Number 493 5 August 2002 Contributions IN Science Systematics of Xantusiid Lizards of the Genus Lepidophyma Robert L. Bezy and Jose L. Camarileo R. Natural History Museum OF Los Angeles County Serial Publications OL THE Natural History Museum oe Los Angeles County Scientific Publications Committee John Heyning, Deputy Director for Research and Collections John M. Liarris, Committee Chairman Brian V. Brown Gordon Hendler Ines Elorovitz Joel W. Martin K. Victoria Brown, Managing Editor The scientific publications of the Natural History Museum of Los Angeles County have been issued at irregular in- tervals in three major series; the issues in each series are numbered individually, and numbers run consecutively, re- gardless of the subject matter. # Contributions in Science, a miscellaneous series of tech- nical papers describing original research in the life and earth sciences. # Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth scienc- es. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now published by the Museum in Contributions in Science. # Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a miscel- laneous series containing information relative to scholarly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum’s Sci- entific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Technical Reports may be obtained from the relevant Section of the Museum. Natural History Museum OF Los Angeles County 900 Exposition Boulevard Los Angeles, Galieornia 90007 Printed at Allen Press, Inc., Lawrence, Kansas ISSN 0459-8113 Systematics of Xantusiid Lizards of the Genus Lepidophyma Robert L. Bezy^ and Jose L. Camarillo R.^ ABSTRACT. Multivariate analyses of variation in 30 scale characters among 129 locality samples of members of the genus Lepidophyma (A. Dumeril in Dumeril and Dumeril, 1851) identify the presence of 6 morphological groups in Mexico below the latitude of 19°N. Univariate comparisons among the samples diagnose a total of 17 morphological species in the genus, 10 of which occur in southern Mexico. Three species of Lepidophyma have broad (but discontinuous) geographic distributions: L. flavimaculatum A. Dumeril in Dumeril and Dumeril, 1851 (Veracruz to Panama), L. smithii Bocourt, 1876 (Guerrero to El Salvador), and L. sylvaticum Taylor, 1939 (Nuevo Leon to Veracruz). The other 14 members of the genus seem to have relatively small ranges, and 7 of these are known from only one or two localities. The greatest species diversity occurs in Oaxaca and Chiapas (10 species) and the northern Sierra Madre Oriental of Queretaro and San Luis Potosi (4 species). RESUMEN. Un analisis multivariado de variacion en 30 caracteres de escamas de 129 localidades con muestras de miembros del genero de Lepidophyma (A. Dumeril in Dumeril y Dumeril, 1851) identifica la presencia de 6 grupos morfologicos en Mexico por debajo de la latitud de 19°N. Comparaciones univar- iadas entre todas las muestras diagnostican un total de 17 especies morfologicas en el genero, 10 de las cuales se encuentran en el sureste de Mexico. Tres especies de Lepidophyma tienen una amplia (pero discontinua) distribucion geografica: L. flavimaculatum A. Dumeril in Dumeril y Dumeril, 1851 (Veracruz a Panama), L. smithii Bocourt, 1876 (Guerrero a El Salvador) y L. sylvaticum Taylor, 1939 (Nuevo Leon a Veracruz). Los otros 14 miembros del genero paracen tener areas de distribucion relativamente pequenas y 7 de estas se conocen unicamente de una o dos localidades. Las regiones con maxima diversidad de especies estan en Oaxaca y Chiapas (10 especies) y el norte de la Sierra Madre Oriental en Queretaro y San Luis Potosi (4 especies). INTRODUCTION The living members of the family Xantusiidae range from the southwestern U.S. to Panama and Cuba and comprise three genera: Xantusia Baird, 1859, Lepidophyma A. Dumeril in Dumeril and Dumeril, 1851, and Cricosaura Gundlach and Peters in Pe- ters, 1863. The majority of the diversity exists in the Middle American genus Lepidophyma, with 15 to 20 nominal species scattered from Nuevo Leon, Mexico, to Panama. Like other xantusiid lizards, members of the ge- nus Lepidophyma are secretive or reclusive and are seldom observed outside rock crevices, caves, and decaying tree stumps and logs. Most species are un- common and/or difficult to collect. Their geograph- ic distribution is spotty, with known localities often separated by considerable distances and represent- ed by few museum specimens. The distributional gaps may be the result of several factors, including narrow habitat requirements, rugged topography, forest destruction, and inadequate field sampling. That many of the disjunctions may be real rather 1. Herpetology, Natural History Museum of Los An- geles County, Los Angeles, California, 90007. 2. Laboratorio y Coleccion de Herpetologia, CyMA, Escuela Nacional de Estudios Profesionales Iztacala, A.P. 314, Tlalnepantla, Estado de Mexico, Mexico. Deceased 2002. Contributions in Science, Number 493, pp. 1-41 Natural History Museum of Los Angeles County, 2002 than artifactual is suggested by high incidence of morphological divergence among the localities. Since the description of Lepidophyma (Dumeril and Dumeril, 1851), a total of 25 taxa have been named in the genus, most of them based on one or two specimens from a single locality. Because many of the known localities are disjunct, represented by small sample sizes, and have various degrees of morphological differentiation, it remains unclear how many valid species exist in the genus, partic- ularly in Mexico (e.g., Flores-Villela, 1993; Liner, 1994; Smith, 1973). In previous studies, we have delineated species of Lepidophyma in northeastern Mexico (Bezy, 1984), Central America (Bezy, 1989), and within the clus- ter of species containing L. gaigeae Mosauer, 1936 (Bezy and Camarillo, 1992, 1997). In this paper we focus on delineating the morphologically diagnos- able species found in Mexico below the latitude of 19°N and synthesize the scattered information on the genus by providing summaries, diagnoses, and a key for all species of Lepidophyma. MATERIALS AND METHODS Data were taken for 31 scalation characters from a total of 1,420 specimens (including all holotypes) from 160 lo- calities throughout the range of Lepidophyma (see Speci- mens Examined). As in previous studies (Bezy, 1984, 1989; Bezy and Camarillo, 1992, 1997), we used discrim- 2 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma inant function analysis as an exploratory tool to visualize overall patterns of variation in these characters among lo- calities. For these analyses, the individual locality samples constituted the a priori groups, but for clarity of presen- tation in the resulting plots, envelopes were constructed to enclose all individuals from larger geographic areas (usually countries of Central America and states of Mex- ico). After delineation of the multivariate groups, the ob- served limits of variation of individual univariate charac- ters were examined to identify discrete (nonoverlapping) differences among the locality samples and sample clus- ters. Thus, the species are diagnosed by univariate features and are not dependent on multivariate analyses. All anal- yses were performed with Systat 7.0 (SSPS, Chicago, IL, 60611 USA). It is recognized that ratios violate the assumptions of a normal distribution and can influence the results of mul- tivariate analyses, particularly principal components anal- yses (e.g., Atchley et ah, 1976). In analyses of data, we found the results of discriminant analyses that used ratios generally to be congruent with those from analyses based on the residuals from regression of the morphometric var- iables on snout-vent length (SVL). Despite their statistical drawbacks, we employ ratios because we found them to be more useful in recognizing and diagnosing morpho- species. We embrace the evolutionary species concept (e.g., Wi- ley, 1978) and its operational equivalent, the phylogenetic species (e.g., Cracraft, 1983). We thus recognize as sepa- rate species any locality sample or group of samples that has 1 or more discrete difference(s) from each of the other samples or groups (i.e., for each pairwise comparison there are 1 or more characters for which the observed limits of variation do not overlap). In most cases these morpho-species are composed of disjunct populations and probably are not united by gene flow. Whether they ac- tually represent monophyletic groups of isolated popula- tions is difficult to determine with the quantitative scale features utilized in this study, but the question is being addressed by phylogenetic analyses of mitochondrial DNA sequences combined with morphology (R.L. Bezy and col- laborators, unpublished). In our analyses of variation, we make three exceptions to the requirement that species of Lepidophyma have dis- crete (nonoverlapping) differences in morphology. Two of the exceptions involve species pairs that are 100% diag- nosable by univariate differences in areas where they oc- cur in sympatry, but that have a small overlap in the ob- served limits of variation for geographically distant local- ities (where a combination of 2 univariate characters serves to distinguish the species). This is the operational equivalent of considering that “biological species” are a subset of “evolutionary species.” The third exception in- volves 2 allopatric species which have slight overlap in 2 morphological characters but have discrete differences in karyotypes throughout their ranges. We have chosen not to recognize subspecies in the genus because of the large number of geographically isolated samples with morphological features that are statistically different but that overlap the observed limits of variation in other samples. STANDARD CHARACTERS The standard data taken on all specimens consist of 31 scale characters: 21 meristic, 9 mensural, and 1 presence/ absence. We employ the scale terminology of Savage (1963) rather than that of Smith (1973) in that we con- sider the parietals to be the scales on each side of the I 23456789 10 II 12 13 14 25 26 27 Figure 1 Dorsal and lateral views of the head of a gen- eralized specimen of Lepidophyma. Numbered points were used in scale measurements and counts (see Materi- als and Methods). interparietal rather than the pair posterior to it. In the descriptions below, “at midbody” indicates that the count was made midway between the gular fold and the vent, above a point corresponding to half the total number of transverse rows of ventrals. Pretympanics (PTMP). The minimum number of small scales (pretympanics) separating the postoculars (Fig. 1: 26, 27) from the second postorbital supralabial (sum of right plus left side of head). The second postorbital su- pralabial (Fig. 1: 28) is usually the seventh and is the sec- ond labial scale posterior to the eye that does not enter the orbital ring of scales. Supralabial scales are skirted and not included in the count. For example, there are a min- imum of 4 pretympanics in Fig. 1, between points 27 and 29. Gulars (GUL). The number of small scales (gulars) along the ventral midline between the posterior gular fold and the second pair of infralabials (Fig. 2, between points A and B). Gulars Contacting First Infralabials (GCIIL). The num- ber of gulars in contact medially with first pair of infral- abials. (Fig. 2, 1 gular at point C). Dorsals (DOR). The number of scales along the mid- dorsal line between the posterior edge of the postparietals and a point directly above the vent. Dorsals Between Paravertebral Rows (DBPVR). The minimum number of dorsal scales separating the rows of large paravertebral tubercles (at midbody; Fig. 2, 5 dorsals separate tubercles D and F) Dorsals Along Paravertebral Row (DAPVR). The num- Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 3 Figure 2 Ventral view of head and dorsal view of body (axilla to groin) of a generalized specimen of Lepidophy- ma. Lettered points were used in scale counts (see Mate- rials and Methods). Figure 3 Dorsal (left) and ventral (right) views of the pre- anal area and the proximal portion of the tail of a gen- eralized specimen of Lepidophyma with characters indi- cated by letters (see Materials and Methods). ber of dorsals (from above the axilla to above the groin) in the row immediately above (i.e., medial to) the para- vertebral tubercle row (Fig. 2, G to H). Dorsals Between Paravertebral Tubercles (DBPVT). The number of middorsal scales equivalent to the distance between 2 consecutive large tubercles (largest size class) within the paravertebral row (at midbody; Fig. 2, the dis- tance between tubercles E and F is equal to 4 dorsals). Large Paravertebral Tubercles (PVTL). The number of large tubercles (largest size class) in the paravertebral row (from above the axilla to above the groin; Fig. 2, between I and J there are 18 large tubercles). Paravertebral Row (PVR). The total number of scales (all sizes) in the paravertebral row (from above the axilla to above the groin; Fig. 2, there are 54 scales between I and J). Paravertebrals, Small (PVS), The number of scales (from above the axilla to above the groin) in the paravertebral row that are shorter than 1.5 dorsal scales. Paravertebrals, 1.5 (PVT1.5). The number of scales (from above the axilla to above the groin) in the paraver- tebral row that are equal to or longer than 1.5 middorsal scales. Paravertebrals, 2 (PVT2). The number of scales (from above the axilla to above the groin) in the paravertebral row that are equal to or longer than 2 middorsal scales. Paravertebrals, 3 (PVT3). The number of scales (from above the axilla to above the groin) in the paravertebral row that are equal to or longer than 3 middorsal scales. Lateral Tubercle Rows (LTR). The number of large tu- bercular scales (largest size class) on the side of the body between the axilla and groin. The count is made along a longitudinal line that is parallel to the paravertebral row and that passes through the second large lateral tubercle below the paravertebrals (Fig. 2, there are 16 tubercle rows along line K to L). Dorsal Interwhorls (IWD2). The maximum number of annuli of small scales separating the second from the third whorl of enlarged caudal scales (at middorsum; Fig. 3, three dorsal interwhorls between M and N). Ventral Interwhorls (IWV2). The minimum number of ventrally complete annuli of small scales separating the second from the third whorl of enlarged caudal scales (Fig. 3, 3 complete ventral interwhorls between O and P). Ventrals, Longitudinal (VL). The number of transverse rows of large ventral scales between gulars and the vent, including the preanals. Ventrals, Across (VA). The number of longitudinal rows of ventrals across midbody. Longitudinal rows were ex- cluded if (at midbody) they contain 2 more scales that are m contact with a single scale of the medially adjacent lon- gitudinal row of ventrals. Femoral Pores (FPT). The total number of pores (right plus left femoral surface). In females and juveniles some or all of the pores may be vestigial or “undeveloped” (marked by shallow depressions rather than discrete glan- dular openings). On all specimens the counts include both “developed” and “undeveloped” pores. Fourth Toe Lamellae (FTL). The number of scales along the ventral midline of the fourth toe between the base and the claw. Divided Fourth Toe Lamellae (FTLD). The number of scales under the fourth toe with midventral sutures. Parietal Spot (PS). The presence (1) or absence (0) of a pale spot on the interparietal scale. The spot marks the approximate position of the parietal foramen (which may be roofed over by bone in adults) and is located along the dorsal midline, usually in the posterior half of the scale (Fig. 1, point 11). Prefrontal Length (RPFL). Ratio of the length along the midline of the lateral prefrontal (Fig. 1, points 4 to 6) divided by the length along the lateral border of the lateral prefrontal (Fig. 1: 3 to 7). Median Prefrontal Length (RML). Ratio of the length of the median prefrontal (Fig. 1: 5 to 8) divided by the length of the midline suture of frontals (Fig. 1: 8 to 9). 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O V. -2 q C/> S rS .u o V) V5 ■2 ro s o 04 ro VO H-i VO d ro d T-l d 8 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Table 2, Variation in 10 scale characters among samples of group S of Lepidophyma. The observed limits of variation are followed by the mean (in parentheses). N FPT LTR GUL DOR L. tarascae Michoacan 4 14-18 (16.8) 16-25 (18.8) 40-43 (43.8) 145-159 (149.8) L. lineri Oaxaca El Pacifico 2 16 23-24 (23.5) 51-52 179-184 (181.5) Alemania 1 17 24 — — L. smithii Veracruz 1 20 18 51 198 Guerrero 17 26-30 (27.3) 15-19(16.7) 50-57 (53.6) 176-194 (185.6) Oaxaca 78 15-26 (20.2) 15-20 (17.8) 44-59 (52.0) 162-213 (191.4) Puerto Escondido 2 15-16 (15.5) 17-18 (17.5) 44-49 (46.5) 173-189(181.0) Tehuantepec 30 16-26 (21.5) 15-20 (17.4) 44-57 (51.0) 164-209 (187.1) Tapanatepec 20 16-21 (19.3) 15-19 (16.8) 49-59 (54.4) 190-213 (200.8) Chiapas 150 16-27 (21.1) 16-22 (17.9) 44-58 (51.9) 182-224 (202.7) Tonala 9 16-22 (18.9) 17-18 (17.4) 46-58 (52.3) 189-219 (204.1) Escuintla 136 17-27 (21.3) 16-22 (18.0) 44-58 (51.8) 182-224 (202.5) Guatemala 54 17-24 (20.8) 15-21 (17.4) 46-61 (52.9) 173-217(198.3) El Salvador 3 16-24 (18.7) 16-18 (17.0) 46-51 (48.3) 196-208 (203.0) Median Prefrontal Width (RMW2). Ratio of the width of the median prefrontal (Fig. 1: 6 to 21) divided by height of the second postorbital supralabial (Fig. 1: 28 to 29). Specimens lacking a median prefrontal were scored as 0. Nasal Length (RNL). Ratio of the length of the midline suture of the nasals (Fig. 1: 1 to 2) divided by length of the midline suture of the postparietals (Fig. 1: 15 to 17). Anomalous Postparietal Suture Length (RAPPSL). Ratio of the total length of all anomalous sutures on both post- parietals (Fig. 1: 13 to 14 plus 16 to 18) divided by length of the midline suture of postparietals (Fig. 1: 15 to 17). Postnasal Height (RPNH). Ratio of the height of the postnasal (Fig. 1: 22 to 23) divided by height of the an- terior loreal (Fig. 1: 22 to 24). Postocular Length (RPOL). Ratio of the maximum length of the postoculars (Fig. 1: 26 to 27) divided by length of the orbit (Fig. 1: 25 to 26). Supralabial Height (RSLH). Ratio of the height of the second postorbital supralabial (Fig. 1: 28 to 29) divided by height of the first postorbital supralabial (Fig. 1: 30 to 31). The first postorbital supralabial is usually the sixth and is the first labial scale posterior to the eye that does not enter the orbital ring of scales. Preanal Width (RPAW). Ratio of the maximum width of the lateral posterior preanal (Fig. 3: S) divided by max- imum width of the median posterior preanal (Fig. 3: R). AUXILIARY CHARACTERS The following 5 additional characters were used for com- parisons among the samples in one of the clusters (group S). Median Prefrontal (M). Presence (1) or absence (0) of a median prefrontal. Median Prefrontal Contact (MC). Presence (1) or ab- sence (0) of a contact between the median prefrontal and the frontal. Specimens lacking a median prefrontal were scored as 0. Median Prefrontal Length 2 (RML2). Ratio of the length of the median prefrontal (Fig. 1: 5 to 8) divided by length of the midline suture of postparietals (Fig. 1: 15 to 17). The character was scored as absent in specimens lack- ing a median prefrontal. Prefrontal Length 2 (RPFL2). Ratio of the length along the midline of the lateral prefrontal (Fig. 1: 4 to 6) divided by length of the midline suture of the nasals (Fig. 1: 1 to 2). Frontal Length 2 (RFL2). Ratio of the length of the midline suture of frontals (Fig. 1: 8 to 9) divided by length of the midline suture of postparietals (Fig. 1: 15 to 17). RESULTS AND DISCUSSION ANALYSIS OF VARIATION AND DELINEATION OF SPECIES Southern Groups An initial discriminant function analysis was per- formed with the 30 basic meristic and mensural scale characters (excluding the presence/absence of the parietal spot; Table 1) for all 129 locality sam- ples from southern Mexico (below 19°N) and Cen- tral America. The analysis revealed three nonover- lapping multivariate clusters (groups S, T, and F), that each contain multiple locality samples (Fig. 4). For these three groups, variation in individual char- acters among the samples is analyzed, the diagnos- able units are identified, and species names are as- signed. In addition to these three multilocality groups, the initial discriminant analysis identified three clusters (D, L, and R), which each contain a single locality sample. Previous work (Bezy and Camarillo, 1992, 1997) demonstrated that each of the three samples represents a morphologically dis- crete species (L. dontomasi Smith, 1942; L. lowei Bezy and Camarillo, 1997; and L. radula Smith, Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 9 Table 2. Extended. RPAW M MC RML2 RPFL2 RFL2 0.41-0.71 (0.62) 0 0 1.44-4.88 (2.53) 0.60-0.89 (0.77) 0.81-0.87 (0.84) 1 0-1 (0.50) 0.14-0.21 (0.18) 1.55-2.49 (2.02) 1.39-1.48 (1.44) 0.56 1 — — — — 0.54 1 0 0.62 3.64 0.88 0.91-1.35 (1.09) 0 0 — 2.56-4.24 (3.45) 0.39-0.67 (0.50) 0.40-1.10 (0.68) 0-1 (0.47) 0-1 (0.08) 0.13-0.80 1.00-4.50 (2.45) 0.23-1.70 (0.79) 0.64-0.65 (0.64) 0 0 — 3.40-3.62 (3.51) 0.47-0.67 (0.57) 0.40-1.10 (0.65) 0-1 (0.33) 0 0.13-0.47 (0.31) 1.00-3.50 (2.33) 0.29-1.62 (0.75) 0.53-0.90 (0.74) 0-1 (0.20) 0-1 (0.20) 0.26-0.80 (0.53) 1.46-4.50 (2.43) 0.35-1.35 (0.86) 0.33-1.01 (0.63) 0-1 (0.99) 0-1 (0.36) 0.36-0.99 (0.56) 1.79-15.15 (3.55) 0.24-1.32 (0.57) 0.44-1.01 (0.73) 1 1 0.38-0.62 (0.50) 2.41-3.86 (3.17) 0.35-0.73 (0.56) 0.33-1.24 (0.62) 0-1 (0.99) 0-1 (0.37) 0.04-0.99 (0.56) 1.79-15.15 (3.59) 0.24-1.32 (0.58) 0.42-1.24 (0.65) 1 0-1 (0.95) 0.48-1.08 (0.74) 2.09-7.95 (3.58) 0.33-1.02 (0.62) 0.47-0.71 (0.57) 1 0 0.31-0.54 (0.44) 3.28-4.12 (3.61) 0.38-0.57 (0.46) 1942, respectively), and these analyses are not re- peated here. However, all species of Lepidophyma recognized as valid are included in the summary and key. GROUP S. The group contains 307 specimens from 33 localities on the Pacific slope from El Sal- vador to Michoacan (see Specimens Examined for a list of the names of Mexican states that are used in the text without reference to country). Members of the group are characterized by the absence in adults (over 75 mm SVL) of an externally visible parietal spot (PS; Table 1). C.V.1 Figure 4 Plots of individual specimen scores on the first two canonical variates for 129 locality samples of Lepi- dophyma from southern Mexico and Central America. Envelopes enclose the specimens for the six nonoverlap- ping groups present. Discriminant function analysis of variation in the 30 characters among the samples of group S re- vealed that the specimens from Michoacan and Guerrero each constitute a discrete cluster (SI and S2, respectively), with a third cluster (S3) composed of multivariately overlapping samples from Oaxaca to El Salvador (Fig. 5). The Michoacan cluster (SI) contains 4 specimens from 2 localities. Univariate comparisons among samples in group S indicate that the specimens from Michoacan have discrete (nonoverlapping) differ- ences in number of dorsals and gulars from all oth- Figure 5 Plots of individual specimen scores on the first two canonical variates for locality samples of group S of Lepidophyma (from Fig. 4). Three nonoverlapping groups (Si, S2, and S3) are present. 10 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Table 3. Variation in 12 scale characters among samples of group T of Lepidophyma. The observed limits of variation are followed by the mean (in parentheses). N FPT LTR IWD2 PTMP PVTL L. mayae Guatemala 16 29-35 (32.3) 33-46 (0.83) 3-4 (3.9) 1-2 (1.9) 21-46 (34.5) L. pajapanense Veracruz 22 30-36 (0.33) 36-43 (40.0) 3-4 (3.1) 6-10 (7.5) 39-51 (43.5) L. tuxtlae Veracruz Los Tuxtlas 85 20-29 (24.5) 30-37 (34.0) 2-5 (3.6) 4-9 (5.5) 30-49 (39.2) Jesus Carranza 1 25 34 3 5 41 Oaxaca Vista Hermosa 6 18-26 (23.5) 35-40 (36.5) 4-5 (4.2) 4-8 (5.7) 34-45 (39.3) La Gloria 1 26 35 4 5 46 Matias Romero 1 23 — 3 5 — Chiapas Ocote 4 25-26 (25.5) 31-33 (31.5) 3-4 (3.8) 4-6 (4.8) 40-46 (43.3) er members of the group (Table 2). The cluster con- tains the holotype of L. tarascae Bezy, Webb, and Alvarez, 1982, and this name is assigned to these specimens. The Guerrero cluster (S2) contains 17 specimens from 1 locality, and these have a higher average number of femoral pores than other members of group S (Table 2). However, the observed limits of variation for femoral pores in the Guerrero sample (S2) slightly overlap those of specimens from Oa- xaca and Chiapas (S3), and thus clusters S2 and S3 are judged to be conspecific. Comparisons among the samples of S2 and S3 reveal that 2 localities appear to share a scale fea- ture that is nonoverlapping with other members of Figure 6 Number of longitudinal rows of ventral scales bearing keels plotted against snout-vent length for speci- mens of Lepidophyma lineri (filled circles) and L. smithii (vertical lines). the group. The two individuals from Cafetal El Pa- dfiico (LACM 134844, UCM 48452) and one dam- aged specimen from the adjacent locality of Cafetal Alemania (AMNH 15977) in the Sierra Madre del Sur of Oaxaca differ in number of lateral tubercle rows from all other specimens of clusters S2 and S3 (Table 2). The 3 specimens include the holotype of L. flavimaculatum lineri Smith, 1973, and they are recognized as a distinct species, L. lineri. Smith (1973) originally described this taxon as a subspecies of L. flavimaculatum, characterized by the following features (abbreviations in parentheses denote equivalent characters in Table 2): a minute median prefrontal (RML2), subtriangular lateral prefrontals (RPFL2), a large frontal (RFL2), and large lateral preanals (RPAW). The 2 specimens of L. lineri from the type locality appear to have av- erage differences from other samples of clusters S2 and S3 in these 4 characters, but there is extensive overlap in the observed limits of variation (Table 2). As noted by Smith (1973) the diagnostic value of the presence of keeled ventrals in this taxon is questionable because the known specimens have a small snout-vent length and this feature is nega- tively correlated with body size (Fig. 6). Although the original diagnostic features used by Smith (1973) appear to be overlapping with other sam- ples, the 2 specimens from the type locality (El Pa- cifico, Oaxaca) have a distinctly higher number of lateral tubercle rows, a feature shared with the specimen from the adjacent locality, Cafetal Ale- mania (Table 2). Lepidophyma lineri is here rec- ognized as a separate species on the basis of its dis- crete difference in number of lateral tubercle rows from other members of group S, but this conclusion must be considered tentative because the taxon is represented by only 3 specimens. Although significant differences exist among some of the other samples in S2 and S3, there is Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 1 1 Table 3. Extended. FTL FTLD DOR DAPVR MC RML RPAW 23-28 (25.1) 10-19 (14.8) 162-188 (174.8) 72-91 (79.7) 0-1 (0.81) 1.09-2.16 (2.49) 0.38-0.77 (0.50) 26-30 (28.5) 10-18 (14.2) 152-177 (165.1) 59-82 (70.0) 0-1 (0.32) 1.00-2.81 (1.73) 0.38-0.69 (0.54) 23-29 (26.0) 26 13-21 (16.6) 15 158-190 (173.5) 169 69-98 (80.2) 76 0-1 (0.89) 0 0.49-3.15 (1.32) 1.24 0.56-1.12(0.83) 0.83 22-26 (23.3) 25 12-15 (13.8) 14 150-171 (159.2) 163 71-84 (76.3) 77 0-1 (0.67) 0 0 0.54-2.20 (1.34) 0.98 0 0.70-0.87 (0.80) 0.73 0.74 26-27 (26.5) 14-16 (14.8) 165-177(171.0) 73-80 (77.8) 0-1 (0.25) 0.00-1.66 (0.91) 0.59-0.68 (0.63) overlap in the observed limits of variation for all scale characters examined and the remaining sam- ples in these clusters are considered to represent a single species. Cluster S3 includes the holotypes of L. smithii Bocourt, 1876; Akleistops guatemalensis Muller, 1877; and L. smithii tehuanae Smith, 1942. The oldest available name, L. smithii, is assigned to the remaining members of clusters S2 and S3. Smith (1942, 1973) distinguished L. flavimacu- latum tehuanae from L. flavimaculatum smithii by its lack of a median prefrontal. The presence (M) and size (RML2) of the median prefrontal and the presence of a contact between the median prefron- tal and the frontal (MC) appear to vary geograph- ically (Table 2). A median prefrontal is present in 33% of the specimens from the vicinity of Tehuan- tepec, Oaxaca, which encompasses the type locality -6.0 -1.5 3.0 7.5 12.0 C.V.1 Figure 7 Plots of individual specimen scores on the first two canonical variates for locality samples of group T of Lepidophyma (from Fig. 4). Three nonoverlapping groups (Tl, T2, and T3) are present. of L. f. tehuanae. No discrete differences between L. f. tehuanae and L. smithii are apparent in any of the scale characters examined in this study (Ta- ble 2) and the two are considered conspecific. The relationship of L. smithii to L. flavimaculatum is discussed under group F, below. On the basis of the above comparisons it is con- cluded that group S contains 3 morphologically dis- crete species: L. tarascae (Michoacan), L. lineri (two localities in the Sierra Madre del Sur, Oaxaca), and L. smithii (Pacific slope from Guerrero to El Salvador). GROUP T. The group contains 136 specimens from 13 localities from Veracruz and Oaxaca to Guatemala. Discriminant function analysis of var- iation in the 30 characters indicates that the sam- ples form three discrete clusters (Tl, T2, and T3; Fig. 7). Cluster Tl is composed of 16 specimens from 4 localities in Guatemala. Univariate comparisons identify no discrete differences among the 4 locality samples. Cluster Tl differs from all specimens of clusters T2 and T3 in pretympanics and from L. smithii, L. lineri, and L. tarascae in lateral tubercle rows (Table 1). The cluster contains one holotype, that of L. mayae Bezy, 1973, and this name is ap- plied to the species. Cluster T2 contains 22 specimens from 3 regions in Veracruz. Univariate comparisons identify no discrete differences among the samples from these regions. The specimens in cluster T2 differ from all individuals of cluster T3 in femoral pores and fourth toe lamellae and from L. smithii, L. lineri, and L. tarascae in lateral tubercle rows and large paravertebrals (Table 1). The cluster contains the holotype of L. pajapanense Werler, 1957, and this name is applied to the species. Cluster T3 includes 98 specimens from 6 locali- ties in Veracruz, Oaxaca, and Chiapas. Univariate comparisons among the six samples do not reveal 12 ■ Contributions in Science, Number 493 Figure 8 Number of longitudinal rows of ventral scales bearing keels plotted against snout-vent length for speci- mens of Lepidophyma tuxtlae from the Ocote region of Chiapas (filled circles) and the Tuxtlas region of Veracruz (vertical lines). Bezy and Camarillo: Systematics of Lepidophyma C.V.1 Figure 9 Plots of individual specimen scores on the first two canonical variates for locality samples of group F of Lepidophyma (from Fig. 4). Three nonoverlapping groups (FI, F2, and F3) are present. any discrete differences. The specimens in cluster T3 differ from L. smith ii, L. lineri, and L. tarascae in lateral tubercle rows and large paravertebrals. The cluster includes the holotypes of L. tuxtlae Werler and Shannon, 1957; L. saivini Smith, 1973; and L. alvarezi Smith, 1973. The oldest available name, L. tuxtlae, is assigned to the members of T3. Smith (1973) described L. alvarezi based on a specimen from the Ocote region of Chiapas. It was distinguished from L. tuxtlae by having keeled ven- trals, smaller lateral preanals, and a smaller median prefrontal. The 4 specimens (of group T3) exam- ined from the Ocote region have slight average dif- ferences from the sample of L. tuxtlae from the Tuxtlas in relative sizes of the median prefrontal and lateral preanals and in number of femoral pores, lateral tubercle rows, pretympanics, large paravertebrals, and dorsals along the paravertebral row, but there is overlap in the ranges of variation for all scale characters examined (Table 3). The number of longitudinal rows of ventrals bearing keels is negatively correlated with snout-vent length and only one of the 4 Ocote specimens is outside of the range of variation found in the L. tuxtlae from the Tuxtla region (Fig. 8). Lepidophy- ma alvarezi is considered conspecific with L. tux- tlae. A second species, L. sawini, was described by Smith based on a specimen from near Vista Her- mosa, Oaxaca. It was distinguished from L. tuxtlae by having relatively large keeled scales bordering the paravertebrals medially. Compared to L. tux- tlae from the Tuxtla region, the 6 specimens now available from the Vista Hermosa region have slightly larger (fewer) scales bordering the para- vertebrals medially (DAPVR; Table 3), but there is extensive overlap in the observed limits of varia- tion. The Vista Hermosa specimens have virtually the same numbers of femoral pores as L. tuxtlae from the Tuxtlas and the observed limits of varia- tion do not overlap those of L. pajapanense (Table 3). The Vista Hermosa specimens resemble L. tux- tlae (rather than L. pajapanense) from the Tuxtla region in color pattern (Figs. 23, 30). Although there are slight average differences between the Vis- ta Hermosa sample and L. tuxtlae from the Tuxtla region (Table 3) in lateral tubercle rows, dorsal cau- dal interwhorls, fourth toe lamellae, divided fourth toe lamellae, dorsals, and dorsals along paraverte- bral row, there is overlap in all characters examined and L. sawini is considered conspecific with L. tux- tlae. Three species are recognized as valid in group T: L. mayae (Guatemala), L. pajapanense (Veracruz), and L. tuxtlae (Veracruz, Oaxaca, and Chiapas). GROUP F. The group contains 507 specimens from 80 localities in Veracruz and Oaxaca to Pan- ama. Discriminant function analysis with the 30 characters indicates that the samples form three clusters: FI, F2, and F3 (Fig. 9). Cluster FI contains 4 specimens from 1 locality in Chiapas. The specimens differ from members of clusters F2 and F3, as well as from all other Lepi- dophyma, in having a greater number of fourth toe lamellae (Table 1). All 4 specimens have 12 longi- tudinal rows of ventrals at midbody (contra Alvar- ez and Valentin, 1988), a condition that is rare within Lepidophyma, but present in one or more individuals of 6 other species of the genus (Table 1). The cluster contains the holotype of L. chicoas- Contributions in Science, Number 493 ense Alvarez and Valentin, 1988, and this name is applied to the species. Cluster F2 contains 2 specimens from 1 locality (Rio de La Venta) in Chiapas. Although the speci- mens share some distinctive features, there are no- table differences between them in several scale characters (Fig. 9) and the 2 are discussed sepa- rately. One of the specimens, the holotype of L, lipetzi Smith and Alvarez del Toro, 1977 (UCM 51425), has a higher number of large paravertebral tubercles than do all members of cluster F3. In ad- dition, it differs in having a higher number of dor- sal scales and in having a pale color pattern from all F3 specimens from Chiapas, including those from the sympatric (Rio de La Venta) and nearby (Selva Ocote) localities. It resembles the 4 speci- mens of L. chicoasense in having 12 longitudinal rows of ventrals and a pale color pattern, but dif- fers in number of fourth toe lamellae. The second F2 specimen (ENEPI 3792) from Rio de La Venta resembles the holotype of L. lipetzi in number of lateral tubercle rows and dorsals and in having a pale color pattern, clearly differing from sympatric (La Venta) and nearby (Ocote) specimens of group F3 in these characters. However, it differs from the holotype in having 11 (vs 12) longitudinal ventral rows, the sixth row from the midline being clearly larger than the adjacent tubercles on only one side of the body. It also differs from the holo- type in number of large paravertebral tubercles, re- sembling F3 specimens in this feature. It is tenta- tively grouped with the holotype in L. lipetzi. The 2 specimens thus assigned to L. lipetzi differ from L. chicoasense in fourth toe lamellae; from L. smi- thii, L. lineri, L. tarascae, and L. tuxtlae in femoral pores; and from L. mayae and L. pajapanense in gulars (Table 1). They differ in color pattern from all members of group F3 and in number of dorsal scales from sympatric (La Venta) and nearby (Oco- te) localities for that group. Variation among Central American locality sam- ples of cluster F3 was examined in detail previously (Bezy, 1989) and resulted in the recognition of L. reticulatum Taylor, 1955 (Pacific slope of Costa Rica) as distinct from L. flavimaculatum Dumeril, 1851 (Atlantic slope from Mexico to Panama); these analyses are not repeated here. The discrimi- nant function analyses indicate that all samples of L. flavimaculatum from southern Mexico (except those from Chiapas) cluster with samples from northern Central America (Guatemala to Hondu- ras). The samples from Chiapas broadly overlap both the southern (Nicaragua to Panama) and northern Central American clusters. No discrete univariate differences are apparent among the sam- ples of L. flavimaculatum from southern Mexico. Lepidophyma flavimaculatum has discrete differ- ences in scalation (Table 1) from L. lineri and L. tarascae (femoral pores), L. chicoasense (fourth toe lamellae), and L. tuxtlae and L. pajapanense (large paravertebral tubercles). It differs from sympatric L. mayae in lateral tubercle rows and pretympan- Bezy and Camarillo: Systematics of Lepidophyma ■ 13 Figure 10 Plots of individual specimen scores on the first two canonical variates for locality samples of Lepidophy- ma sylvaticum and L. flavimaculatum. Of the 2 specimens of L. sylvaticum from Veracruz (VC), the more southern one (a) is placed within L. sylvaticum, whereas the more northern one (b) is placed intermediate between the two species. ics, and from sympatric (Chiapas) L. lipetzi in num- ber of dorsals. Lepidophyma flavimaculatum and L. smithii comprise multivariately discrete clusters (groups F and S in Fig. 4), including the specimens from where their ranges are in close proximity in Oa- xaca. Throughout the extensive ranges of the 2 spe- cies they differ in lateral tubercle rows. In addition, L. flavimaculatum has an externally visible pale pa- rietal spot that is absent in adult L. smithii (SVL greater than 75 mm). The 2 also differ in karyotype (Bezy, 1972) and are recognized as discrete species. Four species are recognized as valid in group F: L. chicoasense (Chiapas), L. lipetzi (Chiapas), L. reticulatum (Costa Rica), and L. flavimaculatum (Veracruz and Oaxaca to Panama). Comparisons with Species in Northeastern Mexico Bezy (1984) recognized 4 species of Lepidophyma as valid in northeastern Mexico: L. gaigeae Mo- sauer, 1936; L. micropholis Walker, 1955; L. oc- culor Smith, 1942; and L. sylvaticum Taylor, 1939. Lepidophyma micropholis differs in number of dor- sals (Table 1) from all southern species delineated above. Lepidophyma gaigeae also differs in dorsals from all southern species except L. radula, from which it differs in femoral pores. Lepidophyma oc- culor differs in gulars from all southern species ex- cept L. smithii, from which it differs in having a parietal spot in adults and a different karyotype. Lepidophyma sylvaticum has discrete differences (Table 1) from L. tarascae and L. lineri (femoral pores), L. tuxtlae (dorsals separating paravertebral 14 ■ Contributions in Science, Number 493 tubercles), L. pajapanense (pretympanics), L. chi- coasense (fourth toe lamellae), L. smithii (presence of a pale parietal spot in adults), L. lipetzi (height of second postorbital infralabial), and L. mayae (median prefrontal width). Lepidophyma sylvaticum and L. flavimaculatiim form discrete multivariate clusters (Fig. 10). Where their ranges are in close proximity in Veracruz, the 6 specimens of L. flavimaculatum and the south- ernmost specimen of L. sylvaticum (Fig. 10, point a\ MZFC 1722 from Alvarado) are placed well within their respective species clusters. However, the more northern Veracruz specimen of L. sylva- ticum (Fig. 10, point b; KU 26909 from Tlapacoy- am) is placed somewhat intermediate between the 2 species clusters. Nevertheless, the 2 L. sylvaticum and 6 L. flavimaculatum from Veracruz are non- overlapping in 5 scale characters (femoral pores, lateral tubercle rows, caudal interwhorls, pretym- panics, gulars contacting the first infralabials, and height of the second postorbital supralabial). For the 545 specimens examined from throughout the ranges, the 2 species generally differ in number of pretympanics and in the relative height of the sec- ond postorbital supralabial. All but one L. flavi- maculatum (UCM 49278 from Chiapas) have 4 or more pretympanics and all but 2 L. sylvaticum (AMNH 107273, UMMZ 101374, both from Ta- maulipas) have 3 or fewer pretympanics. For the relative height of the second postorbital supralabi- al, all specimens of L. sylvaticum have a RSLH of 0.84 or greater, and all but 3 specimens of L. flav- imaculatum (UMMZ 113777 from Tobasco, MVZ 159933 from Guatemala, AMNH 107589 from Panama) have a RSLH of less than 0.84. No spec- imens overlap in both characters. The 2 also have consistent differences in the number and morphol- ogy of chromosomes (Bezy, 1984) and are consid- ered to represent discrete species. SUMMARY OF THE EXTANT SPECIES OE THE GENUS LEPIDOPHYMA Lepidophyma A. Dumeril Lepidophyma A. Dumeril, in Dumeril and Dumeril, 1851:137. Type species: Lepidophyma flavima- culatus A. Dumeril. Poriodogaster Smith in Gray, 1863:154, Plate XXL Type species: Poriodogaster grayii Smith. Akleistops Muller, 1877:390, Tafel I, II. Type spe- cies, Akleistops guatemalensis Muller. Gaigeia Smith, 1939:24. Type species: Lepidophy- ma gaigeae Mosauer. DIAGNOSTIC CHARACTERS. The living members of the genus Lepidophyma differ from Cricosaura typica (condition in parentheses) in hav- ing 1 frontonasal (2), 2 frontals (1), and 2 parietals (none); and from living members of the genus Xan- tusia (condition in parentheses) in having a round pupil (vertically elliptical); in having the nostrils placed in the suture between the nasal and post- Bezy and Camarillo: Systematics of Lepidophyma nasal (in the suture between the nasal, postnasal, and rostral); and no postmentals (postmentals dis- tinct from infralabials; Savage, 1963; R.L. Bezy and J.L. Camarillo, personal observatons). CONTENT. A total of 17 extant species of Lep- idophyma are recognized as a result of this study: L. chicoasense, L. dontomasi, L. flavimaculatum, L. gaigeae, L. lineri, L. lipetzi, L. lowei, L. mayae, L. micropholis, L. occulor, L. pajapanense, L. rad- ula, L. reticulatum, L. smithii, L. sylvaticum, L. tarascae, and L. tuxtlae. The validity of L. arizel- oglyphum (Langebartel, 1953) and other fossil taxa (Estes, 1983) not included in this study requires ad- ditional investigation focusing on osteology. KARYOTYPES. The diploid chromosome num- bers of species in the genus range from 32 to 38; numbers of macrochromosomes, from 16 to 18; and numbers of microchromosomes, from 16 to 20 (Bezy, 1972; Bezy and Camarillo, 1992, 1997). The karyotypes of members of Lepidophyma differ from those of members of Xantusia (Bezy, 1972; Bezy and Flores Villela, 1999) in having lower dip- loid numbers (32-38 vs 40) and fewer microchro- mosomes (16-20 vs 22); and from that of Crico- saura typica (Hass and Hedges, 1992) in having higher diploid numbers (32-38 vs 24), and higher numbers of macrochromosomes (16-18 vs 12) and microchromosomes (16-20 vs 12). SIZE. The maximum snout-vent length of the 12 species for which 10 or more individuals were ex- amined ranges from 56 mm (L. dontomasi, N = 23) to 127 mm (L. flavimaculatum, N = 432). DISTRIBUTION, HABITAT, AND NATURAL HISTORY. Extant species of Lepidophyma are found from Nuevo Leon and Michoacan, Mexico, to Panama, (Figs. 11, 12), excluding most of the Yucatan peninsula (where the extinct L. arizelo- glyphum was found). The greatest species diversity occurs in the region of Oaxaca and Chiapas, Mex- ico (10 species), and in the ranges and valleys as- sociated with the northern Sierra Madre Oriental in Queretaro and San Luis Potosi, Mexico (4 spe- cies). Members of the genus occur in a broad range of vegetation types from semiarid thorn scrub to tropical wet forest, cloud forest, and pine and pine- oak woodland. The lizards are found under a va- riety of cover, including rocks, logs, tree stumps, bark, and forest litter; in rock crevices, caves, cul- verts, and walls; and within human dwellings. Both diurnal and nocturnal activity have been observed, and diet includes fruit and arthropods (Campbell, 1998; Mautz and Lopez-Forment, 1978). The 5 species for which reproductive information is avail- able are viviparous. REMARKS. Lepidophyma dontomasi, L. gai- geae, L. lowei, and L. radula form a distinct phe- netic cluster within the genus characterized by small body size, absence of microtubercles on the scale surfaces (Bezy and Peterson, 1988); absence (or small size) of tubercular scales on the body (ex- cept L. radula); poorly differentiated caudal whorls and interwhorls; and relatively large (and thus few) Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 15 dorsal, ventral, and gular scales (Bezy and Cama- rillo, 1992, 1997). Whether the 4 species comprise a clade {Gaigeia; Smith, 1939, 1973) or a grade is being addressed by phylogenetic analyses of DNA sequences combined with morphology (R.L. Bezy and collaborators, unpublished). ETYMOLOGY. The name Lepidophyma (Greek: lepidos, scale; phyma, enlarged or swollen object) is neuter (Smith, 1973) and refers to the presence of enlarged tubercular scales. COMMON NAMES. Tropical night lizards; la- gartijas nocturnas tropicales; escorpiones noctur- nos. Lepidophyma chicoasense Alvarez and Valentin Group FI; Figure 13 Lepidophyma chicoasensis Alvarez and Valentin, 1988:125, Fig. 1. Holotype: IPN 11132; type lo- cality, “16.3 Km N, 1.6 Km E de Tuxtla Gutier- rez, 600 m., Chiapas, Mexico.” DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma in having 35 or more fourth toe lamellae. It differs from all except L. smithii, L. flavimaculatum, L. micropholis, and L. sylvaticum in having 53 or more gulars; from L. gaigeae, L. dontomasi, L. lowei, L. radida, L. tar- ascae, and L. pajapanense in having 177 or more dorsals; and from L. reticulatum, L. micropholis, and L. occulor in having 192 or fewer dorsals (Ta- ble 1). KARYOTYPE. Unknown. SIZE. The maximum snout-vent length for the 4 specimens examined is 78 mm. SEX RATIO. Of the 3 specimens sexed, 2 (67%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known only from the type locality located in the Canon Sumidero (Fig. 11). REMARKS. Lepidophyma chicoasense resem- bles L. lipetzi in color pattern and in many features of scalation, including the presence of 12 longitu- dinal rows of ventrals at midbody (contra Alvarez and Valentin, 1988). However, it has a higher num- ber of fourth toe lamellae than all other members of the genus, including L. lipetzi, L. flavimacula- tum, and L. tuxtlae, which occur in the region. ETYMOLOGY. The name chicoasense is an ad- jective and refers to La Presa de Chicoasen, Chia- pas, located in the region of the type locality. The original name is emended in this paper to modify the neuter generic name. COMMON NAMES. Sumidero Tropical Night Lizard; Lagartija Nocturna del Sumidero. 16 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Lepidophyma dontomasi (Smith) Group D; Figure 14 Gaigeia dontomasi Smith, 1942:374. Flolotype: USNM 111473; type locality, “Lachiguiri, Oa- xaca, at 7,100 feet.” Lepidophyma dontomasi: Savage, 1963:35. DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. lowei and L. radula in having enlarged caudal whorls that are separated dorsally by 2 rows of inter- whorls, only one of which is complete ventrally; from all except L. gaigeae, L. radula, and L. tar- ascae in having 149 or fewer dorsals; from L. gai- geae in having 21 or fewer femoral pores; from L. tarascae in having 35 or fewer gulars (Table 1); and from L. radula in lacking vertical rows of distinctly enlarged tubercles on the side of the body. KARYOTYPE. Lepidophyma dontomasi has a diploid chromosome number of 32 with 16 macro- chromosomes and 16 microchromosomes (Bezy and Camarillo, 1992). The macrochromosomes are very similar to those of L. smithii. The species has the lowest number of total chromosomes and of microchromosomes in the genus. SIZE. The maximum snout-vent length for the 29 specimens examined is 56 mm, smaller than any other member of the genus except L. radula (known only from the holotype). SEX RATIO. Of the 22 specimens sexed, 9 (41%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known only from the type locality on Cerro Lachiguiri, Oaxaca (Fig. 11), where it has been found under logs in pine forest (Smith, 1942) and in rock crevices and under rocks and dead Agave and Nolina in oak savanna (Bezy and Camarillo, 1992). REMARKS. Additional field work is needed to determine if the species occurs elsewhere in the Si- erra de los Mijes. ETYMOLOGY. The name dontomasi is a noun in the genitive singular case and honors Thomas (Don Tomas) Baillie MacDougall (1896-1973), a naturalist who collected plants and animals (includ- ing the holotype of L. dontomasi) from remote ar- eas in Mexico, particularly the Tehuantepec region (Smith, 1974). COMMON NAMES. MacDougall’s Tropical Night Lizard; Lagartija Nocturna de MacDougall. Lepidophyma flavimaculatum complex Group F3; Figure 15 Lepidophyma flavimaculatus A. Dumeril in Du- meril and Dumeril, 1851:138. Holotype: MNHN 782; type locality, “Province du Peten (America centrale)” (restricted to Rio de la Pa- sion, Guatemala, by Smith and Taylor, 1950). Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 17 Figure 13 The holotype of Lepidophyma chicoasense (IPN 11132; snout-vent length, 78 mm). Figure 14 Living individual of Lepidophyma dontomasi (ENEPI 3016; snout-vent length, 50 mm). 18 1 Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Figure 15 Living individual of Lepidophyma flavimaculatum (ENEPI 3793; snout-vent length, 107 mm). Lepidophyma flavimaculatum-. A.H.A. Dumeril, 1852, Plate 17. Poriodogaster grayii Smith in Gray, 1863:154, Plate XXL Holotype: BMNH 1946.8.30.44; type locality, unknown (originally conjectured to be “Lower California” and subsequently thought by Peters, 1874, to be Panama). Lepidophyma flavomaculatum obscurum Barbour, 1924:10. Holotype: MCZ 17747; type locality, “Rio Chilibrillo, Panama.” Lepidophyma flavomaculatum flavomaculatum: Smith, 1942:379. Lepidophyma anomalum Taylor, 1955:554, Fig. 14. Holotype: KU 34207; type locality, “Los Di- amantes, Guapiles, Limon Province, Costa Rica.” Lepidophyma ophiophthalmum Taylor, 1955:558, Fig. 15. Holotype: KU 36250; type locality, “5 km. NNE Tilaran, Guanacaste, Costa Rica.” DIAGNOSTIC CHARACTERS. The species dif- fers from L. dontomasi, L. radula, L. tarascae, L. lineri, and L. occulor in having 25 or more femoral pores; from L. dontomasi, L. gaigeae, L. radula, and L. tarascae, in having 170 or more dorsals; from L. dontomasi, L. lowei, and L. radula in hav- ing 40 or more gulars; from L. pajapanense and L. tuxtlae in having 27 or fewer large paravertebral tubercles; from L. smithii in having 23 or more lat- eral tubercle rows; from L. micropholis in having 225 or fewer dorsals; from L. chicoasense in having 31 or fewer fourth toe lamellae; from L. lipetzi in having 180 or more dorsals (99.3% of all L. flav- imaculatum, 100% of Chiapas specimens) and a darker color pattern (Fig. 15); from L. reticulatum in lacking a boldly reticulated color pattern on the gular surface; from L. mayae in having 33 or fewer lateral tubercle rows (99.6% of all L. flavimacula- tum, 100% of Guatemala specimens) and 3 or more pretympanics (99.8% of all L. flavimacula- tum, 100% of Guatemala specimens); from L. syl- vaticum in having 4 or more pretympanics (99.3% of all L. flavimaculatum) and a lower second post- orbital supralabial (0.83 or less; 99.3% of all L. flavimaculatum: Table 1); and from L. reticulatum in lacking a bold reticulations on the gular surface. KARYOTYPE. Lepidophyma flavimaculatum has a diploid chromosome number of 38 with 18 macrochromosomes and 20 microchromosomes (Bezy, 1972). One individual from an all-female population appeared to be a diploid-triploid mo- saic. The karyotype is identical to that of L. paja- panense, L. tuxtlae, and L. reticulatum. It differs from that of L. sylvaticum in having 18 rather than 16 microchromosomes. SIZE. The maximum snout-vent length for the 432 specimens examined is 127 mm. Contributions in Science, Number 493 SEX RATIO. Of the 308 specimens sexed, 60 (19%) are males. No males were present in samples from Panama and most populations in Costa Rica. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is found primarily on the Atlantic versant from Veracruz and Oaxaca to Pan- ama, but absent from the outer Yucatan Peninsula (Fig. 12). It occurs in sympatry with L. pajapanense in Veracruz; with L. tuxtlae in Veracruz, Oaxaca, and Chiapas; with L. lipetzi in Chiapas; with L. mayae in Guatemala; and with L. reticulatum in Costa Rica. The species occurs at low and moder- ate elevations in tropical moist forest, tropical dry forest, and subtropical wet forest, where it is found in rotting logs and tree stumps; in forest litter; un- der bark; under rocks; and in rock crevices, caves, and ruins (Alvarez del Toro, 1982; Burt, 1935; Campbell, 1998; Duellman, 1963; Henderson and Hoevers, 1975; Lee, 1996; Meyer and Wilson, 1973; Scott, 1976; Slevin, 1942; Stuart, 1948; Tel- ford and Campbell, 1970; R.L. Bezy and col- leagues, personal observations). It is reported to consume termites, ants, crickets, spiders, scorpions, centipedes, and millipedes, and to be preyed upon by the colubrid snakes Dryadophis melanolomus Cope, 1868, and Drymarchon corais Boie, 1827 (Campbell, 1998; Lieberman, 1986). In Chiapas, five to eight young are born in June and July (Al- varez del Toro, 1982), and in Panama, a female gave birth in June to five neonates, all females (Tel- ford and Campbell, 1970). REMARKS. Lepidophyma flavimaculatum is un- usually variable in scalation because of geographi- cal variation among its widely scattered localities. It is most similar to L. mayae, L. lipetzi, and L. sylvaticum. In areas where it occurs with L. mayae and L. lipetzi, individual scale characters segregate all specimens, indicating reproductive isolation. Univariate scale characters separate 99% of the specimens of L. flavimaculatum from L. sylvati- cum, and the 2 species differ in karyotype for all localities examined (L. flavimaculatum from Chia- pas to Panama and L. sylvaticum from Nuevo Leon to Hidalgo). The term complex is used for the aggregation of Lepidophyma flavimaculatum populations to de- note that it may contain more than 1 species. The populations in Panama and most of Costa Rica lack males but do not appear to differ significantly in allozymes, karyotype, scalation, or color pattern from the populations to the northwest that contain males (Bezy, 1972, 1989; Bezy and Sites, 1987). ETYMOLOGY. The name flavimaculatum (Lat- in: flavus, yellow; maculatum, spotted) is an adjec- tive and refers to the dorsal color pattern. The orig- inal name was justifiably emended by A.H.A. Du- meril (1852) to modify the neuter generic name. COMMON NAMES. Yellow-spotted Tropical Night Lizard; Lepidofima; Escorpion Nocturno Puntos Amarillos; Reina de Culebra; Perrozompo- po Atlantico. Bezy and Camarillo: Systematics of Lepidophyma ■ 19 Lepidophyma gaigeae Mosauer Group G; Figure 16 Lepidophyma gaigeae Mosauer, 1936:3, Plate II. Ho- lotype: MCZ 42145 (Mosauer Mexican Collection 971); type locality, “Durango, State of Hidalgo, Mexico.” Gaigeia gaigeae: Smith, 1939:24. DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. tarascae in having enlarged caudal whorls that are separated dorsally by 2 rows of interwhorls both of which are complete ventrally; from all except L. donto- masi, L. radula, L. tarascae, L. tuxtlae, and L. syl- vaticum in having 151 or fewer dorsals; from L. dontomasi, L. radula, and L. tarascae in having 28 or more femoral pores; from L. tuxtlae in lacking paravertebral rows composed of a continuous string of subequal tubercles; and from L. sylvati- cum in lacking distinctly enlarged tubercles on the side of the body (Table 1). KARYOTYPE. Lepidophyma gaigeae has a dip- loid chromosome number of 38 with 18 macro- chromosomes and 20 microchromosomes (Bezy, 1972). The karyotype is similar to that of L. flavi- maculatum, L. tuxtlae, and L. pajapanense, but dif- fers in centromere position for 2 pairs of chromo- somes. SIZE. The maximum snout-vent length for the 177 specimens examined is 66 mm. SEX RATIO. Of the 147 specimens sexed, 55 (37%) are males. DISTRIBUTION. The species occurs in Quere- taro and Hidalgo (Fig. 11), where it is found pri- marily in limestone crevices in pine-oak woodland along the Sierra Madre Oriental (Dixon et ah, 1972; Taylor, 1939; R.L. Bezy and colleagues, per- sonal observations). It has also been found on an adobe wall in thorn scrub in the Jalpan Valley, Que- retaro (Dixon et ah, 1972), and in a cave in xero- phytic scrub near the Rio Tula, Hidalgo (Gonzalez, 1995). It occurs in sympatry with L. sylvaticum and L. occulor in Queretaro. A captive female gave birth to one young in March (Dixon et ah, 1972). The species appears to be diurnal based on meta- bolic data (Mautz, 1979) and it is more resistant to water loss than is L. smithii (Mautz, 1982). REMARKS. The populations found in the Jalpan Valley differ slightly in scalation (Bezy and Cama- rillo, 1992). ETYMOLOGY. The name gaigeae is a noun in the genitive singular case and honors Helen Thompson Gaige (1890-1976), former curator at the University of Michigan Museum of Zoology. COMMON NAMES. Gaige’s Tropical Night Lizard; Lagartija Nocturna de Gaige. Lepidophyma lineri Smith Group SI; Figure 17 Lepidophyma flavimaculatum lineri Smith, 1973: 118, Plates 1-2. Holotype: UCM 48452; type lo- 20 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Figure 16 Living individual of Lepidophyma gaigeae (ENEPI 4088; snout-vent length, 63 mm). cality, “Cafetal Padfico, 2.1 mi. N and 0.4 mi. E Candelaria Loxicha, Oaxaca, on the trail to Caf- etal Hedweges (about 17 km. N Pochutla).” DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. smithii and L. tarascae in lacking a pale parietal spot; from L. smithii in having 23 or more lateral tubercle rows; from L. tarascae in having 179 or more dor- sals and 51 or more gulars; from L. occulor in hav- ing 53 or fewer gulars and 184 or fewer dorsals; from all except L. dontomasi, L. smithii, L. taras- cae, and L. occulor in having 17 or fewer total fem- oral pores; and from all except L. radula, L. smi- thii, L. tarascae, L. flavimaculatum, L. occulor, and L. sylvaticum in having 24 or fewer lateral tubercle rows (Table 1). KARYOTYPE. Unknown. SIZE. The maximum snout-vent length for the two undamaged specimens examined is 37 mm. SEX RATIO. Both specimens sexed are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known only from the type locality and nearby Cafetal Alemania in the Sierra de Miahuatlan of Oaxaca (Eig. 12). REMARKS. Lepidophyma lineri is most similar to L. smithii. Additional material is needed to clar- ify its variation, distribution, and geographic rela- tionship with L. smithii, which occurs ca. 80 km to the west along road to Puerto Escondido, Oaxaca. ETYMOLOGY. The name lineri is a noun in the genitive singular case and honors Ernest A. Liner, one of the herpetologists who collected the holotype. COMMON NAMES. Liner’s Tropical Night Liz- ard; Lagartija Nocturna de Liner. Lepidophyma lipetzi Smith and Alvarez del Toro Group E2; Eigure 18 Lepidophyma lipetzi Smith and Alvarez del Toro, 1977:37, Figs. 2-4. Holotype: UCM 51425; type locality, “Lago de Mai Paso, headwaters of Rio de La Venta, 30 km N Cintalapa (straight line), Chiapas, Mexico.” DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. smithii, L. lineri, L. tuxtlae, L. chicoasense, L. flavimacu- latum, L. reticulatum, L. micropholis, L. occulor, and L. sylvaticum in having 46 or more gulars; from L. micropholis and L. occulor in having 52 or fewer gulars; from all except L. mayae, L. pa- japanense, L. chicoasense, L. flavimaculatum, L. reticulatum, L. micropholis, and L. sylvaticum in having 35 or more femoral pores; from all except L. gaigeae, L. smithii, L. pajapanense, L. tuxtlae, L. chicoasense, L. flavimaculatum, and L. reticu- latum in having 6 or more pretympanics; from L. chicoasense in having 28 or fewer fourth toe la- Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 21 Figure 17 The holotype of Lepidophyma lineri (UCM 48452; snout-vent length, 36 mm). mellae; from L. reticulatum in having 35 or more femoral pores, 14 or fewer divided fourth toe la- mellae, and 179 or fewer dorsals; and from L. flav- imaculatum in having 179 or fewer dorsals (99.3% of all L. flavimaculatum, 100% of Chiapas speci- mens; Table 1) and a pale dorsal color pattern (Fig. 18). KARYOTYPE. Unknown. SIZE. The maximum snout-vent length for the two specimens examined is 55 mm. SEX RATIO. Unknown DISTRIBUTION, FIABITAT, AND NATURAL FUSTORY. The species is known only from the type locality (Fig, 11), where it occurs with L. flavimacu- latum. The 2 known specimens were collected on the walls of caves (Smith and Alvarez del Toro, 1977). REMARKS. The distributional relationship in Chiapas of L. lipetzi, found on the Rio de La Venta, and L. chicoasense, known only from the Canon Sumidero of the Rio Grijalva, may never be re- solved because the vast Lago Netzahualcoyotl now lies at the confluence of these rivers. ETYMOLOGY. The name lipetzi is a noun in the genitive singular case and honors Milton L. Lipetz, a University of Colorado faculty member admired by Flobart Smith. COMMON NAMES. Lipetz’s Tropical Night Lizard; Lagartija Nocturna del Ocote. Lepidophyma lowei Bezy and Camarillo Group L; Figure 19 Lepidophyma lowei Bezy and Gamarillo, 1997:1, Figs. 1-3. Holotype: IBH 7500; type locality, “4.0 km (by rd) SE San Bartolome Zoogocho, Munici- pio Zoogocho, former Distrito Villa Alta, Oaxaca, Mexico (17°14'N, 96H5'W; ca. 2200 m elev.).” DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. donto- masi and L. radula in having enlarged caudal whorls that are separated dorsally by 2 rows of in- terwhorls, only one of which is complete ventrally; from all except L. dontomasi, L. gaigeae, L. radula, L. tuxtlae, and L. pajapanense in having 37 or few- er gulars; from L. dontomasi, L. gaigeae, and L. radula in having 158 or more dorsals; and from L. tuxtlae, L. pajapanense, L. mayae, L. chicoasense, L. lipetzi, L. flavimaculatum, L. reticulatum, and L. micropholis in having 7 or fewer divided fourth toe lamellae (Table 1). KARYOTYPE. Lepidophyma lowei has a diploid chromosome number of 36 with 16 macrochro- mosomes and 20 microchromosomes (Bezy and Camarillo, 1997). The karyotype appears virtually identical to that of L. smithii. SIZE. The maximum snout-vent length for the 16 specimens examined is 60 mm. SEX RATIO. Of the 10 specimens sexed, 6 (60%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known only from the vi- cinity of the type locality (Fig. 11), where it is found in rock crevices in pine-oak woodland. A female gave birth to four offspring in April (Camarillo, 1999). REMARKS. Additional work is needed to estab- lish the geographic distribution of the species. ETYMOLOGY. The name lowei is a noun in the genitive singular case and honors Charles H. Lowe, a herpetologist at the University of Arizona. COMMON NAMES. Lowe’s Tropical Night Lizard; Lagartija Nocturna de Lowe. Lepidophyma mayae Bezy Group Tl; Figure 20 Lepidophyma mayae Bezy, 1973:1, Figs. 1-3. Ho- lotype: KU 59554; type locality, “near Chinaja, elev. 140 m., Depto. Alta Verapaz, Guatemala.” 22 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Figure 18 Living individual of Lepidophyma lipetzi (ENEPl 3792; snout-vent length, 53 mm). Figure 19 Eiving individual of Lepidophyma lowei (IBH 7500; snout-vent length, 53 mm). Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 23 Figure 20 Living individual of Lepidophyma mayae (UTA-R 38022; snout-vent length, 78 mm). DIAGNOSTIC CHARACTERS. The species dif- fers from L. radula, L. tarascae, L. lineri, L. paja- panense, L. tuxtlae, L. chicoasense, L. lipetzi, and L. reticulatum in having 2 or fewer pretympanics; from L. dontomasi, L. gaigeae, L. radula, and L. tarascae in having 162 or more dorsals; from L. reticulatum, L. micropholis, and L. occulor in hav- ing 188 or fewer dorsals; from L, lowei, L. radula, L. smithii, L. tarascae, L. lineri, L. chicoasense, and L. occulor in having 33 or more lateral tubercle rows; from L. sylvaticum in having a wider median prefrontal (RMW2 greater than 1.6); and from L. flavimaculatum in having 33 or more lateral tuber- cle rows and two or fewer pretympanics (99.6% of all L. flavimaculatum, 100% of Guatemalan spec- imens; Table 1). KARYOTYPE. Unknown. SIZE. The maximum snout-vent length for the 16 specimens examined is 75 mm. SEX RATIO. Of the 10 specimens sexed, 4 (40%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The specimens examined are from 3 ar- eas in Guatemala: the southern El Peten near the base of the Sierra de Chinaja (border of Deptos. El Peten and Alta Verapaz), the Sierra del los Cuchu- matanes (Depto. Huehuetenango), and the valley of the Rio Cahabon near the lower slopes of the Sierra Xucaneb (Finca Volcan, Depto. Alta Verapaz; Fig. 11). They were found in lowland rain forest (Chi- naja, 140 m; Duellman, 1963) and in an area sup- porting cloud forest (Finca Chiblac, ca. 1300 m.; Wake and Elias, 1983). At Chinaja the specimens were taken within the rain forest under rocks and logs or active on the forest floor during the day (W.E. Duellman, 1960 field notes). The species is sympatric with L. flavimaculatum at Chinaja and Finca Volcan. Recently, Campbell (1998) recorded the species from additional localities in the Montanas del Mico, Sierra de Santa Cruz, and Maya Mountains north into Belize. The specimens were found in tropical and subtropical wet forest in limestone crevices and under rocks and logs. REMARKS. Although Lepidophyma mayae can be distinguished from all L, flavimaculatum only by a combination of 2 characters (lateral tubercle rows and pretympanics), where the two species are found in sympatry each of the two characters is nonover- lapping, indicating reproductive isolation. ETYMOLOGY. The name mayae is a noun in the genitive singular case and refers to the Maya, the indigenous people of the region of the type lo- cality. COMMON NAMES. Mayan Tropical Night Lizard; Escorpion Nocturno Maya. 24 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Figure 21 Living individual of Lepidophyma micropholis (LACM 106767; snout-vent length, 87 mm). Lepidophyma micropholis Walker Figure 21 Lepidophyma micropholis Walker, 1955:6. Fio- lotype: UMMZ 101298; type locality, “cave at El Pachon, about 5 miles NNE of Antigua Morelos, Tamaulipas, estimated elevation 600-700 feet.” DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. occulor in having 230 or more dorsal scales. It differs from L. occulor, L. dontomasi, L. radula, L. tarascae, and L. lineri in having 28 or more femoral pores; and from all except L. smithii, L. flavimaculatum, L. chicoasense, L. occulor, and L. sylvaticum in having 55 or more gulars (Table 1). KARYOTYPE. Lepidophyma micropholis has a diploid chromosome number of 36 with 16 macro- chromosomes and 20 microchromosomes. The kar- yotype appears identical to that of L. sylvaticum. SIZE. The maximum snout-vent length for the 31 specimens examined is 111 mm. SEX RATIO. Of the 23 specimens sexed, 12 (52%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known from four locali- ties in Sierra del Abra of Tamaulipas and San Luis Potosi (Pig. 11) where it is found in limestone caves and crevices. The vegetation near the type locality consists of tropical deciduous forest. Within the caves, the lizards have been observed active during the day near the mouth, where they are found on ledges and in crevices in the walls and mud floor (Walker, 1955; R.L. Bezy and colleagues, personal observations). REMARKS. Lepidophyma micropholis ap- pears to be a facultative troglodyte, because a number of individuals have been observed to be active outside caves (J.R. Dixon, personal com- munication). ETYMOLOGY. The name micropholis (Greek: mikros, small; pholis, scale) is a noun in apposition and presumably refers to the small size of the dorsal scales. COMMON NAMES Cave Tropical Night Eiz- ard; Lagartija Nocturna de Cueva. Lepidophyma occulor Smith Pigure 22 Lepidophyma smithii occulor Smith, 1942:378. Holotype: USNM 47133; type locality, “Jalpan, Queretaro.” Lepidophyma flavimaculatum occulor: Walker, 1955:5. Lepidophyma occulor: Bezy, 1972:15. DIAGNOSTIC CHARACTERS. The species dif- Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 25 Figure 22 Living individual of Lepidophyma occulor (TCWC 35605; snout-vent length, 90 mm). fers from all other Lepidophyma except L. smithii and L. micropholis in having 58 or more gulars; from all except L. radula, L. smithii, L. lineri, L. tarascae, L. flavimaculatum, and L. sylvaticum in having 25 or fewer lateral tubercle rows; and from L. smithii in having 22 or more lateral tubercle rows (99%; Table 1) and in having a pale parietal spot throughout life. KARYOTYPE. Lepidophyma occulor has a dip- loid chromosome number of 36 with 18 macro- chromosomes and 18 microchromosomes (Bezy, 1972). The karyotype is unique within the Xantu- siidae. SIZE. The maximum snout-vent length for the 11 specimens examined is 105 mm. SEX RATIO. Of the 11 specimens sexed, 7 (64%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species occurs in thorn scrub in the Jalpan valley of Queretaro and San Luis Po- tosi (Pig. 11), where the lizards have been found under rocks or active at night near limestone out- crops and in rock walls and ruins (Dixon et ah, 1972; R.L. Bezy and J.L. Camarillo, personal ob- servations). East of the Jalpan region, it has been taken in a cave in tropical forest along the Rio Moctezuma near Tamazunchale, San Luis Potosi (Camarillo, 1993). It occurs in sympatry with L. gaigeae. REMARKS. Lepidophyma occulor is most simi- lar to L. smithii, from which it differs in karyotype and in the presence of a parietal spot. ETYMOLOGY. The name occulor (Latin: one who hides) is a noun in apposition and refers to the secretive or reclusive behavior of the members of the genus (H.M. Smith, personal communication). COMMON NAMES. Jalpan Tropical Night Liz- ard; Lagartija Nocturna de Jalpan. Lepidophyma pajapanense Werler Group T2; Figure 23 Lepidophyma pajapanensis Werler, 1957:223, Plate 1, Fig. 1. Holotype: FMNH 78382; type locality, “southeast slopes of Volcan San Martin Pajapan, Veracruz, Mexico; elevation 3,500 feet.” Lepidophyma pajapanense: Perez-Higareda, 1978: 69. DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. tuxtlae in having the paravertebral rows composed of a nearly uninterrupted string of subequal tubercles; from L. tuxtlae, L. dontomasi, L. gaigeae, L. rad- ula, L. tarascae, L. lineri, and L. occulor in having 30 or more femoral pores; from all except L. may- ae, L. tuxtlae, and L. sylvaticum in having 39 or more large paravertebral tubercles; and from L. dontomasi, L. lowei, L. radula, L. tarascae, L. li- neri, L. mayae, L. micropholis, L. occulor, and L. sylvaticum in having 6 or more pretympanics (Ta- ble 1) KARYOTYPE. Lepidophyma pajapanense has a diploid chromosome number of 38 with 18 macro- chromosomes and 20 microchromosomes (Bezy, 1972). The karyotype appears to be identical to that of L. tuxtlae, L. flavimaculatum, and L. reti- culatum. 26 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Figure 23 Living individual of Lepidophyma pajapanense (LACM 135510; snout-vent length, 57 mm). SIZE. The maximum snout-vent length for the 22 specimens examined is 82 mm. SEX RATIO. Of the 16 specimens sexed, 5 (31%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known only from Vera- cruz and occurs in the Sierra de Los Tuxtlas (Volcan San Martin Tuxtla to Volcan San Martin Pajapan) and from the vicinity of the Rio Coatzacoalcos near Jesus Carranza on the Isthmus of Tehuantepec. It is sympatric with L. tuxtlae in the Tuxtlas region, and with L. tuxtlae and L. flavimaculatmn near the Rio Coazacoalcos (Fig. 11). The locality data for the specimen reported from Huayacacotla, Vera- cruz (Aguilar Cortes et ah, 1990), is now consid- ered questionable because extensive collecting in the area has yielded numerous L. sylvaticum, but no additional specimens of L. pajapanense. In the Tuxtlas the lizards are nocturnal and found beneath the bark and between the buttresses of trees within the rain forest from sea level to 1500 m (Vogt et ah, 1997). Both L. pajapanense and L. tuxtlae have been taken from the same tree stump in recent clearings near the forest edge in the Tux- tlas (R.L. Bezy and C.J. Cole, personal observa- tions). The male and female reproductive cycles are asynchronous and four to eight young are born in late May or early June (Mendez-de la Cruz et ah, 1999). REMARKS. This morphologically distinctive species has a rather restricted distribution, which fortunately includes the preserve at Estacion de Biologia Tropical “Los Tuxtlas.” ETYMOLOGY. The name pajapanense is an ad- jective and refers to the type locality, Volcan San Martin Pajapan. The original name was justifiably emended by Perez-Higareda (1978) to modify the neuter generic name. COMMON NAMES. Pajapan Tropical Night Lizard; Lagartija Nocturna de Pajapan. Lepidophyma radula (Smith) Group R; Figure 24 Gaigeia radula Smith, 1942:374. Holotype: USNM 111472; type locality, “San Jose Manteca, 5 ki- lometers from San Carlos Yautepec, Oaxaca.” Lepidophyma radula: Savage, 1963:35. DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. donto- masi and L. lowei in having enlarged caudal whorls that are separated dorsally by 2 rows of inter- whorls, only one of which is complete ventrally; from all except L. gaigeae and L. dontomasi in hav- ing less than 140 dorsals; from L. dontomasi in Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 27 Figure 24 The holotype of Lepidophyma radula (USNM 111472; snout-vent length, 41 mm). having distinctly enlarged tubercles on the side of the body; and from L. lowei and L. gaigeae in hav- ing fewer than 22 femoral pores (Table 1). KARYOTYPE. Unknown. SIZE. The snout-vent length of the only known specimen is 41 mm. SEX RATIO. The holotype is a female. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known only from the type locality (Fig. 11), where the vegetation consists of tropical deciduous forest. REMARKS. Lepidophyma radula remains known only from the holotype. Considerable field work conducted by the authors in the region of the type locality has failed to produce additional spec- imens. Lepidophyma smithii occurs nearby (Porti- llo Nejapa, Oaxaca) and differs extensively in sca- lation. Lepidophyma dontomasi, L. lowei, and L. rad- ula are restricted to eastern Oaxaca and are unique in the genus in having most caudal whorls separat- ed by only 2 interwhorls, one of which is complete ventrally. ETYMOLOGY. The name radula (Latin: scrap- er) is a noun in apposition and refers to the body surface, which is rough, or filelike because of the presence of large tubercular scales (H.M. Smith, personal communication). COMMON NAMES. Yautepec Tropical Night Lizard; Lagartija Nocturna de Yautepec. Lepidophyma reticulatum Taylor Group F3; Figures 25, 26 Lepidophyma reticulatum Taylor, 1955:551, Fig. 14. Holotype: KU 36245; type locality, “Agua Buena, Puntarenas Province, Costa Rica.” Lepidophyma flavimaculatum reticulatum: Wer- muth, 1965:196. DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. smithii, L. flavimaculatum, L. lipetzi, L. micropholis, L. oc- culor, and L. sylvaticum in having 193 or more dor- sals; from L. micropholis in having 229 or fewer dorsals; from L. dontomasi, L. radula, L. tarascae, L. lineri, and L. occulor in having 27 or more fem- oral pores; from L. lipetzi in having 33 or fewer femoral pores; from L. smithii, L. tarascae, L. li- neri, and L. occulor in having 28 or more lateral tubercle rows; from L. sylvaticum in having a lower second postorbital supralabial (RSLH 0.71 or less; Table 1); and from L. flavimaculatum in having a boldly reticulated color pattern on the gular surface (Fig. 26). KARYOTYPE. Lepidophyma reticulatum has a diploid chromosome number of 38 with 18 macro- chromosomes and 20 microchromosomes (Bezy, 1972, as L. flavimaculatum). The karyotype ap- pears to be virtually identical to that of L. paja- panense, L. tuxtlae, and L. flavimaculatum. SIZE. The maximum snout-vent length for the 65 specimens examined is 103 mm. SEX RATIO. No males are present among 29 specimens sexed. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. Lepidophyma reticulatum ranges on the Pacific slope of Costa Rica from the area of Tilaran (Guanacaste Province) to the Peninsula de Osa and San Vito de Java (Puntarenas Province; Fig. 12). In the Tilaran area it occurs in sympatry with a pop- ulation of L. flavimaculatum that includes males. The species has been collected within and under decaying logs and beneath rocks (R.L. Bezy and K. Bolles, personal observations). REMARKS. This unisexual species may range into southwestern Panama. ETYMOLOGY. The name reticulatum (Latin: netlike or reticulated) is an adjective and refers to the gular color pattern. COMMON NAMES. Costa Rican Tropical Night Lizard; Lagartija Nocturna de Costa Rica. Lepidophyma smithii Bocourt Groups S2, S3; Figure 27 Lepidophyma smithii Bocourt, 1876:402. Syntypes: MNHN 76.95, 4323, A4323, B4323, 4968, A4968; type locality, “Tehuantepec” and “Gua- temala occidental” (restricted to Guatemala by Smith, 1942, and to Mazatenango by Smith and Taylor, 1950). Akleistops guatemalensis Muller, 1877:390, Tafel I, 28 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Figure 25 Living individual of Lepidophyma reticulatiim (LACM; snout-vent length, 90 mm). II. Syntypes: NMBA 3748-50, 3752, 8039-40; type locality, “Mazatenango (costa grande von Guatemala).” Lepidophyma smithii tehuanae Smith, 1942:377. Holotype: USNM 111488; type locality, “Cerro Figure 26 Gular pattern of a living individual of Lepido- phyma reticulatiim (LACM; snout-vent length, 90 mm). Arenal, 30 kilometers west of Tehuantepec, Oa- xaca.” Lepidophyma smithii smithii: Smith, 1942:380. Lepidophyma flavimaculatum smithii: Walker, 1955:5. DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. tarascae and L. lineri in lacking a pale spot on the parietal scale in adults (snout-vent length greater than 75 mm); from all except L. tarascae, L. occulor, and L. sylvaticum in having 22 or fewer lateral tubercle rows; and from L. tarascae in having 162 or more dorsals and 44 or more gulars (Table 1). KARYOTYPE. Lepidophyma smithii has a dip- loid chromosome number of 36 with 16 macro- chromosomes and 20 microchromosomes (Bezy, 1972). The macrochromosomes are similar to those of L. dontomasi (Bezy and Camarillo, 1992), but the 2 species differ in number of microchromo- somes (20 vs 16). SIZE. The maximum snout-vent length for the 291 specimens examined is 112 mm. SEX RATIO. Of the 234 specimens sexed, 91 (39%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species ranges along the Pacific ver- sant from Guerrero to El Salvador (Fig. 12). One specimen (TCWC 26716) represents a strange geo- Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 29 Figure 27 Living individual of Lepidophyma smithii (LACM 135511; snout-vent length, 92 mm). graphical anomaly. The specimen bears a field tag with the locality “2 mi Sontecomapan, Vera- cruz.” The species is otherwise unrecorded from the Atlantic versant and no other specimens are present among the many individuals of L. tuxtlae and L. pajapanense from the Los Tuxtlas region. The spec- imen is clearly assignable to L. smithii (based on femoral pores, lateral tubercle rows, and large par- avertebrals; Table 2). It has a parietal spot, a fea- ture that is absent in adult L. smithii (Table 2), but the snout-vent length (41 mm) of the specimen is within the range of the few juvenile L. smithii in which the spot is visible (28-75 mm). The locality is here considered questionable for L. smithii, but the possibility that the species occurs in the Tuxtlas cannot be completely ruled out in view of the many biogeographic enigmas presented by xantusiids. The species occurs in a variety of habitats. At the northernmost known locality, near Puerto Marques, Guerrero, it occurs in a system of caves formed by piles of large granitic boulders in tropi- cal semideciduous forest. Within the caves, the liz- ards have been found to be active day and night in both the dry and wet seasons, to evidence territo- riality, and to consume the fruit of native figs, Ficus petiolaris (Mautz, 1976, 1982; Mautz and Lopez- Forment, 1978). In Oaxaca, specimens have been found in logs and rock crevices on Cerro Quiengola (Hartweg and Oliver, 1940), and in caves and road culverts on the Isthmus of Tehuantepec (R.L. Bezy and colleagues, personal observations); in Chiapas, in logs and tree stumps in savanna (R.L. Bezy and colleagues, personal observations); and in Guate- mala, on the rain forest floor and in rotting logs where the stomachs of specimens contained ter- mites (Slevin, 1942, as L. flavomaculatum). REMARKS. Walker (1955) concluded that L. smithii is conspecific with L. flavimaculatum and this arrangement has been followed by several sub- sequent workers (e.g., Smith, 1973). The ranges of the two forms are separated by only ca. 20 km on the Isthmus of Tehuantepec, with L. smithii ex- tending north to Santo Domingo de Guzman ( = Santo Domingo Petapa; Goodwin, 1969) and L. flavimaculatum south to El Mogone. In this region, the two species differ in number of femoral pores, lateral tubercle rows, and the presence of a pale interparietal spot. Although geographical variation among the localities of these 2 widely distributed species produces overlap in number of femoral pores, the two species consistently differ in two oth- er scale features (Table 1) and in chromosome num- ber (Bezy, 1972). The population of L. smithii found near Puerto Marques, Guerrero, is the most divergent in scala- tion and its systematic status warrants further study. ETYMOLOGY. The name smithii is a noun in the genitive singular case and honors Sir Andrew Smith, M.D. (1797-1872; founder of the South Af- rican Museum), whose description and figure of Poriodogaster grayi (a junior synonym of L. flavi- 30 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma Figure 28 Living individual of Lepidophyma sylvaticum (LACM 106744; snout-vent length, 75 mm). maculatmn) was used by Bocourt (1876) to com- pare with L. smithii. COMMON NAMES. Smith’s Tropical Night Lizard; Lagartija Nocturna de Smith. Lepidophyma sylvaticum Taylor Figure 28 Lepidophyma sylvatica Taylor, 1939:1, Figs. 1, 2. Holotype: FMNH 100102 (EHT-HMS 16259); type locality, “7 mi. north of Zacualtipan, Hi- dalgo.” Gaigeia sylvatica: Smith, 1942:380. Lepidophyma sylvaticum: Walker, 1955:9. Lepidophyma flavimaculatum tenebrarum Walker, 1955:1. Holotype: UMMZ 101374; type locality, “± 5 miles NW (by road) of Gomez Farias, in the Sierra Madre Oriental at ‘Rancho del Cielo,’ + 3600 feet.” DIAGNOSTIC CHARACTERS. The species dif- fers from L. dontomasi, L. radula, L. tarascae, and L. lineri in having 22 or more femoral pores; from L. gaigeae in having distinctly enlarged lateral tu- bercles forming 15 to 38 rows; from L. tuxtlae and L. pajapanense in having the paravertebral rows composed of tubercles that are heterogeneous in size; from L. lowei in having 40 or more gulars; from L. occulor in having 56 or fewer gulars; from L. smithii in having a pale parietal spot throughout life; from L. lipetzi and L. chicoasense in having four or fewer pretympanics; from L. micropholis in having 217 or fewer dorsals; from L. reticulatum in having a higher second postorbital supralabial (RSLH of 0.84 or greater); and from L. flavima- culatum in having 3 or fewer pretympanics (99.3%) and a higher second supralabial (RSLH of 0.84 or greater; 99.3%; Table 1). KARYOTYPE. Lepidophyma sylvaticum has a diploid chromosome number of 36 with 16 macro- chromosomes and 20 microchromosomes. A micro- chromosomal heteromorphism was found in some individuals from Sierra de Cucharas (Bezy, 1984). SIZE. The maximum snout-vent length for the 110 specimens examined is 113 mm. SEX RATIO. Of the 86 specimens sexed, 24 (28%) are males. Males appear to be rare in the populations found in the Sierra de Cucharas, Ta- maulipas (3/29; 10%), and at El Lobo, Queretaro (2/13; 15%). DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is found along the Sierra Madre Oriental from Nuevo Leon to Veracruz (Fig. 12). Outlying populations occur in canyons below the Cumbre de Monterey (Nuevo Leon), in the Si- erra de Tamaulipas (Tamaulipas), the Sierra Alvar- ez (San Luis Potosi), and near Tlapacoyan and Al- Contributions in Science, Number 493 varado, Veracruz, It has recently been reported from Puebla, near Tepango de Rodriguez (Cama- rillo, 1995), and in the Sierra Norte (Canseco-Mar- quez et ah, 2000). The species occurs primarily in pine-oak woodland, humid pine forest, and cloud forest where it is found under bark, logs, and lime- stone rocks; and in rock crevices, walls, and build- ings (Canseco-Marquez et ah, 2000; Dixon et ah, 1972; Martin, 1958; Walker, 1955; R.L. Bezy and J.L. Camarillo, personal observations). It is sym- patric with L. gaigeae in Queretaro, REMARKS. The occurrence of L. sylvaticum at Alvarado on the coast of Veracruz is somewhat enigmatic and brings the range of the species to within ca. 85 km of that of L, flavimaculatum. The two species appear to maintain their morphological differences in this area, with the Alvarado specimen (MZFC 1722) differing from the northernmost L. flavimaculatum (ENEPI 2104 from the Tuxtlas re- gion) in pretympanics (2 vs 4) and the height of the second postorbital supralabial (RSLH 0.98 vs. 0.58h Within L. sylvaticum, the morphologically most distinctive group of populations is composed of the samples from widely disjunct localities in the Sierra Alvarez of San Luis Potosi and in the northern base of the Sierra Madre Oriental of Nuevo Leon. ETYMOLOGY. The name sylvaticum (Latin: of woods) is an adjective and presumably refers to the occurrence of the species in humid pine and fir for- est. COMMON NAMES. Madrean Tropical Night Lizard; Lagartija Nocturna de Montana. Lepidophyma tarascae Bezy, Webb, and Alvarez Group SI; Figure 29 Lepidophyma tarascae Bezy, Webb, and Alvarez, 1982:1, Figs. 1-3. Holotype: IPN 9221; type lo- cality, “near Mexiquillo, Aquila District, Mi- choacan, Mexico.” DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. smithii and L. lineri in lacking a pale parietal spot in adults; from L. smithii, L. lineri, and L. occulor in having 159 or fewer dorsals and 43 or fewer gulars; from all except L. dontomasi, L. smithii, L. lineri, L. tuxtlae, and L. occulor in having 18 or fewer total femoral pores; and from all except L. radula, L. smithii, L. lineri, L. flavimaculatum, L. occulor, and L. sylvaticum in having 25 or fewer lateral tu- bercle rows (Table 1). KARYOTYPE. Unknown. SIZE. The maximum snout-vent length for the 4 specimens examined is 93 mm. SEX RATIO. Of the 4 specimens sexed, 3 (75%) are males. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known only from two localities in Michoacan (Fig. 12): the type locality on the Pacific coast and the Canon del Rio Chilitos Bezy and Camarillo: Systematics of Lepidophyma ■ 3 1 in the northern sector of the Sierra Coalcoman (Guzman- Villa et ah, 1998). The type series was reported to have been collected in a rocky ravine in tropical semideciduous forest (Bezy et ah, 1982), but Alvarez and Diaz-Pardo (1983) indicated that the specimens were found in encinar (evergreen oak woodland). The specimen from the Rio Chilitos was taken from a cave in a transition zone between tropical deciduous forest and oak woodland (Guz- man-Villa et ah, 1998). REMARKS. Lepidophyma tarascae is most sim- ilar to L. smithii, and additional material may bridge the differences between the two in dorsals and gulars. All known specimens of L. tarascae have an uninterrupted row of enlarged tubercles ex- tending from the posteroventral margin of the ear opening to above the gular fold (fig. 3 in Bezy et ah, 1982). However, in the Coalcoman specimen (MZFC 8603) the row is not as distinct as in the type series (i.e., it is less clearly separated from the scattered large tubercles of the upper nuchal area) and approaches the condition seen in some L, smi- thii. ETYMOLOGY. The name tarascae is a noun in the genitive singular case and refers to the Tarasca, the indigenous people of the region of the type lo- cality (Michoacan). COMMON NAMES. Tarascan Tropical Night Lizard; Lagartija Nocturna de Tarasca. Lepidophyma tuxtlae Werler and Shannon Group T3; Figure 30 Lepidophyma tuxtlae Werler and Shannon, 1957: 119, Plate I. Holotype: UIMNH 67064 (FAS 10550); type locality, “lower slopes of Volcan San Martin, Veracruz, elevation 2,500 feet.” Lepidophyma flavimaculatum tuxtlae: Savage, 1963:35. Lepidophyma sawini Smith, 1973:112, Plates 1-2. Holotype: UCM 49280; type locality, “Vista Hermosa, Comaltepec, Ixtlan, Oaxaca, Mexico.” Lepidophyma alvarezi Smith, 1973:115, Plates 1- 2. Holotype: UCM 49281; type locality, “43 km on the road between Ocozocoautla and Mai Paso, 620 m Chiapas.” DIAGNOSTIC CHARACTERS. The species dif- fers from all other Lepidophyma except L. paja- panense in having the paravertebral rows com- posed of a nearly uninterrupted string of subequal tubercles; from L. pajapanense, L. lipetzi, and L. chicoasense in having 29 or fewer femoral pores; from all except L. mayae, L. pajapanense, L. lip- etzi, and L. sylvaticum in having 30 or more large paravertebrals; and from L. radula, L. smithii, L. tarascae, L. lineri, and L. occulor in having 30 or more lateral tubercle rows (Table 1). KARYOTYPE. Lepidophyma tuxtlae has a dip- loid chromosome number of 38 with 18 macro- chromosomes and 20 microchromosomes (Bezy, 1972). The karyotype appears to be identical to that of L. pajapanense, L. flavimaculatum, and L. reticulatum. 32 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma 'Ad Figure 29 The holotype of Lepidophyma tarascae (IPN 9221; snout-vent length, 93 mm). SIZE. The maximum snout-vent length for the 97 specimens examined is 97 mm. SEX RATIO. Of the 82 specimens sexed, 34 (41%) are males. Castillo-Ceron and Lopez-Gon- zalez (1990) found only one male (9%) in the 11 specimens collected at their study site on Volcan Santa Marta, suggesting that the species may have a highly skewed sex ratio at certain localities. DISTRIBUTION, HABITAT, AND NATURAL HISTORY. The species is known from the Sierra de los Tuxtlas (Zapoapan to Volcan San Martin Tux- tla), Veracruz; the Sierra de Juarez, Oaxaca; the Rio Coatzacoalcos and Sierra Atravesada on Isthmus of Tehuantepec, Veracruz and Oaxaca; and the Selva del Ocote, Chiapas (Fig. 11). It is sympatric with L. flavimaculatum in the Rio Coatzacoalcos and the Ocote areas and with L. pajapanense in the Tuxtlas and Rio Coatzacoalcos regions. Vogt et al. (1997) indicated that in the Tuxtlas the species is nocturnal and inhabits trees within the rain forest from sea level to 1500 m. Individuals have been found within rotting logs and beneath the bark of tree stumps within or near the rain for- est (R.L. Bezy and C.J. Cole, personal observa- tions). Greene (1970) reported that a female col- lected near Lago Catemaco in June contained 6 Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma ■ 33 Figure 30 Living individuals of Lepidophyma tuxtlae from the Tuxtlas region of Veracruz (upper; LACM 136358; snout- vent length, 62 mm) and the Vista Hermosa region of Oaxaca (lower; UMMZ 125870; snout-vent length, 97 mm; photo by Kraig Adler). 34 ■ Contributions in Science, Number 493 Bezy and Camarillo: Systematics of Lepidophyma nearly full-term embryos. At their study site located at 900 m on Volcan Santa Marta, Castillo-Ceron and Lopez-Gonzalez (1990) found that females gave birth to three to five young in late June and July; that activity occurred in the day; that stomach contents consisted of insects, amphipods, gastro- pods, and plant material; and that only 1 of 11 specimens preserved was male. Bursey et al. (1998) found that specimens harbored 2 species of nema- tode parasites, one of which has not been recorded from other hosts. REMARKS. Lepidophyma tuxtlae is most simi- lar in scalation to L. pajapanense, with which it is sympatric, but the two differ markedly in color and color pattern (Figs. 23, 30). The widely scattered populations of L. tuxtlae have differences in the means of several features of scalation, but there is broad overlap in the observed limits of variation and they are highly similar in color pattern (Fig. 30). The ecological and biogeographic relationship between L. tuxtlae and L. pajapanense in the Tux- tlas merits additional study. Castillo-Ceron and Lo- pez-Gonzalez (1990) noted that L. pajapanense is the more abundant at the Estacion de Biologia Tropical “Los Tuxtlas,” whereas L. tuxtlae predom- inates at their study site on Volcan Santa Marta. ETYMOLOGY. The name tuxtlae is a noun in the genitive singular and refers to the region of type locality and its indigenous people, the Tuxtla. COMMON NAMES. Tuxtla Tropical Night Liz- ard; Lagartija Nocturna de Los Tuxtlas. KEY TO THE EXTANT SPECIES OE THE GENUS LEPIDOPHYMA la. Scales on at least anterior half of side of body not conspicuously heterogeneous in size, lack- ing conspicuously enlarged pyramidal tuber- cles 2 b. Scales on most of side of body conspicuously heterogeneous in size, with small granular scales interspersed with conspicuously en- larged pyramidal tubercles often arranged in vertical rows 3 2a. Total femoral pores (right + left and includ- ing vestigial indentations) 28 or more L. gaigeae b. Total femoral pores 21 or less L. dontomasi 3a. Most of the enlarged scale rings on tail sep- arated ventrally by only 1 complete ring of smaller scales 4 b. Most of the enlarged scale rings on tail sep- arated ventrally by 2 complete rings of small- er scales 5 4a. Dorsal scales (occiput to rump) 158 or more L. lowei b. Dorsal scales fewer than 158 ... . L. radula 5a. Lamellae under fourth toe 32 or more L. chicoasense b. Lamellae under fourth toe 31 or less .... 6 6a. Paravertebral scale row relatively homoge- neous, composed of tubercular scales that are similar in size and nearly continuous .... 7 b. Paravertebral row heterogeneous, composed of tubercular scales of various sizes inter- spersed with small granular scales 8 7a. Total femoral pores 29 or less; dorsal color- ation dark brown to black with small pale spots (Fig. 30) L. tuxtlae b. Total femoral pores 30 or more; dorsal col- oration pale brown with dark rectangular blotches (Fig. 23) L. pajapanense 8a. Pale parietal spot absent on the interparietal scale (occasionally present in juveniles with a snout-vent length less than 75 mm) .... 9 b. Pale parietal spot present on posterior third of interparietal scale 11 9a. Dorsal scales (occiput to rump) 159 or less L. tarascae b. Dorsal scales 160 or more 10 10a. Vertical rows of enlarged tubercles on side of body 23 or more L. lineri b. Vertical rows of enlarged tubercles on side of body 22 or less L. smithii 11a. Gulars usually 57 or more; if gulars are 55 to 56, then dorsals (occiput to rump) are 230 or more 12 b. Gulars usually 54 or fewer; if gulars are 55 to 56, then dorsals are 229 or fewer ... 13 12a. Total femoral pores 28 or more L. micropholis b. Total femoral pores 22 or less . . L. occulor 13a. Second postorbital supralabial (which usually is the seventh supralabial) lower than the first postorbital supralabial (which usually is the sixth supralabial) and separated from the pos- toculars by a minimum of 2 or more pretym- panics on each side of head 14 b. Second postorbital supralabial usually higher than or equal in height to first postorbital su- pralabial (if it is lower than first postorbital supralabial it is separated from the postocu- lars by less than 2 pretympanics on at least one side of head) 16 14a. Dorsal scales (occiput to rump) 179 or fewer; more than 10 longitudinal rows of ventrals on posterior half of body L. lipetzi b. Dorsal scales usually 180 or more; if fewer than 180, 10 or fewer longitudinal rows of ventrals on posterior half of body 15 15a. Gular surface boldly reticulated with a dark brown network enclosing rounded cream spots (Fig. 26) L. reticulatum b. Gular surface lacking bold reticulation, light with or without dark spots L. flavimaculatum complex 16a. Median prefrontal shorter than frontal or ab- sent; most of the enlarged caudal rings sepa- rated dorsally by 3 smaller scale rows L. sylvaticum b. Median prefrontal equal to or longer than frontal; most of the enlarged caudal rings sep- Contributions in Science, Number 493 arated dorsally by 4 smaller scale rows .... L. mayae SPECIMENS EXAMINED The 1,420 specimens of Lepidophyma examined are listed below. The numbers (in parentheses) pre- ceding the localities denote separate locality sam- ples used in the analyses of variation. The speci- mens are listed generally northwest to southeast, with the states of Mexico (Nuevo Leon, Tamauli- pas, San Luis Potosi, Queretaro, Hidalgo, Guerre- ro, Michoacan, Veracruz, Oaxaca, Tobasco, Chia- pas, and Quintana Roo) followed by the countries of Central America. Museum abbreviations follow Leviton et al. (1985) with the addition of: BCB (B.C. Brown collection at the Strecker Museum), CRE (J.M. Savage Costa Rica collection, now at LACM); EAL (E.A. Liner collection, now at AMNH), JEC (J.E. Copp collection, now at CAS); ENEPI (Escuela Nacional Estudios Profesionales, Iztacala, UNAM), MZEC (Museo de Zoologia, Ea- cultad de Ciencias, UNAM), OMNH (Oklahoma Museum of Natural History), and SAM (Sherman A. Minton Collection, now at AMNH). L. chicoasense CHIAPAS: (1): 16.3 km N, 1.6 km E Tuxtla Gu- tierrez (IPN 11130-33). L. dontomasi OAXACA: (1): D. Tehuantepec: Lachiguiri: Cer- ro de Lachiguiri (ENEPI 3010-17, UCM 39887, 41147, 44446-62); Lachiguiri (USNM 111473- 74). L. flavimaculatum UNKNOWN: (0): Locality unknown (BMNH 1946.8.30.44). VERACRUZ: (1): 35 km SW Jesus Carranza (KU 26919); 25 km SE Jesus Carranza (KU 26920-21); 20 km E Jesus Carranza (KU 24453); (2): Rio de las Playas (USNM 118638); (3): Catemaco: Colonia Adalberto Tejada (ENEPI 2104). OAXACA: (1): Rio Mono Blanco, Juchitan (UIMNH 36832); (2): La Gloria (UIMNH 35515); (3): Mogohe (UIMNH 40811); (4): 50.5 mi S Aca- yucan, Hwy 185 (TNHC 25182); Palomares, Juch- itan (UCM 39888); 2.8 mi N Rio Sarabia (UMMZ 115096); 2 mi S Tolosita (KU 39676). TOBASCO: (1): Teapa (BMNH 1913.7.19.95, LACM 61260- 61, LSUMZ 6878-79, UIMNH 47883, UMMZ 113777); (2): Soledad (UIMNH 47884); (3): Mpo. Tacotalpa: Cueva o Gruta Costachica (IBH 3676). CHIAPAS: (1): Ruinas de Palenque (EAL 3030-31, UE 32915-16, KU 94104-05); Palenque (LACM 65117-19); San Juanito, Palenque (USNM 111486-87); 6.5 km S Palenque (MZEC 4186); (2): 4.5 km S Pichucalco (KU 94106); (3): El Estoracan (ca 50 km N Cintalapa) (AMNH 73468); (4): Chia- pa, 1 mi W (TNHC 27517-18); (5): 32 km NW Ocozocoautla, Selva del Ocote (JEC); 16.1 mi NW Bezy and Camarillo: Systematics of Lepidophyma ■ 35 Ocozocoautla (LACM 61259); 25 mi (by rd to Malpaso) NW Ocozocoautla (UAZ 28764, 28805- 07); Ocozocoautla Selva El Ocote (MCZ 54321- 22); 12 km N Berriozabal (MZEC 4134, UTEP 5365-66); 26 km N Ocozocoautla (UTEP 5367- 68); (6): ca. 5 km S Solusuchiapa (UAZ 31635); (7): Lago Miramar, near San Quintin (JEC); (8): La- canja (LACM 114244); (9): Rancho Alejandria Juarez (UCM 49275-79); (11): Salto del Agua: Cascada Misol-ja (IBH 3461); (12): Lago del Mal- paso: Boca de Rio de La Venta (ENEPI 3793). QUINTANA ROO: (1): 4.1 km NE Eelipe Carrillo Puerto (UMRC 79-252). BELIZE: (1): Middlesex, near “Blue Hole” of Staum Creek (EMNH 4455); (2): State Eorest, 4 m. from 12 m. Sta. on Staum Cr. R.R. (EMNH 4453-54); (3): Bokowma (BMNH 1973.2537-42, EMNH 49271-75); (4): Double Ealls (EMNH 49276); (5): Baking Pot (MCZ 18953); (6): Toledo Dist.: ca. 1/2 mi from Village of Blue Creek (CM 117260; (7): Cayo: Xunantunich Ruins (KU 171679-80). GUATE- MALA: (0): Province du Peten (Amer. centrale) (MNHN 782); (1): El Peten: 15 km NW Chinaja (KU 55864, 59552); Alta Verapaz: Chinaja (KU 59555); Rio de Pasion, nr. Peten-Alta Verapaz bor- der (UMMZ 79072); (2): Izabal: Quirigua (CAS 68479-82, USNM 58384-85); (3): Peten: Piedras Negras (USNM 111475-83, UIMNH 48875); De- sempeho (USNM 111484); (4): Alta Verapaz: Finca Volcan (UMMZ 91145-47); Finca Volcan, elev. 875 m, 25 km NW (by rd) Senahu (MVZ 159932- 33). HONDURAS: (1): Dept. Cortes: Finca Fe, N Lago de Yojoa (LACM 45260); El Copito, 1 mi SE El Jaral (LACM 47831-33); 1 mi W El Jaral (LSUMZ 24217); Amapa Research Sta. (AMNH 70459); (2): Atlantida: mts above Corozal, 15 km E La Ceiba (LACM 47825-30, LSUMZ 21514- 21); Piedra Pintada, 6.5 mi E La Ceiba (LACM 47834-35); (3): Olancho: E.N.A., 4.5 km SE Ca- tacamas (LACM 47823-24); (4): nr. Chamelecon (MCZ 29886-87); (5): Fortuna D.: Puerto Arturo (MCZ 21741); Tela: Port Town (MCZ 21814); Tela (MCZ 29394-95, USNM 94417-22, 101278; (6): Los Indios, Tela R.R. (EMNH 21788); Los In- dios Farm, 76 km W Tela R.R. (MCZ 38903-22); Rio Ulua (UMMZ 77840); about 50 mi from Tela (EMNH 13002-03); (7): San Pedro, Montana San- tana (MCZ 29396); (8): Subirana Valley (MCZ 38901-02). NICARAGUA: (0): Nicaragua (MCZ 6250); (1): Rio San Juan (USNM 19543); San Juan del Norte (USNM 15644); nr. Greytown (USNM 19739); (2): 10 mi above Recreo, Rio Mico (UMMZ 79846); (3): Chontales, Tule Creek (AMNH 16402); (4): Departamento Rio San Juan, appx. 15 km S El Castillo on north bank of the Rio San Juan at Isla el Diamante (OMNH 38244-47). COSTA RICA: (0): Costa Rica (CAS 54645); (1): Limon Prov.: ca. 50 mi NW Limon, ca. 1.75 mi S mouth Rio Tortuguero (AMNH 89257-8); SW face of Cerro Tortuguero (UF TC30, KC923); (2): He- redia Prov.: Finca La Selva, jnct, of Rio Viejo and Rio Sarapiqui (CRE 483, 6300); E of Puerto Viejo 36 B Contributions in Science, Number 493 on La Selva field station, river road trail (UMMZ 145862); Puerto Viejo Sarapiqui (CRE 2890, KU 67382, 104079); 2 km S Puerto Viejo (CRE 2891); 1.5 km N Puerto Viejo (KU 67383-5); 9.7 km (by rd) W Puerto Viejo (LACM 128534-54); 13 km (by rd to Guapiles) SSE Puerto Viejo (LACM 128555- 59); 10 km WSW Puerto Viejo Sarapiqui (MVZ 217606); (5): Lower Reventazon River, La Castilla (ANSP 23682); (6): Limon Prov.: La Lola, Experi- mental Cacao Einca of I.I.C.A. (CRE 500); Batan (KU 34210); La Lola, ca. 4 mi W Batan (KU 34211-12, 34214-16); (7): Cartago Prov.: Turrial- ba: S. Bank of Reventazon (CRE 504); Morehouse Einca, 5 mi S Turrialba (KU 34208); (8): Guana- caste Prov.: El Silencio (CRE 3057); San Bosco (CRE 6282, EMNH 176530); 4-5 km NNE Tilar- an (KU 36250); Lago de Arenal (KU 102446-48); 2.5 km W Tilaran (CRE 734); Tilaran (UE 33137); 2 km E Arenal along rd to Tilaran (EACM 137499, UMMZ 137543); (9): Alajuela Prov.: 11 mi N Elo- rencia (CRE 2624); 5 mi N of Quesada (CRE 7156); (14): Limon Prov.: 1 1/2 mi E of Los Dia- mantes (CRE 8049); Eos Diamantes, Guapiles (KU 34205-7, 34213); (16): Guanacaste: Tenorio, Los Elores, Las Canas (KU 34217-19); (17): Limon Prov.: Suretka, (KU 36251-55); (18): Limon Prov.: Puerto Viejo (KU 36256-57); (19): Alajuela Prov.: Poco Sol, San Carlos drainage (LACM 114858); (21): Limon Prov.: La Colonia, 23 km NE Guapiles (MVZ 82876); (22): Limon Prov.: ca. 29 km (air) W Tortuguero (LACM 131067-111, 132392); (23): La Emilia (ANSP 21403); (24): Puerto Viejo (LACM 115744-45). PANAMA: (1): Canal Zone: 3 mi SE Achiote (Colon Prov.) (UP 32919-22, 32924-48, UAZ 27637-44, 28610-18, 28826-27, 31777); 8 mi (air line) NNW Escobal (LACM 128560-87, 131149); (2): Canal Zone: Barro Col- orado Id. (AMNH 75985, ANSP 21089-90, 23024, 24579-80, CAS 93853, CM 6638, EMNH 13374, KU 80524, MCZ 18861, UMMZ 61611); (3): Panama: Chilibnllo Cave “c” (AMNH 71688- 89, EMNH 60161); Chilibrillo Cave, near Chilibre (KU 113496-97); Canal Zone: Chilibre Bat Caves near Chagres River on Madden Dam Rd (SDSNH 5697); Chilibrillo River (MCZ 17747); (4): Canal Zone: vie. George Green Park, 3 mi (air line) NE Summit (UAZ 28607-09); (5): Colon: Santa Rita Lumber Rd, nr. Agua Clara (MCZ 133640); (6): Panama Prov.: N base Cerro Trinidad, El Aguacate (UP 32923); (7): Ciricito (CAS 71470); (8): Cabi- ma. Pacific slope (USNM 48503); (9): Bocas de Toro: ca. 5 km W Almirante (AMNH 107589); 4.8 km W Almirante (KU 96924); 12.8 km W Almi- rante (KU 96925); 12 mi W of Almirante (USNM 65809); Almirante (USNM 193444). L. gaigeae QUERETARO: (1): 4 mi SW El Lobo (BCB 68- 720, 774-779); 2.8 mi W El Madrono, 11.9 mi W Xilitla (CU 9690-91); 0.6 mi N of Mex 120 on dirt rd to Tres Eagunas, 0.8 mi W El Madrono (CU Bezy and Camarillo: Systematics of Lepidophyma 9684-86, 9770-71); 4.1 mi SW El Eobo (UP 28331-52); 2.2 mi (by Hwy 120) SW El Madrono (LACM 106770); 3.5 mi (by rd) W El Lobo (LACM 106807-09, 127169-70); N of Mex Hwy 120, nr El Eobo (MVZ 111459); 2.2 mi SW El Lobo (SAM 1105); 2.2 mi W Madronos (TCWC 27790-829); 3.2 mi W El Lobo (TCWC 29619, 29650-82, 30664-74, 35665-66); 2.4 mi W El Madrono (TCWC 29620-32, 29634-40, 29642, 29644-47, 29649, 29683-90); 4.9 mi W El Ma- drono (TCWC 29633); 2.4 mi W, 1 mi S El Ma- drono (TCWC 29648, 29643); 3.2 mi W, 2.3 mi S El Lobo (TCWC 30675); 3.2 mi W, 1 mi S El Lobo (TCWC 30676); 3.8 mi W El Lobo (TCWC 32292-93, 32296-363, 33052-62); 0.9 mi E El Lobo (TCWC 32294); 3.8 mi W, 1.8 mi S El Lobo (TCWC 38608-10); El Lobo (TCWC 45511-18, 45521-24, 45633-34); (2): 3 mi E Jalpan (TCWC 35608-10); 6 mi E Jalpan (TCWC 35611-64); 2 mi NE Jalpan (UMMZ 129746); (3): 10.2 mi W San Joaquin (LACM 109153, TCWC 40686-87, 40975); vie Lizaro Vega, 7.5 mi (by rd) N jet. Mex 120 on San Joaquin Rd (TCWC 53024); (4): 8.5 mi S Arroyo Seco (TCWC 32364); 3.7 mi S Arroyo Seco (TCWC 37677-79); 1 mi S Arroyo Seco (TCWC 38611); (5): 5.7 mi (by rd) E Tilaco, ca. 1 mi W Santa Inez (CSL 6863-64); 5.2 mi (by Tilaco Rd) S La Lagunita (LACM 127065-66, 127343- 46, 127418-20, 131143); (7): 16.4 mi E Landa de Matamoros (TCWC 29641); 15.5 mi E Landa de Matamoros (TCWC 32295). HIDALGO: (1): La Placita, 8 km S Jacala (UIMNH 26180-210); (2): Durango (AMNH 66063, 100769-70, EMNH 25885, KU 70550-51, MCZ 42145-48, UAZ 28880-915, UCM 49849-50, UIMNH 26220-23, 48010-11, USNM 111471, UMMZ 80951); 2 mi N Durango, 13 mi (by Mex 85) S Jacala (AMNH 100768, UAZ 28865-79); nr Minas Viejas (CAS- SU 13879, UIMNH 26211-19); 21 mi N Zimapan on Mex 85 (LACM 67530-34); 4 mi N Durango, 11 mi (by Mex 85) S Jacala (UAZ 28862-64). L. lipetzi CHIAPAS: (1): Lago de Mai Paso, headwaters of Rio de La Venta, 30 km N Cintalapa (UCM 51425); Lago de Malpaso, Boca de Rio de La Venta (ENEPI 3792). L. lineri OAXACA: (1): Cafetal El Pacifico, 2.1 mi N and 0.4 mi E Candelaria Loxicha on trail to Cafetal Hedweges (LACM 134844, UCM 48452); (2): Caf- etal “Alemania” (AMNH 15977). L. lowei OAXACA: (1): Zoogocho (IPN 5671-72); 4.0 km (by rd) SE San Bartolome Zoogocho (ENEPI 3732-33, 3804-07, IBH 7497-7500, EACM 143732-33); 5.8 km (by rd) S San Bartolome Zoo- gocho (ENEPI 3738); 1.0 km SE San Bartolome Zoogocho (ENEPI 3739). Contribotions in Science, Number 493 L. mayae GUATEMALA: (1): Alta Verapaz: Chinaja (KU 55863, 59553-=54, 59556-59, LACA4 75194-95); (2) : Vera Paz (NMBA 3751); (3): Huehuetenango: Finca Yulaxac, 15 km N Barillas {MVZ 134702); Finca Chiblac, 12 km NE Barillas (MVZ 143448- 49, 161798); Sierra de los Cuchumatanes, Finca Chiblac, 4.4 km (by rd) S Casa Grande on rd to “La Galera” (UTA R-38021); Finca Chiblac, ca. 22.0 km NNE Barillas (UTA R-38022-23); (4): Finca Volcan, 25 km NW (by rd) Senahu, Alta Ver- apaz (MVZ 159931). L. micropholis TAMAULIPAS: (0): cave (TTU R-7311); (1): Cave at Ei Pachon, ca. 5 mi NNE Antigua Morelos (LACM 106767-68, 136369-70, UAZ 28762, 28767-69, UIMNH 37296, UMMZ 101298-99, 102886-88); 11.3 mi S Ciudad Mante, Hwy 85 (TCWC 57256); (2): 1.5 mi SW Quintero (AMNH 93409); Gruta de Quintero (LACM 66662, SAM 885). SAN LUIS POTOSI: (1): 6 mi E Valles (BCB 13837-42); (2): 5.4 mi S, 1.4 mi E Valles (TCWC 60621, 60766-69, 60836). L. occulor SAN LUIS POTOSL (1): Hda. Capulin (LSUMZ 2379-80); (2): Taman (ENEPI 2946). QUERETA= RO: (1): Jaipan (USNM 47133-35); (2): 2.5 mi S Conca, Hda. Conca (TCWC 35605-06, 48499); (3) : 1.5 mi E Landa (TCWC 33063); 1.2 mi E Lan- da de Matamoros (TCWC 29691). L. pajapanense VERACRUZ: (1): SE slope Volcan San Martin Pajapan (FMNH 78382); SE slope Volcan San Martin (KU 97290, UMMZ 126362); S slope Vol- can San Martin (LACM 135510, UMMZ 118220); Colonia de Bastonal, above Quezalapam (TCWC 19133); 4 mi SE Tebanca (TCWC 21364); 1 mi NW Sontecomapan (TCWC 21365); Univ. Mex. Biol. Exp. Sta. ca. 33 km ENE Catemaco (TCWC 53351); Coyame, 9 mi (by rd) SE Catemaco (UAZ 28804); 2 mi (by rd) SE Sontecomapan, 14 mi (by rd) NE Catemaco (UAZ 28808-11, 28765); Lagu- na Catemaco, nr Cuezalapam (UMMZ 126363- 64); 2.1 mi NW (by rd) Sontecomapan (UTAR 3107, 3110, 3116); (2): 35 km SE Jesus Carranza (KU 26913); (3): Mpo. Huayacocotla: San Anto- nio, Ixtatetla (LACM 137605). L. radula OAXACA: (1): San Jose Manteca (USNM 111472). L. reticulatum COSTA RICA: (0): Pacific coast (KU 34209); (3): Puntarenas Prov.: Coto, km 47 on rail to Golfito (CRE 178); (4): San Jose Prov.: 6 km ESE San Isidro Bezy and Camarillo: Systematics of Lepidophyma ■ 37 del General (CRE 666); 2 mi E San Isidro del Gen- eral (KU 34189-93); 10-12 mi WSW San Isidro del General (KU 34194-202, 34204); 8.8 mi SW San Isidro del General (UMMZ 117474); 7.3 mi SW San Isidro del General (UMMZ 11475); (11): Pun- tarenas Prov.: 2 km SSW Canas Gordas (CRE 7227); Agua Buena (KU 36238-49, 56064); Finca Loma Linda (ca 2 km SW of Canas Gordas) (UMMZ 128639); (12): Puntarenas Prov.: 4 mi S San Vito de Java near Wilson’s Finca (CRE 3119); 6 km S San Vito de Java (LACM 72323); San Vito: ca. 1 mi SW Finca Las Cruces (LACM 76160-62, 76164-70); Copal; 4.5 mi SSE San Vito, vie. S Las Cruces (LACM 76163); jet. rd to Rio Java & San Vito- Agua Buena Rd, 1 mi S Las Cruces (LACM 106794); (13): Puntarenas Prov.: 1 and 7/8 mi SW of Rincon de Osa (CRE 7237); 6 km SSW Rincon (CRE 3158); 4.5 km W Rincon de Osa (KU 102449-50); (15): San Jose Prov.: Salitral de Santa Ana (CRE 229); (25): Guanacaste Prov.: 4-5 km NNE Tilaran (KU 36230-37). L. smithii GUERRERO: (1): 2 km W Puerto Marques (CU 9676-79, 9691-93, 9772); 2.5 km W Puerto Marques (IBH 1651, 2243-45); 1 km (airline) NW Puerto Marques (LACM 128590, 130027-29). VE- RACRUZ: 2 mi NW Sontecomapan (TCWC 26716). OAXACA: (0): Tehuantepec (MNHN 76.95, 4323, A4323, B4323); (1): 3 mi NW Te- huantepec (KU 38026-28); 6 mi WNW Tehuante- pec, Rio Tejas (KU 42289); 8.2 mi (by Hwy 190) \TOW Tehuantepec (LACM 128588-89, 130030- 32); nr Tehuantepec (MCZ 66974, USNM 46998, 47000); Tehuantepec (UIMNH 40745); Cerro Guiengola, 1/2 way up (FMNH 106312-15); Cerro Guiengola (UIMNH 26231-32); Quiengola Mt., 8 km NW Tehuantepec (UMMZ 82419-22); Cerro Arenal (USNM 111488); (2): 39 mi SW Cmtalapa (BCB 13400); 6 mi SE Tapanatepec (KU 42290); 2.7 mi E Tapanatepec (KU 75585); 95 mi N Tuxtla Gutierrez (LACM 100671); Municipio de Juchitan: 3.2 km NE Tapanatepec (LACM 134468-76, 136677); Tapanatepec (MCZ 27900); 1.5 mi (by Mex 190) E Tapanatepec (UAZ 28794,30279); Rio Novillero, 2.7 mi N Tapanatepec (UIMNH 83160); 3 mi SW of Chiapas border (UMMZ 118721); San- ta Efigenia (USNM 46687); (3): Rincon Bamba (UIMNH 35514, 35528); Rincon Bamba (UIMNH 37348); (5): 2 mi S La Ventosa (UIMNH 42835- 36); (6): 7.7 mi SE Camaron (UMMZ 115097); Portillo Nejapa (AMNH 98120); (9): Cerro Atra- vesado, El Palmar (AMNH 66890, 66955); (11): along Oaxaca-Puerto Escondido Rd., km [?] S of Oaxaca (CAS 101381); along Oaxaca-Puerto Es- condido Rd,, km 233 S of Oaxaca (CAS 101383); (13): El Limon (USNM 111490); (15): Tres Cruces (FMNH 100570, USNM 111489); La Concepcion (USNM 111491-92); (17): 29 mi W Tapanatepec (BCB 13397-99); (18): Santo Domingo de Guzman (BMNH 90.10.10.2); Mts nr Santo Domingo 38 ■ Contributions in Science, Number 493 (USNM 48105). CHIAPAS: (1): 9 mi (by Mex 200) NW Escuintla (AMNH 100767, 102295, UAZ 28797-802); La Esperanza (CAS-SU 13877; EMNH 123942-55, 124937-52, 125127-31, 178164-77, MCZ 46786-91, UCM 39685, UIMNH 10952-11005, USNM 111499-523); 30 mi NW Huixtla, Hwy 200 (LACM 100666-70, 101263); Cerro Ovando (LACM 135511); 2.5 mi (by rd) NE Acacoyaga (LACM 136359-64); 14 mi SE Mapastepec (TCWC 55523); 15.3 mi SE Ma- pastepec, Hwy 200 (TCWC 57502, 57504); 14 mi (by Mex 200) NW Escuintla (UAZ 28783-87, 28803); 2-3 km E (rd) El Triunfo (MVZ 191448- 49); 4 mi NW Mapastepec, 24 mi (by Mex 200) NW Escuintla (UAZ 28795, 28812-17); 4 mi (by Mex 200) NW Escuintla (UAZ 28796); ca. 1/2 mi (by Mex 200) NW Escuintla (UAZ 28788-93); Re- gion Soconusco (UIMNH 33604-05); Dist. Socon- usco, 6 km NE Escuintla (UMMZ 87251-54, 87257-300, 87303-22); Dist. Soconusco, vie Es- cuintla (UMMZ 88289, 88313); lake nr Acacoya- gua (USNM 111493-96); La Magnolia, 8 km NW La Esperanza (USNM 111497); (2): Tonola (EMNH 106949, 106951-52, 179365, USNM 111498); 15 mi (by Mex 200) SE Tonola (UAZ 28766); Nr Tonola (UIMNH 26226-29); (3): 6.8 mi N Arriaga (EAL 1746); 9.5 mi NE Arriaga (LACM 61258); crest above Arriaga (UMMZ 94825); (4): “km 945” between Tapanatepec and Las Cruces on Mex Hwy 190 (UMMZ 126867); (5) : near Asuncion (EMNH 106950, 106953). GUATEMALA: (28): (0): Guatemala occidental (MNHN 4968, A4968); (1): Escuintla: Escuintla (CAS 71914); Emca El Zapote (EMNH 57032); Einca El Salto, 2 km E Escuintla (MVZ 88420, 88422-23, 93174, 98023-24, 104181); 10 km N Escuintla, Einca Vista Hermosa (UMMZ 127839- 40); (2): Einca San Carlos Miramar, 12.5 km SW Yepocapa (by rd), Chimaltenango (MVZ 88426); (3): 1 mi SE Monogoy, Jutiapa (KU 42288); (4): Depto Jutiapa: Einca La Trinidad (UMMZ 107646-51); (5): Depto. Santa Rosa: Einca El Ca- cahuito (UMMZ 107070); 7 mi W Taxisco, “Km 99” near Cartago no. 1 bridge (UMMZ 126931); (6) : Depto. Santa Rosa: Einca La Gloria (UMMZ 118263); (7): 10 mi N Chicacao, Suchitepequez (UIMNH 46514-15); (8): Suchitepequez: Mazaten- ango (CAS 68484-86, 68488-89, 68491-504, CM 39734, NMBA 3750, 3748-49, 3752, 8039-40); (9): Suchitepequez: Volcan Zunil (CAS 68505-06, 68483); (11): Solola, Einca Olas de Moca (AMNH 43225). EL SALVADOR: (1): Dept. Sonsonate: Ha- cienda Chilata (MVZ 40071-73). L. sylvaticum NUEVO LEON: (1): La Boca (KU 92612-13); ca. 7 km NE Santiago, Presa La Boca (LACM 106781-92); (2): 5 mi N Las Ajuntas (EAL 4644); (3): San Roque, ca. 6 km S Cerro de la Silla (LACM 138179-80). TAMAULIPAS: (1): Rancho del Cielo [and vie.], ca. 5 mi NW Gomez Earias (AMNH Bezy and Camarillo: Systematics of Lepidophyma 107273, BMNH 1958.1.2.40, LACM 106751-56, UIMNH 37295, UMMZ 101301, 101374-75, 102978-81, 109763-64); El Elephante, ca. 7 km (airline) W Gomez Earias (LACM 106757-60); Ju- lilo, 12.7 km (airline) NW Gomez Earias (LACM 106762-65); W Rancho del Cielo (LSUMZ 10989); Gomez Earias-La Joya de Salas trail at 4600 ft. (UMMZ 102977); Gomez Earias-La Joya de Salas trail below Agua Zarca (UMMZ 109765); 2.5 mi ESE La Joya (UMMZ 109766); cloud forest above Encino (TCWC 65549); (2): 10 mi W, 2 mi S Piedra (KU 33993-94); Sierra de Tamaulipas, Santa Maria (UMMZ 102889-90); (3): 8 mi S, 6 mi W Victorio, Sierra Madre Otiental (KU 33992). SAN LUIS POTOSL (1): Alvarez (58 kilo) (MCZ 24507-08); between San Erancisco and Alvarez (MCZ 157826); Valle de los Eantasmos (SDSNH 60482); (3): Buenavista (20 mi NE Cerritos) (AMNH 64025); (4): 27 km (by Mex 80) W El Naranjo (LACM 131145-48); 3.8 mi (by Hwy 80) NNE Ciudad del Maiz (LACM 131144); 5 mi NE Ciudad del Maiz (TCWC 35582); (5): Huichihuay- an (EMNH 39631). QUERETARO: (1): El Madro- no, 3.5 mi W (rd) El Lobo (LACM 109771); 3.5 mi SW El Lobo (SAM 1104); 2.4 mi W El Madrono (TCWC 29692-95, 29701-07); 1.1 mi S El Ma- drono (TCWC 29696); 2.4 mi W, 1 mi S El Ma- drono (TCWC 29697); 2.4 mi W, 1.8 mi S El Ma- drono (TCWC 29698-700); 3.8 mi W El Lobo (TCWC 32291, 33064); 3.2 mi W El Lobo (TCWC 35607); 2 mi SW El Lobo (UMMZ 129749); (2): Mpo. Pinal de Amoles: 1 km S Pinal de Amoles (IBH 4060). HIDALGO: (1): 7 mi N Zacualtipan (EMNH 100102, 100356, 100411); 4.0 mi (by Hwy 105) S Tianquistengo (LACM 106721); 5.8 mi (by Hwy 105) S Tianquistengo (lAcM 106741-48, 136365-68); 3 mi S Tianquistengo (UIMNH 26230). VERACRUZ: (1): 4 km W Tla- pacoyam (KU 26909); (2): Alvarado Mpo.: Ciudad de Alvarado (MZEC 1722). L. tarascae MICHOACAN: (1): Aquila District, near Mexi- quillo (IPN 9221-22; LACM 134226); (2): Aquil- illa Mepo.: Rio Chilitos Canon, Cueva La Virgen (MZEC 8603). L. tuxtlae VERACRUZ: (1): Sontecomapan (CM 41470); Coxoapan (Sontecomapan) (UE 32914); Montepio (UE 32917); 18 mi NNE Catemaco (JCL 67); South slope Volcan San Martin Tuxtla (KU 59560); E of Lago Catemaco, 12.7 mi from Catemaco by rd (LACM 106795); ca. 5 mi N Catemaco off rd to Sontecomapan (LACM 134845); 12.9 mi (by rd) NNW Sontecomapan (LACM 136352-58); forest above Playa Escondida, 30 km NNE Catemaco (MVZ 171872); Salto de Eyipantla, Eyipantla (TCWC 19134); 1 mi W Zontecomapan (TCWC 19135); lower falls on Rio Quezalapam, Quezala- pam (TCWC 19136); Colonia de Bastonal (TCWC Contributions in Science, Number 493 19137); 4 mi SE Tebanca (TCWC 21366); Rio Quezalapan, 1 mi E Lago Catemaco (TCWC 21367); Rio Quezalapan, 2 mi E Lago Catemaco (TCWC 21368-69); Dos Arroyos, 5 mi E Zapoa- pan (TCWC 21370-71); Volcan San Martin (TCWC 22102-03); 2 mi NW Sontecomapan (TCWC 26717); Volcan San Martin E lower slope (TNHC 29792-93); Univ. Mex. Bio. Exp. Sta., ca. 33 km ENE Catemaco (TCWC 53352-53); Coya- me, 9 mi (by rd) SE Catemaco (UAZ 28763); 2 mi (by rd) SE Sontecomapan, 14 mi (by rd) NE Cate- maco (UAZ 28770-79); San Martin Mt., El Tular Sta. (UIMNH 35449); San Martin Mt. (UIMNH 35463-64); Rancho El Tular, ca. 5 mi N San An- dres Tuxtla (UIMNH 40285); Volcan San Martin (UIMNH 67064, 80695-99); Rio Tecolapan, 2.4 mi NNW Tapalapan (UMMZ 115098-99); S slope Volcan San Martin (UMMZ 118219); midway be- tween Coyame-Tebanca along shore Laguna Cate- maco (UMMZ 121164); SE slope, Volcan San Mar- tin (UMMZ 121165, 122112, 126360-61); be- tween Laguna Catemaco and Volcan San Martin (UMMZ 121166); Rancho El Tular, 15 mi N San Andres Tuxtla (USNM 139731); 2.1 mi NW (by rd) Sontecomapan (UTAR 3101, 3103, 3104, 3108, 3109, 3111-13, 3115); 2.5 mi SSW Sontecomapan (UTAR 3127); ca. 3 mi E of Cuetzalapan (UTAR 3133); Rio Cuetzalapan S of Tebanca (UTAR 3139-40); 5.6 mi ESE Tebanca (UTAR 3156); 7.7 mi NW Sontecomapan (UTAR 3728-30, 3733-34); (2): 25 km SE Jesus Carranza (KU 26912). OA- XACA: (1): Vista Herm.osa (KU 87396-98, UCM 49280); 30 mi (by rd) NE Llano de las Elores (UMMZ 125870); Valle Nacional Mpo.: alrede- dores de Metates, Sierra Juarez (MZEC 2262); (2): Mts nr La Gloria (UIMNH 37236); (3): Finca San Carlos, Matias Romero Oaxaca, Rio Coatzacoal- cos, at the mouth of Rio Sarabia (UF 32918). CHIAPAS: (1): 43 km on rd between Ocozocoautla and Malpaso (UCM 49281); 25 mi (by rd to Mal- paso) NW Ocozocoautla (UAZ 28780-82). ACKNOWLEDGMENTS The following individuals facilitated this study by provid- ing access to specimens in collections that are or were under their care: AMNH (C.J. Cole), ANSP (TM. Uzzell), BCB (B.C. Brown; material at Strecker Museum), BMNH (E.N. Arnold), CAS (R.C. Drewes, A.E. Leviton, J. Vin- dum), CM (C.J. McCoy), CRE (J.M. Savage; material now at LACM), CU (EH. Pough), EAL (E.A. Liner; ma- terial now at AMNH), EMNH (R.E Inger, H. Marx, H.K. Voris), IBH (G. Casas A., A. Ramirez B.), IPN (T. Alvar- ez), JFC (J.F. Copp; material now at CAS), KU (W.E. Duellman), LACM (K. Beaman, D. A. Kizirian), LSUMZ (D.A. Rossman), MCZ (J. Rosado, E.E. Williams), MNHN (A. Dubois, J. Guibe), MVZ (H.W Greene, D.B. Wake), MZEC (O. Flores V.), NMBA (E. Kramer), OMNH (J.P. Caldwell, L.J. Vitt), SAM (S.A. Minton; ma- terial now at AMNH), SDSNH (G.K. Pregill); TCWC (J.R. Dixon), TNHC (R.E Martin), TTU (J. Mecham), UAZ (C.H. Lowe), UCM (S.-K. Wu), UF (W Auffenberg), UIMNH (D.F. Hoffmeister), UMMZ (A.G. Kluge), UMRC (J.C. Lee), USNM (R.W McDiarmid, G.R. Zug), Bezy and Camarillo: Systematics of Lepidophyma ■ 39 UTA (J.A. Campbell, W.E Pyburn), and UTEP (C.S. Lieb, R.C. Webb). Over the years many people have aided us by providing live material, field assistance, and/or infor- mation about field localities for Lepidophyma, including Miguel Alvarez del Toro, Kit Bezy, Kathryn Bolles, Janalee Caldwell, Duke Campbell, Jonathan Campbell, Jay Cole, Salvador Contreras Balderas, James Dixon, R. Durtsche, Alfred Gardner, Jerry Johnson, John Karges, Carl Lieb, Robert Lonard, Charles Lowe, James Lynch, Roy Mc- Diarmid, Jose Ortiz, Martin Ruggles, Norman Scott, Wade Sherbrooke, Hobart Smith, Sam Telford, Wayne VanDevender, Laurie Vitt, Richard Vogt, David Wake, Barbara Warburton, John Wright, P. Zani, and members of the 1977 Tapezco Earth Watch Expedition. Scientific collecting permits were issued by the Direccion General de Vida Silvestre. Kathryn Bolles and Jessie Easley helped with data entry. Photographs and/or photographic assis- tance were provided by Kraig Adler, Kit Bezy, Kathryn Bolles, and John DeLeon. Kit Bezy, Kathryn Bolles, and Tina Ross produced the illustrations. Hobart Smith gen- erously shared his extensive knowledge of the genus Lep- idophyma. Kathryn Bolles, Jonathan Campbell, Jay Cole, Gordon Hendler, Inez Horowitz, and David Kizirian re- viewed an earlier version of the paper and made valuable suggestions. Financial support was provided by the Nat- ural History Museum of Los Angeles County Foundation and a grant from the Ralph J. Weiler Foundation. LITERATURE CITED Aguilar Cortes, R., J.L. Camarillo R., and R.E. Bezy. 1990. Distribution, species status, and reproductive mode of the xantusiid lizard Lepidophyma pajapa- nensis. Southwestern Naturalist 35:273-274. Alvarez, T, and E. Diaz-Pardo. 1983. Estudio de una co- leccibn herpetofaunistica de la costa de Michoacan, Mexico. Anales de la Escuela Nacional de Ciencia Biologta Mexico 27:129-147. Alvarez, T, and N. Valentin. 1988. Descripcion de una nueva especie de Lepidophyma (Reptilia: Xantusi- idae) de Chiapas, Mexico. Anales de la Escuela Na- cional de Ciencia Biologta Mexico 32:123-130. Alvarez del Toro, M. 1982. Los reptiles de Chiapas. Ter- cera Edicion. Tuxtla Gutierrez, Chiapas, Mexico: In- stituto de Historia Natural. 248 pp. Atchley, W.R., C.T. Gaskins, and D. Anderson. 1976. Sta- tistical properties of ratios. I. 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