Systematics of the Onychoteuthidae Gray, 1847 (Cephalopoda: Oegopsida) K.S. Bolstad A thesis submitted to the Earth & Oceanic Sciences Research Institute Auckland University of Technology in fulfilment of the requirements for the degree of Doctor of Philosophy supervised by Dr Steve O’Shea 2008 i TABLE OF CONTENTS List of Tables............................................................................................................... iii List of Figures..............................................................................................................iv Abstract....................................................................................................................... 1 Introduction ................................................................................................................. 3 Foreword — Generic and Familial Nomenclature....................................................... 3 Introduction ................................................................................................................. 4 Materials & Methods ................................................................................................. 13 Checklist of Species.................................................................................................. 24 Quick Visual Reference to the Onychoteuthidae ...................................................... 25 Systematics............................................................................................................... 31 Onychoteuthis ....................................................................................................... 33 Onychoteuthis banksii ....................................................................................... 36 Onychoteuthis bergii ......................................................................................... 42 Onychoteuthis aequimanus............................................................................... 48 Onychoteuthis mollis (new comb.) .................................................................... 57 Onychoteuthis compacta................................................................................... 61 Onychoteuthis borealijaponica .......................................................................... 69 Onychoteuthis meridiopacifica .......................................................................... 78 Onychoteuthis lacrima....................................................................................... 84 Onychoteuthis prolata ....................................................................................... 90 Onykia ................................................................................................................... 96 Onykia (Onykia) carriboea ................................................................................ 99 Onykia (Onykia) robusta (new comb.)............................................................. 106 Onykia (Onykia) loennbergii (new comb.) ....................................................... 118 Onykia (Onykia) aequatorialis (new comb.) .................................................... 125 Onykia (Onykia) robsoni (new comb.)............................................................. 131 Onykia (Onykia) indica .................................................................................... 140 Onykia (Moroteuthopsis) ingens (new comb.)................................................. 145 Ancistroteuthis..................................................................................................... 159 Ancistroteuthis lichtensteinii ............................................................................ 161 Kondakovia ......................................................................................................... 170 Kondakovia longimana.................................................................................... 171 Notonykia ............................................................................................................ 178 Notonykia africanae ........................................................................................ 179 Notonykia nesisi .............................................................................................. 187 Gen. nov. 1.......................................................................................................... 193 Gen. nov. 1 knipovitchi (new comb.) ............................................................... 195 Gen. nov. 2.......................................................................................................... 200 Gen. nov. 2 rancureli (new comb.) .................................................................. 202 Gen. nov. 2 sp. 1............................................................................................. 206 Gen. nov. 2 sp. 2............................................................................................. 209 Gen. nov. 2 sp. 3............................................................................................. 212 Discussion............................................................................................................... 216 Conclusions............................................................................................................. 223 Acknowledgements................................................................................................. 225 References.............................................................................................................. 226 Appendices ............................................................................................................. 240 Appendix A — Translation, Lichtenstein 1818 .................................................... 240 Appendix B — Translation, Appellöf 1891 .......................................................... 242 Appendix C — Translation, Thiele 1920 ............................................................. 246 Appendix D — Translation, Adam 1962.............................................................. 249 Appendix E — Systematic Summary .................................................................. 256 i ATTESTATION OF AUTHORSHIP I hereby declare that this submission is my own work and that, to the best of my knowledge and belief, it contains no material previously published or written by another person (except where explicitly defined in the acknowledgements), nor material that, to a substantial extent, has been submitted for the award of any other degree or diploma from a university or institute of higher learning. _________________________________________ Kathrin S. Bolstad ii LIST OF TABLES 1. Comparative table of onychoteuthid genera 2. Comparative table of species in the genus Onychoteuthis 3. Measurements and counts of Onychoteuthis banksii 4. Measurements and counts of Onychoteuthis bergii 5. Measurements and counts of Onychoteuthis aequimanus 6. Measurements and counts of Onychoteuthis compacta 7. Measurements and counts of Onychoteuthis borealijaponica 8. Measurements and counts of Onychoteuthis meridiopacifica 9. Measurements and counts of Onychoteuthis lacrima 10. Measurements and counts of Onychoteuthis prolata 11. Comparative table of species in the genus Onykia 12. Measurements and counts of Onykia (Onykia) carriboea 13. Measurements and counts of Onykia (Onykia) robusta 14. Measurements and counts of Onykia (Onykia) loennbergii 15. Measurements and counts of Onykia (Onykia) aequatorialis 16. Measurements and counts of Onykia (Onykia) robsoni 17. Measurements and counts of Onykia (Onykia) indica 18. Measurements and counts of Onykia (Moroteuthopsis) ingens 19. Sexually dimorphic characters in the lower beaks of Onykia (Moroteuthopsis) ingens 20. Measurements and counts of Ancistroteuthis lichtensteinii 21. Measurements and counts of Kondakovia longimana 22. Measurements and counts of Notonykia africanae 23. Measurements and counts of Notonykia nesisi 24. Measurements and counts of Gen. nov. 1 knipovitchi 25. Measurements and counts of Gen. nov. 2 rancureli and Gen. nov. 2 spp. 1–3 iii LIST OF FIGURES 1. Distribution of material examined in this study 2. Measures and counts used in text 3. Onychoteuthid hook morphology and terminology 4. Paralarval sucker morphology and terminology 5. Radular morphology and terminology 6, 7. Onychoteuthis banksii 8, 9. Onychoteuthis bergii 10–13. Onychoteuthis aequimanus 14. Onychoteuthis mollis 15, 16. Onychoteuthis compacta 17–19. Onychoteuthis borealijaponica 20, 21. Onychoteuthis meridiopacifica 22, 23. Onychoteuthis lacrima 24, 25. Onychoteuthis prolata 26–28. Onykia (Onykia) carriboea 29–33. Onykia (Onykia) robusta 34–36. Onykia (Onykia) loennbergii 37, 38. Onykia (Onykia) aequatorialis 39–41. Onykia (Onykia) robsoni 42, 43. Onykia (Onykia) indica 44–48. Onykia (Moroteuthopsis) ingens 49–52. Ancistroteuthis lichtensteinii 53–55. Kondakovia longimana 56–58. Notonykia africanae 59, 60. Notonykia nesisi 61, 62. Gen. nov. 1 knipovitchi 63. Walvisteuthis virilis 64–67. Gen. nov. 2 rancureli and Gen. nov. 2 spp. 1–3 iv ABSTRACT Squids in the family Onychoteuthidae Gray, 1847 have been reported from every ocean but the Arctic, are taken frequently in deep-sea fisheries bycatch, and are ecologically important in the diets of many marine predators including cetaceans, pinnipeds, sharks, and seabirds. However, the diversity and systematics of the family have remained poorly understood. Of the 60+ nominal species, 12–14 have generally been accepted in recent