Catalogue of American Amphibians and Reptiles. Ambystoma

AMPHIBIA: CAUDATA: AMBYSTOMATIDAE AMBYSTOMA DUMERlLll Catalogue of American Amphibians and Reptiles.

Brandon, RA. 1992. Ambystoma dumerilii. Ambystoma dumeriffi (Dug&) Achoque, Achoque de Agua, Lake Patzcuaro Salamander

Sired~nDum~liiDugPs, 1870:241. Type-locality, "... la Laguna de Pitzcuaro..." Michoacin, Mhico. Dug& designated no types, but four specimens - ANSP 13862, USNM 16201-2, and a specimen in the Museo Alfredo Dug&, Universidad de Guanajuato, M6xico-are considered to be syntypes (Dunn, 1939; Taylor, 1940; Maldonado-Koerdell, 194&, b; Smith and Necker, 1944). Descriptions and illustrations in Dug& (1870,1872) indicate at least one syntype of each sex. Synrypes not examined by au- thor, but photographs of USNM syntypes examined; existence ofsyntype ANSP 13862 confumed by John E. Cadle. See Com- ment. Siredon dumeriIi: Smith, 187756. Amblystoma dumerilfi Cope, 1887:7. Siredon]. dum&li: Cope, 1889:7. Ambystoma dumwili: Bailey, 1904:69. Amblystoma dumerilii: Galindo y Villa, 1927:209. Ambystoma dumerilii: Lafrentz, 1927:162. Map. Thetype-locality, Lake Pitzcuaro, Michoacin, indicated by the Ambystoma dumereli: Scharlinski, 1939:54. large open circle, is the only confirmed population of Ambystoma ~alb~siredondumerilii: Dunn, 19391. Placed in monotypic new dumerilii. San Juan del Rio, Queretaro, be only other purponed genus. localityand type-locality of Maldonado-Koerdell's A. d.queretarensis Batbysiredon dumerili: Manin del Campo y Sinchez, 1940:486. (see comment), is indicated by [he smaller solid circle. Amblystoma dumereli: Taylor, 1947:576. Amblystoma dumerli: Taylor, 1947576. Batbysiredon dumerilii dumm'lii: Maldonado-Koerdell, 1948a:195. os"[both are well-developed larvae, whose cloacas do not Batbysiredon dumerilii querelarensis Maldonado-Koerdell, 1948a: have swollen lips]. Syntypes not examined, but see Comment. 199. Type-locality, "San Juan del Rio (Qro.)." Two apparently Ambystoma dumerili queretaremb Tihen, 1958:44. unnumbered syntypes collected by Alfredo Dug& are in the Ambystoma dumen'li dumwili: Duellman, 1961:15. Museo Alfredo Dug&, UniversidaddeGuanajuato, Mexico. Ac- Ambystoma dumerilii dumerilii: Brame, 1967:lO. cording to Maldonado-Koerdell, 1948a:200, "Arnbos son larvas Ambystoma dumerilii queretarensis: Brame, 1967:lO. bien desarrolladas, cupscloacan no tienen 10s labios hinchad- Ambystoma dumerillii: Collette and Kig, 1973:391.

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, I . .' I I. I.. 1 - :x-'~--ckE1 *& Figure 1. Adult female Ambystoma dumerilii, 140 mm SVL (from Brandon, Copeia 1970385, fig. 1; used with permission). Ambystoma dumen'llii dumen'llii: Gorham, 19742% sis and changes in body morphology. Shaffer and Lauder (1985a, b) Ambystoma dumen'llii quemtamsis: Gorham, 197428. and Lauder and Shaffer (1985) examined the functional morphology Ambystomaquemlamis: Smithandsmith, 1976:C-A-2. Considered of feeding. Shaffer (1986) referred to A. dumPrilii in a case study of a different species from A. dumerilii. character weighting in phylogenetic analysis. Brandon (1989) and Ambystoma queretame: Frost, 1985:554. Shaffer (1989) mentioned A. dum&lii within the context of other Mexican species of Ambystoma, particularly A. mericanum. Content. Two subspecies have been recognized by some authors, A. d. dumerilii and A. d. qumlamk, but the laner Remarks. Dunn (1939) considered this speciesso distinctive probably was described from specimens with incorrect locality data. that he established a new genus, Batbysitdon, for it. Workers from See Comment. Tihen (1958) on, however, have preferred to treat it as a species of Ambysloma Breeding experiments (Brandon, 1972, 1977) and DefWtion andDiagnosis. This large (reach maturity when genetic distance data (Shaffer, 1984a) showed that it might best be. 115-120 mm SVL and reach 170 rnm SVL), robust, broad-headed, considered a member of the Ambystoma tigrinum species group. paedomorphic, and perennibranchiate (does not transform in na- Prematurely considered to have been brought to extinction by ture) species of Ambystoma is distinguishable from all other Am- introduced game and food fshes (Smith and Taylor, 1948), A. bystoma by: a broad and flanened head; uniformly tan to brown dumerilii was present in the 1950-60's (Duellman, 1961) and was coloration; hyperfdamentous gills (the distinctive dorsal gill fimbriae abundant in the early 1970's. Lazcano-Barrero et al. (1988) consid- are acquired by larvae at 30-35 mm SVL); short, extensively webbed ered the species in danger of extinction from habitat destructionand digits; three phalanges in the fourth hind toe; a conspicuous keel on local exploitation by fishermen, and Flores Vilela and Gerez (1988) the posterior margin of the hind feet; and moderately few (1626, X- also listed it as endangered. Siltation of Lake Pitzcuaro has been 21) gill rakers on the anterior face of the third gill-bearing arch. suggested as a major threat by Shaffer (1989). Ambystoma dumwilii has been listed in CITES Appendix I1 since 1975 (Code of Federal Descriptions. Duges (1870, 1872) provided descriptions Regulations, 1991). and sketches of external morphology and internal anatomy. Taylor The Achoque is a familiar animal to local residents and fisher- (1940) provided tooth counts, various head and body dimensions, men who exploit it for food and folk medicine. Recipes by which and descriptions of tail and foot morphology for three USNM speci- they are prepared for consumption were given by Liner (1978) and mens, including two syntypes. Tihen (1958) reviewed the distinctive Calder6n and Rodriguez (no date). featuresofthespecies. Brandon (1970a, b) described size range, size at maturity, spermatophores, and eggs. Brandon (1972, 1977) de- Etymology. The species was named in honor of Auguste scribed larval development and laboratoryhybrids with A. m'canum DumPril, professorofherpetolo~yandichthyology, Mu&umd'Histoire and a variety of other species of Ambytoma Brandon et al. (1974) ~aturelle,Paris. The common name "~choque"is the Tarascan described sperm morphology of this and three other species, and of language equivalent ofthe Nahuatl 'Axolotl," andthe Spanish "Ajolote." hybrids among them. Brandon (1976) described induced and spon- Like the common name Axolotl, the name Achoque often is applied taneously metamorphosed animals. Shaffer (1984b) provided mor- locally to any large branchiate salamander. Sometimes the species is phometric data and growth trajectories. Zalisko et al. (1984) com- called "Achoque de agua" to distinguish it from terrestrial forms. pared diameters of spermatozoan cytoplasmic droplets in spermatophores with those of a variety of other salamanders. Krebs and Comment. The status of A. d. queretarensis (Maldonado- Brandon (1984) compared foot morphology, number of gill rakers, Koerdell, 1948a) has been questioned and is puzzling on two points metamorphic response, and egg size and color in A. d~m~liiwith(summarized in Frost, 1985). Confusion has surrounded both the several other Mexican Ambystoma. Gomez Rios (1989) described the identity and the collection locality of two San Juan del Rio, Quere- male reproductive tract and histological details of the spermatogenic taro, specimens labeled Siredon dumerilii in the Museo Alfredo cycle. Dug&, Universidadde Guanajuato, that were designatedsyntypes of Bat~~tdondumerilii~tamisbyMaldonado-Koerdell(1948a). Illustrations. DugPs' (1870,1872) descriptions were accom- Smith and Necker (1944) examined the specimens and identified panied by sketches. Maldonado-Koerdell(1948a, b) photographed them as larvae of Ambystomq rather than Batbysiredon dumerilii. the syntype in the Dug& Museum. An excellent photograph of an According to Maldonad~Koerdell(194&), Smith and Necker left a entire animal, showing distinctive features of the feet and gills, note in the jar indicating the specimens belonged to "... una posible appeared inFauna(1971:37). Brandonillustrated a liveadult (1970a), especie nueva del g6nero Ambystoma, ..."(a possibly new species of aiarva (1976), a hatchling (1972), a spermatophore and eggs (1970b); the genus Ambystoma), thus confirmingthey both had examined the individuals underaoin~metamorphosis (1976). and F, hybrids with same specimens. Tihen (1958) treated A. (Batbysiredon) d. A. mexicanurn, ~Tti~tkum,and'A. talpideum (1972; 1977). Krebs gumtarensisas a valid subspecies but said Batbysiredonwas known and Brandon (1984) included comparative sketches of foot morphol- only from Lake Ptitzcuaro. Smith and Smith (1976) considered the ogy. Lauder and Shaffer (1985) illustrated the skull and musculature occurrence of A. dumeriliinear San Juan del Rio highly unlikely and of the head and pectoral region. Duellman and Trueb (1986) suggested that the Dugts specimens either bear erroneous locality included a photograph of the head and gills. Gomez Rios (1989) data (actually coming from Lake Pitzcuaro) or represent another provided excellent color photographs of the testis and histological species of Ambystoma. They suggested treating Ambystoma details of the spermatogenic cycle. gumtamis provisionally as a separate species, a position with which Frost (1985) disagreed. This taxon was not recognized by Distribution. Ambystoma dumerilii is known only from Floresand Gerez (1988). Direct examinationshouldestablish whether Lake Pitzcuaro, a large (ca. 111 km2), moderately shallow (to 15m), or not the two Dug& Museum specimens represent the distinctive A. relatively highelevation (2035 m) lake (Buen, 1944) at the southern dumerilii. Nor would understanding how they might have become edge of the Mexican Plateau in Michoadn, Mexico. Some DugPs associated with erroneous locality information be difficult. A similar specimens sometimes said to be from San Juan del Rio, Queretaro error occurred with syntypes USNM 16201-16202. In the USNM they (not Guanajuato, as pointed out by Maldonado-Koerdell, 1948a) were catalogued originally as being from Guanajuato, from where probably originated fromLake Ptitzcuaro (Taylor, 1940;Tien, 1958). Dug& sent them. A copy of Dug&' original list of specimens sent to See Comment. the USNM (provided by Robert P. Reynolds), however, lists the specimensas being from 'Lacde Pitzcuaro," andthe USNMcatalogue Fossil Record. None. entry subsequently was corrected. Taylor(1940) thought "... it seems certain that this specimen [syntype USNM 162011 originated in Lake Perdnent Literature. Brandon (1970a) described the Ptitzcuaro and was sent to the U.S. National Museum from the breeding season (December through spring) and (1970b) courtship Guanajuato locality by Duges." Further evidence of early confusion behavior. Gallien and Aimar (1974) examined nucleocpoplasmic iscope's (1889) comment that 'Asecond species, the S. dum&rili,has incompatibility in hybrids obtained by transplanting A. dumerilii been described by Dug& [sic] from Lake Patzcuaor [Pitzcuaro] in the nuclei into A. mPxicanum eggs. Brandon (1976) described the fatal State of Guanajuato." consequencesofmetamorphosisin the laboratory. Federet al. (1982) According to Reynolds (pers. comm.), female syntype USNM listed bodytemperatures in the field. Shaffer(l984a, b) examined the 16201 (Dugb field number 168) has head and body dimensions evolutionary relationships of A. dumerilii and other Mexican exactly as reponed by Dug& (1870) and is full of tiny eggs as shown ambystomat ids within the context of the evolution of paedomorpho- in his figure 1, plate 5; and syntype USNM 16202 is a male measuring ca. 130 mm snout to anterior end of vent. Without more convincing Gomez Rlls, J.G. 1989. Estudio histol6gico del aparato reproductor evidence to the contrary, all specimens of this distinctive species masculino del 'achoque" (Ambystoma dumen'lii, Dug&), uro- should be considered to have come from Lake Pitzcuaro. delo del Iagode Pitzcuaro. Bi6logoTesis, Universidad Nacional Aut6noma de Mexico. Literature Cited Gorham, S.W. 1974. Checklist of world amphibians. The New Bmns- wick Museum. St. John, New Bmnswick. Bailey, V. 1904. 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Hybridation nuclCocytoplasmique inters+cifique realisee par greffe nucleaire entre los eseces Published30 November 1992andCopyrightO 1992bythe Society for Ambystoma mem'canumShawet Ambystomadumerilii, Dugb the Study of Amphibians and Reptiles. (Amphibiens, Urodeles). C.R. Acad. Sci. Paris D 279:595-598.