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Oedipina (Caudata: Plethodontidae)

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Oedipina (Caudata: Plethodontidae) Rev. Biol. Trop., 45(3): fl 85-1208, 1997 Phylogenetic and taxonomic implications of protein variation in the Mesoamerican salamander genus Oedipina (Caudata: Plethodontidae) David A. Goodl andDavid B. Wake2 1 Museum oC Natural Science, 119 FosterHall, Louisiana State University, Baton Rouge, LA 70803, U.S.A. Museum oC Vertebrate Z00logy and Department oC lntegrative Biology, University of California, Berkeley, CA 94720- 3160, U.S.A. (Recibido: 10-lV-96 Revisado: 14'-I1I-97 Aceptado: 14'-I1I-97) Abstraet: An analysis of protein variation was conducted among 28 population samples of salamanders of .the genus Oedipina. collected in Guatemala, Honduras, Costa Rica, and Panama, in order to assess the validity oC currently recognized specles and to evaluate their phylogenetic relationships. A set oC eight species of the related genus Nototriton served as an outgroup. There are four major clades, with Nei geneuc distances between elades ranging frorn 0.46 to 2.67. Patteins of geographic variation within O.u.niformis as currently recognized indicate thatpopulations withinit should be placed ioto fourspecies: O. alfatoiis a valid species, and O. gracilis and O. pacificensis are �surrected from the synonymy of O. uniformis, while O, bonitaensis, O, inusitata, O.longissima, and O. syndactyIqremain in synonymy. Geographic variation also is apparent io O.cyclocauda, O. poelzi, and O. pseudouniformis, but.no taxonomic changes arewarranted at thistime. The speciesstatus of O. altura andO. grandis issupported. The lour samplesrepresenting the parvipes group are the rnostdistantly related to the other samples, andgenetic variation within thegroup is sufficientto warrant resurrectingthe Costa Rican O.alleni from the synonyrny of O.parvipes of Panama and Colombia All motphologically differentiated samples are also genetically differentiated. Our results demonstrate that 12 specieswere sampled (O.alfaroi, O. allelli, O.a.ltura, O. cyclocauda, O.gr acilis, O.grandis , O.ignea, O.pac ificensis, O.parvipes, O.poelzi, O. pseudouniformis, and O. uniformis).no samples were available for O.éarablanca, O� collarís, O.complex, O. elongata, O. gephyra, O. paucidentata. O.stenapodia, O.stuarti, or O.taylori, but all are recognizable as distinct species on motphological grounds. , Key words: Caudafa, Oedipina, Allozymes, Proteins, Variation, Phylogeny, Taxonomy The great radiation of plethodontid and intrapopulational variation in this character. salamanders in the New World tropics has Amongthe tropical species in the supergenus, produced a large number of species that· are onlythe elongated, attenuate salamanpers in the diverse in botb morphology and ecology. These genus Oedipina have increased vertebral species, aH members of the monophyletic numbers, and hence Oedipina constitutes one of supergenus Bolitoglossa, comprise about 45% the best supported monophyletic groups of of the living species of caudate amphibians tropical salamanders. Although a few species of (Wake.19 87). Members of this supergenus are Bolitoglossa occur farther south, Oedipina has unique in that the vast majority have a fixed the most southerly overaH distribution of any number (14) of trunk vertebrae, regardless of salamandergenus. The species are ecological whether the body is elongated or noto Most specialists for the IDOSt part, having fossorial other élades ofsalamanders display both inter- habits even in the lowlands where other 1186 REVISTA DE BIOLOGIA TROPICAL salamanders typically are arboreal (Wake and views are generally concordant with those of Lynch 1976). Frost and Hillis (1990), and we accept their Many of the species of Oedipina are main recommendations. Briefly, we consider confusingly similarin structure and coloration. species to be genetically cohesive units that are Most of .the 18 species currently recognized are evolutionarily independent of other such units. dark gray to brown, a1though sorne species In the case of allopatric populations, we are have lightly pigmented dorsal patterns, concerned with historical processes that have especially on the head. They all have very long led to the allopatry, and with estimates of the tails (sometimes 3 times the snout-vent likelihood of future genetic interchange. These length), short legs, and reduced hands and feet. estimates are based on ecology and geography, This high degree of similarity has led to much as well as on measured differences in confusion with regardto the identification and morphological and bíochemical traits. We systematics of species in the genus. The most attemptto estimate relative length of separation difficultcomplex is the taxon currently known on the basis of these characters (primarily as O. uniformis,which ineludes seven described morphology and biochemistry). We assume entities (Brame 1968. O. bonitaensis, O. that populations that have been in isolation gracilis, O. inusitata, O. longissima. O. sufficiently long to permit substantial pacificensis, O. syndactyla, and O. uniformis). morphological and biochemical differentiation It ranges from sea level to over 2000 m are more likely to be reproductively elevation, resulting in its occurrence in a incompatible than are more recently isolated broader range of hábitats than any other populations in which little differentiation has salamander. occurred. This approach is necessarily Two workers, TayJor (1948, 1949, 1952, subjective, but it avoids both the acceptance of 1954) and Brame(1960, 1963, 1968, Brame extremely polytypic species on the one hand andDuellman 1970), have been responsible for and the acceptance of species status for all most of the species descriptions and current undifferentiated (or poorly differentiated) . taxonomy in Oedipina. In the most recent and geographic isolates on the other. in cases in comprehensive reVISlon, Brame (1968) which tests of sympatry are impossible, our recognized 15 species in two species groups. species conceptinvolves an inference regarding He described five new speciesand reduced nine the probability of reproductive isolation. names to synonymy. Brame and Duellmart Sympatry tests probably are rare among (1970), Brodie and Campbell (1993), and relatively young species, but when they are McCranie et al. (1993) each added an additional available the degree of differentiation .among species. closely related sympatric species can guide In the course of field work in Central decisions concerning allopatric populations. In Americaover thelast 15 years(see Wake 1987, general, we believe thatit is better to err in the Wake el al. 1992), an attempthas been made to direction of too many rather than too few obtain samples of these elusive animals for species, and when evidence for differentiation is morphological and biochemical studies. Recent present andwe have sufficientmaterial to make work has concentratedon Costa Rica, where the a reasonable judgement, we choose to recognize majority of species occurs. This report taxa as full species. summarizes a re-examination of the species status of several populations of Oedipina using biochemical information unavailable to MATERIALS AND METHODS previousworkers. In recent papers we have discussed our A total of 26 populations (Table 1) species concept and our operational taxonomic representing 11 specíes (see discussion below) procedures (Good and Wake 1992, 1993). Our were analyzed for variation at all of the 22 Good &Wake: Mesoamerican salamander genus Oedipina 1187 protein loci listed in Table 2. The distribution unavoidable in this analysis because many of of alle1es al these loei is listed in Table 3, as is the species are very rare (e.g., our specimen of the outgroup state, where appropriate, based on O. altura was only the second specimen found the alleles present in eight species of since the species was firstcollected in 1964). Nototriton, a close relative of Oedipina (Wake AH genetic distances reported in this paper and Elias 1983). In addition, one individual of are calculated according to Nei (1978). O. cyclocauda from Honduras al1d five O. FuU details of synonym for the taxa grandis from Costa Rica were analyzed (Tabie discussed in this paper are provided by Brame 1), but only a subset of the loci were scorable. (1968). data for these specimens are presented in the text onIy. Liver and intestine samples were collected RESULTS from freshly killed specimens and stored at - 76°C until used. Tissues were combined and We found 129 alleles distributed among fue homogenized in approximately equal parts 22 proteins examined in the 26 samples of tissueand deionized water, and fuen subjected to Oedipina (Table 3). No proteins are horizontal starch-gel electrophoresis using monomorphic among all samples, although standardtechniques (Selander et al. 1971, Harris Icdhl and Icdh2 vary only in fue presence of andHopkin son 1976, Richardson et al. 1986, polymorphisms involving unique alleles in Hillis and Moritz 1990). The allele patterns single populations. Alleles per protein range resultingfrom this analysis were subjected to a fromtwo (Icdh2 and Pk) to ten (Ldh and Pepl). variety of phenetic and phylogenetic analyses The maximum number of alleles per protein in both by hand and using the BIOSYS· (Swofford a single sample is three (Pepl in sample 20. and Selander 1981) and PAUP (Swofford 1985) see Table 1). computer packages. Voucher speeimens were There areeight distinct morphotypes (Figs. deposited in the Museum of Vertebrate 1 and 2) among the populations of Oedipina Zoology, University of California (MVZ), the surveyedfor aH loei in this analysis, and
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