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1984

Feeding strategies and functional morphology of demersal deep-sea ophidiid fishes

H. Jacque. Carter College of William and Mary - Virginia Institute of Marine Science

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Recommended Citation Carter, H. Jacque., "Feeding strategies and functional morphology of demersal deep-sea ophidiid fishes" (1984). Dissertations, Theses, and Masters Projects. Paper 1539616600. https://dx.doi.org/doi:10.25773/v5-v7m8-1a83

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University MicixSilms International 300 N. Z eeb Road Ann Arbor, Ml 48106

8500636

Carter, Herbert Jacque

FEEDING STRATEGIES AND FUNCTIONAL MORPHOLOGY OF DEMERSAL DEEP-SEA OPHIDIID FISHES

The College of William and M ary in Virginia PH.D. 1984

University Microfilms International300 N. Zeeb Road, Ann Arbor, Ml 48106

FEEDING STRATEGIES AND FUNCTIONAL MORPHOLOGY

OF DEMERSAL DEEP-SEA

OPHIDIID FISHES \

A Dissertation

Presented to

The Faculty of the School of Marine Science The College of William and Mary in Virginia

in Partial Fulfillment

of the Requirements for the Degree of

Doctor of Philosophy

by

H. Jacque Carter

1984 APPROVAL SHEET

This dissertEtior is suciritted in partial fulfillment of

the re^uire.-nsnts for the degree of

Doctor of Philosophy

t h

Approved* May 1934

n A. u s i c k

Daniel ,M. Cohen ' os An5a/tes County Museum

John/V.J Merriner Nat i/o nail Msrine Fisher^re's Service

Ray S. Sirdsong / Old Domindfcn University

yrne Dedication

This work is dedicated to my parents, my wife Judy and my son Jared. Table of Contents Page

Acknowledgments......

List of Tables ...... viii

List of Figures...... xii

List of Appendices...... xxiii

Abstract...... xiv

Introduction...... 2

Study Area...... 13

Material and Methods...... 17

Sampling...... 17

Gut content analysis...... IS

Morphological analysis...... 20

Results...... *...... 27

Food habits...... *...... 27

Feeding associations...... 73

Morphological associations...... 78

Description of the feeding mechanism...... Ill

Osteology...... Ill

Myology...... 116

Biomechanics...... 119

Discussion...... 158

Characterization of feeding strategies...... 158

Relationship between diet and morphology...... 172

Relationship between feeding strategies and regional primary productivity...... 178

Literature Cited...... 183

Appendices...... 199

Vita...... 236

iv Acknowledgments

Hy interest in ichthyology began while I was an undergraduate at

Northern Illinois University. I express my sincere gratitude to Dr.

David W. Greenfield, then professor of biology, who offered much inspiration and guidance in the formative years of my academic career.

Support and direction for this present Btudy were provided at the

Virginia Institute of Marine Science by Dr. John A. Musick. I thank

John Musick for the opportunity to conduct research on deep-sea fishes, for unfettered freedom in the direction of this research, and for his comments, assistance and encouragement throughout this work.

I gratefully acknowledge the influence of a very special group of scientists and students at VIMS whose enthusiasm for ichthyology and disregard for conventional wisdom created the synergism essential for the success of the VIMS Deep-Sea Program. I thank this motley crew from

VIMS for their unselfish assistance in the field and laboratory and for the many long hours of philosophical discussion, that help form many of the idea6 incorporated in my dissertation. Meriting special mention are these individuals: Roy E. Crabtree, M. Eric AnderBon, Thomas A. Munroe,

George R. Sedberry, John Gartner, William Raschi and James

Colvocoresses. In addition to those individuals just listed, others

contributing significantly in this regard include: John Olney, Jeff

Govoni, Joseph Smith, and John Gourly. I thank Bill Jenkins and Kay Stubblefield for their pbotographic and illustrative technical assistance respectively. I thank Ruth Klinger for help with typing the manuscript and James Colvocoresses, Ginny Shav and Patricia Hall for their advice and assistance with computer programs and procedures.

Special thanks to C. R. Robins who allowed me access to specimens deposited in the University of Miami fish collection. For loan of material I thank, Dr. W. Eschmeyer of the California Academy of

Sciences, San Francisco; Dr. R. K. Johnson of the Field Museum of

Natural History, Chicago; Drs. K. F. Liem, W. Fink and Karsten Hartel of the Museum of Comparative Zoology, Cambridge; Mr. N. Merrett of the

Institute of Oceanographic Sciences, Wormely, England; Dr. J. Nielsen of the Zoological Museum of the University of Copenhagen, Denmark; Drs. J.

Russo, B. Collette and S. Jewett and J. Goman of the United States

National Museum, Washington. I thank E. L. Wenner and J. Tan Montfrans and G. R. Sedberry for help in identifying prey organisms.

I am grateful to the members of my committee, John Merriner, Bob Byrne,

Ray Birdsong, Dan Cohen and my committee chairman, John Musick for their comments on the manuscript.

I am grateful for support by NSF Predoctoral Grant OCE 8104574 and the Raney Award from the American Society of Ichthyologists and

Herpetologists. Additional support was provided by NSF Grants NSF OCE

20507 and NSF OCE 7600729, Dr. John Musick, principal investigator, and

NSF GAA 38834, Dr. C. R. Robins, principal investigator.

I am especially appreciative of two individuals whose advice has been indispensable throughout the course of this study and who have significantly shaped the course of my career in science. The first of

these individuals is Dr. Kenneth J. Sulak, a gifted biologist and good

vi friend. His constant encouragement, critical but alway6 constructive reviews, practical knowledge of deep-sea trawling and seamanship, and

commitment to deep-sea biology serve as an example worth emulating. The

second of these individuals is Dr. Daniel M. Cohen. His generous

support and guidance throughout this study is deeply appreciated. His

advice and comment have always helped me communicate my ideas more

clearly and effectively.

Finally, Z am grateful to my wife, Judy, for her many sacrifices

and constant encouragement during the course of my studies.

vii List of Tables

Page

Table 1. Ophidiid fishes examined in food habit study...... 39

Table 2. Size range of predators and corresponding prey...... 40

Table 3. Gut contents of Dicrolene introniera. percent frequency of occurrence (F), percent numerical abundance (N), percent weight (w), total number (ATN) and absolute total weight...... 44

Table 4. Gut contents of Dicrolene kanazawai. percent frequency of occurrence (F), percent numerical abundance (n ), percent weight (w), total number (ATN) and absolute total weight...... 46

Table 5. Gut contents of Bathvonus pectoralis. percent frequency of occurrence (F), percent numerical abundance (N), percent weight (w), total number (ATN) and absolute total weight...... 48

Table 6. Gut contents of Barathrodemua manatinus. percent frequency of occurrence (F), percent numerical abundance (n ), percent weight (W), total number (ATN) and absolute total weight...... 32

Table 7. Gut contents of Bassozetus normalis. percent frequency of occurrence (F), percent numerical abundance (n ), percent weight (ff), total number (ATN) and absolute total weight...... 54

Table 8. Gut contents of Bassozetus taenia. percent frequency of occurrence (F), percent numerical abundance (N), percent weight (W), total number (ATN) and absolute total weight...... 56

Table 9. Gut contents of Xvelacvba mversi. percent frequency of occurrence (F), percent numerical abundance (N)f percent weight (w), total number (ATN) and absolute total weight...... 59

Table 10. Gut contents of Acanthonus armatus. percent frequency of occurrence (f), percent numerical abundance (N)> percent weight (w)( total number (ATN) and absolute total weight...... 61

viii TableB cont. Page

Table 11. Gut contents of Poroeadua miles. percent frequency of occurrence (F), percent numerical abundance (N), percent weight (w), total number (ATN) and absolute total weight...... 63

Table 12. Gut contents of Poroeadua situs. percent frequency of occurrence (f )» percent numerical abundance (N), percent weight (w), total number (ATN) and absolute total weight...... 66

Table 13. Gut contents of Poroeadua catena. percent frequency of occurrence (F), percent numerical abundance (N), percent weight (w), total number (ATN) and absolute total weight...... 68

Table 14. Gut contents of Barathrites parri. percent frequency of occurrence (f ), percent numerical abundance (N), percent weight (W)* total number (ATM) and absolute total weight...... 70

Table 15. Gut contents of Penopus macdonaldi. percent frequency of occurrence (F), percent numerical abundance (N)* percent weight (W)* total number (ATN) and absolute total weight...... 72

Table 16. Ophidiid fishes examined in food habit study...... 87

Table 17. Mean values of morphological characters related to feeding for each species in group I...... 88

Table 18. Morphometric* meristic and qualitative characters related to feeding in Dicrloene kanazawai...... 89

Table 19. Morphometric* meristic and qualitative characters related to feeding in Dicrolene intronigra...... 90

Table 20. Morphometric* meristic and qualitative characters related to feeding in Bathvonus nectoralis...... 91

Table 21. Morphometric* meristic and qualitative characters related to feeding in Monomitopus agassizii...... 92

ix Tables cont. Page

Table 22. Mean values of morphological characters related to feeding for each species in group II...... 93

Table 23. Morphometric, meristic and qualitative characters related to feeding in Anagesoma edentatum...... 94

Table 24. Morphometric, meristic and qualitative characters related to feeding in Xvelacvba mversi. .... 95

Table 25. Morphometric, meristic and qualitative characters related to feeding in BassozetuB norma lie ...... 96

Table 26. Morphometric, meristic and qualitative characters related to feeding in Bassozetus taenia...... 97

Table 27. Morphometric, meristic and qualitative characters related to feeding in Holcomvcteronus sauamosus...... 98

Table 28. Mean values of morphological characters related to feeding for each species in group III and IV...... 99

Table 29. Morphometric, meristic and qualitative characters related to feeding in Barathrodemus manatinus (male)...... 100

Table 30. Morphometric, meristic and qualitative characters related to feeding in Barathrodemus manat inus (female)...... 101

Table 31. Morphometric, meristic and qualitative characters related to feeding in Barathrites narri...... 102

Table 32. Morphometric, meristic and qualitative characters related to feeding in Barathrites iris...... 103

Table 33. Morphometric, meristic and qualitative characters related to feeding in Abvssobrotula galatheae...... 104

x Tables cont. Page

Table 34. Mean values of morphological characters related to feeding for each species in group V and VI...... 105

Table 35. Morphometric, meristic and qualitative characters related to feeding in Poroeadus silus...... 106

Table 36. Morphometric, meristic and qualitative characters related to feeding in Poroeadus catena...... 107

Table 37. Morphometric, meristic and qualitative characters related to feeding in Poroeadus mileB...... 108

Table 38. Morphometric, meristic and qualitative characters related to feeding in Penopus macdonaldi...... 109

Table 39. Morphometric, meristic and qualitative characters related to feeding in Acanthonus armatus ...... 110

xi List of Figures

Page

Porogadus silus. VIMS 7820, 185 mm SL B. Poroeadus catena. VIMS 7821, 125 mm SL C. Porogadus miles. VIMS 7819* 321 mm SL D. Penopua macdonaldi. VIMS 7813 188 ...... 7

A. Bassozetus normalia. VIMS 7814, 695 mm SL B. Bassozetus taenia. VIMS 7818, 482 mm SL C. Holcomvcteronus squamoaus. VIMS 7822 295 mm S L ...... 8

A. Dicrolene introniera. VIMS 7829, 218 mm SL B. Dicrolene kanazawai. VIMS 7828, 266 mm SL C. Honomitopus aeassizii. VIMS 7830, 210 m m SL D. Bathyonus pectoralis. VIMS 7818 132 mmSL ...... 9

A. Barathrites parri. VIMS 7816, 302 mm SL B. Xvelacvba mversi. VIMS 7824, 220 mm SL C. Acanthonus armatus. VIMS 7825 315 mm S L ...... 10

C. Abyssobrotula ealatheae. VIMS 7826, 186 mm SL A. Barathrodemus manatinua. Male, VIMS 7823, 186 mm y™ SL B. Barathrodemus manatinus. Female, VIMS 7852 129 mmSL...... 11

A. Barathrites iris. VIMS 7817, 802 mm SL B. Apaeesoma edentatum. VIMS 7827, 733 mm S L ...... 12

The Bahamas Study Region including the Blake Plateau and Outer Rise and Abyss areaB...... 15

The four major Bahama in deep-sea basins (Tongue of-the-Ocean, Exuma Sound, Columbus Basin, Inagua Basin), and with Providence Channel and a portion of the adjacent abyssal plain...... 16

Mean prey length versus mean standard lengths of ophidiid species...... 41 Figures continued Page

10. Length frequency histograms for examined species of D. kanazawai. D.. intronigra and B.. pectoralis ...... 42

11. Mean prey length versus standard length groups of D.. intronigra. D. kanazawai and B.. pectoralis...... 43

12. Percent frequency occurrence, percent number, percent weight, and index of relative importance (iRl) of higher taxonomic groups of food in the diet of Dicrolene intronigra...... 45

13. Percent frequency occurrence, percent number percent weight, and index of relative importance ((IRl) of higher taxonomic groups of food in the diet of Dicrolene kanazawai...... 47

14. Percent frequency occurrence, percent number percent weight, and index of relative importance (IRl) of higher taxonomic groupB of food in the diet of Bathvonus pectoralis...... 49

15. Length frequency histograms for examined species of JB. manat inus. B. normal is and B, taenia...... 50

16. Mean prey length versus standard length groups of B_. manat inus. B.. normal is and P. silus...... 51

17. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Barathrodemus manatinuB ...... 53

18. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Bassozetus normalis...... 55

19. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Bassozetus taenia...... 57

xiii Figures continued Page

20. Length frequency histograms for examined species of P. miles. X. mversi and armatua...... 58

21. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groupB of food in the diet of Xvelacvba myersi...... 60

22. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Acanthonus armatua ...... 62

23* Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Porogadus miles ...... 64

24. Length frequency histograms for examined species of P.. silus. P.. macdonaldi. P. catena and B. narri...... 65

25. Percent frequency occurrence, percent number, percent weight, and index of relative importance (XRl) of higher taxonomic groups of food in the diet of Poroeadus silus...... 67

26. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Porogadus catena*...... 69

27. Percent frequency occurrence, percent number, percent weight, and index of relative importance (XRl) of higher taxonomic groups of food in the diet of Barathrites parri...... 71

28. Dendogram depicting diet similarity among ophidiid predators...... 77

29. Dendogram depicting morphological similarity among deep-sea ophidiids...... 87

xiv Figures continued Page

30. A. Bathvonus pectoralis. V IMS 7836, 132 mm SL, lateral facing bones B. Porogadus catena. VIMS 7838, 185 mm SL, lateral facing bones C. Poroeadus silus. VIMS 7839, 125 mm SL, lateral facing bones D. Penonus macdonaldi. VIMS 7838, 188 mm SL, lateral facing bones E. Poroeadus miles. VIMS 7837, 321 mm SL, lateral facing bones...... 124

31. A. Dicrolene kanazawai. VIMS 7846, 226 mm SL, lateral facing bones B. Dicrolene intronigra. VIMS 7847, 218 mm SL, lateral facing bones C. Monomitopus aeassizii. VIMS 7849, 210 mm SL, lateral facing bones D. Barathrodemus manatinus. VIMS 7841, 186 mm SL, lateral facing bones E. Barathrites parri. VIMS 7834, 302 mm SL, lateral facing bones F. Abvssobrotula ealatheae. VIMS 7844, 186 mm SL, lateral facing bones...... 125

32. A. Acanthonus armatus. VIMS 7843, 315 mm SL, lateral facing boneB B Xvelacvba mversi. VIMS 7842, 220 mm SL, lateral facing bones C. Bassozetus normalis. VIMS 7831, 695 mm SL, mm SL, lateral facing bones D. Bassozetus taenia. VIMS 7832, 482 mm SL, SL, lateral facing bones...... 126

33. A Poroeadus silus. VIMS 7839, 125 mm SL, suspensorium B. Penopus macdonaldi. VIMS 7838, 188 mm SL, suspensorium C. Poroeadus catena. VIMS 7838, 185 mm SL, suspensorium D. Poroeadus miles. VIMS 7837, 321 mm SL, suspensorium E. Bathvonus pectoralis. VIMS 7836, 132 mm SL, SL, suspensorium...... 127

xv Figures continued Page

Abvssobrotula galatheae. VIMS 7844, 186 nnn SL, 123 nnn SL, suspensorium Barathrites parri. VIMS 7834, 302 nnn SL, suspensorium Barathrodemus manatinus. VIMS 7841, 186 m m SL, mm SL, suspensorium Monomitopus agassizii. VIMS 7849, 210 mm SL, SL, suspensorium Dicrolene kanazawai. VIMS 7846, 226 mm SL, suspensorium Dicrolene intronigra. VIMS 7 847 , 218 mm SL, SL, suspensorium...... 128

35. A. Bassozetus normalis. VIMS 7831, 695 mm SL, suspensorium B. Bassozetus taenia. VIMS 7832, 482 mm SL, suspensorium C. Xvelacvba mversi. VIMS 7842, 220 mm SL, suspensorium D. Acanthonus armatua. VIMS 7843, 315 nnn SL, suspensorium...... 129

36. A. Penonus macdonaldi. VIMS 7838, 188 mm SL, opercular series B. Porogadus silus. VIMS 7839, 125 mm SL, opercular series C. Porogadus catena. VIMS 7838, 185 mm SL, SL, opercular series D. Porogadus miles. VIMS 7837, 321 mm SL, SL, opercular series E. Bathvonus pectoralis. VIMS 7836, 132 mm SL, opercular series...... 130

37. A. Dicrolene intronigra. VIMS 7847, 218 mm SL, opercular series B. Dicrolene kanazawai. VIMS 7846, 226 mm SL, opercular series C. Barathrodemus manatinus. VIMS 7841, 186 m m SL, SL, opercular series D. Monomitopus agassizii, VIMS 7849, 210 m m SL, SL, opercular series E. Barathrites parri. VIMS 7834, 302 mm SL, opercular series F. Abvssobrotula galatheae. VIMS 7844, 186 mm SL, SL, opercular series ...... 131

xvi Figures continued Page

38. A. Bassozetus normalis. VIMS 7831, 695 m m SL, SL, opercular series B. Bassozetus taenia. VIMS 7832, 482 mm SL, SL, opercular series C. Xvelacvba mversi. VIMS 7842, 220 mm SL, opercular series D. Acanthonus armatus. VIMS 7843, 315 mm SL, opercular series ...... 132

39. A. Penonus macdonaldi. VIMS 7838, 188 mm SL, jaw apparatus B. Poroeadus silus. VIMS 7839, 125 mm SL, jaw apparatus C. Bathvonus pectoralis. VIMS 7836, 132 mm SL, jaw apparatus D. Poroeadus catena. VIMS 7838, 185 mm SL, jaw apparatus E. Poroeadus miles. VIMS 7837, 321 mm SL, jaw apparatus...... 133

40. A. Dicrolene intronigra. VIMS 7847, 218 mm SL jaw apparatus B. Barathrodemus manatinus. VIMS 7841, 186 mm SL, mm SL, jaw apparatus C. Dicrolene kanazawai. VIMS 7846, 226 m m SL, jaw apparatus D. Abvssobrotula galatheae. VIMS 7844, 186 mm SL, SL, jaw apparatus E. Monomitopus aeaBsizii. VIMS 7849, 210 mm SL, SL, jaw apparatus F. Barathrites parri. VIMS 7834, 302 mm SL, jaw apparatus...... 134

41. A. Bassozetus normalis. VIMS 7831, 695 TTTTTI SL j jav apparatus B. Bassozetus taenia. VIMS 7832, 482 mm SL, jaw apparatus C. Acanthonus armatus. VIMS 7843 , 315 mm SL, jaw apparatus D. Xvelacvba mversi. VIMS 7842, 220 mm SL, jaw apparatus...... 135

42. A. Poroeadus catena. VIMS 7838, 185 mm SL, hyoid arch B. Poroeadus silus. VIMS 7839, 125 mm SL, hyoid arch C. Bathvonus pectoralis. VIMS 7836, 132 mm SL, hyoid arch D. Porogadus miles. VIMS 7837, 321 mm SL, hyoid arch E. Penonus macdonaldi. VIMS 7838, 188 mm SL, SL, hyoid arch...... 136

xvii Figures continued Page

43. A. Dicrolene intronigra. VIMS 7847, 218 mm SL, hyoid arch B. Dicrolene kanazawai. VIMS 7846, 226 mm SL, hyoid arch C. Monomitopus agassizii. VIMS 7849, 210 mm SL, hyoid arch D. Abvssobrotula galatheae. VIMS 7844, 186 mm SL, SL, hyoid arch E. Barathrodemus manatinus. VIMS 7841, 186 TTTTT1 SL j SL, hyoid arch F. Barathrites parri. VIMS 7834, 302 mm SL, hyoid arch...... 137

44. A. Bassozetus taenia. VIMS 7832, 482 mm SL, hyoid arch B. Bassozetus normalis. VIMS 7831, 695 mm SL, hyoid arch C. Xvelacvba mversi. VIMS 7842, 220 mm SL, hyoid arch D. Acanthonus armatus. VIMS 7843, 315 mm SL, hyoid arch...... 138

45. A. Porogadus catena. VIMS 7838, 185 mm SL, lover branchial series B. Porogadus silus. VIMS 7839, 125 mm SL, lower branchial series C. Porogadus miles. VIMS 7837, 321 mm SL, lover branchial series D. Bathvonus pectoralis. VIMS 7836, 132 mm SL, lower branchial series E. Penopus macdonaldi. VIMS 7838, 188 mm SL, lower branchial series ...... 140

46. A. Dicrolene intronigra. VIMS 7847, 218 mm SL, lower branchial series B. Dicrolene kanazawai. VIMS 7846, 226 mm SL, lower branchial series C. Abvssobrotula galatheae. VIMS 7844, 186 mm SL, SL, lower branchial series D. Monomitopus agassizii. VIMS 7849, 210 mm SL, lower branchial series E. Barathrites parri. VIMS 7834, 302 mm SL, lower branchial series F. Barathrodemus manatinus. VIMS 7841, 186 mm SL, SL, lower branchial basket...... 140

xvlii Figures continued Page

47. A. Bassozetus normalis. VIMS 7831, 695 mm SL, lower branchial series B. Bassozetus taenia VIMS 7832, 482 mm SL, lower branchial series C. Xvelacvba mversi. VIMS 7842, 220 mm SL, lover branchial series D. Acanthonus armatus. VIMS 7843, 315 mm SL, lower branchial series...... 141

48. A. Poroeadus silus. VIMS 7839, 125 mm SL, upper branchial series B. Porogadus miles. VIMS 7837, 321 mm SL, upper branchial series C. Porogadus catena. VIMS 7838, 185 mm SL, upper branchial series D. Bathvonus pectoralis. VIMS 7836, 132 mm SL, upper branchial series E. Penopus macdonaldi. VIMS 7838, 188 mm SL, upper branchial region...... 142

49. A. Barathrites parri. VIMS 7834, 302 mm SL, upper branchial series B. Dicrolene kanazawai. VIMS 7846 , 226 mm SL, upper branchial series C. Monomitopus agassizii. VIMS 7849, 210 mm SL, SL, upper branchial series D. Abvssobrotula galatheae. VIMS 7844, 186 mm SL, upper branchial series E. Barathrodemus manatinus. VIMS 7841, 186 mm SL, upper branchial series F. Dicrolene intronigra. VIMS 7847, 218 mm SL, upper branchial series...... 143

50 A. Xvelacvba mversi. VIMS 7842, 220 mm SL, upper branchial series B. Acanthonus armatus. VIMS 7843, 315 mm SL, upper branchial series C. Bassozetus normalis. VIMS 7831, 695 mm SL, upper branchial series D. Bassozetus taenia. VIMS 7832, 482 mm SL, upper branchial series...... 144

51. A. Penopus macdonaldi. VIMS 7838, 188 mm SL, pectoral girdle B. Bathvonus pectoralis. VIMS 7836, 132 mm SL, pectoral girdle C. Porogadus miles. VIMS 7837, 321 mm SL, pectoral girdle D. Porogadus silus. VIMS 7839, 125 mm SL, pectoral girdle E. Porogadus catena. VIMS 7838, 185 mm SL, pectoral girdle...... 145

xix Figures continued Page

52. A. Monomitopus agassizii. VIMS 7849, 210 mm SL, SL, pectoral girdle B. Abvssobrotula galatheae. VIMS 7844, 186 mm SL, SL, pectoral girdle C. Barathrodemus manatinus. VIMS 7841, 186 mm SL, SL, pectoral girdle D. Barathrites parri. VIMS 7834, 302 mm SL, pectoral girdle E. Dicrolene intronigra. VIMS 7847, 218 mm SL, pectoral girdle D. Dicrolene kanazawai. VIMS 7846, 226 trnn SL, pectoral girdle...... 146

53. A. Bassozetus taenia. VIMS 7832, 482 mm SL, pectoral girdle B. Bassozetus normaliB. VIMS 7831, 695 mm SL, pectoral girdle C. Xvelacvba mversi. VIMS 7842, 220 mm SL, pectoral girdle D. Acanthonus armatus. VIMS 7843, 315 mm SL, pectoral girdle < < ...... 147

54. A. Porogadus catena. VIMS 7838, 185 mm SL, neurocranium, dorsal, lateral, ventral aspects, top to bottom respectively B. Penopus macdonaldi. VIMS 7838, 188 mm SL, neurocranium, dorsal, lateral, ventral aspects, top to bottom respectively C. Bathvonus pectoraliB. VIMS 7836, 132 mm SL, neurocranium, dorsal, lateral, ventral aspects respectively D. Porogadus silus. VIMS 7839, 125 mm SL, neurocranium, dorsal, lateral, ventral, aspects, top to bottom respectively E. Porogadus miles. VIMS 7837, 321 mm SL, neurocranium, dorsal, lateral, ventral, aspects respectively...... 148

55. A. Abvssobrotula galatheaeu VIMS 7844, 186 mm SL, neurocranium, dorsal, lateral, ventral, aspects respectively B. Barathrites parri. VIMS 7834, 302 mm SL, neurocranium, dorsal, lateral, ventral views respectively C. Dicrolene kanazawai. VIMS 7846, 226 mm SL, neurocranium, dorsal, lateral, ventral view respectively D. Dicrolene intronigra. VIMS 7847, 218 mm SL, neurocranium, dorsal, lateral, ventral view respectively

xx Figures continued Page

E. Barathrodemus manatinus. VIMS 7841, 186 mm SL, SL, neurocranium, dorsal, lateral, ventral views respectively F. Monomitopus agassizii. VIMS 7849, 210 mm SL, SL, neurocranium, dorsal, lateral, ventral views respectively ...... 149

56. A. Acanthonus armatus. VIMS 7843, 315 mm SL, neurocranium, dorsal, lateral, ventral views respectively B. Xvelacvba mversi. VIMS 7842, 220 mm SL, neurocranium, dorsal, lateral, ventral viewB respectively C. Bassozetus taenia. VIMS 7832, 482 mm SL, neurocranium, dorsal, lateral, ventral views respectively D. Bassozetus normalis. VIMS 7831, 695 nim SL ^ SL, neurocranium, dorsal, lateral, ventral views respectively...... 150

57. A. Poroeadus miles. VIMS 7837, 321 mm SL, first left gill arch B. Poroeadus catena. VIMS 7838, 185 mm SL, first left gill arch C* Penonus macdonaldi. VIMS 7838, 188 m m SL, first left gill arch D. Bathvonus pectoralis. VIMS 7836, 132 m m SL, first left gill arch E. Poroeadus silus. VIMS 7839, 125 mm SL, first left gill arch...... 151

58. A. Abvssobrotula ealatheae. VIMS 7844, 186 m m SL, first left gill arch B. Dicrolene introniera. VIMS 7847, 218 mm SL, first left gill arch C. Barathrites parri. VIMS 7834, 302 mm SL, first left gill arch D. Monomitopus agassizii. VIMS 7849, 210 mm SL, first left gill arch E. Barathrodemus manatinus. VIMS 7841, 186 mm SL, SL, first left gill arch F . Dicrolene kanazawai. VIMS 7 846 , 226 mm SL, first left gill arch...... 152

59. A. Bassozetus taenia. VIMS 7832, 482 mm SL, first left gill arch B. BassozetuB normalis. VIMS 7831, 695 m m SL, first left gill arch C. Acanthonus armatus. VIMS 7843, 315 m m SL, first left gill arch D. Xvelacvba mversi. VIMS 7842, 220 mm SL, first left gill arch...... 153

xxi Figures continued Page

60. Ligamentous attachments of the upper jaw of Dicrolene intronigra. (dorsal view) VIMS 7847, 218 mm SL...... 154

61. Lateral view of the superficial cheek muscles of Dicrolene intronigra. VIMS 7847, 219 mm S L ...... 155

62. Lateral view of the deep cheek muscles of Dicrolene intronigra. VIMS 7847, 218 mm SL...... 156

63. Ventral view of cranial muscles of Dicrolene intronigra. VIMS 7847, 218 mm SL...... 157

xx ii List of Apendices Page

Appendix X. Abbreviations used in text and figures...... 199

Appendix 2. Morphological analysis data record sample...... 201

Appendix 3. Station data listing for Bahamas study region (Modified from Sulak, 1982)...... 205

Appendix 4* Cluster analysis output based on morphological characters...... 221

Appendix 5. Cluster analysis output based on food habits...... 229

xxiii Abstract

The hypothesis is advanced that ophidiid fishes prevalent beneath oligotrophic tropical seas have evolved trophic specializations appropriate for existence in regions of low primary productivity where ultimate energy available for food production in benthic communities is low. The feeding strategies of these tropical dwelling deep-sea fishes are discussed within the context of this hypothesis. The feeding strategies of tropical dwelling deep-sea ophidiid fishes are examined through an analysis of food habits and examination of morphological features related to feeding. Three main modes of feeding are evident among these fishes: (1) predation upon small benthopelagic or planktonic organisms hy small-bodied species of limited locomotory ability; (2) predation upon benthic-benthopelagic organisms with increasing reliance upon larger nektonic prey by mobile large­ bodied species; (3) predation upon benthic organisms with diminished reliance upon benthopelagic and planktonic organisms. The first feeding mode is shown to be more widespread among deep-sea ophidiids studied. Ontogenetic feeding changes in select species show a shift from small benthic-benthopelagic prey in small fish to more mobile nektonic prey in larger fish. Smaller fish utilize such small-sized prey as calanoid copepods, gammarid amphipods and cumaceana, while larger fish show increasing reliance on decapods and fish. In contrast several small­ bodied species studied show no ontogenetic feeding changes in relation to prey 6ize. Variation and select interrelations of morphological features related to feeding are studied in 18 species of ophidiids. Cluster analysis is employed as a verification procedure to determine the correspondence between feeding morphology and diet. Six morphological species groups are identified among species examined based on similarity of characters related to feeding. The results show poor correspondence between morphologically structured guilds based on characters assumed related to feeding and feeding guilds based on stomach content analysis. In addition morphological characters are interpreted ecologically based on information in the literature and gut content analysis. In several instances there is strong support of the hypothesized adaptations reported in the literature while in other cases there was little or no support. The functional relationship of the feeding apparatus is considered to better understand trophic specializations and feeding adaptations in these fishes. Dicrolene intronigra serves to illustrate the feeding mechanism in deep-sea ophidiid fishes. A basic description of the mechanical units of the head and their movements involved in feeding are presented.

xx iv Feeding Strategies and Functional Morphology of Deep-Sea Ophidiid Fishes Introduction

Neobythitine fishes of the family Ophidiidae are an important

component of the demersal deep-sea fish fauna in the Bahamas and nearby

Sargasso Sea (Sulak et al.. 1979). These fishes are numerous, diverse and have distinct morphological features that have been used to unite

them in a fairly well defined group within the Ophidiiformes (Cohen and

Nielsen, 1978). Host species are small and oviparous. However, little

is known of their anatomy and functional morphology (Mead et al.. 1964;

Cohen and Nielsen, 1978), particularly as it relates to feeding habits.

The purpose of this study is to characterize feeding strategies of

these tropical dwelling deep-sea fishes through an analysis of food

habits and examination of morphological features related to feeding.

The hypothesis is advanced that ophidiid fishes prevalent beneath

oligotrophic tropical seas have evolved trophic specializations

appropriate for existence in regions of low primary productivity where

ultimate energy available for food production in benthic communities is

low. Species examined in this study are depicted in figures 1-6.

Much of our present knowledge of the biology of deep-sea

neobythitine fishes stems from descriptive or faunal works on fiBhes

obtained from expeditions around the turn of the century (Agassiz, 1886;

German, 1899; Goode and Bean 1883, 1885, 1896; Gunther, 1878, 1887;

Koefoed, 1927; Roule, 1913, 1916; Vaillant, 1888; Zugmayer, 1911).

Although these early studies on neobythitine fishes were primarily

descriptive they also provided valuable ecological information.

2 Morphological descriptions of the head, dentition, gill rakers, body size, etc. are especially useful since such features are important elements of the feeding apparatus of fishes. Nybelin (1954, 1957) in his study of neobythitine fishes obtained from the Swedish Deep-Sea

Expedition, reported extensively on distributional patterns and discussed specializations in species dentition. Recent investigators have examined morphological features of the jaw apparatus of neobythitine fishes in an effort to elucidate the phylogentic relationships of the ophidioids as a whole to other teleost groups

(Gosline, 1968, 1971, 1980; Rosen and Patterson, 1969). Additional information available on the functional anatomy of the feeding apparatus of neobythitine fishes has been reported incidentally in the taxonomic

literature (Cohen, 1961, 1966, 1974, 1982; Gosline, 1953; Grey, 1958;

Mead et a l . . 1964; Nielsen, 1964, 1965, 1969, 1971, 1975, 1977; Wenner,

1978).

Little information is available concerning the diets of neobythitine fishes. Rayburn (1975) found that polychaetes were the

primary food items found in the stomachs of Dicrolene intronigra while

ostracods were the primary food of Monomitopus agassizii. He classified

D. intronigra and M. agassizii as generalized grazers since they seemed

adapted to a browsing type of feeding. He further suggested that the

dietary differences between them were due to competition between morphologically similar species occupying the same area. Bright (1968)

found the primary food items of D. intronigra to be foraminifera and

crustaceans whereas polychaetes and ostracods were the primary food

items of M. agassizii. Wenner (1978) reported that D.. intronigra fed

primarily on benthic infauna and epifauna with considerable quantities 4 of mud being found in the intestine. Additional data on the food habits of neobythitine fishes has been reported cursorily in the taxonomic

literature (Nielsen, 1965, 1975, 1977). The available data on diets of

these fishes provides useful information but is insufficient to

characterize their feeding repertoires or infer that competition occurs

among species.

The functional anatomy and feeding strategies of other deep~sea

fishes have been examined in detail by several investigators. Marshall

and Bourne (1964) analyzed fin pattern and body shape of macrourids.

Okamura (1970) correlated several aspects of the anatomy of macrourids

with stomach contents. Geistdoerfer (1972, 1973, 1975) examined

pharyngeal teeth, jaw mechanics, stomach contents, and the digestive

system of several species of macrourids. Chao and Musick (1978) have

established correlations between diet and morphology in several species

of sciaenids. Tchernavin (1953) described the functional anatomy of the

viperfiBh, Chauliodus sloani. and was able to explain how C.. sloani is

capable of capturing both very small and very large prey organisms.

Differences in mouth size and gill raker structure of several

melamphaids have been compared with differences in predatory strategy

among species occurring at different depths (Ebeling and Cailliet,

1974). The mechanics of the head of the Chimera, Callorhvnchus

canensis. and the contents of stomachs have been studied together to

understand its evolutionary biology and ecology (Ribbink, 1971).

McDowell (1973) related head morphology to diet in the deep-sea

notacanthiform fishes.

Mclellan's (1977) study of the head morphology and feeding

strategies of macrourids is particularly important to this research 5 since macrourids occur in the Mid-Atlantic Bight at similar depths inhabited by neobythitines in the Bahamas. A comparison of neobythitine fishes in the Bahamas with those of macrourids in the Mid-Atlantic Bight provides a basis for an ecological characterization of these two groups feeding strategies. Her study concentrated primarily on differences in diet and morphology among extremely specialized macrourids* She found that extremely specialized fishes with very long rostra have a diet consisting primarily of benthic prey whereas species with large terminal mouths and blunt snouts have stomach contents of primarily pelagic and benthopelagic prey items. Species intermediate between the two extremes fed on various food sources depending on size of the fish, depth of occurrence and ability to migrate off the bottom into midwaters.

Most data available on diets of deep-sea fishes have been reported in the taxonomic literature and in a few ecological studies (Bright,

1968; Macpherson, 1979; Marshall and Iwamoto, 1973; Marshall and

Merrett, 1977; McDowell, 1973; Mclellan, 1977; Rayburn, 1975; Robins,

1968; Sedberry and Musick, 1978). Bright (1968) examined the stomachs of several species of deep-sea fishes from the Gulf of Mexico and found that polychaetes and crustaceans were the predominant prey item. He also recognized three feeding modes: (1) predation upon small benthic organisms; (2) predation upon small benthopelagic or planktonic organisms; and (3) predation upon large macrobenthic, planktonic or nektonic animals. Macpherson (1979) noted dietary differences among

several species of macrourids in the western Mediterranean Sea. The fraction of small crustaceans found in diets of all species decreased as

fiBh sized increased. Also the mean size of prey increased with the body size of the fish. Marshall and Merrett (1977), in their review of 6 a deep-sea benthopelagic fauna, concluded that conditions of life near and on the deep-sea floor have favored the evolution of fish utilizing a mixed diet of benthic and pelagic organism. Sedberry and Musick (1978) examined the stomachs of 729 fishes comprising 16 species from the continental elope and rise off the Mid-Atlantic Bight. Two main feeding modes among demersal fishes were evident: those feeding primarily on pelagic food items and those feeding on benthic invertebrates.

Conclusions reached in many of these studies (Bright, 1968; Rayburn,

1975) are based on very small sample sizes and as a consequence should be interpreted with caution until more detailed knowledge is provided by future research. Figure 1. A. Porogadus situs. VIMS 7820, 185 mm SL

B. Porogadus catena. VIMS 7821, 125 mm SL

C. Poroeadus miles. VIMS 7819, 321 mm SL.

D. Penoous macdonaldi. VIMS 7813, 188 mm SL

8

Figure 2. A. Bassozetus normalie. VIMS 7814, 695 mm SL

B. Bassozetus taenia. VIMS 7818, 482 mm SL

C. HolcomvcteromiB sauamosus. VIMS 7822, 295 m m SL

Figure 3. A. Dicrolene intronigra. VIMS 7829, 218 mm SL

B. Dicrolene kanazawai. VIMS 7828, 266 mm SL

C. Monomitopua aeassizii. VIMS 7830, 210 mm SL

D. Bathvonus pectorals. VIMS 7818, 132 mm SL* /

mni

:iO mm Figure 4. A. Barathritea parri. VIMS 7816, 302 xsm SL

B. Xyelacyba mversi. VIMS 7824, 220 imn SL

C. A.captbonus armatua. VIMS 7825, 315 mm SL B Figure 5. A. Baratbrodemus manatinus. (male) VIMS 7823, 186 mm SL.

B. BaratbrodemuB manatinus. (female), VIMS 78523 129 mm SL.

C. Abvssobrotula ealatheae. VIMS 7826 186 mm SL *

Figure 6. A. garathrifceg Iris. VIMS 7817, 802 nnn SL (Figure from Roule, 1916)

B. Apagesoma edentatum. V IM S 7827, 733 mm SL (Figure from Carter, 1982)

Z

Study Area

The Bahamas region includes that portion of the North Atlantic subtended to the southwest by the arc of the Bahamas and the Turks and

Caicos islands (fig. 7). It is characterized by tropical, halostatic, oligotrophic surface waters.

The topography of the slope and rise in the Bahamas region is narrow and precipitous. Except in the region of the Blake Plateau, the distance between the 1000 and 4500m isobaths is no more than 20-50 km along the outside of the Bahamian island arc. The steep wall of the slope is due to the intrusion into the shallow Bahamas banks of several oceanic basins, including the Tongue-of-the-Ocean, Exuraa Sound, Columbus

Basin and Inagua Basin (figs. 8). These basins are basically steep­

sided and level-bottomed; they represent, in the order listed, a stepwise BerieB of increasing depths between 1200 and 3000m.

Bottom sediments in the Bahamas Study Region (including the

southern Blake Plateau) are characterized as silts and clays and consist predominately of calcareous oozes ( Athearn, 1962; Pilkey and Rucker,

1966; Gibson and Schlee, 1967; Emery and Uchupi, 1972). Such sediments

in the Bahamas originate primarily from deposition of foraminifera and

pteropods tests. Organic carbon content of Bahamian slope sediments is

generally low, ranging between 0.5 and 1.0 percent (Emery and Uchupi,

1972). The rate of pelagic sedimentation of particulate organic carbon

(5.7 mg/m^/day) at 2000 m in the Bahamas (Weibe et a l .. 1976) is

13 2 relatively low compared to values (12.8 mg/m /day) reported at comparable depth in the Middle Atlantic Bight (Rowe and Gardner, 1979).

For a more detailed account and general review of the study area the reader is referred to Sulak (1982). Figure 7. The Bahamas Study Region including the Blake Plateau and Outer Rise and Abyss areas. (Modified from Sulak, 1982) T. %

vcnacnj j q 82*00' 80*00' 78*00’ 76*00* 74*00* 72*00* TO'Otf Figure 8, The four major Bahamain deep-sea basins (Tongue of-the-Ocean, Exuma Sound, Columbus Basin, Inagua Basin), and with Providence Channel and a portion of the adjacent abyssal plain. (Modified from Sulak, 1982) m ------26*00 \ S K li t i . ,\ Njg\; /^gr_.

-9 •-' ^& -V! ( ^ ( V $ 4 5

2 5 * 0 0

s %

24*00' x ^ C ^ x " %v

\ + + A \ TONGUE3 U ^ OF THE OCEAN 23*00 f » * " j i • COLUMBUS \ 1. “ SIN?

22*00

INAGUA BASIN

fim* r s-'J'S p p ^ a f

70*00’ 7 7 * 0 0 76*00 7 5 * 0 0 ' 7 4 * 0 0 ' Materials and Methods

Sampling:

This research represents a portion of a much larger study conducted by Dr. John A. Husick to compare the structure and function of demersal deep~sea fish assemblages in tropical and temperate ecosystems. From

1972 to 1981 several cruises vere undertaken in the Bahamas region by the University of Miami, Rosenstiel School of Marine and Atmospheric

Science (RSMAS) and the Virginia Institute of Marine Science (VIMS).

All Bahamian station sites lay within a zone bounded by latitude 21-30°

N. Sampling was concentrated inside four major Bahamian basins:

Tongue-of-the-Ocean, Exuma Sound, Columbus Basin and Inagua Basin (fig.

8). Additional sampling was conducted on the southeastern portion of

Blake plateau, in Providence Channel at the entrance to Tongue-of-the-

Ocean, and on the abyssal plains flanking the Bahamian island arc (fig.

7).

Sampling was conducted with 12.5 m and 13.7 m semi-balloonotter trawls. All fishes captured were identified, counted, measured and weighed. Standard and/or gnathoproctal length was taken on all species.

Each fish was dissected and its stomach excised if not conspicuously empty. Each Btomach was labled, individually wrapped in cheesecloth and fixed in 10% seawater formalin.

Got content analysis:

17 After proper fixation, stomachs were soaked in water and transferred to either 402 isopropanol or 70% ethanol. Gut contents throughout the entire length of the digestive tract were examined in all individuals which were studied morphologically. Gut contents were identified as completely as possible, sorted and counted. Wet weight of the food items was measured as accurately as possible by using an analytical balance. Wet weight determinations were preferred over other more precise techniques (i.e. dry weight) in an effort to maintain prey itemB intact for reference collections. A substantial number of invertebrate prey items obtained in this study are as yet undescribed in the taxonomic literature. Size of prey items (along greatest linear axis) was measured directly from intact items and was estimated from the dimensions of an identified part in the case of broken or digested organisms. The presence of bottom sediment and the incidence of parasitism in the gut were also recorded.

Since methods of food habit analysis are variously biased (Hynes,

1950; Finkas et al., 1971; Windell, 1971), the relative contribution of different food items to the total diet was determined using three methods: (1) the number of stomachs in which a food item occurred was expressed as percentage of the total number of stomachs containing food

(F=percent frequency of occurrence); (2) the number of individuals of each type of food was expressed aB a percentage of the total number of food itemB from all stomachs (NBpercent numerical abundance); (3) the weight of food items was expressed as a percentage of the total weight of food from all stomachs examined (W“percent weight).

From these three measurements an index of relative importance, IRI

(Finkas et al.. 1971), was calculated for each prey species and higher 19

taxon as follows:

IRI-(N+W)F

This index has been useful in evaluating the relative importance of

different food items found in fish stomachs (Pinkas et al.. 1971;

McEachran et al.. 1976, Sedberry and Musick, 1978; Sedberry, 1980).

Similarity and overlap in diet among predators was measured using

numerical classification techniques (cluster analysis, Clifford and

Stephenson, 1975). Stomachs of predators were treated as collections

and were subjected to normal cluster analysis on the basis of prey

similarity, using index of relative importance values (ZRl). In normal

analysis predator species are the entities and prey species are the

attributes by which predators are classified. Cluster analysis was also

performed using percent numerical abundance and percent frequency. The

results from all three analyses were similar, suggesting that the IRX,

although biased, is a reliable measure of prey item importance in this

study. Flexible sorting (Lance and Williams, 1967; Clifford and

Stephenson, 1975), with Bs-0.25, was used based on resemblance expressed

by the Bray Curtis measure (Bray and Curtis, 1957). This is a measure

of dissimilarity and the compliment is used to yield a similarity measure (Clifford and Stephenson, 1975). The Bray-Curtis similarity measure can be expressed as:

where SjkBsimilarity between the entities (predator species) j

and K

Xij*abundance of the ith attribute (prey species) for

entity ij

Xik**is tbe abundance of the ith attribute for entity K. 20

Horphological analysis:

Freshly caught and preserved museum specimens of all taxa were dissected and examined. Representatives of each taxa were cleared and stained for osteological study following procedures presented by Taylor

(1967). The skull was divided into functional units and discussed on a comparative basis following Liem (1967). The nomenclature of bones follows Carter and Sulak (1984). Possible movements of the head were determined by manipulating freshly caught and frozen specimens following procedures presented by Mclellan (1977). Muscles and ligaments were examined from fresh and frozen specimens according to the method of Bock and Shear (1972).

Samples of 30 individuals were measured for each species for which

I had collected sufficient material. Determinations of meriBtic and morphometric characters were made on each individual fish* Measurements were obtained to the nearest 0.1 mm using dial calipers and were expressed as percent standard length. Measurements less than 0.5 mm were obtained with the aid of an ocular micrometer. For all qualitative characters, the various states of the characteristics were coded numerically using integers. These character state codes for qualitative characters are indicated in parentheses below where relevant.

Superficial body and body shape characters:

1. Standard length was recorded as a straight line distance

from the tip of the snout to the caudal peduncle.

2. Completeness of the lateral line canal was recorded as

being complete (1), incomplete (2), or lacking (3).

3. Position of the lateral line canal was recorded as

curving dorsally (1), horizontal (2), curving ventrally (3), or all of the above (4).

4. Body depth was maximum body depth at level of vent.

5. Index of trunk shape was the perpendicular distance from

the anterior tip of the head to an imaginary vertical

line at tthe point of maximum body depth divided by SL

Paired fin characters:

6. Pectoral fin length was the distance from the base of the

pectoral fin to the extreme tip of the fin at its

longest point.

7. Aspect ratio of the pectoral fin was estimated as a

length to width ratio.

8. Pectoral fin shape was coded based on a subjective

evaluation of whether the fins were (1) rounded, (2)

intermediate, (3) pointed.

9. Position of pectoral fin relative to the center of

gravity was an assignment of how the pectoral fin when

adpressed against the lateral surface of the body

relates to a transverse plane through the point CG

(center of gravity). This point CG was determined by

balancing the fish on the tip of a dissecting needle.

Four possible relationships were recognized: (1) fin

wholly anterior to the plane, (2) fin originating

anterior to CG and extending posterior to the plane, (3)

fin originating at the level of the CG plane, and (4)

fin originating posterior to CG.

10. Position of the dorsal fin relative to the center of

gravity was an assignment using the same four 22

character states as were used in the analogous

character for pectoral fin (#9 above). Assignment was

made according to the position of the entire dorsal fin

base relative to CG plane.

11. Position of the anal fin relative to the center of

gravity is an assignment using the same four character

states as were used in the character for pectoral fin

(#9 above).

12. Pectoral fin rays were classified into one of three

categories: (1) entire, (2) lower half free, (3)

completely free.

Head characters:

13. Lateral mouth gape ("mouth width) was measured as the

distance between the left and right posterior junctions

of the upper and lower jaws with the mouth in the fully

open position.

14. Medial mouth gape ("mouth height) was measured as the

distance between the anterior midpoints of the upper and

lower jaws with the mouth in the fully open position.

15. Upper jaw length was measured from the tip of the

premaxilla to the posterior edge of the maxilla.

16. Jaw protrusibility was the ratio of snout length

with the mouth open to snout length with the mouth

closed.

17. Snout length was the distance from the interior surface of

the anterior edge of the bony orbit of the eye to the

anterior margin of the upper jaw at its midpoint. 18. Eye size was the diameter of the eye between fleshy orbits

along an anterior-posterior axis.

19. Position of the eyes involved assigning character states

depending upon whether the eyes were placed (1)

laterally on the head, (2) dorsolaterally, (3)

ventrolaterally.

Dentition characters:

20. Upper and lower jaw dentition were recorded with reference

to three shape descriptors as follows: (1) absent, (2)

villiform (numerous, very small conical teeth arranged

in bands), (3) somewhat larger conical teeth arranged in

single rows, sometimes slightly recurved.

21. Upper and lower pharyngeal dentition were an assignment

using the same three descriptors as were used in the

similar character for jaw dentition (#20 above).

22. Shape vomerine tooth patch was recorded with

reference to the following 4 descriptors listed as

follows: (1) absent, (2) small irregular-shaped patch,

(3) T-shaped flush with palate, (4) T-shaped prominently

raised.

23. Number of basibranchial tooth patches was a count of

these structures which support the floor of the mouth.

Gill raker characters:

24. Number of developed long gill rakers was a total count

on the first gill arch.

25. Inter-raker gap distance was measured as the mean distance between mid-points of the bases of six individuals

anterior rakers.

26. Raker length was measured as the length of the longeBt raker

on the lower limb of the first arch.

27. Filter area, the effective filtering surface area of the

branchial sieve, waB the product of absolute raker length

times lower limb length

Internal body characters:

28. Stomach length was measured as the distance

posterior to the pharynx and anterior to the pylorus.

29. Intestinal length was measured as the distance

immediately following the stomach and terminating at the

anus.

30. Gut length was measured as the length of the

entire alimentary tract posterior to the pharynx.

31. Number of pyloric caeca was a count of the caeca at the

junction of the stomach and intestine.

32. Length of the swimbladder was the ratio of the

length of the swimbladder to the standard length.

33. Width of the swimbladder was the ratio of the

width of the swimbladder to the standard length.

34. Presence of a swimbladder was recorded as a binary

character of presence/absence for each species.

35. Presence of drumming muscles was recorded as a binary

character of presence/absence for each species.

Configuration and shape of feeding apparatus:

36. Species were assigned one of five categories based on overall gross similarity in the configuration and shape of

bony elements comprising the five functional units of the

feeding apparatus as follows: (1) Porogadus pattern.

(2) Bassozetus pattern, (3) Dicrolene pattern,

(4) Barathrites pattern, (5) Acanthonus pattern.

Overall similarity in feeding morphology among species was measured using numerical classification techniques, (cluster analysis, Clifford and Stephenson, 1975). This multivariate technique partitions observations (species) into "clusters" or groupB based on composite similarities or dissimilarities among their attributes (morphological characters). The morphological characters described above and used in this analysis represent character states. Flexible sorting (Lance and

Williams, 1967; Clifford and Stephenson , 1975) with a was used based on a resemblence expressed by the Canberra metric coefficient.

This is a hierarchical, agglomerative clustering strategy in the sense that it determines linkages between individual observations and the entire set of observations by successive fusions or fissions.

Agglomerative refers to the usage of fusions to create groupings.

Fusions are first made between the most similar (or least dissimilar) observations. Repeated fusions are made until all observations form a single group. With each successive fusion new resemblence measures are calculated between the newly formed group and the remaining groups and/or individual observations (Boesch, 1977). Flexible strategies allow the intensity of clustering (i.e. the tendency to form new groups rather than to add on to existing groups) to be altered by changing the value of B, a computational coefficient within the algorithm equations.

The Canberra metric coefficient was used as a resemblence measure in 26 this analysis because equal weighting of all characters is important and quantitative differences between species are to be stressed (Boesch,

1977). The coefficient is calculated as follows:

" in i'S jait1- Where Jik^species compared

N“total number of attributes (characters) considered

Xijavalues of ith character for species j

Xik“values of the ith character for species K Results

Food habits:

Dicrolene introniera. an abundant eurybathic species, occurred on

the Blake plateau and Tongue-of-the-Ocean region between 1086-24Q4m

depths (table 1). A moderate sized fish among species examined, JL*

introniera selected relatively large prey organisms (table 2, figs. 9,

10). Nearly half of the individuals examined contained small amounts of

sediment in the digestive tract. Heavy digene trematode parasite

infestation and light nematode infestation were noted throughout the

alimentary tract.

Dicrolene introniera exhibited one of the most diverse diets of all

species of ophidiids examined. Isopod crustaceans were the most

important food item followed by decapods, polychaetes and copepods respectively. Isopods were first in numerical dominance and frequency of occurrence whereas decapods dominated percent weight. Fish remains and ostracods were less important to the diet. Gastropods, pelecypods and mysids were of minor importance. Although D.. introniera selected relatively large prey in comparison to other ophidiids, ontogenetic

feeding changes of D. introniera shoved a shift from small-sized prey

items in small fish to large-sized prey items in larger fish (table 2,

fig, 11). Smaller-bodied fish utilized smaller ostracods, copepods, mysids, and amphipods to a greater extent while larger fish showed

27 28

increasing reliance on larger decapods isopods and fish. Data depicting

food categories of D.. intronigra and related information are presented

in table 3, fig. 12.

Dicrolene kanazawai. an abundant lower slope species* occurred in

Tongue-of-the-Oceanf between 1298-2250m depths (table 1). This species,

slightly smaller in size than D.. intronigra. selected moderate sized

prey organisms in the diet (table 2, figs. 9, 10). Sediment was

conspicuously absent in all stomach and intestinal tracts examined.

Heavy digene trematode parasite infestation occurred throughout the

alimentary tract in nearly half of all specimens examined. Nematode

parasites were absent.

The diet of D. kanazawai was quite different from that of its morphologically similar congener, D.. introniera. Calanoid copepods and

tube-dwelling polychaetes were the most important food items. Copepods were first in numerical dominance and frequency of occurrence whereas

polychaetes contributed more substantially to percent weight.

Benthopelagic gammaridean and hyperiid amphipods, mysids and to a lesser

extent tanaids comprised the second most important group of prey items

in the diet. Isopods and decapods were less important. Fish remains and ostracods made only minor contributions to the diet. Ontogenetic feeding changes of P.. kanazawai showed a much less noticeable shift from small-sized prey in small fish to large-sized prey in larger fish than was observed in D. intronigra (table 2, fig. 11). All size groups of D. kanazawai utilized similar prey categories and prey sizes, particularly calanoid copepods, however larger fish showed increasing reliance on decapods and larger gammaridean amphipods and isopods. Although P.. kanazawai and D. intronigra shared many food categories in common, the 29 conspicuous absense of sediment, heavy reliance on calanoid copepods, occurrence of pelagic hyperiid amphipods, and decreasing importance of benthic isopods and decapods, suggest the former fed further off the bottom on more pelagic prey than D.. introniera. Data depicting food categories of D. kanazawai and related information are presented in table 4, fig. 13,

Bathvonus nectoralis. a very common and broadly eurybathic species occurred on the lower slope and rise throughout Exuma Sound, Columbus

Basin, and Inagua Basin between l438-5345m depths (table 1). Bathvonus nectoralis. a relatively small species, selected relatively small-sized prey organisms in the diet (table 2 f figs. 9, 10). Sediment occurred in few of the digestive tracts examined. Light digene trematode parasite infestation was observed throughout the digestive tract. Nematode parasites were absent.

Calanoid copepod crustaceans were the most important food item.

Copepods were first in numerical dominance and frequency of occurrence and second in percent weight. Mysids were first in percent weight but third and fourth in numerical dominance and frequency of importance respectively. The relative importance of isopods and mysids in the diet were nearly equal. Amphipods were less important whereas polychaetes and gastropods made only minor contributions to the overall diet. In contrast to species of Dicrolene. ontogentic feeding changes of B.. nectoralis were not evident (table 2, fig. 11). Individuals of all size groups of JB. nectoralis fed predominately on small calanoid copepods and to a lesser degree on other small bentbopelagic crustaceans. Data depicting food categories of B., pectoralis and related information are presented in table 5, fig. 14. Barathrodemus manatinus■ an abundant species, occurred on the lower

slope in TOTO between 1257-2337m depths (table 1). This species is also

relatively small-bodied and similar in size and shape to JB. nectoralis

(figs. 3d, 5a,b,c, 15). Barathrodemus manatinus. fed on relatively

small-sized prey organisms (table 2, fig. 9). Sediment was present in

5Z of the digestive tracts examined. Light digene trematode parasite

infestation was noted in the digestive tract in nearly half of all

specimens examined. Nematode parasites were absent.

Barathrodemus manatinus was unique among species examined in that

small-sized benthic tanaid crustaceans were the most important food

item. Tanaids were first in numerical abundance, frequency of occurrence and percent weight. Calanoid copepods followed tanaids in

importance ranking second in numerical abundance, frequency of occurrence and third in percent weight. Tube dwelling polychaetes were

third in relative importance, frequency of occurrence and second in

percent weight. Gammaridean amphipods and ostracods were also important food items ranking third and fourth respectively in relative importance.

Cumaceans and fish remains were less important ranking sixth and seventh respectively. Pelecypods and mysids were only of minor importance.

Ontogenetic feeding changes were not observed in various size groups of

B.. manatinus (table 2, fig. 16). Individuals of all size groups fed predominately on small tanaids and calanoid copepods and to a lesser extent on other small benthopelagic crustaceans. Data depicting major food categories of this species and related information are presented in table 6, fig. 17.

BassozetuB normalis. an abundant species, occurred on the lower slope of the Tongue-of-the-Ocean between 1271-2960m depths (table 1). This species, which attained a relatively large body size, selected

larger prey organisms over a greater size range than most species examined (table 2, figs. 9, 15). Sediment occurred rarely in the stomach and intestinal tract. Very heavy nematode infestation was noted

in the majority of digestive tracts examined. Nematode infestation in some individuals was so extensive that the parasites nearly occluded the alimentary canal. In contrast, digene trematode parasite infestation was relatively light.

Decapod crustaceans were the most important food item. However, the IRI for decapods was greatly influenced by the extremely high value for percent weight, a component used to calculate the IRI index.

Decapods were also first in numerical dominance and second in frequency of occurrence. Gammaridean amphipods ranked second in relative importance followed closely by mysids. Mysids were second in percent weight but contributed significantly less to diet in this category than decapods. Fish remains, copepods, polychaetes and gastropods were less important to the diet respectively and made similar contributions to the

IRI. Tanaids, cumaceans, and isopods were of minor importance to the diet. Ontogenetic feeding changes of B.. normal is from the Bahamas showed a sharp increase from small benthopelagic prey organisms in smaller-sized fish to larger prey organism in larger fish (table 2, fig.

16). Smaller fish utilized small benthopelagic crustaceans such as calanoid copepods, and gammaridean amphipods to a greater extent while larger fish showed an increasing reliance on large decapod crustaceans and fish. The occurrence of pelagic hyperiid amphipods, benthopelagic copepods and gammaridean amphipods, and benthic cumaceans, isopods and tanaids, suggest B.. normalis is a highly mobile somewhat generalized 32 predator that fed on a wide variety of kinds, shapes and sizes of prey organisms. Data depciting major food categories of B.. normal is and related information are presented in table 7, fig. 18.

Bassozetns taenia, a common and extremely eurybathic species, occurred on the lover rise in 1559-5267m depths (table 1). This species, which also attained a relative large size, selected smaller prey items over a narrower range than did closely related and morphologically similar B. uormalis (table 2, 9, 15). Differences between these two species diet were probably due in large part to inadequate sample size in B,. taenia. Only 18 specimens of this species were available for examination in contrast to 77 specimens of B.. normalis examined. Sediment did not occur in any of the digestive tracts examined. Nematode parasite infestation was very extensive throughout the digestive tract in a majority of specimens examined.

Digene trematode parasite infestation was by comparison very light.

Decapods and gammaridean amphipods were the most important food itemB in the diet. Decapods were first in numerical dominance and percent weight and third in frequency of occurrence. Gammaridean amphipods were second in relative importance, first in frequency of occurrence and second in percent weight and numerical dominance.

Tanaids and mysids were third and fourth respectively in relative importance and possessed similar values for percent weight, frequency of occurrence and numerical dominance. Tube-dwelling polychaetes, cumaceans, isopods and calanoid copepods respectively were the least important prey items in the diet. Ontogenetic feeding changes in the diet of B taenia could not be determined due to inadequate sample size.

However of those few specimens examined decapods primarily occurred in 33 the diets of the large-bodied individuals and were absent from the diets of small-bodied individuals. Bassozetus taenia exhibited a feeding strategy similar to that displayed by its congener B. normalis. Highly mobile, these species fed on a wide variety of shapes, kinds and sizes of prey organisms. Data depicting food categories for B.. taenia and related information are presented in table 8, fig. 19.

Few specimens of the abyssal non-abundant species, A. edentatum were available for study. However, the remains of squid beaks from the epipelagic family Ommastrephidae and fish bones in the intestine suggest this species may Bcavange the bottom for food (Carter, 1984). Its large mouth opening may enable it to utilize a great size range of prey items.

In this respect its feeding strategy is similar to large-bodied individuals of Bassozetus normalis.

Xvelacvba mversi. a non-abundant eurybathic species, occurred on the lower slope between 1296-1977m depths (table 1). This species, which attained a large body size, fed on relatively large sized prey organisms over a wide size range (table 2, figB. 9, 20). Sediment did not occur in any of the digestive tracts examined. Digene trematode and nematode parasite infestation were light throughout the digestive tract in a small percentage of specimens examined.

Benthic isopod crustaceans were the moBt important food item.

Isopods were first in numerical dominance, percent weight and frequency of occurrence. Fish remains, tube-dwelling polychaetes calanoid copepods and decapods respectively were next in importance as food items. Ostracods, mysids and tanaids were less important as food.

Ontogenetic feeding changes in the diet of this species could not be determined due to inadequate sample size. However, of the eight 34 specimens examined containing food items, only the larger individuals

(>300 mm SL) contained decapods and fish remains in the digestive tract.

These results suggest that this species, like other ophidiids that attained a fairly large size, showed increasing reliance on larger more mobile prey in larger individuals. Data depicting major food categories of X. mversi and related information are presented in table 9, fig. 26.

Acanthonus armatus. an abundant and broadly eurybathic species, occurred on the lower slope and rise in Tongue-of-the-Ocean, Exuma

Sound, Columbus and Inagua Basins between 1586-3094m depths (table 1).

This species, which attained a large size and possessed a massively swollen head, selected moderate sized prey over a wide size range (table

2, figs. 9, 20). Sediment did not occur in any of the digestive tracts examined. Digene trematode and nematode parasites were absent.

Tube-dwelling polychaetes were the most important food item in the diet. Polychaetes were first in numerical dominance, frequency of occurrence and percent weight. Gammaridean amphipods, ostracods and calanoid copepods respectively were also important food items. Copepods and ostracods shared similar IRI values whereas the IRI for amphipods was somewhat higher. Mysids were less important as food and gastropods, tanaids, isopods and cumaceans were only occasionally taken in the diet.

Ontogenetic feeding changes in the diet of A. armatus were not evident in the specimens examined. Individuals of all size groups fed on similar size prey items. Data depicting major food categories of A* armatus. and related information are presented in table 10, fig. 22.

Poroeadus miles, a non-abundant and eurybathic species, occurred on the lower slope region between 1360-3032m depths (table 1). Poroeadus miles. which attained a relatively large size, selected moderate-sized 35 prey organisms (table 2, figs. 9, 20). Sediment was not observed in any of the digestive tracts examined. Digene trematode and nematode parasite infestations were moderate to heavy throughout the digestive tract in a majority of specimens examined.

Gammaridean amphipod crustaceans were by far the most important food item for Poroeadus miles. Gammaridean amphipods ranked first and predominated in numerical abundance, frequency of occurrence and percent weight contributions in the diet. Decapods and mysids were nearly equal in their relative importance and together made only a minor contribution to the diet. Ontogenetic feeding changes in the diet of P.. miles could not be determined due to inadequate sample size. Data depicting major food categories of P. miles and related information are presented in table 11, fig. 23.

Poroeadus ailus. an abundant and broadly eurybathic species, occurred on the lower slope and rise of Exuma Sound, Columbus and Inagua

Basins in 1586-2728m depths (table 1). Poroeadus silus. a relatively small-sized species, selected small prey organisms as food items (table

2, figs. 9, 24). Sediment was not present in any of the digestive tracts examined. Digene parasite infestation was very light throughout the digestive tract in a small percentage of specimens examined.

Nematode parasites were absent.

Porogadus siluB displayed a somewhat specialized feeding habit in that small-sized calanoid copepods dominated the diet. Copepods were first in numerical dominance, percent weight, and frequency of occurrence. Mysids, cumaceans, tanaids and ostracods decreased respectively in importance as food and were of only minor importance to the overall diet. Ontogenetic feeding changes were not observed in 36 various size groups of P.. silus (table 2, figs. 9, 16). Small sized calanoid copepods dominated the diet of individuals of all size classes.

This species heavy reliance on small copepods, an absence of sediment in the digestive tract and the minor importance of benthic prey in the diet, all suggest anoff-bottom euryphagous planktivorous feeding habit.

Data depicting food categories of P. situs and related information are presented in table 12, fig. 25.

Poroeadus catena, a non-abundant and eurybathic species, occurred on the lower slope and rise in 2 4 0 4 - 4 9 3 0 m depths (table 1). This species, which is very similar in morphology to P.* Silas.* differed in that it attained a larger body size (table 2, fig. 24). PorogaduB catena also selected relatively small-sized prey organisms. Sediment did not occur in any of the digestive tracts examined. Digene trematode and nematode parasites were absent.

Poroeadus catena also displayed a somewhat specialized feeding habit similar to that reported for for P. silus. Small sized calanoid copepods were the most important food item. Copepods were first in numerical dominance, percent weight and frequency of occurrence.

Cumaceans were second in numerical dominance, frequency of occurrence and percent weight. Ostracods were of less importance and tanaids and mysids were of only minor importance. Ontogenetic feeding changes in the diet of this species could not be determined due to inadequate sample size. However, this species probably followed the same pattern described for closely related P.. silus. The diet of P^. catena suggests an off-bottom euryphagous planktivorous feeding habit. Data depicting food categories of P.. catena and related information are presented in table 13, fig. 26. Barathrites parri. an abundant species, occurred on the lower slope of the Tongue-of-the-Ocean in 1271-2337m depths (table 1). This

species, which attained a moderate size, selected relatively large prey

items over a wide range (table 2, figs. 9. 24). Sediment occurred in

252 of all the digestive tracts examined. Moderate digene parasite and

light nematode infestation was noted throughout the digestive tract in

two-thirds of all specimens examined.

Barathrites parri exhibited the most distinct and specialized diet among all species examined. Tube-dwelling polychaetes and elasipod holothurians dominated the diet respectively. Tube-dwelling polychaetes were first in numerical dominance, percent weight and frequency of

occurrence. Elasipod holothurians, also important prey items, were

second in numerical dominance, percent weight and frequency of

occurrence. Copepod crustaceans were of only minor importance.

Ontogenetic feeding changes in the diet of B.. parri were not observed in

the specimens examined. Frey size may not be an important factor since

it appears that this species has evolved a distintive feeding complex

that enables it to tear bits and pieces of tissue away from whole prey organism. Nearly all prey organisms found in the digestive tract of

this species were torn and shredded. Data depicting food categories B.. parri and related information are presented in table 14, and fig. 27.

Penopus macdonaldi. a non-abundant and eurybathic species, occurred on the middle slope between 1257-2700m depths (table 1). This species, which attained a small to moderate-sized body, selected relatively small-aized prey organisms as food (table 2, fig. 9, 24). Sediment was found in the digestive tract of all specimens that contained food items.

Unfortunately the majority of specimens examined contained little food in the digestive tract. Of those that did only small benthic isopods,

sediment and unidentified crustacean fragments comprised the diet.

However, the presence of benthic isopodB and sediment in the digestive

tracts suggest this species fed on or near the bottom. Little more can

be said concerning the diet of P.* macdonaldj until more data are

available. Data depicting food categories for P.. macdonaldj and related

information are presented in table 15. Table 1. Ophidiid Fishes Examined in Food Habit Study.

Size Percent Percent Range Depth Range Number with with Species (SL mn) Mim'mm Maarimm Examined Food Sediment

Dicrolene intronigra 127-363 1086 2404 77 79.5 45

Dicrolene kanazawai 64-279 1298 2250 100 94.3 0

Bathvonus nectoralis 71-178 1438 5345 130 86.1 3

Bassozetus normalis 178-519 1271 2960 77 80.7 4

Bassozetus taenia 137-386 1559 5267 18 78.3 0

Xvelacvba nrversi 88-422 1296 1977 17 50.0 0

Barathrites narri 115-336 1271 2337 68 32.0 25

Ibroaadus silus 90-160 1586 2728 124 72.8 0

Ibroeadus catena 130-250 2404 4930 29 55.0 0

Poroeadus miles 181-439 2115 3083 12 41.6 0

Penopus macdonaldi 105-237 1257 2700 29 13.7 100

Barathcodpmta manatinus 78-172 1257 2337 122 86.5 5

Acanthoma armatus 86-357 1586 3094 100 67.0 0 Table 2. Size Range of Predators and Corresponding Prey.

Size Range Prey Size Snecies (SL mn) X NSD Ranee (mn) X N SD

Bassozetus normalis 178-519 368 77 84 2.0-56.1 13.6 68 2.1

Acantbonus armatus 86-357 211 100 127 1.0-32.0 6.1 78 2.4

Bnrfithroderus mnnatinufl 78-172 138 122 18 0.3-11.2 2.65 283 1.1

Dicrolene kanazawai 64-279 188 100 55 1.2-16.2 4.1 204 1.5

Bassozetus taenia 137-386 259 18 67 1.5-15.0 4.9 43 2.0

Dicrolene intronisra 127-363 254 77 57 1.2-70.5 8.3 84 3.5

Xvelacvba mversi 88-422 225 17 94 1.5-60.0 12.0 24 6.3

Barathrites narri 115-336 213 68 46 3.2-18.5 12.2 29 1.8

Penorus macdonaldi 105-237 183 29 33 1.1-10.3 3.6 4 4.4

Bnroeadus miles 181-439 318 12 85 2.5-18.2 6.6 12 2.4

Rsrofcadus Bilus 90-160 128 124 13 1.0-15.2 3.3 201 1.0

Poroeadus catena 130-250 187 29 38 1.5- 8.2 2.8 26 1.5

Bathroms nectoralis 71-178 134 130 24 1.0-14.2 4.5 151 1.3 Figure 9. Mean prey length versus mean standard length of ophidiid species. MEAN PREY SIZE (mm)

_ — ro 4 * o 00 01 o m o o o o ro_i______I______I______I____ — I— I— I— I— I— I

—c£)------Porogadus si/us 1 | i Bothyonus pec tor ah's ~ -C p Barathrodemus manatinus

oo- , 0 i [ i— Penopus macdonaidi Porogadus catena «o- “T—M Dicrotene kanazawai o ro 8' ro q~| r I I—— ------Acanthonus armatus -Cp Barathrites parri ro ro- o ro 1 Xyelacyba myersi oi- O * = > Dicrotene intronigra ro 8 " ' Bassozetus taenia

w. o ' oj — I r I------Porogadus miles ro - o os- Bassozetus normalis Figure 10. Length frequency histograms for examined species of D. introniera. D. kanazawai and ]i. pectoralis. Dicrotene intronigra n = 7 7

2 0 -

10 -

0 ~T~ I 150 200 2 5 0 3 0 0 3 5 0

24 Dicrotene kanazawai n = 100

0 “ 1— I— f— |— |— I— | I | I 50 100 150 200 250 300 —

3 0 - Bathyonus pectora/is n = 1 3 0

2 0 -

10 - 1 ,.

0 —] 1 1 1 1 1 1 1 1 1 60 72 84 96 108 120 132 144 156 168 180 STANDARD LENGTH (mm) Figure 11 . Mean prey length versus standard length groups of D. intronigra. D.. kanazawai and B_. pectoralis. o ■ CO

* 4

- d > N 1 ■ fO ttl <0 ■O §

00 dp- ~r~ "T" "T* ~i 1--- 1--- r “ I I- to (0 CM O CO l£> 9J" CM o CO E E

4> o |- ! o a N z I X - 4 3 - to I i-8 iz

3- T" T- ~i i i i i <0 9* CM O CD (O ^ CM o

-8 IO

f to • intronigra. Percent frequency of occurrence (F), percent nurerical abundance (N), percent weight (V), index of relative importatance (IRI) of food items, absolute total nurber (AIN) and absolute total weight (A3V).

TAXTN F W N m i A3N Altf

Polychaeta 30 3.5 9.0 370 47 0.8644

Mollusca Gastropoda 5 0 . 1 3.4 19 18 0.0120 Belecypoda 3 0 . 1 3.8 9 0.1 0.0048 Unidentified mollusca 5 0 . 1 1.2 6 0.1 <•0005 Total Mollusca 13 0 . 1 8.2 106 43 0.0173

Crustacea Ostracoda 16 0.1 4.1 69 22 0.0417 Gopepoda 30 0.6 9.7 306 50 0.1580 Calanoid Mysida 3 0 . 1 0.7 2 4 0.0060 Isopoda 57 22.8 48.3 4033 252 5.6772 Anthuridae Neoanthurus caeca Asellota Amphipoda 24 4.4 9.0 324 47 1.088 Gamnaridea Synopiidae Decapoda 16 68.0 4.8 1182 25 16.9624 Glvobocraneon s d . Unidentified Crustacea 11 0.4 2.8 34 15 0.0892 Total Crustacea 84 96.3 7 9 3 14713 412 24.005

Teleostei 19 0.2 4.8 95 25 0.0420

Nurber of fish examined 77 Nunfeer of prey items 522 Percent with food 79.5 pprrpnt with Rpciimpnt 45 Percent with digene parasite infestation 85 Percent with nematode parasite infestation 26

Digene infestation heavy, nematode infestation light Figure 12. Percent frequency occurrence, percent number, percent weight, and index of relative importance (iRl) of higher taxonomic groups of food in the diet of Dicrolene intronigra. POl. YCHAETA }70 11 ir r n"ii ii i r i i i i i i " n i r i—r ii ’c . 2 * L ' ^ S'® rs ? X - X o h* Q C Ow C C T 83BNM % a ~ > M Z * 5 = 1UOI3M % I £ S

16% i I s 3 Table 4. Gut Contents of Dicrolene kHimmmai. Percent frequency of occurrence (F), percent mmerical abundance (N), percent weight (w), index of relative importance (iRl) of food items, absolute total nuofeer (AIN), and absolute total weight (AlW).

TAXON______F w N mi A3M Am

Polychaeta 67 49.3 18.4 4518 158 2.1616 Eblynoidea

Crustacea Ostracoda 1 0 . 1 0.2 1 2 0.0160 Cbpepoda Calanoid 70 16.3 41.4 4059 355 0.7136 Hysida 31 7.1 7.4 449 66 0.3120 Cunacea 4 0.6 0.7 5 6 0.0282 Tanaidacea 25 2.1 9.6 287 82 0.0898 Isopoda 11 1.1 2.9 44 25 0.0496 Anphipoda 31 6.4 9,3 487 79 0.2818 Hyperidae Gmmaridgfl Eusiridae Fbachotropia s d . Pardaliacidae Synopiidae Syj-rhoe sp. Lysianasidae Hyperiopsidae BvDeriosis s d .

Decapoda 10 7.0 1.9 88 16 0.2740 Glvnhocran&on aculeata Glvnhocraneon so. Unidentified Crustacea 41 9.7 7.9 716 68 0.4236 Total Crustacea 90 50.4 81.3 11873 698 2.1886

Nurber of fish examined 100 Nicber of prey items 859 Percent with food 94.3 Percent with sediment 0 Percent with digene parasite infestation 42 Percent with nematode parasite infestation 0

Digene infestation heavy Figure 13. Percent frequency occurrence, percent number percent weight, and index of relative importance C(lRl) of higher taxonomic groups of food in the diet of Dicrotene kanazawai. Q U - "“ I LU U- O Q O A.UPMPODA n i n I Q ft* O T § ft* O Q O c! CO 3 CQ * CB

•Jj7* 5 3 < \j u \~i 5 8 g O Co n 5f•n s cb U30HnN% T I r LU a £ O O ii r i—i ii i l I I i

Table 5. Gut Contents of Bathvonus pectoralis. Percent frequency of occurrence (F), percent miner ical abundance (N), percent weight (w), index of relative importance (mi) of food it erne, absolute total m uter (AIN), and absolute total weight (AIW).

taxcm ______f w n n r m i aw

Bolychaeta 4 5.1 4.0 41 49 0.3720

Mollusca Gastropoda 2 <0.1 3.0 7 37 0.0601

Crustacea Copepoda Calanoid 38 23.9 31.3 2087 389 1.7400 Mysida 22 27.3 12.1 875 150 1.9824 Isopoda 24 22.6 17.2 972 213 1.6416 Anphipoda 20 9.2 12.2 426 151 0.6672 Ganmaridea PhoxDcephalidae Harrnnin excavata Harmnia s d . Unidentified Crustacea 44 11.8 20.2 1424 251 0.8604 Total Ccuatacea 100 94.8 92.9 18770 1124 6.8916

Number of fish examined 130 Number of prey items 1243 Percent with food 86.1 Percent with sediment 3 Percent with digene parasite infestation 34 Percent with nematode parasite infestation 0

Digene infestation lig h t Figure 14. Percent frequency occurrences percent number percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Bathvonus oectoralia. % % "WCHT % NUMBER 100 0 0 1 20 60 10 80— 90 — SO — 40 70 30 — 20 - Q I 30 — 40 — SO— 90 — 60— 80 — 70— 0 — — — — — — — —

38% . etrls _ pectoralis B. ------24% B FREQUENCY % ------2 2 c % C C A A D D B E F F MHPD 426 AMPHIPODA OYHEA 41 GASTROPODA POLYCHAETA ISOPODA ISOPODA OEOA 2087 COPEPODA YIA 875 MYSIDA 20 D % 4% - I 2 % IRI 972 7 Figure 15 • Length frequency histograms for examined species of B.. manatinus. B. normalis and B.. taenia. NUMBER OF INDIVIDUALS 0 2 30 0 2 - 0 1 10- - 4 - 3 2 6 - 5 7

- - - - 120 - 100 aszts taenia Bassozetus aszts normatis Bassozetus Borathrodemus — i— — 62 manat manat inus 122 n : 200 180 STANDARD LENGTH (mm) n= 77 102

300 210

~I— 122

0 360 300 0 500 400 ~I— 142

162 i 420 600 Figure 16 . Mean prey length versus standard length groups of B.. manat inus . B. normalis and P. silus. r* 2 -C t>

■c Jd — CM «05» • ro H = t > -

{ -S l —I | I— , O ~1 | i i i i i r~ <7* tD N O C ID ^ N

in -8 E

O b — r z Ul ■ mo -I 5:I K> — I | h X CO I» N C> o b» ■«n CM I — i [ t— r~ t V 1 I I o o ~o r o in o to o CJ — — 00 to c*cw

CM -to

CM • IO 3

. CM S'* < tj . CM I5 §5 o» “i i ------1 1------r CMf- CM o co to

(uiuj) 3Z IS A3Hd NV3W Table 6. Gut Contents of Barathrodaras manatims. Percent frequency of occurrence (F), percent numerical abundance (n ), percent weight (W), index of relative importance (Utl) of food it erne, absolute total timber (/IN) and absolute total weight (AIW).

TAKEN______F W N IRI ATO AIW

Bolychaeta 27 24.0 5.1 788 66 0.6300 Pblynoidea Riyllodocidae

Mollusca Pelecypoda 4 <0.1 1.0 3 13 0.0021 Total Mollusca 4 0 .1 1.0 3 13 0.0021

Crustacea OBtracoda 20 2.5 5.8 165 75 0.0600 Copepoda Calanoid 40 11.4 19.4 1233 252 03100 Mysidacea 1 1.0 0 .1 1 2 0.0260 Ckmacea 7 4.1 3 3 52 43 0.1050 Tanaidacea 69 29.5 44.6 5148 578 0.7752 Anphipoda 22 10.6 4.9 347 63 0.2787 fhrrmnr-iflpn Phoxocephalidae Harpinia sp. Lysianasidae Unidentified Crustacea 75 16.4 14 3 2311 185 0.4200 Total Crustacea 99 75.5 92.6 16612 1201 1.9815

Teleostei 8 0 .1 1.6 17 20 0.0123

Nuoier of fish examined 122 Nmber of prey items 1297 Percent with food 86.5 Percent with sediment 5 Percent with digene parasite infestation 42 Percent with nematode parasite infestation 0

Digene infestation light Figure 17. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Barathrodemus manatinus. — I y

* i r

* 2 £.

58

£ 22

i i i — i— r - "TT t i i r— / ! f f I ■ i

j //*>/.v .ii i, Table 7. Gut contents of Bassozetus normalis. Percent frequency of occurrence (f) , percent numerical abundance (n) > percent weight (w), index of relative importance (HU) of food items, absolute total nudber (A3N) and absolute total weight (AIW).

TAXCN F W N IRI AIK Arw

Polychaeta 3 2.5 2.0 16 8 0.9272

Mollusca Gastropoda 3 0.1 0.3 11 1 0.0284 Unidentified items 3.5 <0.1 0.1 0.7 - 0.0194 Total Mollusca 6.5 0.2 0.3 11.7 1 0.0478

Crustacea Ob traced 2 0.1 1.0 2 4 0.0100 Copepoda Calanoid 5 0.8 3.0 20 12 0.2984 Ifysida 34 5.5 22.2 122.4 87 2.0472 Cunacea 2 0.3 1.0 2 4 0.1232 Tanaidacea 2 0.5 1.0 3 4 0.1904 Isopoda 2 0.1 1.5 2 6 0.0276 Amphipoda 16 4.0 15.2 297 59 1.4840 Hyperidae Garmaridpfl Synopiidae Lysianassidae Decapods 33 84.8 24.2 3573 95 31.5324 Benthisienus bartletti GlvDhocranston loneirostris Hvmenonaneus laatis Solenoceridae Unidentified Crustacea 36 1.3 21.2 816 83 0.4920 Total Crustacea 95 97.4 89.9 17762 352 36.2052

Teleostei 9 0.2 5.1 44 0.1 0.0092

Nmber of fish examined 77 limber of prey items 392 Percent with food 80.7 Percent with sediment 4.0 Percent with digene parasite infestation 38 Percent with nematode parasite infestation 80

Digene infestation light, nematode infestation heavy Figure 18. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Bassozetus normalis. IRi

A DECAPODA 2 5 'J

B MYS1DA 956

C AM PH IPOD A 29' too D TELEOSTEI 44 90. E COPEPODA 20 SO F POLYCHAETA 16

70 G GASTROPODA II

60 H t a n a i d a c e a K

50 I CUMACEAS B. normalis J ISOPODA 40 K OSTRACODA JO

20'

10

0 9% 1 0 - 16% J% 2% J4% 5%

2 0-

JO'

40- % FREQUENCY 50•

60 •

70-

SO­

HO' JJ% too< Table 8. Gut Contents of Bassozetus taenia. Percent frequency of occurrence (F), percent numerical abundance (N), percent weight (W), index of relative inportance (iRl) of food items, absolute total number (AIN) and absolute total weight (ATW).

TAX® F W N 1RI AIN ATW

Polychaeta 5 2.4 6.2 43 6 0.03948 Polynoidea

Crustacea Amphipoda 24 9.5 21.5 744 20 1.5629 Ganmaridea Synopiidae Tanaidacea 23 8.8 10.4 441 10 1.4477 Hysida 16 8.2 11.4 313 10 1.3490 Cbpepoda Calanoid 2 2.5 5.1 15.2 5 0.4113 Cunacea 8 1.0 2.1 24 2 0.1645 Isopoda 5 1.2 2.8 20 3 0.1974 Decapoda 22 53,2 35.6 1953 38 8.7524 Glvnhocranson sn. Panaeidea ttvmenonaneus sn. Unidentified Crustacea 68 13.2 4.9 1230 5 2.1716 Total Crustacea 99.6 97.6 93.8 19063 86 16.0568

Nwber of fish examined 92 Number of prey items 18 Percent with food 78 .3 Percent with sediment 0 Percent with digene parasite infestation 59 Percent with nematode parasite infestation 82

Digene infestation light, nematode infestation heavy Figure 19. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Bassozetus taenia. B. taenia IRI 100 A DECAPODA 1606 90 B AMPHIPODA 744 80 C TANAIDACEA 441 70 D MYSIDA 60 E COPEPODA I f 50 F POLYCHAETA 43 40 A G CUMACEAN 24 30. ISOPODA 20 B H 20 D 10

0 f f 10' 16% 8% 5% 22 % 23% 20 2'% 5%

30-

40.

50 % FREQUENCY

60- 2 4 %

70-

80-

90-

100 Figure 20. Length frequency histograms for examined species of P. miles. X. mversi and A. armatus. Porogadus mites n= 12

—i------1------1------1------1------T------1------1 200 3 00 400 500 600 Xyetacyba myersi n = 17

n 1--- 1---- 1— ~\-- 1----1------r — r 100 200 300 400 500 Acanthonus armatus n s 100

1 1 1 1 1 1 1 1 1 100 160 220 280 340 STANDARD LENGTH (mm) Table 9. Gut Contents of Xvelacvba mversi. Percent frequency of occurrence (f ), percent numerical abundance (n ), percent weight (W), index of relative importance (IRI) of food items, absolute total nurber (AIN) and absolute total weight (A3W).

w a n F W N IRI AIN AW

Polychaeta 36 4.0 16.1 723 7 0.2800

Crustacea OBtracoda 7 0.3 6.5 47 4 0.0220 Gopepoda Calanoid 21 0.7 12.9 285 7 0.0510 Ifysida 7 0.1 3.2 23 2 0.0028 7 0.1 3.1 23 1 0.0020 Isopoda 57 11.5 48.4 3414 26 0.8038 Asellota sn. Decapoda 7 20.1 3.2 163 2 1.3968 GlvDhocraneon loneirostris GlvDbocranmon aculeata Glvphocraneon sp. Unidentified Crustacea 10 0.1 4.8 49 3 0.0050 Total Crustacea 79 32.8 80.6 8958 43 2.2752

Teleostei 7 67.2 3.2 492.8 2 4.6587

Nmber of fish examined 17 Nucber of prey items 54 Percent with sediment 0 Percent with food 58,8 Percent with digene parasite infestation 20 Percent with nematode parasiteinfestaion 10

Digene and nematode infestation light Figure 21. Fercent frequency occurrence, percent number, percent weight, and index of relative importance (XRl) of higher taxonomic groupB of food in the diet of Xvelacyba mversi. % % w e ig h t * NUMBER 100 80— SO— — . yersi m X. FREQUENCY % C G E A H D B F ASI CEa AC SAID TA SD Y A SID M POLYCHAETA DECAPODA OSTRACODA TELE0STE1 COPEPODA SPDA 1S0P0D F H G F E

¥ '¥ ■

□

7 % 28ST 219 IRI 603 23 49 47 68 23

Table 10. Gut Contents of Acanthoma armatus. Percent frequency of occurrence (f ) , percent mmerical abundance (N), percent weight (W), index of relative importance (IRI) of food item, absolute total umber (AIN) and absolute total wight (AlW).

TARNFW N m i A3N AlW

Folychaeta 80 82.0 34.3 9384 166 7.7928 Polynoidea

Molluscs Gastropoda 2 0.1 2.8 6 14 0.0065

Crustacea Ostracoda 37 3.2 16.8 737 83 0.2995 Copepoda 33 4.1 14.0 591 68 0.2995 Calanoid Mysida 15 2.6 6.3 136 31 0.2488 Cunacea 2 0.2 0.7 2 5 0.0187 Tanaidacea 4 0.2 1.4 7 7 0.0153 Isopoda 2 0.1 0.7 2 5 0.0139 Amphipoda Ganmaridea 46 6.7 21.7 1296 107 0.6351 Synopiidae Unidentified Crustacea 5 0.1 0.1 1 0.0340 Tbtal Crustacea 85 17.2 61.7 6630 296 1.6561

Nunber of fish examined 100 Nuiber of prey items 486 Percent with food 67.0 Percent with sediment 0 Percent with digene parasite infestation 0 Percent with nematode parasite infestation 0 Figure 22. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Acanthonus armatus. m n n u t/ t ww ca < ? V W \ V . f t b tfi'>U II &

Ml** Table 11. Out Contents of Borogadus miles. Percent frequency of occurrence (F), percent numerical abundance (N), percent veight (w ) , index of relative importance (iRl) of food items, absolute total number ( A m ) , and absolute total weight (AIW).

TASK F W N IRI AIN AIW

Crustacea Mysida 14 2.2 5.3 107 2 0.0096 Anphipoda Garmaridea 57 81.3 73.7 8858 21 0.3502 Steeocebbalidae Decapods 14 4.6 5.3 140 2 0.0196 Hvmenonaneus nereus Unidentified Crustacea 43 11.9 15.8 1186 4 0.0512 Total Crustacea 100 100 100 20000 29 0.4306

Ninber of fisb examined 12 Umber of prey items 29 Percent with food 41.6 Percent with sediment 0 Percent with digene parasite infestation 83 Percent with nematode parasite infestation 80

Digene and nematode infestation moderate Figure 23. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Poroeadus miles. IRI P. miles A AMPHIPODA 8858 B DECAPODA 140

C M YSID A 107 A

B C

14% l4%

% FREQUENCY Figure 24. Length frequency histograms for examined species of P. silus. P,. macdonaldi. P., catena and B.. pectoralis. in 10

CM

CM

<4 CO CM 5> u>

CVJ GO

CM

CM STANDARD LENGTH (mm)

to

©

sivnaiAiam jo asswriN Table 12, Gut Contents of Boreadus silus. Percent frequency of occurrence (F), percent nunerlcal abundance (N), percent weight (w), index of relative importance (iRl) of food items, absolute total number (AIN) and absolute total weight (AItf).

TAXON F W N IRI A m ATW

Crustacea Ostracoda 4 2.1 5.7 30 56 0.0558 Copepoda Calanoid 77 64.0 75.5 10732 745 1.6956 Mysidacea 4 16.7 3.8 79 37 0.4410 Cunacea 8 5.6 3.8 72 38 0.1494 Tanaidacea 4 8.0 3.8 45 37 0.2106 Unidentified Crustacea 15 3.6 7.5 172 74 0.0954 Total Crustacea 100 100 100 20000 987 2.6478

Number of fish examined 126 Number of prey items 987 Percent with food 72.8 Percent with sediment 0 Percent with Higpnp parasite infestation 11 Percent with nematode parasite infestation 0

Digene infestation light Figure 25. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Poroeadus silus. IRI

A c o p e p o d a 10732

B CUMACEAN 72

C OSTRACODA 30

D TANAIDACEA 45

E m y s i d a 79 100-

80

X, £ so I § 40 J* JO

4%

% FREQUENCY

77%

80

100 Table 13. Out Contents of Bproaadus catena. Percent frequency of occurrence (f) » percent nunerlcal abundance (n ) , percent weight (W), index of relative inportance (mi) of food item* absolute total nuuber CAIN) and absolute total weight (ATW).

F W N IRI AIN AIW

Crustacea Oatracod 8 11.8 6 3 144 5 0.02147 Copepoda Galanoid 65 51.6 38.4 5850 32 0.0942 Mysida 3 3.1 4.1 21 3 0.0056 (Xmacea 10 18.3 21.7 400 18 0.0334 Tanaidacea 5 8.4 4 3 65 5 0.0153 Unidentified Crustacea 35 6.8 25.2 1120 20 0.0124 Total Crustacea 100 100 100 20000 83 0.1816

Nmber of fish examined 29 Nurber of prey items 83 Percent with food 55 Percent with sediment 0 Percent with digene parasite infestation 0 Percent with nematode parasite infestation 0 Figure 26. Percent frequency occurrence! percent number, percent weight, and index of relative importance (IRI) of higher taxonomic groups of food in the diet of Porogadus catena. B catena 100' IRI

90- A C0PEP0DA 5850

80' B CUMACEAN 4 0 0

7 0 C OSTRACODA 144

60- D TANAIDACEA 6 5

50- E MYSIDA 21

40-

30. B 20.

10. C —1 D 0-

IQ- 3% 20. 'o 8% 30. 10%

4 0 65% 5 0

60. % FREQUENCY

7 0

8 0

90-

100- Table 14. Gbt Contents of Barathrites parri. Percent frequency of occurrence (f), percent numerical abundance (N), percent weight (W), index of relative importance (ntl) of food items, absolute total nurber (AIN), absolute total weight (ATW).

______T&KCN F W H IRI A3K ATW

Elasipoda 52 63 28 4732 30 3.0648

Rjlychaeta 98 37 72 10682 21 1.8792 Ouuphidae. Hvalinocia ap.

Crustacea Copepoda Calanoid 2 <0.1 35 7 2 0.0049 ftjnfcer of fish examined 68 Nunfcer of prey it one 51 Percent with food 32 Percent with sediment 25 Percent with digene infestation 38 Percent with nematode infestation 15

Digene infestation moderate, nematode infestation light Figure 27. Percent frequency occurrence, percent number, percent weight, and index of relative importance (IRl) of higher taxonomic groups of food in the diet of Barathrites parri. % WEIGHT % NUMBER 100 100 - 0 5 0 — 40 20 30 80 O—— JO 90 60 0 I — 90 20 40 SO 60 SO - SO- 10 0 • 70 0 H - 70 10 0 - — - - — - — — • - —— - - —1

. parri B. 98% — ------FREQUENCY %

3 ------52 B % A C B POLYCHAETA HOLOTHURIANS COPEPODA - 3 2%

10682 IRI 4732 II Table 15. Out Contents of Penonus macdcoaldi. Percent frequency of occurrence (F), percent nuaerical abundance (N)> percent weight (W), index of relative importance (iRl) of food items, absolute total lumber (A3ft) and absolute total weight (ATW).

TA350N FWH IRI ATO A3W

Crustacea Isopoda 100 88 88 17000 7 0.0336 Asellota Unidentified crustacea 25 12 12 3000 1 0.0046 Tbtal Crustacea 100 100 100 200000 8 0.0382

Nircber of fish examined 29 Ntnber of prey items 8 Percent with food 13 Percent with sediment 100 Percent with digene parasite infestation 0 Percent with nematode parasite infestation 0 73

Feeding associations:

Cluster analysis was employed in an attempt to organize species into groups based on overall diet similarity. Six groups were established (fig* 28).

Group I consisted of B. pectoralis. I), kanazawai. P.. silus and P.. catena. Members of group I may be characterized as small to moderate­ sized fishes that fed predominately on small-sized calanoid copepods in addition to other Bmall benthopelagic crustaceans (tables, 4, 5, 12,

14). These fishes selected smaller-sized prey over a narrower than did members of other species groups examined (fig. 9). These fishes also probably fed more frequently off the bottom. Bathyonus pectoralis and

D. kanazawai fed heavily on calanoid copepods, gammaridean amphipods and mysids although both species fed heavily on other prey species

(polychaetes for I), kanazawai; isopods for B.. pectoralis. Dicrolene kanazawai"s diet was more diverse than B.. pectoralis in that the former fed on more kinds of prey items over a greater size range. Poroeadus silua and P. catena are two very closely related and morphologically similar species that fed heavily on small-sized calanoid copepods.

These two species exhibited a much higher dietary similarity than did B. pectoralis and I). kanazawai.

Group II was comprised of the single species B.. manatinus.

Although I), manatinus was distantly associated with members in group I, its unique predatory habits suggest that it be considered separatly.

Barathrodemua manatinus could be characterized as a small-sized fish unique in that it fed predominatley on small-sized benthic tanaids.

Calanoid copepods, tube-dwelling polychaetes, gammaridean amphipods and ostracods were also included in the diet but were much less important (fig. 9, table,66). In contrast to species included in group I, the presence of sediment in the digestive tract and a heavy reliance on more benthic prey (i.e. tanaids) suggests B,. manatinus employed a feeding stategy that specialized on food organisms nearer the bottom.

Group III could be characterized as moderate to large-bodied fishes that fed predominately on polychaetes (tables, 10, 14) Included in this group were two unrelated and morphologically distinct species, A. armatus and B., parri. Tube-dwelling polychaetes were the most important food source for both species and were responsible for their high degree of similarity in the dendogram. Although both species share polychaetes as a major food source, their diets differed significantly in other respects. Food items important in the diet of A. armatus. but nearly absent from the diet of B,. parri were amphipods, ostracods, mysids and copepods. In contrast, B_. parri was the only species examined in this study that fed heavily on holothurians. In fact the actual feeding stategies employed by these two fishes are probably quite different.

Barathrites parri is a firm bodied muscular fish that has evolved a distinctive feeding complex specialized to tear and shred bits of tissue away from prey organisms. In striking contrast, A. armatus captures and ingests prey whole or intact prey organisms. Acanthonus armatus has also evolved energy conserving morphological adaptations that enable it to maintain neutral buoyancy in the absence of a swimbladder. This species probably expendB little energy feeding by hovering at various depths off the bottom in search of prey organisms. These morphological specializations related to feeding are discussed in greater detail in subsequent sections. Group ZV included two closely related and morphologically similar species, B.. normalis and 13. taenia. These species may be characterized as fishes that fed predominately on large decapod crustaceans although both species also fed heavily on amphipods and mysids (fig. 9, tables,

7, 8). In contrast to small-bodied ophidiids these large fisheB probably possessed greater mobility effectively increasing the search range of their prey environment. This idea is supported by the fact that large individuals increasingly rely on larger more mobile prey organisms. Bassozetus normalis and B.. taenia exhibited a diverse diet and were unique in that they fed on much larger prey than other species examined in this study (fig. 9). Tanaids were more important in the diet of B.. taenia whereas fish were more important in the diet of B. normalis. Bassozetus normalis also fed on ostracods and gastropods which were absent from the diet of B. taenia.

Group V is comprised of a single species, P.. miles. Although P.. miles was distantly associated with B.. normalis and B_. taenia of group

IT, its rather restricted diet (due in part to small sample size) and its peculiar morphology suggest that is be considered separately.

Although P.. miles shared food items in common with B. normalis and B. taenia. it differed in that gammaridean amphipods rather than decapods were the most important prey organism (table 11). Furthermore its smaller body size, extremely anteriorly depressed head and very attenuate body suggest that it employed a different feeding strategy.

Unfortunately too few specimens of P. miles contained enough food items to sufficiently characterize its predatory habits. However, based on its general feeding morphology, its unlikely that P. miles was as * 76 heavily reliant on large mobile prey as were large individuals of B.. normalis and B.* taenia*

Group VI may be characterized as fishes that fed predominately on isopods, polychaetes and copepods (tables 3,9, 15). Group VI consisted of three unrelated and morphologically distinct species P. macdonaldi.

X. mversi and D.* introniara. Xvelacvba mversi and D. introniara fed predominately on benthic isopods, tube-dwelling polychaetes and calanoid copepods respectively* D. introniara also fed occasionally on lesser important gastropods and pelecypods, food items absent from the diets of

X. mverBi and P. macdonaldi* whereas tanaids were occasionally taken by

X. mverBi. macdonaldi. an isopod feeder, was tentatively placed within this group based on stomach contents of the few specimens examined. Although morphologically members of group VI are quite distinct, the predominance of benthic prey organisms and the occurrence of sediment in the digestive tract suggest that these species employed a more active benthic predatory habit than members of other groups. This is particularly evident when the diets of 1). introniara and D. kanazawai were compared and contrasted (tables 3, 4). Figure 28. Dendogram depicting diet similarity among ophidiid predators. DIET C L U S T E R t > CtC < m rvj fy. »n 'O +‘ + \ + 'O <“n +* + fN j’ f

SIMILARITY 78

Morphological associations:

Cluster analysis was employed as a verification procedure to determine the correspondence between species groups based on morphology and species groups based on diets. Since it is unlikely that any single morphological character will provide a reliable predictor of food items utilized by a predator it is necessary to consider a combination of characters in concert. Cluster analysis enabled the construction of species groups or "guilds" based on morphological characters associated with feeding. A guild is defined by Root (1967) as a group of species that exploit the same class of environmental resources in a similar way.

In this case species possessing similar morphological characters within the same habitat comprise a morphological guild.

Interpretation of cluster analysis results was a necessary step in the construction of morphological guilds. Cluster procedure itself does not determine what constitutes an operational group i.e. guild.

Criteria were established that require all guilds to consist of species members having similar resemblence coefficient values. The appropriateness of species treatment as a group or as individuals was determined subjectively using the best information available on the biology of the species concomitant with results of the character analysis.

Six groups were established by clustering morphological characters related to feeding in all species (fig. 29). Group I consisted of D.. intronigra. D. kanazawai. M. aeassizii. and B. pectoralis (fig. 29, table 17). With the possible exception of M. agassizii. group I could be characterized as small to moderate-sized fishes that possessed the following combination of key characters; large eyes, terminal mouth, 79 blunt snout, pectoral fins with lower rays free and longer than upper rays, high number of developed gill rakers on the anterior first arch, long and slender rakers, moderate filtering capacity of the branchial sieve, large gape, moderate upper jaw length, intestinal tract of moderate length with several pyloric caeca, greater degree of upper jaw protrusibility, moderate body depth and similar shape and configuration of bony elements of the feeding apparatus.

Monomitopus aeassizii. shared many features in common with the previous three species but differed in that it possessed a longer snout, an inferior mouth, increased body depth, pectoral fin with lower rays joined, increased number of long and slender gill rakers on anterior arch with increased filtering capacity of the branchial sieve, greater number of pyloric caeca, reduced upper jaw protrusibility and pectoral fins placed wholly anterior to the center of gravity rather than midway as in other species.

The morphological characters described above were those features primarily responsible for inclusion of B. pectoralis. M. aeassizii. D. introniera and 1). kanazawai in group I. These characters considered in combination represent the overall morphological pattern related to feeding in these four species. Values for meristic, morphometric and qualitative characters for each species in group I are presented in tables 18-21. Drawings of bony elements of the feeding apparatus for each species are presented in figures 30-59.

Group II is comprised of five species (fig. 29, table 22). Included in this group were B.. normalis. B.. taenia. H. souamoBus. A. edentatum and X. mversi. With the possible exception of X. mversi and edentatum group II could be characterized as relatively large mobile fishes that possessed the following combination of key characters; a

deep body, similar shape and configuration of bony elements of the

feeding apparatus, large gape, moderate upper jaw protrusibility,

slightly rounded swollen snout, small eye, numerous developed gill

rakers on anterior first arch with moderate filtering capacity of the branchial sieve, intestinal tract of moderate length with no pyloric

caeca, a short, slightly pointed pectoral fin with joined rays, and

terminal mouth.

Apagesoma edentatum and X. mversi were more distantly associated with B.. normalis. B. taenia, and H. souamosus. Xvelacvba mversi differed in that it possessed a longer snout, increased upper jaw

length, greater number of longer and more slender gill rakers on anterior arch, rounded pectoral fin, and dorsal fin originating at level of the center of gravity rather than midway through the plane. In addition the shape and configuration of bony elements of the feeding apparatus in X. mversi more closely resemble those of A. armatus. especially elements in the hyoid, suapensorium and anterior neurocranium regions.

Apagesoma edentatum differed from other species in group II in that it possessed the following important characters; a much smaller eye, fewer and shorter gill rakers on the anterior arch with decreased filtering capacity, a rounded pectoral fin and a slightly inferior mouth.

The morphological characters described above were those features primarily responsible for inclusion of B.. normalis. B. taenia. H. souamosus. A. edentatum and X. mversi in group II. These characters considered in combination represent the overall morphological pattern 81 related to feeding in these five species* Values for meristic, morphometric and qualitative characters for each species in group II are

presented in tables 23-27. Drawings of bony elements of the feeding apparatus for each species are presented in figures 30-59.

Group III consisted of B.* Parri and B.. iris (fig. 29, table 16).

These highly distinct fishes were characterized by the following combination of key characters; short head, small eyes, long abdomen, small gape with short upper jaw and much reduced upper jaw protrusibility, similar shape and configuration of bony elements of the feeding apparatus, few number of short gill rakers on anterior arch, much reduced filtering capacityof the branchial sieve, extremely long

intestinal tract, prominent vomerine tooth patch, pectoral fin short, rounded and placed wholly anterior to center of gravity, high pectoral

fin aspect ratio, subterminal mouth with enlarged outer row of conical

teeth in jaws and well developed jaw musculature.

The morphological characters described above were those features primarily responsible for inclusion of B.. parri and B.. iris in group

III. These characters considered in combination represent the overall morphological pattern related to feeding in these two species. Values for meristic, morphometric and qualitative characters for each species in group III are presented in tables 31-32. Drawings of bony elements of the feeding apparatus for B.. narri are presented in figures 30-59.

Group IV consisted of B. manatinus and A. galatheae (fig. 29, table 28). This group, best typified by B,. manatinus. could be characterized by the following combination of key characters; small gape, reduced upper jaw protrusibility, short upper jaw, long snout, small eyes, moderate number of gill rakers, rakers short, reduced 82 filtering capacity of the branchial sieve, inferior mouth, two basibrancbial tooth patches and a similar shape and configuration of bony elements of the feeding apparatus.

Abvssobrotula ealatheae shared many features in common with B,. manatinus but differed in that it possessed longer more pointed pectoral fins, decreased pectoral fin aspect ratio, shorter snout, fewer number of gill rakers on anterior arch, shorter intestinal tract and subtenninal mouth.

Although many of the above characters possessed by B.. manatinus and

A. Ealatheae were also shared by B.« parri and B., iris of group 111, the absense of several key characters Buch as specialized dentition, extremely long abdomen, and muscular jaws suggest that these two groups be treated separately*

The morphological characters described above were thoBe features primarily responsible for inclusion of B.. manatinus and A. ealatheae in group IV. These characters considered in combination represent the overall morphological pattern related to feeding in these two species.

ValueB for meristic, morphometric and qualitative characters for both species are presented in tables 29, 30, 31. Drawings of bony elements of the feeding apparatus for each species are presented in figures 30-

59.

Group V consisted of P. situs. P., catena. P. miles and .P.

Tnflodmmidf (fig. 29, table 34). This group best typified by P. silus and P. catena. could be described by the following combination of characters; slender and highly attenuate body, narrow and short pectoral fins with low aspect ratios, moderate to large gape, slightly reduced upper jaw protrusibility, moderate to large eye, greater number of gill 83 rakers on the anterior arch, rakers of moderate length with high filtering capacity of the branchial sieve, short intestinal tract with pyloric caeca present (P.. miles and P. catena only), similar configuration and shape of bony elements of the feeding apparatus and a

lateral line represented by three rows of circular organa.

Poroeadus miles and P. macdonaldi were more distantly associated with P.. silus and P. catena. Poroeadus miles. which attained a much

larger body size, differed principally in that it possessed a more anteriorly depressed head and longer snout, subterminal mouth and a prominent vomerine tooth patch. Penonus macdonaldi differed in that it possessed an extremely long snout and anteriorly depressed head, much reduced eye, fewer developed gill rakers on anterior arch, reduced filtering capacity of the branchial sieve, and a highly inferior mouth with two basibranchial tooth patches. The anteriorly depressed head and long snout of these two species were the result of a general elongation of bony elements comprising the upper and lower jaws and frontal bones of the neurocranium. Penonus macdonaldi achieved an even longer snout through the elongation of paired nasal bones just forward of the frontals and ethmoid region. In contrast the elements comprising the upper and lower jaw in P. silus and P.. catena were much shorter. Also the frontal bones in both these latter species ascend steeply just above the eye rather than gradually as they do in P. miles and P. macdonaldi.

The morphological characters described above were those features primarily responsible for inclusion of P. silus. P. catenar P. miles and

P.. macdonaldi in group T. These characters considered in combination represent the overall morphological pattern related to feeding in these four species. Values for meristic, morphometric and qualitative 84

characters for each species in group T are presented in tables 35-38.

Drawings of bony elements of the feeding apparatus for each species are presented in figures 30-59.

Group V was comprised of a single morphologically distinct species,

A. armatus (fig. 29, table 34). Acanthonua armatus could be

characterized based on the following combination of key characters; and extremely deep body and short trunk shape, short pectoral fin with a high aspect ratio, large gape, elongate upper jaw, numerous long developed gill rakers on the anterior arch, relatively high filtering

capacity of the branchial sieve, short stomach, prominent vomerine tooth

patch, pectoral, dorsal and anal fins all originating posterior to

center of gravity, absense of swimbladder and associated drumming muscles, massively swollen head, reduced body musculature, and a bifid

frontal spine. In addition to these characters the shape and

configuration of bony elements of the feeding apparatus closely

resembled those of X. mversi. In fact both thes species were

taxonomically quite distinct from all other species examined. Vithin

the subfamily neobythitinae, X. mversi and A. armatus were referred to

the tribe sirembini whereas the remainder of the species examined in

this study were placed within the neobythitini (Cohen and Nielsen,

1 9 7 8 ) .

The morphological characters described above were those features

primarily responsible for inclusion of armatus in group TI. These

characters considered in combination represent the overall morphological

pattern related to feeding in this species. Values for meristic, morphometric and qualitative characters for this species are presented

in table 39. Drawings of bony elements of the feeding apparatus for Jl.. armatus are presented in figures 30-59. Figure 29. Dendogram depicting morphological similarity among deep-sea ophidiids. MORPHOLOGY CLUSTER similarity

*1 .97 .95 .90 .5(5 .55 .79 .75 .72 .68 .55 L_ _L I I _L _1_

M. agassizii

D. intronigra D. kanazawai B. pectoralis

A. edentatum,

H. squamosus B. taenia B. normalis 3 X . myerst

B. parri B. iris 111 B. manatinus . B. manatinus ? — * A. galatheae, IV P. maci7oncWi___

P. r!ll,r P. mites P, catena — VI /I. armatus—

_ r r T T T T T T I n *} ,97 .95 .90 .55 .55 .79 .75 .72 .55 .55

similarity Table 16. Ophidiid Fishes Examined in Morphological Study

Size Range S p e c ie s ______(SL ran) X SD N

1 Dicrolene intrtmigra, 210-304 268 15 J 30

2 TriCrolene fomnrattai 208-254 227 11.3 30

3 Hathymnua pectoralis 125-166 148 13.6 30

4 RflBBftzetua normalis 305-472 408 8.5 30

5 RaBsnzetus taenia 137-386 259 57 18

6 Xvelacyba mversi 88-422 225 94 17

7 Byrflrtirilea parri 175-250 220 31 30

8 Boroeadua silus 118-132 125 5.4 30

9 Borogadua cafera 130-250 187 38 29

10 Boraeadua milea 181-439 318 85 12

11 Bmnptis macdonaldi 105-237 183 33 29

12 Barafhrodfrttia iromapinua 122-162 148 4.2 30

13 jt-anfWimn nrmafus 165-320 285 10.8 30

14 Bprathritea iris 530-530 530 - 1

15 Abvasobrotula ealatheae 128-128 128 - 1

16 MmondtOWa agassizii 153-190 178 6.8 30

17 HnlconWCteroTMB arpumnmm 150-291 214 14.6 3

18 Arfpf>r*m sdentatwn 720-726 723 4.2 2

19 BnrthrodenitB ranaHniiB 128-153 142 5.7 30 Table 17. Mean Values of Iforphological Characters Belated to Feeding for Each Species in Group I* MA,4foncciitopus agaasizii. DI=3)icrolene intronigra. DRt$jerglene kanazawai. and KP^Bathvonus pectoralis.

Charflcter/Species______MA PI IK BP

Body Depth 20.0 15.0 14.8 13.7 Trunk Shape Index 38.1 35.0 33.0 38.0 Pectoral Fin length 10.1 26.0 20.0 17.9 Pectoral Fin Aspect Ratio 33.0 26.0 25.1 16.0 Lateral Gape 9.2 11.0 10.0 10.2 Medial Gape 11.6 13.0 12.0 12.5 Upper Jaw length 10.2 10.0 9.8 10.6 Upper Jaw Protrusibility 82.0 66.0 66.0 73.0 Snout Length 5.4 3.9 4.3 4.8 Bye Dimeter 4.9 4.6 5.2 2.9 No. Developed Bakers 24.0 14.6 11.0 12.3 Inter Baker Gap Distance 0.42 0.59 0.68 0.75 Greatest Baker Length 4.5 3.1 3.4 3.7 Branchial Filter Area 159.0 186.0 175.0 152.0 Stomach Length 10.6 10.4 7.5 9JS Intestinal Tract length 57.7 86.8 62.3 59.6 Total Gut Length 68.2 95.4 70.2 70.4 Swinbladder length 16.2 17.1 12.1 12.9 Rete length 9.6 6.8 8.2 11.2 No. Pyloric Caeca 9.2 4.0 4.0 4.0 Pectoral Fin Shape 2 3 3 3 Pectoral Fin Condition 1 2 2 2 Pectoral Fin Relative to OG 1 2 2 2 Dorsal Fin Relative to OG 2 2 2 2 Anal Fin Relative to OG 4 4 4 4 Mouth Orientation 3 2 2 2 Jaw Dentition 2 2 2 2 Riaryngeal Dentition 2 2 2 2 No. Basibranchial patches 1 1 1 2 Vomer Shape 4 3 3 3 Swdmbladder Presence/Absence 2 2 2 2 Gonpletion of lateral line 1 1 1 1 Position of Lateral line 2 2 2 2 Eye Position 1 1 1 1 No. of lateral Lines 1 1 1 1 Drunuing Muscles Presence/Absence 2 2 2 2 Feeding Apparatus Pattern 3 3 3 3 Table 18. Morpheme trie, Meristic and Qualitative Characters Belated to Feeding in Dicrolene kanazawai.

CHARACTER N RANGE X S.D. MwDfrxnetrics %SL

Standard Length 30 208-254 227 1 1 3 Body depth 30 12.2-16.1 14.8 1.8 Index of Trunk Shape 30 29.2-35.1 33.0 1.2 Pectoral Fin Length 30 18.1-22.3 20.0 2.1 Aspect Ratio of Pectoral Fin 30 18.3-35.5 25.1 1.0 lateral Mouth Gape 30 9.3-10.8 10.0 0.71 Medial Mouth Gape 30 10.2-13.5 12.0 1 3 Upper Jaw length 30 8.8-11.0 9.8 0.91 Upper Jaw Protrusibility 30 53-79 66 4.5 Shout Length 30 3.2-5.1 4 3 0.8 Inter Baker Gap Distance 30 0.58-0.72 0.68 0.01 Greatest Baker length 30 3.0-4.8 3.4 0.55 Branchial Seive Filter Area 30 148-199 175 15 Stomach length 30 6.2-7.9 7.5 1.1 Intestinal Tract Length 30 56.0-65.0 6 2 3 3.0 Total Gut Length 30 63.0-75.1 70.2 4.2 Swinbladder length 30 10.1-13.2 12.1 0.8 Swinbladder Width 30 7 3 - 8 . 2 7.8 0,7 Bete Length 30 8 . 0 - 9 3 8 3 0.8 Bye Dimeter 30 4.9-6.5 5.2 0.93

Meristics

Number of Developed Rakers 30 9-12 11 0.75 Number of Pyloric Caeca 30 4-4 4 limber of lateral Lines 30 1-1 1 Huber of Basibranchial Patches 30 1-1 1 Swinbladder Presence/Absence 30 2 Drumming Hiscles Presence/Absence 30 2

Qualitative Characters

Conpleteneaa of lateral Line 30 1 Position of Lateral Line 30 2 Pectoral Fin Shape 30 3 Pectoral Fin Condition 30 2 Position of Pectoral Fin Relative to Center of Gravity 30 2 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 2 Position of Byes 30 1 Upper and Lower Jaw Dentition 30 2 Upper and Lower Riaryngeal Dentition 30 2 Shape of Vomer 30 3 Feeding Apparatus Pattern 30 3 Table 19. Morphometric, Meristic and Qualitative Characters Related to Feeding in Dicrolene intronigra.

CHARACTER N RANGE X S.D. Morohometries ZSL

Standard length 30 210-304 268 15.3 Body depth 30 14.0-17.3 15.0 1.2 Index of Trunk Shape 30 32.3-36.8 35.0 1.0 Pectoral Fin length 30 19.8-30.0 26.0 3.2 Aspect Ratio of Pectoral Fin 30 24.0-27.1 26 1 3 Lateral Mouth Gape 30 10.3-12.0 11 1.1 Medial Mouth Gspe 30 11.1-14.2 13 1.5 Upper Jaw length 30 9.4-11.2 10.0 0.4 Upper Jaw Protrusibility 30 61.2-69.3 66 3.9 Snout Length 30 3.5-4.2 3.9 0 3 Inter Raker Gap Distance 30 0.55-0.62 0.59 0.6 Greatest Raker Length 30 2.8-3.4 3.1 0.2 Branchial Seive Filter Area 30 100.1-175 186 10.8 Stomach Length 30 9.3-13.0 10.4 1.2 Intestinal Tract length 30 70.4-98.1 86.8 8.0 Tbtal GUt Length 30 88.2-106.1 95.4 8 3 Swinbladder Length 30 15.7-19,1 17.1 3.0 Swinbladder Width 30 9.5-14.1 12.5 2.5 Rete Length 30 5.4-9.1 6.8 1.0 Eye Diameter 30 4 3 - 5 . 2 4.6 0.5

Meristics

Huber of Developed Rakers 30 13-15 14.6 0.6 H u b e r of Pyloric Caeca 30 4-4 4 Huber of Lateral Lines 30 1-1 1 Huber of Basibranchial Patches 30 1 Swinbladder Presence/Absence 30 2 Dnmning Hiscles Presence/Absence 30 2

Qualitative Characters

Completeness of lateral Line 30 1 lbs it ion of lateral line 30 2 Pectoral Fin Shape 30 3 Pectoral Fin Condition 30 2 Position of Pectoral Fin Relative to Center of Gravity 30 2 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 2 Position of Eyes 30 1 Upper and Lower Jaw Dentition 30 2 Upper and Lower Pharyngeal Dentition 30 2 Shape of Vomer 30 3 Feeding Apparatus Pattern 30 3 Table 20. Morpheme trie, Meristic and Qualitative Characters Related to Feeding in Bathvoms pectoralis.

CHARACTER______;______N RANGE X S.D. Morpheme tries %SL

Standard length 30 125-166 148 13.6 Body depth 30 10.1-16.2 13.7 1.5 Index of Trunk Shape 30 32.5-41,2 38.0 2.1 Pectoral Fin Iangth 30 14.1-21.2 17.9 2.1 Aspect Ratio of Pectoral Fin 30 13.5-20.1 16 1 3 lateral Mouth Gape 30 7.2-13.5 10.2 1.2 Medial Mouth Gape 30 11.0-15.2 12.5 1.8 Upper Jaw length 30 10.1-153 10.6 1.0 Upper Jaw Protrusibility 30 67-62 73 5.1 Snout length 30 4.2-5.0 4.8 0.8 Inter Raker Gap Distance 30 0.64-1.7 0.75 0.2 Greatest Raker length 30 2 . 2 - 1 0 3 3.7 1.0 Branchial Seive Filter Area 30 138-182 152 8 3 Stomach length 30 7 . 5 - 1 2 3 9.6 1.9 Intestinal Tract length 30 52.1-59.6 59.6 7.0 Tbtal GUt length 30 9.8-16.2 12.9 1.6 Swinbladder length 30 7 . 8 - 1 2 3 9 3 1.0 Swinbladder Width 30 4.0-15.1 11.2 2.7 Rete length 30 2.0- 2.5 2 3 0.60 Eye Diameter

Msristics

Haber of Developed Rakers 30 12-13 12 0.4 Huber of Pyloric Caeca 30 4-4 4 H u b e r of lateral Lines 30 1-1 1 Huber of Basibranchial Patches 30 2-2 2 Swinbladder Presence/Absence 30 2 Dnnming Hiscles Presence/Absence 30 2

Qualitative Characters

Goopleteness of lateral line 30 1 Position of Lateral Line 30 2 Pectoral Fin Shape 30 3 Pectoral Fin Condition 30 2 Position of Pectoral Fin Relative to Center of Gravity 30 2 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 2 Position of Eyes 30 1 Upper and lower Jaw Dentition 30 2 Upper and Irwer Pharyngeal Dentition 30 2 Shape of Vomer 30 3 Feeding Apparatus Pattern 30 3 Table 21. Morphotnetric, Meristic sad Qualitative Characters Related to Feeding in Mapcmitonus agasssizii.

CHARACTER______N RANGE X SA Morphometries ZSL

Standard Length 30 153-190 178 6.8 Body depth 30 19.1-21.2 20.0 0.9 Index of Trunk Shape 30 36.1-39.8 38.1 1 3 Pectoral Fin Length 30 9.1-11.2 10.1 0.8 Aspect Ratio of Pectoral Fin 30 32.1-35.6 33 1.1 Lateral Mouth Gape 30 8.8-9.9 9.2 0.8 Medial Mouth Gape 30 9.0-12.1 11.6 1 3 Upper Jaw Length 30 9.7-11.4 10.2 0.71 Upper Jaw Protrusibility 30 77.1- 8 5 3 82 3 j6 Snout Length 30 5.0-5.8 5.4 0 3 6 Inter Raker Gap Distance 30 039-0.43 0.42 0 3 Greatest Baker length 30 43-4.9 4.5 0.8 Branchial Seive Filter Area 30 134-182 159 12.8 StcnBch Length 30 9.8-11.1 10.6 0.5 Intestinal Tract Length 30 52.0-63.1 57.7 4.1 Total Gut Length 30 61.0-70.4 6 8 3 5.4 Swinbladder length 30 15.0 - 1 7 3 1 6 3 0.91 Swinbladder Width 30 8.1-113 93 1.2 Rete Length 30 9.0-10.7 9.6 0.8 Eye Diameter 30 4 3 - 5 3 4.9 0.4

Her is tics

Ntcfcer of Developed Rakers 30 23-25 24 0.9 Nunber of Pyloric Caeca 30 9-10 9 3 0.4 Nunber of Lateral Lines 30 1-1 1 Mnber of Basibranchial Patches 30 1-1 1 Swinbladder Presence/Absence 30 2 Drtnming Hiscles Presence/Absence 30 2 ftmlitntive Qwrncfprs

Completeness of Lateral Line 30 1 Position of lateral Line 30 2 Pectoral Fin Shape 30 2 Pectoral Fin Condition 30 1 Position of Pectoral Fin Relative to Center of Gravity 30 1 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 3 Position of Eyes 30 1 Upper and Lower Jaw Dentition 30 2 Upper and Lower Pharyngeal Dentil] 30 2 Shape of Vomer 30 4 Feeding Apparatus Pattern 30 3 Table 22. Mean Values of Morphological Characters Related to Feeding for Each Species in Group II, AE*Apagesana edentatun. BDS^golccmvcterorus flouamosus. EAZW=Baflsozetus normalis. BA23>=Bassozetus fj»»nia and Xtf*=Xyalacvba mversi.

Character/Species______AE BOS BA2N BAZX XM

Body Depth 23.1 17.6 13.2 15.4 25.0 Trunk Shape Index 37.0 41.0 32.0 33.1 38.0 Pectoral Fin length 12.5 20.0 10.1 8.8 11.2 Pectoral Fin Aspect Ratio 45.0 18.0 25.0 26.0 51.0 Lateral Gape 10.7 12.0 10.1 10.1 10.0 Medial Gape 11.9 15.0 12.9 13.2 1 5 3 Upper Jaw Length 11.6 10.6 9.5 9 3 1 3 3 Upper Jaw Protrusibility 75.0 81.0 75.0 76.0 78.5 Snout Length 5 3 5.1 3.9 4 3 7.8 Eye Diameter 0.84 2.7 2.0 1.8 2.6 No. Developed Rakers 8.5 9.5 13.0 14.0 17.0 Tnt-gr Baker Gap Distance 0.67 0.68 0.58 0. 56 0.58 Greatest Raker Length 1.5 2.7 2.7 2.5 3.6 Branchial Filter Area 95.0 1 0 5 3 241.0 250.0 285.0 Stomach Length 16.5 7.1 1 0 3 10.1 8.4 Intestinal Tract length 57.0 68.0 47.0 62.0 52.9 Tbtal Gdt Length 74.0 71.0 58.0 67.1 61.4 Swinbladder Length 1 0 3 1 4 3 10.7 9.6 11.6 Rete length 6.1 13.8 1 2 3 11.0 15.8 No. Pyloric Caeca 0 0 0 0 0 Pectoral Fin Shape 1 2 2 2 1 Pectoral Fin Condition 1 2 1 1 1 Pectoral Fin Relative to OG 2 2 2 2 2 Dorsal Fin Relative to OG 2 2 2 2 3 Anal Fin Relative to OG 4 4 4 4 4 Mouth Orientation 2 2 2 2 2 Jaw Dentition 2 2 2 2 2 Pharyngeal Dentition 2 2 2 2 2 No. Basibranchial patches 1 2 1 1 2 \fcmer Shape 1 3 3 4 4 Swinbladder Presence/Absence 2 2 2 2 2 Gonpletion of Lateral line 1 1 1 1 1 Position of Lateral Line 2 2 2 2 2 Eye Position 1 1 1 1 1 No. of Lateral Lines 1 1 1 1 1 Drumnng Hiscles Presence/Absence 2 2 2 2 2 Feeding Apparatus Pattern 2 2 2 2 5 Table 23. Morphemetrie, Meristic and Qualitative Characters Belated to Feeding in Anagesana edentatum

CHARACTER______N RANGE_____ X S.D. Morphometries ZSL

Standard length 2 720-726 723 4 3 Body depth 2 21.3-25.0 23.1 4.8 Index of Trunk Shape 2 36.2-37.1 37 0.8 Pectoral Fin Length 2 12.5-12.6 12,5 0.1 Aspect Ratio of Pectoral Fin 2 43.2-% .5 45 2.1 Lateral Mouth Gape 2 9.6-11.8 10.7 1.5 Medial Mouth Gape 2 11.0-12.3 11.85 0.9 Upper Jaw Length 2 1 1 .1-12.8 11.6 1.4 Upper Jaw Protrusibility 2 73.1-76.4 75 2 3 Snout Length 2 43-6.2 5.3 1.34 Inter Raker Gap Distance 2 0.59-0.75 0.67 0.11 Greatest Raker Length 2 1.3-1.7 1.5 0 3 Branchial Seive Filter Area 2 93-96 95 2.1 Stomach Length 1 - 16.5 - Intestinal Tract Length 1 - 57.0 - Tbtal Gut Length 1 - 74.0 - Swiirb ladder Length 1 - 1 0 3 - Swini)ladder Width 1 - 6 3 - Rete Length 1 - 6.1 - Eye Diameter 2 0.48-1.2 0.84 0.5

Meristics

Nuober of Developed Rakers 2 7.0-10.0 8.5 2.1 Ntmber of Pyloric Caeca 2 0 Nmber of Lateral Lines 2 1 ftafcer of Basibranchial Patches 2 1-1 1 Swiirb ladder Presence/Absence 2 2 Dnnming Miscles Presence/Absence 2 2

Oualitative Characters

Cbnpleteness of Lateral line 2 1 Position of Lateral Line 2 2 Pectoral Fin Shape 2 1 Pectoral Fin Condition 2 1 Position of Pectoral Fin Relative to Center of Gravity 2 2 Position of Dorsal fin Relative to Center of Gravity 2 1 Position of Anal Fin Relative to Center of Gravity 2 4 Mouth Orientation 2 2 Position of Eyes 2 1 Upper and lower Jaw Dentition 2 2 Upper and lover Pharyngeal Dentition 2 2 Shape of Vomer 2 1 Feeding Apparatus Pattern 2 Table 24. Mbipbometric, Meristic and Qualitative Characters Related to Feeding in Xyelacvba nwersi.

CHARACTER______H RANGE X SJ). Marphcmetrica %SL

Standard Length 17 88-422 225 94 Body depth 17 21.5-28.2 25 1 A Index of Trunk Shape 17 29.9-44.8 38 2 3 Pectoral Fin Length 17 9.0-13.8 11.2 1 3 Aspect Ratio of Pectoral Fin 17 41.4-623 51.0 3.9 Lateral Mouth Gape 17 8 3 - 1 2 3 10.0 1.0 Medial Mouth Gape 17 143-18.9 1 5 3 1.4 Upper Jaw Length 17 12.0-16.1 13.2 1.2 Upper Jaw Protrusibility 17 683-88.9 78.5 5.6 Snout length 17 63-9.2 7.8 1.0 Inter Raker Gap Distance 17 0.42-0.73 0.58 0.4 Greatest Raker Length 17 2.4-43 3.62 0.9 Branchial Seive Filter Area 17 268-313 285 21 Stomach Length 17 6.8-11.3 8.4 1.0 Intestinal Tract Length 17 48.2-63.4 52.9 3.5 Ibtal Gut Length 17 55.7-75.4 61.4 4.0 Swinbladder length 17 9.7-13.4 11.6 1.0 Swimbladder Width 17 6 3 - 1 0 3 7.4 1.0 Rete length 17 13.9-17.8 15.8 1 3 Eye Diameter 17 1.9-3.5 2.6 0.6

Meristics

Nuiber of Developed Rakers 17 15-19 17 1.8 lumber of fyloric Caeca 17 0 Nmber of Lateral Lines 17 1 Number of Basihranchial Patches 17 2 Swimbladder Presence/Absence 17 2 Dnmning Miscles Presence/Absence 17 2

Qualitative Characters

Coopleteness of Lateral Line 17 1 Position of lateral Line 17 2 Pectoral Fin Shape 17 1 Pectoral Fin Condition 17 1 Position of Pectoral Fin Relative to Center of Gravity 17 2 Position of Dorsal fin Relative to Center of Gravity 17 3 Position of Anal Fin Relative to Center of Gravity 17 4 Mouth Orientation 17 2 Position of Eyes 17 1 Upper and lower Jaw Dentition 17 2 Upper and lower Pharyngeal Dentition 17 2 Shape of Vcmer 17 4 Feeding Apparatus Pattern 17 5 Table 25. Morpheme trie, Meristic and Qualitative Characters Related to Feeding in Bassozetus normalis.

CHARACUR N RANGE X S.D. MarDhonetrics %SL

Standard Length 30 305-472 408 8.5 Body depth 30 12.9-19.8 15.4 1.8 Index of Trunk Shape 30 26-35.6 33.1 2,0 Pectoral Fin Length 30 6.2-10.3 8.8 1.5 Aspect Ratio of Pectoral Fin 30 22.2-30.1 26.0 1.1 lateral Mouth Gape 30 8 .0 -12.1 10.1 1.0 Medial Mouth Gape 30 9.4-15.3 13.2 1.5 Upper Jaw Length 30 8.0-12.4 9.3 1.0 Upper Jaw Protrusibility 30 70-84 76 3.0 Shout Length 30 3.4-5.6 4.2 0.5 Later Raker Gap Distance 30 0.37-0.62 0.56 0.13 Greatest Raker Length 30 2.0-3.1 2.5 0.3 Branchial Seive Filter Area 30 226-288 250 15 Stomach Length 30 8 .8-11.1 10.1 1.0 Intestinal Tract Length 30 48-81 62.0 2.6 Total Gut length 30 58-88 67 1.9 Swiribladder Length 30 7.4-12.9 9.6 2.0 Swimbladder Width 30 4.9-8.2 6.1 1.0 Rete length 30 8.0-14 11.0 2.0 Eye Dimeter 30 1 .2-2.6 1.8 0.4

Meristics

Number of Developed Rakers 30 12-16 13 1.7 Number of Pyloric Caeca 30 0 Huber of lateral Lines 30 1-1 1 Huber of Basibranchial Patches 30 1-1 1 Swinb ladder Presence/Absence 30 2 Drunning Hiscles Presence/Absence 30 2

Qualitative Characters

Completeness of Lateral line 30 1 Position of Lateral Line 30 2 Pectoral Fin Shape 30 2 Pectoral Fin Condition 30 1 Position of Pectoral Fin Relative to Center of Gravity 30 2 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 2 Position of Eyes 30 1 Upper and Lower Jaw Dentition 30 2 Upper and Lower Pharyngeal Dentition 30 2 Shape of Vomer 30 4 Feeding Apparatus Pattern 30 2 Table 26. Morpheme trie, Meristic and Qualitative Characters Related to Feeding in Bassozetus rnmia.

CHARACTER______N RAWS X S.D. Morphometries %SL

Standard length 18 137-386 259 57 Body depth 18 12.0-14.1 13.2 1.1 Index of Trunk Shape 18 26.1-37.4 32 3.1 Pectoral Fin length 18 7.5-13.1 10.1 2.0 Aspect Ratio of Pectoral Fin 18 21.1-29.6 25 1.9 lateral Mouth Gape 18 8.8-11.3 10 1.2 Medial Mouth Gape 18 9.6-14.4 12.9 0.9 Upper Jaw length 18 8.0-10.5 9.5 0.73 Upper Jaw Protrusibility 18 6 6 .1-86.2 75 3.4 Snout length 18 3.0-4.8 3.9 0.4 Inter Raker Gap Distance 18 0.50-0.62 0.58 0.06 Greatest Raker Length 18 2.1-3.3 2.7 0.55 Branchial Seive Filter Area 18 219-282 241 20 Stomach Length 18 9.8-10.6 10.2 1.3 Intestinal Tract Length 18 43-52 47 1.7 Ibtal Gut Length 18 53.1-63.2 58 2.1 Swinbladder length 18 8.9-12.3 10.7 1.0 Swinbladder Width 18 4.6-B.5 6.4 1.3 Rete length 18 10.0-16.2 12.3 1.9 Eye Diameter 18 1 .5-2.4 2.0 0.2

Meristics

Huber of Developed Rakers 18 12-13 13 1.0 Huber of Pyloric Caeca 18 0 Huber of lateral Lines 18 1-1 1 H u b e r of Basibranchial Patches 18 1-1 1 Swinbladder Presence/Absence 18 2 Drumming Muscles Presence/Absence 18 2

Qualitative Characters

Completeness of Lateral L i w a 18 1 Position of Lateral Line 18 2 Pectoral Fin Shape 18 2 Pectoral Fin Condition 18 1 Position of Pectoral Fin Relative to Center of Gravity 18 2 Position of Dorsal fin Relative to Center of Gravity 18 2 Position of Anal Fin Relative to Center of Gravity 18 4 Mouth Orientation 18 2 Position of Eyes 18 1 Upper and lower Jaw Dentition 18 2 Upper and lower Pharyngeal Dentition 18 2 Shape of Vomer 18 3 Feeding Apparatus Pattern 18 2 Table 27. Morpbanetric, Her is tic and Qualitative Characters Related to Feeding in Bolcanvcteronus sauamosus.

CHARACTER NRANGE X S.D. Mbrnhauetrics ZSL

Standard length 3 150-291 214 14.6 Body depth 3 17-18 17.6 1.0 Index of Trunk Shape 3 37 .(Hi8 .3 41.0 3.5 Pectoral Fin Length 3 19.0-23.0 20.0 2.7 Aspect Ratio of Pectoral Fin 3 16.3-20.5 18 1.2 lateral Mouth Gape 3 11.3-12.9 12 1.0 Medial Mouth Gape 3 14.0-16.2 15 1.4 Upper Jaw length 3 10.1-10.9 10.6 0.2 Upper Jaw Protrusibility 3 78.0-86.4 81 3.2 Snout length 3 4.9-S.3 5.1 0.2 Inter Raker Gap Distance 3 0.64-0.72 0.68 0.1 Greatest Raker length 3 2.6-2.9 2.7 0.2 Branchial Seive Filter Area 3 98.1-113.2 105.3 7.5 Stonach Length 3 5.2-9.5 7.1 1.2 Intestinal Tract Length 3 62-75 68.0 2.4 Tbtal Gut length 3 54.3-76 71.1 3.5 Swinbladder Length 3 14.1-14.9 14.3 0.1 Swinbladder Width 3 7.8-9.0 8.2 0.4 Rete Length 3 13.2-14.1 13.8 0.7 Eye Dimeter 3 2 .6-2.8 2.7 0.1

Meristics

Nuiber of Developed Rakers 3 9-10 9.5 0.7 Ruber of Pyloric Caeca 3 0 Ruber of Lateral lines 3 1-1 1 Ruber of Basibranchial Patches 3 2-2 2 Swinbladder Presence/Absence 3 2 Dnnming Miscles Presence/Absence 3 2

Qualitative Characters

Completeness of lateral line 3 1 Position of lateral Line 3 2 Pectoral Fin Shape 3 2 Pectoral Fin Condition 3 2 Position of Pectoral Fin Relative to Center of Gravity 3 2 Position of Dorsal fin Relative to Center of Gravity 3 2 Position of Anal Fin Relative to Center of Gravity 3 4 Mouth Orientation 3 2 Position of Eyes 3 1 Upper and lower Jaw Dentition 3 2 Upper and lower Pharyngeal Dentition 3 2 Shape of Vomer 3 3 Feeding Apparatus Pattern 3 2 Table 28. Mean Values of Morphological Characters Related to Feeding for Each Species in Group i n and IV. M WBarathrites pgrri. BAI^Barathries iris. BAM=Barathrodaras pynatinus. and A & Abys_s6brotula ealatheae.

Character/Species BAI BAP BAM AG ______(group III) (group I?)

Body Depth 16.9 23.0 12.0 15.0 Trunk Shape Index 46.1 41.0 31.1 34.0 Pectoral Fin length 7.7 10.0 11.4 25.0 Pectoral Fin Aspect Ratio 30.0 48.0 23.0 12.0 lateral Gape 5.7 7.1 5.4 7.6 Medial Gape 6.8 9.2 6.6 11.5 Upper Jaw Length 6.1 7.2 6.5 9.6 Upper Jaw Pcotrusibility 92.0 90.2 88.0 79.0 Snout length 3.6 4.8 11.1 9.6 Eye Dimeter 2.0 0.66 2.7 1.7 Mo. Developed Rakers 6.0 7.0 14.0 9.0 Inter Raker Gap Distance 0.59 0.45 0.52 0.72 Greatest Raker Length 0.71 0.75 2.2 3.1 Branchial Filter Area 5.8 45.0 42.1 88.5 Stanach Length 8.9 13.2 5.9 5.8 Intestinal Tract length 71.0 132.0 42.4 21.0 Total Gut Length 80.0 138.0 48.1 26.0 Swinbladder length 14.2 9.4 7.8 5.2 Rete Length 17.8 11.3 11.3 14.1 No. lyloric Caeca 0 0 0 0 Pectoral Fin Shape 2 3 1 1 Pectoral Fin Condition 1 1 1 1 Pectoral Fin Relative to OG 1 2 1 1 Dorsal Fin Relative to OG 2 2 2 2 Anal Fin Relative to OG 4 4 4 4 Mouth Orientation 3 3 4 3 Jaw Dentition 3 3 2 2 Pharyngeal Dentition 2 2 2 2 No. Basibranchial patches 1 1 2 2 Vainer Shape 4 4 3 3 Swinbladder Presence/Absence 2 2 2 2 Gonpletion of Lateral T.i u p 1 1 1 1 Position of Lateral Line 2 2 2 2 Eye Position 1 1 1 1 No. of Lateral Lines 1 1 1 1 Drumdng Miscles Presence/Absence 2 2 2 2 Feeding Apparatus Pattern 4 4 4 4 Table 29. Iforphocetric, Meristic and Qualitative Characters Related to Feeding in Barathrodama manatinus (male)

CHARACTER______M RANGE X S.D. jfarphcmetrics ZSL

Standard Length 30 132-152 148 4.2 Body depth 30 10.5-13.8 12.0 1 3 Index of Trunk Shape 30 26.1-35.0 31.1 2.0 Pectoral Fin length 30 8 .8-12.2 11.4 0.9 Aspect Ratio of Pectoral Fin 30 18.5-27.6 23.0 2.0 Lateral Mouth Gape 30 4.5-9.2 5.4 1.0 Medial Mouth Gape 30 5.1-9.6 6.6 1.2 Upper Jaw Length 30 5.9-10.1 6.5 1.0 Upper Jaw Protrusibility 30 75.1-99.4 88 5 3 Snout length 30 10.8-12.3 11.1 0.9 Titer Raker Gap Distance 30 0.37-0.82 0.52 0.09 Greatest Raker Length 30 1.6-33 2.2 0.38 Branchial Seive Filter Area 30 263-69.1 42.1 9.3 Stomach Length 30 4.4-73 5.9 0.73 Intestinal Tract Length 30 28.7-52.2 42.4 6.6 Total Gut Length 30 343-56.1 48.1 6.2 Swinbladder length 30 5.7-103 7.8 1 Swinbladder Width 30 3.0-8.4 4 3 1.2 Rete Length 30 6.5-14.1 1 1 3 2 Eye Diameter 30 23-3.1 2.7 0.12

Meristics

Ruber of Developed Rakers 30 13-15 14 1 Ruber of Pyloric Caeca 30 0 Ruber of Lateral Lines 30 1-1 1 Ruber of Basibranchial Patches 30 2-2 2 Swinbladder Presence/Absence 30 2 DnMining Miscles Presence/Absence 30 2

Qualitative Characters

Gcnpleteness of Lateral Line 30 1 Position of Lateral Line 30 2 Pectoral Fin Shape 30 2 Pectoral Fin Condition 30 1 Position of Pectoral Fin Relative to Center of Gravity 30 1 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 4 Position of Eyes 30 1 Upper and lower Jaw Dentition 30 2 Upper and lower Pharyngeal Dentition 30 2 Shape of Vomer 30 3 Feeding Apparatus Pattern 30 4 Table 30. Mnphometric, Meriatic and Qualitative Characters Related to Feeding in Barathrodams manatinus (female).

CHARACTER N RANGE X S.D. Mb rohdce tries %SL

Standard length 30 128-153 142 5.7 Body depth 30 13.7-18.4 15.4 1 3 Index of Trunk Shape 30 31.3-40.1 36.1 2.5 Pectoral Fin Length 30 11.0-14.1 12.5 0.65 Aspect Ratio of Pectoral Fin 30 18.7-243 21.1 1.8 Lateral Mouth Gape 30 4.4-93 6.1 2.0 Medial Mouth Gape 30 53-10.9 7.9 1.7 Upper Jaw length 30 5.2-8.9 7.0 0.4 Upper Jaw Protrusibility 30 78.1-93.3 86 4 3 Shout Length 30 10.8-143 12.2 0.9 Inter Raker Gap Distance 30 0.42-0.75 0.54 0.08 Greatest Raker length 30 1.5-3.6 2.5 0.5 Branchial Seive Filter Area 30 293-58.4 45.5 6.9 Stomach length 30 4.5-7.9 6.1 0.9 Intestinal Tract Length 30 34.5-52.2 40.1 5.1 Tbtal Gut length 30 38.2-55.4 4 6 3 3.8 Swinbladder Length 30 5.9-9.1 7.0 0.9 Swinbladder Width 30 3.0-7.5 4.5 1.0 Rete Length 30 11.1-17.2 13.1 2 3 Eye Diameter 30 1.5-2.5 1.9 0.22

Meristics

Ruber of Developed Rakers 30 13-15 14 1 Ruber of fyloric Caeca 30 0 Ruber of lateral Li ms 30 1-1 1 R u b e r of Basibranchial Patches 30 2-2 2 Swinbladder Presence/Absence 30 2 Dimming Muscles Presence/Absence 30 2

(Xialitative Characters

Goqpleteness of Lateral Line 30 1 Position of lateral Line 30 2 Pectoral Fin Shape 30 2 Pectoral Fin Condition 30 1 Position of Pectoral Fin Relative to Center of Gravity 30 1 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 4 Position of Eyes 30 1 Upper and Lower Jaw Dentition 30 2 Upper and lower Pharyngeal Dentition 30 2 Shape of Vomer 30 3 Feeding Apparatus Pattern 30 4 Table 31. Morphemetrie, Meristic and Qualitative Characters Related to Feeding in Barathrites narri.

CHARACTER______N RANGE X S.D. Morphometries %SL

Standard Length 30 175-250 220 31 Body depth 30 13.8-19.8 16.9 1.9 Index of Trunk Shape 30 41.3-52.1 46.1 2.1 Pectoral Fin Length 30 7.0-11.3 7.7 1.0 Aspect Ratio of Pectoral Fin 30 27.1-33.4 30 2.2 Lateral Mouth Gape 30 5.1-6.3 5,7 1.1 Medial Mouth Gape 30 6 .0-7.4 6,8 0.9 Upper Jaw Length 30 5.6-6.6 6.1 0.3 Upper Jaw Protruaibility 30 84.1-102.3 92.0 3.9 Snout length 30 3.1-3.8 3.6 0.2 Inter Raker Gap Distance 30 0.38-0.79 0.59 0.1 Greatest Raker Length 30 0.60-0.83 0.71 0.6 Branchial Seive Filter Area 30 14-24 18 3.1 Stomach length 30 6 .2-9.2 8.9 0.3 Intestinal Tract length 30 68-74 71 2.9 Total Gut length 30 71-83 80 3.1 Swinbladder Length 30 12.2-16.8 14.2 1.1 Swinbladder Width 30 6 .2-9.9 8.1 1.0 Rete Length 30 15.8-19.2 17.8 1.0 Eye Diameter 30 1.6-2.5 2.0 0.3

Meristics

Ruber of Developed Bakers 30 4-8 6 1.7 Ruber of Pyloric Caeca 30 0 Ruber of Lateral Lines 30 1-1 1 - R u b e r of Basihranchinl Patches 30 1 Swinbladder Presence/Absence 30 2 Dnnming Riscles Presence/Absence 30 2

(Xialitative Characters

Ctaupleteness of Lateral Line 30 1 Position of Lateral Line 30 2 Pectoral Fin Shape 30 1 Pectoral Fin Condition 30 1 Position of Pectoral Fin Relative to Center of Gravity 30 1 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 3 Position of Eyes 30 1 Upper and lower Jaw Dentition 30 3 Upper and Lower Iharyngeal Dentition 30 2 Shape of Vomer 30 4 Feeding Apparatus Pattern 30 4 Table 32. Morpheme trie, Meristic and Qualitative Characters Belated to Feeding in Barathritea iris.

CHARACTER M RANGE X S.D. Morphometries XSL

Standard Length 1 530 Body depth 1 23.0 Index of Trunk Shape 1 41.0 Pectoral Fin Length 1 10.0 Aspect Ratio of Pectoral Fin 1 48 Lateral Mouth Gape 1 7.1 Medial Mouth Gape 1 9.2 Upper Jaw Length 1 7.3 Upper Jaw Frotrusibilify 1 90.2 Snout Length 1 4.8 Inter Raker Gap Distance 1 0.45 Greatest Raker length 1 0.75 Branchial Seive Filter Area 1 45 Stomach Length 1 13.2 Intestinal Tract Length 1 132.0 Tbtal Gut Length 1 138.9 Swinbladder Length 1 9*4 Swinbladder Width 1 4.5 Rete Length 1 11*3 Eye Diameter 1 0.66

Meristics

Nurber of Developed Bakers 1 7 Ruber of Pyloric Caeca 1 0 Ninber of lateral Lines 1 1 Ruber of Basibranchial Patches 1 1 Swinbladder Presence/Absence 1 2 Dnnming Muscles Presence/Absence 1 2

Rialitative Characters

Ccapleteness of Lateral Line 1 1 Position of lateral Line 1 2 Pectoral Fin Shape 1 1 Pectoral Fin Condition 1 1 Position of Pectoral Fin Relative to Center of Gravity 1 1 Position of Dorsal fin Relative to Center of Gravity 1 2 Position of Anal Fin Relative to Center of Gravity 1 4 Mouth Orientation 1 3 Position of Eyes 1 1 Upper and Lower Jaw Dentition 1 3 Upper and Lower Iharyngeal Dentition 1 2 Shape of Vomer 1 4 Feeding Apparatus Pattern 1 4 Table 33* Morpbometric, Meristic and Qualitative Characters Related to Feeding in AbyssnbTotula BAlnt-h^ae.

CHARACTER N RANGE X S.D. Morphometries ZSL

Standard Length 1 128 Body depth 1 15.0 Index of Trunk Shape 1 34 Pectoral Fin length 1 25 Aspect Ratio of Pectoral Fin 1 12 Lateral Mouth Gape 1 7 .6 Medial Mouth Gape 1 11.5 Upper Jaw length 1 9.6 Upper Jaw Protrusibility 1 79 Shout Length 1 9.6 Inter Raker Gap Distance 1 0.72 Greatest Raker length 1 3.12 Branchial Seive Filter Area 1 88.5 Stomach Length 1 5.8 Intestinal Tract Length 1 21.0 Total Gut Length 1 26.0 Swinbladder Length 1 5.2 9winbladder Width 1 3.7 Rete length 1 14.1 Eye Diameter 1 1.7

Meristics

Ruber of Developed Rakers 1 9 Ruber of fyloric Caeca 1 0 Ruber of lateral Lines 1 1 Ru b e r of Basibranchial Patches 1 2 Swinbladder Presence/Absence Dimming Muscles Presence/Absence

Oialitative Characters

Completeness of Lateral Line 1 1 Position of lateral Line 1 2 Pectoral Fin Shape 1 3 Pectoral Fin Condition 1 1 Position of Pectoral Fin Relative to Center of Gravity 1 2 Position of Dorsal fin Relative to Center of Gravity 1 2 Position of Anal Fin Relative to Center of Gravity 1 4 Mouth Orientation 1 3 Position of Eyes 1 1 Upper and lower Jaw Dentition 1 2 Upper and lower Pharyngeal Dentition 1 2 Shape of Vomer 1 3 Feeding Apparatus Pattern 1 4 Table 34. Mean Values of Marphological Characters Related to Feeding for Each Species in Groups V and VI. PCtHbrogadus miles. FEM=Rpnonis mardnrmlrii. POS^Rjrogadus silus. . and AA=Acanthonu8 annatus.

Character/Species P® MH POS POC M ______(eroun V)______(Vi)

Body Depth 10.5 9.8 9.6 8.4 22.8 Trunk Shape Index 33.0 36.0 29.1 27.0 28.0 Pectoral Fin length 9.3 8.4 5.3 9.4 8.7 Pectoral Fin Aspect Ratio 16.1 21.0 35.0 18.1 54.0 Lateral Gape 7.5 8.0 7.6 6.5 15.1 Medial Gape 9.9 11.2 11.5 8.2 18.6 Upper Jaw length 9.0 9.2 9.6 7.7 14.4 Upper Jaw Protrusibility 87.3 91.0 80.0 82.0 80.1 Snout Length 4.9 8.4 4.6 3.6 7.5 Eye Diameter 3.2 1.1 3.6 2.7 2.8 No. Developed Rakers 15.0 9.0 16.0 15.0 18.5 Later Raker Gap Distance 0.54 0.68 0.65 0.51 0.60 Greatest Raker Length 2.9 2.5 2.9 3.2 4.1 Branchial Filter Area 202.0 86.0 141.0 246.0 385.0 Stomach length 8.9 5.7 7.4 5.1 5.7 Intestinal Tract Length 46.0 39.0 22.0 28.0 30.8 Total Gut Length 55.0 44.8 29.0 34.0 36.5 Swinbladder Length 8.6 11.0 11.4 9.6 0.0 Rate Length 10.3 13.0 22.4 12.8 0.0 Pyloric Caeca 4 0 0 6 0 Pectoral Fin Shape 2 2 2 2 2 Pectoral Fin Condition 1 1 1 1 1 Pectoral Fin Relative to OG 2 2 1 2 4 Dorsal Fin Relative to OG 2 2 2 2 4 Anal Fin Relative to OG 4 4 4 4 4 Mouth Orientation 3 4 2 2 3 Jaw Dentition 2 2 2 2 2 Pharyngeal Dentition 2 2 2 2 2 No. Basibranchial patches 1 2 1 1 2 faner Shape 4 3 2 2 4 Swinbladder Presence/Absence 2 2 2 2 2 Ooopletion of lateral line 1 1 1 1 1 Position of Lateral Line 4 4 4 4 1 Eye Position 1 1 1 1 1 No. of Lateral Lines 3 3 3 3 1 Drunmng Miscles Presence/Absence 2 2 2 2 1 Feeding Apparatus Pattern 1 1 1 1 5 Table 35. Ebrphanetric, Msristic and Qualitative Characters Related to Feeding in Boroeadus silus.

CHARACTER______N RANGE X S.D. Morphometries %SL

Standard Length 30 118-132 125 5.4 Body depth 30 7.3-10.8 9.6 1.1 Index of Trank Shape 30 23.1-34,3 29.1 2 3 Pectoral Fin Length 30 43-5,8 5 3 0.47 Aspect Ratio of Pectoral Fin 30 30.1-38.4 35 2.0 lateral Mouth Gape 30 6.1-10.3 7.6 1.0 Medial Mouth Gape 30 9.4-13.1 11.5 1.0 Upper Jaw length 30 8.0-11.5 9.6 0.9 Upper Jaw Protrusibility 30 69.3-91.2 80 4.5 Snout Length 30 3.8-6.0 4.6 0.5 Inter Raker Gap Distance 30 0.48-0.72 0.65 0.03 Greatest Raker Length 30 1.8-3.4 2.9 0 3 Branchial Seive Filter Area 30 103-162 141 1 8 3 Stomach length 30 5.6-9.1 7.4 1.1 Intestinal Tract Length 30 203-26.2 22 2.0 Tbtal Gut Length 30 26.4-32.1 29 2.0 Swinbladder length 30 93-12.9 11.4 0.9 Swinbladder Width 30 6 .1-10.2 8 3 1.1 Rete Length 30 18.4-263 22.4 2.0 Eye Diameter 30 3.2-3.8 3.6 0.02

Meristics ttaber of Developed Rakers 30 14-18 16 0.9 Nurber of Pyloric Caeca 30 0 Niraber of Lateral Lines 30 3-3 3 Nmber of Basihranchial Patches 30 1-1 1 Swinbladder Presence/Absence 30 2 Swimning Miscles Presence/Absence 30 2

Qualitative Characters

Completeness of Lateral Line 30 1 Position of Lateral Line 30 4 Pectoral Fin Shape 30 2 Pectoral Fin Condition 30 1 Position of Pectoral Fin Relative to Center of Gravity 30 1 Position of Dorsal fin Relative to Center of Gravity 30 2 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 2 Position of Eyes 30 1 Upper and lower Jaw Dentition 30 2 Upper and lower Pharyngeal Dentit: 30 2 Shape of Vomer 30 2 Feeding Apparatus Pattern 30 1 Table 36. Efcrphanetric, Meristic and Qualitative Characters Related to Feeding in Borogadus catena.

CHARACTER N RANGE X S.D, M»rbbametrics %SL

Standard Length 29 130-250 187 38 Body depth 29 7.9-113 8.4 1.1 Index of Ttunk Shape 29 23.1-31.4 27 2 3 Pectoral Fin length 29 7.8-10.5 9.4 1.2 Aspect Ratio of Pectoral Fin 29 15.1-26.4 18.1 2.5 lateral Mouth Gape 29 53-7.2 6.5 1.2 Medial Mouth Gape 29 7.1-93 8 3 1.0 Upper Jaw Length 29 6.9-10.0 7.7 1.0 Upper Jaw Protrusibility 29 75-92 82 3.9 Snout Length 29 2.9-4.5 3.6 0.4 Inter Raker Gap Distance 29 038-0.68 0.51 0.2 Greatest Raker length 29 2.9-4.1 3.2 0.2 Branchial Serve Filter Area 29 238-267 246 12 Stomach Length 29 4.1-5.9 5.1 0.4 Intestinal Tract length 29 14.4-32.1 28.0 0.2 Tbtal Gut Length 29 28.0-35.0 34 2.1 Swinbladder length 29 73-11.8 9.6 1.0 Swinbladder Width 29 2.9-53 4.1 1.0 Rete Length 29 10.1-14.1 12.8 1.5 Rye Dimeter 29 23-3.0 2.7 0.2

Meristics

Nmber of Developed Bakers .29 13-18 16 2.2 Nmber of Pyloric Caeca 29 6-6 6 Nmber of lateral Lines 29 3-3 3 Nmber of Basibranchial Patches 29 1 Swinbladder Presence/Absence 29 2 Drusoing Miscles Presence/Absence 29 2

Qualitative Characters

Conpleteness of Lateral Line 29 1 Position of Lateral Line 29 4 Pectoral Fin Shape 29 2 Pectoral Fin Condition 29 1 Position of Pectoral Fin Relative to Center of Gravity 29 2 Position of Dorsal fin Relative to Center of Gravity 29 2 Position of Anal Fin Relative to Center of Gravity 29 4 Mouth Orientation 29 2 Position of Eyes 29 1 Upper and lower Jaw Dentition 29 2 Upper and lower Pharyngeal Dentition 29 2 Shape of Vomer 29 3 Feeding Apparatus Pattern 29 1 liable 37. Morpheme trie, Meristic and Qualitative Characters Related to Feeding in Pacogadus miles.

CHARACTER N RANGE X S.D. MomhometricB ZSL

Standard Length 12 181-439 318 85 Body depth 12 9.8-123 10.5 1.6 Index of Trunk Shape 12 29.1-36.4 33 2.1 Pectoral Fin Length 12 83-11.2 9 3 0.9 Aspect Ratio of Pectoral Fin 12 13.7-18.5 16 0.8 Lateral Mouth Gape 12 63-9.8 7.5 0.9 Medial Mouth Gape 12 83-11.4 9.9 1.0 Upper Jaw Length 12 8.5-11.3 9.0 1.0 Upper Jaw Protrusibility 12 86.1-93 8 7 3 3.1 Snout Length 12 3.8-6.1 4.9 0.9 Inter Raker Gap Distance 12 0.48-0.68 0.54 0.1 Greatest Raker Length 12 1.8-3.1 2.9 0 3 Branchial Seive Filter Area 12 173-231 202 20 Stomach Length 12 6.9-11.1 8.9 1.7 Intestinal Tract length 12 38-54 46 1.8 Tbtal Gut Length 12 49-61 55 2 3 Swinbladder length 12 7.1-9.2 8.6 0.9 Swinbladder Width 12 3.8-63 4.9 1.0 Rete length 12 9.2-123 1 0 3 1.0 Eye Diane ter 12 2.7-3.5 3.2 0.8

Meristics

Huber of Developed Rakers 12 12-16 15 2.2 Huber of Pyloric Caeca 12 4-4 4 Huber of Lateral Lines 12 3-3 3 Huber of Basibranchial Patches 12 1 Swinbladder Presence/Absence 12 2 Pnuming Miscles Presence/Absence 12 2

Qualitative Characters

Completeness of Lateral Line 12 1 Position of Lateral Line 12 4 Pectoral Fin Shape 12 2 Pectoral Fin Condition 12 1 Position of Pectoral Fin Relative to Center of Gravity 12 2 Position of Dorsal fin Relative to Center of Gravity 12 2 Position of Anal Fin Relative to Center of Gravity 12 4 Mouth Orientation 12 3 Position of Eyes 12 1 Upper and Lower Jaw Dentition 12 2 Upper and lower Miazyngeal Dentition 12 2 Shape of Vomer 12 4 Feeding Apparatus Pattern 12 1 Table 38. Morpheme trie, Meristic and Qualitative Characters Belated to Feeding in Benonus macdonaldi.

CHARACTER N RANGE X S.D. Morohometries ZSL

Standard I^ngfh 29 105-237 183 33 Body depth 29 8 .1-12.0 9.8 1.1 Index of Trunk Shape 29 36.0 Pectoral Fin Length 29 7.0-10.0 8.4 1.5 Aspect Ratio of Pectoral Fin 29 18.1-25.2 21 0.9 lateral Mouth Gape 29 63-9.9 8.0 1.0 Medial Mouth Gepe 29 9.2-13.1 1 1 3 0.8 Upper Jaw length 29 73-10.1 9 3 1.1 Upper Jaw Protrusibility 29 78-101 91 6 3 Snout Length 29 7.9-103 8.4 1.1 T n tp r Baker Gap Distance 29 0.41-0.72 0.68 0.08 Greatest Raker Length 29 2.1-3.0 2.5 0.4 Branchial Seive Filter Area 29 713-102 86 7.5 Stomach Length 29 53-6.1 5.7 0.8 Intestinal Tract length 29 36.1-423 39.0 2.6 Total Git Length 29 41.1-483 44.8 4.4 Swinbladder Length 29 9 .0 -12.2 11.0 1 Swinbladder Width 29 5.9-9.8 73 1.3 Rete Length 29 11.8-13.8 13.0 0.8 Eye Diameter 29 0.8-1.4 1.1 0.1

Meristics

Nmber of Developed Rakers 29 8 .1-10.2 9 1 Nmber of Pyloric Caeca 29 0 Nmber of Lateral Lines 29 3-3 3 Nuaber of Basibranchial Patches 29 2-2 2 Swinbladder Presence/Absence 29 2 Damning Miscles Presence/Absence 29 2

Q ytlitativp OwrflcterB

CmpletenesB of lateral Line 29 1 Position of lateral Line 29 4 Pectoral Fin Shape 29 2 Pectoral Fin Condition 29 1 Position of Pectoral Fin Relative to Center of Gravity 29 2 Position of Dorsal fin Relative to Center of Gravity 29 2 Position of Anal Fin Relative to Center of Gravity 29 4 Mouth Orientation 29 4 Position of Eyes 29 1 Upper and lower Jaw Dentition 29 2 Upper and lower Pharyngeal Dentition 29 2 Shape of Varner 29 3 Feeding Apparatus Pattern 29 1 Table 39. Mirphometric, Meristic and Qualitative Characters Related to Feeding in Acanthoma armatus.

CHARACTER______N RANGE X S.D. Morphometries %SL

Standard Length 30 165-320 285 10.8 Body depth 30 18.1-25.3 22.8 1.4 Index of Trunk Shape 30 25-33 28 1.2 Pectoral Fin Length 30 6.9-103 8.7 1.0 Aspect Ratio of Pectoral Fin 30 48.1-623 54 4.0 Lateral Mouth Gape 30 13.1-17.9 15.1 1 3 Medial Mouth Gape 30 15.8-22.1 18.6 1.5 Upper Jaw Length 30 12.8-15.7 14.4 1.0 Upper Jaw Protrusibility 30 72.1-853 80.1 3.5 Snout Length 30 43-9.0 7.5 1.3 Inter Raker Gap Distance 30 0.52-0.66 0.60 0.2 Greatest Raker length 30 33-4.8 4.1 0.4 Branchial Seive Filter Area 30 328-436 385 32 Stanacb Length 30 3.0-7.0 5.7 1.0 Intestinal Tract Length 30 21.1-383 30.8 4.1 Total Gut Length 30 253-42.1 36.5 4.0 Swinbladder length 30 - Swinbladder Width 30 - Rete Length 30 - Eye Dimeter 30 1.9-3.2 2.8 0.3

Meristics

Nmber of Developed Rakers 30 17-21 18.5 1 3 Nmber of Pyloric Caeca 30 0 Nmber of Lateral Lines 30 1 Nmber of Basibranchial Patches 30 2-4 1.0 Swinbladder Presence/Absence 30 1 Drraming Miscles Presence/Absence 30 1

Qualitative Characters

Goqpleteness of Lateral Line 30 1 Position of Lateral Line 30 1 Pectoral Fin Shape 30 Pectoral Fin Condition 30 1 Position of Pectoral Fin Relative to Center of Gravity 30 4 Position of Dorsal fin Relative to Center of Gravity 30 4 Position of Anal Fin Relative to Center of Gravity 30 4 Mouth Orientation 30 Position of Eyes 30 1 Upper and Lower Jaw Dentition 30 2 Upper and lower Pharyngeal Dentition 30 2 Shape of Vomer 30 4 Feeding Apparatus Pattern 30 5 Ill

Feeding apparatus Model:

Although morphological differences exist in various components of the feeding apparatus among deep-sea ophidiids examined, these Bpecies comprise a relatively uniform group with respect to the mechanisms involved in jaw movements related to feeding. Dicrolene intronigra illustrates the representative condition of the feeding apparatus in deep-sea ophidiids and serves as a model to describe the structures and mechanisms involved in feeding.

Osteology:

Neurocranium. The neurocranium of D. intronigra comprises the dorsal elements of the skull (figs. 31b, 53d). The olfactory region includes the lateral ethmoids, dermetbmoids, vomer and the nasals. The orbital region is composed of the frontals, pterosphenoid, basisphenoid, and the circumorbitalB. The otic region includes the prootics, sphenotics, epioccipitals, exoccipitals, supraoccipital, and the parietals. The basicranial region consists of the basioccipital and the elongated parasphenoid.

Circumorhitals. .These bones enclose the orbit ventrally and are composed of six loosely connected semi-enclosed elements (fig.

31b). The most anterior elongated element is the lacrymal. It partially overlies the maxilla, premaxilla, and covers the palatine.

The sixth suborbital partially overlies the sphenotic.

Susnenaorium .The bones of the suspensorium attach dorsally to the neurocranium, anteriorly to the jaws, and posteriorly to the opercular boneB (fig. 34f). The suspensorium consist of the palatine, entopterygoid, metapterygoid, ectopterygoid, preopercle, quadrate, symplectic, and hyomandibula. The suspensorium is attached to the 112 neurocranium anterior to the eye by the palatine and entopterygoid, and posterior to the eye by the metapterygoid and hyomandibula . The quadrate articulates with the articular by means of a ball and socket joint.

Opercular series .These bones provide a lateral wall to the branchial region (fig. 37a). A ball and socket joint connects the dorsal-most opercle to the hyomandibula. The preopercle, functionally included with the suspensorium, is attached to the quadrate ventrally by a ligament, and it partially overlies and is attached to the hyomandibula. The interopercle is attached to the angular by a ligament as well as medially to the epihyal.

Jaws .The lower jaw (mandible) consist of the tooth-bearing dentary, the articular and angular (fig. 40a). A ball and socket joint between the articular and quadrate allows a hinge-like movement.

The articular is attached to the angular which itself is bound by a ligament to the interopercle. A notch in the dentary and ligaments between it and the articular facilitate a firm connection.

The upper jaw consists of a tooth-bearing premaxilla, maxilla and supramaxilla (fig. 4 0 a ) . premaxilla are fused at their symphysis by connective tissue and are similarly attached at their posteroventral edges to the rostral cartilage. This cartilage rest on the dermethmoid and vomer during protrusion of the upper jaw. The length of the premaxilla is due to its relatively long ascending process. The maxilla has an internal, dorsally directed condyle that articulates with the premaxilla. The palatine overlies the dorsal section of the maxilla laterally. Gill arches .The anterior first gill arch possesses 3-4 rudimentary knob-like rakers and two developed rakers on the upper limb, 1 developed at the angle plus eleven-thirteen developed rakers on lower limb (fig.

46a, 49f).

Hyoid apparatus .The hyoid apparatus consist of a basihyal, dorsal and ventral hypophyals, epihyals, ceratohyals, and small interhyals (fig.

39a, 46a). A keel-shaped urohyal lies between these bones. Eight branchiostegal rays lie lateral to and overlie the epihyal and ceratohyal.

Branchial apparatus .The branchial bones are connected to and continuous with the hyoid bones. From ventral to dorsal, the branchial series is composed of a pair of basibranchials, the second with teeth, three bilateral pairs of hypobranchials, the third with a small tooth patch, five pairs of ceratobranchials, with the fifth pair having lower pharyngeal teeth, four pairs of epibranchials; pharyngobranchial one attached to the first epibranchial, and fused pharyngobranchials two- four bearing the upper pharyngeal teeth (figs. 46a, 49f).

Girdles .The pectoral girdle consist of the posttemporal, extrascapulars, supracleithrum, cleithrum, scapula, coracoid, and postcleithrum (figs. 52e). The pectoral fin is attached to the scapula and coracoid by bony radials. Dorsally the posttemporal is atttached to the skull. The basipterygium of the pelvic girdle has its anterior edge wedged within the cleithra and coracoids of the pectoral girdle.

Connective tissue elements:

Connective tissue elements can be distinct ligaments or tendons or thickened regions of the integument composed mostly of collagenous fibers. The more important elements are listed by region. 114

Snout region .A thickened ring of dermis encircles the mouth. This

tissue overlies and is attached to the ventral portion of the maxilla and from there continues to attach to the dorsal edge of the articular.

When the jaw is protruded, the sheath is pulled taut, filling in the sides of the mouth.

The ventropremaxillomaxillary ligament is a short stout ligament that runs from the posteroventral end of the descending process of the premaxilla to the anteroventral portion of the maxilla medially (fig.

60, vlpm).

The articulodentomaxillary ligament is broad ligament that attaches the medial posteroventral aspect of the maxilla to the dentary and articular in the region of their symphysis. The palatomaxillary ligament is a short, stout ligament that ascends from the anterior tip of the palatine, and inserts on the anterolateral face of the dorsal process of the maxilla (fig. 60, 1 pm).

The nasomaxillary ligament unites the anterior tip of the nasal to the posterodorsal aspect of the maxilla. These fibers plus fibers that unite the dorsal tips of the maxilla across the midline are part of the thick sheath that encapsulates the ascending process of the premaxilla and stretches from lacrymal to lacrymal.

The premaxillary palatine ligament (lpp) and the ethmomaxillary ligament (lem) together form the "cross" ligament. The premaxillary palatine ligament extends from the distal end of the ascending process of the premaxilla posteriorly to the base of the maxillary process of the palatine. The ethmomaxillary ligament extends from the lateral surface of the head of the maxilla to the anterior surface of the ethmoid (fig. 60). 115

The maxillomaxillary ligament is a broad ligament that extends from

the lateral surface of the rostral cartilage to the medial surface of

the head of the maxilla (fig. 60, 1mm ) .

The lacrimomaxillonasofrontal tissue sheath is a thick, wide band

of dermis that originates on the lacrymals, fuses with the dorsolateral

portions of the maxilla, wraps around the anterodorsal border of the maxilla, and attaches to the nasals extending completely over from one

side to the other. Its posterodorsal attachment is to the frontals.

Two apertures pierce this thickened integument on either side. This

entire structure forms a sheath over and lateral to the ascending

process of the premaxilla.

A band of elastin fibers attaches to the dorsal aspect of the

premaxilla. Anteriorly they arise from the dermethmoid just posterior

to the premaxilla. They run over the dorsal surface of the premaxilla.

They insert on the anterodorsal symphysis of the premaxilla, and the

ligamentous sheath enclosing the premaxilla.

Opercular region .The interoperculomandibular ligament unites the

anteroventral corners of the interopercular to the angular.

The anteroventral process of the preopercle is joined by the

preoperculoquadrate ligament both to the ball of the quadrate and to the

posterior ridge of the quadrate that abuts the preopercle.

The adductor mandibulae tendon attaches divisions Al of the ^

adductor mandibulae to the maxilla (fig. 61). This strong tendon originates approximately midway as tendinous sheath and inserts as a

thick, round tendon into a notch ventral to the cranial condyle of the maxilla. 116

Hyoid region .The interoperculoephihyal ligament is a short flat ligament that attaches laterally to the posterodorsal section of the epihyal and runs anteroventrally to attach to the interopercle medially uniting the hyoid apparatus to the jaw mechanism.

The interhyal is united to the epihyal by short connective fibers and is united to the medial side of the metapterygoid by a short metapterygointeryal ligament.

The ceratohyoglossohyal ligaments lie medial to the left and right protractor hyoideus muscle bundles. They unite the anterior end of the glossohyal to the anterolateral aspects of the ceratohyal on both sides.

Myology:

Lateral head muscles .The various divisions of the adductor mandibulae are very difficult to determine and even the path of the Ramus

Mandibularis of the V nerve which innervates the muscle is an unreliable character. As a result the terminology is often varied and confused

(Winterbottom, 1974). The nomenclature of the various divisions found in Dicrolene intronigra is based on the generalized origins and insertions described by Tetter (1878) and Gregory (1933) and comparisons with other studies (Alexander, 1967a; Osse, 1969; Elshoud-Oldenhave and

Osse, 1976; Liem and Osse, 1975).

Divisions of the adductor mandibulae are the most superficial bundles and are exposed as the skin iB removed in the cheek region

(figs, 61, 62). This large subdivided muscle originates primarily from the lateral surface of the palatal arch and preopercle and inserts on the lower and upper jaws. Innervation is by branches of the fifth cranial nerve. 117

Division Al originates on the lateral surfaces of the pterygoid bones (ectopterygoid, mesopterygoid and metapterygoid). The muscle is further subdivided into a ventrolateral section Al and a more dorsal section Al (fig. 61, 62). Al lies completely medial to A2 and is the only major muscle serving the upper jaw.

Division A2 occupies the ventrolateral region of the cheek. It possesses a tendinous insertion on the posterior face of the coronoid process of the dentary and shares a sheath-like connection vith fibers of Aw inserting in the Meckelian fossa. Two subdivisions of this muscle are present. A2 and A2 refer to the dorsal and ventral subdivisions respectively (figs. 61, 62).

Division A3 of the adductor mandibulae lies on the lateral surface of the palatal arch beneath A2. It inserts on the medial face of the dentary and to the articular (figs. 61, 62).

Division Aw, frequently referred to as the intramandibularis muscle, inserts in and fills the Meckelian fossa on the medial face of the dentary and attaches to the articular as well (figs. 61, 62).

The adductor arcus palatini fills the fissura infraorbitalis forming the floor of the orbit (figs. 61, 62; AAP). It originates on the parasphenoid and posteriorly on the prootic wings. The fibers insert on the most medial edge of the metapterygoid, and posteriorly on the hyomandibula.

The levator arcus palatini originates from the sphenotic and inserts on the hyomandibula (figs* 61, 62; LAP).

T h e dialator operculi lies deep to the levator arcus palatini; the base points posteroventrally (figs. 61, 62; DO). Fibers originate from the sphenotic and hyomandibula and insert on the anterodorsal edge of 118

the opercle. Fibers of the levator operculi originate anteriorly very

close to the dorsal origin of the dialator operculi (figs. 61, 62; L O ) .

T he levator operculi fibers originate mainly on the pterotic and

posttemporal. All fibers insert on the dorsomedial side of the opercle,

just posterior to the ball and socket joint with the hyomandibula.

Fibers of the adductor operculi and arcus palatini originate from

the prootic. Fibers originate medial to those of the levator operculi, merge with those of the levator operculi, and insert on the opercle.

Ventral head muscles .The intermandibularis runs transversely from the dentary in the region anterior to the cranial tip of the basihyal (fig.

63; 1MD). A thick sheath of connective tissue dorsal to this muscle forms the floor of the buccal cavity.

The protractor hyoideus connects the hyoid arch to the lower jaw

(fig. 60; P R H Y ) . Posteriorly this muscle is attached to the lateral face of the epihyal and the lateral and posteroventral edge of the ceratohyal. Anteriorly both sides of thiB muscle fuse to attach to the

inner base of the dentary and the connective tissue covering the posterior margin of the intermandibularis.

The hyohyoides inferioris lies covered by the hyohyoidei abductors.

Anteroventrally it meets in a raphe that iB attached to the ventral hypohyal and the glossohyal. Another insertion runB from the anterior end of the left muscle bundle and inserts on the right ventral hypohyal.

Posteriorly, this muscle attaches to the ceratohyal.

The hyohyoidei abductores originates ventrally from a caudally directed protruberance of the ventral hypohyal (fig. 63; HAB). Fibers course dorsocaudally, external to the hyohyoidei inferioris, to insert 119

on the ventral most branchiostegal ray. It then continues a as sheet to

attach to subsequent branchiostegal rays.

The sternohyo'ideus muscle binds the cleithrum to the hyoid

apparatus by attaching to the urohyal dorsolaterally (fig. 63; S T H ) .

Posteriorly this muscle borders the pelvic fin musculature. The

dorsal most fibers of this division attach to the dorsal edge of the

urohyal.

Extrinsic branchial musculature .The pharyngohyoideus muscle extends

from the fifth ceratobranchial to the urohyal. The pharyngocleithralis muscle is differentiated into two heads. The pharyngocleithralis

externus muscle that runs from the anterior tip of the horizontal arm of

the cleithrum and continues dorsally to insert on the ventral Burface of

the fifth ceratobranchial. The second head, the pharyngocleithralis

internus, originates dorsally from the anterior surface of the

cleithrum.

Intrinsic branchial musculature .As noted by Liem (1970:60), "All

intrinsic branchial muscles are adductors. Although in a few instances

* adduction in one part of the branchial apparatus may result in abduction of another region." According to Liem (1970:60) "the attachment is

substituted for origin and insertion since in all cases there is great mobility of the bony elements on either side of the muscle, making it

impossible to distinguish between origin and insertion.”

Biomechanics:

Since Schaeffer and R o s e n ' s (1961) review of the evolution of the feeding mechanism in ray~finned fishes our understanding of the relation between structure and function has increased substantially. While phylogenetic analyses of actinopterygian evolutionary patterns have 120

provided information on the historical sequence of structural change

(Greenwood et al. 1966, 1973,; Rosen, 1982) experimental functional

morphology has furthered our knowledge of the relationship between form

and function in fishes (Alexander, 1966, 1967, 1970; Anker, 1974;

Lauder, 1979; Liem, 1970; Osse, 1969).

Motta (1984) contends that despite growing knowledge on the

phylogeny of the protrusible jaw in fishes studies on mechanics of jaw

protrusion are highly variable and conflicting. Be contrasts four basic

mechanisms of upper jaw protrusion among teleost fishes. Two of theBe

mechanisms are most common among teleost fishes examined to date. Type

"A" where depression of the mandible pulls or pushes the premaxilla

passively into the protruded condition, and type "B" where twisting of

the maxilla around its long axis pulls the premaxilla into the protruded

position. The first mechanism is apparently more prevalent among

teleosts studied than was previously suspected. The twisting maxilla

model proposed by Alexander (1967a, b) may have been overemphasized due

to simplistic experimental design.

In order to accurately determine which mechanism is responsible for

upper jaw protrusion in deep-sea ophidiids it would be necessary to

employ such sophisticated techniques as high speed cinephotograpy and

radiography, eletrical muscle stimulation experiments, connective tissue

severance experiments and electromyography. Unfortunately such

techniques were beyond the scope of this study since it was based on manual manipulation of freshly killed and/or frozen specimens.

In this study the skull was defined to comprise discreet units that

function together in complex movements to perform the function of gill

ventilation and feeding (Alexander, 1966, 1967a,b, 1970; Anker, 1974; 121

Ballintijn and Hughes, 1965; Gans, 1969; Liem, 1967, 1970; Lauder,

1979, 1980; Motta, 1982; Osse, 1969; Tchernavin, 1948, 1953).

Th e functional relationship of these mechanical units and couplings must be considered to fully understand trophic specializations and feeding adaptation in these fishes. A basic description of the mechanical units and their movements followed by a description of the couplings serve to illustrate the feeding mechanism in deep-sea neobythytines. Dicrolene introniera possessed those structural features

typically associated with the generalized teleost feeding mechanism

(Lauder, 1982) such as the division of the premaxilla into a lateral toothed portion and a medial portion closely associated with the ethmoid

(palatine) complex (Patterson, 1973).

Mouth opening .Several major couplings are responsible for opening the mouth. The Bternohyoides-hyoid-interopercle-mandible coupling is responsible for lowering the mandible. Forcibly abducting the sternohyoides by pulling caudally on its posterior edge results in

t pulling the hyoid caudally. This contraction causes the hyoid bars to spread laterally and rotate moving the symphysiB ventrally. The interopercle via the epihyal is pulled posteroventrally which causes the interoperculo-mandibular ligament to pull on the posteroventral corner of the mandible. The mandible rotates around the quadrate depressing the anterior end of the lower jaw.

Cranial elevation is accomplished by the action of the epaxial muscles-cranium-suspensorium coupling. Contraction of the epaxial muscles lifts the cranium and associated suspensorium. This actions lifts the anterior corner of the palatine, maxilla and premaxilla. The quadrate rotates anteriorly depressing the anterior portion of the lower 122 jaw ventrally. The overall effect of lifting the cranium is to depress the lower jaw slightly and expand the oral cavity.

The levator-operculi-opercular coupling is also involved in depressing the lower jaw. Forcibly abducting the opercle by pulling dorsolaterally on its posterior edge results in a slight depression of the lower jaw and very little premaxillary protrusion. This action which simulates contraction of the levator operculi pulls the opercle which inturn pulls on the subopercle, interopercle and interopercular mandibular ligament. This ligament is firmly attached to the mandible causing it to rotate around the quadrate resulting in a slight depression of the lower jaw.

Lateral expansion of the mouth (oral cavity) is accomplished by the action of the levator arcus palatini-suspensorium coupling. Contraction of the levator arcus palatini muscle causes the suspensorium to move laterally. A corresponding lateral movement of the quadrate causes the dentaries to move apart from each other and the lower jaw to drop. The action of other muscle couplings (i.e. hyoid depression) also results in oral cavity expansion but to a lesser degree than the levator arcus palatini.

Upper jaw protrusion .Upper jaw protrusion in teleost fishes has been the focus of much research over the years (Alexander, 1967; Eaton, 1935;

Gregory, 1933; Lauder and Liem, 1981; Liem, 1970, 1979, 1980; Nyberg,

1971; Pietsch, 1978; Schaeffer and Rosen, 1961; Van Dobben, 1937).

Theorectically both mechanisms are possible among deep-sea ophidiids. The articular process of the premaxilla, a morphological prerequisite for the screw mechanism of protrusion postulated by

Alexander (1967a) is present in D. intronigra. The maxilla can either swing about a transverse axiss through the head of the maxilla or it can

rotate or "twist" along its long axis. Manual twisting of the maxilla

in dead specimens causes the jaws to protrude. However manual

depression of the lower jaw also causes the upper jaw to protrude with

very little maxillary twisting. Apparently the maxilla swings forward due to its attachment to the mandible which in turn causes the rostral

cartilage to press against the ethmoid cartilage effecting protrusion.

Mouth closing .Closing of the mouth is basically a reverse of opening but not in the same sequence since that would result in expulsion of water and food previously drawn into the mouth (Nyberg, 1971).

Contraction of the A2 division of the adductor mandibulae and the

adductor mandibulae w results in bringing the distal end of the mandible dorsally and posteriorly. Contraction of the protractor hyoideus causes

the dentaries and hyoid bars to draw together resulting in closing of

the lover jaw. Contraction of the adductor arcus palatini muscle returns the suspensorium to a closed position. Finally the cranium is returned to its resting position through the action of the body musculature.

The exact movements of the head of D. introniera during feeding are more complex than the simple model presented herein. Hopefully future research in this area using closely related "cusk-eels" of the genus

Ophidion may provide additional support and confirmation of the generalized Ophidiid feeding mechanism described above. Figure 30. A. Bathvonua pectoralis. VIMS 7836, 132 mm SL lateral facing bones.

B. Poroeadus catena. VIMS 7821, 125 mm SL lateral facing bones.

C. Poroeadus silus. VIMS 7839, 125 mm SL lateral facing bones.

D. Penonus macdonaldi. VIMS 7838, 188 mm SL lateral facing bones.

E. Poroeadus miles. VIMS 7837, 321 mm SL lateral facing bones. A !Smm

B c !Smm !Smm

E D !Smm !Smm Figure 31. A. Dicrolene kanazawai. VIMS 7846, 226 mm SL lateral facing bones

B* Dicrolene introniera. VIMS 7847, 218 mm SL lateral facing bones

C. Honomitonus aeassizii. VIMS 7849, 210 mm SL lateral facing bones

D. Barathrodemus manatinus. VIMS 7841, 186 mm SL lateral facing bones

E. Baratbrites parri. VIMS 7834, 302 mm SL lateral facing bones

F. Abvssobrotula galatheae. VIMS 7844, 186 mm SL lateral facing bones 10 mm B 10 mm

m m 10mm

10mm 10 mm Figure 32. A. Acanthonus a p n a t u s . VIMS 7843, 315 ran SL lateral facing bones

B. Xvelacvba invars i . VIMS 7842, 220 mm SL lateral facing bones

C. Bassozetus normalis. VIMS 7831, 695 mm SL mm SL, lateral facing bones.

D. Bassozetus ^aenia. VIMS 7832, 482 mm SL SL, lateral facing bones. B S mm 10mm

10 mm 10mm Figure 33. A. Poroeadus ailua. VIMS 7839, 125 mm SL suspensorium

B. Penopus macdonaldi. VIMS 7838, 188 mm SL suspenBorium

C. Poroeadus catena. VIMS 7838, 185 mm SL suspensorium

D. Poroeadus miles. VIMS 7837, 321 mm SL suspensorium

E. Bathvonus pectoralis. VIMS 7836, 132 mm SL, suspensorium. HM

PAL" ME:

POP

EC

QU‘ SY

2mm

B 2mm 5 mm

5 mm 5mm Figure 34. A. Abyssobrotula ealatheae. VIMS 7844, 186 mm SL 123 mm SL, suspensorium

B. Barathrites parri. VIMS 7834, 302 mm SL suspensorium.

C. Barathrndemufl manatinus. VIMS 7841, 186 mm SL mm SL, suspensorium.

D. Monomitopus aeassizii. VIMS 7849, 210 mm SL SL, suspensorium.

E. Dicrolene kanazawai. VIMS 7846, 226 mm SL suspensorium.

F. Dicrolene introniera. VIMS 7847, 218 mm SL SL, suspensorium. 9 mm B 5mm

5mm 9mm

5mm 5 mm Figure 35. A. Bassozetus normalis. VIMS 7831, 695 m m SL suspensorium B. Bassozetus taenia. VIMS 7832, 482 mm SL suspensorium

C. Xvelacvba mversi. VIMS 7842, 220 mm SL suspensorium.

D. Acanthonus armatus. VIMS 7843, 315 mm SL suspensorium. B 5mm 5mm

5mm Figure 36. A. P^nopus macdonaldi. VIMS 7838, 188 m SL opercular series.

B. Poroeadus silus. VIMS 7839, 125 mm SL opercular series.

C. Poroeadus catena. VIMS 7838, 185 mm SL SL, opercular aeries.

D. Poroeadus miles. VIMS 7837, 321 mm SL SL, opercular aeries

E. Bathvonus uectoralis. VIMS 7836, 132 mm SL opercular series. OP

SOP

I OP-

C 5mm D 5mm Figure 37. A. Dicrolene introniera. VIMS 7847, 218 mm SL opercular series

B* Dicrolene kanazawai. TIMS 7846, 226 mm SL opercular series.

C. Barathrodemus manatinus. VIMS 7841, 186 mm SL SL, opercular series

D. Monomitopus aeassizii. VIHS 7849, 210 mm SL SL, opercular series.

E. Barathrites parri. VIMS 7834, 302 mm SL opercular series.

F. AbvBsobrotula ealatbeae. VIMS 7844, 186 mm SL SL, opercular series. A 5 mm B 5 mm

2 mm D 5mm

5mm 2 m m Figure 38. A. Bassozetus normalis. VIMS 7831, 695 mm SL SL, opercular series.

B. Bassozetus taenia. VIMS 7832, 482 mm SL SL, opercular series

C. Xvelacvba mversi. VIMS 7842, 220 mm SL opercular series

D. Acanthonua armatus. VIMS 7843, 315 mm SL opercular series. 5mm A 5mm B

D 5mm c 5mm Figure 39. A. Penopus macdonaldi. VIMS 7838, 188 mm SL jaw apparatus

B. Poroeadus silus. VIMS 7839, 125 mm SL jaw apparatus

C. Bathvonus pectoralis. VIMS 7836, 132 mm SL jaw apparatus

D. Poroeadus miles. VIMS 7837. 321 mm SL jaw apparatus til

I utuf 5 Figure 40. A. Dicrolene introniera. VIMS 7847, 218 mm SL jaw apparatus

B. Barathrodemus manatinus. VIMS 7841, 186 mm SL mm SL, jaw apparatus.

C. Dicrolene kanazawai. VIMS 7846, 226 mm SL jaw apparatus

D. Abvssobrotula ealatheae. VIMS 7844, 186 mm SL SL, jaw apparatus

E. Monomitopus aeassizii. VIMS 7849, 210 mm SL SL, jaw apparatus

F. Barathrites parri. VIMS 7834, 302 mm SL jaw apparatus. 5mm Figure 41, A. Bassozetus normalie. VIMS 7831, 695 mm SL jaw apparatus.

B. Bassozetus taenia. VIMS 7832, 482 mm SL jaw apparatus.

C. Acanthomis flpnatus. VIMS 7843, 315 mm SL jaw apparatus.

D. Xvelacvba mversi. VIMS 7842, 220 mm SL jaw apparatus. B 5 mm 5 mm

lOmm 5 m m Figure 42. A. Poroeadus catena. VIMS 7838. 185 nun SL hyoid arch.

B. Poro&adus silus_, VIMS 7839, 125 ram SL hyoid arch.

c. Bathvonus pectoraliB. VIMS 7836. 132 mm SL hyoid arch.

D. Porosadus miles.. VIMS 7837, 321 mm SL hyoid arch.

E. PenoDus macdonaldi. VIMS 7838, 188 mm SL SL, hyoid arch. EH ■CH DHH

BR

5 mm

B c 5 mm 5 mm

10 mm Figure 43. A. Dicrolene introniera. VIMS 7847, 218 mm SL hyoid arch.

B. Dicrolene kanazawai. VIMS 7846, 226 mm SL hyoid arch.

C. Monomitopua agaasizii. VIMS 7849, 210 mm SL hyoid arch.

D. Abvssobrotula ealatheae. VIMS 7844, 186 mm SL SL, hyoid arch.

E. Barathrodemus manatinus. VIMS 7841, 186 mm SL SL, hyoid arch.

F. Barathritea parri. VIMS 7834, 302 mm SL hyoid arch. A B - Smm

c Smm

E Figure 44. A. Bassozetus taenia. VIMS 7832, 482 mm SL hyoid arch.

B. Bassozetus normalis. VIMS 7831, 695 mm SL hyoid arch.

C. Xvelacvba raversi. VIMS 7842, 220 mm SL hyoid arch.

D. Acanthonus ppnatus. VIMS 7843, 315 mm SL hyoid arch. E E E o E to

CD

E E E E to 0 to Figure 45. A. PoroRaduB catena. VIMS 7838, 185 imn SL lower branchial series.

B. Porogadus situs. VIMS 7839, 125 mm SL lower branchial series.

C. Porogadus miles. VIMS 7837, 321 mm SL lower branchial series.

D. Bathvonus pectoralis. VIMS 7836, 132 mm ; lower branchial series.

E. Penopus macdonaldi. VIMS 7838, 188 mm SL lower branchial series. 84

5mm

9mm 9mm Figure 46. A. Dicrolene intronigra. VIMS 7847, 218 ran SL lower branchial aeries.

B. Dicrolene kanazawai. VIMS 7846, 226 mm SL lower branchial series.

C. Abvssobrotula galatheae. VIMS 7844, 186 mm SL SL, lower branchial series.

D. Monomitonua aeaaaizii. VIMS 7849, 210 mm SL lower branchial series.

E. Barathrites parri. VIMS 7834, 302 mm SL lower branchial series.

F. Barathrodemus manatinus. VIMS 7841, 186 mm SL SL, lower branchial series. A

c ~mm D ~mm

F 5mm Figure 47. A. Bassozetus normalis. VIMS 7831, 695 mm SL lover branchial series.

B. Bassozetus taenia. VIMS 7832, 482 mm SL lower branchial series.

C. Xvelacvba mversi. VIMS 7842, 220 mm SL lover branchial series.

D. Acanthonus ar ma tus. VIMS 7843, 315 mm SL lower branchial series. c 5mm D 5 mm Figure 48. A. Poroeadus silus. VIMS 7839, 125 nnn SL upper branchial series.

B. Poroeadus miles. VIMS 7837, 321 nnn SL upper branchial series.

C. Poroeadus catena. VIMS 7838, 185 mm SL upper branchial series.

D. Bathvonus nectoralis. VIMS 7836, 132 nnn SL upper branchial series.

E. Penoous macdonaldi. VIMS 7838, 188 mm SL upper branchial series. PB2+P PB3+P

Z m m

Zmm Figure 49. A. Barathrites parri. VIMS 7834, 302 mm SL upper branchial series.

B. DicroXene kanazawai. VIMS 7846, 226 mm SL upper branchial series.

C. Monomitopus agaasizii. VIMS 7849, 210 mm SL SL, upper branchial series.

D. Abvssobrotula ealatheae. VIMS 7844, 186 mm SL upper branchial series.

E. Barathrodemus manatinus. VIMS 7841, 186 mm SL upper branchial series.

F. Dicrolene introniera. VIMS 7847, 218 mm SL upper branchial series. S mm B 2 mm mm

2 mm 2 mm

E 5 mm F S mm Figure 50. A. Xvelacvba mversi. VIMS 7842, 220 mm SL upper branchial series.

B. Acanthonus armatus. VIMS 7843, 315 mm SL upper branchial series.

C. Bassozetus normalie. VIMS 7831, 695 mm SL upper branchial series.

D. Bassozetus taenia. VIMS 7832, 482 mm SL upper branchial series. A 2mm B 2mm

2mm 2mm Figure 51* A* Penoous macdonaldi. VIMS 7838, 188 nnn SL pectoral girdle.

B. Bathvonua pectoralis, VIMS 7836, 132 mm SL pectoral girdle,

C. Poroeadus miles. VIMS 7837, 321 mm SL pectoral girdle.

D. Poroeadus silus. VIMS 7839, 125 mm SL pectoral girdle,

E. Poroeadus catena. VIMS 7838, 185 mm SL pectoral girdle. CO

to Figure 52. A. Monomitopus aeassizii. VIMS 7849, 210 mm SL SL, pectoral girdle*

B. Abvsaobrotula ealatheae. VIMS 7844, 186 mm SL SL, pectoral girdle*

C. Barathrodemus manatinua. VIMS 7841, 186 mm SL SL, pectoral girdle*

D. Barathrites parri. VIMS 7834, 302 mm SL pectoral girdle.

E* Dicrolene introniera. VIMS 7847, 218 mm SL pectoral girdle*

D. Dicrolene kanazawai. VIMS 7846, 226 mm SL pectoral girdle* u

E E "'

E E on Figure 53. A. Bassozetus taenia. VIMS 7832, 482 mm SL pectoral girdle.

B. Bassozetus normalis. VIMS 7831, 695 mm SL pectoral girdle.

C. Xvelacvba mversi. VIMS 7842, 220 mm SL pectoral girdle.

D. Acanthonus armatus. VIMS 7843, 315 mm SL pectoral girdle. 5 mm B 5 mm

D c 5mm 5mm Figure 54. A. Poroeadus catena. VIMS 7838, 185 mm SL neurocranium, dorsal, lateral, ventral views respectively.

B. Penopus macdonaldi. VIMS 7838, 188 mm SL neurocranium, dorsal, lateral, ventral views respectively*

C. Bathvonufl oectoralis. VIMS 7836, 132 mm SL neurocranium, dorsal, lateral, ventral views respectively*

D. Poroeadus silus. VIMS 7839, 125 mm SL neurocranium, dorsal, lateral, ventral, views respectively*

E. Poroeadus miles. VIMS 7837, 321 mm SL neurocranium, dorsal, lateral, ventral, views respectively* ·~

v

BOC

EOC LE

A B ~ ....

D E IC)oooft Figure 55. A. Abvssobrotula ealatheae. VIMS 7844, 186 mm SL SL, neurocranium, dorsal, lateral, ventral, views respectively.

B. Barathrites parri. VIMS 7834, 302 mm SL neurocranium, dorsal, lateral, ventral views respectively.

C. Picrolene kanazawai. VIMS 7846, 226 mm SL neurocranium , dorsal, lateral, ventral views respectively.

P. Picrolene introniera. VIMS 7847, 218 mm SL neurocranium, dorsal, lateral, ventral views respectively.

E. Barathrodemus manatinus. VIMS 7841, 186 mm SL SL, neurocranium, dorsal, lateral, ventral views respectively.

F. Monomitopus aeassizii. VIMS 7849, 210 mm SL SL, neurocranium, dorsal, lateral, ventral views respectively. 8

0 o:;;;;- E --;;;;;;-- Figure 56. A. Acanthonus armatufl. VIMS 7843, 315 mm SL neurocranium, dorsal, lateral, ventral views respectively.

B. Xvelacvba mversi. VIMS 7842, 220 mm SL neurocranium, dorsal, lateral, ventral views respectively.

C. Bassozetus taenia. VIMS 7832, 482 mm SL neurocranium, dorsal, lateral, ventral views respectively.

D. Baasozetus normalis. VIMS 7831, 695 mm SL neurocranium, dorsal, lateral, ventral views respectively. A

o 3 m m C lOinm Figure 57. A* P^fngyduB miles. VIMS 7837, 321 nun SL first left gill arch.

B. Pf>rngfldus catena. VIMS 7838, 185 mm SL first left gill arch.

C* Penopus. macdonaldi. VIMS 7838, 188 mm SL first left gill arch.

D. Bafrhvonus pectoralis. VIMS 7836, 132 mm SL firBt left gill arch.

E. Pp^rpgpdus silus. VIMS 7839, 125 mm SL first left gill arch. LG SGR-

Cfl EB GF

2mm Omm Figure 58. A. Abvssobrotula galatheae. VIMS 7844, 186 mm SL first left gill arcb.

B. Picrolene intronigra. VIMS 7847, 218 mm SL first left gill arch.

C. BarathriteB narri. VIMS 7834, 302 mm SL first left gill arcb.

P. Monomitopus aeassizii. VIMS 7849, 210 mm SL first left gill arch.

E. Barathrodemus manatinus. VIMS 7841, 186 mm SL SL, first left gill arch.

F. Picrolene lcanazawai. VIMS 7846, 226 mm SL first left gill arch. A

E F !lmm Figure 59 A. Bassozetus taenia. VIMS 7832, 482 mm SL first left gill arch.

B. Bassozetus normalis. VIMS 7831, 695 mm SL first left gill arch.

C. Acanthouua annatus. VIMS 7843, 315 mm SL first left gill arch.

D. Xvelacvba mversi. VIMS 7842, 220 mm SL first left gill arch. 5 mm

5 mm

10 mm

5 mm Figure 60. Ligamentous attachments of upper jaw of Picrolene intronigra. (dorsal view) VIMS 7847, 218 mm SL. LPP Figure 61 . Lateral viev of the superficial cheek muscles of Picrolene intronigra. VIMS 7847, 218 mm SL ...... ' Figure 62. Lateral view of the deep cheek muscles of Dicrolene intronigra. VIMS 7847, 218 mm SL. CM

—I

I CL Figure 63 .. Ventral view of cranial muscles Dicrolene intronigra. VIMS 7847, 218 mm SL HAD DISCUSSION

Characterization of feeding strategies:

Fishes occupy the highest trophic levels in the food chain and should as a consequence reflect major differences in energy transport to the deep-sea ecosystem (Sedberry and Husick, 1978).

These differences may be manifested in various ways such as diet, morphology or behavior, or by differences in community structure

(Husick, 1976). The hypothesis is advanced, but still remains untested, that fishes prevalent beneath oligotrophic tropical seas have evolved appropriate energy conserving morphological and behavioral adaptations of which trophic specializations play a major role. The purpose of this study was to characterize feeding strategies of tropical dwelling deep- sea ophidiid fishes through an analysis of food habits and examination of morphological features related to feeding and to interprete these results within the context of the above hypothesis.

The ophidiids, prevalent beneath oligotrophic tropical seas, can best be characterized as benthopelagic fishes that fed predominately on small-sized benthopelagic prey (e.g. copepods, mysids, amphipods, tanaids) although there was a tendency for species attaining a large size to occasionally capture fish and more mobile invertebrate prey.

Although the majority of species fed on small benthopelagic prey, they do not represent what might be called a typical "trophic

158 159 generalist" (Dayton and Hessler, 1972; Ivlev, 1961; Schoener, 1971) in the sense that they fed non-selectively in the prey environment. In fact when the diets of individual species were examined more closely, it was clearly evident that considerable differences existed among species in the kinds, numbers and frequencies of prey items ingested. This evidence suggests that different feeding strategies employed by these fishes may in part represent trophic specializations that enable these deep-sea ophidiids to successfully exploit food resources in deep waters off the tropical Bahamas.

Three main modes of feeding were recognized: (1) predation upon small benthic-benthopelagic or planktonic organisms; (2) predation upon small benthic-benthopelagic or planktonic organism with increasing reliance upon larger planktonic or nektonic prey in large-bodied individuals; and (3) predation upon small benthic organisms with less reliance on benthopelagic and planktonic organisms. The majority of species examined exhibited the first mode of feeding which best typified the overall feeding strategy for these tropical dwelling taxa of fishes.

Included in this group were P. silus. P.. catena. P. milesf A. armatus.

B.. oectoralis. and D., kanazawai. These species employed a predominately off-bottom foraging and facultative benthic-benthopelagic feeding strategy. Bassozetus normalis and B. taenia displayed the second main mode of feeding. Smaller individuals of these species fed in a similar manner as those species in group I. However larger individuals displayed an increased reliance on larger more mobile prey organisms such as fish and decapodB. Bassozetus normalis and B,. taenia attained a much greater size at maturity than most species examined in this study.

Dicrolene introniera. Barathrites oarri. X. mversi and P. macdonaldi 160

displayed the third main mode of feeding. These species employed a relatively more active, near bottom foraging and benthic-benthopelagic

feeding strategy.

A closer examination of the diets and morphology among select members of species groups clustered by diet reveals some interesting ecological relationships. Poroeadus silus and P. catena. two morphologically similar species, best exemplify the hovering, passive off-bottom foraging and facultative benthopelagic feeding strategy.

These species fed predominately on small calanoid copepodB. They possessed a small attenuate body at maturity, large head with a large

terminal mouth, large eyes, numerous and elongate gill rakers, branchial sieve with high filtering capacity, and weakly developed body musculature, all features which may facilitate euryphagous planktivorous

feeding habits. The predominance of numerous small calanoid copepods in

the diet suggest that these fishes may hover passively not far off the bottom foraging on swarms of copepods and other small planktivorous benthopelagic crustaceans.

Bathvonus nectoralis also exhibited the off-bottom foraging and

facultative benthopelagic feeding strategies similar to that described for P. silus and P.. catena. Abundant on the lower slope and rise, B.. pectoralis also displayed the greatest overall depth distribution of any species, occurring between 1200-5300 m. This eurybathic species was

characterized by a large terminal mouth, well developed eyes, high number of long gill rakers, branchial basket with high filtering

capacity and slightly procurrent pectoral fin rays, this species fed

primarily on small-sized calanoid copepods and other small benthic and benthopelagic crustaceans, e.g. isopods, mysids, amphipods. 161

In contrast B_, manatinus. which is similar in size and shape to B.. pectoralis. displayed a much different feeding strategy* Barathrodemua manatinus was more restricted in depth distribution than B., pectoralis and occurred only between 1200-2300 m depth. This species lacked the wide terminal mouth typical of many ophidiidB and was characterized instead by a small inferior mouth with reduced jaw protrusibility, long snout} small eyes, fewer gill rakers, and reduced filtering capacity of the branchial sieve, all features which may represent adaptations that facilitate feeding on benthic organisms. B.. manat inus was unique among its fellow ophidiids in that it specialized to a great extent on benthic tanaids as its major food source. The presence of Bediment} fish remains and more benthic crustaceans in the digestive tract of B.* manatinus suggest that this species may have fed nearer the bottom than

B.. pectoralis .

In B. manatinus. the small and inferior mouth with reduced jaw protrusibility is in marked contrast to the more protrusible terminal wider mouth of B.. pectoralis. Upper jaw protrusibility may be related to the ability of a fish to accommodate different prey sizes and to enable prey capture at a wide selection of mouth opening angles

(Alexander* 1967, Gosline, 1973, Motta, 1984). Protrusible jaws minimize a prey's ability to escape and increases suction efficiency by decreasing the dimensions of the orifice through which water enters.

Several other studies have suggested that the index of protrusion is greatest in fishes that prefer smaller prey since the strength of bite is inversely related to the degree to which the premaxilla is protruded and that a stronger bite is required to capture a large prey item (Al- hussaini, 1949; Aleev, 1969; Gosline, 1973), McLellan (1971, 1977) 162 found a significant relationship between a highly protrusible mouth and benthic feeding in several species of deep-sea macrourids. In this study a similar correlation between increased jaw protrusibility and benthic feeding was not evident. In fact the opposite condition appeared to be true. Gatz (1979) also did not support the above assumption based on correlation results from his study of stream fishes.

Mouth position w s b probably much more important in relation to feeding in deep-sea opbidiids than the degree of upper jaw protrusibility. Mouth position (placement of the mouth on the head), and mouth orientation (angle of the open mouth's gape in relation to the horizontal body axis) in B. manatinus was considered to be related to the location of food items relative to the position of the fish.

Several other studies have reported similar findings relating mouth position and orientation in fishes to location of food items (Aleev,

1969; Al-Hussani, 1949; Gatz, 1979; Mclellan, 1976; Schmitz and Baker,

1969; Schultz and Northcote, 1972). Chao and Musick (1977) demonstrated that above bottom feeding sciaenida possessed oblique, upturned mouths, whereas infaunal/epifaunal feeders possessed subterminal mouths.

Alexander (1967) suggested that the inferior mouths of benthic feeding fishes allowed prey to be captured without shifting the fish's body out of the horizontal plane. McLellan (1971, 1977) presented a similar argument for deep-sea macrourids possessing inferior or subterminal mouthB. Gatz (1979) determined that both mouth position and orientation bad a significant relationship with reBpect to food. DeGroot (1971) indicated that the asymmetrical mouths of flounder allow the protrusible jaws to be directed downward during feeding. Dicrolene intronigra and D. kanazawai were two morphologically similar species that served nicely to illustrate two contrasting feeding strategies among species examined. These two species also provided the strongest evidence for an example of trophic resource partitioning among all species examined. D. introniera is important in both the Mid-

Atlantic Bight and tropical Bahamas regions whereas D, kanazawai is present in only the Bahamas region. Both species occur sympatrically in the the Bahamas area with equivalent frequency and abundance (Sulak,

1982). Such a pattern between morphologically similar species suggests an absence of competition. Data from food habits analysis clearly demonstrated that these species specialized on different prey items.

Isopod crustaceans were the most important food item for D. intronigra whereas calanoid copepods and polychaetes dominated the diet of D. kanazawai.. Sediment and mixed infaunal prey items, e.g. pelecypods, gastropods occurred in the diet of D. introniera but were noticeably absent in the diet of D. kanazawai. D.. intronigra selected relatively larger prey items over a greater prey size range than did D.. kanazawai.

Further differences between the two species were noted in the parasite fauna of the digestive tract. D. intronigra was heavily infested with digene trematodes and nematode parasites whereas D. kanazawai had only a light infestation of digene trematodes. Other studies reported the diet of D. intronigra to include mixed infaunal prey (Rayburn, 1975, Wenner,

1978, 1984) with apparent specialization on ampbipods (Campbell et al..

1980). These results provide good inferential evidence that D_. intronigra preferred feeding on larger prey organisms occurring near or on the bottom whereas D. kanazawai specializes in smaller prey organisms found further up in the water column. 164

Trophic resource partitioning appears in part to be a mechanism enabling sympatry between these species of similar size, morphology and depth distribution. Although the concept of resource partitioning as an alternative outcome of competition (Schoener, 1974) has recently been challenged (Levin, 1983a, b), it offers a plausible and satisfactory explanation for the observed pattern and relationship between these two species.

The feeding morphology of D. introniera and I), kanazawai were similar although they fed on different prey organisms. Both species possessed similar dentition patterns consisting of small villiform teeth in the jaws and oral branchial cavities. The upper jaw mechanism including the articulating processes of the maxilla and premaxilla were nearly identical in both species. Other components of the feeding apparatus such as the hyoid, suspensorium and neurocranuim were also similar. Both species possessed a digestive tract and svimbladder of similar shape and configuration. Dicrolene kanazawai differed from D.. intronigra in that it possessed a relatively larger eye, a strongly curved and elongate opercular spine, lacked patches of teeth on the paired bypobranchials and possessed a small tooth patch on the third epibranchial. The pectoral fin of D. kanazawai was directed at a more downward angle and was placed lover on the body relative to the center of gravity than in D. intronigra. Dicrolene kanazawai also possessed slightly fewer gill rakers on the anterior arch. These rakers are shorter, more widely spaced and exhibit less filtering capacity than those of D. intronigra.

Gill rakers, which form a "sieve" to retain food particles, were considered related to the size and type of prey (e.g. benthic vs. 165

epipelagic) eaten. Many studies suggest that the gill raker number is

inversely correlated with presence of larger and more benthic prey in

the diet (Kliewer, 1970; Himberg, 1970; Nilsson, 1958). Kliewer (1970)

found a positive correlation between gill raker number and proportion of

benthic prey consumed by Coreconus cluneaformis. Chao and Musick (1977)

reported that an increased number of rakers was an indicator of filter

feeding in sciaenid fishes. Data from food habits of D. intronigra and

D.* kanazawai disagreed with these general assumptions and demonstrated a

positive correlation between prey size and number of gill rakers.

Similar findings were reported by Gatz (1979) in his study of stream

fishes. Be suggested that a strong taxonomic component negates the

utility of this character in certain species.

In this study species interaker distance was considered to be

related to the size of prey eaten. Raker spacing to some extent

determines the ability of the branchial sieve to retain food particles

of a given size range. Clarke (1980) indicated that gill raker gaps

observed among species of deep-water myctophids were positively related

to the size of food items eaten. Magnuson and Beitz (1971) also showed

that the proportion of smaller food items in the diets of pelagic

carnivores decreased as gill raker gap distance increased. Food habit data from species of Dicrolene also disagreed with these assumptions and demonstrated an inverse correlation between prey size and interaker distance.

Gill raker length in D. intronigra and D . kanazawai was found to be directly proportional to prey size. This relationship also stands in contrast to several studies that consider gill raker length to be

inversely proportional to prey size. Fishes e.g. menhaden, adaptive for feeding predominately on very small prey items such as phytoplankton and copepods, possess finely meshed branchial baskets with relatively long rakers. Conversely fishes, e.g. trout feeding on larger food items, possess coarser mesh branchial baskets with relatively shorter gill rakers. Gatz (1979) and Keast (1980) found similar relationships between gill raker length and prey size. Among sciaenids, Chao and

Uusick (1977), reported numerous long gill rakers in mid-water filter- feeders to shorter, less numerous gill rakers in infaunal/epifaunal predators.

D. intronigra and D. kanazawai possessed relatively long pectoral fins with free elongated lower rays. Relative pectoral fin area and aspect ratio were also high for these two species. Pectoral fin length in fishes has been considered to increase as a function of the degree of low speed maneuverability (Grey, 1868; Kanep, 1971; Starck and

Schroeder, 1970). Whereas the relative pectoral fin shape, area and aspect ratio were considered to be directly proportional to the capacity of the fins to function in braking, fanning to maintain position and acceleration from a resting position (Gosline, 1971). Such functional explanations for fish fin morphology were based primarily on studies of shallow-water fishes that frequently encounter currents. However, this interpretation of fin morphology may be inappropriate in the deep-sea habitat where low water velocity is generally the rule. Factors other than strong current, such as the development of pectoral and pelvic fins as gustatory and tactile sensory appendages, are probably more important to fin design in species of Dicrolene and other ophidiid fishes inhabiting the deep-sea. Baasozetus normal is and B.. taenia were two morphologically similar

species that displayed the second type of feeding mode. These species fed oportunistically in the prey environment by utilizing both benthic

and benthopelagic food organisms. Smaller individuals of B.. normalis fed on smaller more passive benthopelagic prey such as copepods, amphipods, and mysids whereas larger individuals showed an increased reliance on more active decapods and fish. Increased body size probably enabled these fishes to cover a greater area in search of food thereby increasing the chance of encountering more active larger prey. In fact these species selected the largest prey items over the greatest prey

size range of any species examined. Decapods, amphipods and mysids were

the major food items eaten.

Houth size in B. normalis and B.. taenia fishes was considered to relate to the maximum physical size of potential intact prey items

(Aleev, 1969; Starck and Schroeder, 1970; Werner, 1974; Northcote,

1974; Thomas, 1962). Stoner (1980) reported that mouth size was

linearly related to standard length of pinfish, Laeodon rhomboides.

Larger pinfish possessing larger relative mouths were able to utilize a broader range of prey than smaller pinfish. Gatz (1979) and Keast and

Webb (1966) found positive correlations between mouth size and the size of prey items eaten in stream temperate and lake fishes respectively.

Childress and Nygaard (1973) suggested that mouths of predatory fishes in the upper bathypelagic zone tend to converge to a size large enough for handling any of an assemblage of energy-rich vertically-migrating mesopelagic fishes. The importance of mouth size in bathypelagic fishes has been further demonstrated by Ebeling and Caillet (1974) who suggest 168

that bathypelagic fishes should be equipped to eat auy large or small

food item that they should chance to encounter*

The occurrence of sediment, scavenged fish remains and more benthic

prey organisms in the diet of B.. normalis and its absence in the diet of

B. taenia, suggest the former may feed more frequently near the bottom.

Unfortunately too few specimens of B. taenia were available for study in

order to draw any definate conclusion. However it is interesting to

note that B. taenia differed from B.. normalis in that it possessed a weaker less dense body musculature. Increased fluid content in the body

tissues of deep-sea fish have been considered energy conserving morphological adaptations associated with a more pelagic exsistence

(Horn et al., 1978; Nielsen, 1969).

Acanthonus armatus, a highly unusual fish, employed a passive,

facultative benthic-benthopelagic feeding strategy. This species fed on

prey organisms above near and on the bottom. Acanthonus armatus fed

primarily on tube-dwelling polychaetes and other small benthic and benthopelagic crustaceans. Nielsen (1966) reported finding polychaete

filt, copepods, ostracods and amphipods in the stomachs of five

specimens he examined. According to Horn et al. (1978) this species has

been observed to swim slowly just above the deep-sea floor in a horizontal to slightly head-up attitude. Acanthonus armatus probably

feeds as it drifts passively with gentle bottom currents.

In addition A. armatus displayed the most extreme energy conserving morphological and behavioral adaptations among deep-sea ophidiids examined. A. armatus lacked a swimbladder, possessed reduced tissues and components (muscle, bone, brain, gills) and exhibited an enlarged 169 cranial cavity filled with dilute fluids, all of which are energy conserving features to attain neutral buoyancy (Horn et al.. 1978).

Aside from incidental notes in the taxonomic literature (Cohen,

1961; Cohen and Nielsen, 1978) very little is known concerning the feeding habits and basic life history of X. mversi. This species, which possessed a very large head, extremely wide gape and relatively short body, exhibited a more active near bottom feeding strategy similar to that described for D. intronigra. The diet of X. mversi consisted of isopods and small benthic and benthopelagic prey organisms; however, it should be noted that only small individuals were available for examination. Since ontogentic dietary shifts are common as a fish matures (Ross, 1978) I would expect larger individuals of this species to feed on larger more mobile active prey (e.g. fish, large decapods, etc.)

Among all deep-sea ophidiids examined A. armatus and X. mversi possessed the most distinct combination of morphological characters.

Acanthonus armatus is most closely related to X. mversi with which it shares some similarities (Cohen, 1961; Cohen and Nielsen, 1978). Both species possessed relatively large thick heads, small eyes set close to th surface and similar dentition patterns consisting of small villiform teeth in jaws and oral branchial cavities. The upper jaw mechanism including the articulating process of the maxilla and premaxilla and palatine are very similar in both species. Acanthonus armatus and X. mversi also possessed a very short lower jaw relative to other ophidiids examined. Both species possessed well developed spines at the lower angle of the preopercle. The symplectic bone of the suspensorium is somewhat forked posteriorly in both species in contrast to the typical 170 narrow cone shaped symplectic bone observed in all other ophidiids.

Acanthonus armatus differed from X. mversi in that it possessed a prominent protruding bifed frontal spine on the snout, a long tapering body, and thick transparent skin loosely attached to the body.

Acanthonus armatus also possessed a greater number of long closely- spaced gill rakers on the anterior arch with higher filtering capacity than did X. mversi.

Penopus macdonaldi is unusual among ophidiids in that it possessed a spatulate sensory snout and small inferior mouth (figs. Id, 26d, 35a).

Although the occurrence of isopods and sediment in the diet of of P. macdonaldi suggests bottom or near bottom feeding, there is little data to accurately characterize its feeding strategy. However, with respect to morphological and feeding adaptations characteristic of fishes feeding on small prey in oligotrophic regions, the halosaurs of the genus Aldrovandia and Halosauropsis may present an instructive ecological parallel to morphologically similar P.. macdonaldi. Halosaurs displayed great behavioral and dietary plasticity ranging from active benthic foraging in the eutrophic waters of the MAB to passive hovering just off the bottom in the oligotrophic waters of the Bahamas region

(Cohen and Pawson, 1977; Grassle et al.. 1975; Sedberry and Musick,

1978; Sulak, 1977, 1982). The diet of P. macdonaldi has not been sufficiently investigated in the Bahamas region. When it is, it will be interesting to discover if this species like its halosaur cognate has adopted feeding strategies appropriate for oligotrophic energy poor regions.

There is also little data available to accurately characterize the feeding stratey of P.. miles. The occurrence of amphipods, decapods and 171

mysids in the stomachs of the few specimens examined suggest a feeding strategy similar to that reported for species of Bassozetus. Large individuals of P.. miles were capable of capturing large decapods.

Unfortunately not enough specimens were available to determine if ontogenetic feeding shifts occur in diet. Given its' peculiar morphology it's unlikely that small individuals of P. miles fed on large mobile prey such as decapods and fish. It is more probable that young individuals fed on small benthic-benthopelagic organisms. Poroeadus miles is similar in morphology to P. macdonaldi in that it possessed an elongate anteriorly depressed head, subterminal mouth and long slender whip-like body.

Barathrites narri displayed the most specialized and restricted diet of all species examined. This morphologically distinct fish fed almost exclusively on holothurians and polychaetes. In addition B. narri was distinct among species examined in this study in that it possessed a very short head, small eyes, short and highly placed pectoral fins, reduced gill rakers and an enlarged row of teeth in the outer jaws. In addition to these characters B.. narri possessed very firm and well developed head and body musculature in striking contrast to the flaccid poorly developed musculature of many other ophidiid species. Such characteristic features of these fishes may be part of a distinctive feeding complex specialized to feed on rather sedentary benthic holothurians and polychaetes. The presence of only fragments and shreds of food items in the stomach-suggest these fishes cut off small pieces of food rather than ingesting whole intact prey organisms.

Also the high placement on the body of the pectoral fins and their slightly oblique orientation might confer a better attitude for 172 approaching prey. Such a feeding strategy vould be energy conserving and represent an adaptation appropriate for the oligotrophic waters off

the Bahamas. A similar distinctive feeding complex is reported for the closely related species Enchelvbrotula gorooni (Cohen. 1982), He suggests E,. eomoni nips at coelenterate tentacles or at other fishes.

Unfortunately the several stomachs examined in his study were empty.

Relationship between diet and morphology

Once species feeding guilds were established, the fundemental assumption that diet as handled by the numerical analysis was directly related to morphology remained untested. Cluster analysis of morphological characters was employed as a verification procedure to certify that the feeding guilds did indeed represent morphological guilds. Such a comparative approach was based on the assumption that most characters of an integrated organism are of adaptive value even

though their functional significance may not be known (Hecht, 1965).

Each character chosen was based on some inferred functional and/or

ecological significance. In most cases there was strong evidence in support of the hypothesized adaptations reported in the literature although experimental substantiation was rare (Aleev, 1969).

Comparison of morphologically structured guilds with feeding guilds demonstrated a poor correspondence between morphologically structured guilds based on characters assumed related to feeding and feeding guilds based on stomach content analysis. This was not suprising since cluster analysis did not discriminate against characters superflous or confounding to the morphological analysis. The weaknesses and deficiencies of such a purely phenetic approach, as discussed at great lengths in the taxonomic literature (Hayr, 1969); also see recent volumes of systematic zoology) are inherent to ecological applications as well. Additional statistical investigation of the morphological data, such as correlation and discriminate analysis may prove helpful.

It was not uncommon in studies on relationships of ecology with morphological functions to be unable to relate form and function. For example, results from Suyehiro's (1942) study of the digestive system and feeding habits of 150 fish species were not as "interesting" as he had expected them to be but still provided important ecological information. In Davis and Birdsong's (1973) valuable review of the morphology and ecology of coral reef fishes they state that "confusion in attempts to correlate ecology and morphology can arise from several factors." Several of these factors, listed below, were also pertinent to studies concerning deep-sea fishes. (1) Sufficient information concerning the ecology of the species throughout its life history or even over short periods of time was unavailable. (2) The species we observe today was the successful end product of different selection pressures operating at different rates and with varying intensity throughout evolutionary time. Consequently the structures we study could have served more than one function making interpretation quite difficult (3) Host fishes will feed opportunistically (Margalef, 1974) making it very difficult to generalize predatory habits based either on classical comparative anatomy or incidental observations (e.g. submersible dive observations). In addition commonly employed numerical techniques may mask the true relationship between morphology and ecology.

Although morphological characters used in this study were of assumed adaptive value they probably did not adequately describe the feeding morphology of these fishes. The most serious flaw in this technique was the equal weighting of all characters. Such procedures were flawed because they were based on the assumption that during evolution the genotype as a whole changes harmoniously and that all components of it change at approximately equal rates (Mayr, 1969).

Recent studies of patterns of evolution in the feeding mechanism of fishes have provided evidence to the contrary (Lauder, 1981, 1982;

Schaeffer and Rosen, 1961). In fact Liem (1973) argued quite convincingly that the phenomenal colonizing and diversifying success of the percoid fish family Cichlidae was the result of just a minor alteration of the genotype. In studies of this nature, a purely quantitative procedure , by which a few useful characters are diluted by a large number of useless one, can only lead to confusing and misleading results (Stafleu, 1965).

The deep-sea ophidiids examined comprised a relatively uniform group with reBpect to feeding morphology when compared to the diversity of feeding morphologies displayed by coral reef fishes (Hobson, 1973) and tropical freshwater fishes (Lowe-McConnell, 1975). In other groups of fishes characters such as marked differences in dentition or premaxilla configuration, have been used successfully to correlate diet and morphology. However, among the ophidiids examined, little differentiation existed in these traditionally useful characters. There was probably a strong taxonomic component to head morphology in these deep-sea fishes that imparts a considerable amount of evolutionary constraint. This seeming lack of morphological specializations in deep- sea ophidiids is probably more a result of their phylogeny with respect to structural change (see Lauder, 1982) than a reflection of their 175 degree of trophic specialization. Although morphologically less diverse these fishes are just as successful at colonization and resource exploitation in the deep-sea as are structurally more complex fishes inhabiting coral reefs.

In many instances when single characters were examined and interpreted based on information in the literature, good correlation was observed between character and function. For example standard length was considered to he an indicator of prey size (Gatz, 1979; Lindstrom,

1955; Nilsson, 1955, 1958; Thomas, 1962, Swynnerton and Worthington,

1940). In ophidiid species in which sufficient material was available for study, prey size increased as a function of body size. Ontogenetic dietary shifts commonly occur in fishes. Juvenile fishes often pass through several planktivorous stages before adopting omnivorous, herbivorous, or carnivorous feeding habits as adults (Hall, 1970;

Sheridan, 1979; Stoner, 1980). Ross (1978) has shown that in searobins

(family Triglidae)) morphological changes in dentition and mouth size also accompany dietary changes during growth. Relative head length among ophidiids, like body length, was considered to be related to prey size. For example species such as B. normalis that possessed a relatively large head, were able to capture larger prey than those species that possessed smaller heads. Similarly Gatz (1979) demonstrated a positive correlation between prey size and head length in the large headed pirate perch, Anhredoderus savanua. Schoener (1968) has demonstrated a similar trend for certain lizards.

A positive correlation between an elongate depressed snout, inferior mouth and benthic feeding was observed among P. macdonaldi. P.. miles. P.. catena and P.. silus. Among these four species, P.. silus and 176

P.. catena possessed greatly compressed heads, blunt snouts, and terminal mouths with wide gapes. Poroeadus silus and P.. catena fed predominately on small planktivorous benthopelagic crustaceans. In contrast, P. miles and P. macdonaldi respectively, possessed increasingly elongate, depressed snouts with subterminal to inferior mouths. These species fed on larger and more benthic prey organisms respectively than did P.. silus or P.. catena. Increased relative snout length has been considered related to increased benthic feeding in fisheB (Chao and Husick, 1977).

They suggested that this character is related to depth of penetration into the substrate achieved during feeding. Menticirrhus saxatilis and

Micropoeoniaa undulatus possessed relatively long snouts and fed more on epifaunal than infaunal prey, whereas spot possessed a short snout and fed predominantly on infaunal prey. Conversely McLellan (1971, 1977) suggested that macrourids with long rostrums probably use them in probe­ like fasion while feeding on benthic prey.

Nearly all deep-sea ophidiids examined in this study possessed pads of numerous villiform teeth along the jaws, and inner surfaces of the mouth. Among the ophidiids examined very little variation existed in the size, shape, number and arrangement of dentition in both the oral cavity and pharyngeal basket. Such dentition is well suited for feeding on small benthopelagic prey organisms. Gatz (1979) and Gosline (1973) both reported that the shape of the dentary, premaxillary and maxillary teeth were significantly correlated with the importance of suction in feeding. Gatz (1979) also demonstrated a positive correlation between shape of jaw teeth and size of prey eaten. Chao and Musick (1971) reported that sciaenid species feeding on infaunal and epifaunal prey had very Bmall jaw teeth in bands with only the outer rows somewhat 177 enlarged. Conversely mid-water sciaenids possessed bands of fairly large, sharp jaw teeth often accompanied by enlarged canines. Sciaenids feeding just above the bottom possessed jaw teeth of intermediate form.

Keast and Webb (1966) reported similar findings among lake fishes.

In contrast to the characters above that appeared to show a relationship to feeding several other characters demonstrated no correlation with an assumed function. For example the number of pyloric caeca was considered to be correlated with the protein richness of the diet, as these structures function as an enzyme source and an area of absorption for protein and nitrogen (Beamish, 1972; Gatz, 1979;

Phillips, 1969). Six species in this study, M. aeassizii. JD. intronigra. D. kanazawai. P.. miles. P. catena and B . pectoralis possessed pyloric caeca. However, there was nothing in the food habit data to suggest these species consumed more protein in their diets than other species that lacked pyloric caeca.

Among ophidiids there was also little variation in pharyngeal teeth configuration. All species possessed pads of small villiform teeth with occasional clusters of somewhat larger teeth on the medial surface of the epibranchials. There were minor differences in the arrangement of these pads among species examined. For example, B,. pectoralis possessed a tooth patch on the second upper surface of the second epibranchial whereas P. macdonaldi lacked such a patch. There was considerable variation among species examined in the number and location of teeth on the basibranchial series. These differences in tooth pad arrangement were considered to be characters related to species taxonomic affinities and not the results of feeding specializations. There was very poor correspondence between tooth pad arrangement and type of prey eaten. 178

Gatz (1979) reported no correlation between the shape of pharyngeal teeth and the dominant prey item eaten. However Keast (1978) related a wide diversity of pharyngeal forms to prey types in centrarchid fishes.

He concluded that generalized feeders have generalized dentition whereas specialized feeders have divergent forms of dentition. A similar pattern waB observed for young sciaenids (Chao and Musick, 1977). Mid­ water feeders, largely piscivores, had reduced small, sharp pharyngeal

teeth. Near bottom feeders had somewhat stronger blunter teeth whereas

infaunal/epifaunal feeders possessed pharyngeal pads with conical teeth of mixed size and strength.

In many studies relative body depth has been considered to be inversely related to habitat water velocity and directly related to the capacity for making vertical turns on the axis of pitch (Aleev, 1969;

Gatz, 1979; Nikolskii, 1933). Gatz (1979) provided good evidence for

this interpretation hy demonstrating intraspecific variations in relative body depth in centrarchids inhabiting pools with different water velocities. Gromov (1973) reported similar findings when he compared lake and river populations of carp. In deep-sea ophidiid fishes, relative body depth may be related to factors other than those described above. Deep-sea fishes rarely encountered the strong currents common to freshwater stream fishes. Therefore in deep-sea fishes relative body depth as well as fin design may be related to functional designs that conserve energy and increase feeding and reproductive potentials.

Relationship between feeding strategies and regional primary productivity: 179

In contrast to the active foraging and benthic feeding strategies

typical of many taxa of fishes that dominate the eutrophic waters of the

Mid-Atlantic Bight (Middleton, 1979, Musick, 1976; Sedberry and Musick,

1978), the ophidiids, prevalent beneath oligotrophic tropical seas,

employed a more passive off-bottom foraging and benthopelagic feeding

strategy. The hypothesis iB advanced that the feeding strategies

employed by these tropical dwelling fishes constitute trophic

specializations appropriate for existence in oligotrophic waters.

Detailed studies of bathyal (2000-3000 m) and abyssal (3000-5000 m)

fish assemblages off the Mid-Atlantic Bight (MAB) of the U.S.A. (Musick,

1976, 1979; Musick and Sulak, 1979; Musick et al.. 1975) compared with analyses of data collected in the Tongue-of-the-Ocean, Bahamas, and adjacent SargaBso Sea (Sulak et al.r 1979; Sulak, 1982) suggests that

feeding strategies of fishes are quite different between the two areas.

Large mobile euryphagous macrourid and morid fishes, prevalent beneath eutrophic surface waters in temperate latitudes exhibit active foraging modes whereas in contrast small planktivorous and euryphagous ophidiids,

common to the oligotrophic deep-waterB off the Bahamas, employed energy

conserving passive feeding strategies.

Sulak (1982) forwards the hypothesis that the fundemental dichotomy

in taxonomic composition of the demersal deep-sea fish fauna in the

Atlantic results from the life history requirements (feeding and reproduction) of major component taxa in relation to regional primary productivity. He argues that Gadiformes, having undergone evolution in high latitudes where eutrophic conditions prevail, display energetically expensive life histories such as extensive ontogenetic migrations and off bottom foraging excursions (Haedrich and Henderson, 1974; Pearcy and 180

Ambler, 1974; ffenner and Musick, 1977). For example aggregated feeding

(Isaacs and Swbartzlose, 1975) and active foraging on benthic epifauna and infauna prevailed as the predominate feeding strategy in the

Macrouridae and Moridae of higher latitudes (Geistdoerfer, 1978;

Macpherson, 1979). In contrast to the macrourids, morids and gadiids, the tropical dwelling ophidiids, chlorophtalmids, aphyonids and alepocephalids displayed sedentary demersal and passive benthopelagic feeding modes (Sulak, 1982).

Since feeding strategies employed by deep-Bea ophidiid fishes may have evolved in part in response to the kinds and abundances of food available it is important to understand the mechanism for food transport to the Bahamian region. Food transport in the deep-sea may be of continental or oceanic origin (Sedberry and Musick, 1978). Organic matter of continental origin may enter the deep-sea through migration of organisms from continental to oceanic water masses, floating terrestrial and neritic plant material, and turbidity currents. Organic matter of oceanic origin may reach the deep-sea through settling of dead plankton,

(Menzies, 1962, or larger organisms (issacs, 1969; Dayton and Hessler,

1972; Isaacs and Schwartzlose, 1975) and through overlapping, vertically migrating food chains (Vinogradov, 1953 as cited by Vinogrodova, 1962;

Sedberry and Musick, 1978). All of these pathways are ultimately dependent on oceanic primary productivity. Available evidence suggests that photosynthetic oceanic phytoplankton production is the most important basic source of organic matter for deep-sea ecosystems (Rowe et al., 1974; Sokolova, 1972; Grice and Hart, 1972; Jahn and Backus,

1976). Although considerable quantities of organic matter from sinking floating plant material (Sareassum spp.) and its associated fauna 181

reaches the bottom of these Bahamian basins, recent studies suggest most

of this matter is refractory in nature and of little use to fishes.

Primary productivity in the surface waters of the Bahamas region is

quite different from that reported for the MAB. Although precise

primary production values are unavailable for the Bahamas study region

per se Kobletz-Mishke et al. (1970) indicated a general range in the

Bahamas-Antilles region of ca. 35-90g carbon/m /yr. In contrast average

primary production estimates for the Mid-Atlantic Bight (HAB) range from 2 100-200 g carbon/m /yr for waters overlying the shelf and slope to 50-70

g carbon/m /yr for waters overlying the rise and abyss (Riley, 1957;

Ryther and Tentsch, 1958; Menzel and Ryther, 1960; Rowe, 1971a;

Koblentz-Mishke et al.. 1970; Volkovinsky and Kabanova, 1970; Smith,

1978). In addition sediments in the Bahamas originate primarily from

particle deposition of foraminfera and pteropod tests resulting in

generally low organic carbon content ca. 0.5 to 1.0 percent (Emery and

Uchupi, 1972). Also the rate of pelagic sedimentation of particulate 2 organic carbon (5.7 mg/m /day) at 2000 m in the Bahamas (Weibe et al.. 2 1976) is half that reported (12.8 mg/m /day) at equivalent depths in the

MAB (Rowe and Gardner, 1979).

In contrast to high primary productivity in the eutrophic waters off

the MAB, diminished primary productivity in the oligotrophic surface waters of the Bahamas region should be reflected in the composition of benthic invertebrate megafauna and macrofauna. Sulak (1982) reviewed

the literature in this area and characterized the invertebrate megafauna

and macrofauna represented in the two regions. Megafaunal invertebrates

that employed an off-bottom suspension feeding habit (hexactennilid

sponges, pennatulids) were more prevalent in the oligotrophic Bahamian region, in contrast to the predominance of on-bottom deposit invertebrate

feeding (echinoids, ophiuriords) in the MAB region* Unfortunately, data

on the macrofauna of the Bahamas region were not available* However,

data from other low productivity regions of the tropical western North

Atlantic suggest both abundance and biomass of benthic macrofauna are

less by approximately an order of magnitude than in the MAB (Rowe, 1971,

Rowe et al., 1974). Xn additon the predominance of benthopelagic

feeding on micronekton among demersal ophidiid fishes and other fish

taxa of the region probably reflects the importance of benthopelagic megafaunal and macrofaunal organisms.

In conclusion the successful diversification of ophidiids in oligotrophic waters appears in part to lie in energy conserving morphological and behavioral adaptations of which trophic

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Abbreviations

cp cartilage pad Inin maxillcmaxillary ligament 1pm palatomaxillary ligament lpp prenaxillary palatine ligament vlpm ventral premaxillcmaxillary ligament lem etbmomaxillary ligament

MISCLES

Al, A2 A3, subdivisions of the adductor mandihulae AAP adductor arcus palatini AO adductor opercuii Ah intramandibularis DO dilatator opercuii HAB hyohyoidei abductores HAD hyohyoidei adductores HYP hypaxials ICARA infracarinalis anterior 1MD intezmandibularis LAP levator arcus palatini LO levator opercuii ML nuscles of pelvic fin PHHY protractor hyoidei STH sternobyoideus STB 8 te mobranchials BCHES

AN angular ART articular BB basibranchial EH basibyal ER branchiostegals BOC basioccipital CB ceratobranchial CH ceratobyal CL deitbrun CM centrum 00 coracoid COB cixcumzbital D dentary DE dermetbnoid uni dorsal bypohyal

199 appendix 1. continued

EB epibranchial ECT ectopterygoid ES epihyal EO epiotic ECK exoccipital ES extrascapular EL frontal GF gill filaments BB hypobranchial * hyomandibula INC intercalar JH interhyal IQP interopercle ISP ischial process IE lateral etlmoid IGR long gill raiser MES mesopterygoid MET metapterygoid MX maxilla HA nasals HS neural spine CJP opercle P pbaryngeals PA parasphenoid PAT. palatine PB pharyngeobranchials PC prootic PCL postcleithrua PG pterygiophore PL parietal HJRB pleural rib UK premaxilla POP preopercle PP parapophysis m pterospbenoid PS posttenporal p r pterotic POP pubic plate QJ quadrate BA radiala SA scapula sc spbenotic SCL supracleitbmn SL supraoccipital 9tX supramaxilla SOP subopercle SI symplectic VHH ventral hypohyal VO vomer OBO urobyal Appendix 2. Morphological Analysis Data Record

SPECIES STATION

CRUISE SPECIMEN #

SUPERFICIAL BCD? SHAPE CHARACTERS

1. Specimen Standard Length (ran) ______(1)

2. Completeness of Lateral Line

(1) Complete

(2) Inccrplete

(3) Lacking (2)

3. Position of Lateral T.inp

(1) Curving Dorsally

(2) Horizontal

(3) Owing Ventrally

(4) All Above (3)

4. Nurber of Later Lines (4)

5. Body Depth (ran) (5)

6 . Index of Trunk Shape (6)

BATRTO FIN CHARACTERS

7. Pectoral Fin length (ran) (7)

8 . Aspect Ratio of Pectoral Fin (8)

9. Pectoral Fin Shape

(1) Rounded

(2) Intermediate

(3) Pointed (9)

201 10. Pectoral Fin Rays Condition

(1) Hit ire

(2) Lower Half Free

(3) Completely Free (10)

11. Position of Pectoral Fin Relative to the Center of Gravity

(1) Fin Wholly Anterior to Plane

(2) Fin Originating Anterior to OG and Extending Posterior to Plane

(3) Fin Originating at Level of 0G Plane

(4) Fin Originating Posterior to Plane (11)

12. Position of Dorsal Fin to OG

(1) Fin Wholly Anterior to Plane

(2) Fin Originating Anterior to OG and Extending Posterior to Plane

(3) Fin Originating at Level of OG Plane

(4) Fin Originating Posterior to Plane (12)

13. Position of Anal Fin to OG

(1) Fin Wholly Anterior to Plane

(2) Fin Originating Anterior to OG and Extending Posterior to Plane

(3) Fin Originating at Level of OG Plane

(4) Fin Originating Posterior to Plane (13)

HEAD (RARACIERS

14. Mouth Size

a. Lateral Mouth Cape (ran) (14)

b. Medial Mouth Cape (ran) (15)

c. Upper Jaw Length (16)

15. Jaw Protrusibility (17) 203

16. Snout Length £18)

17. Mouth Orientation

(1) Dorsal

(2) Anterior (Terminal)

(3) Downward Oblique (Sub-terminal)

(4) Ventral (Inferior) (19)

18. Eye Diameter (20)

19. Position of Eyes

(1) lateral

(2) Dorsalateral

(3) Ventrolateral (21)

GILL RAKER CHARACTERS

20. Nmfcer of Anterior Rakers (Developed) (22)

21. Inter-Raker Gap Distance (mn)

ARD “ (d + ... d ) N (23)

22. Longest Raker Length (mn) (24)

23. Filter Area (mn )

(Raker Length)(Lower Linb Length) (25) mTHttttL BCD? CHARACTERS

24. Stomach Length (mn) (26)

25. Intestinal Length (mn) £27)

26. Total Gut Length (mn) (28)

27. Nuriber of Pyloric Caeca £29)

28. Svimbladder (1) Absent

(2) Present £30) 29. Dimming Muscles

(1) Absent

(2) Present

30. Swinb ladder Length (ran)

31. Svinb Ladder Width (nm)

32. Rete Mirabilia Length (mn)

DENTUICK CHARACTERS

33. Upper and Lower Jaw Dentition

(1) Absent

(2) Villifonn

(3) Conical

34. Upper and Lower Pharyngeal Dentition

(1) Absent

(2) Villifonn

(3) Coocial

35. Shape of Vomerine Tooth Patch

(1) Absent

(2) Staall Irregular Patch

(3) V-shaped Flush with Palate

(4) V-shaped Prominently Raised

36. Nmber of Basibranchial Tooth Patches (38)

37. Configuration of Ooteological Components of the Feeding Apparatus

(1) Poroeadus miles pattern

(2) Bassozetus noimalis Pattern

(3) Dicrolene int-mm'gra pattern

(4) Barathrites parri pattern

(5) Acanthoma armatus pattern Appendix 3. Station Data Listing (mod. from Sulak, 1982)

Dicrolene introniera

MEAN X SAMPLE LATITUDELONGITUDE DATE TEMP DEPTH CODE DD MM DD MM YYMMDD C M

1 CIT 42 23 31 N 76 44 W 73 223 4.4 1280 4 CIT 46 23 30 N 77 05 W 73 224 4.5 1257 8 CIT 58 24 14 N 77 18 W 73 227 4.2 1408 2 CIT122 24 06 N 77 19 W 73 924 4.2 1434 5 CIT123 24 12 N 77 18 W 73 924 4.2 1435 1 CIT145 24 14 N 77 19 W 74 203 4.2 1440 1 CIT147 23 56 N 77 14 W 74 203 4.2 1356 1 CIT149 23 52 N 77 18 W 74 204 4.2 1388 1 CIT156 23 43 N 76 50 W 74 205 4.3 1332 2 CIT163 23 32 N 77 10 V 74 206 4.3 1345 2 CIT168 24 26 N 77 23 V 74 207 4.0 1577 2 CIT250 23 52 N 76 52 V 741031 4.4 1291 1 CIT251 23 41 H 76 44 V 741031 4.3 1315 1 CIT252 23 39 N 76 45 w 741101 4.3 1327 1 CIT253 23 44 N 76 49 w 741101 4.3 1332 2 CIT254 23 34 N 76 43 V 741101 4.3 1303 1 CIT255 23 30 N 76 57 w 741101 4.3 1320 1 CIT267 24 24 N 77 22 w 741103 4.1 1542 1 CIT301 23 52 H 76 52 V 75 403 4.4 1286 1 CIT309 23 44 N 76 47 w 75 405 4.3 1316 2 CIT311 23 38 N 77 14 w 75 403 4.2 1356 2 CIT314 24 46 N 77 15 w 75 406 4.2 1367 1 CIX316 24 24 H 77 27 tf 75 407 4.0 1559 1 CIT317 24 16 N 77 20 w 75 407 4.1 1464 3 C1T322 23 40 N 77 04 w 75 408 4.2 1363 2 CIT326 23 51 N 77 16 w 75 410 4.2 1383 4 CIT350 24 02 N 77 24 tf 75 420 4.2 1430 1 CIT351 24 19 N 77 21 w 75 420 4.1 1519 5 CIT359 23 53 H 77 10 w 75 821 4.2 1358 1 CIT361 23 30 K 76 55 tf 75 821 4.3 1300 2 C1X362 23 39 N 77 07 V 75 821 4.2 1363 2 CIT363 23 51 N 77 55 tf 75 822 4.3 1320 1 CIT366 23 49 N 77 06 w 75 822 4.2 1366 7 CIT368 23 42 N 76 52 tf 75 822 4.3 1347 3 CIT370 23 34 N 76 42 tf 75 823 4.4 1282 1 CIT402 22 53 N 75 27 tf 75 904 3.5 2404 1 CIT405 24 28 N 77 20 w 76 228 4.1 1464 1 CIT410 23 30 N 76 51 w 76 229 4.3 1323 1 CIT416 23 40 N 77 16 tf 76 302 4.3 1338 1 CIT422 23 53 N 77 07 tf 76 302 4.3 1339 1 CIT431 24 16 N 77 22 tf 76 306 4.1 1478 1 CIT435 23 50 N 77 16 w 76 309 4.2 1380 2 CIC041 23 53 N 77 08 tf 80 915 4.2 1341 3 CIC063 29 06 N 77 08 w 80 921 5.7 1093

205 Dicrolene kanazawai

MEAN COUNT SAMPLE LATITUDE LONGITUDE DATE TEMP DEPTH A CODE DD MM DD MM YYMMDD C M

1 CIT 40 23 47 N 76 59 W 73 222 4.3 1312 2 CIT 41 23 43 N 76 49 W 73 222 4.4 1298 1 CIT 47 23 41 N 77 08 W 73 224 4.2 1372 2 CIT 48 23 44 N 77 13 W 73 224 4.2 1390 3 CIT 57 24 05 N 77 23 W 73 227 4.2 1404 6 CIT 59 24 14 N 77 25 W 73 227 4.2 1435 4 CIT 60 24 24 N 77 27 tf 73 228 4.0 1545 1 CIT 70 24 30 N 76 13 tf 73 303 3.9 1677 1 CIT 71 24 20 N 75 58 tf 73 304 3.9 1682 13 CIT 76 24 39 N 76 28 W 73 305 4.0 1586 1 CIT 78 24 25 N 76 10 tf 73 306 3.9 1692 5 CIT103 24 08 N 77 22 w 73 921 4.2 1436 3 CIT105 23 48 N 77 04 tf 73 921 4.2 1368 1 CIT113 23 41 N 77 06 w 73 923 4.2 1368 8 CIT114 23 41 N 76 50 tf 73 923 4.3 1338 2 CIT122 24 06 N 77 19 tf 73 924 4.2 1434 1 CIT123 24 12 N 77 18 tf 73 924 4.2 1435 2 CIT124 24 25 N 77 25 w 73 925 4.0 1593 7 CIT144 24 30 N 77 22 tf 74 203 4.1 1494 1 CIT145 24 14 N 77 19 V 74 203 4.2 1440 6 CIT147 23 56 N 77 14 tf 74 203 4.2 1356 3 CIT149 23 52 N 77 18 tf 74 204 4.2 1388 1 CIT150 23 48 N 77 03 tf 74 204 4.2 1358 2 CIT156 23 43 N 76 50 tf 74 205 4.3 1332 1 CIT163 23 32 N 77 10 tf 74 206 4.3 1345 2 CIT167 23 38 N 77 17 w 74 207 4.1 1515 14 CIT168 24 26 N 77 23 tf 74 207 4.0 1577 1 CIT175 24 32 N 76 18 w 74 210 3.9 1704 4 CIT176 24 39 N 76 29 tf 74 210 4.0 1632 2 CIT178 24 13 N 76 06 tf 74 210 3.8 1794 1 CIT184 23 55 N 75 26 tf 74 211 3.5 2188 1 CIT187 24 00 N 75 49 w 74 212 3.7 1887 1 CIT191 24 10 N 75 56 tf 74 213 3.8 1810 2 CIT192 24 21 N 75 59 tf 74 213 3.8 1768 2 CIT193 24 25 N 76 11 w 74 213 3.9 1748 2 CIT251 23 41 N 76 44 tf 741031 4.3 1315 1 CIT252 23 39 N 76 45 tf 741101 4.3 1327 4 CIT253 23 44 N 76 49 w 741101 4.3 1332 1 CIT261 23 36 N 77 08 tf 741102 4.2 1353 3 CIT267 24 24 N 77 22 w 741103 4.1 1542 1 CIT270 24 23 N 77 26 tf 741104 4.1 1550 2 CIT305 23 44 N 76 51 tf 75 404 4.3 1350 4 CIT314 24 46 N 77 15 tf 75 406 4.2 1367 7 CIT316 24 24 N 77 27 tf 75 407 4.0 1559 1 CIT317 24 16 N 77 20 tf 75 407 4.1 1464 4 CIT326 23 51 N 77 16 w 75 410 4.2 1383 1 CIT331 24 32 N 76 17 tf 75 412 3.9 1678 Dicrolene kanazawai continued

MEAN COUNT SAMPLE LATITUDE LONGITUDE DATE TEMP DEPTH SP A CODE DD MM DD MM YYMMDD C M

9 CIT332 24 39 N 76 27 W 75 412 4.0 1627 1 CIT333 24 24 N 76 08 W 75 412 3.8 1772 2 CIT334 24 15 N 76 06 W 75 413 3.8 1792 1 CIT336 23 48 N 75 46 W 75 413 3.8 1848 1 CIT339 24 10 N 75 54 W 75 414 3.8 1842 1 CIT340 24 20 N 76 00 W 75 415 3.9 1748 7 CIT341 24 39 N 76 26 W 75 415 4.0 1628 1 CIT343 23 48 N 75 47 W 75 416 3.7 1854 1 CIT350 24 02 N 77 24 W 75 420 4.2 1430 3 CIT354 24 34 N 77 31 w 75 819 3.9 1724 5 CIT356 24 23 N 77 26 w 75 820 3.9 1654 X CIT360 23 38 N 76 49 w 75 821 4.3 1338 1 CIT362 23 39 N 77 07 w 75 821 4.2 1363 1 CIT363 23 51 N 77 55 tf 75 822 4.3 1320 1 CIT365 23 50 N 77 14 tf 75 822 4.2 1372 6 CIT366 23 49 N 77 06 tf 75 822 4.2 1366 1 CIT378 24 21 N 76 04 tf 75 826 3.8 1761 1 CIT380 24 39 N 76 26 w 75 827 4.0 1640 3 CIT387 24 16 N 76 16 tf 75 829 3.8 1762 1 CIT392 22 48 N 75 38 tf 75 831 3.5 2250 4 CIT405 24 28 N 77 20 w 76 228 4.1 1464 4 CIT408 23 50 N 76 51 w 76 229 4.3 1346 1 CIT416 23 40 N 77 16 tf 76 302 4.3 1338 1 CIT417 23 52 N 77 16 w 76 302 4.2 1387 1 CIT418 23 50 N 77 17 tf 76 302 4.2 1386 4 CIT431 24 16 N 77 22 w 76 306 4.1 1478 3 CIT435 23 50 N 77 16 w 76 309 4.2 1380 1 CIC012 23 51 N 75 50 w 80 9 3 3.7 1856

Monomitonua aeaasizii

2 CIC063 29 06 N 77 08 w 80 921 5.7 1093

Holcomvcternous sauamoBus

1 CIT395 22 01 N 75 04 w 75 901 3.4 2686

Xyelacvba taverBi

3 CIT 85 24 23 N 77 23 tf 73 309 4.1 1528 2 CIT124 24 25 N 77 25 w 73 925 4.0 1593 1 CIT125 24 30 N 77 22 tf 73 925 4.1 1496 4 CIT168 24 26 N 77 23 tf 74 207 4.0 1577 1 CIT270 24 23 N 77 26 w 741104 4.1 1550 1 CIT305 23 44 N 76 51 tf 75 404 4.3 1350 Xvelacvba mversi continued

MEAN COUNT SAMPLE LATITUDELONGITUDE DATE TEMP DEPTH SP A CODE DD MM DDMMYYMMDD CM

1 CIT317 24 16 N 77 20 W 75 407 4.1 1464 1 CIT320 23 40 N 76 43 W 45 408 4.3 1322 1 CIT354 24 34 N 77 31 tf 75 819 3.9 1724 1 CIT405 24 28 N 77 20 W 76 228 4.1 1464 3 CIT409 23 37 N 76 43 W 76 229 4.3 1296 1 CIT428 24 49 N 77 40 tf 76 305 3.6 1977 1 CIC011 23 46 N 75 48 W 80 9 3 3.7 1790

Porotsadus miles

2 CIT353 25 16 N 77 39 W 75 421 3.3 3032 1 CIT414 23 40 N 76 59 W 76 301 4.2 1360 2 CIC017 22 55 N 75 25 tf 80 9 7 3.4 2423

PoroKadus silus

4 CIT 70 24 30 N 76 13 tf 73 303 3.9 1677 36 CIT 71 24 20 N 75 58 W 73 304 3.9 1682 17 CIT 72 24 00 N 75 44 tf 73 304 3.7 1884 19 CIT 73 23 46 N 75 42 W 73 304 3.8 1792 2 CIT 76 24 39 N 76 28 W 73 305 4.0 1586 6 CIT 78 24 25 N 76 10 W 73 306 3.9 1692 1 CIT175 24 32 N 76 18 W 74 210 3.9 1704 5 CIT176 24 39 N 76 29 tf 74 210 4.0 1632 48 CIT178 24 13 N 76 06 W 74 210 3.8 1794 4 CIT182 23 56 N 75 58 W 74 211 3.8 1769 3 CIT183 23 45 N 75 39 W 74 211 3.8 1807 85 CIT184 23 55 N 75 26 W 74 211 3.5 2188 11 CIT186 23 45 N 75 42 tf 74 212 3.8 1849 5 CIT187 24 00 N 75 49 W 74 212 3.7 1887 12 CIT191 24 10 N 75 56 tf 74 213 3.8 1810 4 CIT193 24 25 N 76 11 W 74 213 3.9 1748 3 CIT276 24 21 N 76 10 tf 741107 3.8 1772 7 CIT279 23 50 N 75 50 tf 741108 3.8 1838 5 CIT280 23 52 N 75 16 tf 741108 3.5 2354 2 CIT281 23 54 N 75 30 tf 741109 3.6 2141 4 CIT282 24 00 N 75 47 tf 741109 3.7 1905 7 CIT284 23 57 N 75 59 tf 741109 3.8 1776 3 CIT331 24 32 N 76 17 tf 75 412 3.9 1678 7 CIT332 24 39 N 76 27 tf 75 412 4.0 1627 8 CIT333 24 24 N 76 08 tf 75 412 3.8 1772 4 CIT334 24 15 N 76 06 tf 75 413 3.8 1792 45 CIT335 23 58 N 76 01 W 75 413 3.8 1766 13 CIT336 23 48 N 75 46 W 75 413 3.8 1848 2 CIT337 23 55 N 75 28 tf 75 414 3.6 2152 3 CIT338 23 59 N 75 48 tf 75 414 3.7 1890 4 CIT339 24 10 N 75 54 tf 75 414 3.8 1842 209

Poroeadus silus continued MEAN INT SAMPLELATITUDE LONGITUDE DATETEMP DEPTH A CODE DD MM DD MM YYMMDD C M

3 CIT340 24 20 N 76 00 W 75 415 3.9 1748' 13 CIT341 24 39 N 76 26 W 75 415 4.0 1628 4 CIT343 23 48 N 75 47 w 75 416 3.7 1854 2 CIT346 22 55 N 75 17 tf 417 3.5 2422 3 CIT376 23 55 N 75 59 tf 75 826 3.8 1762 2 CIT378 24 21 N 76 04 tf 75 826 3.8 1761 3 CIT380 24 39 N 76 26 V 75 827 4.0 1640 3 CIT382 23 57 N 76 01 tf 75 828 3.8 1763 1 CIT383 23 50 N 75 49 tf 75 828 3.8 1830 8 CIT384 24 09 N 75 55 tf 75 828 3.8 1844 2 CIT386 23 59 N 75 46 tf 75 829 3.8 1922 4 CIT387 24 16 N 76 16 tf 75 829 3.8 1762 11 CIT388 23 56 N 75 32 tf 75 829 3.6 2146 11 CIT390 23 50 N 75 14 w 75 830 3.5 2354 4 CIT391 22 54 N 75 14 tf 75 830 3.5 2427 1 CIT395 22 01 N 75 04 tf 75 901 3.4 2686 1 CIT399 22 29 N 75 11 w 75 902 3.4 2577 3 CIT400 21 59 N 75 06 H 75 903 3.4 2681 67 C1T403 23 57 N 75 27 tf 75 905 3.5 2166 2 CIT426 25 16 N 77 49 tf 76 304 3.4 2700 1 CIT428 24 49 N 77 40 tf 76 305 3.6 1977 25 CIC011 23 46 N 75 48 tf 80 9 3 3.7 1790 7 CIC012 23 51 N 75 50 tf 80 9 3 3.7 1856 2 CIC017 22 55 M 75 25 tf 80 9 7 3.4 2423 19 CID004 21 57 N 74 46 tf 811116 3.0 2728 8 CID005 22 03 N 74 48 tf 811116 * 2728 7 CID009 21 27 N 74 21 tf 811118 3.0 2714 1 CID010 21 24 N 74 05 w 811118 3.0 2703

Porogadus catena

1 CIT402 22 53 N 75 27 tf 75 904 3.5 2404 2 CIT426 25 16 N 77 49 W 76 304 3.4 2700 1 C16004 28 43 1 75 49 7 780212 • 4930

Barathrodemua manatinus

1 CIT 40 23 47 N 76 59 tf 73 222 4.3 1312 1 CIT 41 23 43 N 76 49 W 73 222 4.4 1298 1 CIT 46 23 30 N 77 05 V 73 224 4.5 1257 1 CIT 47 23 41 N 77 08 tf 73 224 4.2 1372 4 CIT 48 23 44 N 77 13 tf 73 224 4.2 1390 2 CIT 51 23 38 N 76 49 tf 73 225 4.4 1290 1 CIT 53 23 48 N 77 03 V 73 226 4.3 1325 3 CIT 57 24 05 N 77 23 w 73 227 4.2 1404 9 CIT 59 24 14 N 77 25 tf 73 227 4.2 1435 2 CIT 60 24 24 N 77 27 tf 73 228 4.0 1545 3 CIT 61 24 26 N 77 20 tf 73 228 4.0 1490 Barathrodemus manatinus continued

MEAN INT SAMPLE LATITUDE LONGITUDE DATE TEMP DEPTH A CODE DD MM DD MM YYMMDD C M

1 CIT 70 24 30 N 76 13 W 73 303 3 .9 1677 1 C1T 78 24 25 N 76 10 W 73 306 3 .9 1692 1 CIT 81 23 37 N 77 01 W 73 308 4.3 1326 6 CIT 85 24 23 N 77 23 W 73 309 4.1 1528 7 CIT101 24 27 N 77 28 W 73 920 4.0 1594 5 CIT103 24 08 N 77 22 w 73 921 4.2 1436 24 CIT104 23 56 N 77 14 w 73 921 4 .2 1350 1 CIT113 23 41 N 77 06 w 73 923 4.2 1368 2 CIT121 23 55 N 77 20 w 73 924 4.2 1394 9 CIT122 24 06 N 77 19 w 73 924 4.2 1434 3 CIT123 24 12 N 77 18 w 73 924 4.2 1435 2 CIT124 24 25 N 77 25 w 73 925 4 .0 1593 7 CIT125 24 30 N 77 22 w 73 925 4.1 1496 30 CIT126 24 41 N 77 34 w 73 925 3.7 1872 4 CIT144 24 30 N 77 22 w 74 203 4.1 1494 1 CIT145 24 14 N 77 19 w 74 203 4 .2 1440 2 CIT147 23 56 N 77 14 w 74 203 4.2 1356 8 CIT149 23 52 N 77 18 w 74 204 4.2 1388 1 CIT150 23 48 N 77 03 tf 74 204 4.2 1358 1 CIT163 23 32 N 77 10 tf 74 206 4.3 1345 15 CIT168 24 26 N 77 23 w 74 207 4.0 1577 55 CIT169 24 35 N 77 32 w 74 207 3 .9 1742 2 CIT176 24 39 N 76 29 V 74 210 4.0 1632 2 CIT177 24 23 N 76 09 V 74 210 3 .8 1770 1 CIT178 24 13 N 76 06 V 74 210 3 .8 1794 3 CIT182 23 56 H 75 58 w 74 211 3 .8 1769 1 CIT183 23 45 N 75 39 w 74 211 3 .8 1807 2 CIT186 23 45 N 75 42 w 74 212 3 .8 1849 2 CIT192 24 21 N 75 59 ff 74 213 3.8 1768 1 CIT193 24 25 N 76 11 tf 74 213 3 .9 1748 26 CIT247 24 39 N 77 32 w 741030 3.9 1724 1 CIT253 23 44 N 76 49 w 741101 4.3 1332 9 CIT256 23 37 N 77 06 w 741101 4.2 1360 2 CIT261 23 36 N 77 08 w 741102 4.2 1353 4 CIT267 24 24 N 77 22 w 741103 4.1 1542 2 CIT268 24 31 N 77 22 w 741104 4.1 1490 2 CIT276 24 21 N 76 10 w 741107 3 .8 1772 1 CIT284 23 57 N 75 59 w 741109 3 .8 1776 28 CIT299 24 39 N 77 33 w 75 402 3 .8 1778 2 CIT302 24 11 N 77 23 w 75 403 4.1 1458 4 CIT314 24 46 N 77 15 w 75 406 4.2 1367 7 CIT316 24 24 N 77 27 w 75 407 4 .0 1559 11 CIT317 24 16 N 77 20 V 75 407 4.1 1464 1 CIT326 23 51 H 77 16 w 75 410 4 .2 1383 4 CIT331 24 32 N 76 17 w 75 412 3 .9 1678 3 CIT332 24 39 N 76 27 w 75 412 4 .0 1627 3 CIT335 23 58 N 76 01 w 75 413 3 .8 1766 1 CIT343 23 48 N 75 47 w 75 416 3.7 1854 2X1

Barathrodemua manatinus continued

MEAN INI SAMPLE LAXIXODE LONGIXODE DAXE XEMP DEPXH A CODE DD MM DD MM YYMMDD C M

72 CIT349 24 36 N 77 33 W 75 419 3 .9 1748 2 CIT350 24 02 N 77 24 w 75 420 4 .2 1430 2 CIT351 24 19 N 77 21 w 75 420 4.1 1519 60 CIT354 24 34 N 77 31 w 75 819 3 .9 1724 3 CIT356 24 23 N 77 26 w 75 820 3 .9 1654 9 CIT357 23 55 N 77 17 w 75 820 4.2 1386 9 C1T359 23 53 N 77 10 w 75 821 4.2 1358 1 CIT360 23 38 N 76 49 w 75 821 4.3 1338 2 CIT361 23 30 N 76 55 w 75 821 4.3 1300 4 CIT362 23 39 N 77 07 w 75 821 4.2 1363 1 CIT363 23 51 N 77 55 w 75 822 4.3 1320 4 CIT365 23 50 N 77 14 w 75 822 4.2 1372 31 CIT366 23 49 N 77 06 w 75 822 4.2 1366 CIT376 23 55 N 75 59 w 75 826 3 .8 1762 1 CIT378 24 21 N 76 04 w 75 826 3.8 1761 1 CIT379 24 33 H 76 17 w 75 827 3 .9 1659 3 CIT382 23 57 N 76 01 w 75 828 3 .8 1763 1 CIT387 24 16 N 76 16 w 75 829 3 .8 1762 CIT392 22 48 N 75 38 w 75 831 3.5 2250 4 CIT405 24 28 N 77 20 w 76 228 4.1 1464 1 CIT407 23 53 N 77 05 w 76 229 4.3 1336 CIT408 23 50 N 76 51 w 76 229 4.3 1346 1 CIX409 23 37 N 76 43 w 76 229 4.3 1296 1 CIX410 23 30 N 76 51 W 76 229 4.3 1323 2 CIT413 23 51 N 77 13 w 76 301 4.2 1372 2 CXI414 23 40 H 76 59 w 76 301 4.2 1360 2 C1X418 23 50 N 77 17 w 76 302 4.2 1386 1 CXX422 23 53 N 77 07 w 76 302 4.3 1339 11 C1X426 25 16 N 77 49 w 76 304 3 .4 2700 1 C1X428 24 49 N 77 40 w 76 305 3.6 1977 20 CIX430 24 32 N 77 31 w 76 306 3.9 1661 2 CIX431 24 16 N 77 22 w 76 306 4.1 1478 1 CIX435 23 50 N 77 16 w 76 309 4.2 1380 2 C1X437 23 32 N 76 52 w 76 309 4.3 1333 7 CIC011 23 46 N 75 48 w 80 9 3 3.7 1790 1 CIC012 23 51 N 75 50 V 80 9 3 3.7 1856 1 CIC041 23 53 N 77 08 w 80 915 4.2 1341

Barathrites parri

1 CII 41 23 43 N 76 49 V 73 222 4 .4 1298 1 CIX 48 23 44 N 77 13 w 73 224 4.2 1390 1 CIX 50 23 42 N 76 43 w 73 225 4.5 1271 1 CII 51 23 38 N 76 49 V 73 225 4 .4 1290 1 CII 57 24 05 N 77 23 w 73 227 4.2 1404 3 CIX 58 24 14 N 77 18 w 73 227 4.2 1408 2 CII 59 24 14 N 77 25 w 73 227 4.2 1435 212

Barathrites parri continued

MEAN T SAMPLE LATITUDE LONGITUDE DATE TEMP DEPTH I CODE DD MM DD MM YYMMDD C M

1 CIT 60 24 24 N 77 27 W 73 228 4 .0 1545 2 CIT 61 24 26 N 77 20 W 73 228 4.0 1490 1 CIT 76 24 39 N 76 28 w 73 305 4 .0 1586 3 CIT 85 24 23 N 77 23 w 73 309 4.1 1528 3 CIT101 24 27 N 77 28 w 73 920 4 .0 1594 9 CIT103 24 08 H 77 22 w 73 921 4.2 1436 1 CIT104 23 56 N 77 14 w 73 921 4 .2 1350 1 C ITlll 23 37 N 77 14 w 73 922 4.3 1338 1 CIT114 23 41 N 76 50 w 73 923 4.3 1338 4 CIT117 23 33 N 76 45 w 73 924 4.3 1308 5 CIT122 24 06 N 77 19 w 73 924 4 .2 1434 5 CIT123 24 12 N 77 18 w 73 924 4 .2 1435 4 CIT124 24 25 N 77 25 w 73 925 4 .0 1593 3 CIT125 24 30 N 77 22 w 73 925 4.1 1496 3 CIII44 24 30 N 77 22 w 74 203 4.1 1494 1 CIT145 24 14 N 77 19 w 74 203 4.2 1440 1 CIT147 23 56 N 77 14 w 74 203 4 .2 1356 2 CIT149 23 52 N 77 18 w 74 204 4.2 1388 7 CIT168 24 26 N 77 23 w 74 207 4 .0 1577 4 CIT169 24 35 N 77 32 w 74 207 3 .9 1742 2 CIT176 24 39 N 76 29 w 74 210 4 .0 1632 1 CIT178 24 13 N 76 06 w 74 210 3 .8 1794 1 CIT184 23 55 N 75 26 w 74 211 3 .5 2188 1 CIT186 23 45 N 75 42 w 74 212 3.8 1849 1 CIT191 24 10 N 75 56 w 74 213 3 .8 1810 2 CIT247 24 39 N 77 32 w 741030 3 .9 1724 1 CIT254 23 34 N 76 43 w 741101 4.3 1303 1 CIT255 23 30 N 76 57 w 741101 4.3 1320 2 CIT261 23 36 N 77 08 w 741102 4 .2 1353 3 CIT267 24 24 N 77 22 w 741103 4.1 1542 2 CIT268 24 31 N 77 22 w 741104 4.1 1490 1 CIT270 24 23 N 77 26 H 741104 4.1 1550 1 CIT272 24 05 N 77 23 w 741104 4 .2 1438 1 CIT284 23 57 N 75 59 w 741109 3 .8 1776 1 CIT305 23 44 N 76 51 w 75 404 4.3 1350 2 CIT309 23 44 N 76 47 w 75 405 4.3 1316 1 CIT311 23 38 N 77 14 w 75 403 4.2 1356 1 CIT314 24 46 N 77 15 w 75 406 4.2 1367 1 CIT316 24 24 N 77 27 w 75 407 4 .0 1559 5 CIT317 24 16 N 77 20 w 75 407 4.1 1464 1 CIT320 23 40 N 76 43 w 45 408 4.3 1322 1 CIT326 23 51 N 77 16 w 75 410 4 .2 1383 1 CIT336 23 48 N 75 46 w 75 413 3 .8 1848 3 CIT349 24 36 N 77 33 w 75 419 3 .9 1748 2 CIT351 24 19 N 77 21 w 75 420 4.1 1519 Barathritea parri CONTINUED MEAN COUNT SAMPLELATITUDELONGITUDE DATE TEMP DEPTH SF A CODE DD MM DD MM YYMMDD C M

3 CIT354 24 34 N 77 31 V 75 819 3.9 1724 4 CIT356 24 23 N 77 26 W 75 820 3.9 1654 5 CIT357 23 55 N 77 17 W 75 820 4.2 1386 2 CIT359 23 53 N 77 10 W 75 821 4.2 1358 1 CIT361 23 30 N 76 55 w 75 821 4.3 1300 2 CIT364 23 38 N 77 07 w 75 822 4.2 1363 1 CIT365 23 50 N 77 14 V 75 822 4.2 1372 2 CIT366 23 49 N 77 06 w 75 822 4.2 1366 1 CIT376 23 55 N 75 59 w 75 826 3.8 1762 1 CIT380 24 39 N 76 26 w 75 827 4.0 1640 2 CIT387 24 16 N 76 16 w 75 829 3.8 1762 1 CIT403 23 57 N 75 27 w 75 905 3.5 2166 3 CIT405 24 28 N 77 20 w 76 228 4.1 1464 1 CIT407 23 53 N 77 05 V 76 229 4.3 1336 2 CIT408 23 50 N 76 51 w 76 229 4.3 1346 1 CIT413 23 51 N 77 13 w 76 301 4.2 1372 1 CIT414 23 40 N 76 59 w 76 301 4.2 1360 2 CIT422 23 53 N 77 07 w 76 302 4.3 1339 12 CIT430 24 32 N 77 31 w 76 306 3.9 1661 5 CIT431 24 16 N 77 22 w 76 306 4.1 1478 1 CIT435 23 50 N 77 16 w 76 309 4.2 1380 1 CIT437 23 32 N 76 52 w 76 309 4.3 1333 1 CIC012 23 51 N 75 50 w 80 9 3 3.7 1856 1 CIC018 22 46 N 75 34 w 80 9 7 3.3 2337 4 CIC041 23 53 N 77 08 w 80 915 4.2 1341

PenoDus macdonaldi

1 CIT 46 23 30 N 77 05 w 73 224 4.5 1257 1 CIT 85 24 23 N 77 23 If 73 309 4.1 1528 2 CIT103 24 08 N 77 22 w 73 921 4.2 1436 3 CIT104 23 56 N 77 14 w 73 921 4.2 1350 1 CITlll 23 37 N 77 14 w 73 922 4.3 1338 1 CIT122 24 06 N 77 19 w 73 924 4.2 1434 2 CIT123 24 12 N 77 18 w 73 924 4.2 1435 1 CIT144 24 30 N 77 22 w 74 203 4.1 1494 1 CIT163 23 32 N 77 10 w 74 206 4.3 1345 1 CIT167 23 38 N 77 17 V 74 207 4.1 1515 1 CIT168 24 26 N 77 23 w 74 207 4.0 1577 1 CIT169 24 35 N 77 32 w 74 207 3.9 1742 3 CIT261 23 36 N 77 08 w 741102 4.2 1353 1 CIT272 24 05 N 77 23 tf 741104 4.2 1438 1 CIT302 24 11 N 77 23 w 75 403 4.1 1458 1 CIT314 24 46 N 77 15 w 75 406 4.2 1367 3 CIT317 24 16 N 77 20 w 75 407 4.1 1464 2 CIT354 24 34 N 77 31 w 75 819 3.9 1724 Penooua macdonaldi continued

MEAN COUNT SAMPLE LATITUDELONGITUDE DATETEMP DEPTH SP A CODE DD MM DD MM YYMMDD C M

2 CIT366 23 49 N 77 06 V 75 822 4.2 1366 1 CIT370 23 34 N 76 42 W 75 823 4.4 1282 1 CIT405 24 28 N 77 20 W 76 228 4.1 1464 1 CIT414 23 40 N 76 59 W 76 301 4.2 1360 4 CIT426 25 16 N 77 49 W 76 304 3.4 2700 1 CIT437 23 32 N 76 52 W 76 309 4.3 1333 1 CIC041 23 53 N 77 08 W 80 915 4.2 1341

Bassozetus nonnalis

1 CIT 41 23 43 N 76 49 W 73 222 4.4 1298 2 CIT 42 23 31 N 76 44 W 73 223 4.4 1280 1 CIT 48 23 44 N 77 13 W 73 224 4.2 1390 2 CIT 50 23 42 N 76 43 W 73 225 4.5 1271 1 CIT 51 23 38 N 76 49 W 73 225 4.4 1290 3 CIT 53 23 48 N 77 03 W 73 226 4.3 1325 2 CIT 57 24 05 N 77 23 W 73 227 4.2 1404 1 CIT 58 24 14 N 77 18 W 73 227 4.2 1408 5 CIT 59 24 14 N 77 25 W 73 227 4.2 1435 1 CIT 60 24 24 N 77 27 W 73 228 4.0 1545 1 CIT 78 24 25 N 76 10 W 73 306 3.9 1692 5 CIT 85 24 23 N 77 23 W 73 309 4.1 1528 1 CIT 97 25 17 N 77 45 W 73 312 3.4 2748 4 CIT103 24 08 N 77 22 H 73 921 4.2 1436 1 CIT104 23 56 N 77 14 W 73 921 4.2 1350 7 CIT105 23 48 N 77 04 W 73 921 4.2 1368 1 CITlll 23 37 N 77 14 W 73 922 4.3 1338 2 CIT114 23 41 N 76 50 W 73 923 4.3 1338 1 CIT115 23 38 N 76 48 W 73 923 4.3 1316 1 CIT121 23 55 N 77 20 W 73 924 4.2 1394 2 CIT124 24 25 N 77 25 W 73 925 4.0 1593 1 CIT126 24 41 N 77 34 W 73 925 3.7 1872 2 CIT144 24 30 N 77 22 W 74 203 4.1 1494 1 CIT145 24 14 N 77 19 W 74 203 4.2 1440 1 CIT149 23 52 N 77 18 W 74 204 4.2 1388 3 CIT150 23 48 N 77 03 W 74 204 4.2 1358 2 CITl54 23 32 N 76 46 W 74 205 4.3 1311 3 CIT156 23 43 N 76 50 W 74 205 4.3 1332 1 CIT159 23 39 N 76 50 W 74 205 4.3 1346 8 CIT168 24 26 N 77 23 tf 74 207 4.0 1577 1 CIT169 24 35 N 77 32 W 74 207 3.9 1742 1 CIT175 24 32 N 76 18 ff 74 210 3.9 1704 1 CIT176 24 39 N 76 29 W 74 210 4.0 1632 2 CIT177 24 23 N 76 09 W 74 210 3.8 1770 3 CIT178 24 13 N 76 06 W 74 210 3.8 1794 2 CITl87 24 00 N 75 49 W 74 212 3.7 1887 1 CIT250 23 52 N 76 52 W 741031 4.4 1291 3 CIT251 23 41 N 76 44 W 741031 4.3 1315 Bassozetus normalis continued

MEAN COUNT SAMPLE LATITUDE LONGITUDE DATE TEMP depth SP A CODE DD MM DD MM YYMMDD CM

2 CIT252 23 39 N 76 45 ff 741101 4.3 1327 8 CIT253 23 44 N 76 49 W 741101 4.3 1332 4 CIT256 23 37 N 77 06 W 741101 4.2 1360 2 CIT268 24 31 N 77 22 W 741104 4.1 1490 1 CIT270 24 23 N 77 26 W 741104 4.1 1550 3 CXT272 24 05 N 77 23 W 741104 4.2 1438 1 CIT281 23 54 N 75 30 W 741109 3.6 2141 1 CIT282 24 00 N 75 47 W 741109 3.7 1905 1 CIT314 24 46 N 77 15 W 75 406 4 .2 1367 3 CIT317 24 16 N 77 20 W 75 407 4.1 1464 1 CIT320 23 40 N 76 43 W 45 408 4.3 1322 2 CIT324 23 31 N 76 56 ff 75 409 4.3 1333 4 CIT331 24 32 N 76 17 W 75 412 3.9 1678 4 CIT333 24 24 N 76 08 W 75 412 3 .8 1772 4 CIT335 23 58 N 76 01 W 75 413 3.8 1766 1 CIT340 24 20 N 76 00 V 75 415 3.9 1748 2 CIT349 24 36 N 77 33 W 75 419 3.9 1748 4 CIT350 24 02 N 77 24 W 75 420 4.2 1430 2 CIT351 24 19 N 77 21 W 75 420 4.1 1519 1 CIT354 24 34 N 77 31 W 75 819 3.9 1724 2 CIT356 24 23 N 77 26 W 75 820 3.9 1654 1 CIT360 23 38 N 76 49 W 75 821 4.3 1338 1 CIT361 23 30 N 76 55 W 75 821 4.3 1300 5 CIT362 23 39 N 77 07 W 75 821 4.2 1363 1 CIT363 23 51 N 77 55 W 75 822 4.3 1320 3 C1T364 23 38 N 77 07 W 75 822 4.2 1363 2 CIT365 23 50 N 77 14 W 75 822 4.2 1372 4 CIT366 23 49 N 77 06 W 75 822 4.2 1366 2 CIT370 23 34 N 76 42 W 75 823 4 .4 1282 1 CIT374 25 16 N 77 42 ff 75 824 3.3 2960 2 CIT378 24 21 N 76 04 ff 75 826 3.8 1761 1 CIT379 24 33 N 76 17 ff 75 827 3.9 1659 2 CXT382 23 57 N 76 01 W 75 828 3 .8 1763 2 CIT384 24 09 N 75 55 W 75 828 3 .8 1844 1 CIT392 22 48 N 75 38 W 75 831 3.5 2250 1 CIT400 21 59 N 75 06 w 75 903 3 .4 2681 1 CIT405 24 28 N 77 20 W 76 228 4.1 1464 4 CIT408 23 50 N 76 51 W 76 229 4.3 1346 1 CIT410 23 30 N 76 51 ff 76 229 4.3 1323 6 CIT414 23 40 N 76 59 W 76 301 4 .2 1360 1 CIT416 23 40 N 77 16 ff 76 302 4.3 1338 4 CIT417 23 52 N 77 16 W 76 302 4 .2 1387 1 CIT418 23 50 N 77 17 H 76 302 4.2 1386 1 CIT422 23 53 N 77 07 ff 76 302 4.3 1339 1 CIT428 24 49 H 77 40 W 76 305 3.6 1977 4 CIT431 24 16 N 77 22 ff 76 306 4.1 1478 6 CIT435 23 50 N 77 16 ff 76 309 4.2 1380 2 CIT437 23 32 N 76 52 ff 76 309 4.3 1333 Baasozetus taenia

MEAN COUNT SAMPLE LATITUDE LONGITUDE DATE TEMP DEPTH SP A CODE DD MM DD MM YYMMDD C M

3 CIT288 25 16 N 77 43 W 741111 3.4 2870 3 CIT316 24 24 N 77 27 W 75 407 4.0 1559 2 CIT332 24 39 N 76 27 W 75 412 4.0 1627 1 CIT353 25 16 N 77 39 W 75 421 3.3 3032 2 CIT426 25 16 N 77 49 W 76 304 3.4 2700 1 CIC015 23 50 N 73 14 W 80 9 5 2.3 5267 1 CIC021 24 57 N 74 25 W 80 9 9 2.2 4846 1 CIC027 26 04 N 74 03 W 80 910 2.2 5043 1 CIC028 26 27 N 74 03 W 80 910 2.2 4581 10 CIC034 26 09 N 75 26 W 80 911 2.3 4539 1 CIC060 29 36 N 74 32 W 80 919 2.2 4553 1 CI6003 28 27 N 76 31 W 78 211 • 4960 2 CID004 21 57 N 74 46 W 811116 3.0 2728

Abvssobrotula ealathese

1 CIC021 24 57 N 74 25 W 80 9 9 2.2 4846 1 CIC034 26 09 N 75 26 W 80 911 2.3 4539 1 CIC056 28 18 N 74 12 W 80 918 2.2 4380 1 CIC060 29 36 N 74 32 W 80 919 2.2 4553 2 CI6004 28 43 1 75 49 7 780212 • 4930 1 CID018 23 02 » 22 55 W 811124 2.1 5345 Bnthvonus ppet-oralis

MEAN INT SAMPLE LATITUDE LONGITUDE DATE temp depth A CODE DD MM DD MM YYMMDD C M

24 CIT 71 24 20 N 75 58 W 73 304 3 .9 1682 4 CIT 72 24 00 N 75 44 W 73 304 3.7 1884 7 CIT 73 23 46 N 75 42 W 73 304 3 .8 1792 13 CIT 76 24 39 N 76 28 W 73 305 4 .0 1586 4 CIT 78 24 25 N 76 10 w 73 306 3 .9 1692 5 CIT 97 25 17 N 77 45 w 73 312 3 .4 2748 2 CIT168 24 26 N 77 23 w 74 207 4 .0 1577 1 CIT172 25 17 N 77 46 w 74 208 3 .4 2836 6 CIT175 24 32 N 76 18 w 74 210 3 .9 1704 4 CITl76 24 39 N 76 29 w 74 210 4.0 1632 4 CIT177 24 23 N 76 09 w 74 210 3.8 1770 31 CIT178 24 13 N 76 06 w 74 210 3 .8 1794 3 CITl82 23 56 N 75 58 w 74 211 3 .8 1769 4 CITl83 23 45 N 75 39 w 74 211 3 .8 1807 10 CIT184 23 55 N 75 26 w 74 211 3 .5 2188 2 CITl86 23 45 N 75 42 w 74 212 3 .8 1849 5 CIT187 24 00 N 75 49 w 74 212 3.7 1887 7 CIT191 24 10 N 75 56 w 74 213 3 .8 1810 3 CITl92 24 21 N 75 59 w 74 213 3 .8 1768 6 CIT193 24 25 N 76 11 w 74 213 3 .9 1748 1 CIT272 24 05 N 77 23 w 741104 4.2 1438 8 CIT276 24 21 N 76 10 w 741107 3 .8 1772 7 CIT279 23 50 N 75 50 w 741108 3 .8 1838 4 CIT280 23 52 N 75 16 w 741108 3 .5 2354 5 CIT281 23 54 N 75 30 w 741109 3.6 2141 2 CIT282 24 00 N 75 47 w 741109 3.7 1905 3 CIT284 23 57 N 75 59 w 741109 3 .8 1776 7 CIT331 24 32 N 76 17 w 75 412 3 .9 1678 22 CIT332 24 39 N 76 27 V 75 412 4 .0 1627 11 CIT333 24 24 N 76 08 w 75 412 3.8 1772 15 CIT334 24 15 N 76 06 w 75 413 3.8 1792 27 CIT335 23 58 N 76 01 w 75 413 3 .8 1766 6 CIT336 23 48 N 75 46 w 75 413 3 .8 1848 5 CIT337 23 55 N 75 28 w 75 414 3.6 2152 218

Bathvonus pectoralis continued

MEAN COUNT SAMPLE LATITUDE LONGITUDE DATE TEMP DEPTH SP A CODE DD MM DD MM YYMMDD* CM 3 CIT339 24 10 N 75 54 W 75 414 3 .8 1842 1 CIT340 24 20 N 76 00 W 75 415 3 .9 1748 14 CIT341 24 39 N 76 26 tf 75 415 4 .0 1628 6 CIT343 23 48 N 75 47 W 75 416 3 .7 1854 16 CIT346 22 55 N 75 17 tf 417 3 .5 2422 4 CIT347 22 51 N 75 38 W 75 417 3 .5 2246 4 CII353 25 16 N 77 39 W 75 421 3 .3 3032 3 CIT374 25 16 N 77 42 W 75 824 3.3 2960 3 CIT376 23 55 N 75 59 W 75 826 3 .8 1762 4 CIT378 24 21 N 76 04 W 75 826 3 .8 1761 2 CIT379 24 33 N 76 17 tf 75 827 3 .9 1659 4 CIT380 24 39 N 76 26 W 75 827 4 .0 1640 1 CIT382 23 57 N 76 01 W 75 828 3 .8 1763 3 CIT383 23 50 N 75 49 W 75 828 3 .8 1830 3 CIT384 24 09 N 75 55 W 75 828 3 .8 1844 1 CIT386 23 59 N 75 46 W 75 829 3 .8 1922 8 C1T387 24 16 N 76 16 W 75 829 3 .8 1762 12 CIT388 23 56 N 75 32 W 75 829 3 .6 2146 1 CIT390 23 50 N 75 14 W 75 830 3 .5 2354 13 CIT391 22 54 N 75 14 W 75 830 3 .5 2427 9 CIT392 22 48 N 75 38 W 75 831 3 .5 2250 2 CIT395 22 01 N 75 04 tf 75 901 3 .4 2686 28 CIT399 22 29 N 75 11 tf 75 902 3 .4 2577 11 CIT400 21 59 N 75 06 W 75 903 3 .4 2681 36 CIT402 22 53 N 75 27 W 75 904 3 .5 2404 8 CIT403 23 57 N 75 27 W 75 905 3 .5 2166 2 CIT442 25 17 N 77 38 W 76 311 3 .3 3094 9 CIC011 23 46 N 75 48 W 80 9 3 3.7 1790 10 CIC017 22 55 N 75 25 W 80 9 7 3.4 2423 4 CIC018 22 46 N 75 34 tf 80 9 7 3.3 2337 2 CIC019 22 34 N 75 35 W 80 9 8 3.3 2565 3 CIC027 26 04 N 74 03 tf 80 910 2.2 5043 5 C1C034 26 09 N 75 26 tf 80 911 2.3 4539 1 CIC045 26 38 N 76 05 W 80 917 2.2 4777 1 CIC056 28 18 N 74 12 tf 80 918 2.2 4380 1 CIC057 29 20 N 73 19 W 80 919 2.2 4253 1 CIC060 29 36 N 74 32 tf 80 919 2.2 4553 1 CIC061 29 48 N 76 02 tf 80 920 2.3 4246 35 CID004 21 57 N 74 46 W 811116 3 .0 2728 5 CID005 22 03 N 74 48 tf 811116 * 2728 3 CID006 21 51 N 74 52 W 81117 3 .0 2748 3 CID007 21 50 N 74 56 tf 811117 3 .0 2758 5 CID008 21 34 N 74 32 tf 811118 3.0 2745 1 CZD009 21 27 N 74 21 tf 811118 3 .0 2714 2 CID018 23 02 N 22 55 W 811124 2.1 5345 Acanthonus armatus

MEAN fNT SAMPLE LATITUDE l o n g i t u d e DATE TEMP DEPTH A CODE DD MM DD MM YYMMDD C M

1 CIT 70 24 30 N 76 13 W 73 303 3.9 1677 11 CIT 71 24 20 N 75 58 W 73 304 3.9 1682 4 CIT 72 24 00 N 75 44 w 73 304 3.7 1884 10 CIT 73 23 46 N 75 42 w 73 304 3.8 1792 11 CIT 76 24 39 N 76 28 w 73 305 4.0 1586 1 CIT 78 24 25 N 76 10 w 73 306 3.9 1692 3 CITl72 25 17 N 77 46 w 74 208 3.4 2836 8 CIT177 24 23 N 76 09 w 74 210 3.8 1770 34 CIT178 24 13 N 76 06 w 74 210 3.8 1794 1 CITl82 23 56 N 75 58 w 74 211 3.8 1769 5 CITl83 23 45 N 75 39 w 74 211 3.8 1807 21 CIT184 23 55 N 75 26 w 74 211 3.5 2188 8 CITl86 23 45 N 75 42 w 74 212 3.8 1849 16 CIT187 24 00 N 75 49 w 74 212 3.7 1887 6 CIT191 24 10 N 75 56 w 74 213 3.8 1810 1 CITl92 24 21 N 75 59 w 74 213 3.8 1768 5 CITl93 24 25 N 76 11 w 74 213 3.9 1748 20 CIT276 24 21 N 76 10 w 741107 3.8 1772 1 CIT279 23 50 N 75 50 w 741108 3.8 1838 1 CIT280 23 52 N 75 16 w 741108 3.5 2354 1 CIT284 23 57 N 75 59 w 741109 3.8 1776 6 CIT332 24 39 N 76 27 w 75 412 4.0 1627 5 CIT333 24 24 N 76 08 w 75 412 3.8 1772 3 CIT334 24 15 N 76 06 V 75 413 3.8 1792 34 CIT335 23 58 N 76 01 w 75 413 3.8 1766 11 CIT336 23 48 N 75 46 w 75 413 3.8 1848 9 CIT337 23 55 N 75 28 w 75 414 3.6 2152 8 CIT338 23 59 N 75 48 w 75 414 3.7 1890 4 CIT339 24 10 N 75 54 w 75 414 3.8 1842 6 CIT341 24 39 N 76 26 w 75 415 4.0 1628 3 CIT343 23 48 N 75 47 V 75 416 3.7 1854 1 CIT353 25 16 N 77 39 w 75 421 3.3 3032 1 CIT374 25 16 N 77 42 w 75 824 3.3 2960 2 CIT376 23 55 N 75 59 w 75 826 3.8 1762 2 CIT377 24 00 N 75 46 w 75 826 3.7 1917 11 CIT378 24 21 N 76 04 V 75 826 3.8 1761 1 CIT380 24 39 N 76 26 w 75 827 4.0 1640 6 CIT382 23 57 N 76 01 w 75 828 3.8 1763 7 CIT383 23 50 N 75 49 w 75 828 3.8 1830 4 CIT384 24 09 N 75 55 w 75 828 3.8 1844 25 CIT387 24 16 N 76 16 w 75 829 3.8 1762 8 CIT388 23 56 N 75 32 w 75 829 3.6 2146 6 CIT390 23 50 N 75 14 tf 75 830 3.5 2354 2 CIT391 22 54 N 75 14 tf 75 830 3.5 2427 4 CIT392 22 48 N 75 38 tf 75 831 3.5 2250 3 CIT399 22 29 N 75 11 tf 75 902 3.4 2577 12 CIT403 23 57 N 75 27 tf 75 905 3.5 2166 2 CIT426 25 16 N 77 49 H 76 304 3.4 2700 Acanthonus armatua continued

MEAN COUNTSAMPLELATITUDE LONGITUDEDATE TEMPDEPTH SP A CODE DDMM DD MM YYMMDD C M

1 CIT442 25 17 N 77 38 ff 76 311 3.3 3094 29 CIC011 23 46 N 75 48 ff 80 9 3 3.7 1790 8 CIC012 23 51 N 75 50 ff 80 9 3 3.7 1856 2 CIC017 22 55 N 75 25 ff 80 9 7 3.4 2423 3 CIC018 22 46 N 75 34 W 80 9 7 3.3 2337 8 CID004 21 57 N 74 46 w 811116 3.0 2728 1 CID005 22 03 N 74 48 w 811116 • 2728 8 CID009 21 27 N 74 21 w 811118 3.0 2714 Appendix 4. Cluster analysis outpuc based on morphological characters. -d •- Ci •J \ 1X 11 or ■tl y y • U ft X J " *- ,"1 • n cv Mi 11 ft j* X Ul - «. «-» il4 lA o i/i X* n 1 1 l3U 3 Cl ift-Hi u« a p- J fHIif 14, X U HI n *• H * M J O. U U X y H a Hi U i/i a h- IU * 9 a * ►* U Ml 4 4 A O 9 41 a Ui O f— t Wl ft. *1 444-4 M *1 n ■T* )4 () ►- !» n» H © Ui JXu X UJ 1 ■-» -1J*- » • *l 4# «• o li i/I r1C•r m» ■ or ►4 Hi pi •* X •1 HI m t* tr *4 144X 4-4 *1 *■ V J «xv» x U« C.I • « «n a ft i_J Ui 4 «* u ♦-u j in * ft. 221 IU »•t ►» 44 I4* -I o -Mlp- Cl '.1 4 X 44O o mi p U* p- HI m Cl9 «« (M 1*1 1' M' 4 I J if (ft f l H & o M H "J |\J f I r>J M N N fJ N ( I ( N fJ N r>JI f N M |\J "J H u in H •A a u* LI o m» ft -J 4*1 I1>LI U 1/1 a a a Hi o A 9 14* Ul O C* L* •* X ■»» 4 1 nXi X ’1*n •i) If u L» t4 rt IN Ul «J Mi n O _J ru M U> ft X -o ;* X p * i n l*» 4 J Ul *- *> U» > *1 ml *c >U © IP POI HP O p .> *■ UifIU im A i/i 4 u u, r n n u> u t/» -J _J 1J u u <*1 */• £ M *"t X P (*lH Pi n m p O «l •i 2“ . Pi * r* i» n J 4/1 HI m a m • I mi mi U a <41 Cl i/i * 4 4*1 Jft‘J ft ■J -4 1 ■ 11 14 i»i 11 »Hir» U •9r* it it* t 4 • i u a

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•a a 3 5 4J oa u VITA

Herbert Jacque Carter

Born in Kankakee, Illinois, March 18, 1953. Received high school diploma from Bradley High School, Bradley, Illinois, 1971. Bachelor of

Science from Northern Illinois University, 1975; Master of Science from

Northern Illinois University, 1977. Entered the graduate school of the

Virginia Institute of Marine Science, College of William and Mary in

1978. Admitted to candidacy for degree of Doctor of Philosophy, College of William and Mary, 1983. Presently employed as a joint research fellow of the New York Zoological Society and Smithsonian Institution,

Museum of Natural History.

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