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Novitatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2951, 74 pp., 61 figs., 6 tables August 10, 1989 Systematics and Biogeography of the Genus Gambusia (Cyprinodontiformes: Poecilidae) MARY RAUCHENBERGER CONTENTS Abstract .......................... 2 Introduction .......................... 2 Historical Review .......................... 3 Methods ........................... 6 Abbreviations .......................... 8 Materials Examined .......................... 9 Acknowledgments ........ ...... ............................................. 12 Osteology .......................... 13 Systematics .......................... 32 Outgroup Relationships ........................... 32 Diagnosis of the Genus Gambusia .......................... 35 Diagnoses of Subgenera .......................... 36 Relationships Among Subgenera .......................... 40 Relationships Within the Subgenus Heterophallina ................ .............. 42 Relationships Within the Subgenus Arthrophallus ............................... 43 Relationships Within the Subgenus Gambusia ................................... 49 Biogeography ......................................... 54 Distribution of the Subgenus Heterophallina .................................... 54 Distribution of the Subgenus Arthrophallus ..................................... 56 Distribution of the Subgenus Gambusia ....................................... 59 Geographic Areas ....................................... 63 References ....................................... 67 Appendix ....................................... 71 1 Postdoctoral Fellow, Division of Fishes, National Museum of Natural History, Smithsonian Institution. Copyright © American Museum of Natural History 1989 ISSN 0003-0082 / Price $8.40 2 AMERICAN MUSEUM NOVITATES NO. 2951 ABSTRACT The systematic and biogeographic relationships ofGambusia hispaniolae also is included, for com- of the poeciliid genus Gambusia, a monophyletic parison and reference. group of 45 species, are examined in detail. As it Biogeographic analysis of the species groups of focuses primarily on the interrelationships ofsub- Gambusia concentrates on three areas. In southern genera and species groups, this work is not in- Texas and northern Mexico, members of the af- tended as a formal revision ofthe genus; however, finis, nobilis, and senilis species groups overlap; areas where revisionary work are needed are point- their area cladograms are compared. The Rio Pa- ed out. The genus has a sister-group relationship nuco basin of eastern Mexico is a meeting point with the genus Belonesox, and these two in turn for members of the subgenus Heterophallina, the are related to the genus Brachyrhaphis. Gambusia senilis species group, and the afjinis species group. vittata and the members of the rachowi species A repeated pattern indicated a relationship be- group, each previously assigned to monotypic gen- tween the Panuco basin and the Cuatro Cienegas era, do possess the synapomorphies described for basin of northern Mexico, with further ties to the genus Gambusia, and are thus included. Three southern Mexico. Finally, links between Central subgenera within Gambusia are diagnosed: the America and the Greater Antilles are explored subgenus Heterophallina, the subgenus Arthro- through the distribution patterns and area clado- phallus, and the subgenus Gambusia. The subgen- grams of the nicaraguensis, puncticulata, and era Arthrophallus and Gambusia are sister groups. punctata species groups. Repeated patterns in- Within the subgenus Heterophallina, the panuco clude a relationship between Nuclear Central and rachowi species groups are more closely re- America (specifically Belize) and the Antilles, be- lated to each other than either is to G. vittata. In tween Cuba and the Cayman Islands, and between the subgenus Arthrophallus, the nobilis and senilis eastern Cuba, Hispaniola, and the Bahamas. species groups are sister taxa, and in the subgenus Whereas most of the Caribbean patterns do not Gambusia, the puncticulata and punctata species strictly reiterate each other, the patterns are all groups are sister taxa. Relationships within species consistent with each other. It is inferred that they groups also are described. This scheme of rela- represent different fragments of the complex his- tionships, and the new classification derived from tory of Caribbean biogeography, and are com- it, is a product ofcladistic analysis ofmorpholog- pared with some published distributional infor- ical characters. A complete descriptive osteology mation for other taxa. INTRODUCTION Much attention was drawn to the poeciliid levels; however, areas of the genus that are genus Gambusia Poey, with around 45 in need of revision are noted. species, in 1963, when two independent works Poeciliid taxonomy traditionally has fo- were published dealing with the systematics, cused on the specialization ofthe male's anal and to a lesser extent, the biogeography, of fin, the gonopodium, often to the exclusion this group (Rivas, 1963; Rosen and Bailey, of other characters. In fact, detailed descrip- 1963). Since that time, the conflicts between tions of other parts of the anatomy of any the two schemes have never been resolved, poeciliid generally are not available, with and describers have had difficulty classifying some notable exceptions, such as Rama- their new Gambusia species (Miller, 1975). swami's 1945 description ofthe chondrocra- Systematic studies have emerged that focus nium of Gambusia affinis, and Weisel's sim- on higher (Parenti, 1981) and lower (Rivas, ilar account ofthe guppy, Poecilia reticulata, 1969; Fink, 197 la, 197 lb; Greenfield, 1985; in 1967. My osteological account ofthe mem- Echelle and Echelle, 1986) systematic levels; bers of the genus Gambusia uses a male G. this work aims to classify the entire genus, hispaniolae as an exemplar, but discusses the and to establish its close relatives, through range of variation in the genus for all char- the principles and methods of cladistic sys- acters. It will serve both as an introduction tematics. This paper is not intended as a for- to the characters important for systematic mal revision of the genus, as it concentrates analysis of the genus, and as a general ref- primarily on the subgenus and species group erence for the complete osteology of a poe- 1989 RAUCHENBERGER: GAMBUSIA 3 TABLE 1 TABLE 1-(Continued) Proposed Classification of Gambusia GAMBUSIA Poey, 1854 G. wrayi Regan, 1913 Genus puncticulata species group: Gambusia Poey, 1854: 382 (type species Gambusia G. yucatana Regan, 1914 punctata Poey, by subsequent designation by Jordan G. hispaniolae Fink, 197 lb and Copeland, 1876: 142) G. manni Hubbs, 1927 Heterophallus Regan, 1914: 66 (type species Hetero- G. hubbsi Breder, 1934 phallus rachovii, by monotypy) G. bucheri Rivas, 1944 Flexipenis Turner, 1940: 89 (nomen nudem for Gam- G. baracoana Rivas, 1944 busia vittata Hubbs, which was subsequently desig- G. monticola Rivas, 1971 nated as the type species ofFlexipenis, by C. L. Hubbs, G. puncticulata Poey, 1854 in Rivas, 1963: 334) G. oligosticta Regan, 1913 Dicerophallus Alvarez, 1952: 95-97 (type species Dicero- G. caymanensis Regan, 1913 phallus echeagayari, by original designation) G. howelli Rivas, 1944 Subgenus Heterophallina Hubbs, 1926:26 (type species punctata species group: G. regani, by original designation) G. luma Rosen and Bailey, 1963 G. vittata Hubbs, 1926 G. beebei Myers, 1935 panuco species group: G. pseudopunctata Rivas, 1969 G. marshi Minckley and Craddock, in Minkley, G. punctata Poey, 1854 1962 G. rhizophorae Rivas, 1969 G. panuco Hubbs, 1926 G. xanthosoma Greenfield, 1983 G. regani Hubbs, 1926 rachowi species group: G. rachowi (Regan, 1914) ciliid, valuable for higher-level comparative G. echeagayari (Alvarez, 1952) work. Subgenus Arthrophallus Hubbs, 1926 (type species The species of Gambusia live in fresh and Heterandriapatreulis Baird and Girard, a subjective brackish water, and range from the eastern synonym of Heterandria affinis Baird and Girard; by original designation) and southern United States through Mexico affinis species group: and Central America, with a single species G. affinis (Baird and Girard, 1854) along the South American coast. Particularly G. holbrooki Girard, 1859b interesting are the 16 taxa found on islands G. speciosa Girard, 1859c in the West Indies. Speculations about how G. lemaitrei Fowler, 1950 these fishes found their island homes, invok- G. aurata Miller and Minckley, 1970 ing phenomena such as land bridges, migra- nobilis species group: tion, and hurricanes, have abounded in the G. nobilis (Baird and Girard, 1854) past, but subsequent developments in the sci- G. heterochir Hubbs, 1957 have made it G. krumholzi Minckley, 1963 ence of biogeography possible G. georgei Hubbs and Peden, 1969 to critically assess hypotheses concerning dis- G. eurystoma Miller, 1975 tribution patterns. A necessary prerequisite G. sexradiata Hubbs, 1936 for such a biogeographic analysis is a sound senilis species group: systematic hypothesis for the fishes involved. G. senilis Girard, 1 859c Only then can the nature of the relationship G. gaigei Hubbs, 1929 between the organisms and the areas they G. alvarezi Hubbs and Springer, 1957 inhabit be determined. The major objective G. hurtadoi Hubbs and Springer, 1957 of this work, therefore, is to describe rela- G. amistadensis Peden, 1973b tionships within the genus, which are sum- G. geiseri Hubbs and Hubbs,
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