

provided by Theil (1983). A total of 43 mayor groups were identified ; of 

these, representatives of five taxa of permanent meiofauna (Nematoda, 

Harpaeticoidea, Polychaeta, Ostracoda, and Binorhyncha) along with 

naupliar larvae (temporary meiofauna) comprised 98% of the collections. 

Table 4-1 shows the total occurrence of the dominant groups in the entire 

collection . (A detailed tabulation of densities at individual stations 

can be found in Appendix Tables C1-2 through C1-6) The abundanees in 

Table 4-1 have also been presented as densities per 10 cm2 according to 

the total area of the replicates (Theil 1983) ; and as approximations of 

relative wet weight following published estimates of group weights (Rowe, 

et al. 1971, Fauble 1982). The nematodes, harpactacoids, and nauplii were 

numerically dominant throughout the collection and in the mean densities 

per replicate; however, because of their much greater size, the less 

abundant polyehaetes and ostracods probably contributed significant 

biomass. 
N 

Table 4-1 . Overall abundance and estimated biomes of the meiofaunal collection. 

Overall Mean Density Standard Wet Weight Approximate 

Taxa Abundance per 10 sg . em Error Multiplier Wet Weight-ug 

Nematoda 135,167 414 .62 10 .25 0 .85 11,892 

Harpactacoida 41,826 128 .3 3 .11 2 .8 117,113 

Nauplii 30,119 92 .39 2 .75 2 .15 6,756 

Polychaeta 9,648 29 .6 1 .56 55 .25 533,052 

Ostracoda 6,118 18 .77 0 .58 39 .15 239,520 

Kinorhyneh 2,222 6 .82 0 .41 2 .8 6,222 

All other taxa 5,648 0 .91 0 .13 NA 75,730 

Total 230,748 707 .82 16 .2 1,151,283 
143 

4 .1 .2 SEASONAL COMPARISONS 

The occurrence of all permanent meiofauna groups and of naupliar 

meiofauna was compared among transects and cruises by plotting the median 

and quartile values for the density of specimens per replicate (Fig. 4-1) . 

The plots were arranged to show seasonal differences between samplings . 

The median densities of the permanent meiofauna were similar for all of 

the transects with the exception of the Eastern and Western Transect 

samples collected during Cruise II; no seasonal differences were apparent . 

The density of nauplii collected during the spring and summer cruises was 

generally higher than the densities observed in th two fall collections, 

although the Cruise II Western and Eastern Transact collections were 

depressed for this group as well. 

Seasonality in abundance of both permanent and temporary meiofauna 

has been reported for coastal and estuarine collections (Rudnick et al . 

1985) ; however, deep sea collections have generally not shown clear 

seasonal patterns (Theil 1983) . The apparent seasonal differences in 

nauplii densities may also have resulted from differenes in the depth 

ranges of the transacts and individual variability between stations . 

Rigorous comparison of seasonal differences would require a level of 

replication that was not obtained in this study. In the absence of gross 

differences, the meiofauna data presented below were treated as though 

season fluctuations did not exist. 

4 .1 .3 COMPARISON OF MEIOFAIINA WITH MACROFAU NA BY DEPTH 

The occurrence of all meiofaunal groups was compared with the total 

macrofauna collection at the 59 sampling stations . The upper panel of 

Figure 4-2 shows regressions of the log of the numbers of meiofauna and 

macrofauna per square meter plotted against depth (see Theil 1983, pg . 215 

for similar treatment). Both regressions were significantly different 

from zero (p -_ 0 .01) . The density of meiofauna was approximately two 

orders of magnitude greater than the density of macrofauna throughout the 

depth range of the sampling stations. The range in density of meiofauna 

among the sampling stations was approximately one order of magnitude ; the 

range of macrofaunal densities was somewhat narrower. Densities of both 
144 

Cr I Cr I I Cr IV Cr Y Cr III 

U 

.= 2000 n 

L 

n 

1000 

E 

z 

0 

Permanent meiofeuna 

C 

G 

C 

E o 

W 

E 

C 

0 

Nov/Dec Jan/Feb Mar/Apr May/Jun Jul/Aug Sep/Oct Nov/Dec 

Cr I Cr I I Cr IV Cr V Cr III 

°' 300 

U 

n 

200 L 

N 

d 

L 

E 100 

z 

0 

Nauplii meiofauna 0 

WC 

C 

C 

0 

C 

w 

E 

Nov/Dec Jan/Feb Mar/Apr May/Jun Jul/Aug Sep/Oct Nov/Dec 

Figure 4-1 . Comparison of median numbers of permanent and nauplii meiofauna arranged by season collected . The boxes show the range 

of the 25th to 75th percentiles ; the median is indicated by 

the heavy line . Whiskers are 1.5 times the range of the boxes . 
145 

7 

y = 5.87 - 0 .000183x 

. , 

- ~ 

M eiotauna Q s 

. " 

" N L 

C 4 

y = 3.52 - 0.000109x 

Z 

13 CP 

1923 O UO p ~ O 

° o a o 0 

o 0 Macrofaune 0 

2 

0 500 1000 1500 2000 2500 3000 

8 

t 

v> 

a> 

'4 

a M e i o f 8 U 118 y = 6 .4400 - 0.000200x % 

6 

-4 4 

o q 

°° °' 

~ 

4 

Q 

-47 D E ° ~ o ° 

x 

v> 

o 

o 

o a o 

° L ^ 5 Macrofeuna 0 o 

a y = 5.88 - 0.000227x 

a 

4 

0 500 1000 1500 2000 2500 3000 

Depth (m) 

Figure 4-2. Relative occurrence of meiofauna and macrofauna with depth . 

at all sampling stations . Upper plot shows log of density per 

square meter . Lower plot shows log of approximate biomass per 

square meter . 
146 

meiofauna and macrofauna decreased with increasing depth . The rate of 

decrease was slightly less for the macrofauna than for the meiofauna; 

however, analysis of covariance would be required to test the difference 

between the two slopes. The regression plots in Figure 4-2 suggest that 

there was a threefold decrease in the density of meiofauna and a twofold 

decrease in the density of macrofauna between 300 and 3000 m depths. 

The lower panel of Figure 4-2 shows regressions of the log of 

approximate wet weight for meiofauna and macrofauna. Approximate wet 

weights per m2 were calculated by multiplying the densities of each of 43 

meiofaunal and 44 macrofaunal groups, respectively, by appropriate 

estimates of wet weight per individual (Rowe et al . 197 4 , Fauble 1982) . 

As with density, the approximate wet weight of both groups decreased with 

increasing depth . In this case, however, the rate of decrease for 

macrofauna was slightly greater than for meiofauna. The regression plots 

in Figure 4-2 suggest that there was a three-and-one-half-fold decrease in 

the biomass of meiofauna and a fourfold decrease in the biomass of 

maerofauna between 300 and 3000 m depths . 

4 .1 .4 CHANGES IN DENSITY WITH DEPTH 

The variation in density of all meiofaunal groups among replicated 

samples was compared in plots of mean density (individuals/10 em2) for the 

sampling transects in the western, central, and eastern sections of the 

study area (Figs. u-3 through 4-6). The comparison showed that there were 

some significant differences (p = 0 .05) among stations within transects. 

In general, the shallowest stations had significantly higher densities 

than the deepest stations; however, no clear pattern of significant change 

was evident. 

On Cruise I, stations on the Central Transact those deeper than 1000 

m in depth had significantly lower meiofauna densities than those 

shallower than 1000 m depth (Fig. 4-3) " This pattern was not repeated 

among the Cruise II Central Transact stations but a decrease was 

suggested . Densities observed on the Central Transact during Cruise III 

showed a high level of variability and stations at 492, 1021, 1465, and 
147 

900 

E 

U 

c~ 700 

h 

h 500 

W 

7 

300 

c 

100 

0 

900 

E 

V 

P 700 

A 

O 

h 500 

a 

.~ 300 

v 

c 

100 

0 

900 

E 

U 

h 700 

0 

500 0 

v 

300 

c 

100 

0 500 1000 1500 2000 2500 3000 

Depth (m) 

Figure 4-3 . Mean densities of all meiofaunal groups collected at Central 

Transect stations . Bars show -F2 standard errors of the means, 

500 1000 1500 2000 2500 3000 

500 1000 1500 2000 2500 3000 
148 

600 

500 V 

P 

h 

O 

400 

300 

v 

c 

200 

100 

0 

Depth (m) 

Et a E1 E1b E1c 

Cruise IV 

600 

E2a E2b E2 E2c E2d E2e 

E3a E3d E3c E3 E3b 

500 

U 

400 

300 

c 

200 

100 

350 620 850 

Sampling 13obath3 ( m) 

E5 

3000 

Figure 4-4 . Mean densities of all meiofaunal groups collected at Eastern 

Transect stations . Bars show + standard errors of the means. 

The Cruise IV stations are arranged by sampling isobath to 

show differences between stations at the same approximate depths . 

1000 2000 3000 
E 

U 

Q 

h 

O 

Il 

A 

7 

V 

.; 

V 

C 

U 

O 

W 

7 

. ; 

V 

C 

500 m laobath 

1 

93' 92° 91° 90' 

Longitude (degrees) 

500 

E 

V 

" 400 

h 

300 

.; 

C 

93* 90° 

E 

U 

Q 

0 

h 

W 

7 

C 

. ; 

V 

C 

Figure 4-5 . Mean densities of all meiofaunal groups collected at Western-Central Transect stations . Bars 

show +2 standard errors of the means . Stations are arranged by sampling isobath to show dif- 1.0 ferences between stations at the same approximate depths . 

93° 92° 91' 90' 

Longitude (degrees) 

93° 92° 91° 90° 

Longitude (degrees) 

92° 91° 

Longitude (degrees) 
150 

450 

E 

350 

P 

h 

O 

250 

150 

c 

50 

Depth (m) 

Figure 4-6. Mean densities of all meiofaunal groups collected at Cruise 

II, Western Transect stations . Bars show +2 standard errors 

of the means . 

0 1000 2000 3000 
151 

2518 m depth (Stations C6, C7, C4, and C5, respectively) were anomalous 

with the trend of decreasing density with increasing depth. 

There were no significant differences in meiofaunal densities among 

the Cruise II Eastern Transect stations (Fig. 4-4) . Sampling the 350, 

620, 850, and 3000 m isobaths during Cruise IV indicated, with one 

exception, that the three shallower depths had significantly greater 

meiofaunal densities than the 3000 m station. Although there were some 

significant differences within the isobaths as well as between the 

isobaths . Densities observed during Cruise II were generally markedly 

lower than during Cruise IV. 

The sample collection from the Western-Central Transeet during Cruise 

V (Fig . 4-5) compared densities within depth isobaths over distances of 

approximately 50 to 100 nm . Although mean densities varied along this 

transect, they were not significantly different within isobaths, nor were 

there apparent trends in the means . 

Meiofaunal densities in the Cruise II Western Transact stations were 

similar to the Cruise II Central Transact stations (Fig . 4-6) . The 

densities observed at the deepest station were significantly different 

from the shallowest station. There were no significant differences 

between the stations at intermediate depths . 

4 .1 .5 VARIATION IN COMPOSITION WITH DEPTH 

Mean densities of the six most abundant meiofauna groups at 

individual stations were plotted for Cruises I-V (these will be shown in 

Figs . 4-7 through 4-11) . Numerical dominance of the nematodes and 

harpaeticoids was apparent throughout the collections ; however, 

accompanying plots of approximate wet weight were provided as an 

indication of differences in biomass . As expected, the polychaetes and 

ostracods Were the dominants with respect to weight. The densities of the 

six dominant meiofauna groups declined overall with increasing depth. The 

trend was not distinct, however, nor were there consistent patterns in the 

changes in density for the individual groups . Patterns apparent in 

changes in density were often reversed when the occurrence of meiofauna 

was expressed as approximate wet weight . 
152 

4 .1 .5 .1 Cruise I 

On Cruise I, the density of nematodes declined only slightly with 

depth with even a slight increase in density going from Station C4 to 

Station C5, the deepest station (Fig. 4-7) . Changes in density were most 

apparent as a decline in the abundance of harpacticoids and nauplii . 

Abundance of polychaetes, ostracods, and kinorhynes decreased as well, 

although this decrease was poorly resolved when plotted as density. The 

approximations of wet weight, however, showed a much sharper decline with 

depth. Decrease in the larger but less numerous polychaetes and ostracods 

was readily apparent when expressed as weight, while the contribution of 

the nematodes was more or less constant . 

4 .1 .5 .2 Cruise II 

Differences in meiofaunal composition between the Western, Central, 

and Eastern Transeets were clearly shown in the plots for the Cruise II 

stations (Fig. 4-8) . Sampling effort was not equal among the transects; 

three replicates were collected at Western Transact stations, six at 

Central Transect stations, and three or four replicates at the Eastern 

Transact stations. However, the differences in effort do not explain the 

substantial differences between mean densities at stations of similar 

depths shown in Figure 4-8. The Central Transact station densities were 

consistently greater than the Western Transact stations of similar depths 

by a factor of two or greater. The density of nematodes was particularly 

noteworthy at Station C1, which also provided the highest overall 

densities of meiofauna seen during the five cruises. 

Decline in both density and approximate wet weight occurred with 

increasing depth in the Central Transact stations . This decline with 

depth was less apparent in the Western Transact stations and did not occur 

in the Eastern Transact stations. The occurrence of meiofauna decreased 

sharply between Stations K1 and W2, both in terms of density and biom ass; 

however, the subsequent stations showed little decrease with depth. On 

the Eastern Transeet stations, the overall density remained generally 

constant with increasing depth except for a slight decrease in the density 

of nematodes that registered at Station E4. When expressed as approximate 
153 

500 

400 

U 

Q 3 

h 

O 

200 

D v 

v 

100 

0 

Station 

2000 

.~ 

1500 

a> 

U 

q 

O 

CD 1000 

.~ 

v% 

500 

0 

Depth (m) 

Kinorhyncha 

p Ostracoda 

Polychaeta 

NAUnlii 

Figure 4-7 . Densities and approximate wet weights for the six most abundant 

meiofaunal groups collected during Cruise I, Central Transect . 

C1 C2 C3 C4 C5 

344 511 852 1375 2475 
154 

700 

600 

500 

Q 

400 

a) -Z 300 

D 

v 

200 

v c 

100 

0 

Station 

3000 

,., 2500 

t 

E 

2000 

c 

.r e 

O 

1500 

a) 

1000 

X 

L y 

a a ~ 500 

a 

0 

Depth (m) 

Ki norhyncha 

Q Ostracoda 

Polychaeta 

Nauplii 

Wornnn+inniiln 

Figure 4-8. Densities and approximate wet weights for the six most abundant 

meiofaunal groups collected during Cruise II, Western, Central, 

and Eastern Transects . 

W1 W2 W3 W4 W5 C1 C2 C3 C4 C5 E1 E2 E3 E4 E5 

359 604 854 1410 2506 354 598 838 1390 2389 354 627 846 1350 2827 
155 

wet weight, the occurrence of meiofauna on the Eastern Transact stations 

actually increased with depth between Stations E1 and E3 (354 to 846 m 

depth). This increease was due to increased abundance of polyehaetes at 

Stations E2 and E3. 

4 .1 .5 .3 Cruise III 

Occurrence of meiofauna on Cruise III (Central Transact) stations was 

variable across the depth gradient sampled (Fig . 4-9) . Densities of 

meiofauna decreased, in general, between the shallow and deep stations; 

however, noteworthy exceptions to this trend occurred at Stations C7, C4, 

and C5 . The reversals were primarily due to increased abundance of 

nematodes, although the abundances of the other mayor groups were also 

elevated in comparison with adjacent stations. When density was expressed 

in terms of approximate wet weight the predominance of polychaetes at 

Station C7 became evident. 

Comparison of meiofauna occurrence at Cruise III stations with the 

Central Transact collections made during Cruises I and II (Figs . 4-7 

through 4-9) showed the relative composition of the six mayor groups to 

have been mostly constant between samplings. Nematodes and harpacticoids 

were the most abundant groups at all stations. Nauplii were more abundant 

than polychaetes except at Station C7 on Cruise III . Ostracods and 

kinorhyneh were the least abundant groups at all stations. There was also 

good agreement among the stations that were sampled on each of the three 

Central Transact cruises (C1-C5), although elevated abundance, 

particularly of nematodes, was observed during the Cruise II sampling. 

4 .1 .5 .4 Cruise IV 

The Cruise IV sampling returned to Stations E1 , E2, E3, and E5 that 

were sampled during Cruise II. In addition, more stations in the same 

respective isobaths as E1, E2, and E3 were also sampled . Figure 4-10 

shows the isobathie stations arranged northwest to southeast. Occurrence 

of the nematodes across this station array was variable; however, there 

was no trend of abundance either with increasing depth or with longitude 

within the three sets of shallower stations . Station E5 (2901 m depth) 
156 

500 

400 

U 

P 

a o 300 

p ~ 200 

v .j 

v 

c 

- 100 

0 

Station 

4000 

++ 3500 t 

~ E 3000 

y U 

Q 

2500 

c) ~ 2000 

'v1500 

"X c 

1000 

n 

n 

Q 500 

0 

Depth (m) 

Ki norhyncha 

Q Ostracoda 

Polychaeta 

Nauplii 

da 

Figure 4-9 . Densities and approximate wet weights for the six most abundant 

meiofaunal groups collected during Cruise III, Central Transect . 

C1 C6 C2 C3 C7 C8 C9 C4 C11 C5 C12 

357 492 633 881 1021 1192 1430 1465 2101 2518 2945 
157 

600 

500 

400 

a o 

300 

°) 

v 200 

v 

c 

100 

0 

Station (NW to SE) 

3000 

.~ 

t 

2500 

E 

i ~ 2000 

P 

N O 

1500 

v1000 

n 

n 

a 

500 

0 

Depth (m) 

Kinorhyncha 

Q Ostracoda 

Polychaeta 

Nauplii 

Harpacticoida 

Figure 4-10 . Densities and approximate wet weights for the six most abundant meiofaunal groups collected during Cruise IV, Eastern 

Transect . 

E1 A E1 EtB PC E2 A E2B E2 E2C E2D E2E E3 A E3D E3C E3 E3B E5 

352 353 346 354 624 627 621 619 630 622 856 849 848 819 860 2902 
158 

had the lowest abundance of nematodes and all other groups. The abundance 

of nematodes was generally lower in the 620 m isobath stations (E2A-E2 E) 

than it was in the 350 m and 850 m isobath stations (E1A-E1C and E3A-E3D) . 

In contrast with the Central Transect samplings, harpacticoids were 

generally less abundant than nauplii in the Cruise IV collection. The 

plots of approximate wet weight showed a greater representation by 

ostracods than was seen in the Central Transeet stations. Stations E1 and 

E1C had the greatest abundance of ostracods seen during the project . 

Abundance of meiofauna was substantially greater for the shallow- and 

intermediate-depth stations (350 to 850 m) sampled during Cruise IV than 

for the samples taken during Cruise II. The deepest station, E5, had 

similar abundances during both cruises. The differences in abundanees 

between Cruises IV and II were reflected in all six mayor groups to a 

similar degree . 

4 .1 .5 .5 Cruise V 

The station array in Cruise V allows comparison of meiofauna 

abundance along isobaths between the Western and Central Transacts (Fig. 

4-11) . No distinct trends in abundance were evident, either with changes 

in depth or longitude, nor were any of the collections from individual 

stations anomalously dense or depauperate. The occurrence of nauplii and 

ostracods was greater in the Western-Central Transact than in either the 

Western or Central Transacts. Overall occurrence of meiofauna was less 

than the Central Transact collections, but substantially greater than the 

West Transact collection. 

4 .1 .6 DISCUSSION OF COMMUNITY ASPECTS 

The decrease in occurrence of meiofauna with increasing depth has 

been reported in the Gulf of Mexico by Rowe (1971, 197) and elsewhere by 

Theil (1983)" The intercept and the slope of the regression of meiofaunal 

densities (see Fig . 4-2) agreed remarkably well with the decrease of 

meiofauna from different eastern Atlantic regions reported by Theil (1983 ; 

pg. 215) . (Note that his description includes reports of densities from 

stations at 4000 m and 5000 m depths in addition to the slope depths.) 
159 

600 

500 

V 400 

P 

h 

O 

300 

d W 

0 200 . ; 

C 

0 

Station 

2000 

.^ 

a> 

3 

1500 

o 

,, .. 

1000 

v+ 

X 

O 

L 

d 

soo 

0 

Depth (m) 

Kinorhyncha 

Q Ostracoda 

Polychaeta 

Nauplii 

Harpacticoida 

Nematode 

Figure 4-11 . Densities and approximate wet weights for the six most abundant meiofaunal groups collected during Cruise V, WesternCentral Transect . 

WC1 WC5 WC2 WC4 WC6 WC7 WC8 WC3 WC9 WC10 WC11 WC12 

339 298 550 547 563 453 547 750 759 748 1226 1236 
160 

Theil also compares occurrence of macrof auna in the northern Gulf of 

Mexico (data were obtained from Route and Menzel 1971) with the Eastern 

Atlantic meiofauna densities. He reports that the ratio of meiofauna to 

macrofauna increases kith depth. One explanation for this change would be 

an overall decrease in the size of infaunal organisms with depth. 

This hypothesis was not supported by the current findings. As Figure 

u-2 shows, the ratio of numbers of meiofauna to numbers of macrofauna on 

the northern Gulf slope actually decreases with increasing depth . 

Approximations of biomass (Fig . 4-2) also showed a relative increase in 

macrofauna, although it was less pronounced than for density . This may 

indicate that the macrofauna as a group decreased in size with depth. 

Trends in relative abundance of meiofauna and macrofauna, and in 

overall infaunal biomass may have been influenced by the representation of 

individual taxonomic groups. A cursory look at four groups consistently 

found in both the meiofauna and macrofauna (Nematoda, Harpaeticoida, 

Polyehaeta, and Ostracoda) indicated that the relative numbers of 

meiofaunal and maerofaunal representatives for each group undergo 

consistent changes with depth (Fig. u-12) . These patterns were distinct 

for each group. For example, the number of macrofaunal nematodes 

increases or remains constant with depth, while meiofaunal nematodes 

decrease with depth. Among the Harpacticoida, the densities of meiofauna 

generally decreased with depth, while the macrofauna shouted the greatest 

densities at intermediate depths. Meiofaunal and macrofaunal polychaetes 

shouted similar changes with depth, while maerofaunal ostracods shouted an 

initial increase in density with depth. Because densities of nematodes 

were high, an increase in maerofaunal nematodes would increase the biomass 

of macrofauna relative to the biomass of meiofauna. A detailed study of 

these data might yield additional information concerning the relative 

occurrence of meiofauna and maerofauna. 

Variability among replicate samples at a station and between stations 

at similar depths is clear from the plots shown in Figures 4-3 through 

4-6 . These data suggest that the density of meiofauna was not simply a 

function of depth . Consider the collections from the Central Transact 

(Fig. 4-3) . This transact was sampled three times during two seasons; the 

tread of decreasing density with depth was apparent in all samplings. 

However, differences between Cruise II densities and those seen in Cruises 
Cruise I 

E280 1.6 U 

O 

C 

7 O 

O .- 

O \Z200'9 

L 

d 

E 

Z160 01 

,E , 140 0.5 

o v 

c m 

o O 

90 0.3 

d 

D 

E 

40 0. 1 

E 20 1 .6 

u 

c 

O y 

10 1.1 

E n 

E 

z 0 0.6 

12 0.70 

o Q 

C y 

O O 

w 

~ 7 0.35 

L y 

v 

z z 0.00 

0 1000 2000 3000 

Heiofauna and Hacrofauna 

Central Transect 

Cruise II 

500 

300 1 .0 

100 0 .6 

120 1 .0 

80 ~ 0 .5 

40 0 .0 

30 3 .0 

is t .s 

0 0.0 

12 0 .6 

7 0 .3 

2 0 .0 

0 1000 2000 3000 

Depth (m) 

b- Mecrofeune 

Cruise III ---- Metofeune 

400 3.0 

C 

7 

O 

200 

U 

V 

0 ~6 0.0 

100 0.4 

0 

c 

7 

60 0.2 L 

U 

O 

L 

20 0.0 

60 3.0 

0 

C 

0 

30 1 .5 L 

U 

E 

0 I , 0 .0 

16 0.4 

\\l C 

9 f 0.2 

I 

i ~ ," " F 0.0 

0 1000 2000 3000 

Nematode 

Harpacticoida 

Polychaeta 

OStf'GCOda 

Figure 4-12 . Comparison of macrofauna and meiofauna densities for four major infaunal groups . Meiofauna all densities are one to two orders of magnitude greater than macrofauna densities . 
162 

I and III are as great as the down-slope differences . Moreover, during 

Cruise III, which sampled the slope at finer depth-intervals, significant 

exceptions to the trend of decreasing density were observed (depths of 

1021, 165, and 2518 m; Stations C7, C4, and C5, respectively) . 

Sediments from two of these stations, C7 and C5, had markedly lowered 

values of the carbon preference index (see Section 3 .2 .2 .3, this volume) 

indicating hydrocarbon enrichment in the surface sediments. Benthie 

photographs at Station C7 revealed a dense community of chemosynthetic 

clams in the surface sediments at this location (Rosman et al. in press) . 

Other investigations of Gulf of Mexico meiofaunal densities have shown 

that localized chemical enrichment of surface sediments can provide the 

basis for a chemosynthetic food-web that causes a general increase in 

inPaunal density and diversity (PoWell et al . 1986) . Similar processes 

seem to have been observed during the Cruise III sampling. 

Comparison of the Eastern Transact collection from Cruise IV (Fig . 

4-4) shows that the variability within isobaths can be as great as the 

differences between isobaths . Although the depth range of the Eastern 

Transact stations was identical to that of the Central Transact stations, 

the decreasing trend is much less clear in the Eastern Transact 

collections. The primary differences between these two transacts were 

variability in sediment grain size, steepness of the slope, and the 

presence of hydrocarbon seeps on the Central Transact. The slope of the 

Central Transact was more or less constant between 300 and 3000 m depth; 

sediments were predominantly clay, becoming gradually coarser with 

increasing depth. The slope of the Eastern Transact was quite gradual to 

a depth of 1000 m and then dropped abruptly at the edge of the Florida 

Escarpment ; sediments were variable, but generally coarser than on the 

Central Transact . The variation in sediment grain size within the Cruise 

IV sampling isobaths was as great as was seen over the entire depth range 

on the Central Transact (see Section 3.2) . The exact relationship between 

sediment grain size and meiofauna abundance is not defined by this 

comparison ; however, variability in grain size within isobaths was 

accompanied by variable meiofauna abundance. 

Comparison of the composition of the meiofauna (Figs . 4-7 through 4-

11) shows that although the ranking of the six most abundant groups 

generally remained constant, there was substantial variation in their 
163 

relative abundance. Sediment characteristics may influence the 

composition of the meiofauna in predictable ways. Studies in coastal 

habitats have indicated that sediment grain size influences the relative 

abundance of nematodes and harpaeticoids (Tietjen 1969, Coull 1970) . 

Because these were the most abundant groups found in the present 

collections, changes in their relative density would influence the overall 

meiofaunal density. Figure 4-13 shows a comparison of the abundance of 

nematodes and harpacticoids plotted against percent sand and percent clay. 

Linear regression of these points shows that the abundance of both groups 

decreased as the sediments became coarser, and increased as they became 

finer. The rate of change was more acute for the nematodes than for the 

harpacticoids, particularly in the case of increasingly sandy sediments. 

The fit of these regressions are poor, however, which suggests that other 

factors also influenced the composition of the meiofauna. More powerful 

statistical techniques, such as principal component analysis, may prove 

useful for analysis of these data. 
164 

Meiofauna 

Abundance vs Sediments 

1500 

y = 463.651 - 4 .342x R = 0 .23 

1000 

N 

la 

U 

500 

Q 

o 

1500 

.j 

C 

. .,. 

y = 150.731 - 1 .991 x R = 0.35 

%I--

0 1000 

N 

z 

500 

0 

: .~ . :.' 

7,- .. .w.° ~.~.. 

0 10 20 30 

Percent Sand 

40 25 45 65 85 

Percent Clay 

y = 241 .247 + 2.646x R = 0 .17 

. . :~; 

- ~ . . .~ ; 

y = 8.687 + 1 .g26x R = 0.38 

R 

O 

lc~ 

C 

N 

Z 

A 

O 

V 

U 

ld 

a 

l0 

2 

Figure 4-13 . Abundance of Nematoda and Harpacticoida plotted against 

percent sand and clay . 

4.2 MACROFAUNA 
167 

4 .2 MACROFAONA 

4 .2 .1 INTRODUCTION 

Macrofauna was defined for the purposes of this study as those 

organisms collected with box corers and retained on a 0 .3 mm sieve and 

thus represent much smaller sizes than one typically finds in other 

macrofauna samplings which have used 0 .5 mm or 0 .42 mm sieves . Three 

general sampling strategies were employed to examine macrofaunal 

distributions : 

1 . Cruises I through III sampled down-slope (across isobaths) 

in three different areas (central, eastern, and western 

transects) and two seasons, in order to study depthrelated changes in macrofauna in various parts of the 

Gulf; 

2 . Cruise IV sampled parallel to isobaths of the eastern 

transact, in order to focus upon variability independent 

of depth ; and 

3 . Cruise V also sampled along isobaths with emphasis placed 

on areas of special biological interest in the area 

between the western and central transacts. 

Samples from the macrofaunal component of box cores contained 18 

phyla, which were separated into 1569 differentiable taxa. Specimens were 

routinely identified to the lowest possible taxon, except for those 

belonging to four mayor groups in which severe taxonomic difficulties were 

encountered : Nematoda, Copepoda (primarily harpacticoid copepods), 

Aplacophora, and Scyphozoa. 

Nematodes and copepods, although often retained in large numbers in 

the macroPaunal sieve, more appropriately should be treated with the 

meiofauna. In this report we describe the nematode and copepod densities 

contained within the macrofaunal collections but do not include their 

numbers in the overall density or diversity estimates . Neither do we 

attempt to identify them below nematode or copepod designations . 
168 

Most scyphozoans were collected in the strobila stage . Taxonomic 

keys for the strobila of deep-sea scyphozoans are virtually nonexistent. 

Aplacophorans require complex preparatory procedures before they can be 

examined properly, and are poorly described at best . Balancing the 

potential costs against the chances of successful identification of lower 

taxa within these groups of organisms, LGL and M MS decided to allocate the 

project's resources in favor of more tractable groups. 

Species-level identification was attempted for the remaining groups 

of macrofauna. For these groups, approximately 89% of the individuals 

were identified either to genus or to species ; 71% of the taxa were 

identified to species. Specimens that could not be identified to species 

were usually either a) juveniles that had not yet developed the requisite 

taxonomic characteristics, or b) specimens that no longer possessed in an 

intact form the morphological structures necessary for positive 

identification. Loss through autotomy or damage to some structures (e.g. 

microscopic appendages on crustaceans) is an unavoidable consequence of 

collection and preservation for some of the more delicate species, no 

matter how carefully the samples are handled. The small sizes of the 

macrofaunal specimens from this study were remarked upon by most of the 

taxonomic specialists, making identification to the lower taxonomic levels 

much more difficult because in most cases the small sizes represented 

immature specimens . 

4 .2 .2 COMMUNITY STRUCTURE 

Table 4-2 lists the 40 major macrofaunal groups in decreasing order 

of total numerical abundance over all five cruises, for all stations 

together. It also gives information on number of taxa identified in each 

group, and relative numbers of taxa identified to the specific, generic, 

and higher taxonomic levels. 

Specimens belonging to the first 20 groups accounted for 99% of the 

total number of individuals collected. Six taxonomic groups (Polyehaeta, 

Ostracoda, Bivalvia, Tanaidacea, Bryozoa, and Isopoda) accounted for 86¢ 

of the total numerical abundance . Counts of bryozoans mainly included 

single, more-or-less intact colonies . An attempt was made to avoid 

overestimating bryozoan abundance by not counting multiple, broken pieces 
169 

Table 4-2 . Relative abundance of major macrofaunal groups . 

Total No . No . Iden . 

No . Individual No . Iden . No . Idea . To Family Or 

Taxonomic group Individuals Forms To Species To Genus Higher Taxa 

POLYCHAETA 24313 626 414 163 49 

OSTRACQDA 4960 19 18 1 0 

BIYALYIA 3645 55 41 10 4 

TANAIDACEA 3610 186 168 13 5 

BRYOZQA 3049 99 82 12 5 

ISOPODA 2327 133 119 8 6 

AMPHIPODA 1285 79 50 11 18 

APLACOPtORA 886 1 0 0 1 

NEMERTEA 630 21 20 0 1 

OPHI UR01 DEA 603 17 13 3 1 

SIPUNCULA 570 37 31 3 3 

CUWEA 521 86 76 8 2 

PORIFERA 424 39 22 10 7 

SCAP'HOPODA 382 10 5 2 3 

SCYPtOZOA 331 1 0 0 1 

GASTROPODA 276 53 8 27 18 

HOLOTHUR01 DEA 250 13 4 4 5 

OLIG4CHAETA 247 9 6 1 2 

ASCI DIACEA 136 18 1t 3 4 

HYDROZOA 103 15 8 3 4 

BRACHIOPODA 79 2 2 0 0 

ARACHNI DA:ACARI hIA 62 1 0 0 1 

KINORHYNCHA 60 3 0 2 1 

ECNIt+OIDEA 49 6 3 1 2 

PRIAPULIDA 33 1 0 0 1 

SCLERACTINEA 33 6 4 1 1 

DECAPODA 25 13 10 1 2 

MYSTACOCARIDA 12 1 0 0 1 

POGONOPHORAfYESTIMENT . 11 1 0 0 1 

EGHIURA 9 1 0 0 1 

ACTINIARIA 8 3 0 0 3 

ALCYOhARIA S . 2 1 0 1 

TURBELLARIA 8 1 0 0 1 

CRINOIDEA 7 2 Z 0 0 

PYCNOGONIDA 7 3 3 0 0 

NEMICHORDATA 4 1 0 0 1 

ASTEROIDEA 2 2 0 1 1 

MISC . ANTHOZOA 2 1 0 0 1 

MYSIDACEA 2 1 0 D 1 

ARCHIANNELIDA 1 1 0 0 1 

Total 48970 1569 1121 288 160 
170 

of colonies. At the species level, most macrofaunal taxa were represented 

by very few individuals (Fig. u-14) . For example, over 550 species, 

excluding the polychaetes, were collected five or fewer times (only 1 to 5 

individuals total, for all samples combined) . 

Table 4-3 lists in rank order the dominant 50 species occurring at 

all macrofaunal stations combined . The list includes 28 species of 

polychaetes, six bivalve mollusks, three species of ostracods, two 

isopods, two amphipods, one sponge, and one nemertean worm. 

4 .2 .3 MACROFAUNA ABUNDANCE PATTERNS 

Based upon preliminary calculations, the overall density levels of 

maerofauna among study area sampling stations (not including nematodes, 

non-myodocopan ostracods and copepods, considered meiofauna) ranged from 

about 518 specimens/m2 (Station C12, Cruise III) to 5369 specimens/m2 

(Station E2E, Cruise IV) . As noted, these overall density estimates are 

preliminary. Other than excluding some obvious meiofaunal forms retained 

on occasion in the macrofauna sieve, we have not, as yet, examined the 

collection lists and deleted other forms that should not, by convention, 

be included in our maerofauna designation. These changes, once made, are 

not expected to bane any detectable effects on the overall abundance 

trends . 

The overall region, season, year by depth patterns of macrofaunal 

abundance are shown by Figure u-15. From inspection, abundance appears 

somewhat higher on the Central Transect (a=3156/m2) than on either the 

Eastern (x.2695/ m2) or Western Transacts (g=2100/ m2) . Based upon the data 

from the Central Transact, spring abundance levels (3156/m2) appeared 

higher than fall abundance levels (x ranged from 1657 to 1987/m2) " The 

annual differences between the fall collections of 1983 and 1984 on the 

Central Transact and the spring collections of 1984 and 1985 on the 

Eastern Transact were 330 and 169/m2, respectively. This compares to 

regional differences ranging between 461 and 1056/m2, and seasonal 

differences ranging from 1169 to 1 4 99/m2 . Annual differences appear less 

than regional and seasonal variation in abundance. 

On both the Eastern and Western Transacts, an overall decline of 

macrofaunal density with depth is clearly indicated, even though there are 
171 

soo 

Qoo 

M 

v 

w a 

0 

w 

ion 

0 

o soo Qoo 90o zoo 

NUMBER OF INDIVIDUALS 

Figure 4-14 . Plot of number of macrofaunal species versus number of 

individuals showing the great number of species that 

were collected few times compared to the fewer species 

that occurred in large numbers . 
172 

Table 4-3. Rank order of dominant macrofaunal species . 

OVERALL 

DENSI Y N0. OF N0. OF DEPTH 

SPECIES GROUP no INDIVIDUALS STATIONS RANGE m 

Iblella monniotae Bryozoa 112 1772 27 298-2901 

r tocorsa antennata Polychaeta : Pilargidae 78 1209 30 298- 881 

rows io ranc ata Polychaeta : Spionidae 77 1190 49 298-2827 

ane sp . Polychaeta : Maldanidae 46 715 15 298-1465 

ode ic~ra sp . Polychaeta : Paraonidae 44 679 53 298-2945 

ach t ane sp . A Polychaeta : Opheliidae 37 571 51 298-2518 

o apes ~ er ~ kele,~ orum Polychaeta : Spionidae 33 505 52 298-2901 

o9one sp. A" Polychaeta : Syllidae 30 467 53 346-2945 

Aricidea suecica Polychaeta : Paraonidae 27 421 54 298-2945 

oe sp. 

CII Polychaeta : Sigalionidae 27 417 29 346-1021 

A~~e "sp. A" Polychaeta : Maldanidae 26 402 16 298-1465 

onos io ehlersi Polychaeta : Spionidae 24 380 37 344-1465 

ev nsen a graciiis Polychaeta : Paraonidae 24 371 53 298-2945 

u~lan~e a~es sp. A Crustacea : Ostracoda 21 332 23 453- 860 

u ame ranchia sp. F Molluscs : Bivalvia 20 

19 

319 

299 

34 

52 

298- 860 

298-2945 Pseudotanais sp. 1 

mama cavalieri 

Crustacea : 

Bryozoa 

Tanaidacea 

18 289 37 298-2506 

auve o s ssp.-B Polychaeta : Fauveliopsidae 18 283 49 298-2945 

u orostrum sp . A Crustacea : Ostracoda 18 

18 

279 

276 

44 

49 

298-1465 

298-1465 rsonu s artmanae Polychaeta : Onuphidae 

p, Bryozoa 17 273 22 353-2945 

onos io cirrifera Polychaeta : Spionidae 18 272 30 298-2506 

Ksp. 

R s E 

Molluscs : 

Molluscs : 

Bivalvia 

Bivalvia 

17 

15 

262 

239 

40 

39 

344-2389 

298-1381 e a p . 

T~-marjpnj Polychaeta : Cirratulidae 15 231 52 298-2945 

r nerides day; Polychaeta : Lumbrineridae 15 229 48 298-2945 

rene a sp. A Molluscs : Bivalvia 14 225 34 547-1465 

atxe hale.oculata Polychaeta : Nereidae 14 220 45 339-2901 

ara ac on a paraJoxa Polychaeta : Lacydoniidae 14 211 37 298-1390 

u a sp. Molluscs : Bivalvia 13 208 46 339-1465 

~iTanedes sp . A Crustacea : Ostracoda 13 203 19 346- 838 

tosco o tos sp. A Polychaeta : Orbiniidae 13 196 20 

42 

298-1430 

298-2901 r c . annulosus Polychaeta: Cirratulidae 12 191 

ahinome "e fre sii Polychaeta : Amphinomidae 12 188 47 298-2475 

otana s sp. 1 Crustacea : Tanaidacea 11 181 34 339-1390 

natFia sp . 15 Crustacea : Tanaidacea 11 173 37 344-2901 

ere es stroemi Polychaeta : Trichobranchidae 11 173 35 298-2518 

~ x rrus ca e ~ nsis Polychaeta : Flabelligeridae 11 168 42 298-2389 ~ _ 

ono llis sp N° Polychaeta : Syllidae 10 161 27 298-1021 

eu ae A Crustacea : Tanaidacea 10 154 19 359-1465 

Aricidea sp. Polychaeta: Paraonidae 10 155 41 298-2518 

ere TTides atlantis 

. 

Polychaeta : Trichobranchid ae 10 153 32 339-1465 

t a sp. 201 Crustacea : Isopoda 9 149 9 344-1226 

ar s no is n, sp: 1 Crustacea : Amphipoda 9 145 36 547-2901 

xocep a idae sp. 1 Crustacea : Amphipoda 9 

9 

143 

141 

30 

28 

298-1465 

453-1381 Isthnamesus sp . 208 

s ar a sp . 

Crustacea : 

Molluscs : 

Isopoda 

Bivalvia 9 141 33 298-2945 

etta sp. Porifera 8 134 25 298-165 

rtes sp. CL4 Nemertea 8 130 47 298-2827 

Spionidae Genus B Polychaeta: Spionidae 10 130 14 298- 759 
MACROFAUNA DENSITY, TOTAL 

REGION -SEASON ~YEAR COMPARISONS 

Station -Moon Depth (m) 

1 -355 

2 -620 

3 -850 

4 -1400 

5- ,2602 

soon 

.. 

aooo 

g 

sooo 

H 

zooo a 

W 

000 0 

0 

Figure 4-15 . Comparative levels of macrofaunal densities by region, season, year and selected depth 

w interval . 

i r .s 4r s i P .s f a i r J i .s ~sreriaW .vuatwA 

~YE.3'T CENTi~.IL "Sr -A]E6YON 
174 

some exceptions at the shallower of the sampled depths (Fig. 4-15) " On 

the Central Transect, the observed trend of abundance decrease with depth 

is interrupted by an apparent abundance peak at the 1400-m deep station. 

Increased sampling intensity on the Central Transect, not only validated 

the 100-m peak, but yielded data suggesting even higher abundance was 

present at about 1000-m depths (Fig . 4-16) . These apparent differences 

will be examined in detail in the Year 4 Final Report. Suffice it to say 

now that we believe the "anomalous" abundance peaks to be related to 

physiography and/or proximity to ehemosynthetic seep communities. 

Results of sampling along isobaths in both the Eastern (Fig. 4-17) 

and Western (Fig. 4-18) Transeets yielded consistent results in terms of 

overall density, except for some of the planned contrasts of sediment 

type, proximity to seep communities and topographic features . The 

exceptions were unexpected low densities at Stations E2D and E3 on the 

Eastern Transact. All of these differences (those accountable by design 

and those unexpected) will be evaluated in the Year 4 Annual Report . 

4 .2 .4 MACROFAUNA DIVERSITY PATTERNS 

The diversity evaluations made in this report are based upon 

comparisons of the H" index applied to data for species only. From review 

of these data and supporting indices (evenness and richness) it is 

apparent that most of the differences in the overall index are 

attributable to the richness aspect, which is greatly influenced by sample 

size . In the Year 4 report, rarefaction analysis will be used to offset 

the confounding effects of sample size on the diversity evaluations. 

The regional, seasonal and yearly patterns of macrofaunal diversity 

by depth are presented in Figure u-19" Although the trends are not 

pronounced, diversity appears to decrease from east to west and to have 

been somewhat higher in fall than in spring on the Central Transact . 

Differences in fall diversity levels between years on the Central Transact 

were negligible, but spring 1985 diversity levels on the Eastern Transact 

were marginally higher than spring 1981 levels (Fig. 4-19) . 

The most consistent depth trend was a marked decrease in diversity 

between the 1400-m deep and 2600-m deep stations on each transact. There 

also appears a tendency of a slight diversity increase between the 
MACROFAUNA DENSITY, TOTAL 

CENTRAL REGION - FALL 1984 4000 a 

3000 

2000 

z 

1000 0 

~, /SOO 

rooo 

P.SOO 

i 

6 

JOr 

S 

Figure 4-16 . Comparative levels of macrofaunal density by depth on the Central Transect, Fall 1984 . 
V 

MACROFAUNA DENSM, TOTAL 

EASTERN REGION - SPRING 1985 

' " 5000 

-7 0 

,. 

3000 >- 

I2000 z 

W O 

1000 

. 

tv~l~v~ 

' 

:~ s3 

in 

. 

f . 

~ . 

is~o 

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: f . 

., 

1 

3D 

~ . 3 C 

. . C 30 

38 2E 

Idqp /CM Idw *WOO `9w AM IA! .lAw 

OEPT/I /Tl 

Figure 4-17 . Comparative levels of macrofaunal density along selected isobaths in the Eastern Gulf of 

Mexico during Spring 1985 . 
MACROFAUNA DENSITY, TOTAL 

WESTERN to CENTRAL REGION - SUMMER 1985 

0 c 

n 

z 

93°00' 9J °G10' 9/°00' 

ta~wrvaE 

Figure 4-18 . Comparative levels of macrofaunal density along isobaths between the Central and Western Gulf 

of Mexico sampling transects, Summer 1985 . 

v 
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DIVERSITY (H~) 

8Li 
(,H) Alisa3nia 

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180 

shallowest station ("'350 m) and some of the sequentially deeper stations, 

yielding somewhat skewed, dome-shaped diversity curves over the depth 

range sampled. 

The data obtained from sampling a higher density of stations on the 

Central Transact in fall 1984 enabled a more detailed examination of 

macrofauna diversity levels over the sampled interval (Fig . 4-20) . 

Diversity appeared to increase slightly from Station 1 (355 m deep) to 

Station 3 (850 m deep), and from there decreased with depth down to 

Station 9 (1428 m deep) . A slight peak was observed at Station 4 (1465 m 

deep), after which the diversity level once more declined gradually over 

the depth interval between 1465 m and 2945 m (Fig. 4-20). 

As shown by Figures u-21 and 4-22, macrofauna diversity levels were 

rather constant along the sampled isobaths in both the eastern and western 

regions where these studies were conducted. Additionally, there appeared 

very little variation by depth, at least within the intervals sampled . 

4 .2 .5 MAJOR GROUP ACCOUNTS 

The 32 macrofaunal groups discussed below account for 99.9% of the 

total macrofauna by abundance. The groups are arranged essentially in 

phylogenetie order (i.e., in order of NODC code numbers) and not in order 

of greatest abundance in the macrofauna. For this report, some groups 

which are not usually treated with macrofauna are included for convenience 

as they will not be discussed elsewhere. The following 32 groups are 

discussed below in this order: Porifera, Hydrozoa, Scyphozoa, Actiniaria, 

Alcyonaria, Scleraetinia, Nemertea, Kinorhyncha, Nematoda, Polychaeta, 

Oligochaeta, Gastropods, Aplacophora, Bivalvia, Seaphopoda, 6carina, 

Ostracoda, Copepods, Cumacea, Tanaidacea, Isopoda, Amphipoda, Decapoda, 

Pogonophora/Vesti mentifera, Sipuncula, Priapulida, Bryozoa, Brachiopods, 

Ophiuroidea, Echinoidea, Holothuroidea, and Ascidiacea. 

Of the 18 taxonomic groups for which highest and lowest total 

transact diversities (i .e., number of taxa at all stations on the 

transact) Ware figured, 61% (11) had their greatest diversity at the 

Eastern Transact on Cruise IV and 61% (11) had their lowest diversities at 

the Western Transact. Likewise, of the 26 macrofaunal groups for which 

mean transact densities were calculated, 54% (14) of the groups had their 
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highest mean transect densities at the Eastern Transect on Cruise IV while 

6% (12) had their lowest mean densities on the Western Transact . Another 

314 (8) had their lowest densities on the Central Transact on Cruise III . 

The reasons for this can only be speculated at this time, and will be 

dealt with in more detail in the synthesis report. 

4 .2 .5 .1 PoriPera 

Sponges ranked 15th in overall abundance in macrofaunal samples and 

ninth in total number of taxa. Some 39 taxa were identified, 22 of which 

were identified to the species level, 10 to the generic level, and seven 

to family or above. 

The number of sponge species ranged from 0-10 taxa at individual 

stations . Highest species diversity (10 taxa) occurred at Station E5, 

Cruise IV (2902 m) . Highest transact species diversity (21 taxa) was at 

the Central Transeet on Cruise III in November 1984. Lowest transact 

species diversity for sponges was five taxa at the Central Transeet on 

Cruise I in November 1983 " 

Sponges in the box core samples consisted of very small, intact 

colonies or protocolonies and not of pieces broken off from larger 

sponges . Therefore, the sponge counts and densities are quantitatively 

reliable . 

Sponge densities for each station on all transacts from all five 

cruises are shown in Figure 4-23 " Sponge densities ranged from 0 to 

137/m2 with the highest density found at Station E5, Cruise IV (2902 m) 

(Fig . 4-23) " Highest mean transact density, 48/m2, was at the East 

Transact on Cruise IV; lowest, 4/m2, was at the Central Transact on Cruise 

I . 

Mean densities arranged by depth shoal the following : 

Depth m Mean Density (No ./m?) 

298- 492 16 

500- 900 28 

1000-1500 22 

2000-2945 50 
oa 

G 

rt 

j Density (no. /m2) Density (no./m2) Density (no ./m2) Density (no./m2) Density (no. /m2) Density (no ./m2) 

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185 

The sponges are one of the few macrofaunal groups (along with the 

Oligoehaeta, Priapulids, and Braehiopoda) that show increased densities at 

the deepest levels sampled, i.e. below 2000 m depth. 

The ten most abundant sponges were : 

Overall Density No . of No . of Depth 

Species (No ./m2) Specimens Stations Range (m) 

Stelletta sp . 8 .47 134 25 298-1165 

Suberitidae sp . B 3 .92 62 19 298-2901 

Sycon sp . 2 .28 36 18 352-2901 

Thenea sp . B 1 .77 28 8 621-2901 

Stylocordyla sp . 1 .64 26 6 339-2901 

Thenea sp . C 1 .20 19 7 1350-295 

tale sp . A 0 .57 9 7 339-2901 

tale sp . B 0 .51 8 6 613-2945 

Suberitidae sp . C 0 .51 8 6 298-1430 

Mierociona sp . 0 .51 8 3 298- 759 

Most of the more abundant sponge taxa were distributed over a wide 

depth range, except for Microciona sp ., which was limited to depths 

shallower than 800 m. 

4 .2 .5 .2 Hydrozoa 

Hydroids ranked 22nd in overall abundance among the macrofaunal 

groups and 15th in total number of taxa . Fifteen teas acre identified, 

eight of which were identified to the species level, and three to the 

generic level. The others were to family level and above. 

The greatest diversity of hydroids was three taxa observed at each of 

Stations C1, WC5, and WC7 " The transact with the greatest number of 

hydroid taaa was the East Transact on Cruise IV where seven total taxa 

were distributed among 14 of the 16 stations sampled. No hydroids were 

collected on the Central Transact on Cruise I or the West Transact on 

Cruise II . 

Hydroids, like the bryozoans, are colonial animals and only colonies 

or pieces of colonies were counted. As with the bryozoans and sponges, in 
186 

most cases the hydroids encountered in the macrofauna were small, intact 

colonies and not broken pieces of larger colonies. 

Hydroid densities ranged from 0-98/m2, with the highest station 

density observed at Station E1A, Cruise IV at 352 m . Mean transect 

densities ranged from 0 at the Central Transect (Cruise I) and the West 

Transect (Cruise II) to 14/m2 at the East Transeet on Cruise II. Overall, 

hydroid densities were highest at the shallower depths (298-492 m) and 

lowest at depths greater than 2000 m . 

Hydroids for which more than one specimen was collected included : 

Overall Density No . of No . of Depth 

Taxon (No ./ m2) Specimens Stations Range (m) 

Eucuspidella sp . 1 .07 17 7 352-2901 

Halecium sp . 0 .44 7 1 453 

Corymorphidae sp . 1 0 .32 5 3 351- 838 

Opercularella sp . 0 .32 5 2 8u6-1021 

AEtlaophenia 

latecarinata 0 .13 2 1 354 

Obelia bidentata 0 .13 2 1 633 

Cladocarpus flexuosus 0 .13 2 2 298- 860 

x .2 .5 .3 Scyohozoa 

Scyphozoans were encountered in the macrofauna samples in the benthic 

strobila stage . They ranked 17th in total abundance of macrofaunal 

groups. No attempt was made to identify these asexual stages to lower 

taxonomic levels . 

Densities ranged from 0-182/m2, with the highest station density 

observed at Station E1A, Cruise IV (352 m) . Highest mean transact density 

for scyphozoans, 39/m2, occurred on the East Transect on Cruise IV and the 

lowest, 1/m2, was encountered on the West Transact. 

Mean densities for scyphozoans arranged by depth were: 

Depth (m) Mean Density (No ./m2) 

298- 492 56 

500- 900 10 
187 

Depth m Mean Density (No ./m2) 

1000-1500 16 

2000-2945 10 

All of the higher station densities (greater than 100/m2) occurred at 

four of the shallower stations (less than 500 m depth). Even within the 

shallow depth range of highest mean density, seyphozoan strobila 

distribution appeared to be very patchy. For example, four of the shallow 

stations had densities ranging from 0-7/m2 . 

4 .2 .5 .4 Aetiniaria 

Sea anemones in the macrofauna exist as very small juvenile forms 

that cannot be identified beyond the family, or higher level . Actiniaria 

ranked 33rd in overall macrofaunal abundance, with only eight individuals 

taken, primarily at stations deeper than 800 m. They were found only on 

the Central and East Transeets with none taken on the West or West-Central 

Transects . Highest station density for anemones in the macrofauna was 

7/m2 at Station C4 (1390 m) and E4 (1350 m) . 

4 .2 .5 .5 Alcyonaria 

Aleyonarian corals ranked 34th in overall macrofaunal abundance and 

occurred only at the East Transact on Cruise IV. Macrofaunal aleqonarians 

consisted of small, young colonies that were not identifiable except for 

one specimen of the isidid species, Lepidisis longiflora, taken at Station 

E3D (849 m) . Highest alcyonarian station densities of 7/m2 were at 

Station E2A (624 m) and E3 (819 m) . 

4 .2 .5 .6 Saleractinia 

Scleraetinian corals ranked 28th in overall macrofaunal abundance and 

20th in number of taxa. Six ta$a were identified, four of which were 

identified to the species level. The most abundant taxa of Seleractinia 

were : 
188 

Overall Density No . of No . of Depth 

Tazon (No./ m2) Specimens Stations Range (m) 

Deltocyathus sp . 0 .76 12 4 298-1465 

Schizocyathus 

fissilis 0 .70 11 6 298- 621 

Deltocyathus calcar 0 .38 6 1 298 

Highest species diversity was three taxa at Station WC5 (298 m) . 

Highest transact mean diversity for scleractinians, four taxa, occurred on 

the Central Transact on Cruise III and on the blest-Central Transact on 

Cruise Y. No scleractinians were taken on the Central Transact during 

Cruises I and II, and none were taken on the West Transact. 

Scleractinian station densities ranged from 0-63/m2 with the highest 

station density occurring at Station WC5, Cruise V (298 m) . Highest 

transeet mean density (6/m2) was observed on the west-Central Transact . 

Scleractinians reached their highest mean densities (6/m2) at the 

shallowest depths (298-192 m) and then dropped off drastically until there 

were none taken at stations deeper than 2000 m. 

4 .2 .5 .7 Nemertea 

Nemertean worms ranked 11th in overall abundance of all macrofaunal 

groups and 12th in total number of taxa. Twenty-one taxa were recognized, 

20 of which were identified to the species level . 

The nemertean species have been designated by letters rather than 

definitive scientific names until it can be determined with certainty 

which species have been previously described and which are new to science. 

The top nine nemertean species are listed in order of overall abundance 

below : 
189 

Overall Density No . of No . of Depth 

Taxon (No ./ m2) Specimens Stations Range (m) 

Nemertea sp . C14 8 .22 130 47 298-2827 

Nemertea sp . D 5 .88 93 36 298-2901 

Nemertea sp . B 4 .93 78 34 298-2475 

Nemertea sp . H 1 .33 21 13 298-1390 

Nemertea sp . E 1 .33 21 11 346- 856 

Nemertea sp . CL5 1 .07 17 12 298- 860 

Nemertea sp . J 0 .89 14 10 298-1165 

Nemertea sp . Z 0 .89 14 9 352-1021 

Nemertea sp . 1 0 .82 13 9 298-1350 

The three most abundant species were found across all depth levels 

sampled, while three other species appeared restricted to depths shallower 

than 1500 m and two were found only at depths shallower than 900 m. 

The number of Nemertean species per station ranged from 1-10 taxa, 

with the highest station diversity occurring at WC5 on Cruise V (298 m) . 

Lowest (one taxon) occurred at four stations, all of which were at depths 

greater than 1400 m. Highest total transact diversity, 17 taxa, occurred 

on the East Transact on Cruise IV. Lowest transact diversity, six taxa, 

was at the West Transact on Cruise II . 

Nemertean station densities are plotted for all transacts and all 

cruises in Figure 4-24. They ranged from 4/m2, at Station C12 (295 m) to 

158/m2 at Station WC5 (298 m) . Mean transact density was highest, 57/m2 , 

for the East Transact on Cruise IV and lowest, 20/m2, for the Central 

Transact on Cruise III. 

Nemerteari mean densities by depth are as follows : 

Depth (m) Mean Density(No ./m2) 

298- 492 51 

500- 900 44 

1000-1500 30 

2000-295 13 

There was a decided break in abundance at depths greater than 2000 m 

as compared to depths 1500 m and shallower . 
190 

NEMERTEA 

n 160- 

80-- 

E Cruise 1 Central 

120 - 

g nears-3e 

40- 

0 

0 500 1000 1500 2000 2500 3000 

N 160E 

120 - 

Cruise II Central 

80- Mean-43 .. 

r 40-

w 

G O 5N 1X00 1500 2000 2500 300 

^y 160- 

120 7 Cruise II West 

80 Man-36 

-a' 40 - 

C r 

0 500 1000 1500 2000 2500 3000 

M ~~ 

120- Cerise II East 

Mean=31 

r 

a 40- J~~\ 

~ \ 

NC ~ F 

O 

0 500 1000 1500 2000 2500 3000 

160- 

E 120 Cruise III Central 

Msan=20 C 80- 

a p0

0 

° 0 500 1000 1500 2000 2500 3000 

n 160 

Cruise IV East 120-

Man-57 80- 

40-

0 

0 500 1000 1500 2000 2500 3000 

N 160 

` 120 

- 

Cruise V West-Control 

Mwn=39 

.. 

a 

v 40-

0 -

O 0 500 1000 1500 2000 25OO 3000 

Depth (m) 

Figure 4-24 . Station densities (no/m2) for Nemertea at all transects 

on all five cruises . 
191 

u .2 .5 .8 ginorhyncha 

Kinorhynchs are tiny, externally segmented, worm-like organisms 

which, because of their small size, were found in both the maerofaunal and 

meiofaunal components of box core samples of this study . Rinorhynchs 

ranked 25th in overall macrofaunal abundance . Three taxa have been 

identified from macrofaunal samples, two of which were to the genus level : 

Overall Density No . of No . of Depth 

Taxon (No./ m2) Specimens Stations Range (m) 

Pycnophyes sp . 2 .66 42 6 346- 849 

Condyloderes sp . 0 .06 1 1 345 

Total kinorhynch densities ranged from 0-60/m2, with the highest 

individual density found at Station E1, Cruise IV (353 m) . Highest mean 

transact density was 10/m2 noted for the East Transeet on Cruise IV. No 

kinorhynchs were taken on the West or West-Central Transects. 

Mean densities of kinorhynch arranged by depth show the following 

pattern : 

Depth (m) Mean Density (No ./m2) 

298- 492 14 

500- 900 0 .5 

1000-1500 3 

2000-295 

Rinorhynchs loin the hydroids, scqphozoans, and scleraetinian corals 

in being the only macrofaunal groups having considerably higher mean 

densities in the shallowest depths sampled, with abundance dropping 

drastically at deeper depths . 

4 .2 .5 .9 Nematoda 

Nematode worms ranked second in abundance after the polychaetes in 

the macroPauna samples obtained by this study. Nematodes were not 
192 

scheduled for identification to more specific levels so there are no 

comparative data for species diversity considerations. 

Nematode population densities by station for all five cruises are 

presented in Figure 4-25 " Their station densities ranged from 144-2937/m2 

with the highest density occurring at Station C9, Cruise III (1430 m 

depth) and the lowest at Station C6, Cruise III (492 m) . 

The highest mean transact density for nematodes (1520/m2) was 

observed for the East Transeet on Cruise IV in May 1985 and the lowest 

(750/m2) was noted for the Central Transect on Cruise III in November 

1984 . 

Mean densities for nematodes by depth shoal the following : 

Depth Range (m) 

298- 492 

500- 900 

1000-1500 

2000-2945 

Mean Density (No ./W 

1025 

1198 

1156 

719 

Again, density for depths exceeding 2000 - m was markedly lower than 

density at depths above 1500 m. 

4 .2 .5 .10 Polychaeta 

A total of 24,313 polyehaetes were collected during five cruises 

sampling at 59 stations. Identifications of the polqchaetes included 414 

(66%) at the specific level, 163 (26%) to the generic level and 49 (8%) at 

some higher taxonomic category (usually family), for a total of 626 taxa. 

Overall, there was an average of 75 .7 polychaetes per box core sample 

representing a density of approximately 1330-1594/m2 " 

Densities of polqchaetes ranged from 267 to 2905/m2 at C12 (2951 m, 

Cruise III) and C2 (597 m, Cruise II), respectively. Densities varied 

with depth of collection, geographic area and cruise (sampling period) . 

Bathqmetric variations were tested specifically on Cruise III When 11 

stations were sampled along the central region transact (Fig . 4-26) . 

Within the depth range of 357-1465 m densities ranged from 899 to 2665/m2 

with minimum and maximum at 633 and 1021 m, respectively. Below 2000 m 
193 

NEMATODA 

3°°° 

2000-

1000 

i 0 0 

0 

N 3000 

E 

0 2000 

e 

r 1000-

MC 

0 

N 3000 

E 

o ZOW 

5 

1000 -

0 -

0 

0 

3000 

c° 2000 

.. 

1000-

A 0 --

0 

3000 

°c 2000 

1000- 

i 0- 0 

0 

n 3000 

E 

e 200 

c .. 

a 1000-

x 

0--

0 

3000 

4 Y000 

b 

y~ 1000-

u C- 0-

0 

c.rvlse I Central 

Hran=821 

500 1000 1 500 2000 2500 

Cruisell Central 

Mean--950 

500 X000 1500 2000 

Cr'utselt West 

Mean-834 

500 1000 1500 2000 2500 

Cruise il East 

Mean=1455 

500 1000 1500 2000 2500 

_ / 1 Cruise III Central 

Mean=750 

3000 

3000 

3000 

3000 

500 1000 1500 2000 2500 3000 

Cruise IV East 

hfean=1520 

500 1000 1500 2000 2500 3000 

Crake V West - Central 

Man--W 

500 1000 1500 2000 2500 3000 

Depth (m 

Figure 4-25 . Station densities (no/m2) for Nematode at all transects 

on all five cruises . 
194 

POLYCHAETA 

boon 

Cruim i JIM : 3000 

2000-

1000 -- 0 . 

o 5oo 1ooo 15oo 2ooo 2soo aooo 

6000 - 

5000- Gviss 11 west X000 

3000 

2000 -

1000 . 

0 

0 300 1000 1500 2000 2500 3000 

6000- 

5000- Cruise 11 Gnlril 

4000 - 

3000- 

2000 - 

1000- 

0 . 

0 500 1000 1500 2000 2500 3000 

.. 

N 

6000 . Cruisc 11 East 

4000 

.r 

3000 

2000 - 

ao 1000 . 

0 m 

0 500 1000 1500 2000 2300 3000 

5000 

4m - Cnrim III Can" 

4000 

3000-

2000 -

1000-

0 .- 

0 500 1000 1500 2000 2500 3000 

6000-

5000 . Cniism N East 

X000 

3000-

2000 -

1000 . . . 

0 .-

0 500 1000 1500 2000 2500 3000 

6000- Cruise V West-Centnl 3000 

4000 -

3000-

2000 -

1000- 

0 

0 500 1000 1500 2000 2500 3000 

Depth (m) 

Figure 4- 26. . Polychaete densities (no/m2) at all stations, all transects, 

and all five cruises . 
195 

low densities were found (167-386/m2) . On all cruises polyehaete 

densities were lowest at depths greater than 2300 m. (Usually no samples 

were taken between 1500 and 2300 m.) For each of the transacts sampled on 

Cruises I-V, maximum densities were observed at 613 359 354, 597, 1021, 

630 and 298 m (Fig. 4-26) . Excluding the deepest stations which always 

had the lowest densities the following depths on each transact sampled had 

the minimum density estimates: 1381, 1410, 354, 1355, 633, 819, and 1236 m 

(Fig. 4-26) . From these data, polyehaete densities usually reached their 

maximum at 300-630 m depths then decreased with increased depth. 

Mean polychaete densities arranged by depth for all stations sampled 

on all transacts show the following pattern of decreasing polychaete 

densities with depth : 

Depth Range (m) 

298- 492 

500- 900 

1500-2000 

2000-2845 

Density (No ./m2) 

1982 

1787 

1 441 

482 

Once more, densities were appreciably lower at depths greater than 

2000 m than at shallower depths. 

The effects of geographic variation within the Gulf were tested 

during Cruise II when five stations along each of three transacts located 

in the western, central and eastern Gulf were occupied . Depths sampled 

were 354-359, 597-627, 838-854, 1355-1410, and 2386-2827 m (Fig . u-26) . 

Except for the shallowest depth range sampled, the maximum densities were 

observed along the Central Transact, while minimum values were found along 

the Western Transact . These extremes were reversed for the shallowest 

depth sampled. Polqchaete densities on the Eastern Transact were 

intermediate. 

On Cruises IV and Y some similar depth ranges were sampled in the 

eastern Gulf and West-central Gulf . Mean densities by depth range are 

shown below. 
196 

Mean Density (No ./m2) 

Depth Ranse West-Central Eastern 

298-353 m 2269 2131 

547-630 m 1876 233 

748-860 m 1069 1709 

These results indicate polychaete densities may be higher in the 

eastern than west-central Gulf at depths in the two ranges of 547-630 m 

and 7u8-860 m, but not shallower, in the range of 298-353 m . These 

results are similar to the ones discussed for Cruise II which showed that 

western stations had lower densities than either central or eastern 

stations, except for the shallow depth 359 m which had the maximum density 

in this depth range contour. 

Density comparisons among cruises can be compared using Central 

Transact stations C1-C5, that were each sampled three times (Cruises IIII) . Densities (no/m2) by station (depth) for each cruise are shown 

below: 

Cruise 

Station (Depth m) I Nov II Mar III Nov 

C1 (354-365) 1242 209 1177 

C2 (597-633) 1523 2905 899 

C3 (838-881) 1108 2263 989 

C4 (1381-11465) 1085 2150 1564 

C5 (2386-2518) 639 872 386 

The highest densities were observed on Cruise II While the lowest 

densities were on Cruise III (except for Station C4) . 

Four stations in the eastern Gulf were occupied in consecutive years 

during the same "season" (May-June) . Polychaete densities found on 

Cruises II and I0 are shown below : 
197 

Station (Depth m) 

E1 (353-35u) 

E2 (619-627) 

E3 (819-846) 

E5 (2827-2901) 

Density (No ./m2) 

Cruise 

11 (198) IV (1985) 

2333 2239 

1927 2601 

2051 1381 

473 515 

These data show considerable variability between years as well as between 

"seasons" or cruises with few patterns or trends relative to increases or 

declines in polyehaete density. 

Polychaete Feeding Types 

Polychaetes were classified according to feeding type on the basis of 

taxonomic family (Attachment A2 .4) . The total abundance and number of 

taxa within each feeding type was computed. A summary of this tabulation 

is shown in Figure 4-27 . Deposit feeders (selective [SDF] and nonselective [NSDF]) Were the most abundant polyehaetes in terms of total 

counts and number of taua. More families (16 with 9g%total taxa) of 

carnivores (C) were collected than any other single category although the 

abundance of this group was less than either deposit feeders (NSDF and 

SDF) or omnivores (0) . Suspension feeders (SF) and scavenger (S) Were 

rarely found ; the two categories combined representing only 8$ of the 

polyehaetes collected and identified. Non-selective deposit feeders were 

the most diverse group with 196 taxa in this category, while only 37 taxa 

were classified as scavengers. 

Polychaete Depth Ranges 

A total of 150 polyehaete species were represented by 10 or more 

individuals (264 species occurred fewer than 10 times) . The depth ranges 

of these 150 species (Attachment A2 .5) showed an interesting pattern: the 

minimum depth of occurrence for 83% (124) of the species was in the range 

of 291-366 m and the deepest minimum depth of occurrence was only 639 m. 

The maximum depth of occurrence was more variable but indicated that depth 
198 

240 

220 

200 

180 

160 

140 

1`V 

S 

O 

100 

VV 

60 

40 

20 

0 

BOOO 

L 6000 

4000 

2000 

O 

No. Families 

p No. Taxa 

~"" 

C=S 7 

~~7 V 

PHY 

HSDF SDF 0 C SF 

4SDf SDF 0 C SF S 

FEEDING TYPE 

Figure 4-27 . Polychaete feeding types showing number of families, taxa, 

and individuals in each feeding type . NSDF = non-selective 

deposit feeder ; SDF = selective deposit feeder ; 0 = omnivore ; 

C = carnivore ; SF = suspension feeder ; S = scavenger. 
199 

per se was not limiting to many of the species. Forty-four of the 150 

species were found to depths in the 2118-2959 m range . Only 27 species 

(18%) Were restricted to less than 1000 m depths. 

There were 50 species arbitrarily designated as abundant (i.e., >50 

total individuals) . Of these abundant polychaetes (Table 4-4), 64% were 

found from shallow ("'300-350 m) to the deepest depths (>2000 m) . Only two 

of these species were restricted to less than 1000 m depths, and the 

remaining 16 species had maximum depths of occurrence in the 1007-1507 

m range (Table 4-4) . 

Polyehaete Diversity 

Polychaetes were the most diverse group of macroinfauna sampled. A 

total of 626 taxa, with 414 at the species level, were identified . 

Species identification was enabled by use of a diverse literature--e.g, we 

used some 43 references which are appended as Attachment A2 .6 . Most of 

the species were uncommon occurrences with low abuadanees and restricted 

distribution. Only 150 of the 414 species bad a total count of 10 or 

greater individuals out of the total of 24,313 polychaetes counted . 

Further, only 156 of the 626 total tana were observed in 10 or more of the 

59 station collections. 

Polychaete diversity as described by the mean number of species per 

replicate in a station collection was variable with depth, geographic 

location and cruise (season and year of collection) . A test of depthrelated differences was conducted during Cruise III When 11 stations were 

located at various depths along a single transect in the central Gulf. 

The number of species at each station ranged from 18 to 82 (Fig . 4-28) . 

The highest number of species were found from 357 to 165 m (range . 48-

82, mean -_ 48) . Kith the exception of Station CT, a possible area of 

hydrocarbon seepage, (number species -_ 82) this range was relatively 

consistent at 48-63 (mean = 55) " At the deeper stations (>2101-2951 m) 

the mean number of species per station was 20, less than half the number 

at the shallower stations . Station C7 appeared to be anomalous with 

respect to the high number of species observed there . On other cruises 

where fewer stations were sampled over approximately the same depth range, 
200 

Table 4-4. A list of the 50 most abundant polychaetes . 

DEPTH (m) 

Speoies Min mauc Number 

PHOLOE "SP .C" 343 1007 417 

PARAHPHINOME JEFFREYSII 291 2377 188 

PARAMPHINOME SP . A 298 250 130 

PARALACYDONIA PARADO%A 302 1236 211 

3IGAMBRA TENTACULATA 324 2953 60 

3YNELMIS KI.ATTI 343 2540 51 

LITOCORSA ANTENNATA 291 $70 1209 

PIONOSYLLIS "SP . H" 298 1032 161 

PIONOSYLLIS "SP .B" 344 860 56 

SYLLIS (EHL.ERSIA) CORNUTA 336 1198 74 

BXOGONE SP .B 291 252 104 

BXOGONE "SP.A" 34? 2959 467 

CERATOCEPHALE OCULATA 335 290 220 

1GLAOPIiAMUS CIRCINATA 349 2524 57 

Cd.YCERA PAPILLOSA? 344 2959 78 

SARSONUPHIS HARTMANAE 298 1507 276 

LUMBRINERIS YERRILLI 302 1507 113 

IAMBRINERIDES DAYI 298 2915 229 

LEITOSCOLOPLOS SP .A 2.91 1389 196 

ARICIDEA SUECICA 291 2959 421 

IIRICIDEA (ARICIDEA) FRAGILIS 302 295 69 

ARICIDEA LOPEZI LOPEZI 298 1414 88 

ARICIDEA (ACMIRA) SIMPLEX 335 1222 61 

CIRROPHORUS LYRA 340 2945 116 

LBVINSENIA GRACILIS 291 2959 371 

LBVINSENIA UNCINATA 335 2953 91 

SABIDIUS CORNATUS 344 2540 62 

PRIONOSPIO CIRRIFERA 291 2524 272 

PRIONOSPIO EHLERSI 320 1506 380 

SPIOPHANES BOMBYX 298 290 118 

SPIOPHANES BERKELEYORUM 291 2902 505 

AIIROSPIO DIBRANCHIATA 29$ 2853 1190 

THARYX ANNULOSUS? 298 2904 191 

THARYX MARIONI 291 2959 231 

DIPLOCIRRUS CAPENSIS 302 2377 168 

DIPLOCIRRUS "SP .A" 291 2900 116 

FAWELIOPSIS SP .B 291 2959 283 

OPHELINA SP .A 320 1507 70 

TACHYTRYPANE JEFFREYSII 444 290 50 

TACHYTRYPANE SP .A 291 2540 571 

gESUN SP . A 335 2897 88 

NOTOMASTUS LATERICEUS 291 2959 52 

NOTOMASTUS AMERICANUS 291 1440 59 

BARANTOLLA "SP . A" 335 2540 86 

NALDANE "SP .A" 302 1507 402 

MYRIOCHELE HEERI? 320 1507 61 

AMPHARETE "SP .A" 291 2118 75 

TEREBELLIDES STROEMI 291 2533 173 

TEREBELLIDES ATLANTIS 336 1507 153 

EUCHONE INCOLOR? 298 1433 101 
zoi 

too 

80 . 

r 

O 

60-

rO 

6 40 

20 

0 500 1000 1500 200 2500 300 

Depth (m) 

C6 

C2 C5 

C I C3 C4 

C 11 

C12 

C5 

Figure 4-28 . Numbers of polychaete species at each station 

(depth) along the Central Transect (bruise III) . 
202 

"deep" stations (2386-2901m ) consistently had fewer species (mean = 27) 

than shallower stations (<1410 m) Where the mean number of species was 62 . 

Abundant Polyehaete Species 

Polychaete species which had a total count of at least 50 were 

considered to be "abundant" and therefore potentially more important to 

understanding the deep-sea processes relating to polychaetes. The species 

listed above in Table 4-4 were selected (along with a few related species) 

for annotations relative to geographic distribution, depth range 

preferences, and feeding type. The results of this effort are shown in 

the annotated species list (Attachment A2 .7) and maps (Attachment A2 .8) 

showing collection location during this study. 

4 .2-5-11 Olixochaeta 

Oligoehaetes ranked 20th in overall abundance of the macrofaunal 

groups and 19th in total number of taxa, with only nine taxa. Six of the 

taxa were identified to the species level. Most of these are new records 

of distribution in the Gulf of Mexico, and three of them are new species 

being described by Dr. Christer Erseus of the University of Goteborg in 

Sweden . 

Highest station species diversity for oligochaetes, four taxa, 

occurred at two stations on the East Transact, E3B and E3D (860 and 819 m 

depth) . Highest oligoehaete transact diversities, five taxa, were present 

on the East Transact on both sampling efforts for that transact (Cruise II 

and Cruise IV) ; lowest, one taxon, was at the West Transact. 

Population densities for oligochaetes ranged from 0-63/m 2. The high 

of 63/m2 occurred at two stations, E2C (619 m) and E5 (2827', m) . 

Oligochaetes reached higher densities on the East Transact than on any of 

the other transacts. Mean densities of oligoehaetes on the East Transact 

were 34/m2 on Cruise II and 27/m2 on Cruise IV. Lowest transact mean 

density, 1/m2, was at the West Transact on Cruise II. 

Oligoehaete mean densities arranged by depths are as follows : 
203 

Depth Range (m) Mean Density (No ./m2) 

298- 492 11 

500- 900 21 

1000-1500 6 

2000-295 19 

Their increase in density deeper than 2000 m is different from any of 

the other macrofaunal groups studied except the sponges and brachiopods. 

Most of the other groups in the macrofauna showed depths of highest 

density at shallower levels with a marked drop in density below 2000 m. 

The two dominant species of oligochaetes were two species of 

Bathydrilus , B. n. sp . (35 individuals taken at 13 stations) and B . 

lonAiatriatus (12 individuals taken at nine stations) . The top five 

oligochaete species are listed below in order of overall abundance : 

Ove ral 1 

Density No . of No . of Depth 

Species No ./ Specimens Stations Range (m) 

Bathydrilus n. sp . 2 .21 35 13 613-295 

Bathgdrilus lon giatriatus 0 .76 12 9 621-2475 

Tubificoides n . sp . 0 .19 3 2 339- 344 

Phallodrilus n . sp . 0 .19 3 3 627- 860 

Limnodriloides monothecus 0 .19 3 1 346 

4 .2 .5 .12 Gastropoda 

Gastropod mollusks ranked 18th in overall numerical abundance in the 

maerofauna and 8th in total number of taxa with 53 " Eight of the taxa 

which acre identified to the species level, 27 to the generic level, and 

18 to family level or higher. 

The relatively lox number of identifications to the species level 

reflects the great number of immature or juvenile specimens of very tiny 

sizes that were not easily identifiable to more specific levels than 

family or genus . 

Numbers of gastropod taxa at individual stations ranged from 0 to 8, 

with the highest number of taxa found at Station C1 , Cruise I at 344 m 
204 

depth. Gastropod transact diversities were greatest (22 taxa) at the 

West-Central Transact on Cruise V, followed closely by 21 taxa on the East 

Transact on Cruise IV. Lowest transact diversity for gastropods was only 

three taxa at the West Transact on Cruise II. 

Gastropod population densities ranged from 0 to 70/m2, with the 

highest density occurring at Station WCS, Cruise V at 298 m depth. Mean 

transact density for gastropods was highest (30/m2) for the Central 

Transact on Cruise II in April 1981 . Two transacts tied for lowest mean 

gastropod density at 7/m2 : the West Transact on Cruise II and the Central 

Transact on Cruise III. 

Gastropod mean densities arranged by depth are as follows : 

Depth Range (m) Mean Density (No ./m2) 

298- 492 20 

500- 900 20 

1000-1500 15 

2000-295 7 

This contrasts with the bivalves, whose overall densities are more than 10 

times greater than the gastropods and whose densities at the 1000-1500 m 

depth is relatively much higher. 

The top seven taxa of gastropods from all macrofaunal samples are 

listed in order of overall abundance : 

Overall 

Density No . of No . of Depth 

Species No ./m2 Specimens Stations Range (m) 

Circa sp . 0 .57 9 6 547-143 

Phileae sp . 0 .51 8 8 563-2506 

Melanella sp . 0 .38 6 5 35u-2475 

Corinnaeturris sp . 0 .25 4 3 344- 853 

Crenilabium sp . 0 .25 4 2 344- 759 

Turbonilla sp . 0 .25 4 3 352- 622 

Solariella sp . 0 .25 4 4 298-856 
205 

x .2 .5 .13 gAlacophora 

The aplacophora ranked 10th in overall abundance for the macrofaunal 

groups collected in this study. They were not identified to the species 

level . 

Aplacophora densities ranged from 0-211/m2, with the highest density 

occurring at Station C7, Cruise III (1021 m depth) . Aplacophorans were 

absent from only three stations, all of which were deeper than 2000 m. 

Aplacophoran mean transact densities were highest, 88/m2, at the East 

Transact on Cruise IV and lowest, 34/m2, at the West Transact . 

Mean densities for aplacophorans by depth are as follows : 

Depth Range (m) Mean Density (No ./m2) 

298- 492 73 

500- 900 65 

1000-1500 50 

2000-29u5 7 

Markedly lower densities occurred at depths greater than 2000 m than 

at shallower depths . 

4 .2 .5 .14 Bivalvia 

The bivalves ranked fifth in overall abundance and seventh in total 

number of taxa. Some 55 taxa were identified, 41 of which were identified 

to the species level and 10 to the generic level . Bivalves reached their 

greatest diversity (19 taxa) at Station E1B, Cruise IV (346 m) . Lowest 

bivalve station diversity was four taxa at Station E5, Cruise II. Highest 

total transact diversity (u2 taxa) occurred at the East Transact on Cruise 

IV. Lowest transact diversity was 24 taxa at the west Transact on Cruise 

II . 

The top 10 bivalve species arranged in order of overall abundance are 

listed below : 
206 

Species 

Eulamellibranahia sp . F 

Vesicomya sp . 

Malletia sp . B 

Crenella sp . A 

Nucula sp . A 

Cuspidaria sp . 

Lucina sp . 

Limopsis sp . 

Eulamellibranehia sp . B 

Voldiella sp . A 

Density 

Overall 

Density No . of No . of Depth 

( No ./m2) Specimens Stations Range (m) 

20 .17 319 34 298- 860 

16 .57 262 40 344-2389 

15 .11 239 39 298-1381 

14 .23 225 34 547-165 

13 .15 208 46 339-165 

8 .92 141 33 298-2945 

7 .15 113' 23 298- 881 

6 .77 107 29 298-1465 

6 .64 105 21 402-295 

6 .64 105 9 856-2389 

Bivalve population densities ranged from 68-484/m2 . The highest 

station densities of greater than 400/m2 occurred at the East Transact on 

Cruise IV at depths between 400-484 m. Highest mean density by transact 

was 296/m2 at the East Transact on Cruise IV. Lowest transact mean 

density for bivalves (157/m2) occurred at the West Transact. 

Bivalve mean densities arranged by depth over all stations, all 

cruises are as follows : 

Depth Range (m) Mean Density (No ./ m2) 

298- 492 n1 

500- 900 227 

1000-1500 260 

2000-2945 124 

Markedly lower density occurred at stations in the 2000-3000 m depth range 

than at shallower depths . 
zoo 

x .2 .5 .15 Scaphoaoda 

Seaphopods ranked 16th in overall abundance in the macrofauna and 

18th in number of taxa, with 10 taxa identified. Five were identified to 

the species level and two to the generic level . Scaphopods were 

identified from the first three cruises only, and their taxa are listed 

below in order of overall abundance in the macrofauna: 

Overall 

Density No . of No . of Depth 

Species ( No ./m2 ) Specimens Stations Range (m) 

Episiphon sp . 1 .01 16 7 613-2506 

Dentalium perlongum 0 .38 6 5 359-2475 

Cadulus sp . 0.32 5 3 881-2475 

Dentalium didymum 0 .25 4 1 275 

Heteroschizmoides callithrix 0.19 3 1 1381 

Dentalium callithrix 0 .19 3 2 1390-130 

Pulsellum pressum 0 .06 1 1 1430 

Three of the seven taxa listed above, Dentalium prolonAUm , Cadulus 

sp., and Pulsellum pressum , were also present in the trawl megafaunal 

samples as more mature and larger sized individuals. 

On the first three cruises, highest scaphopod station diversity, six 

taxa, was observed at Station C4, Cruise I (1381 m) . Highest transact 

diversity of eight taxa was the Central Transact on Cruise I ; lowest 

transact diversity of three taxa was at the East Transact, Cruise II. 

Seaphopod population densities have been calculated for all five 

cruises and ranged from 0-105/m2 With the highest station density at 

Station E2, Cruise IV (353 m). Highest transact mean density of 36/m2 Was 

at the Salt Transact on Cruise IV ; lowest mean transact density, 8/m2 was 

at the East Transact on Cruise II. 

Scaphopod mean densities arranged by depth are as follows : 
zos 

Depth Range ( m) 

298- 492 

500- 900 

1000-1500 

2000-295 

Mean Density (No ./m2) 

36 

20 

21 

17 

Highest density was observed at the shallowest depth range . 

4 .2-5 .16 Acarina 

The marine mites of the family Halacaridae ranked 24th in order of 

total abundance in the macrofaunal samples taken by this study. They were 

not identified to lower levels. 

Halacarid densities ranged from 0-46/m2 with the highest station 

density occurring at Station WCS, Cruise V (298 m) . Highest mean transact 

density for halacarids, 7/m2 was at the West-Central Transact on Cruise Y. 

No halacarids were found on the entire Central Transact on Cruise I nor on 

the West Transact on Cruise II. 

Mean depth-specific densities follow : 

Depth Range (m) 

298- 492 

500- 900 

1000-1500 

2000-2945 

4 .2 .5 .17 Ostracoda 

Mean Density (No ./m2 

5 

u 

3 

Ostracods ranked third in overall abundance and eleventh in species 

diversity, with 22 taxar-18 of which were identified to the species level. 

Only the myodocopan ostracodes were identified to the species level. The 

podocopans and other non- myodooopan ostracodes were not identified to more 

specific levels. 

The maximum number of species of Ostracoda at any boxcore station 

(eight) occurred at fine stations, viz., C2 (Cruise II), E3, E3C, and E3D 

(Cruise IV), and WC6 (Cruise V) . These stations were all between 
209 

approximately 550-850 m depth. Lowest number of species were seen at the 

deepest stations (greater than 2000 m) . Highest overall transect 

diversity of ostracods, 17 taxa, was at the East Transact on Cruise IV ; 

lowest, six taxa, was at the West Transact. 

In general, highest population densities for ostracodes, i.e., 

greater than 500/m2 occurred at depths of 550-850 m on all transacts . 

Lowest densities of less than 100/m2 occurred at the shallowest stations 

(<500 m) and at stations deeper than 2000~m. Mean ostracod densities by 

depth follow : 

Depth Range (m) Density (No ./m2) 

298- 492 131 

500- 900 446 

1000-1500 273 

2000-2945 135 

The seven most abundant species of Ostracoda are listed in order of 

decreasing overall abundance below : 

Overall 

Density No . of No . of Depth 

Species No ./m2 Specimens Stations Range (m) 

Euphilomedes sp . A 20 .99 232 23 453- 860 

Anaulorostrum sp . A 17 .64 279 44 298-1u65 

Philomedes sp . A 12 .84 203 19 346- 83 8 

Pseudophilomedes sp . A 8 .03 127 18 346- 849 

Harbansus sp . A 6 .20 98 18 353-1236 

ene sp . A 2 .21 35 11 563-2518 

Metavarptula sp . A 1 .71 27 9 622- 860 

4 .2 .4 .18 CoDepoda 

Copepods, primarily harpacticoids, ranked 4th in overall abundance of 

macrofaunal groups. No species identifications were scheduled for the 

copepods from this project . 
210 

Station densities ranged from 56-807/m2 with the highest density 

found at Station Cu, Cruise II (1390 m) in April 1984. Lowest copepod 

density of 56/m2 was at Station C12, Cruise III (2945 m) in November 1984. 

Highest transect mean density for copepods, 499/m2 was at the Central 

Transact on Cruise II in April 1984, lowest of 168/m2 was at the West 

Central Transect on Cruise V in June 1985 followed closely with 169/m2 at 

the Central Transact on Cruise III in November 1984 . 

Mean densities of copepods by depth show the following patterns : 

Depth Raarte ( m) 

298- 492 

500- 900 

1000-1500 

2000-295 

Mean Density (No ./m2) 

154 

262 

348 

ZZs 

Copepods were one of the few maerofaunal groups, along with the 

isopods, that had their greatest mean densities in the 1000-1500 m depth 

range . 

4 .2 .5 .19 Cumacea 

Cumaceans ranked 14th in overall abundance and fifth in total numbers 

of taxa with 86 taxa, 76 of which were identified to the species level and 

eight to the generic level . Cumaceans reached their greatest species 

diversity of 14 teas at two stations, viz ., Station C1, Cruise I (344 m) 

and Station E3C, Cruise IV (849 m) . Highest transact diversity of 53 taxa 

was at the East Transact on Cruise IV. Lowest transact diversity for 

cumaceans, 10 taxa, occurred on the West Transact . The most abundant 

cumaceans were : 

Overall 

Species 

Procampylasyis acanthomma 

Cumella antipai 

Campylaspis spinosa 

Density 

No ./m2 

2 .09 

2 .09 

1 .96 

No . of No . of 

Specimens Stations 

33 16 

33 13 

31 16 

Depth 

Range (m) 

3u4-1021 

598-2901 

3u6-1381 
zii 

Overall 

Density No . of No . of Depth 

Species No ./m? Specimens Stations Range (m) 

Cumella erecta 1 .14 18 11 563-2827 

Cumella compacta? 1 .14 18 5 298-1192 

Eudorella n . sp . C 1 .01 16' 8 344-1430 

Cumella bishops 0 .95 15 9 547-1u65 

Epileucom tenuirostris 1 .39 22 16 344-2389 

Cumella daqae 1 .26 20 13 298-2389 

Density 

Cumacean population densities ranged from 0-130/m2 . The highest 

density of cumaceans occurred at Station WCS, Cruise Y at 298 m depth, and 

the lowest (0) at the following three stations : C6 and C11, Cruise III 

(492 and 2101 m) and WC12, Cruise V (1236'm) . 

Highest mean transact density for cumaceans (48/m2) occurred on the 

East Transact on Cruise IV in May 1985. Lowest transact density (20/m2) 

Was at the West Transact on Cruise II in April 198 " 

Cumacean mean densities by depth were : 

Depth Range (m) Mean Density (No ./m2) 

298- 492 36 

500- 900 38 

1000-1500 28 

2000-295 15 

This pattern is similar to those of the amphipods and tanaidaceans, 

with highest densities between the 500-900 m depth range, dropping off 

slightly in the 1000-1500 depths and finally declining drastically in 

depths of greater than 2000 m . This is in contrast to the other mayor 

crustacean group, the isopods, which reached their highest densities at 

deeper levels; i.e., between 1000-1500 m. 
212 

4 .2 .5 .20 Tanaidacea 

Tanaidaceans ranked sixth in total abundance and second in diversity 

of species with 186 taxa, of which 168 were identified to species and 13 

identified to the generic level . Tanaidaceans reached their greatest 

numbers of taxa (u7) at Station EN (Cruise IV, Eastern Transact) and the 

lowest, five taxa, at Station C1 (Cruise III, Central Transect) . In 

general, number of species of tanaidaceans was greatest between 600 and 

1000 m depth and lowest at the deepest (>2000 m) and shallowest (<500 m) 

depths of this study. Highest tanaidacean transact 'diversity", 124 taxa, 

was at the East Transact on Cruise IV ; lowest, 50 taxa, was at the West 

Transact . 

The ten most abundant species of tanaidaceans were, in order of 

decreasing abundance : 

Overal l 

Density No . of No . of Depth 

Species No ./m2 Specimens Stations Range (m) 

Pseudotanais sp . 1 18 .91 299 52 298-295 

Mesotanais sp . 1 11 .45 181 34 339-1390 

LeptoRnathia sp . 15 10 .94 173 37 344-2901 

Apseudidae A 9 .74 154 19 359-165 

Tariaella sp . 1 7 .52 119 34 298-295 

Leptognathiidae genus A 5 .82 92 28 298-1465 

Mesotanais sp. 2 5 .56 88 26 547-1390 

Apseudes sp . 1 5 .50 87 2 550- 624 

Leytoanathia sp . 5 .44 86 33 298-2389 

Typhlotanais sp . 5 4 .17 66 24 298-2901 

Density 

Densities of tanaidaceans ranged from 28/m2 to 512/m2 . Lowest 

density occurred at Station C1, Cruise III (357 m) and the high of 512/m2 

occurred at Station C7, Cruise III (1021 m), both on the Central Transact . 

Highest average density by transact (325/m2) occurred on the Eastern 

Transact on Cruise IV in May 1985 . Lowest average density (134/m2) Was at 
213 

the Central Transact on Cruise III in November 1984 . In general, 

tanaidaceans reached their highest densities (>400/m2) on the Eastern 

Transact in May 1985 (Cruise IV) at depths ranging between approximatly 

620-850 m. 

Tanaidacean mean densities by depth were: 

Depth Ranpce (m) Mean Density (No ./m2) 

298- 492 11 

500- 900 330 

1000-1500 201 

2000-295 63 

Overall density (all replicates pooled and divided by the total area 

sampled) of tanaids was higher at stations located in the middle of the 

depth range (e .g. from 500 to 1500 m) for most transacts than at either 

deeper or shallower stations. There appeared to be a pronounced decline 

in tanaid densities below 2,000 m. On the Central Transact, tanaids were 

most abundant at Stations C2, C3, C4, C7, C8, and C9 (from 598 m to 1 ,465 

m (Figs . 4-46) . On the Western Transact, tanaid densities on Cruise II 

were highest at Station W3 (depth 854 m) . On the Eastern Transact, 

tanaids were most abundant on Cruise II at Station E3 (depth 846'm), and 

on Cruise IV at stations between the 500 m and 1000 m isobaths. There was 

no clear east-west difference seen in tanaid density on the West-Central 

Transact ; high and low densities were each seen at both ends of the 

transact . 

4 .2 .5 .21 Isoyoda 

Isopods ranked eighth in overall abundance and third in diversity of 

species with 133 tans, 119 (89x) of which were identified to species and 

eight of which were identified to the generic level . Most of the Zsopoda 

from this study are new species, hence their identification designation of 

a species code number following the generic name. It is hoped that the 

isopods from this study can be worked up into a series of monographs on 

deep-sea isopods of the Gulf of Mexico by Dr. George Wilson of the Scripps 

Institution of Oceanography. 
214 

The ten most abundant species of isopods are listed in order of 

overall abundance were: 

Overall 

Density No . of No . of Depth 

Sp ecies ( No ./m? ) Specimens Stations Range (m) 

Gnathia sp . 201 9 .42 149 9 344-1226 

Ischnomesus sp . 208 8 .92 111 28 453-1381 

Prochelator sp . 202 7 .46 118 5 344- 492 

Proehelator sp . 408 6 .13 87 21 316- 860 

Maerostylus sp . 256 5 .94 94 15 630-2945 

Prochelator sp . 235 5 .50 87 28 339-2518 

Chelator sp . 237 4 .68 74 27 339-2518 

Prochelator sp . 209 4 " 43 70 19 492-2518 

Macrostylus sp. 223 4 .36 69 19 547-1381 

Torholia sp . 203 3 .86 61 25 339-2901 

Isopods achieved their highest number of species (3u) at Station WC9, 

Cruise V (759 m) in June 1985 . Lowest diversity of isopod species, two 

taxa, occurred at Station C1, Cruise III (357 m) in November 1984 . In 

general, the highest isopod species diversities (greater than 20 taxa at a 

station) occurred at the Central and West Central Transects between 600 

and 1300 m depth. Lowest diversities of less than 10 taxa at a station 

were found at depths greater than 2000 m and at the shallowest stations 

(less than 500 m depth) . 

Highest transect diversity, 87 taxa, occurred at the East Transect on 

Cruise IV ; lowest, 35 taxa, was at the West Transact . 

Isopod population densities ranged from 28-580/m2. Highest density 

of 580/m2 occurred at Station C1, Cruise I (344 m) ; lowest of 28/m2 Was at 

Station W5, Cruise II (2506 m) . 

Mean densities by transect were highest (245/m2) at the Central 

Transact on Cruise II in April 1981 and lowest (93/m2) at the Central 

Transect on Cruise III in November 1984 . 

Mean densities by depth showed the following patterns : 
215 

Depth Range (m) Mean Density (No ./m2) 

298- 492 117 

500- 900 152 

1000-1500 178 

2000-295 66 

Isopods differ from the other crustacean groups of the maerofauna, 

i .e ., the amphipods, cumaceans, ostracods, and tanaidaceans in having 

their greatest densities at the deeper depths of 1000-1500 m instead of at 

500-900 m . However, a marked decrease in density was noted for the 

deepest depth range. 

4 .2 .5 .22 Amphipoda 

The Amphipoda ranked ninth in overall abundance and sixth in total 

number of taaa with 79 tasa identified--50 of which were identified to 

species. Amphipods reached their greatest species diversity of 21 taxa at 

Station C1, Cruise I (344 m depth) in November 1983 " Lowest diversities 

were 0 at two stations : E1 (354 m), Cruise II and C12 (2945 m), Cruise 

III. Highest transact diversity (45 taxa) was found at the East Transact 

on Cruise IV in May 1985 . Lowest transact diversity was 11 taxa at the 

West Transeet on Cruise II in April 1984. 

The top ten species of amphipod species were, in order of decreasing 

abundance : 

Overall 

Density No . of No . of Depth 

Species No ./m2. Specimens Stations Range (m) 

Harpiniinae 11 .76 186 37 298-2518 

Pardisynopia n . sp . 1 9 .17 115 36 517-2901 

PhoYOCephalidae sp. 1 9 .04 143 30 298-1465 

Byblis n . sp . 1 3 .10 49 16 298-1130 

Melita sp . 2 2 .72 43 18 298-2101 

Metayhoaus n . sp . 2 .02 32 13 59a-1394 
216 

Overall 

Density 

Species ( No ./mg). 

Melita sp . 1 1 .90 

Synopiidae sp . 7 1 .14 

Melita sp . 5 1 " 

Synopiidae sp . 5 0 .95 

No . of No . of 

Specimens Stations 

30 15 

Depth 

Range (m) 

298-1192 

18 13 346-1236 

16 3 295- 849 

15 10 352-1465 

Amphipod population densities ranged from O to 232/m2, with the high 

occurring at Station C7, Cruise III at 1021 m depth in November 1984 . 

Highest transact mean density was 101/m2 at the West Central Transact, 

Cruise Y in June 1985 . Lowest transact density for amphipods was 28/m2 at 

the west Transeet on Cruise II in April 1984. 

In general the highest amphipod station densities of greater than 

150/m2 occurred at stations located between depths of 500-900 m. Amphipod 

mean densities (all stations, all transeets) arranged by depths are as 

follows : 

Depth Ranae (m) 

298- 492 

500- 900 

1000-1500 

2000-295 

Mean Density (No ./m2) 

68 

102 

78 

12 

Looking in more detail at the amphipod station densities we find that 

overall densities of amphipods were typically highest at stations 

shallower than 1000 m . On the Central Transeet, amphipod density was 

highest at Station C1 (depth 344 m) on Cruise I and Stations C2 and C3 

(depths 598 and 838 m, respectively) on Cruise II. On Cruise III, which 

included many more stations down-slope on the Central Transact, amphipod 

densities were highest at Station C7 (1021 m), and declined rapidly at 

stations deeper than C7 . 

On the Eastern Transact, amphipod overall densities showed a similar 

trend, with highest densities at Stations E2 and E3 (depth 627 and 816 m, 

respectively) on Cruise II; and at various stations between 624 m (E2A) 

and 819 C E3 ) on Cruise IV. 
217 

On the Western Transact, amphipod overall densities were highest at 

Stations W2 (604 m deep) and W3 (854 m), and decreased at stations seaward 

of the 1000 m contour as well as the one station (W1) inshore of Station 

W2 . 

On the West Central Transact, there was no obvious east-west trend in 

amphipod abundance ; the stations deeper than 1000 m had relatively low 

densities compared to nearly all the shallower stations. 

4 .2 .5 .23 Decapoda 

The decapod crustaceans comprised only a minor part of the macrofauna 

as compared with their great abundance and diversity in the megafauna from 

this project . Decapods ranked 29th in overall macrofaunal abundance out 

of the 44 macrofaunal groups encountered in this study, i.e., in the lower 

1% . They ranked 16th in number of taxa, with a total of 13 taga 

identified from macrofauna samples, 10 of which were identified to the 

species level. The most abundant decapod species from macrofauna samples 

were : 

Overall 

Density No . of No . of Depth 

Species No ./m2 Specimens Stations Range (m) 

Bathyplaz tYphla 0 .51 8 7 547- 622 

Callianassia sp. A 0 .19 3 3 298- 353 

Callianassia marxinata 0 .13 2 2 352 

Pontophilus gracilis 0 .13 2 2 619- 622 

Axiopsis ( Calocarides) sp . A 0 .13 2 2 453- 622 

Some of these species, i.e., Bathyplax typhla and Poatouhilus 

gracilia , represent younger stages of species that were common in the 

trawl megafauna samples. Others, such as the thalassinids ( Callianassa 

and Axiopsis) are burroWers in the sediments and are present only in the 

macroinfaunal samples from this study. 

Highest decapod species diversity was three taxa at Station E2 E, 

Cruise ID (622 m) . This was the same station where the decapods reached 

their greatest density of 11/m2 . 
218 

4 .2 .5 .24 Pogonophora/Vestimentifera 

Eleven specimens of Pogonophora were taken in maerofaunal samples. 

The specimens have been sent to specialists in Great Britain for further 

study . They were found only on the Central Transeet (Cruise III), the 

East Transact (Cruise IV), and the West Central Transact (Cruise V) . 

Highest station density for pogonophorans was 18/m2 at the Station C7 

(1021 m). They were taken in macrofauna samples only between the depths 

of 622-1226 m . 

4 .2 .5 .25 Siyuncula 

Sipunculan worms ranked 13th in overall abundance of the maerofaunal 

groups. They were identified from the first three cruises only, from 

which 37 taxa were identified, 31 of which to the species level . The 10 

most abundant taga of sipunculans are listed below in order of abundance : 

Overall 

Density No . of No . of Depth 

Species No ./m? Specimens Stations Range (m) 

GolflnAia sp . B 1 .01 16 7 354-2827 

Golfingia sp . F 0 .57 9 3 351- 604 

Onehnesoma steenstruy II 0 .51 8 2 354- 359 

Golfingia sp . G 0 .44 7 2 359- 604 

Golfingia sp . 0 .38 6 5 357-110 

Go1PinAia sp . J 0 .38 6 4 633-165 

Golfingia sp. E 0 .25 4 2 359- 604 

Golfinaia sp . R 0 .19 3 1 1309 

Golfingia sp . I 0 .19 3 1 627 

Phascolion ap . A 0 .13 2 2 344- 613 

On the first three cruises, the greatest diversity of sipuneulans, 

seven taxa, was at Station w1 (359 m) ; greatest transact diversity, 14 

taxa, was at the Central Transeet on Cruise I . 

Population densities, ranging from 0-182/m2, were calculated for 

sipunculans over all fine cruises. The highest station density (182/m2) 
219 

occurred at Station E1, Cruise IV (353 m) . The highest transact mean 

density was 81 / m2 on the East Transect on Cruise IV. The lowest, 8/m2, 

was the mean density for the Central Transect on Cruise III. 

Sipuneulan mean densities by depth are as follows : 

Depth Range (m) Mean Density (No ./m2) 

298- 492 78 

500- 900 35 

1000-1500 15 

2000-295 u 

Sipunculans, along with the hydroids, seleractinians, and kinorhynchs 

showed highest densities at the shallowest depths, with densities 

declining over all depths. 

4 .2 .5 .26 PriaDUlida 

Priapulids are small, worm-like, coelomate animals that live buried 

in sediments in shallow and deep water. Only a few species are known 

world-wide. A total of 33 specimens were taken from macrofaunal samples 

in this study, ranking them 27th in overall abundance . They were not 

scheduled for identification to more specific levels because of their 

relatively minor contribution to the total macrofauna. 

Priapulid population densities ranged from 0-28/m2 with the highest 

density at Station C5, Cruise II (2389 m) . Highest overall transact mean 

density, 8/m2, was at the Cental Transact on Cruise II. Lowest transact 

mean density for priapulids of <1/m2 Was at the West-Central Transact on 

Cruise V . 

Mean densities for priapulids arranged by depth are as follows : 

Depth RanEte (m) Mean Density (No ./m2) 

298- 492 

500- 900 1 

1000-1500 <1 

2000-295 7 
220 

The priapulids were one of the few macrofaunal groups, along with the 

sponges and brachiopods, that showed highest mean densities at depths 

greater than 2000 m. However, their numbers were few overall. 

4 .2 .5 .27 Bryozoa 

The bryozoans ranked seventh in overall abundance and fourth in total 

number of taxa. A total of 99 taxa were identified, 82 of which were 

identified to species and 12 to the generic level. 

Station species diversities of bryozoans ranged from 0-14 species . 

Highest diversity was at Station C4, Cruise II (1390 m) in April 1984 . 

Brqozoans were absent from the two shallow stations on Cruise III, C1 and 

C6 (357 acid 492 m depth) in November 1984. In general the higher station 

species diversities for bryozoans (greater than 10 taxa at a station) show 

no particular depth patterns. Highest transect diversity for bryozoans, 

34 taxa, occurred at the Central Transact on Cruise II in April 1984. 

Lowest transact diversity of 11 taxa was also at the Central Transact, but 

on Cruise I in November 1983" 

The top ten bryozoan species arranged in order of overall abundance 

are shown below : 

Overall 

Density No . of No . of Depth 

Species ( Now/ m2 Specimens Stations Range (m) 

Nolella monniotae 112 .05 1772 27 298-291 

E~ZAinoma erualieri 18 .27 289 37 298-2506 

Nolella sp . 17 .26 273 22 353-295 

Setosellina elexaatula 6 .89 109 24 339-2101 

Sphaerulobryozoon 

pendunculatum 5 .19 82 25 298-1465 

inoma n . sp . 4 .24 67 10 547-2901 

Heloidoma impliQata 3 .73 59 7 339- 860 

Paehpzoon atlariticum 2 .91 46 lb 298-291 

Metrarabootomornha 

aenigmatistes 2 .34 37 8 550-1465 

Bathyalazooa forests 2 .21 35 11 547-2901 
221 

Five of the 10 species showed very wide depth ranges from near the 

shallowest to near the deepest depths sampled . TWO of the species were 

found only deeper than 500 m, two Were found only at depths shallower than 

1500 m, and one species was restricted to depths shallower than 900 m. 

Bryozoans are colonial animals thereby making counts of individual 

specimens very difficult. In the macrofauna samples of tnls sLUay, counts 

mere made of brqozoan "pieces". However, in most cases, these were 

intact, small colonies and not broken pieces of larger colonies. 

Bryozoan population densities ranged from 0-1168/m2, with the highest 

density occurring at Station E2 B, Cruise IV (627 m) in May 1985 " In 

general, the higher station densities (greater than 500/m2) showed no 

particular depth patterns ; i.e., high densities were found at all depths 

between 298-2101 m. Bryozoan densities were consistently lower at depths 

greater than 2500 m, however. 

Highest transeet mean density for bryozoans, 412/m2, was at the East 

Transact on Cruise IV in May 1985; lowest transact mean density of 50/m2 

Was at the west Transact . 

Bryozoan mean densities arranged by depths show the following 

patterns : 

Depth Range (m) 

298- 492 

500- 900 

1000-1500 

2000-2945 

Mean Density (No ./m2 

213 

256 

91 

121 

A closer look at individual station densities for bryozoans showed 

typically very high densities at a few stations on each transeet--usually 

from 500 m to 1500 m--and relatively low densities elsewhere. 

4 .2 .5 .28 BrachioDOda 

Brachiopods ranked 23rd in overall abundance in the macrofauna. Only 

two species of brachiopods occurred in the maerofaunal samples, one of 

which was a new species ArAyrotheca . 
222 

Overall 

Density No . of No . of Depth 

Species ( No ./ m2) Specimens Stations Range (m) 

Cryptopora rectimarginata 4 .93 78 16 354-2518 

ArRgrotheca n . sp . 0 .06 1 1 2101 

Brachiopod population densities ranged from 0=158/m2 with the highest 

density at Station C11, Cruise III (2101 m) . Highest transact mean 

density, 18/m2, was at the Central Transact on Cruise III; lowest, <1/m2, 

was at the East Transact on Cruise IV. 

Mean densities by depth for brachiopods were: 

Depth Range (m) Mean Density (No ./m2) 

298- 49 

500- 900 2 

1000-1500 

2000-2945 26 

Brachiopods are one of the fax groups, along with the Porifera and 

Priapulida, that have their highest densities at depths greater than 2000 

m. 

4 .2 .5 .29 Ophiuroidea 

Ophiuroids in the macroPauna ranked 12th in overall abundance . 

Ophiuroids were identified to lower taxonomic levels from Cruise I samples 

only . There were 16 taza, 13 of which were identified to the species 

level, from Cruise I macrofauna. Because so many of the specimens were 

juvenile forms that could be identified to genus or species only with 

great effort and expenditure of time, the ophiuroid specialist, Dr. Gordon 

Handler, concluded that it was impractical to continue the effort on the 

remaining samples . On Cruise I, the highest species diversity for 

ophiuroids, seven tasa, was at Station C4 (1381 m) and the lowest, three 

taxa, was at C5 (275 m) . The most common ophiuroid at the Cent al 

Transact on Cruise I Were Ophiotholia sp . (at 853 and 1381 m) and 

AmyhileDis sp. (at 2475 m) . 
223 

Ophiuroid population densities ranged from 0-133/m2 with the highest 

station density found at W31(85u m) . Highest mean transect density, 

52/m2, was found at the East Transact on Cruise IV ; lowest, 26/m2, was at 

the Central Transact on Cruise III. 

Ophiuroid mean densities by depth were : 

Depth Range (m) 

298- 492 

500- 900 

1000-1500 

2000-2945 

Mean Density (No ./m2 

X10 

45 

37 

18 

u .2 .5 .30 Eehinoidea 

The echinoids ranked 26th in overall abundance of macrofaunal groups. 

This puts them in the lowest 1%. They ranked 20th in number of taxa, with 

only six taxa identified, three of which were identified to the species 

level and one to the generic level as shown below : 

Overall 

Density No . of No, of Depth 

Taucon No./ m2 Specimens Stations Rarxe ( m) 

Aceste bellidifera 2 .28 36 25 298-2945 

Hemiaster experAitus 0 .44 7 4 344-2485 

Aceste sp . 0 .19 3 3 352-1236 

Sehizaster orbignganus 0 .06 1 1 2475 

The two most abundant taxa of echinoids showed no particular depth 

limitations in the macrofaunal samples, being present from near the 

shallowest to the deepest depths sampled . Comparing with the megafaunal 

echinoids taken by trawl, we find that no Hemiaster exuergitus were taken 

in trawl samples and only limited numbers of Aceste bellidifera were taken 

by trawl on Cruise IQ at the E2 stations (615-630 m depth). By contrast, 

the small specimens of this species occurred in the macrofauna samples on 

all transacts and at all depth levels. 
224 

4 .2 .5 .31 Holothuroidea 

Holothuroids ranked 19th in overall abundance in the maerofauna and 

17th in total number of taxa. A total of 13 taxa were identified, four to 

the species level, four to the generic level, and five to family or higher 

levels . 

Greatest number of holothuroid taxa, four, occurred at three 

stations: E2B and E3D, Cruise IV (627 and 849 m) and WC11, Cruise V (1226 

m) . Highest transect diversity, eight taxa, occurred on the West-Central 

Transact on Cruise IV. No holothuroids were taken at the Central Transact 

on Cruise I . 

Holothuroid densities ranged from 0-84/m2 With the highest density 

(84/m2) reached at Station E2D, Cruise IV at 630 m. Highest transact 

average density, 37/m2, was at the East Transact on Cruise IV ; lowest, 

0/ m2, was at the Central Transact on Cruise I. Holothuroids occurred in 

considerably higher densities on the East Transact on both cruises to that 

transact (Cruises II and IV) compared to all the other transacts . 

Mean densities for holothuroids by depth were: 

Mean Density Mean Density 

Depth Range (m) (All Transacts) (East Transact Only) 

298- 492 5 13 

500- 900 27 48 

1000-1500 11 21 

2000-2945 5 7 

The eight holothuroid taxa identified to genus or lower in 

macrofaunal samples are listed below: 

Overall 

Density No . of No . of Depth 

Taaon ( Nod m2 Specimens Stations Range (m) 

Protankyra sp . 6 .96 110 25 352-2518 

Myriotrochus sp . 5 .37 85 23 34b-2901 

Echinocucumis hispida 0 .25 u 3 819-2101 

Pseudostichopus sp . 0 .25 4 2 1430-2101 
zzs 

Overall 

Density No . of No . of Depth 

Tauon No./ m2) Specimens Stations Range (m) 

Molpadia aAassizi? 0.19 3 3 598-1390 

Benthodytes sp . 0 .06 1 1 1465 

Molpadia sp . 0 .06 1 1 849 

Molpadia barbouri 0 .06 1 1 1226 

Whereas Protankyra sp. Was the most abundant holothuroid in the 

maerofauna samples taken at nearly all depths sampled, it was taken only 

twice in trawl samples, both times at Station C5 .on Cruise I and Cruise 

III (2400 and 2504 m depths) . Mgriotroehus sp., the second most abundant 

holothuroid in the macrofauna was not taken in trawl samples. Adults of 

this genus are in the maerofaunal size range and are a very common and 

diverse deep-sea genus (Robert J. Carney, pers. comm.) . The remaining six 

taxa of macroPaunal holothuroids were also fairly well represented in 

trail samples, the macrofaunal specimens most likely being juvenile forms. 

The most abundant megafaunal holothuroid, Mesothuria lactea , was not taken 

in macrofaunal samples. 

x .2 .5 .32 Aseidiacea 

Tunicates of the class Ascidiacea, ranked 21st out of the 44 

maerofaunal groups in overall abundance in this study. They ranked 12th 

in total number of taxa kith 18 taxa, 11 *of which were identified to the 

species level and three to the genus level. 

Ascidian diversities ranged from 0-5 taxa per station. The two 

stations showing the highest tunicate diversities were Station C9, Cruise 

III (1430 m) and E3C, Cruise IV (849 m) . Mean transeet diversities for 

tunicates ranged from two taxa at the Central and West Transacts on 

Cruises I and II, respectively, to 10 taxa at the East Transeet on Cruise 

IV. 

Ascidean population densities ranged from 0 to 49/m2, with the 

highest density at Station C4, Cruise II at 1390 m depth. The greatest 

mean transeet density, 13/m2, was at the Central Transact on Cruise II . 

The lowest mean transact density for ascidians was 5/m2 at the West- 
226 

Central Transact on Cruise Y. Ascideans were absent from 25 stations, 

most of which were located at the shallowest and the deepest depths. None 

were found at stations shallower than 550 m. 

The species of aseideans from this study are listed in order of 

decreasing abundances as follows : 

Overall 

Density No . of No . of Depth 

Species No./ mz Specimens Stations Range (m) 

Dicarpa simplex 5 .37 85 26 550-2101 

Bathystyeloides n . sp . 0 .57 9 ? 846-1465 

Miniyera uapillosa 0 .32 5 3 759-1165 

HeuacrobYlus indicus 0 .32 5 3 849-2901 

Polycarya pseudoalbatrossi 0 .32 5 3 748- 759 

Minnipera pedunculata 0 .25 u 3 2518-2941 

Synoicum draucum 0 .25 4 3 622- 849 

A new species of aseidean and many new distribution records are being 

described in a publication by Drs. Claude and Fraricoise Monniot, who note 

that the ascidean density on the northern Gulf of Mexico continental slope 

is the highest known for aseideans in the deep sea. They also report that 

the 10 previously described tunicate species all live in the South 

Atlantic Ocean and many in the basins bordering South America. It is 

concluded that the deep Gulf ascidean fauna is partly cosmopolitan and 

partly of a South Atlantic origin. 

4 .3 MEGAFAUNA 
229 

4 .3 MEGAFAUNA 

4 .3 .1 INTRODUCTION 

The megafauna discussions below are based upon the fish and 

invertebrate collections made by trawling. Complete listings of the catch 

data can be found in Volume III: Appendices, specifically Appendices C3-1 

through C3-11 . Life history information (e.g ., length-weight) was taken 

for selected species, these raw data can also be found in Volume III, 

Appendices C4-1 through C4-26 . The life history information along with 

food habit and distributional data were used to prepare species or life 

history accounts for selected megafaunal invertebrates and fishes . 

Collectively, these data help set the stage for the comprehensive 

synthesis of the final year. 

Below we first discuss the megafauna in terms of overall community 

structure and diversity and then discuss each mayor group, particularly 

emphasizing comparisons to the TerEco baseline and bathymetrie 

distributions . 

4 .3 .2 COMMUNITY STRUCTURE AND DIVERSITY 

In this section we describe the relative abundance patterns of 

fishes and invertebrates by transact and depth. Based upon previously 

defined lack of any definitive seasonal or annual patterns, these topics 

do not warrant special discussion. 

4 .3 .2 .1 Numerical Abundance by Transact 

Megafauaal invertebrates, based upon data from the five cruises, were 

some four to five times more abundant than fishes on each transact in 

terms of average density : 
230 

Number/ha 

TRANSECT : Western Central Eastern 

Fish 1222 620 1511 

Invertebrates 505 2621 7952 

Combined 6267 321 965 

The Central Transect was characterized by the lowest density of megafauna 

(3241/ha) which was some 50% lower than density observed on the Western 

Transect (6267/ha) and only about one-third of the density found for the 

Eastern Transect (94b3/ha) . 

4 .3 .2 .2 Numerical Abundance by Depth 

The total and mean densities (number per hectare) of demersal fish 

and invertebrates are presented by depth intervals on the three mayor 

transects in Table 4-5 " Note that there are six depth intervals 

beginning with 328-423 m and ending with 2100-2855 m on each of the three 

mayor transacts. The number of stations within each depth interval are 

not the same in the three transacts . For instance, in the case of the 

328-423 m interval, there are three stations on the West and Central 

Transects and five on the East Transact. Hence in the case of fish on the 

West Transect the sum of the densities at the three stations was 476 with 

an average number of individuals of 159/ha. Three mayor points stand out 

very clearly : (1) the densities of the invertebrates are consistently the 

larger, (2) densities are highest on the Eastern Transact and lowest on 

the Central Transact as described above, and (3) the densities tend 

to drop at and below the 1550 m isobath. A similar pattern was found for 

many of the macrofaunal groups . 

In Tables u-6 and 4-7, we show the relative total densities of the 

two dominant megafaunal invertebrate groups and species within groups, 

respectively . The point to be made is that only one or two species 

dominate the overall relative abundance patterns. One should note that 

the data in Tables 4-6 and 4-7 have not been adjusted by number of trawl 

tows and are only for comparison to one another, not to Table 4-5 . 
231 

Table 4-5 . Densities (individuals per neetare) o 

invertebrates and demersal and benthopelagic 

depth intervals on three mayor transects . 

adjusted by bottom trawl and trawl size ; CPE 

haul .) 

Depths 328-423 m 

Western Transact 

Fish Inverts 

Tot . 476 897 

CPE . 159 + 299 + 

Central Transact 

Fish Inverts 

197 345 

66 + 115 + 

r megafaunal 

fishes at six 

(Tot . = Catch 

Density/trawl 

Eastern Transact 

Fish Inverts 

704 2030 = 

141 +. 406 -_ 

Depths X65-750 m 

Western Transeet 

Fish Inverts 

Tot . 469 1960 

CPE . 67 + 280 + 

Central Transeet 

Fish Inverts 

175 732 

58 + 241 + 

Eastern Transeet 

Fish Inverts 

347 4368 

50 + 624 

Depths 751-1000 m 

Western Transact 

Fish Inverts 

Tot . 142 605 

CPE . 47 + 202 + 

Central Transact 

Fish Inverts 

171 900 

34 + 180 + 

Eastern Transact 

Fish Inverts 

239 819 

40 + 117 = 

Depths 1050-1500 m 

Western Transact 

Fish Inverts 

Tot . 135 1511 

CPE . 45 + 514 + 

Central Transact 

Fish Inverts 

56 273 

19 + 68 + 

Eastern Transact 

Fish Inverts 

219 715 

109 + 357 = 

Depths 1550-2050 m 

Western Transeet 

Fish Inverts 

Tot . 0 0 

CPE . 0 0 

Central Transeet 

Fish Inverts 

>> >s2 

+ 61 

Eastern Transact 

Fish Inverts 

0 0 

0 0 

Depths 2100-2855 m 

Western Transact Central Transeet Eastern Transact 

Fish Inverts Fish Inverts Fish Inverts 

Tot . 0 42 10 189 2 20 

CPE . 0 21 + 3 + 63 0 0 

Mean 

198 

Mean 

1342 

221 

Mean 

412 

103 

Mean 

490 

185 

Mean 

QZ 

11 

Mean 

44 

15 
N 

W 

N 

Table 4-6 . Total densities of the most abundant megafaunal invertebrate groups for sampled 

depth intervals . 

23 8-423 465-750 751-1000 1050-1500 1550-2050 2100-2855 

Decapoda 

Penaeidea 318 8 .5 8 .0 12 .5 0 4 .15 

Brachyura 64 195 .5 12 .5 6 .o 18 1 .5 

Galatheidea 31 .5 102 9 6 .5 3 1 

Caridea 32 202 .5 80 293 11 2 . 

Anomura 3 7 9 4 3 2 .5 

Echinodermata 

Asteroidea 5 22 24 9 64 34 

Holothuroidea 3 .5 6 .5 15 6 1 " 

Echinoidea 3 .0 2 .5 2 .5 1 .5 0 

Ophiuroidea 13 " 0 140 .0 24 .5 26 2 .5 82 

Crinoidea 0 2 .5 2 .5 2 0 
Table 4-7 " Densities of the most abundant species of deoapoda that account for the density data assn in Table X1-6 . 

Ave: 375 608 876 1275 1800 247th 

Ta:on Range : (328-423) ( 465-75o ) (751-1000) (1050-1500) (1550-2050) (2100-2855 

PENAEIDEA (All app .) 318 9 8 12 ? 5 

PenaeoDSis serrata 261 2 0 0 0 0 

CARIDEA (All app.) 32 243 80 293 12 3 

Nematocaraiaus rotundus 0 90 39 188 1 0 

Plesionika holthuisi 1 94 3 0 0 0 

Glgphocrangon aculeata 0 0 8 25 

HRACHYIIRA (All app .) 64 196 13 6 18 2 

Benthochascon schmitti 31 62 1 0 0 0 

BathSrplaz tgphla 11 129 7 2 0 0 

Ger9on cuinauedens 1 3 4 3 12 

GALATHEIDBA (1111 sPP .) 32 102 9 7 3 

Munida lonttipes 17 0 0 0 0 0 

Munida valida 0 94 2 1 0 0 

Munidopsis s sbei 0 1 5 1 1 1 

N 

W 

W 
234 

x " 3 .2 .3 Species Diversity by Region grid Depth 

Species diversity (H" and number of species) of megafaunal 

invertebrates and fishes, by region and depth, are presented in Figures 4- 

9 through 4-31, and 4-32 through 4-34, respectively. For invertebrates, 

there were no clear trends in the diversity index by depth nor were there 

any apparent substantive differences in invertebrate diversity among the 

three regions. Using 30 species as a reference point, 80% of the 

collections taken on the Central Transect had fewer than 30 invertebrate 

species whereas only about 30% of the collections on both the Eastern and 

Western Transacts had fewer than 30 species (Figs. u-29 through 4-31) . At 

least part of this difference is attributable to more samples being taken 

at depths below 1250 m on the Central Transact than on either the Western 

or Eastern Transacts. Depths below 1250 m appeared characterized by faker 

species than shallower depths. 

Examination of the fish diversity data suggests a decline in 

diversity as well as number of species at depths exceeding 1250 m (Figs. 

4-32 through 4-34) . There were no obvious differences in fish diversity 

among the three regions . 

4 .3 .2 .1 Bathymetric Zonation of MeRafauna 

The results in the first two Annual Reports emphasized vertical 

zonation discussions drawing upon data from Cruises I through III. The 

preliminary conclusions were that a ease could be made supporting the 

zonation scheme proposed in the TerEco baseline (Pequegnat 1983) . 

Cluster Analyses 

The data derived from Cruises IV and V have provided support for a 

conception presented in earlier reports as to how faunal assemblages are 

distributed over the upper and middle portions of the continental slope . 

For the past several years, oceanographers have differed as to whether or 

not benthic animals are arrayed on the slope in discernible vertical bands 

or zones. Although there is general agreement that the composition of the 

faunal assemblages changes with changes in depth, there are those who 
235 

Diversity of Invertebrate Species on the Eastern Transect 

a 

3 

a r 

,L 4 

I 

O 

1 

0-- 

250 

Number of Invertebrate Species on the Eastern Transect 

120 

100 

h 

80 

v 

4 

d 

N 

v . 60 0 

r 

40 

0 

s 

20 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (1n) 

Figure 4- 29, Species diversity and number of species of megafauna 

invertebrates for the Eastern Transect, Cruises I-V. 

500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (m) 
236 

4 

3 

a 

1A 2 

4 

I 

O 

1 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (m) 

Number of Invertebrate Species on the Central Transect 

60 

50 

N 

~ ao 

"H 

v G 

N 

y. 30 

e 

V 

20 

0 

10 

Figure 4- 30 . Species diversity and number of species of megafauna 

invertebrates for the Central Transect, Crusies I-V. 

Diversity of Invertebrate Species on the Central Transect 

r--~ 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (m) 
237 

Diversity of Invertebrate Species on the Western Transect 

4 

3 

T 

.N r 

L 

4 

C 

2 

Number of Invertebrate Species on the Western Transect 

goo 

so 

N 

4 

60 

N 

w 

0 

40 

a 

E 

s 

20 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth 

Figure 4-31, Species diversity and number of species of megafauna 

invertebrates for the Western Transect, Cruises I-V. 

t 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth 
238 

Diversity of Fish Species on the Eastern Transect 

5 

4 

a 3 

r 

N 

L 

4 

1 

O 

I 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth 

Number of Fish Species on the Eastern Transect 

40 

30 

H 

4 

V 

4 

a 

w 20 

0 

L 4 

a 

E 

i to 

0 +-

250 

Figure 4-32 . Species diversity and number of species of fishes for 

the Eastern Transect, Cruises I-V. 

500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth 
239 

Diversity of Fish Species on the Central Transect 

5 -

4 

a 3 

w 

r 

N 

L 

4 

O 2 

1 

0 

250 

30 

~ 20 

v 

4 

d 

N 

O 

L 

a 10 

E 0 

0+- 

250 

Number of Fish Species on the Central Transect 

Figure 4- 33 . Species diversity and number of species of fishes for 

the Central Transect, Cruises I-V. 

500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth 

500 750 1000 1250 1300 1750 2000 2250 2500 2750 3000 

Depth 
240 

5 

4 

a .. 

1A 

L 

4 

0 

t 

Diversity of Fish Species on the Western Transect 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (m) 

Number of Fish Species on the Western Transect 

40 

30 

VI v 

V 

4 

a 

N 

20 

e 

4 

a 

E 0 

z 10 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 

Depth (m) 

2750 3000 

Figure 4-34 . Species diversity and number of species of fishes for 

the Western Transect, Cruises I-V. 
241 

believe that the so-called zones are artifacts of sampling, i.e., that the 

limits of the zones are wholly related to the depths at which sampling was 

done . One must point out, however, that Haedrich et al . (1975, 1980) 

identified four faunal zones on the U.S. North Atlantic slope, and in 1983 

Hecker et al . in an MMS report designated five faunal zones on the same 

slope . In the same pear, Pequegnat (1983), again in an M MS report, 

identified five mayor zones on the slope and rise of the Gulf of Mexico . 

More recently the 1986 report on the MMS study of the U.S. North Atlantic 

benthic fauna carried out by Battelle, Lamont-Doherty, and Woods Hole 

contains descriptions of six faunal zones on the slope. These findings, 

which are based upon both classification and ordination analyses, reveal 

that the replacement of species with depth is not uniform. In other words 

the "rate of replacement" is not the same on all vertical aspects of the 

slope . Accordingly, one may regard zones as being large areas of 

relatively small faunal change with depth that are separated by smaller 

areas where the rate of exchange of species is high. These separations 

have been referred to as "breaks", but this is not a wholly acceptable 

term because it tends to ignore the fact that few or many taxa may be 

shared between neighboring zones. 

It was reasoned that if this concept of zones is correct, then there 

should be far fewer exchanges of faunal species sampling along isobaths 

than sampling up or down the slope. And, by the same token, sampling 

along successively deeper isobaths should show increasing degrees of 

difference when compared with the samples from the shallowest isobath. 

Thus, it was decided to trawl isobathqmetrieally on consecutively deeper 

isobaths along two transects, namely, the East Transect during Cruise IV 

(Fig . 4-35) and the West-Central Transact during Cruise V, as shown in 

Figure 4-36 " Note that Stations E1, 1a, 1b, and 1c are arrayed slightly 

above or below one isobath at a depth of about 350 m, whereas 2, 2a, etc., 

and 3, 3a, 3b, etc . are on isobaths having depths of about 625 and 840 m, 

respectively. Somewhat the same depth relationships were established on 

the West-Central Transact but the trawling tended to stray off the isobath 

to a greater degree than on the East Transact (Fig . 4-36) . In particular 

note the large depth ranges at Stations WC6 and WC11 . 
242 

CRUISE IV EASTERN TRANSECT STATIONS 

300 - 

400 - 

500 - 

E 

z 

600 - W 0 

o800- 

900- 

EI 1A 1B IC E2 2A 28 2C 2D 2E E3 3A 38 3C 3D 

I 

I = 

1 I 

Figure 4-35 . The range of depth (length of vertical lines) covered by 

trawling isobathymetrically as nearly as possible 

along three "horizontal transects" of increasing depth 

(E1 to E3) in the Eastern sampling area . 
243 

CRUISE V WEST-CENTRAL TRANSECT STATIONS 

500 

400 

5 / ti 4 L 6 IV y 3 11 IG 

t 

500 

I 

600 

TOO 

E 

z 

a 800 

w 0 

900 

1 000 

1100 

1200 

1300 

I 

I 

Figure 4- 36 . The range of depth covered at individual stations by 

trawling isobathymetrically as nearly as possible along 

three "horizontal transects" of increasing depth in the 

West-Central sampling area . 
244 

The Normalized Expected Species Shared (NESS) similarity measure 

(Grassle and Smith 1976) was used to analyze data derived from Cruises IV 

and V. Figure 4-37 shows the dendrogram that results from the application 

of NESS to all of the megafaunal invertebrates taken during Cruise IV . 

There is a clear separation of the faunal clusters from one isobath to 

another. Moreover, the E2 and E3 series of stations are more closely 

related to one another than to the E1 series. The separation of Station 

E2E from the others on the isobathresults from the fact that it lacked a 

few species and had others in reduced numbers as compared with other 

stations on the isobath. This gave it a pattern that resembled those more 

typical of the E3 depth series. Essentially the same mayor clusters occur 

when the fish data are analyzed (Fig . 4-38), but there are some 

differences in station pairings. Note, however, that in the E1 series of 

four stations that E1, 1B, and 1C are closely related and E1 A is the 

outlier in both figures . In the E2 series Stations E2 B, 2C, and 2D are 

clustered together with E2 as an outlier in both figures. The E3 series 

of five stations is much more tightly clustered for invertebrates than 

fishes and the pairings do differ except that Stations E3A and E3C are 

tightly clustered in both, although with different partners . 

The dendrograms (NESS) for Cruise Y invertebrates and fishes in the 

West-Central region, as shown in Figures 4-39 and 4-40, again show a 

tendency toward isobathic clustering with two mayor clusters, each of 

which is subdivided into two. Reference to Figure 2-1 (map of stations) 

and Figure 4-36 shows that the trawling was not carried out as 

effectively here as on Cruise I0. Stations are closely clustered where 

the range between the shallowest and deepest parts of the sampling was 70 

m or less. In the ease of WC6, the range was over 240 m . This depth 

difference will not always make much difference unless it crosses a zone 

of species flux. The clusters derived from the invertebrate and fish data 

are remarkably similar. Note, for example, that in both Figures 4-39 and 

4-40 Stations WC1 and WC5 are paired and Stations WC2, u, 7, and 8 form a 

cluster, as do NC3 and 9 and WC11 and 12, and in both WC10 is an outlier. 

It is only Station WC6 that shifts its position, clustering with the 
245 

Dissimilarity (NESS (m=30)) 

158 .445 .733 1 .000 

Station ElA (351 m) 

Station E1B (344-346 m) 

Station E1 (351-357 m) 

Station E1C (347-353 m) 

Station E2 (613-618 m) 

Station E2C (616-620 m) 

Station E2A (624 m) 

Station E2B (600-625 m) 

Station E2D (624-631 m) 

Station EZE (629 m) 

Station EM (783-841 m) 

Station E3 (871 m) 

Station E3B (823-845 m) 

Station E3C (838-847 m) 

Station E3D (845-856 m) 

Figure 4-37 . All invertebrates taken by trawl from the 15 stations 

of Cruise IV on the Eastern Transect clustered by NESS 

at 30 individuals . Similarity values between pairs of 

stations equal the reciprocal of the values given at 

the top of the figure . The major isobathic clusters 

are easily separated . Outlier stations such as ElA 

result less from depth than from sampling . 
246 

Dissimilarity (NESS (m=30)) 

.188 .465 .742 1 .000 

Station ElA (351 m) 

Station E1 (351-357 m) 

Station E1B (344-346 m) 

Station E1C (347-353 m) 

Station HE (629 m) 

Station E2A (624 m) 

Station E2 (613-618 m) 

Station E2D (624-631 m) 

Station E2B (600-625 m) 

Station E2C (616-620 m) 

Station E3B (823-845 m) 

Station E3 (871 m) 

Station EX (838-847 m) 

Station BA (783-841 m) 

Station E3D (845-856 m) 

Figure 4-38 . All fishes taken by trawl from the 15 stations of 

Cruise IV on the Eastern Transect clustered by NESS 

at 30 individuals . Comparison with invertebrate 

clusters in Figure 4- 37 shows a high level of 

similarity . 
247 

Dissimilarity (NESS (m=30)) 

214 .470 .725 .980 

Station WC1 (344-393 m) 

Station WC5 (402-444 m) 

Station WC8 (472-486 m) 

Station WC7 (457-472 m) 

Station WC4 (516-527 m) 

Station WC2 (518-585 m) 

Station WC10 (741-752 m) 

Station WC6 (543-783 m) 

Station WC9 (695-807 m) 

Station WC3 (768-781 m) 

Station WC11 (1024-1247 m) 

Station WC12 (1170-1236 m) 

Figure 4-39 . All invertebrates taken by trawl from the 12 stations of 

Cruise V clustered by NESS at 30 individuals . 
248 

Dissimilarity (NESS (m=30)) 

265 .518 .770 1 .000 

+ a--f f +-r-~ 

Station WC1 (344-393 m) 

Station WC5 (402-444 m) 

Station WC6 (543-783 m) 

Station WC2 (518-585 m) 

Station WC7 (457-472 m) 

Station WC4 (516-527 m) 

Station WC8 (472-486 m) 

Station WC9 (695-907 m) 

Station WC3 (768-781 m) 

Station WC10 (741-752 m) 

Station WC11 (1024-1247 m) 

Station WC12 (1170-1236 m) 

Figure 4- 40 . All fishes taken by trawl from the 12 stations of 

Cruise V clustered by NESS at 30 individuals . Comparison 

with the dendrogram of Figure 4-39 shows that WC6 is 

the only station that shifts position between the two 

major branches . Otherwise note WCl and 5, and WC11 and 

12, etc . are very similar . 
249 

deeper invertebrate stations and the shallower fish stations . This 

suggests that on the West-Central Transect invertebrates are more abundant 

at deeper stations and fishes at shallower stations . Thus, it would 

appear that during the vertical transit of 240 m (from 543 to 783 m) of 

trawling at Station KC6, relatively more fishes were taken at shallow 

depths and more invertebrates during the deeper part of the haul . This 

suggests that a zonal change occurs somewhere between 540 and 780 m depth. 

NESS analysis was applied to all of the megaf aunal invertebrates 

taken by trawl during Cruise III on the Central Transact, and Cruises IV 

and V on the East and West-Central Transacts (Fig. 4-41) . There is very 

evident isobathic clustering, but Stations C9 and C10 stand apart . 

Reference to the cruise log for Cruise III revealed that in both cases the 

trawl was filled with a huge mud lump that prevented proper sampling. In 

spite of the inclusion in the NESS of some stations where the trawling was 

not successful, there are some reasonably good clustering with possible 

zonal separations. Stations C5 and C11 tend to reflect the Mesoabyssal . 

Stations WC1, E1 B, E1C, E1, E1A, C1, and WC5 compare well with the ShelfSlope Transition. Most of the rest of the stations appear to be horizons 

of the Arehibenthal Zone with the exception of Stations C4, C8, WC11, and 

WC12, which are very likely Upper Abyssal, as their species lists also 

demonstrate . 

The bathymetric distributions of some of the dominant species of 

fishes and invertebrates found on the East Transact that account for the 

clusters shown above are displayed in histograms in Figures 4-42, u-43, 

and 4-4u. Recall that the depth range of the E1 isobath is about 350 m, 

of E2 is 620 m, and of E3 is about 850 m . There is little overlap of 

isobaths by the dominant species, even though some species may have wide 

bathymetrie ranges (note Dibranchus atlanticus in particular) . The 

decapods shown in Figure 4-43 appear to have sharply limited bathymetric 

distributions; however, it should be noted that the penaeid shrimp 

Penaeopsis serrata lines at shallower depths in the Shelf-Slope Transition 

Zone, whereas Benthesicymus bartletti penetrates much deeper than the 850 

m depth of the E3 series of stations. The carideans in Figure 4-43 on the 

other hand, have distinctive depth limits that fall within the station 

depths shown Essentially the same may be said about the braehyuran crabs 
zso 

Dissimilarity [NESS (m=30)J 

STATION Depth Range 

C5 (2486-2523 m) 

C11 (2063-2085 m) 

WC10 (741-752 m) 

WC1 (344-393 m) 

E1B (344-346 m) 

E1C (347-353 m) 

E1 (351-357 m) 

E1A (351 m) 

C1 (326-346 m) 

WC5 (402-444 m) 

WC6 (543-783 m) 

C2 (632 m) 

WC7 (457-472 m) 

WC8 (472-486 m) 

WC4 (516-527 m) 

C6 (448-501 m) 

WC2 (518-585 m) 

E2 (613-618 m) 

E2A (624 m) 

E2D (624-G31 m) 

E2B (600-6?_5 m) 

E2C (616-620 m) 

E2E (629 m) 

C3 (764-841 m) 

C7 (896-1033 m) 

C4 (1420-1600 m) 

CS (1064 m) 

WC3 (768-781 m) 

WC9 (695-807 m) 

WCil (1024-1247 m) 

WC12 (1170-1236 m) 

E3A (783-841 m) 

E3B (823-845 m) 

E3 (871 m) 

E3C (838-847 m) 

E3D (845-856 m) 

C9 (1309-1317 m) 

C10 (1600-1790 m) 

Figure 4-41 . All invertebrates from Cruises III (Central Transect), 

IV (East Transect), and V (West-Central Transect) . 

.338 .502 .665 .829 
251 

60 

40 

20 

0 

BEMBROPS GOBIOIDES 

60 

40 

0 

20 

0 0 

W 

m 

40 

20 

DIBRANCHUS ATLANTICUS 

BATHYGADUS MELANOBRANCHUS 

0 

20 

DICROLENE SP. 

E I ABC 2 A B C D E 3 A B C D 

STATION 

Figure 4-42, Numbers of individuals of four common fishes 

collected from the 15 isobathymetric stations of 

Cruise IV . All stations in each of the three groups 

are about the same depth (El series around 350 m ; 

E2 series about 620 m; and the E3 series about 850 m) . 
252 

600 

J 

Q 

400 

D 

J 

D 

Z 

- 200 

O 

W 

m p 

20 

10 

O 

STATION 

PLESIONIKA TENUIPES (Corideon shrimp) 

PLESIONIKA HOLTHUISI lCorideon shrimp) 

O E 1 ABC 2 a s c V t a A o V v 

STATION 

Figure 4- 43 . Results of isobathymetric trawling during Cruise IV 

on the Eastern Transect . Histograms show the number 

of individuals of two penaeid shrimp species (upper 

pair) taken on three isobaths ; the lower pair show 

the same discrete depth distributions of two caridean 

species at the various stations on the same three 

isobaths as shown above . Depths as in Figure 4-87 . 

30 

Q 20 

10 

O 

D 

40C 

30C 

200 

IOC Z 

E I ABC 2 A B C D E 3 A B C D 
253 

J 40 

a 

0 20 

0 0 z 

300 

w 

200 

100 

0 

BENTHOCHASCON SCHM ITT I 

60 

BATHYPLAX TYPHLA 

E I ABC 2 A B C D E 3 A 8 C 0 

STATION 

Figure 4-44 . Numbers of individuals of two common brachyuran crab 

species collected from the 15 isobathymetric stations 

of Cruise IV . Depths as in Figure 4-87 . 
254 

shown in Figure 4-44 . The few individuals of the very common Bathyplax 

typh1a shown in the E3 series were very young individuals. The young of 

brachyurans are often found in reduced numbers at depths deeper than those 

occupied by adults. 

It bas long been suspected that the chirostylid Uroptychus nitidus 

lives among the branches of the gorgonian coral Chrysogorgia aAassizii . 

The concordance in their station and isobathic distributions, as seen in 

Figure 4-45, lends credence to the belief . Also, the finding of a whole 

flat-backed lobsterette ( Stereomastis sculpta) in the stomach of an eel 

( Synaphobranchus oregoni) led to a debate as to whether the event was 

happenstance or a normal predator/prey relationship. The concordance in 

their bathymetrie and numerical distributions seen in Figure 4-46 makes 

the latter relationship plausible . 

Chi-Square Analyses 

A summary of the results for fishes of the isobathymetrie sampling on 

the Eastern Transact during Cruise IV is shown in Table u-8. Here we see 

a separation of the "starts and stops" for species which form the basis 

for the chi-squared analysis of zonal distribution discussed in the next 

section. Results Prom Stations WC11 and 12 are added for comparison with 

a deeper depth band. 

For detecting faunal boundaries we have utilized the method devised 

by Baekus et al . (1965) and employed more recently by Gage (1986) in 

examining bathy metric zonation of the benthic fauna in the Rockall Trough 

south of the Wyville Thomson Ridge. In essence the method uses 

presence/ absence data to plot the distribution of the sum of first-pluslast captures at between-station intervals on a transact having a gradient 

of increasing depth. If indeed the zonal concept is an acceptable way of 

describing bathymetric distribution, where areas of faunal homogeneity are 

separated by narrow regions of species change, upper and lower limits 

should occur concurrently more frequently than expected on the basis of 

chance . Comparisons were made between expected values for collection 

intervals and those actually observed using the chi-square test and 

obtaining probability levels . The chi-squared values, which can be 

thought of as indices of faunal change for each depth interval, are then 
255 

160 

120 

co 80 

J 

Q 

0 40 

0 0 

z 

UROPTYCHUS NITIDUS (Chirostylid onomuron) 

400 

w 

g 300 

z 

200 

100 

0 

I CHRYSOGORGIA AGASSIZII (Gorconion coral) 

E I A B C 2 A B c v E 

STATION 

3 A C V u 

Figure 4-[,5, The depth and station distribution patterns of a 

gorgonian coral and chirostylid decapod as shown 

in the figure from 15 stations of Cruise IV on the 

Eastern Transect lend further credence to the 

conclusion that the anomuran lives in close association with the gorgonian which builds its holdfast in 

the soft sediments . 
256 

10 

11 ABC, 1 2 A B C D E i ~ 3 A 8 C D ~ 

344-357 m 600-631 m 783-871 m 

O 

SYNAPHOBRANCHUS OREGONI - Predator (Eel) 

STEREOMASTIS SCULPTA ~ Prey (Polychelid decapod) 

Figure 4-46 . The belief that the eel preys upon the polychelid is 

reinforced by the concordance in their distribution 

among 15 stations of Cruise IV on the Eastern Transect . 

Note also that at most stations the prey is more abundant 

than the presumed predator . 

N 

50 CRUISE IV EASTERN TRANSECT STATIONS 

40 

30 

4- 0 

S- 20 

-71 
Table ". sunaary of starts and stops of fishes trawled on the laobathyaeLrio series of 15 stations on the Eastern 

Transact during Cruise IV. 

IsobathSrmetric Traasect Analysis 

Fish E1 (A.B,C) Transact E2 (A .B,C,D,E) Transact E3 (A.B .C .D) Transact 

a) Transact depth range 344-357 m 613-629 m 783-871 m 

b) No . of fish species taken on transact 38 36 30 

c) No . of species taken at 3 or more (2 on WC) 

stations on traasect 17 13 13 

Percent of total 40% 36% 43% 

d) No . of species present at E1 but not E2 30 

"Stops" 

e) No . of species present at E2 but not E1 28 

"Starts" 

f) No . of species common to E1 and E2 8 

Percent of total species 14% 

g) No . of species present at E2 but not E3 17 

"Stops" 

b) No . of species present at E3 but not E2 15 

i) No . of species common to E2 and 83 14 

Percent of total species 30% 

3) No. of species common to E1 and E3 2 2 

N 

V 
258 

plotted against depth on the abeissa . Peaks in the graph mark depth 

intervals of maxima in the rate of faunal change . Valleys on the graph 

are interpreted as being relatively homogeneous, a fact that our 

isobathymetrie sampling tended to demonstrate . 

The chi-squared analysis of the distribution of fish species together 

with species recruitment curves are shown in Figures 4-u7 through 4-52 . 

All of the peaks above the alpha line are significant and indicate depth 

intervals of high species flux . In other words, they mark the depthlocation of boundaries between zones. The valleys on the other hand 

position the zones where the species composition of the slope is uniform. 

On the Central Transect there are strong peaks at depths around 450-550 m, 

675-750 m, 1000 m, and 2200 m (Fig . 4-48) . These are close to those 

designated as faunal breaks for the northern Gulf of Mexico by Pequegnat 

(1983) . Note the corresponding upsweeps in species numbers in the 

recruitment curve in Figure 4-47 . Chi-squared and species recruitment 

values for fishes on the Western Transact are shown in Figures 4-49 and 4-

50 . Again there are peaks around 450, 700, and 1000 m, and deeper, but 

the latter depths are not reliable because of the limited amount of 

sampling at deeper depths on this transact . For the Eastern Transact, 

peaks occur around 300, 500, 1000 and 2000 m (Fig . 4-52) ; again, little 

sampling occurred at depths at or greater than 1000 m. 

Results for the invertebrate chi-square analysis are presented in 

Figures 4-53 through 4-58. The chi-squared analysis of invertebrate 

species on the Central and Western Transacts differs from that for fishes 

on these transacts in that only one mayor peak is found on each transact . 

On the Central Transact the mayor peak lies between 850 and 1050 m depth, 

with minor peaks at 650-700, 1350, and one beginning at 2250 m (Fig. u54) . The recruitment curve shows a weak increase at 625-650, a strong one 

at 775 m, one beginning at 1000 m, and others at 1400 and 2400 m (Fig . 4-

53)- 

The invertebrate chi-squared values on the Western Transact are much 

more clearly defined than those on the Central Transact at depths less 

than 1300 m (Fig . 4-56) . The first peak is at 450 m, followed by a very 

strong one between 700 and 750 m, a smaller one at 1000 m and another near 

1300 m. The very limited sampling at depths greater than 1250 or so m on 
259 

70 

60 

H 

v 

50 

N 

w 

e 

v 40 

a 

E 

x 

30 

20 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth 

Figure 4-47 . Species recruitment curve for fish species taken at all 

stations on the Central Transect . 

20 

15 

v 

4 

L 

A 

10 P 

N 

t 

S 

Figure 4-48 . Chi-squared values for fish species taken on the Central 

Transect . Peaks above alpha line are depth intervals of 

high rate of species change . 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (m) 
260 

70 

60 

z ,50 

4 

i 

A 

40 

E 0 

U 

30 

20 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (M) 

Figure 4-1+9 . Species recruitment curve for fish species trawled on the 

Western Transect . 

20 

15 

v 

L A 

C IU 

N 

c G .1 

5 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (M) 

Figure 4-50 . Chi-squared values for fish species taken on the Western 

Transect . 
261 

70 

60 

~ 50 

V 

a 

'" 40 0 

a 

E 30 

z 

20 

10 

230 300 750 1000 1230 1300 1730 2000 2250 2500 2730 3000 

Depth 

Figure 4-51 . Species recruitment curve for fish species trawled on 

the Eastern Transect . 

zo 

15 

v 

4 

L 

A 

10 

t 

V 

5 

0 

250 

Figure 4-52 . Chi-squared values for fish species taken on the Eastern 

Transect . 

500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (m) 
262 

160 

Sao 

 120 

d 

V 

a 100 

N 

v 

e 

80 

r 

a 

E 

, 60 

z 

ao 

20 

215o 5oo 75o 1ooo 125o 15oo 1750 2000 2250 2500 2750 3000 

Depth (m) 

Figure 4-53 . Species recruitment curve for invertebrate species trawled 

from the Central Transect . Compare with Chi-square values 

below. 

40-

30 -

v 

v 

w 

P 20 - 

N 

U 

10-

0-- 

250 

Figure 4-54 . Chi-squared values for invertebrate species trawled from 

the Central Transect . 

500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (M) 
263 

200 

175 

150 

H 

4 

y 125 

4 

D 

w 100 

0 

v 75 d 

E 

z 50 

zs 

0 

25o 15oo 75o 1ooo 12so 15oo 175o 2ooo 225o 25oo 275o 3000 

Depth 

Figure 4-55 . . Species recruitment curve for all invertebrates trawled 

on the Western Transect . 

80 

60 

v 

r 

w 

40 

L 

v 

20 

O 

250 500 750 1000 1250 1500 1750 

Depth (M) 

Figure 4-5G . Chi-squared analysis of invertebrate species trawled on the 

Western Transect . 

2000 2250 2500 2750 3000 
264 

200 

v 150 

u v 

n 

N 

100 

u 

a 

E 

z 

50 

Figure 4- 57 . Species recruitment curve for all invertebrates trawled 

on the Eastern Transect . 

150 

125 

100 

4 

L 

A 

Q 75 

N 

r 

v 50 

25 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (m) 

Figure 4-53 . Chi-squared analysis of invertebrate species trawled on the 

Eastern Transect . 

0 

250 500 750 1000 1250 1500 1750 2000 2250 2500 2750 3000 

Depth (M) 
265 

this transact make further analysis unreliable. The species recruitment 

curve has very pronounced increases that aid one in interpreting the chisquared values (Fig. 4-55) . 

For the Eastern Transact, the mayor faunal discontinuities are 

indicated for the 500 and 625 to 750-m depth zones (Fig . 4-58) . The 

species recruitment curve has the most pronounced increase at about 625 m 

(Fig . 4-57) . 

x " 3 .3 MAJOR INVERTEBRATE GROUP DESCRIPTIONS 

Below we discuss the data obtained for the mayor invertebrate groups 

collected by trawling. Species accounts are provided for selected decapod 

crustaceans in Section 4.3 .4. Distribution and life history descriptions 

are given for selected individual fish species (rather than group) in 

Section u .3 .5 . 

x " 3 .3 .1 PoriPera 

Seventeen species of sponges were collected in the trawl. The five 

most abundant species were the small Polymastia sol (329-2857 m), Thenea 

sp . (612-1203 m), Thenea sp . A (621-2504 m), Tethya sp. (612-1203 m), and 

Hyalonema sp . (479-1510 m) . 

x " 3 .3 .2 Coelenterata 

Hydrozoa 

Five species of hqdrozoan polyps were collected in the trawl. The 

three most abundant were Lytocarpia bispinosa (345-871 m), Eudendrium 

euimium (618-871 m), and Cladocarpus tenuis (618-629 m) . 
266 

Anthozoa 

Antipatharia and Ceriantharia . Three undeseribed species of 

Antipatharia and one species of Cerianthid were taken in small numbers 

from the upper slope. 

Alcyonaria. Thirteen species of aleyonarians were collected in the 

trawl . The majority of these were gorgonacean corals of which 

ChrysoAOrgia aRassizii and Acanella eburnea were most abundant. 

ChrysogorAia was found at 15 stations ranging in depth from 350 to 1170 m, 

and Acanella occurred at 11 stations ranging in depth from 600 to 2857 m" 

Aetiniaria and Zoanthidia. Among the sea anemones the species 

ActinauAe longicornis was by far the most abundant. It has a very wide 

depth range from 329 to 2500 m and a wide geographic range, being an 

important component of the upper slope fauna in the North Atlantic 

(Maciolek et al . 1986) . 

Zoanthidians were common epizoans on pagurid decapods. 

Scleractinia. Seven species of stony corals were collected in the 

trawl. The most abundant of these were Deltocyathus italicus (551-1203 

m) ; Caryophyllia ambrosia caribbeana (774-1203 m) ; and SteDhanocyathus 

diadema (964-1510 m) . 

x " 3 .3 .3 Annelida (Polychaeta) 

Sixty-five species of polychaetes were collected in the trawl. The 

two most abundant species by far were the onuphid Hyalinoecia tubicola 

(612-1510 m) and the eunicid Eunice noeAica (350-1064 m), followed by the 

ampharetid Isolda pulchella (612-663 m), the polynoid Eunoe sp. A (474-

1510 m), the sigalionid Fhlersileanira incisa (479-?51 m), and the 

polynoid Eunoe hubreehti (368-663 m) . 
267 

4 .3 .3 .4 Mollusca 

Bivalvia 

Forty species of bivalves were collected in the trawl . Fifteen of 

these species were represented by only one or two individuals, and only 12 

of the remaining 25 species were represented by 10 or more individuals. 

The most abundant of the 12 were Propeamussium sp . A (774-1510 m), sp. C 

(612-200 m), and sp. D (338-1438 m) ; Amygdalum of. politum (345-1135 m), 

Anodontia ef. philippiana (342 m), Lithophaga ef. antillarum (62?-1203 m), 

and Propeamussium dalli (338-1438 m) " 

Gastropoda 

Forty-five species of gastropods were collected in the trawl. Only 

eight species were represented by 10 or more individuals. In fact 17 

species were represented by a single individual . The most abundant 

gastropod species were Leucosyring tenoceras (774-1510 m), Buceinum 

eanetae (612-964 m), Hyalorisia gales (338-774 m), Solariella lamellosa 

(612-629 m), Oocorys sulcata (2074-2504 m), and Scaphander Watsoni (329- 

346 m) . 

Cephalopods 

Thirteen species of cephalopods were collected in the trawl. Most of 

the species were represented by one or two individuals, but five species 

are worthy of specific mention: Alloposus mollis (354-663 m), Rossia 

bullisi (342-621 m), Benthoeropus anuari (u64-551 m), and Pholidoteuthis 

adami (627-1170 m) . 

Scaphopoda 

Five species of tooth shells were collected in the trail. The most 

abundant species were Dentalium perlongum , which has a wide depth range 

from 840-2400 m, Laevidentalium callipeplum that occurs only at shallow 
268 

depths from 315 to 368 m, and Dentalium eircumeinetum , which occurs at 

intermediate depths around 840 m . 

u " 3 .3 .5 Arthropoda 

One hundred and sixty-three species of benthic and benthopelagic 

crustacea and other arthropods were collected in the trawl from the mayor 

transects and isobathymetric station-lines established in this study (see 

Table 4-9) " Fifty-nine species or 36% of the 163 species were collected 

for the first time in Cruises IV and V. Forty of the 59 species were 

decapods. Most of these were either galatheid anomurans or brachyuran 

crabs, both of which doubled their species totals over those of the first 

three cruises. Several of these are either new species or represent 

first-time collections in the Gulf or the entire western Atlantic. 

About 79% of the arthropods were decapods (Table 4-9), and of these 

Anomura yielded the most species, followed by the Caridea. Although some 

Isopoda and Amphipoda were collected with the trawl, the overwhelming 

majority of these crustaceans, many of which are new species, were 

collected by the box corer and are therefore discussed in the Macrofauna 

section of this report. 

Whereas we observed from the first three LGL cruises that 53% of the 

arthropod species then collected extend their bathymetric range deeper 

than the 1000-m isobath, an analysis of data from all cruises reveals that 

only 36% are found at or below this isobath. This marked decrease may 

have considerable ecological significance. The principal sampling effort 

during Cruises IV and Y was directed toward isobathymetrie station 

placement, i.e., stations were arranged along horizontal lines that varied 

in depth as little as possible. Moreover, most of these isobathymetrie 

station-lines (essentially horizontal transects) were run at depths 

considerably less than 1000 m. Since all of these depths were sampled in 

the three previous cruises without obtaining the first-time species found 

in Cruises IV and Y, it is suggested that the distribution of these 

species is best described as patchy . The patches are of unknown size but 

are probably related to local factors such as sediment texture and 

chemistry . The existence of such patches is best revealed by isobath 
269 

Table 4-9 " Megafaunal arthropods collected by trawl during Cruises I, II, 

III, In, and V of the LGL study . (Numbers in parentheses are 

the number of species from Cruises I-III only.) 

No . of % of all No . of Species at % of Species 

Taxa Species Arthropods or below 1000 m below 1000 m 

CRUSTACEA 

Cirripedia 16'.( 7) 10 5 31 

Isopoda 8 ( 2) 5 2 25 

Amphipoda 8 ( 5) 5 6 75 

Decapoda 129 (89) 79 

Penaeidea 13 (11) 7 54 

Caridea 33 (24) 12 36 

Anomura 41 (28) 

Paguridae 10 ( 9) 5 50 

Porcellanidae 1 ( 1) 0 0 

Chirostylidae 2 ( 2) 2 100 

Lithodidae 1 ( 1) 1 100 

Galatheidae 

Munida 10 ( 6) 3 30 

Munidopsis 17 C 9) 5 29 

Macrura 11 C 9) 

Polychelidae 4 ( 4) 3 75 

Nephropidae 5 C 3) 2 40 

Seyllaridae 1 ( 1) 0 0 

Axiidae 1 ( 1) 0 0 

Brachyura 30 (17) 5 17 

Stomatopoda 1 ( 1) 1 0 0 

CHELICERATA 

Pycnogonida 2 ( 1) 1 1 50 

163 (104) 100 59 36 
270 

sampling. Beyond doubt these results are another example of the speciessample-effort curve, but it does not follow that isobathy metric sampling 

below 1000 m would reverse the above percentages. It would not be 

expected, for example, that such sampling would add substantial numbers of 

species of such groups as Brachyura and Munida galatheids whose mayor 

population centers are known to be shallower than the 1000-m isobath. 

Moreover, it is commonly believed that ecological conditions are more 

uniformly distributed below 1000 m than above . And most groups of 

megafaunal invertebrates exhibit a marked decrease in diversity with 

increasing depth, as is noted later in this section. 

Pycnogonida 

Two species of pycnogonids were collected in the trawl . Only one 

individual of each species ( Pallenopsia tydemani caraibica and Pallenopsis 

scoDaria) was taken . 

Cirripedia 

Sixteen species in 12 genera of barnacles were collected in trawl 

samples . TWO species live attached to the carapace of crabs . Of these 

two Trilasmis kaempferi inaeQUilaterale , which was found on three crab 

species ( Chacellus filiformis , Acanthocarpus alexandri , and Geryon 

cluinguedens), was much more common than Octolasmis gerYOnophilia which 

apparently occurs only on Geryon guinQUedens. The four most abundant 

species, aside from the above, in descending order were Verruca floridana 

(depth range from 328-750 m), Arcosealpellum portoricanum intonsum (depth 

range from 328-750 m), AmiAdosealaellum semisculptum (depth range u65-2050 

m), and Yerum idioplax with a depth range of 750-1500 m . Only one 

specimen of 1Terruca euAhypta was collected in the west-central region, but 

it proved to be a first record for the Gulf of Mexico . 

Isopoda 

Eight species of isopods were collected by the trawl. Of these only 

one species, Bathynomus giRanteus (479-1170 m) has been described. There 
271 

were three undescribed species in the genus Aega and one species each in 

the genera Cirolana , Meinertia , Rocinela , and Lironeca. The latter, 

Lironeca sp. 286, was found to be a parasite of fish. 

Amphipoda 

Eight species of amphipods were collected by the trawl. Six of these 

were undescribed species in the genera Anonyx , Epimera , Euronyx , and 

Valettiopsis, while two were given tentative identifications, Oediceroides 

ef. abyssorum (1170-1440 m), and Trisehizostoma ef. lorgirostre (1170-2857 

m) . 

Stomatopoda 

Only one species of stomatopod, Sguilla edentata, was collected in 

the trawl. Only five individuals were taken from a single station (C1 at 

329 m) on the Central Transect . It is known to be more abundant at 

shallower depths . 

Decapoda 

Below we provide a summary of the mayor groups of decapods . These 

accounts are supplemented by representative species accounts which can be 

found in Section 4 .3 .4 . 

Natantia :Penaeidea. Thirteen species of benthonic penaeid shrimps 

were collected in this study. Penaeopsis serrata was by far the most 

abundant penaeid species yielding 2896 individuals at 11 stations. The 

depths at 10 of these stations average 375 m and the species was not found 

below 465 m. The species averaged 260 individuals at the shallow stations 

and only two at the deeper station (Fig . 4-59) " It was found on all 

three major transacts. The population density of P. serrata ranged from 4 

to 255 (average 138) per hectare at the shallow stations and was 1 4 per 

hectare at 465 m where it occurred at only one of 17 sampling stations of 

that average depth (Table 4-10) . Perhaps a more realistic view of its 

abundance is the fact that overall it occurred at only 39% of the 28 
300 

too 

W 

G1 

ioo 

Q 

0 

r---= Bathyplux typhla 

Benthochoscon schmJtti 

Munido lonyipes 

~ 

IWunidv valido 

Nemotocarcinua rotundua 

CZ3:= Penaeopsis serrafc 

Plesioniko holthuisl 

375 Goo $76 1275 

3zs-493 465_75o 751-1ooo 1o5o-15oo 

DEPTH (m) 

isoo 

1550-2050 Pioo-zee 

N 

v 

N 

Figure 4-59 . Average number of individuals of selected decapods at collecting stations in the depth 

intervals shown on abscissa . 
273 

stations sampled within its depth range. The dramatic reduction in its 

population is seen in Figure 4-104. Other more detailed facts about the 

biology of Penaeopsis serrata may be found in Section 4 .3 " 4 of this 

report . 

Table u-10 . Comparison of the five most abundant penaeid species from the 

LGL and TerEco studies. 

14L 

Total lie _Dsasity DepLtLW 

4oecien Collected 

Papapopsip aerrata 2896 

Per Hectare 

138 

Hay, Pop, 

375 

ffim rob~atus 184 6' . 605 

yathpsicygus bsrtlettl 166 5 1275 

Emcaeenaeya 1pnaitcaLrla 80 9 3T5 

gTaUpMp~ Lhilla 69 . 605 

rerRm 

Total DepttL.(n) 

Scecies Collected P 

Penaeoonia aerrata 255 

May, OD, 

375 

h~rtlettl 6E6' 1050 

Paraoeciaeun lonairoatria 601 250 

gy,aftnaRapapya robuatua 500 

gh~ Ampuu 296 600 

Only four other penaeid species were of sufficient abundance as to be 

of interest here . Hymenopenaeus robustus was the second most abundant 

penaeid shrimp in numbers collected . One hundred and eighty-four 

specimens were taken at 17 stations out of the 28 established within its 

depth range from 328 to 750 m (Table 4-10) . This establishes it as one of 

the most common shrimps on the northern Gulf slope since the chances of 

getting one or more H. robustus within its depth range are about two out 

of three trawls . Its population density ranges from 1 to 20 per hectare 

with an average of six at those stations where it is found. Benthesicymus 

bartletti was third in abundance, but only a total of 166 individuals was 

collected, but it did occur at 20 stations ranging in depth from about 875 

to 2050 m. The density of this species ranged from 1 to 39 per hectare 

with an average of five per hectare where it occurred . Thus, although B. 

bartletti did not produce as many individuals as P. serrata , it did occur 

at 69% of the 29 stations sampled within its depth range . A more detailed 

treatment of the biology of Benthesicymus bartletti may be found in 

Section 4 .3 .4 of this report . Parapenaeus lonAirostris was fourth in 

abundance with a total of 80 individuals, and it was taken only at four 

stations having an average depth of 375 m . Its density ranged from 1 to 

15 per hectare where found. Hymenopenaeus debilis was represented by 69 

individuals collected at 10 stations ranging in depth from 600 to 1275 
274 

m. The density of this species ranged only from 1 to 14 per hectare . 

Although the actual numbers collected of P . longirostris and H . debilis 

are not far apart, the chances of obtaining one or more of the latter 

within its depth range are about one in four, as compared with better than 

one in 2 .5 trawls for P. lonEtirostris. 

The least abundant penaeids in the present collection are Solenocera 

necopina and Hemipenaeus earpenteri , the first is rare in the present 

collection because the shallowest LGL station is the deepest record for 

the species, and H. carpenters is not well represented in part because 

its population maximum is considerably deeper (3300 m) than the deepest 

LGL station . 

The genus Hymenopenaeus is represented in the collection by three 

sibling species that have some overlap in their distributions but have 

marked differences in population density (Fig . 4-60) . 

Natantia :Caridea. Thirty-three species of caridean shrimps were 

collected during the fine cruises . Nine species were added to the list 

during Cruises IV and V. All were taken at depths less than 1000 m in 

both the west Central and Eastern Transects. Two of the nine are 

undescribed species, viz., a hippolytid, Bythocaris sp. A, taken on the 

West Central Transect at a depth of 113 m, and a palaemonid in the genus 

Palaemonella, which was collected on the same transact at depths of 479 

and 522 m. In addition, we have obtained first records in the Gulf for 

five other caridean species: the pandalid Heterocarvus laevis (Cruise V 

at 663 m) ; the crangonid Metacrangon iaegueti aQassizi (Cruise III at 965 

m) ; the crangonid Prionocrangon peetinata (all five cruises at depths 

ranging from 613 to 1206'm) ; the processid Processa proflinda (Cruise IV at 

367 m and Cruise V at 368 m) ; and the palaemonid Periclimenes pandionis 

taken on Cruise III at a depth of 632 m. In addition we have taken both a 

new species of Stenopodidea, Spongilocoides n. sp., and eight individuals 

of the very rare species Richardina spinicineta. 

The most abundant of the caridean shrimps on the slope of the 

northern Gulf is Nematocarcinus rotundus , which has a wide bathymetric 

range from about 500 to 1550 m (Table 4-11) . Its frequency of occurrence 
Hymenopenoeus ophotlcus 

H. deb!/is 10 - 

H. robustus 

W 

v 

W 

Z 

W 

2 

t7 

W 

1 

0 

376 609 "7d 1176 1000 2478 

328-423 4E5-780 751-1000 1080-1000 1550-2050 2100-2655 

DEPTH (m) 

Figure 4-60 . Average density (no ./hectare) of three sibling species of the genus Hymenopenaeus (penaeid 

shrimp) . N 

v 
276 

is also high having been collected at 34 of the 38 stations (89K) located 

within its depth range (Fig. 4-59) . 

Table 4-11 . Comparison of the five most abundant caridean species from 

the LGL and TerEco studies. 

IAL 

Total Collected 

(A )' 

Ave . Density 

P H t 

Depth (a)' 

Sceciea ll 3tationn M P 

wLtecarelmw rpyypdus 3445 

er ec are 

45 

ax, on, 

12f5 

Pleslpnllm holtAulai 1568 30 607 ". 

917ohocranaee ~~ 333 12 1250 

91XnhcBCaci= aCYlOaLs 314 10 1170 

tarapapdalua id113a1 240 9 375 

Tartan 

total Collected 

3 (All 3t ti )I 

Depth (a) 

oeciea a ona M P p 

Wema .onarni = rofypQyp 1488 

ax, o , 

1050 

Pleaio 1Ica bQjthyipl 222 650 

Li],yohooranaon npbjllm 746 1050 

9112hoaranaon aouleata 371 1150 

m yillyy 255 .00 

Over 3400 specimens of N. rotundus were collected in such a pattern 

as to produce a bimodal curve of abundance, with one peak occurring 

between 465 and 750 m and the other between 1050 and 1500 m (Fig. 4-59). 

Its maximum population density of 334 individuals per hectare is found at 

1135 m on the West Central Transect ; however, it maintains an average 

density of only 45 per hectare (0 .005 individuals/m) . A more detailed 

treatment of the biology of Nematocarcinus rotundus may be found in 

Section 4 .3 .4 of this report . 

There are four species of Nematocareinus that exist in the offshelf 

waters of the Gulf of Mexico. They display interesting bathymetrie and 

geographic distribution. LGL collected two of the four species, viz ., N. 

rotundus and N. ensifer ; however, only six individuals of the latter 

species were taken . This is not unexpected because ensifer ranges in 

depth from 1650 to 3750 m with a maximum population near 2100 m (Pequegnat 

1983) which are depths LGL sampled only a few times. Although the 

geographic ranges of N. rotundus and N. ensifer (sibling species) overlap, 

their bathymetric ranges do not. However, the distribution of the other 

two species is such as to achieve quite effective isolation. Thus, the 

depth range of N. cursor overlaps that of N. rotundus but it is limited to 

the southeastern Gulf where the latter does not occur, and whereas the 

depth range of N . acanthitelsonis overlaps that of ensifer, the two 

species do not compete because acanthitelsonis is limited to the 

southwestern Gulf. 
277 

The second most abundant caridean in the northern Gulf is the 

pandalid Plesionilta holthuisi (Table 4-11) . Its bathy metric range extends 

from about 365 to 850 m. Here it attains an average population density of 

30 individuals per hectare, with a maximum density of 334 per hectare on 

the Eastern Transact at a depth of 600-620 m (Fig. 4-59) . There are four 

siblings of P . holthuisi in the collection, but they are not so common. 

Thus, although Plesionika acanthonotus , tenuipes , lonAipes , and edwardsii 

overlap the bathymetric range of P . holthuisi , they either have shallow 

population centers or occur in larger numbers in geographic parts of the 

Gulf that the latter does not occupy (Fig. 4-61) . The figure shows very 

graphically the predominance of P. holthuisi in the northern Gulf. Amore 

detailed treatment of the biology of Plesionika holthuisi may be found in 

Section 4 .3 .4 of this report. 

The glyphoarangonids GlyphocranAOn nobilis and G. aculeata are the 

third and fourth most abundant carideans on the upper and middle slope of 

the northern Gulf (Table 4-11) . These two species have about the same 

depth range (750-2858 m) and do occur at times at the same station, but G. 

aculeata has a greater frequency of occurrence, 62% of stations within its 

depth range as compared with 45$ for G. nobilis. In fact, G. aculeata has 

a much more uniform distribution of population than nobilis in that its 

density ranges only from 1 to 76, whereas the latter ranges from 1 to 105. 

Were it not for this latter density at Station 1 1 (1136 m) on the West 

Central Transact, G. nobilis would probably be outranked by both aculeata 

and alispina. It is suggested that G . nobilis in particular (perhaps 

other species as well) may have developed a colony way of life. As in the 

ease of the caridean genus Plesionika, Glyphocraagon is polyspecific . The 

present collection contains four species : G. nobilis , aculeata, alispina , 

and lonRlegi (Fig. 4-62) . Both G . nobilis and aculeata seem to achieve 

highest average densities in the same depth unit between 1150 and 1250 m, 

but nobilis is more abundant to the west and aculeata to the east. Amore 

detailed treatment of the biology of Glyphoerargoa aculeata may be found 

in Section 4 .3 .4 of this report. 

The fifth most abundant caridean is the pandalid shrimp ParaDandalus 

Willisi (Table 4-11) . Its population density averages between 9 and 10 

per hectare in its depth range of 250 to 465 m. Its density ranges from 3 

to 28; it has a remarkable frequency of occurrence, having been captured 

26 

Glyphocronyon oculeota 

24- G. ollspina 

G. longlsy! 

E2 - G. nobllis 

Paropandalus wllliaJ 20- 

77 

LSD 16- 

a 

a 14 - 

w 10- 

a - 

<1 6- 

4 - 

2- 

0 

soe 976 1275 isoo 2476 375 

32e-423 4e5-?50 751-1000 1050-1500 15ao-2050 :100-20aa 

DEPTH (m) 

Figure 4-62 . Average density (no ./hectare) of four species of the genus Glyphocrangon and of 

Paranandalus willisi . 

N 

V 
280 

at 10 of the 11 stations sampled in its depth range (Fig . 4-62) . It 

appears to be somewhat more common on the Eastern Transeet than on either 

the Central or Western Trarisect. 

Anomura. The anomuran decapod crustaceans have five families that 

have species found on the northern slope of the Gulf of Mexico. The 

Paguridae, as hermit crabs, are represented by four genera, one of which, 

Paraaagurus , has six species of which two are new to science. One of 

these, which is to be named Parapagurus alaminos , is being described by 

Dr . R. Lemaitre . This species has not been found east of DeSoto Canyon. 

The second family, Galatheidae, is represented by three genera, one of 

which, PhylladiorhYnehus has an undescribedspecies. This genus, first 

found on Cruise V, is its first record in the Gulf. The other genera, 

Munida and Munidopsis , have been discussed in earlier reports. But Cruises 

IV and V collected a new species of Munidopsis (to be named "paraseabra"), 

first records for the Gulf, for Munida constrieta and Munidopsis 

bahamensis , and three very rare species: Munidoosis spinoculata, first 

described from the Gulf but with only two specimens having been collected 

since from the Caribbean Sea (Pequegnat and Pequegnat 1981), Munidopsis 

ef. arias, and Munidopsis serratifrons. The third family, Chirostylidae, 

has two species in the Gulf -- Oroptychus nitidus and Gastronhyehus 

spinifer. The king crab family Lithodidae is represented by Lithodes 

aQassizi in the Gulf, and the Family Porcellanidae by Porcellana 

siASbeiana . 

Paeuridae. Ten species of hermit crabs were collected from the 

slope of the northern Gulf. The two most abundant species in rank order 

are ParapaAUrus Dietus and Parapagurus pilosimanus (Table 4-12) . The 

third and fourth species in abundance, Parapagurus alaminos and Pasurus 

rotundimanus , yielded about a quarter of the individuals of the two 

leaders. Parapagurus is represented by fine species (50x of all pagurid 

species) . Interestingly two of the five species are represented by a 

single individual, so the two most abundant and the two least abundant 

species are siblings. Five Parapagurus species were taken at a total of 

33 stations but only one of those was on the Eastern Transact (Station E4) 

while 16 were on the West Central and 16 on the Central Transacts. Ninety 
zsi 

of the 102 specimens of P. ip etus were collected at eight stations on the 

West Central Transeet of Cruise V. The depth ranges P. victus extends 

from 342 to 775 m, and that of P. pilosimanus from about 603 to 100 m. 

Table 4-12 . The most abundant anomurans on the slope of the northern Gulf 

of Mexico as determined by LGL and TerEco projects. 

LaL 

total Collected 

3o8016- (All 3t tiom)' 

Ave . Desidty 

Per Banters 

Depth of 

a Nat Pop 

102 9 

. , 

465 

pup,-p- 85 5 650 

MARL" ygLULe 1601 25 625 

lkalda lnnaipnn 230 11 368 

26-4dopsts robysya 125 5 465 

Mkold=" ar~naceua 101 4 620 

lmttZChua ylLldua 340 7 625 

LtLLflden tiaaalLl ; 

Emoa1Lr alYhafar 3 

t t l C ll t d 

Depth of 

Soeniea M ion 

p, DJ qyya 

o a o an a 

Not Reported 

ax_ _ 

p. y1logi-Apum 18 600 

b, ygu4g 114 650 

gyp 226 100 

b. robynyg 19 500 

n . erieaeeua 14 600 

]L, e1t1dus 37 954 

J,. AffAftsiyl 8 1100 

t. ilwhaSaY e3 200 

According to Lemaitre (1986), Parapagurus ( SympaAUrus) pictus 

primarily lives symbiotically with a large actinian that serves as a 

shelter for the hermit crab . The actinian, SaAartia , forms a soft 

pseudoshell that covers the interior of the shelter . A similar 

relationship is observed between ParapaAUrus Dilosimanus and zoanthids of 

the genus Epizoanthus. 

Galatheidae . Three genera of galatheids (rock crabs) were 

collected. Two of them are polyspecific : Munida with nine species from 

the northern Gulf slope, and Munidopsis with 16 species. In general, 

species of Munida tend to occupy shallower depths than those of 

Munidopsis , but Munida microyhthalma is an exception living at depths 

averaging 1100 m, whereas the more typical species, Munida valida , lives 

at depths averaging 685 m. Interestingly, the two most abundant species 

of Munidopsis occur at about the same depths as the leading species of 

Munida (Table 4-12) . 

The two most abundant species of Munida are M . valida and M. lonAipes 

(Table 4-12 and Fig . 4-59) " Munida valida far outnumbers all other 

species in the genus ; in fact, more specimens were collected of it than 

all other galatheids combined . Again, after the first two leaders, the 
282 

maximum number of individuals in the remaining nine species is 11 . The 

two rarest species of Munida are M. evermanni, taken only on the West 

Central Transact and M. miles taken at E1B on the shallow isobath. Amore 

detailed treatment of the biology of Munida valida may be found in 

Section 4 .3 .4 of this report. 

The two most abundant species of the genus Munidopsis are M. robusta 

and M. erinaceus. A more detailed treatment of the biology of Munidopsis 

robusta may be found in Section 4.3 .E of this report. Over one hundred 

individuals of both of these species were collected in the program, but an 

average of only 13 individuals of the remaining 14 species were 

collected--again illustrating the typical population drop among siblings. 

The isobathymetric sampling carried out on the Eastern Transact during 

Cruise IV collected Munidoysis species that had not been collected in the 

Gulf since being described, e .g., M. ramahtaylorae (Pequegnat and 

Pequegnat 1971). Five species occurred on the Central Transact and these 

also occurred on both East and West Transacts (these were the population 

leaders) ; three species were restricted to the Eastern Transact, all were 

rare and distributed in the Caribbean; four species were found only on the 

West Central area ; these also were rare and not collected previously in 

the northern Gulf; the remaining four species of the 16 were found on both 

East and West or West Central Transacts but were not found on the Central 

Transact . 

Eleven specimens of the third galatheid genus, Phylladiorhyachus , 

were collected at Station 6 in the west Central region at a depth of 663 

m. This is the first record of the genus in the Gulf and it belongs to an 

undescribed species. 

The fourth galatheid genus taken was Eumunida (E. iD eta) " Its record 

at Station 6 in the West Central region is a first for the LGL study. 

Station WC-6 is near a seep area, which may account for both 

Phylladiorhynchus and Eumunida being taken there and at no other station. 

TWO species of ehirostqlid anomurans were collected in the program : 

Uroptgchus nitidus is more common, being collected from all transacts, as 

was Gastrophyehus spinifer but in smaller numbers (Table 4-12) . The 

habits of the latter are unknown, but U . nitidus is assumed to live only 

on the gorgonian coral Chrgsoaoritia , as is discussed later. 
283 

Five specimens of the king crab Lithodes a.Qassizi were collected in 

the program and all were juveniles. The species is much more common in 

the western Gulf and the adults live at depths not frequently sampled in 

this program. Likewise the porcellanid crab Porcellana siASbeiana is rare 

in the collection because it generally occurs at depths shallower than 

sampled in this program. 

Maerura. The macruran decapods are represented in the LGL collection 

by 10 species in three families : Polychelidae, Nephropidae, and 

Scqllaridae (Table 4-13) " Stereomastis sculpta, the most abundant of the 

macruran decapods in this study, is treated in more detail in Section 

4.3 .4 of this report. Cruises IV and V produced some valuable new records 

and some mayor increases in numbers of individuals of species already in 

the collection. This was especially true of Cruise IV (Eastern Transeet) 

where the nephropids Acanthocaris caeca and Thaumastocheles zaleucus were 

taken for the first time in the program. The taking of three specimens of 

Thaumastocheles is particularly significant because they are the first 

complete specimens seen by these authors. Moreover, the isobathymetric 

sampling of these two cruises more than doubled the number of individuals 

of every macruran species except Scyllarus ehacei, which is a species that 

is more common at shallow depths in the western Gulf, and Polycheles 

validus , which inhabits much deeper waters than those sampled in this 

program . 
284 

Table 4-13 " The most abundant maeruran decapods on the slope of the 

northern Gulf of Mexico, as noted by the LGL and TerEco 

projects . 

_ rrsoo 

total Ave. Density Depth WI 

Soaeles 

Total 

Collected 

Depth (a) 

Haz Pm Soeaiea Collented 

StectamSIItil IIGlilptfl 732 

Per Heetare 

11 

Hay, Poo . 

1000 $" Sculpts 1409 

- _ 

1050 

Polvehelea ,fin loos 77 3 465 P" tJfDtiloos 90 500 

281 6 .65 Y" ioUlSita 65 S00 

Braehyura. There are 28 species of braehyuran crabs now in the LGL 

collection, and of these, 11 were added by collections made during Cruises 

IV and V. Four of the 11 are worthy of special mention--two are 

undeseribed species of the genus Cymonomus , and two are rare species ; one 

individual of Robertsiella mystica found at Station WC-8 (479 m) ; and two 

individuals of Dorhynchus thomvsoni , one taken at Station E2D (628 m) and 

the other at Station WC-6 (near a seep area at 663 m) . It should be noted 

also that Geryon fenneri was collected at Stations E1B (3u5 m) and E1C 

(350 m) on the Eastern Transect . This is likely the northernmost 

extension of its range. 

The 28 braehyuran species are bathy metrically separable into three 

groups of unequal size : (1) those like Acanthocarpus alexandri , 

Bathynectes sunerba , Callinectes similis , and others which are common on 

the shelf but spill over onto the upper slope ; (2) a larger group led by 

such species as BathYplax typhla, Benthochascon schmitti , Rochinia crassa 

and Triehopeltarion nobile that live on the slope above the 1000 m 

isobath ; and (3) the small group led by Geryon guinguedens , HomoloRenus 

rostratus , and a species or two not collected by the program that 

penetrate below 1000 m. This observation does not contradict the 

conclusion that brachyurans are denizens of the shelf and upper slope and 

are markers of high levels of organic production. 

The five most abundant brachyurans on the slope of the northern Gulf 

are listed in Table 4-14 . 
285 

Table 4-1u . The five most abundant brachyuran crabs collected by the LGL 

and TerEco projects. 

Total 

LfiL 

Collected Ave . Density Depth (e) : 

3-Ma . U11 Stations)' Per Hentrre Naz_ POD, 

AtuYDlai y=Dhlt 2027'. 38 1175 

JWpippcpgappp iphmiLLi 1 .95 35 .75 

dew 124 2 1170 

L.E.ieua Mir= 98 6 342 

&rsOala Acaphoa 74 3 338 

IN - 

Total Collected 

f l 

Deptb (a) 

3oaaien lh M 

. t=phjj 

ill 3tatione ' 

603 

z . me . 

550 

J, . balrQU 262 300 

Q. qgipauedana 107'. 950 

g. Appm1fS,1 114 400 

t" "eaCboR 36 200 

Note also the distribution of Bathyplax and Benthochascon in Figure 4-59" 

These latter two species are treated in more detail in Section 4 .3 .4 of 

this report. 

ComDOSition Abundance and Bathymetric Distribution of the Mayor 

Groups of Decanod Crustaceans. The penaeid shrimps are the only group of 

decapods that are most abundant in the shallowest depth interval shown in 

Figure 4-63 (328-423 m) . The observed pattern is determined primarily by 

Penaeopsis serrata (see Fig. 4-59) " The galatheids and brachyurans reach 

peak abundance at stations lying in the depth interval, 465-750 m. These 

patterns are determined primarily by Munida valida (galatheid) and 

Bathyplax typhla (brachyuran), as shown in Figure 4-59 " The caridea have 

a secondary peak at stations in this depth interval, but the major peak of 

abundance is found at stations in the 1050-1500 m depth interval. It is 

clear from an examination of Figure 4-59 that this bimodal pattern is 

determined by Nematocareinus rotundus. The macruran curve follows the 

abundance curve of the flatbaeked lobsterette, Stereomastis sculpts. 

u " 3 .3 .6 PoAOnonhora/Yestimentifera 

Two species of pogonophorans, viz ., Galathealinum n. sp. (464 m) and 

Lamellibraehia sp . (464 m) were taken by the trawl on the west-Central 

Transect at Station WC7 only . 
325- 

PENAEIDEA 300 - 

-CARIDEA 

BRACHYURA 

GALATHEIDEA~ 

250 - Other ANOMURA 

MACRURA 

200- 

U 

150- 

Or 

W 

100- 

w ~a1 i:~l 50 

40 - 

30- 

20 - 

10 - 

p L I 

376 004 $76 1t76 1 .00 p47i 

328-423 4e5-75o r51-1ooo 1o5o-1501o 155o- :o5o :ioo-sacra 

DEPTH W 

N 

00 

QN 

Figure 4-6'3 . Relative abundance (ave . no ./station) of the major groups of decapods . 
287 

x " 3 .3 .7 Brachiopoda 

Only two species of brachiopods were taken in the trawl, both in 

small numbers Ecnomiosa gerda was taken at a single station on the Central 

Transeet (964 m), Platidia anomioides was taken at two stations on the 

West-Central Transact at about 700 m depth. The latter is a widespread 

species that is not normally common to U .S. waters . 

4 .3-3 .8 Echinodermata 

One hundred twenty-six species of megafaunal echinoderms were 

collected by trawling from the three transacts. Forty-two of these 

species were collected for the first time during Cruises IV and V. These 

were distributed among the echinoderm classes, as follows : 13 asteroids, 

15 ophiuroids, 5 echinoids, 5 holothuroids, and 4 crinoids. Overall the 

ophiuroids and asteroids account for about 70% of all the echinoderm 

species collected in the project (Table 4-15 and Fig. 4-6u) . 

Table 4-15 " Total numbers of the five classes of megafaunal echinoderms. 

Class 

Ophiuroidea 

Asteroidea 

Holothuroidea 

Echinoidea 

Crinoidea 

No . of Species 

45 

41 

21 

13 

6 

126 

Percent of all Echinoderm Species 

36 

32 

17 

10 

5 

100 

Asteroidea 

A total of 41 species of asteroids was collected during the five 

cruises. Among the 13 species that were collected for the first time 

during Cruises IV and V Were the rare genus Caymanostella sp. and the 
N 

00 

00 

150- 

ASTEROIDEQ 

CRINOIDEA 

ECHINOIDEA 

HOLOTHUROIDEA 

OPHIUROIDEA 

i00 

o. 

a 

w 

m 

s° 

0 mom 

375 a08 $76 1275 1800 2478 

326-423 461-750 731-1000 1030-1500 1530-2030 Y100-285.7 

DEPTH (m) 

Figure 4-64 . Total individuals collected of the five echinoderm classes. 
289 

species Cireeastes americanus , as well as Odontaster setosus and 

Sclerasterias contorta. 

The fine most abundant asteroids are shown in Table 4-16 of which the 

leader is considered to be Nymphaster arenatus because it was taken at 

such a large number of stations. 

Table 4-16 . Abundance of the five most abundant asteroid species 

collected from the three mayor transacts. 

Total No . of Depth Range 

Species Collected Stations on Transects 

Nymphaster arenatus 45 24 375-1800 

Pectinaster gracilis 63 13 465- 750 

Perseohonaster echinulatus 81 13 465- 750 

Plinthaster dentatus 25 16 465-2100 

Plutonaster intermedius 41 9 876-2100 

As shown in Figure 4-64, the abundance of asteroids in this project 

follows an irregular trend of increase from 328 m to around 1800 m, after 

which there appears to be a reversal of the trend. In part the latter 

reduction results from the fact that the project's sampling effort did not 

effectively cover the lower slope where an abyssal assemblage having such 

species as Dgtaster insiAnis, Benthopecten simplex , Ampheraster alaminos , 

and Litonotaster intermedius is to be found. 

Although 61 species of starfishes were identified in the TerEco 

project (20 more than LGL), it is interesting to note that exactly 20 of 

the 61 reach maximum populations either shallower or deeper than sampled 

in the present project. 

Ophiuroidea 

Forty-fine species of ophiuroids were collected by trawling in this 

project. This compares with 30 species of ophiuroids collected during the 

first three cruises and 43 species listed in the TerEco study . The 15 
290 

"new" species of Cruises IV and V Were distributed unequally among the 

transeets : eight were taken only on the West Central Transect, four only 

on the Eastern Transact, and three on both. The majority (9 of the 15) 

were represented by a single individual and that those three species found 

on both transacts accounted for a disproportionate one-third of the total 

individuals taken . The six most abundant ophiuroids in the project are 

listed in Table 4-17 " 

Table 4-17 " The six most abundant ophiuroid species collected from the 

three mayor transacts. 

Species 

Ophiernus adspersus 

Bathypectinura heros 

Ophiochiton grandis 

Ophioliyus agassizii 

Amphiophiura seulptilis 

Amvhilepis ef . ingolfiensis 

Total No . of Stations Depth Range 

Collected Where Collected on Transacts 

986 19 423- 805 

211 18 615-2400 

175 17 420- 850 

147 16 329- 625 

116 11 750-1440 

492 3 750-2855 

Both Ophieraus adspersus and Bathypectinura heros were among the five 

most abundant brittle stars in the TerEeo study . 

The abundance of ophiuroids has a marked bimodal characteristic with 

one peak around 600 m, which is probably determined primarily by Ophiernus 

adspersus, and the other much deeper peak (Fig. 4-6u) was probably caused 

by species of the genus Amyhilepis. 

Echinoidea 

Thirteen species of sea-urchins were collected from the three major 

transeets. The echinoid species were most common on the Central Transeet, 

where 8 of the 13 species were found (sin found only on that transact) . 

Three species were found only on the Eastern Transeet, two species were 
291 

restricted to the West Central region, and one species each occurred on 

the Central and Eastern or on the Central and Western Transeets. 

The most abundant echinoid species in the present collection is 

Plesiodiadema antillarum followed by Homolampas fragilis and Brissoysis 

alta (Table 4-18) . 

Table 4-18 . Total numbers of the five most abundant sea-urchins trawled 

from the mayor transeats. 

Total No . of Depth Range 

Species Collected Stations (m) 

Plesiodiadema antillarum 70 6 750-1500 

Homolampas fragilis 17 6 465- 750 

Brissopsis alta 19 3 328- 750 

Aceste bellidifera 7 3 465- 750 

Brissopsis sp . 13 1 328- 423 

The totals shown in Table 4-18 are small, reflecting the fact that 

the project's sampling depths were too deep to get typical numbers of 

Brissopsis and not extensive enough in deep waters to get representative 

numbers of either Plesiodiadema or Phormosoma placenta placenta. Also, 

the low numbers of echinoids (Fig . 4-6u) is related in part to the fact 

that some species like Plesiodiadema antillarum and Phormosa placenta 

occur in isolated "herds", as determined by benthic photography. 

Holothuroidea 

Twenty-one species of sea cucumbers were collected by trawling in 

this project. These species were distributed very unequally among the 

transects--18 of the 21 species were found on the Central Transeet (10 

were found only on this transact) ; nine species were collected from the 

Eastern Transact (only one species was restricted to this transact) ; nine 

species were collected from the Western or West Central Transact (only one 

species was restricted to this transact) ; and six species were found on 
292 

all transacts. Three of the four most abundant species were found on all 

transeets and all four were found on the Central Transact (Table 4-19) " 

The flout from the Mississippi River moves westward across this transact 

where it deposits some of its load of organic matter. This is important 

to the sea cucumbers found there in that all are thought to be deposit 

feeders . 

Table 4-19 " The four most abundant holothuroid species collected from the 

three mayor transacts. 

Total No . of Stations Depth Range 

Species Collected Where Collected on Transacts 

Mesothuria laetea 184 18 603-1170 

Molpadia barbouri 44 15 840-2400 

Molpadia musculus 38 9 838-2400 

Benthodytes tqplCg 101 3 2050-2500 

As shown in Table 4-19 and Figure u-64 the sea cucumbers tend to 

have both shallow and deep assemblages. 

Crinoidea 

Six taxa of crinoids (four definitive species) were collected in the 

project, four being added for the first time during Cruises IV and V. The 

latter four, however, were found only in very small numbers ; hence the two 

most abundant species are the same as reported for Cruises I to III. 

The most abundant now is Democrinus brevis followed by Atelecrinus 

balanoides (Table 4-20) . 
293 

Table 4-20 . Complete inventory of erinoids collected in the project . 

Species 

Demoerinus brevis 

Atelecrinus balanoides 

Caryometra alOpe 

Cargometra lisa 

Caryometra sp . 

Comatulida 

Total No . 

Collected 

280 

67 

7 

11 

1 juv . 

1 

No . of Stations 

Where Found 

17 

15 

3 

2 

1 

1 

Depth Range 

on Transacts 

620-200 

620-1200 

522- 663 

600- 775 

740 

615 

u " 3 .3 .9 Ascidiacea (Chordata) 

Only one individual of one species of sea squirt was collected in the 

trawl. Bathgstyeloides enderbyanus was taken at a depth of 200 m on the 

Central Transact . 
294 

4 .3 .4 DECAPOD SPECIES ACCOUNTS 

4 .3 .4 .1 Bathyplax typhlg (A " Milne Edwards, 1880 Family Goneplacidae) 

Distribution 

The reaper crab, Bathyplax typhla , is one of the most abundant deepsea brachyurans in the Gulf of Mexico (Pequegnat and Chace 1970) . It has 

been reported from North Carolina to Brazil, and throughout the Gulf of 

Mexico on the upper continental slope (Pequegnat et al . 1971), in depths 

of 402 to 878 m (Powers 1977) . In the Straits of Florida, it is 

restricted to water colder than 10°C, and deeper than 315 m (Soto 1985) . 

It has usually been found over muddy substrates (Soto 1985, Powers 1977), 

although Polders indicated that it also has been taken from "coral 

bottoms ." Powers was probably referring to calcareous rubble, 

foraminiferans, or sand, rather than reef corals, which would not be 

present at the depths indicated. 

During the Alaminos cruises, Bathgplax typhla was the numerical 

dominant in the deep Gulf, having been collected by benthic skimmer in 

depths from 512 to 860 m at densities ranging from 16 to 135 per ha, with 

the highest density in Pequegnat's (1983) Archibenthal Zone, between 658 

and 860 m . Pequegnat listed the depth of peak population density for 

Bathyplax typhla as being 550 m . Specimens apparently contaminated by 

petroleum from natural oil seeps have been described as common by 

Pequegnat and Chace (1970) . 

During this program, Bathyplax typhla was collected on all five 

cruises, at depths ranging from 366 to 1,438 m (Fig . 4-65) " Specimens 

were taken on the Central Transect from 474 to 1,438 m ; on the Eastern 

Transact from 366 to 1,170 m ; on the Western Transact from 654 to 828 m ; 

and on the West Central Transact from 464 to 751 m. 

Measurements 

Bathyplax typhla males collected during the Alaminos cruises reached 

28 m m (carapace width) ; females reached 25 mm, with ovigerous specimens 

from 16 to 22 mm in width taken in August, November, and December. 
295 

Figure 4-65 . Locations where Bathyplax typhla were collected (closed 

circles) . 
296 

Crabs taken during this program had a unimodal, bell-shaped curve of 

size frequencies (Fig. 4-66A) total of 581 individuals was measured . They 

ranged between 11 and 32 mm in width, and averaged 20 .6 mm. Both males 

and females had similar widths (Fig. 4-67) . Ovigerous females averaged 

22 .9 mm, within a range of 18 to 28 mm . About one-fifth of the females 

collected (49 out of 273) were ovigerous. Non-ovigerous females ranged 

from 10 to 31 mm in width, with a mean of 20.3 mm. Males averaged 19.3 mm 

wide, within a range of 11 to 32 mm. The size distribution of individuals 

remained very similar from cruise to cruise, without apparent cohorts or 

age class peaks or seasonal trends (Fig. 4-68) . There was a positive 

linear correlation (r = 0 .95) between the logs of width and wet weight 

(Fig. 4-66B) for all specimens together. Weights ranged from 0.3 to 11 .9 

g, with a mean of 2 .9 g . 
297 

120 -

100- 

goU 

G 

4J 

Gocu 

w 40-

20 -

0 

0 

3 

n 

00 

v 

t 

a0 

0 

-1 

-2 

2 .2 

Bstbyplsx typbls 

Figure 4-66 . Frequency histogram (A) and size-weight relationship (B) 

for carapace widths of Bathyplax typhla (all stations 

pooled) . 

BATHYPrAX TYPHLA 

Ovlqerous Female 

Female 

Male 

0 ~10 20 30 40 

IIIIDTM (am) 

Figure 4-67 " Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace widths of Bathyplaz typhlg (small squares mark 

outliera) . 

4 e 12 1e 20 24 2e 32 36 40 

Carapace Width (mm) 

2.4 2.6 2A 3A 32 E.4 3.6 

Ln Carapace Width (mm) 
298 

Beldyplex lypdle 

12 

10 

u g 

N 

6 

w 

2 

0 

0 

50-

40 -

30-

v 

v 20 -

44 

10 

0 

0 

20 -

U 

10-

v 

w 

0 + 

40 0 

40 

30 

T 

u 

G 

20 

a 

w 

10 

0 

10 20 30 40 

Carapace Width (mm) 

Crrise Y 

Figure 4-68 . Frequency histograms for carapace widths of Bathyplax 

typhla from individual cruises . 

10 20 30 40 

Carapace Width (mm) 

10 20 30 40 

Carapace Width (mm) 

10 20 30 

Carapace Width (mm) 

crrlsa IV 
299 

4 .3 .4 .2 Benthesicymus bartletti (Smith, 1882 Family Penaeidae) 

Distribution 

Benthesicymus bartletti , Bartlett's short-horned prawn, has been 

collected on both sides of the Atlantic Ocean, in the Pacific, and in the 

Caribbean and Gulf of Mexico, although there is some question about its 

presence in the eastern Indo-Pacific (Crosnier 1978) . It appears to be 

exclusively benthic, seldom if ever venturing much above the bottom 

(Wenner 1980, Pequegnat and Chace 1970) . Its depth range in the Caribbean 

and the Gulf Alaminos collections was between 731 and 1,773 m . Some 

specimens may have been taken by the Blake at 1,934 and 2,866 m, although 

Pequegnat and Chace (1970) questioned the authenticity of the deeper 

record. Pequegnat (1983) indicated a depth range of 700 to 2250 m in the 

Gulf, with a population peak in the Upper Abyssal Zone near 1050 m . 

Wenner (1980) noted an early record of Benthesicymus bartletti from 5,77' 

m, but her collections on the OS Atlantic continental slope produced 

relatively few specimens shallower than 1,000 m, or deeper than 2000 m. 

During this program, Benthesicymus bartletti was collected on all 

fine cruises, at depths from 746 to 2,07 m m (Fig. 4-69). Specimens were 

taken on the Central Transeet from 802 to 2,07 m ; on the Eastern Transect 

from 834 to 1,170 m; on the Western Transact from 828 m to 1,413 m ; and on 

the West Central Transact from 746 to 1,206 m. 

Measurements 

Benthesicymus bartletti collected on the continental slope of the 

Middle Atlantic Bight by Wanner (1970) ranged between about 7 and 30 m m in 

carapace length (from the posterior margin of carapace to the base of the 

rostrum) . Males were slightly smaller than females (means -_ 19 .9 mm and 

21 .5 mm, respectively) . Females over 20 mm long with ripe gonads were 

present at various times of year, but the sample size was too small to 

determine if breeding took place seasonally or throughout the year. 

In this study, 95 measured specimens of Benthesicymus bartletti were 

from 6 to 27 mm in carapace length, with a mean of 19 " u mm . The female 

size range overlapped the male size range at both ends (Fig . 4-70) . 
300 

Figure 4-69 " Locations where Benthesicymus barteletti were collected 

(closed circles) . 

I BENTHES/CYMUS BARTL ET T/ 

0 

Female 

Mule - 

0 10 20 30 

LENGTH (mm) 

Figure 4-70 . Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Benthesieymus barteletti (small squares 

mark outliera . 
301 

Males were between 12 and 22 mm long, with a mean of 19 .3 mm . Females 

averaged 19 "4 in length, within a range of 6 to 27 mm . The overall size 

frequency distribution showed no obvious cohorts (Fig . 4-71A), but sample 

size on each cruise was inadequate to detect seasonal trends (Fig. 4-72) . 

There was a positive linear relationship (r -_ 0 .97) between the logs of 

weight and carapace length (Fig . 4-71B) . Weights ranged from 0 .3 to 8.0 

grams, with a mean of 3 .4 g . 
302 

70 

Y 

Gp 

p 10 

O 

Y 

L 

O 

0 

3-

'. 2-

a 

v 

r 

A 

P 

by 

i 

C 

J p 

BentAesicymus bsrtelettf 

10 20 

Carapace LealtA (mm) 

30 

24 26 2I SA 32 EA 

Le Carapace Length (0110) 

Figure 4-71 . Frequency histogram (A) and size-weight relationship (B) 

for carapace lengths of Benthesicymus barteletti (all 

stations pooled) . 

BsntAssicyn»as bsrleJelti 

5-1 r_ . . .__ I

. 3 r . .. : .. ., III 

s 

e 3 

g 

v 

O 

0 

10-

8 

a 

't` 6-

v 

s 

e 

` 4 

W 

2 

0 

0 

a 2 

a 

e 

v 

0 

0 

4 

3 

a 

s 2 

e 

v L 

1 

0 

0 

30 

Figure 4-72 . Frequency histograms for carapace lengths of Benthesiayaus 

barteletti from individual cruises . 

10 20 30 

Carapace Length ( mm) 

10 2(1 30 

Carapace Length, (1w ") 

10 20 30 

Carapace Length (nn) 

10 20 

Carapace Length (mm) 

^------ 
303 

4 " 3 " 4 " 3 Benthochascon schmitti (Rathbun, 1931 Family Portunidae) 

Distribution 

Sehmitt " s benthie portunid, Benthoehascon sehmitti , was thought to be 

an endemic species in the Gulf of Mexico (Pequegnat and Chace 1970, 

Pequegnat et al . 1971) . However, there is a single record off southern 

New England (Wigley and Messersmith 1976, from Powers 1977), and many 

records from the Straits of Florida in water from 183 to 366'm deep (Soto 

1985) . Within the Gulf, it has been collected from the Dry Tortugas to 

the northern Gulf, within a depth range of 200 to 650 m . Its maximum 

population density occurs near 400 m (Pequegnat 1983). It thus inhabits 

the Shelf/ Slope Transition and Archibenthal zones defined by Pequegnat 

(1983) " Densities reported by Pequegnat and Chace (1970) were from 2 to 

40 individuals per hectare. Benthochaseon prefers mud and silt-clay 

substrates (Polders 1977) " 

During this program, Benthochascon schmitti was captured on all five 

cruises, at depths clustering around the 400 m isobath and extending from 

329 to 840 m (Fig . 4-73) " Collections at 751 m and 840 m represent 

substantial depth range extensions for this species. Specimens were taken 

on the Central Transact from 239 to 632 m ; on the Eastern Transact from 

366 to 840 m ; on the Western Transact at 342 m ; and on the West Central 

Transact from 368 to 751 m. 

Measurements 

Immature Benthochascon males and females were collected from March 

through October during the Alaminos cruises (Pequegnat and Chace 1970, 

Pequegnat 1983), although their size was not reported. 

During this program, 305 measured specimens were from 12 to 86 mm in 

carapace width . The size range of females was similar to that of the 

males (Fig. 4-74)" There was only one ovigerous female collected, 59 mm 

wide . Non-ovigerous females were from 12 to 65 mm wide, with a mean of 

40 .4 mm. Mean width for males was 26 .9 mm, within a range of 11 to 86 mm. 

Two peaks were evident in the overall size frequency distribution 

(Fig . 4-75A), suggesting the seasonal passage of cohorts or age classes, 
304 

Figure u-73 . Locations where Benthochaseon schmitti were collected 

(closed circles) . 

rBENTHOCHASCON SCHM/TT/ 

Oviparous Female 

Usrnyi. oe.wvotron=asmm 

Female 

Mate =WC 0 

0 20 40 60 80 100 

WIDTH (mm) 

Figure 4-74 " Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace widths of Benthochaacon achmitti (small squares 

mark outliers) . 
305 

so 

ao 

O' 40 

70 

g 

W 20 

t0 

O 

O 

s 

a P 

v 

c 

w 

r 

_z 

BenlAocAsscon SCAN/tot/ 

z s 4 s 

L~ Grahes Width (we) 

Figure 4-75 . Frequency histogram (A) and size-Weight relationship (B) 

for carapace widths of Benthochascoa schmitti (all stations 

pooled) . 

BNrIAocAiscan rckArit 11 

43- Cruise 11 30-1 Cruise III 

6 

s 

a 

z 

0 

0 

zo 

ya 

C 

10 

g 

a~ 20 

i 

S 

,~ I o 

0 

0 

40 -

30 -

a 

Y 

a 20-

0 0 

2(1 40 60 80 100 

Carapace wins (mm) 

Figure 4-76 . Frequency histograms for carapace widths of Benthochasoon 

schmitti from individual cruises . 

20 40 60 YO 100 

Carapace Width (mrs) 

20 40 eo so too 

Carapace Width (mow) 

0 

a 20 ~O , 6'0 9`0 10,0 

CKop~ Width (mow) 

20 40 60 00 100 

Gees Width (no) 
306 

or size-specific sex differences. Specimens taken on Cruises II and IV 

(April 1984 and May 1985) were larger than those collected on Cruises III 

and V (November/ December 198 and June 1985) (Fig . 4-76) . Most 

individuals collected during Cruise III and V collections were between 20 

and 40 mm wide, whereas Cruise II and IV collections were dominated by 

specimens from 40 to 60 mm in width. 

There was a positive linear correlation (r . 0.99) between the log of 

specimen width and weight (Fig . 4-75B) for all specimens together. 

Weights ranged from 0.2 to 65 .7 B . with a mean of 9 .9 g. Larger specimens 

were disproportionately heavy, perhaps as a result of increasing 

calcification and decreasing molt frequency, as is common in other 

crustaceans . 

One possible explanation for the differences between cruises would be 

the ability of larger specimens (e .g . those greater than 60 mm in width) 

to escape capture, proposed by Pequegnat and Chace (1970) . This factor, 

combined with an annual spring recruitment, could be responsible for an 

apparent fall disappearance of large size classes from collections. Large 

specimens--similar to those taken in the spring on Cruises II and IV--

might have grown enough by summer to avoid capture during Cruises III and 

V. Individuals that were newly recruited in the spring might have been too 

small to be taken on Cruises II and IV, but could have been large enough 

to be vulnerable to the net by summer. 

However, the presence of many small individuals in the 

November/ December sample (Cruise III) is problematical, and implies more 

than one annual peak of recruitment. This scheme is presented only as a 

hypothesis for further examination, and alternative hypotheses (e.g. heavy 

adult mortality, emigration, seasonal differences among transeets, or 

segregation by sex or size class) are also plausible but have not yet been 

examined . 
307 

4 .3 .4 .4 Glyphocrangon aculeata (A. Milne Edwards, 1881 Family 

Glyphocrangonidae) 

Distribution 

The sharp-spined armored shrimp, Glyphocrangon aculeata, is a widely 

distributed deep-sea caridean, ranging from the western Atlantic (Delaware 

to Brazil), the Straits of Florida, the Gulf of Mexico, and the southern 

Caribbean (Pequegnat and Chace 1970) . Pequegnat et al . (1971) noted that 

it was far more abundant in the western Gulf than the eastern Gulf, and 

typical of the upper continental slope, within the Upper Abyssal Zone 

(Pequegnat 1983) " It has been reported from depths of 732 to 1,767 m, 

with an apparent abundance maximum around 1150 m (Pequegnat 1983, Holthuis 

1971) . It has been taken over various bottom types, including grey ooze, 

yellow ooze, G1obiAerina ooze, mud of various colors, clay, fine sand with 

black specks, forams, etc. 

Glyphocrangon aculeata was collected during this program on all fine 

cruises, at depths ranging from 751 to 2,857 m (Fig . 4-77) " The latter 

represents a depth extension of almost 1,100 m over published reports . 

Specimens were taken on the Central transact from 964 to 1,510 m ; on the 

Eastern Transact from 812 to 2,857 m ; on the Western Transact at 828 m ; 

and on the blest Central Transact from 751 to 1,206 m. 

Measurements 

Specimens of Glyphocrangon aculeata taken during the Alaminos 

cruises were between 5 and 33 m m in carapace length, excluding the rostrum 

(Pequegnat and Chace 1970) . Ovigerous females were collected in August 

and November, and were at least 20 m m long. The Alaminos specimens from 

the Gulf of Mexico were smaller than many that have been collected in the 

Atlantic. Holthuis (1971) examined a large number of specimens from 

various locations, and measured ovigerous females at 36 to 49 mm carapace 

length, including the rostrum. The largest male Holthuis measured was 38 

mm in carapace length, including the rostrum . 

During this program, 268 measured specimens of Glyphocrangon aculeata 

were between 5 and 31 mm in carapace length, excluding the rostrum (Fig. 
308 

Figure 4-77 . Locations Where Glyphocraniton aculeata were collected 

(closed circles) . 

,o 

r 

c, 30 

e 

n 0 

L p 

10 

a 

z 

v 

r 

G 

e~ a 

yY 

i 

C -1 

J 

_Z 

Figure 4-78. Frequency histogram (A) and size-Weight relationship (B) 

for carapace lengths of GlyphocranAOn aculeata (all 

stations pooled). 

O 

O f0 ~ i0 ~0 

Carspsce Le"tA (mm) 

-31 

S 2.0 2.3 &-0 3.3 1 .0 

La Greha UstA (ow) 
309 

4-78A) . There were no obvious individual peaks representing separate 

cohorts. Female lengths overlapped the male length range at both ends, 

and ovigerous females were, on the average, considerably larger than were 

non-ovigerous females or males (Fig. 4-79) . About a fourth of the females 

collected (33 out of 126) were ovigerous . Mean length for ovigerous 

females was 24 .8 mm, within a range of 1 4 to 30 mm. Mean length for nonovigerous females was 17 .7 mm, within a range of 9 to 31 mm. Males were 

between 10 and 22 mm long, with a mean of 15 .9 mm . No clear seasonal 

differences in size were noted (Fig . 4-80) . There was a positive linear 

correlation (r = 0 .99) between the logs of weight and carapace length 

(Fig. 4-78B) for all specimens together. Weights ranged from 0 .9 to 16 .7 

g, with a mean of 3 .4 g. 
310 

GLYHYOCRAN60N ACULEATA 

p 1_- 1-~7 1 Oviqerous Female 

I female ~- 

Mole ~ 

0 10 20 30 40 

LENGTH (mm) 

Figure 4-79 " Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Glyphoerangon aculeata (small squares 

mark outliers) . 

an 

a 

a to -

Y. 

o 4 

0 

20 -

a 

v 

c 

10-

V 

au 

a 20 

s 

Y 

10 

0 0 

0 10 20 30 40 - 1- . . --40 

Grapace Length (on) Greha Length (mow) 

Figure 4-80 . Frequency histograms for carapace lengths of GlyphocranAOn 

aculeata from individual cruises . 

6lypbocrongon eculeeta 

"__i-- " to ^----- . . . 

s 

a 

6-

s 

` 4 -

0 

2 

10 20 30 40 0 

Carapace Length (mm) 

10 20 30 40 

Carapace Length (mm) 

__ .. 
311 

4 .3 .4 .5 Munida valida (Smith, 1883 Family Galatheidae 

Distribution 

The long-armed crab, Munida valida , has been collected along the 

Atlantic coast from New England to Florida, at many locations in the Gulf 

of Mexico, and in the southern Caribbean and the coast of Colombia 

(Williams and Wigley 1977) . It has been taken swarming near the surface 

(0-9 m), but has been described more commonly from 90 to 1823 m (Pequegnat 

and Chace 1970, Williams 1980 " There is also a collection record from 

2,297 m (Wenner, unpub. ms.) . Its maximum population density in the Gulf 

has been reported at 650 m, in the Archibenthal Zone (Pequegnat 1983)" 

During this program, Munida valida was collected on all five cruises, 

at depths ranging from 464 to 1,170 m (Fig . 4-81). Specimens were taken 

on the Central Transeet from 474 to 850 m; on the Eastern Transact from 

612 to 1,170 m; on the Western Transact from 654 to 828 m; and on the West 

Central Transact from 464 to 774 m. 

Measurements 

Munida valida is the largest species of Munida in the Gulf of Mexico 

(Pequegnat and Chace 1970) . Specimens collected during the Alaminos 

cruises ranged from 9 to 33 mm in carapace length. The smallest ovigerous 

female was 21 mm long, taken in November. Other collections have produced 

ovigerous females almost year-round (Williams 1984 ; Wanner, unpub. ms.) . 

In this program, 516 measured specimens ranged from 6 to 37 mm in 

carapace length, with the largest number of specimens between 10 and 30 mm 

long (Fig. 4-82A). Females overlapped males in length at both ends of the 

size range (Fig. u-83) " Only eight of the 307 females measured were 

ovigerous. Mean length for ovigerous females was 22 mm, within a range of 

11 to 33 mm. Non-ovigerous females were between 6 and 37 mm long, with a 

mean length of 16 .6 mm. Males averaged 15 .7 mm, and were between 6 and 36 

mm long. The overall size frequency distribution formed a bimodal curve 

as a result of a large number of small specimens (10-15 mm long) having 

been collected during Cruise V (June 1985) Fig . 4-84) . There was a 

positive linear correlation (r -_ 0 .99) between the logs of specimen width 
312 

Figure u-81 . Locations where Munida valida were collected (closed 

circles) . 

A 

'j, 60 . 

S, 

40, 

2 0- 

0 L 

0 

_ j 

10 

s 

z 

a 

.. 

c 

o 

r 

/fanide velide 

Zp gp 10 

Carapace Length (now) 

Figure 4-82 . Frequency histogram (A) and size-Weight relationship (B) 

for carapace lengths of Munida valida (all stations 

pooled) . 

-s 

~a 2n :s s.~ ss 40 

Le Carapace L.eWA ~~) 
313 

MYIN/DA VAL/DA 

Ovlqeous Female 

Female 

Male l IT1 L o 

0 b 20 30 40 50 

LENGTH (mm) 

Figure 4-83 . Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Munida valida (small squares mark 

outliers) . 

Iyunldo volide 

6-

a 4 -

s 

v 

3-

U. 

s 

L 2-

1 

0 

0 

40 

30 

a 

v a 

~ 20 

gt 

10 

0 

10 20 30 

Carapace Length (aim) 

Cruise IV 

60 

50 

a 40 -

a 3U 

P 

v 

20-

'001 

10 20 30 

Carapace Length (mm) 

r. .. . .. v 

40 

0 10 20 30 40 0 10 20 30 40 

Carapace length (mow) Carapace Length (mm) 

Figure 4-84 . Frequency histograms for carapace lengths of Muaida valida 

from individual cruises . 

20 

a 

10 . - 

J~ 0 

40 0 
314 

and weight (Fig . 4-82B) . Weights ranged between 0 .1 and 33 .3 g, with a 

mean of 3 .9 8 " 
315 

4 .3 .4 .6 Munidopsis robusta (A. Milne Edwards, 1880 Family Galatheidae) 

Distribution 

The robust spiny-tailed crab, Munidopsis robusta, has been reported 

throughout the Gulf of Mexico, the western Atlantic, the Straits of 

Florida, and in the southern Caribbean (Mayo 197 4 , Pequegnat and Chace 

1970) . In the Gulf, it has been collected from depths of 110 to 1000 m, 

inhabiting the Shelf/Slope Transition and Archibenthal Zones, with a 

population maximum at about 500 m (Pequegnat 1983) . 

During this program, Munidopsis robusta was taken on Cruises 2 

through 5, in depths ranging from 342 to 746 m (Fig . 4-85) . Specimens 

were captured on the Central Transect from 346 to 747 m ; on the Eastern 

Transact from 366 to 627 m ; on the Western Transact at 342 m ; and on the 

West Central Transact from 423 to 746 m. 

Measurements 

Munidopsis robusta specimens collected during the Alaminos cruises 

were between 7 and 20 mm in carapace length. Ovigerous females between 13 

and 20 mm long were present in August and November. 

Specimens measured during this program were similar in size to those 

collected during the Alaminos cruises. The 81 individuals measured were 

between 10 and 20 mm in length, and averaged 15 .6' mm long. Their 

frequency distribution was unimodal, with the greatest number of 

individuals between 14 and 18 mm long (Fig. 4-86A). Females tended to be 

slightly larger than did males (Fig. 4-87) " Mean carapace length for 

ovigerous females was 16 .6 mm, within a range of 14 to 19 mm . Nonovigerous females averaged 15 .3 mm in length, and were betWeeen 8 acid 20 

mm long. Males ranged from 7 to 19 mm long, with a mean of 14 .7 mm. There 

were no obvious seasonal differences in length frequency distributions 

(Fig . 4-88) . There was a positive linear correlation (r = 0 .93) between 

the logs of specimen width and weight (Fig . 4-86B), although smaller 

specimens were proportionately heavier per unit length. Mean weight was 

3 .0 g, within a range of 0 .4 to 5 .8 g . 
316 

Figure 4-85 . Locations where Munidopais robusta were collected (closed 

circles) . 

20 

u 

e 

v u 

W 

0 

0 

z 

.. 

P 

v 1 

r 

a 

a 

O 

C 0 

J 

Figure 4-86 . Frequency histogram (A) and size-Weight relationship (B) 

for carapace lengths of Muaidopsis robusta (all stations 

pooled) . 

iS 2.4 iA 2." 3.0 

LO GrOMSIS LOht" (MM) 

io 20 

carapace LeuOtw (yam) 
317 

MUN/OOPS/S ROBUSTA 

Ovig*r4ma Female a i 1 F~y O 

Female a 

Mole 0 1 

0 2 4 6 8 10 12 14 16 18 20 22I 

LENGTH (mm) 

Figure 4-87 . Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Munidopsis robusta (small squares mark 

outliera) . 

6-

5-

4 -

C 

3 

1 

a V 

2 -

0 

O 

12 

10 

a 8 

H 

e 

6 

4 

a 

0 

6-

5-

4 -

C 

3 

U. 

a 

v 

2 -

1 

0 

0 

CarsNCe Length too) 

Figure 4-88 . Frequency histograms for carapace lengths of NunidoDSis 

robusta from individual cruises. 

10 20 30 

Carapace Length ( mm) 

10 20 30 

Carapace Length (mm) 
318 

4 .3 .4 .7 Nematocarcinus rotundus (Crosnier & Forest, 1973 Family 

Nematocarcinidae) 

Distribution 

Nematocareinus rotundus , the thread-footed shrimp, was not separated 

taxonomically from N. cursor , a closely related species, until 1973 

(Crosnier and Forest 1973) " Therefore, descriptions of N. cursor prior to 

that date may refer to either (or both) species, as well as a third (N. 

africanus) not discussed here. 

Nematocarcinus rotundus is a very widely distributed caridean in the 

Caribbean and Atlantic. It is known from depths of 330-380 m to 1,240 m 

(Crosnier and Forest 1973) having been collected in the southern 

Caribbean, the Gulf of Mexico, along the coast of South America, and the 

southwestern Atlantic coast of the United States (Williams and Wigley 

1977) . 

Specimens collected in the Gulf of Mexico during the Alaminos cruises 

were taken across the upper continental slope from 512 to 1,829 m, with a 

population density peak in the upper portion of the Upper Abyssal Zone, 

near 1,050 m (Pequegnat et al . 1971, Pequegnat 1983) . Nematocareinus 

rotundus was the most abundant and widespread caridean collected. 

During this program, Nematocarcinus rotundus was collected on all 

five cruises, at nearly every station between 500 m and 1,000 m, and 

spanning a depth range of 521 to 1,510 m (Fig . 4-89) . Specimens were 

taken on the Central Transact from 603 to 1,510 m ; on the Eastern Transact 

from 612 to 1 ,170 m ; on the Western Transact from 654 to 1 ,113 m ; and on 

the west Central Transact from 521 to 1,206 m. 

Measurements 

Specimens of Nematocarcinus rotundus taken during the Alaminos 

cruises were between 9 and 27 mm carapace length, with the smallest 

ovigerous female 20 mm long . Ovigerous females were collected during 

July, August, and November. 

During this program, 1,060 measured Nematocareinus rotundus were 

between 9 and 30 mm in carapace length, and averaged 19 .1 mm (Fig. 4-90A) . 
319 

Figure 4-89. Locations Where Nematocarainus rotundus were collected 

(closed circles) . 

100 

eo 

Figure 4-90 . 

a 

so 

ZO 

a 

z 0-4 

E 

.. 

r 

e 

a o 

r 

-i 

e 

J 

Nwwilocorciiws retoAdas 

22 :.~ 2s 2a sn 32 3.4 3.6 

Le Gripn UhtA (me) 

Frequency histogram (A) and size-Weight relationship (B) 

for carapace lengths of NematocarQinus rotundus .(all 

stations pooled) . 

O 

O 10 20 30 

Carepwe LeaytA (aim) 
320 

There was little difference in the size ranges of males and females, 

although males did tend to be slightly smaller than females (Fig. 4-91) . 

Ovigerous females averaged 21 .7 mm long, and ranged between 10 and 29 mm . 

More than half of the females measured (284 out of 543) Were ovigerous . 

Non-ovigerous females had a mean carapace length of 18 .7 mm, within a 

range of 9 to 30 mm. Males were between 9 and 28 mm long, with a mean 

length of 17 .1 mm. There appeared to be a break in the frequency 

distribution near 20 mm, which could also be seen in the cruise-by-cruise 

size frequency distribution graphs (Fig . 4-92) . The break was probably 

due to size differences between ovigerous and non-ovigerous females . 

There was a positive linear relationship (r . 0 .95) between the logs of 

weight and carapace length (Fig. 4-90B) . Mean weight was 3.0 g, within a 

range of 0 .1 to 11 .5 g . 
321 

NEMATOCARC/AKJS RIOTUNDiUS J 

Oviqerous FemaN o c ~ T 1 

Female oo h T7 ~ O 

Male 

0 10 20 30 40 

LENGTH (mm) 

Figure 4-91 . Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Nematocarciaus rotundus (small squares 

mark outliers) . 

20 

a 

v 

C 

10 

E 

V 

W 

0 

0 

60-

50-

a 40 -

v 

7p 30-

Y 

20-

10- 

0 

10 20 30 

Carapace Length ( mm) 

10-

8 

a 

s 6 -

0 

v 

6- 

v 

4 -

2-

0 

0 

30 

a 20 

v 

a 

Y 

7 

0 

0 10 20 30 0 10 20 30 

Carapace Lemok (mm) Carapace Length (mm) 

Figure 4-92 . Frequency histograms for carapace lengths of Nematocarcinus 

rotundua from individual cruises . 

10 20 30 

Carapace Length (mm) 
322 

4 .3 .4 .8 Penaeopsis serrata (Bate, 1881 Family Penaeidae) 

Distribution 

The big-eyed shrimp, Penaeopsis serrata , is the most abundant deepwater penaeid in the Gulf of Mexico, being especially common near the 

upper edge of the continental slope (Pequegnat and Chace 1970, Pequegnat 

et al . 1971) . Large numbers have been taken in the De Soto Canyon in 

water about 3T0 m deep, and off the Rio Grande (about 275 m deep) . It is 

found in the subtropical Atlantic north of the equator, from the Caribbean 

to South Carolina, and along the west coast of Africa . Pequegnat (1983) 

described it as present in the Alaminos samples from 200 to 550 m depths, 

with a population peak in the Shelf/Slope Transition Zone at 300 m. 

During this program, Penaeopsis serrata was collected on all five 

cruises, within a fairly narrow depth range : from 329 to 464 m (Fig. ug3) . Specimens were taken on the Central Transact from 329 to 346 m ; on 

the Eastern Transect from 345 to 366 m ; on the Western Transect at 342 m ; 

and on the blest Central Transact from 368 to 464 m. 

Measurements 

Specimens of 520 Penaeopsis serrata measured in this program ranged 

from 8 to 32 mm in carapace length (Fig . 4-94A) . The overall size 

frequency distribution formed a bell-shaped, unimodal curve. Mean length 

was 19.3 mm. Males tended to be somewhat smaller than did females (Fig. 4-

95), acid females achieved greater maximum sizes. Mean length for females 

was 23 .1 mm, within a range of 12 to 32 mm. Males were between 12 and 26 

mm, with a mean length of 19 .3 mm . No obvious seasonal trends in size 

frequencies were evident (Fig. 4-96) . There was a positive linear 

correlation (r -_ 0 .89) between the logs of specimen length and weight 

(Fig . 4-94B)" Mean weight was 6 .0 g, within a range of 0 .2 and 13 .2 8" 
323 

Figure 4-93 " Locations Where Penaeopais serrate were collected (closed 

circles) . 

t0 

SP 60 

u 

e 

V 

40 v 

Y 

V 

20 

a 

va 2 

v 

.r 

v 

o 

C 

J 

oensBnnsis ser~ets 

Figure 4-94 . Frequency histogram (A) and size-Weight relationship (B) 

for carapace lengths of Penaeopsis serrate (all stations 

pooled) . 

0+ 

0 10 ------ 20 30 40 

Carapace Length (mm) 

_2 

2. 2.3 3.0 13 410 

Ln CsrsNce Length (on) 
324 

PEN.4EOPS/S SER4ATA 

Oaq O 

Female 

Male 

0 10 20 30 40 , 

LENGTH (gym) I 

Figure 4-95 . Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Penaeopsis serrata (small squares mark 

outliers) . 

POWesopsis sanrste 

20 

a 

s 10 

v 

e 

O 

50 

40 

a 

30 

V 

s 

s 

201 L 

10 

0 

10 20 3() 

Csrspme Length (gym) 

LV 

G 

tU 

r O 

L 

0 

0 10 20 30 40 0 10 20 30 40 

Care Lent" (on) CareNcs LshtA (mm) 

Figure 4-96 . Frequency histograms for carapace lengths of Penaeopsis 

serrata from individual cruises . 

10- 

e 

~' 6- 

a 

e 

2 

0 

40 O 10 20 30 40 

Carapace len4tb (mm) 
325 

4 .3 .4 .9 Plesionika holthuisi (Crosnier & Forest, 1967 Family Pandalidae) 

Distribution 

Holthuis's pandalid shrimp, Plesionika holthuisi is a widely 

distributed caridean that has been collected on the west coast of Africa 

in water 500 m deep, and in the northern Gulf of Mexico between 479 and 

864 m (Pequegnat and Chace 1970) . It has been described as the most 

abundant deep-water pandalid in the Gulf above 800'm (Pequegnat et al . 

1971) . Pequegnat (1983) ascribed a depth range of 500 to 900 m to 

Plesionika holthuisi, with a population maximum in the Archibenthal Zone 

at 650 m. 

During this program, Plesionika holthuisi Was collected on all five 

cruises, within a depth range of 366 to 871 m (Fig. 4-97) " Specimens were 

taken on the Central Transect from 474 to 850 m; on the Eastern Trarisect 

from 366 to 871 m ; on the Western Transact at 654 m; and on the West 

Central Transact from 464 to 774 m" 

Measurements 

Specimens of Plesionika holthuisi collected during the Alaminos 

cruises were between 6 and 18 mm in carapace length. Ovigerous females 

were 9 to 1 4 mm long, and mere collected in July, October, and November 

(Pequegnat and Chace 1970) . 

The 459 individuals of Plesionika holthuisi measured in this program 

averaged 11 .2 mm in carapace length, and ranged from 5 to 17 mm (Fig. 4-

98A) . Males and females were quite similar in size (Fig. 4-99) . Mean 

length for ovigerous females was 11 .8 mm, within a range of 8 to 17'.mm. 

About two thirds (216 out of 333) of the females that were measured were 

ovigeroua. Non-ovigerous females averaged 11 mm in length, and were 

between 6 and 16 mm long . Males averaged 10 .7 mm in length, and were 

between 6 and 16 mm long. The overall frequency distribution for P. 

holthuisi formed a bell-shaped, uni modal curve, without obvious seasonal 

trends (Fig . 4-100) . There was a positive linear correlation (r = 0 .89) 

between the logs of specimen width and weight (Fig. 4-98B). Mean weight 

was 1 .1 g, within a range of 0 .2 to 3 .9 g" 
326 

Figure 4-97 . Locations Where Pleaionika holthuisi were collected (closed 

circles) . 

020 

100 

6p 

V C 

v 

v 60 

Q 

q 

L 40 

ZO 

z 

a 

v 

r C 

a 

'u ° 

e 

J 

P1oc3nisf/~w An1lAiiici 

=0 .89 

Figure 4-98 . Frequency histogram (A) and size-weight relationship (B) 

for carapace lengths of Plesionika holthuisi (all stations 

pooled) . 

0 

O 10 20 

Carapace Length ( mm) 

_2 

1 A 2A T~ iA 

le Carapace length (mow) 
327 

fLE37AN/KA HOLTM/S/ 

ovtQ*rous Female a 

Female 0 1 O 

Male p ~ 

T 1 0 

0 2 4 6 8 10 12 14 16 18 

LENGTH (mm) 

Figure 4-99 . Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Plesionika holthuisi (small squares 

mark outliers) . 

20 

u C 

10 

C 

a 

0 

60 

u 

e 

40 

v 

Y 

L 

20 

0 

o%ston/Re Ao/1/ra/s~ 

20-1 

u 

C 

10 

O 

Y 

10 

Carapace Length (mm) 

40 

30 

a 

u 

e 

Q 

a 

L 

W 

10 

O 

10 

Carapace length (mat) 

~-. . .; -- v 

20 

0 10 20 0 10 20 

Grahee Length (am) carepece Length (am) 

Figure 4-100 . Frequency histograms for carapace lengths of Pleaionjka 

holthuisi from individual cruises . 

20 0 
328 

4 .3 .4 .10 Rochinia erassa (A . Milne Edwards, 1879 Family Majidae) 

Distribution 

Roehinia crassa , the short-clawed spider crab, is found in water 

along the Atlantic coast of the US from New England to the Straits of 

Florida, and throughout the Gulf of Mexico except the southwest portion 

(Pequegnat and Chace 1970) . The Alaminos collection includes six 

specimens from between 382 and 728 m ; deeper records exist off Cuba (Chace 

1940) . In the Straits of Florida, it has a somewhat shallower 

distribution, from 180 to 642 m, and is apparently confined to water 

colder than 100C (Soto 1982). Soto states that it is "exclusively (a) mud 

dweller .* 

During this program, Rochiaia erassa was collected on Cruises II 

through V, over a depth range of 338 to 1,170 m (Fig. 4-101) . Specimens 

were taken on the Central Transact from 338 to 632 m ; on the Eastern 

Transecst from 351 to 1,170 m; on the Western Transact at 342 m; and on the 

West Central Transact from 36$ to 751 m. 

Measurements 

Those few specimens of Rochinia crassa collected by the Alaminos were 

between 11 and 72 m m in carapace width (Pequegnat and Chace 1970). 

In this study, 51 measured specimens of Rochinia crassa Were between 

9 and 126 mm in carapace width. Males had a mean width of 37 .6 mm, within 

a range of 14 to 126 mm. Non-ovigerous females were from 10 to 56 mm 

wide, with a mean of 30 .6 mm. Ovigerous females had a mean width of 63 .1 

mm, and were between 57 and 70 mm wide. 

The size frequency distribution showed some evidence of bimodality 

(Fig. 4-102A), but sample size was too small to permit the differentiation 

of seasonal trends (Fig. u-103) " Most specimens were less than 50 mm 

wide, but there was a second apparent peak in size, with very large males 

making up the second peak (Fig. 4-10u) . This may have been due to sexual 

dimorphism, although biased handling of specimens on deck (catch 

selection) Was known to have taken place for this species . There was a 

positive linear relationship (r . 0 .97) between the logs of weight and 
329 

12 

10 

e 

ar 

6 

v 

N 

L 

Z 

e 

6 

.. 

.. 

t 

-a 

e o 

J 

Figure 4-102 . Frequency histogram (A) and size-weight relationship (B) 

for carapace widths of Rochinia erassa (all stations 

pooled) . 

Figure 4-101 . Locations where Rochinia erassa were collected (closed 

circles) . 

0 

O 100 20O 

Csrspsce Width (mm) 

1 

Z 

L . csr.pa Width (no) 
330 

3 

a 2 

v C 

v v 

v Y 

0 -t 

Cruise 11 3-

s 2 

v C 

u 

v Y 

2O 

Carapace Width (mm) 

100 200 

Carapace Width (mm) 

^------ " V g ^----- 

4 - 

a 4 - a 

~ 3- 

3- 

i 

2- 

2 

' l O 0- 

0 1oo 2oo o 1oo 2oo 

Carapace Width (mu) Carapace Widtb (MM) 

Figure 4-103 . Frequency histograms for carapace widths of Rochinia crassa 

from individual cruises. 

I ROA51/N/A CRASSA 

Rochinte cresse 

Ovlq*rous Female 

Female 

Mals o 0 0 

0 20 40 60 60 100 120 140 

WIDTH (mom) 

Figure 4-104 . Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace widths of Rochinia crassa (small squares mark 

outliera) . 
331 

carapace length (Fig . u-102B) Mean weight was 41 .6 g, within a range of 

0 .1 to 454 .9 8 " 
332 

4 .3 .4 .11 Stereomastis sculpta (S. Smith, 1882 Family Polychelidae) 

Distribution 

The flat-backed lobsterette, Stereomastis sculyta, is one of the most 

widely distributed polychelids. The subspecies Stereomastis seulata 

seulpta has been reported from the eastern Atlantic, Mediterranean, and 

the Indo-Pacific near Indonesia, the Arabian Sea, as well as the Gulf of 

Mexico and the Caribbean (Firth and Pequegnat 1971, gensley 1981, Williams 

and Wigley 1977) . Another subspecies, S. sculpta oacifica, replaces it in 

the eastern Pacific. The depth range of S. sculpta sculpta is from 230 to 

4,000 m . It is the most frequently captured polychelid in the Gulf and 

the Caribbean, having been taken by the Alaminos in depths of 476 to 1875 

m. It is abundant enough to have been considered for commercial fisheries 

(BCF 1969) " Its depth range in the Gulf has been given by Pequegnat 

(1983) as from 500 to 2,750 m, with maximum population density in the 

Upper Abyssal Zone at 1,050 m. 

Stereomastis sculpta was collected during this program on all fine 

cruises, at depths from 603 to 2,857 m (Fig. 4-105)" Specimens were taken 

on the Central Transect from 6Q3'to 1,510 m ; on the Eastern Transect from 

612 to 2,857 m ; on the Western Transact from 828 m to 1 ,413 m ; and on the 

West Central Transact from 643'to 1,206 m. 

Measurements 

Specimens of Stereomastis sculpta collected during the Alaminos 

cruises were from 17 to 70 m m in carapace length, with ovigerous females 

43 to 58 mm long, and ovigerous females with spermatophores attached 

between 41 and 70 mm long. 

During this program, 430 measured individuals of Stereomastis sculpta 

ranged from 15 to 61 mm in carapace length, averaging 33 .4 mm (Fig. 4-

1061) . Mean length for males was 34.1 mm, within a range of 17 to 42 mm. 

Non-ovigerous females were between 16 and 58 mm long, with a mean of 32 .5 

mm. Ovigerous females had a mean length of 51 .8 mm, and were between 44 

and 61 mm long. Females overlapped males at both ends of their size range 

(Fig. 4-107) " Their overall frequency distribution formed a bell-shaped, 
333 

Figure 4-105 . Locations Where Stereomastis sculpta were collected (closed 

circles) . 

801 A 

a `° 

V 

Q 

b 

a ,o 

v 

v. 

zo 

a 

a 

v 

... , 

u 

e 

J 

O 

Stereomastis scdJpts 

Figure 4-106 . Frequency histogram (A) and size-weight relationship (B) 

for carapace lengths of Stereomastis sculpta (all stations 

pooled) . 

0 

O 20 ~0 60 60 

Carapace Length (aim) '' 

Z3 7A 3,3 "D 4A 

Le Carapace lsMt" (ow) 
334 

STEfTfOMAST/S SCULPTA 

Oviparous Female 

Female 

Male 

0 10 20 30 40 50 60 70 

L LENGTH mm 

Figure 4-107 " Median size (dark vertical bars), 25th and 75th percentiles 

(boxes), and approximate confidence intervals (whiskers) on 

carapace lengths of Stereomastis sculpta (small squares 

mark outliers) . 

1o 

s 

c 

a 

r q 

2 

0-- 

0 

40 

30 

a 

u 

ep 

7 20 

v 

v 

10 

0 

0 

20 40 60 

Carapace Length ( mm) 

Ste~eomestis sculpte 

10 -

a -

e 6-

v 

'` 4 

W 

2 

0 

80 O 

30 

a 20 

v 

G 

u 0 

v 

v 

10 

0 

0 

Figure 4-108 . Frequency histograms for carapace lengths of Stereomastis 

sculDta from individual cruises . 

ZO 40 60 80 

Carapace Length (mm) 

20 40 60 so 

Carapace Length (111111) 

^---=-- 

20 40 60 90 

Carapace Leith (mm) 
335 

unimodal curve. No obvious seasonal trends were evident (Fig. 4-108) . 

There was a positive linear correlation (r = 0 .97) between the logs of 

specimen width and weight (Fig. 4-106B). Mean weight was 10.5 g, within a 

range of 0 .7 and 54 g. 
336 

4 .3 .5 FISH SPECIES ACCOUNTS 

4 .3 .5 .1 Dibranehus atlanticus 

The batfish, Dibranehicus atlanticus , occurs in the eastern and 

western Atlantic and is the only member of the genus that has been 

reported in the western North Atlantic (Bradbury 1967, 1977) . In the 

northern and central Gulf of Mexico, Bright (1968) collected this species 

from 165-869 m in conjunction with temperatures of 18 .9-9 .3°C. Rayburn 

(1975) extended the maximum depth to 1353 m and reported that the greatest 

concentrations of D . atlanticus occurred from 360-500 m. 

D . atlanticus was the most abundant fish species collected during 

this project. During Cruises II-Y, a total of 575 specimens were taken in 

32 trawls at 29 stations (Fig. 4-109). Based on the average depth of each 

trawl, D . atlanticus were collected at depths ranging from 351-964 m . 

Temperatures at these depths ranged from 10-6-5-10C (extrapolated from 

nearest hydrographie stations) . 

This species was most abundant at depths ranging from 423-871 m. The 

27 trawls (46% of total) made outside of this depth range accounted for 

only 1 .4% (n -_ 8) of all specimens collected . This trend was consistent 

among transects and cruises. Depth related abundance levels also varied 

in conjunction with fish size . The 519 specimens examined ranged in total 

length from 30-145 mm (mean = 77 .7, S.E. -_ 1 .3) (Fig. 4-110A) . However, 
30' 

28' 

Z6° 

Figure 4-109 . Locations where Dibranchus atlanticus were collected (closed circles) . w 
338 

Dibrencl4us etl&Wcus 

60 -

50 -

a 40, 

v 

C 

30 Q 

as 

Li 20 

10 

0 

4 

3 

a 

v 

r 

a .^ 

as 

C 

J 

-2 

Figure 4-110 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Dibranehus atlanticus (all stations 

pooled) . 

0 100 200 

Length (mm) 

3 4 5 

Ln Length (mm) 
339 

specimens measuring <90 m m total length occurred at depths ranging from 

603-663 m (Fig. u-111), whereas the majority of fish >90 mm were taken at 

depths of 802-871 m (Cruises II, III and IV) and X423-479 m (Cruise V) 

(Fig . 4-111) . The influence of depth ' and fish size coupled with 

disproportionate sampling efforts (by depth and among cruises) obscured 

any geographic and yearly distinctions in abundance levels. 

The In length/ln weight relationship for D. atlanticus was linear (y 

= 11 .20 + 2.985x, r = 0.98) across the entire length spectrum (40-149 mm) 

and indicates a uniform exponential increase in condition for all size 

cohorts (Fig. u-110B) . 

D. atlanticus was the most opportunistic feeder of all the species 

examined. Thirty-three stomachs taken from the Western, Central and 

Eastern Transects contained a total of 19 identifiable prey items. 

Amphipods, polychaetes, isopods and cumaceans were the most frequently 

occurring food items; however, paguroidea, brachyura and thecosom ata were 

also dominant in terms of weight and volume (Fig. 4-112) . Comparisons of 

small (< median fish weight) and large (> median fish weight) individuals 

indicated size related differences in feeding habits . In terms of 

frequency of occurrence, small fish showed a greater preference for 

isopods (40 .0¢ vs 5.6¢) and pyenogonids (20x vs 2%) . 

Rayburn (1975) examined D . atlanticus taken from the northern Gulf of 

Mexico and found that bivalves (43%), gammarid amphipods (26%), and 

ostracods (22%) Were the most frequently occurring of 17 food items. The 

difference in results is undoubtedly due, in part, to size related feeding 

habits and trophic diversity exhibited by this species. 
340 

6 

5 

a 4 

v e 

v 0 3 

v 

t1 

L 2 

0 

0 

50-

40 -

a 

30 -

0 

v 

20 

01 

0 

100 - 

80 -

v 

v 60-

a 

6. 

a 

C 

d 

40 W 

20 -

0 + 

0 

100 

Length (mm) 

Dibrenchus etlenticus 

20 

a 

v 

C 

10 

v 

V 

L 

0 

200 0 

30 

a 20 

v 

C 

u 

e 

v 

H 

10 

200 0 

100 -

so - 

a 

60-

tf 

D 

P 

40 -

L6 

20 -

0 

200 p 

Figure 4-111 . Frequency histograms for total lengths of Dibranchus 

atlanticus from individual cruises . 

100 

Length (mm) 

100 200 

Length (mm) 

100 

Length (mm) 

100 200 

Length (mm) 

100 200 

Length (mm) 
341 

Dibre,7cl4us atlenticus 

229 

269 

7 % 9 % 

5W; 

12% 

Volume 

Gastropods 

Ostracoda 

My sidacea 

Crangonidae 

Paguroidea 

Pycnogonida 

Brachyura 

a Foraminifera 

o, Scyphozoa 

Polychaeta 

v Tanaidacea 

e Cumacea 

,i Harpacticoida 

Isopoda 

Amphipoda 

Trematoda 

Nematoda 

Ophiuroidea 

Bivalva 

Crustacea Parts 

Unident . Mat. 

289 

D Polychaeta 

Amphipoda 

Paguroidea 

Brachyura 

Thecosomata 

D 13 Catagories 

12% 

0 20 40 60 80 100 

Frequency of Occurrence ( Z) 

Figure 4-112 . Stomach contents found in Dibranchus atlanticus plotted by 

volume and weight (upper) and by percent frequency of 

occurrence . 

7 % 

41% 

Weight 
342 

u " 3 .5 .2 Nezumia aeQualis 

The smooth grenadier, Nexumia aegualis , is found in the eastern 

Atlantic from Faroe Bank to northern Angola and into the Mediterranean, 

and in the western Atlantic from Davis Straights to northern Brazil and 

through the Gulf of Mexico. Its bathymetric range has been listed as 200-

1000 m (Marshall and Iwamato 1973) " Recent collections in the Norfolk 

Canyon off Virginia found N. aegualis at depths of 367-1109 and in 

temperatures of 4 .3-8 .0°C (Middleton and Musick 1986) " 

A total of 236 N. aegualis was collected in 26 ',trawls and at 20 

stations during all five cruises of this project (Fig. 4-113) . Specimens 

were collected over a relatively wide bathymetrie range (474-871 m), based 

upon average depth of trawl . Relative abundances of this species 

indicated no obvious center of distribution either by depth or by 

geographic region within the Gulf. 

Head lengths for the 201 specimens measured during this project 

ranged from 14-45 mm (mean = 35 .6 mm, S. E. -_ 0 .4 mm ; Fig. 4-114A) . The 

length-frequency distribution was modal at 36-38 mm and skewed toward the 

higher end of the length spectrum. These results are biased in favor of 

Cruise IV catches since they constituted 65% of the total sample size 

(Fig . 4-115) " 

Middleton and Musick (1986) found a positive correlation between size 

and depth-distribution for N. aegualis collected on the east coast of the 
M 

M 

" (saTD.ziD pasoTD) palOaTToO azari stZsn aE Vztunzapl aaauM suoiIPDo7 " £TT_~ azn$z3 

v9e 

.8Z 

.0£ 
344 

Nezumie eequalis 

50 

40 

a 30 

C 

d 

7 

Q 

6. 20 

v. 

10 

0 

0 

5 

4 

a 

v 

rr C 

a 

.,. 

y3,1 

i 

e 

J 

2 .6 

Figure 4-114 . Frequency histogram (A) and size-weight relationship (B) 

for total lengths of Nezumia aegualis (all stations 

pooled) . 

10 20 30 40 50 

Length (mm) 

2 .8 3.0 3 .2 3.4 3.6 3 .8 4 .0 

Ln Length ( mm) 
345 

Nezu1we eequelis 

4 

3 

a 

v 

e 

0 

P 

L 

0 

0 

40 -

30-

a 

e 

~ 20-

P 

4) 

66 

10 - 

01 

0 

10 20 30 40 

Length (mm) 

50 

10-

aa 

6- v C 

w 

0 

v 

c+ 

L 4 -

21 

0 

0 

5 -

4 -

a 

v e 3-

7 

P 

V 2-

1 

0-

0 

10 20 30 40 

Length (mm) 

50 

Figure 4-115 . Frequency histograms for total lengths of Nezumia aequalis 

from individual cruises. 

10 20 30 40 50 

Length (mm) 

10 20 30 40 50 

Length (mm) 
346 

United States . For fish collected at depths of 600-700 m and 800-900 m, 

mean head lengths were 35 .0 and 39 .0 mm, respectively. These data are 

consistent with the findings of Middleton and Musiek (1986) . However, 

while larger fish (>4O mm) were found at both depth ranges, few small fish 

(<35 mm) were caught at 800-900 m stations . 

Only thirteen stomachs of N . aegualis were examined, all of which 

contained some prey items. Nantantia was the most important of the nine 

identifiable food categories taken in terms of volume, weight and 

frequency of occurrence (Fig. 4-116) . Amphipoda was the only other mayor 

food item by volume and weight while calanoid copepods and tanaidaceans 

represented mayor food items based upon frequency of occurrence. 

Rayburn (1975) also examined the diets of N. aegualis collected in 

the Gulf of Mexico. Based upon frequency of occurrence, he found 

polyehaetes (88%) to be the most important food item followed by gammarid 

amphipods (60%), tanadacians (WOK), ostracods (36'x) and cumaceans (30K) . 

More recently, Mauchline and Gordan (1984) performed detailed trophic 

analysis on N. aequalis collected in the northeastern Atlantic. Based 

upon the examination of 1013 individuals, they found amphipods to be the 

dominant food item by volume (50-60$) and by frequency of occurrence 

(80%) . Calanoid copepods, mysids, polychaetes, isopods, decapods and 

brittle stars also contributed substantially to the diet. They concluded 

that N. aegualis consumed a wide variety of food items, all of which were 

organisms closely associated with sediment surface . 

The In transformed length/weight regression yielded a strong linear 

relationship (y . 8 .202 + 3 .195x, r = 0 .96), indicating a uniform 

exponential increase in condition factor with increasing length (Fig. u114B) . 
347 

12% 

639 

Volume 

a 

0 

a 

v 

v 0 0 

Poly chaeta 

Tanaidacea 

Calanoida 

Isopoda 

Amphipoda 

Natantia 

Brachyura 

Ophiuroidea 

Trematoda 

Crustacea Parts 

Unident . Mat . 

NezuD?ie eEquelis 

Amphipoda 

0 Natantia 

0 6 Catagories 

4% 149 

829 

Weight 

Figure 4-116 . Stomach contents found in Nezumia aequalis plotted by 

volume and weight (upper) and by percent frequency of 

occurrence . 

0 20 40 60 80 

Frequency of Occurrence M 
348 

u " 3 .5 .3 Bembrons Aobioides 

The percophid, Bembrops gobioides , occurs in the western North 

Atlantic from New York to Florida, throughout the Caribbean and in the 

northern Gulf of Mexico . Its depth distribution ranges from 118-594 m 

with corresponding temperatures of 1u .4-8 .6°C (Bright 1968) . 

B. gobioides was the third most abundant fish species collected 

during this project. A total of 366 specimens were caught in 15 trawls at 

13 stations during Cruises II-V (Fig. 4-117) . Based on average depth of 

trawl, B. gobioides were collected primarily between 338-551 m. The 42 

trawls (72$, of total) made below 511 m resulted in the capture of one 

specimen. With the exception of Cruise I, all trawls made within the 338-

551 m depth range resulted in the capture of B. gobioides regardless of 

transact or cruise. 

Specimens of B. gobioides ranged in total length from 70-262 m m (mean 

-_ 190 .4, S.E. -_ 2 .4) (Fig . 4-118A) . The dominant size cohort during 

Cruises II, IV and V was around 200-240 mm (Fig. 4-119) " The small sample 

size (n -_ 10) obtained during Cruise III prevented interpretation of 

length-frequency data. Multimodal distributions indicated the presence of 

additional size cohorts at 100-120 mm, 160-170 m m and possibly 180-200 mm. 

We found no clear distinctions in depth distribution based upon fish size. 
30° 

28' 

z6° 

Figure 4-117 w 

. Locations where Bembrops gobioides were collected (closed circles) 41 . 
350 

Bembrops gobivides 

50 

40 

a 

30 

0 

20 

v. 

10 

0 

5 

4 

a 

v 

*~ 3 r 

a 

C 

J 

Length (mm) 

Figure 4-118 . Frequency histogram (A) and size-weight relationship (B) 

for total lengths of Bembrops gobioides (all stations 

pooled) . 

0 100 200 300 

5 6 

Ln Length (mm) 
351 

Beinbrops gobioides 

12-

10 - 

a6 -

L6 

a 

C 

P 

61 

4 -

2-

0 - 

0 

30 

a 20 

Gf 

e 

P N 

~i 10 

0 

0 

4 

3 

a 

V 

C 

v 

V 

L 

0 

0 

15 

a 10 

c> 

e V 

7 

P 

u 

~i 5 

0 

Figure 4-119 . . Frequency histograms for total lengths of Bembrops gobioides 

from individual cruises . 

100 200 300 

Length (mm) 

100 200 300 

Length (mm) 

100 200 300 

Length (mm) 

100 200 300 

Length (mm) 
352 

The In length/ln weight relationship (Fig. 4-118B) for all specimens 

combined was linear (y = -13 " 043 + 3 .13x, r . 0 .98) indicating a uniform 

increase in condition across size cohorts (70-262 mm) . 

B. gobioides appears to be an active predator that feeds primarily on 

large nektonic organisms. Seventeen of the 30 stomachs taken from fish 

collected in the Eastern, Central and Western Transects contained only 

three identifiable organisms : Nantantia, Osteichtys and Forminifera. 

Nantantia was the most abundant food item in terms of weight, volume and 

frequency of occurrence (Fig. 4-120) . Because the dominant prey items are 

large motile organisms, it is likely that Foraminifera were consumed 

incidentally. Bright (1968) examined three specimens of B. gobioides and 

reported similar results . In addition to the predominance of large 

macruran decapods, he also found one gammarid amphipod, one calinoid 

copepod and one mysid. 

Comparison of stomach contents between small (< median fish weight) 

and large (> median fish weight) individuals indicate similar diets with 

one exception. Six of the seven large individuals examined contained 

Nantantia while only two of the 10 small specimens contained this prey 

item . 
353 

Be1"brops gnbioidas 

9 % 

Volume / Foraminifera Weig ht 

D Natantia 

Q Osteichthys 

Foraminifera 

a Natantia 

0 

a 

as 

Osteichthy es 

V 

v 0 

Crustacea Parts 

Unident . Mat . 

19~ 

Frequency of Occurrence (%) 

47.0 

Figure 4-120 . Stomach contents found in Bembrops gobioides plotted by 

volume and weight (upper) and by percent frequency of 

occurrence . 

0 10 20 30 40 50 
354 

u " 3 .5 .4 Coelorinchus caribbaeus 

:e.' ~.' . . ,. ' .~ .~ . ~~ . : . : .. . . .' . . . . . . , 

Coelorinehus caribbaeus is a shallow water grenidier found in the 

western North Atlantic from Virginia (Middleton and Musick 1968) to 

northeastern South American waters and throughout the Caribbean and Gulf 

of Mexico . Its depth range is listed at 183-732 m with populations 

apparently centered between 329-412 m (Marshall and Iwamoto 1973) " 

C . caribbaeus was collected during all five cruises but the 177 

individuals were taken in only eight trawls at six stations (Fig. 4-121) . 

The most productive station was C1 where specimens were taken on all three 

sampling occasions (Cruises I, II, III) . This station accounted for 71% 

(n -_ 125) of all C. caribbaeus caught during this project. The other five 

stations were C2 (n = 1), W1 (n = 3), 1A (n . 20), WC1 (n = 10), and WC5 

(n -_ 12) . Except for Stations C2 (786 m) and WC5 (423 m), all fish were 

collected within a narrow depth range of 329-368 m . This bathymetric 

range overlaps well with that reported by Marshall and Iwamato (1973) . C. 

caribbaeus was not present in all areas based upon trawl depth. The 

average trawl depths at Stations E1, 1, 1B, and 1C were between 315-366 m, 

yet no specimens were taken. 

The 124 specimens of C . caribbaeus examined ranged in head length 

from 25-72 mm (mean = 54 .2, S.E. -_ 1 .0) (Fig . 4-122A) . For each of the 

five cruises there was a dominant size cohort in the 55-65 mm area (Fig. 

4-123). The only signs of a distinct multimodal distribution was during 
Figure 4-121 . Locations where Coelorinchus caribbaeus were collected (closed circles) . 
356 

Coelorinc/4us caribbaeus 

30 

a 20 

C 

d 

7 

P 

L 

10 

0 

5 

 4 

a 

v 

rr 

r 

d 3 

e 

2 

32 

Figure 4-122 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Coelorinchus caribbaeus (all stations 

pooled) . 

20 40 60 80 

Length (mm) 

3.4 3.6 3.8 4 .0 4 .2 4.4 

Ln Length (mm) 
357 

10-

8 -

a 

v 6 -

C 

V 

D 

P 

u 4 -

L 

W 

2 -

0--

0 

5 -

4 -

a 

w 

e 3 -

d 

e 

v d 2- 

1 

0--

0 

10- 

8 -

a 

c+ 

c 6 -

7 

Q 

d 4 -

2 -

0 -- 

0 

Cvelorincbus c6pribboeus 

^------ 1 8-1 

20 40 60 

Length (mm) 

6 

a 

C 

4 

Q 

31 

L 

W 

2 

80 0 

8 

6 

a 

v 

C 

4 

P 

u 

u. 

2 

80 

r_. .t__ tI 

Figure 4-123 . Frequency histograms for total lengths of Coelorinchus 

caribbaeus from individual cruises . 

20 40 60 80 

Length (mm) 

20 40 60 80 

Length (mm) 

20 40 60 

Length (mm) 

0 

0 20 40 60 80 

Length (mm) 
358 

Cruise I in which there was a secondary size aggregation at about 30 mm. 

The In length/ln weight relationship was linear (y = 8.429 + 3.013x, 

r = 0 .99) throughout the entire length spectrum (25-72 mm) and indicated a 

uniform exponential increase in condition across at least two size cohorts 

(Fig. 4-122B) . 

All 15 stomachs examined came from collections at Station C1 . 

Fourteen identifiable food items were taken by this species, indicating 

that it exploits a wide array of prey (Fig . 4-124) . Polychaeta was the 

most important food category by volume and weight but ranked third behind 

Amphipoda and Calinoida in frequency of occurrence . While Natantia 

occurred in only 26 .7 of the stomachs examined, it was the second most 

dominant food item by volume and weight (Fig. 4-121) . 
359 

12% 

109 

a 

L 

0 

a 

c.~ 

0 0 

Foraminifera 

Scyphozoa 

Polychaeta 

Tanaidacea 

Harpacticoida 

Calanoida 

Isopoda 

Amphipoda 

Ostracoda 

Mysidacea 

Natantia 

Brachyura 

Coleoidea 

Osteichthy es 

Nematoda 

Crustacea parts 

Unident. Mat . 

Polychaeta 

D Natantia 

D 10 Catagories 

Calanoida 

Amphipoda 

Brachyura 

Weight 

5535 

Figure 4-124 . Stomach contents found in Coelorinchus caribbaeus plotted by 

volume and weight (upper) and by percent frequency of 

occurrence . 

14% 

Volume 

0 20 40 60 80 100 120 

Frequency of Occurrence ( %) 
360 

u " 3 .5 .5 Poecilopsetta beani 

The right-eyed flounder Poecilopsetta beani occurs in the western 

Atlantic from New England to northeastern South America and throughout the 

Caribbean and Gulf of Mexico. It has been reported over an extremely wide 

bathymetrie range (53-1637 m) but is apparently most common between 180-

550 m (Bright 1968) . 

P. beam was collected during every cruise in a total of 13 trawls at 

11 stations (Fig. 4-125) . Specimens were generally restricted to the 

shallowest areas sampled during this project . Of the 155 individuals 

taken, 93$ (n = 142) were collected at depths ranging from 329-368 m. The 

only station that fell within this depth range and where no specimens were 

found was E1 (366'.m) . 

P. beam ranged in total length from X10-122 mm for the 137 ; gPecimens 

examined (mean -_ 83 .3, S.E. . 1 .7) (Fig . 4-126A) . The largest sample was 

collected during Cruise IV, and the length-frequency distribution 

indicates the presence of at least three size groups : a dominant mode at 

50-60 mm and secondary modes at 85T90 mm and 105-110 mm (Fig. 4-127) " 

Samples were considerably smaller for the remaining cruises but there was 

evidence of dominant size cohorts at 90-100 mm during Cruises I, II, and V 

(Fig . 4-126B) . 
Figure 4-125 . Locations where Poecilosetta beans were collected (closed circles) . 

30° 

28° 

Z6° 

w 

a~ 
362 

Poecilopset to bee,7i 

Zo 

a 

C 

d 

10 

Q 

d 

L 

0 4B 

40 

4 -

3-

a 

2- C 

a 

d 

1 

e 

J 

01 

3 .6 

Figure 4-126 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Poecilosetta beans (all stations 

pooled) . 

60 80 100 120 

3.8 4 .0 4.2 4 .4 4.6 4.8 5.0 

Ln Length (mm) 
363 

poecilopsetta besni 

10 -

8 -

a 

c, 6 

e 

C1 

P 

4 - L 

21 

0 

0 

15 

a 10 

v 

e 

0 

P 

5 

10 -

8 -

a 

v e 6-

V 

7 

Q 

v 4 L 

2 -

. 0--

0 

4 

3 

a 

v 

c 

2 

Q 

u 

1 

0 

0 

0 

Figure 4-127 . Frequency histograms for total lengths of Poecilosetta beani 

from individual cruises . 

50 100 150 

Length (mm) 

50 100 150 

Length (mm) 

50 100 150 

Length (mm) 

50 100 150 

Length (mm) 
364 

The In transformed length weight relationship was linear (y = -11 .778 

+ 3.062, r _- 0.98) for this species and indicates an exponential increase 

in condition for fish within the length range sampled (22-122 mm) (Fig. 

4-126B) . 

Stomach analysis was performed on 21 specimens of P. beani collected 

from Stations W1 and C1 ; however, 12 of these were empty. Amphipods were 

the preferred prey item in terms of volume, weight (Fig. 4-128) and 

frequency of occurrence (Fig. 4-128) . Calanoid copepods and mysids were 

also substantial dietary components with polychaetes being the only other 

identifiable item. 
365 

poecilopswt to b~eiri 

1% 

Polychaeta 

Calanoida 

0 Amphipoda 

12 Mysidacea 

Poly chaeta 

Calanoida 

Amphipoda 

0 

a 

My sidacea 

Nematoda 0 0 

Crustacea Parts 

Unident . Mat. 

129 1 % 109 

Figure 4-128 . Stomach contents found in Poecilosetta beam plotted by 

volume and weight (upper) and by percent frequency of 

occurrence . 

669 

Volume 

779 

Weight 

0 20 40 60 80 

Frequency of Occurrence ( S6) 
366 

x " 3 .5 .6 Gadomus longifilis 

Gadomus lonAifilis occurs over s wide bathymetric range (630-2165 m) 

in tropical and sub-tropical regions of the Atlantic . In the east it is 

found from Portugal to the Gulf of Guinea and off the Azores and Canary 

Islands. In the western Atlantic, it has been reported from the east 

coast of Florida, throughout the Gulf of Mexico and the Caribbean Sea 

(Iwamoto 1970, Marshal and Iwamoto 1973) . 

G. lonAifilis were collected during all five cruises at depths 

ranging from 812-1203 m. The 191 specimens were taken in 11 trawls at 11 

stations (Fig . 4-129), of which 110 (58$) were collected at Station WC11 

(1135 m) and 31 (16%) at Station E4 (1170 m) . Thus, nearly three-fourths 

of all specimens were taken within a narrow depth range of 1135-1170 m. 

Specimens ranged in head length from 16-46 mm (mean . 32 .3, S.E. -_ 

0 .5) (Fig. 4-130A) . The largest sample of fish measured came from Cruise 

V (n = 85) and the length-frequency distribution was bimodal at 28 and 35 

mm (Fig. 4-131) . Specimens collected during Cruise II (n -_ 27) were 

unimodal at 22 mm. This differnee is apparently not depth related since 

most of the specimens were taken in the aforementioned 1135 and 1170 m 

trawls. Whether the difference is a yearly or geographic phenomenon or 

dust an artifact of sample size is unknown. 
30° 

28' 

260 

w 

o, 

v 

Figure 4-129 . Locations where Gadomus longifilis were collected (closed circles) . 
368 

6adnirrus 1olrgiTilis 

30 

20 

a 

v 

e 

w 

P 

d 

L 

L6 

10 

0 

4 -

3 -

r 

a 2-

C 

J 

0-7-

2 .6 

Figure 4-130 . Frequency histogram (A) and size-weight relationship (B) 

for total lengths of Gadomus lonAifilis (all stations 

pooled) . 

10 20 30 40 50 

Length (mm) 

2.8 3.0 3 .2 3 .4 3.6 3.8 4.0 

Ln Length (mm) 
369 

6adomus longililis 

6 

5 

a 4 

v 

C 

3 

P 

d 

21 

0 

0 

5 

4 

a 

v 3 c 

w 

e 

d 2- LL6 

0 

0 

r- .. :se I 

10 20 30 40 

Length (mm) 

SO 

6 

5 

a 4 

v 

e 

3 

Q 

L6 21 

0 

0 

20 

15 

a 

c 

10 

v 

31 

L 

5 

0 

0 

r.. . : . . . . all 

Figure 4-131 . Frequency histograms for total lengths of Gadomus longifilis 

from individual cruises . 

10 20 30 40 50 

Length (mm) 

r . . . . . . v 

10 20 30 40 50 

Length (mm) 

10 20 30 40 50 

Length (mm) 
370 

A regression of log transformed length-weight data showed a linear 

correlation (y -_ -8.037 + 3 .034x, r . 0 .95), indicating a uniform increase 

in condition across all specimens examined (Fig. 4-130) . 

Examination of 23 G. lonAifilis stomachs taken from three central and 

one eastern station indicated that this species feeds primarily on 

crustaceans. Calanoid copepods were the dominant food category consumed 

by frequency of occurrence and by volume (Fig. 4-132) " Nantantia 

dominated in terms of weight . Amphipods and mysids also constituted 

substantial portions of the diet. 

In relating diets to fin structure and haw placement, Marshall (1965, 

1977 ; p . 276) pointed out that members of the subfamily Bathygadinae 

(e .g ., Gadomus spp.) are best suited for the consumption of mobile 

benthopelagie prey items . Our results generally tend to support 

Marshall's conclusions although small proportions of tanaidaceans and 

harpaeticoid copepods do represent organisms that are typically benthic in 

nature . 
371 

6edomus longirflis 

3 % 

21 

16% 

Tanaidacea 

Harpacticoida 

Calanoida 

a 

Amphipoda 

as 

o Ostracoda 

c.~ 

°e My sidacea 

0 

Natantia 

Crustacea Parts 

Unident . Mat . 

389 

449 

Calanoida 

Amphipoda 

m Ostracoda 

Mysidacea 

D Natantia 

2 Catagories 

1% 

1 1 gS 

S% 

Figure 4-132 . Stomach contents found in Gadomus longifilis plotted by volume 

and weight (upper) and by percent frequency of occurrence . 

10% 13% 

Volume 

209 

Weight 

0 20 40 60 80 

Frequency of Occurrence (%) 
372 

x "3 .5 .7 Chaunax pictus 

A total of 150 C . pictus were collected during all five cruises and 

occurred in 21 trawls at 19 stations (Fig . 4-133) " While C . ip etus were 

taken at depths ranging from X423-786 m, nearly 82% , (n . 123) of all 

specimens were collected between 603-663 m. This depth concentration 

appears to be geographically independent as specimens were present in all 

trawls made within this depth range, regardless of cruise or station. 

Fish ranged in length from 30-170 mm with the size distribution being 

bimodal at 40 and 125-150 m m (Fig . 4-134A)" The length-frequency 

distributions of specimens taken during Cruises IV and Y suggest that 

depth distribution may be size related (Fig. 4-135). These two cruises 

accounted for 89% .of all specimens examined . During Cruise IV, all but 

one of the 89 fish taken were collected between 615-629 m . The 21 

specimens examined from Cruise V ranged in length from 50-220 mm and the 

size distribution was less distinct (Fig . 4-135) . In terms of average 

depth of trawl nearly all C. in etus <100 mm were taken within the 603-663 

m depth range. 

The linear regression of In transformed length weight data showed a 

strong correlation (y -_ -9 .681 + 2 .842x, r . 0 .97) and again suggests a 

uniform increase in condition across the length spectrum analyzed (Fig. 

134B) . 
Figure 4-133 . Locations where Chaunax p; , were collected (closed circles) . 

w 

V 

W 
374 

30 - 

20- 

a 

v 

e 

u 

P 

G1 

6. 

10- 

6 ] 

5 

v 4 

r 

a 3 . .. 

C J 

3 

CABUirax pictus 

Length (mm) 

Ln Length (mm) 

Figure 4-134 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Chaunaz ip etus (all stations pooled) . 

100 200 300 

4 5 6 
CAeunex pictus 

30 

a 20 

e 

u 0 

v 

Li 10 

0 

0 

30 

a 20 

v 

e 

u 

0 

v 

w 

~ 10 

0 

0 

r~. . ;se Iv 5 

4 

a 

v e 3 

w 

0 

P 

u 

0 

0 

5-

4 -

a 

c 3-

w 

0 

v 

2 -

1 

0 - 

0 

f ruiea V 

375 

Figure 4-135 . Frequency histograms for total lengths of Chaunax pictus 

from individual cruises . 

100 200 300 

Length (mm) 

100 200 300 

Length (mm) 

100 200 300 

Length (mm) 

100 200 300 

Length (mm) 
376 

4 .3 .5 .8 Coelorinchus coelorhynchus 

s/ , ~~~. . . . . . . . 

Coelorinchus coelorhynehus were collected during all five cruises and 

occurred in 17 trawls at 16 stations (Fig . 4-136) " Of the 205 specimens 

collected, nearly 93% .(n = 190) occurred at depths below 479 m . The 

remaining 15 individuals were taken at depths ranging from 521-663 m. Of 

the 15 trawls taken at depths below 557 m, 13 contained specimens of C . 

coelorhynchus. These results appeared independent of geographic location 

and season. 

The 192 specimens of C. coelorhYnchus were examined and they ranged 

in head length from 15-75 mm (mean -_ 48 .9, S.E. . 0 .9) Fig. 4-137A) . The 

overall length-frequency distribution was unimodal at X45-55 mm . The 

smallest fish were taken at Station C1 during Cruises I and II (Fig. u138) ; however, sample sizes were extremely small in both instances and 

prevent speculation as to size related differences in distribution. 

As in most other cases, the In length/ln weight regression for C . 

coelorhynchus was highly linear (y -_ -220 + 3.249x, r - 0 .99) indicating 

a uniform exponential increase in condition across the length spectrum of 

fish examined (Fig. 4-1378) " 
3d° 

28° 

26° 

Fibure 4-136. Locations where Coelorinchus coelorhynchus were collected (closed circles) . 

V 

V 
378 

Coelo~i~rc~us cnelo~/4y~rc/4us 

40 

30 

a 

v C 

20 

v 

41 

L 

4. 

10 

5 

4 

v v~ 3 

y. 

2 

C 

J 

0 

2 

Ln Length (mm) 

Figure 4-137 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Coelorinehus coelorhynchus (all 

stations pooled) . 

20 40 60 80 

Length (mm) 

3 4 5 
379 

3 

a 2 

v 

e 

0 

v 

w 

u~. 1 

80 

1 

0 

0 

12-

10 - 

a 8 -

v 

c 

6 

P 

u 

4 -

2 -

0 

0 

12 

10 

a 8 

v 

e 

6 

P 

V 

1i 4 

2 

0 

0 

Coelnrinc/4us coelor/4y~c/9us 

20 40 60 

Length (mm) 

20 40 60 

Length (mm) 

3 

a 

v 

e 

2 

v 

u 

0 

0 

20 -

a 

v 

e 

10-

P 

0 4 

80 0 

Figure 4-138 . Frequency histograms for total lengths of Coelorinchus 

coelorhynchus from individual cruises . 

20 40 60 80 

Length ( mm) 

20 40 60 80 

Length (mm) 

20 40 60 80 

Length (mm) 
380 

4 .3 .5 .9 Chloropthalmus agassizi 

The 305 specimens of C. aAassizi were collected during all five 

cruises in 13 trawls at 11 stations . This species was consistently 

collected at depths <479 m, a trend that was evident regardless of station 

or trawl--C. agassizi were caught in all 13 trawls at or below 479 m (Fig. 

u-139) . 

All 305 C. agassizi were examined and they ranged in length from 42-

161 mm (mean . 95 .0, S.E. -_ 1 .2) (Fig . 4-140A) . There were some 

differences in length-frequency distributions for the three most 

productive cruises (II, IV, V) . The mode for Cruise IV was 20 mm lower 

than for Cruises II and V (Fig. 4-141) . There was no indication that this 

difference was related either to depth or season and may be geographic in 

nature . 

The regression of In weight against In length was again highly linear 

(y -_ 12 .919, $" E " = 3 .319%, r -_ 0 .98) (Fig. 4-14OB) . 

The stomach contents of 21 C. aAassizi contained 13 identifiable prey 

items (Fig . 4-142) . In terms of frequency of occurrence, Amphipoda, 

Calanoida and Nantantia were the most dominant organisms present . 

Calanoida (21%), Osteichthys (20%), Nantantia (18K), Amphipoda (15%) and 

Polyehaeta (12$) were most abundant by volume while Osteichthys (47%), 

Calanoida (21%) and Nantantia (14%) dominated by weight. 
Figure 4-139 . Locations where Chloropthalmus agassizi were collected (closed 

circles) . 

30° 

28' 

Z6° 

w 
382 

CI41o~ap/4t/9aln~us egessizi 

40 - 

30- 

a 

v 

c 

20 - 

v 

L 

L6 

10- 

0 

4 

3 

.. 

P 

V 

C 

a 

41 

C 

J 

3 

Ln Length (mm) 

Figure 4-110 . Frequency histogram (A) and size-weight relationship (B) 

for total lengths of Chloropthalmus agassizi (all stations 

pooled) . 

100 200 

Length (mm) 

4 5 6 
4 

3 

a 

v 

e 

u 

v 2 

C1 

L 

0 -f 

4 

3 

a 

e 

7 2 

P 

d 

L 

4. 

1 

0 

0 

25 

20 

a 

C 15 

7 

v 

10 

5 -

0- 

0 

C/41n~op/9t/4elirrus egassizi 

Cruise 1 20 

100 

Length (mm) 

Cruise Y 

a 

v 

c 

~ 10 

Q 

L 

200 0 

bV 

50 

a 40 

v 

C 

30 

Q 

d 

L 

20 

10 

0 

200 p 

383 

Cruise II 

100 200 

Length (mm) 

f ruioa IV 

Figure 4-141 . Frequency histograms for total lengths of Chloropthalmus 

agassizi from individual cruises . 

100 200 

Length (mm) 

100 200 

Length (mm) 

100 

Length (mm) 

Cruise III 
384 

CI4loropl4tl4e1mus egessizi 

14% 

1% 

20% 

18% 

12% 

15% 

219 

Volume 

D 

D 

D 

Foraminifera 

Poly chaeta 

Tanadacea 

Harpacticoida 

a Calanoida 

e Amphipoda 

d Ostracoda 

v Mysidacea 

Ophiuroidea 

° 0 Natantia 

Gastropods 

Osteichthy s 

Nematoda 

Crustacea Parts 

Unident . Mat. 

47% 

Polychaeta 

Calanoida 

Wei ght Amphipoda 

Natantia 

Osteichthy es 

6 Catagories 

4% 

Figure 4-142 . Stomach contents found in Chloropthalmus agassizi plotted by 

volume and weight (upper) and by percent frequency of occurrence . 

0 10 20 30 40 50 

Frequency of Occurrence (96) 

7% 7% 
385 

x " 3 .5 .10 Svnaphrobranchus oregoni 

Synaphrobranchus oreQOni was the fourth most abundant fish species 

collected. A total of 354 specimens were taken during all five cruises in 

27 trawls at 24 stations (Fig. 4-113) " S . oreaoni were taken at depths 

(average depth of trawl) ranging from X179-1203 m. This bathymetric 

distribution was not consistent across trawls and stations as seven (21K) 

of 34 trawls taken within this depth range failed to catch specimens. 

While the 285 examined specimens of S. oreAOni ranged in total length 

from 120-691 mm (mean = 415 .2, S.E. = 5 .4), better than 90$ of all 

individuals exceeded 300 mm (Fig. 4-144A) . The dominant size aggregation 

between 300-600 mm was evident during each of the four most productive 

cruises--II, III, IV, and V (Fig . 4-145) . There was some evidence of 

size-dependent depth distribution in that the vast majority of fish <300 

mm were taken at depths of approximately 750-850 m. 

Length-weight regression analysis on In transformed data yielded a 

highly linear relationship (y -_ -17 .570 + 3 .630, r = 0 .96) across all size 

groups tested. Results included four noticeble outliers (Fig. 4-14uB) . 
W 

W 

Figure 4-143 . Locations where Synaphobranchus oregoni were collected (closed 

circles) . 

300 

28° 

MO 
SPrspl4rnb"irc/4us Ore-9917i 

30 

~, 20 

v 

c 

d 

7 

P 

L 

10 

0 

8 

6 -

.. 

v 

*" 4 -

a 

WC 

.^ 

d 

C 

J 

-2 

4 

Ln Length (mm) 

387 

Figure 4-144 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Synaphrobranehus oreAOni (all stations 

pooled) . 

100 200 300 400 500 600 

Length ( mm) 

5 6 7 
388 

Synap~~ob~anc/4us n~egoni 

7 -

6 

5 

a 

e 4 

61 

7 

v 

61 

1 

0 j 

0 

12 

10 

a 

H 

G 

6 

v 

as 

4 

2 

0 

0 

200 400 

Length (mm) 

Cruise IV 

6 

a 4 

v 

e 

u 

7 

P 

u 

0 

600 0 

10- 

8 

a 

cr C 6-

u 0 

v 

u 4 -

2 -

0-- 

0 

Cruise 111 

Figure 4-145 . Frequency histograms for total lengths of Synaphobranchus 

oregoni from individual cruises . 

200 400 600 

Length (mm) 

200 400 600 

Length (mm) 

Cruise Y 

200 400 600 

Length (mm) 
389 

The stomach contents of 24 specimens of S. orepconi were examined. Of 

the fine identifiable food items, Osteichthys, Coleoidea and Nantantia 

constituted nearly 100% of stomach content by volume and weight--With 

Osteiehthys being most dominant (Fig . 4-146) . Osteiehthys also was the 

most abundant prey item in terms of frequency of occurrence and was 

followed by Coleoidea. 
390 

Sy~rap/4robre~cl4us o~egoni 

8% 

47% 

589 

Trematoda 

Nematod< 

a 

L 

a Natanti< 

as 

a 

v Coleoidea 

v 0 0 

Ostichthye_ 

Unident . Mat 

269 

D Natantia 

Coleoidea 

D Osteichthys Weight 

Figure 4-146 . Stomach contents found in Synaphobranchus oregoni plotted by 

volume and weight (upper) and by percent frequency of 

occurrence . 

Volume 

0 20 40 60 80 

Frequency of Occurrence (96) 
391 

x " 3 .5 .11 Urophysis cirratus 

\. . _,. . . . . . . . .. . 

Urophysis cirratus was the second most abundant fish species 

collected during this project . During all five cruises, a total of 380 

specimens were collected in 19 trawls at 16 stations (Fig. 4-147) . Based 

upon average depth of trawl, this species was collected at depths of 329-

625 m. The greatest concentrations, however, occurred at depths ranging 

from 329-179 m and this trend was consistent among all trawls and 

stations--U. cirratus were present in all 14 trawls taken within the 329-

479 m depth range and accounted for 92% of all specimens collected. 

The 300 specimens of U. cirratus that were examined ranged in total 

length from 230-517 mm (mean -_ 328 .4, S.E. . 3 .1) (Fig . 4-148A) . There 

were differences in length-frequency distributions among cruises (Fig. 4-

149) ; however, much of this was attributable to size differentials at 

individual stations. Distribution in terms of specific size cohorts was 

difficult to characterize by depth, location and season because of the 

numerous differentials among stations. 
U 

U 

4J 

b 

ao 

i 

a 

O 

U 

ca 

4-1 

0 

3 

N 

N 

O 

U 

~+ 

c0 

U 

O 

N 

U 

N 

b 

u 

U 

O 

N 

w 

.14 
393 

Urop/4ycis cirratus 

40 -

30 -

a 

v 

e 

20- 

P 

L 

10-

01 

0 

7 

a 6 

v 

r 

G 

a 

C 

5 

5.4 

Figure 4-118 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Urophysis cirratus (all stations 

pooled) . 

100 200 300 400 500 600 

Length (mm) 

5 .6 5.8 6 .0 6 .2 6.4 

Ln Length (mm) 
394 

a 

6 

a 

v 

e 

4 

v 

d 

2 

0 

20 

a 

v 

G 

10 

P 

u L 

s 

6 

a 

v 

C 

4 

P 

L 

W 

2 

0 

20 

a 

v e 

10 

P V 

V 

0 

0 

UropAycis cirretus 

6-

5-

4 - 

C 

3-

P 

' 

a 

v 

2 -

u 

L 

0 

0 

Figure 4-149 . Frequency histograms for total lengths of Urophysis cirratus 

from individual cruises . 

100 200 300 400 500 600 

Length (mm) 

100 200 300 400 500 600 

Length (mm) 

100 200 300 400 500 600 

Length (mm) 

100 200 300 400 500 600 

Length (mm) 

100 200 300 400 500 600 

Length (mm) 
395 

x " 3 .5 .12 Peristedion arayae 

Peristedion grayae were collected during all five cruises in 17 

trawls at 17 stations (Fig . 4-150) . Of the 128 individuals taken, 127 

were caught at depths ranging from 329-663 m" A single specimen was 

caught at 1107 m. 

The 122 specimens examined ranged in total length from 55-205 m m 

(mean = 121 .0, S. E . . 4 .3) (Fig . 4-151A) . Length-frequency distributions 

indicated a greater preponderance of smaller individuals during Cruises II 

and IV (Fig . 4-152) ; however, these results reflect the difference in 

effort at depth among cruises. Specimens of P. rg aYae <125 mm were more 

abundant at depths of 329-368 m while fish >125 mm were more abundant at 

depths of 47u-663 m. 

The In length/ln weight relationship for P. rA ayae was linear (y = 

-11 .632 + 2 .94x, r . 0.99) across the entire length spectrum and indicates 

a uniform exponential increase in condition for all size cohorts (Fig. u151B) . 
O 

p 

N 

U 

O 

N 

u 

N 

u 

U 

O 

N 

b 

U 

U 

41 

b 

W 

~+ 

.,~ 

w 

60 

0 

a 

O 

U 

co 

u 

O 

G 

3 

v 

v 

W 

41 
397 

f'e'ristediolr greyee 

30 

20 

a 

v 

c 

as 

P 

d 

L 

L6 

10 

5 

4 

a 

, 3 

L 

.

a 

.. 

a> 

3r 2 

e 

J 

Length (mm) 

Figure u-151 . Frequency histogram (A) and size-Weight relationship (B) 

for total lengths of Peristedion grayae (all stations 

pooled) . 

0 100 200 300 

4 S 6 

Ln Length (mm) 
398 

10 

8 

a 

v c 6 

C1 

7 

P 

d 4 

2 

0 

0 

20 

a 

v 

e 

10 

P 

w 

30 

a 20 

e 

u 

D 

v 

v 

10 

Pruiea 11 

15 

a 10 

v e 

O 

7 

v 

u L 

5 

25 

20 

a 

15-

q 

7 

v 

` 10- 

51 

0 

0 

I-eristedion groyoe 

10- 

8 

a 

v 6- e 

u 

7 

Q 

4 -

L6 

as 

2 -

0 

0 

Figure 4-152 . Frequency histograms for total lengths of Peristedion rg ayae 

from individual cruises. 

50 100 150 200 250 

Length (mm) 

Cruise Y 

50 100 150 200 250 

Length (mm) 

Cruise IY 

0 

0 50 100 150 200 250 

Length (mm) 

0 

0 50 100 150 200 250 

Length (mm) 

0 

0 50 100 150 200 250 

Length (mm) 

50 100 150 200 250 

Length (mm) 

5 .0 BENTHIC PHOTOGRAPHY RESULTS 
401 

5 .0 BENTHIC PHOTOGRAPHY RESULTS 

All planned photography transeets were successfully completed at 60 

stations. Four categories of subjects were analyzed in the photographs : 

lebensspuren (tracks, trails and impressions left by animals), 

consolidated materials, man-made artifacts, and megafauna. The total 

number of photographic samples and area analyzed for lebensspuren features 

(including consolidated materials and artificats) and biota for Cruises 

II-V appears in Table 5-1 . It will be noted in Table 5-1 that the sample 

sizes differ between the lebensspuren and biota categories, even on the 

same transeets. This difference is because many of the photographs that 

were of sufficient quality to identify megafaunal biota were not of 

sufficient resolution to identify lebensspuren categories with certainty . 

Table 5-1 . Total area surveyed by benthic photography, Cruises II-n . 

Lebenssuuren 

Cruise 2 Cruise 3 Total 

Number of Frames 1275 1046 2321 

Total Area (m2) 369 3100 6569 

Biota 

Cruise 2 Cruise Cruise 4 Cruise _5 Total 

Number of Frames 1279 2328 3126 2414 9,147 

Total Area (m squared) 385 6709 7398 6988 24,590 

5 .1 LEBENSSPUREN 

Lebensspuren features documented on the surface of the bottom 

sediments ware observed at all benthic photography stations. These 

features, caused by the movements of animals living on or below the 

sediment's surface, were the most abundant category of any observations 

represented in benthic photographs. Lebensspuren classification for this 

study followed Ewing and Davis (1967) and used five of the six mayor 

lebensspuren groups described in that study including ridges, lumps, 
402 

grooves, depressions, and sculptured strips. These mayor categories were 

subdivided by two additional levels of hierarchy as shown in Volume III, 

Appendix D1 . permitting fairly distinct morphological descriptions of all 

lebensspuren features . Additionally, a large data base was created 

containing areas and/or length measurements of most lebensspuren features. 

These data can be later utilized for investigations of long-term changes 

in lebensspuren characteristics and may provide clues to associations of 

specific animal taxa and lebensspuren features. 

General descriptions and densities of lebensspuren features 

documented at each station appear below . 

5 .1 .1 STATION SUMMARIES 

Lebensspuren analyses were based upon photography taken during 

Cruises II and III, only . Summaries of the number of frames taken on 

Cruise II that were suitable for analysis, the total areas sampled by 

these frames and their mean depth is shown by Table 5-2 . A corresponding 

results summary (density of the various features by station) is shown in 

Table 5-3 " The same information is provided for Cruise III in Tables 5-4 

and 5-5, respectively. Detailed supportive data can be found in Appendix 

D1-3- 

5 .1 .1 .1 Cruise II West Transact 

Station W1 

All five Western Transact stations were photographed only during 

Cruise II. Station W1 lebensspuren analyses included a total area of 261 

m2 (Table 5-2) . Total lebensspuren density for this station was 182,464 

features/ha (Table 5-3)" The most abundant lebensspuren type was "groups 

of depressions" represented by a total density of 167,460/ha which also 

represented the most abundant lebensspuren tape at any of the 15 stations 

sampled during Cruise II. The vast majority of depression groups 

(165,388/ba) were characterized by a single type, described as a single 

row of depresions arranged in a partial circle. It is not known what 

organism produced this feature, but the very high density of the feature 
403 

5-2. Design table for Cruise II benthic photography lebensspuren . 

STATION : W1 W2 W'- A W5 _ Total 

Number of frames 97 92 49 99 99 436 

Area (m squared) 260 .6 248 .8 94 .6 223 .3 286 .9 1114 .2 

Mean depth (m) 366 604 814 1396 2456 

STATION : C1 C2 -C3 C11 C5 Total 

Number of frames 79 42 100 99 99 419 

Area (m squared) 190 .1 117 .9 338 " 6 230 .6 262 .6 1139 .7 

Mean depth (m) 358 587 798 1420 2436 

STATION : E1 E2 E3 - E~E - Total 

Number of frames 90 34 99 98 99 420 

Area (m squared) 303 .7 130 304 .4 244 .1 233 .3 1215 .5 

Mean depth (m) 347 612 849 1322 2893 
404 

Table 5-3 " I .ebenaapuren mayor category densities (no./hectare) Cruise II 

stations . 

STATION: 

Individual ridges 

Solitary lumps 

Sets of lumps 

Individual grooves 

Sets of grooves 

Solitary depressions 

Groups of depressions 

Strip of adjacent 

depressions 

Total 

STATION : 

Individual ridges 

Solitary lumps 

Sets of lumps 

Individual grooves 

Sets of grooves 

Solitary depressions 

Groups of depressions 

Strip of adjacent 

depressions 

Total 

STATION : 

W1 W2 iii'- W4 W5 - Total 

0 121 317 2,015 4,914 1,723 

3569 11,857 1903 5329 5124 6031 

0 0 423 179 0 72 

3377 5306 32,886 54,816 2510 16,398 

1036 3618 0 627 1464 1553, 

7022 62,784 27,176 17,018 22,760 27,240 

167,460 21,544 2221 269 0 44,222 

0 0 0 0 0 0 

182 , 464 105,229 64,926 80,253 36,771 97,239 

C1 C2 Cj- C4 C5 Total 

53 764 384 87 267 281 

6786 12,218 1949 7069 1638 4782 

0 0 59 87 724 202 

210 12,897 10,9 2689 ' 24,982 10,932 

0 2461 295 0 2018 807 

21,305 46,581 47,814 12,057 67,291 40,519 

9363 10,012 13,231 347 30 .770 13,688 

0 0 207 0 1028 298 

37,717 84,931 74,926 22,336 128,717 71,509 

E1 E2 _E3 E4 E5 - Total 

Individual ridges 0 3461 493 205 22,37 4829 

Solitary lumps 3095 4999 3088 1639 11,702 4656 

Sets of lumps 0 151 230 205 643 239 

Individual grooves 1841 1161 25,033 33 .470 4072 14,389 

Sets of grooves 0 1077 427 614 3086 938 

Solitary depressions 32,99 33 .453 32,293 15,199 29,104 28,424 

Groups of depressions 128,61 238+ 6669 492 3086 34,750 

Strip of adjacent 

depressions 99 0 0 0 0 25 

Total 166,151 46,988 68,233 51,823 74 .067 88,29 
405 

Table 5-u " Design table for Cruise III benthic photography lebensspuren . 

STATION : C1 C6 C2 C3 C7 - C8 

Number of frames 98 101 100 100 92 60 

Area (m squared) 259 .6 338 300 .7 257 210 151 

Mean depth (m) 353 525 629 866 950 1098 

STATION : C9 C4 C10 C11 C5 C12 Total 

Number of frames 89 87 98 96 57 68 106 

Area (m squared) 274 .9 279 313 .6 272 175 .8 238 3100 

Mean depth (m) 1325 1408 1605 207 2401 2882 
0 

Table 5-5 . Lebensspuren mayor category densities (no ./hectare) Cruise III stations . 

STATION: C1 C6 C2 C3 C C8 

Individual ridges 231 798 1231 1050 1167 2320 

Solitary lumps 6317 16,256 16,031 2411 3666 1193 

Sets of lumps 0 0 100 0 0 0 

Individual grooves 31,241 1537 16,098 11,201 36,661 385 

Sets of grooves 77 680 133 1556 167 66 

Solitary depressions 35,555 101,944 116,111 124,654 31,537 8817 

Groups of depressions 2512 23,801 25,078 13,885 6624 0 

Strip of adjacent depressions 0 0 0 0 0 0 

Total 75,964 144,016 175,908 154,758 79,822 16,242 

STATION : - 

C9 _, C4 C10 C11 C5 C12 Total 

Individual ridges 4947 3800 5295 4296 341 6679 2742 

Solitary lumps 377 4087 36,102 1652 1024 6091 9313 

Sets of lumps 73 0 32 37 0 42 26 

Individual grooves 1237 717 2137 : 1836 398 10,117 9674 

Sets of grooves 0 0 0 441 341 1176 503 

Solitary depressions 25,391 25,17 42,226 25,737 44,833 26,799 5,241 

Groups of depressions 0 789 510 334 35,218 1638 9180 

Strip of adjacent depressions 0 0 0 0 0 0 0 

Total 35,394 34,810 86,31 34,328 82,157 52,812 85,679 
407 

suggests extensive biological activity probably by a specific type of 

organism . 

Four other categories were recorded at this station: solitary lumps 

with 3569 features/ha; individual grooves with 3377/ha; sets of grooves at 

1036/ha; and solitary depressions represented by 7022/ha (Table 5-3) " Two 

of these groups, individual and sets of grooves indicated motile 

organisms. The most abundant solitary groove type was described as short, 

narrow and straight . These grooves are most probably produced by crab 

legs, either due to the crab walking or possibly resulting from feeding 

behavior. The crab, Lyreidus bairdii was the dominant decapod crab 

obtained in trawls at this station with a weighted density of 11/ha 

(Appendix C2-3) . 

Two tapes of groove sets were recorded and were caused by 

echinoderms, one type was asteriod-shaped impressions representing 230/ha, 

and the other distinctly ophiuroid-shaped impressions with a density of 

384/ha . _ 

Solitary depressions were equally distributed between deep and 

shallow depressions (probably indicating age) with 3031 and 3991 

features/ha, respectively. 

Station W2 

Total lebensspuren density from analyses of 248 .8 m2 at Station W2 

was 105,229/ha. The dominant lebensspuren feature was the category 

solitary depressions with 62,784/ha. These depressions were divided 

primarily between deep depressions (35,090/ha) and shallow, with 

26,810/ha.The deeper depressions likely indicate more recent activity . 

Groups of depressions were present at the relatively high density of 

21,544/ha, a value but significantly lower than observed at Station W1 . 

Solitary lumps were the second most abundant lebensspuren group at W2 

With 11,857 features/ha. The majority of these features (8682/ha) were 

described as excavated sediment lumps adjacent to burrows. Other types of 

solitary lumps included a total of 925 lumps of probable holothurid fecal 

deposits although no holothuroids were captured in the trawl at Station 

W2 . However, the holothuroid Mesothuria lactea was obtained on the other 

two transacts at the same depth (Stations C2 and E2; Appendix C2-3) . 
408 

Remaining lebensspuren categories included : a very low density 

(121/ha) of individual ridges ; individual grooves (5306/ha the dominant 

type being closely spaced and almost certainly decapod tracks), and sets 

of grooves (3618/ha) . A portion of these sets of grooves (804/ha) were 

produced by burrowing organisms which feed on the surface of the sediment 

producing a variety of grooves originating from a central point. 

Station W3 

At Station W3, total lebensspuren density from analyzed frames 

occupied an area of 94.6 m2, which was lower than observed at shallower 

Stations W1 and W2 (6,926 features/ha) . The most numerous feature type 

at W3 was individual grooves at 32,886/ha. Within this category, the 

dominant classification by far (21,889/ha) Was short, narrow, closelyspaced grooves, probably representing decapod tracks. 

Only one other lebensspuren category was represented by a noteworthy. 

density. Solitary depressions were observed at an overall density of 

27,176/ha, the majority of these (21,149/ha) being shallow depressions 

indicating a greater age of the structures compared to a density of only 

5922/ha for deep depressions. 

Four other lebensspuren groups were represented, but at relatively 

low densities. These were individual ridges with only 317/ha observed, 

sets of lumps at 423/ha, probable due to a chance arrangement of the more 

common solitary conical lumps (represented by a density of 1903/ha), and 

groups of depressions arranged in a partial circle observed at a density 

of 2221 / ha . 

Station A 

This station showed an overall increase in total lebensspuren density 

(80,253/ha) compared to Station W3 but had fewer features than Stations W1 

and W2 (Table 5-3) " The total area analyzed for lebensspuren features at 

W4 Was 223.3 m2 . The dominant feature type was again individual grooves 

attributable to decapod tracks (54,816 grooves/ha) . Interestingly, very 

few decapods were captured in the trawl at Station W4, although one Geryon 

guinguedens , a very likely track maker, was taken . An overall weighted 
409 

density of only eight decapods/ha, including both crabs and shrimp, were 

trawled at this station. 

Solitary depressions were represented by 17,018 features/ha, the 

great majority of these being shallow depressions (15,674/ha compared to 

only 1299 deep depressions/ha). Other lebensspuren categories occurred at 

relatively low densities, including individual ridges with 2015 

features/ha, solitary lumps and sets of lumps with 5329 and 179/ha, 

respectively, and sets of grooves with 627/ha. Groups of depressions were 

represented by a density of only 269/ha, much lower than the three 

shallower stations of the Western Transact. 

Station W5 

This station, at a depth of 2456 m, had the lowest overall 

lebensspuren density at 36,771 features/ha. The most abundant 

lebensspuren type recorded from the total area of 286 .9 m2 analyzed Was_ 

solitary depressions at a density of 22,760 depressions/ha. Of these, 

nearly all (21,993/ha) were classified as shallow depressions indicating 

burrowing activity in the more distant past and/or a more rapid inf filling 

of excavated burrows . Only 732 depressions/ha were classified as deep 

depressions which had not been filled in by sediments or smoothed over. 

Other categories of mayor lebensspuren types were represented by very 

low densities, all less than 5000/ha. However, the category of individual 

ridges had the highest density of the Western Transact at Station W5 with 

4914 ridges/ha. These ridges indicated a burrowing organism which moves 

dust below the surface of the sediment leaving an elevated ridge 

projecting above the surrounding bottom . Three other lebensspuren 

categories were recorded including solitary lumps at 5124/ha, individual 

grooves at 2510/ha and sets of grooves with 1u64/ha. 

Western Transact Overview 

Figure 5-1 illustrates overall lebensspuren density for all 15 Cruise 

II stations by depth. The Western Transact exhibits a general decrease in 

total lebensspuren density with depth with the exception of Station W4 

where the density increased to 80,253 features/ha before decreasing to the 
410 

Cruise I I Total lebensspurea Densities 

v 

0 

n 

150000 

e 

s 

t OUOM ., . 

e 

r 

D 

o 50000 

0 

C 

Cruise I I Stations 

Figure 5-l. Total lebensspuren densities for Cruise II stations . 

p 1 2 3 4 5 
411 

lowest of the transects at Station W5 with an overall lebensspuren density 

of 36,771/ha. 

Figure 5-2 presents the overall density of all eight mayor 

lebensspuren categories for each of the Cruise II stations separately, the 

Western Transect being in the first column in order of increasing depth. 

Figure 5-3 graphs each mayor lebensspuren category separately for all 15 

Cruise II stations to illustrate how stations and transects related to 

each other within a specific lebensspuren type. One obvious trend was 

that shown by the dominant feature type on the Western Transect, groups of 

depressions which had a very high density of 167,460/ha at Station W1, but 

had decreased to 0 at Station W5 . 

The second ranked lebensspuren feature category, solitary depressions 

(those not arranged in any perceptible pattern), peaked in density at 

Station W2. The general trend was similar to both the Eastern and Central 

Transects showing a sharp drop in density between Station 3 to 4 and an 

increase from Station 4 to 5 . 

Individual grooves were also a significant contribution to the 

overall lebensspuren density on the Western Transect, having a peak at 

Station Wu. The increase in this particular feature with depth resulted 

in the reversal of the general trend of decreasing total lebensspuren 

density with depth. Figure 5-3 illustrates the peak density of individual 

grooves occurred at Station W4 and this peak represented the highest value 

from all 15 Cruise II stations. 

Although sets of grooves did not represent a significant contribution 

to total lebensspuren density at any station, Station W2 exhibited another 

maximum for all Cruise II stations with a density of 3618/ha. The pattern 

for distribution of groove sets appears to be similar for all three Cruise 

II transacts--a density peak at Station 2, a minimum at Station 3 and a 

rise to a secondary peak at Station 5 . 

Individual ridges and sets of lumps were found at very low densities 

throughout the Western Transact. Solitary lumps exhibited a peak in 

density at Station W2 and a minimum density at Station W3. 

There were no recorded strips of adjacent depressions representing 

holothuroid trails in any of the Western Transact stations; however, there 

were a few larger holothuroids collected at Station W5 in the trawl 

(Appendix C2-3) . The two genera known to produce a visible trail, 
412 

soooo 

e 

r 

v 

4 

40000 

r 

h 20000 e 

r 0 

0 

80000 

60000 

400010 

20000 

0 

eoooo 

40000 

20000 

0 

goo ni 

Yi 167460 

vow in 

Y2 

6 14 m 

Y3 

Y4 

I 

60000 

40000 

20000 

0 

soooa 

600010 

40000 

20000 

0 

2436 m 

347 m 

E2 

E3 

1~ 

E4 

2E 

E5 

A B C D E F G H A B C D E F G H ^ ° `' " ` ' 

A. . . . . . . . . . . Individual Rides E. . . . . . . . . .. Sets of Grooves 

B. . . . . . . . . . . Solitary Lumps F. . . . . . . . . . . Solitary Depressions 

C. . . . . . . . . . . Sets of Lumps 0. . . . . . . . . . . Oroups of Depressions 

D. . . . . . . . . . . Individual Orooves H.. . . . .. . . . . Strip of Adjaoent Depressions 

Figure 5-2 . Major lebensspuren category densities by station, Cruise II . 

L456 m 

VS 

C1 

587 

C2 

14 

C4 
30000 

25000 

e 

r 

n 

20000 

Individual Ridges 

Tronsect 

o West 

Central 

a East 

i 15000 

r 

w 10000 

e Y 

D 

5000 

0 

14 

12 

10 

8 

6 

4 

2 

1 't 3 4 5 

6'00 

700 

600 

500 

400 

300 

xro 

1001 

0 

Sets of Lumps 

Tronsect 

o West 

* Central 

East 

60000 

SOOUO 

40000 

30000 

20000 

10000 

0 

1 2 3 4 5 

individual Grooves 

Tronseci 

o West 

A Central 

s East 

4000 

3000 

nib 

1000 

0 

8000 

60000 

40000 

20000 

Q 

Set of Oreeves 

Tronsect 

u West 

A Central 

a East 

1 2 3 4 5 1 2 3 4 v 

Cruise 2 Stations Cruise 2 Stations 

200000 

Cruise I I Station Depths 

St ti 

West Central . EMS. 150000 

a on 

1 366 358 347 

2 604 587 612 100000 

3 814 798 849 

4 1396 1420 1322 

5 2456 2436 2893 

50000 

0 

1 2 3 4 5 

Groups of Depressions 

1 2 3 4 5 

Cruise 2 Stations 

Figure S-3 . Cruise II station densities for separate major lebensspuren categories . 

w 

1 2 .. 4 5 

Solitary Depressions 
414 

Benthodytes , and Paelopatides were not captured in trawls or recorded in 

benthic photography transects at any Western Transect station. 

5 .1 .1 .2 Cruise II Eastern Transeet 

Station E1 

The five stations of the Eastern Transect, E1-E5 were photographed 

during Cruise II, Station E1 being the shallowest at a depth of 347 m. A 

total area of 304 m2 at E1 was analyzed for lebensspuren features 

resulting in the very high overall lebensspuren density of 166,151/ha, 

surpassed only by Station w1 with 182,464 features/ha, also sampled during 

Cruise II . 

Similar to Station W1 and unlike the remaining Cruise II stations, 

the dominant lebensspuren featured at Station E1 was the category "groups 

of depressions" represented by a density of 128,61u/ha. Sediment grain 

size analyses from both Stations W1 and E1 resulted in substantial 

proportions of sand, the only Cruise II stations with more than 35% sand. 

Three other mayor lebensspuren categories were represented at E1 by a 

density of more than 1000 features/ha. The second ranked category was 

solitary depressions (32,499 features/ha) . This feature was the dominant 

type in most all other Cruise II stations. These depressions appeared to 

be largely deep, recently active burrows (27,461/ha compared to 5005/ha 

for shallow depressions) . Other categories represented included solitary 

lumps and individual grooves, with 3095 and 184 features/ha, 

respectively . Very few (99/ha) tracks of holothuroids (represented as 

strips of adjacent depressions) were observed. 

Station E2 

Station E2 Was at a depth of 612 m and analyzed frames constituted a 

total bottom area of 130 m2 " Sediment grain size was also heavily 

influenced by the sand fraction, similar to Station E1 . Overall 

lebensspuren density was 46,988 features/ha, the lowest on the Eastern 

Transect . 
415 

The dominant lebensspuren category was solitary depressions, the 

greater proportion described as shallow (18,918/ha) as opposed to deep 

depressions (14,535/ha) " The second ranked feature type was solitary 

lumps with a density of 4999/ha. Approximately one-third of these had an 

apical round hole (115u/ha compared to 3076/ha without a hole) suggesting 

an active inhabitant. 

Other features represented at Station E2 made up a much smaller 

proportion of the total lebensspuren abundance. These included individual 

ridges with 3461 features/ha, individual grooves with a density of 1461/ha 

and sets of grooves with 1077/ha. 

Station E3 

Lebensspuren analyses at Station E3, at a mean depth of 849 m, 

resulted in an intermediate overall lebensspuren density of 68,233 

features/ha. A total bottom area of 304 m2 Was analyzed . 

The first ranked lebensspuren category was again solitary depressions 

which had a density very similar to that observed at Station E2 . However, 

this category was closely followed in abundance at E3 by individual 

grooves which had a density of 25,033/ha. The majority of these grooves 

were described as short, narrow, and straight, probably resulting from 

decapod movements across the sediment although decapod density resulting 

from trawling was quite low in comparison to the two previous stations, E1 

and E2. The large crab GerSron was captured here, but at a density of only 

2/ha. 

Other lebensspuren categories well represented at Station E3 included 

groups of depressions with 6669/ha, and solitary lumps with a density of 

3o88/ha . 

Station E4 

Station E4 Was at a mean depth of 1322 m and a total area of 233 m2 

was analyzed for lebensspuren features. Total lebensspuren density was 

51,823 features/ha, dominated by individual grooves which represented 

33,470/ha of the total . Station E4 was the only Cruise II station 

dominated by the category of individual grooves . The majority of the 
416 

grooves (28,667/ha) Were short, narrow, and straight, similar to those 

observed at Station E3 . Many large benthic decapods were well represented 

in trawl samples from Station E4, including the large crab GerSron known to 

leave visible short, narrow grooves in the sediment. 

The second-ranked lebensspuren category was solitary depressions, 

dominated by the shallow variety (13,355/ha from a total density of 

15,199). The only other category with a significant density was solitary 

lumps with only 1639 lumps/ha, most all of those (1065/ha) Without a 

visible, apical, round hole. 

Station E5 

The deepest station of the Eastern Transeet, which had a mean depth 

of 2893 m, was essentially off the continental slope due to the difficulty 

photographing the target depth located directly on the Florida Escarpment. 

Sediment grain size analyses indicated Station E5 had the lowest sand 

proportion and highest clay proportion of any of the Eastern Transeet 

stations. Overall lebensspuren density was a relatively high 74,067 

features/ha, second highest of the Eastern Transect. 

The dominant feature type was solitary depressions, virtually all 

(28,27 of 29,104/ha) being classified as shallow, apparently filled in 

depressions. The second ranked feature category, individual ridges, was 

generally very rare throughout all other Cruise II stations. An unknown 

burrowing animal produced narrow grooves with a random path having a 

density of 18,517/ha . Coincidentlp, a species of isopod, AAea sp ., was 

captured in the trawl at Station E5 at a very high density of 992/ha and 

was found at extremely low densities at only three of the other Cruise II 

stations (Appendix C2-3) . 

Solitary lumps were well represented with a density of 11,702/ha but 

only a small portion of that density (1629/ha) Was represented by conical 

lumps common at other stations. The remainder represented fecal material 

of holothuroids (Appendix D1-3)" One type, with a total density of 

6258/ha, was described as an elongate sculptured lump resembling a coiled 

rope lying on its side. The other similar material (having a density of 

3515/ha) was described as a sculptured elongate lump resembling a pile of 

uncoiled rope. The actual traces of holothuroids which would have been 
417 

classified as strips of adjacent depressions were not observed at this 

station, probably due to the fine texture of the sediment and its failure 

to preserve that type of impression. 

Other lebensspuren categories representing a significant density 

included individual grooves with 4072/ha, and two other categories, sets 

of grooves and groups of depressions, both represented by a density of 

3086 features/ha. 

Eastern Transect Overview 

Figure 5-1 shows total lebensspuren density by stations for each of 

the three Cruise II transacts : West, Central and East. The Eastern 

Transact shows a sharp decrease in lebensspuren density below the 

shallowest Station E1 , similar to the Western Transact but there is no 

consistent trend of decreasing density with depth . Station E2 had the 

lowest overall lebensspuren density of the five Eastern Transact stations. 

The combined lebensspuren density total for all Eastern Transact stations 

was 88,29 features/ha, intermediate to the totals of the Western and 

Central Transacts. 

The eight mayor lebensspuren categories are shown in histograms for 

the five Eastern Transact stations individually in Figure 5-2 . Figure 5-3 

plots densities of each lebensspuren category separately for all five 

stations of each transact in separate graphs. The most numerous category, 

groups of depressions, showed a similar pattern as the Western Transact : a 

very high peak occurred at the shallowest station at a depth of 347 m, and 

relatively low densities occurred throughout the other four stations. The 

second ranked category for the transact was solitary depressions . 

Stations E1-3 and E5 showed similar solitary depression densities, the 

lowest value for this category appearing at Station E4. 

Individual grooves, the third most numerous major lebensspuren 

category on the Eastern Transact showed a similar distribution pattern to 

that previously described for the Western Transact . Individual groove 

densities were low at Stations 1 and 2, rose to a peak at Station u and 

decreased dramatically from Station 4 to 5 . 

Other lebensspuren features were not very abundant on the Eastern 

Transact with the notable exception of individual ridges discussed 
418 

earlier. At Statin E5, a peak density of 22,374 individual ridges/ha is 

shown graphically in Figure 5-3 " This value represented the highest 

density of all 15 Cruise II stations. 

5 .1 .1 .3 Central Transect Stations : Cruises II and III (by depth) 

Station C1 

Station C1, centered around a depth of 335 m, was photographed on 

three different cruises : Cruise I, II and III . Only during Cruises II 

and III, which utilized the LGL BUCS camera system were photographs 

obtained with sufficient information to allow quantitative analyses . A 

total area of 190 .1 m2 was analyzed from Cruise II and 259 .6 m2 from 

Cruise III (Tables 5-2 and 5-4) . 

Overall lebensspurea density was higher from Cruise III compared to 

Cruise II ; 75,964 and 37,717/ha, respectively (Tables 5-3 and 5-5) " The 

dominant category from both cruises was solitary depressions with 

35,555/ha during Cruise III and 21,345 from Cruise II. The solitary 

depression types were nearly equally divided between shallow and deep 

during Cruise II (10,363 and 10,78/ha, Appendix D1-4), but the shallow 

type depressions were more numerous during Cruise III With 20,004 compared 

to 13,945/ha for deep solitary depressions. The midpoint of the two 

transacts were relatively close together (separated by 2.22 km, Table 5-6) 

but even this small separation could have placed the transacts into 

significantly different areas with slightly different sediment 

characteristics with different abilities to preserve or fill in 

depressions. However, the sediment grain size analyses did not show 

significant differences between the two cruises (Attachment Figs. 7-2 and 

7-3) " 
419 

Table 5-6 . Distance between Central Transact station photo transacts 

between Cruises II and III. 

Station Cruise II Depth Cruise III Depth Separation Distance (km) 

C1 335 337 2 .22 

C2 589 668 2 .95 

C3 802 852 2 .72 

C4 1407 1394 4 .54 

C5 233 2407 4 .38 

The second ranked lebensspuren feature at Station C1, individual 

grooves, was surprisingly higher in density by more than two orders of 

magnitude (31,2+1 compared to 210/ha) during Cruise III compared to Cruise 

II. The great majority of individual grooves recorded during Cruise III 

were categorized as short, narrow, closely spaced decapod tracks 

(23,074/ha) . It was curious that more short grooves appearing to be 

decapod tracks were not recorded from Station C1 during Cruise II as there 

were similar numbers of large crabs captured in trawls during both of 

these two cruises at this station (Appendices C2-3 and C2-4) . 

Solitary lump density was virtually the same between the two cruises 

with 6786/ha recorded from Cruise II and 6317/ha on Cruise III. 

Individual ridges appeared in very low densities during both cruises with 

only 53/ha on Cruise II and 231/ha from Cruise III. 

Depressions were dominated by solitary observations (21,305/ha during 

Cruise II and 35,555/ha during Cruise III). Depressions arranged in 

groups (single row in partial circle) Were observed at considerably lower 

densities during both cruises. Cruise II with g363/ha and 2542/ ha during 

Cruise III . 

Station C6 

This station at a depth of 525 m was sampled only during Cruise III . 

A total area of 338 m2 was analyzed with an overall lebensspuren desnity 

of 144,016/ha. Sediment characteristics were very similar to Station C1 
420 

but Station C6 had a drastically reduced density of individual grooves, 

1537/ha as compared to 31,241 at Station C1 during Cruing III. 

Solitary depressions exhibited the highest density at Station C6 with 

101,944/ha. Depressions arranged in groups were the second ranked 

lebensspuren category with 23,1 depressions/ha. Solitary lumps were a 

visually important feature but occurred as the third ranked category at 

15,256/ha, primarily due to their relatively large size. The lebensspuren 

categories representing motile biota, i .e., ridges and grooves showed very 

low densities. 

Station C2 

This station was sampled on the first three cruises with the results 

from Cruises II and III presented in the figures. A total area of 118 m2 

was analyzed from Cruise II and 300 m2 from Cruise III. Again, the 

sediment grain size analyses for this station between the two different 

cruises were quite similar. In the photographs, the sediment's surface 

appeared to be very fine grained and almost flocculent, however, grain 

size analyses showed an increase in the sand fraction as compared to 

neighboring Stations C6 and C3. The photographic transact midpoints of 

Cruises II and III were separated by 2 .95 km at depths of 587 and 629 m, 

respectively. 

The dominant lebensspuren category for both cruises was solitary 

depressions, represented by densities of 45,581/ha during Cruise II and a 

substantially higher 116,111/ha during Cruise III. One taxa, the decapod 

crab Bathyplaz typhlg , frequently seen in benthic photographs, appeared to 

be a mayor contributor to the creation of solitary depressions as well as 

short individual grooves left behind as tracks. In several frames, B . 

typhla was photographed inside depressions with apparent tracks 

surrounding the burrow. This species was also captured in trawls at 

Station C2 during both Cruises II and III in the highest densities found 

at all Central Transact stations, 26 and 30/ha, respectively. 

The second ranked category varied between cruises, during Cruise II 

individual grooves were represented by a density of 12,218/ha and during 

Cruise III the second ranked category was groups of depressions with a 

density of 25,078/ha. One other mayor category, solitary lumps, was 
421 

represented during both cruises in significant density, Cruise III being 

greater than Cruise II. 

Station C3 

This station was also sampled during three cruises with quantitative 

data available from Cruises II and III. The sampling locations were 

separated by a distance of 2 .72 km between these two sampling periods, 

Cruise II at a depth of 798 m and Cruise III at a depth of 866 m. 

Analyzed area from Cruise II included 339 m2, and 257 :m2 was analyzed from 

Cruise III . 

The overall lebensspuren density during Cruise III Was more than 

double that of Cruise II, 15,758 features/ha compared to 7u,926/ha, 

respectively . The bulk of this difference was due to a three-fold 

increase in the density of solitary depressions in Cruise III sample 

frames . Looking at the specific depression types, Cruise III solitary:-

depressions were represented by a much higher density (by a factor of more 

than 3 .7) of shallow depressions as compared to deep . During Cruise II, 

this relationship was reversed with deep, apparently active depressions, 

outnumbering shallow ones, but by only a small margin (35,090 compared to 

26,810/ha) . The remaining mayor lebensspuren types were very similar 

between the two cruises. 

Station C7 

Station C7, at a depth of 950 m was sampled only during Cruise III. 

A total area of 240 m2 was analyzed for lebensspuren features. Overall 

lebensspuren density was 79,822/ha, a significant decrease from the 

previous two stations and very similar to Station C1 . In general, Station 

C7 was very anomalous due to at least two extensive areas of apparent 

chemical influence on the color, texture and epifauna occurring on the 

surface layer of the sediment. Of particular note with regard to 

lebensspuren was the very high density of individual grooves, 36,661/ha, 

the highest of all Central Transeet stations from both cruises, and also 

making this station the only one not dominated by the category of solitary 

depressions. The primary contributor to this higher density of individual 
422 

grooves was a single feature described as rough broad grooves formed in 

this case by the surf icial plowing of living vesicomyid clams . At least 

two species were represented, both of which were related to the bivalves 

in chemoautotrophic communities found in the Gulf of Mexico and at 

hydrothermal vents in the Pacific Ocean . Descriptions of these clam 

aggregations and their significance appeared in the previous Annual Report 

(LGL and Texas ABM 1986). 

One other mayor lebensspuren category, solitary depressions, was 

represented by high densities (31,537/ha) at Station C7 . This category 

was divided relatively equally between shallow and deep depressions, 

17,164 and 14,081/ha, respectively. 

Station CS 

Station C8, sampled only during Cruise III at a depth of 1098 m, 

exhibited an overall lebensspuren density of 16,22 features/ha, the 

lowest of all Cruise III Central Transect stations. A total bottom area 

of 151 m2 Was analyzed from this station. 

The dominant feature was solitary depressions with only 8817 

depressions/ha which also represented the lowest density for this category 

from all Central Transect stations. Other categories represented included 

individual grooves with 3845/ha, individual ridges with 2,320/ha and 

solitary lumps with only 1193/ha. The sediment grain size distribution 

(Attachment Fig. 7-3) Was very similar to that of Station C7 but this was 

the first station which did not have depressions arranged in regular 

patterns. No obvious deletion of invertebrate megafauna taxa occurred, 

compared to previously described shallower stations and Station C8 

(Appendix C2-4) Which would explain the lack of groups of depressions. 

Station C9 

This station was also sampled only during Cruise III and had a mean 

transect depth of 1325 m. Overall lebensspuren density was 35,39+ 

features/ha. The dominant category by far was solitary depressions having 

a density of 25,391/ha. In general, the pattern and types of features 

observed were very similar to those noted at Station C8 . Again, as at 
423 

Station C8, there were no groups of depressions and the other three 

lebensspuren categories observed at Station C9 ; individual ridges, 

solitary lumps, and individual grooves appeared in similar low densities 

as had been observed for Station C8. 

Station C4 

Station C4 was sampled during the first three cruises and processed 

quantitatively using samples from Cruises II and III. Total area analyzed 

included 231 m2 from Cruise II at a mean depth of 1420 m, and 279 m2 from 

Cruise III for samples having a mean depth of 1408 m. The photo transect 

midpoints from the two cruises were separated by a relatively large 

distance of 4 .54 km . Although the sediment grain size analyses were 

generally similar between the two cruises, there was a slight increase in 

the proportion of coarser grained sand in Cruise II samples compared to 

Cruise III (Attachment Figs. q=2 and 7-3) " Overall lebensspuren density. 

was higher during Cruise III with 34,810 features/ha compared to 22,236/ha 

during Cruise II. 

Lebensspuren density by categories during Cruise III was very similar 

to that observed for previous Stations C9 and C8 except for the addition 

of a very low density of groups of depressions at only 789/ha. The 

dominant category during both cruises was solitary depressions, the higher 

density of 25,417/ha found during Cruise III, about half that density seen 

during Cruise II (12,057/ha) . One dominant species having an apparent 

significant effect on lebensspuren features was the holothuroid Penia.gone 

sp . This small sea cucumber (2-3 cm) Was seen in virtually every 

photograph from this station. Although not classified as individual 

lebensspuren features, very small lumps were seen in most all frames which 

appeared to be related to the abundance of these holothuroids. 

Station C10 

Station C10 was sampled only during Cruise III at a mean depth of 

1605 m with a total analyzed area of 314 m2. The overall lebensspuren 

density was in the upper range of Central Transeet stations with 86 ;301 

features/ha. The general appearance of the sediment surface was quite 
424 

unusual. The bottom was highly textured throughout most of the transact, 

in some frames looking like a coarse gravel. The texturing throughout the 

area appeared to be due to strong bottom currents scouring away the finer 

surf icial sediments. In many frames, there were recognizable ripple marks 

in the sediments . Unfortunately, box cores were not obtained from this 

station due to the coarse or compact nature of the sediments . The box 

cores attempted at this station could either not penetrate through the 

coarse sediment or the corers could not close completely when tripped and 

lost most or all the sample on the way to the surface. Presumably the 

sediment grain size analyzes would have resulted in a significantly higher 

sand proportion compared to the other Central Transeet stations. Visually 

at least, Station C10 Was by far the roughest in texture and showed the 

most prominent signs of strong bottom water currents in comparison to any 

other Central Transact station. 

TWO categories dominated the lebensspuren counts, solitary 

_ depressions with a density of 42,226/ha and solitary lumps with 36,102/ha. 

Solitary depressions were the dominant lebensspuren features but the 

second ranked category type, solitary lumps, were also very prominent 

appearing in most every frame . The majority (26,694/ha) were conical 

lumps which exhibited an apical round hole indicating an active 

inhabitant. This density represents the highest of all Central Transact 

stations by more than an order of magnitude. An additional 5900/ha were 

classified as conical lumps without apical round holes, even this figure 

approximately doubles that of any other station. Very few animals were 

captured in the trawl at this station and none of those that were obtained 

suggested a probable animal responsible for creating the high density of 

these lumps. 

TWO other lebensspuren categories were represented by significant 

densities at Station C10; individual ridges with 5294/ha and individual 

grooves with 2137/ha. Both of these densities were comparable to adjacent 

Stations C9 and C11 . 

Station C11 

This station was sampled only during Cruise III at a depth of 2074 m. 

The total area analyzed for lebensspuren features was 273 m2 . Overall 
425 

lebensspuren density was relatively low at 34,328 features/ha, reduced 

from the higher density seen at the next shallower Station C10 but very 

similar to Stations C9 and C4. The sediment grain size distribution at 

Station C11 was somewhat confused, with a variety of sediment types 

represented in the different box core replicates (ranged from clay to 

silty clay to sand-silt-clay mixes) . Visually, the bottom texture 

appeared very coarse in some places, but the coarseness seemed to be on 

the surface of the sediment only. The patchiness of coarse grained 

surficial sediments could have accounted for the variability seen in box 

core grain size analyses. 

The great majority of records were solitary depressions (25,737/ha), 

most of those described as shallow depressions (24,599/ha) compared to 

deep with a density of only 1138/ha. The only other significant mayor 

lebensspuren category was individual ridges represented by a density of 

4296/ha . 

Station C5 

Station C5 was the deepest station sampled during all of the first 

three cruises. The separation of the photographic transact midpoints 

between the two data-producing cruises was 4 .38 km. The mean depth during 

Cruise II was 236 m with an analyzed area of 263(m 2 . During Cruise III 

the mean depth was 201 m from an area of 176 m2 . Sediment grain size 

analyses results varied only slightly between the two cruises with all 

replicates falling within the silty-clay size classification. 

Lebensspuren density for this station varied considerably between 

cruises . Cruise II showed a very high overall density of 128,717 

features/ha and 82,157/ha were recorded from Cruise III. Station C5 Was 

the only one of multiply sampled Central Transact Stations C1-5 Where 

Cruise II overall lebensspuren density surpassed that of Cruise III. 

During both cruises, the two dominant lebensspuren categories were 

solitary depresions (ranking first) and groups of depressions (ranking 

second) . The category of solitary depressions represented a slight 

increase in density compared to neighboring stations but the high 

densities observed for groups of depressions (30,770 and 44,833/ha, 

Cruises II and III, respectively) represented an extreme increase from 
426 

other deep Central Transact stations and were the highest recorded for 

this category for all Central Transact stations from both cruises. 

The third ranked mayor lebensspuren category during Cruise II, 

individual grooves with a density of 2u,982/ha, was virtually absent 

during Cruise III (only 398/ha) . No obvious explanation for this 

discrepancy is evident in invertebrate densities resulting from trawl 

samples taken during the two cruises at this station (Appendicies C2-3 and 

C2-4) . In contrast, species capable of producing grooves (holothuroids, 

ophiuroids, etc .) were actually more abundant during Cruise III than 

Cruise II. 

Station C5 sampled during Cruise II was one of the few stations where 

adjacent strips of depressions were recorded. This type of feature is 

produced by the movement of holothuroids and is one of the few which can 

be attributed to a specific type of animal. 

Station C12 

Station C12 was the deepest station of the Central Transeet with a 

mean depth of 2882 m and was sampled only during Cruise III. A total area 

of 238 m2 was analyzed for lebensspuren resulting in a moderate overall 

lebensspuren density of 52,842 features/ha. The sediment surface appeared 

very fine grained in the photographs but the grain size analyses resulted 

in the highest sand and lowest clay proportion of all Cruise III stations. 

What appeared to be fine grained sediments in the photographs may have 

been limited to a thin layer on the surface of the substrate. 

The dominant lebeasspuren category was solitary depressions with the 

great majority of those classified as shallow (23,229/ha) as opposed to 

deep depressions (3570/ha) . The second ranked category was individual 

grooves with a density of 10,417/ha. This classification was primarily 

divided between two types of grooves both of which were produced by the 

plowing action of an animal moving across the bottom. One classification 

was narrow smooth grooves (4537/ha) and a second classification of broad 

sculptured grooves (4327/ha) produced by a larger animal causing ridges of 

sediment to form paralleling ridges along the length of the groove. The 

latter classification was rarely seen at any of the other shallower 

Central Transeet stations. 
427 

Individual ridges and solitary lumps were also well represented by 

densities of 6679 and 6091 features/ha, respectively . 

Central Transact Overview for Cruises II and III 

Total lebensspuren densities for all 12 Cruise III stations are shown 

in Figure 5-4 and combined Central Transeet stations from both Cruises II 

and III in Figure 5-5 . Both cruises show a similar pattern of density 

with increasing depth although there is some variation between cruises . 

Total densities from Stations C1-C4 were all lower during Cruise II 

compared to Cruise III but Station C5 total density was higher during 

Cruise II. 

Maximum lebensspuren density occurred at Station C2 during both 

cruises. The peak density of 15,980 lebensspuren features/ha occurred 

during Cruise III. Unlike Stations E1 and w1 of the Eastern and Western 

Transacts, the shallowest station of the Central Transact, Station C1 Was 

relatively low in overall lebensspuren density . The minimum total 

lebensspuren density on the Central Transact appeared at Station C8 

(16,22/ ha) at a depth of 1098 m. With increasing depth below Station C8, 

total lebensspuren density oscillated at a moderate level between 35,000-

86,000 features/ha. 

Figure 5-6 presents histograms of all six mayor lebensspuren 

categories for each station separately, arranged in columns in order of 

increasing depth. Density of features by depth from all Cruise III 

stations is shown for each lebensspuren category in separate graphs in 

Figure 5-? " The most numerous feature type, solitary depressions (overall 

density of 54,241/ha), had a pattern of distribution by depth similar to 

that of two other category types, groups of depressions and sets of 

grooves (Fig . 5-7) . The shallowest station, C1 had relatively low 

densities for these three categories, but levels increased to maximums for 

the transact at Stations C2 or C3 (629 and 866 m), and then dropped 

sharply at Station C8 at a depth of 1098 m . The category of groups of 

depressions showed a substantial increase from surrounding stations at the 

deep Station C5 (2401 m) representing the highest density of the transact 

for this category. A sharp increase was also seen for sets of grooves at 

the deeper Station C12 (2882 m) . 
428 

Cruise III Total Lebensspuren Densities 

m 

0 

.~ 

m 

Z 

Z 

r 

C 

m 

0 

O 

O 

H 

200000 

15000a 

I 0000c 

5000( 

Ordered by Depth In Graphe 

Station Depth (in) 

CI 353 

C6 525 

C2 629 

C3 866 

C7 950 

C0 1098 

C9 1325 

U 1408 

C10 1605 

C11 2074 

CS 2401 

ciz zoaz 

Figure 5-4 . Total lebensspuren densities for Cruise III stations . 

( :1 b 1 3 / 0 y 9 1 U 1 1 Z) 1 1 

Central Transect Stations 
429 

m 

0 

v 

m 

2 

O 

Z 

... 

w 

C 

m 

C 

r. 

O 

O 

F180000 

Total Lebensspuren Densities Cruises II and I11 

Cruise III 

Cruise ii 

100000 

0 

1 6 2 3 7 8 9 4 10 11 5 12 

Central Transact Stetlons 

Figure 5-5. Total lebensspuren densities for Cruise II and III stations . 
430 

L 

r 

50000 

O 

40000 

o 

20000 r 

e 10000 

Y 

D 

0 

so00c 

40000 

30dOC 

20004 

1000( 

soooo 

400 

30000 

20400 

10000 

0 

50000 

40000 

30000 

20000 

10000 

0 

J5J m 

C 

1 I 

C6 101944 

11 

C8 

629 m 

116111 

8bb m 

C3 (24654 

13 

C9 

14 

lOUJ III 

CIO 

2074 m 

C11 

L4U1 m 

C5 

012 

A B ' C ' D " E . F . G . H . A B C D ~ E " F " G ' H . A B C D E F G H 

A. . . . . . . . . . . Individual Rides E. . .. .. . . . . . Sets of (waves 

B. . . . . . . . . . . Solitary Lumps F. . . . . . . . . . . Solitary Depressions 

(. . . . . . . . . . .Sets of Lumps 0. . . . .. . . . . . Oroups o1 Depressions 

D. . . . . . . . . . . Individual Grooves H.. . . . . . .. .. Strip of Adjacent Depressions 

C4 

Figure 5-6 . Major lebensspuren category densities by station, Cruise III. 
431 

8000 

Individual Ridges 

e 

r 

6000 

4000 - 

a r 

r 

C 2000 

4 

D 

0 

0 500 1004 1500 2000 2500 3000 

40000 -

30000- 

20000 -

10000 

0 

0 500 1000 1500 2000 2500 3000 

120 

Sets of Lamps 

100- 

so60-

40 -

20 

. 0 

0 500 1000 1500 2000 2500 3000 

Individual Grooves 

40000 

300001 4 

20000 -

10000- 

0 

0 500 1000 1500 2000 2500 3000 

Depth (M) 

2000 

1500 

1000 

500 

0 

Sets of Grooves 

0 500 1000 1500 2000 2500 3000 

140000 -------~ --~--__._.. 

120000 - 

100000-

80000-

60000-

40000 -

20000 - 

0-

0 500 1000 1500 2000 2500 3000 

Groups of Depressions 

30000-

20000 -

10000 -

0 

u A. 

0 500 1000 1500 2000 2500 3000 

Depth (M) 

Central Transact Stations 

Ordered by Depth in Grapla 

Station Depth (m) 

C1 353 

C6 525 

C2 629 

C3 866 

C7 950 

C8 1098 

C9 1325 

C4 1408 

C10 1605 

C11 2074 

C5 2401 

C12 2882 

Figure 5-7 . Cruise III density by depth (station) for separate major 

lebensspuren categories . 
432 

Two mayor categories in Figure 5-7, individual ridges and individual 

grooves, represented movement of animals across or dust below the 

sediment's surface. The second ranked category in overall density, 

individual grooves, peaked at Station C7, primarily due to the numerous 

furrows left behind by vesicomyid clams found at this station (Appendix 

D1-4) . Stations occurring between C8 at 1,098 m and C5 at 2101 m showed 

very low densities of individual grooves . Distribution of individual 

ridges was generally opposite to that of individual grooves . With 

increasing depth, individual groove density increased at Station C8 to 

relatively high levels throughout four of the deeper stations, C9, Cu, C10 

and C11 . Station C5 showed very low densities of both individual grooves 

and ridges. Station C12 on the other hand, showed an increase in both 

lebensspuren types compared to the neighboring Station C5. An increase in 

density from Station C5 to C12 was also evident in two other categories 

shown in Figure 5-7, sets of grooves and solitary lumps. 

The distribution pattern of solitary lumps was similar to the overall 

trend of total lebensspuren density with a peak at Station C2 and low 

densities at Station C1 and mid-depths below 700 m with the distinct 

exception of the very high density peak at Station C10 (1605 m) . The 

unusual nature of bottom sediments and evidence of strong bottom currents 

discussed previously may have played a role in the presence of large 

numbers of solitary lumps at this station. 

5 .2 FAUNA 

A total of 9147 benthic photographs were analyzed for biota 

representing a bottom area of 2,590 m2 Table 5-1) . Sampling effort 

varied between project years; for Cruise II, 100 frames from each station 

were analyzed for biota and for Cruises III, IV, and V, 200 frames were 

analyzed from each station. 

Faunal observations in benthic photographs were identified to the 

lowest possible taxon. As with any visual sampling technique, each 

taxonomic group possessed different attributes which contributed to 

variation in the accuracy of the identification. Assignment of taxonomic 

names was also a partially interpretive process, utilizing not only visual 

taxonomic keys, but also drawing upon other sources of information 
433 

considered reliable for eliminating mayor gaps in identification quality . 

Additional sources of information used for making taxonomic assignments 

included subtle colors (visible only in live specimens), taxonomic lists 

of biota captured in trawls as the same station, and records of species 

ranges within the Gulf of Mexico or neighboring deep sea habitats. 

These information sources were used very conservatively to avoid 

misidentifications ; e .g., ignoring the possibility of a new species and 

forcing a particular observation into a specific previously recorded taxa. 

However, the utilization of additional information beyond taxonomic keys 

for identifications allowed relatively reliable, accurate processing of 

the data with greater resolution that would have been possible without 

these considerations. 

As with all visual census techniques, lumping of unidentifiable taxa 

into taxonomic groups was unavoidable. In many cases, the uncertainties 

of visual identification occurs at the species level (e .g., many of the 

species of the macrourid fish Nezumia are visually indistinguishable) but_ 

there were also many groups which could not be identified visually beyond 

families, orders, and even classes or phylums (e .g ., Porifera and 

Pogonophora) . 

5 .2 .1 NUMERICAL ABUNDANCE 

A total of 20,755 individual animals were recorded from the 55 

stations analyzed quantitatively (Cruises II-Y) . Due to the very low 

abundanees of most taxa, and to allow comparisons between relative 

abundances, all data are presented as numbers of individuals per hectare 

or 10,000 m2 . 

Appendices D2-2 to D2-5 detail all benthic photography biota 

densities for Cruises II-V to the lowest level of identification possible. 

In several of the groups such as Porifera or Ophiuroidea, the majority of 

the observations are listed as unidentified species numbers or letters . 

These taxa are believed to represent distinct species due to unique 

morphological characteristics or colors, ete., but could not be properly 

identified at this time. 

Of the 245 total different biota taxa listed in Appendices D2-2 to 

D2-5, it is believed there are a minimum of 190 different species 
434 

represented. Much of the overlap occurs when higher level taxa are 

assigned to a specific observation due to poor resolution or lack of 

identifiable characteristics. In many cases, the unidentified biota is 

assigned a genus, family or higher level name which may also be 

represented by species level identifications appearing in the same 

station's taxonomic list . 

As a general overview of all megafauna observed in benthic 

photographs, Table 5-7 shows a summary of mayor biota group density over 

all 55 stations and four cruises using total abundance and total area 

surveyed of 2,950 m2. Figure 5-8 illustrates the relative abundance of 

these mayor groups as a histogram. Six of the mayor groups appearing in 

Table 5-7 were deleted in this figure--bacteria, algae, plants, a single 

isopod and two unique groups, bivalves and tube worms which only occurred 

at isolated stations representing probable chemosynthetic communities. 

Although these data represent a wide variety of stations ranging in 

depth from 300 to 3000 m, a general picture of the megafauna observable by_ 

a specific method of benthic photography on the continental slope of the 

Gulf of Mexico is illustrated. It is immediately apparent that very few 

megafaunal groups are represented by high densities. The group of animals 

with the highest density by far is the class Holothuroidea, which 

exhibited an overall density of 4645 individuals/ha. This value is over 

one half of the total density for all megafauna observed is benthie 

photography sampling. 

The second ranked group shown in Table 5-7 is the bivalves which had 

an overall density of 1152/ha. However, this group was represented almost 

entirely at a single Station (C7) which had characteristics suggesting the 

presence of a chemosynthetic community as described in detail in the 

previous Annual Report (LGL and TAMU 1986) . 

Other mayor groups contributing a relatively large proportion of the 

overall density included Alcyonaria (primarily Pennatulacea or sea pens) 

with a density of ?09/ha, decapod crustaceans with 612/ha, Porifera with a 

density of 462/ha and demersal fishes with an overall density of 198/ha. 

Tables 5-8 through 5-11 present total biota densities from Cruises 

II-Y summarized by mayor groups. These mayor groups (also used for 

megafauna and macrofauna data) vary in taxonomic level from classes in the 

case of the echinoderms, to phylums in eases such as Pogonophora or 
435 

Table 5-7 " Total grouped densities, Cruises II-Y. 

Taxon Total Abundance Density (no ./ha) 

Sehizonycetes 102 41 .5 

Phaeophyceae 145 59 .0 

Hydrocharitaceae 55 22 " 

Porifera 1135 461 .6 

Alcyonaria 174 79 .2 

Actiniaria 106 x+3 .1 

Scleractinia 10 4 " 

Polyehaeta 264 107 .E 

Gastropoda 40 16 .3 

Bivalvia 2832 1151 .E 

Scaphopoda 70 28 .5 

Ceph alopoda 8 3 .3 

Isopoda 1 o .4 

Decapoda 1505 612 .0 

Pogonophora/ 

Vestimentifera 223 90 .7 

Asteroidea 40 1 6 . 

Ophiuroidea 425 172 .8 

Echinoidea 22 8 .9 

Holothuroidea 11,423 4645 " u 

Crinoidea 127 51 .6 

Ascidiacea 10 4 " 

Demersal Fish 488 798 .5 

Totals 20,775 848 .6 
o 

n 

c 

Poi fere 

o Alcyonaria 

G rt 

ww 

Actiniarie 

cn a 

m 

H U1 Sclerectinie 

I w 

C rr 

Polycheeta 

°, Gastropods 

X Scaphopoda 

0 

n 

Cephalopods 

0 

b Oecepode 

o A steroidee 

Ophiuroidee 

Echinoidea 

w 

o~ Holothuroidea 

Crinoidea 

Ascidiacea 

Demersal fish 

Density No ./Hectare 

0 0 0 0 

0 0 0 0 
Table 5-8. Deathic .fwnal density (!b ./da) : Cruise II Western . 

Station 

Taxon W1 W2 ,-jW _ W4 -il5 C1 -C2 C3 _ C?I g5_ Et E2 E3 E4 E5 - 

Hydroaharitaceae 49 0 0 0 69 105 0 28 0 0 0 0 0 0 42 

Porifera 774 119 0 0 0 0 U 28 42 0 99 0 0 0 168 

llcyonaria 0 0 211 44 941 0. '~ 0 42 76 0 922 0 81 294 

Actiniaria 290 0 0 0 0 0~ 85 0 42 76 66 76 0 0 0 

3cleraQtinia 0 0 0 0 0 0 0 0 0 0 0 0 0 40 0 

Poiychaeta 0 39 0 403 69 53 1372 375 510 0 0 0 65 0 168 

Iia3tropoda 0 0 0 24,765 14,77 0 0 0 0 0 0 0 0 0 301,80 

3oaphopoda 0 0 0 0 243 0 0 0 127 0 0 0 0 0 07 

Cephalopoda 0 0 0 0 0 0 D 28 0 0 0 0 0 0 0 

Decapoda 920 199 317 582 34 1841 943 144 85 0 99 615 32 122 294 

Pogonophora/ 

Yestimentifera 0 279 0 0 0 0 0 0 85 0 0 0 0 0 0 

Astaroidea 0 0 0 0 0 53 85 28 0 0 0 0 32 0 0 

Ophiuroidea ;48 239 0 134 0 0 5662 57 0 38 0 307 32 0 42 

Eohinoidea 0 0 0 0 34 0' 0 0 85 152 0 0 0 40 0 

Holothuroidea 0 199 0 44 69 0 943 722 6215 76 0 76 0 0 168 

Ascidiacea 0 0 0 0 0 0 171 0 42 0 0 0 0 0 0 

Demersal Fish 290 239 0 0 0 684 171 202 0 0 99 230 197 50 0 

UnIcnown 339 15,553 26,012 16,122 1847 316 25,224 3381 127 0 99 0 0 40 0 

Total Biota (less 

pteropods) 2371 1313 528 1207 1459 2631 932 1584 7275 418 363 2226 358 323 1176 

w 

V 
4-1 

w 

00 

Table 5-9 " Henthic faunas density (No ./ha) : Cruise III-Central . 

Tazon 

Sohizomycetea 

Phaeopt~yceae 

flpdrooharitaceae 

Porifera 

Aloyonaria 

Aotiniaria 

3cleractinia 

Polqahaeta 

Gastropods 

Bivalvia 

Soaphopoda 

Cephalopods 

Dsoapoda 

Pogonophora/ 

Yeatimentifera 

kateroidea 

Ophiuroidea 

Sehinoidea 

Holothuroidea 

Crinoidea 

Deaerasl Fish 

Mtifacta 

Consolidated Material 

Dnirnoxn 

Station 

C1 C6 C2 C3 ~C' _ CS C9 C4 C10 C11 C5 C12 

0 0 110 0 366 0 0 0 0 0 0 0 

19 15 0 396 18 232 186 53 175 145 33 49 

0 0 0 0 0 21 37 176 70 18 0 33 

97 265 94 20 0 739 149 16,514 0 0 0 

0 78 2612 20 952 147 186 212 788 802 183 249 

38 265 0 0 0 21 18 17 35 127 50 33 

0 0 15 0 0 0 0 0 0 0 0 0 

0 0 15 229 73 42 37 459 280 0 0 16 

174 0 8450 62,411 641 0 79,585 0 65,550 83,862 26,755 96,281 

0 0 0 41 51,566 42 37 0 227 0 0 0 

0 0 0 0 0 42 93 35 175 200 0 116 

0 0 0 0 0 0 0 0 0 0 0 16 

194 3066 330 626 641 1035 579 212 350 255 83 99 

0 0 94 0 91 0 0 17 0 0 0 0 

0 15 0 104 0 42 0 70 17 91 16 33 

0 78 424 208 238 126 130 0 0 0 50 16 

38 0 0 0 18 21 56 0 0 0 0 0 

0 15 125 146 2877 4649 151,800 40,212 4014 127 250 16 

0 0 31 41 0 21 0 17 0 0 0 0 

232 203 78 271 311 338 18 17 52 36 50 16 

0 0 0 0 91 0 18 0 0 0 0 0 

38 375 62 1253 439 1162 5173 3091 5627 5087 0 713 

58 0 15 104 421 21 0 106 140 72 0 16 

Total Biota (less 

pteropods) 792 4000 3928 2102 57 .370 7518 156,344 41,638 22,697 1801 765 692 
Table 5-10 . Henthia faunal density (No ./ha) : Cruise IT Eastern . 

Station 

Taxon El B El A E1 El C E2A E2 E2E E2B E2D E2C E3 EN E3D _ E3A E 3B 

Phaeophyceae 0 27 0 23 0 4 22 46 0 0 0 34 0 23 0 38 

Hydrocharitaceae 0 27 0 207 0 0 45 0 0 0 46 17 20 47 0 248 

Porifera 0 0 0 161 52 159 0 210 56 80 23 0 41 0 0 1T2 

Illcyonaria 22 0 0 369 3778 1625 30.95 2696 2029 4342 836 331 662 431 701 1129 

Aotiniaria 111 55 0 23 26 79 68 0 42 20 0 0 20 0 0 0 

Seleraetinia 0 0 0 0 0 0 0 0 0 0 0 34 0 0 0 0 

Polychaeta 0 0 0 184 52 106 91 281 42 383 278 52 372 47 160 153 

Gastropoda 0 . 0 0 0 26 0 0 0 0 0 0 0 20 23 20 0 

3oaphopoda 0 0 0 0 0 26 0 0 0 0 0 0 0 0 0 153 

Cephalopods 0 0 0 0 0 0 0 0 56 0 116 0 0 0 20 0 

Iaopoda 0 0 0 0 0 0 22 0 0 0 0 0 0 0 0 0 

Decapoda 66 0 74 992 1433 2478 2246 1453 1000 256 580 488 455 575 481 114 

Pogonophora/ 

Vestimentifera 0 0 0 0 0 26 22 0 0 0 0 0 0 0 0 0 

Aateroidea 22 0 0 23 52 0 22 0 14 0 23 0 0 0 0 0 

Ophiuroidea 0 0 0 0 104 186 183 164 84 100 0 34 0 0 0 172 

Bohinoidea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 19 

Holothuroidea 0 0 0 0 26 26 22 0 42 40 23 0 0 0 0 95 

Crinoidea 0 0 0 0 156 26 710 726 84 80 0 0 0 0 0 0 

Aacidiacea 0 27 0 69 0 0 0 0 0 0 0 0 0 0 0 0 

Demersal Fish 334 444 519 738 234 106 275 140 507 383 255 209 124 119 200 0 

Unknown 0 0 0 0 0 186 0 0 0 0 0 0 0 23 0 19 

Total Biota 555 580 593 2789 5939 4843 6823 5116 3956 7992 2110 1199 1714 1265 1582 2293 

w 
41 0 

Table 5-11 . Benthie fauna density (No ./ha) : Cruise Y West-Central . 

Station 

Taxon WC5 WC1 W-jC _ WC8 WC4 WC2 WC6 WC3 _ WC9 W10 W12 W11 

Schizomycetes 0 0 996 0 0 0 211 0 0 0 0 0 

Phaeophyceae 36 21 33 54 71 34 0 66 106 15 267 483 

Porifera 0 0 0 72 232 213 39 0 15 60 223 60 

Alcyonaria 54 216 50 290 160 417 2264 243 196 165 193 1 

Aetiniaria 0 0 84 200 89 208 105 0 45 15 0 0 

Seleractinia 0 0 33 18 0 0 39 0 0 0 0 0 

Polychaeta 0 108 0 0 0 34 13 132 60 195 0 161 

Gastropoda 91 64 0 0 0 0 0 22 15 45 14 0 

Scaphopoda 109 0 0 0 0 0 0 0 0 15 104 0 

Decapoda 127 454 135 72 357 974 370 819 560 540 506 926 

Pogonophora/ 

Vestimentifera 0 0 3310 0 0 0 52 0 0 0 0 0 

Asteroidea 36 0 16 0 0 34 0 44 0 15 0 20 

Ophiuroidea 18 21 84 181 214 383 119 155 560 210 253 1737 

Eehinoidea 109 0 0 0 0 0 0 0 0 0 0 0 

Holothuroidea 0 0 0 0 0 0 13 0 0 15 14 0 

Crinoidea 0 0 0 0 0 0 463 132 15 0 0 0 

Aseidiacea 0 0 0 54 0 0 0 0 0 0 0 0 

Demersal Fish 438 216 219 236 428 87 198 88 181 195 89 483 

Unknown 0 0 0 0 0 0 13 0 0 0 0 40 

Total 1017 1100 4960 1177 1551 241 3886 1701 1753 185 1663 4004 
441 

Porifera. Total biota densities at the bottom of each column excludes the 

unknown category and non-biota categories of artifacts and consolidated 

material. Very high densities of what were believed to be fragments of 

pteropod shells were recorded at many stations. This record type was also 

subtracted from the total because of the probability that none of these 

records represented living organisms . Figure 5-9 illustrates these total 

biota densities in graphs of the four cruises including all 55 stations . 

5 .2 .2 ABUNDANCE BY DEPTH 

The following table summarizes densities for all transects together, 

by depth increments as used in other sections. 

Depth Mean Density (no ./ha) 

278- 500 1,575 

500-1000 5,278 

1000-2000 26,963 

>2000 1229 

Faunal density was generally low at shallow stations and was lowest 

in water deeper than 2000 m. The depth range of 1000-2000 m showed the 

highest mean density by more than a factor of five compared to the neat 

lower depth increment. This high density was due to the very high numbers 

of the holothuroid Peniagone sp. encountered at four mid-depth stations 

sampled during Cruise III; Stations C8, C9, C4, and C10 . When the density 

contribution of this solitary taza is removed from the total for the 1000-

2000 m depth increment, the overall density is reduced to 4365/ha. The 

contribution of this particular genus is also seen in comparison of 

densities by transact. Peniagone sp. was not collected in the trawls. 
442 

toooo 

6000 

4000 

i 

2000 

0 

Cruise II Total Biota Density 

{* was 

~ Central 

f East 

1 2 3 4 5 

STATION 

160000 

rq 

120000 

80000 rp 

40000 

H 

C1 6 2 3 7 8 9 4 10 11 5 12 

ST"TIAY 

1000 

600 

400 

r 

F ZOO 

Lruise m i ocai Biota venst 

0 

boon 

5000 

m 3000 

2000 

1000 

iMC5 1 7 8 4 2 6 3 9 10 12 11 

STATION 

Figure 5-9 . Total biota density from benthic photography by station, 

Cruises II-V . 

STATION 

Cruise V Total Biota Density 
443 

5 .2 .3 ABUNDANCE BY TRANSECT 

The following table lists mean total densities of all fauna by 

transect : 

Cruise Transact 

2 Western 

Central 

Eastern 

3 Central 

Eastern 

5 West-Central 

Mean Density (no ./ha) 

1,365 

4,268 

889 

24,970 

3,122 

2,226 

Over all mean densities on the Central Transacts were consistently 

higher than at other transacts. Again, the holothuroid PeniaAOne sp. was 

the most significant contribution to these density values. During Cruise 

2, this taxa was encountered at two stations, C3 and C4 . At Station C4, 

the contribution of Peniagone sp. density to the total biota density for 

the entire station was a very significant 85%,as illustrated in Figure 5-

9. The high station density at Station C2 was due primarily to an 

unidentified ophiuroid tasa represented by a relatively high density of 

5662/ha. The very high transact density of 24,970 individuals/ha on the 

Central Transact during Cruise 3 Was due almost entirely from the 

extremely high abundance of the Peniagone sp. holothuroid. Fine stations, 

C7, C8, C9, C4, and C10 exhibited high densities of PeniaAOne sp . 

resulting in the density peaks shown in Figure 5-9" 

The Eastern and West-Central Transacts sampled during Cruises IV and 

V were moderate in overall biota density. The Eastern Transact from 

Cruise IV, with a mean density of 3122 individuals/ha, showed higher 

densities through the six mid-depth stations between Stations EZA and E2C 

representing depths between 624-667 m illustrated in order of depth in 

Figure 5-9" Two mayor taxa contributed to these higher biota densities ; 

several pennatulaeca taxa and a number of decapod crustacean taxa 

(Appendix D2-4). Both the shallower stations in the 33a-360 m depth range 
444 

and the deeper stations between 809 and 2,890 m were all lower in mean 

biota density . 

The West-Central Transeet during Cruise V had mean biota density of 

2,226 individuals/ha. Station WC7, representing the highest density of 

the transect is unique. A single taxa group, Pogonophora/Vestimentifera, 

contributed a density of 3309/ha to the total for the station total of 

X960/ha . The bulk of the records (2905/ha) making up the total 

Pogonophora/Vest imentifera density were recorded as vestimentiferans 

considered to be chemosynthetic (LGL and TAMU 1986) . 

5 .2 .4 DENSITY DIFFERENCES WITHIN TRANSECTS 

TWO seasonal comparisons were possible in benthic photography 

results. The Central Transact was sampled both during Cruise II in March 

1984 , and during Cruise III in November 1984. Five of the 12 total 

Central Transact stations were sampled during both cruises; Stations C1, 

C2, C3, C4, and C5 . Several stations on the Eastern Transact were also 

sampled on two separate cruises. Stations E1, E2, E3, and E5 were sampled 

both during Cruise II in March 1984 and also during Cruise IV in August of 

19$5 " Mean densities for these stations sampled on multiple occasions are 

shown in Table 5-12. 

On the Central Transeet, mean density in spring 198u :(Cruise III) was 

more than double that of winter 198 (Cruise II) (9845/ha and 4268/ha, 

respectively) . A similar decrease in total density from Cruise II 

sampling was found on the Eastern Transept stations shared between Cruises 

II and IV . The mean total biota density of the four replicted Eastern 

Transeet stations was more than double in summer 1985 samples (Cruise IV) 

compared to winter 1984 (Cruise II) (21160/ha and 1031/ha, respectively) . 

The variation in sampling effort between Cruise II (100 frames/station) 

and both Cruises III and IV (200 frames/station) may be the cause of these 

transact variations . 

5 .2 .5 SEASONAL DIFFERENCES IN NUMBER OF TAXA 

Numbers of recognizable taxa in benthic photographs were quite low in 

comparison to other sampling techniques. Use of these data for 
445 

Table 5-12 . Total biota densities for all stations sampled on multiple 

cruises . 

Station Cruise II Cruise III Station Cruise II Cruise IV 

C1 2631 792 E1 363 593 

C2 932 3928 E2 2226 483 

C3 158 2102 E3 358 2110 

C4 7275 41,638 E5 1176 2293 

C5 418 765 

Mean Density 4268 9845 1031 246 

Table 5-13 . Numbers of unique taxa for all stations and transect totals, -

Cruises II-V. 

Cruise II 

Station No . Taxa 

W1 14 

W2 11 

w3 3 

W4 6 

w5 a 

Total 35 

C1 14 

C2 12 

C3 15 

C4 12 

C5 5 

Total 41 

E1 5 

E2 10 

E3 8 

E4 5 

E5 9 

Total 29 

Cruise III Cruise IV Cruise V 

Station No . Taxa Station No . Taxa Station No . Taxa 

C1 14 E1B 11 WC5 11 

C6 22 E1A 11 WC1 19 

C2 20 E1 6 WC7 23 

C3 22 E1C 17 WC8 22 

C7 28 E2A 30 WC4 26 

C8 37 E2 22 EC2 21 

C9 23 E2E 31 WC6 34 

C4 24 E2B 29 WC3 20 

C10 20 E2D 32 WC9 28 

C11 15 E2C 32 WC10 33 

C5 15 E3 19 WC12 16 

C12 20 EN 21 WC11 22 

Total 123 EP 21 Total 118 

E3 A 1 4 

E3 B 18 

E5 26 

Total 123 
446 

calculation of diversity indicies would be of little value due to the wide 

variety of species groups that represented more than a single species . 

Total numbers of taxa recorded in benthie photographs by cruise and 

station are shown in Table 5-13. 

The most noticeable contrast is evident in comparison between numbers 

of taxa recorded from Cruise II (Marsh) compared to the other three 

cruises . Numbers of separate taxa on the three transects of Cruise II 

ranged between 29 on the Eastern Transact to 41 on the Central Transact. 

Numbers of taxa represented on the other three cruises were approximately 

three times as high, ranging between 118 on Cruise V and 123 taxa from 

both Cruises III and IV. The most likely explanation for these 

differences is the sampling intensity--the number of frames analyzed for 

biota during Cruises III, IV, and V was double that of Cruise II . There 

may also be seasonal or other temporal effects involved but this possible 

cause of variation cannot be separated from possible effects of variable 

sampling intensity. The effects of unequal sampling effort upon numbers 

of taua was also evident in the macrofauna data 

The number of different taxa recorded in benthic photographs were 

averaged by depth increments over all transacts and all four cruises 

resulting in the following distribution : 

Depth Range No . of Taua 

278- 500 15 

500-1000 26 

1000-2000 18 

>2000 1 4 

Total number of taxa by depth increment matched the pattern shown by 

maerofauna with the highest number occurring in the intermediate depths 

and lower numbers at the shallower and deeper stations . 

5 .2 .6 SIZE MEASUREMENTS FROM BENTHIC PHOTOGRAPHS 

By the use of digitizing procedures described in Section 2 .3 .1 .u, 

objects appearing in benthic photographs and located on a level bottom 

surface could be measured for length or surface area. A total of 16,049 
447 

length measurements (Appendix D3-2) and 328 area measurements (Appendix 

D3-3) were obtained. Table 5-9 shows several representative examples of 

length measurements. Appendix D3-1 summarizes biota length measurements 

from all 55 stations for each taxa. 

The principal utility of this technique is most evident when taxa are 

measured that are rare or non-existent in trawl samples. The ease of the 

extremely abundant holothuroid, Peniagone sp. is an exceptional example. 

This was the most abundant taxon in all 55 stations sampled by benthic 

photography with a peak density of 154,669 individuals/ha at Station C9 

during Cruise III, but was not recorded from a single trawl sample . A 

total of 11,326 were recorded and 10,581 of these were measured (Table 5-

14) . 

5 .2 .7 SELECTED SPECIES ACCOUNTS 

Several taxa were represented in sufficient numbers and are selected 

for further description in this Annual Report. The holothuroid Peniagone 

sp. (identified from photographs by Dr. R. Carney) Was the most numerous 

organism recorded in all of the 9147 analyzed photographs. It was 

observed at six different stations on the Central Transect during Cruises 

II and III over a relatively narrow depth range, as illustrated in Figure 

5-10. The very high peak density of 154,669 individuals/ha occurred at 

Station C9 at a depth of 1325 m. The very narrow depth and station 

distribution of this species and their close proximity to chemosynthetic 

communities in the same area suggests a possible association. 

The length distribution of all observed Peniagone sp . ranged from 

1 .0-7 " 4 em with a mean of 2 .96 cm (Appendix D3-1) . Figure 5-10 shows the 

length frequency distribution for the 10,581 individuals that were 

measured . 

Bembrops gobioides was one of the more numerous fishes recorded by 

benthie photography. A total of 47 individuals occurred at 12 stations 

(Appendix D3-1) . Although it was relatively low in abundance compared to 

the total of 367 specimens collected by trawling, the total density was 

significantly higher than that estimated from trawling due to the smaller 

area photographed . 
00 

Table 5-14 . Sample of length measurements from benthic photography . 

Taxa 

Thalassia sp . 

Hyalinoeeia tubieola 

Gly,phoerangon sp . 

valida 

Natantia 

Ophiuroidea sp . 6 

Peniagone sp . 

Svnanhobranehus sp . 

Bembrons gobioides 

ChloroDhthalmus aga4gizi 

Abundance 

Number of 

Stations 

Number 

Measured 

49 17 49 

69 12 68 

51 21 46 

42 6 16 

126$ 47 810 

160 6 159 

11,36 9 10,581 

32 14 16 

47 12 45 

49 10 47 

Mean 

Length ( em) 

Minimum 

Length ( em) 

Maxim 

Length ( em) 

13 .9 4 .9 29 .3 

20 .7 3 .9 51 .7 

3 .1 0 .9 5 .3 

3 .4 1 .8 4 .5 

2 .7 0 .5 20 .0 

6 .3 3 .6 17 .4 

3 .0 1 .0 7 .4 

45 .1 12 .0 60 .6 

20 .2 6 .6 34 " 3 

10 .5 4 .3 18 .4 
449 

taoooo 

m 

140000 0 

v tZOOOo m 

~ 00000 

Z 0 80000 

60000 .r 

o qpppp C 

m 

G 

0 

3000 

2900 

v 2000 

e 

m 

Q 

0 1500 L 

1000 

500 

0 

PEN IA60ME SP. Crui~ 11 end 11.1 by Depth 

0 300 1000 1300 2000 

Depth (m) 

23Q0 3000 

.25 .75 125 1 .75 225 2.73 325 3.75 4.23 4.75 5.23 5.75 625 6.75 725 7.73 

Length (cm) 

Figure 5-10 . Distribution by depth and length frequency distribution of 

neniagone sp ., Cruises II and III. 
450 

The distribution by depth and length frequency distribution for B . 

Aobioides are shown in Figure 5-11 . This fish occurred only at shallow 

stations between 274 and 579 m " Length measurements ranged from 6 .6 to 

34 " 3 am with a mean of 20 .2 em . The mean length found from benthic 

photography sampling was slightly higher than that determined from trawl 

samples (202 mm and 190 mm, respectively) which is probably caused by 

measurement of total lengths in benthic photographs as opposed to fork 

length in trawl specimens. There appears to be at least two growth 

cohorts shown in Figure 5-11, although the abundance of the smaller 

individuals producing the first mode was very small. Interestingly, 

additional size cohorts were also seen in the trawl collections. 

Chloropthalmus agassizi was another fish well represented in benthic 

photography samples. A total of 49 individuals were recorded from 10 

different stations. As shown in Figure 5-12, this fish was also 

represented only at shallow stations of the continental slope between 355 

acid 766 m depths . 

A total of 47 individuals were measured with a mean length of 10.5 am 

ranging from a minimum of 4 .3 em to a maximum of 18.3 am (Appendix D3-1) . 

Figure 5-12 presents the length frequency distribution for these total 

length measurements. Similar to the case of B. gobioides above, this 

species appears to be represented by several growth cohorts which were 

also evident in trawl collections. The mean length for trawl samples was 

9.5 am compared to 10 .5 em for benthic photography samples, the difference 

due at least in part to the measurement of total length in photographs as 

opposed to fork length of trawl specimens. 

A total of 37 sYnaphobranchus sp. were observed from 12 stations from 

all cruise samples . All of these observations were most probably S . 

oreROni and were inadvertently divided between the species classification 

and the unknown species listing in the appendices . The mean length for 

these observations was 45 .1 am ranging between 24 .5 and 60 .6 cm. In the 

case of this taxa, benthic photography measurements were also larger than 

the trawl mean of 41 .5 em, but this difference cannot be attributed to the 

measurement of fork lengths for this eel species . 

Distribution by depth is presented in Figure 5-13, Synanhobranchus 

sp. Were seen primarily throughout mid-depths between 604 and 1246'.m. 
451 

eEneROps soerotoES crV#m 11-r bg Depth 

goo 

e 

v 

s 

"' too r 

e 

o 

to 

a 

C 6 

m 

O 

O 

4 

o soo 1000 1soo 

Depth (M) 

2000 2500 3000 

e 

69 82 10.3 12.3 142 16A 17.8 196 214 232 25.0 26 .8 28 .7 305 323 34.1 

Length (cm) 

Figure 5-11 . Distribution by depth and length frequency distribution of 

Bembrops gobioides, Cruises II-V . 
452 

goo 

CHLOROPTNALMUS A6ASS121 Croios II-r 6V Depth 

e 

0 

v 0 

z 

o 

s 

goo 

c 

m 0 0 

6 

v 

c 

m 

2 

0 

Figure 5-12 . 

o soo 1000 1300 

Depth (m) 

zooo 250o aWo 

4 v 

4.5 5.4 6.3 7.3 82 92 10 .1 11A 12A 12.9 13 .8 14 .B 15 .7 16 .7 17b 18.5 

Length (cm) 

Distribution by depth and length frequency distribution of 

Chloropthalmus agassizi , Cruises II-V . 
453 

t2o 

o 100-

.r 

eo 

o 

s 

40-

a 

C 

O 

2D -

0 

SYIIAPN08RAUC1111S SP_ CMIws 11- r OV both 

0 

s 

6 

a 

v 

e 

e 

Q 4 

0 

L 

2 

0 

300 1000 100 2Q00 2300 3000 

Depth (m) 

13.3 16,5 19 .7 22.7 23.8 289 31 .9 33A 38A 41 .1 442 472 50.3 333 56.4 593 

Length (cm) 

Figure 5-13 . Distribution by depth and length frequency distribution of 

Synaphobranchus sp ., Cruises II-V . 
454 

The density of the galatheid crab Munida valida is illustrated in 

Figure 5-14. This galatheid was only recorded during Cruise IV and was 

represented only at the mid-depth stations between 629 and 662 m. Only 16 

measurements were possible from the 42 total observations resulting in a 

mean carapace length of 3 .4 em ranging from a minimum of 1 .8 em to a 

maximum of 4 .5 cm. 

5 .2 .8 COMPARISONS OF BENTHIC PHOTOGRAPHY AND TRAWL DENSITIES 

Tables 5-15 through 5-17 present overall megaf auna group densities 

derived from trawling and benthic photography samples from Cruises III, 

IV, and Y, respectively. Cruise II density data were not included in 

these comparisons because of the fewer number of analyzed frames (100 

compared to 200 for the subsequent three cruises) . Calculation of bottom 

areas sampled by trawling (necessary for density calculations) took into 

account the expected mouth opening of the trawl when fishing normally, and 

length of the trawled area derived from the time the trawl spent on the 

bottom and the logged speed of the vessel. 

Figures 5-15, 5-16 and 5-17 present these data in graphs for selected 

megafauna groups (decapod crustaceans, combined asteroids and ophiuroids, 

and demersal fishes), and total megafauna (all groups combined) . The 

mayor groups were chosen for comparison because of the similar likelihood 

that both trawling and benthic photography would obtain representative 

samples. Some other groups would be expected to be sampled more 

efficiently by benthic photography, e .g., sea pens (Alcyonaria) or other 

biota groups attached to, or buried below the surface of the substrate. 

Yet other groups would be expected to be sampled more efficiently by 

trawling in comparison to a visual technique. These groups would include 

small individuals such as barnacles, solitary corals and other taxa not 

easily observed in photographs. 

The overall results indicated that with very few exceptions, 

megafauna densities obtained from benthic photography greatly exceeded 

those obtained from trawling. During Cruise III, the density differences 

between the two sampling methodologies was the most pronounced with 

variations in some cases being more than four orders of magnitude (Station 

C9, Table 5-15). The dramatic differences between benthie photography and 
455 

1000 m 

1500 m 

2000 m 

%% 2500 m 

SUU m 100 m M1IN1DA VALIDA 

a E 1 A 0 0.00 to 35.50 

El 35.50 to 71 .00 

0 71 .00 to 106.50 

106.50 to 142.00 

142.00 to 177.50 

E2A 

~ 177.50 to 213.00 

Ea QE 1 

E2B 

EX a E1B 

E3 E~E2C aE1C 

E3B . E20 

ES 

E2E 

3000 m 

Figure 5-14 . Station distribution of density for Munida valida , Cruise IV . 
456 

Table 5-15 . Total megafauna density (no ./hectare) from trawling and 

benthic photography : Cruise III. 

Tuce C1 C6 C2 C3 C7 ce C9 C4 CIO C111 C5 C12 

PORIFERA Trawl 0 0 0 1 1 9 3 2 0 1 6 

Photography 9E 264 94 20 0 739 147 140 16,514 0 45 0 

ALCYUNAkIA Trawl U 0 G G 0 0 0 0 0 1 14 x 

Photography 0 78 2611 20 951 147 186 212 7E8 802 1S< <47 

ACTINIARIA Trawl d 1 0 3 4 0 0 0 2 15 5 

Photography 38 265 0 0 0 21 18 17 35 127 50 .,. 

PECAPOUA Trawl 91 189 100 34 52 115 0 16 2 7 10 

Photography 192 3065 328 625 640 1034 577 210 350 254 83 98 

BIVALVIA Trawl 0 1 8 0 1 9 2 10 0 0 1 

Photography 0 0 0 41 51,565 42 37 0 227 0 0 0 

OPHIUROIDEA Trawl 1 8 192 3 1 4 0 2 0 5 30 

Photography 0 77 423 208 237 126 129 0 0 0 49 16 

ECHiN01DEA Trawl 0 0 0 0 0 2 0 1 1 fj 0 

Photography 38 0 0 0 1£s 21 56 0 0 0 0 0 

ASTERCtIDEA Trawl 0 2 " 2 25 7 9 6 18 3 16 12 

Photography 0 15 0 103 0 42 0 70 17 90 16 32 

HOLOTHJROIDEA Trawl 0 19 0 14 10 2 0 2 0 10 34 

Photography 0 15 125 145 2877 4649 151,799 10 .211 4014 127 249 16 

CkIN01DEA Trawl 0 0 0 0 0 1 0 G 0 U U 

Photography 0 0 31 41 0 21 0 17 0 0 0 0 

OTHER INVERTEBRATE Trawl 4 3 2 282 123 12 1 6 55 7 8 

Photography 174 0 124 229 382 84 148 511 454 200 0 148 

DEh1ERSAl FISH Trawl 54 119 35 44 34 50 0 4 5 4 4 

Photography 231 200 76 267 307 336 18 17 52 36 50 16 

TOTALS Trawl 154 342 339 406 243 213 12 61 68 66 124 

Photography 769 3479 3812 1699 56,977 7262 156,115 41,405 22,415 1636 728 606 

* Station C I2 Not Trawled 
Table 5-16 . Total megafauna density (no ./hectare) from trawling and benthic photography : Cruise IV . 

Tan E19 EtA E1 E1C ETA E2 EZE E28 E211) EZC E8 E8C E3D E8A E38 E3 

PORFERA Trawl 1 2 4 2 10 14 5 21 16 10 2 2 1 2 4 + 

Photography 0 0 0 161 52 159 0 210 56 80 23 0 40 0 0 171 

ALCYONARIA Trawl 0 5 2 0 59 101 13 66 193 51 9 8 3 3 11 

Photography 22 0 0 369 3777 1823 3091 2691 2027 4339 835 329 661 429 701 1128 

ACTNIARIA Trawl 1 7 2 20 0 0 0 0 7 0 0 1 0 0 0 

Photography 111 54 0 23 26 79 67 0 42 20 0 0 20 0 0 0 

DECAPODA Trawl 152 167 294 329 329 309 284 272 638 384 47 88 63 101 56 

Photography 66 0 74 992 1432 2478 2243 1451 999 2582 580 486 453 574 481 114 

BIVALVIA Trawl 4 0 1 4 8 0 2 26 18 3 6 9 12 10 2 

Photography 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

OPHIUROIDEA Trawl 6 4 4 4 21 35 29 41 66 40 2 4 7 6 3 

Photography 0 0 0 0 104 185 182 162 84 100 0 34 0 0 0 171 

ECHINOIDEA Trawl 1 0 0 0 1 6 3 1 4 5 0 0 0 0 

Photography 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 19 

ASTEROIDEA Trawl 1 7 0 1 41 20 10 28 16 12 5 10 12 22 2 

Photography 22 0 0 23 32 0 22 0 14 0 23 0 0 0 0 0 

HOLOTHUROIDEA Trawl 0 0 0 2 4 0 1 0 2 1 5 8 11 11 5 

Photography 0 0 0 0 26 26 22 0 42 40 23 0 0 0 0 95 

CRINOIDEA Trawl 0 1 1 0 7 18 7 22 22 14 3 1 1 0 2 +~ 

Photography 0 0 0 0 156 26 710 725 84 80 0 0 0 0 0 0 

OTHER INVERTEBRA TE Trawl 19 13 34 28 28 27 18 39 63 28 8 15 15 10 15 

Photography 0 27 0 253 78 158 135 280 98 382 324 85 391 69 200 305 

DEMERSALFISH Trawl 52 126 191 88 83 42 21 44 57 44 23 40 34 87 27 +~ 

Photography 331 441 518 737 234 105 268 138 506 380 253 206 120 116 200 0 

TOTALS Trawl 237 332 533 478 593 572 371 560 1100 592 110 184 159 252 129 

Photography 552 522 592 2558 5937 4839 6740 5657 3952 7983 2081 1140 1685 1188 1582 2003 

v 

* Station E5 Not Trawled 
458 

Table 5-17 . Total megafauna density (no ./hectare) from trawling and 

benthic photography : Cruise V. 

Ton VCS YC1 YC7 YC8 YC4 VC2 YC6 YC3 VC9 Y10 Y12 Y11 

PORIFERA Trawl 0 4 2 3 5 0 16 20 2 0 611 18 

Photography 0 0 0 72 230 243 39 0 15 60 221 60 

ALCYONARIA Trawl 0 2 7 1 23 1 9 0 0 0 1 1 

Photography 54 215 49 290 157 416 2263 243 195 165 193 140 

ACTINIARIA Trawl 19 44 26 0 27 17 3 0 10 0 0 26 

Photography 0 0 83 199 88 208 105 0 45 15 0 0 

DECAPODA Trawl 100 252 174 484 153 43 146 97 162 25 122 496 

Photography 127 452 134 72 354 973 369 319 558 540 505 925 

BIVALVIA Trawl 28 3 4 2 1 11 11 24 5 3 84 19 

Photography 0 0 0 0 0 0 0 0 0 0 0 0 

OPHIUROIDEA Trawl 2 7 118 149 62 20 47 47 69 8 22 46 

Photography 18 21 83 181 213 382 118 154 560 210 253 1731 

ECHINOR)EA Trawl 0 0 0 0 0 0 3 10 1 0 6 6 

Photography 109 0 0 0 0 0 0 0 0 0 0 0 

ASTEROIOEA Trawl 1 9 4 2 0 3 9 26 13 2 5 1 

Photography 36 0 16 0 0 34 0 44 0 15 0 20 

HOLOTHURODEA Trawl 0 3 2 4 2 8 1 1 1 G 6 4 

Photography 0 0 0 0 0 0 13 0 0 15 14 0 

CRIIVODEA Trawl 0 0 0 1 3 2 7 E 0 1 1 0 

Photography 0 0 0 0 0 0 4E2 132 15 0 0 0 

OTI#RINVERTEBRATE Trawl 9 23 115 19 15 5 101 26 27 0 31 9 

Photography 200 170 3342 72 0 34 104 154 75 . 255 118 160 

DEhffRSAL FISH Trawl 96 78 145 144 54 38 51 42 70 14 24 106 

Photography 437 213 213 234 422 85 195 88 180 195 85 480 

TOTALS Trawl 255 425 597 809 345 148 404 301 306 53 913 732 

Photography 981 1071 3920 1120 1464 2375 3668 1634 1643 1470 1389 3516 
459 

Total r1eiptwas ouisttg 

~eo0oo tooo 

Photography 140000- 

Trawl 90p a 

S 120000 r 

v 

100000 . app So 

o 

app 

40000- o 

r 20000 N O 

p Lo d 

Cl 6 2 3 7 8 9 4 10 11 5 12 

Cruise III Stations by Depth 

3000 

e 

v 

m 

x 2000 

_ 1500 

r lppp 

w 

C ypp 

c 0 

soo 

m 

o qpp 

v 

m 

x 300 

0 

s 

200 a 

r 

C loo 

m 

O 

Total Dacepod Density 

Photography 

" Trawl 

C162379241011512 

Total Asteroid/Ophiuroid Density 

Photography 

" Trawl 

0 

goo 

350 0 

r 

v 

x 

300 

m 

zso 

0 200 

Z 

150 

ap 100 

C 

O 

0 

C16 2 3 7 0 9 410 11 S 1Z 

Total Demersal Fish Density 

Photography 

" Trawl 

C162378941011512 

Central Tneaeet Stations 

Ordered by Depth In Graphs 

Station Depth (m) 

C1 353 

C6 525 

C2 629 

C3 866 

C7 950 

C8 1098 

C9 1325 

C4 1408 

C10 1605 

C11 2074 

CS 2401 

C12 28e2 

Figure 5-15 . Megafauna density comparisons of trawl and benthic 

photography : Cruise III . 
460 

Photography 

" Trawl 

Etb 1a 1 lc 2a 2 2r 7i 21 20 3 3e 31 38 3b 3 

Cruise IV Stations bg Depth 

3000 

m 

L 2500 0 

r 

m 

S 

2000 

1500 

z 

a tooo 

.r 

C 300 

m 

O 

0 

E1b 1s 1 1a 2v 2 2e 7i 21 2e 3 9s 91 8@ 9r 7 

zoo 

Total Docspod Density 

Photography 

" Trawl 

Total Astenfd/Ophtunid Deesitg 

Photography 

" Trawl 

m 

0 

r 130 

H 

m 

O 100 

Z 

N 

50 O 

C 

m 

O 

0 

Etb 1s 1 10 2s 2 2r 7i 21 2a 3 30 3d 30 3b S 

Total Dowenal Fish Density 

Photography 

" Trawl 

b 

- . 

Cruise Iv 

Eastern Transact Stations 

Ordered by Depth Groups 

Station Depth (m) 

EIB 338 

ElA 355 

E1 356 

E1C 360 

E2A 624 

E2 626 

E2E 626 

E28 630 

E2D 656 

E2C 667 

E3 809 

EX 847 

E3D 853 

E3A B54 

E38 867 

ES 2890 

cup. t t.2. z 2.2rti2. a ae3+a.s s 

Figure 5-16 . Megafauna density comparisons of trawl and benthic 

photography : Cruise IV . 

eoo 

e 

v aoo 

Z 

e app 

Z 

r 

c Z°° 0 0 

0 

Total 1leqeinune Density 

e 

r 

u soon 

0 

S 

o X000 

Z 

.+ 

c 

m 

D 

0 
461 

Total rapiwia Doos1ty 

O 

r 

m 

2 

C 

Z 

N 

1000 

C 

0 

O 

0 

Photography 

Trawl 

w 

M517842539101211 

Cnln V StauM M oorta 

1000 

a BOO 

.+ 

v 

m 

S 600 

0 

z 400 

.+ 

e 2°° 

m 0 0 

1800 

Total DoeMN DewN1g 

Photography 

" Trawl 

cruse v 

West-Central Transact 

Stations Ordered by Depth 

Station Depth (m) 

WCs 27e 

WC1 338 

WC7 417 

WCB 449 

WC4 479 

WC2 515 

WC6 552 

WC3 748 

WC9 767 

WCtO 807 

WC12 1217 

WC11 1231 

fC3 1 7 0 . 2 6 8 9 10 12 1 

Total Asteroid/Ophiuroid Densitg 

Photography 

" Trawl 

tboo 

o 1400 .+ 

m 1200 

S 

1000 

o 

Z GOO 

ao 600 

.+ 

400 

C 

m 

D 

200 

0 

1I;3 1 7 0 4 2 6 3 9 10 12 11 

500 -

m 

Photography 

N ,00V m 

S 

300 - 

O 

Z 

e 100 

m 

O 

0 

VC5 1 7 . 4 2 6 3 9 10 12 11 

Figure 5-17 . Megafauna density comparisons of trawl and benthic 

photography : Cruise V. 

Total Demnal Fish Deestto 
462 

trawl densities in these cases was, for the most part due to the small 

holothuroid, PeniaAOne sp . discussed previously. The density peak at 

Station C7 was due primarily to large aggregations of bivalves of the 

family vesicomyiadae not sampled by trawling. 

Total megafauna densities from benthie photography exceeded that of 

trawling at all Cruise III stations, differing by no less than a factor of 

five at Station C1 (769 animals/ha and 154/ha, respectively, Table 5-15) . 

Station C12 was not sampled by trawling. Other groups from Cruise III in 

Figure 5-15 also show benthic photography density estimates higher at all 

stations than those obtained from trawl samples. Total decapod density 

for Cruise III (Fig. 5-15) shows a very similar pattern for both sampling 

methods resulting in density maximums at Station C6 and secondary peaks at 

Station C8 . 

Density plots for combined asteroids and ophiuroids also resulted in 

similar patterns for both photography and trawling . Minimum densities 

appeared at Stations C1 and C10 and the maximum density occurred at_ 

Station C2 for both sampling types. The maximum asteroid/ophiuroid 

density in benthic photography resulted primarily from a single 

unidentified ophiuroid taxa (298 of 423 animals/ha) . Similarly, a single 

species of ophiuroid, Ovhiocamaa faseiculata accounted for a density of 

188 individuals/ha out of the total asteroid/ophiuroid trawl density of 

194/ ha. 

Demersal fish densities at Cruise III stations were variable but 

generally low at Station C9 (1325 m) and deeper stations . Station 

densities calculated from benthic photography differed from trawling by a 

maximum of 12 .5 times that of trawling at Station C5 " The highest 

demersal fish density from photographs was recorded at Station C8 (336 

fish/ha) . 

Figure 5-16 shows Cruise IV densities divided into the three mayor 

depth intervals, E1 stations (338-360 m), E2 stations (624-667 m) and E3 

stations (809-867 m) . Station E5 has data only from benthic photography 

and was not sampled by trawling . 

Total megafauna at the top of Figure 5-16 shows the same general 

trend as Cruise III. Benthic photography densities exceeded trawl 

megaPauna densities at all stations, but only by a small margin at three 

of the shallow stations. The density differences between the techniques 
463 

were most pronounced at the mid-depth station group where all E2 station 

benthic photography densities exceeded those from trawling by a factor of 

at least 3 .5 (Station E2D, Table 5-16) . 

The deeper, E3 station group also shoaled large differences between 

total megafauna densities obtained from photography and trawling. Benthic 

photography densities surpassed trawling values by factors of between 6 

and 18 but overall, the density values were lower for both sample types 

compared to the mid-depth, E2 stations. 

The three taxa selected for comparisons in Figure 5-16 show some 

variation in which trawling resulted in higher density estimates. For the 

decapod crustaceans, trawling samples resulted in higher density values at 

three stations, E1 B, E1 A and El . The remaining Cruise IV stations were 

all represented by higher decapod densities derived from benthic 

photography. 

For the asteroids and ophiuroids combined, a total of five stations 

were represented by higher densities resulting from trawl samples, 

Stations E1A and E1 of the shallow group and Stations E3D, E3A and E3H of 

the deep station group . All five of these differences were due to the 

fact that not a single asteroid or ophiuroid was photographed at these 

stations resulting in zero densities (Table 5-16) . One of the highest 

densities for these groups was recorded from benthic photography at 

Station E5 (171/ha) which was not sampled by trawling. 

Demersal fish densities at Cruise IV stations were all higher from 

benthic photography compared to densities from trawling. No fishes were 

observed in photographs at Station E5 but the remaining station densities 

calculated from benthie photography differed from trawling by factors 

between 1 .3 Station E3 A) and 12 .7 Station E2 E) . The highest demersal 

fish density from photographs was recorded at Station E1C (737 fish/ha) . 

Cruise V total megaPauna densities are shown at the top of Figure 5-

17" The stations are arranged by depth between 278 and 1231 m but do not 

represent regular depth increments. The resulting general comparison of 

the two methodologies was the same as for Cruises III and IV ; at all 12 

Cruise V stations, total megafauna densities obtained from benthic 

photography analyses were higher than densities obtained from trawling. 

Two peaks in megafauna density from benthic photography occurring at 

Stations WC7 and WC6 require special mention. At Station WC7, the bulk of 
464 

the megafauna abundance resulted from a unique group of invertebrates, the 

Pogonophora/Vestimentifera . This group, included within "other 

invertebrates" in Table 5-17, accounted for 85% of the total megafauna 

density at that station (3342/tube worms/ha from a total megafauna density 

of 3920/ha). Tube worms were also obtained by trawling by the addition of 

a special device between the trawl doors designed to snag the animals 

which live partially buried below the surface at the sediment. This trawl 

device, certainly less efficient than photography, resulted in a density 

of only 52/ha for pogonophoran and vestimentiferan tube worms. 

Station WC6 represented another anomalous area, particularly in 

context of the trawl sample at this station. A large volume of living 

scleractinian corals as well as a number of antipatharian corals were 

collected in the trawl . These were difficult to enumerate and are not 

fairly represented in the total megafauna density of 404 animals/ha from 

the trawl sample (Table 5-17) . In contrast, the high total density 

represented for this station from benthic photography in Figure 5-17 was_ 

primarily the result of sea pens (2263/ha) . The very high density of 

these alcyonarians (an order of magnitude higher than any other Cruise V 

station) may be related to the proximity of an area of abnormally high 

productivity sampled by the trawl at this station. 

Decapod crustacean densities for Cruise Y stations in Figure 5-1T 

show generally higher values from benthic photography as compared to 

trawling. Two stations were exceptions, WC7 and WCS where trawl density 

estimates surpassed that of photography . The peak density of 973 

decapods/ha (Table 5-17) occurred at Station WC2. The highest decapod 

density obtained from trawl samples (496/ha) was at the deepest station of 

Cruise V, Station WC11 which also showed a similar peak in density 

resulting from photography (925/ha) . 

Asteroid/ ophiuroid densities were similar between the sampling 

methods at the four shallowest Cruise V stations (Fig. 5-17). Densities 

from photography were higher than from trawls for the remaining deeper 

stations with small peaks in photographs densities at Stations wC2 and 

WC9 . The maximum density for asteroids/ophiuroids was at Station WC11 

Where large numbers of small ophiuroids were photographed (1731/ha, Table 

5-17) but very few ophiuroids were collected in the trawl. 
465 

A similar pattern between benthic photography and trawling density 

estimates is shown for demersal fish in Figure 5-17 " Densities from 

photography samples were higher at all Cruise V stations compared to trawl 

density estimates. The highest density for fish (480/ha) was seen at 

Station WC11 which was reflected by similar density peaks in the trawl 

sample . 

5 .3 QUALITATIVE RESULTS OF CRUISE I PHOTOGRAPHY 

The overall quality of Cruise I photographs obtained by the Woods 

Hole Mini-Angus System did not permit the detailed benthic photography 

analyses as was performed on samples obtained from Cruises II-Y that were 

obtained using the LGL BUCS apparatus . In general, the majority of the 

photographs obtained during Cruise I were at an extreme altitude (above 4 

m) which prohibited both adequate lighting of the bottom and proper 

identification of any biota which appeared within individual frames. An_ 

unexpected problem also arose preventing accurate assessment of the area 

of each frame. During Cruise I, the Woods Hole Mini-Angus system provided 

altitude information as only a telemetered signal to the chart recorder on 

the surface and not as part of the data chamber information exposed onto 

each photograph. Due to the variation in altitude above the bottom from 

one frame to another, it was necessary to establish an exact correlation 

between the frame to be analyzed and the altitude mark on the chart 

recording. The difference in altitude between photographs taken eight 

seconds apart could easily be several meters which would translate into a 

large difference in area. If the wrong altitude was chosen from the 

confusing mass of tic marks representing camera altitudes on the chart 

record, resulting density estimates and size measurements could be grossly 

over- or underestimated . For these reasons it was necessary to utilize 

the photographs obtained from Cruise I, Stations C1-C5 in a qualitative or 

semi-qualitative descriptive manner only. The following brief accounts of 

each station include notable observations of dominant or unusual biota and 

general lebensspuren descriptions. 
466 

5 .3 .1 STATION C1 

A total of 235 of the 800 possible frames were usable for a general 

description of this station during Cruise I. The total transeet length 

was 2 .31 nm. The dominant lebensspuren feature was white colored mounds 

of sediment in the shape of conical lumps. These lumps bad apparently 

been excavated from underlying sediment of a lighter color than the 

surrounding surface. Other lebensspuren were composed of little other than 

these white-colored conical lumps . There were a few frames which had 

numerous small depressions but in general depressions were uncommon. The 

only other notable feature was long, narrow grooves or trails across many 

of the frames. 

The most common animal was the hake UroyhYSis cirratus although only 

four were seen within the 235 usable frames. We observed three flatheads, 

Bembrops gobioides , and one large brachyuran crab (possibly the genus 

Pyromaia ), several small shrimp swimming off the bottom (probably 

Penaeopsis serrata), and two unidentified sea pens within a single frame. 

5 .3 .2 STATION C2 

A total of 250 of the possible 800 frames at this station were 

suitable for interpretation. The transact covered a total distance of 

0 .94 nm . Station C2 Was characterized by a much higher density of large 

depressions or burrows than at the shallower Station C1 . These 

depressions were not generally arranged in any regular pattern (e .g ., 

complete or partial circles) but in many frames, numerous depressions 

appeared in dense groups or clusters. 

A large percentage of the observed lebensspuren appeared to be 

directly caused by a dominant large ophiuroid, possibly of the genus 

ODhiomuaium . In numerous frames these ophiuroids apparently produced 

individual depressions with each of their fine arms. 

Tracks left behind by motile ophiuroids was another common feature. 

As a result o!' crawling along the bottom these ophiuroids produced series 

of short grooves regularly spaced from each other and at an angle to the 

direction of movement. Many photographic frames showed these features 

with the individual ophiuroids responsible in view. 
467 

What appeared to be a single ophiuroid producing the lebensspuren 

features mentioned above was by far the most abundant species. A total of 

131 individual ophiuroids were enumerated from the 250 usable frames. The 

highest density was represented by a group of 10 individuals in a single 

frame with an estimated area of 14 m2. Ophiuroids appeared in two 

separate aggregations along the photographic transect . The first 

aggregation was encountered after the first 29 min of the 110 min total 

transect time had elapsed. This aggregation continued for approximately 

34 min or 31% of the total transact distance of 0 .94 nm equivalent to a 

linear distance of 1770 m. The second aggregation did not occur until 32 

min after the first had ended. The gap between the aggregations was 1656 

m . The second aggregation continued until the end of the transact 

resulting in a minimum aggregation size of 857 m. 

Other megafauna observed at Station C2 included two starfish which 

both appeared to be Persephonaster echinulatus. This species was also 

obtained from trawl samples at Station C2 during Cruise I. Only a single 

shrimp was observed in the 250 frames . Two pagurid crabs, ParaDaaurus , 

were observed with attached com mensal anemones. Other invertebrates 

included a single unidentified zoantharian anemone, a single unidentified 

echinoid and a single holothuroid, Mesothuria. 

Two large macrourid fish were recorded but could not be further 

identified due to the high altitude of the photographs . TWO eels, 

Syriaphobranchus , were recorded. Two other fish were observed but could 

not be distinguished in any detail. One appeared to be in the family 

Alepocephalidae . 

5 .3 .3 STATION C3 

A total of 1 .66',nm were transacted at this station but only 113 

frames were in view of the bottom and even fewer were below an altitude of 

5 m, permitting detailed observation of the substrate or biota. In 

general, the dominant lebensspuren features consisted of rounded 

depressions or burrows in apparent random arrangement without recognizable 

symmetry or patterns. 

The biota observed in the 113 frames included three shrimp, one of 

which was the genus Glyphocrangon , two small ophiuroids with short white 
468 

arms, one holothuroid ( Mesothuria) and four fish. The fish included two 

Synaphobranchus , one Halosaurus , and one large unidentified macrourid, 

probably Nezumia. 

One very unusual photograph was obtained from the supplementary 

second camera equipped with a wide angle 16 mm lens. A fairly large 

holothuroid was photographed while completely off the bottom . Although it 

is common for smaller holothuroids (2-3 em Peniagone) to be found in the 

water column, this larger individual appeared to be on the order of 20 em 

in length. A small sediment plume could be seen below where it had 

launched itself off the bottom. 

5 .3 .4 STATION C4 

A total of 208 frames were acceptable for use in the general 

description of this station. The transact covered a total distance of 

2.32 nm. The substrate was highly bioturbated in comparison to previous_ 

Stations C1-C3 showing considerable texture and a general bumpy nature. 

The major lebensspuren features included depressions and conical lumps but 

these occurred only in low density. Other than small-scale texturing of 

the sediment, most of the substrate surface was relatively unPeatured . 

White colored sediment patches were evident due to their contrast in 

color, though they did not have much relief. Visible evidence of bottom 

water currents was photographed in some areas of the transact. In these 

photographs, small features or objects projecting above the bottom 

produced tails of accumulated sediments (lineations) to one side of the 

features only. These features can be produced by strong bottom currents 

transporting resuspended sediments which are then redeposited on the down 

current sides of objects. The deposition occurs as a result of the 

reduced energy due to turbulence (and ability to keep sediment in 

suspension) as the water current flows over and around an obstruction or 

depression. In one frame, a stalked object, possibly a dead pogonophoran 

worm tube or a sea pen had produced a depression in the sediment where it 

had been pushed back and forth by water movement. 

In some segments of the transact in proximity to those showing 

lineation features, there was also evidence of bottom scouring. These 

frames exhibited a very rough texture in comparison to the fine grain 
469 

nature of the sediment typical of previous Cruise I Stations C1-C3-

Evidence of scouring and resulting sediment lineations in some frames 

appeared to be restricted to relatively small areas. The beginning of the 

transact shoaled the more common fine grain sediment texture indicating the 

absence of significant bottom currents. 

One of the components of the coarse material evident in the majority 

of the transact was small, white colored fragments many of which had the 

shape and features of pteropod shells. These white, presumably calcareous 

objects were very numerous in some frames with estimated densities in the 

range of several hundred fragments per m2 . 

At one point along the transact there were large (1 m) lumps of mud 

projecting above the bottom with relief of approximately 0.2-0 .5 m. These 

large displaced portions of sediment with resulting trenches and large 

depressions map be the result of localized down-slope slumping of unstable 

sediment lagers known to occur in this area of the continental slope south 

of Louisiana. 

Megafauna was relatively abundant at this station with the 

holothuroid Peniagone by far the dominant taxa. The highest density of 

Peniajtone was in a single frame with 30 individuals in an estimated area 

of 2 .0' m2 representing a density of approximately 15 per m2 . A total of 

560 individuals were enumerated from the 208 frames. However, they were 

not evenly dispersed throughout the transact. One cluster of holothuroids 

began about one-third into the 2 .32 nm transeet and continued for 

approximately 266 m over the following 7 :min of the transact drift. The 

second cluster did not occur until after a 647 m gap in the transact . 

This second cluster of Peniagone continued in all visible photographic 

frames for the following 6 min representing a transact distance of 228 m. 

In the last 5 min of the transact (190 m in length) there was one other 

aggregation of Peniaaone , with one frame containing 45 individuals in an 

estimated area of 3 .5 m2 " 

There was also a variety of other biota enumerated in the 208 usable 

frames at this station. Invertebrates included three glass sponges 

Hyalonema sp., three worm tubes, Hyalinoecia tubieola and two other worm 

tubes which appeared to be pogonophorans . There were a total of six 

shrimp observed, four of which were unidentifiable, either caridean or 

penaeid but one was of the genus GlyphoeranAOn and another large shrimp 
470 

was the genus Plesiopenaeus. Echinoderms other than the 560 Peniagone 

included one unidentified crinoid and three ophiuroids of the same species 

with white colored arms. 

Fishes observed at Station Cu included one Synaphobranchus , one hake 

(Gadidae), and two unidentified individuals, one of which was eel-shaped, 

perhaps a small Synaphobranchus . 

5 .3 .5 STATION C5 

A large number of frames were lost at this station due to starting 

the camera at the surface before lowering to the bottom depth of 2470 m at 

this station while continually taking photographs along the way. The 178 

remaining frames were especially significant due to a poor trawl 

deployment at this station. 

The general texture of the flat and unfeatured bottom sediment 

appeared very floceulent with very few burrows. Of the few depressions 

that were observed, many were arranged in very distinct circular patterns 

usually forming a complete ring of between fine and seven burrows, 

typically six. The dominant lebensspuren were the trails left behind, by 

holothuroids, asteroids (complete with the impressions of tube feet), taxa 

not readily identified by their trails and tracks . Fecal deposits from 

holothuroids were relatively common. 

Biota enumerated from the 178 usable frames primarily included 

echinoderms . Only one taxa of holothuroid was actually seen in the 

photographs; Pelopatides , a spectacular specimen in situ. Apparently this 

genus is either lost by passing completely through the net during trawling 

or the remaining animal tissue is so unconsolidated it cannot be 

identified in trawl samples ; there was never a single specimen collected 

from trawls throughout the entire project. 

Other invertebrates were recorded including three unidentified sea 

pens and a single shrimp, Plesiopenaeus armatus . A single asteroid, 

Plutonaster intermedius , was recorded. No fish were seen in any of the 

visible frames at Station C5 . 

6 .0 REFERENCES 
473 

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Bartlett, M .S. 1975 . The statistical analysis of spatial patterns . 

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Berg, C.J., Jr. 1985 . Reproductive strategies of mollusks from abyssal 

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7 .0 ATTACHMENTS 
483 

7 .0 ATTACHMENTS 

ATTACHMENT A1 .0 : Supportive Data, Section 3 .0 

A1 .1 : Hydrographic Profiles by Cruise 

A1 .2 : Sediment Data 

ATTACHMENT A2 .0 : Supportive Data, Section 4 .0 

A2 .1 : Summary of Macrofaunal Densities (no/m2) by Mayor 

Taxonomic Group, Cruises I-V 

A2 .2 : Summary of Macrofaunal Taxa by Station and Cruise 

A2 .3 : Summary of Macrofaunal Diversity Data by Station 

and Cruise 

A2 .4 : Polychaete Feeding Types 

A2 .5 : Depth Ranges of 150 Most Abundant Polychaetes _ 

A2 .6 : Polychaete References 

A2-7 : Annotated List of Abundant Polqchaete Species 

A2 .8 : Distribution of Abundant Polychaete Species 

A2 .9 : Summary of Invertebrate Species Found in Trawl 

A2 .10 : Summary of Fish Species Found in Trawl 

ATTACHMENT A1 .0 : Supportive Data, Section 3 .0 


ATTACHMENT A1 .1 : Hydrographic Profiles by Cruise 
489 

Cerise I 

Ttmparotun (C) 

10 20 30 3 

Salinity (ppt) Dis 

4 35 36 37 

solved Oxygen (ml/1) 

2 3 4 5 6 

0 0 . . . ., 0 . . . 

r 

1 .J . J. 

300 - 300 ~" 500 

.. 

1000 

,. 

1000 1000 

t .j 

d 

1500 1500 1500 

2000 2000 2000 

2500 2500 2500 

Phosphate (y~M) 

0 i 2 

Nitrate (pM) 

0 10 20 30 40 

Silica (Jim) 0 10 20 30 

0 0 0 ,, 

. .~ . ' " + 

. . . ~. .L 

500 500 

. .. 

500 

. . 

, .., 1000 1000 1000 ' 

E ,., 

t 

a 

$ 1500 1500 1500 

2000 2000 2000 

2500 2500 2500 
490 

Cruise II - Western Transect 

Temperature (C) Salinity (ppt) Dissolved Oxygen (ml/1) 

0 10 20 34 35 36 37 2 3 4 5 6 

0 0 ,1 0 . 

1L 

500 "~ 500 , 500 

.' 

,.,1000 1000 1000 

.L+ 

91500 ~ 1500 1500 '. 

2000 2000 2000 -

2500 - ' 25002500' 

Phosphate (AM) 

0.0 1 .0 2.0 

0 

soo 

. . . 

1000 . .. 

V 

.Lr 

91500 . ' . 

2000 

2500 

Nitrate (0) 

0 10 20 30 10 

0- 

.r, .r . 

J 

S 

.: 

500 - 

. " 

1000 . 

1500 " ' 

2000 - 

2500 - 

Silica (yrn) 

4 10 20 30 

0 

. ; 

500- .. 

1000 . 

1500-

2000 -

2500 . 
491 

Cruis e II - Central Tronsect 

~ emperoturo (C) Salinity (ppt) Dls sotved Oxygen W/1) 

10 20 30 3 4 35 36 37 2 3 4 5 6 

0 0 :. p 

t 

soo soo soo 

t ., 

. 

1000 1000 100o 

, 

~ 

v 

. 

t 

a 

~ 

1500 1500 1500 

2000 

zooo 2000 

' zsoo 

2500 2500 ' 

Phosphate (uhl) 

Nitrate (uM) 0 1 2 0 10 20 30 40 Vice (JAM) 0 10 20 30 

. . . ~ . . 

300 500 500 

. . 

1000 1000 1000 

. 

r 

.~ a 

1500 1500 1500 

2000 2000 2000 

2500 2500 2500 
442 

Cruise II - East Tronsect 

~ Temperotun (C) Salinity (ppt) Diss olved Oxygen (mt/1) 

10 20 3 4 35 36 37 2 3 4 5 6 

0 ~' 0 fr . ~ 0 " , . 

r 

soo soo 500- 

1000 - 1ooo 1ooo 

. . 

, ... 

E 

L 

a 1500 

~ 

1500 1500 

2000 2000 2000 - 

2500 2500 2500- 

3000 3000 3000 

Phosphate (uM) 

0 i 2 

Nitrate (uM) 

0 10 20 30 40 

0 

Silica (UM) 

0 10 20 30 

0 0 

soo " ". soo '~ soo 

1o00 '~ 1o00 ' . 1o00 

t 1500 1500 1500 

.r 

O 

2000 2000 2000 

2300 2500 2500 

3000 ~' 3000 3000 "" 
493 

Cruise III 

Temperature (C) Salinity 

pt) Dis solved Oxygen (ml/1) 

0 10 20 30 3 4 36 37 2 3 4 5 6 

0 - 

0 ' =~ 0 

500 

.' 

500 ~ 500 

' . 

1000 1000 _ 1000 

v 

1500 o 1500 1500 

2ooo 2oo0 2000 

2500 2500 2500 , 

0 

Phosphate (uM) 

1 2 3 

Nitrate (yM) 

0 10 20 30 40 

Silica (Jim) 

0 10 20 30 40 

0 0 0 s 

500 " ~ 

. . 

500 

. , 

500 " 

' , 

1000 

r 

1000 

i 

1000 

C 

G 

v 

. . . 

t 

y 

a 

o 1500 1500 1500 

2ooo 2ooo soon 

2500 2500 2500 
494 

Temperature (C) 

0 10 20 30 3 

p 

2 

, . 

i 

I 

1000 1000 

v 

r 

a 

0 

2000 2000 

3000 3000 

Cruise IV 

Salinity (ppt) Dtasolyed Oxygen (ml/1) 

4 35 36 37 2 3 4 5 6 

.: f` . 0 { j 

.L L 

i 

.ti . . 

1000 

" 2000 

3000 . 

Phosphate (uM) Nitrate (uH) S111ce (AM) 

0 1 2 0 10 20 30 40 0 10 20 30 

0 , ~ .~ 0 f, 0 

r '.L 

:S . . 1 . . 

.~ 

1000 1000 1000 

v 

2000 2000 2000 

1 t 

3000 ' 3000 ' 3000 
495 

Temperature (C) 

10 20 30 3 

Cerise Y 

Salinity (ppt) 

4 35 36 37 

Dissolved Oxygen (ml/1) 

2 3 4 5 6 

0 - 0 0 

.~ ~ 

500 , 500 ., 500 

t 

.r 

1000 1000 1000 

1300 1500 1500 

Phosphate (yM) 

0.0 1 .0 2.0 

500 ,, 500 

.' 

E ,. ., , . 

d 

1000 1000 

Nitrate (jiM) Silica (Jim) 

10 20 30 40 0 10 20 30 

0 

500 

1000 

I . . . 

1500 j 1500 J 1500 


ATTACHMENT A1 .2 : Sediment Data by Cruise and Station 
499 

CLAY 

SANDY SILT 

CLAY C 

20 SAND 

CLAYEY SILT 

20 

_ 

CLAY [L "YE " 

SsNO SILT 

20 

SILTY 

SAND SAND 

SANDY 

S ANDY ILT 7 SILT 

b0% 

Figure A1 .2-1 . Sediment triangles Cruise I. 

Station C-1 

Station C-2 

Station C-3 

Station C-4 

Station C-5 
500 

I 

i 

l \ 

Station W-1 Station C-i Station E-1 

Station W-2 Station C-2 Station E-2 

" . 

Station W-3 Station C-3 Station E-3 

Station W-4 Station C-4 Station E-4 

Station 

& 

u-s Station c-s station E-s 

Figure A1 .2-2 . Sediment triangles Cruise II . 
n 

u.. 

Station C-01 

Sta t ion C-02 

Station C-08 

Station C-OS 

vro. SILTY ~ 

CLAY CLAY 

A 

SAND 

m 

CLAYEY SILT -can curt 

'AND 

~ 

NO 541 

20 

L

SILTY ' SANDY 

.0 SAND SILT 110 

Station C-03 

501 

Station C-06 

Sta t ion C-07 

Station C-09 

Station C-11 

Station C-04 

Station C-12 

Figure A1 .2-3 . Sediment trinagles Cruise III. 
502 

A 

A n 

a 

Station E-1 Station E-2 Station E-3 

l`~T-~'\ 

Station E-1R Statiori E-2A Station E-3R 

1 

Station 

\ 1 

E-iB 

/ Z& 

Station E-2B Station E-3B -

Station E-1C Station E-2C Station E-3C 

.o. 

a. 

swr x" 

Station E-2D Station E-30 

L1D 

7.1q 

a 

!KT ' 7Ypi 

frr wo w ru 

w eoa 

1 

Station E- 2E Station E-5 

Figure A1 .2-4 . Sediment triangles Cruise IV . 
503 

Station WC- 1 Station WC-it 

Station WC- 2 Station WC- 3 

station uc- 4 Station UC-12 

Station WC- 5 Station WC- 6 

Station WC- 7 Station WC- 9 

Station WC- 8 Station UC-10 

C 

CLAY 

SA IN SILTY 

CLAY CLAY 

to 

SAND 

CLATEY SILT _CL,, CLAYEY 

SAND SILT 

w 

SILTY 

. 

SANDY 

$AND AZAND-LT U z LA, SOLT . ., , LT , 

Figure A1 .2-5 . Sediment triangles Cruise V. 
504 

Cruise 1 

r 

u 

A 

a 

r 

i .,. 

N 

r 

e 

r 

H 

r 

d 

goo 

so 

60 

40 

20 

0 

Statiew 

Figure A1 .2-6 . 

saw96 

siH9g 

clay96 

C-1 C-2 C-3 C-4 C-5 
505 

Cruise II West Trsnsect 

too 

. . 

." 

v 

C 

w 

e 

g 

r 

so 

60 

40 

zo 

0 

Station 

sand 

Silt 

clay 

Figure A1 .2-7 . 

W-1 W-2 W-3 W-4 W-5 
506 

Cruise 11 Central Trensect 

goo 

so 

60 

40 

20 

0 

Figure X1 .2-8 . 

sand 

silt 

clay 

C-1 C-2 C-3 C-4 C-5 
507 

Cruise 11 East Transact 

too 

so 

60 

40 

20 

sand 

silt 

clay 

Figure A1 .2-9 . 

E-1 E-2 E-3 E-4 E-5 
v, 

0 

00 

too 

.. 

r 

x 

so 

60 

40 

w 

v 

r 

d 

20 

0 

Cruise III Shallow to Deep 

sand 

sill 

clay 

Figure A1 .2-10 . 

C-01 C-06 C-02 C-03 C-OT C-08 C-09 C-04 C-11 C-03 C-12 
v 

L 

.,. 

a 

""" 

8Z-3 

OZ-3 

OZ-3 

3Z-3 

£-3 

V£-3 

8£-3 

3£-3 

4£-3 

sa 

;. 

V Z-3 

11 

St-3 

3t-3 

Z-3 

1-3 

N 

t-3 

w 

a0 

r' 

C ~ ~ ~ N 

r 

QN4}"Ji ins 4800jaa 
v, 

0 

too 

r 

Y 

A 

." 60 

y 40 

r 

C 

r 20 

Cruise V Shallow to Deep 

sand 

silt 

clay 

Figure A1 .2-12 . 

WC- s WC- I WC- 7 WC- a WC- s VC- z WC- 6 WC- 10 WC- a WC- 9 VC- > > WC- 12 

ATTACHMENT A2 .0 : Supportive Data, Section 4 .0 

ATTACHMENT A2 .1 : Summary of Macrofaunal Densities (no/m2) 

by Mayor Taxonomic Group, Cruises I-V 
515 

Cruise I. 

STATION 

C 1 C 2 C 3 C , C 5 Average 

TAXOp 

NEMATODA 469 618 343 1175 1500 821 

06TRACOOA 70 375 630 231 50 271 

COPEPODA 179 310 214 483 252 288 

BIVALVIA 138 91 179 240 141 158 

TANAIDACEA 108 290 170 193 29 158 

BRYOZQA 3 12 21 64 3 21 

ISOPODA 580 182 161 146 64 227 

11hPMIPODA 190 97 85 53 9 87 

APLACOPHORA 56 47 56 26 9 39 

NEhERTEA 44 32 35 59 18 38 

OPHHXt01DEA 23 12 32 44 26 27 

SIPINCILA 18 6 21 18 0 13 

CUMACEA 79 23 18 47 3 34 

PaRIFERA 0 3 3 12 3 4 

SCAPFIOPODA 12 15 23 38 23 22 

SCYPfI0ZDA 18 0 0 3 0 4 

6ASTROPODA 35 9 26 3 3 15 

HOLOTHLJROIDEA 0 0 0 0 0 0 

OLI60CNAETA 6 0 0 0 26 6 

ASCIDIACEA 0 0 9 32 0 8 

HYDROZOA 0 0 0 0 0 0 

BRACNIOPODA 0 9 0 0 0 2 

ARACHlIDA:ACARINA 0 0 0 0 0 0 

Kim 3 0 3 9 0 3 

ECHINOIOEA 3 0 3 3 15 5 

PRIJIPILIDA 0 0 3 0 21 5 

SCLERACTMEA 0 0 0 0 0 0 

OECAPOOA 0 6 0 0 0 1 

h1YSTACOCARIOA 0 0 0 0 0 0 

STIhENT. 0 0 0 0 0 0 

ECHNa2A 0 0 0 0 0 0 

ACTMNARIA 0 0 0 0 0 0 

ALCYQiARIA 0 0 0 0 0 0 

TtRNLLARll1 0 0 0 0 0 0 

q21NODEA 0 0 0 0 0 0 

PYCN09QNIDA 0 0 0 0 0 0 

111SC. A11111MA 0 0 0 0 0 0 

IfhNCFIMATA 0 0 0 0 0 0 

ASTEROIMA 0 0 0 0 0 0 

MYSIDACEA 0 0 0 0 0 0 

ARCHIAMELIDA 0 0 0 0 0 0 

CEPHALOCFIOADATA 0 0 0 3 0 1 

CNRWDIA 0 0 0 0 0 0 

08TEICMTMYES 0 0 0 0 0 0 

TOW 2034 2137 2035 2882 2195 

Without Nwrhtods 1565 1519 1692 1707 695 
516 

Cruise II Western. 

wI wz wa w4 ws Rwrq . 

nEMRr70A 6a5 Laos 12a35 2a 347 93t 4

sTmcaoA z 2z2 M s9 1z25 n 22 

a~oo~ 2D4 201 45a 522 404 3a3 

91tJ1LV1R 218 112 189 102 84 137 

TOMIMKEA 1.0 147 1A4 140 112 1,6 

"IVOM 84 7 70 14 77 30 

18~OOR 106 211 96 161 28 121 

AlHIPOW 21 03 33 14 7 26 

ilLi~OPF10i~1 77 14 44 7 21 34 

tEl'EATER 49 03 42 21 7 30 

OPIl l lM1 DEA 28 77 133 7 0 49 

SIPINULA 140 49 0 14 0 4i 1

IAAEA 21 21 42 7 7 20 

P'QRIPEM 0 7 7 7 84 21 

SCRPHOPOOA 14 0 2S 7 7 11 

9CYRGZM 0 0 7 0 0 1 

mmg 7 21 0 7 7 

FMORMND 1 MR 0 42 14 0 0 11 

OL160qFETR 0 7 0 0 0 1 

fGC 10111 0 14 7 0 7 6 

IllfCPl)M 0 0 0 0 0 0 

111-10MI1 ~1 0 0 0 7 0 1 

1"WCilIIai:ACfRllii 0 0 0 0 0 0 

KIlDF*M'CMi 0 0 0 0 0 0 

OCHlNqlM 0 0 7 0 0 1 

PRI!!lLIa1 0 0 7 0 0 1 

9CL9VICT IIER 0 0 0 0 0 0 

~lP~1 0 7 0 0 0 1 

MIRTICOqrllq! 0 0 0 0 0 0 

IIEMf O 0 0 0 0 0 

ECNIM 0 0 0 0 O 0 

iICTIM1AIlA 0 0 0 0 0 0 

MD~IA 0 0 0 0 0 0 

flUNELLM1A 0 0 0 0 0 0 

CRIlDiM 0 0 0 0 0 0 

rya"oo10ia o 00 0 0 0 

M"18C . lllIM111 0 0 0 0 0 0 

FEMIQIPORR 0 0 0 0 0 O 

ABT0101 M 0 0 0 0 0 0 

f1V81p~B1 0 0 0 0 0 0 

XM1AlNLIpl 0 0 0 0 0 0 

CEPHRLOCHUFMM 0 0 0 0 O 0 

C11"!1MIA 0 0 0 0 0 0 

Oi'fE1G11lIYEB 0 0 0 0 0 0 

TTotal 780 27Z2 2A 11 1?22 id06 

Ni thout tlrotodo 1122 1333 1376 M1 =0 
517 

Cruise II Central . 

STAT 1 al 

C i C Z C 9 C 4 C S Avwow 

T111=1 

main" X04 410 786 1147 1274 9Q0 

067HRCM 42 330 319 323 232 329 

COPEPOOA 168 706 401 A07 323 490 

BItJiLVIR 211 133 263 36e 184 233 

TlMi I OACEA 91 433 337 230 74 233 

EIYaQI 30 +9 35 137 28 39 

18Q~OOA 106 372 16 1 382 112 245 

MW1 PW1 70 134 134 74 32 97 

OIJICOPlM 91 21 53 49 11 43 

low Ta I 33 39 33 45 23 43 

OPNIUl01MA 67 29 29 32 36 42 

81PlNDA.R 11 4 11 23 0 10 

C1MFICE'A 11 63 23 34 18 81 

Pa11FE1W 0 0 4 21 4 6 

8Cir1OPOOiR 0 7 7 14 35 13 

8C1VlDM 7 0 7 . 14 6 

OliB111IRM 42 11 46 35 14 30 

iMbTltM113ER 0 4 14 14 4 7 

OL I OOqFETW 0 7 40 4 18 16 

HOC 1 D 1lM 0 0 14 49 0 13 

HVpOZOA 14 4 11 11 0 8 

ONFICM I 7 11 4 0 23 9 

AVCNIIOA:RCiW ItW 0 0 0 18 0 4 

KIMMOMMMI 0 0 0 7 0 1 

ExH1101DEA 0 0 4 14 14 6 

PRIlfIjLlal 4 0 4 4 28 8 

ACT I tER 0 0 0 0 0 0 

OEC!!WR 4 4 0 0 0 2 

Ml8'ifAOMlal 0 0 0 0 0 0 

r09Q~1~1/1~8TIfEMT 0 0 O 0 0 O 

Ex111UW 0 0 0 0 0 0 

ICTIMiI"IIR 4 0 0 7 0 2 

ML4MM1A 0 0 0 0 0 0 

TtA111111LA11A 0 0 0 0 7 1 

C1111IOIm 0 0 0 7 0 1 

PyCl~~Ilal 0 0 0 0 0 0 

"18C . IMfiM 0 0 0 0 0 0 

IBIIQ~TR 0 0 0 0 0 0 

A676b1M 0 0 0 0 0 0 

INSIaRM 0 0 0 0 0 0 

i11011AlELial 0 0 0 0 0 0 

LOCHWORM 0 0 0 0 0 0 

CIiR1PED1A 0 0 0 0 0 0 

O`7EIQRMIE3 0 0 0 0 0 0 

Total 176 352 1 3090 387T 2337 

W i th~out Ilrolodo 1122 2391 2M7 TM 1253 
518 

Cruise II Eastern. 

6M IN 

111 

E 1 E 2 E 8 E 4 E S f11wrom 

f'EMT~I 1491 1= 17M 1124 774 14W 

06TiFC00A 211 204 4.4 364 174 278 

CQEPM 314 295 434 396 306 334 

91 WLU i A 372 147 267 257 69 222 

TlMl QCEA 03 147 305 202 N 173 

61WOm01i 184 21 91 43 16 83 

1901000111 ,9 77 112 173 95 101 

iMPH I POOA 0 112 96 .6 11 53 

!lL1 182 91 20 23 0 66 

HOERTEA 35 56 21 21 21 31 

QH 1 tAM1 DEA 35 21 84 17 3 37 

S I PISA 77 36 35 17 21 37 

CtRMICEA 21 7 77 34 37 33 

PQIIFE'Ri 0 21 14 25 116 35 

8Cl~1~R 21 0 0 17 0 8 

9CVPIOZOA 106 0 42 13 11 34 

OlArTlOPOOR 21 0 14 13 21 14 

FMGTMFIOIOEA 21 56 35 21 0 27 

0. I OOCHE1 II 0 35 28 21 03 34 

ROCIDIACER 0 7 29 8 11 11 

H1DROZM 42 0 26 0 0 14 

BIWCMIOPODA 0 7 0 0 0 1 

ifwicHl l lDR : RMI lei 0 0 14 0 16 6 

K I IOWV"CHi 35 0 0 0 0 7 

ECH11'OIM 0 7 0 4 S 3 

PRIMILLIDR 0 0 14 0 0 3 

SCLEPW T 1 HER 0 0 7 0 0 1 

~fp~A 0 7 O 0 0 1 

1'wSTl~OCf~110R 0 0 0 0 0 0 

PO0011/«ST I MMT 0 0 0 0 0 0 

EQIIIAI 63 0 0 0 0 13 

WTIM11M1A 0 0 7 4 0 2 

fLCYQM1 R 0 0 0 0 0 0 

TIRNILM1A 7 0 0 0 0 1 

CR1101M 0 0 0 0 0 0 

P1~C106011p1 0 0 0 0 0 0 

1`118c . Mn ~ 0 0 0 0 0 0 

MiaHoMIRrh o 0 0 0 0 0 

AsTeMiM 0 0 0 0 0 0 

nYsiaKMI o 0 0 0 0 0 

ORCHiMISLiM o 0 0 0 0 0 

LOCHWOM 0 0 0 0 0 0 

CIlr11PEDIR 0 0 0 0 0 0 

OSTE I CMMIES 0 0 0 0 0 0 

Tonal 35W 32M 4103 Ml 1= 

W i thout Nrotodo 1914 1374 2314 1761 1066 
Cruise III . 

STATION 

C 1 C 6 C 2 C 3 C 7 C a C 9 C 4 C11 C 5 C 12 Average 

TAXOII 

NEMATOOA 225 144 288 228 1568 884 2937 847 463 449 256 750 

OSTRACOOA 28 42 347 246 393 379 253 168 21 253 77 201 

COPEPODA 60 84 63 88 358 349 312 267 119 144 56 169 

BIVALVIA 112 109 147 218 396 218 337 214 95 193 91 194 

TANAIDACEA 28 46 126 196 512 193 133 116 32 64 35 134 

BRYOZOA 0 0 18 18 7 67 140 179 607 42 4 98 

ISOPOOA 70 98 42 46 232 95 137 154 28 63 56 93 

Af'pHIP00A 46 91 46 42 232 98 112 28 7 11 0 65 

APLACOPHORA 91 25 42 42 211 56 42 32 0 11 0 50 

PEMERTEA 21 14 11 18 46 35 7 42 11 7 4 20 

OPHIUROIDEA 42 0 4 53 32 46 18 39 4 53 0 26 

SIPUNCULA 7 0 4 11 21 21 4 7 4 4 0 8 

CUMACEA 18 0 21 18 74 28 39 7 4 7 18 21 

PORIFERA 0 0 0 4 7 46 28 25 25 7 25 15 

SCAPHOPODA 0 4 4 21 28 18 21 32 28 32 4 17 

SCYPHOZOA 7 0 4 0 14 32 32 53 39 7 0 17 

6ASTROPOOA 7 4 7 0 32 14 11 7 0 0 0 7 

HOLOTKIROIDEA 0 0 0 4 7 18 11 11 7 18 0 7 

aL160CHAETA 0 0 0 4 7 4 0 7 0 11 14 4 

ASCIDIACEA 0 0 0 7 0 4 21 39 7 4 0 7 

HYDROZOA 4 0 7 0 4 0 0 4 4 0 0 1 

BRACHIOPODA 0 0 0 0 4 7 4 0 158 25 0 18 

ARACFNIOA:ACARINA 0 0 4 0 14 11 0 4 4 0 0 3 

KINORHYNCHA 0 0 0 0 4 0 0 7 4 4 0 2 

ECNINOIDEA 0 0 4 0 0 0 4 0 4 4 4 1 

PRIAPULIDA 0 0 0 0 0 0 0 0 0 7 7 1 

SCLERACTINEA 14 0 0 4 0 0 0 7 0 0 0 2 

DECAPODA 0 0 0 0 4 0 0 0 0 0 0 0 

MYSTACOCARIDA 0 0 42 0 0 0 0 0 0 0 0 4 

P060NOPHaRA/VESTIMENT 0 0 0 0 18 0 0 0 0 0 0 2 
N 

O 

Cruiae III (cont'd), 

STATION 

C 1 C6 C2 C3 C7 Ca C9 C4 C11 CS C12 Average 

TAXOII 

ECMIURA 0 0 0 0 0 0 4 0 0 0 0 0 

ACTINIARIA 0 0 0 0 0 0 4 4 4 0 0 1 

ALCYONARIA 0 0 0 0 0 4 0 0 0 0 0 0 

TURBELLARIA 0 0 0 4 0 4 4 0 0 4 0 1 

CRINOIDEA 0 0 0 0 0 0 4 0 0 0 0 4 

PYCN060NIDA 0 0 0 0 0 4 4 4 0 0 0 0 

MISC . ANT110ZOA 0 0 0 0 0 0 0 0 0 0 0 0 

IfMICHORDATA 0 0 0 0 0 0 0 4 0 0 4 0 

ASTEROIDEA 4 0 0 0 4 0 0 0 0 0 0 1 

MYSIOACEA 0 0 0 0 0 0 4 4 0 0 0 0 

ARCHIANNELIDA 0 0 0 0 0 0 0 0 0 4 0 0 

CEPHALOCHORDATA 0 0 0 0 0 0 0 0 0 0 0 4 

CIRRIPEDIA 0 0 0 4 0 0 0 0 0 0 0 0 

OSTEICHTHYES 4 0 0 0 0 0 4 0 0 0 0 0 

Total 788 661 1231 1276 4229 2579 4611 2264 1667 1420 651 

Without Nematode 563 517 943 1048 2661 1695 1674 1457 1204 971 395 
Cruise IV . 

STATION 

E1B E1C ElA E1 E25 E2 E2C E2A E2D E2E E3A E38 E3C E30 E3 ES Aven4e 

TAXqi 

NEMATODA 1400 1298 1326 1561 2011 1558 1639 1119 2098 2246 1428 1644 2172 1144 1228 488 1520 

OSTRACOOA 172 267 102 270 505 389 326 1084 411 512 484 267 428 435 267 144 379 

COPEPOOA 98 175 105 193 344 225 182 151 196 256 277 214 281 182 161 144 199 

BIVALVIA 361 484 323 421 260 211 225 172 218 396 354 235 372 400 179 133 296 

TANADACEA 116 172 81 165 439 393 393 519 491 456 368 295 509 425 298 74 325 

BRYODDA 526 393 95 379 1168 653 891 105 161 747 319 519 288 123 151 74 412 

ISOPODA 35 56 77 25 246 189 179 172 144 214 91 151 151 119 109 81 127 

Af'PHPODA 39 39 49 18 88 91 70 137 91 77 126 119 158 91 151 21 85 

APLACOPHORA 102 67 74 130 105 130 77 109 112 133 102 35 77 84 63 4 88 

NEMERTEA 60 70 56 56 56 70 70 53 60 56 42 70 81 60 42 7 57 

OPHIUROIDEA 39 70 56 74 60 28 25 28 21 46 70 77 95 91 46 0 52 

SIPINCULA 161 175 144 182 60 67 49 46 70 123 28 46 39 63 42 4 81 

CUMACEA 49 81 18 11 60 70 18 39 49 53 49 56 119 35 32 32 48 

PORIfERA 46 32 4 28 70 74 56 4 63 60 46 28 81 70 32 137 52 

SCAPHOPODA 60 63 56 105 28 18 28 0 21 28 46 18 32 56 11 4 36 

SCYPHOZOA 116 53 182 126 32 4 7 4 25 21 14 11 0 11 11 7 39 

6ASTROPODA 14 14 11 21 32 7 . 4 18 25 42 35 25 28 49 14 7 22 

HOIOTHUR0IDEA 18 14 7 7 42 60 63 28 84 56 21 46 46 42 49 14 37 

aL160CHAETA 11 21 0 0 53 53 63 4 35 0 39 42 21 39 35 21 27 

ASCIDIACEA 0 0 0 0 11 0 11 11 4 18 25 11 32 25 21 11 11 

HYDROZOA 4 7 98 7 4 4 7 0 18 4 0 4 7 4 11 7 12 

BRACHIOPODA 0 0 0 0 4 0 0 0 0 0 0 0 4 0 4 0 0 

ARACHNIDA:ACARINA 0 11 0 0 4 7 0 0 0 4 4 14 4 7 14 4 5 

KINORNYNCHA 28 49 4 60 0 0 0 0 0 0 0 0 7 0 4 0 10 

ECFIINOIDEA 0 4 0 4 4 4 4 0 0 4 4 11 0 4 4 4 3 

PRIAPULIDA 7 0 7 4 0 0 0 4 0 0 0 0 0 0 0 0 1 

SCLERACTINEA 0 0 0 0 0 4 4 0 0 0 4 0 0 0 0 0 1 

DECAPODA 4 4 7 4 0 4 7 4 0 11 0 0 0 0 0 0 3 

nYSTACOCARIDA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

POG0.IVEST. 0 0 0 0 0 0 0 0 0 4 0 0 4 0 4 0 1 

N 
Cruise IV (cont'd) . 

STATION 

E16 E1C E1A E1 E28 E2 E2C E2A E20 E2E E3A E36 E3C E30 E3 ES Avenge 

TAXON 

ECHIURA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

ACTINIARlA 0 0 0 0 0 0 0 0 0 0 0 0 4 0 0 0 0 

ALCYONARIA 0 0 0 0 0 0 4 7 4 0 4 0 0 4 7 0 2 

TURBELLARIA 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

CWNOIDEA 0 0 0 0 0 0 0 0 0 4 0 4 0 0 7 0 1 

PYCNOGONIDA 0 0 0 0 0 7 4 0 0 0 4 0 4 0 0 0 1 

fiISC. ANTHOZOA 0 0 0 0 0 0 0 0 0 0 7 0 0 0 0 0 0 

FfMICHORDATA 0 0 0 4 0 0 0 0 0 0 0 0 0 0 0 0 0 

ASTEROiDEA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

nYSIDACEA 0 0 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

ARCHIANNELIDA 0 0 0 0 0 0 0 0 0 4 0 0 0 0 0 0 0 

CEPHALOCHORDATA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

CIRRIPEOIA 0 0 0 0 0 0 0 0 0 0 0 0 4 0 0 0 0 

OSTEICHTHYES 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

Total 3470 3619 2886 3855 5682 4320 4406 3818 4401 5575 3991 3902 5044 3563 2997 1422 

Wfihout Nsmstoda 2070 2312 1560 2294 3671 2762 2767 2699 2303 3329 2563 2298 2872 2419 1769 934 

N 

N 
Cruise V. 

STATION 

- WC I WCS VC7 WCA WC, IrC2 WC6 WCA WC9 WC3 WCd WCC Average 

TAX011 

rEMATODA 986 1758 1200 839 632 407 2053 533 681 589 919 677 940 

OSTRACODA 67 228 182 751 396 372 589 393 705 491 263 277 393 

COPEPOOA 140 147 109 140 316 214 246 151 133 109 240 112 168 

BIVALVIA 305 326 144 246 196 316 270 144 218 211 172 214 230 

TANAIDACEA 77 221 186 365 333 421 347 126 351 295 144 137 250 

BRYOrOA 70 544 239 49 56 263 319 133 133 263 98 81 187 

ISOPOOA 88 39 109 207 207 147 221 56 123 81 154 147 132 

AI'PHIPODA 46 186 84 165 95 102 95 84 130 98 63 60 101 

APLACOPHORA 39 11 4 42 42 46 77 35 70 28 35 14 37 

PEMERTEA 28 158 25 46 32 14 42 18 32 25 18 35 39 

OPHIUROIDEA 32 32 4 4 25 18 63 18 42 35 74 63 34 

SIPUNCULA 35 46 18 21 7 28 70 21 25 4 7 14 25 

CUMACEA 11 130 14 18 28 4 63 11 21 14 7 0 27 

PORIFERA 25 63 11 0 14 39 32 4 25 14 28 21 23 

SCAPHOPOOA 28 53 46 11 28 35 28 18 21 25 18 21 28 

SCYPHOZOA 49 53 14 7 18 7 0 14 21 4 4 14 17 

6ASTROPODA 11 70 11 28 7 32 21 11 18 4 14 21 21 

HOLOTFIUROIDEA 0 4 0 0 18 0 0 0 4 14 14 18 6 

OLIGOCHAETA 7 70 28 7 7 0 32 0 0 0 7 11 14 

ASCIDIACEA 0 0 0 0 0 4 7 11 18 14 11 0 5 

HYDROZOA 14 18 39 0 4 11 4 0 0 4 4 0 8 

BRACHIOPODA 0 0 7 0 0 4 0 0 0 0 0 0 1 

ARACHNIDA:ACARINA 0 46 4 4 4 0 11 14 0 0 0 0 7 

KINORHYNCHA 0 0 0 0 0 0 0 0 0 0 0 0 0 

ECHINOIDEA 0 4 0 0 7 7 0 4 0 7 7 4 3 

PRIAPUIIDA 0 0 0 0 0 0 4 0 0 0 0 0 0 

SCLERACTINEA 0 63 0 0 0 7 4 0 4 0 0 0 8 

DECAPODA 0 4 7 7 4 0 0 0 0 0 4 0 2 

MYSTACOCARIOA 0 0 0 0 0 0 0 0 0 0 0 0 0 

P060NOPHORA/VESTIMENT 0 0 0 0 0 0 0 4 0 0 7 0 1 rlj 

w 
N 

Cruise V (cont'd) . 

STATION 

WIC 1 WXS WK7 WCA WC4 WCZ UIiCb WCA UK9 WiC3 WCd WCC Averse 

TAX011 

ECMIURA 0 0 0 0 0 0 4 0 0 0 0 0 0 

ACTINIARIA 0 0 0 0 0 0 0 0 0 0 0 4 0 

ALCYaWARIA 0 0 0 0 0 0 0 0 0 0 0 0 0 

TURBELLARIA 0 0 0 0 0 0 4 0 0 0 0 0 0 

CRINOIOEA 0 0 0 0 0 0 0 4 0 0 0 0 4 

PYCNOGONIDA 0 4 0 0 0 0 0 0 0 0 0 0 0 

MISC . ANTHOZOA 0 0 4 0 0 0 0 4 0 0 0 7 1 

FEMICIiqi0ATA 0 7 0 0 0 0 0 0 0 0 0 4 1 

ASTEROIDEA 0 0 0 0 0 0 0 0 0 0 0 4 4 

MYSIDACEA 0 0 0 0 0 0 0 0 0 0 4 0 0 

ARCNIANNELIDA 0 0 0 0 0 0 0 0 0 0 0 0 0 

CEPHALOCHORDATA 0 0 0 4 0 0 0 0 0 0 4 0 0 

CIRRIPEDIA 0 0 0 0 0 0 0 4 0 0 0 0 0 

OSTEICHTHYES 0 0 0 0 0 0 0 4 0 0 0 0 4 

Total 2058 4285 2485 2957 2478 2498 4602 1803 2775 2329 2264 1948 

Without Nsmatods 1072 2527 1285 2116 1840 2091 2549 1270 2094 1740 1345 1271 

ATTACHMENT A2 .2 : Summary of Macrofaunal Taxa by Station and Cruise 
527 

Cruise I. 

STATION 

C 1 C2 C3 C4 CS Total 

TAXON 

TANAIDACEA 13 25 27 28 6 65 

ISOPODA 9 16 25 27 8 58 

BRYOZQA 1 2 5 8 1 11 

CUhIACEA 14 7 6 10 1 27 

AMPHIPODA 21 12 16 9 2 36 

BIYALVIA 14 8 16 17 9 35 

OASTROPODA 8 2 6 1 1 1 S 

PORIFERA 0 1 1 2 1 S 

SIPUNCULA 6 2 S 4 0 14 

06TRAOOUA 4 5 4 3 1 10 

NEMERTEA 6 4 4 4 3 8 

ASCIDIACEA 0 0 1 2 0 2 

OPHIUROIDEA 5 4 4 7 3 16 

HYDROZQA 0 0 0 0 0 0 

DECAPODA 0 2 0 0 0 2 

HOLOTHUROIDEA 0 0 0 0 0 0 

SG4PHOPODA 2 4 3 6 4 8 

OL100C1AETA 2 0 0 0 2 4 

ECHINOIDEA 1 0 1 1 2 2 

S( :LERACTINEA 0 0 0 0 0 0 

ACTINIARIA 0 0 0 0 0 0 

K I NORHYNCFIA 1 0 1 1 0 1 

PYCNOOOWIDA 0 0 0 0 0 0 

M I SC. ANT HOZQA 0 0 0 0 0 0 

ALCYONARIA 0 0 0 0 0 0 

ASTEROIDEA 0 0 0 0 0 0 

BRANIOPODA 0 1 0 0 0 1 

COPEPOQA 1 1 1 1 1 1 

CRINOIDEA 0 0 0 0 0 0 

APLAOOPHORA 1 1 1 1 1 1 

ARACHNIQA"INA 0 0 0 0 0 0 

NtqiIANNELIDA 0 0 0 0 0 0 

CEPWILO(:NORDATA 0 0 0 1 0 1 

CIRRIPEDIA 0 0 0 0 0 0 

ECHIURA 0 0 0 0 0 0 

HEMICHORQATA 0 0 0 0 0 0 

MYSI DAL'EA 0 0 0 0 0 0 

MYSTA00(,ARIDA 0 0 0 0 0 0 

NEPIATODA 1 1 1 1 1 1 

OSTEICNTHYES 0 0 0 0 0 0 

POOOPIOPHORA/VESTIMENT . 0 0 0 0 0 0 

PRIAPULIDA 0 0 1 0 1 1 

SCYPHOZOrA 1 0 0 1 0 1 

TURBEILARll1 0 0 0 0 0 0 

Total 111 98 129 135 48 
528 

Cruise II Western. 

STATION 

W 1 W2 W3 W1 WS Total 

TAXON 

TMIAI DACEA 11 15 20 12 8 50 

ISOPODA 3 14 8 17 4 35 

BRYOZQA 8 1 5 2 8 21 

CUMACEA 3 3 5 1 1 10 

AMPHIPODA 3 4 5 2 1 11 

BIYALVIA 8 8 12 15 6 24 

OASTROPOOA 0 1 2 0 1 3 

PORIFERA 0 1 1 1 3 6 

SIPUNCUU 7 5 0 2 0 9 

OSTRAOODA 2 6 4 3 1 6 

NEMERTEA 3 3 2 3 1 6 

ASCI D IACEA 0 1 . 1 0 1 2 

OPHIUROIDEA 1 1 1 1 0 1 

HYDROZOA 0 0 0 0 0 0 

DEG4PODA 0 1 0 0 0 1 

HOLOTHUROIDEA 0 2 1 0 0 3 

SG4PHOPODA 1 0 2 1 1 4 

OLIOOCHAETA 0 1 0 0 0 1 

ECHINOIDEA 0 0 1 0 0 1 

SCIERACTINEA 0 0 0 0 0 0 

ACTINIARIA 0 0 0 0 0 0 

KINORHYNqiA 0 0 0 0 0 0 

PYCNOOONIDA 0 0 0 0 0 0 

MISC. ANTHOZQA 0 0 0 0 0 0 

ALCYONARIA 0 0 0 0 0 0 

ASTEROIDEA 0 0 0 0 0 0 

BRACHIOPODA 0 0 0 1 0 1 

OOPEPODA 1 1 1 1 1 1 

CRINOIDEA 0 0 0 0 0 0 

APLAOOPHORA 1 1 1 1 1 1 

ARAqiNIDA:AGIRINA 0 0 0 0 0 0 

ARCHIANNELIQA 0 0 0 0 0 0 

CEPWILOCHORQATA 0 0 0 0 0 0 

CIRRIPEDIA 0 0 0 0 0 0 

ECHIURA 0 0 0 0 0 0 

HEMICHOROATA 0 0 0 0 0 0 

MYSIDACEA 0 0 0 0 0 0 

MYSTACO(`JIRIQA 0 0 0 0 0 0 

NEMATODA 1 1 1 1 1 1 

06TEICHTHYES 0 0 0 0 0 0 

POOOPIOPHORA/YESTIMENT . 0 0 0 0 0 0 

PRIAPUL1Duh 0 0 1 0 0 1 

SCYPHOZQA 0 0 1 0 0 1 

TURBELURIA 0 0 0 0 0 0 

Total 53 70 75 64 39 
529 

Cruise II Central . 

STATION 

C 1 C2 C3 C4 CS Total 

TAXON 

TMIAIDACEA 8 31 32 31 12 66 

ISOPODA 5 32 18 32 8 62 

BRYOZQA 8 9 9 14 2 34 

CUhIACEA 3 10 6 9 3 23 

AMPHIPODA 10 16 16 12 3 32 

BIYALYIA 9 13 16 17 13 36 

QASTROPODA 4 2 2 1 1 5 

PORIFERA 0 0 1 4 1 6 

SIPUNCt1U 2 1 2 3 0 8 

OSTRACODA 1 8 7 3 1 9 

NEMERTEA 7 3 5 5 3 8 

ASCIDIACfJ1 0 0 1 3 0 3 

OPHI UROI DEA 1 1 1 1 1 1 

HYDROIOrA 3 1 2 1 0 6 

DECAPODA 1 1 0 0 0 2 

HOLOTHUROIDEA 0 1 2 3 1 3 

S(`,~tPHOPDUA 0 1 1 3 2 4 

aiooaHAETn o 1 i 1 z 

ECFlIMOIDEA 0 0 1 1 1 1 

SCLERACTINEA 0 0 0 0 0 0 

ACTINIARIA 1 0 0 1 0 2 

KINORHYNCHA 0 0 0 1 0 1 

PYCNOOOIVIDH 0 0 0 0 0 0 

MISC. ANTHOZQA 0 0 0 0 0 0 

ALCYONARIA 0 0 0 0 0 0 

ASTEROIDEA 0 0 0 0 0 0 

BW1tlilOPODA 1 1 1 0 1 1 

COPEPOOA 1 1 1 1 1 1 

CRINOIDEA 0 0 0 1 0 1 

APLAOOPHORA 1 1 1 1 1 1 

ARACtiNIDA:ACARINA 0 0 0 1 0 1 

ARCHIANNELIQA 0 0 0 0 0 0 

CEPHALO(:MORDIITA 0 0 0 0 0 0 

CIRRIPEDIA 0 0 0 0 0 0 

ECHIURA 0 0 0 0 0 0 

HEMICHORQATA 0 0 0 0 0 0 

MYSIOACEA 0 0 0 0 0 0 

MYSTA00(,ARIDA 0 0 0 0 0 0 

NEPIATODA 1 1 1 1 1 1 

06TEICHTHYES 0 0 0 0 0 0 

POOOWOPHORA/YESTIMENT. 0 0 0 0 0 0 

PRIAPULiDA 1 0 1 1 1 1 

SCYPHOZQA 1 0 1 1 1 1 

TURBELURIA 0 0 0 0 1 1 

Total 69 135 129 153 61 
530 

Cruise II Eastern. 

STATION 

E 1 E2 E3 E4 E5 Total 

TAXON 

TANAIDACEA 8 18 27 27 10 61 

ISOPODA 6 8 9 24 11 45 

BRYOZOA 7 2 6 7 3 18 

CUh1ACFA 3 1 9 7 4 18 

AMPHIPODA 0 9 10 9 2 22 

BIVALYIA 12 11 12 17 4 27 

QASTROPODA 3 0 2 2 1 S 

PORIFERA 0 3 1 4 5 10 

SIPUNa1LA 5 3 1 2 3 11 

06TRACODA 4 6 5 4 1 11 

NEMERTFA 4 3 3 2 3 9 

ASCIDIACEA 0 1 2 1 2 4 

OPHIUROIDEA 1 1 1 1 1 1 

HYDROZQA 1 0 1 0 0 2 

DECARODA 0 1 0 0 0 1 

HOLOTHUROIDEA 1 2 2 3 0 4 

SCqpHOpOpA 1 0 0 2 0 3 

OLIOOCFWETA 0 3 3 2 2 S 

ECHINOIDEJI 0 1 0 1 1 1 

SCLERACT I NEA 0 0 1 0 0 1 

ACTIN111RIA 0 0 1 1 0 2 

KINORHYNCHW 1 0 0 0 0 1 

PYCNOOONIDA 0 0 0 0 0 0 

MISC. M(THOZQA 0 0 0 0 0 0 

ALCYONARIA 0 0 0 0 0 0 

ASTEROfDEA 0 0 0 0 0 0 

BW1a-IIOPODA 0 1 0 0 0 1 

COPEPOOA 1 1 1 1 1 1 

CRINOIDFA 0 0 0 0 0 0 

ApLACOPHORA 1 1 1 1 0 1 

ARACHN I DA:AGR I NA 0 0 1 0 1 1 

ARCHIMINEIIUA 0 0 0 0 0 0 

CEPW1LOCf10RDATA 0 0 0 0 0 0 

CIRRIPEDIA 0 0 0 0 0 0 

ECHIURA 1 0 0 0 0 1 

HEMICIi0R0ATA 0 0 0 0 0 0 

MYSIDACEA 0 0 0 0 0 0 

MYSTACOCARIQA 0 0 0 0 0 0 

NEMATODA 1 1 1 1 1 1 

OSTEICHTHYES 0 0 0 0 0 0 

POOOIJOPHORl1/YESTIMENT . 0 0 0 0 0 0 

PRIAPULIDA 0 0 1 0 0 1 

SCYPHOZpV1 1 0 1 1 1 1 

TURBELLARIA 1 0 0 0 0 1 

Total 63 77 102 120 57 
Cruise III . 

STATION 

C 1 C6 C2 C3 C7 CO Cg C4 CII CS C12 Total 

TAXOq 

TANAIDACEA 

ISOPODA 

BRYOZOA 

CUMACEA 

AhPHIP00A 

BIVALVIA 

6ASTROPOOA 

PORIFERA 

SIPtINCULA 

OSTRACOOA 

NEMERTEA 

ASCIDIACEA 

OPHIUROIDEA 

HYDROZOA 

OECAPODA 

HOLOTH1ROIDEA 

SCAPHOPOOA 

OLIGOCHAETA 

ECMINOIDEA 

SCLERACTINEA 

ACTINIARIA 

KINORHYNCHA 

PYCN060NIDA 

MISC. ANTHOZOA 

ALCYONARIA 

ASTEROIDEA 

BRACHIOPODA 

COPEPODA 

CRINOtDEA 

APLACOPHORA 

5 6 17 18 32 23 23 24 6 12 6 78 

2 5 10 10 29 19 21 15 6 10 8 65 

0 0 3 2 1 5 5 10 8 4 1 20 

2 0 5 3 9 7 8 2 0 2 3 30 

6 6 7 6 10 12 10 5 2 2 0 25 

10 9 11 12 16 18 18 13 11 8 7 42 

2 1 2 0 4 4 3 2 0 0 0 13 

0 0 0 1 2 8 5 7 4 2 5 21 

2 0 1 3 3 4 1 2 1 1 0 10 

1 5 5 5 5 3 2 2 1 2 1 10 

5 3 3 3 6 5 1 6 1 2 1 12 

0 0 0 1 0 1 5 4 2 1 0 8 

1 0 1 1 1 1 1 1 1 1 0 1 

1 0 1 0 1 0 0 0 0 0 0 3 

0 0 0 0 1 0 0 0 0 0 4 1 

0 0 0 1 1 1 1 3 2 2 0 8 

0 1 1 3 1 3 4 1 1 1 1 6 

0 0 0 1 2 0 0 1 0 1 2 3 

0 0 1 0 0 0 1 0 0 1 1 1 

2 0 0 1 0 0 0 2 0 0 0 4 

0 0 0 0 0 0 0 1 1 0 0 1 

0 0 0 0 1 0 0 1 1 1 0 1 

0 0 0 0 0 0 0 1 0 0 0 1 

0 0 0 0 0 0 0 0 0 0 0 0 

0 0 0 0 0 0 0 0 0 0 0 0 

1 0 0 0 1 0 0 0 0 0 0 2 

8 0 0 0 1 1 1 0 2 1 0 2 

1 1 1 1 1 1 1 1 1 1 1 1 

0 0 0 0 0 0 1 0 0 0 4 1 

1 1 1 1 1 1 1 1 0 1 0 1 w 

r 
w 

N 

Cruise III (cont'd) . 

STATION 

C 1 C6 C2 C3 C7 C8 C9 C4 C11 CS C12 Total 

TAX011 

ARACHNIDA:ACARINA 0 0 1 0 1 1 0 0 1 0 0 1 

ARCHIANNELIDA 0 0 0 0 0 0 0 0 0 0 0 0 

CEPHALOCMORDATA 0 0 0 0 0 0 0 0 0 0 0 0 

CIRRIPEDIA 0 0 0 1 0 0 0 0 0 0 0 1 

ECHIURA 0 0 0 0 0 0 0 0 0 0 0 0 

FfMICF10RDATA 0 0 0 0 0 0 0 1 0 0 0 1 

MYSIOACEA 0 0 0 0 0 0 0 1 0 0 0 1 

MYSTACOCARIDA 0 0 1 0 0 0 0 0 0 0 0 1 

NEMATODA 1 1 1 1 1 1 1 1 1 1 1 1 

OSTEICHTHYES 1 0 0 0 0 0 0 0 0 0 0 1 

P060NOPHORA/VESTIP'ENT 0 0 0 0 1 0 0 0 0 0 0 1 

PRIAPULIDA 0 0 0 0 0 0 0 0 0 1 1 1 

SCYPHOZOA 1 0 1 0 1 1 1 1 1 1 0 1 

TURBELLARIA 0 0 0 1 0 0 1 0 0 1 0 1 

Total 45 39 74 76 133 120 116 109 54 60 39 
Cruise IV . 

sTAnoN 

HIS E1C ElA E1 E28 E2 E2C E2A E2D E2E E3A E313 E3C E3D E3 E5 Total 

TAXON 

TANAIDACEA 20 20 10 14 40 42 31 39 38 39 39 33 47 40 30 15 124 

ISOPODA 6 5 6 7 23 25 24 18 21 19 16 22 26 20 15 14 87 

BRYOZOA 6 6 6 10 4 8 5 5 7 5 10 9 8 11 7 6 29 

CUMACEA 11 11 4 2 13 11 4 9 8 8 7 9 14 9 7 4 53 

AhPHIP00A 9 8 7 2 14 10 8 1l 10 11 13 12 17 14 1l 3 45 

BIVALVIA 19 17 15 18 10 9 7 8 10 9 14 12 12 14 9 8 32 

6ASTROPOOA 3 4 2 4 3 2 1 3 3 5 7 2 5 6 2 1 21 

PORIFERA 3 2 1 3 3 3 3 1 4 4 5 3 6 5 4 10 14 

SIPUVCULA 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

OSTRACODA 5 6 5 4 7 6 5 7 9 7 7 7 8 8 8 1 17 

NEMERTEA 7 5 7 7 7 8 6 6 6 4 5 7 7 6 4 2 17 

ASCIDIACEA 0 0 0 0 2 0 2 3 1 3 1 1 5 3 1 2 10 

OPHIUROIDEA 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 

HYDROZOA 1 1 2 1 1 1 1 0 2 1 0 1 1 1 2 2 7 

OECAPODA 1 1 2 1 0 1 2 1 0 3 0 0 0 0 0 0 7 

HOLOTHUROIDEA 1 2 1 1 4 2 3 2 3 3 3 3 3 4 3 3 7 

SCAPHOPODA 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 

O1160CHAETA 1 1 0 0 1 2 2 1 2 0 3 4 1 4 1 2 5 

ECHINOIDEA 0 1 0 1 1 1 1 0 0 1 1 1 0 1 1 1 3 

SCLERACTINEA 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 2 

ACTINIARIA 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 

KINORHYNCHA 2 1 1 1 0 0 0 0 0 0 0 0 1 0 1 0 2 

PYCNOGONIDA 0 0 0 0 0 1 1 0 0 0 1 0 1 0 0 0 3 

MISC. ANTFIOZOA 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 

ALCYONARIA 0 0 0 0 0 0 1 1 1 0 1 0 0 1 1 0 2 

ASTEROIDEA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

BRACMIOPODA 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 

COPEPODA 2 2 1 1 1 2 2 2 1 1 2 2 2 1 2 2 2 

CRINOIOEA 0 0 0 0 0 0 0 0 0 1 0 1 0 0 1 0 1 

APLACOPHORA 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

w 

w 
Cruise IV (cont'd) . 

STATION 

E1B E1C ElA E1 E28 E2 E2C E2A E2D E2E E311 E38 E3C E3D E3 E5 Total 

TAXON 

ARACHNIDA:ACARINA 0 1 0 0 1 1 0 0 0 1 1 1 1 1 1 1 1 

ARCHIANNELIDA 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 

CEPHAIOCHORDATA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

CIRRIPEDIA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

ECHIURA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

HEMICHORDATA 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 

MYSIDACEA 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 

MYSTACOCARIDA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

NEMATODA 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 

OSTEICHTHYES 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 

P060NOPH./VEST. 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 1 

PRIAPULiDA 1 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 1 

SCYPHOZOA 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 

TURBELLARIA 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 

Total 105 100 78 85 141 142 116 123 132 133 144 136 173 155 119 82 

w 

r 
Cruise V. 

TAX011 

TANAIDACEA 

ISOPODA 

BRYOZOA 

Clq'1ACEA 

AMPHIPODA 

BIVALVIA 

GASTROPOaA 

PORIFERA 

SIPUNCUIA 

OSTRACODA 

NEMERTEA 

ASCIDIACEA 

oPHiuaoioEA 

HYDROZOA 

DECAPOOA 

HOLOTHURqDEA 

SCAPMOPOQA 

aicocwAETA 

ECMINOIDEA 

SCLERACTINEA 

ACTINIARIA 

KINORHYNCHA 

PYCNOGONIOA 

MISC . ANTFqZOA 

ALCYONARIA 

ASTEROIDEA 

BRACHIOPODA 

COPEPODA 

CRINOIDEA 

APLACOPHaRA 

STATION 

- WC l WCS WC7 WCA WC4 WC2 WC6 WC 10 WC9 1MC3 Wll W12 Total 

8 25 12 19 24 11 29 22 34 28 20 25 86 

9 8 14 21 22 17 27 14 21 13 2 1 20 76 

6 6 3 5 4 10 9 3 6 7 2 9 28 

3 12 4 4 4 1 12 2 6 4 2 0 35 

6 12 7 9 11 6 10 9 9 6 10 8 34 

11 12 7 10 12 10 10 11 14 11 13 16 27 

2 5 3 5 2 5 5 2 3 1 4 4 22 

5 6 1 0 4 4 5 1 2 2 4 3 15 

1 1 1 1 1 1 1 1 1 1 1 1 1 

1 2 6 6 6 6 8 3 4 5 3 4 13 

5 10 5 4 5 2 3 4 5 3 3 5 11 

0 0 0 0 0 1 2 1 3 4 2 0 7 

> > > > > > > > > > > > > 

1 3 3 0 1 2 1 0 0 1 0 0 6 

0 1 2 1 1 0 0 0 0 0 0 0 4 

0 1 0 0 1 0 0 0 1 2 4 3 5 

1 1 1 1 1 1 1 1 1 1 1 1 1 

1 1 2 1 1 0 1 0 0 0 1 1 3 

0 1 0 0 1 2 0 1 0 1 1 1 3 

0 3 0 0 0 1 1 0 1 0 0 0 4 

0 0 0 0 0 0 4 0 0 0 0 0 0 

0 0 0 0 0 0 0 0 0 0 0 0 0 

0 1 0 0 0 0 0 4 0 0 0 0 1 

0 0 0 0 0 0 0 0 0 0 0 1 1 

0 0 0 0 4 0 0 0 0 0 0 0 0 

0 0 0 0 0 0 0 0 0 0 0 0 0 

0 0 1 0 0 1 0 0 0 0 0 0 1 

1 1 2 1 1 1 2 1 2 1 1 1 2 

0 0 0 0 0 0 0 0 0 0 0 0 0 

1 1 1 1 1 1 1 1 1 1 1 1 1 w 
w 

ON 

Cruise V (cont'd) 

STATION 

- WC 1 WCs vrc7 wca Wc4 vcz WCs WC 10 Wc9 wc3 WII W12 Total 

TAXaII 

ARACHNIDA:ACARNA 0 1 1 1 1 4 1 1 0 4 0 0 1 

ARCHIANNELIDA 0 0 0 0 0 0 0 0 0 0 0 0 0 

CEPHALOCHOROATA 0 0 0 0 0 0 0 0 0 0 0 0 4 

CIRRIPEDIA 0 0 0 0 0 0 0 0 0 0 0 0 0 

ECHIURA 0 0 0 0 0 0 0 0 0 0 0 4 0 

FEMICHORDATA 0 1 0 0 0 0 0 0 0 0 0 0 1 

MYSIOACEA 0 0 0 0 0 0 0 0 0 0 0 0 0 

MYSTACOCARIDA 0 0 0 0 0 0 0 0 0 0 0 0 0 

PEMATOOA 1 1 1 1 1 1 1 1 1 1 1 1 1 

OSTEICHTHYES 0 0 0 0 0 0 0 0 0 0 0 0 0 

P060NOPHORA/VESTInENT 0 0 0 0 0 0 0 1 0 0 1 0 1 

PRIAPULIOA 0 0 0 0 0 0 1 0 0 0 0 0 1 

SCYPHOZOA 1 1 1 1 1 1 0 1 1 1 1 1 1 

lURBELLARIA 0 0 0 0 0 0 1 0 0 0 0 0 1 

Total 65 118 79 93 107 86 133 82 117 95 98 107 

ATTACHMENT A2 .3 : Summary of Maerofaunal Diversity Data by 

Station and Cruise 
Cruise 1: Central Tr 

31-ABP 

C 1 CZ C3 C4 C5 

TAX011 

TANAIDACEA 2.25 2 .59 2 .73 2 .87 1 .70 

ISOPODA 1 .04 2 .17 2 .91 3 .12 1 .87 

BRYOZQA 0.00 0 .69 1 .33 1 .69 0 .00 

CUr1ACEA 2 .19 1 .56 1 .61 1 .93 0 .00 

AMPHIPOOA 2 .23 1 .62 2 .26 1 .74 0.00 

BIVALVIA 2 .11 1 .75 1 .97 2 .30 1 .48 

Cruise 11 : West ern Tnmsect Castrsl Tramsect Esstarm Tramsect 

STATION 

1U1 W2 W3 W4 W5 C 1 C2 C3 C4 CS E 1 E2 E3 E4 E5 

Tsxem 

TANAIDACEA 2 .02 2.39 2.81 2.32 1 .71 1 .69 2.80 2.88 3.09 2.24 2 .04 2.67 2.96 2.97 2.22 

ISOPODA 0.88 228 1 .78 2.71 1 .10 0 .55 2.90 2.67 2.94 1 .32 1 .75 1 .97 1 .97 2.91 2.31 

BRYOZQA 1 .24 0.00 0.69 0.69 1 .83 1 .92 1 .62 1 .79 1 .62 0.69 1 .17 0.04 1 .10 1 .15 1 .10 

CU'1AACEA 0.69 1 .10 1 .56 0.00 0.00 0 .69 1 .99 1 .39 1 .85 0.97 0.69 0 .00 1 .97 1 .75 0.85 

AMPHIPODA 0.69 0.56 0.69 0 .00 0.00 1 .60 2.00 1 .99 1 .92 0.00 0.00 1 .58 1 .55 1 .10 0.00 

BIVALVIA 1 .40 1 .79 2.02 2 .54 1 .21 1 .61 2.20 2.11 2.16 2 .05 2.13 1 .96 2 .40 2.28 0.93 

w 
Cruise 111: CNtral Tranect 

STATION 

C 1 C6 C2 C3 C7 Ce C9 C4 C11 CS C12 

TAX011 

TANAIDACEA 1 .51 1 .61 2.51 2 .27 2 .71 2.63 2.83 3 .01 1 .74 2 .39 1 .71 

ISOPODA 0.43 0.74 1 .99 2.27 3.1a 2.89 2.91 2 .30 1 .69 2 .00 1 .60 

BRYOZQA 0.00 0.00 0.58 0.53 0.00 1 .09 1 .07 1 .52 1 .06 0 .64 0.00 

CIIMACEA 0.53 0.00 1 .35 1 .07 1 .85 1 .92 1 .99 0.69 0.04 0 .00 0.97 

AMPHIPODA 0.99 1 .02 1 .54 1 .04 0.77 1 .66 1 .73 0.91 0 .69 0 .00 0.00 

BIVAIVIA 1 .76 1 .44 1 .92 1 .83 2.04 2.59 2.37 1 .84 1 .99 1 .42 1 .25 

Cruise IV : Essterm Tramsett 

STATION 

Eld EtC ElA Ei E28 E2 E2C E2A E21) E2E E3A E3B E3C E31) E3 ES 

TAX011 

TANAIOACEA 2 .69 2 .63 2.05 2.35 3.32 3.23 2.95 3.06 3.24 3 .25 3.20 3.15 3.39 3.23 3 .08 2.80 

ISOPODA 1 .71 1 .54 1 .68 1 .39 2.60 2.83 2.82 2.55 2.85 2 .47 2.62 2.86 2.82 2.84 2 .51 2.51 

BRYOZQA 0 .56 1 .16 1 .29 1 .23 0.14 0.40 0.55 1 .18 0.60 0 .34 0.95 0.67 1 .19 1 .96 1 .30 1 .48 

CUMACEA 2 .27 2 .17 1 .06 0.66 2.41 2.06 1 .35 1 .76 2.02 1 .78 1 .73 1 .93 2.33 2.06 1 .85 1 .17 

AMPIiIPODA 1 .35 1 .61 1 .27 0.53 2.25 1 .64 0.97 1 .60 1 .46 1 .69 1 .90 1 .61 1 .99 2.07 1 .62 0.58 

BIVALVIA 2 .25 228 2. 16 2.18 1 .63 1 .80 1 .53 1 .41 1 .75 1 .76 2.11 1 .95 2.19 2.18 1 .65 1 .30 

Cruise V " Westerm Central Transect 

STATION 

WC 1 VCS WC7 WCA WC4 WC2 WC6 WIC 10 WC9 IdC3 WIC 11 WC 12 

TAXOII 

TANAIDACEA 1 .64 2 .80 2 .06 

ISOPODA 1 .82 1 .99 2 .16 

2 .38 2 .60 1 .11 2.93 2.83 3 .00 2 .80 2.56 3.09 

2 .59 2 .64 2.52 3.07 2.50 2.82 2 .33 2.51 2.66 

0 

BRYOZQA 1 .53 0 .74 0 .40 1 .08 1 .06 1 .55 1 .35 0.90 1 .23 1 .23 0.00 1 .67 

CUMACEA 1 .10 2 .25 1 .39 1 .35 0 .97 0.00 2.13 0.66 1 .79 1 .39 0.69 0.00 

AMPHIPODA 1 .02 1 .50 1 .32 1 .07 0.96 1 .02 1 .29 0.38 1 .58 0 .66 1 .78 1 .58 

BIVALVIA 2 .00 2 .04 1 .59 1 .53 1 .98 1 .58 1 .71 2.03 2 .02 1 .85 2 .19 2.55 

ATTACHMENT A2 .4 : Polychaete Feeding Types 
543 

Attachment A2 .4 

POLYCHAETE F=E-=E=-:E3 I NC=' TYPES 

FAl"QLY NODC FEEDING TYPE FC 

ACROCIRRIDAE 51 SELECTIVE DEPOSIT FEEDER SDF 

AMPHARETIDAE 67 11 10 11 SDF 

AMF'HINOMIDAE 10 CARNIVORE C 

APHRODITIDAE 01 C 

ARABELLIDAE 33 C 

ARENICOLIDAE 62 NON-SELECT DEPOSIT FEED . NSDF 

BOGUEIDAE 76 to 11 NSDF 

CAPITELLIDAE 60 go 11 NSDF 

CHAETOPTERIDAE 49 SELECTIVE DEPOSIT FEEDER SDF 

CHRYSOPETALIDAE OS CARNIVORE C 

CIRRATULIDAE 50 SELECTIVE DEPOSIT FEEDER SUF 

COSSUF2IDAE 52 NON-SELECT DEPOSIT FEED . NSDF 

CTENODRILIDAE 53 ;1 is It NSDF 

DORVILLEIDAE 36 CARNIVORE C 

EULEPETHIDAE 05 11 C 

EUNICIDAE 30 OMNIVORE 0 

EUPHROSINIDAE 11 SELECTIVE DEPOSIT FEEDER SDF 

"FAMILY C" 99 CARNIVORE C 

FAUVELIOPSIDAE 55 NON-SELECT DEPOSIT FEED . NSDF 

FLABELLIGERIDAE 54 to 1 . .1 NSDF 

GLYCERIDAE 27 CARNIVORE C 

GONIADIDAE 28 11 C 

HARTMANIELLIDAE 77 C 

HESIONIDAE 21 11 C 

HETEROSPIONIDAE 47 SELECTIVE DEPOSIT FEEDER SDF 

IOSPILIDAE 17 CARNIVORE C 

LACYDONIIDAE 16 OMNIVORE 0 

LUMBRINERIDAE 31 CARNIVORE C 

MAGELONIDAE 44 NUN-SELECT DEPOSIT FEED . NSDF 

MAI,DANIDAE 63 is 11 it NSDF 

NEPHTYIDAE 25 OMNIVORE 0 

NEREIDAE 24 11 0 

ONUPHIDAE 29 SCAVENGER S 

OPHELIIDAE 58 NON-SELECT DEPOSIT FEED . NSDF 

ORBINIIDAE 40 to 11 11 NSDF 

OWENIIDAE 64 SUSPENSION FEEDER SF 

PAL1"IYRIDAE 09 CARNIVORE C 

PARAONIDAE 41 NON-SELECT DEPOSIT FEED . NSDF 

PHYLLODOCIDAE 13 CARNIVORE C 

PILAR6IDAE 22 OMNIVORE 0 

POECILOCHAETIDAE 46 SELECTIVE DEPOSIT FEEDER SDF 

POLYNOIDAE 02 CARNIVORE C 

QUESTIDAE 74 NON-SELECT DEPOSIT FEED . NSDF 

SABELLARIIDAE 65 SUSPENSION FEEDER SF 

SABELLIDAE 70 it 11 SF 

SCALIBREGMIDAE 57 SELECTIVE DEPOSIT FEEDER SDF 

SERPULIDAE 73 SUSPENSION FEEDER SF 
544 

Attachment A2 .4 (cont'd) 

POI-YCHAETE FEES I NC=' TYPES 

FPJ`QLY NODC FEEDING TYPE FC 

SIGALIONIDAE 06 CARNIVORE C 

SPHAERODORZDAE 26 NON-SELECT DEPOSIT FEED . NSDF 

SPIONIDAE 43 SELECTIVE DEPOSIT FEEDER SDF 

STERNASPIDAE 59 NON-SELECT DEPOSIT FEED . NSDF 

SYLLIDAE 23 OMNIVORE 0 

TEREBELLIDAE 68 SELECTIVE DEPOSIT FEEDER SDF 

TRICHOBRANCHIDAE 69 SDF 

TROCHOCHAETIDAE 45 SDF 

TYPHLOSCOLECIDAE 19 UNKNOWN ? 

ATTACHMENT 92 .5 : Depth Ranges of 150 Most Abundant Polychaetes 
547 

Attachment A2 .5 . Depth ranges of 150 most abundant polychaetes . (A = NODC 

code, B = minimum depth observed, C - maximum depth 

observed, D = taxes name .) The taxes are ordered by depth 

of maximum occurrence . (Depths are in meters .) 

A ~ d C D 

1 5001250115 320 349 NEPHTYS INCISA 

2 5001430503 505 505 PRIONOSPIO HETEROBRANCHIA? 

3 5001430596 302 615 PRIONO6PI0 SR D 

5001290399 320 615 NOTFRIA SPA 

S 5001310196 351 625 LlR'6RINERIS 'SP . J' 

6 5001230791 444 625 EXUG(lW 'SP .H' 

7 5001430599 320 629 PRION06PI0 SPA 

8 5001430506 343 629 PRION06PI0 STEENSIRUPI 

9 5001600307 302 631 NOT01"U1ST11S FEhfIPWUS 

10 5001360203 298 633 PROTODORVILLEA KEFERSTEINI 

11 5001670501 343 639 P'ELINNA CRISTAi'A 

12 5001220199 3M 639 AWISTROSYLLIS "PA' 

13 5001250303 335 A6LAOPHAMUS VERRIIII 

1, 5001260296 350 034 IS 'SP.E' 

15 5001409898 298 845 LEIT06COLOPL06 FRA6ILIS 

16 5001230790 336 H45 EXOGONE 'SP .O' 

17 5001411198 595 845 SABIOIUS SPA 

18 5001400998 358 847 CALIFIA SPB 

19 5001410222 302 SW ARICIOEA 1'AYLORI 

20 5001700187 298 852 CFIOPE SP. N 

21 5001230295 618 852 PION06YLLIS 'SP. D' 

22 3001 700193 291 060 00E SR.6 

23 5001700199 302 860 CHOW SPA 

24 5001230299 344 860 PION06riLIS 'SPB' 

25 5001290599 580 860 HYALINOECU JI.IVENALIS 

26 5001220999 291 870 LITOCORSA ANTEMATA 

27 5001310116 353 OM LlMDRINERIS 6REVIPES 

28 5001060199 343 1007 PHOLOE 'SP .C' 

29 5001410298 409 . 1007 IIRICIOEA ABRANCHIATA 

30 5001230296 290 1032 PIQM06YLLIS 'SP . H' 

31 5001060101 320 1032 PHOLOE MINUTA? 

32 5001630199 545 1032 ASYCHIS ATLANTICUS 

33 5001440197 344 1147 h1A6Q.ONA SF'. B 

3, 5001580696 580 1147 OPFfLINA SPD 

33 500170139Q 302 1198 FAdRICIA SPB 

36 5001230306 336 1198 SYLLIS (EFLERSIA) CORMJTA 

37 5001310104 291 1222 LUhBR1PEWS IATRIELLi 

30 3001410224 335 1222 ARICIDEA (ACMIRA) SIPPLEX 

39 500158999 349 1222 11P'"'10TRYPANELLA SP . A 

,0 5001410299 351 1222 ARICIDEA SP. A 

41 5001460199 298 1729 POECILOCHIIETI.IS SPA 

42 5001291096 554 1229 PARONLPHIS SPA- 1 

43 500123029 298 1232 PION06YLLIS SPA 

44 5001670303 347 1232 N°PHICTEIS GIR~ERI 

45 5001260599 595 1232 SPHAEREPFESIA SPA 

46 5001231099 616 1235 DRANIELLA 'SPA* 

47 5001299996 298 1236 SARS0NLPHIS SP. A 

40 5001160101 302 1236 PARALACYDONIA PARADOXA 
548 

Attachment A2 .5 (cont'd) 

A s C p 

49 500110102 340 1236 AEUICIRA BELGICAE 

50 5001690197 347 123$ TEREBELI.IDES DISTINCTA 

31 50017210 291 1330 EUCfiONf ROSEA 

52 5001409999 344 1330 MICRORBINIA LINEA 

53 5001230399 347 1330 SYLLI5 (CHLERSIA) SR.A 

54 5001410605 353 1330 CIR20F'FIQftU5 tIW4NCHIATUS 

55 5001400399 554 1330 SCOLORLO5 5V.A 

56 50015898 615 1330 OPHELINA SP.B 

57 5001409899 291 1389 LEITOSCOLOPLOS SP.A 

58 5001600299 358 1390 FETEROMASTUS SP.A 

59 5001230896 639 1390 SPHAEROSYLLIS SP.B 

60 5001280601 547 1419~ PROGONIADA REGIILARIS 

61 5001700204 298 1433 EUCH0NE INCOLOR? 

62 5001600310 291 1440 NOT011ASTUS AMRICANUS 

63 5001310301 298 1440 LUMBRINERIDES ACUTA? 

64 5001410211 344 1440 ARICIDEA CERRI.ITI 

65 5001230898 353 1440 SPHAERa6YLLIS PIRIFEROPSIS 

66 5001580695 35.5 1440 OPHELINA SRI 

67 5001410215 298 1444 ARICIDEA LOPEZI LOREZI 

68 5001430591 340 1482, PRIONO6PI0 SPA 

69 5Q0158010;5 340 1482 OPFELINA CYLINDRICAUDATA 

To soo I640 331 14e2 nYRiocHELE "SP . E71 500 1260891 , S66` 1482 51'HAER06YLLIS RENAUDAE? 

72 5001430594 320 1506 RRIONOSPIO EHLEIt:il 

73 5001 ?30796 335 1506 EXOGONE SP I) 

74 5001700196 622 1506 CHONE SR.D 

75 SM1299999 298 1 507 SAR5UNUPHIS NARTMANAE 

76 5001310124 302 1507 LlN1BRINERIS VERRILLI 

77 5001630399 ' 302 1507 MALDANE 'SP .A' 

70 5001580699 320 1507 QF'FIELINA SP.A 

79 5001607899 320 1507 NEOMFDIOMASTUS SP.A 

00 5001640201 ~ 320 1507 MYRIU(:FIELE HEERI? 

01 5001671901 ; 320 1507 AUCHENOPI.AX CRINITA 

02 5001690196 336 1507 TEREBELLIDES ATLANTIS 

03 5001630302 366 1507 MALDANE GLEBIFEX 

61 5001230701 618 1507 EXOGONE DISPAR 

05 5401670299 291 2118 AMPHARETE 'SP.A' 

96 5001290397 347 2124 NOTFWA I EXTOR 

67 5001100401 291 2377 PARAhPH1NOME JEFFREYSII 

5001540404 302 2377 DIPLOCIRRUS CAPENSIS 

0! 5001590101 335 2377 STERNASPIS SCUTATA 

90 5001410228 351 2377 ARICIDEA TRILOBATA? 

91 5001289999 550 2377 BATHYGLYCINDE 'SP .A" 

92 5001230795 615 2377 EXOGONE LONGICIRF2US? 

93 5001490302 291 2470 SPIOCHAETOPTERIS COSTARUM 

" 5001520103 320 2470 CO55UF2A DELTA 

95 5001 X31 U(Yb ` 335 2,70 SPIOPIiANES WI6LEYI 

16 5001580997 353 2470 TACHYTRYPME SP .C 
549 

Attachment A2 .5 (cont'd) 

A B C D 

97 500150644 455 2470 OPHELINA SP .F 

98 5001230710 505 2470 EXOGONE ATLANTICA 

99 5001230798 291 2524 EXOGOW SP.B 

100 5001430502 291 2524 RRIONOSPIO CIR21FERA 

101 5001130599 347 2524 MYSTIDES SP. A 

102 5001250304 349 2524 A6LAOPHAMUS CIRCINATA 

103 5001060399 605 2524 STFaENELAIS SP.A 

104 5001230/93 618 2524 EXQGONC 'SP. C' 

105 5001690101 291 2533 TEREBELLIDES STROEMI 

106 5001431006 343 2533 SPIOPHANES MISSIONEN515 

107 5001580999 291 2540 TACHYTRYPANE SP.A 

108 5001100499 298 2540 PARAhPHINOh1E SR. A 

109 50016699 335 2540 BARANTOLLA "SP. A" 

110 5001220501 343 2540 SYNELh11S KLATTI 

111 5401411101 344 2540 SABIDIUS CORNATUS 

112 5001400398 351 2540 SCOLOPLOS 'SP.H' 

113 5001520198 353 2540 COSSURA HETEROCHAETA 

114 5001230792 358 2540~ EXOGONE 'SP.E' 

115 5001550199 366 2540 FAUVELIOPSIS 'SR.A' 

116 5001439699 298 2853 AIIROSPIO DIBRANCHIATA 

117 50015495 347 2853 CHAETOZ0NE "SP .H" 

118 5001430201 351 2853 LAONICE CIRRATA 

119 5001672299 366 2853 ECLYSIPPE "SP. B" 

120 5001240601 618 2853 GEItATOCEPHALELOVENI 

121 5001589999~ 335 2897 KESUN SP. A 

122 5001540499 291 ~ 2900 OIPLOCIPRl15 'SG .A' 

123 5001431004 291 2902 SPIOPHANCS BERKELEYORUM 

124 5001700195 291 2904 CRONE SP .E 

125 5001431001 298 2904 SPIOPHANES BOMBYX 

126 5001500306 298 2904 TFIARYX ANNULOSUS? 

127 5001700299 298 2904 EUCHONE 'SR.A' 

120 5001319999 302 2904 AUGENERIA BIDENS 

129 5001240603 

' 

335 2904 CERATOCEPHALE OCIAATA 

130 1 500 1580901 444 2904 TACHYTRYRANE ,EFFREYS11 

131 5001410208 291 2945 ARICIDEA CATFIERINAE 

132 5001600402 291 2945 MEDIOMASTUS CALIFOFWIEN5IS 

133 5001310302 298 294.5 LUMR1NERIDES DAY1 

134 5001410214 302 2945 ARICIDEA (ARICIDEA) FRAGILIS 

135 5001410603 340 2945 CIRROPHORLIS LYRA 

136 5001602099 347 2945 RERESIELLA SPATHULATA? 

131 5001220201 320 2953 SIGAMBRA TENTACULATA 

131 5001410803 335 2953 LEVINSENIA tWINATA 

13! 500 1410201 291 2959 AkICIDEA SIJECICA 

140 5001410801 291 2959 LEVINS'ENIAGRACILIS 

141 5001500307 291 2959 THARYX MARIONI 

1+2 5001550198 291 2959 FAUVELIORSIS SP.B 

143 5001600306 291 

1 

2959 NOTQIASTUS LATERICEUS 

144 , 5001640399 343 2959 1hIYRIOWEMA SP.A 
550 

Attachment A2 .5 (cont'd) 

A B C D 

145 5001270107 344 2959 GLYCERA PAR lO5A? 

146 5001500499 344) 2959 CHAETOZONE 'SR.C' 

147 5001570201 344 2959 ASCLEROCHEILUS BERINGIANUS 

1,8 5001230799 347 2959 EXOG0NE 'SP.A' 

149 5001470101 347 2959 FETEROSPIU LUN61SSIMA? 

130 1 5M 1130501 622 2959 I"IYSTIDE5 BMALIS 

ATTACHMENT A2 .6 : Polychaete References 
553 

Attachment A2 .6 . Polychaete References . 

1 Banae, K_ and K. D. Habson. 1974 . Benthic errantiate 

polychaetes of British Columbia and Washington_ Bull . 

Fish . Res . Bd . Canada, Bull . 185. 

2. Blake, J . A. 1985 . POLYCHAETA FROM THE VICINITY OF DEEPSEA GEOTHERMAL VENTS IN THE EASTERN PACIFIC I : 

EUPHROSINIDAE, PHYLLOUOCIDAF, HESIONIDAE, NEREIDIDAE, 

GLYCERIDAE, DORVILLEIDAE, ORBINIIDAE, AND MALDANIDAE . 

BIOL_ SOC . WASH . BULL . No . 6, 1985, pp .67-101 . 

3. Day, J. H. 1967 . A Monograph on the Polychaetes of South 

Africa, Part I . Errantia, Part II . Sedentaria. 

British Museum (Natural History) London . Pub. No .656 . 

4 . 1973 . New Polychaeta from Beaufort, with a Key 

to All Species Recorded from North Carolina . NOAH 

Technical Report NI'IFS CIRC-375 . 

5. Fauchald, K. 1968 . Onuphidae (Polychaeta) from Western 

Mexico . Allan Hancock Monographs in Marine Biology, 

No .3 . University of Southern California, Los Angeles, 

California . 

6. - 1977 . The Polychaete Worms--Definitions and 

Keys to the Orders, Families, and Genera . Natural 

History Museum of Los Angeles County, Science Series 

28 :1-190 . 

7. 1982a. TWO NEW SPECIES OF ONUPHIS (ONUPHIDAE :POLYCHAETA) FROM URUGUAY. PROC . BIOL . SOC. 

WASH . 95(1), 1982, pp .203-209 . 

8. 1982b. Revision of Onuphis, Nothria, and 

Paradiopatra (Polychaeta:Onuphidae) Based upon Type 

Material . Smithsonian Contributions to Zoology; 

No . 356. 

9. Fauvel, P. 1923 . Faune de France . 5. Yolychetes Errantes . 

1927. Fauna de France .l4. PolychetQS Sedentaires . 

10 . Gardiner, S. L. 1975 . ERRANT POLYCHAETE ANNELIDS FROM NORTH 

CAROLINA. Journal of the Elisha Mitchell Scientific 

Society, Vol . 91, No . 3 . 

11 . Hart man, 0. 1939 . Polychaetous Annelids, Part I . Aphroditidae to Pisionidae . Allan Hancock Pacific Expeditions 

Volume 7, Nos. 1-2. University of Southern California 

Press, Los Angeles, California . 
554 

Attachment A2 .6 (cont'd) 

12 . Hart man, 0. 1940 . 

petalidae to 

ditions, Vol . 

L. A. , Ca . 

Polychaetous Annelids, Part II . ChrysoGoniadidae . Allan Hancock Pacific F_xpe7, No . 3 . Univ. South. Calif. Press, 

13 . 1944a. Polychaetous Annelids, Part V. Eunicea. 

Allan Hancock Pacific Expeditions, Vol . 10, No . 1 . 

Univ. South. Calif . Press, L. A. , Ca . 

14 . 1944b . Polychaetous Annelids from California 

Including the Descriptions of Two New Genera and 

Nine New Species. Polychaetous Annelids, Part VI . 

Paraonidae, Magelonidae, Longosomidae, Ctenodrilidae, 

and Sabellariidae. Allan Hancock Pacific Expeditions, 

Vol . 10, Nos. 2-3. Univ. South. Calif . Press, L. A. ,Ca . 

15 . 1947. Polychaetous Annelids, Part VII . Capitellidae ; Part VIII . Pilargiidae. Allan Hancock Pacific 

Expeditions, Vol . 10, Nos. 4-5. Univ . South. Calif. 

Press, L. A. , Ca . 

16 . 1965 . Deep-water Benthic Polychaetous Annelids Off New England to Bermuda and Other North 

Atlantic Areas . Allan Hancock Foundation Publications 

Occasional Paper No . 28 . Univ. South. Calif . Press, 

L. A. , Ca . 

17 . 1968 . Atlas of the Errantiate Polychaetous 

Annelids from California . Allan Hancock Foundation. 

Univ. South. Calif ., L . A. , Ca . 

18 . 1969 . Atlas of the Sedentariate Polychaetous 

Annelids from California . Allan Hancock Foundation . 

Univ . South. Calif ., L. A. , Ca . 

19 . Hart man, 0. and K. Fauchald_ 1971_ Deep-water Benthic 

Polychaetous Annelids Off New England to Bermuda and 

Other North Atlantic Areas, Part II . Allan Hancock 

Monographs in Marine Biology, No . 6. Univ . South, 

Calif., L. A. , Ca . 

20 . Ima,jima, M. and 0 . Hart ma n. 1964 . The Polychaetous Annelids of Japan, Part I ., Part II . Allan Hancock 

Foundation Publications Occasional Paper No . 26 . 

Univ . South. Calif ., L. A. , Ca. 
555 

Attachment A2 .6 (cont'd) 

21 . Hobson, K. D . and K. Banse . 1981 . Sedentariate and archiannelid polychaetes of British Columbia and Washington. Canadian Bulletin of Fisheries and Aquatic 

Sciences, Bull . 209. 

22 . Hutchings, P. A. , Editor . 1984 . Proceedings of the First 

International Polychaete Conference . Sydney, Australia, July 1983 . Linnean Society of New South Wales . 

23 . Jones, M. L. 1963 . Four New Species of MaAelona (Annelida, Polychaeta) and a Rediscription of Magelona 

longicornis Johnson. American Museum Novitates, 

No_ 2164 . 

24 . Light, W. J. 1978 . "Spionidae (Polychaeta, Annelids)" 

Invertebrates of the San Francisco Bay Estuary System. 

Welton E. Lee, Editor . California Academy of Sciences . 

The Boxwood Press. 

25 . Maciolek, N. J. 1981 . A NEW GENUS AND SPECIES OF SPIONIDAE 

(ANNELIDA:POLYCHAETA) FROM THE NORTH AND SOUTH ATLANTIC. PROC . BIOL . SOC. WASH . 94(l), 1981, pp . 228-239. 

26 . 1985 . A revision of the genus Prionospio 

Malmgren, with special emphasis on species from the 

Atlantic Ocean, and now records of species belonging 

to the genera Apoprionospio Foster and Paraprionospio Caullery (Polychaeta, Annelids, Spionidae) . 

Zoological Journal of the Linnean Society (1985),84 : 

325-383- 

27 . Miura, T. 1980 . Lvmbrineridae (Annelides Polychetes) abyssaux recoltes au cours de campagnes du Centre Oceanologique de Bretagne dens 1'Atlantique et la Mediterranee. Bull . Mus. natn . Hist . nat ., Paris, 4- ser_, 2, 

1980, section A, no 4 : 1019-1057. 

28 . Orensanz, J. M. 1976 . Los Anelidos PoliQuetos de la Provincia Biogaografica Argentina. IX. Poecilochaetidae y 

Cosauridae .Comunicaciones Zoological del Museo de Historia Natural de Montevideo, Vol . X, No . 140 . 

29 . Perkins, T . H. 1979 . LUI`iBRINERIDAE, ARABELLIDAE, AND DORVILLEIDAE (POLYCHAETA), PRINCIPALLY FROM FLORIDA, 

WITH DESCRIPTIONS OF SIX NEW SPECIES. PROC . BIOL . SOC . 

WASH . 92(3), 1979, pp . 415-465. 

30 . 1980a. REVIEW OF SPECIES PREVIOUSLY REFERRED TO 

CERATONEREIS MIRABILIS, AND DESCRIPTIONS OF NEW 

SPECIES OF CERATONEREIS , NEPHTYS , AND GONIADA 

(POLYCHAETA) . PROC . BIOL . SOC. WASH. 93(1), 1980, 

pp . 1-49 . 
556 

Attachment A2 .6 (cont'd) 

31 . Perkins, T . H. 1980b . SYLLIDAE (POLYCHAETA), PRINCIPALLY 

FROM FLORIDA, WITH DESCRIPTIONS OF A NEW GENUS AND 

TWENTY-ONE NEW SPECIES . PROC . BIOL .SOC . WASH . 93(4), 

1980, pp . 1080-1172. 

32 . 1984a. REVISION OF DEMONAX KINBERG, HYPSICOMUS GRUBE, AND NOTAULAX TAUBER, WITH A REVIEW OF 

I'EGALOMMA JOHANSSON FROM FLORIDA (POLYCHAETA: SABELLIDAE ) PROC . BI OL . SOC. WASH . 97(2 ) , 1984, pp . 285-

368 . 

33 . 1984b. NEW SPECIES OF PHYLLODOCIDAE AND 

HESIONIDAE (POI.YCHAETA), PRINCIPALLY FROM FLORIDA_ 

PROL'. BIOL . SOC. WASH . 97(3 ) , 1984, pp . 555-582 . 

34 . Perkins, T . H. and T. Savage . 1975 . A Bibliography and 

Checklist of Polychaetous Annelids of Florida, the 

Gulf of Mexico, and the Caribbean Region_ Florida 

Marine Research Publications, No . 14 . 

35 . Pettibone, M. H. 1962 . NEW SPECIES OF POLYCHAETE WORMS 

(SPIONIDAE ;SPIOPHANES) FROM THE EAST AND WEST COAST 

OF NORTH AMERICA. PROC . BIOL_ SOC. WASH . Vol . 75, 

pp . 77-88. 

36 . Reish, D. J . and K_ Fauchald . 1977_ Essays on Polychaetous 

Annelids in Memory of Dr . Olga Hart ma n. Special 

Publication of the Allan Hancock Foundation . Univ. 

South. Calif-, L. A. , Ca. 

37 . Strelzov, V. E. 1979 . Polychaete Worms of the Family 

Paraonidae Cerruti, 1909 (Polychaeta, Sedentaria) . 

U.S . Department of Commerce, National Technical Information Service, Springfield, Virginia 22161 . 

38 . Uebelacker, J. M. and P . G. Johnson, Editors . 1984 . 

Taxonomic Guide to the Polychaetes of the Northern 

Gulf of Mexico . Final Report to the Minerals Management Service, contract 14-12-001-29091 . Barry A_ Vittor & Associates, Inc., Mobile, Alabama. 

39 . Ushakov, P. V. 1955 . Polychaetes of the Far Eastern Seas of 

the U_ S. S. R. Akademi ya Navk Soyuza Sovetskikh 

Sotsialisticheskikh Respublik_ Keys to the Fauna of 

the U.S .S .R. published by the Zoological Institute of 

the Academy of Sciences of the U.S .S .R., No_ 56 . 

40 . Wainright, S. C. and T . H. Perkins . 1982 . GYT'IIVODORVILLEA 

FLORIDANA, A NEW GENUS AND SPECIES OF DORVILLEIDAE 

(POLYCHAETA) FROM SOUTHEASTERN FLORIDA. PROC . BIOL . 

SOC. WASH . 95(4), 1982, pp . 694-701 . 
557 

Attachment A2 .6 (cont'd) 

41 . Williams, S . J. 1984 . The status ofTerebellides atroemi 

(Polychaata; Trichobranchidae) as a cosmopolitan 

species, based on a worldwide morphological survey, 

including description of new species. Proceedings 

of the First International Polychaete Conference, 

Sydney, edited by P. A. Hutchings, published by the 

Linnean Society of New South Wales, 1984, pp . 118-142. 

42 . Wolf, P . S. 1983 . A REVISION 

FAUCHALD, 1971, AND ITS 

SUBFAMILY OF MALDANIDAE 

SOC. WASH. 96 (2 ) , 1983, 

OF THE BOGUEIDAE HARTf1AN AND 

REDUCTION TO BOGUEINAE, A 

(POLYCHAETA) . PROC_ BIOL . 

pp . 238-249 . 

43 . 1986 . THREE NEW SPECIES OF PILARGIDAE (ANNELIDA :POLYCHAETA) FROM THE EAST COAST OF FLORIDA, 

PUERTO RICO, AND THE GULF OF MEXICO . PROC . BIOL .SOC . 

WASH. 99(3), 1986, pp . 464-471 . 


ATTACHMENT A2 .7 : Annotated List of Abundant Polyehaete Species 
561 

Attachment A2 .7 . 

Lit ocorsa antennata (Pilargidae) Wolf, 1986 . 

Geographic Distribution : Gulf of Mexico (Florida to Texas), 

Puerto Rico . +3* 

Depth hangs : 12-145mf3 . Depth range extension to . 870m 

Feeding Type : Omnivore3B 

Aurospio dibranchiata (Spionidae) Maciolek, 1981 . 

GD_ North and South Atlantic .- 

DR : 300-3600m2 s 

FT : Selective Deposit Feeder3 

Range extension : Gulf of Mexico 

Tachytrypana Sp_ A (Opheliidae) 

6D : Gulf of Mexico 

DR : 291-2540m 

FT : Non-selective Deposit Feeder3 

Spiophanes barkeleyor~ (Spionidae) Pettibone, 1962 . 

GD : North Pacific ; Gulf of Mexicozf, 3e 

D12 : I ntert idal-465m2 t 

FT : Selective Deposit Feeder3 

Depth extension to : 2470m 

Aricidea suacica (Paraonidae) Eliason, 1920 . 

GD_ North Atlantic, North Pacific, Arctic, Gulf of Mexico3a 

DR : 5-2000m4 

FT : Non-selective Deposit Feeder3 

DE : 2959m 

Pholoe "sp_C" (Sigalionidae) Wolf, 1384 . 

GD : Gulf of Mexico (Florida)3B 

DR : 177-189m3a 

FT : Carnivore3e 

RE : Central to western Gulf of Mexico (see fig. ) 

DE : 1007m 

Maldane "op- A" (Maldanidae) Wolf, 1984 . 

GD : Gulf of Mexico (Florida)3e 

DR: 189m38 

FT : Non-selective Deposit Feeder3,3a 

RE : North-central Gulf of Mexico 

DE : 1507m 

Prionosaio ahlarsi (Spionidae) Fauvel, 1928 . 

GD : Widespread (Atlantic, Pacific, Mediterranean)zs 

DR : 524-1700m2 e 

FT : Selective Deposit Feeder3 

RE : Gulf of Mexico 

DE : 320m 
562 

Attachment A2 .7 (cont'd) . 

l.evinsenia gracilis (Paraonidae)(Tauber,1879) 

GD : Cosmopolitan sub-antarctic3e 

DR : 5. 5-3860m3 7 

FT . Non-Selective Deposit Feedc3r3 

Fauveliopsis sp_ B (Fauveliopsidae) 

GD : Gulf of Mexico 

DR : 291-2959m 

FT : Non-selective Deposit Faeder3e 

Sarsonuphis hart manae (Onuphidae) (Kirke$aard, 1980) 

GD : North Atlantic ; Gulf of Mexicoa,3d 

DR : 37-4265me 

FT : Scavenger3,3e 

Prionospio cirrifara (Spionidae) Wiren, 1883 . 

GD : Beaufort Sea; North Atlanticzs 

DR : 11-2900 m2 g 

FT : Selective Deposit Feeder3 

RE : Gulf or Mexico?(previous reports confused) 

Thaw marioni (Cirratulidae) (Saint-Joseph, 1894) 

GD- Widespread (Atlantic, Pacific)ie 

DR : 1-1000m18 

FT : Selective Deposit Feeder3 

RE : Gulf of Mexico 

DE : 2955m 

Lumbrinarides dayi (Lumbrineridae) Perkins, 1979 . 

GD : North Carolina ; Florida; Puerto Rico ; Gulf of Mexico3a 

DR : 20-189m3B 

HT : Carnivore3 

DE : 2945m 

Caratocaphala oculata (Nereidaa) Banse, 1977 . 

GD : North Carolina ; Gulf of Mexico3e 

DR : 15-35m3d 

FT : Omnivore3 

DE : 2904m 

Paralacydonia paradoxes (Lacydoniidae) Fauvel, 1913 . 

GD : Widesproad3e 

DR_ 7-5498m38 

FT : Omriivore3e 

Thar3rx annulosus? (Cirratulidae) Hartman,1965 . 

6D : New England; northeastern South America ; Gulf of 

MBX1C01613B 

DR : 15-4540mi8 

FT : Selective Deposit Feeder3 
563 

Attachment A2 .7 (cont'd) . 

Paramphinoie aaff raysii (Amphinomidae) (McIntosh, 1868) 

GD : North Atlantic ; northeastern South America's 

DR . 36-5304 m1 B 

FT : CarnivorQ3 

RE : Gulf of Mexico (previously reported as P . pulchella) 

Tarebellides stroemi (Trichobranchidae) Sars, 1835 . 

GD : (Previously considered cosmopolitan ; now thought to 

be restricted to the North Atlantic . Specimens in 

this study identified as T- stroemi probably 

should be assigned to T- atlant ia or Tdistinct a)11-1 

Diploci.rrcis capansis (F'labelligeridae) Day, 1961 . 

GD : South Africa ; North Carolina ; Gulf of Mexico3e 

DR : 11-200m3 e 

FT : Non-selective Deposit Feadar3 

DE : 2377m 

Tarebellidas atlantis (Trichobranchidae) Williams, 1984 . 

GD : New England Slope-4-1 

DR : 400-508m41 

FT : Selective Deposit Feeder3e 

RE : Gulf of Mexico 

DE : 1507m 

Terebellidas distincta (Trichobranchidae) Williams, 1984 . 

GD : New England SlopeDR : 466-508mf1 

FT : Selective Deposit Feeder3e 

RE : Gulf of Mexico 

DE : 357-1236m 

Paramphinoie "op. A" (Amphinomidae) Gathof, 1984 . 

GD : Gulf of MexicoDR : 75m3 e 

FT : Carnivore3 

DE : 2540m 

Spiophanas bomb »c (Spionidae) (Claparede, 1870) 

GD- CosmopolitanDR : 0-119m3e 

FT : Selective Deposit Feeder3 

DE : 2904m 

Uiplocirrus "op- A" (Flabelligeridae) Milligan, 1984 . 

GD : Gulf of Mexico3a 

DR : 19-106m38 

FT : Non-selective Deposit Feeder3 

DE : 2900m 
564 

Attachment A2 .7 (cont'd) . 

Cirrophorus lyres (Faraonidae)(Southern, 1914) 

GD : North Atlantic, South Africa, Mediterranean37 

DR . 2-1060m37 

FT : Non-selective Deposit Feeder3 

RE : Gulf of Mexico 

DE : 2945m 

Lumbrineris verri lli (Lumbrineridae) Perkins, 1979 . 

GD : East coast of Florida ; Gulf of Mexico3e 

DR : 3-189m3a 

FT : Carnivore3 

DE : 1507m 

Euchona incolor? (Sabellidae) 

GD : Atlantic and Pacific 

DR : 17. 5-2500m3 e 

FT : Suspension Feeder3 

Hart man, 1965 . 

North America; Gulf of Mexico3e 

Levinsenia uncinata (Paraonidae) (Hartman, 1965) 

GD : Atlantic ; Sea of New Guinea ; central North Pacific37 

DR : 1044-5681m37 

FT : Non-selective Deposit Feeder3 

RE . Gulf of Mexico 

DE : 335m 

Aricides lopezi l opezi (Paraonidae) Berkeley and Berkeley, 

1956 . 

GD : Widespread37 

DR : 6-110Om37 

FT . Non-selective Deposit Feedar3 

DE : 1444m 

Barantolla "sip. A" (Capitellidae) Ewing, 1984 . 

GD : Gulf of Mexico (southern Florida)3e 

DR : 87-91mFT : Hon-selective Deposit Feeder3 

RE : Northwestern and northcentral Gulf of Mexico 

DE : 2540m 

G13ncara aapillosa?_ (Glyceridae) Grube, 1857. 

GD : South Africa ; Peru ; Chile; North Carolina ; Gulf of 

Mexico3e 

DR : Intertidal - 200m3a 

FT : Carnivore3 

DE : 2959m 

Ampharete "sp. A" (Ampharetidae) Uebelacker, 1984 . 

GD : Gulf of Mexico3e 

DR : 10-189m3e 

FT : Selective Deposit Feeder3 

DE : 2118m 
565 

Attachment A2 .7 (cont'd) . 

Syllis (Fhlarsi) cornvta (Syllidae) Rathke, 1843 . 

GD : Cosmopolitan3e 

DR : Intertidal - 2560m36 

FT : Omnivore3,3e 

Aricidea f ragilis (Paraonidae) Webster, 1879 . 

GU : Widespread (Atlantic, Pacific, Mediterranean, Gulf of 

Mexico)37 

DR : 0-54M37 

FT : Non-selective Deposit Feeder3 

DE : 2945m 

Sabidius cornatvs (Paraonidae) (Hartman, 1965) 

GD : Atlantic and PacificDR : 400-3388m37 

FT : Non-selective Deposit Feeder3 

RE : Gulf of Mexico . 

DE : 344m 

Myriochal a haari? (Oweniidae) Malmgren, 1869 . 

GD : Cosmopolitan's 

DR : EurybathiclB 

FT : Suspension Feeder3 

Aricidaa simplex (Paraonidae) Day, 1963 . 

GD : Widespread (Atlantic, Pacific, Gulf of Mexico)3e 

DR : 35-1072m3B 

FT : Non-selective Deposit Feeder3 

DE : 1222m 

Sigambra tentaculata (Pilargidae) (Treadwell, 1941) 

GD : WidespreadDR : Intertidal - 5121m3e 

FT : Omnivore3 

NotomHStus asericanus (Capitellidae) Day, 1973 . 

GD : North Carolina ; Gulf of Mexico3a 

DR : 80-10Om3e 

FT : Non-selective Deposit Eeeder3,3a 

DE : 1440m 

Aulaophamus circinata (Nephtyidae) (Verrill, 1874) 

GD : Boreal and temperate western Atlantic ; Gulf of 

Mexico3a 

DR : 10-189m3a 

FT- Omnivore3,3e 

DE : 2524m 

Notomastus latariceus (Capitellidae) Sars, 11351 . 

GD : Cosmopolitan3e 

DR : Intertidal - 4360m38 

FT . Non-selective Deposit Feeder3,3a 
566 

Attachment A2 .7 (cont'd) . 

S_LsLmis klatti (Pilargidae) (Friedrich, 1951) 

GD : North Sea ; Scotland ; Gulf of Mexico3e 

DR : 11-680m3e 

FT : Omnivore3 

DE : 2540m 

Tachytrypana .iaffreysii (OphQliidae) McIntosh, 1875 . 

GD : North Atlanticlg 

DR : Abyssal ; 78U-4850mis 

FT : Non-selective Deposit Feeder3 

RE : Gulf of Mexico 

DE : 444m 

Notomastus_ hgmipadus (Capitellidao) Hartman, 1945 . 

GD : North Carolina ; California ; Gulf of Mexico3e 

DR: Intertidal - 120m3e 

FT : Non-selective Deposit Feeder3,3e 

DE : 631m 

Heterospio longissisa? (Heterospionidae) Ehlers, 1874 . 

GD : Widespread (North Atlantic ; Mediterranean ; Japan ; Gulf 

of Mexico)3e 

DR : 35-4950m3e 

FT : Selective Deposit Feeder3 

Aglaophamus verrilli (Nephtyidae) (McIntosh, 1£85) 

GD- Temperate and subtropical western Atlantic ; Gulf of 

Mexico3a 

DR : Estuarine - 189m3a 

FT : Omnivore3 

DE : 834m 

s Numbers refer to citations listed in "Polychaete References-" 

ATTACHMENT A2 .8 : Distribution of Abundant Polychaete Species 

(Taxa name is provided on each map, annotational 

notes for each taxa can be found in Attachment 

A2 .70 ) 
569 
570 
571 
572 
573 
574 
575 
576 
577 
578 
579 
580 
581 
582 
583 
584 
585 
586 
587 
588 
589 
590 
591 
592 

ATTACHMENT A2 .9 : Summary of Invertebrate Species Found in the 

Trawl 
595 

Attachment A2 .9 . Summary of invertebrate species found in the 

trawl . (Depth range is in meters .) 

Taxa 

AGTINIARIA 

ZOANTHIOEA 

ACTINAttGE LONGICORNIS 

HORMATHIIDAE 

AMPH I ANTHUS OR ?STEPHEN AIIGE SP 

ISANTHIQAE 

HORMATHIIDAF 'RI-!Ft ! 1 tiRMFR' 

ANT HtAUli A h'tFN IX 

ACT1VOSCYPHM SAGINATA 

ACTINIDAE 

HALCJRIAS PLATUS 

ACT WOSTOI.IDAE 

ALCYONARIA 

CMRYSOGRG M IIG IIS.S Q I I 

AC ANELL A EBUR1E A 

ANTFIOPTILUM GRANDIFLORI" 

ACAhELLA Sam . 

AC AIVEI .L A ARB11SqJl A 

PARANftJRICEA SP. 

SY FT I A EXSERT A 

AcANrHOCORCiA tta 

C ANU lUt.l .LI, . '~F . 

MIIRICEDES HRTA 

PARAMRtCEA PLAt:i:fi" itla .-' 

CALLOGORGIA VERTICLLATA? 

TELESTO W.SP. 

Ar1F'HPOOA 

EPMERA SP. 

AFIONYX SP. 

EPPIER {A sF .1 

YALETTIOPSIS SPA 

TRISCFIIZOSTOMA LONGAt0.STRE? 

EPM Kk 

OEDI(:EROIDES AEM :::tMIJM? 

EIJRONY}{ SP. 

Asc1) i+uT a 

BAIHYS1YttUUkS tR1VEKl3YMwJS 

ASTEROIDEA 

PERSEF'HQNASTER ECHKI.ATUS 

PECTINASTER GRACLIS 

PLIITGN'ASTER INTEItMUJS 

Humber of 

Abundance Stations Depth t'. Ji age 

380 1t 342 - 1135 

204 19 329 - 25(14 

21 3 342 - S 135 

17 1 2400 

16 1 627 

5 423-751 

6 5 329-474 

5 2 751 -2074 

5 1 802 

2 1 342 

1 1 338 

1283 15 351 - 1170 

T1..' 11 61r

. '7 1777 

77 2 2400 - 2bU4 

41 7 479 -2074 

22 3 812- 2401 

13 1 663 

$ 1 464 

s 1 663 

a > 1438 

3 1 4E' 

s S ;sW 

2 1 464 

1 1 6GS 

16 5 834 - 12u3 

11 2 627- $50 

6 e 964 - 1170 

4 4 521 - 1438 

3 2 1170 - 2857 

2 1 106-11, 

2 2 1170-1440 

1 1 62? 

1 1 240Q 

186 15 474 - 774 

158 13 603 - 850 

146 9 850 - ~"t~ : 
596 

Attachment A2 .9 (cont'd) . 

Number u: 

Taxa Abundance Stations Deptf, Range 

Za20ASTER FIAGENS 109 12 612-2400 

NYMPHASTER ARENATUS 102 25 351-1735 

GCIPIICIF''f'f'7r!r ~rrtr ., . , ,., ., `^ F 751 -850 

PLINTH ASTER DENT ATUS 49 19 474- 2504 

DYT ASTER INS ION IS 36 1 2504 

PSEUDARCH naTER. fSF.~A(::ILIS 33 10 Ef - 24ri?? 

ASTROPECTEN AMERICANUS 24 4 329 - 368 

ODONTASTER SET0.SUS 17 10 345 - 842 

BENTHOF'ECTEN SA°it'LLX 10 3 2Cil4 - 2504 

PTERASTER PERSON ATUS 7 1 2074 

BRISWQDAE 7 4 663 - 25U4 

HYMN ASTER MODEST U, 5 4 551 -1203 

GAYhtANit:~~I El_LA SP. 5 2 627-1203 

OOONTItiSTER HISPIt?! K' S 3 342 - 663 

BEHTHOPECTWIDAF 4 1 625 

FENR IC M ART ILL Ak'UM 4 2 1064 - 1170 

DIF'Sr11:A51 U: af' . 4 3 805-2313 

ASTROPECT W ID AE 3 2 338-774 

PSEUDARCH ASi tK yh'. S t 4516 

HYMEN ASTER SR. 3 2 654 - t 17n 

AVTFEhI01DES PIERCEI 3 1 329 

LITUhui ASTEk kuRNOIGRAPNJLUM 2 1 12113 

PLUTONASTER AGASSIZI 2 1 774 

PTER ASTER IDAE 2 1 627 

ASTRCIPECTEN QP. 2 2 625 - 6~~ 

MEDIASTER PEDICELLARl:, 2 1 964 

LITCrNOTASTEF tIYTFF't'1FDIUS 2 2 2074 -2504 

GOUIASTERmAE 2 2 61? - p 

PSILASTER GASSIOPE 2 2 746-850 

SCLERASTERIAS C4NTORTA 1 1 464 

CIRCEASTER AMERICAhW 1 1 615 

DQRASTEl: CQPr:.TE( t NTUS 1 1 351 

TOSIA PARVA 1 1 1313 

NOVOOMIA AHTI.LENSIS 1 1 964 

(HENCASItK MIKI1tfILIS I I iiut 

FIYMENASTERDAE 1 1 621 

CERAMASTER GRENADENSIS 1 1 143 

ASTk!cpErTFN rrtIFITO !S' 1 1 329 

BMALVIA 

PROPS AMIJS& pJM SP. A 304 11 774-1510 

PROPEMMISSUM SR.C 228 18 612-2400 

PROPS Ar1USS IUM D ALL I 161 4 338-1438 

Ah10DOhITIA PHILPPIANA? 117 1 342 

LITHOPHAGA ANT Ill ARUM? 57 2 627 - 1203 
597 

Attachment A2 .9 (cont'd) . 

NixnMr of 

Taxa Abundance Stativrr . Depth Range 

AMYGDALlN1 ROlRlM1? 36 17 345 - 1 f 3c; 

PROREANIUSSAJM SPD 32 3 511 -(312 

PQFrc'"." 13P. A 24 8 618 - 786 

LqHOPSIS AIRITA? 18 3 12u3 - 1 ;1u 

TELL IN A --J-'J : 18 6 751 -2400 

LUC qVHA F SOS A 13 3 342 - 85u 

M ACRODON (BENTH ARC A) ASPERUL A 11 1 663 

LYONS IEt ( k SP. A 9 4 368 - 2S,O4 

PROPS Ah1USS I1JM SP .B 8 1 774 

BAhACIA SP. 8 2 850-1203 

LYONS M SP. A 8 4 551 - C4' 

CLISPIDARIA SP.A 8 5 612 - 629 

MALLETIA SP . O~ 8 1 2400 

TELLWA SP.A 7 4 329 - 632 

MYuNER A P ALIC ISTR I AT A 4 4 464 - 1135 

PARVAMCM^fUM CANCELLATUM 3 1 751 

XYLQPHAGA ABYSSORUM 3 1 625 

POROMY A ROSTR AT A 3 2 345-350 

CUSPiDAf tA 8f' . 3 2 2313-2400 

GALYPTOGENIA PONDEROSA 3 1 $5U 

PROPEAMMAJM HOLhESII 2 1 GEC 

CUSPDARIA OBESA 2 2 345 - t121?? 

CARD101'1YA SP . 2 2 663-1064 

EUL AhEll BR ARCH SP .B 2 1 2400 

CUSPDARIA ROSTRAT A' ' 2 2 329 - 3~1 

hA.ICUL AM IDAE 1 1 812 

LIICINA SP. 1 1 612 

LIMA SF' . t 1 612 

BATHYARCA SP . A 1 1 612 

P ARY M'AJSS NM POIIRT ALES 1 AhNM 1 1 350 

M1ClIL AN A PL ATESS A 1 1 345 

POROMYA SP. 1 1 621 

PkU~Er~ kK,-',.U , "A' . 1 1 603 

I'A.ICUL A CALL ICREDEM A 1 1 342 

VESICCx"t'(A t::Wc!~".' .' . t t K, n 

LMOPSIS SR . 1 1 850 

BRAf`F!H?F`ODA 

ECIWIMMA fWRDA 45 1 9f,4 

PLA1 IDIA MNJMIUIUr_~- ; 4 Y 666 - l51 

CEPHALOPOIDA 

ROSSIA BIILLtSI 12 S 342 - 621 

ALLOPOS(tf; t--U]LLIS 11 7 354-663 

PHOLK)OTEUTHIS ADAMl 7 6 627-1170 
598 

Attachment A2 .9 (cont'd) . 

Nurr` 

Taxa AGun, ~ -rice Stations Depth Ran ; 

BENTFiOCTOPUS JAI~!IAF?1 5 4 464 -5b1 

ROSSIA SP . 4 2 479-812 

STULOTEUTHIS SP. 3 2 551 - 627 

Fl Ekiii: t u;""t L t 3 350 - 603 

SEM ROSS I A EQU AL IS 3 1 329 

OCTOF'ODIDAE 2 1 E?q 

OPISTHOTEUTH IS AG ASR IZ I 2 1 802 

OCTOPUS BURRYI 2 2 329-338 

OCTfFIJ$ SP. 1 1 464 

LOLIGINIDAE 1 1 350 

OCTOPUS JOUBINI 1 1 2313 

QCTrP. 

IS MF It !F' :' I I 1 7X1 Z 

CNZRI'EDM 

TRLASMIS KAEt"l'FERI INAEQUILATERAI 561 14 329 - 1735 

CYPRIS LARVAE 433 2 802 - 173° 

VEhRUCA FLORN)APA 194 6 354-774 

ARMSGALF'FU. Ut `i Pi,iki OR K:Af;"_Efl 1!~! . -~ . L, ~45 - Gt~ 

OCTOLASMt:~ GEf"i'uf+It"klA 4b 1 7b4 

LITOSCALPELLIAM REGINA 26 4 423 - 5~:1 

AMMOSCALPELLUM SEMISCULPTUM 26 6 615 - 1203 

1rERRtICA NEXA 25 1 319 

EISGALPEL1..lM STRATUM 15 4 342-627 

NEFdlN1 ID MJF'l. AX 10 E 774-1510 

LITOSCALPELLUM GIGAFtTElH 7 2 521 -751 

TELOSCALRELLUM ANTILLARlM 7 3 351-621 

MEGALASMA CARWATU'1 4 1 2401 

B AL AMJS IIrPROV t;:U; . 3 2 464-66,4 

AMICiDOSCALF'ELLUM AURIV ill 1I AURNII 3 2 612 - .-f,0 74 

C ATHER WUM ALB ATROSS I AMJM? 3 2 2074 - 2400 

VERRUC A EIIGI .YPT A 1 1 751 

GRNODEA 

DEMOCRMJS BREVIS 226 19 351 -2400 

ATELLCRllrlb IsItiLAhO lUtS 67 1 `r 4?9 . ,^W 

CARYONETRA LISA 11 2 612 - 774 

CARYQP'ETRA ALQPE 7 3 521 -663 

CARYdWTRA SP. 1 1 746 

CdM ATUL ID A 1 1 615 

DECAPODA 

hEMATOCARCMJS ROTUNDUS 339?" 35 521 -1510 

PEI'IAEOPStS SERRATA 2896 11 329-464 

B ATHYPL AX TYPHL A 2297 25 366-1438 

PLESIdIVIKA HOITHUISI 1664 24 366 - 871 
599 

Attachment A2 .9 (cont'd) . 

Nixnbnr of 

Taxa Aburdance Stations Depth Range 

MIMV IDA V AL IDA 1634 28 464 - 1170 

BENTHOCH ASCON SCHM ITT I 1390 18 329-840 

STEREOMASTIS SCULPT A 695 34 603 - 28;~ .' 

lIRllf'T'il:ttll:. .~1C11Lwa,., . :~1~ .:u 

GLYF~F;i-iGRANGOrI r:_ : .iIIS 3,3;. 1 ;~ -11 

t3lYPHOCRANCON ACIILEATA 314 18 751 - 2857 

NEPFIRORStS ACULEATA 243 14 329 - 632 

1-IV171DfSt ~ .^vIS)Ir Cr 233 13 329 - 1135 

oPHILus c;RacLIs Zap av aa~ - 2857 

P AR AP AND ALUS 1af ILL IS I 206 11 329-464 

H1T~tENOPENAEUS ROBUSTUS 184 17 342-654 

BENTFfSICYMJ", F:Ar:TLFTTI 166 20 746 - 2074 

GLYPFIOCRANGOIV AIISPMIA 161 16 423 - 

OER1+Oly QUfXQIIEpENS 125 29 366 -2074 

MLIF! IDOPS IS ROBUST A 108 14 342-746 

P AR AP AGURIIS P ICTUS 102 13 342-2313 

LYRFIDUS BARDII 102 7 329 - 4?3 

MIqVIDOPSIS ERpVACEUS 97 13 474 - 774 

tfTEROCARpUS ORYX 87 7 812-1438 

P AR AP AGURIIS P ILOS IM ANUS 87 12 603-1510 

ROCH IPI I A CR ASS A 82 20 338-1170 

PLESIUNIKn 'iEhJl_nF'ES $1 8 345-423 

PARAAENAEIIS lf1N(+IRRSTRLS 79 4 329 - 3ff+ 

PYRUMAIA AfiACH1+lA 74 11 329- 42S 

PQLYCHELES TYPFILOPS 69 12 338-663 

HYMENOPENAEUS DEEIL1 :=. 69 10 474-1170 

MUVDQPSIS LQN(?MANUS 46 18 3w - 1 2u6 

PtES10NIKA ACANTFIaVOTUS 43 11 464 - 629 

M.AViDOF'SIS SIGS'BEI 41 10 479 1 .~i u 

NEPFROPS 18 POSE A 34 11 603-1170 

GLYPFIOCRAHCON LONGI.EYI 29 12 464 - GS4 

TRICf10PELTARION NQFf[F 28 5 603-786 

PARAPA!wPIQ A! tMMpS 27 7 850- 2504 

ACANTHACARIS CAECA 23 9 521-751 

PACiUtUS ROTUVDiP'IANUS 23 8 342-751 

SYSTELLASF'IS F'CtLUCDA 2? 3 36P-842 

ACANTFEPFIYRA EXMIA 21 2 117u - I': ft' . 

ETHUSA MICROPFiTHALMA 21 6 329-706 

PALICL1S GRACILIS 19 10 319 

MIAVDOPSIS I"POLITA 15 9 345 - 663 

MUFlOOPSIS BAHA1yEFlSIS 14 2 423-663 

CATAPAGtIRODES MICROPS 14 7 751-1440 

PRIONOCRMIOOIV PECTINATA 14 10 521-1203 

MUVDOPSIS SRMVOSA 13 4 751 - 828 

CYhiUNG~°U~ N .SP .A 1 3 8 612-850 
600 

Attachment A2 .9 (cont'd) . 

FkxnMr of 

Taxa Abundance Stations Depth Range 

h1UV DOPS IS ABBREV I AT A 12 6 774-1170 

TH AL ASSOFL AX ANGUST A 12 5 338-464 

ROCHINIA lJMBONATA 12 8 612 - 8`t1 

ACANTNFPHVaA aPHaT,4 12 6 751 - 117 

PHYLL AD IORFIYFCHUS N. SP . 11 1 663 

MIbVIDA MICROPHTHALMA 11 5 964-2401 

to 4 85U - 1510 

AC AhITIiQC ARf'lIS ALEX ANUK I 10 4 329 - 368 

PLESIl1i'l.kkLi1.+ i,iu n i li,r '~ L Lu i4* 

HETEROCARPtlS ENSIFER 9 5 342 - F,$ 

h1M1DOP515 ALAMt~r^ E ,-, 4'..' ~~r~' 

R IqHARD INA SP ANIC pVCT A 8 4 551 -774 

IVEPFfitOPSIS AGASSIZI $ 7 fi t ;' - i 

MIJN DOPS IS POL IT A 8 7 342 - 1170 

NL1f+l lG A C:UFl':.~ t t~ :Ll f~. 7 3 612-62 1-1 

BEHTNESIGYNAiS UREUSIRN)ESCENS 7 1 2504 

9AT F ;'i I' :tiLi,lf-lUi~i.li. n ;kl:ki, i a-' iL1-i  7 4 850 - 1203 

GASTROPTYCHL)S SRNIFER 7 4 345-2401 

FLESIOPENAEUS EDYARDSIANU^ 7 6 621 - 1135 

PS AL IGOF'lK; L AhEC1Uki 7 6 618-786 

FIEMATOCARC9VUS ENSFER 6 3 2074- 2504 

BATRY"!ECTE° r! PER9A 6 4 338- 369 

HOr10LODROMI A P AR ADOX A 5 3 627 -751 

RODiIVIA HYSTRC{ 5 3 351 - 625 

POLYCFELES CRICIFER 5 3 ?51 - 113~~ 

LiTHODES AGASSIZII 5 4 774- 1438 

MUIV ID A FORCEPS 5 3 329-368 

M1AN1DOPSIS N. SR. 4 2 663 - 151 

PANADALIDAE 4 2 479 -521 

FUFtGFf r! ~. ~ 4 T'! "Fww'"71' 4 1 350 

(iERY0F1 FEPNERI 4 2 345 - 3501 

AXIORSIS SP.A 4 4 366 -62V 

HYhENOPENAEUS AM-101 x.143 4 4 1135-2504 

FETEROCARPIIS LAEVIS 3 1 663 

AR ISTEOMORPH A FOL I ACE A 3 2 464 - 5:e 1 

THALIMASTO(:HELES ZALEIICUS 3 2 842 - 871 

CYMOb101`1US SF' .B. 3 3 61 2 - t .'.27 

PAR APAGURUS SP. 3 3 464 - BU2 

PORCELLANA SIANA 3 2 329-338 

ElI'4U'tVr+ ~~'7A 2 1 663 

CALLINECTES SMILIS 2 1 368 

rLIumoPsIS; sPIroaxATa 2 1 871 

pORHYhiCNU~ ~I tu-rami c 2 62( - bbs 

MUNIDOPSIS RAMAHTAYLCIf:KE 2 2 612-62-1 

ETHU.SA SP. 2 2 350-368 
601 

Attachment A2 .9 (cont'd) . 

Number of 

Taxa Abundance Stations Depth Range 

PROCESS A PROFUND A 2 2 354 - 368 

MEl ki~F~APJu0t~4 _l ;-,!-:iWETt AGASSRII 2 1 964 

MUIVIDA SP . 2 2 366-621 

TRIZOCARCpVIJS TACITUS 2 2 345-366 

POIVTOC AR IS C AR IBB AEUS Z 2 329 - ~: -4E, 

HOMRf tt?RE 1 1 1203 

FOMOLOGEIVUS ROSTRATUS 1 1 1203 

BYTHOC AR IS SR. A 1 1 1135 

RCIBERTSELL A MYSTIC A t 1 479 

MUNO0PSIS ?ARCS 1 1 6G? 

h1lqVIDQPSIS SEk'F'RTIFRQP6 1 1 6E' 

TOh10P AGURIJS COICER I 1 1 368 

"DA EVERMANI 1 1 3b$ 

PLESIOIVKA EDWARDSiI 1 1 3cwl 

TR120CARCINUS SR. 1 1 350 

HrP4L`(T fDAE 1 1 350 

MUVDA MILES 1 1 345 

(iWRLACDAE 1 1 345 

POLYqiELES VALDUS 1 1 2Q't4 

HEMPENAEUS CARPEHTERI 1 1 2074 

PAR AP AGURIIS MAUS 1 1 2504 

ACANTFEPHYRA MK:ROPHTHALMA 1 1 2504 

PERICLNHENES PANDIaIsIIS 1 1 632 

SF'OHIGICOLOk?ES N.SF . 1 1 1170 

RATHYPA! AFMfW 1 r, TFYANb f f 1170 

PLESl01V1CA LONGPES 1 1 366 

STEI'JOCIONQPS SFWIMANA 1 1 342 

scnLaRUS cHACEi > > 342 

1-IVNIL}1"Y IRF~Ar" f7 

P AR AP AwllFcliS b k.k la i ATUS 1 1 850 

PAGIRISTES SP. 1 1 329 

S0.Efi(71,'Elt A MElW 1NA 1 1 324 

COLLODES LEPTOCNELES 1 1 329 

CHACELLIiS FILIFORMIS 1 1 329 

ECFIINODEA 

PLESIODIADEMA ANTI.LARLIM 70 6 751-1440 

BttSSOPSIS ALTA 19 3 324- 611-1 

HOMOI. AMP AS FR AG IL IS 17 6 615-629 

BRISSOPSIS SP . 13 1 329 

ACESTE BELLIDFERA 7 3 615 - 

ECH1y0CYMHUS ukMDPORUS 6 4 612 - 629 

PHOWOSOM APL ACErlT A 6 3 1438 - 131 0 

POOOCDARIS SCU0.PTA 4 1 6U1 

BRISSOPSIS ATLANT IC A 2 1 329 
602 

Attachment A2 .9 (cont'd) . 

Number of 

Taxa Abundance Stations Depth Range 

CDARIS RIIGQSA 1 1 663 

ECHMIS TYLODES 1 1 1735 

ECHVOCYAMIJS P?r!,^RQSTOMUS 1 1 1064 

ASP Mr) IanFMA .)ArrlRYI 1 1 1064 

GASTROPODA 

YS SIILCATA 45 3 2074 - 25(14 

BUGC NVUM C ANET AE 29 8 61 2 - 964 

LEUGfKiYku~4n 1 EN'-' ; . . ?~ ~ 774 - 1510 

SQLARIELLA LAMELLOSA 25 6 612-629 

HYALORlSIA QALEA 21 4 338 - 774 

XEh10PFI0RA LOPIGLEYI 17 9 338 - 627 

ADMETE SP. 16 1 627 

LELICOSYR AVX VERB ILL I 7 3 774 - 12M 

CDR INN AETURR IS INC IL IS 7 3 612 - 625 

PLELH20BRANCFIAEA SP. 6 2 464 - 479 

CORMINHEl i II~'"~',': ; Lt:IJi :1 ._l ~ Al P, 6 4 612 - 1203 

TROCHIDAE N. SP. 6 3 612-850 

GAZA SUPERB A 5 4 551 -774 

TROCHIDAE 5 2 615-627 

CERITHIUt°S Iy . SP. 5 1 345 

CANTRAMffA M.SP . 4 1 964 

SC APH AMER CL AY IS 4 4 603-1170 

(:ORALLIOPHILA RICHARDI 3 1 6E.3 

t3YMP10BEl A SP. 3 2 964-1510 

PHILENE ALBA 3 1 329 

(X]CCIIL IN A SP. 2 2 521-1203 

(3Et"IMUL A PER ISCEL Yi h 2 1 368 

SOL AR Fl 1 A PQ 1?T A1 F^. f 7 ? R47 - 1 MA 

MIl RA f3AwiUl 2 1 615 

(ORMIAETL~tRIS SP. 2 2 871 - 1 170 

SC APHELL A DLB I A 2 2 329 - 4 

BEMTHOBIA TRYONI 1 1 751 

SC APFIELL A GOILD I AN A t 1 464 

FMSI.A2ELLA SP. 1 1 663 

SC1LARfEl.I_A TIAPF, 1 1 812 

PLELIROBRANCHDAE 1 1 627 

FECT IIODOhIT A ARCU AT A 1 1 627 

,pRpTRppIS SP . 1 1 618 

pRt.L10LA SP. 1 1 612 

PlElIROTCIMELt A SP. 1 1 62: 

14ODRLLIA SP. 8 1 1 345 

IhIODRILLIA SP. A 1 1 345 
603 

Attachment A2 .9 (cont'd) . 

Nun-~t~i 

1 axa Abundance Stations f?epth range 

QQCORYS BARTCHI 1 1 802 

ARMIN A SP. 1 1 346 

TRoPI .R"J Arl 7 !' :",Tt !^ 1 1 117U 

G AZ A F ISCHEI: t 1 1 828 

GYLICHNKJJM SP ATHA 1 1 240C-1 

SC APH+SNDER BATHYhWHILUS 1 1 850 

HOI.OTHlIRO DE A 

MESOTHUR I A L ACTS A 184 17 474 - 1170 

BEMTFIOOYTES TYF'IGA 101 3 2074 -2504 

ENYF'Pt,'A^TES SP. 74 4 621 - 2504 

MAP AD I A B ARBQIIR I 44 15 5?q- 2400 

MOLPADIA MUSCiJLUS 38 9 479 - 2400 

SPH AEROTHLJR M T AL IShi AN I 26 7 350 - 625 

PSYCHROPOTES DEPRESS A 23 2 2U'=~ ~ ": :_~r I .1 

BATIiYPLQTE-z NATAMS? 12 3 3EE - 25U4 

MOLPADIA BLAKEI 10 6 551 - 2U'l4 

BENTHOOYTES LINGUA 8 3 2074- 2504 

MMHIGYhW" BAHAh11VENSIS 7 4 464 -663 

PSEIK?OST K'.lW1PUS SR 7 2 1064- 2074 

ECHWOqJClIly11S HlSf'IUA 7 5 618 -1440 

PSEUDOST ICFIUFl1.ti SF. h 6 2 1438-2316 

PROTAh11CYRA Sf' . 6 2 2400 - ='t04 

BENIFIWYTES SANGUINLEWTA 4 1 1203 

PSEUDOSTIC:HOP! l,q',qP . R 3 1 2311 

MOLPADIA AGASSIZI? 1 1 551 

MOI.PADIA SP. 1 1 8'50 

pE IM A VALIDlIM 1 1 1510 

MQIPADIA CUBAFIA 1 1 338 

HYDROZOA 

IYTQCARF'IR Ct^F 5' 345 - 871 

EUDENDRIUM EXMIUM 3 2 618 - 871 

(XAppCARpUS TENUIS 3 2 618-629 

CIADOCARFIM fILt.~;i_If1 ;tl : . 1 E29 

ACRYPTOL AR A CQNFERT R 1 1 464 

LOAM A 

AEGA SP .285 1001 7 366 - 28.57 

B ATHYHON0.MS GtG AhITEUS 18 11 479 - 1170 

AEGA SF' .29 1 6 4 625 - 1 14 

ROCKLA gp.29G 5 5 354-625 

CROIMIA SP282 4 2 4?9 - 677 

AEGA SP299 1 1 663 

MEINERTIA SP. 287 1 1 621 
604 

Attachment A2 .9 (cont'd) . 

Taxa 

MISC . ANTHOZOA 

ANTIPATHARIA SP.B 

ANTIPATHARIA SP.A 

ANT IPATHARIA SP.C 

CERIANTHIDAE 

NEP'IERTE A 

NEMERTEA SP .L 

MEMERTEA 

OPH IURO IDE A 

OPHIERPN.IS ADSPER:;lt. 

AMFHUPI~: f F1AMA!! 

OPHIOCAMAX FASCICUI.ATA 

BATHYPECTMJRA HERDS 

QPFI NJR A SK A 

OF'FIIOCHITOIV GRAINS 

QPHqLIPUS AGASSIZII 

AMPH IOPH IUR A SCUPIT IL IS 

OPH IOPR I1JM PERM IXTUM 

OPHIQSPHALMA ARMK3fftlR`i 

OPN{Ct il:~lllll LEI'[ (lElF.'Pd.'l ill.x") 

OPHIACMI7HA SY.ti 

OPHIn(".!~(ur,~'i'' ; ~~nnK"I rTalca 

ORHIACANTHA 1P.tti 

OPNIACAN I HA VALFNC IENMVtbI 

JWEMLE OPHIURINAE 

OPH104`11151UM ACUFERlN1 

OPNAJRA FALCFERA 

H IST AC RICA ADIIF't IL 4 ( A 

OPHIOCHROMIDAE 

AhiPHIURA SEMERMIS 

OPHIACAHTHA ECHMVULAT A 

HOMALOPHIURA qVQRfVATA 

AMPMIOPLUS DALEUS 

OPHIOMUSIUM TESTlDO 

AMPHORA SP.A (CF . FLEMIOSA) 

AMPHIOPLUS WCISUS 

AMPHIORI.US SPB 

OPH 1OM1.IS pJM ELI 

Ar1PHIURA SP. 

QPH IERNUS V ALL IMICOL A 

oPHu_IPa ar_FawATA 

AwTtIL~~~ A INccL F I 

Nun-&r of 

Abundance Stations Depth Range 

35 1 663 

8 1 663 

1 1 663 

1 1 E27 

1 1 1203 

1 1 618 

1 087 19 423-850 

959 3 612 - ?504 

857 3 6U::~ - 14c: 

229 18 366 - 20-4,U 

209 11 563 - 1440 

173 17 423-1170 

153 16 329 - 850 

123 11 618-1 440 

113 15 464-1170 

92 10 612 - 2504 

43 9 612-812 

28 7 612-629 

24 2 621 - 850 

21 1 6E.'" 

19 9 612-1170 

17 3 345 - 354 

17 7 612-tiiZ 

17 6 627 - 1510 

12 4 368 - 62'! 

7 3 612-8.4 

7 6 612-120 

7 3 850- 2504 

6 1 2400 

4 4 627- 774 

4 4 663 - 1064 

2 2 q?9 - GXT 

2 1 346 

2 1 621 

2 2 2074- 2400 

2 1 329 

1 1 464 
605 

Attachment A2 .9 (cont'd) . 

NuritLQ: of 

Taxi Abundance Stations Depth Range 

AMPHIOPLUS CLINEATUS? 1 1 6E3 

OPHIACTUS ABYSSICOI A? 1 1 663 

OPHIACATHA SP.G 1 1 663 

OPFIIOr1YXA TUMIDA 1 1 6E,F 

BAS LuMBRIcus? 1 1 368 

pus SP. A 1 1 sso 

OPHIAMBIX SP. 1 1 627 

AMPH(TARSIlS MIRABILIS 1 1 615 

QFHNl?rPFlF! t A ti"M b f 1 621 

ASTEROSCHEMA TENUE? 1 1 342 

IN I I IL~I' IC'~= Il M" S--. r. 1 1 2T0\.1 

OP+rocREas sRU+ULL ;u> 1 I $5u 

POGONOPHOR A /VEST MENT IFER A 

G Al ATHE AL " N. SP. 

LAI`1ELL1&tACHM SP . 

F'OLYCHAETA 

NYALtPMCV, TMICOLA 

EUNItE NORVEGICA 

ISOLDA PIkCFIELLA 

MIX Iil1BtECHTI 

ElqNUE SE' .A 

S ARSOMIPH IS H ARTM AN AE 

EUN ICE COFKL01'1ER AlVG 

EHLERSLEANIRA IhlCISA 

APMODITA SP.A 

MELNNA CRISTATA 

KEFERSTEINIA CRRATA 

MYRICICHELE HEERI? 

SC IOhELLA SP . A 

hIEREIS (ff AMl HES) SP.A 

RH AMPHOBR ACH AJr1 Sr' . w 

MALDANDAE 

UJI'1ER q'iCF` ' . G'C1M`tF ~% 

TEREBELLA EHtEN6Ef:W ' 

ANELMVA ABYSSALIS 

LA-rMw" bF. 

SYLLIS (TYPOSYLLIS) ALTERNATA? 

MYRIOGLOBIJLA SP. A 

SP 1001 AETOPTERUS COST ARUM 

HAPLOSYLLIS SPONGICOIA 

AhIPHICTEIS SP. A 

TEfiELtLLIUAI. 

POlYNOIDAE 

122 1 464 

5 1 464 

219 17 612- 15111 

65 13 350 - 1064 

27 8 612 - 663 

23 3 368 - 663 

14 474 - 1510 

13 2 834- 2074 

11 2 342 - 464 

10 7 4?9 

9 4 612-627 

8 2 612-621 

7 1 6E.? 

z s 611 - Win 

7 1 2400 

6 2 663-1203 

6 3 627 - 850 

6 3 850 - 2404 

5 3 368 - 464 

5 3 342 - 354 

4 1 621 

3 t 36tt 

3 2 368-464 

3 1 618 

3 2 615-6425 

3 2 342-663 

3 1 2400 

3 3 351-1 440 

2 2 1135-1203 
606 

Attachment A2 .9 (cont'd) . 

Number of 

Taxa Abundance Stations Depth Flange 

EINli:l i au1.w. r, 2 1 464 

AMPH ARET ID AE GENUS F 2 2 629-834 

GHaETO2QNE "SP .A" 2 1 627 

PARA(yWA?RA 2 618- 621 

ARCTONOE SP. A 2 2 479-618 

M1PIi ARET ID AE 2 2 461 G f 

MARRHYSIA SP . A 2 1 612 

LEANfRA HYSTRICUS 2 2 612-871 

GYPT IS SP B 2 2 1064 - S 75;o 

TEREBELLIDES STROEMI 2 2 474-627 

Cil.OEIA VIRDIS Z 2 632 - Ed, :~ 

LAfTMOIVI~.E RFNTHAI IsNA? 2 2 346 -350 

HY AL qVOEC M JUYEN AL IS 2 2 621 -774 

HARMOTHQE SP. 2 2 342 - 464 

S TS P AI If:IRR ANf:HI IS 2 1 24fif1 

ASYCNIS UU 1 UI? 2 2 621 - SW 

ElW2YTH0E SP. A 1 1 12W 

NEPHTYS SP. A 1 1 1203 

MARPFi'i" fp, 3-F .B 1 1 1135 

f,GL AOPH AM= C itC IN AT A 1 1 479 

Ll1MBRINERIS VERRILLI 1 1 464 

CERATOCEPHhi.E OCllIATA 1 1 46" 

SYLL IS (TYPOSYLL IS) "SP .C" 1 1 46=: 

IVEOLEFRIA "SP .A" 1 1 464 

DASYBRANCFUJS LUMULATUS 1 1 6E3 

MET AVERML IA SP. 1 1 663 

LYS ID IGF r: VTn A 1 1 663 

RRIQNOSPIO (MpVOSPIC1) "SP.A" 1 1 521 

TEREBELLA SP.A 1 1 368 

PISTA i :kISTATA 1 1 834 

PLiI ELL A ARM AT A 1 1 627 

HEREIS (PEAIyTFES) Sl1CC0+lEA 1 1 612 

NOTFRIA TEXTOR 1 1 612 

L~INERDAE 1 1 351 

PARCU'JLR't ~ "' ^P. Pr f 1 1 20?4 

SYFELMIS KLATT I 1 1 621 

MALDANI[>AE GENU'~, l? 1 1 621 

M ALD AN ID AE GENUS C 1 1 621 

MALDAFE GLEBFEX 1 1 621 

FIOB.SOIVIA SP . 1 1 621 

Ar1PWAPFTPAE CM1i; 1 f 621 

RFIODNE SPB 1 1 621 

STHEI'n1f.Fr't^ ^P . A 1 1 828 

APFIROp IT ID AE 1 1 654 

SYLLIS (TYPOSYLL IS) PRA HER A 1 1 342 
607 

Attachment A2 .9 (cont'd) . 

Taxa 

OPHf! WO cP p 

PQR HER A 

POLYMAGTIA and 

THENEA SP. 

DEcMAC`FI f P!!F 

THENE A SP. A 

TETHYA SP. 

HYALONEMA SP. 

P ACH ASTRELL A SP . 

REGADRELLA SP. 

TETHY p SP -B 

DESMACELLA SP. 

HYALONEMAT IliAE 

XESTOSPOIVGIA SP . 

TETILLA SP. A 

HEXA!'TKLLIDr1 

PACHASTRELLA ABYSSI? 

STYLOCQRDYLA SP. 

APFROCALLISTES %" . 

cHOvaRosia SP . 

THENEA SP.D 

LELICCIpS AC RS ID AF 

CLATHRIIDAE 

CAULOPFIACDAE 

H AL Ii:HOMDE; E IL, HE 

THENE A SPB 

Mumhar of 

AbUnd3fii~~, Stations Depth Range 

1 1 1440 

6S? . 21 329 -2857 

651 8 612-1203 

596 1 1203 

309 4 621 - 1`d i~ 

282 9 612 - 1 lU3 

84 151 4i'3- 1bSu 

24 1 663 

18 8 834 - 231 3 

15 2 621 -1170 

8 2 368-663 

6 3 812-1135 

5 1 663 

3 2 1413-2400 

5 3 4E4 -25U4 

3 1 66? 

3 2 35U - 805 

2 2 EE2, - ; ;. 

2 1 805 

2 2 805 - 2.61 6 

2 1 342 

1 1 751 

1 1 464 

1 1 663 

1 1 621 

FDA 

PALLEhtOPSfS TYf!Fr1 AMI f+~F . : q ,~,, ~, 

PALLENOPStS SCOPARIA 

SCAF'HOF'ODA 

LAEVDENTALIlM CALLPEPLl1M 

DENTAL IUM PERLOhIGIIM 

pEpa1 pLq,U`1(;u.,CI.IN1GUdl: CUM 

PIJLSELLUM PRESSUM 

S PNOIVODENT AL I ID AE 

CADt1lUS SP. 

SCLEfi ACT III A 

1064 

5 2 345-368 

5 4 842-2400 

2 2 834 - P 4' 

1 1 774 

1 1 850 

1 1 842 

DELTOCYATHUS IT RIIi_US 80 21 551 - 1,2'03 

LC1F'FtELtA t'ROLFERA 48 1 663 

STEPHAhlOCYATHJS DIADEMA 45 6 964 - 1510 

CARYQPFtYLLIA AMBROSIA CARIBBEAFIA 28 9 774 - 1203 
608 

Attachment A2 .9 (cont'd) . 

Number of 

Taxa Abund.J :~ Stations Depth Flange 

CARYOPHYL! !A BERTEI:IA6A 8 2 464-751 

FL +4BELLUM MOSELEYI S 5 612 - t .'y 

DEPiDROPHYLLIA ALTERNATA 4 1 368 

STOMATOPODA 

SQU ILL A EDENT A7 A 5 2 329-346 

ATTACHMENT A2 .10 : Summary of Fish Species Found in Trawl 
611 

Atachment A2 .10 . Summary of fish species found in the trawl . 

Nux+nbar of 

Taxa Abundance Stations Depth Range 

D IBR AFB ATL ANT ICUS 575 32 351 - 964 

UROPHYCIS CIRRATUS 380 19 329-632 

BEMMRF''; ~"Rtf 366 16 338- 654 

SYMAPFIOBRANCHIJS ORFf;ONI 352 30 479 - 1510 

CHLOROPHTHALMUS AGASSRI 305 1 3 329-479 

HYh1EWlK:F.~'iini.ua i i AL KUS 25.? In 7-1 f.1 - .591 

PEZIJM I A AEOJ AL IS 236 27 474 - 1438 

COELORAVCHIJS IXIELORfiIMCLtS 205 16 329 ~- C-63 

G ADOfHIJS LONG F L IS 191 11 751 - 1203 

COELOR AVCF~IS C AR BR AEUS 177 8 329-786 

EPIGONIJS PANDIOFIIS 1?5 17 329 - 828 

POECILOPSETT A BF ail ~ 155 13 329 - SOS 

CHAUNAX PICTUS 150 21 350 -840 

PERISTEDiON GRAYAE 128 17 329 - 117u 

L AEMOFIEM A B ARB ATLLUM 116 14 423 - 663 

M AL ACACEPH ALUS OCC VENT AL IS 111 19 329-663 

NERUICCIUS ALB", 107 21 329-812 

B ATFIYG ADUS M ACROPS 106 17 551-1170 

B ATHYG ADDS MEL ANOBk MIt>0.1S 97 16 603-1064 

SET ARCHES GUENTHER I 94 10 329 - 464 . 

D IPL AC ANTFIOPOM A SP . 94 17 464 - 840 

DICROLENE SP. 93 20 746 - 1438 

hqNOMITppUS SP. 87 16 479-1170 

84 13 746 - 120S 

ETMOPTERUS SUAlLTZI 62 13 464-774 

STEPHANOBERYX MOIVAE 53 5 1064 - 1510 

P AR ASllU IS 1 FtIICULEPI 1 h 52 12 329 - 1170 

COkYPmAEFqDES NEXICANIJS 50 16 751- 2074 

FffZIIMM CYRAPlO 47 11 615-1170 

PSEUDOPIi1CHTHYS LATERODORSALIS 41 11 603 -871 

STEINDACHP1Ef:tA AC;GLP1IEA 38 7 329 - 41 

HAL0.SAUtUS GUEMTHERI 34 10 731 - 117U 

BROSMICLUS MERBIS 33 13 342 - 871 

PONT IMUS Li 114G 1 '14'114 k_ ; 30 4 329- 351 

CORYPHAEMODES COLON 29 15 60.3-964 

SYFIAGROPS BELLA 26 1 u :. : .. _ 

HEM ANTN i A^a V IV ANUS 25 9 329 -368 

BARATHRONUS BICOLOR 23 10 603 - 173b 

GIIRCFS El l A S pVU~:t- K : : IC ANUS 21 13 342- 1135 

EiATHYP1EM1151,'UAM ihL15 1$ S 964-1735 

UROPFIYCIS FLORIIiitiNUS 17 6 329-423 

HOPLOSTETHUS OCCIDENT ALIS 17 10 319- 4;4 

ALDROYMIDIA ORACLtS 17 3 1064- 1733 

VEFIEF I( : A F'I:OCER A 16 7 964- 2857 

ETMOPTERUS GRACILISPINIS 16 2 474 - 632 
612 

Attachment A2 .10~(cont'd) . 

IWxrbor of 

Taxa Abundance Stations Ueptt, oqe 

BATHYGADUS FAYtjWS 1E 4 1064 - 1203 

NE2UMIA ATLANTICk 15 7 612-812 

SYPV,^^'J^S SF'Mt^"A 14 5 342-354 

Pubic iw ;t1, ~u' , i : : v 31y - 423 

GNATHAGNUS EGREGIUS 13 7 329 - 423 

ALDROVANDIA AFFINIS 13 4 f O&1 - 1 77F 

SYMPHUtUS MARGHVATUS 12 9 351-663 

ETMOPTEkl1S VqZENS 12 3 464 - 479 

NETT ASTQM A MEL ANUR A t 1 8 61S - 850 

FAGi:lOLELl 11 5 464-632 

ARDENT IN A STR I AT A 11 6 329-368 

ACROMYCTFR W IpTt PP A.) rQ 1 t z or,4 - i 17i l 

YARELLA BL ACKFQRDI 10 6 603-850 

hIEOSCOPELUS MRCR!lt FPIDOTUS 10 4 621 - S50 

IPFIOPS MURRAY I 10 2 113 - 141 

SFf-~Ar)EMACURIIS GRENADA" 9 2 10164 - 12 07; 

h1EZUMIA SUILLA 8 4 $28 - 1170 

LOPH llt::; G K; f Oi~i!'HYSUS 8 6 350 - 479 

GADQhKJS ARCUATUS 7 6 ri1 - 11"tu 

EP f ATRE I U:. :~F'RliIuu:t 7 6 474-714 

CUMQC AR A SP. 7 2 1170-1203 

BEh16ROPS ANATIR0.STRIS 7 5 345 - 11'0- 

TRACHOIVURUS VLLQSI IS 6 3 746 - 117G 

PERISTEDIOM MWIA1 UM 6 4 329-423 

MnLACf1PA~ IA FAR ~'=lWA f, '? 774-834 

LYCENCHELYS BULL ISI 6 5 629, - 11 ; 5 

EF~C~C1Nlr,-, Rt'.!'.IDENTALIS 6 3 368-850 

ARIOMMA SP. 6 2 751 - 1135 

POLY AGANTFIQNOTt1S I"ERRETTI 5 3 842-1135 

hEZLIMIA SP. 5 5 464 - f CK' 

LOPH IQDES tH[11VQD I 5 2 366-621 

HYDRIILAGU5 SY. S 4 551 -834 

DYS!'1MMlMA POK;DSt 5 4 4E4 - 6?7 

CRla2RAJA RLIGOSA S 4 521 -786 

BATHYPTEROIS GALLATat 5 2 2074 - 25114 

f9rRIJ 1 VRW rRVf VIWWW--1 J J 61 v - i 1. --J 

l ALOGADUS MOD IF IC ATUS 4 2 1170-1203 

NEZUMIA SCLERORHYNCHUS 4 1 474 

M AL ACOR AJ A Plp2PUR IVEAITR AL I .S 4 3 802 - 1438 

LUCinERn~' T xrn^;- . mr0 r P .? ? 52` - 774 

G ADOMIJS D ISP AR 4 2 663-871 

EXECHCIDONTES DAIDALEUS 4 3 479-834 

CRUR IR AJ A C ADEN AT I 4 1 34,'r 

CORYPHAENODES MACROCEPHALUS 4 3 1064 -2504 

CAT AETYX SP . 4 4 964-1510 
613 

Attachment A2 .10 (cont'd) . 

Nurr~Gf t , r 

Taxa Abundance Stations Deptt~ Range 

RATNYPTFPnPQ NpW%Fcc'Fl1q 4 4 627 - 1170 

BAT4`; f' t ER01S PHENAX 4 1 1170 

BATFCfF'T('r^'^ Err, ~t'rr'1 4 2 625-834 

APR ISTRURUS PAR4'V tt:NI-IS 4 4 751 -871 

TR ACHYSCORP I A Clot ISTUL AT A 3 2 612-615 

RAJA GARMAMt 3 3 329-368 

POLYNIETME CORYTHAEOLA 3 2 3:8 - 34.' 

OPHICHTHUS SP . 3 2 464-774 

NOT AC+4N'IHUa"C1JI<<< 3 3 629-7tk1 

H ALOS AIRUS OVEN I I 3 3 603 - 746 

EP * l A1 Kt t u5 wiilkk s t 464 

DPLOPHOS MADERFh1qIG 3 1 346 

CONGRIDAE 3 3 551-632 

BATFfYGA[!! K' (7 . 3 2 663- '74 

ACANTFOBATUS FOLIR0.STRIS 3 3 351 - 474 

XYEC ACYfi A MYERS I 2 1 1170 

SYNAPHOBRAIrCi;U ; SP. 2 2 621 - 14^,tf 

SCYLIORFI", RETIFER ? 1 354 

(+E21Ih11A LOIGEBARBATUS 2 1 1735 

MEL ANn*k1q TIPMAYFRI 2 2 850 - 1203 

MACkOi'I ; .4E 2 2 464 - 654 

UICIOBROTULA SP. 2 2 632-812 

LEPTODERM A MACROFS 2 Z 840-871 

Ha'LOSTETFUS MEDITERRAhEUS 2 2 423-464 

EPIGONUS MACROS 2 1 828 

EP DEMT MATUS 2 2 464-521 

tXIELORqVCi1JS SP. 2 2 871 - 117C' 

COP fp " r- nl''.t'. A 9 1 7 5 1 

BATOR VICpWJ,S Z 2 850 - 1135 

BATHYCLl1Pf A SP. 2 2 346-521 

BASSOZETUS SP . 2 1 250s 

AGMRHOMlfS ARI'1ATU5 2 1 2504 

VENTRFOSSA MACROPOGON 1 1 521 

STUh11AS AFF WIS 1 1 632 

SCIADONUS PEDICELLAkb 1 1 62? 

SCIADONIIS C.F . GALATHEA 1 1 629 

SACCOGASTER STAIGIER 1 1 354 

RNYACTES NAsU'(UN f f 1170 

RAJIDAE BIGELOIdI 1 1 751 

RAJ r; LM"1TtGfft~_'',A 1 1 342 

PRIOIVOTUS STEARHSI 1 1 324 

PENOPLIS SP. 1 1 1135 

QPHICHTNl1S (:it1IE+iTFER 1 1 654 

hIEZII'11 A BUECtpl tS 1 1 632 

NE08YTHITqVAE 1 1 1064 
614 

Attachment A2 .10 (cont'd) . 

Number of 

Taxa Abundance StatXa, - Depth Range 

NEOBYTHRES SF' . S 1 475 

NEMICHTHYDAE 1 1 2074 

h1YXINE GLUTIV0.SA 1 1 850 

M21D AE 1 1 1064 

MAGkUR1li"ct(1 ~,:i1t i t 1 325 

LEPOF'IiDAJM B2EVIBARBE 1 1 329 

FIOWELLA SP. 1 1 46" 

HOF'LtT!t'.'; :7' . 1 1 654 

HILDEBRANDIA Gl1F'PYI 1 1 351 

HEF'"f a pWCH1, %:=; PERLO 1 1 354 

HELIGOLEPIUS DACTYLOPTERUS 1 1 366 

GADONpJS SP. 1 1 $71 

ETMOPTERIIS PUS LLUS 1 1 627 

DEC APTERUS PUNCT ATl1S 1 1 786 

CORYPHIIENOmES SE' . 1 1 1135 

BOROSTOMIAS AMTARCTICI.IS 1 1 632 

BATHPTEROIS SP. 1 1 625 

BATHOPHILUS SP. 1 1 1440 

AP ISTRURIIS L AURUSSUN I I 1 1 1170 
As the Nation's principal conservation 

agency, the Department of the Interior has 

responsibility for most of our nationally 

owned public lands and natural resources . 

The includes fostering the wisest use of our 

land and water resources, protecting our 

fish and wildlife, preserving the 

environmental and cultural values of our 

national parks and historical places, and 

providing for the enjoyment of life through 

outdoor recreation. The Department 

assesses our energy and mineral resources 

and works to assure that their development 

is in the best interest of all our people. The 

Department also has a major responsibility 

for American Indian reservation communities 

and for people who live in Island Territories 

under U.S. Administration. 

y i 

. _ ~

. 

+~e.~..ar~'