Number of Pages: 3Integrative BiologyGeological Science

Chiszar, David

Liner, Ernest A.

Ponce-Campos, Paulino

Smith, Hobart M.

English

Number of Pages: 3Integrative BiologyEarth and Planetary Science

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REPTILIA: SQUAMATA: PHRYNOSOMATIDAE SCELOPORUS HETEROLEPISCatalogue of American Amphibians and Reptiles.Smith, H.M., P. Ponce-Campos, E.A. Liner, and D. Chiszar.2004. Sceloporus heterolepis.Sceloporus heterolepis BoulengerSceloporus heterolepis Boulenger 1894:731. Type locality,“various localities in the State of Jalisco (La Cumbre de LosArrastrados, Real Alto, Riocho [= Rancho] La Berbería [inthe] Sierra de Bolaños) at altitudes varying between 7800 and8500 feet.” Syntypes, Museum of Comparative Zoology,Harvard University (MCZ) 32346, California Academy ofSciences (CAS) 3712–3 (lost), British Museum (NaturalHistory (BMNH) 92.2.8.30, 1946.8.10.20, 1946.8.25–30,1946.8.10.37–43, collected by Dr. A.C. Buller. Inasmuch asthe syntype from Rancho La Berbería represents a differentspecies (S. shannonorum Langebartel) from the one to whichthe name has previously been applied, we here designateBMNH 92.2.8.30 from La Cumbre de los Arrastrados aslectotype. It most closely conforms with Boulenger’s (1894)illustration fide Colin McCarthy, BMNH. MCZ 32346 is atopotypic syntype.Sceloporus heterolepsis: Larsen and Tanner 1974: 6. Lapsus.Sceloporus heterolepis heterolepis: Webb 1969:305.797.1FIGURE 2. Venter of an adult female S. heterolepis from Volcán deTequila, Jalisco, 2876 m (20°47'42.0"N, 102°50'37.0"W), 8 September2001 (photograph by PPC). Note the complete absence of both gularand abdominal semeions, sharply different from sympatric S. grammicus,which has at least abdominal semeions.FIGURE 3. A juvenile female S. heterolepis, 31.8 mm SVL, same lo-cality as in Figure 1, 21 July 2001 (photograph by PPC).FIGURE 1. The presumed lectotype of S. heterolepis (from Boulenger1894), showing dorsal scalation.MAP. Distribution of Sceloporus heterolepis: the circle indicates thetype locality; dots indicate other sites of collection.0 200 km100• CONTENT. No subspecies are recognized.• DEFINITION. Sceloporus heterolepis is a medium-sizedspecies, maximum SVL 70 mm in males, 63 mm in females.Dorsal and lateral scales are irregular in size, small ones scatteredamong larger scales, or enlarged scales among smaller ones. Aparavertebral row of 31–38 enlarged scales form a continuousrow on each side from the neck to the rear margins of the thighs;these are narrowly separated from each other by an irregularseries of 56–80 small vertebral scales. Large dorsolateral nuchalscales are sharply differentiated from small lateral nuchal scales,forming a prominent lateral nuchal fringe and tuft. A group oftiny scales separates the two rows of paravertebral scales on theneck, and another group separates the paravertebral scales fromthe nuchal fringe on each side. Femoral pores number 13–21 ( x= 16, N = 36).Dorsal and lateral surfaces are dark gray, with 4–5 transversedark lines that are light-bordered and usually much expandedlengthwise at middorsum. The first is anterior to the arm and inmales crosses the throat. Males have bluish, medially dark-bordered abdominal semeions, each with a relatively short darkborder tapering at each end; these are in contact medially ornarrowly separated. The gular region is light yellowish in males,with some dark speckling. Females are unmarked ventrally.• DIAGNOSIS. This species is unique in its genus in having acontinuous paravertebral row of 31–38 enlarged scales on eachside, narrowly separated by an irregular vertebral series of 56–80 small scales. Also unique is the presence of three rows ofvery small scales on the nape separating the paravertebral rowsof larger scales from each other and from the lateral nuchalfringes. These features also separate this species from its closestrelative, S. shannonorum. The latter species usually has twosuperimposed preoculars, whereas S. heterolepis usually has one.All scales are imbricate. Sympatric S. grammicus females differin having rather well-developed abdominal semeions.• DESCRIPTIONS. Descriptions are in Boulenger (1894),Smith (1939), Grant and Smith (1960), Duellman (1961), Webb(1969) and Köhler and Heimes (2002).• ILLUSTRATIONS. A black and white drawing of thelectotype is in Boulenger (1894), reproduced herein.• DISTRIBUTION. This species occurs in oak-pine-fir forestsfrom central and eastern Jalisco to western Michoacán, at eleva-tions of 1250–3000 m, south of the Río Santiago Valley in thewestern end of the Neovolcanic Axis and the northern end ofthe Sierra Madre del Sur. Sceloporus shannonorum replaces itin the Sierra Madre Occidental.• FOSSIL RECORD. None.• PERTINENT LITERATURE. Citations in the literature incontexts other than previously indicated include the following:checklists (Flores-Villela 1993; Liner 1994; Smith and Taylor1950b, 1966), common names (Sokolov 1988, Liner 1994),comparisons of S. h. heterolepis and S. h. shannonorum(Langebartel 1959, Duellman 1961, Webb 1969), distribution(Cope 1896; Flores-Villela and Gérez 1988, 1994; McCranieand Wilson 1987, 1990; Nieto-Montes de Oca 1987; Wilsonand McCranie 1979), ecology (Duellman 1965), epidermato-glyphics (Burstein et al. 1974), habitat (Duellman 1961, 1965;Flores-Villela 1993; Webb 1969), karyology (Gilboa 1974, Hall1980, Sites et al. 1992), key (Köhler and Heimes 2002, Smithand Taylor 1950b), literature (Smith and Smith 1976, 1993),localities (Campbell and Murphy 1977; Chrapliwy 1956; Gadow1905, 1930; Grant and Smith 1960; Langebartel 1959; Smithand Taylor 1950b; Webb 1982, 1984), mention only (Wills1977), phylogeny (Flores-Villela et al. 2000; Hall 1980; Larsenand Tanner 1974, 1975; Sites et al. 1992; Wiens and Reeder1997), semeions (Wiens 1999), skull (Larsen and Tanner 1974),speciation (Hall 1980), types (Barbour and Loveridge 1946,Smith et al. 1964), type locality (Smith and Taylor 1950a), andviviparity (Guillette et al. 1980, Méndez-de la Cruz et al. 1998).• REMARKS. This species belongs to the S. grammicus group(Webb 1969, Wiens and Reeder 1997) and, like others of thatgroup, is arboreal.Webb (1969) described what he regarded as intergradesbetween S. shannonorum and S. heterolepis in a sample of threelizards from the western edge of the range of the latter (Sierrade Cuale, Jalisco; see Comment) and of one from Sierra deAutlán, 15 mi SE Autlán, Jalisco. They are not, however, froman area between the ranges of the two species as presentlyunderstood. No evidence suggests that the two taxa intergrade;they are dichopatric so far as is known at present.• ETYMOLOGY. The name heterolepis (Latin, irregular scales)reflects the varied sizes of the dorsal and lateral scales in thisspecies.• COMMENT. We have examined the series from the Sierra deCuale, and regard their morphology as within the range ofvariation of S. heterolepis. The dorsal scales are flattened, buttheir arrangement is more or less typical. Most importantly, thedorsal nuchal scales are of different sizes, with small scalesbetween the paravertebral series, and also between the latterand the lateral nuchal fringes. These specimens are from analtitude (1250 m) considerably lower than others recorded forthe species.• ACKNOWLEDGMENTS. We are much indebted to theauthorities of these institutions for information on their holdingsof this species (acronyms follow Leviton et al. 1985, except forMZFC, Museo de Zoología, Facultad de Ciencias, UNAM):CAS, KU, MZFC, TCWC, UCM, UMMZ, USNM, UTA, andUTEP. We especially thank Dr. Colin McCarthy, who kindlyprovided critical information on the syntypes of S. heterolepis,which includes one (BMNH 92.2.8.30) from Rancho LaBerbería, Sierra de Bolaños, representing S. shannonorum.Whitney C. Johnson and Dr. Sharon K. Collinge, EE Biology,University of Colorado, prepared the map.LITERATURE CITEDBarbour, T. and A. Loveridge. 1946. First supplement to typical reptilesand amphibians. Bull. Mus. Comp. Zool. 96:59–214.Boulenger, G.A. 1894. Second report on additions to the lizard collectionin the Natural-History Museum. Proc. Zool. Soc. London 1894:722–736, pls. 57–59.Burstein, N., K.R. Larsen, and H.M. Smith. 1974. A preliminary surveyof dermatoglyphic variation in the lizard genus Sceloporus. J. Herpe-tol. 8:359–369.Campbell, J.A. and J.B. Murphy. 1977. A new species of Geophis (Rep-tilia, Serpentes, Colubridae) from the Sierra de Coalcomán, Micho-acán, México. J. Herpetol. 11:397–403.Chrapliwy, P.S. 1956. Extension of known range of certain amphibiansand reptiles of Mexico. Herpetologica 12:121–124.Cope, E.D. 1896. The geographical distribution of Batrachia and Reptiliain North America. Amer. Nat. 30:886–902, 1003–1026.Duellman, W.E. 1961. The amphibians and reptiles of Michoacán, Méx-ico. Univ. Kansas Publ. Mus. Nat. Hist. 15:1–148.—. 1965. A biogeographic account of the herpetofauna of Michoacán,México. Univ. Kansas Publ. Mus. Nat. Hist: 15:627–709.Flores-Villela, O. 1993. Herpetofauna Mexicana. Annotated list of the797.2species of amphibians and reptiles of Mexico, recent taxonomicchanges, and new species. Spec. Publ. Carnegie Mus. Nat. Hist. (17):iv+ 73 p.— and P. Gérez. 1988. Conservación en México: Síntesis sobre Verte-brados Terrestres, Vegetación y el Uso del Suelo. Instituto Nacionalde Investigaciones sobre Recursos Bióticos, Xalapa, México.— and —. 1994. Biodiversidad y conservación en México: vertebrados,vegetación y uso del suelo. 2nd ed. Comisión Nacional para el Cono-cimiento y Uso de la Biodiversidad, and UNAM, México, D.F.—, K.M. Kjer, M. Benabib, and J.W. Sites, Jr. 2000. Multiple data sets,congruence, and hypothesis testing for the phylogeny of basal groupsof the lizard genus Sceloporus (Squamata, Phrynosomatidae). Syst.Biol. 49:713–739.Gadow, H. 1905. The distribution of Mexican amphibians and reptiles.Proc. Zool. Soc. London 1905:191–244.—. 1930. The history of the volcano of Jorullo and the reclamation ofthe devastated district by animals and plants. Cambridge Univ. Press,Cambridge.Gilboa, I. 1974. Karyotypes of amphibians and reptiles: a biogeographicreview, p. 91–156. In H.G. Dowling (ed.), 1974 Yearbook of Herpe-tology. New York, Herpetol. Info. Search Systems, Amer. Mus. Nat.Hist., New York.Grant, C. and H.M. Smith. 1960. Herpetozoa from Jalisco, Mexico. Her-petologica 16:39–43.Guillette, L.J., Jr., R.E. Jones, K.T. Fitzgerald, and H.M. Smith. 1980.Evolution of viviparity in the lizard genus Sceloporus. Herpetologica36:201–215.Hall, W.P. 1980. Chromosomes, speciation and evolution of Mexicaniguanid lizards. Nat. Geogr. Soc. Res. Rep. (12):309–329.Köhler, G. and P. Heimes. 2002. Stachelleguane: Lebensweise, Pflege,Zucht. Herpeton, Offenbach, Germany.Langebartel, D.A. 1959. A new lizard (Sceloporus) from the Sierra MadreOccidental of Mexico. Herpetologica 15:25–27.Larsen, K.R. and W.W. Tanner. 1974. Numeric analysis of the lizardgenus Sceloporus with special reference to cranial osteology. GreatBasin Nat. 34:1–41.— and —. 1975. Evolution of the sceloporine lizards (Iguanidae). GreatBasin Nat. 35:1–20.Leviton, A.E., R.H. Gibbs, Jr., E. Heal, and C.E. Dawson. 1985. Stan-dards in herpetology and ichthyology: Part I. Standard symbolic codesfor institutional resource collections in herpetology and ichthyology.Copeia 1985:802–821.Liner, E.A. 1994. Scientific and common names for the amphibians andreptiles of Mexico in English and Spanish. Nombres científicos ycomunes en Inglés y Español de los anfibios y los reptiles de México.SSAR Herpetol. Circ. (23):v + 113 p.McCranie, J.R. and L.D. Wilson. 1987. The biogeography of the herpe-tofauna of the pine-oak woodlands of the Sierra Madre Occidental ofMexico. Milwaukee Pub. Mus. Contrib. Biol. Geol. (72):1–30.— and —. 1990. Annotated bibliography to the herpetofauna of thepine-oak woodlands of the Sierra Madre Occidental, Mexico. Smith-son. Herpetol. Info. Serv. (84):1–16.Méndez-de la Cruz, F.R., M. Villagrán-Santa Cruz, and R.M. Andrews.1998. Evolution of viviparity in the lizard genus Sceloporus. Herpeto-logica 54:521–532.Nieto-Montes de Oca, A. 1987. Variación morfológica en Sceloporusgrammicus (Lacertilia, Iguanidae) entre poblaciones de las partescentral y noroccidental de su área de distribución. Biol. Diss., Esc.Nac. Est. Prof. Zaragoza, México, D.F.Sites, J.W., Jr., J.W. Archie, C.J. Cole, and O. Flores Villela. 1992. Areview of phylogenetic hypotheses for the lizards of the genus Scelop-797.3orus (Phrynosomatidae): implications for ecological and evolutionarystudies. Bull. Amer. Mus. Nat. Hist. (213):1–110.Smith, H.M. 1939. The Mexican and Central American lizards of thegenus Sceloporus. Field Mus. Nat. Hist. Zool. Ser. 26:1–397.—, D.A. Langebartel, and K.L. Williams. 1964. Herpetological type-specimens in the University of Illinois Museum of Natural History.Illinois Biol. Monogr. (32):1–80.— and R.B. Smith. 1976. Synopsis of the Herpetofauna of Mexico. Vol.III. Source Analysis and Index for Mexican Reptiles. John Johnson,North Bennington, Vermont.— and —. 1993. Synopsis of the Herpetofauna of Mexico. Vol. VII.Bibliographic Addendum IV and Index, Bibliographic Addenda II–IV, 1979–1991. Univ. Press Colorado, Niwot.— and E.H. Taylor. 1950a. Type localities of Mexican reptiles and am-phibians. Univ. Kansas Sci. Bull. 33:313–380.— and —. 1950b. An annotated checklist and key to the reptiles ofMexico exclusive of the snakes. Bull. U.S. Natl. Mus. (199):v + 253p.— and —. 1966. Herpetology of Mexico. Annotated checklists and keysto the amphibians and reptiles. A reprint of Bulletins 187, 194 and199 of the U.S. National Museum, with a list of subsequent taxonomicinnovations. Eric Lundberg, Ashton, Maryland.Sokolov, V.E. (ed.). 1988. Dictionary of Animal Names in Five Lan-guages. Amphibians and Reptiles. Russky Yazyk, Moscow.Webb, R.G. 1969. Variation, status, and relationship of the iguanid lizardSceloporus shannonorum. Herpetogica 25:300–307.—. 1982. Distributional records for Mexican reptiles. Herpetol. Rev.13:132.—. 1984. Herpetogeography in the Mazatlán-Durango region of theSierra Madre Occidental, Mexico. Spec. Publ. Univ. Kansas Mus.Nat. Hist. 10:217–241.Wiens, J.J. 1999. Phylogenetic evidence for multiple losses of a sexuallyselected character in phrynosomatid lizards. Proc. R. Soc. Lond. B266:1529–1535.— and T.W. Reeder. 1997. Phylogeny of the spiny lizards (Sceloporus)based on molecular and morphological evidence. Herpetol. Monogr.(11):1–101.Wills, F.M. 1977. Systematics of Sceloporus parvus. Unpubl. M.S.Thesis, Texas A&M Univ., College Station.Wilson, L.D. and J.R. McCranie. 1979. Notes on the herpetofauna oftwo mountain ranges in Mexico (Sierra Fría, Aguascalients, and SierraMorones, Zacatecas). J. Herpetol. 13:271–278.HOBART M. SMITH, Department of EEBiology, Universityof Colorado, Boulder, CO 80309-0334 USA (hsmith@colorado.edu); PAULINO PONCE-CAMPOS, BosqueTropical A. C., A. P. 5-515, Guadalajara, Jalisco, México, 45042(poncecp@hotmail.com); ERNEST A. LINER, 310 MalibouBoulevard, Houma, LA 70364-2598 (eliner@mobiletel.com);and DAVID CHISZAR, Department of Psychology, Universityof Colorado, Boulder, Colorado 80309-0345 (chiszar@clipr.colorado.edu).Primary editor for this account, Andrew H. Price.Published 30 October 2004 and Copyright © 2004 by the Societyfor the Study of Amphibians and Reptiles.

Sceloporus heterolepis

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Sceloporus heterolepis

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