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Home , Adrianichthys, Anablepidae, Anableps anableps, Aphyosemion, Aplocheilidae, Aplocheiloidei

A PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSIS OF CYPRINODONTIFORM (TELEOSTEI, )

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 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 - Group 389 The --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, , , and the Procatopines 450 Group G ,454 , , and the 454 Group I 457 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 470 470 Suborder 471 471 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 477 Subgenus Paludopanchax 479 Subgenus Paraphyosemion 479 Subgenus Gularopanchax 479 Subgenus Callopanchax 479 Subgenus Raddaella 479 Genus Adamas 479 Genus Nothobranchius 479 Family 481 Genus 481 Genus "Rivulus" 483 Genus Trigonectes 484 Genus Pterolebias 485 Genus 486 Genus 487 Genus Neofundulus 489 Genus "Neofundulus" 489 Genus Cynolebias 490 Suborder 491 Section 1 492 Family 492 Genus 492 Section 2 493 Division 1 493 Family 493 Genus Plancterus 493 Genus 494 Genus 496 Genus Leptolucania 497 Genus Adinia 498 Division 2 499 Sept 1 499 Family Valenciidae, New Family 499 Genus 500 Sept 2 501 Superfamily Poecilioidea 501 Family 501 Subfamily Anablepinae 501 Genus Anableps 501 Genus Jenynsia 503 Subfamily Oxyzygonectinae, New Subfamily 504 Genus Oxyzygonectes 504 Family 505 Subfamily 505 Subfamily Aplocheilichthyinae 507 Genus Aplocheilichthys 507 Subgenus 508 Subgenus 508 Subgenus 508 Subgenus Congopanchax 508 Genus Lamprichthys 508 Genus Pantanodon 509 Genus "Aplocheilichthys" 510 Genus 511 Genus Cynopanchax 513 Genus 513 Genus 513 Subfamily Fluviphylacinae 514 Genus Fluviphylax 514 Superfamily Cyprinodontoidea 515 Family Goodeidae 515 Subfamily 516 Genus Empetrichthys 516 Genus Crenichthys 517 Subfamily 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 , 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 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 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 (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 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 the maxilla enters the gape, whereas the (Poecilia reticulata) and -fish maxilla is excluded from the gape in - (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 (1919) who reported that the Microcyprini of the major subgroups. (including the phallostethoids after Regan, Killifishes are typically soft-rayed, and, as 1913) have a derived branchiostegal number such, have historically been aligned with the and arrangement, comparable to those of the more primitive teleost groups. Gill (1874) acanthopterygians; whereas, the Haplomi, aligned the cyprinodontiforms with the esoc- sensu Regan, are primitive in this regard. oids, which together comprised the order Myers (1928a) removed the phallostethoid Haplomi. Starks (1904) divided the Haplomi fishes from the Microcyprini, and suggested into three suborders: the Esocoidei (includ- their close relationship to the Atherinidae, ing the mud-minnow Umbra, and the pike then in the order Percesoces. Esox), the Amblyopsioidei (the cavefishes), The alignment of the amblyopsoids with and the Poecilioidei (=Cyprinodontoidei of the cyprinodontiforms and adrianichthyoids Rosen, 1964). Yet, he admitted there were was never more than tentative; yet it re- no important [unique] characters which de- mained unchallenged until Rosen (1962) re- fined the order. moved the cavefishes from the Microcyprini, Regan (1911) remarked on the killifish and referred to as the Cyprinodontiformes fol- cavefish relationship to more derived teleost lowing Berg (1940), and placed them in the groups while noting that the esocoids were newly created Amblyopsiformes which he 1981 PARENTI: CYPRINODONTIFORM FISHES 343 claimed was more closely related to the Per- TABLE 1 copsiformes. The cyprinodontiforms and ad- Classification of Fishes of the Order rianichthyoids remained as the sole constit- Atheriniformes uents of the order Cyprinodontiformes. (Rosen, 1964) Gosline (1963) continued to support the Superorder naturalness of the order Microcyprini, and Series Atherinomorpha criticized Rosen (1962) for separating the or- Order Atheriniformes der into two groups while giving no hint as Suborder to the placement of the Cyprinodontiformes Superfamily Atherinoidea in a higher classification of teleost fishes. Superfamily Phallostethoidea An answer to this was provided by Rosen Suborder Cyprinodontoidei (1964) when he created the order Atherini- Superfamily Adrianichthyoidea formes to include the cyprinodontiforms, ad- Superfamily Cyprinodontoidea Family Cyprinodontidae rianichthyoids, atherinoids, phallostethoids, Family Anablepidae exocoetoids, and scomberesocoids. Rosen's Family Jenynsiidae (1964) classification is summarized in table 1. Family Goodeidae The alignment of these fishes had casually Family Poeciliidae been suggested earlier by several workers, Suborder Exocoetoidei although, this was done with little formal Superfamily Exocoetoidea taxonomic treatment. Cope (1870) first re- Superfamily Scomberesocoidea marked on the possible close relationship of Series atherinids and cyprinodonts. In addition, Myers (1928a) commented that the structure of the ethmoid region and the mouth sug- Patterson and Rosen (1977) and Rosen and gested the affinity of cyprinodontiforms and Parent! (MS), and is supported by derived members of the Percesoces. Furthermore, features of the gill arches and the jaws and Regan (1911, p. 321), commenting on the jaw suspensorium. possible alignment of his new order, said: Thus, with increased knowledge of inter- "Whereas the Haplomi show relationship to relationships of teleosts, cyprinodontiforms the most generalized isospondylous fishes, have progressed from a primary alignment the Microcyprini bear more resemblance to with the primitive esocoids to a hypothesized the Salmopercae and Synentognathi, espe- close relationship with the advanced perco- cially the latter." morph fishes. The monophyly (in the sense of Hennig, Yet, as stated previously, our knowledge 1966) of the order Atheriniformes, and the of the interrelationships of members of the monophyly and interrelationships of its superfamily Cyprinodontoidea has under- subgroups were not rigorously defined by gone little comparable progress. Relation- Rosen (1964). However, recent evidence in- ships among the" families and among the in- dicates that the Atheriniformes is monophy- cluded genera, have been presented as letic, and problems of its higher order inter- speculation. Workers have either dealt with relationships may easily be summarized the primary groups of oviparous cyprinodon- (Rosen and Parent!, MS). tiforms alone (e.g., Myers, 1931, 1955; Sethi, Rosen (1964, p. 260) suggested that the 1960; Uyeno and Miller, 1962), or one of the atherinomorph fishes: "arose from a group four viviparous families (e.g., Rosen and that stood somewhere in the ancestry of the Bailey, 1963; Hubbs and Turner, 1939; Mil- order Perciformes." This point, which may ler, 1979), never more than casually discuss- be restated as fishes of the Atherinomorpha ing the relationship of one family to another and Percomorpha share a common ancestor, or to a group of oviparous cyprinodontids. was reiterated in the classifications of Green- However, aside from discussions and re- wood, Rosen, Weitzman and Myers (1966), peated speculation on the affinity of one Rosen and Patterson (1969), Rosen (1973a), group of killifishes to another, there has been 344 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 no formal treatment of the interrelationships Fund, American Museum of Natural Histo- of the five families, no statement supporting ry, the City University of New York and the or refuting the monophyly of each of the five Department of Ichthyology, Acad- families, and no formal definition of the su- emy of Sciences. perfamily Cyprinodontoidea. For loan of specimens and accommoda- Knowledge of such interrelationships tions during visits, I particularly thank Drs. could serve as a basis for biogeographic hy- William N. Eschmeyer and Tomio Iwamoto, potheses concerning the history of the pan- Ms. Pearl M. Sonoda, Mr. William C. Ruark tropical and temperate Laurasian regions, and Mr. James Gordon (California Academy could be an invaluable reference for research of Sciences, San Francisco), as well as Dr. scientists and aquarists alike, and form the James Bohlke and Mr. William Saul (Acad- framework for an understanding of the vari- emy of Natural Sciences, Philadelphia), Dr. ety of reproductive modes found within the P. Humphry Greenwood, Mr. James Cham- group. bers and Mr. Gordon Howes (British Mu- Thus, the objectives of this study are to: seum [Natural History], London), Dr. Rob- (1) determine the monophyly of the super- ert K. Johnson and Mr. Garrett S. Glodek family Cyprinodontoidea; (2) determine the (Field Museum of Natural History, Chica- monophyly of each of the five families; (3) go), Dr. Gianna Arbocco (Museo Civico di define the major subgroups of cyprinodonti- Storia Naturale, Genova), Dr. William L. forms, with a concentration on the genera of Fink and Mr. Karsten Hartel (Museum of the family Cyprinodontidae; (4) determine Comparative , Harvard University, the interrelationships of the subgroups; (5) Cambridge), Dr. Robert R. Miller (Univer- present a comprehensive classification of the sity of Michigan, Museum of Zoology, Ann cyprinodontiforms which reflects the inter- Arbor), Drs. Richard P. Vari and Stanley H. relationships; and (6) provide a hypothesis Weitzman (National Museum of Natural His- for the distribution of the group. tory, Washington, D.C.), and Dr. Raul Vaz- Ferreira (Uruguay). For the gift of live or preserved specimens, ACKNOWLEDGMENTS I thank Mr. John S. Brill, Jr., Dr. A. Casi- I thank Dr. Donn E. Rosen for his advice nos, Mr. Thomas Coon, Mr. J. Dry an, Mr. and encouragement throughout the course of Daniel Fromm, Mr. Tim Hardin, Mr. Leon- this study as well as directing my interests ard Mackowiak and Mr. Jerry Shapiro. toward many of the problems in cyprinodon- For access to a text-editor and computing tiform systematics. I also thank Dr. James funds for preparation of the typescript, I S. Farris, and especially Dr. M. F. Micke- thank Drs. Robert F. Rockwell, the City Col- vich for their early encouragement and sup- lege, and Leslie F. Marcus, Queens College, port of my systematic endeavors. both of the City University of New York. For helpful discussions of aspects of the For information on systematic literature, research in progress, I thank Mr. Guido Din- as well as personal observations, I thank Dr. gerkus, Dr. Mickevich, Ms. Nancy A. Neff James W. Atz, Mr. Fromm, Drs. Kenneth J. and Dr. Richard P. Vari. Ms. M. Norma Lazara, F. Douglas Martin, Mr. Eric Quin- Feinberg provided invaluable technical ad- ter, Drs. Alfred C. Radda, Jamie Thomer- vice and assistance for which I am most son, Bruce J. Turner, Stanley H. Weitzman, grateful. E. O. Wiley and John P. Wourms. Financial and logistical support was pro- VERNACULAR NAMES: In a systematic vided primarily by the Department of Ich- study that ends with a reclassification, names thyology, American Museum of Natural His- must be used in the discussion of interrela- tory for which I extend sincere thanks. tionships and character distributions which Additional support was provided by a grant are at once familiar to most workers on the from the Theodore Roosevelt Memorial group, and which unambiguously refer to a 1981 PARENTI: CYPRINODONTIFORM FISHES 345 given group of genera or families. Thus, I TABLE 2 use the following vernacular names through- Current Comprehensive Classification of the out the text. Cyprinodontid Fishes The term acanthopterygian refers to fishes Family Cyprinodontidae of the superorder Acanthopterygii, which in- Subfamily Fundulinae cludes the two series Atherinomorpha and Genus Fundulus, Lucania, Leptolucania, Oxy- Percomorpha (Rosen, 1973a). The series are zygonectes, Cubanichthys, Chriopeoides, Val- termed atherinomorph and percomorph, re- encia, Empetrichthys, Crenichthys, Profundulus, spectively. Hubbsichthys,11 Adinia The series Atherinomorpha contains a sole Subfamily Cyprinodontinae order, the Atheriniformes. Thus, the two cat- Genus Cyprinodon, Megupsilon, Floridichthys, egories are equivalent, and the term atheri- Jordanella, Cualac, Aphanius, Tellia, Kosswig- nomorph describes the membership of both. ichthys, Anatolichthys Subfamily Lamprichthyinae Atheriniform is therefore not used herein to Genus Lamprichthys avoid confusion. Subfamily Orestiatinae Within the order Atheriniformes, vernac- Genus Orestias ular names are used for the major subdivi- Subfamily Pantanodontinae sions listed in the classification of table 1. Genus Pantanodon Fishes of the suborder Atherinoidei are re- Subfamily ferred to as the silversides. The terms ath- Genus Aplocheilichthys, Procatopus, Hypsopan- erinoid and phallostethoid are reserved for chax, Micropanchax, Cynopanchax, Plataplo- members of the superfamilies Atherinoidea chilus, Platypanchax, Hylopanchax, Congopan- and Phallostethoidea, respectively. chax, Poropanchax Subfamily Rivulinae There is no vernacular reference for the Genus Rivulus, Trigonectes, Rivulichthys, Ptero- suborder Cyprinodontoidei of Rosen; com- lebias, Rachovia, Austrofundulus, Terranotus, ponents are referred to separately. The term Cynolebias, , , adrianichthyoid refers to fishes of the super- Simpsonichthys, Aphyosemion, Nothobranchius, family Adrianichthyoidea. The fishes of the Adamas, Epiplalys, Aplocheilus, Pachypanchax, superfamily Cyprinodontoidea, the subject Fundulosoma, Callopanchax of this revision, are referred to alternately as Subfamily Fluviphylacinae the cyprinodontiforms, cyprinodonts, or kil- Genus Fluviphylax lifishes. They are reclassified in this study as " Schultz (1949) described Hubbsichthys laurae, new the order Cyprinodontiformes. Two subor- genus and species of cyprinodontid. The holotype ders are named, the Aplocheiloidei, com- (USNM 120999), the only recorded specimen, was ex- prising those fishes of the Rivulinae, and the amined and determined as a female poeciliid, and most Cyprinodontoidei, comprising all other cy- likely of the species Poecilia caucana (Steindachner). prinodontiforms. These groups will be re- I propose Hubbsichthys be dropped from the subfamily ferred to as the aplocheiloids and cyprino- and placed in synonymy of the genus Poecilia. dontoids, respectively. There is no vernacular reference for the killifishes, whereas the four remaining fami- suborder Exocoetoidei, and its two super- lies are collectively referred to as the vivipa- families are referred to as the exocoetoids rous killifishes. (superfamily Exocoetoidea) and the scom- Vernacular names for the various groups beresocoids (superfamily Scomberesocoi- found within the Cyprinodontidae follow the dea). current classification of the family, listed in Within the Cyprinodontoidea, members of table 2. the five families are normally referred to as The subfamily Cyprinodontinae is referred the cyprinodontid, anablepid, jenynsiid, to as the cyprinodontines, which are further goodeid or poeciliid fishes. The Cyprinodon- divided into the New World cyprinodon- tidae are also referenced as the oviparous tines, comprising Cyprinodon and its imme- 346 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 diate relatives, and the Anatolian or Old which are no longer considered to be mem- World cyprinodontines, comprising Aphan- bers of a monophyletic group containing Fun- ius and its immediate relatives. dulus, are referred to by their formal generic The subfamilies Fluviphylacinae, Orestiat- names (e.g., Oxyzygonectes, Cubanichthys, inae, and Pantanodontinae are referred to as Chriopeoides, Empetrichthys, Crenichthys, the genera Fluviphylax, Orestias, and Pan- and Profundulus). The term funduline refers tanodon, respectively. The Procatopodinae to Fundulus, Adinia, Leptolucania, Lucania and Lamprichthyinae are collectively re- and their nominal subgenera. ferred to as the procatopines. Various other groups of teleosts are dis- Members of the subfamily Fundulinae, cussed using conventional terminology.

METHODS PHYLOGENETIC ANALYSIS The method of phylogenetic analysis neric level, genera that are not monophyletic adopted here is that put forth formally by may have groups of species assignable to Hennig (1950, 1966), alternately referred to other monophyletic groups, leaving the re- as cladistics, cladism, or phylogenetic sys- mainder as a paraphyletic assemblage at the tematics. Within a cladistic scheme, taxa are most plesiomorph position of the more inclu- grouped hierarchically on the basis of their sive monophyletic group. In these cases, the sharing derived characters (termed synapo- traditional generic name will be retained, and morphies), rather than on their overall simi- recommendations for a species-level revision larity. This method of analysis is preferred will be made. over those of evolutionary (e.g., When character conflicts occur at any Mayr, 1969, 1974; Simpson, 1961) and phe- level in the analysis, the principle of parsi- netics (e.g., Sokal and Sneath, 1963; Sneath mony is invoked to choose among alternative and Sokal, 1973) if the goal is the hierarchical explanations of the data. The assumption is grouping of taxa based solely on a hypothesis not made that evolution always, or ever, of common ancestry. Given the assumption must proceed along a parsimonious course; that nature is structured hierarchically, a however, it is concluded that our explanation cladogram that reflects increasing levels of of a hypothesized phylogeny should be the generality of character distributions is con- most parsimonious one since, by definition, cluded to be the best estimate of the one, it is the one which requires that we invoke true phylogeny. the fewest assumptions about character Recognized taxa are those which can be transformations. Similarly, characters are defined as monophyletic groups in the sense not weighted in the analysis since no objec- of Hennig. That is, a monophyletic group tive criteria for weighting could be deter- contains all the descendants, and only the mined. descendants of a common ancestor. Mono- Character transformation series are con- phyletic groups, therefore, are defined by structed among states of homologous char- their members sharing derived characters. acters. Characters are hypothesized to be Such groups are assembled into more inclu- homologous if they are comparable in shape sive monophyletic groups until a hierarchical and position, or present in different forms, arrangement of all the members is achieved. but exhibiting the same ontogenetic se- It is not the intention of a cladistic analysis quence. A homology, therefore, is, at one to recognize paraphyletic groups. However, level, comparable to a derived character or since this study is done primarily at the ge- apomorphy (Wiley, 1975). 1981 PARENTI: CYPRINODONTIFORM FISHES 347

The polarity of a transformation series is readily be transformed into a cladogram of initially determined by comparison to an out- areas occupied by monophyletic groups (Ro- group (as discussed by Lundberg, 1972), or sen, 1978). A pattern of earth history is sug- by comparison with an ontogenetic transfor- gested by the interrelationships of the areas. mation (as discussed in Nelson, 1973). The generality of this pattern and those of In the latter procedure, an ontogenetic monophyletic groups will be tested by com- change in one of two taxa that are hypothe- parison to other established patterns as well sized to share a common ancestor, hence as to each other. termed sister groups, must logically be con- DISPOSITION OF SPECIMENS AND COLLEC- sidered derived if the principle of parsimony TION OF DATA: Counterstained specimens of is applied. That is, one need only make the cyprinodontiforms were prepared according assumption that the transformation was to the alcian blue-alizarin Red S method of gained by one taxon, rather than the as- Dingerkus and Uhler (1977) to facilitate the sumptions that the character was present in examination of cartilage as well as bone. the common ancestor, and that it was sub- When possible, at least two males and two sequently lost in the other. females of several species in a genus were In the former procedure, a character state prepared. In some cases, just one pair was is analyzed as being primitive or derived by prepared. When lots were only large enough comparing it to the state within other groups for the preparation of one specimen a male of atherinomorph fishes, within the perco- was chosen since cyprinodontiforms are morphs, or to the teleosts as a whole. Char- markedly sexually dimorphic with males typ- acters or character complexes recognized at ically exhibiting a greater degree of variation once as being unique are analyzed as de- than females. rived. The general state of a character in an Additional specimens, which were cleared outgroup or in the cyprinodontiforms is ini- and solely alizarin-stained, were available tially assessed as primitive. However, a from the collection of fishes in the Depart- character which may be described in the ment of Ichthyology, AMNH. same manner as that of the general state may Radiographs were prepared primarily of be termed secondarily derived within a group species represented only by the material of cyprinodontiforms if this interpretation is in order to facilitate a cursory examination consistent with the most parsimonious inter- of the osteological details. pretation of all the data. That is, the polarity Anatomical illustrations were prepared of a transformation series is not always de- from sketches of structures as viewed termined by the constraint that the general through a camera lucida mounted on a dis- state represents the primitive condition. secting microscope. Primarily, dissected Transformation series treated in this manner cleared and stained material was used for are discussed in detail. this purpose; however, alcohol specimens BIOGEOGRAPHIC ANALYSIS: A hypothesis were partially dissected when necessary. of the historical distribution of cyprinodonti- Most illustrations and descriptions of states form fishes is constructed upon completion of cartilaginous elements are of counter- of the phylogenetic analysis. The distribution stained preparations. of monophyletic groups should reflect the Developmental series of available aquari- history of the areas of distribution if we ac- um representatives of several genera of ath- cept the premise inherent in the works of erinomorphs were bred and reared in the vi- Croizat (1958, 1964) that the world and its varium of the Department of Ichthyology, biota evolved together. This concept forms AMNH. Details of development, including the basis of vicariance biogeography as put structure of the egg, period of time from forth by Croizat, and Croizat, Nelson and spawning to hatching, and age at first - Rosen (1974), Platnick and Nelson (1978) and ing were observed for several genera. Adults Rosen (1976, 1978). were also observed for details of reproduc- The cladogram of cyprinodontiforms may tive behavior. All preserved aquarium spec- 348 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 imens were catalogued in the department's CAS, California Academy of Sciences, San Fran- collection. cisco Measurements and counts were made ac- FMNH, Field Museum of Natural History, Chi- cording to the procedure outlined by Miller cago (1948) for cyprinodont fishes, except as not- IU, Indiana University (now at California Acad- ed. As Miller pointed out, killifishes do not emy of Sciences, San Francisco) MCSN, Museo Civico di Storia Naturale, Genova possess a complete lateral line; therefore, it MCZ, Museum of Comparative Zoology, Cam- is customary to count scales in a lateral se- bridge ries starting from the shoulder girdle to the MNHN, Museum National d'Histoire Naturelle, end of the hypural plate, ascertained by Paris bending the caudal fin. SU, Stanford University (now at California Acad- Names for skeletal structures are those emy of Sciences, San Francisco) traditionally used in a description of teleost UMMZ, University of Michigan, Museum of Zo- anatomy, as updated by Patterson (1975). ology, Ann Arbor Details of the gonopodium of poeciliid fishes USNM, National Museum of Natural History, are described using the terminology of Rosen Washington, DC. ZVC, Zoologia Vertebrados, de la Facultad Cien- and Bailey (1963). cias, Montevideo, Uruguay Patterns of head scales and sensory pores and canals are described according to the ANATOMICAL conventions established by Hoedeman (1958) and Gosline (1949), respectively, to facilitate AC, anterior ceratohyal comparisons among the results of this and ALV, alveolar arm of premaxilla other studies. AMR, middle anal radial Estimates of number of species in groups APL, autopalatine APR, proximal anal radial currently classified in the Cyprinodontidae AR 1, anal ray 1 are from Lazara (1979) unless otherwise stat- ART, articular ed. ASC, ascending process of premaxilla Specimens examined and their catalog BOC, basioccipital numbers appear in the systematic section fol- BR, branchiostegal ray lowing each generic and family diagnosis. CL, cleithrum Catalog numbers followed by an asterisk COR, coracoid (*) indicate lots from which counterstained DEN, dentary specimens were prepared; those followed by DHH, dorsal hypohyal a cross (+) are lots from which solely aliza- DMX, dorsal process of maxilla DPR, proximal dorsal radial rin-stained preparations had been made. The DR1, first dorsal ray number of such specimens prepared is given E 1-4, epibranchial 1-4 in both cases as a fraction of the total spec- END, endopterygoid imens in the lot (e.g., 4 out of 20 is given as EP, epural 4/20). Catalog numbers with no designation EPL, epipleural rib are of alcohol lots from which no special EPO, epiotic preparations were made. EPO-PRO, epiotic processes EXO, exoccipital FRO, frontal ABBREVIATIONS HY 1-5, hypural 1-5 INSTITUTIONAL HYO, hyomandibula HYP, hypaxial musculature AMNH, American Museum of Natural History, ICARM, infracarnalis medius New York ICARP, infracarnalis posterior ANSP, Academy of Natural Sciences, Philadel- IF, inferior pharyngeals phia IH, interhyal BMNH, British Museum (Natural History), Lon- INCLA, inclinatores anales don IS, ischial process 1981 PARENTI: CYPRINODONTIFORM FISHES 349

K, kidney QUA, quadrate MAX, maxilla RAD, radials MDN, medial process of dentary RC, rostral cartilage MET, metapterygoid RET, retroarticular NA-1, neural arch 1 SAC, subautopalatine cartilage NL, nasal SCL, supracleithrum NS, neural spine SOC, supraoccipital PAR, parietal SOC-PRO, supraoccipital processes PAS, parasphenoid SPH, sphenotic PB 1-3, pharyngobranchial 1-3 SYM, symplectic PC, posterior ceratohyal T, testis PCL1, 3, postcleithrum 1, 3 TPB 1-4, pharyngobranchial toothplate 1-4 PHY, parhypural UB, urinary bladder PL, pleural rib UG, urogenital opening PMX, premaxilla UN, uroneural POP, preopercle UR, ureter PRO, prootic VHH, ventral hypohyal PSP, pseudophallus VMX, ventral process of maxilla PTT, posttemporal VO, vomer PU2, preural centrum 2

OVERVIEW OF PAST INTERNAL CLASSIFICATIONS OF CYPRINODONTIFORM FISHES

Cyprinodontiform fishes, as a whole or in goodeid fishes were included in the subfam- part, have been the subject of various revi- ily Cyprinodontinae, along with Neolebias. sionary studies, many of which have includ- The poeciliid fishes were the sole constitu- ed a formal reclassification. Together, these ents of the subfamilies Poeciliinae and Bel- works may be characterized as studies in onesocinae. Jenynsia and Anableps were recognition of diversity, rather than in elu- each placed in their own subfamilies, Jen- cidation of interrelationships. From Gar- ynsiinae and Anablepinae, respectively. man's (1895) summary of all members to Cyprinodontids were divided among the Sethi's (1960) discussion of primarily the Cyprinodontinae and the remaining three oviparous cyprinodontids, reclassifications subfamilies. The monotypic Orestiasinae have focused on the description of differ- contained the genus Orestias; the known cy- ences, rather than of the derived similarities, prinodontines constituted the remainder of among groups of cyprinodontiforms. The ma-, the Cyprinodontinae. jor classifications are summarized in table 3. Carman's Nothobranchiinae consisted of Garman attempted a synopsis of cyprino- two African genera, Haplochilichthys and dontiforms; however, he had included the the aplocheiloid Nothobranchius. The re- characin Neolebias and the cyprinoid Fun- mainder of the cyprinodontids, including the dulichthys in the group. As a result, his di- fundulines and South American aplochei- agnosis was general enough to apply to al- loids and Fundulichthys, together formed most any group of soft-rayed fishes with a the subfamily Haplochilinae. The name of single . Aside from these shortcom- the type genus, Haplochilus, was a correct- ings, however, Carman's summary of cy- ed spelling of Aplocheilus McClelland. This prinodontiform subgroups has remained little spelling change was not valid under the In- changed in subsequent reclassifications. ternational Code of Zoological Nomencla- Garman divided the Cyprinodontes Gill ture and was not used in subsequent revi- (1865) (=Cyprinodontoidea of Rosen, 1964) sions. However, the names Haplochilus and into eight subfamilies. The known genera of Haplochilichthys persist as identifications in ca 73

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many collections. The name Haplochilus has family consisted of the following characters: been used to refer to fishes in such genera as occipital condyles present on both basioccip- Epiplatys, Pachypanchax, Fundulus, Aphy- itals and exoccipitals, no modifications of the osemion, and Oxyzygonectes as well as anal fin into an intromittent organ, ovipa- Aplocheilus. Similarly, Haplochilichthys rous, premaxillaries distinct from maxillaries has been used to reference fishes in any of and protractile or not, and never more than the procatopine genera, not solely Aplochei- 65 scales in a lateral series. lichthys. The definition, however, is not consistent In Regan's (1911) reclassification of the with the distribution of characters among all order Microcyprini, the two subfamilies of cyprinodontids. More importantly, all the poeciliids were united into the subfamily characteristics are primitive, and are found Poeciliinae. He separated the goodeids from generally among teleost fishes. Thus, Myers the cyprinodontines and placed them in their (1931) effectively described the fishes of the own subfamily, the Characodontinae. Thus, family Cyprinodontidae as cyprinodonti- with Regan's work, the four viviparous fam- forms that lack the prominent specializations ilies were separated from the oviparous cy- of the viviparous groups. That is, no derived prinodontids. The precedent for treating [unique] characters of the family were given oviparous and viviparous cyprinodontiforms to unambiguously define it as a monophyletic separately in systematic revisions was estab- group. lished, and has remained virtually unchal- Myers (1931) divided the family into four lenged until the present study. subfamilies, and further divided the largest, The only concrete statements regarding the Fundulinae, into four tribes. the interrelationships of the five currently The tribe Fundulini was restricted to the recognized families were made by Hubbs North American fundulines and their pre- (1924) and Regan (1929), and these were in sumed relatives. The tribe Rivulini is coex- direct opposition. Hubbs grouped Jordan's tensive with the aplocheiloids as discussed Fitzroyidae (which he corrected to Jenynsi- throughout the present paper. idae) and Anablepidae together into the fam- Together, the tribe Aplocheilichthyini and ily Anablepidae, and Jordan's - subfamily Lamprichthyinae are coextensive tidae and Goodeidae into the family with the procatopines. Similarly, the subfam- Goodeidae. His action concerning the Good- ily Orestiatinae consisted solely of the genus eidae has remained unchallenged. However, Orestias. the grouping together of Anableps and Jen- Eigenmann (1920) suggested that the ynsia was rejected by Regan (1929) who North American Empetrichthys was the placed Anableps in the Poeciliidae. Myers closest relative of Orestias based on the fact (1931) effectively avoided the problem by re- that both lack pelvic fins and fin supports, verting to the placement of the two genera and have fleshy bases of the dorsal and anal in monotypic families. Most recently, Miller fins. As a result, the two genera constituted (1979) has criticized the inferred relationship the membership of the Orestiidae and Ores- of the two, and considered Anableps to be tiinae of Jordan (1923) and Hubbs (1924), re- more closely related to the Poeciliidae than spectively. Myers (1931), however, support- to any other group of cyprinodontiforms. ed the idea that Empetrichthys was more Since Hubbs s work, reclassification of cy- closely related to Fundulus, and placed it in prinodontiforms has focused on the division his tribe Fundulini where it has remained un- of the Cyprinodontidae (encompassing the til this study. Cyprinodontinae and Fundulinae) into a va- The final tribe, Aplocheilini Bleeker, con- riety of subgroups which have undergone el- sisted of a single genus, the evations or reductions in rank. which Myers and contemporaries referred to Myers (1931) treated the oviparous cyprin- as Aplocheilus. Fishes now commonly re- odontids and Oryzias. His definition of the ferred to the genus Aplocheilus were re- 352 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 ferred to as members of the genus Panchax, of these groups yet again. The oviparous cy- in the Rivulini. The confusion over the avail- prinodontids were classified in six families. able names for these genera, all of the Indo- Oryzias, included in the study, was also Malaysian region, was eliminated by Smith placed in its own family. The procatopines (1938) who demonstrated that Panchax was and Lamprichthys were grouped together in an objective of Aplocheilus, and the family Aplocheilichthyidae. that Oryzias was the proper name for the Aphanius and its allies were removed from ricefish. However, there was enough time for the subfamily Cyprinodontinae of Myers and the name Panchax to become established as placed in their own family, the Aphaniidae. a common name for most of the aplocheiloid Thus the Cyprinodontidae of Sethi consisted fishes, and it is still casually employed. solely of Cyprinodon and its New World rel- Earlier, Myers (1924c) pointed out that atives. Fundulichthys Bleeker, a name applied to a The Pantanodontinae was inexplicably specimen known only from an illustration, omitted from Sethi's study, as were other referred to a cyprinoid. unique cyprinodontid genera such as Oxy- Berg (1940) substituted the name Cyprino- zygonectes, Chriopeoides, Cubanichthys, dontiformes for the Microcyprini. He divid- and Rivulichthys. Also in 1970, Roberts cre- ed the order into two superfamilies, an ovip- ated yet another subfamily, the Fluviphyla- arous Cyprinodontoidea including the fami- cinae, to include a single genus and species, lies Cyprinodontidae and Adrianichthyidae, Fluviphylax pygmaeus (Myers and Carval- and a viviparous Poeciloidea, including the ho). This genus was also disregarded by Goodeidae, Poeciliidae, Jenynsiidae, and Sethi. Therefore, the most comprehensive Anablepidae. The adrianichthyid fishes, and also most widely accepted classification comprising the genera and of oviparous killifishes is that listed in table , were associated with the Cy- 2. The eight subfamilies are grouped together prinodontiformes since their transfer from in a single family, the Cyprinodontidae. the (=Exocoetoidei) by Weber The relationship of the subfamilies to each and de Beaufort (1922). However, because other and to the families of viviparous killi- of its poor representation in collections (the fishes has never been formally treated before monotypic Adrianichthys known only until this study. In fact, Sethi's grouping of the recently from the single holotype), the family Lamprichthyinae and procatopines repre- has gone virtually ignored in revisions. Ro- sented the only alignment of subgroups of sen (1964) placed it along with Oryzias, cyprinodontids since Garman (1895). Revi- which he elevated to family rank, and the sions of the viviparous killifishes have fo- Horaich thy idae, in the superfamily Adrian- cused on the interrelationships of included ichthyoidea. genera or species, but have presented no Myers (1955), again treating the oviparous more than informal remarks about the rela- killifishes and Oryzias, elevated each of his tionship of the considered family to another tribes of 1931 to subfamily rank. He ac- viviparous family or to the cyprinodontids. knowledged the correction of the use of the Hubbs and Turner (1939), in a revision of name Aplocheilus for Oryzias by elevating the Goodeidae, emphasized the structural the rank of the tribe Aplocheilini to the differences between the family and other cy- subfamily Oryziatinae. A new subfamily prinodontiforms. More recently, Miller and Pantanodontinae appeared to include the sin- Fitzsimons (1971) proposed several defining gle genus and species Pantanodon podoxys characters of the family (some of which are described in Myers (1955) by name only. An found among the oviparous cyprinodonts), eighth subfamily, the Lamprichthyinae, was and synonymies of several genera, yet made omitted from the list presumably inadver- no statement as to the relationship of the tently. goodeids to other cyprinodontiforms. The oviparous killifishes were treated Rosen and Bailey (1963) provided a com- again by Sethi (1960) who elevated the ranks prehensive discussion of the relationships of 1981 PARENTI: CYPRINODONTIFORM FISHES 353 the family Poeciliidae to other cyprinodon- licles during the entire developmental peri- tiforms, yet came to no firm conclusions. od, and the fact that the first three anal rays They suggested that the closest relative of are unbranched in both. However, he made the poeciliids is perhaps another viviparous no formal reclassification and maintained killifish; however, they stressed the fact that that Anableps was so distinct that it should modifications for viviparity among the four remain in its own family. families were not alike, except for the similar Alignments of one subfamily of cyprino- gonopodial structure of Jenynsia and Ana- dontids to another have been suggested by bleps, the intrafollicular development of poe- a number of workers (e.g., Ahl, 1924, 1928; ciliids and Anableps, and the presence of Hoedeman and Bronner, 1951; Miller, 1955a; trophic processes in goodeids and Jenynsia. Uyeno and Miller, 1962). In addition, Rosen and Bailey maintained Ahl (1924, 1928) considered solely the Af- that poeciliids were more like some ovipa- rican cyprinodontid genera of the Rivulinae rous than viviparous cyprinodontiforms in and Procatopodinae, which he believed general body form and osteology; however, formed a natural group. they did not suggest a group of cyprinodon- Hoedeman and Bronner (1951, p. 1) made tids which could possibly be a close relative recommendations for the alteration of cy- of the poeciliids. prinodontiform classification. They con- Rosen and Bailey classified the Poeciliidae structed the tribe Profundulidi to include the in three subfamilies: the Tomeurinae Eigen- Old World genera Kosswigichthys and Va- mann, containing just one genus and species, lencia, and the North and Central American Tomeurus gracilis; the Xenodexiinae Hubbs, fundulines Profundulus and Adinia. The also containing just one genus and species, tribe was regarded as unnatural by Miller Xenodexia ctenolepis; and, the Poeciliinae, (1955a) who suggested that the Fundulinae containing all other members of the family. and Cyprinodontinae be merged. Each of Structurally, Tomeurus is much like other these four genera is regarded as a member of poeciliids; however, it diverges strongly in a different subgroup in the phylogenetic anal- the elaborate modifications of the gonopo- ysis of the present study. dium, and also in that it is the only oviparous Miller (1956), in describing Cualac, a new poeciliid. (Internal fertilization results in the genus of cyprinodontids, stated that it was laying of a fertilized egg.) These differences, intermediate between the Fundulinae and and also the remarkable similarity of the North American Cyprinodontinae. He reit- form of the gonopodium to that of the ovipa- erated his previous suggestion that the two rous Horaichthys, an adrianichthyoid, led groups together comprise the subfamily Cy- Nikol'skii (1954) to propose that the two gen- prinodontinae, claiming that they were prob- era, each classified in its own family, be ably artificially separated on the basis of den- united into one superfamily, the Tomeuroi- tal morphology. dea. Kulkarni (1948) however, suggested Later, however, Uyeno and Miller (1962) that on the basis of overall osteological sim- supported Sethi's conclusion that the two ilarity, Horaichthys was closer to Oryzias groups remain separated in a classification. than Tomeurus. This conclusion has been They listed a series of characters from Sethi supported by all recent workers on both poe- (1960) by which the fundulines could be dis- ciliids and adrianichthyoids (e.g., Rosen, tinguished from the cyprinodontines: the 1964, 1973a; Rosen and Parent!, MS). The presence of parietals; neural arches of the alignment of Tomeurus with the poeciliids is first vertebra not fused to skull, and there- also supported by the present study. fore, taking no part in the articulation of the Miller (1979), in discussing the relation- vertebral column to the skull; the presence ships of Anableps dowi, supported the align- of occipital condyles; and, the lack of a gap ment of Anableps with the poeciliids, citing between the first and second vertebrae. as evidence of close relationship the reten- These characters are, however, as Myers's tion of the embryos in modified ovarian fol- defining characters of the Cyprinodontidae, 354 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

primitive for atherinomorph fishes. Thus, it closely related to the adrianichthyoids. How- is not just the superfamily and families which ever, in spite of such inconsistencies related are poorly defined, but the subfamilies as to the level at which he was approaching the well that lack precise definitions and, there- problem, this work represented for the first fore, need to be supported or refuted as time a precise although informal statement monophyletic groups of genera. about the interrelationships of the four vivip- Foster (1967) presented a summary of his arous families and their relationship to the conclusions regarding atherinomorph phy- subfamilies of the Cyprinodontidae was pre- logeny in a branching diagram. The Pantan- sented. It is noteworthy also for including odontinae was removed from the cyprino- the implicit statement of the nonmonophylet- dontiforms and placed as a close relative of ic nature of the family Cyprinodontidae. the adrianichthyoids. Within the cyprinodon- However, Foster's treatment, like the oth- tids, he recognized six subfamilies (the Flu- ers, grouped the families and subfamilies viphylacinae Roberts obviously omitted), mainly on overall similarity without regard concluding that the Rivulinae and Orestiati- to the primitive or derived nature of char- nae were sister groups primitive to other cy- acters. The family Cyprinodontidae and also prinodontiforms, excluding Pantanodon. the cyprinodontiform fishes as a whole, as The Procatopodinae and Lamprichthyinae indicated by Foster's removal of Pantano- were similarly depicted as sister groups don, are left to be formally defined as mono- primitive to the four viviparous families and phyletic groups or to have their monophyly the Fundulinae and Cyprinodontinae. Foster refuted. Similarly, the viviparous families re- considered the fundulines to be most closely main to be unambiguously defined on the ba- related to the Anablepidae (presumably in- sis of derived characters unique to them and cluding Jenynsia) which together formed the not found in other cyprinodontiforms. sister group of the poeciliids. This subgroup, Thus, to accomplish the stated objectives in turn, was assessed as being most closely of this study, the four viviparous families are related to the Cyprinodontinae and Goodei- treated as four more genera of cyprinodon- dae, represented as sister groups. tiform fishes, the monophyly of each being This analysis was based on an assessment supported or rejected. Furthermore, the re- largely of overall similarity for a group of 16 classification of cyprinodontiforms pre- or more characters, including those of the sented in this study is based on an attempt osteology, behavior and development. to define the major groups of genera and rep- Foster presented no formal reclassification resent their hierarchical relationship, rather of the atherinomorph fishes. He therefore re- than to obscure such relationship by basing tained the subfamily rank for Pantanodon the rank of a taxon on subjective criteria of even though he considered it to be more uniqueness.

DERIVED CHARACTERS OF CYPRINODONTIFORMS The Cyprinodontoidea has been recog- ichthyoid fishes. Inadequate definition of the nized since the definition of the family Cy- Cyprinodontidae also is responsible for the prinodontidae by Gill (1865). However, the unsupported placement of Pantanodon with failure by him and subsequent workers to adrianichthyoids by Foster (1967). define the Cyprinodontidae rigorously has Previous workers attempting to define the resulted in the uncritical inclusion with them superfamily (e.g., Regan, 1911; Hubbs, 1924) of the ricefish genus Oryzias until Rosen have included characters either primitive for (1964) placed it in its own family and sug- atherinomorph fishes or shared by a number gested its close relationship to the adrian- of its subgroups. As a result, the superfamily 1981 PARENTI: CYPRINODONTIFORM FISHES 355 has never been unambiguously denned as a Among other groups of atherinomorphs, monophyletic group (in the sense of Hennig, there are lobate caudal fins exclusively. The 1966). atherinoid Menidia beryllina (fig. 2B) has a The current study has revealed that all caudal skeleton which is asymmetrical in fishes of the superfamily Cyprinodontoidea having two epurals which are relatively may be distinguished from all other teleost smaller than the opposing parhypural. The fishes by the following derived features. divided hypural plate has a larger dorsal seg- CAUDAL FIN: A series of derived charac- ment. Oryzias (fig. 2C) has an asymmetrical ters within the caudal fin is found relatively caudal fin support in which two small epurals unmodified in all cyprinodontiforms. Exter- oppose the single, large parhypural. The hy- nally, the fin is rounded or truncate, although pural plate is divided into subequal dorsal in males of several aplocheiloid genera, pro- and ventral segments. catopines, and the South American Orestias, FIRST PLEURAL RIB: Typically among the there are often extensions of the dorsal and atherinomorph fishes, the first pleural rib ventral caudal rays. In no case are there in- arises on the parapophysis of the third ver- cipient lobes; although, Miller (1979) reports tebra. Occasionally the rib is borne on the that in males of Anableps microlepis, lower parapophysis of the fourth vertebra in males caudal rays are often grouped together form- of the genus Ceratostethus, and both males ing a lobelike structure. Branched caudal and females of the genus Gulaphallus, both rays typically number eight or more. phallostethoid fishes in the suborder Atherin- Internally, the supports of the caudal fin oidei (Roberts, 1971). are symmetrical (fig. 2E). There are two hy- Within the cyprinodontiforms, the first pural plates, one above and one below cor- pleural rib is borne on the parapophysis of responding to fused hypurals 3, 4, and 5 and the second vertebra. In the funduline genus 1 and 2, respectively. In some species of Adinia there is a pleural rib on the parapoph- Epiplatys and Aplocheilus, the upper hypu- ysis of the first vertebra; this condition is ral plate is divided in two, apparently rep- considered to be apomorphic for the genus. resenting the unfused hypurals 4 and 5 (fig. Rosen (1964) followed Myers (1928a) in 2D). stating that the first pleural rib of phallosteth- Within the cyprinodontiforms, fusion of oids arose on the parapophysis of the second the hypural plates into a so-called hypural vertebra, and therefore suggested a close af- fan (following the terminology of Rosen, finity between the phallostethoids and cy- 1964) occurs within several monophyletic prinodontiforms. However, Roberts (1971) groups of genera (e.g., fig. 2F). has shown this to be a misidentification of There is just one epural which mirrors in the state in phallostethoids. Examination of shape and position the autogenous parhy- several genera of phallostethoid fishes as pural. There are no separate ural centra. The part of this and other studies has supported hypochordal musculature is also absent (Ro- Roberts' contention. Therefore, the first rib sen, 1964). arising on the parapophysis of the second This formation of a symmetrical caudal fin vertebra is a characteristic unique to cyprin- in unique among teleost fishes. The esocoid odontiforms. Umbra limi has a caudal fin which is exter- JAW STRUCTURE: The protrusible upper nally unlobed and rounded. Yet, an exami- jaw of the atherinomorph fishes differs from nation of the internal structure reveals that that of other acanthopterygians in the lack of the external symmetry is effected by a com- a ball and socket joint between the autopala- plex of two epurals, one uroneural, five un- tine and the maxilla, and the absence of fused hypurals, and two separate ural centra, crossed rostral ligaments (Rosen, 1964). the second of which is dorsally offset to the The absence of a ball and socket joint pre- first (fig. 2A). In addition, there are fewer vents the premaxillaries from being locked than eight branched caudal rays. in the protruded position by the autopala- 356 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 1981 PARENTI: CYPRINODONTIFORM FISHES 357 tines upon the opening of the mouth. How- bound by connective tissue fibers to the an- ever, the premaxillaries may still be held pro- terior end of the rostral cartilage. tracted by contraction of the superficial The presumed function of the rostral car- division of the adductor mandibulae (Al) tilage is to prevent the independent move- (Alexander, 1967a, 1967b), which inserts on ment of the premaxillaries, and also to pre- the middle of the distal arm of the maxilla. vent their rolling off the cranium when the Crossed rostral ligaments run from the left mouth is opened (Alexander, 1967a, 1967b). autopalatine and the right autopalatine to the However, since the maxilla functions as a heads of the right and left premaxillaries, re- brace during the forward movement of the spectively. These ligaments, along with a premaxillaries in atherinomorphs, and the pair of ethmomaxillary ligaments, typify the rostral cartilage is not present in all cyprin- mechanism of the protrusible upper jaw of odontiforms, it appears that the rostral car- acanthopterygians (Schaeffer and Rosen, tilage serves mainly as a restrainer of the in- 1961). dependent movement of the premaxillaries. The atherinomorphs lack crossed rostral Primitively, within most acanthopterygi- ligaments; thus, the forward movement of ans and most atherinomorphs, in addition to the premaxillaries is limited by contact with being bound to the ascending processes of the maxilla. Among the atherinomorphs, the premaxillaries, the rostral cartilage is at- Alexander (1967b) reports the presence of an tached to the median processes of the max- ethmomaxillary ligament in the atherinoids illary heads by connective tissue fibers. Atherina and Melanotaenia, and the aplo- Alexander (1967b) reports that in atherinoids cheiloid Aplocheilus. He notes its absence there is also often an attachment of the car- in Fundulus and the poeciliid Xiphophorus. tilage to the vomer and to the ethmoid car- The mechanism of protrusion of the upper tilage. jaw of cyprinodontiforms has been described Among cyprinodontiforms, variability in in detail by Rosen (1964) and Alexander the states of degree of attachment of the ros- (1967a, 1967b). It is characterized by a two- tral cartilage, size of the cartilage, its pres- part alveolar process of the premaxillaries. ence or absence, length of the ascending pro- A distal part of the process is joined to an cesses of the premaxillaries and presence or offset proximal part of the process, thus cre- absence of the ethmomaxillary ligaments al- ating a wide bow, as illustrated in the aplo- lows for the description of transition series cheiloid Austrofundulus (fig. 3A). In all cy- of these characters and the delimitation of at prinodontoids, the process is primitively least three distinct mouth forms within the S-shaped (fig. 3B) as a result of the distal group. (Sethi, 1960, referred to both the ros- part of the process being strongly indented tral cartilage and the mesethmoid as the mes- posteriorly. ethmoid; therefore, his descriptions of states At the posterior tip of the ascending pro- of the mesethmoid are unreliable since they cesses of the premaxillaries is a large, free refer to either one or the other.) rostral cartilage. Among acanthopterygians, The aplocheiloids share some similar up- the rostral cartilage is typically firmly at- per jaw characteristics with the atherinoids. tached to the ventral surface of the tips of These are the presence of an ethmomaxillary the ascending processes by connective tissue ligament, the presence of a ligament from the fibers, and, in addition, is sometimes internal hooks of the maxillaries to the ros- wrapped around the tips of the processes. tral cartilage, and the presence of a meniscus The median process of the maxillary head is between the premaxilla and maxilla. These

FIG. 2. Diagrammatic representation of the caudal skeleton of A. Umbra limi (after Rosen, 1974); B. Menidia heryllina; C. Oryzias latipes (after Rosen, 1964); D. Aplocheilus panchax; E. Aphyosemion gardneri; F. Fundulus heteroclitus. 358 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

ASC

ALV B FIG. 3. Diagrammatic representation of premaxillary alveolar arm, lateral view, of A. Austrofun- dulus transilis; B. Profundulus punctatus. Arrows point to areas indented in cyprinodontoids to form S-shaped arm.

ligaments and the meniscus are absent in all Anableps, and held together by connective cyprinodontoids. tissue fibers. The rostral cartilage in cyprinodontiforms, Thus, within the cyprinodontiforms there as stated, is free and not wrapped around the are three basic forms of the upper jaw and ascending processes of the premaxillaries as jaw suspension. The first, and apparently it is in most acanthopterygians. This permits most primitive, is that of the aplocheiloids. the movement of the rostral cartilage relative The rostral cartilage is large and firmly at- to, rather than with, the ascending process- tached to the broad premaxillary ascending es. In all aplocheiloids, the rostral cartilage processes. There are ligamentous attach- is a large, disc-shaped element lying beneath ments of the head of the maxilla to the rostral the flat and broad ascending processes (fig. cartilage and of the maxilla to the ethmoid. 4). In one cyprinodontoid genus, Profundu- The large size of the rostral cartilage and the lus (fig. 5B), the ascending processes are presence of ligaments are assessed as prim- broad and the cartilage large, yet somewhat itive by comparison with an outgroup, the reduced relative to the aplocheiloid condi- atherinoids, and to the percomorph fishes, in tion. It is further reduced in the fundulines which these states are present. (fig. 5C), and Valencia (fig. 5D). There are three distinct states of the pre- Among remaining cyprinodontiforms (figs. maxillary ascending processes in cyprino- 35, 39), the cartilage is present as a minute dontiforms: flat and broad in aplocheiloids disc, or absent, whereas the ascending pro- and Profundulus, long and narrow in Fun- cesses are shortened, or nearly absent as in dulus and related genera, and short and 1981 PARENTI: CYPRINODONTIFORM FISHES 359

opal

FIG. 4. Diagrammatic representation of the upper jaw in A. Aplocheilus panchax; B. Pachypanchax playfairi; C. Aphyosemion petersi; D. Cynolehias whitei.

pointed or triangular in all remaining cyprin- systems clearly indicates that Profundulus is odontiforms. Exceptions occur in Anableps, more closely related to cyprinodontoids than as mentioned, and in Oxyzygonectes in to aplocheiloids. Therefore, the flat and which the processes are enlarged. These ex- broad ascending processes are most parsi- ceptions are discussed in the phylogenetic moniously assessed as the primitive state analyses and generic diagnoses. within the cyprinodontiforms. The short and The size of the ascending processes in oth- pointed or triangular processes coupled with er atherinomorphs and acanthopterygians is an extremely reduced, or in some cases ab- variable; however, the general or most com- sent, rostral cartilage, are defining char- mon state is for the processes to be long and acters of a large group of cyprinodontiforms narrow. This would suggest that the transi- encompassing the poeciliids, goodeids, Je- tion series for ascending processes is from a nynsia, Anableps, Oxyzygonectes, the cy- primitive state of long and narrow to flat and prinodontines, Orestias, Cubanichthys, broad in one lineage, and to short and narrow Chriopeoides, and the procatopines. in another. However, information from other In addition, in this group as well as in the 360 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 5. Diagrammatic representation of the upper jaw in A. Rivulus harti; B. Profundulus punctatus; C. Fundulus diaphanus; D. Valencia hispanica.

Mediterranean Valencia, there is a dorsal branchial and the second pharyngobranchial. extension of the maxilla over the ascending In contrast, the uncinate process is lacking processes, which forms a cuplike process in all cyprinodontiforms. The cartilage is with the ventral extension. The short as- subequal to the epibranchials in aplochei- cending premaxillary processes slide in and loids, and extends between the posterior out of this maxillary cup. In primitive cy- base of the first epibranchial and the second prinodontiforms the twisted maxilla extends pharyngobranchial (fig. 6A). In cyprinodon- ventrally under the premaxilla but not dor- toids, the cartilage and the first epibranchial sally. are both present in the same position; yet, GILL ARCHES: Interarcual cartilages are they are reduced to approximately half their found among the percomorph fishes (Rosen length relative to the size of these elements and Greenwood, 1976). The general condi- in the aplocheiloids (fig. 6B). tion is that found in the atherinoid genus The interarcual cartilage is present in all Melanotaenia. A rod of cartilage extends cyprinodontiforms except groups of proca- between an uncinate process of the first epi- topine and of aplocheiloid genera in which 1981 PARENTI: CYPRINODONTIFORM FISHES 361

B PB2

TPB4

FIG. 6. Diagrammatic representation of dorsal gill arches, ventral view, A. Austrofundulus transilis; B. Profundulus punctatus. other elements of the dorsal gill arches are teleosts. Roberts (1970) referred to it as the present in a typical arrangement, and the in- "first rib?" in his description of the osteol- terarcual cartilage is assumed to have been ogy of the South American Fluviphylax pyg- lost. maeus. Sethi (1960) refers to the element PECTORAL GIRDLE: The cyprinodontiform only in an illustration in which it is labeled shoulder girdle typically has a first post- "PCL [Postcleithrum] 2." cleithrum which is large and scale-shaped Weitzman (1962) illustrated the shoulder (figs. 7C, 8A, B). This is in contrast to the girdle of the characin Brycon meeki which condition in atherinoids and exocoetoids in has three postcleithral elements. The third which the first postcleithrum is typically a postcleithrum is comparable in shape and slender bone (figs. 7A, B). Among cyprino- position to the so-called secondary post- dontiforms the first postcleithrum is absent, cleithrum of cyprinodontiforms. Therefore, and therefore presumed lost, in poeciliids, I interpret these structures as homologues most procatopines, Leptolucania, Orestias, and conclude that the general condition for Rivulus and its South American relatives and a cyprinodontiform is to have a large first one species of Anableps, A. dowi. postcleithrum, and a narrow third postclei- There is another postcleithrum situated thrum, with the second postcleithrum al- medial to the scapula and radials, and ex- ways lacking. tending ventrally beyond the coracoid. This Also, the lowset pectoral fin, effected by long, slender element is present in cyprino- a ventral position of the radials (e.g., fig. 7A, dontiforms. Rosen and Bailey (1963) inter- D), primitively distinguishes the cyprinodon- preted this as a "secondary postcleithrum" tiforms from all other atherinomorphs which in the poeciliids without discussing its ho- have highset pectoral fins and more dorsally mology to the second postcleithrum of lower situated radials (fig. 7A, B). 362 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

pc-3 FIG. 7. Diagrammatic representation of left shoulder girdle of A. ; B. Menidia menidia; C. Aplocheilus panchax; D. Rivulus harti. Cartilage is stippled.

Highset pectoral fins within cyprinodonti- of the shoulder girdle based on (1) a series forms occur in the poeciliids and procato- of uniquely derived characters that indicate pines (fig. 8C, D), and are interpreted as a close relationship between the poeciliids being secondarily derived. This is the most and procatopines, and (2) a series of derived parsimonious interpretation of the condition characters that indicate that these two are 1981 PARENTI: CYPRINODONTIFORM FISHES 363

D

FIG. 8. Diagrammatic representation of left shoulder girdle of A. Profundulus punctatus; B. Cy- prinodon variegatus; C. Procatopus gracilis; D. Tomeurus gracilis. Cartilage is stippled. together more closely related to one group than to another, which also possesses lowset of cyprinodontiforms with lowset pectorals pectorals. 364 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

BREEDING AND DEVELOPMENT: Eggs of sexual maturity, which occurs as early as the oviparous atherinomorph fishes are dis- four to six weeks, until death. tinguished by having long, chorionic fila- Annual fish eggs enter three diapause ments by which they attach to the spawning stages prior to hatching (Wourms, 1972a). substrate, and conspicuous oil droplets (in- The first, termed Diapause I, occurs during ferred to be secondarily lost in the suborder the pre-embryonic stage. The cells of the Exocoetoidei) (Foster, 1967). Cyprinodon- blastodisc separate and disperse around the toid eggs, in turn, are distinguished by their surface of the yolk sphere. Arrest lasts until relatively longer development time and the cells reaggregate to form the embryonic thickened chorion, the outermost egg mem- shield when the anterior-posterior axis of the brane. embryo is established for the first time. Thus, the egg of a typical oviparous cy- Diapause II occurs during the mid-somite prinodontiform may be characterized as stage about the time of formation of the heart large (some over 2.0 mm in diameter), con- tube. Diapause III occurs just prior to hatch- taining several oil droplets and surrounded ing. The embryo is fully formed and capable by a thick, filamentous chorion. A typical of hatching, yet does not. The embryo re- nonannual cyprinodont egg has a develop- mains quiescent; its heart beat slows down ment time of 12 days or longer. Eggs of the and the characteristic turning of the embryo atherinoid Bedotia geayi were observed in and associated beating of the pectoral fins the laboratory to have a development time within the chorion are slowed or cease. The of about nine days. Development time is up duration of each of the diapause stages is to six months for eggs of fishes in true annual controlled either by genetic or environmental genera such as the South American Cyno- factors, or an interplay of the two. Embryos lebias and the African Nothobranchius. The in stage III have remained quiescent for thick chorion permits survival under condi- more than six months (Wourms, 1964). tions of desiccation during an extended de- Previous workers have considered the an- velopment period, such as those typical of nual habit to be uniquely derived within the Fundulus. annual killifish genera therefore suggesting The annual habit (first reported by Myers, that these fishes form a monophyletic group. 1942; then described in detail by Peters, The present study disagrees with this con- 1965, and Wourms, 1963, 1964, 1967, 1972a, clusion for two reasons: (1) On the basis of 1972b, and 1972c) is exhibited by a minority anatomical characters, certain true annuals of the aplocheiloid species of tropical South are assessed as being more closely related to America and Africa. Adults live for no more nonannuals than they are to other true an- than one rainy season during which time they nuals. (2) All cyprinodontiforms have a pro- spawn. The eggs enter diapause and survive longed development time, and within genera the dry period buried in the substrate. The that are not closely related to the aplochei- fertilized eggs normally hatch at the onset of loids, survival of eggs through periods of the subsequent rainy season; however, they desiccation has been demonstrated. have been known to survive dry periods of The first of these reasons is discussed fully several years. in the phylogenetic analysis of cyprinodon- Foster (1967) reports that in the atherinoid tiform genera. The second is given in support Melanotaenia and in Oryzias spawning nor- of the contention that the annual habit is no mally takes place without direct contact with more than an exaggeration, due to extreme a substrate. In addition, a large number of environmental fluctuations, of a capability of eggs are extruded at once. In contrast, all all cyprinodontiforms to survive stress that killifishes, with a few possible exceptions, involves desiccation. spawn in contact with a substrate, and eggs Foster (1967, p. 538) summarized the hab- are extruded one at a time. Spawning in a itats of killifishes as: "If any generalization typical annual occurs daily from the onset of could be made about the ecology of killifish- 1981 PARENTI: CYPRINODONTIFORM FISHES 365 es, it is that they exploit niches mostly in tion in the nonannuals appears to be com- ephemeral waters, places which are tempo- parable to Diapause III. rarily submerged by tides, floods from heavy Turner (1966) reports that a collection of rains, or similar causes." Pantanodon podoxys has been made in Af- The ability of nonannual killifish embryos rica from stagnant pools. The common cy- to survive desiccation has been reported for prinodontiforms in the vicinity were two a number of species within the North Amer- species of the annual Nothobranchius which ican genera Fundulus and Cyprinodon. were present in similar pools. No permanent Harrington (1959) reported that popula- body of fresh water was found that could be tions of Fundulus confluentus in inferred to have originally formed the pools. have survived hatching delays of up to three Such circumstances suggest that Pantano- months. Areas of the salt marsh habitat of don may be an annual, and therefore that this killifish are exposed to the air during the annualism among cyprinodontiforms is not months of October through December. Also, restricted to the aplocheiloids, but is perhaps Taylor, DiMichele and Leach (1977) ob- a general characteristic of those cyprinodon- served that another estuarine Fundulus, F. tiforms which inhabit ephemeral waters. heteroclitus, often spawns during the high The use of a potential annual lifestyle as night tide. Eggs are thus stranded in the sub- a defining character of the cyprinodontiforms strate up to a week after expected hatching is confounded by the fact that the atherinoid time; hatching is delayed until reimmersion Leuresthes tenuis spawns in conjunction occurs. with the tidal water level fluctuations so that In an effort to test the generality of the the eggs are incubated while exposed to the ability of cyprinodont eggs to survive desic- air (Clark, 1925). This suggests that the abil- cation, F. Douglas Martin (personal com- ity to survive desiccation is a derived char- mun.) exposed eggs of Cyprinodon to the air acter for the atherinomorph or some larger and found that they can survive through group of fishes. However, the generality of hatching; although, on the average, they are this condition and its concordant develop- less successful at hatching than Fundulus mental alterations for other groups of fishes species. This may indicate that the ability to awaits further description. Therefore, the survive periods of desiccation is primitive for early and regular breeding habit and long de- cyprinodontiforms and is lost in the more velopmental period, coupled with the ability advanced genera such as Cyprinodon. to survive desiccation, is considered to de- The first two diapause stages have not scribe a unique developmental pattern of cy- been demonstrated in nonannuals; however, prinodontiform fishes. the ability to survive pre-hatching desicca-

PHYLOGENETIC ANALYSIS

The monophyly of each of the five families esis is that some oviparous cyprinodonti- of cyprinodontiform fishes has been tested. forms are more closely related to the vivip- A preliminary examination revealed that arous cyprinodontiforms than they are to each of the four viviparous families is mono- other oviparous forms. phyletic and can be unambiguously defined, Therefore, the genera of cyprinodontid although not with all of the characters pre- fishes are used as a basis for a cladogram of viously used to define them. all cyprinodontiforms. The poeciliids, good- The Cyprinodontidae, however, as cur- eids, Jenynsia, and Anableps are treated as rently constituted, cannot be defined as a additional genera incorporated into the over- monophyletic group. The alternative hypoth- all scheme. 366 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 9. Cladogram of major groups of cyprinodontiforms. Derived characters: Node A: symmetrical caudal fin, externally rounded or truncate, internally with one epural opposing a similarly shaped par- hypural, hypural plates symmetrical; alveolar arm of premaxilla; interarcual cartilage from base of first epibranchial to second pharyngobranchial; first pleural rib on parapophysis of second vertebra; lowset pectoral girdle with scale-shaped first postcleithrum; early breeding habit and long developmental period; Node B: attached orbital rim; cartilaginous mesethmoid; close-set pelvic fin supports; narrow and twisted lacrimal; broad anterior end of basihyal; tubular anterior naris; reduced cephalic sensory pore pattern; pigmentation pattern (see text); Node C: two ossified basibranchials; loss of the dorsal hypo- hyal; reduced interarcual cartilage; head of autopalatine offset to main axis and with posterior flange; anterior ventral extension of the autopalatine; loss of the metapterygoid; premaxilla with a posterior indentation of the alveolar arm; dentary expanded medially; loss of the first dorsal fin ray; loss of an ethmomaxillary ligament; loss of a ligament from the maxillaries to the rostral cartilage; loss of a meniscus from between the premaxilla and maxilla; Node D: premaxillary ascending processes narrow or greatly reduced in adults; rostral cartilage reduced or absent; inner arms of maxillaries not abutting the rostral cartilage; lateral ethmoid with reduced facet for articulation of autopalatine; Node E: maxilla with straight proximal arm; large dorsal process of the maxilla directed over the premaxillary ascending process; Node F: ascending processes of the premaxillaries short and narrow; dorsal processes of maxillaries rounded or reduced; nasal expanded medially (or secondarily reduced); Node G: lateral ethmoid expanded medially and lying perpendicular to the frontal; reduced autopterotic fossa; enlarged inclinators of the anal; Node H: maxilla with expanded distal arm (or secondarily reduced); parasphenoid with expanded anterior arm; dorsal process of maxilla with a distinct lateral indentation; elongate retroarticular; pouch created by scales surrounding urogenital opening of females. Node I: dorsal pro- cesses of the maxillaries expanded medially, nearly meeting in the midline and possessing a distinct groove; lateral arm of maxilla robust; toothplate of fourth pharyngobranchial reduced. Node J: uniserial outer teeth: second pharyngobranchial offset to third; parietal absent; Meekers cartilage expanded posteriorly; transverse processes of vertebrae reduced and cup-shaped. For defining characters and relationships within terminal taxa, see text and figures 20, 25, 75, 81, 83, 87, and 89.

In this discussion of phylogenetic relation- gories. At the conclusion of the phylogenetic ships, the vernacular names as summarized analysis, a reclassification is presented. New previously are used for suprageneric cate- group names in the classification are used in 1981 PARENTI: CYPRINODONTIFORM FISHES 367 the systematic account and biogeographic the previous section. These are the unique analysis which follow. formation of a symmetrical caudal fin; the The results of the phylogenetic analysis first pleural rib arising on the parapophysis are best presented in a cladogram (fig. 9). of the second vertebra, rather than on that Limits and definitions of the recognized gen- of the third; a derived type of protrusible era are presented in this discussion and for- jaw; a unique form and position of the inter- mally in the systematic accounts. arcual cartilage; the lowset pectoral fins with The cladogram is a hierarchic representa- a large, scale-shaped first postcleithrum; and tion of the relationships among genera and a unique pattern of breeding and develop- suprageneric categories which are being pro- ment. posed. The representation is of the most par- The cyprinodontiforms are readily divided simonious distribution of derived characters into two subgroups, the currently recognized and character states. Hypothesized conver- cyprinodontid subfamily Rivulinae (the gences are discussed along with the pro- aplocheiloids), and all other cyprinodonti- posed derived characters. forms, termed the cyprinodontoids. Since The characters for the most inclusive node these groups are both large and quite dis- are the derived characters of the cyprino- tinct, their interrelationships are discussed dontiforms (Group A of fig. 9) discussed in separately.

APLOCHEILOIDS (GROUP B) The* aplocheiloid killifishes comprise over and Pachypanchax initially appears to be an 500 species in 44 nominal genera and sub- intermediate state between the completely genera. Within the aplocheiloids, there are free rim and the fully attached rim. How- two groups of genera, the Old World aplo- ever, because of its apparent folding under cheiloids comprising Epiplatys, Aplocheilus, the frontals, and also because the Old World Pachypanchax, Nothobranchius, Aphyo- aplocheiloids are assessed as a monophyletic semion and their included subgenera; and, group on the basis of a series of other char- the New World or Neotropical aplocheiloids acters, the partially attached rim is most par- comprising the genera Rivulus, Trigonectes, simoniously assessed as secondarily derived Rivulichthys, Rachovia, Pterolebias, Simp- in Aplocheilus and Pachypanchax. sonichthys, Campellolebias, Cynolebias, Mesethmoid: The atherinomorph fishes Austrofundulus, Cynopoecilus, Terranotus exhibit a derived condition of the ethmoid and their included subgenera. region as described by Rosen (1964). The CHARACTER ANALYSIS: Orbital rim: The mesethmoid is typically represented by two orbital rim is attached to some degree in all ossified discs which are angled toward each aplocheiloids. In all the Neotropical aplo- other at their anterior limit to create a wedge. cheiloids and in the African genera Aphy- In all aplocheiloids examined, the meseth- osemion, Fundulosoma, Nothobranchius moid is totally cartilaginous, except for the and Epiplatys, the covering of the eye is con- presence of some small ossification centers tinuous with that of the head along the pe- in several larger specimens of Cynolebias. rimeter of the orbit. In the remaining aploch- The mesethmoid is generally a large ossi- eiloids, those species of the genera fied structure in most other cyprinodonti- Aplocheilus and Pachypanchax, the rim is forms; however, it is cartilaginous in the pro- attached on the lower half of the orbit, and catopines, and in the cyprinodontines of the is apparently folded under the expansion of Anatolian region (e.g., in Aphanius and Kos- the orbit dorsally. In the cyprinodontoids, swigichthys). Because the last two groups and other atherinomorph fishes, the orbital are members of the well defined, monophy- rim is free all along its perimeter. letic cyprinodontoids, the cartilaginous mes- The partially attached rim of Aplocheilus ethmoid is hypothesized to be independently 368 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

derived several times within the evolution of cyprinodontiform fishes. Pelvic Girdle: The pelvic girdles of ather- inomorphs are found united, as in Menidia and other atherinoids, or widely separated, as in Oryzias. When united, the pelvic bones are joined medially by overlapping process- es. In cyprinodontiforms, they are so united. In the cyprinodontoids, as well as in Meni- dia, the anterior part of the girdle is perpen- dicular to the medial overlapping processes (fig. 10B, C). In contrast, the aplocheiloids (fig. 10A) have pelvic bones which are set close together as a result of the medial pro- cesses being reduced. Gill Arches: Aplocheiloids generally ex- hibit the primitive state of the gill arch char- acters for cyprinodontiform fishes. That is, there is a large interarcual cartilage running from the base of the first epibranchial to the side of the second pharyngobranchial, to which it attaches by a ligament. There are also rosette-shaped gill rakers which have been described by Myers (1927) as being unique to the aplocheiloids. This type of gill raker, however, is found in Menidia and oth- er atherinoids, as well as many cyprinodon- toids. Therefore, it is hypothesized to be a primitive character of a group larger than the cyprinodontiforms and is not a defining char- acter of the aplocheiloids. One characteristic of the gill arches unique to the aplocheiloids is the broad anterior end of the basihyal (fig. 11 A). This is typically a slender bone with a cartilaginous cap which is only slightly flared in cyprinodontoids (fig. 11B) and other atherinomorphs. In aplochei- loids, however, the bone and its associated cartilage approach the shape of an equilateral triangle, especially in the Old World aplo- cheiloids which have just a small ossified ba- sihyal and large cartilaginous segment (fig. 11 A). In the Neotropical aplocheiloids, the ossified segment is much larger. The large cartilaginous anterior end of the basihyal gives the aplocheiloids their characteristic large "tongue" which is readily visible upon opening the mouth. Lacrimal: Another derived character that FIG. 10. Diagrammatic representation of pel- defines the aplocheiloids as a monophyletic vic girdles of A. Aplocheilus panchax; B. Jen- group is the shape of the lacrimal. In all ynsia lineata; C. Aphanius fasciatus. aplocheiloids, the lacrimal is a narrow and 1981 PARENTI: CYPRINODONTIFORM FISHES 369

FIG. 11. Diagrammatic representation of ventral gill arches of A. Nothobranchius melanospilus; B. . Cartilage is stippled. twisted bone which often carries a distinct anterior and dorsal to the orbit. The anterior sensory canal (fig. 12A). This is in contrast naris of all aplocheiloids is surrounded by a to the wide and flat lacrimal typically found distinct tube of skin which projects anterior- in the atherinomorph fishes. In Menidia (fig. ly over the upper jaw (fig. 13). A distinct tu- 12B) the lacrimal is flattened and expanded bular naris is found among the cyprinodon- ventrally. In Poecilia (fig. 12C) the lacrimal toids in Cubanichthys and Anableps, and is is also flat and wide. The lacrimal is reduced considered to be independently derived in among the cyprinodontoids in the genus the aplocheiloids and in each of these ad- Pantanodon; however, the preorbital dis- vanced genera. In all other cyprinodonti- tance is wide compared to that of the aplo- forms, the anterior naris never has such a cheiloids, and the narrow lacrimal of Pan- fleshy extension. tanodon is considered secondarily derived. Cephalic Sensory Pores and Squamation: Among the aplocheiloids, the canal of the Gosline (1949) surveyed sensory pore pat- lacrimal is apparently reduced along with the terns in the cyprinodontiforms as a whole sensory canals of other dermal bones in the with an emphasis on the goodeids and Fun- Neotropical aplocheiloids, as discussed be- dulus. Hoedeman (1958, 1974) in his surveys low. of head-scale patterns among cyprinodonti- Anterior Naris: In all atherinomorph fishes forms was responsible for bringing attention minus the exocoetoids, the anterior and pos- to this associated character. Rosen and Men- terior naris are represented by two separate delsohn (1960) surveyed the sensory pore as openings. The anterior naris is typically a well as head-scale patterns among the poe- small opening just posterior to the fold of ciliids. It has been observed that both sen- skin surrounding the maxilla, whereas the sory pore and head-scale patterns are cor- posterior naris is typically a small slit just related. That is, their separate analyses 370 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 12. Diagrammatic representation of left lacrimal of A. Rivutus harti; B. Menidia menidia; C. Poecilia vivipara. Anterior is to the left. could introduce redundancy into this study. lacrimal pores, four mandibular pores, and Therefore, the two systems are discussed to- seven supraorbital pores. This pattern is gether. judged to be the primitive pattern for cyprin- There is much taxic variability as well as odontid fishes based on personal observa- ontogenetic variation within these systems. tion, and data from Gosline (1949) and Wiley The cephalic sensory system starts out in on- (personal commun.). The numbering system togeny as a series of open grooves over for the pores follows Gosline. which skin grows leaving the grooves open Departures from the general pattern (fig. to the surface only by a series of pores. Gos- 14) occur both with respect to the continuity line also noted for three species of Fundulus of canals between pores, and therefore, the that neuromasts of the juvenile pattern are number of pores, and the number and posi- covered over as the individual grows and re- tion of scales. placed by a covered canal with pores. Among the aplocheiloids, the sensory Hoedeman (1958) referred to replacement pores of the head are reduced to a series of of two scales by one larger scale as a fusion; neuromasts, as in Rivulus harti (fig. 13C), a however, Roberts (1970) suggested instead Neotropical aplocheiloid, and in Epiplatys that it was the formation of fewer scale pre- sexfasciatus (fig. 13B), and Old World spe- cursors that resulted in such changes. With- cies. Among the cyprinodontoids, the sen- out developmental studies on comparative sory pores of the head are often reduced, scale formation, this difference seems moot; but not replaced by neuromasts. however, such replacements will not be re- Sexual Dimorphism: All cyprinodonti- ferred to as fusions because this implies a forms show marked sexual dimorphism. process for which there is no evidence. Males typically are elaborately pigmented As a consequence of variability within and frequently have elongated rays in the these systems, it is only the most general unpaired and the pelvic fins. characteristics which may be incorporated All cyprinodontiforms exhibit sexual di- into a higher level phylogenetic analysis. morphism in size. In aplocheiloids, males are Also, because of the ontogenetic changes, it larger than females. In the cyprinodontoids, is usually the maximum development of the reverse is true: females are larger than pores and scales that is reported as the gen- males. The exception in the cyprinodontoid eral state for a species or genus. The gener- killifishes occurs within the procatopines and alized sensory pore and squamation pattern Pantanodon and Fluviphylax in which the for cyprinodontiforms is as in the cyprino- males are always larger than the females. dontoid, Jenynsia lineata (fig. 14A, B, C). Among the silversides, in genera such as There are seven preopercular pores, three Bedotia, Melanotaenia, Gulaphallus and 1981 PARENTI: CYPRINODONTIFORM FISHES 371

FIG. 13. Diagrammatic representation of cephalic sensory pores and squamation. Epiplatys sexfas- ciatus: A. lateral view of lacrimal and preopercular pores, B. dorsal view of neuromasts and squamation, C. ventral view of mandibular and preopercular pores; Rivulus harti: D. lateral view of reduced pore pattern, E. dorsal view of neuromast and squamation pattern (lettering of scales corresponds with b.), F. ventral view of covered branchiostegal region; Cynolebias elongates: G. lateral view of minute neuromast pattern, H. dorsal view of neuromast pattern, I. ventral view of neuromast pattern and branchiostegal region. others, the males are larger than the females. tion and the small size of the males is a de- In at least one species related to Menidia rived character of cyprinodontoids. The beryllina, females are larger than males. In large male in procatopines is, therefore, a exocoetoids the males are also larger than secondarily derived condition. females, or of approximately the same size. No general pattern of pigmentation has Thus, the most general condition among the been discovered among aplocheiloids. Some atherinomorph fishes appears to be that pigmentation patterns do, however, occur in males are larger than females. If so, the prim- some of the more primitive genera of both itive state for the cyprinodontiforms is for the Old World and Neotropical groups, and males to be larger than females. In this case, these are assessed as derived for the aplo- the aplocheiloids retain the primitive condi- cheiloid fishes. These include a caudal ocel- 372 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 14. Diagrammatic representation of cephalic sensory pores and squamation. Jenynsia tineata: A. lateral view of preopercular and lacrimal pores, B. dorsal view of supraorbital pores and squamation, C. ventral view of mandibular and preopercular pores; A generalized poeciliid (after Rosen and Men- delsohn, 1960): D. lateral view of preopercular and lacrimal pores, E. dorsal view of supraorbital pores and squamation pattern, F. ventral view of mandibular and preopercular pores; Orestias cuvieri: G. lateral view of minute neuromast pattern, H. dorsal view of minute neuromast pattern, I. ventral view of neuromast pattern and embedded urohyal. lus in females; bars running across the ven- sometimes discrete spot (e.g., as in R. mar- tral surface of the head, commonly called moratus) dorsally on the caudal fin base. "throat bars"; and a band, normally of white Such an ocellus, however, is also found in or yellow, on the dorsal and ventral base of some females of the genus Aphyosemion. the caudal. Assuming that the spots are homologous, the The caudal ocellus, or "Rivulus spot" has character is a synapomorphy of the aplochei- been used to diagnose fishes of the genus loids, and no longer defines Rivulus as a Rivulus. In females of all nominal species of monophyletic genus. the genus, there is a black blotch (fig. 54), or Throat bars are most prominent in species 1981 PARENTI: CYPRINODONTIFORM FISHES 373

FIG. 15. Diagrammatic representation of ventral view of head to show throat bars in A. Epiplatys dageti, B. Epiplatys chaperi, C. Aplocheilus panchax. of the genus Epiplatys (fig. 15A, B), although many cyprinodontoids, as well as in many they are also a conspicuous component of aplocheiloids which do not possess conspic- the pigmentation patterns of species of uous throat bars (e.g., in Nothobranchius). Aphyosemion and Aplocheilus (fig. 15C), A horizontal band of yellow or white, rare- and to a lesser extent Rivulus and Pachy- ly blue, on the dorsal and ventral bases of panchax. These are often found in conjunc- the caudal fin is a distinctive component of tion with a line of pigment on the lower lip; the pigmentation patterns of the males of however, this line of pigment is found among most species of Rivulus, Aphyosemion, Fun-

A " " C FIG. 16. Diagrammatic representation of a ventral view of skull in A. Fundulus heteroclitus, B. Tomeurus gracilis, C. Aplocheilichthys johnstoni. Lateral ethmoid is cross-hatched, dermosphenotic blackened, autopterotic stippled. 374 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 17. Diagrammatic representation of a ventral view of skull in A. Aphyosemion occidentale, B. Aplocheilus panchax, C. Rivulus hard. Lateral ethmoid is cross-hatched, dermosphenotic blackened, autopterotic stippled. dulosoma, and to a lesser extent in species color patterns occur within the poeciliid fish- of the genus Epiplatys (e.g., in Epiplatys fas- es among cyprinodontoids. Elaborate male ciolatus). Such a band is found only on the coloration does not occur in the hypothe- ventral base of the caudal in species of the sized primitive poeciliid, Tomeurus gracilis, nominal genera Rachovia and Neofundulus. however. In at least two species of Aphyosemion, A. Position of the Vomer: The posterior ex- celiae and cinnamomeum, the light yellow tension of the vomer typically lies ventral to bars extend along the posterior margin of the the anterior arm of the parasphenoid in all fin to form one continuous band (Radda, cyprinodontoids (fig. 16), as well as other 1979). atherinomorphs. In contrast, the posterior The difficulty in determining the primitive extension of the vomer lies dorsal to the an- or advanced state of these pigment patterns terior arm of the parasphenoid in all aplo- is analogous to determining whether sexual cheiloids (fig. 17), a clearly derived condi- dimorphism involving large males or large tion. females is primitive. The general color pat- terns just described are normally retained in SUMMARY OF DERIVED CHARACTERS preserved specimens since the majority in- 1. Attached orbital rim. volve dark pigments, rather than the unsta- 2. Cartilaginous mesethmoid. ble or water and alcohol soluble pigments of 3. Close set pelvic girdles. yellow, blue and red. In an analysis done at 4. Broad anterior end of the basihyal. the level of the current study, intrageneric 5. Narrow and twisted lacrimal. variation has not been the subject of concen- 6. Tubular anterior nans. trated study. However, elaborate color pat- 7. Reduced cephalic sensory pores. terns of large males, distinguished by the de- 8. Males more elaborately pigmented than rived characters listed here, distinguish aplocheiloids from all other cyprinodonti- females. form fishes. Other types of elaborate male 9. Posterior extension of the vomer dorsal to the anterior arm of the parasphenoid. NEOTROPICAL APLOCHEILOIDS The aplocheiloids are hypothesized to the close-set pelvic fins in all members, but comprise two monophyletic groups of gen- did not compare these with other cyprino- era: one a solely Old World group, and the dontids in an effort to present defining char- second Neotropical and north temperate. acters of the group. Similarly, Taphorn and The latter range from southern Florida and Thomerson (1978) who treated only those the Bahamas, through Central America, and species of the genera Rachovia and Austro- southward to Paraguay (fig. 18). They are fundulus, describing Terranotus as new, currently classified in 13 nominal genera and concentrated on enumerating differences subgenera: Rivulus, Rachovia, Austrofun- among nominal genera, rather than describ- dulus, Trigonectes, Rivulichthys, Pterole- ing derived similarities in an effort to deter- bias, Terranotus, Neofundulus, , mine interrelationships of the genera. Campellolebias, Cynopoecilus, Simpsonich- The question still remains, therefore, if the thys, and Cynolebias, five of which are Neotropical and Old World aplocheiloids monotypic. form distinct monophyletic groups of genera, In all revisions or discussions of all or part or whether some Neotropical genera are of the Neotropical aplocheiloids, they have more closely related to some Old World gen- been treated as a monophyletic group (e.g., era than to other Neotropical genera. Ref- Regan, 1912; Myers, 1927; Weitzman and erences to the overall similarity of the Afri- Wourms, 1967; and Taphorn and Thomer- can Nothobranchius to the South American son, 1978), although the group has never Cynolebias have been made continuously in been formally defined. Regan (1912) listed the aquarium literature (e.g., Stoye, 1947; the following characters to distinguish the Scheel, 1968). Furthermore, and perhaps three then known genera Rivulus, Pterole- more importantly, the idea that annualism is bias, and Cynolebias, and his newly named a uniquely derived character (Wourms, Cynopoecilus from other fishes then classi- 1972a) suggests that Neotropical annuals are fied in the Fundulinae: snout short, margin more closely related to the Old World an- of eyes not free, gill membranes separate, nuals than to the fishes of the predominantly mouth wide and transverse with the premax- nonannual genus Rivulus. illaries protractile, lower jaw prominent and Taphorn and Thomerson (1978) stated oblique, teeth subconical and arranged in they agreed with Weitzman and Wourms's bands, with one large outer row, pectorals (1967) conclusion that the Neotropical placed low, and pelvics not far in advance of aplocheiloids such as Austrofundulus and the anal. Rachovia were derived from a Rivulus-like The majority of these characters are either ancestor, but Cynolebias and Terranotus derived for the aplocheiloids as a group, or were not included in this endorsement of for all cyprinodontiforms. Others, such as Weitzman and Wourms' position. "gill membranes separate" may refer to cer- Thus, the definition of the Neotropical tain characters inferred in this study to be aplocheiloids as a distinct monophyletic derived. group is a step toward our understanding of Myers (1927) in a revision of Neotropical the evolution of the annual lifestyle, as well aplocheiloids cited the attached orbital rim as toward a definition of the problems that and rosette-shaped gill rakers as defining remain in aplocheiloid systematics. characters, characters which clearly define CHARACTER ANALYSIS: Shoulder Girdle: a larger group of fishes. The shoulder girdle of Neotropical aplochei- Weitzman and Wourms (1967) emphasized loids is distinguished from that of other aplo- the ambiguity of the definitions of the genera cheiloids by lacking the first postcleithrum of Neotropical aplocheiloids, and remarked (fig. 7D). Other cyprinodontiforms have a on the states of certain characters, such as large, scale-shaped first postcleithrum, as in 376 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

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40° 90° 75° V\ /60° L 1 FIG. 18. Distributional limits of Neotropical aplocheiloids.

Aplocheilus (fig. 7C). The first postclei- is not united by a membrane to the opercu- thrum is present in all Old World aplochei- lum ventrally, carries a distinct sensory ca- loids examined. nal which is open externally as a series of Cephalic Sensory Pores and Squamation: pores. There is a fold of skin covering the A series of derived characters related to the throat region between the dentary and the sensory canal system of the head, including urohyal. The throat region, including the squamation patterns, opercular and bran- branchiostegal membranes, is generally not chiostegal membranes, and development of covered with scales. In contrast, the mem- dermal bones carrying sensory canals serve branes covering the opercular region and the to define the Neotropical aplocheiloids as a branchiostegal rays are totally united in the monophyletic group. For example, the typi- Neotropical aplocheiloids. In the generalized cal condition of the branchiostegal and oper- state (e.g., in Rivulus, fig. 13F) the bran- cular membranes in atherinomorph fishes is chiostegal rays are not conspicuous exter- that exhibited by the Old World aplocheiloid nally. Scales usually extend over this contin- Epiplatys (fig. 13C). Several branchiostegal uous covering of the ventral surface of the rays are visible through a clear membrane head which obscures the separation between which is separate from that overlying the the preoperculum and operculum. opercular region. The preoperculum, which There are no preopercular pores, which is 1981 PARENTI: CYPRINODONTIFORM FISHES 377 concordant with the weakly developed or neuromast patterns found among the species absent sensory canal in the preoperculum currently placed in those genera. There are (fig. 13D). No Neotropical aplocheiloid ex- a series of neuromasts running posteriorly amined has a complete canal in the preoper- from a position medial to the anterior naris culum; although, there is a short canal in its and extending to the posterior limit of the dorsal extension in all except Terranotus, orbit. The pattern continues in the preorbital Cynolebias, Campellolebias, Simpsonich- region. thys, Cynopoecilus, and Leptolebias. The A derived form of this pattern is exhibited canal is visible externally as an obsolescent by members of the genus Cynolebias (fig. canal, rather than as a pore or series of 13G, H, I). There are a series of minute neu- pores. Similarly, the dermosphenotic is re- romasts which resemble perforations of the duced (fig. 17C) and carries just a small ca- skin. These neuromasts run posteriorly along nal. In the Old World aplocheiloids, the the dorsal surface of the head from a position dermosphenotic is more deeply concave. medial to the anterior naris to a position pos- Also, as noted in the previous section, the terior to the orbit where the line turns lacrimal is a narrow and twisted bone in all abruptly back toward the orbit, then poste- aplocheiloids; it carries a distinct canal in the riorly again for a short distance, curving Old World aplocheiloids, whereas in those of back to the path of the postorbital canal, con- the Neotropics, the canal is obsolescent. tinuing around the eye, and ending near the As expected, a reduction in the sensory posterior naris. canal system in the opercular and infraorbital A similar pattern is exhibited by some cy- region is correlated with that on the dorsal prinodontoids, e.g., the South American ge- surface of the head. The reduction involves nus Orestias (fig. 14G, H, I) and the nominal the substitution of enclosed canals which Anatolian genus Anatolichthys and some open to the surface by pores for neuromasts species of Aphanius. This pattern is consid- or pit organs. ered to be secondarily derived in the cyprin- Gosline (1949) examined just one species odontoids based on the fact that the last of Neotropical aplocheiloid, Rivulus hol- three mentioned genera share a series of de- miae, and concluded that it had not devel- rived characters with other cyprinodonti- oped supraorbital canals, or preopercular, forms that are not shared by the aplochei- mandibular, or preorbital canals. This was loids. The significance of the pattern exhibited found to be true for all Neotropical aplo- by these genera will be evaluated in the dis- cheiloids; however, there is not a progressive cussion of their interrelationships. reduction of neuromast patterns within the The head squamation pattern of figure 14B group, but rather a further elaboration of the is hypothesized to be primitive for cyprino- neuromast pattern in more derived genera. dontiform fishes. Following the convention The pattern of neuromasts is the simplest established by Hoedeman (1958), there is a in primitive members of the genus Rivulus. single A scale, which is identified as the me- For example, in R. marmoratus, there are dian scale lying posterior to a line drawn four supraorbital neuromasts, which may through the posterior limits of the orbits. It correspond to the pores 1, 2a, 2b, and 5 or is preceded by two E scales, one of which 6 of the general pattern. overlaps the other. A single G scale precedes The lacrimal, mandibular and preorbital these, and there are often two or more F pores typically are represented by neuro- scales laterally. masts. The neuromast pattern is progressive- The terminology of the head scales used ly more elaborate in the more derived to describe the general pattern found in cy- species of Rivulus (e.g., R. harti) and in the prinodontiforms was not developed to de- other genera of Neotropical aplocheiloids. In scribe this pattern, but for that typically a review of species of the nominal genera found in Rivulus and other Neotropical Rachovia and Austrofundulus, Taphorn and aplocheiloids. Hoedeman considered the Thomerson (1978) illustrated a variety of pattern of Rivulus (fig. 13B) to be a funda- 378 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 mental arrangement of head scales. The cen- The vomer is broad and typically bears a tral A scale is surrounded by a series of patch of teeth. Its lateral processes extend scales which are coded B through E pro- anteriorly just under the lateral ethmoids. ceeding counterclockwise from the scale just There is a distinct facet for articulation of the posterior to A. This pattern is not fundamen- autopalatine on the anterior edge of the lat- tal for cyprinodontiforms, but is unique to eral ethmoid. This process, which is promi- the Neotropical genera. The unique compo- nent in all species of Profundulus (fig. 57A), nent of the pattern in more derived species is not strongly formed in any other cyprino- of Rivulus and the other Neotropical genera dontoids. It is considered to be a retained is the inclusion of the G scale in series sur- primitive character in Profundulus. rounding the A, thus preventing the overlap The medial face of the lateral ethmoid in of the E scales. Also, scales B through E are Old World aplocheiloids is not expanded to- all approximately the same size, whereas, ward the parasphenoid. Within the Neotrop- generally in cyprinodontiforms, the A, E, ical aplocheiloids, however, the medial pro- and G scales are prominent. cesses of the lateral ethmoids are greatly Hoedeman (1974) illustrated a pattern for expanded and lie just lateral or dorsal to the Cynopoecilus melanotaenia in which there anterior arm of the parasphenoid (fig. 17C). are two small G scales, rather than one, and Maxilla: The maxilla among all aplochei- these are not in series around the A. I have loids, Profundulus and fundulines is narrow not observed this pattern in C. melanotae- and twisted (fig. 5B, C). The anterior arms nia, and conclude that it is possibly part of of the maxilla in aplocheiloids and Profun- intraspecific variation. There is a great deal dulus extend medially toward the large ros- of such variation with head squamation pat- tral cartilage to which they are affixed by terns. The single A scale is not always pres- connective tissue fibers. In the Neotropical ent, and when present can either overlie the aplocheiloids alone, the arm has a process circular arrangement of the B through E on its anterior face (fig. 5A), rather than scales, or lie beneath them. being gently curved as in all other cyprino- There are also a series of smaller scales dontiforms with pronounced anterior arms. anterior to the lettered scales, covering the dorsal surface of the head from a position SUMMARY OF DERIVED CHARACTERS over the middle of the eye to the margin of the frontals. 1. First postcleithrum absent. Taphorn and Thomerson (1978) reported 2. Branchiostegal and opercular membranes much individual variation in the named scale united. patterns of Hoedeman and therefore ques- 3. Obsolescent preopercular and lacrimal tioned the value of this character to distin- canals. guish among species groups within the Neo- 4. Lacrimal, preopercular and mandibular tropical aplocheiloids. Their judgment was canals represented by neuromasts. borne out by this study, and it is concluded 5. Head scales arranged in circular pattern. that the sensory neuromast patterns may be 6. Medial process of lateral ethmoid expand- more readily characterized and incorporated ed. into a phylogenetic analysis. 7. Process on ventral arm of maxilla. Lateral Ethmoid: Another series of char- RELATIONSHIPS OF NEOTROPICAL APLO- acters uniting the Neotropical aplocheiloids CHEILOIDS: The interrelationships of Neo- into a monophyletic group concerns the rel- tropical aplocheiloids have been discussed ative shape and position of the lateral eth- most recently by Weitzman and Wourms moid and the vomer. The orientation and rel- (1967) and Taphorn and Thomerson (1978). ative size of the lateral ethmoid varies among There are currently 13 nominal genera and cyprinodontiforms, but the general condition subgenera in the group as listed above which among aplocheiloids is that exhibited by the together are defined as monophyletic by the Old World genus Aphyosemion (fig. 17A). characters just discussed. 1981 PARENTI: CYPRINODONTIFORM FISHES 379

FIG. 19. Sketch of body form and fin formation in a male Cynolebias (Austrofundulus) dolichopterus. Dotted line approximates base of hypural plates. (After Weitzman and Wourms, 1967.)

Treatments of the interrelationships of elongate fin rays. Taphorn and Thomerson these genera have focused on the overall removed dolichopterus from Austrofundulus similarity of genera without regard to the and placed it in a new monotypic genus, Ter- primitive or derived nature of characters, ranotus. They stressed that it had no clear and, also, on the failure of current generic relationship to any known genus, but that it definitions to distinguish the included species might be more closely related to Cynolebias from those of other genera. The latter prob- than to either Austrofundulus or Rachovia. lem is perhaps the most serious barrier to The creation of monotypic genera for the re- recognizing the monophyletic groups of ception of species whose placement in a species. Because no single character consis- taxonomic scheme is not readily apparent tently distinguished all the species of Pter- has been the trend in aplocheiloid system- olebias, Austrofundulus, and Rachovia, atics. Five of the 13 genera of Neotropical Weitzman and Wourms suggested that they aplocheiloids are monotypic: Terranotus, be included in one genus, although no formal Simpsonichthys, Cynopoecilus, Campellole- taxonomic decisions were made. In the same bias, and Trigonectes. paper, they described a new species of South Redefinitions of aplocheiloid genera based American aplocheiloid which they hesitantly on derived characters rather than on unique placed in the genus Austrofundulus on the combinations of characters will eliminate the basis of overall body shape and coloration. ambiguous placement of species such as dol- The species, A. dolichopterus (fig. 19) is ichopterus. The following phylogenetic anal- readily distinguished from all other members ysis does not involve the revision of all of the group by its small size and extremely species of the genera of Neotropical aplo- 380 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 20. Cladogram of relationships of Neotropical aplocheiloids. Node A: lack of the first post- cleithrum; branchiostegal rays covered; reduced cephalic sensory pore pattern; reduced dermosphen- otic; reduced lacrimal; reduced preopercular; medial processes of the lateral ethmoids expanded; vomer triangular posteriorly; maxilla with anterior process; Node B: cartilaginous interhyal; pelvic fin rays seven or more; Node C: elongate rostral cartilage; extension of the pectoral fin rays to the base of the pelvics; Node D: reduction of anterior ramus of the alveolar arm; derived pigmentation pattern; Node E: lack of the interarcual cartilage; Node F: vertical bar through the eye; thickened anal rays in females; tendency to develop a fatty predorsal ridge in older males; Node G: derived color pattern; Node H: reduced number of scales in lateral series; Node I: dorsal fin rays number 14 or more; Node J: no derived characters; Node K: first proximal radial absent; enlarged spine on first vertebra; reduced fourth cerato- branchial dentition; Node L: pigmented anal papilla; Node M: caudal fin not scaled; preopercular canal closed.

cheiloids which comprise over 110 species to be more closely related to the more de- (Lazara, 1979). Instead, interrelationships rived genera of Neotropical aplocheiloids. based primarily on the type species of nom- The nature of the development of many inal genera are estimated to produce a work- species of Rivulus is unknown, and there are ing model of the phylogeny of this group as probably more annual species included. Fur- in the cladogram of figure 20. No new generic thermore, the annualism of the nominal gen- names are proposed for species excluded era Trigonectes and Rivulichthys is merely from nominal genera, since it is believed that inferred by their locality of capture. Thus, it new generic names should be proposed only has not been determined whether the annual for definable monophyletic groups of species. Neotropical aplocheiloids form a monophy- Rivulus currently contains over 60 species letic group. This possibility remains to be and as such is the largest genus within the tested by a revision of the nominal species group. Traditionally it has been defined on of Rivulus in light of the present findings. the basis of the presence of a caudal ocellus Jaw Structure: All Neotropical aplochei- in females (fig. 54). The presence of such an loids excluding the species of Rivulus ex- ocellus in females of the African genus amined, have an elongate rostral cartilage Aphyosemion suggests that this character is (fig. 4) extending for at least half its length primitive for all aplocheiloid fishes. beyond the tips of the premaxillary ascend- Annualism: Among the Neotropical ing processes. The primitive state of the ros- aplocheiloids, only the species of the genus tral cartilage for cyprinodontiforms as evi- Rivulus, except for R. stellifer Thomerson denced by its occurrence in all other and Turner, are reportedly nonannual. Rivu- aplocheiloids, and in a slightly modified state lus stellifer, in this scheme, is hypothesized in Profundulus, is as a pentagonal block of 1981 PARENTI: CYPRINODONTIFORM FISHES 381 cartilage, the posterior end of which extends ASC just slightly beyond the tips of the ascending processes (e.g., in R. hard, fig. 5A). The monotypic Trigonectes and its includ- ed species strigabundus and the genus Rivu- lichthys are distinguished from all other aplocheiloids by their sharply angled mouth cleft. This is produced by a foreshortening of the anterior ramus of the arm of the pre- maxilla (fig. 21). Such an oblique cleft and reduction of the anterior ramus also occur in a nominal species of Rivulus, R. rogoaguae, considered herein to be a close relative of strigabundus and rondoni. Coloration: Species of Trigonectes and RivulicHthys, as well as R. rogoaguae, also have a derived color pattern consisting of four rows of brown or red reticulations along the sides of the body. The dorsal, anal, caudal, and pectoral fins also have two or three rows of reticulations. A similar color pattern occurs in one other species of South American aplocheiloids, Neofundulus paraguayensis. Neofundulus paraguayensis on the basis of other charac- FIG. 21. Diagrammatic representation of pre- ters, which will be discussed, is apparently maxilla of Trigonectes {Rivulichthys) rondoni, not closely related to strigabundus and ron- lateral view. doni. Rivulus rogoaguae, on the other hand, most likely belongs in a monophyletic group with strigabundus and rondoni, although it anal papillae are only slightly pigmented or is considered to be a synonym of neither. bare in other genera. A vertical bar through the eye is a promi- Predorsal Ridge: Taphorn and Thomerson nent component of the color pattern of (1978) stated that only members of the gen- species of the genera Rachovia, Austrofun- era Rachovia and Austrofundulus as they dulus, Cynolebias, as well as its included defined them, developed a fatty predorsal subgenera, and the genus Neofundulus (fig. ridge, and that this ridge did not develop in 19). This bar is not present in species of Pter- Terranotus. However, in all male Terrano- olebias, Rivulichthys, Trigonectes, nor any tus examined, including types, a prominent species of Rivulus examined. A bar occurs dorsal ridge was found comparable to those in just one other species of aplocheiloids, in Rachovia and Austrofundulus. Therefore, Nothobranchius microlepis of Somalia. The it is proposed that this character is derived occurrence of such a derived pattern in what for this inclusive group. have otherwise been evaluated as two unre- Spine on First Vertebra: The inferred lated groups of aplocheiloids suggests that more primitive genera, including Rachovia, this pattern has been independently derived Pterolebias, Trigonectes, Rivulichthys, and twice within the aplocheiloids. Rivulus, have a relatively short spine on the The two species of Austrofundulus, A. first vertebra and a correspondingly straight- limnaeus and transilis, are distinguished er dorsal profile than in Austrofundulus, Ter- from all other Neotropical aplocheiloids by ranotus and Cynolebias. In these latter gen- having heavily pigmented anal papillae. The era, the dorsal profile is greatly arched (figs. 382 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

13G, 19), apparently as a result of an en- fins are elongate and reach the base of the larged spine on the first vertebra. pelvic fins in rogoaguae and all non-Rivulus Gill Arches: The species of the genus Riv- species. Cynolebias and its relatives have a ulus may be divided into two groups. One caudal which is rounded as a result of a includes the smaller forms such as cylindra- unique orientation of the procurrent rays ceus, marmoratus, and heyei. These are all perpendicular to the vertebral column. characterized by having an interhyal which There are usually six pelvic fin rays in ath- is ossified; that is, there is a perichondrally erinomorph fishes. There are six in all Old ossified element between the posterior cer- World aplocheiloids as well as members of atohyal and hyomandibula. This is the con- the genus Rivulus with an ossified interhyal dition typical of the interhyal of teleost fish- (e.g., R. marmoratus). In the larger species es. The larger species of Rivulus, of which of Rivulus examined, e.g., harti and stelli- hard may be considered typical, have an fer, the number of pelvic fin rays is increased unossified interhyal instead. The character of to seven or eight as it is in all other Neo- an unossified interhyal, however, does not tropical aplocheiloids. This increase is inter- define a monophyletic group of Rivulus preted as a derived character at this level. species, for it is found in all species exam- The pelvic fin rays rarely are increased to ined of all remaining Neotropical aplochei- seven among the cyprinodontoids in some loids. Thus, it indicates that some species of species of the North American cyprinodon- Rivulus are more closely related to the other tines. genera than to some species of Rivulus. The nominal species of Cynolebias and Hence, the genus as currently defined is Terranotus lack scales on the caudal fin. As paraphyletic. mentioned, the scaled caudal fin is apparent- Rachovia has fourth ceratobranchials ly primitive for aplocheiloids, and is present which are covered with teeth. In Austrofun- in all other members. dulus and Terranotus, there are no teeth on Taphorn and Thomerson suggested that the medial expansion of the gill arch ele- Terranotus may be more closely related to ments, but the posterior arm does possess at Cynolebias because its anterior proximal least one tooth. In Cynolebias, the fourth anal radials "articulate" with ribs rather ceratobranchials are devoid of teeth. There- than hemal spines. However, within actinop- fore, the reduction of teeth on the fourth terygian fishes, the proximal anal radials ceratobranchial is assessed as a defining char- never properly articulate with either ribs or acter of a group including Cynolebias, Aus- hemal spines, but are often found intercalat- trofundulus, and Terranotus. ed with the latter. The character to which All characters could not be determined for Taphorn and Thomerson referred could the sole specimens of ornatipinnis and Par- more appropriately be described in terms of aguay ensis studied; however, they do have the position of the anal fin relative to the ab- the primitive character of fourth ceratobran- dominal and precaudal vertebrae. chials covered with teeth. In dolichopterus, there are 14 abdominal The interarcual cartilage is present primi- vertebrae and 12 precaudal; the first proxi- tively in the aplocheiloids (e.g., as in Aus- mal anal radial (equivalent to the second of trofundulus transilis, fig. 6A) and in all Neo- Rivulus since the first is lost) extends be- tropical aplocheiloids except those of the tween the pectoral ribs of the ninth abdom- nominal genus Pterolebias, in which it is ab- inal vertebra. sent. The cartilage is reduced in one species In the one species of Pterolebias which of Rachovia, R. maculipinnis. exhibits this character, there are 15 abdom- Fins: All species of Rivulus examined (ex- inal vertebrae, 18 precaudal, and the first cept rogoaguae) have all fins rounded. There proximal anal radial extends just behind the are never any caudal, pelvic, dorsal or anal ribs of the twelfth abdominal. fin extensions as found in the other Neotrop- Among species of Cynolebias, the verte- ical aplocheiloids. Typically, the pectoral bral number is quite variable. In C. melano- 1981 PARENTI: CYPRINODONTIFORM FISHES 383 taenia, there are 12 abdominal, 17 precaudal, However, de Souza (1979) reported meris- and the first proximal anal radial extends just tic data for what were referred to as nine posterior to the ribs of the tenth abdominal randomly chosen individuals of paraguay- vertebra. In C. whitei, the counts are 14 plus ensis, that included two juveniles. The ho- 15, with the anal radial at the rib of the eighth lotype is a female, and it can reasonably be abdominal. In C. elongatus, the counts are assumed that at least one of the nine speci- 16 plus 21, with the anal radial at the rib of mens was a male; therefore, I conclude that the fourteenth abdominal. ornatipinnis and paraguayensis are two Perhaps a derived character is the extreme species which may be easily distinguished on anterior position of the anal fin which would the basis of meristic characters. indicate that dolichopterus is closely related Furthermore, the disparate meristic data to a group of Cynolebias species that in- indicate the nonmonophyletic nature of the cludes whitei. However, in addition to con- genus. Taphorn and Thomerson separated tradicting the derived characters of Cynole- Rachovia from Austrofundulus the basis of, bias not shared by dolichopterus, it conflicts among other characters, fewer dorsal fin rays with another character that may be of signif- which are generally less than 13 (range 9-14) icance in Cynolebias interrelationships; that in Rachovia, as compared with generally 14 is, the number of anal fin rays. In both elon- or more (range 12-18) in Austrofundulus. gatus and whitei there are more than 20; in Although there is range overlap, the increase dolichopterus they range from 15 to 18 in dorsal fin ray numbers in Austrofundulus (Weitzman and Wourms, 1967). The useful- and N. ornatipinnis is consistent with an in- ness of this character can only be determined crease in dorsal fin ray numbers to 15 or by a survey of its states among all species of more in Cynolebias and Terranotus. Cynolebias. The first proximal anal radial is often fused The genus Neofundulus currently contains to the second at its base. Nonetheless it is two species, paraguayensis Myers, the type, present in the genus Rachovia, as well as in and ornatipinnis Myers. Both species are the other more primitive Neotropical genera. known from only a few specimens, and only The radial is absent (fig. 22) in the genus the holotype of each has been examined as Austrofundulus as well as in Terranotus and part of this study. Cynolebias and its included subgenera. Arambaru, Arambaru, and Ringuelit (in de Scales in a Lateral Series: The Neotropical Souza, 1979) suggested the two species be aplocheiloids typically have a high number synonymized. This is opposed by evidence of scales in a lateral series. Determining the that they are not even closest relatives. De polarity of scale number, however, presents Souza (1979) reported meristic data for a re- certain problems since the scale count is not cent collection of paraguayensis, and listed high in all members. Within primitive mem- these along with values for the holotypes of bers of the group, such as R. marmoratus, each species. The number of dorsal fin rays the scale count is over 40, greater than the in paraguayensis ranges from 10 to 13; in more typical number of 30 for cyprinodon- ornatipinnis it is 15. The number of anal rays tiforms as a group. The scale count is high is 12 to 16, and 18, respectively. The number also in some members of the derived genus of scales in the lateral series ranges from 34 Cynolebias, such as C. elongatus, in which to 38 in paraguayensis, and is 37 in ornati- the scales number over 60. There are mem- pinnis. bers of the genus Cynolebias which have The holotype of paraguayensis is a fe- lower counts in the thirties and forties. male, and that of ornatipinnis a male. There- Among all Neotropical aplocheiloids, how- fore, from the type material alone, it is dif- ever, the count is low only in members of ficult to tell whether the differences are due the genus Rachovia, as the genus is delim- to sexual dimorphism alone. (In a group of ited by Taphorn and Thomerson. In fact, species in the genus Cynolebias, males have they gave as a defining character of the genus higher dorsal and anal fin ray numbers.) a lateral series scale count of less than 32, 384 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 22. Diagrammatic representation of anal fin of Auslrofundulus transilis. Unbranched anal rays are blackened, middle and distal anal radials are stippled. lower than any other Neotropical aplochei- The species of Rivulichthys, Trigonectes loid. Taking all other characters into consid- strigabundus, and Rivulus rogoaguae share eration, the high scale count is derived for a pointed snout (caused by reduction of the some group, the limits of which cannot be anterior ramus of the premaxilla) and a de- readily determined at the level of this study. rived color pattern. I propose, therefore, that Therefore, it is logical to conclude that the the two genera and R. rogoaguae be grouped reduced scale count of Rachovia is a derived in Trigonectes, of which Rivulichthys is a character, or else the increased number of junior synonym. scales has occurred several times which is The nominal genus Neofundulus is consid- an unparsimonious assessment of this char- ered to contain one species paraguayensis. acter. The band of pigment on the ventral edge of SUBGROUP DEFINITION AND COMPOSI- the caudal fin in males is a character it shares TION: The assignment of species currently in with Rachovia. However, this character the genus Rivulus to one group or another may very well be part of a transition series requires examination of all species for the from the pigmentation pattern derived for state of the interhyal and number of pelvic aplocheiloid fishes; that is, a band of white rays. Such a survey is out of the scope of the or yellow pigment on both dorsal and ventral present study. Rivulus cylindraceus is the margins of the caudal. Therefore, although type species of the genus; therefore, it will Rachovia and Neofundulus are considered retain the name. This leaves those species of to be included in a monophyletic group with Rivulus with a cartilaginous interhyal with- Auslrofundulus, Terranotus, and Cynole- out a name. I suggest they be referred to as bias, this relationship is represented as an "Rivulus" rather than propose a new name unresolved trichotomy (fig. 20). at this time since the relationship of all Neofundulus ornatipinnis does not have species of "Rivulus''' to the remaining Neo- the pigmentation pattern, yet, as stated, has tropical genera is uncertain, as is, therefore, an elongate dorsal fin. The condition of the the monophyly of the genus. holotype precludes its inclusion in any of the 1981 PARENTI: CYPRINODONTIFORM FISHES 385

recognized genera. Therefore, I propose that They are distinguished from other Neotrop- it be referred to as "Neofundulus" ornati- ical aplocheiloids with a vertical bar through pinnis until additional specimens are avail- the eye and a fourth ceratobranchial covered able. by teeth by having a lateral series scale count Six nominal species have been described of less than 32 (Taphorn and Thomerson, in the genus Pterolebias: longipinnis Gar- 1978). Terranotus, with an unsealed caudal man, zonatus Myers, peruensis Myers, fin and a closed preopercular canal, is hy- bockermanni Travassos, hoignei Thomer- pothesized to be the sister group of an son, and maculipinnis (Weibezahn). They all assemblage of four genera: Cynolebias, possess a dorsal fin of 10 to 12 rays which is Simpsonichthys, Cynopoecilus, and Cam- set back on the body, normally the first ray pellolebias. The members of these four gen- being over the second half of the anal fin; the era all possess rounded caudal fins in both caudal fin is finely scaled for at least one- sexes and have fourth ceratobranchials with- third its length; and, the caudal peduncle is out teeth. They all have been suggested as strongly compressed laterally. These char- synonyms at one time or another (Myers, acters, however, are typical for all species of 1942; Lazara, 1979), and on the basis of their Neotropical aplocheiloids discussed so far, shared characters, I unite them (along with or, as in the case of the compressed caudal Terranotus) within their senior synonym, peduncle, may be an artifact of preservation Cynolebias. (Weitzman and Wourms, 1967). As such, the CLADISTIC SUMMARY OF NEOTROPICAL genus has never been defined as a monophy- APLOCHEILOIDS: Rivulus and other Neo- letic group. Taphorn and Thomerson (1978) tropical aplocheiloids share the derived char- removed maculipinnis from Pterolebias and acters outlined in the previous section: ab- placed it in the genus Rachovia, after Thom- sence of a first postcleithrum; a unique head erson (1974) stated that three species {bock- squamation pattern; the uniting of the oper- ermanni, maculipinnis, and peruensis) are cular and branchiostegal membranes; reduc- probably not closely related to the other tion of the preopercular and dermosphenotic members of Pterolebias. The removal of canals; the medial expansion of the lateral maculipinnis from a close relationship to ethmoids and posterior extension of the longipinnis seems to be a valid decision vomer; and, the triangular process on the based on the following characters: anterior face of the medial arms of the max- Pterolebias longipinnis and zonatus both illa. lack the interarcual cartilage, a lack that may The genus Rivulus is paraphyletic, and its be considered a defining character of Pter- species are referenced to two genera Rivulus olebias. This element is present in maculi- and "Rivulus." pinnis, although apparently reduced relative Members of "Rivulus" share with other to the generalized state for aplocheiloids as Neotropical aplocheiloids, excluding Rivu- previously discussed. lus, a cartilaginous interhyal and seven or The species maculipinnis shares with the more pelvic fin rays. In addition, all species remaining Neotropical aplocheiloids (species in the genus Rivulus are nonannual, whereas assigned to Rachovia, Austrofundulus, Ter- at least one of the genus "Rivulus," stellifer ranotus, Neofundulus, and Cynolebias): a Thomerson and Turner (1973), is annual. vertical bar running from below the eye to "Rivulus" and Rivulus are excluded from near the dorsal surface of the head; thick- a larger group which is defined by an elon- ened anal rays in females; and the tendency gate rostral cartilage, and extensions of the of males to develop a fatty predorsal ridge. pectoral fin rays to the base of the pelvics. Provisionally, Rachovia may remain as Within this larger group, three subgroups are delimited by Taphorn and Thomerson; how- recognized: ever, the four included species brevis, macu- 1. Trigonectes, of which Rivulichthys is lipinnis, pyropunctata, and hummelincki considered to be a junior synonym, is de- might not constitute a monophyletic group. fined by an oblique mouth cleft formed prin- 386 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 cipally by the reduction of the anterior ramus of cyprinodontiform fishes which have either of the premaxilla. been lost or become more derived in other 2. Pterolebias is defined here by its lack genera. These characters include the arm of of the interarcual cartilage. the premaxilla being free rather than embed- 3. A group including Rachovia, Neofun- ded in the skin on the side of the head, and dulus, Austrofundulus, Terranotus and Cy- the swimbladder extending through several nolebias is defined by the following charac- hemal arches rather than ending at the first ters: a vertical bar through the eye often hemal arch. My own analysis of derived extending on to the top of the head; thick- characters of the aplocheiloids and of those ened anal rays in females; and the tendency which define the Old World aplocheiloids, to develop a fatty predorsal ridge in older indicates that Aplocheilus is a relatively de- males. Rachovia may not be definable as a rived aplocheiloid genus. monophyletic group, but is retained here to Aplocheiloids of the Old World are cur- reference its four included species which rently classified in 29 nominal genera and may be distinguished from other members of subgenera. These include Aplocheilus of the group 3 that have teeth on the fourth cera- Indian subcontinent and Laurasia extending tobranchial by its low number of scales in a along the Indo-Australian archipelago to lateral series. Java; Pachypanchax of Madagascar and the As presently defined, Neofundulus is Seychelles; and the African genera Epiplatys polyphyletic. Neofundulus ornatipinnis is and four proposed subgenera; Aphyosemion considered to be more closely related to the and 11 subgenera; Adamas; Nothobranchius more derived aplocheiloids Cynolebias, Aus- and four included subgenera; and Fundulo- trofundulus, and Terranotus on the basis of soma (fig. 23). Together they comprise near- its increase in dorsal fin rays. It is the sole ly 300 species, over 100 of which are referred constituent of the genus "Neofundulus." to the genus Aphyosemion or one of its sub- Terranotus, Austrofundulus, and Cynole- genera. bias lack the first proximal anal radial; have The definition of the Old World aplochei- a large spine on the first vertebra; and have loids as a monophyletic group again supports reduced dentition on the fourth ceratobran- the contention that annualism is a lifestyle chial. which has either arisen at least twice within Austrofundulus is defined by its darkly pig- cyprinodontiform fishes, or is a characteris- mented anal papilla. tic that in some sense is basic to all members. Terranotus and Cynolebias have a caudal CHARACTER ANALYSIS: Shoulder Girdle: fin which is not finely scaled and lack a pre- The Old World aplocheiloids are distin- opercular canal. A group of species has a guished from all other killifishes by the fu- unique head neuromast pattern and an in- sion of the posttemporal and supracleithrum crease in the number of dorsal and anal fin (fig. 7C) to form one slender bone connecting rays in males. These two characters define the shoulder girdle to the skull. The fusion subgroups of Cynolebias. Since there is no is complete, and no joint lines are visible. In longer reason to maintain Terranotus as a the Neotropical aplocheiloids, the posttem- monotypic genus, I propose that this genus poral and supracleithrum are similarly be considered synonymous with Cynolebias. shaped, however, the two bones may always be separated easily. Gill Arches: Two derived characters of the OLD WORLD APLOCHEILOIDS gill arches distinguish the Old World aplo- cheiloids. One is the reduction of the basi- Members of the aplocheiloids of the Old World have often been referred to as the hyal to a small triangular-shaped bone which is capped by a large cartilaginous wedge (fig. most primitive of all cyprinodontiforms. Myers (1958) stated that the genus Aplochei- 11 A). As stated, in all aplocheiloids, the ba- sihyal is very wide, and forms the basis of a lus represents the most basic characteristics wide "tongue" which is visible upon opening 1981 PARENTI: CYPRINODONTIFORM FISHES 387

FIG. 23. Distributional limits of Old World aplocheiloids. the mouth. In the Neotropical aplocheiloids, Within the Old World aplocheiloids, the as well as in the cyprinodontoids and other premaxillary ascending processes are ta- atherinomorphs, the basihyal is ossified for pered posteriorly to form, in the most de- more than half its length. rived state, the greatly expanded triangular A second character concerns the attach- processes of Aplocheilus (fig. 4A). In Pachy- ment of the interarcual cartilage to the sec- panchax (fig. 4B) and Epiplatys and its in- ond pharyngobranchial. The Neotropical cluded subgenera, the processes are also ex- aplocheiloids exhibit the primitive state for panded, although not to the degree exhibited cyprinodontiforms; that is, the cartilage at- by Aplocheilus. taches to a small flange of bone lateral to the In Aphyosemion (fig. 4C), Nothobran- cartilaginous extension of the pharyngobran- chius and Fundulosoma, the processes are chial (fig. 6A) as it does in the aplocheiloids. tapered posteriorly, but never as widely ex- In the Old World aplocheiloids, the carti- panded as in any of the three above men- lage attaches in the same position, but the tioned genera. bony flange is absent; thus, the cartilage nearly abuts the cartilaginous articulation SUMMARY OF DERIVED CHARACTERS point of the pharyngobranchial (fig. 24A). 1. Supracleithrum fused to posttemporal. In the genus Nothobranchius, the carti- 2. Small, triangular basihyal capped by a lage attaches directly to the cartilaginous wedge of cartilage. head of the pharyngobranchial (fig. 24B). This character is apomorphic for the genus. 3. Interarcual cartilage attached directly to Premaxillary: The premaxillary ascending lateral face of the second pharyngobran- processes are flat and broad in the Neotrop- chial. 4. Premaxillary ascending processes ta- ical aplocheiloids and in Profundulus. This pered. state is most parsimoniously assessed as the most primitive among cyprinodontiforms as RELATIONSHIPS OF OLD WORLD AP- discussed in the section on derived charac- PLOCHEILOIDS: The Old World aplochei- ters of the group. loids, like the Neotropical aplocheiloids, 388 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

PB3

FIG. 24. Diagrammatic representation of dorsal view of dorsal gill arches of A. Aplocheilus panchax, B. Nothobranchius melanospilus. Cartilage is stippled.

have been treated as a natural group of fishes The procatopines do not possess any of (e.g., Scheel, 1968) although their monophy- the derived characters of Old World aplo- ly had never been tested. In addition to con- cheiloids summarized above; they possess sidering Aplocheilus as the most primitive only one of the derived characters for cyprinodontiform (e.g., Myers, 1958), var- aplocheiloids as a whole, the cartilaginous ious authors suggest that the annual Old mesethmoid. This condition is considered to World aplocheiloids are the closest relatives be independently derived in these two of the Neotropical aplocheiloids. groups, as well as in Anatolian cyprinodonts. Furthermore, the Old World aplocheiloids Procatopines, however, do share a series were considered by many workers to be of unique features with the rest of the cy- closely related to the procatopines, the other prinodontoids. Thus, procatopines and aplo- group of cyprinodontiforms which inhabits cheiloids are not considered together further. subsaharan Africa. Ahl (1924, 1928), for ex- Taxonomic revisions of Old World aplo- ample, grouped the two without questioning cheiloids, like those of New World forms, their monophyly; more recently, Huber place emphasis on the recognition of differ- (1979) described a new genus and species, ences among taxa rather than on the discov- Adamas formosus, which he considered to ery and description of derived characters be intermediate between Old World aplo- shared among taxa. Emphasis on differences cheiloids of the genus Aphyosemion and the has led to the naming of four subgenera of procatopines. Such conclusions were a prod- Epiplatys, four subgenera of Nothobran- uct of the ambiguous definitions previously chius, and the division of Aphyosemion into put forth for the subfamilies Rivulinae and 13 genera and subgenera. Aplocheilichthyinae. Nevertheless, as In a recent paper by Radda (1977), four understood here, these two distributionally new subgenera are named, each to encom- similar groups have little more than primitive pass a group of species referable to Aphy- characters in common. osemion. Radda included a phylogenetic tree 1981 PARENTI: CYPRINODONTIFORM FISHES 389 which purports to summarize the relation- would allow for the reference of a particular ships of subgenera within the genus. The species to one monophyletic group or monophyly of Aphyosemion is doubtful, fol- another, and avoid the confusion created by lowing Radda's diagram, for the subgenus the current generic limits. For example, Pronothobranchius and the genus Fundulo- Aphyobranchius janpapi is either more soma, considered here as close relatives, are closely related to species of Nothobranchius included as more closely related to some or to some group of Aphyosemion. A concise subgenera of Aphyosemion than they are to definition of each group would allow such a each other. Furthermore, the genus Notho- decision to be made. branchius is not considered at all; therefore, As for the Neotropical aplocheiloids, this it is unclear whether the implication is that analysis does not include the revision of all Nothobranchius is in turn most closely re- species of each genus. Rather, it is an at- lated to Fundulosoma or Pronothobranch- tempt to identify and define on the basis of ius, to some subgroup of Aphyosemion, or shared derived characters the major mono- whether it need be considered at all in a re- phyletic groups of species and their proposed vision of Aphyosemion. interrelationships (given in the cladogram of The genus Aphyosemion is large, current- fig. 25) which will eventually lead to the def- ly comprising over 110 species; if some mem- inition of monophyletic genera. bers are more closely related to Nothobran- Supraspecific categories of Old World chius species then the two genera must be aplocheiloids may be divided into two major considered together in a phylogenetic anal- monophyletic groups. One is referred to as ysis. If the two genera do not form a mono- the Aphyosemion-Nothobranchius group; phyletic group, then Nothobranchius need and the second is referred to as the Aplo- not be considered in a study of the interre- cheilus-Pachypanchax-Epiplatys group. The lationships of Aphyosemion and its subgen- interrelationships of the members of the two era. The problem of defining monophyletic are discussed separately with reference to genera and subgenera extends to all mem- the states of characters in the other, in the bers of the Old World aplocheiloids. Scheel Neotropical aplocheiloids, and in cyprino- (1972) has recommended that the genera dontiforms as a whole. Epiplatys and Aplocheilus be synonymized. Clausen (1967) has named a new subgenus of THE Aphyosemion-Nothobranchius GROUP Epiplatys to included the species E. duboisi; a subgenus Aphyoplatys is named to indicate Species in this group are currently classi- that this species is intermediate between fied in four genera and fifteen subgenera. Aphyosemion and Epiplatys. Wildekamp These are Aphyosemion Myers with eleven (1977) has recently named a new subgenus subgenera: Archiaphyosemion Radda, Me- of Nothobranchius to include the species N. soaphyosemion Radda, Kathetys Huber, janpapi; it is named Aphyobranchius to re- Fundulopanchax Myers, Paludopanchax flect its intermediacy between Aphyosemion Radda, Chromaphyosemion Radda, Callo- and Nothobranchius. panchax Myers, Raddaella Huber, Diapter- It is of little use, however, to know that all on Huber and Seegers, Paraphyosemion the nominal genera of Old World aplochei- Kottelat and Gularopanchax Radda; No- loids grade into one another from the thobranchius Peters, with four subgenera: Aplocheilus type to the Nothobranchius Adiniops Myers, Pronothobranchius Radda, type. Logically, there is no reason not to Zonothobranchius Radda, and Aphyobran- classify all the species in one genus; how- chius Wildekamp; and Fundulosoma Ahl ever, that too would be avoiding the problem and Adamas Huber, two monotypic genera. of the interrelationships of the included Most of these names are unfamiliar to the species as much as if each species were put majority of workers on cyprinodontiform into its own genus. Unambiguous definitions fishes since nearly all have been just recently of monophyletic groups of nominal genera described in journals that are not widely dis- 390 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 25. Cladogram of relationships of Old World aplocheiloids. Taxa treated as subgenera not included in diagram (see text for further information). Node A: posttemporal fused to supracleithrum; reduction of basihyal to small, triangular wedge; interarcual cartilage attached to second pharyngo- branchial which lacks a bony flange; premaxillary processes tapered posteriorly; Node B: broad, flat- tened upper jaw caused by expanded premaxillary ascending processes; expanded coronoid process on the dentary; bifurcate upper hypural plate in juveniles and some adults; loss of the uncinate process on fourth epibranchial; Node C: lower limb of posttemporal represented by ligament; teeth on third and fourth hypobranchials; dorsal ocellus in females; orbital rim attached only ventrally; darkened caudal fin margin; Node D: premaxillary ascending processes expanded medially and overlapping in the midline; attenuate lower jaw; Node E: hypural fan in adults; lateral scales of male angled away from body; Node F; bifid epipleural ribs; reduced chromosome number; attenuate posterior extension of the vomer; Node G: dorsal fin rays fourteen or more; dorsal origin opposite anal origin; swimbladder not expanded past first arch; Node H: preopercular canal open, not represented by pores; attachment of the interarcual cartilage directly to the second pharyngobranchial; oval eggs.

tributed publicly. Nonetheless, they repre- examination, they are easily determined as sent available supraspecific categories within bifid. For example, in Nothobranchius or- the Aphyosemion-Nothobranchius group; thonotus, the type of the genus, the first six therefore, their references are summarized epipleural ribs are unambiguously bifid. This in the systematic accounts, and they are con- character does not appear to be related to sidered herein. the size or age of specimens, nor is any sex- CHARACTER ANALYSIS: Bifid Epipleural ual dimorphism apparent. Ribs: Bifid epipleural ribs have been used to Vomer: The vomer typically has a broad distinguish species of the genus Aphyose- posterior extension in cyprinodontiform fish- mion from those of the genus Nothobran- es (e.g., in Aplocheilus panchax, fig. 17B). chius. Typically, in Aphyosemion, the first In contrast, in members of the Aphyose- five or six epipleural ribs are strongly bifid mion-Nothobranchius group the posterior distally (fig. 26). This derived character is extension of the vomer is narrow (fig. 17A). unambiguously present in all species of This sets off the anterior extension of the Aphyosemion examined, including A. peter- vomer as a large rectangular element. si, a species which has alternately been Chromosome Number: Fishes of this placed in the genus Aphyosemion, and in group exhibit some of the lowest chromo- Epiplatys. On this basis, peter si should some numbers known for teleost fishes. Te- properly be placed in the Aphyosemion- leosts generally have a haploid chromosome Nothobranchius group. number of 24, and therefore a diploid number In some species of Aphyosemion, for ex- of 48. Gyldenholm and Scheel (1971) listed ample, A. gulare and sjoestedti, and in haploid and diploid chromosome numbers of Nothobranchius, the epipleural ribs are temperate and tropical freshwater fishes in often not as strongly bifid as in most of the 19 families. Included were representatives of species of Aphyosemion; however, on close the percomorph, ostariophysan, atherino- 1981 PARENTI: CYPRINODONTIFORM FISHES 391

FIG. 26. Diagrammatic representation of the posterior region of the skull and attachment of first vertebra, lateral view, of Aphyosemion occidentale. Posttemporal removed. morph, and paracanthopterygian lineages. In species of Aphyosemion (which in- Karyotypes of 53 species within the family cludes the species placed in Roloffia in Gyl- Cyprinodontidae (including Oryzias latipes) denholm and Scheel) and Nothobranchius, and those of 19 species of poeciliid fishes the number ranges from nine to 23 (Scheel, were listed. Within the cyprinodontiforms, 1968). Among the nominal species of Epi- as well as all teleosts, the usual haploid chro- platys, the haploid chromosome number mosome number is 24. In poeciliids the num- ranges from 17 to 25; it is 24 for the type ber ranges from 23 to 25; in cyprinodonti- species of the genus, E. sexfasciatus. forms from nine to 25. Scheel (1968) stated Among the nominal species of Aplocheilus, that only among the aplocheiloid fishes with- the haploid number ranges from 18 to 25; it in the family Cyprinodontidae did the hap- is 18 for the type of the genus, A. panchax. loid chromosome number reach 25; there- Pachypanchax playfairi is reported to have fore, he concluded that this is perhaps the 24 haploid chromosomes, as does Aphyopla- primitive number for aplocheiloids. In light tys duboisi. of present findings concerning the non- Scheel (1968) maintains that the type of monophyletic nature of the cyprinodontids, chromosome reduction differs in the genera inclusion of the poeciliids indicates that (1) Epiplatys and Aplocheilus from that in the the haploid number of 25 is attained in cy- Aphyosemion-Nothobranchius group. That prinodontiforms other than aplocheiloids, is, in the former genera, the reduction in- and that (2) it is logical to conclude on the volves the production of large metacentric basis of outgroup comparison that n = 24 is elements with a subsequent loss of the small- basic for aplocheiloids, as well as cyprino- er metacentrics. In the latter genera, the re- dontiforms as a whole. duction involves the production of large ac- 392 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 rocentric elements, accompanied by loss of branchius group than to Epiplatys, or else the smaller elements. White (1968 in Scheel) the character of bifid epipleural ribs is de- maintained that such so-called superacrocen- rived for Old World aplocheiloids. I suggest trics could arise from pericentric inversions that no synonymies of Old World genera be in metacentric elements from the same size. undertaken unless the generic limits are for- While this could account for the difference mally defined in terms of shared derived in karyological morphology in these two characters drawn from a survey of all species groups of Old World aplocheiloids, Scheel for such characters. correctly maintains that there are no indica- Adamas formosus (new genus and species tions the superacrocentrics were produced in described by Huber, 1979) has not been ex- this manner. amined. It is placed in this group on the basis In one species of Neotropical aplochei- of overall external morphology, color pattern loids, Cynolebias (Cynopoecilus) melano- and sexual dimorphism as noted from a pho- taenia, the superacrocentrics occur and the tograph included in the description. It ap- haploid chromosome number of the current pears to be most closely related to the prim- aquarium strain is 22 (Scheel, 1968). itive species of Aphyosemion as described The occurrence of superacrocentrics with- below. Thus, I cannot accept Huber's sug- in another group of aplocheiloid fishes indi- gestion that Adamas is intermediate between cates that they are not unique to fishes of the the procatopines and Old World aplochei- Aphyosemion-Nothobranchius group, and loids. Its precise placement within one of the also that chromosome reduction is not lim- existing supraspecific subdivisions of Aphy- ited to the Old World aplocheiloids. There- osemion will require an examination of ma- fore, the division of Old World aplocheiloids terial. into two groups on the inferred mode of re- Annualism: Annualism means that the fer- duction is suspect since the polarity of this tilized egg and embryo exhibit diapause. The reduction cannot be determined. We are left species included in the Aphyosemion-No- with the character of reduction of chromo- thobranchius group are both annual somes which is useful in a phylogenetic anal- and nonannual. All members of the Aplo- ysis if it can be correlated with characters cheilus-Pachypanchax-Epiplatys group are from a presumed independent source. Such nonannual. The nonannual members of is the case with the bifid epipleural ribs and Aphyosemion are Archiaphyosemion, Me- slender posterior extension of the vomer soaphyosemion, Kathetys, Chromaphyose- used here. mion, Diapteron, Aphyosemion, and Pare- piplatys. The annual species are in SUMMARY OF DERIVED CHARACTERS Raddaella, Paraphyosemion, Paludopan- chax, Gularopanchax, Callopanchax, Fun- 1. Bifid epipleural ribs. dulopanchax, as well as the subgenera of 2. Attenuate posterior process of the vomer. Nothobranchius and in Fundulosoma. 3. Reduced chromosome number. Members of the genus Chromaphyosemion have been referred to as semiannual (e.g., RELATIONSHIPS OF THE Radda, 1979) because the eggs were ob- Aphyosemion-Nothobranchius GROUP served to tolerate partial drying in the field. Among those species of Epiplatys with However, since all annuals, including so- low haploid numbers, the following have called true annuals such as Austrofundulus been examined and possess weakly bifid transilis can be water-incubated (Wourms, epipleural ribs: E. bifasciatus (n = 20) and 1972a) it is perhaps more appropriate to refer E. spilargyreia (n = 17). Thus, it appears to the semiannual species as nonannual un- that on the basis of these two characters cer- less diapause can be demonstrated. Other- tain species of Epiplatys may be more wise nothing more than the tolerance of des- closely related to the Aphyosemion-Notho- iccation has been demonstrated. 1981 PARENTI: CYPRINODONTIFORM FISHES 393

In addition to being annual, species of the whereas in the closed pattern the two neu- genera Fundulosotna and Nothobranchius romasts lie separated by a ridge of epidermis. have oval rather than the more typical round Scheel (1968) published photographs of the eggs of other aplocheiloids (Scheel, 1968). two conditions, and used this character to Swimbladder: The swimbladder of the place Aphyosemion petersi in the genus Cal- aplocheiloids typically extends posteriorly lopanchax, along with the more apomorphic beyond several hemal arches, as in the occidentale. Radda (1977) recognized the ap- Aplocheilus-Pachypanchax-Epiplatys group parent unnatural status of Callopanchax as and Aphyosemion petersi. Failure of the thus constituted and placed petersi in his swimbladder to extend beyond more than the Archiaphyosemion, where on the basis of first pair of hemal arches in Paludopanchax, the above characters, it more properly be- Gularopanchax, Callopanchax, Fundu- longs. lopanchax, Raddaella, Paraphyosemion, The closed frontal neuromast pattern was Nothobranchius, and Fundulosotna is inter- used again by Clausen (1967) to separate the preted as a derived character of the included species of his Parepiplatys from the rest of species. the Epiplatys species. Scheel (1968) reports Dorsal Fin Position and Ray Number: The that in a brood of Pachypanchax playfairi, most primitive position of the dorsal fin for some individuals developed with the open aplocheiloids is inferred to be the general pattern and some with the closed. Thus, the condition for Old and New World genera in character seems of doubtful significance for which the dorsal fin is set back on the body a phylogenetic study, and therefore, fails as approximately opposite the last third of the a defining character of Callopanchax. anal fin. Dorsal fin rays typically number The preopercular canal is present in all Old seven to 10, though the number can be slight- World aplocheiloids and typically opens to ly higher. These primitive conditions occur the outside by a series of pores (fig. 13A). In in the following subgenera of Aphyosemion: the subgenera of Nothobranchius and in Archiaphyosemion, Mesoaphyosemion, Fundulosoma thierryi, there are no pores, as Kathetys, and Aphyosemion, as well as the canal is open to the outside all along the Adamas. A second group of subgenera have margin of the preoperculum. a dorsal fin which is slightly elongate (gen- SUBGROUP DEFINITION AND COMPOSI- erally from 10-14 rays) and situated over the TION: I conclude that (1) the genus Aphy- first quarter of the anal fin. In this group are osemion as currently constituted is not the subgenera Chromaphyosemion and monophyletic; and (2) the annual species Diapteron. previously assigned to Aphyosemion are A third group of Aphyosemion subgenera most closely related to the species of Noth- (Paludopanchax, Gularopanchax, Callopan- obranchius and Fundulosoma. Annualism is chax, Fundulopanchax, Raddaella, and Par- thus postulated to have arisen just once with- aphyosemion) and Fundulosoma and Noth- in the Old World aplocheiloids, as may also obranchius have an elongate dorsal fin of be true of Neotropical aplocheiloids. over 14 dorsal fin rays the origin of which is Division of species of the Aphyosemion- opposite the anal origin. Nothobranchius group on the basis of dorsal Cephalic Sensory Pores and Squamation: fin position and ray number is problematic. One character which does not seem to be The position of the dorsal fin is variable even useful in separating these subgenera into among individuals of the same species. How- groups is the open versus closed frontal neu- ever, the species of Aphyosemion may be romas! pattern. Clausen (1966) first used the grouped artificially into two categories; those closed pattern as a defining character of a with from seven to 14 dorsal fin rays and the new genus Roloffia (^Callopanchax Myers). dorsal situated no farther forward than op- In the closed pattern, the two frontal neu- posite the first quarter of the anal fin; and romasts are encircled by a rim of epidermis, those with more than 14 dorsal fin rays and 394 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 the dorsal situated over the anal fin origin or onymy of the subgenera is suggested at this just slightly before or after. time. Species groups with a posterior dorsal fin THE Aplocheilus-Pachypanchax- and low dorsal-fin ray number are nonannual Epiplatys GROUP and have a swimbladder extending beyond the first hemal arch. They include Aphyose- Species in this group are currently classi- mion, Archiaphyosemion, Mesoaphyose- fied in three genera (Aplocheilus, Pachypan- mion, Chromaphyosemion, Diapteron, and chax and Epiplatys). Epiplatys, in turn, is Kathetys. Since they share only primitive divided into four subgenera (Lycocyprinus, characters, these subgenera are not consid- Parepiplatys, Aphyoplatys, and Pseudepi- ered to form a monophyletic group. I suggest platys). that they be referred to the genus Aphyose- CHARACTER ANALYSIS: Jaw: Typically, mion, however, they will not be formally the head is greatly flattened, as is the upper synonymized with the genus since the mono- jaw, resulting in a dorsal profile which has phyletic nature of the group is not implied. been referred to as pikelike. (When first de- Species groups with an anterior dorsal fin scribed, Aplocheilus panchax was placed in and high fin ray number are annuals that the genus Esox.) The flattened upper jaw is have a swimbladder which does not extend represented internally by broadly expanded posteriorly beyond the first hemal arch. They premaxillary ascending processes (fig. 4A). include Raddaella, Paludopanchax, In addition to this upper jaw characteristic Gularopanchax, Callopanchax, Fundulo- there is a concordant feature of the lower jaw panchax, Paraphyosemion, Fundulosoma, which contributes to the flattened appear- and Nothobranchius. ance of the mouth. As illustrated for Aplo- Species of the genera Fundulosoma and cheilus panchax (fig. 27) there is a unique, Nothobranchius both share the derived state large coronoid process on the dentary which of the interarcual cartilage as described, oval overlaps the dorsal extension of the articu- eggs, and an open preopercular canal. The lar. There is no such process in the Aphyose- sole species of Fundulosoma may be distin- mion-Nothobranchius group (e.g., fig. 31C). guished from all species included in Notho- Caudal fin: The internal supports of the branchius by the forked posttemporal, and caudal fin differ among adults of the three the caudal fin extensions of the males. How- genera although they are similar in juveniles. ever, since Fundulosoma is monotypic, In Aplocheilus, the upper hypural plate is there is no reason to separate it from the rest divided in two (fig. 2D). In Epiplatys, the of the Nothobranchius species. Therefore, upper and lower hypural plates are separate I consider it to be a junior synonym of Noth- and never fused together to form an hypural obranchius. It may be considered as the fan. In at least one species, E. sexfasciatus, most primitive Nothobranchius species. there is evidence of a line of division in the The remaining subgenera of Aphyosemion upper hypural plate, suggesting the division have no generic reference if they are exclud- seen in species of Aplocheilus. ed from Aphyosemion and hypothesized to In adult Pachypanchax, the hypural plates be more closely related to the species of are fused to form an hypural fan, as is the Nothobranchius as defined above. Among case in Nothobranchius, Fundulosoma, and the names of subgenera within this group, some species of Aphyosemion, a group of Fundulopanchax Myers is the oldest and Neotropical aplocheiloids, and most, but not therefore the name which will be used to ref- all, the cyprinodontoids. However, in juve- erence the annual Aphyosemion species. nile lab-reared Pachypanchax playfairi the However, it is not implied that this group dorsal and ventral hypural plates have an ev- itself is monophyletic since some members ident joint, and there is also such a suture may be more closely related to Nothobran- between the dorsal and ventral portions of chius than to each other. Therefore, no syn- the upper hypural plate. In Fundulus majal- 1981 PARENTI: CYPRINODONTIFORM FISHES 395 is, a funduline, and in Nothobranchius several groups of cyprinodontiforms, the guentheri, species in which adults have a hy- lower limb of the posttemporal extending to pural fan, the juveniles possess a hypural the exoccipital is unossified, and represented fan, even at the stage of a cartilaginous pre- only by a ligament. Within the aplocheiloids, cursor of the hypural elements. Therefore, this occurs in the genera Aplocheilus and given that the aplocheiloids form a mono- Pachypanchax and in Nothobranchius. It is phyletic group, the separate hypurals of the fully forked in all species of Epiplatys and Aplocheilus -Pachypanchax-Epiplatys group Aphyosemion examined, as well as in Fun- are an indication of a secondarily derived dulosoma thierryi. The lower limb being rep- condition. resented by an unossified ligament is most Dorsal Gill Arches: The Aplocheilus- parsimoniously assessed as independently Pachypanchax-Epiplatys group exhibits a derived in Aplocheilus and Pachypanchax derived feature of the dorsal gill arches. Typ- and Nothobranchius. ically among cyprinodontiforms, an uncinate Hypobranchial Teeth: Both Aplocheilus process from the third epibranchial articu- and Pachypanchax have patches of teeth on lates via a cartilage with a corresponding the second and third pair of hypobranchials, process on the fourth epibranchial. The un- as well as on the fourth ceratobranchials. cinate process of the fourth epibranchial, Such teeth are typically found on the fourth however, is absent in these three genera. The ceratobranchials of atherinomorphs except fourth epibranchial is present as a slender when lost or reduced as in a group of the element (fig. 24A) which has no point of ar- Neotropical aplocheiloids. Teeth on the hy- ticulation to the third. pobranchial elements, however, have not been found except in these two genera of SUMMARY OF DERIVED CHARACTERS aplocheiloids and in two cyprinodontoid gen- era Anableps and Oxyzygonectes. Thus, the 1. Broadly expanded premaxillary ascend- presence of hypobranchial teeth is consid- ing processes. ered to be independently derived in these 2. Coronoid process on dentary overlaps two cases. dorsal extension of articular. Dorsal Ocellus: A dorsal ocellus is present 3. Separate upper hypurals at least in juve- in all females of Aplocheilus and Pachypan- niles. chax playfairi. The ocellus is developed also 4. Loss of the uncinate process on the fourth in males of several species of Aplocheilus epibranchial. such as in A. panchax. The dorsal ocellus is Aplocheilus-Pachypanchax-Epiplatys absent in all other Old World aplocheiloids. GROUP RELATIONSHIPS: On the basis of the The genus Pachypanchax contains two following characters, I conclude that Pachy- species, playfairi and homalanotus. The panchax and Aplocheilus are more closely dorsal ocellus is reported to be absent from related to each other than either is to Epi- both males and females of the latter species platys; therefore, placing Epiplatys in syn- (Scheel, 1968). Only one specimen of hom- onymy with Aplocheilus (Scheel, 1972; Rad- alanotus was examined, and the species' da, 1973) and excluding Pachypanchax continued placement in Pachypanchax would create a paraphyletic genus. should perhaps be investigated. Posttemporal: The posttemporal is typi- Orbital Rim: As discussed for the defining cally a forked bone attaching distally to the characters of the aplocheiloids, Aplocheilus supracleithrum and proximally to the epiotic and Pachypanchax have an orbital rim dorsally and the exoccipital ventrally. In Old which is attached ventrally and folded under World aplocheiloids, the supracleithrum is the frontals dorsally. This is in contrast to not a distinct element, thus the posterior ex- the condition in all other aplocheiloids in tension of the posttemporal-supracleithrum which the orbital rim is attached all along its attaches directly to the cleithrum. Among perimeter. 396 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 27. Diagrammatic representation of upper and lower jaw of Aplocheilus panchax, lateral view. Maxilla is stippled.

Caudal Fin Margin: In Pachypanchax The species of Epiplatys considered to be playfairi and a number of species of Aploch- part of this monophyletic group may be dis- eilus, including panchax, there is a dark line tinguished only by its lack of the derived of pigment on the caudal fin. Such a margin characters present in Pachypanchax and is not found elsewhere within the aplochei- Aplocheilus. Epiplatys has a forked post- loids, and as such it is considered uniquely temporal, a completely attached orbital rim, derived. and lacks a dorsal ocellus, darkened caudal SUBGROUP DEFINITION AND COMPOSI- fin margin and teeth on the second and third TION: Pachypanchax may be distinguished hypobranchials. Epiplatys, therefore is not from the species of Aplocheilus and Epiplat- definable as a monophyletic group; it may ys by lateral scales in males which are angled eventually be restricted to the type species, away from the body, and by the fusion of the sexfasciatus Gill, and closely allied species. hypural plates in adults into an hypural fan. CLADISTIC SUMMARY OF OLD WORLD The former character refers to a long-known APLOCHEILOIDS: Old World aplocheiloids feature of playfairi. The scales stand away are divisible into two groups. The Aplo- from the body in live adult males and give cheilus-Pachypanchax-Epiplatys group is the impression that the individual is suffering distinguished from the other Old World aplo- from dropsy. cheiloids by the following derived charac- Aplocheilus may be distinguished from the ters: a broad, flattened, upper jaw effected species of Pachypanchax and Epiplatys by by expanded premaxillary ascending pro- an attenuate lower jaw and medially greatly cesses and an expanded coronoid process on expanded premaxillary ascending processes. the dentary, a bifurcate upper hypural plate 1981 PARENTI: CYPRINODONTIFORM FISHES 397 in juveniles and some adults; and the loss of possess a dorsal fin of from seven to 14 rays the uncinate process on the fourth epibran- which is situated no farther anterior than op- chial. posite the first quarter of the anal fin origin, Aplocheilus and Pachypanchax are as- and have a swimbladder which extends pos- sessed as sister genera on the basis of the teriorly to the first one or two hemal spines. following derived characters: teeth on the The Fundulopanchax group comprising second and third hypobranchials; lower limb the subgenera Fundulopanchax, Gularopan- of the posttemporal represented by an unos- chax, Raddaella, Callopanchax, Paraphy- sified ligament; a dorsal ocellus in females; osemion, and Paludopanchax which shares an orbital rim attached only ventrally; and a with the species of Nothobranchius and dark caudal fin margin. Fundulosoma the following derived charac- Pachypanchax is defined by two derived ters: dorsal fin rays increased to 14 or more; characters: fusion of the upper and lower dorsal situated opposite the anal fin origin or hypural plates into a hypural fan in adults; just slightly in front or behind the origin; and and, lateral scales of males angled away from swimbladder not expanded past the first the body. hemal arch. All included species are annual. Aplocheilus is defined by an attenuate low- Monophyly of Aphyosemion and Fundulo- er jaw and premaxillary ascending processes panchax is not implied. expanded medially and overlapping. Fundulosoma and Nothobranchius share The genus Epiplatys as recognized here the following derived characters: preoper- cannot be defined as a monophyletic group. cular canal open, not represented by pores; Some species currently referred to the group a derived position of the interarcual cartilage may prove to be more closely related to and oval eggs. The species of Nothobran- forms of Aphyosemion and Nothobranchius. chius and its included subgenera may be sep- The Aphyosemion-Nothobranchius group arated from Fundulosoma thierryi on the ba- is defined by the following derived charac- sis of the following derived characters: lower ters: bifid epipleural ribs; attenuate posterior limb of posttemporal represented only by an expansion of the vomer; and a reduced chro- unossified ligament and all fins rounded with mosome number. no caudal fin extensions. However, thierryi The subgenera of Aphyosemion may be is considered to be the primitive member of grouped into the following two categories: the genus Nothobranchius since the recog- The Aphyosemion group comprising the nition of a monotypic genus at this position subgenera Aphyosemion, Archiaphyose- in the phylogenetic analysis is uninformative mion, Mesoaphyosemion, Kathetys, Diap- with respect to the interrelationships of in- teron, and Chromaphyosemion, and the ge- cluded species. nus Adamas. They are all nonannual,

CYPRINODONTOIDS (GROUP C) The cyprinodontoids as the term is used in gether as a group without including the this study refers to the fishes of the four vi- aplocheiloids. However, together they form viparous families, the Poeciliidae, Goodei- one of the most well-corroborated monophy- dae, Jenynsiidae and Anablepidae, and the letic groups of fishes. cyprinodontid subfamilies Fundulinae, Lam- Together these groups comprise nearly 400 prichthyinae, Fluviphylacinae, Cyprinodon- species, slightly less than its sister group, the tinae, Aplocheilichthyinae, Orestiatinae and aplocheiloids. Their diversity includes ovi- Pantanodontinae (see table 2). The subgroups parity, ovoviviparity to viviparity; unicus- are referred to using the vernacular names pid, bicuspid, tricuspid, or no teeth in the as defined previously. Prior to this study, jaws; and a size range from the diminutive these fishes have not been considered to- male formosa of the poeciliid 398 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

fishes which matures at a standard length of most teleosts, the hyoid bar is composed an- approximately 8 mm. (Rosen and Bailey, teriorly of two hypohyals. The two elements, 1963) to the large females of the viviparous a dorsal and a ventral hypohyal, articulate Anableps which reach a standard length of with the anterior process of the anterior cer- over 300 mm. (Miller, 1979). They are found atohyal. Typically among aplocheiloids, and in fresh, brackish and salt water, and are dis- most other cyprinodontiforms, there is an tributed pantropically as well as in temperate extension of the anterior ceratohyal under Laurasia from and as far east the ventral hypohyal. This is the case as il- as Iran. lustrated for Pachypanchax playfairi (fig. CHARACTER ANALYSIS: Gill Arches: Sev- 28 A). eral derived characteristics of the gill arches In all cyprinodontoids, the dorsal hypo- distinguish the cyprinodontoids from the hyal is absent. The anterior ceratohyal typi- aplocheiloids and all other atherinomorph cally retains its anterior extension under the fishes. The first of these is the presence of ventral hypohyal, as in Oxyzygonectes dowi just two basibranchials in the ventral gill arch (fig. 28B). However, in the poeciliid fishes, skeleton. In aplocheiloids, as in all other ath- Fluviphylax and procatopines there is no dis- erinomorphs, as well as most other acan- tinct anterior extension of the anterior cera- thopterygian fishes, there are three ossified tohyal, and the remaining ventral hypohyal basibranchials. These lie medially in a is present as a cap of bone over the end of straight line behind the basihyal and extend the anterior ceratohyal as in Procatopus gra- posteriorly to the angle created by the fifth cilis (fig. 28C). In Pantanodon madagascar- ceratobranchials (fig. 11). The basihyal and iensis, there is no extension of the anterior basibranchials are initially represented in on- ceratohyal under the ventral hypohyal; how- togeny by a continuous rod of cartilage ever, there appear to be two ossification cen- known as the copula. This precursor is re- ters in the cap of cartilage present on its an- placed in ontogeny by separate basihyal and terior face. These would probably be basibranchial ossifications. In the aplochei- interpreted as a dorsal and a ventral hypo- loids, as illustrated for Nothobranchius me- hyal; however, in the light of the other evi- lanospilus (fig. 11 A), the basihyal is followed dence which clearly places Pantanodon as posteriorly by three ossified basibranchials. a member of the cyprinodontoids with a de- In contrast, within all cyprinodontoids, the rived state of the anterior ceratohyal, I in- first ossified basibranchial is absent, whereas terpret the cartilaginous cap with its two os- the second and third are present in much the sification centers as a secondarily derived same position as those of the aplocheiloids condition which is most like that described (fig. 11B). for the poeciliids, procatopines and Fluvi- Two ossified basibranchials occur else- phylax. where in the acanthopterygian fishes, nota- In the aplocheiloids, the typical state of bly in synbranchid eels. Rosen and Green- the interarcual cartilage is as an elongate rod wood (1976) report that the condition of two approximately equal in length to the epibran- ossified basibranchials is effected by the fu- chials. It is absent among aplocheiloids in the sion of the first basibranchial with the basi- genus Pterolebias and was found reduced in hyal. In the cyprinodontoids, however, there Rachovia maculipinnis. In all cyprinodon- is no such apparent fusion of the first basi- toids, the interarcual cartilage is reduced to branchial to either the basihyal or the second approximately one half the length of the epi- basibranchial. In addition, the section of the branchials (fig. 6B). (The reduced condition cartilaginous precursor of the first basi- in maculipinnis is considered secondarily branchial is absent in adult cyprinodontoids; derived within the aplocheiloids.) thus, the condition of the two ossified basi- Jaw and Jaw Suspensorium: In cyprino- branchials may be described as the loss of dontiforms as a whole, as is true for many the first basibranchial. other, but not all atherinomorphs, there are Typically among atherinomorphs, as for no dermal jaw suspensorium elements. Sim- 1981 PARENTI: CYPRINODONTIFORM FISHES 399

A

FIG. 28. Diagrammatic representation of hyoid bar of A. Rivulus harti; B. Oxyzygonectes dowi; C. Procatopus gracilis. Cartilage is stippled. ilarly, the ectopterygoid is also lacking in the ventral extension of the autopalatine as many atherinomorphs including cyprinodon- the ectopterygoid, although it is not present tiforms, although the identification of this as a distinct bone, and no joint lines are vis- state has not been made consistently in ath- ible between it and the autopalatine. erinomorph studies. Rosen (1964) illustrated In some exocoetoids in which the ecto- a section of the jaw suspensorium of Xi- pterygoid is a separate bone (e.g., in Par- phophorus helleri, a poeciliid, and identified exocoetus brachypterus) there is also a ven- 400 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

HYO

FIG. 29. Diagrammatic representation of jaw suspensorium of Cynolebias whitei. tral extension of the autopalatine. Thus the merhead. In a group of the cyprinodontoids, ectopterygoid is considered to be lost in cer- which include goodeids, Empetrichthys, tain atherinomorphs including the cyprino- and Crenichthys, the head of the autopala- dontiforms. The degree of the extension on tine is reduced to a nubbin, a condition con- the autopalatine varies among cyprinodonti- sidered to be secondarily derived. It is readi- forms. In the aplocheiloids, as in most other ly distinguished from the aplocheiloid atherinomorphs, the autopalatine extension condition in that the head is blunt, rather is short and does not reach the quadrate. In than slender. contrast, in the cyprinodontoids, as illustrat- A third derived character of the jaw sus- ed for Procatopus gracilis (fig. 30), the ex- pensorium of the cyprinodontoids is the loss tension of the autopalatine is enlarged and of the metapterygoid (see figs. 29 and 30). covers part of the quadrate. The metapterygoid is also absent in the ad- In addition to having an enlarged ventral rianichthyoids Oryzias and Horaichthys; process, the head of the autopalatine is set but, since they possess none of the derived at an angle to its arm. In the aplocheiloids characters for cyprinodontiforms, their loss (fig. 29), as is true generally for atherino- of the metapterygoid is inferred to be inde- morphs, the head of the autopalatine is pendent. straight, whereas in the cyprinodontoids (fig. The cyprinodontiforms exhibit a derived 30) the head of the autopalatine is distinctly state of the premaxilla characterized by the offset. There is also a bony flange which ex- two-part alveolar process. The superficial tends posteriorly giving the anterior exten- division of the adductor mandibulae inserts sion of the autopalatine the shape of a ham- via a tendon to the middle of the arm of the 1981 PARENTI: CYPRINODONTIFORM FISHES 401

t-HYO

POP

FIG. 30. Diagrammatic representation of jaw suspensorium of Procatopus gracilis.

maxilla, whereas the more anterior layers in- 33) expanded medially, and therefore, car- sert on the posterior extension of the alveo- rying the sensory canal along its midline. lar arm. There are no ethmomaxillary ligaments In the cyprinodontoids, the alveolar arm present in cyprinodontoids as there are in is distinctly S-shaped (fig. 3B), as a result of aplocheiloids. Similarly, there are no liga- bending and enlarging of the post-maxillary ments extending from the interior arms of the process. This is the condition typical of cy- maxillaries to the middle of the rostral car- prinodontoids, and although the arm under- tilage. In addition, there is no meniscus be- goes modifications in several of its subgroups, tween the premaxilla and the maxilla. These it can always be distinguished from that of elements are present, however, in the the aplocheiloids by the posterior indenta- aplocheiloids and atherinoids (Alexander, tion. 1967b). Hence, their absence in cyprinodon- In the aplocheiloids, the dentary is a rel- toids is considered derived. atively thin bone, which carries a distinct Vomer: The vomer bears teeth in all sensory canal (fig. 31C). In Menidia (fig. aplocheiloid species. The state of this char- 3IB), as in many other atherinoids, there is acter is variable, however. When present, a large coronoid process on the dentary; yet, the teeth are usually in a round patch at the ventrally, the bone is unexpanded as in the anteromedial extension of the vomer, as in aplocheiloids. Similarly, in Oryzias (fig. 31 A) Rivulus (fig. 17C). In Aplocheilus (fig. 17B), the dentary is unexpanded. In all cyprino- the teeth extend across the anterior edge of dontoids, the dentary is a robust bone (fig. the vomer. In cyprinodontoids and in ather- 402 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 31. Diagrammatic representation of lower jaw, lateral view: A. Oryzias javanicus; B. Menidia menidia; C. Aphyosemion occidentale. Cartilage is stippled, Meekers cartilage is the elongate central element.

inoids the vomer does not possess a medial acter, however, is dubious because its extension and there are never any teeth on distribution coincides with no other known the vomer. Vomerine teeth might, therefore, character. be derived for aplocheiloids and lost inde- Loss of First Dorsal Fin Ray: Another de- pendently in some aplocheiloids and in cy- rived character which defines the cyprino- prinodontoids. The usefulness of this char- dontoids as a monophyletic group pertains to 1981 PARENTI: CYPRINODONTIFORM FISHES 403

FIG. 32. Diagrammatic representation of first two proximal dorsal radials articulating with two dorsal fin rays in A. Pachypanchax playfairi; with one dorsal fin ray in B. Adinia xenica.

the number of dorsal fin rays. In all aplo- ment of the first epibranchial closer to cheiloids, there is one dorsal fin ray articu- the second pharyngobranchial. lating with each of the first two dorsal radials 4. Autopalatine with its anterior extension (fig. 32A). The first dorsal ray is often rudi- bent sharply and hammer-shaped. mentary; nonetheless, it is present. In all cy- 5. Extension of the autopalatine ventrally prinodontoids (fig. 32B) the first dorsal ray forming an anterior covering of the is apparently lost, and the second remaining quadrate. ray articulates with the first two proximal 6. Metapterygoid absent. dorsal radials. 7. Alveolar arm of premaxilla S-shaped. 8. Dentary expanded medially and robust. SUMMARY OF DERIVED CHARACTERS 9. Loss of an ethmomaxillary ligament. 1. Two basibranchials in the ventral gill 10. Loss of a ligament from the interior arms arch skeleton. of the maxillaries to the middle of the 2. Loss of the dorsal hypohyal. rostral cartilage. 3. Reduction of interarcual cartilage to one 11. Loss of a meniscus from between the half its length, relative to that of the premaxilla and the maxilla. aplocheiloids, and the associated place- 12. Loss of an anterior dorsal fin ray result- 404 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 33. Diagrammatic representation of lower jaw, lateral view, A. Profundulus punctatus; B. Characodon lateralis; C. Crenichthys baileyi; D. pahrump.

ing in the articulation of the first dorsal duced relative to the condition found in Pro- fin ray with first two proximal radials. fundulus. Alexander (1967b) stated that in Fundulus the rostral cartilage is Y-shaped RELATIONSHIPS OF THE CYPRINODON- and therefore comes in contact with the TOIDS: Jaw and jaw suspensorium: The most hooks on the interior arms of the maxillaries. primitive type of jaw structure within cy- With the benefit of the counterstaining tech- prinodontoids is that found in the Central nique employed throughout this study, it has American genus Profundulus. The alveolar been determined that the rostral cartilage is arm of the premaxilla is indented posteriorly, not Y-shaped but is represented by, at most, forming an S-shaped distal process. The den- four small discs of cartilage in the fundulines; tary (fig. 33A) is expanded medially forming one is situated posterior, and two smaller a robust lower jaw. There are no large pro- elements anterior, to a larger medial cartilage cesses on the dentary or the articular, and located between the internal hooks of the the retroarticular is of moderate size. maxillaries (fig. 5C). These bits of cartilage Premaxillary ascending processes in Pro- are held together and to the maxillary by fundulus are flat and broad (fig. 5B). At their connective tissue fibers, forming what is pre- tips sits the large, rectangular rostral carti- sumably the "Y-shaped" rostral cartilage of lage. The interior arms of the twisted max- Alexander. Thus, in fundulines as well as in illaries abut the rostral cartilage and are all other cyprinodontoids (excluding Profun- bound to it by collagen fibers. No ligament from the interior arms to the cartilage has dulus) there has been a loss of contact be- tween the inner arms of the maxillaries and been found, as present in the aplocheiloids and atherinoids, as reported by Alexander the rostral cartilage, a condition associated (1967b). Similarly, there is no ethmomaxil- with reduction of the cartilage. lary ligament, nor is there a meniscus be- In both the fundulines and the Mediterra- tween the premaxilla and maxilla. In other nean genus Valencia which has heretofore cyprinodontoids, the rostral cartilage is re- been classified in the same subfamily as the 1981 PARENTI: CYPRINODONTIFORM FISHES 405

FIG. 34. Diagrammatic representation of upper and lower jaw, lateral view, of Fundulus diaphanus. Maxilla is stippled. fundulines, the premaxillary ascending pro- aplocheiloids to the short and narrow pro- cesses are narrow and elongate (fig. 5D). cesses of the large subgroup of cyprinodon- Narrow premaxillary processes are charac- toids comprising the following: Jenynsia, teristic of the group of cyprinodontoids ex- Anableps, Oxyzygonectes, the poeciliids, cluding Profundulus. The elongate processes procatopines, Pantanodon, Fluviphylax, the of fundulines and Valencia are considered as goodeids, Empetrichthys, Crenichthys, Or- stages in a transition series from the broad estias, the cyprinodontines, Cubanichthys processes typical of Profundulus and the and Chriopeoides. This large group, plus 406 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 35. Diagrammatic representation of the upper jaw in A. Anableps dowi; B. Oxyzygonectes dowi; C. Aplocheilichthys johnstoni; D. Jenynsia lineata.

Valencia, has attenuate interior arms of the considered derived for the fundulines, and maxillaries, rather than the broad tips asso- therefore define them as a monophyletic ciated with Profundulus and the aplochei- group. loids. In addition, they have a maxilla which Valencia shares the derived characters of is straight rather than characteristically the rest of the cyprinodontoids as described twisted as in fundulines, aplocheiloids, and above; that is, a straight maxilla with atten- Profundulus. The arm does not have a pro- uate interior arms, and the development of nounced bend anterior to the autopalatine a dorsal extension over the premaxillary as- (as in Fundulus diaphanus, fig. 5C), but, it cending processes. Valencia is unique is rather straight and often has a pronounced among cyprinodontiforms in having very flat dorsal process which extends anteriorly long attenuate dorsal processes of the max- over the premaxillary ascending processes illaries (fig. 5D). In other cyprinodontoids of (fig. 5D). the large group delimited above, the dorsal The fundulines are unique among cyprin- processes are rounded when present. Since odontiforms in having pronounced hooks on there are no such dorsal processes in the fun- the interior arms of the maxillaries (figs. 5C, dulines, Profundulus or aplocheiloids, the 34). In addition, the interior arms are direct- polarity of the character is ambiguous. The ed anteriorly, rather than medially as in other elongate dorsal processes of Valencia may cyprinodontoids. These characters may be represent the primitive state of the processes 1981 PARENTI: CYPRINODONTIFORM FISHES 407

FIG. 36. Diagrammatic representation of upper and lower jaw, lateral view, of Oxyzygonectes dowi. Maxilla is stippled. which are further reduced in the large processes. The dorsal process is found in this subgroup; or, the reverse may be true. The state, as well, in Anableps (fig. 35A), Oxy- polarity of this character may be resolvable zygonectes (fig. 35B), the remaining proca- with an ontogenetic series of Valencia. topine genera, and the majority of the poe- The large subgroup minus Valencia is de- ciliids (e.g., as illustrated in Rosen and fined by having narrow and shortened pre- Bailey, 1963). In both Pantanodon and Flu- maxillary ascending processes, and the ros- viphylax the dorsal processes are weakly tral cartilage reduced or absent. The dorsal formed; yet, on the basis of characters to be processes of the maxilla are rounded when discussed they are considered to be part of present. This subgroup may itself be subdi- this monophyletic group, and their weakly vided into two monophyletic groups. formed processes are considered to be sec- The poeciliids, procatopines, Fluviphylax, ondarily derived. Pantanodon, and Oxyzygonectes, Jenynsia The distal arm of the maxilla is enlarged and Anableps form a monophyletic group at its most ventral extension (e.g., as in Oxy- based on three derived jaw characters: the zygonectes dowi, fig. 36) in all members of dorsal processes of the maxillaries are in- the group excluding the procatopine genera dented laterally to form nearly fan-shaped Procatopus and Hypsopanchax, and a group processes; the distal arm of the maxilla is of species of Aplocheilichthys (e.g., as in expanded; and the retroarticular is enlarged. Procatopus gracilis, fig. 37), in which the The dorsal process of the maxilla, as in the distal arm of the maxilla is shortened relative procatopine Aplocheilichthys johnstoni (fig. to its condition in the other members of this 35C) and for Jenynsia lineata (fig. 35D), has group. a distinct lateral indentation. The result is a Similarly, the retroarticular is extremely distinct fan-shaped process which projects elongate in Anableps, Jenynsia (fig. 38B), over the triangular premaxillary ascending Oxyzygonectes (fig. 36), and the procatopine 408 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 37. Diagrammatic representation of upper and lower jaw, lateral view, of Procatopus gracilis. Maxilla is stippled. genera Aplocheilichthys (fig. 38C) and Lam- ondarily reduced in Hypsopanchax, Proca- prichthys, and moderately elongate in To- topus, Pantanodon, and Fluviphylax. meurus, the presumed primitive poeciliid. Within this large group, the poeciliids, Within Pantanodon, Fluviphylax, and Pro- Fluviphylax, Pantanodon and the procato- catopus (fig. 37) and Hypsopanchax, the pines are distinguished by the formation of retroarticular is reduced. a greatly enlarged dentary. The less expand- Thus, the premaxillary and retroarticular ed dentary of Jenynsia (fig. 38B), Oxyzygo- characters appear to be correlated. The elon- nectes (fig. 36) and Anableps is the condition gate retroarticular and expanded arm of the primitive for all cyprinodontoids. premaxilla are a general characteristic of the In the poeciliids, procatopines, Pantano- group, but both of these elements are sec- don and Fluviphylax, the dentary is much 1981 PARENTI: CYPRINODONTIFORM FISHES 409

FIG. 38. Diagrammatic representation of lower jaw, lateral view, of A. Valencia hispanica; B. Jenynsia lineata; C. Aplocheilichthys johnstoni. Cartilage is stippled, Meckel's cartilage is elongate medial element. more expanded, especially at its most ante- short and attenuate, rather than triangular as rior end, e.g., in Procatopus gracilis (fig. in the former group. 37). The dentary in this case continues to In the goodeids and the two North Amer- carry a sensory canal; however, the ossified ican genera Empetrichthys and Crenichthys, enclosure of the canal is reduced relative to the dorsal process of the maxilla is present that in other cyprinodontiforms. yet weakly formed (fig. 39). The result is a The second division of these cyprinodon- maxilla which has a small cup-shaped pro- toids comprises the genera Empetrichthys, cess medially to receive the premaxillary as- Crenichthys, Cubanichthys, Chriopeoides, cending process. Because these three taxa Orestias, the goodeids, and cyprinodontines. share other characters with the more derived The mouth of this group is smaller than that cyprinodontiforms, the dorsal process is in- in any other group of cyprinodontiforms. ferred to be reduced rather than primitively The premaxillary ascending processes are unformed. 410 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

ASC

ALV

FIG. 40. Diagrammatic representation of pre- maxilla, lateral view in A. Characodon lateralis; B. Crenichthys baileyi; C. Empetrichthys latos pahrump.

A third unique jaw characteristic of Em- FIG. 39. Diagrammatic representation of the petrichthys, Crenichthys, and the goodeids upper jaw in A. Empetrichthys merriami; B. is the greatly reduced articular that possess- Crenichthys baileyi; C. Characodon lateralis. es no medial extension to carry the sensory canal (fig. 33B, C, D). A fourth unique char- acter, mentioned previously, is the reduction Among these three, the distal arm of the of the anterior arm of the autopalatine, with premaxilla is straight, rather than S-shaped, no anterior or posterior extensions. although the posterior indentation of the al- These four jaw and jaw suspensorium veolar arm is well-formed (fig. 40). characters of goodeids, Empetrichthys, and 1981 PARENTI: CYPRINODONTIFORM FISHES 411

FIG. 41. Diagrammatic representation of the upper jaw in A. Cubanichthys cubensis; B. Aphanius fasciatus.

Crenichthys, along with characteristics of single outer row independently in one other other systems, unite them into a monophy- species of cyprinodontiform, the funduline letic group. Lucania parva. In the nominal genera Cubanichthys, In addition to a single row of outer teeth, Chriopeoid.es, Orestias and in the cyprino- the cyprinodontines and Orestias also have dontines, the dorsal processes of the maxil- a derived lower jaw which is characterized laries are expanded medially, and nearly by the posterior expansion of Meckel's car- meet in the midline (fig. 41). There is also a tilage (e.g., as in Aphanius fasciatus, fig. distinct groove running down the middle of 43A). The cartilage is expanded so that it the dorsal process. The large distal arm of covers a large portion of the articular, in con- the maxilla is correlated in this group (fig. trast to the state of the cartilage in other cy- 42) with the development of a robust upper prinodontiforms (e.g., figs. 27, 38) in which jaw. the cartilage is present as a rod of uniform Cubanichthys and Chriopeoides are hy- width. pothesized to be the primitive members of In Anatolian cyprinodontines (e.g., this assemblage because they possess two Aphanius, fig. 43A) there is a medial exten- primitive characteristics of the jaws found sion of the dentary which projects anteriorly. modified in the remaining members. Both In the South American genus Orestias (fig. nominal genera possess several rows of teeth 43B) the dentary is even further expanded to on the upper and lower jaw; there is a prom- form a medial shield of bone. The condition inent outer row with smaller, scattered inner in Orestias is considered to be the most de- jaw teeth not forming regular rows. Also, rived condition of this transition series (i.e., Meekers cartilage is narrow posteriorly from the typical condition in Chriopeoides, where it inserts into the medial articular pro- fig. 42, to the expanded condition of Aphan- cess (e.g., as in fig. 38). ius, to the fully expanded condition of Or- In Orestias and the cyprinodontines, the estias). The dentary characteristics are cor- teeth are present in a single outer row on related with those of the gill arches, shoulder both the upper and lower jaws. These teeth girdle and pattern of squamation, to be dis- are unicuspid and bicuspid in Orestias, uni- cussed. The characteristic lower jaw of these cuspid in Kosswigichthys, and tricuspid in two groups is characterized by a robust den- remaining cyprinodontines. Teeth occur in a tary and recession of the urohyal and branch- 412 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 42. Diagrammatic representation of upper and lower jaw, lateral view of Cubanichthys (Chrio- peoides) pengelleyi. iostegal rays (fig. 141). In lateral view, the group of cyprinodontiforms, fishes of the mouth cleft is nearly vertical (fig. 14G). genera Jenynsia, Anableps, and Oxyzygo- Aphanius fasciatus has tricuspid jaw nectes. The teeth of Jenynsia are distributed teeth, as do other Old World and all New in one large outer row and several smaller World cyprinodontines. The character trans- scattered in indistinct inner rows. All the jaw formation series described above for the den- teeth are tricuspid. However, the shape of tary indicates that if tricuspid teeth can be the outer teeth of Jenynsia varies from dis- used as a derived character, it is only at the tinctly tricuspidate with the inner cusp just level of defining the cyprinodontines and slightly longer than the middle (fig. 44B) to Orestias as a monophyletic group, with a re- a faintly tricuspidate form in which the lat- version to unicuspid teeth in some Orestias eral shoulders are only weakly formed (fig. and in Kosswigichthys. 44 A). Tricuspid outer teeth occur in one other In Oxyzygonectes (fig. 36) adults have a 1981 PARENTI: CYPRINODONTIFORM FISHES 413

mdn

FIG. 43. Diagrammatic representation of lower jaw, lateral view, of A. Aphanius fasciatus; B. Orestias sp. Cartilage is stippled, Meckel's cartilage is the enlarged medial element. row of very large recurved unicuspid teeth, The jaw dentition of an adult Anableps and a dense inner patch of teeth which ap- consists of one large outer row of recurved pear to be distributed in about five or six teeth and several smaller scattered inner rows. These inner jaw teeth are all tricuspid rows of unicuspid teeth. The inner jaw teeth in both juveniles and adults. The teeth are so closely packed that on a cursory examination have what appear to be weakly formed lat- they appear to be villiform. The outer teeth eral shoulders. of juvenile Oxyzygonectes are weakly tricus- The upper jaw of an adult Anableps is very pidate. Thus, the jaw dentition of Oxyzygo- derived (fig. 35A). There are only weakly nectes and Jenynsia is apparently very much formed premaxillary ascending processes the same, with Oxyzygonectes losing the lat- and the premaxillaries form an arc. The max- eral cusps of the outer row, and having more illa is elongated medially, however, the dor- inner jaw teeth. sal process of the maxilla and expanded dis- 414 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 44. Sketch of two forms of tricuspid teeth in the genus Jenynsia (see text for discussion).

tal arm distinctive of the monophyletic group to which it is assigned are prominent. Another unique feature is the dumbbell shape of the rostral cartilage, unknown in other cyprinodontiforms. Also, a block of cartilage sits between the autopalatine and the maxilla (fig. 35A) termed here the subau- topalatine cartilage. Such a block is often found in this position in atherinomorph fish- es; its presence in Anableps is therefore con- sidered primitive. Juvenile Anableps show all the specializa- tions of the adults, therefore, an embryo of Anableps dowi, the presumed most primitive species of the genus (Miller, 1979) was ex- amined. The yolk sac was removed and the specimen counterstained. The outer teeth of the embryo have distinct lateral shoulders and there is a very narrow medial cusp, they FIG. 45. Diagrammatic representation of dor- differ little from the weakly tricuspidate sal gill arches, ventral view, of Pantanodon mad- teeth in Jenynsia. Furthermore, in the em- agascariensis. Cartilage is stippled. bryo, triangular-shaped ascending processes like those in Jenynsia and Oxyzygonectes are present on the premaxillaries. and quite variable among them (Nelson, Hence, on the basis of dentition, the three 1969; Rosen, 1973). Except in the unusual genera, Jenynsia, Anableps, and Oxyzygo- Pantanodon (fig. 45) and certain poeciliids, nectes are hypothesized to form a monophy- dorsal gill arch anatomy among cyprinodon- letic group. toids varies little from the basic structure ex- GILL ARCHES: The structure of the bran- hibited by Profundulus punctatus (fig. 6B). chial skeleton has been used in recent years In Profundulus, the interarcual cartilage is to deduce phylogenetic relationships be- reduced relative to the condition in aplo- cause it is both constant within large groups cheiloids. The three pharyngobranchial 1981 PARENTI: CYPRINODONTIFORM FISHES 415

FIG. 46. Diagrammatic representation of dorsal gill arches, ventral view, of A. Fundulus diaphanus; B. Lucania parva. Cartilage is stippled. toothplates (associated with pharyngobran- however, toothlike structures lie above it chials 2,3, and 4) are separate elements. The suspended in connective tissue. The third cartilaginous points of articulation are rela- and fourth pharyngobranchial toothplates tively narrow. Species in the subgenera Zyg- are fused into one large toothbearing ele- onectes, Xenisma, and Fundulus of the ge- ment. The teeth are arranged in discrete nus Fundulus differ from Profundulus in rows, with tricuspid teeth being found on the having the cartilaginous point of articulation posterior five rows. Epibranchials one of the second pharyngobranchial toothplate through three are absent, as is the interarcual greatly expanded laterally to produce a cartilage. (Also, the hypobranchials of Pan- broad head for the articulation of the inter- tanodon are reduced or absent, as illustrated arcual cartilage (fig. 46A). In the subgenus by Rosen, 1965.) The expanded second pha- Plancterus, and in the funduline genera Ad- ryngobranchial toothplate has been found in inia, Leptolucania and Lucania (fig. 46B), no other atherinomorph genus examined. the cartilaginous point of articulation is not Fusion of the third and fourth pharyngobran- enlarged. This is also the case in Valencia chial toothplates occurs within a group of (fig. 47A) which exhibits the primitive con- cyprinodontines, but otherwise their struc- dition for the cyprinodontoids. ture is basically that of the general form. Among the more derived cyprinodontoids, Among the poeciliids, teeth on the third the structure of the dorsal gill arches differs and fourth pharyngobranchial toothplates most from the general condition in Pantan- are often arranged in discrete rows, and even odon and some derived poeciliid genera. tricuspid teeth are present. In these poe- In Pantanodon madagascariensis (fig. 45), ciliids however, the epibranchials and inter- the second pharyngobranchial toothplate is arcual cartilage are present, as is a more greatly expanded into a sheet of bone. There primitively shaped second pharyngobranchi- are no teeth in sockets on the toothplate; al toothplate. In this study, Pantanodon is 416 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 47. Diagrammatic representation of dorsal gill arches, ventral view, of A. Valencia hispanica; B. Empetrichthys latos pahrump. Cartilage is stippled.

PB3

FIG. 48. Diagrammatic representation of dorsal gill arches, ventral view, of A. Procatopus gracilis; B. Tomeurus gracilis. Cartilage is stippled. 1981 PARENTI: CYPRINODONTIFORM FISHES 417 considered to be a close relative of the poe- ciliids; however, its close relationship is not based on gill arch morphology. Tomeurus has dorsal gill arches of the structure primi- tive for cyprinodontoids (fig. 48B), although the epibranchials are reduced and no inter- arcual cartilage has been found. These char- acters are considered derived for the genus, since the cartilage and more robust epibran- chials are found in more derived poeciliids. Therefore, either Pantanodon is a poeciliid which lost its gonopodium, or the similar gill arch structure of the poeciliids and Pantan- odon are independently derived. On the ba- sis of the distribution of all derived charac- ters, the latter hypothesis is accepted here. Empetrichthys (fig. 47B) and Crenichthys exhibit a peculiar shape of the first epibran- chial. The bone is nearly Y-shaped resulting from an indentation at its base. This type of first epibranchial has not been found else- where within cyprinodontiforms. In the cyprinodontines and Orestias, the FIG. 49. Diagrammatic representation, dorsal second pharyngobranchial is offset to the gill arches, ventral view, of Cyprinodon vari- third, as in Cyprinodon variegatus (fig. 49). egatus. Cartilage is stippled. This change in orientation of the pharyngo- branchial excludes the cartilaginous point of articulation from the ventral toothed surface teeth. In Cualac tessellatus (fig. 50B) there of the pharyngobranchial toothplates. In ad- is no toothplate yet there is a distinct sepa- dition, the fourth pharyngobranchial tooth- rate cartilage which sits at the anterior tip of plate is reduced. However, such a reduction the first epibranchial cartilage. is not unique to this group, as the toothplate An element in this position has been found is also reduced in Procatopus (fig. 48A) and in only one other species of cyprinodonti- other procatopines. form, Cynolebias elongatus. In this Neo- In Cyprinodon (fig. 49) the third and fourth tropical aplocheiloid, there is a distinct car- pharyngobranchial toothplates are fused into tilage as well as a bony toothbearing a single toothbearing element. The teeth are element. Its condition is comparable to that arranged in rather discrete rows. Such fusion of Floridichthys. occurs in many, perhaps most, individuals of Among atherinomorph fishes, no first the New World cyprinodontine genera Cy- pharyngobranchial toothplate is found ex- prinodon, Jordanella, Garmanella, Megup- cept in the genera Cynolebias and Flori- silon, Floridichthys, and Cualac, and in Or- dichthys. An ossified first pharyngobranchial estias. Although the occurrence of some is present among some atherinoid fishes, in- individuals with unfused toothplates makes cluding species of the genera Melanotaenia the upper pharyngeal character difficult to and Menidia. Otherwise among atherino- use, the regular arrangement of the teeth is morphs, first pharyngobranchial elements a constant defining character of all of these are absent. Their appearance within these genera. two rather unrelated groups of cyprinodon- In Floridichthys carpio (fig. 50A) there is tiforms poses a problem for interpretation. a distinct first pharyngobranchial cartilage as If the condition is a retained primitive char- well as a toothplate which bears a patch of acter, then the most parsimonious interpre- 418 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

TPBl? S^)

FIG. 50. Diagrammatic representation of dorsal gill arches, dorsal view, of A. Floridichthys carpio; B. Cualac tessellatus. Cartilage is stippled. tation in light of all other data would be that Within this group, the pectoral fins are dis- the elements are lost individually in all other tinctly highset (e.g., as in Tomeurus gracilis, groups of cyprinodontiforms. So many in- fig. 51; and Heterandria bimaculata, fig. 52; dependent losses, however, presuppose far and the procatopines Procatopus glaucicau- more evolutionary events than if it is as- dus, fig. 53) as opposed to the lowset fins of, sumed that these elements are uniquely de- for example, Rivulus (fig. 54), Cualac (fig. rived twice among cyprinodontiforms, once 55) and Lucania (fig. 56). The highset pec- in C. elongatus and again in Floridichthys toral fins are related to the placement of the and Cualac, thereby supporting a sister radials in a dorsal position on the scapulo- group relationship of the last two genera. coracoid, and a gently arched dorsal limit of The condition may also define a subgroup of the scapula and cleithrum (fig. 8C, D). This Cynolebias species. is correlated with a loss of the first post- PECTORAL GIRDLE: The pectoral fins of cleithrum which is wanting in all members of cyprinodontiforms are described as typically the group except for some nominal species lowset, with the corresponding radials situ- of the genus Aplocheilichthys. The structure ated ventrally rather than dorsally. There is of the shoulder girdle of Profundulus (fig. a large, scale-shaped first postcleithrum and 8A) is the general condition, as in the aplo- a thin third postcleithrum. The posttemporal cheiloids with the radials situated ventrally. may have an ossified lower limb, or a limb The pectoral fins are distinctly lowset in represented solely by a ligament; this char- the genera Anableps, Jenynsia, and Oxyzy- acter is used only at the lower levels of phy- gonectes, as well as other cyprinodonti- togeny reconstruction since it is not corre- forms; however, as stated, the pectoral fins lated with larger sets of characters used to are generally highset in most other groups of delimit major groups. afherinomorph fishes. Since the derived form Shoulder girdles are lowset in all cyprin- of the pectoral fins has been interpreted as odontiforms except the poeciliids, Fluviphy- lowset within cyprinodontiforms, the highset lax, Pantanodon, and the procatopines. pectoral fins may only be interpreted as sec- 1981 PARENTI: CYPRINODONTIFORM FISHES 419

FIG. 51. Sketch of body form and fin position of Tomeurus gracilis, male above, female below. Dotted line approximates base of hypural plate. (After Rosen and Bailey, 1963.)

ondarily derived in poeciliids, procatopines, amined. The presence of this character in all Pantanodon and Fluviphylax. aplocheiloids and Profundulus suggests that SKULL ANATOMY: A constant feature of it is a primitive character for cyprinodonti- the skull of aplocheiloids is the presence of forms. Thus, its absence or reduction in all a lateral facet on the anterior surface of the other cyprinodontoids is evaluated as a de- lateral ethmoid which articulates with the rived character supporting their monophyly. head of the autopalatine (fig. 17). Such an The generalized state of the size and po- extension is present in the cyprinodontoids sition of the lateral ethmoid is exemplified by only in the genus Profundulus (fig. 57A). Tomeurus gracilis (fig. 16B). This character was one of several Farris Among procatopines, the lateral ethmoid (1968) used to separate the species of Fun- is expanded medially under the broad arm of dulus from Profundulus; he reported the pro- the parasphenoid, as in Aplocheilichthys cess as absent in all species of Fundulus ex- johnstoni (fig. 16C). (Compare this expan- 420 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 52. Sketch of body form and fin position of Heterandria bimaculata, male above, female below. Dotted line approximates base of hypural plate. (After Rosen, 1979.) sion of the lateral ethmoid with that of the not only by the anteriorly projecting ventral Neotropical aplocheiloids, fig. 17C.) arms of the maxillaries, but by the extension Medial expansion of the lateral ethmoid is of the autopalatines to a position lateral to accompanied by a change in its orientation the enlarged vomer as well. relative to the frontal bones in Empetrich- The mesethmoid is cartilaginous in aplo- thys, Crenichthys, the goodeids, Cuban- cheiloids. In addition, it is cartilaginous ichthys, and Chriopeoides, the cyprinodon- among the cyprinodontoids in Pantanodon tines and Orestias. As in the goodeid, and the procatopines (it is ossified in Fluvi- Characodon lateralis (fig. 57D), the lateral phylax) as well as in the Anatolian cyprino- ethmoid is oriented such that the greater part dontines. The cartilaginous mesethmoid is of the element lies anterior to the limit of the considered a derived condition defining the frontals. This may be compared with the gen- aplocheiloids; among the cyprinodontoids, eral condition in cyprinodontoids as in Pro- its independent occurrence within two un- fundulus punctatus (fig. 57A), in which the related groups is convergent. outer flange of the lateral ethmoid is expand- The group consisting of Empetrichthys, ed, rather than narrow as in Characodon. Crenichthys, the goodeids, Cubanichthys, Among the fundulines, the lateral ethmoid and Chriopeoides, the cyprinodontines and is also expanded under the parasphenoid (fig. Orestias possess another derived feature of 16A). However, the lateral ethmoid not only the skull; viz., a reduced autopterotic fossa lacks the facet for articulation of the auto- (fig. 57B, C, D). Uyeno and Miller (1962) palatine, but the autopalatine does not come used the narrow fossa to separate Empe- in contact with the lateral ethmoid. Rather, trichthys and Crenichthys from Profundulus the fundulines' pronounced snout is effected which has an extremely wide fossa (fig. 57A). 1981 PARENTI: CYPRINODONTIFORM FISHES 421

FIG. 53. Sketch of body form and fin position of Procatopus glaucicaudus, male above, female below. Dotted line approximates base of hypural plate. (After Clausen, 1959.)

However, they did not compare this condi- nectes and Jenynsia. As illustrated for Oxy- tion to its state in other cyprinodontiforms. zygonectes dowi (fig. 59) the crests are great- The fossa of Profundulus is wider than in ly elongate and are separated by a distinct any other cyprinodontiform and may be con- notch from the dome over the foramen mag- sidered an autapomorphy of the genus. num. In contrast, supraoccipital crests are Enlarged supraoccipital and epiotic pro- present in Profundulus (fig. 60), yet they cesses occur among many groups of acan- abut the dorsal wall of the foramen magnum, thopterygian fishes. The general condition of rather than being separated from it by a the supraoccipital crests among atherino- notch. morph fishes is paired (Rosen, 1964); among The states of the first vertebra in oviparous cyprinodontiforms this is the case except in cyprinodontiforms have been described, al- the two monotypic genera Cubanichthys and though somewhat erroneously, by Sethi Chriopeoides. In these genera the supraoc- (1960). All the aplocheiloids have a complete cipital crest is a large, single process which neural spine on the first vertebra (fig. 26). extends above the dorsal profile (fig. 58). Among the cyprinodontoids, the neural arch Thus, the sister group relationship of these of the first vertebra is open, and therefore, two genera is supported again. does not form a neural spine (fig. 60) in Pro- Another unique form of the supraoccipital fundulus, Valencia, Empetrichthys, Crenich- processes is shared by Anableps, Oxyzygo- thys, the fundulines, and goodeids. 422 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 54. Sketch of body form and fin position of Rivulus beniensis, male above, female below. Dotted line approximates base of hypural plate. (After Klee, 1965.)

Among the procatopines, as well as in Flu- genus Poecilia, the neurapophyses are even viphylax, the neurapophyses meet in the more expanded anteriorly and applied to, midline; no distinct spine is formed, how- and fused with, the skull. This characteristic ever. The condition is interpreted as a re- attachment of the first vertebra must be con- duction in the neurapophyses and a second- sidered independently derived in both Pan- arily derived medial fusion. In this group as tanodon and the poeciliids if the monophyly well as in the aforementioned cyprinodonti- of the poeciliids based on the presence of a forms, the basioccipital and exoccipital con- gonopodium and other reproductive spe- dyles are all well-formed. cializations is accepted. In all poeciliids, as well as Pantanodon, A superficially similar condition of the at- there are no exoccipital condyles. The at- tachment of the first vertebra to the skull oc- tachment of the first vertebra to the skull in curs in the New World cyprinodontines of Tomeurus (fig. 61) involves the forward ex- the nominal genera Cyprinodon, Megupsi- pansion of the neurapophyses around the lon, Jordanella, Floridichthys, Cualac, and base of the foramen magnum. The arch is Garmanella. There is no spine formed by the open and the first vertebra articulates with neurapophyses of the first vertebra. Instead, the skull only via the basioccipital condyle. the neurapophyses are slightly expanded, In Pantanodon, as well as some of the brought forward, and applied to the skull more derived poeciliids such as those of the (fig. 62). Th.i exoccipital condyles are lacking 1981 PARENTI: CYPRINODONTIFORM FISHES 423

FIG. 55. Sketch of body form and fin position of Cualac tessellatus, male above, female below. Dotted line approximates base of hypural plate. (After Miller, 1956.) and, in addition, the supraoccipital forms the The vomer is absent in Pantanodon, roof of the foramen magnum. In all other cy- Fluviphylax, the procatopines (except Po- prinodontiforms, as well as in the poeciliids ropanchax, Lamprichthys and species of and Pantanodon, the supraoccipital is ex- Aplocheilichthys), and the South American cluded from formation of the foramen mag- Orestias. The vomer is hypothesized to be num. Also, the form and position of the lost independently at least twice among cy- neurapophyses is quite different between the prinodontiforms, once in Orestias, and once poeciliids and New World cyprinodontines. in the procatopines, Fluviphylax and Pan- In poeciliids, they are greatly expanded and tanodon. The significance of its distribution form a trough in which the supraoccipital re- is discussed in the following section. gion of the skull sits; in cyprinodontines, the Parietals are absent in two groups of cy- neurapophyses are simply applied to the prinodontoids. They are lacking in Orestias skull and provide reinforcement yet form no and the cyprinodontines, as well as the pro- trough similar to that of the poeciliids. catopines, Fluviphylax and Pantanodon. In Orestias and the Anatolian cyprinodon- Their absence in these two groups is consid- tines the exoccipital condyles are present as ered to be an independent loss. In addition, in Profundulus, yet reduced. The neur- their absence from more derived members of apophyses of the first vertebra are also re- the poeciliids is secondarily derived since duced, as in the New World cyprinodon- parietals are present in Tomeurus. tines, and may or may not meet in the Axial Skeleton: The first pleural rib arising midline. on the parapophyses of the second vertebra 424 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 56. Sketch of body form and fin position of Lucania parva, male above, female below. Dotted line approximates base of hypural plate. (After Hubbs and Miller, 1965.) has been described as a derived character of a furrow in the posterior face of the process. cyprinodontiforms. This state occurs in all Within Orestias and the cyprinodontines, members of the group except the funduline the transverse processes are reduced to cup- genus Adinia in which the rib arises on the shaped processes (Sethi, 1960) into which parapophyses of the first vertebra. Since the the pleural ribs insert. This reduction is con- general state among acanthopterygians is for sidered as another derived character of Or- the rib to be on the third vertebra, this case estias and the cyprinodontines. in Adinia is hypothesized to be a further de- Pleural ribs by definition occur only on rived state in the transition series, and serves parapophyses of abdominal vertebrae and as a defining character of the genus. not on caudal vertebrae. However, within The parapophyses themselves are gener- Pantanodon, the procatopines and some ally robust, with the pleural rib inserting into poeciliids including Xiphophorus and Poe-

FIG. 57. Diagrammatic representation of the skull, ventral view, in A. Profundulus punctatus; B. Cyprinodon variegatus; C. Empetrichthys latos pahrump; D. Characodon lateralis. Lateral ethmoid is cross-hatched; lacrimal is blackened; autopterotic stippled. 1981 PARENTI: CYPRINODONTIFORM FISHES

C D 426 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 58. Diagrammatic representation of the posterior region of the skull and attachment of first vertebra, lateral view, of Cubanichthys {Chriopeoides) pengelleyi. Posttemporal removed.

cilia, at least one or two pleural ribs are nectes, and many but not all goodeids (see found on the first and second caudal verte- Miller and Fitzsimons, 1971). In Anableps, brae. This character, absent in Fluviphylax the central row of pores (2b through 4a) is and certain poeciliids, is ambiguous. reduced to two pores which are referred to Sensory Pores and Cephalic Squamation: as pores 3 and 4a. Departure from the gen- The general pattern of the head scales and eral squamation pattern also occurs in Ana- sensory pore patterns of cyprinodontiform bleps in which there are many scales ar- fishes is exhibited by the genus Jenynsia (fig. ranged in a scattered pattern which cannot 14B). There are seven preorbital, three or readily be interpreted using Hoedeman's ter- four lacrimal, four mandibular, and six or minology. seven supraorbital pores. Since the pattern of Jenynsia is postulated In Jenynsia, there is a break between the as the plesiomorphic sensory pore pattern, anterior and posterior section of supraorbital departures from this pattern are of interest pore 2, termed 2a and 2b. A series of three in defining monophyletic groups of cyprino- pores (2b, 3, and 4) follow the section formed dontoids. However, patterns discussed here by pores 1 and 2a. There is another break are only the most common ones found within between sections of pore 4 referred to as a group of genera. A rigorous analysis of pores 4a and 4b. The section 4b through 7 head pore and scale patterns requires a sur- completes the supraorbital series. Gosline vey of inter- and intraspecific variation that (1949) figured an identical pattern for Fun- is outside the scope of this study. dulus chrysotus and stated that this was the Supraorbital pores of the poeciliids, pro- common pattern among Fundulus species. It catopines, Fluviphylax and Pantanodon was also observed that such a pattern is typ- show an apparently unique modification. The ical of cyprinodontoids such as Profundulus maximum development of sensory pores of (also reported by Miller, 1955a), Oxyzygo- poeciliids (fig. 14D, E, F) was based on a 427 1981 PARENTI: CYPRINODONTIFORM FISHES

soc-pro epo-pro

FIG. 59. Diagrammatic representation of the posterior region of the skull and attachment of first vertebra, lateral view, of Oxyzygonectes dowi. Posttemporal removed.

FIG. 60. Diagrammatic representation of the posterior region of the skull and attachment of first vertebra, lateral view, of Profundulus punctatus. Posttemporal removed. 428 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

soc-pro

FIG. 61. Diagrammatic representation of the posterior region of the skull and attachment of first vertebra, lateral view, of Tomeurus gracilis. Posttemporal removed.

survey of such patterns in all major supra- In the New World cyprinodontine Jordan- specific categories of the family (Rosen and ella and some goodeids (see Gosline, 1949), Mendelsohn, 1960). The supraorbital pores the canal is continuous except for a break lie in groupings similar to the plesiomorphic between pores 4a and 4b. This pattern is con- pattern for cyprinodontiforms. The unique sidered to be independently derived in Jor- feature is the recessed neuromasts in the danella and among a group of goodeids. middle section (pores 2b through 4a) forming Reduction of the pore system to pores 6 a small trough. {Poropanchax was defined and 7 only, occurs in the fundulines Adinia on the basis of its embedded neuromasts and Leptolucania. A cephalic sensory pore which open as a series of pores.) The pattern system is absent in the monotypic New is only weakly shown by the diminutive Flu- World cyprinodontine, Megupsilon. viphylax and by Pantanodon. In Cubanichthys a canal is present be- The connection of the canal between pores tween what appear to be pores 1 and 3 only, 4a and 4b (forming just one pore 4) occurs in although Gosline stated they were present Empetrichthys and Crenichthys, both of between pores 2 and 3 as well as 6 and 7. which retain the disrupted canal between There are no pores posterior to what is iden- pores 2a and 2b. tified here as pore 3 in Cubanichthys; how- The pattern Gosline termed the simplest ever, since Gosline reports pores 6 and 7 among cyprinodontiforms (in the cyprino- present they must be considered part of the dontines Cyprinodon, Floridichthys, and maximally developed pattern. Because of Garmanella, and the funduline Lucania) in their position, I interpret the first two pores addition to connection of canals between 4a as 1 and 3, even though by definition the pore and 4b involves a connection between 2a and anterior to 3 should be 2b or 2. The ambi- 2b (resulting in one pore 2). Thus, the canal guity of the numbering system is evident in is continuous between pores 1 through 7. such a case. 1981 PARENTI: CYPRINODONTIFORM FISHES 429

FIG. 62. Diagrammatic representation of the posterior region of the skull and attachment of first vertebra, lateral view, of Cyprinodon variegatus. Posttemporal removed.

Pore 3 of Cubanichthys is large, as it is in 14) and of the aplocheiloid Cynolebias (fig. Chriopeoides, and is considered a synapo- 13). This character within cyprinodontoids morphy of the two monotypic genera. cannot always be distinguished from that in Replacement of the two E scales by one Cynolebias (except for the fact that the large E scale also occurs within the New preorbital area is smaller in the aplochei- World cyprinodontines, Lucania, Cuba- loids; yet this character is independent of the nichthys, Chriopeoides, Empetrichthys, and preorbital line of neuromasts). A line of Crenichthys. Reduction of the number of preorbital neuromasts is not peculiar to these pores is apparently correlated at some level genera, as it is also found among the fundu- with the reduction in the number of head lines; therefore, the generality of the pattern scales. cannot yet be determined. Consequently, it Gosline reported that Aphanius dispar, an is assessed as a convergence between mem- Anatolian cyprinodontine, has a canal be- bers of the genus Cynolebias and Orestias tween pores 2 and 4 and 6 and 7, and also and a group of Anatolian cyprinodontines. noted the lack of mandibular canals. Speci- INTERNAL FERTILIZATION AND VIVIPARI- mens of A. dispar also possess pore 1, and TY: Previous workers (e.g., Rosen and Bai- three neuromasts apparently corresponding ley, 1963; Miller, 1979) have assumed that to pores 5 through 7. viviparity defines a monophyletic group of Another species, A. mento, lacks cephalic cyprinodontiforms, and have therefore fo- sensory pores and has a series of minute neu- cused on describing the similarities and dif- romasts arranged in a lyre-shaped pattern. ferences of adaptations for viviparity among Neuromasts ring the orbit and a line of mi- the families in an effort to determine which nute neuromasts replaces the preopercular viviparous family was more closely related and mandibular canals. The entire system is to which other such family. In the present strikingly like that of the genus Orestias (fig. study, this presumption was discarded at the 430 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 64. Diagrammatic representation of first several rays of the anal fin of a male Cynolebias (Campellolebias) brucei. External view.

true gonopodium. Both cases are inferred to represent modifications for internal fertiliza- tion, yet both species are oviparous as well as annual. FIG. 63. Diagrammatic representation of first Females of brucei isolated after being in several rays of the anal fin of a male Cynolebias (Cynopoecilus) melanotaenia. External view. contact with males have laid fertilized eggs (Vaz-Ferreira and Sierra, 1974). Presumably, once the eggs are laid, they develop in a fash- outset. The simple division of cyprinodonti- ion typical of their annual relatives. forms into oviparous and viviparous groups One other case of internal fertilization oc- is an artificial one, and grossly oversimplifies curs within aplocheiloids in Rivulus mar- the question of cyprinodontiform interrela- moratus. Populations of this species have tionships. been found consisting of self-fertilizing her- Internal Fertilization and Anal Fin Modifi- maphrodites and possess color patterns in- cation: Internal fertilization occurs in groups distinguishable from females of the species of atherinomorph fishes with and without an (Harrington, 1961). This self-fertilization, of anal fin modified into a gonopodium. Devel- course, involves no modification of the anal oping embryos have been found in the body fin. The fertilized eggs of marmoratus are cavity of the ricefish Oryzias (Amemiya and laid as in C. brucei and melanotaenia, thus Murayama, 1931), yet no modifications in the there are no known cases of embryo reten- anal fin structure of this genus have been re- tion among the aplocheiloids. ported. Among cyprinodontoids, internal fertiliza- Among aplocheiloids, a group of Neotrop- tion has been demonstrated only among the ical genera are distinguished by the thick- viviparous families; however, its discovery ening of the anal rays of the females. All in- in an oviparous cyprinodontoid would not be cluded species are annual, and this has been surprising, considering the generality of the suggested as an aid to the depositing of eggs condition. in the substrate during fertilization (Weitz- Structure of the gonopodium: Among the man and Wourms, 1967). viviparous families there are three basic The anal fin of aplocheiloids is typically types of anal fin modifications of the male unmodified, except in two species placed which effect internal fertilization. These are here in the genus Cynolebias: melanotaenia the gonopodia of poeciliids, the tubular gon- (Regan) and brucei (Vaz-Ferreira and Sierra). opodia of Jenynsia and Anableps, and the In melanotaenia, the first six anal rays of the muscular internal organ and slightly modified male are crowded together (fig. 63) and anal fin of the goodeids. slightly offset from the rest of the fin. The The structure and development of the gon- rays are covered with small contact organs. opodium of the poeciliid fishes has been dis- In brucei (fig. 64) the first three anal rays are cussed in detail (Rosen and Bailey, 1963; drawn together to form what is effectively a Rosen and Gordon, 1953; Rosen and Kail- 1981 PARENTI: CYPRINODONTIFORM FISHES 431

FIG. 65. Diagrammatic representation of gonopodium and gonopodial suspensorium of Poecilia vivipara. Anal radials are blackened. man, 1959). The gonopodia and gonopodial of poeciliid fishes; it is primarily on these suspensoria vary among taxonomic groups structures that such groups are defined. 432 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 66. Sketch of body form and fin position of Anableps microlepis, male above, female below. Dotted line approximates base of hypural plate. (After Rosen, 1973b.)

The poeciliid gonopodium (fig. 65) is (fig. 66). In Anableps, the sheath is covered formed principally from the third, fourth, with scales; in Jenynsia it is bare. and fifth anal rays. Transformation from an Internally, the structures are similar in that undifferentiated anal fin begins with a thick- the anal rays are twisted around each other ening of the third anal ray. In all poeciliids (figs. 67, 68). Similarly, there is an enlarge- the first three anal rays are unbranched. ment of the proximal anal radials, as well as There is a rapid growth of rays three an elongation of the hemal spines. Gonopo- through five to form the so-called 3-4-5 com- dial development in Jenynsia and Anableps plex. Further elaboration and growth occur, differs considerably; neither resembles that resulting in a gonopodium which is often of the 3-4-5 complex typical of poeciliids, adorned with various spicules, hooks, and however. spines. In Anableps, the gonopodium is formed Internal supports are modified within poe- from the 12 anal fin rays, counting each ray ciliids to a greater degree than in the other separately. The first ray is rudimentary, viviparous families. Again, as in Poecilia vi- nonetheless it will be referred to as ray 1, vipara (fig. 65), the proximal anal radials two contrary to the convention established by through five are elongated. Histolysis of the Turner (1950). first hemal arch results in an ossified remnant The first four rays are unbranched. Rays termed the ligastyle, which migrates ante- three through six are enlarged and twisted riorly. In addition, the second, third, and around each other (fig. 67), whereas seven sometimes fourth hemal arches are expand- through nine are also enlarged but lie ed, the distal tips of which project anteriorly straight. Rays 10 through 12 are drawn for- to meet the anteriorly projecting tips of the ward in the formation of the tubular sheath, proximal anal radials. but otherwise undergo little differentiation. Anableps and Jenynsia have a tubular The proximal radials are also enlarged, gonopodium formed from enlarged anal fin drawn together and angled anteriorly. The rays covered anteriorly with a fleshy sheath first four or five proximal radials are offset 1981 PARENTI: CYPRINODONTIFORM FISHES 433

^

FIG. 67. Diagrammatic representation of gonopodium and associated elements of Anableps. Anal radials are blackened. (After Turner, 1950.)

FIG. 68. Diagrammatic representation of gonopodium and associated elements of Jenynsia lineata. Sixth middle anal radial is stippled; all other radials are blackened. 434 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 69. Diagrammatic representation of the anal fin of a female Jenynsia lineata. Sixth middle anal radial is stippled; all other radials are blackened. to either the left or the right of the midline In Jenynsia (fig. 68) the gonopodium is in sinistral or dextral males, respectively; formed from 10 anal rays; however, its de- they extend to just beneath the vertebral col- velopment involves primarily reductions of umn. Typically, there is histolysis of the last some elements found typically in the female several pleural ribs. The hemal spines, es- anal fin. Therefore, the anal fin of the female pecially the first three, extend ventrally and will be described first so that a comparison are situated between the proximal radials. At with the structures within the gonopodium their bases these radials typically have bony may be made easily. flanges which project dorsally. Similarly, the In an adult female Jenynsia (fig. 69) there bases of the anal fin rays are greatly enlarged are 10 anal rays, counting the last two sep- and have similar flanges which overlap on arately. The first two rays are unbranched. adjacent rays. A full complement of middle The first six rays are crowded together; there and basal radials appears to be present, al- is a corresponding crowding and reduction though Turner (1950) did not illustrate all of of the first five middle anal radials. The first these for his specimen. two proximal radials are fused at their base. 1981 PARENTI: CYPRINODONTIFORM FISHES 435

FIG. 70. Diagrammatic representation of the internal structure of the abdominal cavity, anal fin rays and vertebral column of Characodon lateralis.

There appears to be no separate proximal In an adult male Jenynsia the sixth middle radial for the first anal ray; however, it is anal radial is the first unreduced radial as it possible that the radial has become fused to is in the female (fig. 69), and as in the female, the base of the recognized first radial which five rays precede and four rays follow this has a small bony knob projecting anteriorly. radial. Of the first six rays, all but rays 3 and This interpretation is supported by the fact 6 are extremely reduced. The seventh and that there are three middle radials present eighth rays, as well as 3 and 6, are elongate corresponding to the large proximal radial. and thickened; together with the relatively Five proximal radials all lie anterior to the unelaborated segments of the ninth ray, they elongate first hemal spine. constitute the principal rays of the gonopo- 436 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 71. Anterior rays and supports of the anal fin of a male A. Characodon lateralis; B. Crenichthys bailey/; C. Empetrichthys merriami. Cartilage is stippled. 1981 PARENTI: CYPRINODONTIFORM FISHES 437 dium. Ray 6, the thickest and the longest, is The rudimentary anal ray is formed to hooked at its tip. The proximal radials, es- varying degrees among members of the fam- pecially those of rays 2 through 6, are crowd- ily (Miller and Fitzsimons, 1971). The first ed together more so than in the female. four to seven middle anal radials are not These radials appear to be fused in part, al- present as distinct structures in all goodeids though they have not been observed com- examined (e.g., as in Characodon lateralis, pletely fused. All radials appear to be pres- fig. 71A). However, among cyprinodonti- ent, but identification of individual segments form fishes, middle anal radials are fused to is difficult. the proximal radials in the two North Amer- In male and female Jenynsia, there are ican genera suggested as close relatives of bony flanges on the base of the rays and the the goodeids, Crenichthys and Empetrich- proximal radials. The proximal radials are thys. In Crenichthys baileyi (fig. 76), the first offset to the midline in males, corresponding five middle radials are lacking. In Empetrich- to the laterality of the individual as in Ana- thys merriami (fig. 71C) the first proximal ra- bleps; a dextral male is illustrated. A liga- dial is fused and there is no first or second style has been found in Jenynsia males, as middle radial; the third middle radial is rep- in poeciliid males. In addition, the gonopo- resented by a minute ossification at the base dium of Jenynsia is similar to that of Ana- of the third proximal radial. bleps and differs from that of the poeciliids The proximal radials corresponding to the in having the proximal radials enlarged and shortened anal fin rays are greatly elongate angled forward to the left or right; there are in goodeids (fig. 70). They are not fused to- never enlarged hemal arches which project gether as in the other viviparous families, anteriorly to meet the proximal radials of the however. The proximal radials of Empe- anal fin which migrates anteriorly in its de- trichthys and Crenichthys are slightly elon- velopment within the poeciliids. gate; however, they differ little from that of The structure of the jenynsiid gonopodium a typical oviparous cyprinodont (fig. 22). differs from that described by Turner (1950) In the ontogeny of anal fin rays, all are who stated (p. 352): "most of the rays in the formed unbranched and then successively anterior part of the fin undergo absorption become branched. In both Oryzias and Men- . . . ." It agrees more with that of de Gil idia, the number of unbranched anal rays is (1949) who illustrated several variations in two, as it is in many cyprinodontoids. the formation of the fin, but in each case in- Among the aplocheiloid fishes which have dicated that all the fin rays were present. lost the first proximal anal radial, there are The anal fin of the goodeid males is rela- often three unbranched anal rays. The num- tively unmodified compared with those of the ber of unbranched rays varies among cyprin- poeciliid, jenynsiid, and anablepid fishes. odontiforms from no rays unbranched in an The structure is not properly termed a gon- occasional specimen of Profundulus (Miller, opodium, as the modifications of the anal fin 1955a) and in members of the genus Orestias elements themselves appear to have little to to all but one unbranched in some fundulines do directly with the transfer of sperm. and cyprinodontines. In all poeciliids, as well Goodeid males, however, are diagnosable as the genus Pantanodon and at least one on the basis of anal fin structure. The first nominal species of Aplocheilichthys, A. six or seven fin rays are shortened and un- johnstoni, there are three unbranched anal branched, and offset from the rest of the fin rays. In Anableps there are four in males and rays (fig. 70). The first anal fin ray is rudi- three in females, whereas there are two in mentary, and the middle radials of the first both males and females of Jenynsia. Among six or seven rays are fused to the base of the the goodeids, the unbranched anal fin rays proximal radials. Taken together with the typically number more than four. presence of trophotaeniae in embryos, these Considering the ontogeny of anal fin rays, characters were used by Miller and Fitzsi- it could be argued that a high number of un- mons (1971) to define the Goodeidae. branched rays is primitive, while successive- 438 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

HYP

HYP INCL

CARP ICARP INCLA

1CARM ICARM

FIG. 72. Diagrammatic representation of the generalized primitive state of the anal fin muscu- FIG. 73. Diagrammatic representation of the lature in cyprinodontiforms. derived anal fin musculature of cyprinodontiforms. ly lower numbers are derived. Conversely, Nelson (1975) discussed the mechanism of a certain number of rays could be primitive sperm transfer in the goodeids, and illustrat- for a group, and the suppression of branching ed the anal fin muscles. Such muscles in a derived modification. I accept the latter goodeids diverge most from the generalized viewpoint since it is the description of char- type in forming a large muscular mass sur- acters in their adult form whose distribution rounding the vas deferens and urinary tract must be analyzed without recourse to pre- in a structure which was termed a pseudo- sumptions about varying ontogenies. Thus, phallus by Mohsen (1961a). Along with this the increase in unbranched anal fin rays urogenital organ, is an elaboration of the in- above one or two is hypothesized to be a clinators of the anal fin. The inclinators, derived character among cyprinodontiforms. which arise between the hypaxial muscula- Anal Fin Musculature: The anal fin mus- ture and insert on the bases of the anal fin culature of the aplocheiloids which exhibit rays distal to the insertion of the erectors and internal fertilization, C. melanotaenia and depressors (Winterbottom, 1974), arise just C. brucei, is of the primitive type for cyprin- below the division of the epaxial and hypax- odontiforms (fig. 72). That is, there is a set ial musculature (fig. 73). Such elaborate in- of external inclinators as well as erectors and clinators, however, are not restricted to the depressors corresponding to the individual goodeids. They are found also in a group de- anal fin rays. One broad band of muscle, the fined above on the basis of skull and jaw infracarnalis medius runs from the base of specializations, viz., Empetrichthys, Cren- the pelvic fins to the first anal radial; a sec- ichthys, Cubanichthys, Chriopeoides, Ores- ond, the infracarnalis posterior runs from the tias, and the cyprinodontines. last anal radial to the distal tip of the last SPERM TRANSFER: Sperm transfer occurs hemal spine (Winterbottom, 1974). in poeciliids when the gonopodium is swung In male poeciliids, the anal inclinators are forward and folds over to one side or the drawn together into a fan-shaped mass of other to form a groove. Sperm pass down the muscle. In Anableps and Jenynsia, there are groove in unencapsulated bundles termed no such fan-shaped masses of muscles. The spermatozeugmata and are transferred to the inclinators are thickened, otherwise the mus- female by application of the gonopodial tip cles differ little from those of the generalized to, or within, the genital pore. type. In some poeciliids the pelvic fins are also 1981 PARENTI: CYPRINODONTIFORM FISHES 439 modified in the males. Clark and Kamrin and Jenynsia the phenomenon of dextral (1951) report that in a number of poeciliids males pairing with sinistral females (and vice tested, the pelvic fin of one side is swung versa) has been reported tentatively in the forward together with the gonopodium. They literature (e.g., Rosen and Bailey, 1963) al- speculated that the pelvics in such species though its occurrence is still doubtful (Miller, contribute to the formation of the sperm 1979). In theory, there are two kinds of groove. males, sinistral and dextral and correspond- In the poeciliids, procatopines, Pantano- ing types of females. A dextral male suppos- don and Fluviphylax, pelvic fins are set far edly has a gonopodium which is offset to the forward and are often under the pectoral fin right; therefore, he can only copulate with a bases (e.g., in the poeciliid Heterandria bi- sinistral female. The sidedness of a female is maculata, fig. 52 and in the procatopine Pro- determined by the placement of one or two catopus glaucicaudus, fig. 53). The thoracic scales over one side of the urogenital open- or subthoracic position of the pelvic fins in ing; hence a scale covering the left side of these groups results from an ontogenetic for- the opening defines a female as dextral. In- ward migration during sexual differentiation. dividuals, although they may easily have This fact has been well known within the their laterality determined, have not been poeciliids for some time (Clark and Kamrin, observed to be either exclusively dextral or 1951; Rosen and Kallman, 1959); however, sinistral in their mating (Miller, 1979). Thus, the phenomenon in the procatopines and oth- the significance of the anatomical modifica- er genera is little known. Trewavas (1974) tions related to sidedness remain specula- reported data for several species of Proca- tive. topus from West Cameroon to support the Although laterality is not evident among fact that the pelvic fins indeed migrate for- young males of Anableps examined, adult ward in ontogeny. The extent of this phe- males could easily be classified as left or nomenon among procatopines is not known right. Females' sidedness is also generally and could properly be explored with labo- easy to determine; however, some large ratory developmental series of representa- adult female Anableps seem to be neither left tives of all procatopine genera as well as Flu- nor right. viphylax and Pantanodon. In the oviparous Oxyzygonectes, a hy- Within Anableps and Jenynsia the tubular pothesized close relative of Jenynsia and gonopodium is associated with a distinctive Anableps, males have a distinct anal papilla mode of sperm transfer. In both genera, all which in preserved specimens has been ob- the rays of the anal fin are brought close to- served to be offset to the left or to the right. gether and surrounded by a fleshy tube (fig. The significance of this character is equivo- 66). The sperm duct enters the tube at the cal since it may simply be an artifact of pres- base of the first anal ray and follows the ven- ervation. Females offer no clue since they tral edge of the tube to its tip. Sperm do not possess a fleshy pouch surrounding the gen- travel down the tube in sperm bundles, but ital opening. Both male and female Oxyzy- individually. Grier, Burns and Flores (MS) gonectes, however, have scales around the report that partial sperm bundles are formed anterior region of the anal fin much like the in Anableps dowi, the presumed primitive pocket of scales surrounding the anus and species of the genus (Miller, 1979) but break first several anal fin ray bases considered to down before they enter the sperm duct; only be a diagnostic character of the procatopines free spermatozoa were observed in both the (Clausen, 1967). This pocket of scales ap- efferent and main testes ducts of A. anableps parently is a derived character of the larger and Jenynsia lineata. The abdominal pelvics group including the poeciliids, Jenynsia, An- of Jenynsia and Anableps are inferred to ableps, the procatopines, Fluviphylax and have no function in sperm transfer. Pantanodon as well as Oxyzygonectes. If so, Since Carman (1895) described the pres- these scales have been modified or reduced ence of sexual lefts and rights in Anableps in the viviparous forms. 440 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

Sperm bundles are formed in the goodeids forms. Among other poeciliids, the egg (Grier, Fitzsimons and Linton, 1978), al- retains an extremely reduced chorion (Zahnd though the precise mechanism of sperm and Porte, 1962; Flegler, 1977). Also among transfer is still unknown. Mohsen (1961a, the poeciliids are found some of the smallest 1961b) described a muscular organ surround- vertebrate eggs (Scrimshaw, 1946). ing the vas deferens and urinary canal and Ovoviviparity may be identified in certain believed that sperm bundles were ejected groups of poeciliids, for example Brachy- during copulation by a contraction of the or- rhaphis episcopi (Turner, 1938). The yolk gan. Nelson (1975) reported that during a sac is relatively large, and although devel- copulation attempt, the male clasps the fe- opment is internal, nutritional support is de- male: the anterior portion of the anal fin rived primarily from the yolk. formed by the shortened rays is wrapped Development of fertilized eggs in the rest around the genital opening of the female of the true viviparous poeciliids is of two while the notch in the fin is placed near the major types (Turner, 1939). In Heterandria anterior margin of the anal fin of the female. formosa, for example, a so-called pseudo- Thus, the urogenital organ of the male comes chorion and pseudoamnion are formed by very close to the female's urogenital open- the folding around the embryo of extraem- ing. bryonic somatopleure in the pericardia! re- True spermatophores, that is, encapsulat- gion. The outer membrane thus formed is ed sperm bundles, occur only in the adrian- termed the pseudochorion; it is highly vas- ichthyoid Horaichthys (Kulkarni, 1940). cularized, whereas the inner pseudoamnion Sperm bundles, however, occur among sev- is nonvascular. The pseudochorion is used eral teleost groups including the cyprinodon- as an organ for respiration and nutrition tiforms just mentioned and the exocoetoid throughout development. In species of Poe- Dermogenys whose sperm bundles are indis- ciliopsis, which also have small yolk re- tinguishable from those of the poeciliids serves, the lateral region of the somatopleure (Grier, Burns and Flores, MS). Free sper- is poorly developed. The ventral region be- matogonia, like oviparity, must be consid- comes highly vascularized and invades the ered a primitive character. However, the oc- region between villi formed in the follicular currence of spermatozeugmata with viviparity membrane; together they form what is among different relatively unrelated groups termed a follicular pseudoplacenta. Thus, suggests that spermatozeugmata have arisen the embryos of Heterandria formosa and independently along with viviparity. Poeciliopsis and related species derive the Fertilization and Development: The distri- greater part of their nutrition from maternal bution of characteristics related to egg reten- tissues rather than from stored yolk. Poecil- tion and maternal contribution to develop- iids also exhibit the phenomenon of super- ment precludes the ready division of members fetation, that is, the overlapping of devel- of the four so-called viviparous families into oping broods in the ovary (Scrimshaw, oviparous, viviparous and ovoviviparous 1944). groups. Anablepid, jenynsiid, and goodeid fishes Among the poeciliids, one genus and exhibit more elaborate adaptations for vivi- species, Tomeurus gracilis, is oviparous and parity. just facultatively viviparous (Rosen and Bai- In Anableps, both fertilization and devel- ley, 1963). Fertilization takes place as in all opment are intrafollicular as in poeciliids. In other poeciliids, within the follicle; however, addition, a follicular-pseudoplacenta more the developing embryo is quickly released elaborate than that of the poeciliids is also from the ovary into the oviduct and then developed. The ventral portion of the so- passed to the outside for the remainder of matopleure expands to form highly vascular- the developmental period. The egg of To- ized projections; the surrounding follicle is meurus has a thick chorion with adhesive fil- covered with vascular villi. In Anableps aments like that in oviparous cyprinodonti- dowi, the species presumed to be most prim- 1981 PARENTI: CYPRINODONTIFORM FISHES 441

itive, the large intestine expands and nearly ovigerous tissue, the structure of which is as fills this sac. Follicular fluid is absorbed by two folded masses, one in each section of the the embryo across intestinal villi. At birth, ovary. The first of these is apparently the the follicle ruptures and the expanded belly primitive state of the fused ovaries, and the sac eventually undergoes shortening. second the more derived state with oviger- In jenynsiids, although fertilization is in- ous tissue excluded from the walls and sep- trafollicular the embryo is evacuated from tum and confined to the middle of the ovarian the follicle and development takes place cavities. In one genus and species, Chara- within the ovary. Development is viviparous codon lateralis, there is an intermediate type rather than ovoviviparous since the yolk sup- of ovary which has ovigerous tissue both in ply is consumed at an early stage, and nutri- a short section of the septum and in weakly tion is derived mainly from maternal fluids. formed tissue extensions into each of the Respiration occurs across an expanded ven- ovarian cavities. This so-called intermediate tral somatopleural sac as in the anablepids. condition, however, may be more accurately Maternal fluids enter the developing embryo assessed as more closely related to the de- through its mouth or through the opercular rived type 2; that is, it forms a transition be- openings (Turner, 1940b). Flaps grow out tween the distinctly primitive and derived from the wall of the ovary and invade the types. opercular region and an intimate connection Trophotaeniae occur in three types: ro- between embryo and mother is provided as sette or ribbon-like, and when ribbon-like, the flaps of tissue invade the pharyngeal and sheathed or unsheathed. In a sheathed pro- buccal cavities. cess the external epithelium is separated In goodeids, as in jenynsiids, the eggs are from the internal connective tissue by a wide fertilized in the follicles and then released space, thus giving the external epithelium the into the ovarian cavity for development. appearance of a thin external covering Goodeids are characterized by the posses- (Wourms and Cohen, 1975). sion of trophotaeniae (Turner, 1937), elabo- Mendoza (1965) studied the ontogeny of rate outgrowths in the perianal region, the trophotaeniae and found that all are first epithelium of which possesses villi and is in- formed as rosette perianal outgrowths; these distinguishable from intestinal epithelium are then elongated in those goodeids which (Wourms and Cohen, 1975). Their function have ribbon-shaped trophotaeniae. Thus, as absorbers of nutritive ovarian fluids is in- based on ontogenetic information, the ro- ferred from their structure. sette type of trophotaeniae is primitive, Morphology of the trophotaeniae and of whereas the ribbon-like processes are de- the ovary served as the principal character for rived. The significance of a sheathed versus the last general revision of the Goodeidae by unsheathed process is questionable since the Hubbs and Turner. However, more recently, rosette is histologically identical to the Miller and Fitzsimons (1971) have reviewed sheathed process (Wourms and Cohen, the classification and concluded that the 1975). great degree of variability among these struc- Hubbs and Turner (1939) classified the tures makes them of little importance in phy- goodeids in four subfamilies: the Ataeniobi- logenetic studies. Miller and Fitzsimons did nae consisting of one genus and species, not propose a reclassification. Ataeniobius toweri which lacks trophotaen- Goodeid ovaries (median organs formed iae altogether and possesses the primitive by fusion of right and left anlagen) fall into type of fused ovary; the Goodeinae which one of two main types (Hubbs and Turner, possesses the primitive rosette type of tro- 1939): (1) an ovarian septum and outer wall photaeniae and the primitive type of fused composed of ovigerous tissues, the inner ovary; the Characodontinae, consisting sole- septum is often folded down the middle of ly of Characodon lateralis, which has a type the joint ovarian cavity; and (2) an ovarian of ovary distinctly intermediate between the septum and outer wall which is devoid of primitive and derived type, and sheathed rib- 442 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 bon-like trophotaeniae; and the Girardinich- of poeciliids would be judged as indepen- thyinae which has the derived type of ovary dently derived in two groups, once in the and sheathed ribbon-like trophotaeniae. poeciliids with intrafollicular gestation and If, as suggested, Characodon and the Gir- once in all other poeciliids. ardinichthyinae can be considered sister Furthermore, the tubular gonopodia groups on the basis of their sharing derived formed internally by enlarged and twisted trophotaeniae and ovaries, then the Good- anal rays with laterality determined internal- einae and possibly Ataeniobius represent an ly by offset proximal anal radials would have intermediate assemblage between the com- to be considered independently derived in mon ancestor of the goodeids and these two Anableps and Jenynsia. Neither of these de- more derived subfamilies. Thus, the Good- cisions is warranted by the information cur- einae is a paraphyletic subfamily as it has no rently known about adaptations for vivipar- defining characteristics. The status of Atae- ity, including anal and pelvic fin structures. niobius is still problematic since its lack of Therefore, Anableps and Jenynsia are con- trophotaeniae may be assessed as a loss sidered to be sister genera; this decision is character if, on the basis of other character- supported by the above argument and char- istics, it is assessed as more closely related acters from other systems as discussed. to the more derived goodeids. The problem Similarly, the development of trophotaen- with goodeid classification is that it is cur- iae in both the goodeids and the ophidioids- rently based only on transition series of two zoarcids does not support the close characters and does not take into account the relationship of these three groups. Taken in- apparent osteological and internal and exter- dependently without knowledge of other nal morphological differences among genera. characters, trophotaeniae in these three vi- SUMMARY OF VIVIPARITY: The peculiarity viparous groups would indeed suggest their of cyprinodontiform reproduction in general close relationship. However, it is only after is a long developmental period associated an assessment of all apparently derived with an early breeding habit. structures that a hypothesis of convergent Viviparity occurs in teleosts outside cy- characters such as the trophotaeniae may be prinodontiforms in the Hemirhamphidae and made. Oryziatidae in atherinomorphs, and for ex- Thus, hypotheses of convergences con- ample, zoarcids, scorpaenoids, and ophid- cerning internal fertilization and viviparity ioids in the percomorphs. In addition, tro- , are the following: photaeniae are not unique to the goodeids; they occur in nearly the same form and 1. Internal fertilization occurs indepen- therefore have the same inferred function in dently among the cyprinodontiforms at least embryos of the zoarcids and ophidioids five times; in Rivulus and Cynolebias among (Wourms and Cohen, 1975; Cohen and the aplocheiloids, and in poeciliids, good- Wourms, 1976). The similarity of the gono- eids, and in Jenynsia-Anableps among the podia of Tomeurus and Horaichthys has cyprinodontoids. long been recognized. However, even if it is 2. Gonopodia occur independently three allowed that each of these characteristics times; in Cynolebias brucei among aplo- when it appears in cyprinodontiforms is pos- cheiloids and, in poeciliids and in Jenynsia- sibly a uniquely derived character, this pos- Anableps among the cyprinodontoids. sibility may only be evaluated with the use 3. Intrafollicular gestation occurs indepen- of data from other systems. For example, in- dently in a group of poeciliids and in Ana- trafollicular gestation in poeciliids and Ana- bleps. bleps, if evaluated as a derived character 4. Spermatozeugmata, which occur in all uniting them into a monophyletic group, viviparous cyprinodontiforms except Jen- makes the family Poeciliidae as currently ynsia and outside the cyprinodontiforms, are constituted a paraphyletic group. That is, the considered to be independently derived in complex gonopodia and derived pelvic fins poeciliids, Anableps and goodeids. 1981 PARENTI: CYPRINODONTIFORM FISHES 443

These assessed convergences are support- GROUP D: Hypothesized to be monophy- ed by the following conclusions: letic because its members share the following 1. The poeciliid gonopodium and gonopo- derived characters: premaxillary ascending dial suspensorium as described herein is a processes narrow or absent in adults, while unique complex of characters defining this at least weakly formed in embryos or juve- group. niles; rostral cartilage greatly reduced or ab- 2. The gonopodia of Anableps and Jen- sent; inner arms of the maxillaries do not ynsia represent a unique form among teleosts abut the rostral cartilage, yet remain at- and support the sister group relationship of tached to it, when present, by connective tis- these two genera. sue; and the lateral ethmoid having a greatly 3. The goodeid manner of internal fertil- reduced facet for articulation of the auto- ization with its associated modifications of palatine. the anal fin and presence of a copulatory or- FUNDULINES: The term as used through- gan as well as a derived ovary and presence out this study refers to species of the follow- of trophotaeniae defined this family as a ing nominal genera: Fundulus Lacepede, monophyletic group. Other modifications- with four included subgenera Zygonectes support the relationship of goodeids to two Agassiz, Xenisma Jordan, Plancterus Gar- North American genera, Empetrichthys and man, and Fundulus; Lucania Girard, Adinia Crenichthys, and then to a larger group of Girard, and Leptolucania Myers. The last oviparous cyprinodontoids. two genera are monotypic, Lucania com- prises three species, whereas the genus Fun- dulus, the largest of this group, comprises CLADISTIC SUMMARY OF THE from 30 to 35 species (Miller, 1955b; Lazara, CYPRINODONTOIDS 1979). Profundulus (of northern Central America They are hypothesized to form a mono- and southern Mexico, fig. 74) is hypothe- phyletic group on the basis of the following sized to be the most primitive cyprinodon- derived characteristics of the jaw and skull: toid genus. inner arms of the maxillaries directed ante- Five species are currently recognized in riorly and often with pronounced hooks; and two subgenera (Miller, 1955a): subgenus snout pointed and drawn anteriorly with the Profundulus Hubbs with two species, punc- autopalatine projecting anteriorly and not ar- tatus (Giinther) and guatemalensis (Giin- ticulating with the lateral ethmoid. ther); and Alvarez and Carranza with Relationships of the recognized genera as three species, labialis (Giinther), candalar- described below are summarized in the ius Hubbs and hildebrandi Miller. The genus cladogram of figure 75; their distribution is is defined by a high number of gill rakers on given in figure 76. the first arch. The number ranges from 14 to Two species, Fundulus kansae and zebri- 23 (modally 16), whereas there are four to nus, currently placed in the subgenus Planc- 14, typically fewer than 12, in Fundulus terus are distinguished from other fundulines (Miller, 1955a), as well as other fundulines, by having a posttemporal with an ossified, and a majority of the cyprinodontoids. In the rather than ligamentous, ventral limb. This aplocheiloids, the highest number observed character is primitive for cyprinodontiforms is 21 in Nothobranchius microlepis. The and may be used to separate Plancterus from number of gill rakers is increased in some other fundulines. The two nominal species cyprinodontoids, for example, in Lam- are regarded as probable conspecifics (Mil- prichthys and Pantanodon. However, the ler, 1955b). The genus Plancterus may also high number in Profundulus is considered to be distinguished by a derived character be a unique increase and therefore a derived shared by the two species; that is, a rather character of the genus. An additional aut- long and convoluted intestine (Garman, apomorphy of Profundulus is the relatively 1895). large autopterotic fossa. Nominal species of the genera Fundulus 444 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

\ GULF OF h I MEXICO y^" 9

16"

°0c- ^-^^ri

98° 90°

FIG. 74. Distributional limits of Pwfundulus. (After Miller, 1955a.)

(excluding zebrinus and kansae), Lucania, significance of the sequence; he therefore did Leptolucania, and Adinia are linked by the not recognize the subgeneric divisions of absence of a ventral limb on the posttem- Fundulus. poral. Farris (1968) performed a phylogenetic In the most recent review of Fundulus, analysis of the species of Fundulus and Pw- Miller (1955b) listed 26 species in what he fundulus and concluded that Fundulus was called an approximate phylogenetic se- a monophyletic genus comprising the follow- quence, although he did not elaborate on the ing monophyletic subgenera: Plancterus,

PLANCTERUS FUNDULUS LUCANIA ADINIA LEPTOLUCANIA •i

FIG. 75. Cladogram of relationships of the fundulines. Node A: inner arms of the maxillaries directed anteriorly, often with pronounced hooks; snout pointed and drawn anteriorly with the autopalatine projecting anteriorly and not articulating with the lateral ethmoids; Node B: convoluted intestine; Node C: Posttemporal lacks an ossified lower limb; Node D: expanded articular process of the second phar- yngobranchial; Node E: epipleural ribs meet the parapophyses of the abdominal ribs at their tips; reduction of the supraorbital pores; Node F: Block of cartilage between the interarcual cartilage and articulation point of the second pharyngobranchial; Node G: No supraoccipital processes; Node H: Quadrangular body form, first pleural rib arises on parapophysis of first vertebra; Node I: three bran- chiostegal rays, first postcleithrum absent, large black ocellus at midbody and on the caudal peduncle. 1981 PARENTI: CYPRINODONTIFORM FISHES 445

FIG. 76. Distributional limits of the fundulines.

Xenisma, Zygonectes, and Fundulus. Unique to be determined. However, since the genus defining characters of the genus and subgen- is considered to be monophyletic without era were not enumerated. Plancterus, the subgenera Xenisma and Brown (1957) divided the species of Fun- Zygonectes for convenience are treated as dulus into the subgenera Fundulus, Fontin- synonyms of Fundulus in this study. us, Plancterus, and Zygonectes without giv- The remaining funduline genera (Lucania, ing defining characters of each. Adinia, and Leptolucania) lack the broad With Plancterus removed from the genus, articular process of the second pharyngo- Fundulus can be defined in having a broad branchial of Fundulus, but are defined by articular surface on the second pharyngo- having epipleural ribs attaching directly to branchial (fig. 46A). parapophyses rather than to pleural ribs. In The interrelationships of the species of the other cyprinodontiforms the epipleural ribs subgenera Xenisma, Zygonectes, and Fun- attach ventrally along the proximal extent of dulus have yet to be formally investigated. the pleural ribs rather than being in contact Wiley and Hall (1975) and Wiley (1977) have with the parapophyses (e.g., in Aphyose- presented the only such revision of a group mion occidentale, fig. 26). of Fundulus species, the nomV-complex and A reduction in the number of supraorbital three other species all of which would be in- pores might be an additional derived char- cluded in but not totally comprise the sub- acter of the three genera; however, the dif- genus Zygonectes. As such, the interrela- ficulty of postulating transition series within tionships of species and of the subgenera as the system has been noted already. well as the limits of both categories remain There is evidence that each of the three 446 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 77. Sketch of body form and fin position of Adinia xenica, male. Dotted line approximates base of hypural plate. (After Rosen, 1973b.) genera is monophyletic. Lucania possesses The retention of five generic categories an apparently unique modification of the dor- comprising the fundulines is deemed appro- sal gill arches (fig. 46B). There is a small priate since although Fundulus is considered block of cartilage between the interarcual to be monophyletic, all its species have not cartilage and the articulation point of the sec- been examined. It is possible a more parsi- ond pharyngobranchial. No such indepen- monious interpretation would place some dent cartilage has been found in other cy- species of Fundulus as more closely related prinodontiforms, yet it is present in the to Lucania, Leptolucania or Adinia; there- species of Lucania examined. fore, a synonymy of all genera now would The genus Adinia is readily separated obscure interrelationships. from the other funduline genera by its later- CLADISTIC SUMMARY OF FUNDULINES: ally compressed, deep-bodied form (fig. 77), Fundulines share several derived characters described as diamond-shaped or quadrangu- of the jaw and skull: inner arms of the max- lar. In addition, Adinia is unique among cy- illaries are directed anteriorly and often have prinodontiforms in having the first pleura! rib pronounced hooks, the snout is pointed and arising on the parapophysis of the first ver- drawn anteriorly, while the autopalatine tebra rather than the second. does not articulate with the lateral ethmoid. Adinia and Leptolucania lack epiotic and Plancterus is defined by its long and con- supraoccipital processes found among all voluted intestine. fundulines and generally among cyprinodon- The four remaining funduline genera, Fun- tiforms. This condition is considered second- dulus, Lucania, Adinia, and Leptolucania, arily derived within these two genera. are defined as a monophyletic group by their Leptolucania ommata is a diminutive fun- sharing a posttemporal with a ligamentous duline rarely reaching over 20 mm. standard rather than ossified ventral limb. length. It is unique among fundulines and Fundulus is defined by an expanded artic- apparently all cyprinodontiforms in possess- ular surface of the second pharyngobranchi- ing just three branchiostegal rays. These rays al. The subgenera Xenisma and Zygonectes number from four to six among other cyprin- are treated as synonyms of Fundulus. odontiforms. Leptolucania alone among fun- Lucania, Adinia, and Leptolucania are dulines also lacks the first postcleithrum. defined as a monophyletic group by having The pattern of coloration is also unique. epipleural ribs attaching directly to par- There is a large black ocellus on the caudal apophyses rather than to pleural ribs. peduncle and another at midbody. Lucania is defined by a small block of car- 1981 PARENTI: CYPRINODONTIFORM FISHES 447

FIG. 78. Distributional limits of Jenynsia, Anableps, and Oxyzygonectes.

tilage between the interarcual cartilage and in aplocheiloids, Profundulus and the fun- articulation point of the second pharyngo- dulines, and (2) an enlarged dorsal process branchial. of the maxilla directed over the premaxillary Adinia has a unique quadrangular body ascending process. form as well as the first pleural rib arising on Valencia: Unique among cyprinodonti- the parapophysis of the first rather than the forms in having greatly expanded dorsal pro- second vertebra. cesses of the maxillaries. The posttemporal Adinia and Leptolucania lack epiotic and also has a ligamentous ventral limb. supraoccipital processes. GROUP F: Members of Group F share Leptolucania has just three branchiostegal three derived features: (1) the ascending pro- rays, and lacks the first postcleithrum, and cesses of the premaxillaries are short and has a derived color pattern characterized by narrow; (2) the dorsal processes of the max- a large black ocellus on the caudal peduncle illaries are rounded or greatly reduced; and and another at midbody. (3) the nasals are expanded medially in near- GROUP E: TWO derived characters define ly all members. Group E: (1) maxilla with a straight proximal GROUP H: The members of Group H share arm, rather than the overtly twisted arm as the following derived characters: maxilla 448 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

AN ABLE PS JEN YN SI A O X YZ YG ON EC T E S epiotic and supraoccipital processes, and outer and inner jaw teeth with lateral cusps in at least embryos and juveniles. A third possible derived character is laterality, ex- pressed as a shift in the gonopodium (in the case of Anableps and Jenynsia) or the gen- ital papilla (in the case of Oxyzygonectes) to either the left or the right. Anableps and Jenynsia are hypothesized close relatives (fig. 79) based on their sharing specializations of the gonopodium and gon- opodial suspensorium. The thickened and FIG. 79. Cladogram of relationships of Jen- elongated anal rays are twisted around each ynsia, Anableps, and Oxyzygonectes. Node A: other and covered by a fleshy tube. The enlarged supraoccipital and epiotic processes; sperm duct enters the tube at the anterior outer and inner teeth with lateral cusps in at least base of the anal fin and opens to the outside juveniles and embryos; sexual laterality; Node B: at the distal tip of the gonopodium which, in thickened and elongated anal rays in male, twisted turn, is offset to either the left or the right of around each other, covered by a fleshy tube; tu- the midline, whereas the expanded proximal bular sperm duct; gonopodium offset either to the radials are offset to either the left or right of left or the right; proximal anal radials enlarged, nearly reaching vertebral column and offset to the the vertebral column. left or the right; derived pigmentation pattern (see Females of Jenynsia and Anableps do not text); Node C: eyes divided horizontally; supraor- have a complete pouch of scales surrounding bital processes of frontals expanded; greatly re- the urogenital opening. Instead, there are duced premaxillary ascending processes in adults; just one or a few scales covering the left or dumbbell shaped rostral cartilage; pectoral fins the right side of the opening defining a female lowset and rays increased to 20-23; vertebrae in- as either dextral or sinistral, respectively. creased to 45-54; supraorbital pores reduced; in- The pigmentation pattern of several trafollicular gestation; formation of spermato- species of Jenynsia (e.g., eigenmanni) zeugmata; gonopodium formed principally from closely approaches that of Anableps micro- anal rays 3 through 9; Node D: gonopodium formed principally from anal rays 3, 6, and 7; oth- lepis (von Ihering, 1931). On the sides of the er anal rays undergo degeneration; tubular gono- body are several rows of dashes of dark pig- podium lacks scales; embryo receives nourish- ment over a pale yellow-green background. ment via ovarian flaps; tricuspid outer jaw teeth If the very regular pattern of A. dowi (as in adults; Node E: greatly expanded premaxillary shown in Miller, 1979) consisting of a longi- ascending processes; no rostral cartilage; five or tudinal yellow stripe on a dark dorsal surface six rows of tricuspid inner jaw teeth; fleshy sheath and light ventral surface is defined as derived over urogenital opening in females; males with for that species, then Jenynsia and Anableps large anal papilla; preopercular pores covered share some derived features of color pat- with enlarged scales; males with from four to five terns. precaudal bars. Jenynsia has a unique formation and de- velopment of the rays of the gonopodium. The principal rays of the gonopodium (3, 6, with an expanded distal arm; a parasphenoid and 7) are elongated and elaborated. All oth- with an expanded anterior arm; dorsal pro- er anal rays undergo degeneration; they are cess of the maxilla with a distinct lateral in- weakly formed and may easily be reported dentation; an elongate retroarticular; and a as absent in a cursory examination of the fin. pouch created by scales surrounding the uro- In addition, the outer covering of the gono- genital opening of females. podial tube is scaleless, whereas in Anableps Jenynsia, Anableps, AND Oxyzygonectes: it is fully scaled. If the presence of scales is These three genera (fig. 78) share enlarged primitive in the development of the gono- 1981 PARENTI: CYPRINODONTIFORM FISHES 449 podial tube, then their absence in Jenynsia ity in Anableps from that of Jenynsia, par- is derived. ticularly the intrafollicular gestation and The development of Jenynsia embryos is formation of sperm bundles in at least one also unique in the formation of the large species, A. dowi. These characters appear in ovarian flaps which enter the pharyngeal and other viviparous cyprinodontiforms and ath- buccal cavities and are believed to supply erinomorphs, however, they must properly nourishment to developing embryos. be regarded as independently derived in An- In addition, the species of Jenynsia may ableps within this scheme. be identified by their outer tricuspid teeth in The monotypic Oxyzygonectes, previ- adults. However, given that the character is ously classified in the subfamily Fundulinae, expressed to some degree by Anableps and exhibits a further derived state of the upper Oxyzygonectes, it should be considered jaw exhibited by Jenynsia and the procato- primitive in Jenynsia and unicuspid teeth de- pines. The premaxillary ascending processes rived in Anableps and Oxyzygonectes. are greatly expanded in adults (fig. 35B). The genus Anableps is defined by a num- Also, no rostral cartilage has been found in ber of derived features. The eyes are divided the specimens examined. The difference of horizontally such that the fish has simulta- this condition relative to that in Aplocheilus, neous aerial and aquatic vision (fig. 66). The which also has expanded premaxillary as- frontals are greatly expanded dorsally to ac- cending processes, may be observed by com- commodate the enlarged orbits. The derived paring the lateral views of jaws of the two. upper jaw of Anableps is also readily distin- In Aplocheilus (fig. 27) the ascending pro- guished from all other cyprinodontiforms: cesses are longer relative to the length of the there are weakly formed premaxillary as- dentary than they are in Oxyzygonectes (fig. cending processes in adults; the remaining 36) in which the ascending processes are inner arms of the premaxillaries meet in the shortened. midline to form an arc. The ascending pro- In addition there are five or six rows of cesses are present in embryos, therefore, the tricuspid teeth on both the upper and lower adults represent the derived condition of this jaws, three to four more rows than found in state. Also, the rostral cartilage is dumbbell Jenynsia and Anableps. shaped. Female Oxyzygonectes may be distin- The pectoral fins are set ventrally; inter- guished from males by the presence of a nally this is effected by the radials being set fleshy sheath covering the urogenital opening ventrally. Pectoral rays are increased to 20- which itself is partially covered with scales. 23, as opposed to 15 in Jenynsia. Males have a large anal papilla which, as The number of vertebrae is higher than in stated, has been found in preserved speci- any other cyprinodontiform genus. The num- mens offset to the left or the right. ber ranges from 45 to 54 (Miller, 1979); In life, males have from four to five pre- whereas the highest number in other cyprin- caudal bars and faintly mottled unpaired fins, odontiforms is 41 in the genus Lamprich- as compared with the more drab females thys. (Daniel Fromm, personal commun.). Back- Gill rakers on the first arch are also in- ground coloration in both males and females creased to 21-30 compared to 10-11 in Jen- is a drab dark brown in preservation. ynsia. The preopercular sensory pore system is The supraorbital pore system is reduced represented by the primitive number of sev- from the general condition exhibited by Jen- en pores. These are set apart and covered ynsia to what have been identified as pores slightly by a series of large scales. 3 and 4a and, 6 and 7; in addition, the head CLADISTIC SUMMARY OF Jenynsia, Ana- scales are arranged randomly, rather than in bleps, AND Oxyzygonectes: Enlarged epiotic the primitive pattern. and supraoccipital processes, outer and in- Other derived characters related to repro- ner jaw teeth with lateral cusps in at least duction distinguish the condition of vivipar- embryos and juveniles, and sexual laterality 450 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 80. Distributional limits of poeciliids, procatopines, Fluviphylax and Pantanodon. (After Rosen and Bailey, 1963; Turner, 1966.) are shared by Jenynsia, Anableps and Oxy- ascending processes, and a dumbbell shaped zygonectes. rostral cartilage; pectoral fins set ventrally, Jenynsia and Anableps share derived and an increase in pectoral fin ray number to characters of the gonopodium and gonopo- more than 20; reduced supraorbital pore sys- dial suspensorium (elaborate anal rays are tem and head scales randomly arranged; for- twisted around each other and covered by a mation of spermatozeugmata; intrafollicular fleshy tube; the sperm duct enters the tube gestation; and vertebrae increased to 45 or at the anterior base of the anal fin and opens more. to the outside at the distal tip of the gono- Oxyzygonectes is defined by enlarged pre- podium; gonopodium is offset to the left or maxillary ascending processes; the absence right of the midline; proximal radials are off- of a rostral cartilage; an increase in the num- set to the left or right of the vertebral col- ber of rows of outer tricuspid teeth; males umn) and of general pigmentation. with anal papilla; females with anal pouch; Jenynsia is defined by its unique form of males with four to five precaudal bars and the gonopodium (anal rays 3, 6, and 7 are faintly mottled unpaired fins; and large scales elaborate, whereas all other anal rays covering the preopercular canal. undergo degeneration; and the gonopodium POECILIIDS, Fluxiphylax, Pantanodon AND is scaleless) and development (ovarian flaps THE PROCATOPINES: This group (fig. 80) is enter pharyngeal and buccal cavities to pro- hypothesized to be monophyletic by its vide nourishment). members sharing five derived characters: (1) Anableps is defined by enlarged eyes, di- pectoral fins set high on the sides caused by vided horizontally and accommodated by ex- the dorsal placement of the radials; (2) a de- panded frontals; weakly formed premaxillary rived hyoid bar with no ventral extension of 1981 PARENTI: CYPRINODONTIFORM FISHES 451

POECILIIDS PIOCHEIUCHTHYS LAMPRICHTHYS PANTANODON '• APL OC H E I L IC H T H YS " H YPS O P ANC H AX PRQCATOPU!

FIG. 81. Cladogram of relationships of poeciliids, procatopines, Fluviphylax and Pantanodon. Node A: pectoral fins set high on the sides, caused by the dorsal placement of the radials; hyoid bar with no ventral extension of the anterior ceratohyal accompanied by an expanded ventral hypohyal; pleural ribs on the first several hemal spines; anterior placement of the pelvic fins and their anterior migration during growth; recessed supraorbital pores 2b through 4a. Node B: gonopodium in males formed principally from anal rays 3, 4, and 5; modified hemal arches to provide support for the gonopodium; expansion of the inclinators of the anal fin to form a fan-shaped mass; modified pelvic fins of males; expansion of fourth epibranchial to become main support of the dorsal gill arch elements; exoccipital condyles absent; neural arches open. Node C: Enlarged eyes and reduced preorbital space; vomer unossified. Node D: mesethmoid cartilaginous. Node E: first postcleithrum absent; anal fin of 14 rays or more; expanded swimbladder extends beyond the first two to five hemal spines. Node F: increased number of vertebrae; ctenoid scales; caudal fin lyre-shaped; posttemporal with unossified lower limb. Node G: vomer tends to be unossified; no interarcual cartilage. Node H: tricuspid pharyngobranchial teeth; enlarged second pharyngobranchial toothplate; epibranchials one through three absent; hypobranchials absent; exoccip- ital condyles absent; neural arches of first vertebra expanded and applied to skull; fin spines present. Node I: robust dentary; articular reduced. Node J: reduction of alveolar arm; teeth extend to distal tip of premaxilla; reduction of the spatulate distal arm of the maxilla. Node K: Deep-bodied, caused by enlarged pleural ribs. Node L: Branchiostegal rays of males free from the membrane and extend pos- teriorly. the anterior ceratohyal accompanied by the The interrelationships of the poeciliids and ventral hypohyal forming a bony cap over its other genera included in this group (fig. 81) anterior facet; (3) pleural ribs on the first sev- are considered tentative because of the na- eral hemal arches; (4) the thoracic or sub tho- ture of some evidence, as discussed below, racic pelvic fins that migrate anteriorly dur- the scarcity of some material (particularly of ing growth; and (5) recessed supraorbital the genus Pantanodon) and the large number sensory pores 2b through 4a. Also, Clausen of species in a polyphyletic genus (Aplo- (1959) described what he considered to be a cheilichthys). behavioral convergence between the proca- The poeciliid fishes are easily distin- topines and the poeciliid subfamily Poecili- guished from the other fishes of this group inae. This included an encounter in which by the presence of a gonopodium in males the male "dances" before the female, formed principally from rays 3, 4, and 5 of spreads his fins and moves backward and the anal fin, modifications of the hemal arch- forward. He also noted similarities in shape es to provide support for the gonopodium, and coloration, without qualification. expansion of the inclinators of the anal fin to 452 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 form a fan-shaped mass of muscles, and cheilichthys Bleeker: Micropanchax Myers, modifications of the pelvic fins of males and and its subgenus Lacustricola Myers; Con- their inferred function during copulation. gopanchax Poll, Poropanchax Clausen, Cy- Also, the dorsal gill arches express a de- nopanchax Ahl, Plataplochilus Ahl, and rived character in the expansion of the fourth Platypanchax Ahl. epibranchial to become the main support of Platypanchax is apparently closely related the dorsal gill arch elements. to another procatopine genus, Hypsopan- As discussed for aplocheiloid interrela- chax, for reasons discussed below. The re- tionships, the polarity of sexual dimorphism maining genera and subgenera contain some related to size cannot be determined. How- species that possess characters derived for ever, it is generally true that among cy- more exclusive groups of procatopines. prinodontoids females are larger than males. (Specimens of Cynopanchax and Plataplo- This is the case for the poeciliids and is chilus were not available for study.) therefore considered primitive. Among the All species of Aplocheilichthys examined procatopines, Fluviphylax and Pantanodon, possess a cartilaginous mesethmoid, there- males are generally much larger than fe- fore, are properly included in the group. males. Given the general nature of this char- Lamprichthys, Procatopus, Hypsopanchax, acter among cyprinodontoids, it may only be Hylopanchax, Platypanchax and their in- interpreted as a secondarily derived charac- cluded subgenera are distinguished from ter suggesting the close relationship of these more primitive species of Aplocheilichthys three groups. by lacking the first postcleithrum as well as Another derived character which all three having an elongate anal fin of 14 rays or share is the absence of parietals. Parietals more, and an expanded swimbladder which are absent from some derived poeciliids, extends beyond the first two to five hemal however, they are present in Tomeurus. spines. Therefore, their presence must be consid- The first postcleithrum is present in ered primitive for poeciliids. Aplocheilichthys spilauchena, the type The interrelationships of the poeciliids, species, yet is lacking in A. johnstoni. It is Fluviphylax, and the procatopines and Pan- present in Poropanchax. tanodon is depicted as a trichotomy (fig. 81). Within this group of procatopines and Larger males and absence of parietals sug- Pantanodon, Lamprichthys may be consid- gest the close relationship of Fluviphylax to ered the relatively primitive member since it Pantanodon and the procatopines. How- possesses a fully ossified vomer and an in- ever, the diminutive South American Fluvi- terarcual cartilage, as does A. spilauchena. phylax has a derived resemblance to the poe- The interarcual cartilage is absent, there- ciliid Tomeurus in two respects: (1) the fore, presumed lost in Procatopus, Hypso- dorsal fin is small, composed of just four to panchax, Pantanodon, and A. johnstoni. six rays and is set rather far back on the Thus, members of the genus Aplochei- body, and (2) the pectoral fins are extremely lichthys are related at different levels of gen- reduced, being composed of just nine or ten erality to the other procatopine genera. rays. It is proposed that the more derived mem- A derived character of the procatopines bers of the genus, of which A. johnstoni may and Pantanodon is the presence of a cartilag- be considered representative, be referenced inous rather than an ossified mesethmoid. by the genus "Aplocheilichthys." The mesethmoid is ossified in both Fluvi- The state of the ossified versus unossified phylax and the poeciliids. vomer is rather ambiguous. The vomer is ful- The genus Aplocheilichthys as currently ly ossified in the specimens of Lamprichthys constituted is polyphyletic. It contains over examined. However, in the genus Hypso- 60 nominal species (Lazara, 1979). These panchax, one species, platysternus, has an species have been divided into the following ossified vomer, whereas another species, ze- genera and subgenera in addition to Aplo- bra, has an unossified vomer. Nonetheless 1981 PARENTI: CYPRINODONTIFORM FISHES 453 these two species of are considered to be The dentary (fig. 37) is greatly expanded me- members of the same genus. Therefore, the dially, more so than in any other group of character may be more properly character- cyprinodontoids, while the retroarticular is ized as the tendency to not ossify the vomer. reduced. The vomer is similarly unossified in Fluvi- Within this group, Procatopus, Hylopan- phylax, therefore, this may be an additional chax, and Hypsopanchax together are de- derived character indicating Fluviphylax is fined as monophyletic by the reduction of the more closely related to the procatopines and alveolar arm of the premaxilla and the exten- Pantanodon than to the poeciliids. The am- sion of teeth to near the distal tip (fig. 37). biguous nature of the character precludes This is accompanied by a reduction in the this conclusion at this time, however. spatulate distal arm of the maxilla, distinc- The diminutive Fluviphylax pygmaeus is tive in other members of the larger group in- denned by its greatly enlarged eyes. This cluding Jenynsia, Anableps, and Oxyzygo- condition is accompanied by a reduced nectes. preorbital distance as well as a narrow rather The species of Procatopus exhibit a de- than wide lacrimal. rived character by which they may be readily The genus Lamprichthys, endemic to distinguished from all other cyprinodontoids. Lake Tanganyika is a pelagic cyprinodonti- In males, the first two branchiostegal rays form. It is one of the largest members of the are prolonged and extend beyond the oper- group, attaining a standard length of over cular margin (fig. 53). Clausen (1959) divided 150 mm. Vertebrae typically number 41, the species into two subgenera, Procatopus whereas the average number for this group is Boulenger and Andreasenius Clausen. The 30. pelvic fins in members of Andreasenius are Males are distinguished from females by set farther back on the body than those of the presence of distinct ctenoid scales. the subgenus Procatopus. However, both A third derived character is the shape of possess the distinctive branchiostegal ray the caudal fin. Typically in cyprinodonti- character, therefore, Andreasenius is treated forms, the caudal fin is rounded or truncate. as a synonym of Procatopus. Similarly, the In Lamprichthys, the upper and lower cau- genus Hylopanchax Poll and Lambert com- dal fin rays are extended to form a lyre- prising just one species, silvestris, also has shaped caudal fin. Internally the supports are prolonged branchiostegal rays; therefore, it typical of those for cyprinodontiforms, how- is also considered to be a synonym of Pro- ever. catopus. (Poll and Lambert [1965] consid- A fourth derived character is a straight ered it to be intermediate between Hypso- posttemporal. As in other groups with such panchax and Procatopus; however, the posttemporals the ventral limb is represented presence of this derived character clearly in- by an unossified ligament. dicates its closer relationship to Procato- Among the genera that lack a vomer and pus.) an interarcual cartilage, Pantanodon is de- Hypsopanchax is distinguished from all fined by the following derived characters: tri- other procatopines by its deep abdominal cuspid pharyngobranchial teeth, greatly en- keel effected internally by enlarged ribs. larged second pharyngobranchial toothplate, Based on its description, the sole species of outer pelvic rays of males curved and elon- the genus Platypanchax is placed in Hyp- gate, exoccipital condyles absent, neur- sopanchax; however, specimens have not apophyses of the first vertebra expanded and been examined. closely applied to the skull, fin spines pres- CLADISTIC SUMMARY OF POECILIIDS Flu- ent, the lacrimal reduced, and the absence of viphylax, Pantanodon, AND PROCATOPINES: hypobranchials. This group is defined by five derived char- The remaining genera "Aplocheilichthys," acters: (1) pectoral fins highset, caused in- Procatopus, Hylopanchax, and Hypsopan- ternally by dorsally placed radials; (2) ven- chax share an extremely robust lower jaw. tral hypohyal forming a bony cap over the 454 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 anterior facet of the anterior ceratohyal; (3) cartilage, and the tendency for the vomer to pleural ribs on the hemal arches; (4) thoracic be unossified. or subthoracic pelvic fins which migrate or Pantanodon is defined by seven derived are inferred to migrate in ontogeny from a characters: tricuspid pharyngobranchial more posterior position; and (5) recessed su- teeth; enlarged second pharyngobranchial praorbital pores 2b through 4a. toothplate; epibranchials one through three The poeciliids, Fluviphylax and the pro- absent; hypobranchials absent; exoccipital catopines and Pantanodon form an unre- condyles absent; neural arches of first ver- solved trichotomy. Fluviphylax and Tomeu- tebra expanded and applied to the skull; and rus share a dorsal fin of four to six rays set fin spines present. far back on the body, and pectoral fins of "Aplocheilichthys," Procatopus and just nine or 10 rays. Fluviphylax shares an Hypsopanchax all have a robust dentary and absence of parietals and the condition of reduced articular. males larger than females with Pantanodon Procatopus and Hypsopanchax have a re- and the procatopines. duced alveolar arm of the premaxilla, with The poeciliids are defined by the following teeth extending to its distal tip, and a reduc- derived characters: internal fertilization by tion of the distal arm of the maxilla. a gonopodium formed principally from anal Procatopus is defined by several branchio- rays 3,4, and 5; modified hemal arches pro- stegal rays free from the branchiostegal viding support for the gonopodium; expand- membrane and extending posteriorly in ed inclinators of the anal fin; modified pelvic males. Hylopanchax is treated as its junior fin rays in males; expansion of the fourth synonym. epibranchial to become the main support of Hypsopanchax, of which Platypanchax is the dorsal gill arch elements; exoccipital con- considered a junior synonym, is defined by dyles absent, and neural arches open, not its deep abdominal keel. forming a spine. The two nominal genera Cynopanchax Fluviphylax is defined by enlarged eyes and Plataplochilus were not examined, and and reduced preorbital space, as well as a are simply listed in the systematic accounts. possibly secondarily derived unossified vo- GROUP G: The members of this group mer. share the following derived characters: lat- The procatopines and Pantanodon are de- eral ethmoid expanded medially and oriented fined by a cartilaginous mesethmoid. so that it lies roughly perpendicular to the Aplocheilichthys is polyphyletic as cur- frontal; reduced autopterotic fossa; and in- rently constituted. The genus is maintained clinators of the anal fin greatly enlarged. to comprise the most primitive procatopines. Empetrichthys, Crenichthys, AND THE The more derived members of the genus, re- GOODEIDAE: The three taxa (fig. 82) share ferred to the genus "Aplocheilichthys" share four derived characters: (1) the first two to three derived characters with the remaining seven middle anal radials are fused to the procatopine genera: absence of the first proximal radials; (2) dorsal processes of the postcleithrum; anal fin rays increased to 14 maxillaries are greatly reduced; (3) the distal or more; and a swimbladder extending be- arm of the premaxilla is straight, and (4) the yond the first two hemal spines. articular is reduced. Lamprichthys is defined by four derived Empetrichthys and Crenichthys are hy- characters: increased number of vertebrae; pothesized to be sister taxa (fig. 83) based on ctenoid scales; a lyre-shaped caudal fin; and, their lack of pelvic fins and fin supports, and a posttemporal with a ligamentous ventral the derived shape of the first epibranchial. limb. Empetrichthys Gilbert is distinguished The genera "Aplocheilichthys," Procato- from Crenichthys Hubbs as well as other cy- pus, Pantanodon, and Hypsopanchax to- prinodontiform genera by its enlarged infe- gether are defined by a lack of the interarcual rior pharyngeals as figured by Uyeno and 1981 PARENTI: CYPRINODONTIFORM FISHES 455

FIG. 82. Distributional limits of goodeids, Empetrichthys and Crenichthys. (After Miller and Fitz- simons, 1971; Miller, 1948.)

Miller (1962). It comprises two Recent ing character of the cyprinodontine genus species, Empetrichthys merriami (which is Cualac is the dense conical inferior pharyn- reportedly extinct) and E. latos which is di- geal teeth, fig. 84; however, these are much vided into three subspecies. The genus smaller than those in Empetrichthys.) name, meaning "fish with rocks within" re- Crenichthys is readily distinguished by its fers to the enlarged molariform pharyngeal unique arrangement of outer jaw teeth (fig. teeth found in merriami, the type species. 33C). There is one large outer row of bicus- Uyeno and Miller (1962) illustrated these ele- pid teeth and several scattered inner rows of ments for both species indicating that the unicuspid teeth. Replacement teeth are teeth were only slightly enlarged in latos. prominent on the outer surface of the pre- However, the large conical teeth of both are maxilla and dentary. A similar arrangement unique among cyprinodontiforms. (A defin- is found in several goodeid genera (e.g., Skif- 456 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

GOODEIDS EMPETRICHTHYS CRENICHTHYS

FIG. 84. Diagrammatic representation of infe- FIG. 83. Cladogram of relationships of good- rior pharyngeals of Cualac tessellatus. Cartilage eids, Empetrichthys, and Crenichthys. Node A: first two to seven middle anal radials absent or is stippled. fused to proximal radials; distal arm of the pre- maxilla straight; dorsal process of the maxillaries greatly reduced; articular reduced. Node B: vi- set off from the rest of the fin by a notch; viparous; first five to seven anal rays of males first anal ray rudimentary in adult males; a unbranched, shortened and set off from the rest muscular urogenital organ or pseudophallus of the anal fin by a notch; first anal ray rudimen- present in males; trophotaeniae of either a tary in males; muscular urogenital organ in males; trophotaeniae on embryos; ovary with ovigerous rosette or ribbon-like configuration in all but tissue partly or completely eliminated from ovar- one species, and ovaries united medially ian walls. Node C: No pelvic fins or fin supports; with ovigerous tissue partly to completely Y-shaped first epibranchial. Node D: enlarged eliminated from ovarian walls. outer teeth; enlarged pharyngeal teeth; fleshy bas- CLADISTIC SUMMARY OF Empetrichthys, es of dorsal and anal fins. Node E: Bicuspid outer Crenichthys, AND THE GOODEIDAE: This teeth; high number of gill rakers on first arch. group is defined by four derived characters: first two to seven middle anal radials absent or fused to proximal radials; distal arm of the fia and ); however, a close re- premaxilla straight; dorsal processes of the lationship of Crenichthys to the goodeids is maxillaries greatly reduced; and, a reduced not postulated on the basis of such a char- articular. acter. A proposed close relationship of Cren- Empetrichthys and Crenichthys both lack ichthys to some more derived group of good- pelvic fins and fin supports, and also share eids would render the goodeids as now a derived form of the first epibranchial. constituted polyphyletic and such a conclu- Empetrichthys is defined by enlarged outer sion is not supported. In addition, the genus and pharyngeal teeth, and fleshy bases of the is distinguished from Empetrichthys by its dorsal and anal fins. high number of gill rakers on the first arch Crenichthys is defined by bicuspid outer which is 20 or more as opposed to 12-13 in teeth, and an increase in the number of gill Empetrichthys. The latter genus is further rakers on the first arch. distinguished by having a fleshy base of the The viviparous goodeids are defined by the anal fin, whereas the base is fully scaled in following reproductive characters: first five Crenichthys. to seven anal rays of male unbranched, The goodeids are defined as a monophy- shortened and set off from the rest of the letic group by the following derived repro- anal fin by a notch; first anal fin ray rudi- ductive characters: the first five to seven anal mentary in males; muscular urogenital or- rays of the male unbranched, shortened and gan, termed a pseudophallus, in males; em- 1981 PARENTI: CYPRINODONTIFORM FISHES 457 bryos with intestinal outpocketings, termed trophotaeniae; and, ovaries with ovigerous tissue partly to completely eliminated from ovarian walls. GROUP I: The genera in this group share three derived characters: (1) the dorsal pro- cess of the maxillaries are expanded medi- ally, nearly meet in the midline, and have a distinct groove; (2) the lateral arm of the maxilla is robust (fig. 42); and (3) the tooth- plate of the fourth pharyngobranchial is greatly reduced. Cubanichthys AND Chriopeoides: These two monotypic genera are hypothesized to be sister taxa since they share a supraocci- pital crest, an elongate dorsal process of the autopalatine, a supraorbital sensory pore pattern consisting of a large third pore, and a posttemporal lacking an ossified ventral limb. Thus, Chriopeoides Fowler, the youn- ger name is treated as a junior synonym of Cubanichthys Hubbs. FIG. 85. Distributional limits of Orestias. GROUP J: Members of this group share the following derived characters: uniserial outer jaw teeth; second pharyngobranchial offset present and with a distinct trough for the sen- to the third; parietals absent; Meckel's car- sory canal. tilage expanded posteriorly; and the trans- In A. mento, as well as in Orestias, Kos- verse processes of the vertebrae reduced and swigichthys, and Anatolichthys, the inter- cup-shaped. hyal is cartilaginous and the urohyal is Orestias AND THE ANATOLIAN CYPRINO- embedded as the lower jaw is set at an angle DONTINES: The Andean Orestias (fig. 85) almost perpendicular to the body axis. In ad- shares with the cyprinodontines of the Old dition, there is the distinctive neuromast pat- World (fig. 86) a medial extension of the den- tern on the dorsal surface of the head shared tary which is enlarged in Orestias to form a by the species of Aphanius related to mento bony shield in the upturned lower jaw. and Orestias. The pattern is only weakly ex- The mesethmoid is cartilaginous in all hibited by the genus Anatolichthys and not specimens of Anatolian cyprinodontines ex- at all in Kosswigichthys which lacks scales amined in all four nominal genera: Aphanius altogether. Nardo, Aphaniops Hoedeman, Kosswig- Species of Orestias and those of Anato- ichthys Sozer, and Anatolichthys Kosswig lichthys and Kosswigichthys are hypothe- and Sozer. It is ossified in all specimens of sized to form a monophyletic group within Orestias examined. However, the uniquely this assemblage (fig. 87). They have in ad- derived status of the cartilaginous meseth- dition to the derived characters they share moid is refuted in the analysis of other char- with A. mento, the following: the ventral acters present in these five nominal genera. limb of the posttemporal is represented by Aphanius is nonmonophyletic and is only an unossified ligament; there is a reduction defined by a set of primitive characters: os- or total absence of scales (Ermin, 1946); and, sified interhyal; body fully scaled; posttem- the number of vertebrae is increased to 28 or poral forked; urohyal not embedded, jaw not more, as opposed to the general number of upturned (as discussed for Orestias and 24 found among the species of Aphanius, Aphanius mento); and the dermosphenotic Aphaniops, and other cyprinodontines, as 458 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 86. Distributional limits of Anatolian cyprinodontines.

well as in Cubanichthys. The type species of to the body axis; and a derived neuromast Aphanius, A. fasciatus, is considered a cephalic sensory pattern. primitive member of the Anatolian cyprino- Kosswigichthys, Anatolichthys, and Or- dontines. I propose to reference derived estias have a posttemporal with a ligamen- species as "Aphanius." tous lower limb, a reduction or absence of Orestias has the following derived char- scales, and an increase in the number of ver- acters: no vomer, no pelvic fins or fin sup- tebrae to 28 or more. ports, and no first postcleithrum. Anatolichthys is treated as a junior syn- The absence of a dermosphenotic supports onym of Kosswigichthys which is defined by the monophyly of, and hence, the synonymy the absence of the dermosphenotic. of Anatolichthys in Kosswigichthys. Orestias has no vomer, no pelvic fins or CLADISTIC SUMMARY OF Orestias AND fin supports, and no first postcleithrum. THE ANATOLIAN CYPRINODONTINES: This NEW WORLD CYPRINODONTINES: New group is defined by an expanded medial pro- World cyprinodontines (of the nominal gen- cess of the dentary. era Cyprinodon Lacepede, Megupsilon Mil- Aphanius is polyphyletic; within a phylog- ler and Walters, Cualac Miller, Florid- eny of the group, its derived members are ichthys Hubbs, Jordanella Goode and Bean, referred to the genus "Aphanius." and Garmanella Hubbs) are hypothesized to "Aphanius" and the nominal genera Kos- form a monophyletic group of genera based swigichthys, Anatolichthys, and Orestias on their sharing a derived form of the attach- share a cartilaginous interhyal; an embedded ment of the first vertebra to the skull. This urohyal and lower jaw at nearly a right angle is characterized by the loss of exoccipital 1981 PARENTI: CYPRINODONTIFORM FISHES 459

APHAN1US "APHANIUS" KOSSWiGICHTHYS OREST1AS d 9*e

FIG. 87. Cladogram of relationships of Orestias and the Anatolian cyprinodontines. Node A: medial extension of dentary. Node B: interhyal cartilaginous; urohyal embedded and lower jaw nearly at right angle to body axis; neuromast sensory pore pattern. Node C: posttemporal lacks ossified lower limb; reduction or absence of scales; increase in number of vertebrae to 28 or more. Node D: loss of the dermosphenotic. Node E: no vomer; no pelvic fins or fin supports; no first postcleithrum. condyles, the supraoccipital forming the dor- The position of Megupsilon in a phylogeny sal wall of the foramen magnum rather than of all New World cyprinodontines is unre- being excluded from it, and neurapophyses solved. Megupsilon lacks pelvic fins and fin of the first vertebra angled anteriorly and supports, as do several species of Cyprino- firmly applied to the skull. In addition, the don. Miller (1956) stated that Cyprinodon pharyngobranchial teeth are arranged in dis- alone possessed an enlarged humeral scale; crete rows. however, the variability of the size of such The taxonomy of the New World cyprin- a scale in Cyprinodon and other genera pre- odontines has been dominated by the naming cludes the use of this character for defining of monotypic taxa which in some way depart Cyprinodon as a monophyletic genus. Cy- from the general form of Cyprinodon. This prinodon does have an enlarged extension of genus currently comprises 36 species and the scapula (fig. 8B). The scapula of Meg- subspecies (Lazara, 1979), whereas all the upsilon is slightly enlarged, whereas that of rest are monotypic. Jordanella, Floridichthys, and Cualac is Interrelationships of New World cyprino- even less so. dontines are summarized in the cladogram of Some Cyprinodon and Jordanella floridae figure 89; their distribution in figure 88. have a thickened first dorsal ray that resem- Garmanella is treated as a junior synonym bles a spine. The absence of a spine in Jor- of Jordanella. Both share elongate dorsal danella {Garmanella) pulchra suggests that fins of 15 rays or more. They also both pos- this character is derived for some larger sess a discrete blotch at midbody and a black group. Thus Megupsilon is parsimoniously suborbital bar. assessed as forming an unresolved trichoto- Megupsilon aporus was originally distin- my with Cyprinodon and Jordanella (fig. guished from all other species of the group 89). It shares the absence of pelvics with by its possession of an enlarged Y-chromo- Cyprinodon and the midlateral blotch with some in the male, a sexually dimorphic chro- Jordanella. mosome number, the absence of pores in the Floridichthys and Cualac have an en- cephalic sensory pore system, blackened larged element in the position of the first scales on the sides of the body of breeding pharyngobranchial. Floridichthys has an ac- males, also lack a terminal band on the cau- tual toothplate with a patch of teeth, whereas dal fin, and not present in all Cyprinodon in Cualac the element is cartilaginous and (Miller and Walters, 1972). Several breeding devoid of teeth. characteristics reportedly differ from those Floridichthys is unique among cyprino- of Cyprinodon. dontines in having a pectoral fin of 18 to 460 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 88. Distributional limits of New World cyprinodontines.

20 rays, whereas Cualac has the typical low- first vertebra open and applied to the skull; er range of 11 to 13. supraoccipital included in the formation of Cualac is defined by expanded inferior the foramen magnum; and, pharyngobran- pharyngeals (fig. 84) and their close-set, vil- chial teeth arranged in discrete rows. liform teeth, as well as an increase in the Garmanella is treated as a junior synonym number of gill rakers on the first arch to 17. of Jordanella which is defined by a dorsal It could be argued that all New World cy- fin with more than 15 rays, and a suborbital prinodontines should be placed in one genus. bar. However, since monophyletic groups of Megupsilon is defined by an enlarged species can be defined, it is suggested that Y-chromosome in males; sexually dimorphic they be recognized until a phylogenetic anal- chromosome number; and no cephalic sen- ysis of all species is presented. sory pores. CLADISTIC SUMMARY OF NEW WORLD Cyprinodon is tentatively defined by an CYPRINODONTINES: New World cyprinodon- enlarged scapular process; yet, it is stressed tines share the following derived characters: that this genus is probably polyphyletic. no exoccipital condyles; neural arches of the Megupsilon, Jordanella, and Cyprinodon 1981 PARENTI: CYPRINODONTIFORM FISHES 461

CYPRINODON MEGUPSILON JORDANtLLA F 1_C R I D IC H T H YS CUALAC

FIG. 89. Cladogram of relationships of New World cyprinodontines. Node A: no exoccipital con- dyles; neural arches of first vertebra angled anteriorly and applied to skull; supraoccipital included in formation of foramen magnum; pharyngobranchial teeth arranged in discrete rows. Node B: first dorsal spine often present. Node C: Dark, midlateral blotch. Node D: Pelvic fins and fin supports often absent. Node E: greatly enlarged scapular process. Node F: enlarged Y-chromosome in male, sexually dimor- phic chromosome number; no cephalic sensory pores. Node G: elongate dorsal fin of more than 15 rays; suborbital bar. Node H: first pharyngobranchial present. Node I: first pharyngobranchial toothplate present with a patch of teeth. Node J: increased number of gill rakers on the first arch; inferior pha- ryngeal teeth packed closely together.

form an unresolved trichotomy. Megupsilon Cualac and Floridichthys have a first pha- shares the absence of pelvic fins and fin sup- ryngobranchial element. Floridichthys has ports with some species of Cyprinodon, the inferred derived state, an ossified tooth- whereas it shares the presence of a midlat- plate with a patch of teeth. The genus is also eral blotch with Jordanella. Some Cyprino- defined by a high number of pectoral fin rays don and Jordanella floridae have a thick- (18-20). Cualac is defined by expanded in- ened first dorsal fin ray resembling a true ferior pharyngeals with close-set villiform spine. teeth, and a high number of gill rakers on the first arch.

CLASSIFICATION

The phylogenetic analysis just presented At the same time some stability in nomen- reveals that the present classification of cy- clature is desirable for reasons which are ap- prinodontiform fishes is not based on defin- parent. However, stability ceases to be de- able monophyletic groups. The purpose of a sirable when a taxon as currently constituted, classification in a cladistic system is to sum- such as the Cyprinodontidae, is unnatural. marize the hierarchy of relationships of the Retaining the name to reference all ovipa- defined monophyletic groups of taxa (e.g., rous killifishes (minus Tomeurus) would be Nelson, 1973). It is also desirable but not to ignore the overwhelming evidence against necessary (see Farris, 1976) to give sister the monophyletic nature of the group. groups the same rank so that a cladogram or The system of interrelationships proposed scheme of interrelationships may be inferred in this study is far too complicated to be sum- easily from a written classification. marized easily using the existing system of 462 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 five family names grouped together in one Genus Nothobranchius Pe- ters, 1868 superfamily. Therefore, I propose that the 1 rank of the superfamily be raised to an order, Family Rivulidae Myers, 1925 which shall be known as the Cyprinodonti- Genus Rivulus Poey, 1860 Genus "Rivulus" (see pp. formes Berg, a well-known and still widely 483-484) used term for this group of fishes. In order Genus Trigonectes Myers, that this classification conform to that for all 1925 members of the series Atherinomorpha, Ro- Genus Pterolebias Gar man, sen and Parenti (MS) have written a new clas- 1895 sification of atherinomorph fishes. Genus Rachovia Myers, 1927 Thus, the following classification of the Genus Neofundulus Myers, cyprinodontiform fishes is proposed: 1924b Genus "Neofundulus" (see pp. 489-490) Order Cyprinodontiformes Berg, 1940 Genus Austrofundulus Suborder Aplocheiloidei, new usage Myers, 1932 Family Aplocheilidae Bleeker, 1860 Genus Cynolebias Steindach- Genus Aplocheilus Mc- ner, 1876 Clelland, 1839 Suborder Cyprinodontoidei Genus Pachypanchax Myers, Section 1 1933b Family Profundulidae Hoedeman Genus Epiplatys Gill, 1862 and Bronner, 1951 Subgenus Lycocyprinus Genus Profundulus Hubbs, Peters, 1868 1924 Subgenus Parepiplatys Section 2 Clausen, 1967 Division 1 Subgenus Pseudepiplatys Family Fundulidae Jordan and Gil- Clausen, 1967 bert, 1882 Subgenus Aphyoplatys Genus Plancterus Gar man, Clausen, 1967 1895 Genus Adamas Huber, 1979 Genus Fundulus Lacepede, Genus Aphyosemion Myers, 1803 1924b Genus Adinia Girard, 1859 Subgenus Archiaphyose- Genus Lucania Girard, 1859 mion Radda, 1977 Genus Leptolucania Myers, Subgenus Chromaphyose- 1924b mion Radda, 1971 Division 2 Subgenus Diapteron Hub- Sept 1 er and Seegers, 1977 Family Valenciidae, new family Subgenus Kathetys Huber, Genus Valencia Myers, 1928b 1977 Sept 2 Subgenus Mesoaphyose- Superfamily Poecilioidea, new usage mion Radda, 1977 Family Anablepidae Garman, 1895 Genus Fundulopanchax Subfamily Anablepinae Garman, Myers, 1924b 1895 Subgenus Callopanchax Myers, 1933c Subgenus Paraphyose- 1 Within the Lepidoptera, the subfamily Rivulinae mion Kottelat, 1976 was named by McDunnough (1938); the type genus is Subgenus Paludopanchax the North American moth Rivula Guenee [in Radda, 1977 Duponchel, 1835]. Following the International Code of Subgenus Gularopanchax Zoological Nomenclature, article 55a, this is a case of Radda, 1977 homonymy of family group names. The family group Subgenus Raddaella Hu- name is older in the Cyprinodontiformes and should be ber, 1977 dropped from use in the Lepidoptera. 1981 PARENTI: CYPRINODONTIFORM FISHES 463

Genus Anableps (Gronow) Subfamily Cyprinodontinae Gill, Scopoli, 1777 1865 Genus Jenynsia Giinther, Tribe Orestiini Bleeker, 1860 1866 Genus Orestias Valenci- Subfamily Oxyzygonectinae, new ennes, 1839 subfamily Genus Kosswigichthys Soz- Genus Oxyzygonectes Fowl- er, 1942 er, 1916 Genus Aphanius Nardo, 1827 Family Poeciliidae Carman, 1895 Genus "Aphanius" (see pp. Subfamily Poeciliinae Garman, 522-524) 1895 Tribe Cyprinodontini Gill, 1865 Subfamily Fluviphylacinae Rob- Genus Cyprinodon Lace- erts, 1970 pede, 1803 Genus Fluviphylax Whitley, Genus Megupsilon Miller 1965 and Walters, 1972 Subfamily Aplocheilichthyinae Genus Jordanella Goode and Myers, 1928a Bean, 1879 Genus Aplocheilichthys Genus Floridichthys Hubbs, Bleeker, 1863 1926 Subgenus Micropanchax Genus Cualac Miller, 1956 Myers, 1924a Subgenus Lacustricola Myers, 1924a Familiar group names have been retained Subgenus Congopanchax only if they can be used in the same manner Poll, 1971 as in previous classifications, or, if the mem- Subgenus Poropanchax bership of such categories could be slightly Clausen, 1967 expanded or contracted to include close rel- Genus Lamprichthys Regan, atives or eliminate unrelated taxa, respec- 1911 tively. An example in which the practice has Genus "Aplocheilichthys" (see pp. 510-511) been applied is in the family Poeciliidae. As Genus Procatopus Boulen- previously denned, it included only those ger, 1904b members which possessed a gonopodium in Genus Hypsopanchax Myers, males formed principally from anal rays 3, 4, 1924a and 5. This group is retained here as the Genus Pantanodon Myers, subfamily Poeciliinae, whereas the family 1955 Poeciliidae has been expanded to include the Genus Cynopanchax Ahl, procatopines and Pantanodon (subfamily 1928 Aplocheilichthyinae), and Fluviphylax Genus Plataplochilus Ahl, (subfamily Fluviphylacinae). The relation- 1928 Superfamliy Cyprinodontoidea, new ships of these three groups are expressed in usage the cladogram as an unresolved trichotomy, Family Goodeidae Jordan, 1923 therefore, each of the three groups is given Subfamily Empetrichthyinae Jor- equal rank. Similarly, the family Goodeidae dan, Evermann and Clark, has traditionally been limited to the vivipa- 1930 rous forms of the Mexican Plateau. The rank Genus Empetrichthys Gil- of this group of genera has been reduced to bert, 1893 a subfamily, the Goodeinae, whereas the two Genus Crenichthys Hubbs, genera proposed as its sister group, Empe- 1932 Subfamily Goodeinae Jordan, 1923 trichthys and Crenichthys, are placed in the Family Cyprinodontidae Gill, 1865 subfamily Empetrichthyinae. The family Subfamily Cubanichthyinae, new name Goodeidae is therefore used to encom- subfamily pass these two subfamilies. Genus Cubanichthys Hubbs, Three informal categories (division, sec- 1926 tion, and sept) have been employed both to 464 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 provide stability in the nomenclatorial some fossil specimens currently placed in the scheme and also to minimize the number of genus Fundulus may in fact be closely relat- empty categories. Thus, the genus Profun- ed to Profundulus. If so, another taxon (fos- dulus, represented as the primitive sister sil or Recent) may be added to the system as group of all other cyprinodontoids (classified a sister group of Profundulus without dis- here as the suborder Cyprinodontoidei) is rupting the existing higher categories. This placed in its own family, for the sake of tra- type of change to a classification may be dition alone, the Profundulidae. However, termed a "nondisruptive" change. In con- rather than placing the family Profundulidae trast, a "disruptive" change would occur in in its own superfamily, and similarly all other a classification if, for example, a family was members of the suborder in another super- found to be nonmonophyletic as in the case family, I have elected to use the informal cat- of the family Cyprinodontidae as the group egory of section for this purpose. Logically, name is used in the beginning of this study. there is no difference between the two ap- A "nondisruptive" change, by definition, proaches. However, the use of an informal may always be incorporated into a system category at this point and between the next with the use of informal categories if it is not two divisions of the suborder, leaves the cat- possible to work it into the existing nomen- egory superfamily to reference those groups clatorial system. A revisor of the subfamily that contain more than one family. Thus, Goodeinae (containing approximately 36 there is no alteration of the definitions of species in 16 genera) may wish to present, in higher categories except in the cases where the form of a cladistic classification, the in- families contain just one genus, but the rea- terrelationships of the species with a redefi- son for this has already been stated. nition of the genera. Below the subfamily The informal categories have not been level there are eight traditional hierarchical named since their names would be trivial ad- categories into which species may be ditions to the classification. For example, grouped: species group, superspecies, sub- names of Section 2, Division 2 and Sept 2 genus, genus, supergenus, subtribe, tribe, would all have Cyprinodon as a root and and supertribe. A completely dichotomous some arbitrary ending. The names of Section sytem of interrelationships of 512 species 1, Division 1, and Sept 1 would all be mod- could be accommodated within such a sys- ifications of the family names already includ- tem. If there were not sufficient categories, ed in them. informal categories could be applied. These The generic groups within most families may be named or numbered at the discretion are those in current use. For the most part, of the revisor. In such cases, I propose that their interrelationships and composition numbered informal categories be used in were not dealt with formally here. In the conjunction with traditional names. Most classification, the genera are simply listed; taxonomists work at the level of revising my expectation is that future work will suc- families or their subgroups. A revision of a ceed in classifying these groups in a hier- superfamily is not frequently done in con- archy of suprageneric categories and that a junction with the revision of an order, as is revisor of the Rivulidae, for example, could the present case, so that all ranks may be very well introduce such categories into the adjusted accordingly. system without altering the existing classifi- Ideally, we should have a system of no- cation. menclature adaptable to all changes. A sys- The reason for this expectation is that this tem of prefixes and suffixes could be agreed classification system is designed to be both upon (as in Farris, 1976) to accomplish just flexible and minimally disruptive to currently this task; however, group names would named monophyletic groups. As constituted, quickly become unwieldy and therefore ig- Section 1 of the suborder Cyprinodontoidei nored in favor of the existing names. A group contains just one genus, Profundulus. numbering system may be more usable. Uyeno and Miller (1962) have remarked that However, introduction at this time of a nu- 1981 PARENTI: CYPRINODONTIFORM FISHES 465 merical classification, in stark contrast to all If so, the traditional use of the term Cy- other existing classifications of fishes, would prinodontidae would have been retained, and not have the desired effect. That is, it would the genus Jomeurus moved from the Poeci- not prompt the adoption of the new classifi- liidae to that oviparous family. The philoso- cation. phy adopted in this study, however, is not A "disruptive" change requires, by defi- one of subjective weighting of characters for nition, a reclassification unless a group of expression in a classification, but rather of taxa can be conveniently moved from one the incorporation of all available evidence higher taxon to another. The taxonomy of into a scheme of interrelationships which re- many groups is in such a state; however, a flect the genealogy of the group under revi- more likely case is one such as that pre- sion. It is concluded that the scheme pre- sented here. A "disruptive" change in the sented here, in being rigorously cladistic, is classification was judged to be necessary. a better estimate of the one true phylogeny Such decisions are always subjective. For of the cyprinodontiform fishes than others, example, mode of reproduction could have past or current (Garman, 1895; Regan, 1911; been judged as the single most important Jordan, 1923; Hubbs, 1924; Myers, 1931, character to be expressed in a classification. 1955; and Sethi, 1960).

KEY TO GENERA AND SUPRAGENERIC CATEGORIES The following key is provided to aid in maxilla present; basihyal expanded ante- identifying the generic and suprageneric cat- riorly (fig. 10A); anterior arm of autopal- egories of cyprinodontoid fishes. It is based atine straight (fig. 29); exoccipital and bas- on the cladogram of figure 9 and the clado- ioccipital condyles not reduced (fig. 26); grams of monophyletic groups; however, pelvic fin supports set close together with medial processes reduced (fig. 10A) since the key is dichotomous, the more de- Suborder Aplocheiloidei 2A rived states of transition series could not be 2A Supracleithrum fused to posttemporal; first represented. In addition, categories referred postcleithrum present; opercular and to in the classification as subgenera are not branchiostegal membrane not covered represented since, for the most part, these with scales; head-scales not arranged in are either paraphyletic assemblages or circular pattern; preopercle without ven- groups which have not been studied in detail. tral expansion (fig. 30); dermosphenotic The key is presented in the hope that it will large, with distinct canal (fig. 17B); pre- be useful in recognizing the major differ- maxillary ascending processes tapered ences among groups many of which have posteriorly (fig. 4A, B, C); no flange on second pharyngobranchial at point of ar- long been confused. ticulation of the interarcual cartilage (fig. 1A Three basibranchials; metapterygoid pres- 24) Family Aplocheilidae 3A ent; a dorsal ray on each of the first two 3A Epipleural ribs not bifid; premaxillary as- dorsal radials; dorsal hypohyal present; cending processes expanded posteriorly alveolar arm of premaxilla not strongly in- (fig. 4A, B); expanded coronoid process dented posteriorly (fig. 3A); autopalatine on dentary (fig. 27); no uncinate process process small, not reaching quadrate (fig. on fourth epibranchial for articulation of 29); orbital rim attached on lower half of third epibranchial (fig. 24A) 4A orbit; lacrimal narrow and twisted (fig. 12A); dentary not expanded medially (fig. 4A Posttemporal straight, lower limb not pres- 31C); rostral cartilage large and disc- ent; ocellus at anterior base of dorsal fin shaped (fig. 4); ligament from the interior in at least females and juveniles; orbital arms of the maxillaries to the rostral car- rim indented under frontals; teeth on the tilage present; ethmomaxillary ligament second and third hypobranchials .... 5A present; meniscus between premaxillary 5A Posterior edges of scales in males stand ascending processes and interior arms of away from body; hypural plates fused into 466 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

a hypural fan in adults; premaxillary as- rays 6 Rivulus cending processes do not meet in the mid- (Caribbean, North, Middle and line (fig. 4B) Pachypanchax South America) (East Africa, Madagascar, and 8B Interhyal cartilaginous; dorsal and anal fins the Seychelles) elongate in males; pelvic fin rays 7 .. 9A 5B Scales of males close to body; hypural plates 9A Pectoral rays not reaching base of pel vies; separate in adults, upper plate often divid- rostral cartilage not elongate ed in two (fig. 2D); premaxillary ascending ' 'Rivulus'' processes meet and overlap in the midline (Caribbean, Middle and South America) (fig. 4A) Aplocheilus 9B Pectoral rays extended, reaching to or be- (Indo-Malaysian region) yond the base of the pelvic fins; rostral 4B Posttemporal forked; no ocellus at anterior cartilage elongate (fig. 4D) 10A base of dorsal fin; orbital rim not indented; 10A No vertical bar through eyes; origin of dorsal no teeth on the second and third hypo- posterior to origin of anal; anal rays of fe- branchials Epiplatys males not thickened 11A (West Central Africa) 11A Mouth cleft oblique, anterior ramus of pre- 3B First five or six epipleural ribs bifid; pre- maxilla reduced (fig. 21); snout pointed; maxillary ascending processes tapered interarcual cartilage present posteriorly, but not expanded (fig. 4C); Trigonectes coronoid process on dentary not enlarged (, Paraguay, ) (fig. 31C); extension present on fourth 11B Mouth cleft not oblique, anterior ramus of epibranchial for articulation with third premaxilla not reduced (fig. 3A); snout epibranchial (fig. 24B) 6A not pointed; interarcual cartilage absent 6A Dorsal origin posterior to anal origin; dorsal Pterolebias fin rays less than 14; swimbladder extends (Brazil, ) past the first two or three hemal arches 10B Vertical bar through eyes, often reaching top Aphyosemion of head; thickened anal rays in (West and Central Africa) females 12A 6B Dorsal origin opposite that of anal, or more 12A Dorsal fin not elongate, rays generally less anterior; dorsal fin rays 14 or more; swim- than 14 13A bladder does not extend past the hemal 13A Less than 32 scales in a lateral series arches 7A Rachovia 7A Interarcual cartilage attaches to bony flange (Coastal llanos of Colombia and on second pharyngobranchial (fig. 24A); Venezuela) predpercular canal represented by 13B More than 34 scales in a lateral series pores Fundulopanchax Neofundulus (Central Africa) (Paraguay) 7B Interarcual cartilage attaches directly to car- tilage of second pharyngobranchial (fig. 12B Dorsal fin elongate, rays greater than or 24B); preopercular canal represented by equal to 14 14A open groove Nothobranchius 14A First proximal anal radial present; teeth (West, Central and East Africa) on fourth ceratobranchial not reduced; 2B Supracleithrum not fused to posttemporal; neural spine on first vertebra not enlarged first postcleithrum absent; opercular and "Neofundulus" branchiostegal membrane united and cov- (Paraguay) ered with scales; head scales small, in se- 14B First proximal anal radial absent; teeth on ries around the central "A" scale (fig. the fourth ceratobranchial reduced; en- 13E); preopercle with ventral expansion larged neural spine on the first vertebra and obsolescent sensory pore canal (fig. 15A 13D); dermosphenotic small, often with- 15A Heavily pigmented anal papilla; caudal out distinct sensory canal (fig. 17C); bony fin finely scaled for more than one-third its flange present on second pharyngobran- length; preopercular canal open chial at point of articulation of interarcual Austrofundulus cartilage (fig. 6A) .. Family Rivulidae 8A (Coastal llanos, Colombia, 8A Interhyal ossified; all fins rounded; pelvic fin and Venezuela) 1981 PARENTI: CYPRINODONTIFORM FISHES 467

15B Anal papilla bare or only lightly pigmented; articular surface; epipleural ribs meet dis- caudal fin scaled only to its base; pre- tal tips of parapophyses of the abdominal opercular canal closed Cynolebias vertebrae 20A (Venezuela, Brazil, Argentina) 20A Epiotic processes present; supraoccipital ca- IB Two basibranchials; metapterygoid absent; nal system present between pores 1 and one dorsal ray articulating with the first 7 Lucania two dorsal radial s; dorsal hypohyal ab- (East Coast of North America and sent; alveolar arm of premaxilla strongly Cuatro Cienegas Basin) indented posteriorly (fig. 3B); autopalatine 20B No epiotic processes; supraorbital canal sys- process large, reaching quadrate (fig. 30); tem present between pores 6 and 7 21A orbital rim free; lacrimal flat and wide (fig. 21A Body quadrangular; branchiostegal rays 5; 12C); dentary expanded medially (fig. 33); first postcleithrum present; first pleural rib rostral cartilage reduced; no ligament from on parapophyses of first vertebra; no cau- the interior arms of the maxillaries to the dal or midbody ocellus Adinia rostral cartilage; no ethmomaxillary liga- (Florida to ) ments; no meniscus between maxilla and 21B Body elongate, not trapezoidal; branchioste- premaxilla; basihyal narrow, not expand- gal rays 3; no first postcleithrum; first ed anteriorly; anterior arm of autopalatine pleural rib on parapophysis of the second angled sharply (fig. 30); exoccipital and vertebra; an ocellus on the caudal pedun- basioccipital condyles reduced; pelvic fin cle and one at midbody ... Leptolucania bases not set close together and medial (Florida and Georgia) processes not reduced (fig. 10B, C) 17B Interior arms of maxillaries attenuate, di- Suborder Cyprinodontoidei 16A rected medially, never with pronounced 16A Premaxillary ascending processes flat and hooks; dorsal process directed over the broad; inner arms of maxillaries united premaxillary ascending processes, or pro- with large and rectangular rostral cartilage cess absent; maxilla straight, not twisted (fig. 5B); lateral ethmoids with anterior (fig. 3!% 22A flanges (fig. 57A); autopterotic fossa en- 22A Ascending processes of premaxillaries long larged (fig. 57A).. Family Profundulidae and thin, not shortened; dorsal processes Profundulus of maxillaries elongate (fig. 50) (Highlands of western Middle America) Family Valenciidae 16B Premaxillary ascending processes narrow or Valencia absent; rostral cartilage small and disc- (Spain, Italy and Corfu) shaped or absent; inner arms of maxillar- 22B Ascending processes of premaxillaries short- ies not in direct contact with rostral car- ened; dorsal processes of the maxillaries tilage; lateral ethmoids without anterior rounded when present 23A flanges (fig. 57B); autopterotic fossa not 23A Distal arm of maxilla expanded; anterior arm enlarged (fig. 57B) 17A of parasphenoid spatulate; lateral eth- 17A Interior arms of maxillaries directed ante- moids not expanded, dorsal processes lie riorly, often with pronounced hooks (fig. under frontals; premaxillary ascending 34), no dorsal process directed over the processes with distinct lateral indentation premaxillary ascending processes; maxilla (fig. 35); autopterotic fossa not reduced twisted, not straight (fig. 5C) (fig. 16); inclinators of the anal fin not en- Family Fundulidae 18A larged (fig. 72) 24A 18A Posttemporal forked; intestine convoluted 24A Dentary not enlarged; no pouch created by Plancterus scales of female around anus and first few (Central North America) anal rays; no pectoral ribs on hemal 18B Posttemporal straight; intestine straight spines; pectoral girdle set low on the 19A sides, radials situated ventrally or poste- 19A Second pharyngobranchial with expanded riorly (fig. 8); pelvic fins not set forward; articular surface; epipleural ribs meet pec- hyoid bar with ventral extension of ante- toral ribs rather than the parapophyses of rior ceratohyal; enlarged epiotic and su- the abdominal vertebrae Fundulus praoccipital processes; outer teeth with (North and Middle America) distinct lateral cusps in embryos or 19B Second pharyngobranchial without expanded juveniles Family Anablepidae 25A 468 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

25A Tubular gonopodium associated with sperm rays 3, 4, and 5 associated with expanded duct in males formed from anal rays anal radials and anteriorly porjecting he- crowded together and twisted around each mal arches (fig. 65); males smaller than fe- other; inner rows of jaw teeth unicuspi- males; parietals present or absent date or tricuspidate, in two uneven rows; Subfamily Poeciliinae adult males with gonopodium offset to left (North, Middle and South America) or right of midline; females with no pouch 27B No gonopodium, anal fin normal; males larg- around urogenital opening or first few anal er than females; parietals absent ... 28A rays; females with one or a few scales on 28A Mesethmoid ossified; body fusiform; anal fin left or right side of urogenital opening rays 7 or 8; dorsal fin rays 5 or 6; eye large Subfamily Anablepinae 26A in head, preorbital space narrow 26A Eyes normal; vertebrae 31; outer row of jaw Subfamily Fluviphylacinae teeth tricuspid; premaxillary ascending Fluviphylax processes present; frontals flat, not ex- (Amazon Basin) panded; tubular sperm duct not covered 28B Mesethmoid cartilaginous; body and pedun- with scales; radials set posteriorly; ante- cle compressed; anal fin rays 11 or more; rior nares not tubular; supraorbital sen- dorsal fin rays 10 or more, eye with small- sory pores represented by pores l-2a, 2b- er preorbital space 4a, and 4b-7; rostral cartilage round; gill Subfamily Aplocheilichthyini 29B rakers on first arch 10-11; gonopodium 29A First postcleithrum present; anal fin rays less formed principally from rays 3, 6, and 7; than 14; swimbladder not extending past sixth middle radial enlarged (fig. 68) the first hemal arch Aplocheilichthys Jenynsia (Central and Eastern Africa) (Southern South America) 29B First postcleithrum absent; anal fin rays Eyes divided horizontally; vertebrae 45 or more than 14; swimbladder extending past more; outer row of jaw teeth unicuspid in first hemal arch 30A adults; premaxillary ascending processes absent in adults, weakly present in em- 30A Interarcual cartilage present; posttemporal bryos; frontals expanded dorsally above straight, without lower limb; vertebrae 41; orbit; tubular sperm duct covered with vomer ossified; cteni on scales; caudal fin scales; radials set ventrally; tubular ante- with upper and lower caudal fin extensions rior nares; supraorbital pore system rep- Lamprichthys resented by pores 1-2, 3, 4a and 6 and 7; (Lake Tanganyika) rostral cartilage dumbbell-shaped; gill rak- 30B Interarcual cartilage absent; posttemporal ers on first arch 21-30; gonopodium forked; vertebrae less than 41; vomer os- formed principally from anal rays 3-6; sified or not; scales not ctenoid; caudal fin sixth middle radial not enlarged (fig. rounded or truncate 31A 67) Anableps 31A Pharyngobranchial teeth tricuspid; three pel- (Central and Southern South America) vic spines in males; enlarged second 25B No gonopodium; inner jaw teeth tricuspi- pharyngobranchial toothplate (fig. 45); date, set in numerous bands (fig. 36); outer pelvic fin rays of males curved and scales covering preopercular canal; fe- elongate; teeth on premaxilla do not ex- males with pouch over urogenital opening tend distally; distal arm of premaxilla and first few anal rays curved; exoccipital condyles absent; no Subfamily Oxyzygonectinae neural spine on first vertebra, neurapo- Oxyzygonectes physes of first arch expanded (Pacific coast of Costa Rica) Pantanodon 24B Dentary enlarged (fig. 38C); pectoral rib on (Dar es Salaam and Madagascar) first hemal spine; pectoral girdle set high 3IB Pharyngobranchial teeth unicuspid; no spines on the sides, radials situated dorsally (fig. present; second pharyngobranchial tooth- 8); pelvic fins set forward, nearly under plate not enlarged; outer pelvic fin rays of pectorals in most; hyoid bar without ven- males normal; exoccipital condyles pres- tral extension of anterior ceratohyal; ent; neural spine on first vertebra .. 32A epiotic and supraoccipital processes not 32A Distal arm of premaxilla curved; teeth do not enlarged; outer teeth without lateral extend distally on the arm cusps Family Poeciliidae 27A "Aplocheilichthys" 27A Gonopodium in males, formed from anal fin (Central Africa) 1981 PARENTI: CYPRINODONTIFORM FISHES 469

32B Distal arm of premaxilla straight; teeth ex- 31C); transverse processes of vertebrae tend distally along the arm 33A not reduced 33A Deep-bodied; first two branchiostegal rays Subfamily Cubanichthyinae of males not free from branchiostegal Cubanichthys membranes Hypsopanchax (Cuba and Jamaica) (East Central Africa) 37B Supraoccipital crest not enlarged; dorsal 33B Not deep-bodied; first two branchiostegal process of autopalatine short; parietals ab- rays of males free from branchiostegal sent; uniserial outer jaw teeth; second membranes (fig. 53) Procatopus pharyngobranchial greatly reduced (fig. (Central Africa) 49); Meckel's cartilage expanded poste- 23B Distal arm of maxilla not expanded; anterior riorly (fig. 43A); transverse processes of arm of parasphenoid thin; lateral ethmoids vertebrae reduced and cup-shaped Tribe expanded medially, dorsal process not un- Orestiini 38A der frontals; attenuate premaxillary as- 38A Medial extension of the dentary (fig. 43A, cending processes; autopterotic fossa re- B); neural spine on first vertebra; supra- duced (fig. 57); inclinators of the anal fin occipital excluded from formation of fo- enlarged (fig. 73) 34A ramen magnum; exoccipital condyles 34A First two to five middle anal radials absent; present; pharyngobranchial teeth random- proximal anal radials of males crowded ly arranged 39A together; dorsal processes of maxilla 39A Urohyal not embedded (fig. 13C); dermo- greatly reduced, no groove; toothplate of sphenotic not reduced Aphanius fourth pharyngobranchial not reduced; (Mediterranean) distal arm of premaxilla straight (fig. 40); 39B Urohyal embedded (fig. 141); dermosphen- articular reduced (fig. 33) otic reduced or absent 40A Family Goodeidae 35 40A Interhyal ossified; body fully scaled; post- 35A Pelvic fins present; first epibranchial not Y- temporal with lower limb ossified; verte- shaped; anal rays 2 to 7 of male shortened; brae 27 or less "Aphanius" first anal ray rudimentary; pseudophallus (Eastern Mediterranean, Iran and Turkey) present Subfamily Goodeinae 40B Interhyal cartilaginous; scales greatly re- (Mesa Central, Mexico) duced or absent; posttemporal straight, 35B No pelvic fins or fin supports; first epibran- without lower limb; vertebrae more than chial Y-shaped (fig. 47B); anal rays of 27 41A males not shortened; first anal ray not ru- 41A Pelvic fins present; vomer present; first post- dimentary; pseudophallus not present ... cleithrum present; dermosphenotic ab- Subfamily Empetrichthyinae 36A sent Kosswigichthys 36A Outer teeth unicuspid; anal and dorsal fin (Fresh water lakes of Turkey) bases fleshy; inferior pharyngeal teeth en- 41B Pelvic fins absent; vomer absent; no first larged; gill rakers on first arch 12- postcleithrum; dermosphenotic greatly 13 Empetrichthys reduced Orestias (Death Valley System) (Lakes of South American continental 36B Outer teeth bicuspid; anal and dorsal fin bas- divide) es fully scaled; inferior pharyngeal teeth 38B No anteriorly directed medial extension of normal; gill rakers on first arch more than dentary; neural arches of first vertebra ap- 20 Crenichthys plied to skull (fig. 62); no neural spine on (Eastern Nevada) first vertebra; supraoccipital bordering the 34B All anal radials present; groove in dorsal dorsal wall of the foramen magnum; ex- processes of maxillae, expanded medially occipital condyles wanting; pharyngo- nearly meeting in midline (fig. 41); tooth- branchial teeth arranged in discrete rows plate of fourth pharyngobranchial re- (fig. 49) Tribe Cyprinodontini. 42A duced; distal arm of premaxilla curved 42A First dorsal spine often present; dorsal ocel- (fig. 42) 37A lus often present in females; first pharyn- 37A Supraoccipital crest enlarged (fig. 58); dorsal gobranchial absent 43A process of autopalatine elongate; parietals 43A Dorsal fin with 14 or fewer rays; no subor- present; biserial outer teeth; second pha- bital bar; prominent scapular process ryngobranchial not reduced; Meekers 44A cartilage not expanded posteriorly (fig. 44A Males without blackened scales on side; 470 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

pores present in cephalic sensory canal 45A First pharyngobranchial toothplate present system; scapular process greatly enlarged with a patch of unicuspid teeth (fig. 50A); (fig. 8B) Cyprinodon pectoral fin rays 18-20; inferior pharyn- (North, Middle and South America) geals not expanded, teeth with slight 44B Males with blackened scales on side; pores shoulders; gill rakers on the first arch 9- absent in cephalic sensory canal system; 10 Floridichthys scapular process not enlarged (Florida and the Yucatan Peninsula Megupsilon south to Honduras) (Nuevo Leon, Mexico) 45B Cartilaginous first pharyngobranchial lacking 43B Dorsal fin with more than 15 rays; suborbital teeth (fig. 50B); pectoral fin rays 11-13; bar present; scapular process not promi- inferior pharyngeals expanded, with nent Jordanella closely set villiform teeth (fig. 84); gill rak- (Florida and the Yucatan Peninsula ers on the first arch 17 Cualac south to Belize) (San Luis Potosi, Mexico) 42B No dorsal spine; no dorsal ocellus; first pha- ryngobranchial present 45A

SYSTEMATIC ACCOUNTS

ORDER CYPRINODONTIFORMES BERG

DIAGNOSIS: Distinguished from all other compressed. Fins soft-rayed, rarely a spi- atherinomorph fishes by the following de- nous first dorsal or first three or four pelvic rived characters: symmetrical caudal fin sup- fin rays. Pectoral and pelvic fin position vari- ported internally by one epural which mir- able, pelvic fins and fin supports present or rors in shape and position an opposing absent. parhypural; premaxilla with a two-part al- Caudal fin rounded or truncate, or with veolar process; first pleura) rib arising on the dorsal and ventral fin extensions in males. parapophysis of the second vertebra rather Caudal skeleton composed of one epural than the third; interarcual cartilage arising similar in shape and position to the opposing from the base of the first epibranchial and parhypural. Hypurals fused into two sub- attaching to the second pharyngobranchial; equal dorsal and ventral segments, or the primitively lowset pectoral girdle with a segments fused to form a hypural fan. In sev- large, scale-shaped first postcleithrum; and eral species, upper hypural plate represented an extended developmental period. by two segments. DEFINITION: Typically small fishes, aver- Infraorbital series represented by a lacri- age adult 80-100 mm. SL; range approxi- mal and dermosphenotic only. Vomer pres- mately 8-300 mm. SL. Distribution pantrop- ent or absent. Scales usually cycloid, some- ical and temperate North and Middle times ctenoid; body generally fully scaled; American, Eurasian, and Indo-Malaysian. scales reduced on venter or totally absent in Inhabitants generally of fresh water, al- some members; head generally fully scaled; though many members enter brackish water. trunk lateral line represented by pitted Markedly sexually dimorphic. Males and scales. Cephalic lateral line system repre- females of the same or unequal sizes. Males sented by canals, exposed neuromasts and often brightly colored, with extensions of pit organs. pelvic, dorsal, and caudal fin rays. Females Jaw teeth uni-, bi- or tri-cuspidate, in one usually drab, rarely with any brightly colored outer and one to several inner rows, or in markings or fin extensions. Males with a gon- single outer row only. opodium in three groups. Premaxillary ascending processes present Body typically fusiform, rarely laterally or absent; when present, flat and broad, nar- 1981 PARENTI: CYPRINODONTIFORM FISHES 471 row and elongate or narrow and short. Ros- to form one large toothplate; pharyngo- tral cartilage present as a large disc, reduced branchial teeth unicuspid or tricuspid, often to a minute disc, or absent. molariform. Interarcual cartilage from base of the first Branchiostegal rays three to seven. epibranchial to cartilage of second pharyn- Scales in lateral series 24-96, vertebrae gobranchial present or rarely absent; when 24-54. Oviparous, ovoviviparous or vivipa- present large and equal or one-half the length rous. Developmental period generally of 10 of the first epibranchial. Pharyngobranchial days or longer. Annual, semiannual, or non- toothplates two, three, and four separate annual reproductive modes. Internal or ex- or, three and four fused; or all three fused ternal fertilization.

SUBORDER APLOCHEILOIDEI DIAGNOSIS: Distinguished from other cy- four subgenera, Pachypanchax Myers, prinodontiforms by the following uniquely Nothobranchius Peters with four subgenera, derived characters: orbital rim attached at Adamas Huber and Aphyosemion Myers least on the lower half of the orbit, pelvic fin with 11 subgenera. bases set close together with medial pro- DISTRIBUTION: (fig. 23) Old World; Africa cesses reduced, a broad anterior end of the south of the Sahara Desert from the lowlands basihyal, a narrow and twisted lacrimal as- of southern Mauritania in western Africa, sociated with a narrow preorbital distance, south through Zaire; the northern limit the and the posterior extension of the vomer dor- Niger River then east to western Sudan, sal to the parasphenoid; distinguished by the south of the Ethiopian Highlands to western following derived characters considered to Somalia, southward to the coastal lowlands be convergent in other cyprinodontiforms: of South Africa; Madagascar; the Sey- mesethmoid cartilaginous, tubular anterior chelles; Indian subcontinent and nans, males always larger than females, and (Ceylon), eastward through the Indo-Malay- a reduced supraorbital sensory pore system. sian Archipelago to Java. COMPOSITION: TWO families, Rivulidae Myers and Aplocheilidae Bleeker, as defined GENUS APLOCHEILUS MCCLELLAND below. Aplocheilus McClelland, 1839, p. 301 (type DISTRIBUTION: Pantropical and Old World species Esox panchax Hamilton-Buchanan, by temperate Laurasian, one family (Rivulidae) original designation). New World, the other (Aplocheilidae) Old Panchax Cuvier and Valenciennes, 1846, p. 380 World, with distributions as detailed below. (type species Esox panchax Hamilton-Buchan- an, by original designation). FAMILY APLOCHEILIDAE BLEEKER Haplochilus Agassiz, 1846, p. 24 (proposed as an emendation of Aplocheilus McClelland). Type Genus Aplocheilus McClelland, 1839. ETYMOLOGY: Aplocheilus from the Greek DIAGNOSIS: Distinguished from all other aplos, meaning single or simple and cheilus, Cyprinodontiformes in having the supra- meaning lip, referring to the thin upper and cleithrum fused to the posttemporal; pre- lower jaw margins. maxillary ascending processes tapered pos- COMPOSITION: Five species: panchax teriorly; basihyal reduced to a small triangular (Hamilton-Buchanan), with seven nominal ossification; and the interarcual cartilage at- subspecies; werneri Meinken; lineatus (Cu- taching directly to the lateral surface of the vier and Valenciennes); blocki (Arnold); and cartilaginous articular surface of the second dayi (Steindachner). pharyngobranchial. DIAGNOSIS: Distinguished from all other COMPOSITION: Six recognized genera: cyprinodontiforms by a derived upper jaw in Aplocheilus McClelland, Epiplatys Gill with which the premaxillary ascending processes 472 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

overlap in the midline, and the lower jaw is toral girdle lowset; first postcleithrum greatly attenuated. present; posttemporal with unossified lower DEFINITION: Anal: Hi, 12-iii, 13; Dorsal: limb; posttemporal fused to supracleithrum. ii, 6; Pelvic: 6; Pectoral: 14; Caudal: 5, 14, Vomer ossified, dentigerous; medial arm 5. Vertebrae: 13+14. Gill rakers on the an- of maxilla twisted with no pronounced dorsal terior arm of the first arch: 12; Branchioste- process; ventral arms gently curved toward gal rays: 6. Scales lateral series: 25-31. and abutting rostral cartilage; outer arm nar- First pleural rib on parapophysis of second row. vertebra; parapophysis not reduced; a pleu- Premaxillary ascending processes flat and ral rib often on first hemal spine; hypural broad, tapered posteriorly and overlapping plates divided, upper plate often divided in in the midline; rostral cartilage large and pen- two. Epipleural ribs not bifid. tagonal; outer arm of premaxilla with alveo- Anal fin not modified into a gonopodium; lar process, not indented posteriorly. Liga- anal fin musculature unmodified; first proxi- ment extending from ventral arms of mal radial present; middle anal radials pres- maxillaries to middle of rostral cartilage; eth- ent. momaxillary ligament present; meniscus Spermatozeugmata not formed; fertiliza- present between premaxilla and maxilla. tion external; development nonannual; ovip- Dentary not expanded medially, not ro- arous. Eggs round. bust; coronoid process on dentary overlap- One dorsal ray articulating with each of ping with that of articular; retroarticular not the first two dorsal radials; dorsal fin on pos- elongate. Autopalatine with straight head, terior third of body. ventral process not elongate, not reaching Autopterotic fossa normal; lateral ethmoid quadrate; metapterygoid present. not expanded medially; not reaching para- Orbital rim free dorsally; anterior nares tu- sphenoid; parasphenoid not expanded ante- bular; supraorbital sensory pores reduced to riorly; weakly formed supraoccipital and a series of neuromasts; seven preopercular epiotic processes; neural spine on first ver- pores, four mandibular pores; two or three tebra; first vertebra articulates with skull via lacrimal pores. basioccipital and exoccipital condyles; su- Males larger than females, often with fin praoccipital excluded from formation of fo- extensions; pigment pattern in many cases ramen magnum; parietals present; nasal not composed of several dark crossbars on the expanded medially. sides of the body; all species with a spot at Mesethmoid cartilaginous; medial pro- the anterior base of the dorsal fin at least in cesses of pelvic fin base and ischial process males; often a darkened caudal margin; mid- reduced; interarcual cartilage large, attaches dle rays of caudal elongate in juveniles and laterally to second pharyngobranchial which adults; throat bars present. lacks bony flange; basihyal broad anteriorly, No fatty predorsal ridge; caudal scaled for triangular ossification posteriorly; tooth one-third its length; swimbladder extending patches on second and third hypobranchials; posteriorly to parhypural. teeth on fourth ceratobranchials; dorsal and DISTRIBUTION: Indian subcontinent and ventral hypohyal present; anterior extension Sri Lanka (Ceylon) eastward along the Indo- of anterior ceratohyal ventral to hypohyals; Malaysian Archipelago to Java. no uncinate process on fourth epibranchial REMARKS: Members of the genus Aplo- to articulate with that of third; first epibran- cheilus as defined herein have been most re- chial narrow at its base. Interhyal ossified; cently reviewed by Radda (1973). He fol- three ossified basibranchials. Vomer with lowed Scheel (1972) in treating species of the posterior extension dorsal to parasphenoid. genus Epiplatys as a subgenus, and there- Lacrimal narrow and twisted carrying dis- fore, referred to the species of Aplocheilus tinct sensory canal; dermosphenotic and pre- as forming a subgenus. In the present study opercular with distinct sensory canal; pec- Aplocheilus and Epiplatys are not consid- 1981 PARENTI: CYPRINODONTIFORM FISHES 473 ered to be synonyms, rather Aplocheilus is mal radial present; middle anal radials pres- considered to be more closely related to the ent. genus Pachypanchax. Spermatozeugmata not formed; fertiliza- Also, for a variety of reasons, Aplocheilus tion external; development nonannual; ovip- has long been considered to be the most arous. Eggs round. primitive cyprinodontiform genus. Derived One dorsal ray articulating with each of characters it shares with other Old World the first two dorsal radials; dorsal fin on pos- aplocheiloids refute this hypothesis. terior third of body. MATERIAL EXAMINED: A. panchax: Gulf Autopterotic fossa normal; lateral ethmoid of Thailand, CAS 37934 (5*/64); India: Ma- not expanded medially, not reaching para- dras, SU 41523 (35); Malaya: SU 32785 (19). sphenoid; parasphenoid not expanded ante- A. lineatus: India: Calicut: SU41516 (3*731). riorly; weakly formed supraoccipital and Aquarium material: AMNH 21498 SW (2+/ epiotic processes; neural spine on first ver- 2). A. blocki: S. India: Cochin: SU 41513 tebra; first vertebra articulates with skull via (14). basioccipital and exoccipital condyles; su- praoccipital excluded from formation of fo- ramen magnum; parietals present; nasals not GENUS PACHYPANCHAX MYERS expanded medially. Mesethmoid cartilaginous; medial pro- Pachypanchax Myers, 1933b, p. 1 (type species cesses of pelvic fin base and ischial process Haplochilus playfairii Gunther, by original designation). reduced; interarcual cartilage large, attaches laterally to second pharyngobranchial which ETYMOLOGY: Pachypanchax from the lacks bony flange; basihyal broad anteriorly, Greek pachy, meaning robust and Panchax, triangular ossification posteriorly; tooth a synonym of a related genus, referring to patches on second and third hypobranchials; the robust appearance of this genus. teeth on fourth ceratobranchials; dorsal and TYPES: Seychelles: Haplochilus playfairii ventral hypohyal present; anterior extension Gunther, Syntypes, BMNH 1864.11.15:91- of anterior ceratohyal ventral to hypohyals; 93(2). no uncinate process on fourth epibranchial COMPOSITION: TWO species: playfairi to articulate with that of third; first epibranch- (Gunther), and homalanotus (Dumeril). ial narrow at its base. Interhyal ossified; DIAGNOSIS: Distinguished from other cy- three ossified basibranchials. Vomer with prinodontiforms in having the posterior edge posterior extension dorsal to parasphenoid. of scales of males angled away from the body Lacrimal narrow and twisted carrying dis- in life; from other members of the Aplo- tinct sensory canal; dermosphenotic and pre- cheilus-Pachy panchax-Epiplatys group in opercular with distinct sensory canal; pec- having the hypural plates fused into a hy- toral girdle lowset; first postcleithrum pural fan in adults. present; posttemporal with unossified lower DEFINITION: Anal: ii, 15-ii, 16; Dorsal: ii, limb; posttemporal fused to supracleithrum. 9-iii, 9; Pelvic: 6; Pectoral: 19; Caudal: 6, Vomer ossified, dentigerous; medial arm 21, 6; Vertebrae: 13+16. Gill rakers on first of maxilla twisted with no pronounced dorsal arch: 10; Branchiostegal rays: 6. Scales lat- process; ventral arms gently curved toward eral series: 25-27. and abutting rostral cartilage; outer arm nar- First pleural rib on parapophysis of second row. vertebra; parapophysis not reduced; no pleu- Premaxillary ascending processes flat and ral ribs on hemal spines; hypural plates fused broad, tapered posteriorly, not overlapping into hypural fan in adults, joint line visible in the midline; rostral cartilage large and pen- in juveniles. Epipleural ribs not bifid. tagonal; outer arm of premaxilla with alveo- Anal fin not modified into a gonopodium; lar process, not indented posteriorly. Liga- anal fin musculature unmodified; first proxi- ment extending from ventral arms of 474 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

maxillaries to middle of rostral cartilage; eth- referring to the flattened dorsal aspect of the momaxillary ligament present; meniscus skull. present between premaxilla and maxilla. TYPES: Gaboon: Haplochilus sexfasciatus Dentary not expanded medially, not ro- Gill, Types, ANSP 7129 to 7141. bust; coronoid process on dentary overlap- COMPOSITION: Over 50 nominal species ping with that of articular; retroarticular not and subspecies as listed in Lazara (1979). elongate. Autopalatine with straight head, DIAGNOSIS: Distinguished from other ventral process not elongate, not reaching members of the Aplocheilus-Pachypanchax- quadrate; metapterygoid present. Epiplatys group by the following primitive Orbital rim free dorsally; anterior naris tu- characters found derived in Aplocheilus and bular; supraorbital sensory pores reduced to Pachypanchax: posttemporal with an ossi- a series of neuromasts; seven preopercular fied lower limb; orbital rim completely at- pores, four mandibular pores; two or three tached; no teeth on the hypobranchials; no lacrimal pores. darkened caudal margin, and no dorsal ocel- Males larger than females, never with fin lus. extensions; pigment pattern not consisting of DEFINITION: Anal: iv, 12; Dorsal: iv, 7; crossbars; males and females with faint red Pelvic: 6; Pectoral: 17; Caudal: 6, 20, 6; Ver- reticulations; juveniles and females with spot tebrae: 11 + 14. Gill rakers on the anterior at anterior base of the dorsal fin (in play- arm of the first arch: 8; Branchiostegal rays: fairi); a darkened caudal and anal margin in 6. Scales lateral series: 26-30. males of play>j"aid; middle rays of caudal nev- First pleural rib on parapophysis of second er elongate; throat bars present. vertebra; parapophysis not reduced; a pleu- No fatty predorsal ridge; caudal scaled for ral rib often on first hemal spine; hypural at least one-third its length; swimbladder ex- plates divided, upper plate often divided in tending posteriorly to parhypural. two. Epipleural ribs not bifid. DISTRIBUTION: Madagascar, the Sey- Anal fin not modified into a gonopodium; chelles, coastal lowlands of eastern Mozam- anal fin musculature unmodified; first proxi- bique, and Zanzibar north of the Zambezi mal radial present; middle anal radials pres- River. ent. REMARKS: The genus Pachypanchax is Spermatozeugmata not formed; fertiliza- considered to contain two species: playfairi, tion external; development nonannual; ovip- the type, and homalanotus. However, a arous. Eggs round. specimen of homalanotus was not available One dorsal ray articulating with each of for osteological examination, therefore, it is the first two dorsal radials; dorsal fin on pos- included with caution in the genus. terior third of body. MATERIAL EXAMINED: P. playfairi: Sey- Autopterotic fossa normal; lateral ethmoid chelles: Syntypes as listed above, AMNH not expanded medially, not reaching para- 20637 (4*/17); Zanzibar: AMNH 20701 (4*/ sphenoid; parasphenoid not expanded ante- 19); Aquarium material: AMNH 38413 (3*/ riorly; weakly formed supraoccipital and 3); P. homalanotus: Aquarium material: SU epiotic processes; neural spine on first ver- 52679(1). tebra; first vertebra articulates with skull via basioccipital and exoccipital condyles; su- praoccipital excluded from formation of fo- GENUS EPIPLATYS GILL ramen magnum; parietals present; nasals not expanded medially. Epiplatys Gill, 1862, p. 136 (type species Haplo- Mesethmoid cartilaginous; medial pro- chilus sexfasciatus Gill, by original designa- tion). cesses of pelvic fin base and ischial process reduced; interarcual cartilage large, attaches ETYMOLOGY: Epiplatys from the Greek laterally to second pharyngobranchial which epi, meaning above, and platys meaning flat, lacks bony flange; basihyal broad anteriorly, 1981 PARENTI: CYPRINODONTIFORM FISHES 475 triangular ossification posteriorly; no tooth tending posteriorly to first or second hemal patches on second and third hypobranchials; spine. teeth on fourth ceratobranchials; dorsal and DISTRIBUTION: West and central Africa: ventral hypohyal present; anterior extension eastward from Senegal, northern limit the of anterior ceratohyal ventral to hypohyals; Niger River, eastward to the Ethiopian High- no uncinate process on fourth epibranchial lands, southward west of the Rift lakes to the to articulate with that of third; first epibran- Katanga Plateau, then eastward to the coast. chial narrow at its base. Interhyal ossified; REMARKS: The genus Epiplatys as consti- three ossified basibranchials. Vomer with tuted in this study is paraphyletic. However, posterior extension dorsal to parasphenoid. rather than create new generic names for Lacrimal narrow and twisted carrying dis- subgroups of the genus which could not be tinct sensory canal; dermosphenotic and pre- incorporated into an overall phylogenetic opercular with distinct sensory canal; pec- scheme, the species group names which al- toral girdle lowset; first postcleithrum ready exist are treated as subgenera of the present; posttemporal with ossified lower genus Epiplatys until such time that they limb; posttemporal fused to supracleithrum. may be defined and their relationships deter- Vomer ossified, dentigerous; medial arm mined. These subgenera are the following: of maxilla twisted with no pronounced dorsal process; ventral arms gently curved toward and abutting rostral cartilage; outer arm nar- SUBGENUS LYCOCYPRINUS PETERS row. Lycocyprinus Peters, 1868, p. 146 (type species Premaxillary ascending processes flat and Epiplatys dageti Poll by monotypy). broad, tapered posteriorly, not overlapping in the midline; rostral cartilage large and pen- SUBGENUS PAREPIPLATYS CLAUSEN tagonal; outer arm of premaxilla with alveo- lar process, not indented posteriorly. Liga- Parepiplatys Clausen, 1967, p. 28 (type species ment extending from ventral arms of Haplochilus grahami Boulenger, by original maxillaries to middle of rostral cartilage; eth- designation [proposed as a subgenusj). momaxillary ligament present; meniscus present between premaxilla and maxilla. SUBGENUS PSEUDEPIPLATYS CLAUSEN Dentary not expanded medially, not ro- bust; coronoid process on dentary not over- Pseudepiplatys Clausen, 1967, p. 30 (type species lapping with that of articular; retroarticular Haplochilus annulatus Boulenger, by original not elongate. Autopalatine with straight designation [proposed as a subgenus]). head, ventral process not elongate, not reaching quadrate; metapterygoid present. SUBGENUS APHYOPLATYS CLAUSEN Orbital rim attached; anterior naris tubu- lar; supraorbital sensory pores reduced to a Aphyoplatys Clausen, 1967, p. 32 (type species Epiplatys duboisi Poll, by original designation series of neuromasts; seven preopercular [proposed as a subgenus]). pores, three mandibular pores; two or three lacrimal pores. MATERIAL EXAMINED: E. sexfasciatus: Males larger than females, often with fin Dahomey, Iquidi R: USNM 218752 (2*721); extensions; pigment pattern often composed E. chaperi: Ghana: SU 64709 (1*712); E.fas- of several dark crossbars on the sides of the ciolatus: Liberia: Tchien: AMNH 32735 (5*7 body; no species with a spot at the anterior 10); E. bifasciatus: Nigeria (Aquarium ma- base of the dorsal fin; middle rays of caudal terial) AMNH 21866SW (1+/1). E. senegal- elongate in juveniles and adults; throat bars ensis: Nigeria (Aquarium material) AMNH present. 21867 (1+/1); E. dageti: Ghana: SU 64640 No fatty predorsal ridge; caudal scaled for (29). Haplochilus annulatus: Type: Sierra at least one-third its length; swimbladder ex- Leone: BMNH 1914. 12. 9: 5-6. 476 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

GENUS APHYOSEMION MYERS Mesethmoid cartilaginous; medial pro- cesses of pelvic fin base and ischial process Aphyosemion Myers, 1924b, p. 2 (type species Aphysemion castaneum Myers, by original reduced; interarcual cartilage large, attaches designation). laterally to second pharyngobranchial which lacks bony flange; basihyal broad anteriorly, ETYMOLOGY: Aphyosemion from the triangular ossification posteriorly; no tooth Greek aphyos, meaning small and semion patches on second and third hypobranchials; meaning flag or banner referring to the lyre- teeth on fourth ceratobranchials; dorsal and shaped caudal fin of males. ventral hypohyal present; anterior extension TYPES: Zaire (Congo): Aphyosemion cas- of anterior ceratohyal ventral to hypohyals; taneum Myers, Type AMNH 8337 (1). uncinate process on fourth epibranchial ar- COMPOSITION: Approximately 60 species ticulates with that of third; first epibranchial as listed in Lazara (1979). narrow at its base. Interhyal ossified; three DIAGNOSIS: Distinguished from other ossified basibranchials. Vomer with atten- members of the Aphyosemion-Nothobran- uate posterior extension dorsal to parasphe- chius group by being nonannual, having a noid. dorsal fin of seven to 14 rays which is situ- Lacrimal narrow and twisted carrying dis- ated no farther anteriorly than opposite the tinct sensory canal; dermosphenotic and pre- first quarter of the anal fin orgin, and pos- opercular with distinct sensory canal; pec- sessing a swimbladder extending posteriorly toral girdle lowset; first postcleithrum to the first one or two hemal spines. present; posttemporal with ossified lower DEFINITION: Anal: iii, 12; Dorsal: i, 6-ii, limb; posttemporal fused to supracleithrum. 12; Pelvic: 6; Pectoral: 16-17; Caudal: 8, 13, Vomer ossified, dentigerous; medial arm 8; Vertebrae: 13 + 15. Gill rakers on anterior of maxilla twisted with no pronounced dorsal arm of the first arch: 8, 9; Branchiostegal process; ventral arms gently curved toward rays: 6. Scales Lateral Series: 29-33. and abutting rostral cartilage; outer arm nar- First pleural rib on parapophysis of second row. vertebra; parapophysis not reduced; no pleu- Premaxillary ascending processes flat and ral rib on first hemal spine; hypural plates broad, tapered posteriorly not overlapping in divided, upper plate never divided in two. the midline; rostral cartilage large and pen- Epipleural ribs bifid. tagonal; outer arm of premaxilla with alveo- Anal fin not modified into a gonopodium; lar process, not indented posteriorly. Liga- anal fin musculature unmodified; first proxi- ment extending from ventral arms of mal radial present; middle anal radials pres- maxillaries to middle of rostral cartilage; eth- ent. momaxillary ligament present; meniscus Spermatozeugmata not formed; fertiliza- present between premaxilla and maxilla. tion external; development nonannual; ovip- Dentary not expanded medially, not ro- arous. Eggs round. bust; coronoid process on dentary not over- One dorsal ray articulating with each of lapping with that of articular; retroarticular the first two dorsal radials; dorsal fin on pos- not elongate. Autopalatine with straight terior third of body. head, ventral process not elongate, not Autopterotic fossa normal; lateral ethmoid reaching quadrate; metapterygoid present. not expanded medially, not reaching para- Orbital rim attached; anterior nans tubu- sphenoid; parasphenoid not expanded ante- lar; supraorbital sensory pores reduced to a riorly; weakly formed supraoccipital and series of neuromasts; seven preopercular epiotic processes; neural spine on first ver- pores, four mandibular pores; two or three tebra; first vertebra articulates with skull via lacrimal pores. basioccipital and exoccipital condyles; su- Males larger than females, often with fin praoccipital excluded from formation of fo- extensions; pigment pattern rarely composed ramen magnum; parietals present; nasals not of several dark crossbars on the sides of the expanded medially. body; no species with a spot at the anterior 1981 PARENTI: CYPRINODONTIFORM FISHES 477 base of the dorsal fin; rarely middle rays of Gery by original designation [proposed as a caudal elongate in juveniles and adults; subgenus]). throat bars weakly present. Males with light dorsal and ventral caudal margins; often light SUBGENUS KATHETYS HUBER anal caudal margins; "wound" spot typically Kathetys Huber, 1977, p. 8 (type species Fun- present. dulus exiguus Boulenger by original designa- No fatty predorsal ridge; caudal scaled for tion [proposed as a subgenus]). at least one-third its length; swimbladder ex- tending posteriorly to first one or two hemal SUBGENUS MESOAPHYOSEMION RADDA spines. DISTRIBUTION: Western central Africa, Mesoaphyosemion Radda, 1977, p. 213 (type concentrated in the Congo Basin, westward species Haplochilus cameronensis Boulenger, by original designation [proposed as a to Gambia along the coast. subgenus]). REMARKS: Aphyosemion Myers cannot be defined as a monophyletic group; nor are the MATERIAL EXAMINED: A. castaneum: interrelationships of its named subgenera Zaire: type as listed above; Paratypes: AMNH within the scope of the present paper. Most 8338 (l*/4); A. petersi: Aquarium material supraspecific categories are recently named AMNH 21572 (2+/2); Ghana: SU 64709 and encompass a large number of species. (2*/28). A. cameronensis: Cameroon: Ntem Therefore, the genus Aphyosemion is divid- R. SU 15713 (8). A. bivittatum: Cameroon: ed into two major groupings, one encom- SU55491 (l*/6). passing the more primitive species and the other encompassing the more derived species GENUS FUNDULOPANCHAX MYERS that are more closely related to Nothobran- Fundulopanchax Myers, 1924b, p. 4 (type species chius than to other species of Aphyosemion. Fundulus sjoestedti Lonnberg, by monotypy The primitive members are referred to the [proposed as a subgenus]). genus Aphyosemion, whereas the derived members are referred to the genus Fundu- ETYMOLOGY: Fundulopanchax from Fun- lopanchax, an available supraspecific cate- dulus and Panchax, two nominal genera, the gory. Huber (1978) grouped all the subgenera former from the New World and the latter within Aphyosemion, therefore not recogniz- from the Old World, between which Fun- ing the closer association of Fundulopan- dulopanchax was thought to be intermediate. chax to Nothobranchius. The subgenera of COMPOSITION: Approximately 50 species Aphyosemion of the more primitive grouping currently referred to the included subgeneric are: categories. DIAGNOSIS: Distinguished from other species of Aphyosemion by elongate dorsal SUBGENUS ARCHIAPHYOSEMION RADDA fin of 14 rays or more, and the swimbladder Archiaphyosemion Radda, 1977, p. 214 (type which does not penetrate beyond the first species Aphyosemion guineense Daget, by hemal spine. All included species are annual. original designation [proposed as a subgenus]). DEFINITION: Anal: iii, 13; Dorsal: i, 13-ii, 18; Pelvic: 6; Pectoral: 17-18; Caudal: 8, 14, SUBGENUS CHROMAPHYOSEMION RADDA 8; Vertebrae: 12+15-14+19. Gill rakers on anterior arm of the first arch: 12, Branchios- Chromaphyosemion Radda, 1971, p. 157 (type tegal rays: 6; Scales lateral series: 33-37. species Fundulus bivittatus Boulenger, by orig- inal designation [proposed as a subgenus]). First pleural rib on parapophysis of second vertebra; parapophysis not reduced; no pleu- ral rib on first hemal spine; hypural plates SUBGENUS DIAPTERON HUBER AND SEEGERS divided, upper plate never divided in two. Diapteron Huber and Seegers, 1977, p. 146 (type Epipleural ribs bifid. species Aphyosemion georgiae Lambert and Anal fin not modified into a gonopodium; 478 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 anal fin musculature unmodified; first proxi- lar process, not indented posteriorly. Liga- mal radial present; middle anal radials pres- ment extending from ventral arms of ent. maxillaries to middle of rostral cartilage; eth- Spermatozeugmata not formed; fertiliza- momaxillary ligament present; meniscus tion external; development annual; ovipa- present between premaxilla and maxilla. rous. Eggs round. Dentary not expanded medially, not ro- One dorsal ray articulating with each of bust; coronoid process on dentary not over- the first two dorsal radials; dorsal fin oppo- lapping with that of articular; retroarticular site or anterior to origin of anal. not elongate. Autopalatine with straight Autopterotic fossa normal; lateral ethmoid head, ventral process not elongate, not not expanded medially, not reaching para- reaching quadrate; metapterygoid present. sphenoid; parasphenoid not expanded ante- Orbital rim attached; anterior naris tubu- riorly; weakly formed supraoccipital and lar; supraorbital sensory pores reduced to a epiotic processes; neural spine on first ver- series of neuromasts; seven preopercular tebra; first vertebra articulates with skull via pores, four mandibular pores; two or three basioccipital and exoccipital condyles; su- lacrimal pores. praoccipital excluded from formation of fo- Males larger than females, often with fin ramen magnum; parietals present; nasals not extensions; pigment pattern rarely composed expanded medially. of several dark crossbars on the sides of the Mesethmoid cartilaginous; medial pro- body; no species with a spot at the anterior cesses of pelvic fin base and ischial process base of the dorsal fin; often middle rays of reduced; interarcual cartilage large, attaches caudal elongate in juveniles and adults; laterally to second pharyngobranchial which "Wound" mark typically present; males lacks bony flange; basihyal broad anteriorly, often with light dorsal and ventral margins of triangular ossification posteriorly; no tooth caudal, as well as dorsal and anal. patches on second and third hypobranchials; No fatty predorsal ridge; caudal scaled for teeth on fourth ceratobranchials; dorsal and at least one third its length; swimbladder ex- ventral hypohyal present; anterior extension tending just to the first hemal spine. of anterior ceratohyal ventral to hypohyals; DISTRIBUTION: Congo Basin. uncinate process on fourth epibranchial ar- REMARKS: Fundulopanchax contains ticulates with that of third; first epibranchial species previously assigned to Aphyosemion narrow at its base. Interhyal ossified; three that are more closely related to Nothobran- ossified basibranchials. Vomer with atten- chius than to the more primitive species of uate posterior extension dorsal to parasphe- Aphyosemion. This usage of the genus Fun- noid. dulopanchax differs from that of Loiselle and Lacrimal narrow and twisted carrying dis- Glasgow (1971) who limited the group to tinct sensory canal; dermosphenotic and pre- large forms such as sjoestedti. opercular with distinct sensory canal; pec- The designation of a type species for the toral girdle lowset; first postcleithrum genus Callopanchax as A. occidentale by present; posttemporal with ossified lower the International Commission places the ge- limb; posttemporal fused to supracleithrum. nus Roloffia as an objective synonym of Cal- Vomer ossified, dentigerous; medial arm lopanchax. This decision has been rejected of maxilla twisted with no pronounced dorsal by many aquarists who believe that Myers process; ventral arms gently curved toward (1924b) had either sjoestedti or another and abutting rostral cartilage; outer arm nar- species in hand when he named the subgenus row. Callopanchax, with sjoestedti as the type, Premaxillary ascending processes flat and even though giving a description of occiden- broad, tapered posteriorly not overlapping in tale. It is interesting to note that the sole lot the midline; rostral cartilage large and pen- catalogued as sjoestedti in the collection of tagonal; outer arm of premaxilla with alveo- the AMNH is an aquarium lot of occiden- 1981 PARENTI: CYPRINODONTIFORM FISHES 479 tale. Therefore, it is likely that Myers had TYPES: Zaire: Village of Ntokon near the specimens of occidentale not sjoestedti in banks of the Likouala-Mossaka. Adamas hand when naming Callopanchax. formosus Huber, Holotype MNHN 1979-199 The subgenera placed in Fundulopanchax (1). are: COMPOSITION: Solely the type species. DIAGNOSIS: Distinguished from other SUBGENUS PALUDOPANCHAX RADDA aplocheiloids by the presence of a brilliant diamond shaped frontal spot. Paludopanchax Radda, 1977, p. 211 (type species DEFINITION: Anal: 15, Dorsal: 8-9; dorsal Fundulus arnoldi Boulenger, by original des- ignation). fin originates opposite the twelfth to four- teenth rays of anal. SUBGENUS PARAPHYOSEMION KOTTELAT Both males and females are characterized by a brilliant diamond-shaped frontal spot. Paraphyosemion Kottelat, 1976, p. 158 (type There are no dark crossbars. species Fundulus gardneri Boulenger, by orig- DISTRIBUTION: That of the type locality inal designation [proposed as a subgenus]). and environs. REMARKS: Specimens of this new genus SUBGENUS GULAROPANCHAX RADDA and species have not been examined. Data Gularopanchax Radda, 1977, p. 210 (type species are from the original description of Huber Fundulus gularis Boulenger, by original des- (1979). ignation [proposed as a subgenus]). Huber considered it difficult to place his new genus in a subfamily of aplocheiloids or SUBGENUS CALLOPANCHAX MYERS of procatopines, citing as evidence of rela- tionship with the former the overall mor- Callopanchax Myers, 1933c, p. 184 (type species Aphyosemion occidentale Roloff, by monotypy phology and color pattern, and with the latter [proposed as a subgenus]). the behavior and biology, on which he did Roloffia Clausen, 1966, p. 388 (type species not elaborate. Aphyosemion occidentale Roloff, by original On the basis of its external morphology designation). and dorsal and anal fin ray number and po- sition, I consider Adamas to represent just SUBGENUS RADDAELLA HUBER one more species allied to Aphyosemion. The concept that the Old World aplochei- Raddaella Huber, 1977, p. 8 (type species Fun- loids and procatopines are closely related dulus batesi Boulenger, by original designation [proposed as a subgenus]). has prevailed, although there is no evidence to support such a relationship. MATERIAL EXAMINED: F. sjoestedti: Aquarium material: AMNH 21575 (4+/4); F. GENUS NOTHOBRANCHIUS PETERS gardneri: Ghana: SU 64693 (l*/7); F. gulare: Aquarium material: AMNH 20563 (3+/13); Nothobranchius Peters, 1868, p. 10 (type species F. arnoldi: Nigeria: Port Harcourt: AMNH Cyprinodon orthonotus Peters, by original des- 21570 (1+/1). F. occidentale: Aquarium ma- ignation). terial: AMNH 14611 (2*/6). Adiniops Myers, 1924b, p. 6 (type species Fun- dulus guentheri Pfeffer, by original designation GENUS ADAMAS HUBER [proposed as a subgenus]). Fundulosoma Ahl, 1924, p. 52 (type species Fun- Adamas Huber, 1979, p. 5, 6 (type species Ada- dulosoma thierryi Ahl, by original designation). mas formosus Huber, by original designation). Pronothobranchius Radda, 1969, p. 4 (type species Nothobranchius kiyawensis Ahl, by ETYMOLOGY: Adamas from the Greek, original designation [proposed as a subgenus]). meaning steel or diamond which refers to the Zonothobranchius Radda, 1969, p. 4 (type species brilliant frontal spot. Nothobranchius rubroreticulatus Blache and 480 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

Miton, by original designation [proposed as a ramen magnum; parietals present; nasals not subgenus]). expanded medially. Aphyobranchius Wildekamp, 1977, p. 327 (type Mesethmoid cartilaginous; medial pro- species Nothobranchius janpapi Wildekamp, cesses of pelvic fin base and ischial process by original designation [proposed as a reduced; interarcual cartilage large, attaches subgenus]). directly to articulation point, second pharyn- ETYMOLOGY: Nothobranchius, from the gobranchial which lacks bony flange; basi- Greek nothos, meaning false and branchia hyal broad anteriorly, triangular ossification meaning gills, in reference to the restricted posteriorly; no tooth patches on second and gill opening. third hypobranchials; teeth on fourth cera- TYPES: Mozambique: Quelimane: Cyprin- tobranchials; dorsal and ventral hypohyal odon orthonotus Peters, Syntype BMNH present; anterior extension of anterior cera- 1861. 5.2: 88-9(1). tohyal ventral to hypohyals; uncinate pro- COMPOSITION: Approximately 35 species cess on fourth epibranchial articulates with as listed in Lazara (1979). that of third; first epibranchial narrow at its DIAGNOSIS: Distinguished from all other base. Interhyal ossified; three ossified basi- cyprinodontiforms by the interarcual carti- branchials. Vomer with attenuate posterior lage attaching directly to the cartilage of the extension dorsal to parasphenoid. second pharyngobranchial and oval, rather Lacrimal narrow and twisted carrying dis- than round eggs; and from other aplochei- tinct sensory canal; dermosphenotic and pre- loids by an uncovered preopercular canal. opercular with distinct sensory canal; pec- DEFINITION: Anal: ii, 13-ii, 17; Dorsal: ii, toral girdle lowset; first postcleithrum 13-ii, 16; pelvic: 6; Pectoral: 16-21; Caudal: present; posttemporal with ossified lower 8, 15, 8; Vertebrae: 12+15-12+17. Gill rak- limb; posttemporal fused to supracleithrum. ers on anterior arm of the first arch: 12-21. Vomer ossified, dentigerous; medial arm Branchiostegal rays: 6-7; Scales lateral se- of maxilla twisted with no pronounced dorsal ries: 26-42. process; ventral arms gently curved toward First pleural rib on parapophysis of second and abutting rostral cartilage; outer arm nar- vertebra; parapophysis not reduced; no pleu- row. ral rib on first hemal spine; hypural plates Premaxillary ascending processes flat and fused into a hypural fan. Epipleural ribs bi- broad, tapered posteriorly not overlapping in fid. the midline; rostral cartilage large and pen- Anal fin not modified into a gonopodium; tagonal; outer arm of premaxilla with alveo- anal fin musculature unmodified; first proxi- lar process, not indented posteriorly. Liga- mal radial present; middle anal radials pres- ment extending from ventral arms of ent. maxillaries to middle of rostral cartilage; eth- Spermatozeugmata not formed; fertiliza- momaxillary ligament present; meniscus tion external; development annual; ovipa- present between premaxilla and maxilla. rous. Eggs oval. Dentary not expanded medially, not ro- One dorsal ray articulating with each of bust; coronoid process on dentary not over- the first two dorsal radials; dorsal fin oppo- lapping with that of articular; retroarticular site or anterior to origin of anal. not elongate. Autopalatine with straight Autopterotic fossa normal; lateral ethmoid head, ventral process not elongate, not not expanded medially, not reaching para- reaching quadrate; metapterygoid present. sphenoid; parasphenoid not expanded ante- Orbital rim attached; anterior nares tubu- riorly; weakly formed supraoccipital and lar; supraorbital sensory pores reduced to a epiotic processes; neural spine on first ver- series of neuromasts; preopercular canal rep- tebra; first vertebra articulates with skull via resented by an open groove, four mandibular basioccipital and exoccipital condyles; su- pores; two or three lacrimal pores. praoccipital excluded from formation of fo- Males larger than females, caudal fin ex- 1981 PARENTI: CYPRINODONTIFORM FISHES 481 tension in thierryi; caudal typically rounded; "Rivulus," Trigonectes, Neofundulus, pigmentation pattern typically consisting of "Neofundulus," Rachovia, Cynolebias, red reticulations, or body uniformly red and Pterolebias and Austrofundulus. blue throughout. Vertical bar through eye in DISTRIBUTION: (fig. 18) New World from one species (microlepis). southern Florida and the Bahamas, Cuba, No fatty predorsal ridge; caudal scaled for Hispaniola, Trinidad, Middle America, south at least one third its length; swimbladder ex- through South America to Uruguay. tending to the first hemal spine. DISTRIBUTION: West central and east Af- rica, northern limit the Niger River, east to GENUS RIVULUS POEY Somalia, and south to northern South Africa. Rivulus Poey, 1860, p. 307 (type species Rivulus REMARKS: The composition of the genus cylindraceus Poey, by original designation). Nothobranchius is increased here by one Cynodonichthys Meek, 1904, p. 101 (type species species, thierryi, formally placed in the Cynodonichthys tenuis Meek, by original des- monotypic Fundulosoma. There is no rep- ignation). resentative of this genus in the Seychelles, Vomerivulus Fowler, 1944, p. 244 (type species and the description of N. seychellensis Rivulus leucurus Fowler, by original designa- tion). seems to have been based on material from Africa. ETYMOLOGY: The genus Rivulus meaning MATERIAL EXAMINED: N. orthonotus: small stream or rivulet, the typical habitat of Mozambique: the type as listed above; Kru- these killifishes. ger National Park: AMNH 22255SW (1+/1). TYPES: Cuba: Stream at Mardaza, near N. thierryi: Ghana: BMNH 1970.10.22.1-13. Havana: Rivulus cylindraceus Poey. Types (l*/6); N. guentheri: Zanzibar: AMNH MCZ 6397 (3). 22252 (19+/19); N. melanospilus: Aquarium COMPOSITION: Smaller nonannual species material: USNM uncat. (l*/8); N. kirki: presently referred to Rivulus (as in Lazara, Aquarium material: AMNH 38407 (2*72); N. 1979) including cylindraceus, heyei, and microlepis: Somalia Meridonele: MCSN marmoratus. 15163 (4) Syntypes, AMNH 20588 (l*/4), DIAGNOSIS: Neotropical aplocheiloids with Syntypes; N. patrizii: Somalia: MCSN 33702 ossified interhyals, fin rays not elongate and (4) Syntypes, AMNH 20587 (4) Syntypes; N. six pelvic fin rays. kiyawensis: Nigeria: Cotypes USNM 92820 DEFINITION: Anal: ii, 9-ii, 12; Dorsal: i, (2). 8; Pelvic: 6; Pectoral: 13-14; Caudal: 8, 14, 8-8, 18, 8; Vertebrae: 14+16-20+20. Gill rakers on anterior arm of the first arch: 8; FAMILY RIVULIDAE MYERS Branchiostegal rays: 6. Scales lateral series: Type Genus Rivulus Poey, 1860. 34-51. DIAGNOSIS: The Rivulidae differ from all First pleural rib on parapophysis of second other aplocheiloids by lacking the first post- vertebra; parapophysis not reduced; no pleu- cleithrum and from other cyprinodontiforms ral rib on first hemal spine; hypural plates by having a preoperculum and lacrimal with divided, upper plate never divided in two. obsolescent sensory canals, lateral ethmoid Epipleural ribs not bifid. expanded medially under the lateral exten- Anal fin not modified into a gonopodium; sion of the vomer, opercular, and branchio- anal fin musculature unmodified; first proxi- stegal membranes united and often covered mal radial present; middle anal radials pres- with scales, urohyal not distinct externally, ent. a unique headscale pattern, and a triangular Spermatozeugmata not formed; fertiliza- flange on the anterior face of the ventral pro- tion external; development nonannual; ovip- cess of the maxilla. arous. Eggs round. COMPOSITION: Nine genera: Rivulus, One dorsal ray articulating with each of 482 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 the first two dorsal radials; dorsal fin on pos- lapping with that of articular; retroarticular terior third of body. not elongate. Autopalatine with straight Autopterotic fossa normal; lateral ethmoid head, ventral process not elongate, not expanded medially, extending under para- reaching quadrate; metapterygoid present. sphenoid; parasphenoid slightly expanded Orbital rim attached; anterior nares tubu- anteriorly; weakly formed supraoccipital and lar; supraorbital sensory pores reduced to a epiotic processes; neural spine on first ver- series of neuromasts; preopercular, mandib- tebra; first vertebra articulates with skull via ular, and lacrimal pores absent, replaced by basioccipital and exoccipital condyles; su- a series of weakly formed neuromasts. Bran- praoccipital excluded from formation of fo- chiostegal rays covered. ramen magnum; parietals present; nasals not Males larger than females, all fins round- expanded medially. ed; females with distinct caudal ocellus; Mesethmoid cartilaginous; medial pro- males with light dorsal and ventral caudal fin cesses of pelvic fin base and ischial process margins; no species with a spot at the ante- reduced; interarcual cartilage large, attaches rior base of the dorsal fin; middle rays of laterally to second pharyngobranchial which caudal never elongate in juveniles and possesses a bony flange; basihyal broad an- adults; throat bars weakly present. teriorly, elongate ossification posteriorly; no No fatty predorsal ridge; caudal scaled for tooth patches on second and third hypobran- at least one-third its length; swimbladder ex- chials; teeth on fourth ceratobranchials; dor- tending posteriorly to first one or two hemal sal and ventral hypohyal present; anterior spines. extension of anterior ceratohyal ventral to DISTRIBUTION: Rivulus and "Rivulus" hypohyals; uncinate process on fourth epi- have distribution patterns nearly coincident branchial articulates with that of third; first with that of the family, as listed above. "Riv- epibranchial narrow at its base. Interhyal os- ulus" is excluded from North America, sified; three ossified basibranchials. Vomer however. with posterior extension dorsal to parasphe- REMARKS: Rivulus as constituted tradi- noid. tionally is not monophyletic. Species as- Lacrimal narrow and twisted with obso- signed to the genus may be divided into two lescent sensory canal; dermosphenotic and groups, one a primitive assemblage of preopercular with obsolescent sensory ca- species which possess an ossified interhyal, nal; pectoral girdle lowset; first postclei- six pelvic fin rays, and dorsal and anal fin thrum absent; posttemporal with ossified rays that are not elongate. These species, lower limb; posttemporal not fused to supra- which include the type cylindraceus and oth- cleithrum. er diminutive forms (including marmoratus Vomer ossified, dentigerous; medial arm and heyei), are referred to the genus Rivu- of maxilla twisted with no pronounced dorsal lus. The other group of species, of which process; ventral arms with pronounced an- harti may be considered representative, terior extension abutting rostral cartilage; have a cartilaginous interhyal, seven pelvic outer arm narrow. fin rays and elongate dorsal and anal fin rays, Premaxillary ascending processes flat and derived characters they share with Neotrop- broad, not tapered posteriorly not overlap- ical aplocheiloids. These species are referred ping in the midline; rostral cartilage large and to the genus "Rivulus" until they are revised pentagonal; outer arm of premaxilla with al- at the species level, and monophyletic veolar process, not indented posteriorly. groups of species recognized within both as- Ligament extending from ventral arms of semblages. maxillaries to middle of rostral cartilage; eth- Hoedeman (1961) presented a key to the momaxillary ligament present; meniscus recognized species and subspecies of the ge- present between premaxilla and maxilla. nus Rivulus (which includes the genera Riv- Dentary not expanded medially, not ro- ulus and "Rivulus" herein). bust; coronoid process on dentary not over- MATERIAL EXAMINED: Cuba: R. cylindra- 1981 PARENTI: CYPRINODONTIFORM FISHES 483 ceus: the types as listed above; SU 32018 (2); possesses a bony flange; basihyal broad an- R. marmoratus: Cuba: USNM 164438 (1*76); teriorly, elongate ossification posteriorly; no Florida: AMNH 16117 (6). R. tenuis: Gua- tooth patches on second and third hypobran- temala: Alta Verapaz: AMNH 32069 (2+/2). chials; teeth on fourth ceratobranchials; dor- sal and ventral hypohyal present; anterior extension of anterior ceratohyal ventral to GENUS "RIVULUS" hypohyals; uncinate process on fourth epi- ETYMOLOGY: "Rivulus" is used to refer- branchial articulates with that of third; first ence the more derived species of the genus epibranchial narrow at its base. Interhyal Rivulus. cartilaginous; three ossified basibranchials. DIAGNOSIS: Neotropical aplocheiloids with Vomer with posterior extension dorsal to an ossified interhyal, elongate pelvic fin rays, parasphenoid. seven or eight pelvic fin rays and known or Lacrimal narrow and twisted with obso- inferred annual habit, characters which they lescent sensory canal; dermosphenotic and share with the more derived genera of the preopercular with obsolescent sensory ca- group. nal; pectoral girdle lowset; first postclei- DEFINITION: Anal: iii, 12; Dorsal: i, 8; Pel- thrum absent; posttemporal with ossified vic: 7-8; Pectoral: 16; Caudal: 8, 15, 8; Ver- lower limb; posttemporal not fused to supra- tebrae: 12+18. Gill rakers on anterior arm of cleithrum. the first arch: 8, 9; Branchiostegal rays: 6; Vomer ossified, dentigerous; medial arm Scales lateral series: 35-36. of maxilla twisted with no pronounced dorsal First pleural rib on parapophysis of second process; ventral arms with pronounced an- vertebra; parapophysis not reduced; no pleu- terior extension abutting rostral cartilage; ral rib on first hemal spine; hypural plates outer arm narrow. divided, upper plate never divided in two. Premaxillary ascending processes flat and Epipleural ribs not bifid. broad, not tapered posteriorly, not overlap- Anal fin not modified into a gonopodium; ping in the midline; rostral cartilage enlarged anal fin musculature unmodified; first proxi- and rectangular; outer arm of premaxilla mal radial present; middle anal radials pres- with alveolar process, not indented poste- ent. riorly. Ligament extending from ventral Spermatozeugmata not formed; fertiliza- arms of maxillaries to middle of rostral car- tion external; development known to be an- tilage; ethmomaxillary ligament present; me- nual in at least one species; oviparous. Eggs niscus present between premaxilla and max- round. illa. One dorsal ray articulating with each of Dentary not expanded medially, not ro- the first two dorsal radials; dorsal fin on pos- bust; coronoid process on dentary not over- terior third of body. lapping with that of articular; retroarticular Autopterotic fossa normal; lateral ethmoid not elongate. Autopalatine with straight expanded medially, extending under para- head, ventral process not elongate, not sphenoid; parasphenoid slightly expanded reaching quadrate; metapterygoid present. anteriorly; weakly formed supraoccipital and Orbital rim attached; anterior nares tubu- epiotic processes; neural spine on first ver- lar; supraorbital sensory pores reduced to a tebra; first vertebra articulates with skull via series of neuromasts; preopercular, mandib- basioccipital and exoccipital condyles; su- ular and lacrimal pores absent, replaced by praoccipital excluded from formation of fo- a series of weakly formed neuromasts. Bran- ramen magnum; parietals present; nasals not chiostegal rays covered. expanded medially. Males larger than females, often with fin Mesethmoid cartilaginous; medial pro- extensions; females with distinct caudal cesses of pelvic fin base and ischial process ocellus; males with light dorsal and ventral reduced; interarcual cartilage large, attaches caudal fin margins; no species with a spot at laterally to second pharyngobranchial which the anterior base of the dorsal fin; middle 484 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

rays of caudal never elongate in juveniles ral rib on first hemal spine; hypural plates and adults; throat bars weakly present. divided, upper plate never divided in two. No fatty predorsal ridge; caudal scaled for Epipleural ribs not bifid. at least one-third its length; swimbladder ex- Anal fin not modified into a gonopodium; tending posteriorly to first one or two hemal anal fin musculature unmodified; first proxi- spines. mal radial present; middle anal radials pres- DISTRIBUTION: See Rivulus above. ent. REMARKS: In the description of the annual Spermatozeugmata not formed; fertiliza- Rivulus stellifer Thomerson and Turner tion external; development inferred to be an- (1973) stated that they believed it served as nual; oviparous. Eggs round. a link between the nonannual forms and the One dorsal ray articulating with each of annual Neotropical aplocheiloids. Their sug- the first two dorsal radials; dorsal fin on pos- gestion has been supported here; however, terior third of body. the position of stellifer is restated as more Autopterotic fossa normal; lateral ethmoid closely related to the annual genera than it expanded medially, extending under para- is to the nonannual species of Rivulus. sphenoid; parasphenoid slightly expanded MATERIAL EXAMINED: R. harti: Trinidad: anteriorly; weakly formed supraoccipital and AMNH 15189 (2*719); Venezuela: Margarita epiotic processes; neural spine on first ver- Is.: AMNH 8354 (2+/2). R. stellifer: Vene- tebra; first vertebra articulates with skull via zuela: Cojedes: Paratopotypes: CAS 27556 basioccipital and exoccipital condyles; su- (2). praoccipital excluded from formation of fo- ramen magnum; parietals present; nasals not GENUS TRIGONECTES MYERS expanded medially. Trigonectes Myers, 1925, p. 371 (type species Mesethmoid cartilaginous; medial pro- Trigonectes strigabundus Myers, by original cesses of pelvic fin base and ischial process designation). reduced; interarcual cartilage large, attaches Rivulichthys Myers, 1927, p. 118 (type species laterally to second pharyngobranchial which Rivulus rondoni Ribeiro, by original designa- possesses a bony flange; basihyal broad an- tion). teriorly, elongate ossification posteriorly; no ETYMOLOGY: Trigonectes from the Greek tooth patches on second and third hypobran- trig, meaning angular and nektos, to swim, chials; teeth on fourth ceratobranchials; dor- referring to the wedge-shaped schools in sal and ventral hypohyal present; anterior which members have been observed. extension of anterior ceratohyal ventral to TYPES: Brazil: Goias: Porto Nacional, hypohyals; uncinate process on fourth epi- Donna Francisquinha into Tocantins. Tri- branchial articulates with that of third; first gonectes strigabundus Myers, Holotype epibranchial narrow at its base. Interhyal CAS 40701 (1). cartilaginous; three ossified basibranchials. COMPOSITION: Four species: strigabundus Vomer with posterior extension dorsal to Myers, rondoni (Myers), rogaoguae (Pear- parasphenoid. son and Myers), and luelingi (Meinken). Lacrimal narrow and twisted with obso- DIAGNOSIS: Distinguished from all other lescent sensory canal; dermosphenotic and cyprinodontiforms by an oblique mouth cleft preopercular with obsolescent sensory ca- caused by a reduction in the anterior ramus nal; pectoral girdle lowset; first postclei- of the alveolar arm of the premaxilla. thrum absent; posttemporal with ossified DEFINITION: Anal: ii, 15; Dorsal: ii, 9; Pel- lower limb; posttemporal not fused to vic: 7; Pectoral: 13-14; Caudal: 6, 16, 6; Ver- supracleithrum. tebrae: 13+15. Gill rakers on anterior arm of Vomer ossified, dentigerous; medial arm the first arch: 11. Branchiostegal rays: 6. of maxilla twisted with no pronounced dorsal Scales lateral series: 35-41. process; ventral arms with pronounced an- First pleural rib on parapophysis of second terior extension abutting rostral cartilage; vertebra; parapophysis not reduced; no pleu- outer arm narrow. 1981 PARENTI: CYPRINODONTIFORM FISHES 485

Premaxillary ascending processes flat and pteros, meaning wings and Lebias, a nomi- broad, not tapered posteriorly not overlap- nal cyprinodontiform genus, referring to the ping in the midline; rostral cartilage enlarged elongate pelvic fin rays in males. and rectangular; outer arm of premaxilla TYPES: Brazil: Santarem: Pterolebias lon- with alveolar process, not indented poste- gipinnis Garman Type. USNM 120429 (1). riorly, anterior ramus reduced; mouth cleft COMPOSITION: Five species: longipinnis oblique. Ligament extending from ventral Garman; zonatus Myers; peruensis Myers; arms of maxillaries to middle of rostral car- bockermanni Travassos; and, hoignei tilage; ethmomaxillary ligament present; me- Thomerson. niscus present between premaxilla and max- DIAGNOSIS: Distinguished from all other illa. aplocheiloids by lacking the interarcual car- Dentary not expanded medially, not ro- tilage. bust; coronoid process on dentary not over- DEFINITION: Anal: ii, 17—ii, 18; Dorsal: ii, lapping with that of articular; retroarticular 8-ii, 9; Pelvic: 7-8; Pectoral: 15-16; Caudal: not elongate. Autopalatine with straight 5, 14, 5; Vertebrae: 15+15-15+18. Gill rak- head, ventral process not elongate, not ers on anterior arm of the first arch: 9; Bran- reaching quadrate; metapterygoid present. chiostegal rays: 6; Scales lateral series: 35- Orbital rim attached; anterior nans tubu- 38. lar; supraorbital sensory pores reduced to a First pleural rib on parapophysis of second series of neuromasts preopercular, mandib- vertebra; parapophysis not reduced; no pleu- ular and lacrimal pores absent, replaced by ral rib on first hemal spine; hypural plates a series of weakly formed neuromasts. Bran- divided, upper plate never divided in two. chiostegal rays covered. Epipleural ribs not bifid. Males larger than females, often with fin Anal fin not modified into a gonopodium; extensions; pigmentation pattern consisting anal fin musculature unmodified; first proxi- of even rows of brown or reddish brown re- mal radial present; middle anal radials pres- ticulations along the side of the body, and ent. extending onto the unpaired fins. Spermatozeugmata not formed; fertiliza- No fatty predorsal ridge; caudal scaled for tion external; development annual; ovipa- at least one-third its length; swimbladder ex- rous. Eggs round. tending posteriorly to first one or two hemal One dorsal ray articulating with each of spines. the first two dorsal radials; dorsal fin on pos- DISTRIBUTION: Paraguay; Brazil; Bolivia terior third of body. (Lake Rogoaga). Autopterotic fossa normal; lateral ethmoid REMARKS: Trigonectes strigabundus is expanded medially, extending under para- represented in collections only by the holo- sphenoid; parasphenoid slightly expanded type and four paratypes. anteriorly; weakly formed supraoccipital and MATERIAL EXAMINED: T. strigabundus: epiotic processes; neural spine on first ver- the types as listed above; Paratypes: CAS tebra; first vertebra articulates with skull via 40702 (1* gill arches only/4); T. rondoni: Par- basioccipital and exoccipital condyles; su- aguay: Makthlawaiya: BMNH 1927.11.23.55- praoccipital excluded from formation of fo- 64 (l*/6). T. rogoaguae: Bolivia: Lake Ro- ramen magnum; parietals present; nasals not goagua: Holotype: CAS 43531 (1); Para- expanded medially. types: CAS 42532 (39). Mesethmoid cartilaginous; medial pro- cesses of pelvic fin base and ischial process GENUS PTEROLEBIAS GARMAN reduced; interarcual cartilage absent; second pharyngobranchial which possesses a bony Pterolebias Carman, 1895, p. 141 (type species flange; basihyal broad anteriorly, elongate Pterolebias longipinnis Garman, by original designation). ossification posteriorly; no tooth patches on second and third hypobranchials; teeth on ETYMOLOGY: Pterolebias from the Greek fourth ceratobranchials; dorsal and ventral 486 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 hypohyal present; anterior extension of an- DISTRIBUTION: Lowlands of Peru, Vene- terior ceratohyal ventral to hypohyals; un- zuela and Brazil. cinate process on fourth epibranchial artic- REMARKS: The composition of Pterolebias ulates with that of third; first epibranchial was changed by Taphorn and Thomerson narrow at its base. Interhyal cartilaginous; (1978) who removed the nominal species P. three ossified basibranchials. Vomer with maculipinnis Radda from the genus and posterior extension dorsal to parasphenoid. placed it in Rachovia. The findings of this Lacrimal narrow and twisted with obso- study agree with such a change. lescent sensory canal; dermosphenotic and MATERIAL EXAMINED: P. longipinnis: the preopercular with obsolescent sensory ca- holotype as listed above; Brazil (Aquarium nal; pectoral girdle lowset; first postclei- material): SU 4782-3 (1*73); Para: AMNH thrum absent; posttemporal with ossified 22466 (24-/2); P. peruensis: Peru: E. Loreto lower limb; posttemporal not fused to Prov: Paratypes: SU 47659 (3); P. hoignei: supracleithrum. Venezuela: Cojedes: Paratopotypes: CAS Vomer ossified, dentigerous; medial arm 27555 (4); P. zonatus: Venezuela: Guarico: of maxilla twisted with no pronounced dorsal AMNH 22467(1+/!). process; ventral arms with pronounced an- terior extension abutting rostral cartilage; GENUS RACHOVIA MYERS outer arm narrow. Premaxillary ascending processes flat and Rachovia Myers, 1927, p. 116, 119 (type species Rivulus brevis Regan, by original designation). broad, not tapered posteriorly not overlap- ping in the midline; rostral cartilage enlarged ETYMOLOGY: Rachovia named in honor of and rectangular; outer arm of premaxilla Arthur Rachow, an aquarist. with alveolar process, not indented poste- TYPES: Colombia: Rivulus brevis Regan, riorly. Ligament extending from ventral Type BMNH 1908.5.14.8 (1). arms of maxillaries to middle of rostral car- COMPOSITION: Four species: brevis (Re- tilage; ethmomaxillary ligament present; me- gan); maculipinnis (Weibezahn); humme- niscus present between premaxilla and max- lincki de Beaufort; and pyropunctata Ta- illa. phorn and Thomerson. Dentary not expanded medially, not ro- DIAGNOSIS: Distinguished from all other bust; coronoid process on dentary not over- Neotropical aplocheiloids with a vertical bar lapping with that of articular; retroarticular through the eye, and a tendency to develop not elongate. Autopalatine with straight a fatty dorsal ridge by having a low number head, ventral process not elongate, not of scales in a lateral series, generally 32 or reaching quadrate; metapterygoid present. less. Orbital rim attached; anterior naris tubu- DEFINITION: Anal: iii, 11-iii, 12; Dorsal: lar; supraorbital sensory pores reduced to a i, 9-i, 11; Pelvic: 7; Pectoral: 15-16; Caudal: series of neuromasts preopercular, mandib- 8, 15, 8; Vertebrae: 12+15-13 + 17. Gill rak- ular and lacrimal pores absent, replaced by ers on anterior arm of the first arch: 9, 10; a series of weakly formed neuromasts. Bran- Branchiostegal rays: 6; Scales lateral series: chiostegal rays covered. 27-33. Males larger than females, often with fin First pleural rib on parapophysis of second extensions; pelvic fins elongate, extending vertebra; parapophysis not reduced; no pleu- beyond first four or five anal rays in some ral rib on first hemal spine; hypural plates species (longipinnis). Throat bars not pres- divided or fused into a fan, upper plate never ent. divided in two. Epipleural ribs not bifid. No fatty predorsal ridge; caudal scaled for Anal fin not modified into a gonopodium; at least one-third its length; swimbladder ex- anal fin musculature unmodified; first proxi- tending posteriorly to first one or two hemal mal radial present; middle anal radials pres- spines. ent; anal rays of female thickened. 1981 PARENTI: CYPRINODONTIFORM FISHES 487

Spermatozeugmata not formed; fertiliza- arms of maxillaries to middle of rostral car- tion external; development annual; ovipa- tilage; ethmomaxillary ligament present; me- rous. Eggs round. niscus present between premaxilla and max- One dorsal ray articulating with each of illa. the first two dorsal radials; dorsal fin origin Dentary not expanded medially, not ro- opposite that of anal. bust; coronoid process on dentary not over- Autopterotic fossa normal; lateral ethmoid lapping with that of articular; retroarticular expanded medially, extending under para- not elongate. Autopalatine with straight sphenoid; parasphenoid slightly expanded head, ventral process not elongate, not anteriorly; weakly formed supraoccipital and reaching quadrate; metapterygoid present. epiotic processes; neural spine on first ver- Orbital rim attached; anterior naris tubu- tebra; first vertebra articulates with skull via lar; supraorbital sensory pores reduced to a basioccipital and exoccipital condyles; su- series of neuromasts in weakly formed lyre- praoccipital excluded from formation of fo- shaped pattern; preopercular, mandibular ramen magnum; parietals present; nasals not and lacrimal pores absent, replaced by a se- expanded medially. ries of weakly formed neuromasts. Branchio- Mesethmoid cartilaginous; medial pro- stegal rays covered. cesses of pelvic fin base and ischial process Males larger than females, often with fin reduced; interarcual cartilage large, attaches extensions; males often with light ventral laterally to second pharyngobranchial which margin of caudal fin; females typically drab; possesses a bony flange; basihyal broad an- vertical bar through eye; males often with teriorly, elongate ossification posteriorly; no dark blotch on the dorsal. tooth patches on second and third hypobran- Fatty predorsal ridge in older males; cau- chials; teeth on fourth ceratobranchials; dor- dal scaled for more than one third; swim- sal and ventral hypohyal present; anterior bladder extending posteriorly to first one or extension of anterior ceratohyal ventral to two hemal spines. hypohyals; uncinate process on fourth epi- DISTRIBUTION: Coastal llanos of Venezue- branchial articulates with that of third; first la and Colombia. epibranchial narrow at its base. Interhyal REMARKS: The composition of the genus cartilaginous; three ossified basibranchials. Rachovia was enlarged by Taphorn and Vomer with posterior extension dorsal to Thomerson (1978) who removed the species parasphenoid. maculipinnis from Pterolebias and placed it Lacrimal narrow and twisted with obso- in Rachovia. This study supports that deci- lescent sensory canal; dermosphenotic and sion. The genus was most recently revised preopercular with obsolescent sensory ca- by Taphorn and Thomerson (1978). nal; pectoral girdle lowset; first postclei- MATERIAL EXAMINED: R. brevis: Colom- thrum absent; posttemporal with ossified bia: the type as listed above; SU 49519 (1*/ lower limb; posttemporal not fused to 2); AMNH 22476 (\+); R. maculipinnis: Co- supracleithrum. lombia: Meta: CAS 36650 (1*710). Vomer ossified, dentigerous; medial arm of maxilla twisted with no pronounced dorsal GENUS AUSTROFUNDULUS MYERS process; ventral arms with pronounced an- terior extension abutting rostral cartilage; Austrofundulus Myers, 1932, p. 159 (type species outer arm narrow. Austrofundulus transilis Myers, by original designation). Premaxillary ascending processes flat and broad, not tapered posteriorly not overlap- ETYMOLOGY: Austrofundulus from aus- ping in the midline; rostral cartilage enlarged tro, meaning southern and Fundulus, a and rectangular; outer arm of premaxilla North American cyprinodontiform genus, in with alveolar process, not indented poste- reference to the genus as the Fundulus of the riorly. Ligament extending from ventral Southern Hemisphere. 488 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

TYPES: Venezuela: State of Guarico, Ori- first epibranchial narrow at its base. Inter- noco Drainage. Austrofundulus transilis hyal cartilaginous; three ossified basibranch- Myers, Holotype USNM 92191 (1). ials. Vomer with posterior extension dorsal COMPOSITION: Two species: transilis to parasphenoid. Myers, and limnaeus Schultz. Lacrimal narrow and twisted with obso- DIAGNOSIS: Distinguished from all other lescent sensory canal; dermosphenotic and aplocheiloids by a darkly pigmented anal pa- preopercular with obsolescent sensory ca- pilla. nal; pectoral girdle lowset; first postclei- DEFINITION: Anal: iii, 12-iii, 14; Dorsal: thrum absent; posttemporal with ossified iii, 9-iv, 10; Pelvic: 7; Pectoral: 15-16; Cau- lower limb; posttemporal not fused to dal: 8, 16, 8; Vertebrae: 13 + 17-13 + 18. Gill supraclei thrum. rakers on anterior arm of the first arch: 13- Vomer ossified, dentigerous; medial arm 15; Branchiostegal rays: 6; Scales lateral se- of maxilla twisted with no pronounced dorsal ries: 28-38. process; ventral arms with pronounced an- First pleural rib on parapophysis of second terior extension abutting rostral cartilage; vertebra; parapophysis not reduced; no pleu- outer arm narrow. ral rib on first hemal spine; hypural plates Premaxillary ascending processes flat and divided or fused into a fan, upper plate never broad, not tapered posteriorly not overlap- divided in two. Epipleural ribs not bifid. ping in the midline; rostral cartilage enlarged Anal fin not modified into a gonopodium; and rectangular; outer arm of premaxilla anal fin musculature unmodified; first proxi- with alveolar process, not indented poste- mal radial absent; middle anal radials pres- riorly. Ligament extending from ventral ent; anal rays of females thickened. arms of maxillaries to middle of rostral car- Spermatozeugmata not formed; fertiliza- tilage; ethmomaxillary ligament present; me- tion external; development annual; ovipa- niscus present between premaxilla and max- rous. Eggs round. illa. One dorsal ray articulating with each of Dentary not expanded medially, not ro- the first two dorsal radials; dorsal fin origin bust; coronoid process on dentary not over- opposite anal fin origin. lapping with that of articular; retroarticular Autopterotic fossa normal; lateral ethmoid not elongate. Autopalatine with straight expanded medially, extending under para- head, ventral process not elongate, not sphenoid; parasphenoid slightly expanded reaching quadrate; metapterygoid present. anteriorly; weakly formed supraoccipital and Orbital rim attached; anterior naris tubu- epiotic processes; enlarged neural spine on lar; supraorbital sensory pores reduced to a first vertebra; first vertebra articulates with series of neuromasts in weakly formed lyre- skull via basioccipital and exoccipital con- shaped pattern; preopercular, mandibular dyles; supraoccipital excluded from forma- and lacrimal pores absent, replaced by a se- tion of foramen magnum; parietals present; ries of weakly formed neuromasts. Branchio- nasals not expanded medially. stegal rays covered. Mesethmoid cartilaginous; medial pro- Males larger than females, often with fin cesses of pelvic fin base and ischial process extensions; females typically drab; anal pa- reduced; interarcual cartilage large, attaches pilla heavily pigmented; vertical bar through laterally to second pharyngobranchial which eye. possesses a bony flange; basihyal broad an- Fatty predorsal ridge in older males; cau- teriorly, elongate ossification posteriorly; no dal scaled for more than one-third its length; tooth patches on second and third hypobran- swimbladder extending posteriorly to first chials; teeth on fourth ceratobranchials re- one or two hemal spines. duced; dorsal and ventral hypohyal present; DISTRIBUTION: Coastal llanos of Colom- anterior extension of anterior ceratohyal ven- bia and Venezuela with greatest concentra- tral to hypohyals; uncinate process on fourth tion between the Rio Orinoco and Rio Mag- epibranchial articulates with that of third; dalena. 1981 PARENTI: CYPRINODONTIFORM FISHES 489

REMARKS: The genus has been most re- sides of the body and extending on to caudal cently revised by Taphorn and Thomerson and anal fins. (1978). No fatty predorsal ridge; caudal scaled for MATERIAL EXAMINED: A. transilis: Ven- at least one third its length. ezuela: the holotype as listed above; Falcon: DISTRIBUTION: That of the holotype. SU 33822 (1*76); AMNH 22000 (5+/5); REMARKS: The meristic data for the genus Cojedes: FMNH 85725 (3*717). A. limnaeus: were recorded from the alcohol specimen Venezuela: Zulia: CAS 39378 (2*740). (the type) and supplemented by the species description. The color pattern described is from de Souza (1979), from which informa- GENUS NEOFUNDULUS MYERS tion on osteology and distribution also was Neofundulus Myers, 1924b, p. 9 (type species obtained. Fundulus paraguayensis Eigenmann and Ken- Arambaru, Arambaru and Ringuelit (in de nedy, by original designation). Souza, 1979) suggested the species previ- ously referred to the genus, paraguayensis ETYMOLOGY: Neofundulus from the Greek and ornatipinnis, be synonymized. This is neos, meaning new and Fundulus, a North not recommended considering their scant American cyprinodontiform genus, meaning sample sizes and the poor condition of mu- a new form of that genus. seum specimens. The holotype of paraguay- TYPES: Paraguay: Arroyo Trematina. ensis is a female and that of ornatipinnis a Neofundulus paraguayensis Eigenmann and male, so from the type material alone, it is Kennedy. Holotype CAS 42533 (1). impossible to determine whether the differ- COMPOSITION: Solely the type species. ences are due to sexual dimorphism. How- DIAGNOSIS: Distinguished from other Neo- ever, data and photographs of paraguayen- tropical aplocheiloids with a vertical bar sis in de Souza (1979) indicate that in both through the eye by a derived pigmentation males and females the meristic counts are pattern consisting of rows of dark red re- lower than in the ornatipinnis holotype. ticulations on the sides and dark maroon MATERIAL EXAMINED: The holotype as margins of the caudal and anal fins. listed above. DEFINITION: Anal: iii, 12; Dorsal: i, 12; Pelvic: 7; Pectoral: 16-17; Caudal: 8, 13, 8; GENUS"NEOFUNDULUS" Gill rakers on anterior arm of the first arch: 8; Branchiostegal rays: 6; Scales lateral se- ETYMOLOGY: "Neofundulus" is used to ries: 34-38. reference the more derived species of the First pleural rib on parapophysis of second genus Neofundulus. vertebra; hypural plates divided, upper plate COMPOSITION: A single species, ornatipin- never divided in two. nis Myers. Anal fin not modified into a gonopodium. TYPES: Paraguay: Neofundulus ornatipin- Spermatozeugmata not formed; fertiliza- nis Myers, Holotype, USNM 94401 (1). tion external; development annual; ovipa- DEFINITION: Anal: iii, 12; Dorsal: ii, 13; rous. Eggs round. Pelvic: 7; Pectoral: 16-17; Caudal: 8, 13, 8; Dorsal fin origin opposite that of anal. Vertebrae: 13 + 15. Gill rakers on anterior Orbital rim attached; anterior naris tubu- arm of the first arch: 8, 9; Branchiostegal lar; supraorbital sensory pores reduced to a rays: 6; Scales lateral series: 37. series of neuromasts; preopercular, mandib- Anal fin not modified into a gonopodium. ular and lacrimal pores absent, replaced by Spermatozeugmata not formed; fertiliza- a series of weakly formed neuromasts. Bran- tion external; development annual; ovipa- chiostegal rays covered. rous. Eggs round. Males larger than females, often with fin Dorsal fin origin opposite that of anal. extensions; pigmentation pattern consisting Orbital rim attached; anterior naris tubu- of row of dark red reticulations along the lar; supraorbital sensory pores reduced to a 490 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

series of neuromasts; preopercular, mandib- on anterior arm of the first arch: 9-17; Bran- ular and lacrimal pores absent, replaced by chiostegal rays: 6; Scales lateral series: 23- a series of weakly formed neuromasts. Bran- 60. chiostegal rays covered. First pleural rib on parapophysis of second No fatty predorsal ridge; caudal scaled for vertebra; parapophysis not reduced; no pleu- at least one-third its length. ral rib on first hemal spine; hypural plates DISTRIBUTION: That of the holotype. fused to form an hypural fan. Epipleural ribs REMARKS: The species in "Neofundulus" not bifid. is more closely related to the more derived Anal fin modified into a gonopodium in a genera of Austrofundulus and Cynolebias sole species {brucei); anal fin musculature than it is to Neofundulus paraguayensis. unmodified; first proximal radial absent; mid- Since the condition of the specimen is poor, dle anal radials present; anal rays of females it is not considered appropriate to create a thickened. new genus based on the specimen to which Spermatozeugmata not formed; fertiliza- the temporary name of "Neofundulus" is tion external in all but two species {brucei applied. and melanotaenia); development annual; oviparous. Eggs round. One dorsal ray articulating with each of GENUS CYNOLEBIAS STEINDACHNER the first two dorsal radials; dorsal fin on pos- Cynolebias Steindachner, 1876, p. 172 (type terior third of body. species Cynolebias porosus Steindachner, by Autopterotic fossa normal; lateral ethmoid original designation). expanded medially, extending under para- Cynopoecilus Regan, 1912, p. 642 (type species sphenoid; parasphenoid slightly expanded Cynolebias melanotaenia Regan, by original anteriorly; weakly formed supraoccipital and designation). epiotic processes; enlarged neural spine on Leptolebias Myers, 1952, p. 140 (type species first vertebra; first vertebra articulates with Cynolebias marmoratus Ladiges, by original designation [proposed as a subgenus]). skull via basioccipital and exoccipital con- Simpsonichthys de Carvalho, 1959, p. 2 (type dyles; supraoccipital excluded from forma- species Simpsonichthys boitonei de Carvalho, tion of foramen magnum; parietals present; by original designation). nasals not expanded medially. Campellolebias Vaz-Ferreira and Sierra, 1974, p. Mesethmoid cartilaginous; medial pro- 1 (type species Campellolebias brucei Vaz- cesses of pelvic fin base and ischial process Ferreira and Sierra, by original designation). reduced; interarcual cartilage large, attaches Terranotus Taphorn and Thomerson, 1978, p. 384 laterally to second pharyngobranchial which (type species Austrofundulus dolichopterus possesses a bony flange; basihyal broad an- Weitzman and Wourms, by original designa- teriorly, elongate ossification posteriorly; no tion). tooth patches on second and third hypobran- ETYMOLOGY: Cynolebias from the Greek chials; teeth on fourth ceratobranchials re- cyno, meaning dog and Lebias, a synonym duced or absent; dorsal and ventral hypohyal of another cyprinodontiform genus, referring present; anterior extension of anterior cera- to the robust outer teeth in the lower jaw tohyal ventral to hypohyals; uncinate pro- referred to as canines and suggesting a dog. cess on fourth epibranchial articulates with COMPOSITION: Approximately 35 species, that of third; first epibranchial narrow at its referable to the above synonyms (Lazara, base. Interhyal cartilaginous; three ossified 1979). basibranchials. Vomer with posterior exten- DIAGNOSIS: Distinguished from all other sion dorsal to parasphenoid. aplocheiloids by having a preopercular canal Lacrimal narrow and twisted with obso- which is completely closed. lescent sensory canal; dermosphenotic and DEFINITION: Anal: ii, 14; Dorsal: iii, 11- preopercular with obsolescent sensory ca- ii, 16; Pelvic: 7; Pectoral: 13-14; Caudal: 7, nal; pectoral girdle lowset; first postclei- 19, 7; Vertebrae: 13 + 16-15+22. Gill rakers thrum absent; posttemporal with ossified 1981 PARENTI: CYPRINODONTIFORM FISHES 491 lower limb; posttemporal not fused to DISTRIBUTION: Cojedes, Venezuela; low- supraclei thrum. lands of Argentina, Brazil and Paraguay. Vomer ossified, dentigerous; medial arm REMARKS: Species of Cynolebias have of maxilla twisted with no pronounced dorsal been isolated in genera or subgenera on the process; ventral arms with pronounced an- basis of their apomorphous characters. Tap- terior extension abutting rostral cartilage; horn and Thomerson (1978) stated that Ter- outer arm narrow. ranotus dolichopterus perhaps was more Premaxillary ascending processes flat and closely related to Cynolebias than to either broad, not tapered posteriorly not overlap- Austrofundulus or Rachovia. ping in the midline; rostral cartilage enlarged A subgroup of species may be defined by and rectangular; outer arm of premaxilla having a higher number of dorsal and anal with alveolar process, not indented poste- fin rays in males. However, subgroups are riorly. Ligament extending from ventral not named since a revision of the genus is arms of maxillaries to middle of rostral car- judged to be a necessary prerequisite to such tilage; ethmomaxillary ligament present; me- taxonomic decisions. Since a name placed in niscus present between premaxilla and max- synonymy herein may be suitable for some illa. subgroup of Cynolebias, new names are not Dentary not expanded medially, not ro- produced in the interest of minimizing syn- bust; coronoid process on dentary not over- onyms. lapping with that of articular; retroarticular AM (1934) reviewed Cynolebias; the defi- not elongate. Autopalatine with straight nition, distribution and composition of the head, ventral process not elongate, not genus again were reviewed by Vaz-Ferreira reaching quadrate; metapterygoid present. and Sierra (1973). Orbital rim attached; anterior nans tubu- MATERIAL EXAMINED: C. belottii: Argen- lar; supraorbital sensory pores reduced to a tina: Buenos Aires: USNM 176105 (17); C. series of neuromasts in distinctive lyre- elongatus: Argentina: Villa Elisa: SU 64048 shaped pattern; preopercular, mandibular, (2*/9); C. melanotaenia: Brazil: Porto Ale- and lacrimal pores absent, replaced by a se- gre: SU 64060 (2*/8), Paranagua: Types: ries of weakly formed neuromasts. Branchio- BMNH 1909.9.5.15-22 (8); C. boitonei: stegal rays covered. Aquarium material: AMNH 38431 (3); C. Males larger than females, rarely with fin whiter. Aquarium material: AMNH 36769 extensions; no species with a spot at the an- (4*/8); C. brucei: Brazil: Santa Catarina: terior base of the dorsal fin; middle rays of ZVC,P. 2123 (1), ZVC, P. 2121 (1); C. ladi- caudal never elongate in juveniles and gesi: Brazil: Paratypes: SU 50177 (2); C. dol- adults; vertical bar through eye. ichopterus: Venezuela: Cojedes: Holotype: Fatty predorsal ridge in at least one USNM 200784 (1); Paratypes: AMNH 22718 species (dolichopterus); caudal fin not (3); FMNH: 85726 (l*/5). scaled; swimbladder extending posteriorly to first one or two hemal spines.

SUBORDER CYPRINODONTOIDEI DIAGNOSIS: Distinguished from the chial; an autopalatine with anterior arm off- aplocheiloid cyprinodontiforms by 12 set to the main axis and a weakly to strongly uniquely derived characters: two basibran- formed posterior flange; a ventral extension chials in the ventral gill arch skeleton; loss of the autopalatine forming an anterior cov- of the dorsal hypohyal; reduction of the in- ering of the quadrate; metapterygoid absent; terarcual cartilage to approximately half its premaxilla with a posterior indentation of the length with the placement of the first epi- alveolar arm to form an S-shaped distal arm; branchial closer to the second pharyngobran- a dentary expanded medially to form a ro- 492 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

bust lower jaw; loss of the first dorsal ray so vertebra; parapophysis not reduced; no pleu- that the first dorsal fin ray articulates with ral ribs on hemal spines; hypural plates the first two proximal radials; loss of the eth- slightly divided in midline. Epipleural ribs momaxillary ligament; loss of a ligament not bifid. from the interior arms of the maxilla to the Anal fin not modified into a gonopodium; middle of the rostral cartilage; and, the ab- anal fin musculature unmodified; first proxi- sence of a meniscus between the premaxilla mal radial present; middle anal radials pres- and maxilla. ent. Spermatozeugmata not formed; fertiliza- SECTION 1 tion external; development nonannual; ovip- arous. Eggs round. FAMILY PROFUNDULIDAE One dorsal ray articulating with the first HOEDEMAN AND BRONNER two dorsal radials; dorsal fin origin opposite Type Genus Profundulus Hubbs, 1924. origin of anal. DIAGNOSIS: Distinguished from all other Autopterotic fossa large; lateral ethmoid cyprinodontoids by a large autopterotic fos- not expanded medially, not reaching para- sa, and a high number of gill rakers on the sphenoid; parasphenoid not expanded ante- anterior arm of the first arch (14-23). riorly; no supraoccipital and exoccipital pro- COMPOSITION: One genus, Profundulus, cesses; neural arches of first vertebra open, with five species: candalarius Hubbs, gua- not forming a spine; first vertebra articulates temalensis (Giinther), hildebrandi Miller, la- with skull via basioccipital and exoccipital bialis (Giinther), and punctatus (Giinther). condyles; supraoccipital excluded from for- DISTRIBUTION: Both Atlantic and Pacific mation of foramen magnum; parietals pres- slopes of Middle America. Atlantic slope: ent; nasals not expanded medially. Lateral Guatemala: Isthmus of Tehuantepec to the ethmoid with reduced anterior facet. Rio Motagua; Pacific slope: Rio Papagaya, Mesethmoid ossified; medial processes of Guerrero, Mexico to Rio Lempa drainage, pelvic fin base and ischial process not re- Honduras (fig. 74). duced; interarcual cartilage reduced, at- taches laterally to second pharyngobranchial with a bony flange; basihyal long and nar- GENUS PROFUNDULUS HUBBS row; no tooth patches on second and third Profundulus Hubbs, 1924, p. 12 (type species hypobranchials; teeth on fourth ceratobran- Fundulus punctatus Giinther, by original des- chials; dorsal hypohyal absent; anterior ex- ignation). tension of anterior ceratohyal ventral to ven- Tlaloc Alvarez and Carranza, 1951, p. 40 (type tral hypohyal; uncinate process on fourth species Fundulus labialis Giinther by mono- epibranchial articulates with that of third; typy). first epibranchial wide at its base; interhyal ETYMOLOGY: Profundulus in reference to ossified; two ossified basibranchials. Vomer its presumed primitive relationship to the ge- with posterior extension ventral to parasphe- nus Fundulus. noid. TYPES: Guatemala: Chiapas: Fundulus Lacrimal flat and broad, carrying a distinct punctatus Giinther; Types, BMNH 1864. sensory canal; dermosphenotic and preoper- 1.26. 187A-C (3). cular with distinct sensory canal; pectoral COMPOSITION: AS for the family. girdle lowset; first postcleithrum present; DIAGNOSIS: AS for the family. posttemporal with ossified lower limb; post- DEFINITION: Anal: (0-iii) 11-18; Dorsal temporal not fused to supracleithrum. (i-iii) 10-16; Pelvic: 5-7; Pectoral: 15-22 Vomer ossified, edentulous; medial arm of Caudal: 17-25; Vertebrae: 15+16-16+23 maxilla twisted with no pronounced dorsal Gill rakers on anterior arm of the first arch process; ventral arms gently curved toward 14-23. Branchiostegal rays: 6; Scales lateral and abutting rostral cartilage; outer arm nar- series: 31-39. row. First pleural rib on parapophysis of second Premaxillary ascending processes flat and 1981 PARENTI: CYPRINODONTIFORM FISHES 493 broad, not tapered posteriorly and not over- DIVISION l lapping in the midline; rostral cartilage large FAMILY FUNDULIDAE JORDAN AND GILBERT and rectangular; outer arm of premaxilla with alveolar process, indented posteriorly Type Genus Fundulus Lacepede, 1803. to form S-shaped arm. No ligament from the DIAGNOSIS: Distinguished from all other ventral arms of maxillaries to middle of ros- members of the suborder Cyprinodontoidei tral cartilage; no ethmomaxillary ligament; by the anteriorly directed ventral arms of the no meniscus between premaxilla and maxil- maxillaries often with pronounced hooks; la. Unicuspid bi- or triserial outer teeth. snout pointed and drawn anteriorly; and au- Dentary expanded medially, robust; cor- topalatine projecting anterior to the lateral onoid process on dentary not overlapping ethmoid. with that of articular; retroarticular not elon- COMPOSITION: Five genera comprising gate. Autopalatine with head angled ante- approximately 40 species (Lazara, 1979): riorly; ventral process elongate reaching Fundulus Lacepede; Planetems Garman; quadrate; metapterygoid absent. Lucania Girard; Adinia Girard; and Lepto- Orbital rim free; anterior naris slightly tu- lucania Myers. bular; supraorbital sensory pores l-2a, 2b- DISTRIBUTION: Lowlands of North and 4a, 4b-7; seven preopercular pores; four Middle America, southward to Yucatan; mandibular pores; 3 lacrimal pores. Bermuda and Cuba (fig. 76). Females larger than males; males never with An extensions; middle rays of caudal GENUS PLANCTERUS GARMAN never elongate; no throat bars present. No fatty predorsal ridge; swimbladder Plancterus Garman, 1895, p. 96 (type species does not extend posteriorly beyond hemal Hydrargyra kansae Garman, by original des- spines. ignation). DISTRIBUTION: AS for the family. Fontinus Jordan and Evermann, 1896, p. 645 (type species Jordan and REMARKS: The genus is the most plesio- Gilbert, by monotypy). morphic cyprinodontoid, a conclusion that supports Hubbs (1924). ETYMOLOGY: Plancterus from the Greek Profundulus has been most recently re- planktos, meaning wandering. viewed by Miller (1955a) from which ranges COMPOSITION: Two species: zebrinus of meristic data have been supplemented. (Jordan and Gilbert) and kansae (Garman). MATERIAL EXAMINED: P. punctatus: DIAGNOSIS: Distinguished from all other Guatemala: the types as listed above; Sa- members of the family Fundulidae by the chitepequez: AMNH 32306 (3*/30); AMNH derived character of a greatly convoluted in- 24432 (1+/1). P. labialis: Guatemala: AMNH testine. 24567 (2+/2); Baja Verapaz: AMNH 22896 DEFINITION: Anal: i, 11; Dorsal: ii, 11; (240). P. guatemalensis: Guatemala: Rio La Pelvic: 6; Pectoral: 17-18; Caudal: 8, 16, 8; Conquista: AMNH 31721 (4). Vertebrae: 14+18; Gill rakers on the anterior arm of the first arch: 8. Branchiostegal rays: 6; Scales lateral series: 60-63. SECTION 2 First pleural rib on parapophysis of second DIAGNOSIS: Distinguished from Profun- vertebra; parapophysis not reduced; no pleu- dulus and the aplocheiloids by the following ral ribs on hemal spines; hypural plates fused derived characters: premaxillary ascending into hypural fan. Epipleural ribs not bifid. processes narrow or absent in adults; rostral Anal fin not modified into a gonopodium; cartilage greatly reduced or absent; inner anal fin musculature unmodified; first proxi- arms of the maxillaries not abutting rostral mal radial present; middle anal radials pres- cartilage, and remaining attached to it by ent. connective tissue; and, the lateral ethmoid Spermatozeugmata not formed; fertil- lacking or with a greatly reduced facet for ization external; development nonannual; articulation of the autopalatine. oviparous. Eggs round. 494 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

One dorsal ray articulating with the first onoid process on dentary not overlapping two dorsal radials; dorsal fin origin opposite with that of articular; retroarticular not elon- origin of anal. gate. Autopalatine with head angled ante- Autopterotic fossa moderate; lateral eth- riorly; ventral process elongate reaching moid not expanded medially, not reaching quadrate; metapterygoid absent. parasphenoid; parasphenoid not expanded Orbital rim free; anterior naris not tubular; anteriorly; normal supraoccipital and exoc- supraorbital sensory pores l-2a, 2b-4a, cipital processes; neural arches of first ver- 4b-7; seven preopercular pores; four man- tebra open, not forming a spine; first verte- dibular pores; three lacrimal pores. bra articulates with skull via basioccipital Females larger than males; males never and exoccipital condyles; supraoccipital ex- with fin extensions; middle rays of caudal not cluded from formation of foramen magnum; elongate; pigmentation pattern consisting of parietals present; nasals not expanded me- several silvery crossbars on the sides. dially. Lateral ethmoid with reduced anterior No fatty predorsal ridge; swimbladder facet. does not extend posteriorly beyond hemal Mesethmoid ossified; medial processes of spines. pelvic fin base and ischial process not re- DISTRIBUTION: Texas and Mexico: Bra- duced; interarcual cartilage reduced, at- zos, Colorado and Pecos drainages; Llano taches laterally to second pharyngobranchial Estacado, Texas; Great Plains of South Da- with a bony flange; basihyal long and nar- kota (possible introduction) to Red and Ar- row; no tooth patches on second and third kansas rivers, Texas and New Mexico, re- hypobranchials; teeth on fourth ceratobran- spectively. chials; dorsal hypohyal absent; anterior ex- REMARKS: The species of the formerly tension of anterior ceratohyal ventral to ven- ranked subgenus Plancterus are treated as tral hypohyal; uncinate process on fourth a genus in this study since they have been epibranchial articulates with that of third; excluded from the monophyletic group in- first epibranchial wide at its base; interhyal cluding the other species of Fundulus and ossified; two ossified basibranchials. Vomer the other funduline genera. Distribution data with posterior extension ventral to parasphe- are from Miller (1955b). noid. MATERIAL EXAMINED: P. kansae: Okla- Lacrimal flat and broad, carrying a distinct homa: Major Co.: AMNH 28600 (2+/12); sensory canal; dermosphenotic and preoper- Kansas: AMNH 27278 (384); P. zebrinus: cular with distinct sensory canal; pectoral AMNH 7781 (2+/2). girdle lowset; first postcleithrum present; posttemporal with ossified lower limb; post- GENUS FUNDULUS LACEPEDE temporal not fused to supracleithrum. Fundulus Lacepede, 1803, p. 37 (type species Vomer ossified, edentulous; medial arm of Cobitus heteroclita Linnaeus, by monotypy). maxilla twisted with no pronounced dorsal Hydrargira Lacepede, 1803, p. 378 (type species process; ventral arms projecting anteriorly Hydrargira swampina Lacepede, by mono- with pronounced hooks, not abutting rostral typy)- cartilage; outer arm narrow. Hydrargyra Rafinesque, 1815, p. 88 (emended Premaxillary ascending processes narrow spelling of Hydrargira Lacepede). and elongate, not tapered posteriorly and not Zygonectes Agassiz, 1853, p. 135 (type species overlapping in the midline; rostral cartilage Poecilia olivacea Storer, by original designa- reduced; outer arm of premaxilla with alveo- tion). lar process, indented posteriorly to form Micristius Gill, 1865, p. 24 (type species Fundulus S-shaped arm. No ligament from the ventral cingulatus Valenciennes, by original designa- tion). arms of maxillaries to middle of rostral car- Xenisma Jordan, 1876, p. 142 (type species Fun- tilage; no ethmomaxillary ligament; no me- dulus stellifer Jordan, by original designation). niscus between premaxilla and maxilla. Uni- Borhorys (Broussonet) Goode and Bean, 1885, p. cuspid bi- or tri-serial outer teeth. 204 (type species Cobitus heteroclita Linnaeus, Dentary expanded medially, robust; cor- by monotypy). 1981 PARENTI: CYPRINODONTIFORM FISHES 495

Gambusinus Jordan and Evermann, 1896, p. 649 taches laterally to expanded articulation (type species Fundulus rathbuni Jordan and point of second pharyngobranchial with a Meek, by original designation). bony flange; basihyal long and narrow; no Galasaceus Fowler, 1916, p. 417 (type species tooth patches on second and third hypobran- Hydragyra similis Baird and Girard, by original chials; teeth on fourth ceratobranchials; dor- designation [proposed as a substitute for Hy- sal hypohyal absent; anterior extension of drargyra as commonly used to reference H. swampina]). anterior ceratohyal ventral to ventral hypo- hyal; uncinate process on fourth epibranchial ETYMOLOGY: Fundulus from the Latin articulates with that of third; first epibran- fundus, meaning bottom referring to the hab- chial wide at its base; interhyal ossified; two itat of the type species. ossified basibranchials. Vomer with poste- COMPOSITION: Approximately 40 species rior extension ventral to parasphenoid. as listed in Lazara (1979). Lacrimal flat and broad, carrying a distinct DIAGNOSIS: Distinguished from all other sensory canal; dermosphenotic and preoper- cyprinodontiforms by the enlarged cartilagi- cular with distinct sensory canal; pectoral nous articulation point of the second pha- girdle lowset; first postcleithrum present; ryngobranchial. posttemporal with unossified lower limb; DEFINITION: Anal: 8-16; Dorsal: i, 11-15; posttemporal not fused to supracleithrum. Pelvic: 6; Pectoral: 15-20; Caudal: 6, 15, 6; Vomer ossified, edentulous; medial arm of Vertebrae: 15+16-16+22; Gill rakers on the maxilla twisted with no pronounced dorsal anterior arm of the first arch: 6-12. Bran- process; ventral arms projecting anteriorly chiostegal rays: 5-6; Scales lateral series: often with pronounced hooks, not abutting 34-48. rostral cartilage; outer arm narrow. First pleural rib on parapophysis of second Premaxillary ascending processes narrow vertebra; parapophysis not reduced; no pleu- and elongate, not tapered posteriorly and not ral ribs on hemal spines; hypural plates fused overlapping in the midline; rostral cartilage into an hypural fan. Epipleural ribs not bifid. reduced; outer arm of premaxilla with alveo- Anal fin not modified into a gonopodium; lar process, indented posteriorly to form anal fin musculature unmodified; first proxi- S-shaped arm. No ligament from the ventral mal radial present; middle anal radials pres- arms of maxillaries to middle of rostral ent. cartilage; no ethmomaxillary ligament; no Spermatozeugmata not formed; fertil- meniscus between premaxilla and maxilla. ization external; development nonannual; Unicuspid bi- or tri-serial outer teeth. oviparous. Eggs round. Dentary expanded medially, robust; cor- One dorsal ray articulating with the first onoid process on dentary not overlapping two dorsal radials; dorsal fin origin opposite with that of articular; retroarticular not elon- origin of anal. gate. Autopalatine with head angled ante- Autopterotic fossa moderate; lateral eth- riorly; ventral process elongate reaching moid not expanded medially, not reaching quadrate; metapterygoid absent. parasphenoid; parasphenoid not expanded Orbital rim free; anterior naris not tubular; anteriorly; normal supraoccipital and exoc- supraorbital sensory pores l-2a, 2b-4a, cipital processes; neural arches of first ver- 4b-7; seven preopercular pores; four man- tebra open, not forming a spine; first verte- dibular pores; 3 lacrimal pores. bra articulates with skull via basioccipital Females larger than males; males never and exoccipital condyles; supraoccipital ex- with fin extensions; middle rays of caudal cluded from formation of foramen magnum; never elongate. parietals present; nasals not expanded me- No fatty predorsal ridge; swimbladder dially. Lateral ethmoid with reduced anterior does not extend posteriorly beyond hemal facet. spines. Mesethmoid ossified; medial processes of DISTRIBUTION: AS for the family. pelvic fin base and ischial process not re- REMARKS: The genus Fundulus as consti- duced; interarcual cartilage reduced, at- tuted herein is a monophyletic group. There 496 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 has been no decision made to recognize in- GENUS LUCANIA GIRARD dividual components of the genus as separate Lucania Girard, 1859, p. 118 (type species Cy- genera or subgenera since this is a problem prinodon parvus Baird, by monotypy). for a future revisor. Chriopeops Fowler, 1916, p. 425 (type species The genus has been most recently re- Cyprinodon goodei Jordan, by original desig- viewed by Miller (1955b) and Brown (1957). nation [proposed as a subgenus]). A rediagnosis and comparison with the gen- era Empetrichthys and Crenichthys as well ETYMOLOGY: The genus Lucania, named as Profundulus was given by Uyeno and by Girard, is of no known significance. Miller (1962). Farms (1968) presented a phy- TYPES: New York: Greenport: Cyprino- logenetic analysis of the species of Profun- don parvus Baird: USNM 15280 (1). dulus and Fundulus. Wiley and Hall (1975) COMPOSITION: Three species: parva and Wiley (1977) have performed phyloge- (Baird); goodei (Jordan); and interiorus netic analyses of the species of the F. nottii- Hubbs and Miller. complex and its close relatives. Ranges for DIAGNOSIS: Distinguished from all other meristic data are supplemented by data from cyprinodontiforms by an independent block Carman (1895) and Miller (1955b). of cartilage between the interarcual cartilage MATERIAL EXAMINED: F. heteroclitus: and articulation point of the second pharyn- New York: Sheepshead Bay: AMNH 21916 gobranchial. (2+/2); Columbia Co. AMNH 26464 (3). DEFINITION: Anal: i, 8-i, 9; Dorsal: i, 11; Aquarium material AMNH 38414 (2*/2). F. Pelvic: 6; Pectoral: 12-13 Caudal: 8, 13, 8; diaphanus: New York: Grassy Pt. AMNH Vertebrae: 12+14; Gill rakers on the anterior 10209SW (2*). F. grandis: Florida: AMNH arm of the first arch: 8-9. Branchiostegal 21915SW (1+); : AMNH 3570 (9). rays: 6; scales lateral series: 27-29. F. similis: Florida: AMNH 21919SW (1+). First pleural rib on parapophysis of second F. chrysotus: Florida: AMNH 21911SW vertebra; parapophysis not reduced; no pleu- (1+); F. catenatus: Oklahoma: AMNH ral ribs on hemal spines; hypural plates fused 28589SW (2+). F. confluentus: Florida: into hypural fan. Epipleural ribs not bifid. AMNH 21913SW (2 + ). F. majalis: New Anal fin not modified into a gonopodium; York: AMNH 28526 (2+); AMNH 28532 anal fin musculature unmodified; first proxi- (11). F. olivaceus: Oklahoma: AMNH 28599 mal radial present; middle anal radials pres- (2+/11). F. parvipinnis: California: AMNH ent. 37743 (7). Zygonectes dispar: Missouri. Spermatozeugmata not formed; fertiliza- Types: USNM 120298 (4). tion external; development nonannual; ovip- FOSSILS: Fossil fundulines generally are arous. Eggs round. placed hesitantly in Fundulus (Lugaski, One dorsal ray articulating with the first 1977; Miller, 1945). The following genus has two dorsal radials; dorsal fin origin opposite also been named for such fossils. origin of anal. Autopterotic fossa moderate; lateral eth- moid not expanded medially, not reaching GENUS PARAFUNDULUS EASTMAN parasphenoid; parasphenoid not expanded anteriorly; normal supraoccipital and exoc- Parafundulus Eastman, 1917, p. 291 (type species cipital processes; neural arches of first ver- Parafundulus nevadensis Eastman, by original tebra open, not forming a spine; first verte- designation). (Pleistocene of Nevada) bra articulates with skull via basioccipital Farris (1968) identified some specimens of and exoccipital condyles; supraoccipital ex- this genus as members of Profundulus; how- cluded from formation of foramen magnum; ever, since he presented no unique charac- parietals present; nasals not expanded me- ters to define either genus, such an identifi- dially. Lateral ethmoid with reduced anterior cation is suspect. facet. 1981 PARENTI: CYPRINODONTIFORM FISHES 497

Mesethmoid ossified; medial processes of with a dark lateral band extending through pelvic fin base and ischial process not re- the eye to base of the caudal fin. duced; interarcual cartilage reduced, at- No fatty predorsal ridge; swimbladder taches laterally by a block of cartilage to sec- does not extend beyond hemal spines. ond pharyngobranchial which possesses a DISTRIBUTION: Coastal lowlands of north- bony flange; basihyal long and narrow; no eastern North America; Cuatro Cienegas Ba- tooth patches on second and third hypobran- sin, Mexico; introduced in California and chials; teeth on fourth ceratobranchials; dor- Texas. sal hypohyal absent; anterior extension of REMARKS: The genus Lucania has often anterior ceratohyal ventral to ventral hypo- been referred to as a presumed close relative hyal; uncinate process on fourth epibranchial of Cubanichthys of Cuba, and possibly of articulates with that of third; first epibran- Chriopeoides of Jamaica (Hubbs and Miller, chial wide at its base; interhyal ossified; two 1965; Rosen, 1976). However, it has been ossified basibranchials. Vomer with poste- found that the two island forms are not rior extension ventral to parasphenoid. closely related to the genus Lucania nor to Lacrimal flat and broad, carrying a distinct any other funduline genus but are rather sensory canal; dermosphenotic and preoper- more derived cyprinodonts related to an as- cular with distinct sensory canal; pectoral semblage including the Old World and New girdle lowset; first postcleithrum present; World cyprinodontines and Orestias. posttemporal with ossified lower limb; post- MATERIAL EXAMINED: L. parva: New temporal not fused to supracleithrum. York: the type as listed above; AMNH Vomer ossified, edentulous; medial arm of 35922SW (4*/1071); AMNH 27462SW (5+); maxilla twisted with no pronounced dorsal L. goodei: Florida: Punta Gorda: ANSP process; ventral arms projecting anteriorly, 91218 (12); Little Springs: AMNH 22082SW with pronounced hooks, not abutting rostral (8+); St. John's R.: Types USNM 23505 (2). cartilage; outer arm narrow. Premaxillary ascending processes narrow GENUS LEPTOLUCANIA MYERS and elongate, not tapered posteriorly and not Leptolucania Myers, 1924b, p. 8 (type species overlapping in the midline; rostral cartilage Heterandria ommata Jordan, by original des- reduced; outer arm of premaxilla with alve- ignation [proposed as a subgenus]). olar process, indented posteriorly to form S-shaped arm. No ligament from the ventral ETYMOLOGY: Leptolucania from the Greek arms of maxillaries to middle of rostral car- lepto, meaning thin or elongate, and Lucan- tilage; no ethmomaxillary ligament; no me- ia, another funduline genus, referring to the niscus between premaxilla and maxilla. Uni- elongate body form. cuspid uni- bi- or tri-serial outer teeth. TYPES: Florida: Heterandria ommata Jor- Dentary expanded medially, robust; cor- dan, Type, USNM 25331 (1). onoid process on dentary not overlapping COMPOSITION: Solely the type species. with that of articular; retroarticular not elon- DIAGNOSIS: Distinguished from all other gate. Autopalatine with head angled ante- cyprinodontiform genera by three branchios- tegal rays and a unique color pattern con- riorly; ventral process elongate reaching sisting of an ocellus on the caudal peduncle quadrate; metapterygoid absent. and one at midbody. Distinguished from all Orbital rim free; anterior naris not tubular: other fundulid genera in lacking the first post- supraorbital canal continuous between pores cleithrum. 1 through 7; seven preopercular pores; four DEFINITION: Anal: ii, 8; Dorsal: i, 6, Pel- mandibular pores; three lacrimal pores. vic: 6; Pectoral: 13-14; Caudal: 6, 10, 6; Ver- Females larger than males; all fins round- tebrae: 12+15; Gill rakers on the anterior ed, never extensions; parva males with an arm of the first arch: 6-7. Branchiostegal ocellus at anterior base of dorsal fin; goodei rays: 6; Scales lateral series: 25-26. 498 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

First pleural rib on parapophysis of second and elongate, not tapered posteriorly and not vertebra; parapophysis not reduced; no pleu- overlapping in the midline; rostral cartilage ral ribs on hemal spines; hypural plates fused reduced; outer arm of premaxilla with alve- into hypural fan. Epipleural ribs not bifid. olar process, indented posteriorly to form Anal fin not modified into a gonopodium; S-shaped arm. No ligament from the ventral anal fin musculature unmodified; first proxi- arms of maxillaries to middle of rostral car- mal radial present; middle anal radials pres- tilage; no ethmomaxillary ligament; no me- ent. niscus between premaxilla and maxilla. Uni- Spermatozeugmata not formed; fertiliza- cuspid bi- or triserial outer teeth. tion external; development nonannual; ovip- Dentary expanded medially, robust; cor- arous. Eggs round. onoid process on dentary not overlapping One dorsal ray articulating with the first with that of articular; retroarticular not elon- two dorsal radials; dorsal fin origin opposite gate. Autopalatine with head angled ante- origin of anal. riorly; ventral process elongate reaching Autopterotic fossa moderate; lateral eth- quadrate; metapterygoid absent. moid not expanded medially, not reaching Orbital rim free; anterior naris not tubular; parasphenoid; parasphenoid not expanded supraorbital sensory pores 6-7; seven pre- anteriorly; no supraoccipital and exoccipital opercular pores; four mandibular pores; processes; neural arches of first vertebra three lacrimal pores. open, not forming a spine; first vertebra ar- Females larger than males; males never ticulates with skull via basioccipital and ex- with fin extensions. Pigmentation pattern occipital condyles; supraoccipital excluded consisting of a dark blotch at midbody and from formation of foramen magnum; pari- a midcaudal ocellus. etals present; nasals not expanded medially. No fatty predorsal ridge; swimbladder Lateral ethmoid with reduced anterior facet. does not extend posteriorly beyond hemal Mesethmoid ossified; medial processes of spines. pelvic fin base and ischial process not re- DISTRIBUTION: Florida and Georgia. duced; interarcual cartilage reduced, at- REMARKS: The genus Leptolucania has taches laterally to second pharyngobranchial been confused with a diminutive poeciliid with a bony flange; basihyal long and nar- Heterandria formosa, also of Florida, the row; no tooth patches on second and third genus in which ommata was placed when hypobranchials; teeth on fourth ceratobran- described. chials; dorsal hypohyal absent; anterior ex- MATERIAL EXAMINED: Florida: the types tension of anterior ceratohyal ventral to ven- as listed above; Leon Co.: AMNH 20383 tral hypohyal; uncinate process on fourth (6+/4*/118). epibranchial articulates with that of third; first epibranchial wide at its base; interhyal GENUS ADINIA GIRARD ossified; two ossified basibranchials. Vomer with posterior extension ventral to parasphe- Adinia Girard, 1859, p. 117 (type species Adinia noid. multifasciata Girard, by original designation). Lacrimal flat and broad, carrying a distinct ETYMOLOGY: The genus Adinia, named sensory canal; dermosphenotic and preoper- by Girard, is of no known significance. cular with distinct sensory canal; pectoral TYPES: Texas: Indianola: Adinia multifas- girdle lowset; first postcleithrum absent; ciata Girard, Cotypes, ANSP 7291, 7292. posttemporal with ossified lower limb; post- COMPOSITION: Two species: multifasciata temporal not fused to supracleithrum. Girard and xenica (Jordan and Gilbert). Vomer ossified, edentulous; medial arm of DIAGNOSIS: Distinguished from all other maxilla twisted with no pronounced dorsal cyprinodontiforms by the first pleural rib on process; ventral arms projecting anteriorly the parapophysis of the first vertebra, rather with pronounced hooks, not abutting rostral than the second and a distinct quadrangular cartilage; outer arm narrow. body form. Premaxillary ascending processes narrow DEFINITION: Anal: ii, 8; Dorsal: ii, 7; Pel- 1981 PARENTI: CYPRINODONTIFORM FISHES 499

vie: 6; Pectoral: 17; Caudal: 8, 14, 8; Verte- process; ventral arms projecting anteriorly brae: 12+15. Gill rakers on the anterior arm with pronounced hooks, not abutting rostral of the first arch: 9-10. Branchiostegal rays: cartilage; outer arm narrow. 5; Scales lateral series: 24-25. Premaxillary ascending processes narrow First pleural rib on parapophysis of first and elongate, not tapered posteriorly and not vertebra; parapophysis not reduced; no pleu- overlapping in the midline; rostral cartilage ral ribs on hemal spines; hypural plates fused reduced; outer arm of premaxilla with alve- into hypural fan. Epipleural ribs not bifid. olar process, indented posteriorly to form Anal fin not modified into a gonopodium; S-shaped arm; No ligament from the ventral anal fin musculature unmodified; first proxi- arms of maxillaries to middle of rostral car- mal radial present; middle anal radials pres- tilage; no ethmomaxillary ligament; no me- ent. niscus between premaxilla and maxilla. Uni- Spermatozeugmata not formed; fertiliza- cuspid bi- or tri-serial outer teeth. tion external; development nonannual; ovip- Dentary expanded medially, robust; cor- arous. Eggs round. onoid process on dentary not overlapping One dorsal ray articulating with the first with that of articular; retroarticular not elon- two dorsal radials; dorsal fin origin opposite gate. Autopalatine with head angled ante- origin of anal. riorly; ventral process elongate reaching Autopterotic fossa moderate; lateral eth- quadrate; metapterygoid absent. moid not expanded medially, not reaching Orbital rim free; anterior naris not tubular; parasphenoid; parasphenoid not expanded supraorbital sensory pores 6-7; seven pre- anteriorly; no supraoccipital and exoccipital opercular pores; four mandibular pores; processes; neural arches of first vertebra three lacrimal pores. open, not forming a spine; first vertebra ar- Females larger than males; males never ticulates with skull via basioccipital and ex- with fin extensions; body form quadrangular; occipital condyles; supraoccipital excluded series of silvery crossbars on sides of body. from formation of foramen magnum; pari- No fatty predorsal ridge; swimbladder etals present; nasals not expanded medially. does not extend posteriorly beyond hemal Lateral ethmoid with reduced anterior facet. spines. Mesethmoid ossified; medial processes of DISTRIBUTION: Brackish water from Flor- pelvic fin base and ischial process not re- ida to Texas. duced; interarcual cartilage reduced, at- MATERIAL EXAMINED: A. multifasciata: taches laterally to second pharyngobranchial Louisiana: USNM 124795 (2*733); A. xenica: with a bony flange; basihyal long and nar- Alabama: AMNH 35673 (2*/19). row; no tooth patches on second and third hypobranchials; teeth on fourth ceratobran- DIVISION 2 chials; dorsal hypohyal absent; anterior ex- DIAGNOSIS: Distinguished from all other tension of anterior ceratohyal ventral to ven- cyprinodontiforms by two derived charac- tral hypohyal; uncinate process on fourth ters: maxilla with a straight proximal arm; epibranchial articulates with that of third; and an enlarged dorsal process of the max- first epibranchial wide at its base; interhyal illa. ossified; two ossified basibranchials. Vomer with posterior extension ventral to parasphe- SEPT l noid. FAMILY VALENCIIDAE, NEW FAMILY Lacrimal flat and broad, carrying a distinct sensory canal; dermosphenotic and preoper- Type Genus Valencia Myers 1928b. cular with distinct sensory canal; pectoral DIAGNOSIS: Distinguished from all other girdle lowset; first postcleithrum present; cyprinodontiforms by an elongate and atten- posttemporal with ossified lower limb; post- uate dorsal process of the maxilla. temporal not fused to supracleithrum. COMPOSITION: One genus, Valencia, and Vomer ossified, edentulous; medial arm of two nominal species: hispanica (Cuvier and maxilla twisted with no pronounced dorsal Valenciennes) and letourneauxi (Sauvage). 500 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

DISTRIBUTION: Fresh waters of southeast- tension of anterior ceratohyal ventral to ven- ern Spain, Italy, and Corfu. tral hypohyal; uncinate process on fourth epibranchial articulates with that of third; GENUS VALENCIA MYERS first epibranchial wide at its base; interhyal ossified; two ossified basibranchials. Vomer Valencia Myers, 1928b, p. 8 (type species Hy- with posterior extension ventral to parasphe- drargyra hispanica Cuvier and Valenciennes, noid. by original designation). Lacrimal flat and broad, carrying a distinct ETYMOLOGY: Valencia to connote the oc- sensory canal; dermosphenotic and preoper- currence of the genus in Valencia, Spain. cular with distinct sensory canal; pectoral COMPOSITION: AS for the family. girdle lowset; first postcleithrum present; DIAGNOSIS: As for the family. posttemporal with ossified lower limb; post- DEFINITION: Anal: i, 13; Dorsal: i, 9; Pel- temporal not fused to supracleithrum. vic: 6; Pectoral: 15; Caudal: 8, 19, 8; Verte- Vomer ossified, edentulous; medial arm of brae: 13 + 16; Gill rakers on the anterior arm maxilla straight with elongate dorsal process; of the first arch: 12-13; Branchiostegal rays: ventral arms not abutting rostral cartilage; 6; Scales lateral series: 28. outer arm narrow. First pleural rib on parapophysis of second Premaxillary ascending processes narrow vertebra; parapophysis not reduced; no pleu- and elongate, not tapered posteriorly and not ral ribs on hemal spines; hypural plates fused overlapping in the midline; rostral cartilage into hypural fan. Epipleural ribs not bifid. reduced; outer arm of premaxilla with alve- Anal fin not modified into a gonopodium; olar process, indented posteriorly to form anal fin musculature unmodified; first proxi- S-shaped arm. No ligament from the ventral mal radial present; middle anal radials pres- arms of maxillaries to middle of rostral car- ent. tilage; no ethmomaxillary ligament; no me- Spermatozeugmata not formed; fertiliza- niscus between premaxilla and maxilla. Uni- tion external; development nonannual; ovip- cuspid bi- or tri-serial outer teeth. arous. Eggs round. Dentary expanded medially, robust; cor- One dorsal ray articulating with the first onoid process on dentary not overlapping two dorsal radials; dorsal fin origin opposite with that of articular; retroarticular not elon- origin of anal. gate. Autopalatine with head angled ante- Autopterotic fossa moderate; lateral eth- riorly; ventral process elongate reaching moid not expanded medially, not reaching quadrate; metapterygoid absent. parasphenoid; parasphenoid not expanded Orbital rim free; anterior naris not tubular; anteriorly; weakly formed supraoccipital and supraorbital sensory pores l-2a, 2b-4a, exoccipital processes; neural arches of first 4b-7; seven preopercular pores; four man- vertebra open, not forming a spine; first ver- dibular pores; three lacrimal pores. tebra articulates with skull via basioccipital Females larger than males; males with en- and exoccipital condyles; supraoccipital ex- larged dorsal and anal fins; pigmentation pat- cluded from formation of foramen magnum; tern consisting of a series of medium brown parietals present; nasals not expanded me- reticulations on the unpaired fins. dially. Lateral ethmoid with reduced anterior No fatty predorsal ridge; swimbladder facet. does not extend posteriorly beyond hemal Mesethmoid ossified; medial processes of spines. pelvic fin base and ischial process not re- DISTRIBUTION: AS for the family. duced; interarcual cartilage reduced, at- REMARKS: The genus Valencia has been taches laterally to second pharyngobranchial confused with both Fundulus and Aphanius, with a bony flange; basihyal long and nar- the former on the basis of shared primitive row; no tooth patches on second and third characters and the latter on its distribution. hypobranchials; teeth on fourth ceratobran- Valencia is not closely related to the other chials; dorsal hypohyal absent; anterior ex- Old World Laurasian cyprinodonts, and its 1981 PARENTI: CYPRINODONTIFORM FISHES 501 similarities to the fundulines have been SUBFAMILY ANABLEPINAE GARMAN shown to be derived for a much larger group. MATERIAL EXAMINED: Spain: V. hispan- DIAGNOSIS: Distinguished from all other ica: AMNH 38401 2*/5; Italy: ANSP 7254 cyprinodontiforms by having thickened and (1); Corfu: Fundulus letourneauxi: Syntypes elongated anal rays in males which are twist- BMNH 1880.9.13:1-6(4). ed around each other and covered by a fleshy tube, tubular sperm duct, gonopodium offset SEPT 2 either to the left or to the right, proximal anal radials enlarged and derived pigmentation DIAGNOSIS: Distinguished from all other pattern. cyprinodontiforms by three derived charac- COMPOSITION: TWO genera, Anableps and ters: ascending processes of the premaxillar- Jenynsia with species as listed for the fami- ies short and narrow; dorsal processes of the ly. maxillaries rounded or greatly reduced; and DISTRIBUTION: AS for the family. nasals expanded medially. GENUS ANABLEPS (GRONOW) SCOPOLI SUPERFAMILY POECILIOIDEA Anableps (Gronow) Scopoli, 1777, p. 450 (type DIAGNOSIS: Distinguished from other cy- species Linnaeus, by orig- prinodontiform fishes by the following de- inal designation). rived characters: maxilla with an expanded Peltatetrops Fowler, 1931, p. 396 (type species distal arm; parasphenoid with expanded an- Anableps microlepis Miiller and Troschel, by terior arm; dorsal processes of the maxillar- original designation [proposed as a subgenus]). ies with a distinct lateral indentation; primi- tively an elongate retroarticular and a pouch ETYMOLOGY: Anableps from the Greek created by scales surrounding the urogenital ana, meaning great or enlarged and bleps, opening of females. meaning eye referring to the enlarged orbits of this genus. COMPOSITION: Three species as listed for FAMILY ANABLEPIDAE CARMAN the family. Type Genus Anableps (Gronow) Scopoli, DIAGNOSIS: Distinguished from all other 1777. cyprinodontiform fishes by enlarged supraor- DIAGNOSIS: Distinguished from all other bital processes of the frontals to accommo- fishes of the order Cyprinodontiformes by date enlarged eyes divided horizontally to robust epiotic and supraoccipital processes; effect vision above and below the water; ver- and an outer row of tricuspidate teeth, at tebrae 45 or more; posterior section or su- least in juveniles or embryos, and several in- praorbital sensory pore pattern represented ner rows of unicuspidate or tricuspidate jaw by pores 6 and 7 only; head scales arranged teeth. in random pattern; gonopodium formed prin- COMPOSITION: Three genera: Anableps cipally from anal rays 3 through 9; upper jaw (Gronow) Scopoli, three species: dowi Gill, blunt with a reduced premaxillary ascending anableps Linnaeus, and microlepis Miiller processes; dumbbell-shaped rostral carti- and Troschel; Oxyzygonectes Fowler, with lage; and from Jenynsia and Oxyzygonectes one species: dowi (Giinther); and Jenynsia by possessing a tubular anterior naris; and Giinther, with four species: lineata (Giin- enlarged pectoral fins with 20 rays or more. ther), eigenmanni (Haseman), maculata DEFINITION: Anal: (ii) 9-10; Dorsal: 7-10; (Regan) and pygogramma (Boulenger). Pelvic: 6; Pectoral: 20-26; Caudal: 6, 16, 6; DISTRIBUTION: Southern Mexico to Nica- Vertebrae: 45-54; Gill rakers on the anterior ragua; northern coast of South America from arm of the first arch: 21-30; Branchiostegal Venezuela to Para, Brazil; southern South rays: 6; Scales lateral series: 50-96. America (southern Brazil, Argentina, Uru- First pleural rib on parapophysis of second guay) (fig. 78). vertebra; parapophysis not reduced; no pleu- 502 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 ral ribs on hemal spines; hypural plates fused limb; posttemporal not fused to supraclei- into hypural fan. Epipleural ribs not bifid. thrum. Anal fin modified into a gonopodium Vomer ossified, edentulous; medial arm of formed principally from anal rays 3 through maxilla straight with pronounced dorsal pro- 9; anal fin musculature unmodified; proximal cess indented laterally; ventral arms not anal radials expanded and positioned ante- abutting rostral cartilage; outer arm spatu- riorly; middle anal radials present; sexual late. dextrality: males with gonopodium offset to Premaxillary ascending processes narrow the left or the right; females with one or two and reduced, not tapered posteriorly and not scales covering left or right side of urogenital overlapping in the midline; dumbbell shaped opening; gonopodium scaled; tubular sperm rostral cartilage; outer arm of premaxilla duct opening at tip of gonopodium. with alveolar process, indented posteriorly Spermatozeugmata in some species; fertil- to form S-shaped arm. No ligament from the ization internal; viviparous; intrafollicular ventral arms of maxillaries to middle of ros- gestation. tral cartilage; no ethmomaxillary ligament; One dorsal ray articulating with the first no meniscus between premaxilla and maxil- two dorsal radials; dorsal fin set back on pos- la. Weakly tricuspid outer teeth. terior third of body. Dentary expanded medially, robust; cor- Autopterotic fossa moderate; lateral eth- onoid process on dentary not overlapping moid not expanded medially, not reaching with that of articular; retroarticular elongate. parasphenoid; parasphenoid expanded ante- Autopalatine with head angled anteriorly; riorly; enlarged supraoccipital and exoccipi- ventral process elongate reaching quadrate; tal processes; neural arches of first vertebra metapterygoid absent. open, not forming a spine; first vertebra ar- Orbital rim free; anterior naris tubular; su- ticulates with skull via basioccipital and ex- praorbital sensory pores 1-2, 3-4a, 6-7; sev- occipital condyles; supraoccipital excluded en preopercular pores; four mandibular from formation of foramen magnum; pari- pores; two to four lacrimal pores. etals present; nasals reduced; supraorbital Females larger than males; males never processes of frontal expanded to accommo- with fin extensions; lower rays of caudal date enlarged eyes possessing divided reti- drawn into a lobe in one species (microle- nas. pis); pigmentation pattern consisting of lon- Mesethmoid ossified; medial processes of gitudinal stripes of varying number and pelvic fin base and ischial process not re- width. duced; interarcual cartilage reduced, at- No fatty predorsal ridge; swimbladder taches laterally to second pharyngobranchial does not extend posteriorly beyond hemal toothplate with a bony flange; basihyal long spines. and narrow; tooth patches on second and DISTRIBUTION: Central America, both At- third hypobranchials; teeth on fourth cera- lantic and Pacific slopes; northern South tobranchials; dorsal hypohyal absent; ante- America. rior extension of anterior ceratohyal ventral REMARKS: The genus has been most re- to ventral hypohyal; uncinate process on cently reviewed by Miller (1979); Grier, fourth epibranchial articulates with that of Burns and Flores (MS) described the forma- third; first epibranchial wide at its base; in- terhyal ossified; two ossified basibranchials. tion of spermatozeugmata in A. dowi. Vomer with posterior extension ventral to Ranges of meristic characters are supple- parasphenoid. mented by data from Miller (1979). Lacrimal flat and broad, carrying a distinct MATERIAL EXAMINED: A. dowi: Guate- sensory canal; dermosphenotic and preoper- mala: Santa Rosa: AMNH 32449 (30); cular with distinct sensory canal; pectoral AMNH 24402 (12); Jutiapa: AMNH 31529 girdle lowset; first postcleithrum present or (25); No data: AMNH 20830 (2*); AMNH absent; posttemporal with ossified lower 38412SW(1+). 1981 PARENTI: CYPRINODONTIFORM FISHES 503

GENUS JENYNSIA GUNTHER tal processes; neural arches of first vertebra open, not forming a spine; first vertebra ar- Jenynsia Gunther, 1866, p. 331 (type species Le- ticulates with skull via basioccipital and ex- bias lineata Jenyns, by original designation). Fitzroyia Gunther, 1866, p. 307 (type species Le- occipital condyles; supraoccipital excluded bias multidentata Jenyns, by original designa- from formation of foramen magnum; pari- tion). etals present; nasals expanded medially. Mesethmoid ossified; medial processes of ETYMOLOGY: Jenynsia, a patronym for pelvic fin base and ischial process not re- Jenyns, author of the "Fish Section" of the duced; interarcual cartilage reduced, at- Voyage of the Beagle. Fitzroyia and Jen- taches laterally to second pharyngobranchial ynsia were proposed in the same work by with a bony flange; basihyal long and nar- Gunther, and although Fitzroyia has page row; no tooth patches on second and third priority, the name Jenynsia should be used hypobranchials; teeth on fourth ceratobran- since it was accepted by earlier writers who chials; dorsal hypohyal absent; anterior ex- regarded the two genera as synonyms. tension of anterior ceratohyal ventral to ven- TYPES: South America (Voyage of the tral hypohyal; uncinate process on fourth Beagle): Lebias lineata Jenyns, Types: epibranchial articulates with that of third; BMNH 1917.7.14.20-23. first epibranchial wide at its base; interhyal COMPOSITION: Four species as listed for ossified; two ossified basibranchials. Vomer the family. with posterior extension ventral to parasphe- DIAGNOSIS: Distinguished from all other noid. cyprinodontiforms by tricuspidate outer jaw Lacrimal flat and broad, carrying a distinct teeth in adults; an unsealed gonopodium sensory canal; dermosphenotic and preoper- formed principally from rays 3, 6, and 7; and cular with distinct sensory canal; pectoral an enlarged sixth middle anal radial. girdle lowset; first postcleithrum present; DEFINITION: Anal: ii, 8; Dorsal: i, 8; Pel- posttemporal with ossified lower limb; post- vic: 6; Pectoral: 15; Caudal: 8, 17, 8; Verte- temporal not fused to supracleithrum. brae: 13+16. Gill rakers on the anterior arm Vomer ossified, edentulous; medial arm of of the first arch: 10-11. Branchiostegal rays: maxilla straight with pronounced dorsal pro- 5; Scales lateral series: 25-28. cess indented laterally; ventral arms narrow, First pleural rib on parapophysis of second not abutting rostral cartilage; outer spatu- vertebra; parapophysis not reduced; no pleu- late. ral ribs on hemal spines; hypural plates fused Premaxillary ascending processes narrow into hypural fan. Epipleural ribs not bifid. and reduced, not tapered posteriorly and not Anal fin modified into a gonopodium overlapping in the midline; rostral cartilage formed principally from anal rays 3, 6, and reduced; outer arm of premaxilla with alveo- 7; anal musculature unmodified; first proxi- lar process, indented posteriorly to form mal radial present; middle anal radials pres- S-shaped arm; No ligament from the ventral ent, sixth enlarged in both males and fe- arms of maxillaries to middle of rostral car- males; gonopodium unsealed; tubular sperm tilage; no ethmomaxillary ligament; no me- duct opening at tip of gonopodium. niscus between premaxilla and maxilla. Tri- Spermatozeugmata not formed; fertiliza- tion internal; viviparous; intraovarian gesta- cuspid bi- or tri-serial outer teeth. tion. Dentary expanded medially, robust; cor- One dorsal ray articulating with the first onoid process on dentary not overlapping two dorsal radials; dorsal fin origin opposite with that of articular; retroarticular elongate. origin of anal. Autopalatine with head angled anteriorly; Autopterotic fossa moderate; lateral eth- ventral process elongate reaching quadrate; moid not expanded medially, not reaching metapterygoid absent. parasphenoid; parasphenoid expanded ante Orbital rim free; anterior naris not tubular; riorly; enlarged supraoccipital and exoccipi- supraorbital sensory pores l-2a, 2b-4a, 504 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

4b-7; seven preopercular pores; four man- Greek oxy, meaning pointed and Zygo- dibular pores; three lacrimal pores. nectes, a presumed close relative in refer- Females larger than males; males never ence to the pointed snout. Miller (1966) has with fin extensions. Pigmentation pattern shown that the specific modifier should be consisting of dark interrupted stripes. dowi, not dovii or dovi since the name is a No fatty predorsal ridge; swimbladder patronym for Capt. J. M. Dow. does not extend posteriorly beyond hemal TYPES: Costa Rica: Punta Arena: Haplo- spines. chilus dovii Giinther, Types BMNH. DISTRIBUTION: Southern South America: 1865.7.20 29-30 (2). lowlands of Brazil, Paraguay, Uruguay, and COMPOSITION: Solely the type. Argentina. DIAGNOSIS: AS for the subfamily above. REMARKS: The genus Jenynsia has been DEFINITION: Anal: ii, 10; Dorsal: i, 6; Pel- confused in past classifications with the cy- vic: 6; Pectoral: 16-17; Caudal: 7, 15, 7; Ver- prinodontines since both possess tricuspi- tebrae: 13 + 15; Gill rakers on the anterior date outer teeth. The variability in the size arm of the first arch: 15-16. Branchiostegal and shape of these teeth in Jenynsia indi- rays: 6; Scales lateral series: 29-30. cates that dental morphological data should First pleura! rib on parapophysis of second be used with caution in a phylogenetic study, vertebra; parapophysis not reduced; no pleu- especially when there are ontogenetic ra! ribs on hemal spines; hypural plates fused changes in tooth structure as in Anableps into hypural fan; Epipleural ribs not bifid. and Oxyzygonectes. Anal fin not modified into a gonopodium; MATERIAL EXAMINED: J. lineata: the anal fin musculature unmodified; first proxi- types as listed above; Brazil: CAS 40706 (4*/ mal radial present; middle anal radials pres- 62); AMNH 12938 (1+/2); J. pygogramma: ent. Cordova: Types: BMNH 1902. 5. 22. 72-81 Spermatozeugmata not formed; fertiliza- (4): /. maculata: Argentina: Salta: Types: tion external; development nonannual; BMNH 1906. 5.31. 62-71 (9); sp.: Brazil: oviparous. Eggs round. Santa Catarina, Rio Pique: USNM uncat. One dorsal ray articulating with the first (Field no. SW 9-22-77 -3) (6); Lago Perto do two dorsal radials; dorsal fin origin posterior mar: USNM uncat. (Field no. SW 9-19-77-1) to origin of anal. (7). Autopterotic fossa moderate; lateral eth- SUBFAMILY OXYZYGONECTINAE, moid slightly expanded medially, not reach- NEW SUBFAMILY ing parasphenoid; parasphenoid expanded anteriorly; enlarged supraoccipital and ex- TYPE GENUS: Oxyzygonectes Fowler, occipital processes; neural arches of first 1916 vertebra open, not forming a spine; first ver- DIAGNOSIS: Distinguished from all other tebra articulates with skull via basioccipital cyprinodontiforms by having an inner series and exoccipital condyles; supraoccipital ex- of tricuspidate jaw teeth arranged in four or cluded from formation of foramen magnum; five closely packed rows; a series of scales parietals present; nasals expanded medially. overlying the preorbital canal; males with Mesethmoid ossified; medial processes of anal papilla and enlarged premaxillary as- pelvic fin base and ischial process not re- cending processes. duced; interarcual cartilage reduced, at- COMPOSITION: Solely the type. taches laterally to second pharyngobranchial DISTRIBUTION: Pacific coast of Costa toothplate with a bony flange; basihyal long Rica. and narrow; tooth patches on second and GENUS OXYZYGONECTES FOWLER third hypobranchials; teeth on fourth cera- Oxyzygonectes Fowler, 1916, p. 425 (type species tobranchials; dorsal hypohyal absent; ante- Haplochilus dovii Giinther, by original desig- rior extension of anterior ceratohyal ventral nation [proposed as a subgenus]). to ventral hypohyal; uncinate process on ETYMOLOGY: Oxyzygonectes, after the fourth epibranchial articulates with that of 1981 PARENTI: CYPRINODONTIFORM FISHES 505 third; first epibranchial wide at its base; in- No fatty predorsal ridge; swimbladder terhyal ossified; two ossified basibranchials. does not extend posteriorly beyond hemal Vomer with posterior extension ventral to spines. parasphenoid. DISTRIBUTION: AS for the subfamily. Lacrimal flat and broad, carrying a distinct REMARKS: The genus Oxyzygonectes had sensory canal; dermosphenotic and preoper- previously been placed in the subfamily Fun- cular with distinct sensory canal; pectoral dulinae. girdle lowset; first postcleithrum present; MATERIAL EXAMINED: Costa Rica: the posttemporal with ossified lower limb; post- syntypes, as listed above; Golfito: AMNH temporal not fused to supracleithrum. 17657 (l*/l+/8); AMNH 37733 (2*/25). Vomer ossified, edentulous; medial arm of maxilla straight with pronounced dorsal pro- FAMILY POECILIIDAE CARMAN cess indented laterally; ventral arms narrow, Type Genus Poecilia Bloch and Schnei- not abutting rostral cartilage; outer arm spat- der, 1801. ulate. DIAGNOSIS: Distinguished from all other Premaxillary ascending processes narrow cyprinodontiforms by the following derived and reduced, not tapered posteriorly and not characters: pectoral fins set high on the sides overlapping in the midline; rostral cartilage effected by the dorsal placement of the ra- absent; outer arm of premaxilla with alveolar dials; a hyoid bar with no ventral extension process, indented posteriorly to form of the anterior ceratohyal accompanied by S-shaped arm; no ligament from the ventral the ventral hypohyal typically forming a arms of maxillaries to middle of rostral car- bony cap over its anterior facet; pleural ribs tilage; no ethmomaxillary ligament; no me- on the first several hemal arches; the anterior niscus between premaxilla and maxilla. placement of the pelvic fins and their inferred Weakly tricuspid outer row of outer teeth. anterior migration during growth; and re- Several inner rows of closely packed tricus- cessed supraorbital sensory pores 2b through pid teeth. 4a. Dentary expanded medially, robust; cor- COMPOSITION: Three subfamilies: Poecili- onoid process on dentary not overlapping inae Gar man, Fluviphylacinae Roberts, and with that of articular; retroarticular elongate. Aplocheilichthyinae Myers, as defined be- Autopalatine with head angled anteriorly; low. ventral process elongate reaching quadrate; DISTRIBUTION: (fig. 80) North and Middle metapterygoid absent. America, Caribbean, South America to Orbital rim free; anterior nans not tubular; southern Uruguay; Africa (Congo Basin and supraorbital sensory pores l-2a, 2b-4a, the African rift lakes); Dar es Salaam and 4b-7; seven preopercular pores; four man- Madagascar. dibular pores; three lacrimal pores. Females larger than males; middle rays of SUBFAMILY POECILIINAE CARMAN caudal never elongate; no throat bars pres- DIAGNOSIS: Distinguished from all other ent. cyprinodontiform fishes by the gonopodium Males and females dark brown overall; of males formed principally from anal rays males with spotted dorsal and anal fins, and 3, 4, and 5, enlarged hemal arches providing several weak crossbars just anterior to the its support; expanded inclinators of the anal caudal peduncle; females' unpaired fins pale. fin to form a fan-shaped mass; pelvic fins of In life, males with a blue tinge on the dorsal males with curved rays inferred to function surface, and both males and females with a during copulation. shiny white spot on dorsal aspect of orbit. COMPOSITION: Approximately 200 species Males with urogenital papilla offset to the left in 16 genera. or right in preservation; females with scaled, DISTRIBUTION: North America through fleshy pouch covering first several anal fin Central America, the Caribbean, through rays. South America to Uruguay. 506 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

DEFINITION: Anal: iii, 6; Dorsal: 4-14; Vomer with posterior extension ventral to Pelvic: 6; Pectoral: 9-16; Caudal: 8,11, 8-8, parasphenoid. 15, 8; Vertebrae: 11+26. Gill rakers on the Lacrimal flat and broad, carrying a distinct anterior arm of the first arch: 6-27. Bran- sensory canal; dermosphenotic and preoper- chiostegal rays: 5; Scales lateral series: 30- cular with distinct sensory canal; pectoral 34. girdle highset; first postcleithrum absent; First pleural rib on parapophysis of second posttemporal with ossified lower limb; post- vertebra; parapophysis not reduced; no pleu- temporal not fused to supracleithrum. ral ribs on hemal spines; hypural plates fused Vomer ossified, edentulous; medial arm of into hypural fan; Epipleural ribs not bifid. maxilla straight with pronounced dorsal pro- Anal fin modified into a gonopodium cess indented laterally; ventral arms not formed principally from anal rays 3, 4, and abutting rostral cartilage; outer arm narrow. 5; hemal arches expanded and projecting an- Premaxillary ascending processes narrow teriorly; proximal radials enlarged; anal fin and reduced, not tapered posteriorly and not musculature expanded into fan-shaped mass; overlapping in the midline; rostral cartilage first proximal radial present; middle anal ra- reduced or absent; outer arm of premaxilla dials present. with alveolar process, indented posteriorly Spermatozeugmata formed; fertilization to form S-shaped arm. No ligament from the internal; development nonannual; ovipa- ventral arms of maxillaries to middle of ros- rious, ovoviviparous, viviparous. Gestation tral cartilage; no ethmomaxillary ligament; intra- or extra-follicular; superfetation oc- no meniscus between premaxilla and maxil- curs in some. la. Unicuspid bi- or tri-serial outer teeth. One dorsal ray articulating with the first Dentary expanded medially, robust; cor- two dorsal radials; dorsal fin origin opposite onoid process on dentary not overlapping origin of anal. with that of articular; retroarticular not elon- Autopterotic fossa moderate; lateral eth- gate. Autopalatine with head angled ante- moid not expanded medially, not reaching riorly; ventral process elongate reaching parasphenoid; parasphenoid not expanded quadrate; metapterygoid absent. anteriorly; weakly formed supraoccipital and Orbital rim free; anterior nans not tubular; exoccipital processes; neural arches of first supraorbital sensory pores l-2a, recessed vertebra open and often expanded, not form- 2b-4a, 4b-7; seven preopercular pores; four ing a spine; first vertebra articulates with mandibular pores; three or four lacrimal skull via basioccipital condyles; exoccipital pores. condyles absent; supraoccipital excluded Females larger than males; males often from formation of foramen magnum; pari- with fin extensions; males drab to elaborate- etals present or absent; nasals expanded me- ly pigmented; females typically drab. dially. No fatty predorsal ridge; swimbladder Mesethmoid ossified; medial processes of does not extend posteriorly beyond hemal pelvic fin base and ischial process not re- spines. duced; interarcual cartilage absent or re- REMARKS: The subfamily Poeciliinae as duced attaching laterally to second pharyn- the term is used here is equivalent to the fam- gobranchial with a bony flange; basihyal long ily Poeciliidae of previous authors. and narrow; tooth patches on second and Ranges for meristic characters are for third hypobranchials present or absent; teeth specimens examined. on fourth ceratobranchials; dorsal hypohyal MATERIAL EXAMINED: Jomeurus graci- absent; anterior extension of anterior cera- lis: Aquarium material: AMNH 22685 (16+); tohyal reduced or absent; ventral hypohyal Poecilia vivipara: Brazil: ; expanded; uncinate process on fourth epi- AMNH 20708 (5+/20); Belonesox belizanus branchial articulates with that of third or ab- maxillosus: Aquarium material: AMNH sent; first epibranchial narrow at its base; in- 27493SW (5+/5); JT/pAopAorKj Ae/Zen: terhyal ossified; two ossified basibranchials. Aquarium material: AMNH 38409 (3*/3). 1981 PARENTI: CYPRINODONTIFORM FISHES 507

FOSSILS: Upper Tertiary fossil poeciliids mal radial present; middle anal radials pres- are known from Brazil. ent; urogenital opening of female covered with a pocket of scales. SUBFAMILY APLOCHEILICHTHYINAE MYERS Spermatozeugmata not formed; fertiliza- tion external; development nonannual; ovip- Type Genus Aplocheilichthys Bleeker. arous. Eggs round. DIAGNOSIS: Distinguished from other poe- One dorsal ray articulating with the first cilioids by the possession of a cartilaginous two dorsal radials; dorsal fin origin opposite mesethmoid, and lateral ethmoid expanded origin of anal. toward the parasphenoid. Autopterotic fossa moderate; lateral eth- COMPOSITION: Eight genera in over 100 moid expanded medially, reaching parasphe- species: Aplocheilichthys Bleeker, "Aplo- noid; parasphenoid expanded anteriorly; no cheilichthys,," Lamprichthys Regan, Proca- supraoccipital and exoccipital processes; topus Boulenger, Pantanodon Myers, Cy- neural arches of first vertebra open, not nopanchax Ahl, Plataplochilus Ahl and forming a spine; first vertebra articulates Hypsopanchax Myers. with skull via basioccipital and exoccipital DISTRIBUTION: Savanna and forest low- condyles; supraoccipital excluded from for- land regions of central and east Africa south mation of foramen magnum; parietals ab- of the Sahara, the Rift lakes, and Madagas- sent; nasals expanded medially. car. Mesethmoid unossified; medial processes GENUS APLOCHEILICHTHYS BLEEKER of pelvic fin base and ischial process not re- duced; interarcual cartilage reduced, at- Aplocheilichthys Bleeker, 1863, p. 116 (type taches laterally to second pharyngobranchial species Poecilia spliauchena Dumeril, by with a bony flange; basihyal long and nar- monotypy). row; no tooth patches on second and third Haplochilichthys Regan, 1911, p. 323 (emendation hypobranchials; teeth on fourth ceratobran- of spelling of Aplocheilichthys Bleeker). chials; dorsal hypohyal absent; anterior ex- ETYMOLOGY: Aplocheilichthys after tension of anterior ceratohyal reduced; ven- Aplocheilus McClelland, another cyprino- tral hypohyal enlarged; uncinate process on dontiform genus, and ichthys, referring to a fourth epibranchial articulates with that of presumed close relationship between the two third; first epibranchial wide at its base; in- genera. terhyal ossified; two ossified basibranchials. TYPES: Poecilia spilauchena Dumeril, Vomer with posterior extension ventral to BMNH. parasphenoid. DIAGNOSIS: Distinguished from other poe- Lacrimal flat and broad, carrying a distinct cilioid fishes by possessing a first postcleith- sensory canal; dermosphenotic and preoper- rum, and from other procatopines by having cular with distinct sensory canal; pectoral an unexpanded swimbladder, and an anal fin girdle highset; first postcleithrum present; with less than 14 rays. posttemporal with ossified lower limb; post- DEFINITION: Anal: ii, 10; Dorsal: i, 9; Pel- temporal not fused to supracleithrum. vic: 6; Pectoral: 13-15; Caudal: 6, 14, 6; Ver- Vomer ossified, edentulous; medial arm of tebrae: 12+13; Gill rakers on the anterior maxilla straight with pronounced dorsal pro- arm of the first arch: 7-8. Branchiostegal cess indented laterally; ventral arms not rays: 5; Scales lateral series: 28-30. abutting rostal cartilage; outer arm expand- First pleural rib on parapophysis of second ed. vertebra; parapophysis not reduced; often a Premaxillary ascending processes narrow pleural rib on hemal spines; hypural plates and reduced, not tapered posteriorly and not fused into hypural fan. Epipleural ribs not overlapping in the midline; rostral cartilage bifid. reduced or absent; outer arm of premaxilla Anal fin not modified into a gonopodium; with alveolar process, indented posteriorly anal fin musculature unmodified; first proxi- to form S-shaped arm. No ligament from the 508 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 ventral arms of maxillaries to middle of ros- MATERIAL EXAMINED: A. spilauchena: tral cartilage; no ethmomaxillary ligament; the types as listed above; Ghana: SU 63440 no meniscus between premaxilla and maxil- (2*125); SU 64629 (6); A. baudoni: West Af- la. Unicuspid bi- or tri-serial outer teeth. rica: AMNH 20936 Paratype (1): A. pumilus: Dentary expanded medially, robust; cor- Tanganyika: AMNH 8274 (1); Aquarium ma- onoid process on dentary not overlapping terial: AMNH 27465 (4+); A. macrophthal- with that of articular; retroarticular elongate. mus: Niger Delta: BMNH 1977. 12.6: 1-10 Autopalatine with head angled anteriorly; (1*/10); A. myersi: no data: ANSP (2 and ventral process elongate reaching quadrate; young). metapterygoid absent. Orbital rim free; anterior nans not tubular; GENUS LAMPRICHTHYS REGAN supraorbital sensory pores l-2a, recessed 2b-4a, 4b-7; seven preopercular pores; four Lamprichthys Regan, 1911, p. 325 (type species mandibular pores; three lacrimal pores. Haplochilus tanganicanus Boulenger, by orig- Males larger than females; males often inal designation). with extension of pectoral and caudal fins; Mohanga Boulenger, 1911, p. 261 (type species body straw-colored overall. Haplochilus tanganicanus Boulenger, by orig- inal designation). No fatty predorsal ridge; swimbladder does not extend posteriorly beyond hemal ETYMOLOGY: Lamprichthys from the Lat- spines. in lampas, meaning bright, and ichthys, re- DISTRIBUTION: Savanna and forest regions ferring to the large, bright eyes typical of of central and eastern Africa. procatopines. REMARKS: The genus Aplocheilichthys TYPES: Lake Tanganyika: Haplochilus cannot be defined as a monophyletic group. tanganicanus Boulenger, Type: BMNH The more derived members of the genus are 1898.9.9.82 (1). placed in the genus group "Aplocheilich- COMPOSITION: Solely the type species. thys." This decision is made here since a DIAGNOSIS: Distinguished from other poe- revision of the entire genus is necessary to cilioids by having up to 41 vertebrae, ctenoid define its major monophyletic subgroups. scales, posttemporal with a ligamentous low- The following named taxa are treated as sub- er limb, and lyre-shaped caudal fin. genera in this analysis: DEFINITION: Anal: ii, 26; Dorsal: iii, 13; Pelvic: 6; Pectoral: 17; Caudal: 8, 18, 8; Ver- SUBGENUS MICROPANCHAX MYERS tebrae: 14+26-14+27. Gill rakers on the an- Micropanchax Myers, 1924a, p. 42 (type species terior arm of the first arch: 27. Branchioste- Haplochilus schoelleri Boulenger, by original gal rays: 5; Scales lateral series: 30-33. designation). First pleural rib on parapophysis of second vertebra; parapophysis not reduced; one or SUBGENUS LACUSTRICOLA MYERS two pleural ribs on hemal spines; hypural plates fused into hypural fan. Epipleural ribs Lacustricola Myers, 1924b, p. 43 (type species not bifid. Haplochilus pumilus Boulenger, by original designation [proposed as a subgenusj). Anal fin not modified into a gonopodium; anal fin musculature unmodified; first proxi- SUBGENUS POROPANCHAX CLAUSEN mal radial present; middle anal radials pres- ent; urogenital opening of female covered Poropanchax Clausen, 1967, p. 12 (type species, with pocket of scales. Aplocheilichthys macrophthalmus Meinken, Spermatozeugmata not formed; fertiliza- by original designation). tion external; development nonannual; oviparous. Eggs round. SUBGENUS CONGOPANCHAX POLL One dorsal ray articulating with the first Congopanchax Poll, 1971, p. 303 (type species two dorsal radials; dorsal fin origin opposite Aplocheilichthys myersi Poll, by original des- origin of anal. ignation). Autopterotic fossa moderate; lateral eth- 1981 PARENTI: CYPRINODONTIFORM FISHES 509 moid expanded medially, reaching parasphe- supraorbital sensory pores l-2a, recessed noid; parasphenoid expanded anteriorly; 2b-4a, 4b-7; seven preopercular pores, four weakly formed supraoccipital and exoccipi- mandibular pores; three lacrimal pores. tal processes; neural arches of first vertebra Males larger than females; males and fe- open, not forming a spine; first vertebra ar- males with upper and lower rays of caudal ticulates with skull via basioccipital and ex- extended; body in life with light blue tinge; occipital condyles; supraoccipital excluded color brownish in alcohol. from formation of foramen magnum; pari- No fatty predorsal ridge; swimbladder ex- etals absent; nasals expanded medially. tends posteriorly beyond 12 hemal spines. Mesethmoid cartilaginous; medial pro- DISTRIBUTION: Lake Tanganyika. cesses of pelvic fin base and ischial process MATERIAL EXAMINED: Lake Tanganyika: not reduced; interarcual cartilage reduced, the types as listed above; FMNH 62958 (1*/ attaches laterally to second pharyngobran- 4); FMNH 62959 (l*/4); AMNH 11732 (1); chial with a bony flange; basihyal long and AMNH 11728(2). narrow; no tooth patches on second and third hypobranchials; teeth on fourth cera- tobranchials; dorsal hypohyal absent; ante- GENUS PANTANODON MYERS rior extension of anterior ceratohyal re- Pantanodon Myers, 1955, p. 7 (type species Pan- duced; ventral hypohyal expanded; uncinate tanodon podoxys Myers, by original designa- process on fourth epibranchial articulates tion). with that of third; first epibranchial wide at its base; interhyal ossified; two ossified ba- ETYMOLOGY: Pantanodon from the Greek sibranchials. Vomer with posterior extension pantos, meaning all and anodon, meaning ventral to parasphenoid. without teeth referring to the absence of Lacrimal flat and broad, carrying a distinct teeth in the jaws in the type species. sensory canal; dermosphenotic and preoper- TYPES: Dar es Salaam: Pantanodon po- cular with distinct sensory canal; pectoral doxys Myers; Holotype: SU 50194 (1). girdle highset; first postcleithrum absent; COMPOSITION: TWO species: podoxys posttemporal with unossified lower limb; Myers and, madagascariensis (Arnoult). posttemporal not fused to supracleithrum. DIAGNOSIS: Distinguished from all other Vomer ossified, edentulous; medial arm of cyprinodontiforms by the greatly enlarged maxilla straight with pronounced dorsal pro- second pharyngobranchial toothplate; three cess indented laterally; ventral arms not pelvic spines in males; the absence of hy- abutting rostral cartilage; outer arm expand- pobranchials; and from other aplocheilich- ed. thyids by the presence of tricuspid inner Premaxillary ascending processes narrow teeth; the absence of exoccipital condyles; and reduced, not tapered posteriorly and not the application of the neurapophyses of the overlapping in the midline; rostral cartilage first vertebra to the skull; and an increase in reduced or absent; outer arm of premaxilla the number of gill rakers on the first arch to with alveolar process, indented posteriorly 45. to form S-shaped arm. No ligament from the DEFINITION: Anal: iii, 14; Dorsal: i, 8; Pel- ventral arms of maxillaries to middle of ros- vic: iii, 3-6; Pectoral: 9; Caudal: 5, 16, 5; tral cartilage; no ethmomaxillary ligament; Vertebrae: 14+16; Gill rakers on the anterior no mensicus between premaxilla and maxil- arm of the first arch: 45; Branchiostegal rays: la. Unicuspid bi- or tri-serial outer teeth. 5; Scales lateral series: 29-31. Dentary expanded medially, robust; cor- First pleural rib on parapophysis of second onoid process on dentary not overlapping vertebra; parapophysis not reduced; no pleu- with that of articular; retroarticular not elon- ral ribs on hemal spines; hypural plates fused gate. Autopalatine with head angled ante- into hypural fan. Epipleural ribs not bifid. riorly; ventral process elongate reaching Anal fin not modified into a gonopodium; quadrate; metapterygoid absent. anal fin musculature unmodified; first proxi- Orbital rim free; anterior nans not tubular; mal radial present; middle anal radials pres- 510 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

ent; urogenital opening of female covered no meniscus between premaxilla and maxil- with pocket of scales. la. Unicuspid bi- or tri-serial or no outer Spermatozeugmata not formed; fertiliza- teeth. tion external; development nonannual; ovip- Dentary expanded medially, robust; cor- arous. Eggs round. onoid process on dentary not overlapping One dorsal ray articulating with the first with that of articular; retroarticular not elon- two dorsal radials; dorsal fin origin opposite gate. Autopalatine with head angled ante- origin of anal. riorly; ventral process elongate reaching Autopterotic fossa moderate; lateral eth- quadrate; metapterygoid absent. moid not expanded medially, not reaching Orbital rim free; anterior naris not tubular; parasphenoid; parasphenoid not expanded supraorbital sensory pores weakly formed; anteriorly; no supraoccipital and exoccipital preopercular canal represented by an open processes; neural arches of first vertebra groove. open, not forming a spine; first vertebra ar- Males larger than females; males with no ticulates with skull via basioccipital con- fin extensions; caudal fin truncate; body gen- dyles; exoccipital condyles absent; supra- erally straw-colored. occipital excluded from formation of foramen No fatty predorsal ridge; swimbladder ex- magnum; parietals absent; nasals not ex- tends posteriorly beyond five hemal spines. panded medially. DISTRIBUTION: Tanzania, Mozambique, Mesethmoid unossified; medial processes and Madagascar. of pelvic fin base and ischial process not re- REMARKS: The genus Pantanodon was duced; interarcual cartilage absent; second placed in its own subfamily, the Pantanodon- pharyngobranchial toothplate greatly en- tinae, by Whitehead (1962) on the basis of an larged; basihyal long and narrow; no tooth absence of teeth in the jaws and other de- patches on second and third hypobranchials; rived specializations of podoxys, the only teeth on fourth ceratobranchials; dorsal hy- species known at that time. pohyal absent; anterior extension of anterior MATERIAL EXAMINED: P. podoxys: Dar ceratohyal reduced; ventral hypohyal ex- es Salaam: the holotype as listed above; panded; uncinate process on fourth epibran- paratype: SU 50195 (1); SU 61761; BMNH chial; first three epibranchials absent; inter- 1962.4.4.: 1-12 (2). P. madagascariensis: hyal ossified; two ossified basibranchials. Paratypes: AMNH 20526 (l*/4). Vomer with posterior extension ventral to parasphenoid. Lacrimal narrow, carrying a distinct sen- GENUS "APLOCHEILICHTHYS" sory canal; dermosphenotic and preopercu- ETYMOLOGY: "Aplocheilichthys" is used lar with distinct sensory canal; pectoral gir- to reference the more derived species of the dle highset; first postcleithrum absent; genus Aplocheilichthys. posttemporal with ossified lower limb; post- COMPOSITION: The more derived species temporal not fused to supracleithrum. of the genus Aplocheilichthys of which john- Unossified vomer; medial arm of maxilla stoni may be considered typical. straight, reduced dorsal process; ventral DIAGNOSIS: Distinguished from other arms narrow and straight, no abutting rostral members of the genus Aplocheilichthys and cartilage; outer arm narrow. resembling the more derived procatopines by Premaxillary ascending processes narrow lacking the interarcual cartilage, possessing and reduced, not tapered posteriorly and not an unossified vomer, an anal fin of 14 rays or overlapping in the midline; rostral cartilage more, swimbladder extending beyond the reduced or absent; outer arm of premaxilla first three hemal arches, and having a robust with alveolar process, indented posteriorly lower jaw. to form S-shaped arm. No ligament from the DEFINITION: Anal: iii, 11; Dorsal: ii, 13; ventral arms of maxillaries to middle of ros- Pelvic: 5; Pectoral: 12; Caudal: 6, 9, 6; Ver- tral cartilage; no ethmomaxillary ligament; tebrae: 14+16. Gill rakers on the anterior 1981 PARENTI: CYPRINODONTIFORM FISHES 511 arm of the first arch: 8. Branchiostegal rays: cess indented laterally; ventral arms not 5; Scales lateral series: 31. abutting rostral cartilage; outer arm expand- First pleural rib on parapophysis of second ed. vertebra; parapophysis not reduced; one or Premaxillary ascending processes narrow two pleural ribs on hemal spines; hypural and reduced, not tapered posteriorly and not plates fused into hypural fan. Epipleural ribs overlapping in the midline; rostral cartilage not bifid. reduced or absent; outer arm of premaxilla Anal fin not modified into a gonopodium; with alveolar process, indented posteriorly anal fin musculature unmodified; first proxi- to form S-shaped arm; No ligament from the mal radial present; middle anal radials pres- ventral arms of maxillaries to middle of ros- ent; urogenital opening of female covered tral cartilage; no ethmomaxillary ligament; with pocket of scales. no meniscus between premaxilla and maxil- Spermatozeugmata not formed; fertiliza- la. Unicuspid bi- or tri-serial outer teeth. tion external; development nonannual; ovip- Dentary expanded medially, extremely ro- arous. Eggs round. bust; coronoid process on dentary not over- One dorsal ray articulating with the first lapping with that of articular; retroarticular two dorsal radials; dorsal fin origin opposite not elongate. Autopalatine with head angled origin of anal. anteriorly; ventral process elongate reaching Autopterotic fossa moderate; lateral eth- quadrate; metapterygoid absent. moid expanded medially, reaching parasphe- Orbital rim free; anterior nans not tubular; noid; parasphenoid expanded anteriorly; supraorbital sensory pores l-2a, recessed weakly formed supraoccipital and exoccipi- 2b-4a, 4b-7; seven preopercular pores; four tal processes; neural arches of first vertebra mandibular pores; three lacrimal pores. open, not forming a spine; first vertebra ar- Males larger than females; males and fe- ticulates with skull via basioccipital and ex- males with truncate caudal fin; body straw- occipital condyles; supraoccipital excluded colored overall. from formation of foramen magnum; pari- No fatty predorsal ridge; swimbladder ex- etals absent; nasals expanded medially. tends posteriorly beyond four hemal spines. Mesethmoid ossified; medial processes of DISTRIBUTION: See Aplocheilichthys pelvic fin base and ischial process not re- above. duced; interarcual cartilage absent; second REMARKS: "Aplocheilichthys" is not for- pharyngobranchial possesses a bony flange; mally named since its limits cannot be readi- basihyal long and narrow; no tooth patches ly determined as the genus Aplocheilichthys on second and third hypobranchials; teeth on is large (containing over 100 species) and is fourth ceratobranchials; dorsal hypohyal ab- polyphyletic. sent; anterior extension of anterior cerato- MATERIAL EXAMINED: A. johnstoni: No hyal reduced; ventral hypohyal expanded, data: ANSP 54348-57 (2*/10). uncinate process on fourth epibranchial ar- ticulates with that of third; first epibranchial wide at its base; interhyal ossified; two ossi- GENUS PROCATOPUS BOULENGER fied basibranchials. Vomer with posterior Procatopus Boulenger, 1904b, p. 20 (type species extension ventral to parasphenoid. Procatopus nototaenia Boulenger, by original Lacrimal flat and broad, carrying a distinct designation). sensory canal; dermosphenotic and preoper- Andreasenius Clausen, 1959, p. 264 (type species Procatopus aberrans Ahl, by original desig- cular with distinct sensory canal; pectoral nation [proposed as a subgenus]). girdle highset; first postcleithrum absent; Hylopanchax Poll and Lambert, 1965, p. 623 (type posttemporal with ossified lower limb; post- species Hypsopanchax silvestris Poll and Lam- temporal not fused to supracleithrum. bert, by original designation). Vomer ossified, edentulous; medial arm of ETYMOLOGY: Procatopus from the Greek maxilla straight with pronounced dorsal pro- pro, meaning if front of, catos, meaning in- 512 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

ferior or ventral, and op, meaning opening, first epibranchial wide at its base; interhyal referring to the anterior position of the pel- ossified; two ossified basibranchials. Vomer vies and anus. with posterior extension ventral to parasphe- TYPES: Cameroon, Lobi River: Procato- noid. pus nototaenia Boulenger, Types: BMNH Lacrimal flat and broad, carrying a distinct 1904.7.1.141-160. sensory canal; dermosphenotic and preoper- DIAGNOSIS: Distinguished from all other cular with distinct sensory canal; pectoral cyprinodontiforms by having several rays girdle highset; first postcleithrum absent; free from the branchiostegal membrane and posttemporal with ossified lower limb; post- extending posteriorly in males. temporal not fused to supracleithrum. DEFINITION: Anal: (i) 14-17; Dorsal: 7-8; Unossified vomer; medial arm of maxilla Pelvic: 6; Pectoral: 13; Caudal: 8,11,8; Ver- pronounced dorsal process indented lateral- tebrae: 12+17; Gill rakers on the anterior ly; ventral arms narrow and straight, not arm of the first arch: 12-13. Branchiostegal abutting rostral cartilage; outer arm reduced. rays: 5; Scales lateral series: 24-28. Premaxillary ascending processes narrow First pleural rib on parapophysis of second and reduced, not tapered posteriorly and not vertebra; parapophysis not reduced; no pleu- overlapping in the midline; rostral cartilage ral ribs on hemal spines; hypural plates fused reduced or absent; outer arm of premaxilla into hypural fan; Epipleural ribs not bifid. with alveolar process, indented posteriorly Anal fin not modified into a gonopodium; to form S-shaped arm. No ligament from the anal fin musculature unmodified; first proxi- ventral arms of maxillaries to middle of ros- mal radial present; middle anal radials pres- tral cartilage; no ethmomaxillary ligament; ent; urogenital opening of female covered no meniscus between premaxilla and maxil- with pocket of scales. la. Unicuspid bi- or tri-serial outer teeth. Spermatozeugmata not formed; fertiliza- Dentary expanded medially, extremely ro- tion external; development nonannual; ovip- bust; coronoid process on dentary not over- arous. Eggs round. lapping with that of articular; retroarticular One dorsal ray articulating with the first not elongate. Autopalatine with head angled two dorsal radials; dorsal fin origin opposite anteriorly; ventral process elongate reaching origin of anal. quadrate; metapterygoid absent. Autopterotic fossa moderate; lateral eth- Orbital rim free; anterior naris not tubular; moid expanded medially, reaching parasphe- supraorbital sensory pores l-2a, recessed noid; parasphenoid expanded anteriorly; no 2b-4a, 4b-7; seven preopercular pores; four supraoccipital and exoccipital processes; mandibular pores; three lacrimal pores. neural arches of first vertebra open, not Males larger than females; often with cau- forming a spine; first vertebra articulates dal fin extensions; one or two branchiostegal with skull via basioccipital and exoccipital rays free from opercular membrane and pro- condyles; supraoccipital excluded from for- jecting posteriorly. mation of foramen magnum; parietals ab- No fatty predorsal ridge; swimbladder sent; nasals expanded medially. does not extend posteriorly beyond hemal Mesethmoid unossified; medial processes spines. of pelvic fin base and ischial process not re- DISTRIBUTION: Congo Basin. duced; interarcual cartilage absent; second pharyngobranchial toothplate possesses a REMARKS: The anterior displacement of bony flange; basihyal long and narrow; no the pelvic fins has been demonstrated in the tooth patches on second and third hypobran- genus Procatopus (Trewavas, 1974). chials; teeth on fourth ceratobranchials; dor- MATERIAL EXAMINED: P. nototaenia: sal hypohyal absent; anterior extension of Cameroon: the types as listed above; SU anterior ceratohyal reduced; ventral hypo- 47714 (l*/8). P. gracilis: Aquarium material: hyal expanded; uncinate process on fourth AMNH 38406 (2*/2). P. silvestris: Zaire epibranchial articulates with that of third; (Congo): UMMZ 188727 (l*/4). 1981 PARENTI: CYPRINODONTIFORM FISHES 513

GENUS CYNOPANCHAX MYERS GENUS HYPSOPANCHAX MYERS Cynopanchax Ahl, 1928, p. 115 (type species Hypsopanchax Myers, 1924a, p. 41 (type species Haplochilichthys bukobanus Ahl, by original Hypsopanchax platysternus Myers, by original designation). designation). ETYMOLOGY: Cynopanchax from the Platypanchax Ahl, 1928, p. 116 (type species Greek cyno, meaning dog and Panchax, a Haplochilus modestus Pappenheim, by original commonly used generic reference for all Old designation). World cyprinodonts, referring to the pres- ETYMOLOGY: Hypsopanchax, from the ence of an outer row of enlarged, recurved Greek hypso meaning deep and Panchax, a teeth. common generic reference for Old World cy- COMPOSITION: Solely the type species. prinodonts, referring to the deep body. TYPES: Zaire (Congo): Hypsopanchax GENUS PLATAPLOCHILUS AHL platysternus Myers, Type: AMNH 6299 (1). COMPOSITION: Approximately 15 species Plataplochilus Ahl, 1928, p. 116 (type species as listed in Lazara (1979). Haplochilichthys ngaensis Ahl, by original DIAGNOSIS: Distinguished from all other designation). aplocheilichthyids by being deep-bodied as ETYMOLOGY: The genus Plataplochilus a result of expanded pleural ribs. from the Greek platys meaning flat and Hap- DEFINITION: Anal: (ii) 15-19; Dorsal: 12- lochilus, a general reference for African cy- 16; Pelvic: 6; Pectoral: 16; Caudal: 8, 16, 8; prinodonts, referring to the species as a cy- Vertebrae: 12+16; Gill rakers on the anterior prinodont with a laterally compressed body arm of the first arch: 11-12. Branchiostegal form. rays: 5; Scales lateral series: 29-30. COMPOSITION: Solely for the type species. First pleural rib on parapophysis of second REMARKS: Specimens of Cynopanchax vertebra; parapophysis not reduced; often a bukobanus and Plataplochilus ngaensis pleural rib on hemal spines; pleural ribs ex- have not been examined. Ahl (1928) separat- panded ventrally; hypural plates fused into ed the species from other Aplocheilichthys hypural fan. Epipleural ribs not bifid. species placing them in monotypic genera on Anal fin not modified into a gonopodium; the basis of dental characteristics. Cynopan- anal fin musculature unmodified; first proxi- chax possesses an outer row of greatly en- mal radial present; middle anal radials pres- larged teeth in the jaws, the lateral pair sug- ent; urogenital opening of female covered gesting canines and therefore the with pocket of scales. characteristic on which the genus was Spermatozeugmata not formed; fertiliza- named. However, from the description, bu- tion external; development nonannual; ovip- kobanus seems to differ little from primitive arous. Eggs round. Aplocheilichthys species. The nonmonophy- One dorsal ray articulating with the first letic nature of the latter genus does not help two dorsal radials; dorsal fin origin opposite in placing Cynopanchax or Plataplochilus in origin of anal. synonymy of any recognized genus. Lambert Autopterotic fossa moderate; lateral eth- and Clausen (1967) redefined Plataplochilus moid expanded medially, reaching parasphe- using a series of primitive and derived char- noid; parasphenoid expanded anteriorly; no acters none of which was unique. Therefore, supraoccipital and exoccipital processes; the genera are listed here as available generic neural arches of first vertebra open, not categories whose membership may be ex- forming a spine; first vertebra articulates panded after a revision of the genus Aplo- with skull via basioccipital and exoccipital cheilichthys. Both Myers (1938) and Clausen condyles; supraoccipital excluded from for- (1967) were unable to treat these genera in mation of foramen magnum; parietals ab- their reviews of the procatopines because of sent; nasals expanded medially. the unavailability of material. Mesethmoid unossifed; processes of pel- 514 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 vie fin base and ischial process not reduced; DISTRIBUTION: Congo Basin. interarcual cartilage absent, second pharyn- REMARKS: The genus has been recently gobranchial toothplate possesses a bony reviewed by Poll and Lambert (1965). flange; basihyal long and narrow; no tooth MATERIAL EXAMINED: H. platysternus: patches on second and third hypobranchials; Zaire (Congo): the types as listed above; teeth on fourth ceratobranchials; dorsal hy- paratypes: AMNH 6078 (1*/12). H. zebra: pohyal absent; anterior extension of anterior West Africa: Cotype: USNM 92965 (1); ceratohyal reduced; ventral hypohyal ex- USNM 191521 (l*/22). H. deprimozi: Con- panded; uncinate process on fourth epibran- go: SU 17480(6); BMNH 1974.9.18:522-527. chial articulates with that of third; first epi- branchial wide at its base; interhyal ossified; SUBFAMILY FLUVIPHYLACINAE ROBERTS two ossified basibranchials. Vomer with pos- Type Genus Fluviphylax Whitley, 1965. terior extension ventral to parasphenoid. DIAGNOSIS: Diminutive species of the fam- Lacrimal flat and broad, carrying a distinct ily Poeciliidae distinguished from all other sensory canal; dermosphenotic and preoper- cyprinodontiforms by extremely large eyes cular with distinct sensory canal; pectoral and reduced preorbital distance. girdle highset; first postcleithrum absent; COMPOSITION: One genus and species: posttemporal with ossified lower limb; post- Fluviphylax pygmaeus (Myers and Carval- temporal not fused to supracleithrum. ho). Ossified or unossified vomer; medial arm DISTRIBUTION: Brazil: Amazon Basin. of maxilla pronounced, dorsal process in- dented laterally; ventral arms narrow and GENUS FLUVIPHYLAX WHITLEY straight, not abutting rostral cartilage; outer arm reduced. Potamophylax Myers and Carvalho, in Myers, Premaxillary ascending processes narrow 1955, p. 7 (type species Potamophylax pyg- and reduced, not tapered posteriorly and not maeus Myers and Carvalho, by original desig- overlapping in the midline; rostral cartilage nation [name preoccupied in the Insecta]). reduced; outer arm of premaxilla with alve- Fluviphylax Whitley, 1965, p. 25 (type species olar process, indented posteriorly to form Potamophylax pygmaeus Myers and Carvalho, by monotypy). S-shaped arm. No ligament from the ventral arms of maxillaries to middle of rostral car- ETYMOLOGY: Fluviphylax from the Latin tilage; no ethmomaxillary ligament; no me- fluvius, meaning a stream or river, and from niscus between premaxilla and maxilla. Uni- the Greek phylax, meaning a guarder. cuspid bi- or tri-serial outer teeth. TYPES: Brazil: Rio Madeira at Borba: Po- Dentary expanded medially, very robust; tamophylax pygmaeus Myers and Carvalho, coronoid process on dentary not overlapping Paratypes SU 50196 (3). with that of articular; retroarticular not elon- COMPOSITION: As for the subfamily. gate. Autopalatine with head angled ante- DIAGNOSIS: AS for the subfamily. riorly; ventral process elongate reaching DEFINITION: Anal: ii, 6; Dorsal: 5-6; Pel- quadrate; metapterygoid absent. vic: 6; Pectoral: 10-11; Caudal; 4, 10, 4; Ver- Orbital rim free; anterior naris not tubular; tebrae: 12+14, 13 + 13; Gill rakers on the supraorbital sensory pores l-2a, recessed anterior arm of the first arch: 9-10. 2b-4a, 4b-7; seven preopercular pores; four Branchiostegal rays: 5; Scales lateral series: mandibular and three lacrimal pores. 24-26. Males larger than females; caudal fin trun- First pleural rib on parapophysis of second cate; pigmentation pattern often consisting vertebra; parapophysis not reduced; no pleu- of several dark crossbars. ral ribs on hemal spines; hypural plates fused No fatty predorsal ridge; swimbladder ex- into hypural fan. Epipleural ribs not bifid. tends posteriorly beyond three or four hemal Anal fin not modified into a gonopodium; spines. anal fin musculature unmodified; first proxi- 1981 PARENTI: CYPRINODONTIFORM FISHES 515 mal radial present; middle anal radials pres- arms of maxillaries to middle of rostral car- ent; urogenital opening of female covered tilage; no ethmomaxillary ligament; no me- with pocket of scales. niscus between premaxilla and maxilla. Uni- Spermatozeugmata not formed; fertiliza- cuspid bi- or tri-serial outer teeth. tion external; development nonannual; ovip- Dentary expanded medially, robust; cor- arous. Eggs round. onoid process on dentary not overlapping One dorsal ray articulating with the first with that of articular; retroarticular not elon- two dorsal radials; dorsal fin origin on pos- gate. Autopalatine with head angled ante- terior third of body. riorly; ventral process elongate reaching Autopterotic fossa moderate; lateral eth- quadrate; metapterygoid absent. moid not expanded medially, not reaching Orbital rim free; anterior naris not tubular; parasphenoid; parasphenoid not expanded supraorbital sensory pores reduced; seven anteriorly; no supraoccipital and exoccipital preopercular pores; four mandibular pores; processes; neural arches of first vertebra three lacrimal pores. open, not forming a spine; first vertebra ar- Males larger than females; all fins round- ticulates with skull via basioccipital and ex- ed; body generally straw-colored overall; occipital condyles; supraoccipital excluded with faint midlateral reticulations. from formation of foramen magnum; pari- No fatty predorsal ridge; swimbladder etals absent; nasals expanded medially. does not extend posteriorly beyond hemal Mesethmoid ossified; medial processes of spines. pelvic fin base and ischial process not re- DISTRIBUTION: AS for the subfamily. duced; interarcual cartilage reduced, attach- REMARKS: The genus and species was re- es laterally to second pharyngobranchial viewed by Roberts (1970) who reiterated with a bony flange; basihyal long and nar- Myers's (1955) statement that it was most row; no tooth patches on second and third likely related to the procatopines, although hypobranchials; teeth on fourth ceratobran- its precise placement within a phylogeny of chials; dorsal hypohyal absent; anterior ex- procatopines was not possible. Range of me- tension of anterior ceratohyal reduced; ven- ristic characters is supplemented by data tral hypohyal expanded; uncinate process on from Roberts (1970). fourth epibranchial articulates with that of MATERIAL EXAMINED: Brazil: Rio Madei- third; first epibranchial wide at its base; in- ra at Borba: the paratypes as listed above; terhyal ossified; two ossified basibranchials. SU 50196 (3); Manaus: near Rio Negro: MCZ Vomer with posterior extension ventral to 46714 (11); Lago Hyanuary: MCZ 41367 (5); parasphenoid. Manaus area: MCZ 49958 (4+/4); MCZ Lacrimal reduced, carrying a distinct sen- 46712 (21); MCZ 46713 (3*/64). sory canal; dermosphenotic and preopercu- lar with distinct sensory canal; pectoral gir- SUPERFAMILY CYPRINODONTOIDEA dle highset; first postcleithrum absent; DIAGNOSIS: Distinguished from other cy- posttemporal with ossified lower limb; post- prinodontiforms by the three derived char- temporal not fused to supracleithrum. acters: lateral ethmoid expanded medially Unossified vomer; medial arm of maxilla and oriented so it lies perpendicular to the straight with reduced dorsal process; ventral frontal; reduced autopterotic fossa; and in- arms narrow and straight, not abutting ros- tral cartilage; outer arm narrow. clinators of the anal fin greatly enlarged. Premaxillary ascending processes narrow and reduced, not tapered posteriorly and not FAMILY GOODEIDAE JORDAN overlapping in the midline; rostral cartilage Type Genus Jordan, 1880. reduced; outer arm of premaxilla with alve- DIAGNOSIS: Distingushed from other cy- olar process, indented posteriorly to form prinodontiform fishes by four derived char- S-shaped arm No ligament from the ventral acters: first two to seven middle anal radials 516 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 fused to the proximal radials; dorsal pro- vertebra; parapophysis not reduced; no pleu- cesses of the maxillaries greatly reduced; ral ribs on hemal spines; hypural plates fused distal arm of the maxilla straight, rather than into hypural fan. Epipleural ribs not bifid. curved; and articular greatly reduced. Anal fin not modified into a gonopodium; COMPOSITION: TWO subfamilies: Goodei- anal fin inclinators enlarged; first proximal nae Jordan and Empetrichthyinae Jordan, radial present; first middle anal radial absent; Evermann and Clark, as defined below. second fused to second proximal radial. DISTRIBUTION: Nevada; Death Valley sys- Spermatozeugmata not formed; fertiliza- tem; Mesa Central, Mexico (fig. 82). tion external; development nonannual; ovip- arous Eggs round. SUBFAMILY EMPETRICHTHYINAE JORDAN, One dorsal ray articulating with the first EVERMANN AND CLARK two dorsal radials; dorsal fin origin opposite origin of anal. Type Genus Empetrichthys, Gilbert, 1893. Autopterotic fossa reduced; lateral eth- DIAGNOSIS: Distinguished from other cy- moid expanded medially, reaching parasphe- prinodontiforms by a derived first epibran- noid and lying perpendicular to frontal; para- chial and from other fishes of the Goodeidae sphenoid not expanded anteriorly; no by lacking pelvic fins and fin supports. supraoccipital and exoccipital processes; COMPOSITION: Two genera, Empetrich- neural arches of first vertebra open, not thys with two species, merriami Gilbert and forming a spine; first vertebra articulates latos Miller, the latter with three subspecies with skull via basioccipital and exoccipital (latos, pahrump Miller, concavus Miller); condyles; supraoccipital excluded from for- Crenichthys with two species, nevadae mation of foramen magnum; parietals pres- Hubbs and baileyi (Gilbert). ent; nasals expanded medially. DISTRIBUTION: Death Valley system and Mesethmoid ossified; pelvic fins and fin eastern Nevada. supports absent; interarcual cartilage re- duced, attaches laterally to second pharyn- GENUS EMPETRICHTHYS GILBERT gobranchial with a bony flange; basihyal long and narrow; no tooth patches on second and Empetrichthys Gilbert, 1893, p. 233 (type species third hypobranchials; teeth on fourth cera- Empetrichthys merriami Gilbert, by original tobranchials; dorsal hypohyal absent; ante- designation). rior extension of anterior ceratohyal ventral ETYMOLOGY: Empetrichthys from the to ventral hypohyal; uncinate process on Greek em, meaning within; petros, meaning fourth epibranchial articulates with that of rocks; and ichthys, meaning fish with rocks third; first epibranchial with indentation; in- within, referring to the large molariform pha- terhyal ossified; two ossified basibranchials. ryngeal teeth. Vomer with posterior extension ventral to TYPES: Nevada: Ash Meadows: Empe- parasphenoid. trichthys merriami Gilbert, Type, USNM Lacrimal flat and broad, carrying a distinct 131151 (1). sensory canal; dermosphenotic and preoper- COMPOSITION: AS listed for the subfamily. cular with a distinct sensory canal; pectoral DIAGNOSIS: Distinguished from other girdle highset; first postcleithrum absent; members of the Goodeidae by having fleshy posttemporal with ossified lower limb; post- bases of the dorsal and anal fins, and greatly temporal not fused to supracleithrum. enlarged pharyngeal teeth. Vomer ossified, edentulous; medial arm of DEFINITION: Anal: i, 13; Dorsal: i, 11; Pel- maxilla straight with reduced dorsal process; vic: 0; Pectoral: 17; Caudal: 6, 18, 6; Verte- ventral arms not abutting rostral cartilage; brae: 15+16; Gill rakers on the anterior arm outer arm narrow. of the first arch: 12-13. Branchiostegal rays: Premaxillary ascending processes narrow 6; Scales lateral series: 29-30. and reduced, not tapered posteriorly and not First pleural rib on parapophysis of second overlapping in the midline; rostral cartilage 1981 PARENTI: CYPRINODONTIFORM FISHES 517 reduced; outer arm of premaxilla with alveo- COMPOSITION: As listed for the subfamily. lar process, indented posteriorly to form DIAGNOSIS: Distinguished from Empe- S-shaped arm. No ligament from the ventral trichthys by an outer row of bicupid teeth, arms of maxillaries to middle of rostral car- and an increase in the number of gill rakers tilage; no ethmomaxillary ligament; no me- on the first arm to 20-22. niscus between premaxilla and maxilla. En- DEFINITION: Anal: i, 13; Dorsal: i, 11; Pel- larged biserial outer teeth. vic: 0; Pectoral: 16; Caudal: 9, 18, 9; Verte- Dentary expanded medially, robust; cor- brae: 11 + 17; Gill rakers on the anterior arm onoid process on dentary not overlapping of the first arch: 20-22. Branchiostegal rays: with that of articular; articular reduced; ret- 6; Scales lateral series: 26-28. roarticular not elongate. Autopalatine with First pleural rib on parapophysis of second head angled anteriorly; ventral process elon- vertebra; parapophysis not reduced; no pleu- gate reaching quadrate; metapterygoid ab- ral ribs on hemal spines; hypural plates fused sent. into hypural fan. Epipleural ribs not bifid. Orbital rim free; anterior naris not tubular; Anal fin not modified into a gonopodium; supraorbital sensory pores l-2a, 2b-4, 5-7; anal fin inclinators enlarged; first proximal seven preopercular pores; four mandibular radial present; first five middle anal radials pores; three lacrimal pores. absent. Females larger than males; fins rounded or Spermatozeugmata not formed; fertiliza- truncate. Bases of anal and dorsal fins fleshy. tion external; development nonannual; ovip- No fatty predorsal ridge; swimbladder arous. Eggs round. does not extend posteriorly beyond hemal One dorsal ray articulating with the first spines. two dorsal radials; dorsal fin origin opposite DISTRIBUTION: Death Valley, Nevada sys- origin of anal. tem. Autopterotic fossa reduced; lateral eth- REMARKS: The genus Empetrichthys was moid expanded medially, reaching parasphe- formerly assigned to the subfamily Funduli- noid and lying perpendicular to frontal; para- nae on the basis of primitive characters. Of sphenoid not expanded anteriorly; no the named taxa, all but one, latos pahrump, supraoccipital and exoccipital processes; are extinct (Soltz and Naiman, 1978). neural arches of first vertebra open, not MATERIAL EXAMINED: E. merriami: Ne- forming a spine; first vertebra articulates vada: Ash Meadows: the type as listed with skull via basioccipital and exoccipital above; Paratypes: USNM 46102 (2); Pah- condyles; supraoccipital excluded from for- rump: SU 35966 (1*/15). E. latos pahrump: mation of foramen magnum; parietals pres- Nevada: Pahrump Ranch: CAS 22990 (1*/ ent; nasals expanded medially. 16). Mesethmoid ossified; pelvic fin and fin FOSSILS: One species, E. erdisi Uyeno and supports absent; interarcual cartilage re- Miller from the Pleistocene of Nevada, has duced, attaches laterally to second pharyn- been described (Uyeno and Miller, 1962). gobranchial with a bony flange; basihyal long and narrow; no tooth patches on second and GENUS CRENICHTHYS HUBBS third hypobranchials; teeth on fourth cera- tobranchials; dorsal hypohyal absent; ante- Crenichthys Hubbs, 1932, p. 1 (type species rior extension of anterior ceratohyal ventral Crenichthys nevadae Hubbs, by original des- to ventral hypohyal; uncinate process on ignation). fourth epibranchial articulates with that of ETYMOLOGY: Crenichthys from the Greek third; first epibranchial wide at its base; in- cren, meaning spring, and ichthys referring terhyal ossified; two ossified basibranchials. to the spring habitat of the genus. Vomer with posterior extension ventral to TYPES: Nevada: Railroad Valley: Cren- parasphenoid. ichthys nevadae Hubbs, Holotype MCZ Lacrimal flat and broad, carrying a distinct 32948 (1). sensory canal; dermosphenotic and preoper- 518 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

cular with distinct sensory canal; pectoral ened, unbranched anal fin rays in males girdle highset; first postcleithrum absent; crowded together and separated by a notch posttemporal with ossified lower limb; post- from the rest of the fin; first anal fin ray of temporal not fused to supracleithrum. males rudimentary; males with a pseudo- Vomer ossified, edentulous; medial arm of phallus; embryos with nutritive trophic pro- maxilla straight with reduced dorsal process; cesses; and ovaries united medially with ventral arms not abutting rostral cartilage; ovigerous tissue partly to completely elimi- outer arm narrow. nated from the ovarian walls. Premaxillary ascending processes narrow COMPOSITION: Approximately 36 species and reduced, not tapered posteriorly and not in 16 genera. overlapping in the midline; rostral cartilage DISTRIBUTION: Mesa Central, Mexico reduced or absent; outer arm of premaxilla with a concentration of species in the Rio with alveolar process, indented posteriorly Lerma Basin. to form S-shaped arm. No ligament from the DEFINITION: Anal (i, ii) 11-13; Dorsal: i, ventral arms of maxillaries to middle of ros- 14—i, 15; Pelvic: 6; Pectoral: 15-16; Caudal: tral cartilage; no ethmomaxillary ligament; 8,12,8; Vertebrae: 16+21; Gill rakers on the no meniscus between premaxilla and maxil- anterior arm of the first arch: 27-29. Bran- la. Bicuspid, biserial outer teeth. chiostegal rays: 4-6; Scales lateral series: Dentary expanded medially, robust; cor- 30-35. onoid process on dentary not overlapping First pleural rib on parapophysis of second with that of articular; retroarticular not elon- vertebra; parapophysis not reduced; no pleu- gate. Autopalatine with head angled ante- ral ribs on hemal spines; hypural plates fused riorly; ventral process elongate reaching into hypural fan. Epipleural ribs not bifid. quadrate; metapterygoid absent. Anal fin not modified into a gonopodium; Orbital rim free; anterior naris not tubular; anal fin inclinators enlarged; first proximal supraorbital sensory pores l-2a, 2b-4, 5- radial present; first five middle anal radials 7; seven preopercular pores; four mandiublar absent. First anal ray rudimentary in males. pores; three lacrimal pores. Spermatozeugmata formed; fertilization Femlaes larger than males; all fins rounded internal; males possess a muscular organ re- or truncate. Pigmentation pattern typically sponsible for the transfer of sperm; ovary consisting of a series of midlateral dark fused with a concentration of ovigerous tis- blotches. sue in the outer regions; embryos without- No fatty predorsal ridge; swimbladder pocketings of the intestine termed tropho- does not extend posteriorly beyond hemal taeniae; fertilization in the ovary. spines. One dorsal ray articulating with the first DISTRIBUTION: Nevada: Railroad Valley two dorsal radials; dorsal fin origin opposite and White River Valley. origin of anal. REMARKS: Crenichthys, as well as Empe- Autopterotic fossa reduced; lateral eth- trichthys, has been removed from a classifi- moid expanded medially, reaching parasphe- cation with the fundulines and placed in a noid and lying perpendicular to frontal; para- monophyletic group with the goodeid fishes sphenoid not expanded anteriorly; no for the first time in this study. supraoccipital and exoccipital processes; MATERIAL EXAMINED: C. nevadae: the neural arches of first vertebra open, not types as listed above: Nevada: Twin Springs forming a spine; first vertebra articulates Ranch: SU 48125 (2); C. baileyi: Nevada: with skull via basioccipital and exoccipital Ash Springs: CAS 22980 (l*/42); USNM condyles; supraoccipital excluded from for- 11750 (2*/25); Aquarium material: AMNH mation of foramen magnum; parietals pres- 38408SW (4*/4). ent; nasals expanded medially. Mesethmoid ossified; pelvic fin and fin SUBFAMILY GOODEINAE JORDAN supports absent; interarcual cartilage re- DIAGNOSIS: Viviparous killifishes distin- duced, attaches laterally to second pharyn- guished from cyprinodontiforms by short- gobranchial with a bony flange; basihyal long 1981 PARENTI: CYPRINODONTIFORM FISHES 519

and narrow; no tooth patches on second and and Fitzsimons (1971). In addition, several third hypobranchials; teeth on fourth cera- recommendations for a reclassification of the tobranchials; dorsal hypohyal absent; ante- goodeids are made in this paper, including rior extension of anterior ceratohyal ventral the suggestion that Ataentiobius is a derived to ventral hypohyal; uncinate process on member of the group and has secondarily fourth epibranchial articulates with that of lost trophotaeniae. third; first epibranchial wide at its base; in- MATERIAL EXAMINED: Ataeniobius tow- terhyal ossified; two ossified basibranchials. eri: San Luis Potosi: SU 9396 (2); Chara- Vomer with posterior extension ventral to codon lateralis: Durango: CAS 40705 (2*1 parasphenoid. 40); innominatus: Lerma Lacrimal flat and broad, carrying a distinct River Basin: SU 47063 (45); Goodea luitpol- sensory canal; dermosphenotic and preoper- di: AMNH 18622 (1+/7); whiter. cular with distinct sensory canal; pectoral Michoacan: CAS 16044 (2*/10); ler- girdle highset; first postcleithrum absent; mae: Patzcuaro: SU 22342 (2*/20); Xenen- posttemporal with ossified lower limb; post- dum xaliscone: : Paratypes, SU 6207 temporal not fused to supracleithrum. (4); variata: Jalisco: ex UMMZ Vomer ossified, edentulous; medial arm of 179760 (uncat at CAS) (15): Zoogoneticus maxilla straight with reduced dorsal process; diazi: Michoacan: USNM 218752 (2*/34). ventral arms not abutting rostral cartilage; FOSSILS: Goodeid fossils are known from outer arm narrow. Pleistocene and Miocene deposits, Mesa Premaxillary ascending processes narrow Central, Mexico (Alvarez and Arreola, 1972; and reduced, not tapered posteriorly and not Smith, Cavender and Miller 1975). overlapping in the midline; rostral cartilage reduced or absent; outer arm of premaxilla FAMILY CYPRINODONTIDAE GILL with alveolar process, indented posteriorly Type Genus Cyprinodon Lacepede, 1803. to form S-shaped arm; No ligament from the DIAGNOSIS: Distinguished from other cy- ventral arms of maxillaries to middle of ros- prinodon tiforms by three derived characters: tral cartilage; no ethmomaxillary ligament; dorsal processes of the maxillaries expanded no meniscus between premaxilla and maxil- medially nearly meeting in the midline, and la. Uni- or bi-cuspid outer teeth. possessing a distinct groove; lateral arm of Dentary expanded medially, robust; cor- the maxilla greatly expanded; and, the tooth- onoid process on dentary not overlapping with that of articular; retroarticular not elon- plate of the fourth pharyngobranchial greatly reduced. gate. Autopalatine with head angled ante- COMPOSITION: Two subfamilies, Cuban- riorly; ventral process elongate reaching ichthyinae, new subfamily, and Cyprinodon- quadrate; metapterygoid absent. Orbital rim free; anterior naris not tubular; tinae Gill, as described below. DISTRIBUTION: North, South, and Middle supraorbital sensory pores l-2a, 2b-4a, 4b- 7; seven preopercular pores; four mandibular America, the Caribbean; Mediterranean An- atolian regions, as detailed below. pores; three lacrimal pores. Females larger than males; all fins rounded or truncate. SUBFAMILY CUBANICHTHYINAE, No fatty predorsal ridge; swimbladder NEW SUBFAMILY does not extend posteriorly beyond hemal Type Genus Cubanichthys Hubbs, 1924. spines. DIAGNOSIS: Distinguished from other cy- REMARKS: It is stressed that the subfamily prinodontiforms by four derived characters: Goodeinae in this present study is equivalent an enlarged supraoccipital crest; an elongate to the family Goodeidae of former authors. dorsal process of the autopalatine; a supraor- Workers on the subfamily generally follow bital sensory pore pattern characterized by the classification of the group presented by a large third pore; and from other primitive Hubbs and Turner (1939) and suggestions for cyprinodontids in lacking an ossified lower modifications of that classification by Miller limb of the posttemporal. 520 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

COMPOSITION: A single genus, Cuban- row; no tooth patches on second and third ichthys Hubbs, with two species: cubensis hypobranchials; teeth on fourth ceratobran- Eigenmann and pengelleyi (Fowler). chials; dorsal hypohyal absent; anterior ex- DISTRIBUTION: Cuba and Jamaica. tension of anterior ceratohyal ventral to ven- tral hypohyal; uncinate process on fourth epibranchial articulates with that of third; GENUS CUBANICHTHYS HUBBS first epibranchial wide at its base; interhyal Cubanichthys Hubbs, 1926, p. 4 (type species ossified; two ossified basibranchials. Vomer Fundulus cubensis Eigenmann, by original des- with posterior extension ventral to parasphe- ignation). noid. Chriopeoides Fowler, 1939, p. 4 (type species Lacrimal flat and broad, carrying a distinct Chriopeoides pengelleyi Fowler, by original sensory canal; dermosphenotic and preoper- designation). cular with distinct sensory canal; pectoral ETYMOLOGY: Cubanichthys, after Cuba to girdle lowset; first postcleithrum present; which the species cubensis is endemic. posttemporal with unossified lower limb; TYPES: Cuba: Pinar del Rio: Fundulus posttemporal not fused to supracleithrum. cubensis Eigenmann; IU 9887 (3). Vomer ossified, edentulous; medial arm of COMPOSITION: AS for the subfamily. maxilla straight with pronounced dorsal pro- DIAGNOSIS: AS for the subfamily. cesses with a groove, nearly meeting in the DEFINITION: Anal: i, 9; Dorsal: ii, 9; Pel- midline; ventral arms narrow, not abutting vic: 6; Pectoral: 18; Caudal: 7, 10, 7; Verte- rostral cartilage; outer arm robust. brae: 11 + 16; Gill rakers on the anterior arm Premaxillary ascending processes narrow of the first arch: 9. Branchiostegal rays: 6; and reduced, not tapered posteriorly and not Scales lateral series: 24-26. overlapping in the midline; rostral cartilage First pleural rib on parapophysis of second reduced; outer arm of premaxilla with alveo- vertebra; parapophysis not reduced; no pleu- lar process, indented posteriorly to form ral ribs on hemal spines; hypural plates fused S-shaped arm; No ligament from the ventral into hypural fan. Epipleural ribs not bifid. arms of maxillaries to middle of rostral car- Anal fin not modified into a gonopodium; tilage; no ethmomaxillary ligament; no me- anal fin inclinators enlarged; first proximal niscus between premaxilla and maxilla. Un- radial present; middle anal radials present. icuspid bi- or tri-serial outer teeth. Spermatozeugmata not formed; fertiliza- Dentary expanded medially, robust; cor- tion external; development nonannual; ovip- onoid process on dentary not overlapping arous. Eggs round. with that of articular; retroarticular not elon- One dorsal ray articulating with the first gate. Autopalatine with head angled ante- two dorsal radials; dorsal fin origin opposite riorly, elongate dorsal process; ventral pro- origin of anal. cess elongate reaching quadrate; meta- Autopterotic fossa reduced; lateral ethmoid pterygoid absent. expanded medially, reaching parasphenoid Orbital rim free; anterior naris not tubular; and lying perpendicular to frontal; para- supraorbital sensory pores l-2a, an enlarged sphenoid not expanded anteriorly; supra- 3, 4b-7; seven preopercular pores; four man- occipital crest present; neural arches of first dibular pores; three lacrimal pores. vertebra open, not forming a spine; first ver- Females larger than males; males with dor- tebra articulates with skull via basioccipital sal fin enlarged; caudal truncate; pigmenta- and exoccipital condyles; supraoccipital ex- tion pattern consisting of a dark lateral band cluded from formation of foramen magnum; extending from the eye onto the caudal pe- parietals present; nasals expanded medially. duncle. Mesethmoid ossified; medial processes of No fatty predorsal ridge; swimbladder pelvic fin base and ischial process not re- does not extend posteriorly beyond hemal duced; interarcual cartilage reduced, at- spines. taches laterally to second pharyngobranchial DISTRIBUTION: AS for the subfamily. with a bony flange; basihyal long and nar- REMARKS: Both cubensis and pengelleyi 1981 PARENTI: CYPRINODONTIFORM FISHES 521 were classified in the subfamily Fundulinae Tellia Gervais, 1853, p. 15 (type species Tellia prior to this study. The generic name Chrio- apoda Gervais, by original designation). peoides refers to the resemblance of pengel- Micromugil Gulia, 1861, p. 11 (type species Aphanius faciatus Nardo, by monotypy). leyi to the North American Lucania (Chrio- Aphaniops Hoedeman, 1951, p. 2 (type species peops) goodei. Also, cubensis had been Lebias dispar Ruppell, by original designa- considered an island form of the wholly tion). North American Lucania (Hubbs and Miller, 1965; Rosen, 1976). Foster (1969) provided ETYMOLOGY: Aphanius from the Greek observations on the habitat and locality of aphanes, meaning secret or unknown in ref- pengelleyi. erence to the cyprinodonts of the Anatolian MATERIAL EXAMINED: Cuba: Pinar del region. Rio, Syntypes as listed above, C. cubensis COMPOSITION: Approximately 30 species, ANSP 60283-87 (l*/4); Jamaica: St. Eliza- as listed in Lazara (1979), minus those of the beth's Parish; Black River Drainage: C. pen- men/o-complex that are referred to the ge- gelleyi ANSP 112908 (l*/64). nus "Aphanius.." DIAGNOSIS: Distinguished from other SUBFAMILY CYPRINODONTINAE GILL members of the tribe Orestiini by three prim- itive characters: cephalic sensory pore pat- DIAGNOSIS: Distinguished from all other tern represented by pores rather than re- cyprinodontiforms by three uniquely derived duced to neuromasts; a urohyal that is not characters: second pharyngobranchial offset embedded in the urohyal membranes; and an to the third; Meckel's cartilage expanded ossified interhyal. posteriorly; and, transverse processes of the DEFINITION: Anal: ii, 8; Dorsal: i, 9; Pel- vertebrae reduced and cup-shaped; and by vic: 6; Pectoral: 15-16; Caudal: 7, 14, 7; Ver- two independently derived characters: pari- tebrae: 11 + 15; Gill rakers on the anterior etals absent, and uniserial outer jaw teeth. arm of the first arch: 8-10. Branchiostegal COMPOSITION: TWO tribes, Cyprinodontini rays: 5; Scales lateral series: 23-26. Gill and Orestiini Bleeker, as described be- First pleural rib on parapophysis of second low. vertebra; parapophysis reduced; no pleural DISTRIBUTION: AS for the tribes listed be- ribs on hemal spines; hypural plates fused low. into hypural fan. Epipleural ribs not bifid. TRIBE ORESTIINI BLEEKER Anal fin not modified into a gonopodium; anal fin inclinators enlarged; first proximal DIAGNOSIS: Distinguished from all other radial present; middle anal radials present. cyprinodontiform fishes by an extremely ro- Spermatozeugmata not formed; fertiliza- bust lower jaw caused by a medial extension tion external; development nonannual; ovip- of the dentary. arous. Eggs round. COMPOSITION: Four genera and approxi- One dorsal ray articulating with the first mately 65 species: Aphanius Nardo, two dorsal radials; dorsal fin origin opposite "Aphanius," Kosswigichthys Sozer, and origin of anal. Orestias Valenciennes. Autopterotic fossa reduced; lateral eth- DISTRIBUTION: (fig. 86) Mediterranean re- moid expanded medially, reaching parasphe- gion: North Africa, Spain, Italy, Turkey, noid and lying perpendicular to frontal; para- Greece, and Mediterranean islands, as well sphenoid not expanded anteriorly; no as the Saudi Arabian Peninsula and Iran. supraoccipital and exoccipital processes; neural arches of first vertebra open, not GENUS APHANIUS NARDO forming a spine; first vertebra articulates Aphanius Nardo, 1827, p. 48 (type species with skull via basioccipital and exoccipital Aphanius fasciatus Nardo, by subsequent des- condyles, exoccipital condyles reduced; su- ignation). praoccipital excluded from formation of fo- Lehias Cuvier, 1817, p. 119 (type species Aphan- ramen magnum; parietals absent; nasals ex- ius fasciatus Nardo, by monotypy). panded medially. 522 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

Mesethmoid unossified; medial processes No fatty predorsal ridge; swimbladder of pelvic fin base and ischial process not re- does not extend posteriorly beyond hemal duced; interarcual cartilage reduced, at- spines. taches laterally to second pharygobranchial DISTRIBUTION: Mediterranean, along the with a flange and oriented dorsally; basihyal north coast of Africa, Spain, Italy, along the long and narrow; no tooth patches on second periphery of the Saudi Arabian Peninsula; and third hypobranchials; teeth on fourth Turkey and Greece. ceratobranchials; dorsal hypohyal absent; REMARKS: The genus Aphanius as the anterior extension of anterior ceratohyal term is used here references those species of ventral to ventral hypohyal; uncinate pro- the genus assessed as the most plesiomor- cess on fourth epibranchial articulates with phic of the Anatolian cyprinodonts and Or- that of third; first epibranchial wide at its es tias. The more apomorphic species are base; interhyal ossified; two ossified basi- placed in "Aphanius." branchials. Vomer with posterior extension MATERIAL EXAMINED: A. fasciatus: ventral to parasphenoid. Aquarium material: AMNH 36770SW (3*/7); Lacrimal flat and broad, carrying a distinct BMNH 1958.3.3:551-580 (30); A. dispar: sensory canal; dermosphenotic and preoper- Abyssinia: Syntype: BMNH 1860.11.9:152 cular with distinct sensory canal; pectoral (1/3); Saudi Arabia: Persian Gulf: USNM girdle lowset; first postcleithrum present; 147834 (41); BMNH 1977.12.13:1-490. (2*/ posttemporal with ossified lower limb; post- 490); A. apoda: Algeria: BMNH 1958.4.22: temporal not fused to supracleithrum. 1-7(7). Vomer ossified, edentulous; medial arm of FOSSILS: Fossil cyprinodontiforms of the maxilla straight with pronounced dorsal pro- Old World primarily in the Mediterranean cesses with a groove, nearly meeting in the region have been assigned to the genus midline; ventral arms narrow, not abutting Aphanius or to new fossil taxa: rostral cartilage; outer arm robust. Premaxillary ascending processes narrow GENUS PROLEBIAS SAUVAGE and reduced, not tapered posteriorly and not overlapping in the midline; rostral cartilage Prolebias Sauvage, 1874, p. 187 (type species Le- reduced; outer arm of premaxilla with alve- bias cephalotes Agassiz, by original designa- olar process, indented posteriorly to form tion). (Oligocene and Miocene of Western Eu- S-shaped arm. No ligament from the ventral rope.) arms of maxillaries to middle of rostral car- tilage; no ethmomaxillary ligament; no me- GENUS PACHYLEBIAS WOODWARD niscus between premaxilla and maxilla. Tri- cuspid, uniserial outer teeth. Pachylebias Woodward, 1901, p. 294 (type species Lebias crassicaudus Agassiz, by orig- Dentary with medial extension, robust; inal designation ). (Miocene of Italy and Crete.) coronoid process on dentary not overlapping with that of articular; retroarticular not elon- gate. Autopalatine with head angled ante- GENUS BRACHYLEBIAS PRIEM riorly; ventral process elongate reaching Brachylebias Priem, 1908, p. 21 (type species quadrate; metapterygoid absent. Brachylebias persicus Priem, by original des- Orbital rim free; anterior naris not tubular; ignation). (Miocene of Iran.) supraorbital sensory pores l-4a, 4b-7; seven preopercular pores; four mandibular pores; three lacrimal pores. GENUS "APHANIUS" Females larger than males; males often ETYMOLOGY: The genus "Aphanius" is with enlarged dorsal and anal fins; caudal fin used as a reference for the Aphanius mento- rounded or truncate. complex hypothesized to be more closely re- 1981 PARENTI: CYPRINODONTIFORM FISHES 523 lated to Kosswigichthys and Orestias than with that of third; first epibranchial wide at to other Aphanius. its base; interhyal ossified; two ossified ba- COMPOSITION: Species of the Aphanius sibranchials. Vomer with posterior extension mento-complex including mento (Heckel) ventral to parasphenoid. and chantrei (Gaillard). Lacrimal flat and broad, carrying a distinct DIAGNOSIS: Distinguished from other sensory canal; dermosphenotic and preoper- Aphanius species by a urohyal embedded in cular with distinct sensory canal; pectoral the fold of the branchiostegal membranes girdle lowset; first postcleithrum present; and a derived head pore pattern. posttemporal with ossified lower limb; post- DEFINITION: Anal: i, 11; Dorsal: i, 9; Pel- temporal not fused to supracleithrum. vic: 6; Pectoral: 15-16; Caudal: 7, 14, 7; Ver- Vomer ossified, edentulous; medial arm of tebrae: 12+14; Gill rakers on the anterior maxilla straight with pronounced dorsal pro- arm of the first arch: 8-10. Branchiostegal cesses with a groove, nearly meeting in the rays: 5; Scales lateral series: 23-26. midline; ventral arms narrow, not abutting First pleura! rib on parapophysis of second rostral cartilage; outer arm robust. vertebra; parapophysis reduced; no pleural Premaxillary ascending processes narrow ribs on hemal spines; hypural plates fused and reduced, not tapered posteriorly and not into hypural fan. Epipleural ribs not bifid. overlapping in the midline; rostral cartilage Anal fin not modified into a gonopodium; reduced; outer arm of premaxilla with alveo- anal fin inclinators enlarged; first proximal lar process, indented posteriorly to form radial present; middle anal radials present. S-shaped arm. No ligament from the ventral Spermatozeugmata not formed; fertiliza- arms of maxillaries to middle of rostral car- tion external; development nonannual; ovip- tilage; not ethmomaxillary ligament; no me- arous. Eggs round. niscus between premaxilla and maxilla. Tri- One dorsal ray articulating with the first cuspid, uniserial outer teeth. two dorsal radials; dorsal fin origin opposite Dentary with medial extension, robust; origin of anal. coronoid process on dentary not overlapping Autopterotic fossa reduced; lateral eth- with that of articular; retroarticular not elon- moid expanded medially, reaching parasphe- gate. Autopalatine with head angled ante- noid and lying perpendicular to frontal; para- riorly; ventral process elongate reaching sphenoid not expanded anteriorly; no quadrate; metapterygoid absent, urohyal supraoccipital and exoccipital processes; embedded in branchiostegal membranes. neural arches of first vertebra open, not Orbital rim free; anterior naris not tubular; forming a spine; first vertebra articulates cephalic sensory pores reduced to a series of with skull via basioccipital and exoccipital neuromasts. condyles; supraoccipital excluded from for- Females larger than males; males often mation of foramen magnum; parietals ab- with enlarged dorsal and anal fins; caudal fin sent; nasals expanded medially. rounded or truncate. Mesethmoid unossified; medial processes No fatty predorsal ridge; swimbladder of pelvic fin base and ischial process not re- does not extend posteriorly beyond hemal duced; interarcual cartilage reduced, at- spines. taches laterally to second pharyngobranchial DISTRIBUTION: The Mediterranean (Spain, with a bony flange and oriented dorsally; ba- Italy) and western Turkey. sihyal long and narrow; no tooth patches on REMARKS: The genus "Aphanius'''' is used second and third hypobranchials; teeth on as a reference for derived species of Aphan- fourth ceratobranchials; dorsal hypohyal ab- ius. It is not formally named as a new genus sent; anterior extension of anterior cerato- since its limits cannot readily be defined and hyal ventral to ventral hypohyal; uncinate may be expanded to include more species process on fourth epibranchial articulates still referenced in the genus Aphanius. 524 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

MATERIAL EXAMINED: A. mento: Aquar- ramen magnum; parietals absent; nasals ex- ium material: AMNH 28610 (2+/l*/9). panded medially. Mesethmoid unossified; medial processes of pelvic fin base and ischial process not re- duced; interarcual cartilage reduced, at- GENUS KOSSWIGICHTHYS SOZER taches laterally to second pharyngobranchial Kosswigichthys Sozer, 1942, p. 308 (type species with a bony flange and oriented dorsally; ba- Kosswigichthys asquamatus Sozer, by original sihyal long and narrow; no tooth patches on designation). second and third hypobranchials; teeth on Anatolichthys Kosswig and Sozer, 1945, p. 77 fourth ceratobranchials; dorsal hypohyal ab- (type species Anatolichthys splendens Kosswig sent; anterior extension of anterior cerato- and Sozer, by original designation). hyal ventral to ventral hypohyal; uncinate ETYMOLOGY: Kosswigichthys, in honor of process on fourth epibranchial articulates Curt Kosswig, prominent ichthyologist of with that of third; first epibranchial wide at fishes of the Anatolian region. its base; interhyal unossified; two ossified COMPOSITION: Four species: asquamatus basibranchials. Vomer with posterior exten- Sozer, burdurensis (Askiray), splendens sion ventral to parasphenoid. (Kosswig and Sozer), transgrediens (Aski- Lacrimal flat and broad, carrying a distinct ray). sensory canal; dermosphenotic absent; pre- DIAGNOSIS: Distinguished from other opercular with distinct sensory canal; pec- members of the tribe Orestiini by lacking the toral girdle lowset; first postcleithrum pres- dermosphenotic. ent; posttemporal with unossified lower DEFINITION: Anal: i, 10; Dorsal: i, 9; Pel- limb; posttemporal not fused to supraclei- vic: 6-7; Pectoral: 12; Caudal: 8, 10, 8; Ver- thrum. tebrae: 11 +17-12+17; Gill rakers on the an- Vomer ossified, edentulous; medial arm of terior arm of the first arch: 10-12. maxilla straight with pronounced dorsal pro- Branchiostegal rays: 5; Scales lateral series: cesses with a groove, nearly meeting in the 0-30. midline; ventral arms narrow, not abutting First pleural rib on parapophysis of second rostral cartilage; outer arm robust. vertebra; parapophysis reduced; no pleural Premaxillary ascending processes narrow ribs on hemal spines; hypural plates fused and reduced, not tapered posteriorly and not into hypural fan. Epipleural ribs not bifid. overlapping in the midline; rostral cartilage Anal fin not modified into a gonopodium; reduced or absent; outer arm of premaxilla anal fin inclinators enlarged; first proximal with alveolar process, indented posteriorly radial present; middle anal radials present. to form S-shaped arm. No ligament from the Spermatozeugmata not formed; fertiliza- ventral arms of maxillaries to middle of ros- tion external; development nonannual; ovip- tral cartilage; no ethmomaxillary ligament; arous. Eggs round. no meniscus between premaxilla and maxil- One dorsal ray articulating with the first la. Uni- or tri-cuspid uniserial outer teeth. two dorsal radials; dorsal fin origin opposite Dentary with medial extension, robust; origin of anal. coronoid process on dentary not overlapping Autopterotic fossa reduced; lateral eth- with that of articular; retroarticular not elon- moid expanded medially, reaching parasphe- gate. Autopalatine with head angled ante- noid and lying perpendicular to frontal; para- riorly; ventral process elongate reaching sphenoid not expanded anteriorly; no quadrate; metapterygoid absent. supraoccipital and exoccipital processes; Orbital rim free; anterior naris not tubular; neural arches of first vertebra open, not cephalic sensory pores reduced to derived forming a spine; first vertebra articulates neuromast pattern or absent. with skull via basioccipital and exoccipital Females larger than males; males never condyles, exoccipital condyles reduced; su- with fin extensions; caudal fin truncate; praoccipital excluded from formation of fo- scales reduced or absent. 1981 PARENTI: CYPRINODONTIFORM FISHES 525

No fatty predorsal ridge; swimbladder Spermatozeugmata not formed; fertiliza- does not extend posteriorly beyond hemal tion external; development nonannual; ovip- spines. arous. Eggs round. DISTRIBUTION: Freshwater lakes of Tur- One dorsal ray articulating with the first key. two dorsal radials; dorsal fin origin opposite REMARKS: Kosswigichthys has been syn- origin of anal. onymized with Aphanius by many authors; Autopterotic fossa reduced; lateral eth- it is hypothesized, however, that the genus moid expanded medially, reaching parasphe- is more closely related to Orestias than to noid and lying perpendicular to frontal; para- other Anatolian cyprinodontines. This asso- sphenoid not expanded anteriorly; no ciation produces one of the more unusual supraoccipital and exoccipital processes; distributional patterns within the group. neural arches of first vertebra open, not MATERIAL EXAMINED: K. asquamatus: forming a spine; first vertebra articulates Aquarium material: AMNH 28622 (2+15); with skull via basioccipital and exoccipital Turkey: ANSP 89883 (l*/6); BMNH condyles, exoccipital condyles reduced; su- 1948.3.15:40-43 (l*/3); AT. fra/wgredfgm: praoccipital excluded from formation of fo- Acy Gol- Akpinar ANSP 89890 (6); K. splen- ramen magnum; parietals absent; nasals ex- dens: Turkey: SU 15830 (2*/15). panded medially. Mesethmoid ossified; pelvic fin and fin GENUS ORESTIAS VALENCIENNES supports absent; interarcual cartilage re- duced, attaches laterally to second pharyn- Orestias Valenciennes, 1839, p. 118 (type species gobranchial with a bony flange; basihyal long Orestias cuvieri Valenciennes, by subsequent and narrow; no tooth patches on second and designation). third hypobranchials; teeth on fourth cera- Protorestias Allen (in Eigenmann and Allen), tobranchials; dorsal hypohyal absent; no an- 1942, p. 353 (hypothetical ancestral genus of terior extension of anterior ceratohyal ven- Orestias, no type designated). tral to ventral hypohyal; uncinate process on ETYMOLOGY: Orestias, after the Greek fourth epibranchial articulates with that of mythological figure Orestes, said to have third; first epibranchial wide at its base; in- been hidden in the mountains, in reference terhyal not ossified; two ossified basibran- to the unique distribution of the genus in the chials. Vomer with posterior extension ven- high-altitude lakes of the . tral to parasphenoid. TYPES: Peru: : Orestias cu- Lacrimal flat and braod, carrying a distinct vieri Valenciennes, MHNH. sensory canal; dermosphenotic reduced; pre- DIAGNOSIS: Distinguished from other opercular with reduced sensory canal; pec- members of the family Cyprinodontidae by toral girdle lowset; first postcleithrum ab- an absence of pelvic fins and fin supports, sent; posttemporal with unossified lower absence of a vomer, and absence of a first limb; posttemporal not fused to supraclei- postcleithrum. thrum. DEFINITION: Anal: i, 13: Dorsal: i, 13; Pel- Vomer absent; medial arm of maxilla vic: 0; Pectoral: 15-17; Caudal: 8, 15, 8; Ver- straight with pronounced dorsal processes tebrae: 15+16-15+18; Gill rakers on the with a groove, nearly meeting in the midline; anterior arm of the first arch: 9-24. ventral arms narrow, not abutting rostral car- Branchiostegal rays: 5-6; Scales lateral se- tilage; outer arm robust. ries: 0-54. Premaxillary ascending processes narrow First pleural rib on parapophysis of second and reduced, not tapered posteriorly and not vertebra; parapophysis not reduced; no pleu- overlapping in the midline; rostral cartilage ral ribs on hemal spines; hypural plates fused reduced or absent; outer arm of premaxilla into hypural fan. Epipleural ribs bifid or not. with alveolar process, indented posteriorly Anal fin not modified into a gonopodium; to form S-shaped arm. No ligament from the anal fin inclinators enlarged; first proximal ventral arms of maxillaries to middle of ros- radial present; middle anal radials present. tral cartilage; no ethmomaxillary ligament; 526 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 no meniscus between premaxilla and maxil- of the first vertebra to the skull with supra- la. Uni- or bi-cuspid uniserial outer teeth. occipital forming, rather than excluded from, Dentary with medial extension, robust; the dorsal wall of the foramen magnum, and coronoid process on dentary not overlapping neurapophyses of the first vertebra angled with that of articular; retroarticular not elon- forward and firmly applied to the skull, and gate. Autopalatine with head angled ante- no exoccipital condyles; and pharyngobran- riorly; ventral process elongate reaching chial teeth in discrete rows. quadrate; metapterygoid absent. COMPOSITION: Five genera and approxi- Orbital rim free; anterior nans not tubular; mately 40 species: Cyprinodon Lacepede, cephalic sensory pore system represented by Megupsilon Miller and Walters, Jordanella derived, lyre-shaped pattern of minute neu- Goode and Bean, Cualac Miller and Flori- romasts. dichthys Hubbs. Females larger than males; all fins round- DISTRIBUTION: North and Middle America ed; scales present or absent. to Honduras; the West Indies southward to No fatty predorsal ridge; swimbladder Venezuela (fig. 88). does not extend posteriorly beyond hemal FOSSILS: In addition to the species referred spines. to the genus Cyprinodon (Miller, 1945) of DISTRIBUTION: (fig. 85) High-altitude lakes California, a fossil genus, Carrionellus of the South American continental divide. (White, 1927) from the Lower Miocene of Known range from Lago Ascotan, Chile Ecuador has been referred to the cyprino- north to Lago Llascha, Peru. dontines on the basis of its overall fin posi- REMARKS: The distribution above repre- tion and the possession of tricuspid teeth in sents a range extension of the genus. Speci- the jaws. The outer teeth, however, are in mens were collected by Tim Hardin of Col- two rows, not one as in the cyprinodontines. orado State University on June 13, 1979. The I suggest that Carrionellus is perhaps a char- placement of the genus Orestias in the tribe acoid rather than a cyprinodont since in that Orestiini along with the Anatolian cyprino- group biserial tricuspid teeth are not uncom- dontines represents the first statement con- mon, and the condition is unknown in the cerning its relationship to other cyprinodon- cyprinodontines. tiforms since the tentative statements by Eigenmann (1920) and Foster (1967). GENUS CYPRINODON LACEPEDE MATERIAL EXAMINED: O. cuvieri: Lake Titicaca: SU 9331 (1); BMNH 1944.6.6.: 1-6 Cyprinodon Lacepede, 1803, p. 486 (type species (6); O. pentlandi: Lake Titicaca: Puno Bay: Cyprinodon variegatus Lacepede by original AMNH 1117 (1), BMNH 1944.6.6.=22-25 (3); designation). Prinodon Rafinesque, 1815, p. 88 (type species O. agassi: Peru: Rio Caminaque, near Lake Cyprinodon variegatus Lacepede, by original Titicaca: CAS 42534 (20); O. mooni: Lake designation [proposed as a substitute for Cy- Titicaca: Puno Bay: Syntype, USNM 133139 prinodon which was considered to be too long (1); O. polonorum: Peru: : Type, a name]). BMNH 1944.6.6.:223 (1); species unidenti- Encrotes Gistel, 1848, p. 9 (type species Cyprin- fied: Peru: Challhuacocha: CAS 40700 (2*1 odon variegatus Lacepede, by original desig- 42); Bolivia: tributary of Lake Titicaca: nation [proposed as a substitute for Lebia or AMNH 20355SW (2*/60); AMNH 20353SW Lebias Cuvier]). (2+/8); Peru: Lago Llascha: AMNH 38411(3). Trifarcius Poey, 1860, p. 306 (type species Tri- farcius riverendi Poey, by original designation).

TRIBE CYPRINODONTINI GILL ETYMOLOGY: Cyprinodon from Cyprinus and odon, meaning teeth, referring to the DIAGNOSIS: Distinguished from all other genus as a minnow with teeth. cyprinodontiforms by the following derived COMPOSITION: Approximately 36 species characters: a derived form of the attachment as listed in Lazara (1979). 1981 PARENTI: CYPRINODONTIFORM FISHES 527

DIAGNOSIS: Cyprinodontines with an en- sensory canal; dermosphenotic and preoper- larged scapular process. cular with distinct sensory canal; pectoral DEFINITIONS: Anal: (i), 9-10; Dorsal: ii, 9; girdle low-set; first postcleithrum present; Pelvic: 0-7; Pectoral: 14-16; Caudal: 7, 12, scapular process enlarged; posttemporal 7; Vertebrae: 12+12-12+14. Gill rakers on with ossified lower limb; posttemporal not the anterior arm of the first arch: 14-23. fused to supracleithrum. Branchiostegal rays: 5; Scales lateral series: Vomer ossified, edentulous; medial arm of 24-28. maxilla straight with pronounced dorsal pro- First pleural rib on parapophysis of second cesses with a groove, nearly meeting in the vertebra; parapophysis not reduced; no pleu- midline; ventral arms narrow, not abutting ral ribs on hemal spines; hypural plates fused rostral cartilage; outer arm robust. into hypural fan; Epipleural ribs not bifid. Premaxillary ascending processes narrow Anal fin not modified into a gonopodium; and reduced, not tapered posteriorly and not anal fin musculature unmodified; first proxi- overlapping in the midline; rostral cartilage mal radial present; middle anal radials pres- reduced or absent; outer arm of premaxilla ent. with alveolar process, indented posteriorly Spermatozeugmata not formed; fertiliza- to form S-shaped arm; No ligament from the tion external; development nonannual; ovip- ventral arms of maxillaries to middle of ros- arous. Eggs round. tral cartilage; no ethmomaxillary ligament; One dorsal ray articulating with the first no meniscus between premaxilla and maxil- two dorsal radials; dorsal fin origin opposite la. Tricuspid, uniserial outer teeth. origin of anal. Dentary expanded medially, robust; cor- Autopterotic fossa reduced; lateral eth- onoid process on dentary not overlapping moid expanded medially, reaching parasphe- with that of articular; retroarticular not elon- noid and lying perpendicular to frontal; para- gate. Autopalatine with head angled ante- sphenoid not expanded anteriorly; no riorly; ventral process elongate reaching supraoccipital and exoccipital processes; quadrate; metapterygoid absent. neural arches of first vertebra open and ap- Orbital rim free; anterior naris not tubular; plied to skull; first vertebra articulates with supraorbital sensory pores 1-7; seven pre- skull via basioccipital condyles; exoccipital opercular pores; two mandibular pores; condyles absent; supraoccipital included in three lacrimal pores. formation of foramen magnum; parietals ab- Females and males of equal sizes or males sent; nasals expanded medially. larger; males without fin extensions; males Mesethmoid ossified; medial processes of often with a dark caudal margin; usually a pelvic fin base and ischial process not re- spot at the base of dorsal fin. duced, or pel vies absent; interarcual carti- No fatty predorsal ridge; swimbladder lage reduced, attaches laterally to second does not extend posteriorly beyond hemal pharyngobranchial with a bony flange; phar- spines. yngobranchial teeth arranged in discrete DISTRIBUTION: North and Middle Ameri- rows; basihyal long and narrow; no tooth patches on second and third hypobranchials; ca, the West Indies to Venezuela. teeth on fourth ceratobranchials; dorsal hy- REMARKS: Since all species have not been pohyal absent; anterior extension of anterior examined, Cyprinodon is only tentatively ceratohyal ventral to ventral hypo- identified as monophyletic by a greatly en- hyal; uncinate process on fourth epibranchial larged scapular process. articulates with that of third; first epibran- MATERIAL EXAMINED: C. variegatus: chial wide at its base; interhyal ossified; two New York: Long Island: AMNH 36072 (4*/ ossified basibranchials. Vomer with poste- 97); AMNH 21800 (1+); Alabama: AMNH rior extension ventral to parasphenoid. 35750 (4). C. diabolis: Nevada: Devil's Hole: Lacrimal flat and broad, carrying a distinct CAS 22994 (16); C. bondi: Haiti: Etang Sau- 528 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 matre AMNH 377341 (12); C. macularius: Mesethmoid ossified; medial processes of Nevada: Nye Co., AMNH 20232 (17). pelvic fin base and ischial process not re- duced; interarcual cartilage reduced, at- taches laterally to second pharyngobranchial GENUS JORDANELLA GOODE AND BEAN toothplate with a bony flange; pharyngobran- Jordanella Goode and Bean, 1879, p. 117 (type chial teeth arranged in discrete rows; basi- species Jordanella floridae Goode and Bean, hyal long and narrow; no tooth patches on by original designation). second and third hypobranchials; teeth on Garmanella Hubbs, 1936, p. 218 (type species fourth ceratobranchials; dorsal hypohyal ab- Garmanella pulchra Hubbs, by original desig- sent; anterior extension of anterior cerato- nation). hyal ventral to ventral hypohyal; uncinate ETYMOLOGY: Jordanella in honor of the process on fourth epibranchial articulates American ichthyologist . with that of third; first epibranchial wide at TYPES: Florida: Jordanella floridae Goode its base; interhyal ossified; two ossified ba- and Bean, Types, USNM 22903 (3). sibranchials. Vomer with posterior extension COMPOSITION: Two species floridae Goode ventral to parasphenoid. and Bean, and pulchra (Hubbs). Lacrimal flat and broad, carrying a distinct DIAGNOSIS: Distinguished from other cy- sensory canal; dermosphenotic and preoper- prinodontoids by an elongate dorsal fin of 15 cular with distinct sensory canal; pectoral or more rays, and a dark suborbital bar. girdle lowset; first postcleithrum present; DEFINITION: Anal: i, 10; Dorsal: (i or I) posttemporal with ossified lower limb; post- 14-17; Pelvic: 6-7; Pectoral: 15-16; Caudal: temporal not fused to supracleithrum. 6, 18, 6; Vertebrae: 12+14; Gill rakers on the Vomer ossified, edentulous; medial arm of anterior arm of the first arch: 10-12. Bran- maxilla straight with pronounced dorsal pro- chiostegal rays: 5; Scales lateral series: 24- cesses with a groove, nearly meeting in the 26. midline; ventral arms narrow, not abutting First pleural rib on parapophysis of second rostral cartilage; outer arm robust. vertebra; parapophysis not reduced; no pleu- Premaxillary ascending processes narrow ral ribs on hemal spines; hypural plates fused and reduced, not tapered posteriorly and not into hypural fan. Epipleural ribs not bifid. overlapping in the midline; rostral cartilage Anal fin not modified into a gonopodium; reduced or absent; outer arm of premaxilla anal fin musculature unmodified; first proxi- with alveolar process, indented posteriorly mal radial present; middle anal radials pres- to form S-shaped arm. No ligament from the ent. ventral arms of maxillaries to middle of ros- Spermatozeugmata not formed; fertiliza- tral cartilage; no ethmomaxillary ligament; tion external; development nonannual; ovip- no meniscus between premaxilla and maxil- arous. Eggs round. la. Tricuspid, uniserial outer teeth. One dorsal ray articulating with the first Dentary expanded medially, robust; cor- two dorsal radials; dorsal fin origin opposite onoid process on dentary not overlapping origin of anal. with that of articular; retroarticular not elon- Autopterotic fossa reduced; lateral eth- gate. Autopalatine with head angled ante- moid expanded medially, reaching para- riorly; ventral process elongate reaching sphenoid and lying perpendicular to frontal; quadrate; metapterygoid absent. parasphenoid not expanded anteriorly; no Orbital rim free; anterior naris not tubular; supraoccipital and exoccipital processs; supraorbital sensory pores 1-7 or 1-3, 4-7; neural arches of first vertebra open and ap- seven preopercular pores; no mandibular plied to skull; first vertebra articulates with pores; three lacrimal pores. skull via basioccipital condyles; exoccipital Females and males of equal sizes or males condyles absent; supraoccipital included in larger; all fins rounded. Pigmentation pattern formation of foramen magnum; parietals ab- consisting of a prominent midlateral blotch, sent; nasals expanded medially. and a suborbital bar. 1981 PARENTI: CYPRINODONTIFORM FISHES 529

No fatty predorsal ridge; swimbladder anal fin musculature unmodified; first proxi- does not extend posteriorly beyond hemal mal radial present; middle anal radials pres- spines. ent. DISTRIBUTION: Florida and the Yucatan Spermatozeugmata not formed; fertiliza- peninsula, south to Belize. tion external; development nonannual; ovip- REMARKS: All workers on the cyprinodon- arous. Eggs round. tines have remarked on the derived similarity One dorsal ray articulating with the first of the two nominal genera Jordanella and two dorsal radials; dorsal fin origin opposite Garmanella but preferred to keep them dis- origin of anal. tinct. Since each genus is monotypic, it is in Autopterotic fossa reduced; lateral eth- the interest of having generic categories de- moid expanded medially, reaching parasphe- fine derived groups, rather than recognize in- noid and lying perpendicular to frontal; para- dividual differences, that these two genera sphenoid not expanded anteriorly; no are synonymized. supraoccipital and exoccipital processes; MATERIAL EXAMINED: J. floridae: Flori- neural arches of first vertebra open and ap- da: the types as listed above; AMNH 2769 plied to skull; first vertebra articulates with (3); Collier Co. AMNH 22060 (6+); Aquari- skull via basioccipital condyles; exoccipital um material: AMNH 38410SW (3*). J. pul- condyles absent; supraoccipital included in chra: Yucatan: Paratypes: USNM 117542 formation of foramen magnum; parietals ab- (2); USNM 192329 (1*/19); Belize: FMNH sent; nasals expanded medially. 82343 (l*/3). Mesethmoid ossified; medial processes of pelvic fin base and ischial process not re- GENUS CUALAC MILLER duced; interarcual cartilage reduced, at- taches laterally to second pharyngobranchial Cualac Miller, 1956, p. 1 (type species Cualac with a bony flange; pharyngobranchial teeth tessellatus Miller by original designation). arranged in discrete rows; basihyal long and ETYMOLOGY: Cualac derived from a Mex- narrow; no tooth patches on second and ican place name of Nahuatl origin meaning third hypobranchials; teeth on fourth cera- where there is good water (Miller, 1956). tobranchials; dorsal hypohyal absent; ante- TYPES: Mexico: San Luis Potosi: outlet rior extension of anterior ceratohyal ventral ditch of La Media Luna, 7 mi. SSW of set- to ventral hypohyal; uncinate process on tlement of Rio Verde: Cualac tessellatus fourth epibranchial articulates with that of Miller, Holotype, UMMZ 17135 (1). third; first epibranchial wide at its base; in- COMPOSITION: Solely the type species. terhyal ossified; two ossified basibranchials. DIAGNOSIS: Distinguished from all other Vomer with posterior extension ventral to cyprinodontiform fishes by having closely parasphenoid. packed villiform inferior pharyngeal teeth, Lacrimal flat and broad, carrying a distinct and from other New World cyprinodontines sensory canal; dermosphenotic and preoper- by an increase in gill rakers on the first arch cular with distinct sensory canal; pectoral to 17. girdle lowset; first postcleithrum present; DEFINITION: Anal: (i) 9-10; Dorsal: (i) 9- posttemporal with ossified lower limb; post- 11; Pelvic: 6-8. Pectoral: 12-13; Caudal: 7, temporal not fused to supracleithrum. 14,7; Vertebrae: 14+15; Gill rakers on the Vomer ossified, edentulous; medial arm of anterior arm of the first arch: 14-23. Bran- maxilla straight with pronounced dorsal pro- chiostegal rays: 5; Scales lateral series: 26- cesses with a groove, nearly meeting in the 29. midline; ventral arms narrow, not abutting First pleural rib on parapophysis of second rostral cartilage; outer arm robust. vertebra; parapophysis not reduced; no pleu- Premaxillary ascending processes narrow ral ribs on hemal spines; hypural plates fused and reduced, not tapered posteriorly and not into hypural fan. Epipleural ribs not bifid. overlapping in the midline; rostral cartilage Anal fin not modified into a gonopodium; reduced or absent; outer arm of premaxilla 530 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 with alveolar process, indented posteriorly subspecies: carpio (Giinther), barbouri to form S-shaped arm. No ligament from the Hubbs and polymnus Hubbs. ventral arms of maxillaries to middle of ros- DIAGNOSIS: Distinguished from all other tral cartilage; no ethmomaxillary ligament; cyprinodontoids by having an ossified first no meniscus between premaxilla and maxil- pharyngobranchial toothplate which bears a la. Tricuspid, uniserial outer teeth. patch of teeth. Dentary expanded medially, robust; cor- DEFINITION: Anal: i, 8; Dorsal: ii, 9; Pel- onoid process on dentary not overlapping vic: 6-7; Pectoral: 18; Caudal: 9, 13, 9; Ver- with that of articular; retroarticular not elon- tebrae: 10+13; Gill rakers on the anterior gate. Autopalatine with head angled ante- arm of the first arch: 9. Branchiostegal rays: riorly; ventral process elongate reaching 5-6; Scales lateral series: 23-25. quadrate; metapterygoid absent. First pleural rib on parapophysis of second Orbital rim free; anterior nans not tubular; vertebra; parapophysis not reduced; no pleu- supraorbital sensory pores l-2a, 2b-6a, 6b- ral ribs on hemal spines; hypural plates fused 7; seven preopercular pores; mandibular and into hypural fan. Epipleural ribs not bifid. lacrimal pores replaced by neuromasts. Anal fin not modified into a gonopodium; Females and males of equal sizes; all fins anal fin musculature unmodified; first proxi- rounded or truncate; pigmentation pattern mal radial present; middle anal radials pres- characterized by a lateral band and faint re- ent. ticulations on the dorsal and anal fins, more Spermatozeugmata not formed; fertiliza- pronounced in males. tion external; development nonannual; ovip- No fatty predorsal ridge; swimbladder arous. Eggs round. does not extend posteriorly beyond hemal One dorsal ray articulating with the first spines. two dorsal radials; dorsal fin origin opposite DISTRIBUTION: San Luis Potosi, Mexico. origin of anal. REMARKS: Cualac was proposed by Miller Autopterotic fossa reduced; lateral eth- (1956) as an intermediate between the moid expanded medially, reaching parasphe- subfamilies Fundulinae and Cyprinodontinae noid and lying perpendicular to frontal; para- because it has a rather elongate body form sphenoid not expanded anteriorly; no as in the fundulines, and possesses tricuspid supraoccipital and exoccipital processes; outer teeth and the firm attachment of the neural arches of first vertebra open and ap- first vertebra to the skull as in the cyprino- plied to skull; first vertebra articulates with dontines. The characters it shares with the skull via basioccipital condyles; exoccipital cyprinodontines are derived while those it condyles absent; supraoccipital included in shares with the fundulines are primitive. formation of foramen magnum; parietals ab- Therefore, the genus is placed in the tribe sent; nasals expanded medially. Cyprinodontini. Mesethmoid ossified; medial processes of MATERIAL EXAMINED: Mexico: San Luis pelvic fin base and ischial process not re- Potosi: Paratopotypes: SU 50213 (1*716). duced; interarcual cartilage reduced, at- taches laterally to second pharyngobranchial GENUS FLORIDICHTHYS HUBBS with a bony flange; pharyngobranchial teeth arranged in discrete rows; basihyal long and Floridichthys Hubbs, 1926, p. 16 (type species narrow; no tooth patches on second and Cyprinodon carpio Giinther, by monotypy). third hypobranchials; teeth on fourth cera- ETYMOLOGY: Floridichthys, after Florida, tobranchials; dorsal hypohyal absent; ante- to which the genus was believed to be en- rior extension of anterior ceratohyal ventral demic. to ventral hypohyal; uncinate process on TYPES: Florida: Cyprinodon carpio Giin- fourth epibranchial articulates with that of ther, Type, BMNH 1855.9.19:821-825 (1). third; first epibranchial wide at its base; first COMPOSITION: The type divided into three epibranchial toothplate with patch of unicus- 1981 PARENTI: CYPRINODONTIFORM FISHES 531 pid teeth; interhyal ossified; two ossified ba- Florida: Key West: AMNH 2610 (1); Sara- sibranchials. Vomer with posterior extension sota: 17079 (2); no data: AMNH 21908SW ventral to parasphenoid. (1+/1), AMNH 21905SW (1+/1), AMNH Lacrimal flat and broad, carrying a distinct 21907SW (3+/3). sensory canal; dermosphenotic and preoper- cular with distinct sensory canal; pectoral GENUS MEGUPSILON MILLER AND WALTERS girdle lowset; first postcleithrum present; posttemporal with ossified lower limb; post- Megupsilon Miller and Walters, 1972, p. 2 (type temporal not fused to supracleithrum. species, Megupsilon aporus Miller and Wal- Vomer ossified, edentulous; medial arm of ters, by original designation). maxilla straight with pronounced dorsal pro- ETYMOLOGY: Megupsilon from the Greek cesses with a groove, nearly meeting in the megas, meaning great, and upsilon, the midline; ventral arms narrow, not abutting name of the greek letter Y, in reference to rostral cartilage; outer arm robust. the large Y-chromosome in the male. Premaxillary ascending processes narrow TYPES: Mexico: Nuevo Leon: El Potosi: and reduced, not tapered posteriorly and not Megupsilon aporus Miller and Walters, Ho- overlapping in the midline; rostral cartilage lotype: UMMZ 189018 (1). reduced or absent; outer arm of premaxilla COMPOSITION: Solely the type species. with alveolar process, indented posteriorly DIAGNOSIS: Distinguished from all other to form S-shaped arm. No ligament from the cyprinodontiform fishes by having an en- ventral arms of maxillaries to middle of ros- larged Y-chromosome in the male; sexually tral cartilage; no ethmomaxillary ligament; dimorphic chromosome numbers; and a lack no meniscus between premaxilla and maxil- of cephalic sensory pores. la. Tricuspid, uniserial outer teeth. DEFINITION: Anal: (0,1) 9-11; Dorsal: 9- Dentary expanded medially, robust; cor- 11; Pelvic: 0; Pectoral: 13-15; Caudal: 18; onoid process on dentary not overlapping Vertebrae: 11-12+13-16. Gill rakers on the with that of articular; retroarticular not elon- anterior arm of the first arch: 8-13; Bran- gate. Autopalatine with head angled ante- chiostegal rays; 5; Scales lateral series: 24- riorly; ventral process elongate reaching 26. quadrate; metaperygoid absent. First pleural rib on parapophysis of second Orbital rim free; anterior nans not tubular; vertebra; parapophysis not reduced; no pleu- supraorbital sensory pores 1-7; seven pre- ral ribs on hemal spines; hypural plates fused opercular pores; three mandibular pores; into hypural fan. Epipleural ribs not bifid. three lacrimal pores. Anal fin not modified into a gonopodium; Females and males of equal sizes or males anal fin musculature unmodified; first proxi- larger; fins rounded or truncate; pigmenta- mal radial present; middle anal radials pres- tion pattern characterized by a series of gold- ent. en crossbars. Spermatozeugmata not formed; fertiliza- No fatty predorsal ridge; swimbladder tion external; development nonannual; ovip- does not extend posteriorly beyond hemal arous. Eggs round. spines. One dorsal ray articulating with the first DISTRIBUTION: Florida and the Yucatan two dorsal radials; dorsal fin origin opposite peninsula, south to Honduras. origin of anal. REMARKS: Floridichthys is the only Autopterotic fossa reduced; lateral eth- cyprinodontoid to possess an ossified first moid expanded medially, reaching parasphe- pharyngobranchial toothplate that also has noid and lying perpendicular to frontal; para- teeth. A similar but possibly not homologous sphenoid not expanded anteriorly; no state occurs in one species of Cynolebias. supraoccipital and exoccipital processes; MATERIAL EXAMINED: Yucatan: the types neural arches of first vertebra open and ap- as listed above; Belize: AMNH 24632 (3); plied to processes; neural arches of first ver- 532 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

tebra open and applied to skull; first vertebra and reduced, not tapered posteriorly and not articulates with skull via basioccipital con- overlapping in the midline; rostral cartilage dyles; exoccipital condyles absent; supra- reduced; outer arm of premaxilla with alveo- occipital included in formation of foramen lar process, indented posteriorly to form magnum; parietals absent; nasals expanded S-shaped arm; No ligament from the ventral medially. arms of maxillaries to middle of rostral car- Mesethmoid ossified; pelvic fins and fin tilage; no ethmomaxillary ligament; no me- supports absent; interarcual cartilage re- niscus between premaxilla and maxilla. Tri- duced, attaches laterally to second pharyn- cuspid, uniserial outer teeth. gobranchial with a bony flange; pharyngo- Dentary expanded medially, robust; cor- branchial teeth arranged in discrete rows; onoid process on dentary not overlapping basihyal long and narrow; no tooth patches with that of articular; retroarticular not elon- on second and third hypobranchials; teeth on gate. Autopalatine with head angled ante- fourth ceratobranchials; dorsal hypohyal ab- riorly; ventral process elongate reaching sent; anterior extension of anterior cerato- quadrate; metapterygoid absent. hyal ventral to ventral hypohyal; uncinate Orbital rim free; anterior nans not tubular; process on fourth epibranchial articulates supraorbital sensory pores absent or re- with that of third; first epibranchial wide at placed by neuromasts; no preopercular its base; interhyal ossified; two ossified ba- pores; no mandibular pores; no lacrimal sibranchials. Vomer with posterior extension pores. ventral to parasphenoid. Females larger than males; fins rounded or Lacrimal flat and broad, carrying a distinct truncate; males with blackened scales on sensory canal; dermosphenotic and preoper- sides of body. cular with a distinct sensory canal; pectoral No fatty predorsal ridge; swimbladder girdle lowset; first postcleithrum present; does not extend posteriorly beyond hemal posttemporal with ossified lower limb; post- spines. temporal not fused to supracleithrum. DISTRIBUTION: Nuevo Leon, Mexico. Vomer ossified, edentulous; medial arm of REMARKS: Meristic data ranges for this ge- maxilla straight with pronounced dorsal pro- nus are supplemented by data from Miller cesses with a groove, nearly meeting in the and Walters (1972). midline; ventral arms narrow, not abutting MATERIAL EXAMINED: Mexico: Nuevo rostral cartilage; outer arm robust. Leon, spring-fed pond at Potosi: Paratypes: Premaxillary ascending processes narrow AMNH 38405 (ex. UMMZ 189020) (2*/10). HISTORICAL BIOGEOGRAPHY Historical distributions of organisms are and its biota evolved together. In striking interpreted traditionally in terms of the iden- contrast to both the dispersalist model and tification of a center of origin or dispersal. the Progression Rule, the aim of vicariance This is defined as the area of greatest den- biogeography is to interpret the distribution sity, or of the location of the most advanced of a group in terms of its relationship to a members. Once such a center is chosen, the general pattern exhibited by all other organ- distribution of a group of organisms may be isms inhabiting the same area. Dispersalists explained by its dispersal from that center and Progression Rule biogeographers typi- throughout its range (Darlington, 1957). The cally interpret the distribution of a group as ability of an organism to disperse a given dis- if it evolved not only in isolation from other tance is termed that organism's vagility, and organisms, but with little relation to the geo- is typically estimated from the observed ac- logical or geographic history of the earth as tivity. well. Aquatic organisms are supposedly restrict- Vicariance biogeography as outlined by ed in ability to disperse by their ability to Croizat and other workers deals with the rec- survive in marine, fresh, or brackish waters. ognition of tracks of organisms which are Cyprinodontiform fishes have been termed defined by the coincident distributions of secondary freshwater fishes since many many groups. A large number of groups shar- members typically enter brackish water ing the same distribution are logically in- either seasonally or throughout the year. The ferred to have shared an ancestral distribu- widespread distribution of the group (fig. 1), tion as outlined by the limits of the biota. therefore, has been explained by dispersalist The assessment of a dispersal is viewed as biogeographers (Myers, 1933a; Kosswig, a parsimony problem; that is, if there were 1943; Sethi, 1978) as a result of a dispersal 10 taxa exhibiting the same distribution and from a Mediterranean (Tethys Sea) center of one exhibiting a slightly different one, the origin during the late . unique distribution is postulated to be caused One alternative method of biogeographic by a dispersal of that taxon away from the analysis has been proposed by Hennig (1966) rest of the biota (Rosen, 1976). and exemplified for the midges by Brundin Vicariance biogeography has been com- (1966). This method, termed the Progression bined with the theories of phylogenetic sys- Rule, assumed that the most primitive mem- tematics to become what is termed cladistic bers of a group occupy the group's center of biogeography. As outlined by Rosen (1978) dispersal, and the more derived members and Platnick and Nelson (1978), the aspects have arrived at their present distribution by of a group to be compared are not solely the dispersing from that center. Although it distributions of those organisms, but their takes into consideration the phylogeny of a phylogenies as well. group of organisms, the Progression Rule is Introduced into the analysis was the trans- essentially a center of origin/dispersal model. formation of cladograms of taxa into clado- It assumes that cladistically apomorph or- grams of areas, which in turn could be ana- ganisms are better at dispersing than their lyzed using the parsimony method. An area cladistically plesiomorph relatives, a general is then treated as an area state, and its rela- assumption that is unsupportable. tion to other area states, as expressed by the A third method of analysis has been pro- phylogeny, are treated as character states in posed principally by Croizat and others the analysis (Parent!, 1981; Rosen, 1978). (Croizat, 1958, 1964; Croizat, Nelson and Once this is done for seveal groups of organ- Rosen, 1974). The method, now alternately isms with coincident distributions, a pattern termed vicariance or cladistic biogeography of earth history is suggested. The pattern is (although it was not proposed as such) has independent of geological hypotheses; how- as its main premise the idea that the world ever, a proposed geological model may fit 534 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 such a pattern. If none is found, this does land mass which began its disruption in the not necessarily suggest that the phylogeny of late Triassic. An alternative theory of earth these organisms is incorrect, but that the history has been advanced by Nur and Ben- geological models may be inappropriate. Avraham (1977) and Shields (1979) in which The role of fossils in biogeographic studies another ancient continent, Pacifica, existed has traditionally been as the estimator of the until starting to break up during the , age of origin or earliest time of dispersal into opening the Pacific for the first time in mod- a region (Darlington, 1957). Proponents of ern history. Transpacific distributions would the Progression Rule also supported this ar- support such a theory; however, none exists gument. Vicariance biogeographers have for cyprinodontiform fishes. That cyprino- emphasized that a fossil can only give the dontiforms are uninformative with respect to minimum estimate of the age of a group, and the existence of a Pacific continent could be that most groups must be older than their an indication of a difference in the absolute oldest fossil representative. age of Pacific versus Pangean groups. This The precise role fossils play in biogeo- suggests that such competing theories of graphic studies has been summarized most earth history may not be in competition at recently and debated by Patterson (1981) and all, but rather that they explain the distri- Parent! (1981). Patterson concurred with oth- bution of groups that originated at different ers recently addressing the problem (e.g., times. Rosen, 1976) that a fossil may be used as an The late Triassic is given as an estimate of arbiter between dispersal and in the minimum age of cyprinodontiforms since lieu of parsimony. For example, if an incon- by that time the Laurasian and Gondwanian gruent distribution was expressed by a land masses had begun their separation. A group, and such member fossils, then one more precise estimate of a minimum age could know that the fossils represented an would be the last time these land masses ancestral distribution rather than a dispersal. were still in contact. The late Triassic, there- Parent! argued that even though an organism fore, is used as an estimate only until a more could not have dispersed since the time it precise date can be given. was fossilized, previous to that time it could The oldest described cyprinodont fossil is have dispersed as well as any other member a species referred to the genus Prolebias, an of the group; therefore, fossils did not auto- Anatolian cyprinodontine from the Oligo- matically indicate the limits of an ancestral cene of Europe (Sauvage, 1874). The distri- biota unless an estimate of the age of that bution of the group, however, supports the biota was specified. contention that cyprinodontiforms could be GENERAL PATTERN OF CYPRINODONTI- much older. The oldest fossil does give a FORMS: The present-day distribution of cy- minimum age of the group; however, a more prinodontiforms (fig. 1) seems to be an ex- reliable indicator is the minimum age of the panded Gondwanian or reduced Pangean group's distributional pattern. This may be pattern with members absent from Australia, estimated as the age of the oldest fossil of a Antarctica, and the Orient but present in group exhibiting the pattern, or, without North America. No members are found east such evidence, reference to a particular geo- of Wallace's line, a classic line of demarca- logical pattern. For example, cichlids have tion in the Indo-Australian region. The dis- a distribution corresponding, in part, to that tribution is approximated roughly in fishes of the cyprinodontiforms, and their pattern by the Ostariophysi minus the Oriental cyp- of relationships falls into the same general rinids and Australian plotosids (Nelson, pattern. Therefore, one could also estimate 1976) and the synbranchids minus the Aus- the age of cichlids as at least the late Trias- tralian and Oriental components (Rosen, sic. If a fossil of an older age was found for 1976). either group, then the minimum age of both The pattern is interpreted as Pangean in groups should be re-estimated at that greater part based on a reconstruction of the ancient age. This conclusion follows the most par- 1981 PARENTI: CYPRINODONTIFORM FISHES 535

WESTERN

NORTH, CENTRAL & ,w?B!r» CENTRAL NORTH & SOUTH AMERICA & CENTRAL 4 AMERICA JAMAICA & SOUTH AMERICA & NORTH & REGION AMERICA CENTRAL AMERICA MEDITERRANEAN AFRICA SOUTH AMERICA &MEXICO CUBA EURASIA CENTRAL AMERICA

FIG. 90. Cladogram of areas for all cyprinodontiforms (after fig. 9).

simonious assumption; that is, if two groups ichthys and the goodeids. The South Amer- share a pattern, their history of distribution, ican-Caribbean track is associated with the and hence age of origin, must correspond distribution of Jenynsia, Anableps, and Oxy- (see Parenti, 1981). Also, the oldest fossil is zygonectes, and the Neotropical aplochei- an unreliable indicator of the age of cyprin- loids. The poeciliids exhibit properties coin- odontiforms since, were there no fossil rec- cident with both patterns (Compare figs. 9 ord, the estimated minimum age would be and 90). the same. The association of Jamaica with Cuba (fig. PATTERNS WITHIN CYPRINODONTIFORMS: 90) representing the distribution of the genus To describe the association of sister groups Cubanichthys does not coincide with distri- with respect to their distribution, it is useful butions on these two Caribbean islands as to transform the cladogram of interrelation- discussed by Rosen (1976); however, pur ships (fig. 9) into a cladogram of areas (fig. poor knowledge of relationships of organ- 90), following the method suggested by Ro- isms in this region, as well as the sister group sen (1978). An association between South relationship of Cubanichthys to Old World America and Africa is repeated twice on the and New World cyprinodontines and Ores- overall cladogram, once corresponding to tias, precludes a conclusion concerning the the distribution of the tropical aplocheiloids, dispersal of one or the other species of this and again corresponding to the distribution genus. of the poeciliids, Fluviphylax, procatopines, The placement of the Mediterranean Val- and Pantanodon. encia on the cladogram of cyprinodontiforms The associations among North, Middle, indicates the polyphyletic nature of the Eur- and South America correspond to those asian killifishes. Valencia, although previ- found by Rosen (1976). His North American- ously classified as a funduline, has also been Caribbean track is associated with the North referred to the Anatolian cyprinodontines, to American-Central American distributions both of which it is inferred in this study to exhibited by the fundulines; New World cy- have no close relationship. prinodontines; and Empetrichthys, Cren- The aberrant pattern of the general distri- 536 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 bution is that expressed by the relationship 1981), and therefore appears to have some of the western South American Orestias and generality. the Anatolian cyprinodontines. Is the relationship of South American taxa Previous to this study, the sister group of to North American-European taxa dispersal Orestias had been speculated on by several of such taxa? Such is postulated by Marshall workers including Foster (1967). The south- (1979) who, on the basis of geological and ern South American aplocheiloid Cynolebias radioisotopic as well as fossil plant and ani- was postulated as its sister group based on mal data, concludes that South American cri- apparently derived head pore pattern and cetine rodents entered South America 7 Mill. osteological similarities. However, it is hy- YBP, at least four million years before the pothesized that these similarities represent presumed formation of the Panamanian land- convergences, and that Orestias is most bridge. Therefore, as a dispersalist, Marshall closely related to the Anatolian genus Kos- was forced to postulate waif dispersal for the swigichthys. rodents. The genus Orestias is endemic to the high- Assuming that waif dispersal is possible, altitude lakes of the Andes. Its distribution any group could have arrived in South Amer- (fig. 85) was believed (Villwock, 1963) to be ica at any time in history, leaving the dis- restricted to the region between Lake Junin, persalist's theory untestable. There are no Peru and northern Chile; however, recent fossil Orestias on which to base such a the- collections of the genus indicate it is distrib- ory. Time of formation of the Andes, how- uted much farther north (the known limit ever, provides an estimate of a minimum age Lago Llascha, Peru), thus supporting the of the group, or latest time of dispersal into idea that it is distributed extensively in the South America, if that occurred. The uplift lakes all along the continental divide. The of the modern Andes occurred during the genus has no fossil record. period from the Late Jurassic through the Anatolian cyprinodontines are currently Late (Dott and Batten, 1976). widely distributed in the Mediterranean re- The modern Andes were preceded by high- gion, along the periphery of the Saudi Ara- lands in western Venezuela, Colombia, Ec- bian Peninsula and into Turkey and Iran (fig. uador, and Peru which were a result of uplift 86). Fossils referable to the group are rela- and intrusion during the Permo-Triassic. If tively abundant; the oldest is from the Olig- we assume that Orestias was uplifted along ocene of Europe. with the Andes, then we must conclude they The extra-continental sister-group rela- are at least as old as the Late Cretaceous and tionship of South America-Eurasia is not lim- possibly older if Orestias is most closely re- ited to the Orestias-AnaXoli&n group. In the lated to a fish from the Anatolian region. dog family (Canidae) the Eurasian Nycte- The Anatolian cyprinodontines have a dis- reutes has a hypothesized sister group, the tribution corresponding to the ancient land genus Cerdocyon, found in the savanna- connections of Iran and Arabia, Turkey- grasslands of western South America from Greece and North Africa, Italy, Sardinia, Venezuela to Paraguay. There is also a pos- Corsica and southern France, and eastern sible fossil form referable to Cerdocyon in Spain, and northwestern Africa and Spain. North America (R. H. Tedford, personal The distribution is most simply explained by commun.). However, the close relationship the break-up of these land masses as the Te- of Cerdocyon to a North American taxon thys Sea became obliterated. If Orestias is would serve as a refutation of the close re- closely related to a member of the group, lationship of South America to Eurasia. A however, and the distribution is related to third set of relationships involving taxa of earth history at some level, two explanations western South America and a North Ameri- are possible: (1) there was once a connection can-European sister pair was supported by between western South America and the An- over 10 cladograms of areas (Patterson, atolian region, or (2) a monophyletic group 1981 PARENTI: CYPRINODONTIFORM FISHES 537

FIG. 91. Distributional limits of Nothobranchius and Aphyosemion. Darkly stippled area represents the distribution of Nothobranchius minus its most plesiomorphic species, thierryi. Cross-hatched area represents distribution of thierryi. Lightly stippled area represents distribution of Aphyosemion (in- cluding Fundulopanchax as in this study). (After Huber, 1978.) including Orestias and the Anatolian cyprin- This distribution could be interpreted as a odontines was once more extensive and has dispersal, although the relationship of the undergone widespread extinction. land masses today makes such an explana- Little support exists for the first pro- tion highly unparsimonious. posal. The second, however, is supported by We see the pattern as unusual because the distribution of canids, as well as of pleth- there is no general explanation for it. Rela- odontid salamanders (see Rosen, 1976). tionships of organisms consistent with a Pan- 538 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

EAST AFRICA, INDO MALAYSIAN MADAGASCAR, CENTRAL CENTRAL CENTRAL CENTRAL, EASTERN & REGION THE SEYCHELLES WESTERN AFRICA WESTERN AFRICA WESTERN AFRICA SOUTH AFRICA

FIG. 92. Cladogram of areas for Old World aplocheiloids (after fig. 25). gean break-up may not be part of the most exclusively, the more derived members of a general pattern of earth history; however, pair found in southern South America (e.g., the only way of discovering this is to discov- as for Austrofundulus and Cynolebias). Cen- er patterns of distribution that suggest some tral America is the area plesiomorphic to other explanation of earth history. The Pan- these two. gean model is not considered an adequate Such a pattern is repeated, in part, by Jen- explanation for the historical distribution of ynsia, Anableps, and Oxyzygonectes (fig. 94) cyprinodontiforms for it does not explain sat- in which the plesiomorphic Oxyzygonectes isfactorily the relationship among South inhabits the Pacific coast of Costa Rica, America, Eurasia, and North America sug- whereas the more derived pair of Anableps gested by cyprinodontiforms, canids, pleth- and Jenynsia inhabit Central America and odontids, and perhaps many other groups. northern South America and southern South Current geological hypotheses are uninfor- America, respectively. Anableps, however, mative with respect to relationships of such is sympatric with Oxyzygonectes suggesting areas. a more complicated distributional history PATTERNS OF MONOPHYLETIC SUBGROUPS: within Central America. Poeciliids inhabit all Within Africa, there is an east-west as well three areas, as well as part of North Ameri- as a north-south dichotomy (fig. 91). Noth- ca. However, the phylogeny of its subgroups obranchius thierryi inhabits western Africa, has not been worked out. whereas all other Nothobranchius species Given that the associations between South are found in eastern and southern Africa. America and Africa are supported twice The same is the case for species of Aphy- (figs. 93, 95), resolution of the relationship of osemion, the most primitive of which are Fluviphylax as the sister group of either the found west of the Dahomey Gap, whereas poeciliids or the procatopines or Pan- the more derived members and those more tanodon would not produce an astounding closely related to Nothobranchius and re- biogeographic pattern. Simpson (1977) pos- ferred to the genus Fundulopanchax are tulates that the two continental masses began found in the Congo Basin (fig. 92). rifting 130 mill. YBP. The final separation of Similarly, Neotropical aplocheiloids (fig. South America and Africa at the equatorial 93) show a repeated north/south South shear zone is estimated at 80 mill YBP, in American dichotomy, with typically, but not the Upper Cretaceous. Therefore, estimates 1981 PARENTI: CYPRINODONTIFORM FISHES 539

NORTHERN S.A. SOUTHERNS.A. NORTHERNS. 4 SOUTHERNS.A. NORTHERN S.t SOUTHERN $.t

FIG. 93. Cladogram of areas for Neotropical aplocheiloids (after fig. 20). of the ages of trans-South Atlantic relatives Valley system, and Crenichthys of eastern should be no earlier than 80 mill YBP, based Nevada are closely related to Fundulus, on Simpson's hypothesis. Confirmation to which in turn, are close relatives of Profun- some other, older geological hypothesis dulus. All four genera are postulated to be could support an older estimate, however. derived from a funduline "ancestral stock" Among the procatopines and Pantanodon (Uyeno and Miller, 1962). Since the oldest (fig. 95) several of the associations exhibited fossil of New world cyprinodonts is a Mio- by the Old World aplocheiloids are repeated. cene specimen referable to the genus Fun- That is, the close relationship of Madagascar dulus (Lugaski, 1977), all differentation of to east Africa, and a split between east and these genera has been postulated to have oc- west central Africa. Because procatopines curred since the Miocene (Uyeno and Miller, are found in the African Rift lakes the asso- 1962; Hubbs, Miller and Hubbs, 1974). The ciation of these fishes with the poeciliids and conclusion that the Cyprinodontidae is a Fluviphylax of the New World resembles, in nonmonophyletic group has led to the ex- part, the distribution of cichlids (Nelson, amination of other cyprinodontoids as pos- 1976). sible close relatives of subgroups of the fam- Interrelationships of cyprinodontiform ily. Two results of this study are the present fishes distributed exclusively in North and hypotheses that the Mexican goodeids are Central America (including Mexico and the the closest relatives of Empetrichthys and Caribbean) present a particular opportunity Crenichthys, and that Profundulus is primi- for interpretation of the biogeographic pat- tive to all other cyprinodontoids. This ne- terns previously discussed for these regions. cessitates a redefinition of biogeographic pat- No subfamily listed in table 2 has been dis- terns and hence a new estimate of the time rupted in the new classification more so than of differentiation of these genera. the Fundulinae; here, the group is elevated The area cladogram for New World cy- to family rank and restricted to five genera prinodontines (fig. 97) simply repeats the whose interrelationships are summarized in close relationship of Floridian and Yucatan- figure 75 and biogeographic associations in Central American taxa to taxa in central figure 96. Mexico. The North, Middle, and South A prevalent theory among cyprinodont American distribution of Cyprinodon is unin- workers is that Empetrichthys of the Death formative with respect to the three area pat- 540 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

SOUTHERN SOUTH CENTRAL AMERICA, PACIF IC COAST OF AMERICA NORTHERN S. A. COSTA RICA

FIG. 94. Cladogram of areas for Jenynsia, Anableps and Oxyzygonectes (after fig. 79). tern. However, a subspecies of C. variega- North and Middle America as well as Ber- tus, a species typically found in Florida and muda, Cuba, and Hispaniola, there are sev- along the East Coast of North America, has eral species associations between Florida been described from Yucatan (Hubbs, 1936). and the Yucatan including the similis-per- One fossil genus, Carrionellus from the similis group and the grandis-grandissimus Miocene of Ecuador, has been referred to group (Miller, 1955b). the cyprinodontines. As stated in the system- Adinia and Leptolucania, of Florida and atic section, however, this genus is probably coastal regions along the Gulf of Mexico are referable to the characoids rather than to the closely related to Lucania of the East Coast cyprinodontiforms. of North America and Mexico. The associ- Fundulines (fig. 96) share some distribu- ation of Cuba and Florida, supported several tion patterns with the cyprinodontines. With- times by the distribution of poeciliid species in the genus Fundulus, distributed through (Rosen and Bailey, 1963) as well as Rivulus,

CENTRAL AFRICA CENTRAL AFRICA CENTRAL AFRICA

FIG. 95. Cladogram of areas of poeciliids, procatopines, Fluviphylax and Pantanodon (after fig. 81). 1981 PARENTI: CYPRINODONTIFORM FISHES 541

MEXICO, CENTRAL NORTH NORTHS, MIDDLE EAST COAST TEXAS TO GEORGIA AMERICA AMERICA NORTH AMERICA FLORIDA FLORIDA,

FIG. 96. Cladogram of areas of fundulines (after fig. 75). is not supported here by Cubanichthys cu- Cyprinodon, Fundulus and Empetrichthys bensis which has no supported close rela- and Parafundulus (Miller, 1945; Uyeno and tionships to the Floridian , Miller, 1962). As stated, the oldest fossil is as suspected by Hubbs and Miller (1965). Miocene; therefore, differentiation of these Cyprinodontiforms west of the Rocky genera have been interpreted as resulting Mountains are represented by four Recent from events since the Miocene. However, it genera: Fundulus, Empetrichthys, Crenich- is again emphasized that a Miocene fossil thys, and Cyprinodon. [Lucania has been in- presents only a minimum estimate of the age troduced into San Francisco Bay and other of the group. Therefore, an age estimated by localities in California, Hubbs and Miller, the pattern of interrelationships as correlated 1965.] Fossils are referable to the genera with a geologic event will give a more useful

CARIBBEAN, NORTH & MIDDLE NUEVO LEON, CENTRAL AMERICA, CENTRAL AMERICA, SAN LUIS POTOSI, AMERICA MEXICO FLORIDA, YUCATAN FLORIDA, YUCATAN MEXICO

FIG. 97. Cladogram of areas of New World cyprinodontines (after fig. 89). 542 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

FIG. 98. Fossil affinities across western North and Middle America (after Axelrod, 1979). Open circled denote localities of fossils with affinities in the dry tropic forest (stippled) and temperate rain- forest (blackened). Present-day desert regions are cross-hatched.

estimate. The association of Empetrichthys cene-Pliocene also occur for various groups and Crenichthys with the goodeids of the of desert plants (Axelrod, 1979). Mexican Plateau enlarges the area for such The species of Cyprinodon, including the an analysis. nevadensis complex, diabolis, salinus, ra- Pleistocene and Miocene goodeids (Alva- diosus, and milleri are probably most closely rez and Arreola, 1972; Smith, Miller and related to the Cyprinodon species of eastern Cavender, 1975) allow for an estimate of the Mexico. Miocene for a minimum age of the ancestral The age of the regions of the Cordillera biota of these three taxa. inhabited by cyprinodontiforms may be es- The two Recent Californian species of timated from either a hypothesis of the age Fundulus, parvipinnis, and lima, the last of mountain building events, or age of fossils only of Baja California, are considered to be of groups showing similar distributions and sister species and in turn most closely related interrelationships. to diaphanus of east of the Rocky Mountains The recent formation of the Cordillera, a (Farris, 1968). Sister group pairs across the result of the Late Jurassic subduction of the Gulf of California, which opened in the Mio- Pacific plate, began near the Pacific Coast 1981 PARENTI: CYPRINODONTIFORM FISHES 543 and then moved eastward. In the Rocky MEXICAN PLATEAU DEATH VALLEY EASTERN SYSTEM NEVADA Mountains, evidence points toward a Late Cretaceous, early Cenozoic period of defor- mation (Dott and Batten, 1976). The last major epeiric seas of which the maximum transgression occurred about 100 mill. YBP was caused by a worldwide rise in sea-level during the Cretaceous. The earliest uplift of the Mexican Plateau and the high- lands of western Guatemala and southern FIG. 99. Cladogram of areas for goodeids, Mexico occurred by the Late Cretaceous. By Empetrichthys, and Crenichthys (after fig. 83). the early Cenozoic, most of North and Cen- tral America was approaching its present- day geological features. Thus, geological evi- formation of desert regions is inferred to dence does not preclude an age of origin of have disrupted larger, more widespread an- these cyprinodontiform fishes by Late Cre- cestral biotas. taceous times. The flora of western North America is rep- However, owing to the prevalent marine resented by Eocene taxa from central Cali- transgression, support from other organisms fornia northward, the Green River and as- is considered almost a prerequisite to push- sociated areas. These have their closest ing back the estimates of latest time of origin relatives in the temperate rain and dry trop- of the killifishes. Another group of organisms ical forests of Mexico which are not repre- with a similar distribution and yet much old- sented by fossils. Fossil taxa of the south- er fossils would help to support a more an- eastern also have affinities cient origin. with Recent plants of these regions in Mex- Axelrod (1979) has recently summarized ico (fig. 98). Thus, an ancient, widespread the available information on distribution of dicot forest spanning the western Cordilleran plants, both fossil and Recent, in western from northwestern North America south- North and Middle America in order to esti- ward into the highlands of Mexico and nu- mate a time of origin of the Sonoran Desert clear Central America existed prior to the flora. Also summarized were the major floral late Eocene. Members of this forest included associations of the other North and Middle representatives of the dicot genera Anemia, American desert regions. Ficus, Magnolia and Platanus. Axelrod sug- Ancient associations across desert envi- gested that the "pathway" for such a con- ronments, which Axelrod termed links nection was the Sierra Madre Occidentale, across the desert, led him to conclude that the eastern range not having been formed by the Sonoran Desert vegetation as well as as- that time. sociated floras throughout the Cordilleran re- Disjunct distributions of killifishes across gion is as old as the Late Cretaceous and the region are of course supported by the early Tertiary, having originated before mod- sister group relationship of Empetrichthys, ern desert formation. Crenichthys, and the goodeids (fig. 99), as Distributions of sister-group pairs between well as the Cyprinodon and Fundulus California and Arizona-Texas across the des- species, and the genus Lucania. One species ert imply at least a late Eocene-Oligocene of Lucania, interiorus, occupies the Cuatro connection. Cienegas basin, an aquatic environment cen- The development of the Sonoran Desert is tered in the Chihuahuan desert with a divei se seen by Axelrod (p. 11) as a: "gradual des- fish fauna (Minckley, 1969). Other species iccation during the Tertiary, changing grad- inhabit the coastal lowlands of the south- ually from a well-watered area to the present eastern United States. region of extreme drought." Therefore, the Soltz and Naiman (1978) summarized the 544 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168 distribution of the cyprinodont fishes of cies of Empetrichthys described, all but one Death Valley with regard to their relation- have become extinct in modern history. ship to the past connections of Pleistocene It could be argued that the distribution of lakes. The disruption of past aquatic connec- plants has nothing to do with that of fishes tions across much of the region inhabited by or that fishes are much younger than plants, these fishes no doubt produced much of the and therefore have distributions with little in specific differentiation. However, the pos- common. However, the point taken here is tulated sister-group relationship of Empe- first that the distribution of biotas rather than trichthys and Crenichthys to the goodeids in- individual groups of organisms should be in- dicates that generic differentiation among the vestigated in a biogeographic analysis, and cyprinodonts of the West is much older. second that the minimum age of a group may The distribution of Empetrichthys, Cren- be estimated by its oldest fossil representa- ichthys, and the goodeids corresponds with tive or the oldest fossil of a group which that of the ancient dicot forest (fig. 98). Em- shares the pattern. An early estimate for the petrichthys is limited to the Death Valley age of origin of cyprinodonts of the West is system today, whereas Crenichthys has a further supported by the assessment of Pro- limited distribution in southeastern Nevada fundulus, distributed on the west coast of (fig. 82). Implied is an early Tertiary forma- southern Mexico and Guatemala, as the most tion of the ancestral biota which was dis- primitive cyprinodontoid. It must be at least rupted by the formation of deserts, wth an as old as the rest of the cyprinodontoids, apparent extinction of centrally distributed found througout most of temperate and trop- forms. Desert formation (as well as intrusion ical Africa, North America, and Eurasia, the by humans) continues to cause the extinction distribution of which suggests a still older of elements of this once more widespread history. biota; of the two species and three subspe-

SUMMARY

The cyprinodontiform fishes, formerly derived characters: (1) symmetrical caudal classified in five families, the oviparous Cy- fin, externally rounded or truncate (or with prinodontidae and four viviparous families, fin extensions) and internally with a single the Poeciliidae, Jenynsiidae, Anablepidae, epural opposing a similarly shaped parhy- and Goodeidae constitute a well-known and pural, with the dorsal and ventral hypural widely distributed group of acanthopterygian plates symmetrical and separate or fused into fishes. This study was carried out with the an hypural fan; (2) a derived form of the up- following six objectives: (1) determine the per jaw, the distinctive feature of which is a monophyly of the cyprinodontiforms as a two-part alveolar arm on the premaxilla; (3) group; (2) determine the monophyly of each a derived form of the interarcual cartilage, of the five families; (3) define the major primitively rodlike and equal in length to the subgroups of cyprinodontiforms, concentrat- epibranchials, from the base of the first epi- ing on the genera of the family Cyprinodon- branchial to a cartilage of the second phar- tidae; (4) determine the interrelationships of yngobranchial; (4) first pleural rib arising on the subgroups; (5) present a comprehensive the parapophysis of the second vertebra (or classification of the cyprinodontiforms which rarely the first) rather than the third; (5) a reflects the interrelationships; and (6) pro- primitively lowset pectoral girdle possessing vide a hypothesis for the present-day distri- a large, scale-shaped first postcleithrum; bution of the group. and, (6) a derived reproductive pattern char- The cyprinodontiforms are a monophyletic acterized by a long developmental period. group based on their sharing the following The family Cyprinodontidae previously 1981 PARENTI: CYPRINODONTIFORM FISHES 545 comprised oviparous fishes in eight subfam- genera and the Neotropical genera are each ilies which were listed in sequence in clas- recognized as monophyletic groups. Annual- sifications giving no hint of interrelation- ism is therefore not considered a uniquely ships. One subfamily, the Rivulinae, is derived character. hypothesized to be the primitive sister group The recognized groups dealt with in this of all other cyprinodontiforms, both ovipa- study are summarized in the comprehensive rous and viviparous, based on the following classification as follows: derived characters which define all cyprino- Order Cyprinodontiformes Berg, 1940 dontoids as a monophyletic group: (1) two Suborder Aplocheiloidei, new usage ossified basibranchials, rather than three, in Family Aplocheilidae Sleeker, 1860 the ventral gill arch skeleton; (2) the loss of Family Rivulidae Myers, 1925 the dorsal hypohyal; (3) a reduced interar- Suborder Cyprinodontoidei, new usage cual cartilage; (4) an autopalatine with an an- Section 1 terior extension offset to the main axis, and Family Profundulidae Hoedeman a posterior flange together creating a ham- and Bronner, 1951 mer-shaped head of the autopalatine; (5) ex- Section 2 tension of the autopalatine ventrally to form Division 1 Family Fundulidae Jordan and Gil- an anterior covering of the quadrate; (6) loss bert, 1882 of the metapterygoid; (7) premaxilla with a Division 2 posterior indentation of the alveolar arm Sept 1 forming an S-shaped distal arm; (8) dentary Family Valenciidae, new family expanded medially to form a robust lower Sept 2 jaw; (9) loss of the first dorsal fin ray result- Superfamily Poecilioidea, new usage ing in the articulation of the first dorsal fin Family Anablepidae Carman, 1895 ray with the first two proximal radials; (10) Subfamily Anablepinae Garman, loss of an ethmomaxillary ligament; (11) loss 1895 of a ligament from the interior arms of the Subfamily Oxyzygonectinae, new maxillaries to the middle of the rostral car- subfamily Family Poeciliidae Garman, 1895 tilage; and, (12) loss of a meniscus from be- Subfamily Poeciliinae Garman, tween the premaxilla and the maxilla. 1895 The aplocheiloids (Rivulinae) are hypoth- Subfamily Fluviphylacinae Rob- esized to be a monophyletic group based on erts, 1970 their sharing the following uniquely or sec- Subfamily Aplocheilichthyinae ondarily derived characters: (1) attached or- Myers, 1928b bital rim; (2) cartilaginous mesethmoid; (3) Superfamily Cyprinodontoidea, new close-set pelvic fin supports; (4) narrow and usage twisted lacrimal; (5) broad anterior end of the Family Goodeidae Jordan, 1923 basihyal; (6) tubular anterior nares; (7) re- Subfamily Empetrichthyinae Jor- dan, Evermann and Clark, duced cephalic sensory pore pattern; (8) as- 1930 pects of the pigmentation pattern; and, ten- Subfamily Goodeinae Jordan, 1923 tatively (9) a size dimorphism with males Family Cyprinodontidae Gill, 1865 larger than females. Subfamily Cubanichthyinae, new Thus, the cyprinodontiforms are divided subfamily into two major subgroups, the aplocheiloids Subfamily Cyprinodontinae Gill, and the cyprinodontoids. In the new classi- 1865 fication, the entire group is raised to the rank Tribe Orestiini Bleeker, 1860 of an order and these subgroups are classi- Tribe Cyprinodontini Gill, 1865 fied in the suborders Aplocheiloidei and Cy- A combination naming and numbering sys- prinodontoidei, respectively. tem of classification is adopted in order to Among the aplocheiloids, the Old World represent most dichotomies while keeping 546 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

the number of empty categories to a mini- ableps; (3) intrafollicular gestation occurs in- mum. The adoption of such numbering sys- dependently in a derived group of poeciliids tems is encouraged in order to represent di- and in Anableps; and (4) spermatozeugmata chotomies while maintaining certain aspects are considered independently derived in poe- of traditional classifications and not prolif- ciliids, Anableps and goodeids. erating a series of rarely used names. Within the classification, the following Of the genera included previously in the genera are placed in synonymy: Fundulinae (table 2) only Fundulus, Lucan- Rivulichthys in Trigonectes. ia, Adinia, Leptolucania, and the previously Simpsonichthys, Campellolebias, Cynopo- classified subgenus Plancterus remain to- ecilus, and Terranotus in Cynolebias. gether as a monophyletic group of genera. In Platypanchax in Hypsopanchax. addition, (1) Profundulus is hypothesized to Hylopanchax in Procatopus. be the most primitive cyprinodontoid; (2) Garmanella in Jordanella. Valencia the sister group of all cyprinodon- Anatolichthys in Kosswigichthys. toids minus Profundulus and the fundulines; Fundulosoma in Nothobranchius. (3) Oxyzygonectes the sister group of Ana- Hubbsichthys is considered a synonym of bleps and Jenynsia; (4) Chriopeoides and Poecilia. Cubanichthys, now synonyms, a sister An analysis of biogeographic patterns ex- group of the more derived cyprinodontines pressed by cyprinodontiform fishes indicates and Orestias, and (5) Empetrichthys and that they are distributed in a manner consis- Crenichthys closest relatives of the good- tent only in part with the break-up of the eids. ancient supercontinent Pangea. The one sis- The four viviparous families are retained ter-group relationship inconsistent with this as monophyletic groups, each undergoing a pattern is that of the Andean Orestias and change in rank. Anableps and Jenynsia as- sessed as sister genera primarily on their the Eurasian Kosswigichthys. This pattern has either been caused by dispersal, extinc- shared derived reproductive characters, to- tion, or by the vicariance of an ancient land gether with Oxyzygonectes comprise the connection. family Anablepidae. Therefore, the family Cyprinodonts of western North America Jenynsiidae is dropped from usage. have a close relationship with those of Mid- The poeciliids are reduced to subfamily dle America. Correlations with patterns ex- rank and included with Fluviphylax and the Pantanodon-proc&topine group in the family hibited by ancient plants indicate that there was once a more widespread biota extending Poeciliidae. from northwestern North America through The goodeids are similarly reduced to a Central America, the flora of which was subfamily and together with Empetrichthys characterized by a dicot forest. Disruption of and Crenichthys comprise the family Good- the forest, and hence of the biota, by desert eidae. The following conclusions concerning vi- formation caused throughout viparity and internal fertilization are pre- much of the range. Fossils across desert give an estimate of the minimum age as the Late sented:. (1) internal fertilization has arisen in- Cretaceous-Early Tertiary. Therefore, ge- dependently among the cyprinodontiforms at neric differentiation of cyprinodonts of the least five times, in Rivulus, Cynolebias, poe- West is hypothesized to have begun at least ciliids, goodeids, and Jenynsia-Anableps; (2) gonopodia occur independently three times, by this period. in Cynolebias, poeciliids and Jenynsia-An- 1981 PARENTI: CYPRINODONTIFORM FISHES 547

LITERATURE CITED Agassiz, L. Verb. Holland Maatsch. Wetensch, ser. 1846. Nomenclatoris Zoologici. Index Univ- 18, pp. 1-136. ersalis. Soloduri, Jent and Gassmann, Bloch, M. E. and J. O. G. Schneider. 393 pp. 1801. Systema ichthyologiae. Berlin, vols. 1, 1853. Recent researches of Professor Agassiz. 2, 584 pp. Amer. Jour. Sci. Arts, vol. 16, pp. 134- Boulenger, G. 136. 1904a. Fishes, systematic account of Teleostei. Ahl, E. In, The Cambridge Natural History, 1924. Zur Systematik der altweltlichen Zahn- vol. 7, pp. 540-727. karpfen der Unterfamily Fundulinae. 1904b. III. Descriptions of new West African Zool. Anz., vol. 40, pp. 49-55. freshwater fishes. Ann. Mag. Nat. 1928. Beitrage zur Systematik der afrikan- Hist., ser. 7, vol. 14, pp. 16-20. ischen Zahnkarpfen. Ibid., vol. 79, pp. 1911. Descriptions of new African cyprino- 113-123. dont fishes. Ibid., ser. 8, vol. 8, pp. 1934. Eine revision der Zahnkarpfengattung 260-268. Cynolebias. Ibid., vol. 108, pp. 304- Breder, C. M., and D. E. Rosen 310. 1966. Modes of reproduction in fishes. Garden Alexander, R. McN. City, New York, Nat. Hist. Press, pp. 1967a. Functional design in fishes. London, i-xv, 1-941. Hutchinson and Co., Ltd., 160 pp. Brown, J. L. 1967b. Mechanisms of the jaws of some ath- 1957. A key to the species and subspecies of eriniform fish. Jour. Zool. London, vol. the cyprinodont genus Fundulus in the 151, pp. 233-255. United States and Canada east of the Alvarez, J., and J. Arreola continental divide. Jour. Washington 1972. Primero goodeido fosil procedente del Acad. Sci., vol. 47, no. 3, pp. 69-77. Pliocene Jaliscense (Pisces, Teleosto- Brundin, L. mi). Bol. Soc. Cien. Nat. Jalisco, vol. 1966. Transantarctic relationships and their 6, pp. 6-15. significance as evidenced in midges. Alvarez, J., and J. Carranza Kungliga Svenska Vetensk-akad. 1951. Description de un genero y especie neu- Handl., vol. 11,472 pp. vos de peces Ciprinodontidos prece- Carvalho, A. L. de dences de Chiapas (Mexico). Ciencia, 1959. Novo genero e nova especie de peixe vol. 9, nos. 1-2, pp. 40-42. anual de Brasilia, com una nota sobre Amemiya, I., and S. Murayama os peixes anuais da Baixaad fluminense, 1931. Some remarks on the existence of de- Brasil. Bol. Mus. Nac. (Brasil), Zool., veloping embryos in the body of an no. 201, pp. 1-10. oviparous cyprinodont, Oryzias Clark, E., and R. P. Kamrin (Aplocheilus) latipes (Temminck and 1951. The role of the pelvic fins in the copu- Schlegel). Proc. Imp. Acad. Japan, vol. latory act of certain poeciliid fishes. 7, no. 4, pp. 176-178. Amer. Mus. Novitates, no. 1509, pp. 1- Axelrod, D. I. 14. 1979. Age and origin of Sonoran Desert vege- Clark, F. N. tation. Occas. Papers California Acad. 1925. The life history of Leuresthes tenuis, an Sci., no. 132, 74 pp. atherine fish with tide controlled spawn- Berg, L. ing habits. California Div. Fish and 1940. Classification of fishes both Recent and Game Fish. Bull., no. 10, pp. 1-51. fossil. Ann Arbor, Michigan, Edwards Clausen, H. S. Bros., Inc., 517 pp. [Reproduced in 1959. Description of two subgenera and six Russian and English in 1947.] new species of Procatopus, Boul., a lit- Bleeker, P. tle known West African genus of cypri- 1860. Ordo Cyprini, Karpers. Act. Soc. Sci. nodont fishes. Vidensk. Medd. Dansk Indo-Neerl. vol. 7, pp. 1-492. Naturh. Foren. Copenhagen, vol. 121, 1863. Familia Cyprinodontoidei. Memoire sur pp. 261-291. les Poissons de la Cote de Guinee, Nat. 1966. Definition of a new cyprinodontid genus 548 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 168

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BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY

Volume 168 1981

PUBLISHED BY ORDER OF THE TRUSTEES NEW YORK: 1981 Edited by FLORENCE BRAUNER CONTENTS OF VOLUME 168

Article 1. Breeding Birds and Old Field Succession on Fallow Long Island Farmland. By Wesley E. Lanyon. Pages 1-60, figures 1-31, tables 1-24. February 26, 1981 Price $3.90 Article 2. Results of the Archbold Expeditions. No. 104. Systematic Revision of the Mar- supial Dasyurid Genus Sminthopsis Thomas. By Michael Archer. Pages 61-224, figures 1-74, tables 1-3. April 10, 1981 Price $10.95 Article 3. Results of the Archbold Expeditions. No. 105. Notes on Systematics of Indo- Malayan Murid Rodents, and Descriptions of New Genera and Species from Ceylon, Sulawesi, and the Philippines. By Guy G. Musser. Pages 225-334, figures 1-51, tables 1-12. June 19, 1981 Price $7.90 Article 4. A Phylogenetic and Biogeographic Analysis of Cyprinodontiform Fishes (Teleo- stei, Atherinomorpha). By Lynne R. Parent!. Pages 335-557, figures 1-99, tables 1-3. September 3, 1981 Price $14.35