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Systematic Status of Pseudoeurycea Unguidentis

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Systematic Status of Pseudoeurycea Unguidentis 46 HERPETOLOGICA [Vol. 33, No. 1 FERM, V. H., AND S. J. CARPENTER. 1968. Mal- SANDERS, H. 0. 1970. Pesticide toxicities to formations induced by sodium arsenate. J. tadpoles of the westem chorus frog Pseudacris Reprod. Fertil. 17:199-201. triseriata and Fowler's toad Bufo woodhousii FERM, V. H., A. SAXON, AND B. M. SMrTH. 1971. fowleri. Copeia 1970:246-251. The teratogenic profile of sodium arsenate in SOKAL, R. R., AND F. J. ROHLF. 1969. Biometry. the golden hamster. Arch. Environ. Health W. H. Freeman and Co., San Francisco. 22:557-560. SWIGGART,R. C., C. J. WHITEHEAD, JR., A. HAZELWOOD, E. 1970. Frog pond contaminated. CURLEY, AND F. E. KELLOG. 1972. Wildlife British J. Herpetol. 4:177-185. kill resulting from the misuse of arsenic acid HOOD,R. D., AND S. L. BISHOP. 1972. Terato- herbicide. Bull. Environ. Contam. Toxicol. 8: genic effects of sodium arsenate in mice. Arch. 122-128. Environ. Health 24:62-65. 1976 MATSUMURA, F. 1972. Current pesticide situa- Received: 16 June tion in the United States, p. 33-60. In: F. Accepted: 23 September 1976 Matsumura,G. M. Boush, and T. Misato [eds.] Environmental toxicology of pesticides. Aca- Department of Biology, Pan American demic Press, New York. University,Edinburg, Texas 78539, USA STUDIES OF NEOTROPICAL SALAMANDERS OF THE GENUS PSEUDOEURYCEA, I: SYSTEMATIC STATUS OF PSEUDOEURYCEA UNGUIDENTIS JAMES F. LYNCH, SUH YUNG YANG, AND THEODOREJ. PAPENFUSS ABSTRACT: The systematic status of Pseudoeurycea unguidentis, an enigmatic Mexican plethodontid salamander described by Taylor (1941), is reviewed in the light of morpho- metric and biochemical analysis of recently collected material. It is concluded that P. un- guidentis is a valid species which differs from Pseudoeurycea smithi, a superficially similar sympatric form, in numerous phenetic and genetic characters. Far from being conspecific, as suggested by an earlier investigator, these two salamanders are more different genetically than are most congeners in other animal groups. SOME 35 yr ago, E. H. Taylor (1941) smithi, influenced Bogert (1967) to syn- described Bolitoglossa unguidentis from a onymize the two forms in his treatment series of plethodontid salamanders col- of OaxacanPseudoeurycea. lected in 1938 on Cerro San Felipe and Field parties from the Museumof Verte- nearby Cerro San Luis in central Oaxaca, brate Zoology,University of California,and Mexico. The type series of the new species the Smithsonian Institution visited Cerro (which subsequently was assigned to the San Felipe on four occasions in 1974 and genus Pseudoeurycea) was taken in local 1975 in conjunction with studies of the sympatry with Pseudoeurycea smithi, a salamanderfauna of the region. On two of closely similar salamander. Except for the these trips (January and November 1974) mention of one additional specimen of P. smithi was the only Pseudoeuryceaen- Pseudoeuryceaunguidentis by Taylor and countered,but in August 1974 and August Smith (1945), the species has not been 1975 one of us (TJP) obtained series of reported since Taylor's original description. what appeared to be a different species in The lack of fresh comparative material, sympatrywith P. smithi. Both of the latter combined with the close resemblance be- collections were made at the same locality tween P. unguidentis and the common P. along the crest of a gently sloping ridge HERPETOLOGICA 33:46-52. March 1977 March 1977] HERPETOLOGICA 47 more vigorous in its attempts to escape and tended to occur off the ground more fre- quently, the loose bark of fallen logs being the favored microhabitat. The complex of morphological, ecological, and behav- ioral differences distinguishing the two Pseudoeurycea is reminiscent of that sepa- rating the slender, active, bark-dwelling Aneides ferreus from the more robust, sluggish, ground-dwelling Aneides flavi- punctatus where the two species are sym- FIG. 1.-Dorsal vi.ew of living adult P. mithi patric in northwestern California (Steb- *.XX.:ee.e::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~................................. bins, 1951; J. Lynch, personal observation). In his description, Taylor stated (1941: 61) that P. unguidentis differs from P. smithi ". in having longer limbs ... and larger hands and feet. The tail is somewhat longer and narrower." Because these dif- ferences correspond to those we perceived in the Pseudoeurycea from Cerro San Felipe, we provisionally assigned the sec- (top) and P. unguidentis (bottom) a a. Vertical ond form to P. unguidentis. To test our bar = 25 mm. Photograph by James Hendel, hypothesis that two distinct species (as Scientific Photographic Laboratory, University of California,Berkeley. opposed to mere morphological variants) were collected, we employed two inde- pendent modes of analysis: multivariate covered with humid, fairly open oak-pine morphometrics and electrophoretic screen- forest. Based on several altimeter readings ing of isozymes. taken over a 2-yr period, the elevation of this locality is 3050 + 25 m. Taylor (1941: METHODS 57) gives the elevation of the type locality of P. unguidentis as "about 2200 m", but The specimens reported in this paper in view of the rarity of specimens of any are deposited in the collection of the Mu- species of Pseudoeurycea at such low ele- seum of Vertebrate Zoology, Berkeley. A vations in central Oaxaca, and given the sample of living salamanders was shipped restriction of the sympatric P. smithi to to Berkeley for biochemical analysis in areas above 2800 m (Bogert, 1967), we August 1974; other specimens were killed believe Taylor's figure to be in error. by immersion in dilute chloretone solution Relative to sympatric P. smithi (Fig. 1A), within a few hours of capture, fixed in the second Pseudoeurycea (Fig. 1B) ap- 10% Formalin, and later transferred to peared to us to have a somewhat more 70% ethanol for permanent storage. slender habitus, longer legs, and lighter Morphometric Analysis.-The morpho- lateral and ventral pigmentation. These logical analysis is based on a detailed ex- subtle differences appeared to be consis- amination of 49 preserved P. smithi and tent, even though the overall similarity to P. smithi in size, build, and pigmentation 21 P. unguidentis. Discriminant function was indeed striking. Collecting notes sug- analysis was employed to specify patterns gested the presence of behavioral and eco- of interspecific differentiation. The follow- logical differences as well. Relative to P. ing characters were considered (all mea- smithi, the second form was noticeably surements in millimetres): head length 48 HERPETOLOGICA [Vol. 33, No. 1 (HL), the distance from tip of snout to were dissected and samples of blood and center of gular fold; body length (BL), tissue were removed. Liver, kidney, spleen, the distance from center of gular fold to stomach, heart, and skeletal muscle were posteriorangle of vent; head width (HW), homogenized using the methods of Se- the distance across broadest part of head; lander et al., (1971), and extracts were tail length (TL), the distance from pos- stored at -76? C. Combinedtissue extracts terior angle of vent to tip of tail; coupling and hemolysate fractions of blood were ratio (CR), the sum of the lengths of subjected to horizontal starch-gel electro- right fore limb and right hind limb divided phoresis as described by Selander et al., by the axilla-groindistance; maxillary tooth (1971). The following gel buffer systems count (MT), the total number of teeth on were used: (1) tris-hydrochloricacid (pH the maxillae; and vomerine tooth count 8.5) for hemoglobin (Hb); (2) lithium (VT), the total number of teeth on the hydroxide for albumin (ALB), transferrin vomers. Standard length (SL), the dis- (TRF), glutamate oxalate transaminases tance from tip of snout to posterior angle (GOT-1 and GOT-2), and peptidases of vent, was substituted for charactersHL (PEPT-1 and PEPT-2); (3) discontinuous and BL in comparisonsinvolving adult P. tris-citrate (Poulik) for lactate dehydrog- unguidentis, most of which had been dis- enases (LDH-1 and LDH-2); (4) con- sected in the gular region for tissue re- tinuous tris-citrate (pH 8.0) for sorbitol moval. dehydrogenase(SDH), alpha-glycerophos- Discriminantfunction is a multivariatesta- phate dehydrogenase (a GPD), isocitrate tistical technique that defines the weighted dehydrogenases(IDH-1 and IDH-2), man- combinationof characterswhich best sepa- nose phosphate isomerase (MPI), and su- rates the members of two or more pre- peroxidedismutase (SOD); (5) continuous defined groups. Each individual is as- tris-citrate (pH 7.0) for leucine amino- signed group membership a priori; the peptidase (LAP), malate dehydrogenases discriminantprocedure then specifies the (MDH-1 and MDH-2), phosphogluco- optimal set of weighted character coef- mutases (PGM-1 and PGM-2), and phos- ficients that maximizes the ratio of be- phoglucose isomerase (PGI); and (6) tris- tween-groupto within-groupvariance. The maleic EDTA for 6-phosphogluconate reader is referredto Blackithand Reyment dehydrogenase(6PGD). (1971) for a general discussion of the Genetic interpretationsof allozymicvari- applicationof this and related multivariate ation are based on criteria elaborated by techniques to morphometricstudies, and to Selander et al. (1971). Estimates of het- Lynch and Wake (1975) for an example erozygosityare derived from actual counts involving plethodontidsalamanders. of presumedheterozygotic genotypes. Mean Because ontogenetic and sexual variation heterozygosity (H) is defined as the num- in plethodontid salamanders can obscure ber of heterozygotic
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