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<I>POECILIA FORMOSA</I> Evolution, 34(5), 1980, pp. 917-922 INTERSPECIFIC HYBRIDIZATION AND THE EVOLUTIONARY ORIGIN OF A GYNOGENETIC FISH, POECILIA FORMOSA 1 BRUCE J. TURNER ,3 , BETTy-LOU H. BRETT"", AND ROBERT R. MILLER! 'Museum ofZoology and "School of Natural Resources, University of Michigan, Ann Arbor, Michigan 48109 Received November IS, 1979. Revised February 11, 1980 In recent years, largely through the ap­ Atlantic slope of Mexico and Guatemala. plication of allozyme techniques, several The stocks of shortfin mollies used by the unisexual (thelytokous) vertebrates have Hubbses (and by Meyer, who obtained his been shown to be genetically equivalent material from them) were descended to F 1 hybrids of certain related bisexual largely from progenitors collected on the species. The causal relationship, however, Pacific coast of Mexico near Acapulco (L. between interspecific hybridization and C. Hubbs, pers. comm.). They were the origin of unisexuality has not been therefore most likely P. butleri or P. sphe­ elucidated, and the role of hybridization nops (both sensu Schultz and Miller, per se is controversial (Cuellar, 1974, 1971), or perhaps inadvertent laboratory 1977, 1978; Cole, 1978; Wright, 1978). hybrids of one of these species with the The amazon molly, Poeciliaformosa (Gi­ other or with P. mexicana. In other rard), an ovoviviparous gynogentic teleost words, the attempts to produce gynoge­ (family Poeciliidae) was the first known netic P. formosa in the laboratory by in­ unisexual vertebrate (Hubbs and Hubbs, terspecific hybridization were probably 1932). The discoverers of its unisexuality done with what later turned out to be at recognized early that it was probably of least one "wrong" species. hybrid origin, as it was an almost exact To our knowledge, there have been no morphological intermediate between the subsequent attempts to produce gynogens sailfin molly (Poecilia latipinna) and the by interspecific hybridization of Poecilia shortfin mollies (which were then regarded species. Prompted by this, and by the suc­ as a single species, P. sphenops). Labo­ cessful laboratory "synthesis" of hybrido­ ratory hybridization of the presumptive genetic unisexual Poeciliopsis by direct parental species, however, produced only hybridization of bisexual progenitor species bisexual progeny, including fertile males (Schultz, 1973, 1977), we hypothesized that (Hubbs, 1933, 1955, 1961; Meyer, 1938; hybridization of the "correct" Poecilia Hubbs and Hubbs, 19400, 1946b). Sub­ species should produce at least some gyn­ sequent to those early hybridization ex­ ogenetic progeny. Moreover, if gynogens periments, P. "sphenops ," as it was then could be easily synthesized, the interfer­ recognized, has been shown to be an as­ tility of other shortfin mollies in laboratory semblage of morphologically similar but crosses would provide a ready-made sys­ genetically quite distinct species (Hubbs, tem (complete with allozyme markers) in 1961; Schultz and Miller, 1971; Miller, which the genetic bases of hybrid unisex­ 1975; Brett et al., unpubl.). A variety of uality could be studied in detail. The re­ zoogeographic, morphological (Darnell sults presented here, however, indicate and Abramoff, 1968), chromosomal (Prehn that our hypothesis was incorrect or at and Rasch, 1969), and biochemical genetic least incomplete; laboratory hybrid prog­ (Abramoff et aI., 1968; Turner et aI., eny of P. latipinna and P. mexicana are 1980) data suggest that the shortfin species not gynogenetic. involved in the ancestry of P.formosa was P. mexicana, a species restricted to the MATERIALS AND METHODS Our experiments took advantage of the relative ease with which mollies can be 3 Present address: Department of Biology, Virgin­ ia Polytechnic Institute and State University, cultivated and interbred in laboratory Blacksburg, Virginia 24061. aquaria. Hybridization was accomplished 917 918 B. J. TURNER ET AL. by combining males and females in the 1. Some hybrid females were bred to same aquarium and removing broods of "tester" male commercial "black mollies." young as they appeared. Virgin (sperm­ ("Black mollies" are melanistic Poecilia free) females, from established laboratory hybrids that are bred in large numbers by stocks, P. mexicana Veracruz and P. the ornamental fish industry. They have sphenops Veracruz, were used. The field­ had a complex history [e.g., Hubbs, 1936, caught females of other stocks we used p. 247, pI. 8], as separate all-black strains were assumed to have stored homo specific were derived by selective breeding from sperm before capture. In these cases, the partially black P. latipinna, P. velifera first two broods that each female delivered and possibly from P. mexicana and/or P. after being placed with a heterospecific sphenops. These were subsequently hy­ male were considered to be contaminated bridized in various combinations. The by homospecific progeny and discarded; males we used resembled those of P. la­ the third and subsequent broods were pre­ tipinna, and were all from the same sumed to consist of hybrids. All hybrid source. Shortfin-like strains [sometimes broods were reared in individual aquaria. called "Yucatan mollies"] are also avail­ Males were removed as they became evi­ able commercially.) When mated to such dent by their developing fin colors and/or a tester, nongynogenetic ("bisexual") hy­ gonopodia, and reared separately. brid females should produce offspring that Sources of parental specimens used in express some of the paternal genes for hybridization experiments were as fol­ black pigment (see Schroder, 1964, for the lows: P. latipinna Naples-Stock B77-2, genetics of black body color in Poecilia). canal along Florida Hwy 451; 19 km SE Gynogenetic females, however, should Naples, Florida; collected January, 1977. produce progeny that never express the P. latipinna Nueces-Stock T77-1, Nueces "paternal" genes. This method of detect­ R., boat launching inlet at ranger head­ ing potential gynogenesis requires that all quarters, Hazel Bazemore State Park, test broods be reared until black color Robstown, Texas; collected February genes can be expressed (in some cases, up 1977. P. formosa have also been taken at to 16 wk). this locality (W. S. Moore, pers. comm.) 2. Some hybrid females were bred to but not by us. P. mexicana Monterrey­ tester males of the more distantly related Stock M77-39, Ojo de Agua de Apodaca, insular species, Poecilia (Limia) vittata. 32 km W of Hwy 54, near Monterrey, In the laboratory, males of P. vittata are Nuevo Leon, Mexico; collected June, efficient "fathers" of pseudogamous P. for­ 1977. This stock is presumably referable mosa broods (Hubbs and Hubbs, 1946a, to the subspecies P. m. limantouri (Menzel 1946b;]. S. Balsano, pers. comm.). How­ and Darnell, 1973). P. mexicana Vera­ ever, the hybrid cross of P. vittata males cruz-Stock M66-29; trib. to a lagoon on with females of several bisexual molly the Rancho San Gabriel, 38.3 km N of species is apparently associated with sig­ San Jose Cardel, Veracruz, Mexico; line­ nificant zygote mortality (Schroder, 1965). bred laboratory stock, progenitors collect­ We therefore reasoned that in an aquari­ ed 1966. This stock is presumably refer­ um containing F 1 hybrid females and a able to the subspecies P. m. mexicana male P. vittata of known fertility, any fe­ (Menzel and Darnell, 1973). P. sphenops males which became gravid and carried Veracruz-Stock M67-2, pond 1 km S of broods to term were likely to be gynoge­ La Piedra bridge, approx. 35 km S of Cd. netic; the broods of nongynogenetic fe­ Veracruz, Veracruz, Mexico; line-bred lab­ males would fail before parturition, and oratory stock, progenitors collected 1967. most likely be resorbed. This method is Tests for gynogenesis.-We used three potentially very efficient, for, unless gy­ separate testcross procedures in attempts nogens are very common among the hy­ to detect gynogenetic reproduction among brid progeny, relatively few test broods F 1 hybrid females. need be reared. HYBRIDIZATI ON IN POECILI A 919 latipinna x P. sphenops hybrids were pro­ duced (Table 1). The following consider­ ations are note worthy. 1. There is little evidence, if any, of sig­ nificant zygote mortality associated with a any of the hybrid combinations. Brood size was lowest with the two crosses in­ volving P. latipinna Nueces females, but, lacking data on the perform ance of equ iv­ alent females when mated to conspecifics, we do not know if these small broods are characteristic of the particular femal e, the population, the cross, or other factors. b 2. All of the crosses produced repro­ ductively competent males as well as fe­ male s. Two male hybrid P. mexicana Ve­ racruz female x P. latipinna Nueces males, chosen at random, were individ­ ually backcrossed to female P. mexicana ; they produced ten broods totaling 98 prog­ eny. Similarly, three males of the recip­ c rocal cross, when backcrossed to female P. latipinna , produced 19 broods with 122 progeny. T wo P. sphenops x P. latipinna hybrid males produced more than 30 youn g when crossed to P. latipinna fe­ male s. Males wer e less frequent in the d progeny of all the P. latipinna x P. mex­ icana crosses than would be expected on the basis on 1:1 sex ratios (Table 1 "G;" FIG. 1. a. F, hybrid, sexually mature male, 32 values); interbrood heterogeneities (Table mm Sta ndard Length (S.L.); P. latipinna Nueces 1, "G;" values) were not significant. The male x P. mexicana Veracruz female . b. Hybrid as combined apparent sex ratio for all of the in a female, 37 mm S.L. Progeny from the other interspecific crosses are very similar to these in gen­ P . latipinna x P. mex icana hybrid prog­ eral appear an ce. c. Backcross progeny, female 36 eny was 106 males to 231 females or mm S.L.; F, hybrid reciprocal of that in a X male rou ghly 1:2.2. F rom the pooled brood P. mexicana Veracruz. Extern al morphology very data, the sex ratio of the progeny of the similar to that of P.
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