ABSTRACT SYSTEMATICS OF AFRICAN CICHLID FISHES: DETERMINATION OF THE MOST PRIMITIVE TAXON, AND STUDIES ON THE HAPLOCHROMINES OF LAKE MALAWI (TELEOSTEI: CICHLIDAE) Michael Kenneth Oliver Yale University 1984 The cichlids of Africa present many challenges to the taxonomist. Diverse systematic problems at three levels in cichlid phylogeny form the subject matter of this dissertation. At the most general level, I summarize current morphological evidence that supports cichlid monophyly and the most-inclusive groups of cichlids. With this review as a framework, I argue that the most phylogenetically primitive African cichlid is Heterochromia multidens, a little-known large cichlid from the Zaire River basin. I redescribe this monotypic genus and species and illustrate much of its osteology. It possesses a distinctive suite of derived characters, but is uniquely primitive among African cichlids in infraorbital morphology and other features. At a less inclusive phylogenetic level, I examine the controversy over the monophyletic status of the "species flock" of haplochromine cichlids in Lake Malawi. A derived type of anal-fin markings present in a subset of the Malawian fauna is shared with many non-Malawian haplochromines, indicating that the species flock of Lake Malawi is not monophyletic. The two (or more) faunal components of Lake Malawi are ecologically distinct. The remaining major parts of the study are concerned with particular assemblages of haplochromine cichlids in Lake Malawi. I present a detailed phylogenetic analysis of the Cyrtocara liyingstonii species-group of ambush predators, and characterize a new species in the group. I also describe three new species of haplochromines with three lateral spots and present a key to the three-spot assemblage. Intralacustrine dispersion of Malawi cichlids is briefly addressed in appended material. SYSTEMATICS OF AFRICAN CICHLID FISHES: DETERMINATION OF THE MOST PRIMITIVE TAXON, AND STUDIES ON THE HAPLOCHROMINES OF LAKE MALAWI (TELEOSTEI: CICHLIDAE) A Dissertation Presented to the Faculty of the Graduate School of Yale University in Candidacy for the Degree of Doctor of Philosophy by Michael Kenneth Oliver May 1984 l ~ Copyright ~ Michael Kenneth Oliver ~ ALL RIGHTS RESERVED Ex Africa chaos. P. H. Greenwood (pers. comm., 1971) I ACKNOWLEDGMENTS This work was supported by NSF grants DEB 80-05538 and DEB 79-12258, and by grants from the Smithsonian Oceanographic Sorting Center and the Peabody Museum of Natural History, Yale University. The curators of the Department of Ichthyology, American Museum of Natural History and the Fish Division, British Museum (Natural History) gave multifarious kinds of support, encouragement, and training in earlier phases of my graduate career. For permitting me to examine or borrow material in their care, I thank P. H. Greenwood and the Trustees of the British Museum, London; c. L. Smith, American Museum of Natural History, New York; D. Thys van den Audenaerde, Mus~e Royal de l'Afrique Centrale, Tervuren; and s. H. Weitzman and R. P. Vari, US National Museum of Natural History, Washington, DC. I want to thank Frederick Cichocki for sharing with me, a decade ago, his unpublished information that cichlid infraorbitals are phylogenetically informative. This advice led me to survey them independently, and to discover that Heterochromia is among the most plesiomorphic of all cichlids. Cichocki's remarkable dissertation, filled with valuable data as well as provocative discussion, will surely remain indispensable to students of cichlid interrelationships. My final dissertation committee at Yale consisted of J. David Archibald, James E. Rodman, Bruce H. Tiffney, and Keith s. Thomson (chairman). Each contributed precious support, friendship, advice, and/or philosophy. My dear, patient wife, Joan, and parents, Kenneth and Madaline, have supported me all along in every way. I couldn't have done it without them and wouldn't have wanted to try. Zikomo wambiri! TABLE OF CONTENTS LIST OF FIGURES ••••••••••••••••••••••••••••••••••••••••••••••••••• vi LIST OF TABLES .................................................... ix CHAPTER 1. GENERAL INTRODUCTION ••••••••••••••••••••••••••••••••••••••••• 1 CHAPTER 2. RECOGNITION OF HETEROCHROMIS MULTIDENS (PELLEGRIN) FROM THE CENTRAL ZAIRE RIVER SYSTEM AS THE MOST PRIMITIVE AFRICAN CICHLID ••••••••••••••••••••••••••••••• 5 CHAPTER 3. IS THE HAPLOCHROMINE "SPECIES FLOCK" OF LAKE MALAWI A MONOPHYLETIC GROUP? •••••••••••••••••••••••••••••••••••••••• 102 CHAPTER 4. CHARACTERIZATION AND PHYLOGENY OF THE CYRTOCARA LIVINGSTON!! SPECIES-GROUP FROM LAKE MALAWI, INCLUDING A NEW SPECIES WITH HYPERTROPHIED LIPS ........................ 125 CHAPTER 5. THREE NEW HAPLOCHROMINE CICHLIDS WITH THREE LATERAL SPOTS FROM LAKE MALAWI, AND A KEY TO THE SPECIES OF THE THREE-SPOT ASSEMBLAGE •••••••••••••••••••••••••••••••••••••••• 189 APPENDIX 1. FLOATING ISLANDS: A MEANS OF FISH DISPERSAL IN LAKE MALAWI, AFRICA ••••••••••••••••••••••••••••••••••••••• 280 APPENDIX 2. USE OF BEAD TAGS TO STUDY LONG-TERM TERRITORIALITY AMONG CICHLID FISHES OF ROCKY SHORES IN LAKE MALAWI •••••••••••••••• 299 LITERATURE CITED •••••••••••••••••••••••••••••••••••••••••••••••••• 313 v LIST OF FIGURES Figure 1. Heterochromis multidens ••••••••••••••••••••••••••••••• 39 Figure 2. Cladogram summarizing information on character distribution among higher-level groups of cichlids .... 41 Figure 3. Dorsal gill-arch elements in Heterochromis multidens ....................................................... 43 Figure 4. Dorsal view of interorbital region of head of Heterochromis multidens after removal of skin ......... 46 Figure 5. Infraorbital bones of some cichlids .................. 48 Figure 6. Palatine-ectopterygoid region of Heterochromis multidens ••••••••••••••••••••••••••••••• 55 Figure 7. Suspensorium and jaws of Heterochromis mul t idens • • • • . • • . • . • • • . • • . • • • • . • • . • • . • • • • • • • . • . 57 Figure 8. Scales of Heterochromis multidens ••••••••••••••••••••• 60 Figure 9. Posterior part of dorsal fin of young Heterochromis multidens . • • . • . • . • . • . • . 62 Figure 10. Caudal skeletons of some cichlids .................... 64 Figure 11. Caudal skeletons of Astatotilapia calliptera, showing individual variation •••••••••••••••••••••••••• 66 Figure 12. Neurocranium of Heterochromis multidens .............. 68 Figure 13. Head of Heterochromis multidens after removal of skin, eyes, infraorbitals, and nasals ...... 70 Figure 14. Ventral gill-arch elements of Heterochromis multidens ••••••••••••••••••••••••••••••••••••••••••••• 72 Figure 15. Infraorbitals of Heterochromis multidens •••••••••••••• 75 Figure 16. Neurocranial apophysis for upper pharyngeal bones in Heterochromis multidens •••••••••••••••••••••••••••• 77 Figure 17. Hyoid bar and urohyal of Heterochromis multidens ...... 79 Figure 18. Pelvic basipterygium of Heterochromis multidens ....... 81 Figure 19. Pectoral girdle of Heterochromis multidens ............ 83 Figure 20. Geographic distribution of Heterochromis multidens .... 85 Figure 21. Anal fins of some adult male haplochromines with true ocellae .......................................... 115 vi Figure 22. Anal fins of some adult male haplochromines with nonocellate spots ..................................... 117 Figure 23. Thoracic-abdominal scale transition in some non-Malawian haplochromines ••••••••••••••••••••••••••• 119 Figure 24. Thoracic-abdominal scale transition in ocellate Malawian haplochromines ••••••••••••••••••••••••••••••• 121 Figure 25. Live individuals of Cyrtocara milomo, new species . • . • . • . • . • . 162 Figure 26. Live individuals of Cyrtocara iohnstoni, ~. venustus, and~. livingstonii •••••••••••••••••••••• 164 Figure 27. Live individuals of Cyrtocara polystigma, ~. linni, and~. species A •••••••••••••••••••••••••••• 166 Figure 28. Cyrtocara fuscotaeniatus holotype, after Regan •••••••• 168 Figure 29. Variation in color pattern in Cyrtocara iohnstoni, ~. milomo, and~. fuscotaeniatus •••••••••••••••••••••• 170 Figure 30. Variation in color pattern in Cyrtocara yenustus, ~. livingstonii, ~. linni, and~. species A ••••••••••• 172 Figure 31. Variation in color pattern in Cyrtocara polystigm.a •••••••••••.••.•.•••.••••••.•.•••••••••.••.• 174 Figure 32. Neurocranium of Cyrtocara milomo •••••••••••••••••••••• 176 Figure 33. Suspensorium in species of the Cyrtocara liyingstonii species-group and in~. fuscotaeniatus •••••••••••••••• 178 Figure 34. Upper jaw in species of the Cyrtocara livingstonii species-group and in~. fuscotaeniatus •••••••••••••••• 180 Figure 35. Lower pharyngeal bones of Cyrtocara 1ohnstoni, ~. milomo, and~. yenustus •••••••••••••••••••••••••••• 182 Figure 36. Dorsal gill-arch elements in Cyrtocara venustus ••••••• 184 Figure 37. Cladogram of the Cyrtocara livingstonii species-group ....................................................... 186 Figure 38. Diagram of the lower pharyngeal bone of a cichlid, showing counts and measurements ••••••••••••••••••••••• 246 Figure 39. Terminology of spots in "three-spotted" haplochromines ••••••••••••••.•••.••••••.••••.••••••••• 248 Figure 40. Live holotype of Cyrtocara lithobates, new species ••••••••••••••••••••••••••••••••••••••••••• 250 Figure 41. Holotype of Cyrtocara lithobates, new species ......... 252 vii --~~~~-~----- -~--~- Figure 42. Live paratype of Cyrtocara lithobates, new species • . • . • . • . 254 Figure 43. Lower pharyngeal bone of Cyrtocara lithobates ......... 256 Figure 44. Suspensorium and jaws of Cyrtocara lithobates ......... 258 Figure