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W. L Minckley a Revision of the Gambusia Nobilis Species Group, with Descriptions of Three New Species, and Notes on Their Variation, Ecology, and Evolution

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W. L Minckley a Revision of the Gambusia Nobilis Species Group, with Descriptions of Three New Species, and Notes on Their Variation, Ecology, and Evolution 13 Reprinted from THE TEXAS JOURNAL OF SCIENCE, Vol. IX, No. 3, September, 1957, ." 1 W. L Minckley A Revision of the Gambusia nobilis Species Group, With Descriptions of Three New Species, and Notes On Their Variation, Ecology, and Evolution by CLARK HUBBS and VICTOR G. SPRINGER The University of Texas In the last review of the Gambusia nobilis species group, Carl L. Hubbs (1929) recognized four species: G. nobilis (Baird and Girard), G. senilis Girard, G. gaigei Hubbs, and G. affinis Baird and Girard. The species with which the name affinis was associated then only, is here- in accorded a new name, G. geiseri. True G. affinis does not belong to this species group. Two other previously unreported forms, G. hurtadoi and G. alvarezi, are also described. The variation, ecology, and evolution in the several species are discussed. Carl L. Hubbs (1926) included three genera ( Heterophallus Regan, Belonesox Kner, and Gambusia Poey) in the tribe Gambusiini. He divided the genus Gambusia into four subgenera ( Heterophallina Hubbs, Gambusia Poey, Arthrophallus Hubbs, and Schizophallus Hubbs). The primary dichotomy within the subgenus Gambusia in his key divided the forms from southern Mexico and the Antilles from those inhabiting northeastern Mexico and southwestern United States. Later (1929) he called the latter fishes the nobilis group. He distinguished these fishes from the other species of Gambusia by: "The distal spines of ray 3 (of the modified male anal fin) are very long, projecting far beyond the hook at tip of ray 4; the largest segment is about equal to the combined basal length of all the spine-bearing segments. Other common characters more or less diagnostic of the nobilis group are the development of a diffuse lateral band extending from eye to caudal base, and of dusky or black markings on the anal fin of the female. The dorsal rays are usually 8, sometimes 7 or 9." We concur with this description with the following modifications: the largest segment of the spines of ray 3 may even exceed the combined basal length of all spine-bearing segments, and the anal rays are usually 7 to 9, sometimes 10. 279 280 THE TEXAS JOURNAL OF SCIENCE KrumHoltz (1948) stated that Hubbs and Walker (unpublished MS) consider that the two nominal species (affinis and holbrookii) comprising the nominal subgenera Arthrophallus and Schizoph.allus respectively, regularly integrade in nature and therefore are to be assigned to the same species. Haskins and Rosen (unpublished MS) also have demonstrated integradation between the two nominal spe- cies. On the basis of gonopodial types, Rosen and Gordon (1951) di- vided the tribe Gambusiini into five "species groups." Despite their statement of the need for a new revision of the Gambusiini, their five gonopodial types represent the three subgenera of Gambusia and the two other gambusfine genera currently recognized. ACKNOWLEDGEMENTS We are indebted to Drs. Reeve M. Bailey and Robert R. Miller of the Museum of Zoology, University of Michigan, as well as Mr. Loren P. Woods of The Chicago Natural History Museum for the loan of specimens. Dr. Ernest A. Lochner provided information on specimens in the U.S. National Museum. Mr. Oscar f. Wiegand assisted in sev- eral collections and served as a translator during a collecting trip in December, 1954, and January, 1955. Dr. Kirk Stravvn, Dr. William F. Pyburn, Mr. Chesley Woods, Mr. William H. Brown, Dr. David Pettus, Mr. Jerome Dorf, and Mr. Elgin M. C. Dietz, assisted in Collecting specimens in Texas. Mr. Pablo Guzman-Rivas collected specimens of fishes from three localities in the Rio Sota la Marina drainage. Dr. Jose Alvarez del Villar and Sr. Jorge Carranza of the Direccion General de Pesca aided in obtaining a permit for collecting in Mexico. Sr. Ing. Leopoldo Hurtado Olin of the Department° de Economia of Chihuahua assisted the June field trip and informed us of the location of El Ojo de la Hacinda Dolores. Dr. Ross Maxwell, Mr. Lemuel A. Garrison, both former Superintendents of the Big Bend National Park, Mr. George W. Miller, presently Superintendent of the park, and Mr. John Palmer, formerly District Ranger at Bo- quillas Ranger Station, gave information on warm springs in the park. Sr. Eduardo Rojas of Valle de Allende gave much information on the post-1900 changes of water connections resulting from irrigation. He also informed us of the location of El Ojo de San Gregorio. Sr. Jesus Pallares of La Escuela Normal de Salicias gave information on the present flow of irrigation water. Sr. Raul Miramontes of El Sauz informed us on the disappearance of the Rio Sauz. Mr. William E. Humphrey of DeGolyer and MacNaughton, Inc., gave information on the geology of the Estaci6n Troya region. Dr. William J. Koster pro- vided locality data for G. nobilis in New Mexico. Drs. Carl L. Hubbs A REVISION OF THE GAMBUSIA NOBILIS SPECIES GROUP 281 and Royal D. Suttkus have read and criticized this manuscript. Dr. Hubbs has also examined specimens in the U.S. National Museum, University of Michigan Museum of Zoology, and the Museum of Com- parative Zoology, and provided color notes on living Gambusia nobilis. The figures and descriptions of the gonopodial suspensorium in these species are abstracted from an unpublished manuscript by Luis Howell Rivero and Carl L. Hubbs. The other drawings were mare by Mrs. Grace H. Groce and Miss Nancy Walker. Mr. George G. Hender- son, Jr., made the photographs. Dr. Kirk Strawn and Mr. Walter E. Fosberg assisted by feeding the living stocks. Miss Gail Kaufman has conducted the mate preference experiments. The project is supported by two grants from the National Science Foundation. METHODS The measurements and counts used in this work are those defined by Hubbs and Lagler (1947). The rays of the gonopodium are num- bered 3 to 5 and the anterior and posterior branches are indicated by the symbols 4a, 4p, and 5a. All dorsal and anal rays are counted, in- cluding the two small anterior anal rays, and the last two elements are counted as one. The special designations for gonopodial struc- tures are labelled on figure 1. Many of the specimens are catalogued at several museums. The fol- lowing abbreviations for these museums are used; CNHM—Chicago Natural History Museum; ENCB—Escuela Nacional de Ciencias Bio- logicas of Mexico; MCZ—Museum of Comparative Zoology (Harvard University) ; SU—Stanford Natural History Museum; TNHC—Texas Natural History Collection; UMMZ—University of Michigan Mu- seum of Zoology; and USNM—United States National Museum. We have virtually omitted proportional measurements from the species descriptions. Our laboratory experiments with five of the species in this group have shown that these characters may vary con- siderably due to environmental influence. For instance, the caudal peduncle depth of wild G. hurtadoi is contained 1.1 to 1.3 times in the caudal length and 0.8 to 0.9 times in laboratory-raised fish (kept under very favorable nutritional conditions). In general, laboratory-raised stocks have deeper bodies and shorter fin rays (see figure 13 and 14) than do wild fish. However, such morphometric differences between the species are undoubtedly in part of genetic origin. Wild or labora- tory-raised G. gaigei are more deep-bodied than their G. hurtadoi counterparts, although laboratory-raised G. hurtadoi are more deep- bodied than wild G. gaigei. We have not noticed any variation of gonopodial structure between laboratory stocks and wild specimens 3 S 6a Sa Sp :: SERRAE ELBOW SPINE TERMINAL HOOKS Fig. 1. Anal fin rays of laboratory raised Gambusia hurtadoi male (TNHC 45861 showing special designations of gonopodial structures. A REVISION OF THE GAMBUSIA NOBILIS SPECIES GROUP 283 of the same form. Likewise, although laboratory fish are paler than wild ones, their basic colors and color patterns are similar. All color markings used (except in G. nobilis, which has not been kept in our laboratory) are evident in laboratory-raised individuals and are thus considered genetic. Variations in fin-ray numbers attributable to the environment are less than the differences between species. In tables 2 through 5 all fish with no trace of gonopodium on 10 to 45 X magnification are tallied as females. The dorsal rays of some females were not counted. Immature males are those whose gono- podium is more or less elongated but not fully developed. Key to Wild Adults of the Gambusia nobilis Species Group la. First and/or second proximal enlarged spines on ray 3 of gonopodium with recurved hook (Fig. 2, A and B) ; terminal hooks of rays 4p and 5a angular at tip; dorsal rays usually 7 (occasionally 8) ; postanal streak prominent (darker than markings on scale pockets); a median row of spots on caudal fin and some at its base; a median row of spots on dorsal; in females distance from front of dorsal to caudal equal to distance from front of dorsal to first predorsal body scale; caudal peduncle depth in females contained 1.5 to 1.8 times in caudal fin length ............................................................... G. geiseri lb. None of the enlarged spines on ray 3 of gonopodium with recurved hooks; terminal hooks of rays 4p and 5a rounded at tip; dorsal rays S or 9 (occasionally 7 or 10) ; postanal streak weaker than markings on scale pockets; no prominent spots on caudal; either no spots or a subbasal row on dorsal; in females distance from front of dorsal to caudal equal to distance from front of dorsal to second or third predorsal body scale; caudal peduncle depth contained 0.8 to 1.4 times in caudal length ......................................................................................2 2a. Ray 4a of gonopodium extends to or beyond tip of 4b (Fig. 2, C and D) ; lateral stripe thin and threadlike; caudal with dark margin; dark margins on scale pockets the most prominent color; markings on side rounded - specks; predorsal streak strong ...................................G.
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