A PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSIS OF CYPRINODONTIFORM FISHES (TELEOSTEI, ATHERINOMORPHA) LYNNE R. PARENT] BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 168 : ARTICLE 1 NEW YORK : 1981 A PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSIS OF CYPRINODONTIFORM FISHES (TELEOSTEI, ATHERINOMORPHA) LYNNE R. PARENTI Department of Ichthyology American Museum of Natural History SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN THE FACULTY IN BIOLOGY OF THE CITY UNIVERSITY OF NEW YORK BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 168 : ARTICLE 4 NEW YORK : 1981 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 168, article 4, pages 335-557, figures 1-99, tables 1-3 Issued September 3, 1981 Price: $14.35 a copy ISSN 0003-0090 This article completes Volume 168. Copyright © American Museum of Natural History 1981 CONTENTS Abstract 341 Introduction 341 Acknowledgments 344 Methods 346 Overview of Past Internal Classifications of Cyprinodontiform Fishes 349 Derived Characters of Cyprinodontiforms 354 Phylogenetic Analysis 365 Aplocheiloids (Group B) 367 Neotropical Aplocheiloids 375 Cladistic Summary of Neotropical Aplocheiloids 385 Old World Aplocheiloids 386 The Aphyosemion-Nothobranchius Group 389 The Aplocheilus-Pachypanchax-Epiplatys Group 394 Cladistic Summary of Old World Aplocheiloids 396 Cyprinodontoids (Group C) 397 Relationships of the Cyprinodontoids 404 Internal Fertilization and Viviparity 429 Cladistic Summary of the Cyprinodontoids 443 Group D 443 Fundulines 443 Group E 447 Group F 447 Group H 447 Jenynsia, Anableps, and Oxyzygonectes 448 Poeciliids, Fluviphylax, Pantanodon, and the Procatopines 450 Group G ,454 Empetrichthys, Crenichthys, and the Goodeidae 454 Group I 457 Cubanichthys and Chriopeoides 457 Group J 457 Orestias and the Anatolian Cyprinodontines 457 New World Cyprinodontines 458 Classification 461 Key to Genera and Suprageneric Categories 465 Systematic Accounts 470 Order Cyprinodontiformes 470 Suborder Aplocheiloidei 471 Family Aplocheilidae 471 Genus Aplocheilus 471 Genus Pachypanchax 473 Genus Epiplatys 474 Subgenus Lycocyprinus 475 Subgenus Parepiplatys 475 Subgenus Pseudepiplatys 475 Subgenus Aphyoplatys 475 Genus Aphyosemion 476 Subgenus Archiaphyosemion 477 Subgenus Chromaphyosemion 477 Subgenus Diapteron 477 Subgenus Kathetys 477 Subgenus Mesoaphyosemion 477 Genus Fundulopanchax 477 Subgenus Paludopanchax 479 Subgenus Paraphyosemion 479 Subgenus Gularopanchax 479 Subgenus Callopanchax 479 Subgenus Raddaella 479 Genus Adamas 479 Genus Nothobranchius 479 Family Rivulidae 481 Genus Rivulus 481 Genus "Rivulus" 483 Genus Trigonectes 484 Genus Pterolebias 485 Genus Rachovia 486 Genus Austrofundulus 487 Genus Neofundulus 489 Genus "Neofundulus" 489 Genus Cynolebias 490 Suborder Cyprinodontoidei 491 Section 1 492 Family Profundulidae 492 Genus Profundulus 492 Section 2 493 Division 1 493 Family Fundulidae 493 Genus Plancterus 493 Genus Fundulus 494 Genus Lucania 496 Genus Leptolucania 497 Genus Adinia 498 Division 2 499 Sept 1 499 Family Valenciidae, New Family 499 Genus Valencia 500 Sept 2 501 Superfamily Poecilioidea 501 Family Anablepidae 501 Subfamily Anablepinae 501 Genus Anableps 501 Genus Jenynsia 503 Subfamily Oxyzygonectinae, New Subfamily 504 Genus Oxyzygonectes 504 Family Poeciliidae 505 Subfamily Poeciliinae 505 Subfamily Aplocheilichthyinae 507 Genus Aplocheilichthys 507 Subgenus Micropanchax 508 Subgenus Lacustricola 508 Subgenus Poropanchax 508 Subgenus Congopanchax 508 Genus Lamprichthys 508 Genus Pantanodon 509 Genus "Aplocheilichthys" 510 Genus Procatopus 511 Genus Cynopanchax 513 Genus Plataplochilus 513 Genus Hypsopanchax 513 Subfamily Fluviphylacinae 514 Genus Fluviphylax 514 Superfamily Cyprinodontoidea 515 Family Goodeidae 515 Subfamily Empetrichthyinae 516 Genus Empetrichthys 516 Genus Crenichthys 517 Subfamily Goodeinae 518 Family Cyprinodontidae 519 Subfamily Cubanichthyinae, New Subfamily 519 Genus Cubanichthys 520 Subfamily Cyprinodontinae 521 Tribe Orestiini 521 Genus Aphanius 521 Genus "Aphanius" 522 Genus Kosswigichthys 524 Genus Orestias 525 Tribe Cyprinodontini 526 Genus Cyprinodon 526 Genus Jordanella 528 Genus Cualac 529 Genus Floridichthys 530 Genus Megupsilon 531 Historical Biogeography 533 Summary 544 Literature Cited 547 ABSTRACT The cyprinodontiforms, or killifishes, are a forms; (3) each of the four viviparous families is large and diverse group of 900 fresh- and brackish- monophyletic; however, their previous definitions water species with a pantropical and temperate in terms of uniquely derived characters have been Laurasian distribution. Traditionally, it has been altered; (4) the development of an annual habit, classified in five families: the worldwide, ovipa- exhibited by members of the aplocheiloid killi- rous Cyprinodontidae, and four New World vi- fishes and possibly some cyprinodontoids, in- viparous families: the Poeciliidae, Anablepidae, cludes derived reproductive traits exhibited to Jenynsiidae, and Goodeidae. Fishes of the diverse some degree by all killifishes; therefore, the an- Cyprinodontidae, in turn, have been divided into nual habit does not define a monophyletic group as many as eight subfamilies. of killifishes; (5) similarly, viviparity is not hy- The objectives of the present study are to: (1) pothesized to be a uniquely derived character, but determine if the cyprinodontiform fishes as a has apparently arisen at least three times within whole form a monophyletic group; (2) determine the group; and (6) the interrelationships of cy- if each of the five families is monophyletic; (3) prinodontiforms correspond, in part, with a pat- define the major subgroups of cyprinodontiforms, tern of the break-up of Pangea, except for an An- concentrating on the genera of the Cyprinodon- dean-Eurasian sister group pair. tidae; (4) determine the interrelationships of the A scheme of interrelationships of cyprinodon- subgroups; (5) present a comprehensive classifi- tiforms as well as of monophyletic subgroups is cation of the cyprinodontiforms that reflects the presented in the form of cladograms, of which the interrelationships; and (6) provide a hypothesis former is transformed into a comprehensive clas- for the distribution of the group. sification of the group. The fishes under study are The following general results were obtained by recognized as comprising the order Cyprinodon- using the methods of phylogenetic systematics tiformes Berg and divided into two suborders, the and vicariance biogeography: (1) the cyprinodon- Aplocheiloidei (which previously comprised, in tiforms are considered to be monophyletic by part, the Cyprinodontidae), and the Cyprinodon- their sharing derived characters of the caudal toidei (comprising all other cyprinodontiforms as skeleton, upper jaw, gill arches, position of the well as the four viviparous families). In order to first pleura! rib, pectoral girdle, and aspects of minimize the number of named empty categories, breeding and development; (2) the family Cyprin- a numbering system is incorporated into a tradi- odontidae is nonmonophyletic as it contains some tional naming system to create the new classifi- of the most primitive and derived cyprinodonti- cation. INTRODUCTION The cyprinodontiforms, commonly known nominal species in approximately 80 nominal as killifishes, top minnows, or toothcarps, are genera. Included are the popular aquarium a large and diverse group of teleostean fishes fishes, including the annual killifishes of trop- distributed nearly worldwide in temperate ical South America and Africa of the subfam- and tropical freshwaters (fig. 1), with some ily Rivulinae (Myers, 1955), and widely used members regularly entering brackish water. experimental fish such as those of the genus The term cyprinodontiforms as used in this Fundulus. paper refers to fishes of the five families of Of the four viviparous families, two, the the superfamily Cyprinodontoidea, order Mexican, Central American, and northern Atheriniformes (Rosen, 1964). These are the South American Anablepidae, and the south- cosmopolitan and oviparous Cyprinodonti- eastern South American Jenynsiidae, con- dae, and four New World viviparous fami- tain just one genus each with several species. lies, the Anablepidae, Goodeidae, Jenynsi- The Goodeidae are diverse, comprising ap- idae, and Poeciliidae. proximately 35 species in 16 genera, all of The Cyprinodontidae are the largest and which are restricted to the Mexican Plateau most diverse family containing over 650 (Miller and Fitzsimons, 1971). The Neotrop- 342 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 FIG. 1. Present day distribution of cyprinodontiforms. Dotted and dashed line approximates Wal- lace's Line. ical and temperate Poeciliidae comprise ap- relatively primitive. He included as evidence proximately 200 species in 19 genera (Rosen for this distinction the fact that in esocoids and Bailey, 1963; Rosen, 1979), and the guppy the maxilla enters the gape, whereas the (Poecilia reticulata) and mosquito-fish maxilla is excluded from the gape in killifish- (Gambusia affinis) are included. es (including the adrianichthyoids) and cave- Knowledge of the relationships of the cy- fishes. Regan (1909) separated the last two prinodontiforms to other fishes has advanced groups from the rest of the Haplomi and con- considerably, whereas the proposed interre- structed for them a new order Microcyprini. lationships of cyprinodontiforms has pro- This action was supported by Hubbs gressed little since Carman's (1895) outline