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Isolating Mechanisms in Goodeid Fish Ethological

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Isolating Mechanisms in Goodeid Fish Ethological 84 85 BULLETIN SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VOLUME 75 1976 ISOLATING MECHANISMS IN GOODEID FISH Ktumholz, L. A., J. R. Charles, and W. L. Minckley. Power, 13. M. 1970. Geographic variation of red. of these features among cyprinodontiform fishes, 1962. The fish population of the Ohio River. winged blackbilds in Cential North America. Carl Hubbs, working with Turner, used goodeid Pp. 49-89, 143-152, 165-180, 200-210. in: Univ. Kan .Mus. Natur. Hist., 19:1-83. Aquatic.life Resources of the Ohio River. Ohio trophotaeniae and ovarian anatomy in an exhaus- tive family classification (Hubbs and Turner, River Valley Water Sanit. Comm., Cincinnati, Seymour Ohio, 218 pp. Effects of temperature upon 1939). Although a number of taxonomic papers on of vertebrae and fin rays in young including goodeids (deBuen, 1941, 1942; Turner, salmon. Trans. Amer. Fish. Soc., 88: Lindsey. C. C. 1954. Temperature-contmlled me- 69. 1946; Alvarez and Navarro, 1957; Alvarez, 1959, ristic variation in the paradise fish Marino() 1963; Alvarez and Cortes, 1962; and Romero, opercularis (L.). Canadian J. Zool., 32:87- have appeared since the Hubbs and Turner Snedecor, G. W., and W. G. Cochran. 1967. Statis- 1967) tical methods. Iowa St. Univ. Press, Ames, Iowa, revision, only one (Miller and Fitzsimons, 1971) Lindsey, C. C., and R. W. Harrington, Jr. 1972. 593 pp. significantly altered their classificatory scheme. Extreme vertebral variation induced by !Alper- Greatly influenced by Guillermo Mendoza's study attire in a homozygous clone of the self-fertilizing Vining, A. V. 1952. Experimental study of meristic (1965) with Xenotoca eiseni, Robert Miller and cyprinodontid fish !Grail's mormoratus. Cana- I judged trophotaenial and ovarian anatomy of dian J. Zool., characters in fishes. Biol. Rev., Cambridge 50(6):733-744, Philos. Soc., 27:169-193. goodeids likely too variable within a species to justify broader use in a family classification. MacCrimmon, R., and W. H. Kwain. 1969. Uyeno, T., and R. R. Miller. 1962. Relationships Through synonymies, we reduced the Hubbs- Influence of light on early development and of Empeirichihys erdisi, meristic a Pliocene cyprinodontid Turner classification by four genera and one characters in the rainbow trout, Salina fish from California. with remarks on the but in the same report managed to erect a eahdneri Richardson. Canadian J. species Zool., 47: Fundulinae and Cyprinodontinae. Copeia, 1962: new genus and species of uncertain phylogenetic (,3I-637. 520-532. affinity. Mlrir E. 1963. Animal species and evolution. During a revision of the goodeid genera (Fitzsimons, 1972), I Harvard Univ. Press, Cambridge, Mass., 797 pp. Accepted for publication May I, 1976. Characodon and Xenotoca learned that goodeids exhibit elaborate pair- forming rituals during courtship. Ethological Figure I. Males (above) and females (below) of data confirmed taxonomic interpretations of Xenotoca variata (A) 41.5 and 45 mm SL, X. eiseni X. melanosonta (C) 66.5 and 76. i v morphological information and, in many instances, (B) 33 and 38.5, and were better indices to relationships because atten- ETHOLOGICAL ISOLATING MECHANISMS IN GOODEID FISHES OF THE GENUS tion was drawn to features that the animals them- several localities along the Rios XENOTOCA (CYPRINODONTIFORMES, OSTEICHTHYES) selves used in distinguishing species. Although known from the study revealed certain elements of courtship to Tamazula and Tuxpan in southern Jalisco but be generically and specifically invariable, intra- has never been collected in the central Jalisco JOIN MICIIAEL FITZSIMONS' specific differences in two species of Xenotoca basins of the Rio Armeria or Rio Ameca which were unmistakable. This report examines these lie between its northwestern and southeastern Ansrancr: Although no natural hybrids are known, fertile laboratory hybrids were readily differences. limits. Xenotoca melanosoma is known only from obtained from Xenowa eiseni and X. inelanosoina in forced crosses in which a conspecific mate The genus Xenotoca includes X. variata (Bean), Jalisco. Near its southern limit, the type locality was not available. In choice crosses with males and females of both species, sympatric stocks X. eiseni (Rutter), and X. melanosoma Fitz- is in the Rio Tamazula about 16 km south by never hybridized, but allopatric fishes frequently mismated. Ethological data reveal differences simons (Fig. 1). Xenotoca variata, the type species highway from the town of the same name. It in courtship behavior and discriminatory ability in sympatric fishes not observed for allopatric , of the genus and perhaps the most widely distrib- ranges north into streams and ponds about 32 km conspecific stocks nor in fish from two populations of the congener X. variala. These uted goodeid, occurs in a variety of lentic and south of Guadalajara, east to Lago de Chapala, differences, which prevent interbreeding, are offered as evidence for the perfection of premating and west into basins of the Rio Ameca and Rio isolating mechanisms lotic habitats associated with the drainage basins in sympatry. Hybrid inferiority, essential for divergence in sympatry, of the Rios Verde and Aguascalientes in the state Grande de Santiago. Xenotoca eiseni and X. was reflected in reduced survival and inability to compete for males. of Aguascalientes, the Rio Santa Maria of San melanosonta have been collected together at sev- Luis Potosi, the Rio Grande de Santiago of Jalisco eral localities in the Rios Grande de Santiago and the Rio Lerma of Guanajuato, Tamazula. No natural hybrids are known, but in and Michoacan, . la Laja of Queretaro. Members of laboratory crosses fertile F ' hybrids have been The Gooch:id:lc is a family of about and the Rio de 35 species of hausted early in embryogeny and the young receive this species have not been taken sympatrically with produced from all combinations of allopatric and cyprinodontifortn freshwater fishes restricted to nutrition from the female through anal rosette either X. eiseni or X. melanosoma. Forced hybrid sympatric stocks when conspecific mates were the Mesa Central of Mexico. Goodeids differ or ribbon-like structures analogous to the placentae crosses (only heterospecific mates available) and riot available (Fitzsimons, 1972, 1974). In from their livehearing relatives, the anablepids, of mammals. These structures, called tropho- X. variata with its con- choice crosses consisting of males and females of jcnynsiids, and pocciliids, in having true viviparity artificial inseminations of tacniae, were studied in detail by thc embryologist geners have been unsuccessful. The distribution of both species, members of allopatric stocks some- rather than ovoviviparity in which embryos de- Turner (1933, 1937). Realizing the uniqueness X. eiseni is disjunct. In Nayarit, it occurs in the times hybridized but sympatric fishes never mis- velop at the expense of yolk in retained eggs and Rio Compostela and above 600 m in the drainage mated. receive little or no nourishment from the mother. basin of the Rio Grande de Santiago in which it The maintenance of X. eiseni and X. melano- In all Mus. Zoology and Dept. Zoology and Physiology, but one goodeid species, the yolk is ex- Louisiana State Univ., Baton Rouge, Louisiana 70893. ranges into northwestern Jalisco. The species is soma as discrete species depends on the presence 4 87 86 BULLETIN SOUTHERN CALIF° RNIA ACADEMY OF SCIENCES VOLUME 75 1976 ISOLATING MECHANISMS IN GOODEID FISH EL of one or more premating isolating mechanisms aunt species-specificity, the displays of courting RIO SANTA PRESA which prevent their interbreeding. For sympatric males showed a greater promise for use in taxon- GIGANTE populations of these species there are two possibil- omy (Fitzsimons, 1972; Fitzsimons and Le- MARIA ities ( Mayr, 1965): either potential mates do not (irande, 1974) and in the study of ethological meet (seasonal or habitat isolation) or potential isolating mechanisms. The displays are assumed mates meet but do not mate (ethological isola- to be inherited and relatively unmodified by ex- tion). There is no evidence for seasonal or habitat perience rather than learned characteristics because HEAD- isolation. Females of the two species bear young they were identical in animals with conspecific FLICKING 1 '5' throughout the year when kept in aquaria, and experience only, heterospecific experience only, '9 29 2 36 pregnant females and neonates of both species are and in those with no experience at all, i.e., ones 29 9 29 found in collections made during the same times raised in complete isolation. For at least two of the year from sympatric and allopatric popula- goodeid species underwater observations in Me- LATERAL T - tions. Their ecological requirements are similar xico indicated that reproductive behavior in F ORMATION (Fitzsimons, 1972), and there is no apparent dis- aquaria is the same as that which occurs naturally continuity in their distribution at a given locality— (Fitzsimons, 1972). both species can be caught in the same seine haul. 23 17 25 14 Thus, differences in mating behavior are likely alone responsible for preserving the integrity of METHODS these two potentially hybridizable species where L ATERAL they are sympairic. This paper examines the Materials and methods included certain of those TILTING nature of ethological isolating mechanisms opera- used in a recent revision of Xenotoca (Fitzsimons, tive in Xenotoca variata, X. eiseni, and X. melano- 1972). Live stocks for behavioral studies and 9 7 9 ('? 3 soma and, for the latter two species, suggests how hybridization experiments included X. variata: these mechanisms have evolved toward greater Rio Santa Maria, ca. 1.6 km S Villa de Reyes, San efficiency in sympatry. Luis Potosi, R. R. Miller and H. L. Huddle, III: 26:1968; Presa El Gigante, near Santa Maria de Gallardo, 21 km NE (Hwy 45) on road to Loreto ROUND GOODEI D COURTSHIP (at La Dichosa), Aguascalientes, R. R. Miller, H. DANCE L. Huddle, and J. Gomez, IV: 1:1968; X. eiseni: Internal fertilization and viviparity require that Manantial El Sacristan, 1.3 km NW plaza Topic, there be sexual selection and pair-forming for suc- Nayarit, C.
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