(anura: leptodactylidae) from minas gerais, southeastern brazil - BioOne

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ABSTRACT: A new, brightly colored species of Physalaemus similar to P. deimaticus ... Estadual do Itacolomi, Municipality of Ouro Preto, State of Minas Gerais,.

Herpetologica, 59(4), 2003, 519–524 Ó 2003 by The Herpetologists’ League, Inc.





Departamento de Vertebrados, Museu Nacional/UFRJ, Quinta da Boa Vista, 20940-040 Rio de Janeiro, RJ, Brasil 2 Departamento de Biologia Animal, Universidade Federal de Vic¸osa, 36571-000 Vic¸osa, MG, Brasil 3 Instituto Estadual de Florestas, Caixa Postal 116, 35400-000 Ouro Preto, MG, Brasil ABSTRACT: A new, brightly colored species of Physalaemus similar to P. deimaticus and P. rupestris is described from the Parque Estadual do Itacolomi, Municipality of Ouro Preto, State of Minas Gerais, southeastern Brazil. The new species is characterized by its small size (snout–vent length 23.2 mm in male, 26.2–26.6 mm in females) and by having the snout rounded in the dorsal view and protruding in profile; large inguinal glands, ovoid, light brown with a black ocellus outlined with white on their posterior half; light grayish brown dorsum with elongate longitudinal dark grayish brown stripes and blotches; venter white with gray mottling; and bright red on groin and ventral surfaces of arms, hands, thighs, tibiae, and feet. Key words: Amphibia; Anura; Leptodactylidae; Physalaemus erythros sp. nov.; Southeastern Brazil; Taxonomy

THE FROG genus Physalaemus contains 41 recognized species that are distributed from Mexico to southern South America (Frost, 2002). The genus has been arranged into four species groups by Lynch (1970); however, these groupings have been disputed and treated as non-monophyletic (Cannatella and Duellman, 1984; Heyer and Wolf, 1989). Although Caramaschi et al. (1991) considered these groups useful in discussing diversity within the genus, the affinities of several species with the currently recognized species groups is unclear. For instance, P. deimaticus Sazima and Caramaschi, 1988 and P. rupestris Caramaschi, Carcerelli and Feio, 1991 apparently do not belong to any of Lynch’s (1970) species groups. Both species are similar in morphology and habitat, as noted by Caramaschi et al. (1991), but these authors were unwilling to propose a new group for that species without a more thorough study of the genus, although they recognized that it might constitute a distinct morphological and ecological unity. During a survey of anuran species of the Parque Estadual do Itacolomi in the State of Minas Gerais, Brazil, specimens of a brightly colored species morphologically and ecologically similar to P. deimaticus and P. rupestris


CORRESPONDENCE: e-mail, [email protected] 519

were collected. Herein, we describe this new species. MATERIALS AND METHODS Specimens examined are deposited in MNRJ (Museu Nacional, Rio de Janeiro, RJ, Brazil), MZUFV (Museu de Zoologia Joa˜o Moojen de Oliveira, Universidade Federal de Vic¸osa, MG, Brazil), and JJ (Jorge Jim Collection, housed in the Departamento de Zoologia, Universidade Estadual Paulista, Botucatu, SP, Brazil). Specimens examined are listed in Appendix I. Abbreviations used for measurements in millimeters are SVL (snout–vent length), HL (head length), HW (head width), IND (internarial distance), END (eye–nostril distance), ED (eye diameter), UEW (upper eyelid width), IOD (interorbital distance), THL (thigh length), TL (tibia length), and FL (foot length). Specimens were fixed in 10% formalin and preserved in 70% alcohol. Measurements were made to the nearest 0.01 mm with digital calipers under a Zeiss SV-4 stereomicroscope; drawings of the holotype were made using a drawing tube. Physalaemus erythros sp. nov. Holotype.—MNRJ 27986, adult male (Fig. 1) from Parque Estadual do Itacolomi (208 259 S, 438 299 W, approximately 1600 m elevation), Municipality of Ouro Preto, State of Minas



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FIG. 1.—Dorsal and ventral views of Physalaemus erythros sp. nov. (MNRJ 27986, holotype, SVL 23.2 mm).

Gerais, Brazil, 29 October 2001, collected by A. S. Guimara˜es-Neto and J. B. Isaac, Jr. Paratypes.—Collected in the type locality: MNRJ 27539, adult female, 22 October 2001, by R. N. Feio and A. S. Guimara˜es-Neto; MNRJ 30608, adult female, 12–13 December 2002, by A. S. Guimara˜es-Neto, C. A. G. Cruz, R. N. Feio, L. B. Nascimento, and J. P. Pombal, Jr. Diagnosis.—A species ressembling P. deimaticus and P. rupestris was diagnosed by the following characters: (1) relatively small size (SVL 23.2 mm in male, 26.2–26.6 mm in females); (2) snout rounded in dorsal view and protruding in profile; (3) vomerine, premaxillary, and maxillary teeth absent; (4) relative lengths of fingers II , IV , I , III; (5) dorsal skin slightly rugose, ventral skin smooth, and proximal undersurfaces of thighs granular; (6) parotoid glands absent; (7) inguinal glands large, ovoid, light brown with a black, white outlined ocellus on their posterior half; (8) dorsum light grayish brown with elongate longitudinal grayish dark brown stripes and blotches; (9) venter white with gray mottling; (10) ventral surfaces of arms, hands, thighs, tibiae, feet, and inguinal region bright red. Comparisons with other species.—Physalaemus deimaticus, P. erythros, and P. rupestris are excluded from the P. biligonigerus species

group of Lynch (1970) by the smaller size (combined SVL of the three species, 15.9–23.2 mm in males, 17.2–26.6 mm in females; of the species in the P. biligonigerus group, 40–45 mm), relatively slender build (chubby in P. biligonigerus group), tarsal tubercle absent (present in P. biligonigerus group), inner metatarsal tubercle ovoid and outer metatarsal tubercle rounded (enlarged, compressed metatarsal tubercles in P. biligonigerus group), and absence of premaxillary and maxillary teeth (present in P. biligonigerus group, except P. nattereri). From the P. pustulosus group, by the dorsal skin rugose (tuberculate in P. pustulosus group), inguinal glands present (absent in P. pustulosus group), absence of parotoid and flank glands (present in P. pustulosus group), and tarsal tubercle absent (present in P. pustulosus group, except P. freibergi and P. pustulatus). From the P. signifer group, by the dorsal skin rugose (smooth in P. signifer group), snout rounded in dorsal view (acuminate in P. signifer group), no black lateral, defined band from eye to groin (present in P. signifer group), concealed tympanum (evident in P. signifer group, except P. maculiventris and P. olfersi), and inner metatarsal tubercle present (absent in P. signifer group). From the P. cuvieri group, by the absence of tarsal tubercle (inner tarsal

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tubercle present in P. cuvieri group), presence of large inguinal glands (lack or small inguinal glands in P. cuvieri group), and absence of premaxillary and maxillary teeth (present in P. cuvieri group). Physalaemus erythros is distinguished from P. deimaticus and P. rupestris (see Caramaschi et al., 1991; Nascimento et al., 2001; Sazima and Caramaschi, 1988) by the greater size (SVL 21.4 mm in male, 24.2–24.4 mm in females of P. deimaticus; 15.9–18.3 mm in males, 17.2–17.9 mm in females of P. rupestris) and the shape of the snout, which is more elongate, round in dorsal view and protruding in profile (short, slightly acuminate in dorsal view and truncate in profile in P. deimaticus, see figures 2–3 in Sazima and Caramaschi, 1988; short, rounded in dorsal and lateral views in P. rupestris, see figures 2A–B in Caramaschi et al., 1991). The color pattern of the inguinal glands is light brown with a black, white outlined ocellus on the posterior half (inguinal glands dark brown to black with a fine light brown to white border in P. deimaticus and P. rupestris). There is also a bright red color on the ventral surfaces of the arms, hands, thighs, tibiae, feet, and inguinal region (hands, feet, and inguinal region pink to light red in P. deimaticus; palms and soles reddish with scattered black dots in P. rupestris). Description of holotype.—General aspect robust; head slightly longer than wide; head width approximately one-third SVL; snout rounded in dorsal view (Fig. 2A), protruding in profile (Fig. 2B); nostrils slightly protuberant, directed anterolaterally, nearer to tip of snout than to eyes; internarial distance slightly smaller than eye–nostril distance, equal to upper eyelid width; canthus rostralis rounded, slightly concave; loreal region concave; eye lateral, slightly protuberant; eye diameter larger than internarial and interorbital distances; tympanum concealed; supratympanic fold indistinct; vocal sac poorly developed, subgular; vocal slits present; choanae large, rounded, widely separated; tongue narrow, long, not notched posteriorly; vomerine, premaxillary, and maxillary teeth absent; a single, very small odontoid process protrudes dorsally from the symphysis of the mentomeckelian cartilage. Arms robust, forearms slightly hypertrophied; fold and crest on forearm absent; fingers long, slender, lacking webbing (Fig.


2C); lateral sides of fingers weakly fringed; nuptial pad large, subdivided among prepollical region and dorsal surface of finger I; subarticular tubercles single, round; inner metacarpal tubercle absent; outer metacarpal tubercle large, ovoid; supernumerary tubercles present; tip of fingers slightly expanded; relative lengths of fingers II , IV , I , III. Legs robust; tibia length shorter than thigh length; sum of tibia and thigh lengths less than SVL; foot length longer than lengths of both tibia and thigh; toes long, free, weakly fringed (Fig. 2D); tarsal tubercle absent; inner metatarsal tubercle large, ovoid, protruding; outer metatarsal tubercle small, conical, protuberant; subarticular tubercles single, round, protruding; supernumerary tubercles absent; tip of toes slightly expanded; relative lengths of toes I , II , V , III , IV; tarsal fold, crest, and tubercles absent. Inguinal glands large, ovoid, with an ocellus on posterior half. Dorsal skin slightly rugose, ventral skin smooth; proximal undersurfaces of thighs granular. In life, ground color of dorsum grayish light brown with irregular, longitudinal grayish dark brown stripes; an irregular grayish dark brown blotch on each upper eyelid; a dark brown, white outlined, longitudinal and semidivided mark from shoulder to region of urostyle; dorsal surfaces of arms and limbs grayish light brown with grayish dark brown transverse stripes and bars. Loreal region gray dorsally and grayish brown ventrally. A distinct lateral black stripe from posterior corner of eye to groin, dorsally outlined by a white line; small white dots scattered within this lateral black stripe. Inguinal glands light brown with a black, white outlined ocellus on the posterior half. Gular region gray, venter white with gray mottling. Red blotch extending from lateral regions of chest and proximal undersurfaces of arms to elbow. Ventral surfaces of hands and fingers red. Dorsal surfaces of hands and fingers III and IV grayish red, fingers I and II red. A large red blotch extending from inguinal region to middle of ventral surface of thighs and immediately below anal region. Undersurfaces of tibiae and internal area of feet to toes I, II, and III, red; toes IV and V grayish red with subarticular tubercles red; metatarsal tubercles red. Iris yellow with intense black vermiculation. In preservative, general color becomes gray with dark gray



FIG. 2.—Physalaemus erythros sp. nov. (MNRJ 27986, holotype). (A) Dorsal view of head; (B) lateral view of head; (C) ventral view of hand; (D) ventral view of foot.

marks on dorsum, grayish cream on arms and limbs; red color fades to cream. Measurements of holotype.—SVL 23.2; HL 7.8; HW 7.5; IND 1.8; END 1.9; ED 2.4; UEW 1.8; IOD 2.2; THL 10.3; TL 10.0; FL 11.0. Measurements of paratypes.—SVL 26.2– 26.6; HL 7.5–7.9; HW 8.1; IND 1.8–1.9; END 1.9; ED 2.4–2.7; UEW 1.9–2.0; IOD 2.3–2.5; THL 10.7–11.2; TL 10.6–10.8; FL 12.2–12.8. Variation.—Besides the measurements and details of color pattern involving the shape of the dorsal marks and minor differences in the extension of red areas, there is little variation among the type specimens. This small variation in color probably is not associated with sexual dimorphism, as also referred to P. rupestris by Nascimento et al. (2001). The male holotype has slightly hypertrophied arms; the first finger has a nuptial pad; the vocal sac is poorly developed; and the venter has a finer mottling. The female paratype MNRJ 27539 has a slim aspect and the venter is withered, which suggests recent egg laying. Etymology.—The specific name ‘‘erythros,’’ meaning ‘‘red,’’ is a Greek adjective used in reference to the remarkably bright ventral color pattern of the species.

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REMARKS The specimens were collected at the Serra do Itacolomi, in the Espinhac¸o Mountain Range, in an ephemeral pond at approximately 1600 m above sea level. This pond consists of a rain-flooded depression of approximately 800 m2 located in a rocky field called ‘‘campo rupestre,’’ with bushes and small trees on a sandy soil predominantly covered with grasses. Some scattered massifs of quartzitic rocks emerge in the field. The sparse bushes and small trees emerging from the continuous herbaceous layer are typical elements also found in other regions of ‘‘campos rupestres,’’ such as Velloziaceae, Orchidaceae, Bromeliaceae, and Eryocaulaceae (mainly Paepalanthus), and isolated Melastomataceae. The perimeter of the flooded area is densely covered by Juncaceae (mainly Juncus cf. microcephalus), Cyperaceae, and grasses, leaving only a small central portion of standing water, which is a typical ephemeral pond in an open area. During the dry months (April– September), the pond becomes totally dry, and peats and lichens cover the exposed areas between grasses. The regional climate is of CwB type (Ko¨ppen), that is, wet temperate with dry winters and hot and rainy summers. The annual rain mean is 1450–1900 mm (Pedralli et al., 2001). The site, despite its location in a conservation area, undergoes anthropogenic pressures, including frequent visitations by horses (feces were observed in the pond where the frogs were collected). In addition, the area was accidentally burned. Specimens of P. erythros were occasionally collected in October and December, in the rainy season. The advertisement call and tadpole of this species are unknown. In a more extensive survey of anuran species of the Parque Estadual do Itacolomi, conducted between January 1998 and April 1999 in two areas covered by ciliary and seasonal forests and ponds, Pedralli et al. (2001) registered 24 species. These included bufonids (Bufo crucifer), hylids (Hyla albopunctata, H. arildae, H. cavicola, H. decipiens, H. elegans, H. faber, H. martinsi, H. minuta, H. pardalis, H. polytaenia, H. semilineata, Hyla sp. of the circumdata group, Scinax eurydice, S. flavoguttatus, S. longilineus, S. luizotavioi, and S. rizibilis), and leptodactylids (Eleutherodactylus

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izecksohni, Hylodes sp. of the lateristrigatus group, Leptodactylus fuscus, L. labyrinthicus, L. ocellatus, and Physalaemus cuvieri). DISCUSSION The peculiar areas of open vegetation occurring above 1000 m high in montane regions of southeastern Brazil, mainly associated with the Cerrados and Atlantic Rain Forest Morphoclimatic Domains (Ab’Saber, 1977), are called ‘‘campos rupestres’’ (see Rizzini, 1979 for definition). These isolated highland areas commonly contain faunistic and floristic endemics (see Heyer, 1999 for comments), as exemplified by the three species of Physalaemus, respectively associated with the Serra do Cipo´ (P. deimaticus, see Sazima and Caramaschi, 1988), to the Serra do Ibitipoca (P. rupestris, see Caramaschi et al., 1991; Nascimento et al., 2001), and to the Serra do Itacolomi (P. erythros, present study). In addition to sharing similar habitats, the three species share morphological traits such as relatively small size for the genus, poorly developed vocal sacs, large inguinal glands, and the presence of red or reddish areas on fore and hindlimbs. Deimatic behavior, as reported for P. deimaticus by Sazima and Caramaschi (1988), was not observed for P. rupestris (Nascimento et al., 2001) or for P. erythros. The advertisement call and tadpole are known only for P. rupestris (Nascimento et al., 2001). It is possible that, as knowledge of the morphological relationships of the species of the genus Physalaemus grows, a distinct group will be recognized for these three species. RESUMO Uma nova espe´cie de Physalaemus de colorido vistoso, relacionada a P. deimaticus e P. rupestris, e´ descrita do Parque Estadual do Itacolomi, Municı´pio de Ouro Preto, Estado de Minas Gerais, sudeste do Brasil. A nova espe´cie e´ caracterizada pelo tamanho pequeno (comprimento rostro–anal 23,2 mm no macho, 26,2–26,6 mm nas feˆmeas), focinho arredondado em vista dorsal e saliente em perfil, glaˆndulas ingu¨inais grandes, ovo´ides, marrom claro e com um ocelo preto delineado por uma linha branca na sua metade posterior, dorso marrom claro acinzentado com linhas e manchas longitudinais marrom escuro acin-


zentado, ventre branco com marmoreado cinza e forte colorido vermelho nas superfı´cies ventrais dos brac¸os, ma˜os, coxas, tı´bias, pe´s e regia˜o ingu¨inal. Acknowledgments.—We thank C. A. G. Cruz, J. B. Isaac, Jr., J. P. Pombal, Jr., and L. B. Nascimento for help in the field work, H. C. Souza for botanical information on the surveyed area, P. R. Nascimento for the line drawings, and C. A. G. Cruz and J. P. Pombal, Jr. for critically reviewing the manuscript. We acknowledge the Instituto Estadual de Florestas de Minas Gerais for permission to collect in the Parque Estadual do Itacolomi; U. Caramaschi thanks the Conselho Nacional de Desenvolvimento Cientı´fico e Tecnolo´gico (CNPq) for the fellowship.

LITERATURE CITED AB’SABER, A. N. 1977. Os domı´nios morfoclima´ticos na Ame´rica do Sul. Primeira aproximac¸a˜o. Geomorfologia 52:1–21, 1 map. CANNATELLA, D. C., AND W. E. DUELLMAN. 1984. Leptodactylid frogs of the Physalaemus pustulosus group. Copeia 1984:902–921. CARAMASCHI, U., L. C. CARCERELLI, AND R. N. FEIO. 1991. A new species of Physalaemus (Anura: Leptodactylidae) from Minas Gerais, Southeastern Brazil. Herpetologica 47:148–151. FROST, D. R. 2002. Amphibian Species of the World: An Online Reference, Version 2.21. http://research.amnh. org/herpetology/amphibia/index.html HEYER, W. R. 1999. A new genus and species of frog from Bahia, Brazil (Amphibia: Anura: Leptodactylidae) with comments on the zoogeography of the Brazilian campos rupestres. Proceedings of the Biological Society of Washington 112:19–39. HEYER, W. R., AND A. J. WOLF. 1989. Physalaemus crombiei (Amphibia: Leptodactylidae), a new frog species from Espı´rito Santo, Brazil with comments on the P. signifer group. Proceedings of the Biological Society of Washington 102:500–506. LYNCH, J. D. 1970. Systematic status of the American leptodactylid frog genera Engystomops, Eupemphix, and Physalaemus. Copeia 1970:488–496. NASCIMENTO, L. B., R. R. CARVALHO, JR., H. WOGEL, D. S. FERNANDES, AND R. N. FEIO. 2001. Reproduc¸a˜o e descric¸a˜o do girino de Physalaemus rupestris Caramaschi, Carcerelli & Feio, 1991 (Amphibia, Anura, Leptodactylidae). Boletim do Museu Nacional, Nova Se´rie, Zoologia 450:1–10. PEDRALLI, G., A. S. GUIMARA˜ES-NETO, AND M. C. B. TEIXEIRA. 2001. Diversidade de anfı´bios na regia˜o de Ouro Preto. Cieˆncia Hoje 30:70–73. RIZZINI, C. T. 1979. Tratado de Fitogeografia do Brasil. Editora Hucitec and Editora da Universidade de Sa˜o Paulo, Brazil. SAZIMA, I., AND U. CARAMASCHI. 1988 [1986]. Descric¸a˜o de Physalaemus deimaticus, sp. n., e observac¸~oes sobre o comportamento deima´tico em P. nattereri (Steindn.)— Anura, Leptodactylidae. Revista de Biologia 13: 91–101. Accepted: 4 June 2003 Associate Editor: Joseph Mendelson III



APPENDIX I Specimens Examined Physalaemus deimaticus.—BRAZIL: Minas Gerais: Serra do Cipo´, Jaboticatubas (JJ 6057, holotype; JJ 6058, paratype).

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Physalaemus erythros.—BRAZIL: Minas Gerais: Parque Estadual do Itacolomi, Ouro Preto (MNRJ 27539, holotype; MNRJ 27986, 30608, paratypes). Physalaemus rupestris.—BRAZIL: Minas Gerais: Parque Estadual do Ibitipoca, Lima Duarte (MNRJ 10551, holotype; MNRJ 10552–53, paratypes; MNRJ 28812–24; MZUFV 4083–84).

Herpetologica, 59(4), 2003, 524–531 Ó 2003 by The Herpetologists’ League, Inc.




Museo de Zoologı´a, Facultad de Ciencias, Universidad Nacional Auto´noma de Me´xico, A.Pz. 70-399, Me´xico, D.F. 04510, Me´xico ABSTRACT: We describe a new species of lizard of the genus Abronia from the Sierra Madre del Sur of Guerrero, Mexico. This species previously was confused with its presumed sister taxon Abronia deppii. The discovery to the north of the Balsas Basin of additional specimens referrable to A. deppii and a morphological comparison of these specimens to the populations of Abronia inhabiting the highlands of Guerrero to the south of the Balsas Basin clearly indicate that distinct species are present on each side of the Balsas Depression. Examination of the type material of A. deppii reveals that this name should be associated with the northern populations, and the southern population, often allocated to A. deppii, is an undescribed species. This new species is distinguished from A. deppii by having enlarged knob-like posterior head scales, no azygous scale between interparietal and interoccipital, a greater number of scale whorls on unregenerated tail, a greater number of infralabial scales, and dorsal and ventral coloration differences. We discuss the natural history of the new Guerrero species. Key words: Abronia deppii; Abronia martindelcampoi sp. nov.; Anguidae; Guerrero, Mexico; Lacertilia; Taxonomy

THE ZOOLOGICAL literature on the lizard previously known as Abronia deppii has involved considerable confusion, apparently because of inadequate descriptions and the imprecise designation of the type locality. Additionally, a perplexing problem has arisen since the discovery of a disjunct population that was referred to A. deppei by Sa´nchez-H. and Lo´pez-F. (1980). These authors reported Abronia from northwest of the Balsas Basin. They compared external morphological data for their only specimen (now catalogued in the Museo de Zoologia, Facultad de Ciencias, 1

PRESENT ADDRESS: Av. Ixtlahuaca 609, Col. Sa´nchez, Toluca 50040, Estado de Me´xico, Me´xico. 2 CORRESPONDENCE: e-mail, [email protected]

UNAM, as MZFC 764) with data for specimens identified as A. deppii by Bocourt (1878) and Bogert and Porter (1967) from the Sierra Madre del Sur in the State of Guerrero. Because of limited materials, no striking differences in scalation were noted between specimen MZFC 764 from the State of Mexico and descriptions of material from Guerrero. Although they did not examine the type and paratype (Zoologisches Museum, Berlin; ZMB 1149, 1150, respectively), Sa´nchez-H. and Lo´pez-F. (1980) proposed rejection of the restriction of the type locality of A. deppii to Omilteme, Guerrero, suggested by Smith and Taylor (1950). Sa´nchez-H. and Lo´pez-F. (1980) suggested a corrected restriction to Temascaltepec-Real de Arriba, State of Mex-

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ico, in the vicinity of Valle de Bravo. One of the reasons for their proposal was the well documented fact that the collector of the type series of A. deppii, Ferdinand Deppe, never visited Guerrero, but traveled extensively in the State of Mexico (Sibley and Davis, 1946; Stresseman, 1954; Taylor, 1969). During the last few years, several persons at various Mexican institutions have discovered additional specimens of Abronia, both in the mountains north of the Balsas Basin and in the Sierra Madre del Sur of Guerrero (CastroFranco, 1987; Flores-Villela and HernandezGarcia, 1989; Flores-Villela and Mun˜oz Alonso, 1993). These specimens allow a more critical assessment of variation within and among the populations currently assigned to A. deppii. MATERIALS AND METHODS The present study is based on 37 adult specimens of Abronia, 9 and 28 individuals from the northern and southern populations, respectively (Appendix I); a few juveniles were also examined and their identities verified, but their morphological data are not included. We also had available a set of photographs, both black-and-white and color, of the type (ZMB 1149) and paratype (ZMB 1150) of A. deppii (Wiegmann, 1828). We analyzed external morphological characters that traditionally have been used by other workers (Bogert and Porter, 1967; Campbell, 1982; Campbell and Frost, 1993; Good and Schwenk, 1985). Body measurements were taken with vernier calipers to the nearest 0.1 mm. Based on our analysis of morphological characters and bolstered by the striking pattern of allopatry found among most species of the genus Abronia (e.g., Campbell and Frost, 1993, documented only one known case of sympatry), we regard northern and southern samples as representing two distinct species. These species differ from each other in details of scalation, body proportions, and color pattern. Flores-Villela (1993) indicated that the Guerrero population was a distinct species differing from A. deppii. The phylogenetic analyses published by Campbell and Frost (1993), Campbell et al. (1998), and Chippindale et al. (1998) clearly suggest that the populations to the north and south of the Rı´o Balsas Depression represent different species


and represent sister taxa. The type specimen (ZMB 1149) and the paratype (ZMB 1150) of A. deppii possess morphological characteristics that unquestionably associate them with specimens of the northern populations (see Campbell and Frost, 1993), confirming that the restriction of the type locality for this species to Temascaltepec-Real de Arriba, State of Mexico, by Sa´nchez-H. and Lo´pez-F. (1980) is appropriate. In order to more clearly diagnose the new species from Guerrero, a redescription of A. deppii follows. Abronia deppii (Wiegmann, 1828) Gerrhonotus deppii—Wiegmann, 1828 Gerrhonotus deppii—Wiegmann, 1834; Gu¨nther, 1902 Abronia deppii—Gray, 1838; Tihen, 1949; Campbell and Frost, 1993 Gerrhonotus (Abronia) deppii—Bocourt, 1878 Type.—Zoologisches Museum, Berlin (ZMB 1149). One paratype (ZMB 1150: see Fig. 1). Type locality.—Originally given as ‘‘Mexico,’’ restricted to Temascaltepec-Real de Arriba, State of Mexico by Sa´nchez-H. and Lo´pez-F. (1980). Restricted in error to the vicinity of Omilteme, Guerrero, by Smith and Taylor (1950). Diagnosis.—This species has (1) the supraauricular scales weakly developed knob-like, having no indication of spines; (2) supranasals not expanded and not in contact at dorsal midline; (3) frontonasal usually in contact with frontal; (4) posterior internasals almost twice as long as anterior internasals (1.8); (5) no canthals; (6) three anterior temporals in each side; (7) parietal separated from supraoculars; (8) single occipital; (9) posterior head scales weakly developed knob-like; (10) anterior supercilliary not contacting the cantholoreal; (11) posterior suboccular separated from the lower primary temporal by the penultimate scale in the supralabial series; (12) preauricular scales slightly enlarged and in one row; (13) two postmentals; (14) posterior infralabials elongated; (15) preauricular scales slightly enlarged; (16) six longitudinal nuchal scale rows; (17) 24–28 tranversal dorsal scale rows; (18) 10–12 longitudinal dorsal scale rows; (19)



FIG. 1.—Dorsal aspect of the head of type and paratype of Abronia deppii. A: ZMB 1149; B: ZMB 1150. Photograph by George R. Zug.

12–14 longitudinal ventral scale rows; (20) six, sometimes seven, dark dorsal bands. Abronia deppii can be distinguished from the Guerrero population by weakly developed knob-like posterior head scales (Fig. 2); lower anterior temporal in broad contact with antepenultimate supralabial; azygous scale between interparietal and occipital frequently present (about 60% of cases); head width ,22 mm in adult males and ,18 in adult females; ratio of head width to head length ,0.75 in adult males; scale whorls on tail 68–80 ( x 5 77); infralabial scales 7–8; preauricular scales only slightly enlarged; longitudinal dorsal scale rows 10–11; dorsum with well defined bold black crossbands, confluent and more or less symmetrical on vertebral region; dorsal head coloration with pattern of black and white irregular blotches; ventral color in adult males tends to be reddish, occasionally bright red; upper lips marked with black. Differs from other species of the subgenus Abronia (sensu Campbell and Frost, 1993) by having oblique longitudinal rows of dorsal scales, where fuscolabialis, taeniata, and graminea have

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FIG. 2.—Dorsal aspect of the head A: of the holotype of Abronia martindelcampoi, adult male, MZFC 2778; B: of an adult male A. deppii, ENEPI 963. Notice the well developed knob-like posterior head scales of A. martindelcampoi, as well as the presence of an azygous scale between the interparietal and interoccipital in A. deppii (present in 54% of the individuals examined, see text for details). Head length is approximately the same for both specimens.

parallel longitudinal scale rows, where taeniata and graminea do not; has one lower temporal contacting postoccular series; and a single occipital, where mixteca and oaxacae have three occipitals and two primary temporals contacting postocular series; has body coloration that it is not uniformly green, where graminea is; has six nuchals, where graminea has four (only juveniles of graminea have bands on the dorsum of the body). Description.—A species of Abronia having 24–28 ( x 5 26.2) transverse dorsal scale rows; 10–12 longitudinal dorsal scale rows ( x 5 10.6); 34–37 ( x 5 35) transverse ventral rows; 12–14 ( x 5 13.3) longitudinal ventral scale rows; 76–80 scale whorls on unregenerated tails; usually minimum of six nuchal scales (one of 12 specimens had 5); 6–8 ( x 5 6.6) scales between hind legs; 9–10 supralabials; anterior temporals 3/3; posterior temporals 3/3; 5/5

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FIG. 3.—Distribution of Abronia deppii and A. martindelcampoi in South Central Mexico. Triangles represent known localities for A. deppii and squares show known localities for A. martindelcampoi; dots refer to major cities. See Appendix I for specific localities.

medial supraoculars; one temporal in contact with postocular; usually one subocular (2 of 12 specimens had 2, see Good, 1988:20); two postmentals; and a single occipital. Supranasals not expanded; frontonasal and frontal usually in contact; no contact of anterior superciliary and cantholoreal (only 1 of 12 had contact); lateral neck scales enlarged; osteoderms reduced or absent on dorsum of adults. Coloration.—Dorsal ground color whitish or grayish in preservative (white in life); usually with six or occasionally seven dark dorsal bands confluent on mid-dorsum; adult males with tendency toward especially dark dorsum. Tail with 11–15 ( x 5 12.6) dark bands; dorsum of head with sharply defined irregular black and white blotches. Ventral coloration whitish with reddish hues on gular and pectoral regions (in some adult males, venter intense red-orange, as well as plantar and palmar surfaces). Measurements.—Maximumsnout–ventlength in adults 115 mm; maximum head length 31 mm; head width 21.8 mm. Distribution.—This species is known from several localities in the mountain ranges north of the Balsas Basin (Fig. 3), from the Sierra de

Chichinautzin, Morelos, to TemascaltepecReal de Arriba, Mexico, from approximately 1850–2600 m above sea level (see Appendix I for specific localities). The species of Abronia from the highlands of Guerrero was originally allocated to A. deppii (Bocourt, 1878). However, in view of our current understanding of the type material of A. deppii and the variation exhibited by this species, we propose that the populations from Guerrero be known as: Abronia martindelcampoi sp. nov. Abronia deppii—Smith and Taylor, 1950; Tihen, 1954; Bogert and Porter, 1967 [also spelled as deppei]; Good and Shwenk, 1985; Good, 1988 Abronia deppi—Davis and Dixon, 1961 Holotype.—Adult male, Museo de Zoologia, Facultad de Ciencias UNAM (MZFC 02778). Collected in Mexico, Guerrero, Chilpancingo, Omiltemi, Orilla Norte in oak forest at 2250 m on 7 April 1985 by Antonio Mun˜oz Alonso (Field number AMA-033). Paratypes.—Mexico, Guerrero, Chilpancingo, Parque Ecolo´gico Estatal Omiltemi: MZFC 02774–77, 02779–82, 02875; Atoyac,



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FIG. 4.—Abronia martindelcampoi, paratype from 0.8–1.6 km NE Puerto del Gallo, Guerrero, Mexico; now cataloged as UTA-R 12136. Photo by W. W. Lamar, courtesy of J. A. Campbell.

Carrizal de Bravos (MVZ 164922); Atoyac, Carrizal de Bravos, 60 km by road (LACM 109262); Atoyac, Carrizal de Bravos, 1 km SW (MVZ 110941, 164922, 134109); Vicinity of Chilpancingo (AMNH 72543, MVZ 45005); Omiltemi (BMNH 1913.7.19.98, 1913.7.19.99, 1913.7.19.100, 1913.7.19.101; MVZ 57173, UTA-R 4451, 5645, 5646, 5653); Puerto del Gallo (UTA-R 4151); 0.8–1.6 km NE Puerto del Gallo (UTA-R 12136); San Miguel Totolapan, 1 km W of Toro Muerto (MZFC 765). Diagnosis.—This species has (1) supraauricular scales protuberant and knoblike, with no indication of spines; (2) supranasals not expanded and not in contact at dorsal midline; (3) frontonasal present and usually not in contact with frontal; (4) posterior internasals almost twice as long as anterior internasals (1.6–2.0); (5) no canthals; (6) three anterior temporals; (7) parietal not in contact with median supraoculars; (8) a single occipital; (9) posterior head scales strongly convex and knoblike; (10) anterior supercilliary not in contact with cantholoreal; (11) posterior suboccular in broad contact with lower primary temporal; (12) preauricular scales notoriously enlarged and in one row; (13) postmentals usually two (only 1 out of 14 had one); (14) posterior infralabials usually elongated; (15) six

longitudinal nuchal scales; (16) dorsal scales in 23–27 transverse rows; (17) dorsal scales in 10– 12 longitudinal scale rows; (18) ventral scales in 12–14 longitudinal scale rows; and (19) 5–6 brown dorsal markings in adults (Fig. 4). Abronia martindelcampoi differs from A. deppii in having (1) well developed enlarged knob-like posterior head scales (Fig. 2); (2) lower anterior temporal fused with penultimate supralabial; (3) no azygous scale between interparietal and interoccipital; (4) preauricular scales notoriously enlarged; (5) head width in adult males .22 mm and .18 mm in females; (6) ratio of head width to head length .0.75 in adult males; (7) number of scale whorls on unregenerated tail 77–86 ( x 5 80.7); (8) number of infralabial scales 8–10 (only one had seven); (9) number of longitudinal dorsal scale rows 10–12; (10) dorsal ground color grayish to pale olive green in life (grayish in preservative) with dark brown markings which, if confluent on the vertebral region, may be asymmetrical; (11) dorsum of head pale brown to olive green, usually unblotched, individual scales with fine dark vermiculations, some specimens with large dark area on top of head, more, rarely a few, black individual scales (including both upper anterior temporals); (12) ventral region immaculate cream colored

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in adults, rarely with faint suggestion of yellowish orange (see below); and (13) upper lips mostly unmarked. It differs from other species of the subgenus Abronia (sensu Campbell and Frost, 1993) in having oblique longitudinal rows of dorsal scales, where fuscolabialis, taeniata, and graminea have parallel longitudinal scale rows and taeniata and graminea do not; in most specimens, one lower temporal contacts the postoccular series and a single occipital, where mixteca and oaxacae have three occipitals and two primary temporals contacting postocular series; has body coloration that is not uniformly green like graminea; has six nuchals, where graminea has four (only juveniles of graminea have bands on the dorsum of the body). Description of holotype.—Measurements (all measurements in mm): Snout–vent length 115; head length (from tip of snout to posteriormost edge of head) 31.5; head width 25.2; head depth 14.7; tail length 163. The supra-auricular scales are knob-like and without any indication of spines; small supranasals and not in contact at dorsal midline; romboid frontonasal, separated from frontal; posterior internasals 1.6 times larger than anterior internasals; cantholoreal evident from above the head and large; anterior supercilliary not in contact with cantholoreal; and a single occipital. On each side of the head are 10 supralabials; three anterior temporals; three posterior temporals; three scales between occipital and first nuchal row; five median supraoculars; one temporal contacting the postocular; one subocular; parietal not in contact with median supraoculars; preauricular scales in one row and notoriously enlarged; two postmentals; and posterior infralabials enlarged. The following combinations of scales all fail to make contact with one another: frontonasal-frontal, temporal-subocular, superciliary-prefrontal, and superciliary-cantholoreal. The postparietals are not reduced; the lateral neck scales are enlarged. The specimen has 27 transverse dorsal scale rows; 12 longitudinal dorsal scale rows; 2 postmentals; 6 nuchal scales across neck; 34 transverse ventral rows; 14 longitudinal ventral scale rows; 80 scale whorls on unregenerated tail; 6 scales between hind limbs; 20 lamellae under fourth toe. A weak lateral fold is present; the lateral dorsal scales are arranged in oblique


rows, and no osteoderms are present beneath the scales on the trunk. Coloration.—Head olive green with each dorsal scale having fine dark vermiculations, with only a faint indication of dark blotches on the first nuchal row; supralabial region slightly paler than dorsum of the head; black markings behind ear openings. Ground color of dorsum pale olive green with some brownish areas, with six narrow, dark crossbands, each about one scale wide (Fig. 4). Dorsolateral scales with white posterior edge; tail inconspicuously marked (13 bands). The ventral region is immaculate and cream colored (in preservative), and the gular region is pale. Color variation.—In some specimens the dorsal crossbands are interrupted middorsally and are asymmetrical. The ground color may be olivaceous to pale brown, and the tail blotches may be relatively conspicuous (in preservative); the dorsolateral scales may lack white posterior edges, while in other specimens white-tipped scales may cover the dorsum. The ventral coloration in some specimens is pale orange, but usually is not intense (the only specimen that apparently had bright orange ventral coloration was MVZ 45005). The color described by Good (1988) in his account of A. deppii is based on A. martindelcampoi. Comparisons.—For comparisons with other species of the genus, see table 1 in Good and Schwenk (1985) under A. deppii. Distribution.—This species is known only from the Sierra Madre del Sur (Fig. 3), Mexican State of Guerrero from 2100 to 2600 m above sea level (see Appendix I for specific localities). There is one specimen (BMNH 1913.7.19.102) from Amula, Guerrero, tentatively assigned to this taxon. Etymology.—This species is named after the late Rafael Martı´n del Campo y Sa´nchez in recognition of his contributions to the development of modern Mexican herpetology, and particularly for his knowledge of gerrhonotine lizards. Miscellaneous natural history notes.—During field work at Omiltemi, Guerrero, we collected an Ophryacus undulatus whose stomach contained an A. martindelcampoi. A courting pair of A. martindelcampoi was collected in pine-oak forest on 4 August 1985. A female collected May 1989 by Jack Sites at Omiltemi gave birth to six young, and another



specimen (UTA-R 5653), collected August 1975 by J. A. Campbell, gave birth to two young in December 1975. Davis and Dixon (1961) reported that a female taken in June contained enlarged ovarian follicles. The defensive display consists of opening and expanding the lower jaw and biting. Results of an analysis of the gonads of the female suggest that the species is a fall breeder. In a recent visit to Omiltemi, the type locality of A. martindelcampoi, we found that there is no longer a protected state park; instead, logging and other human activities were taking place, even though the city of Chilpancingo (capital of Guerrero) receives high quality water from the mountains around Omiltemi. This new situation may put populations of A. martindelcampoi and the biota of that portion of the Sierra Madre del Sur at risk. RESUMEN Se describe una especie nueva de lagartija del ge´nero Abronia proveniente de la Sierra Madre del Sur, Guerrero, Me´xico. Esta especie fue confundida en el pasado con A. deppii. El hallazgo de especı´menes provenientes de la parte norte de la cuenca del Balsas y su estudio morfolo´gico, indican que las poblaciones de Abronia que habitan las tierras altas al sur de la cuenca del Balsas deben ser reconocidas como un taxo´n diferente. A. martindelcampoi sp. nov. se distingue de A. deppii por presentar las escamas posteriores de la cabeza alargadas y redondeadas a manera de perilla; por la ausencia de una escama a´ziga entre la interparietal y la interoccipital; por presentar un mayor nu´mero de hileras de escamas en la cola, no regenerada; mayor nu´mero de escamas infralabiales y por las diferencias en coloracio´n dorsal y ventral. Se agregan comentarios sobre la historia natural de A. martindelcampoi. Acknowledgments.—We wish to thank A. Nieto, J. W. Sites, J. Johnson, D. A. Good, and D. Frost for help in preparing this manuscript; J. A. Campbell for his review of the manuscript; G. Zug for the photographic material of the type and paratype of Abronia deppii from the Berlin Museum and for arranging the loan of those specimens; C. J. Cole (AMNH), D. B. Wake (UC-MVZ), J. Wright and R. L. Bezy (LACM), E. Godinez (ENEPI), and R. CastroFranco (EBUM) for checking materials in their collections; and A. Mun˜oz, E. Herna´ndez, A. Herna´ndez, and J. Jua´rez for assistance in the field. Part of the field work in Guerrero was possible thanks to the economic support of the Government of the State of Guerrero and the Facul-

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tad de Ciencias, UNAM. Part of this project was financed by a grant from DGAPA-UNAM project IN-201789 to J. Llorente and O. Flores-Villela. This paper is based in part upon work supported by the NSF under grant No. DEB-0102383.

LITERATURE CITED BOCOURT, M. F. 1878. Etudes sur les Reptiles. Mission scientifique au Mexique et dans l’Ame´rique Centrale. Recherches Zoologiques:325–327. BOGERT, C. M., AND A. P. PORTER. 1967. A new species of Abronia (Sauria, Anguidae) from the Sierra Madre del Sur of Oaxaca, Mexico. American Museum Novitates (2279):1–21. CAMPBELL, J. A. 1982. A new species of Abronia (Sauria, Anguidae) from the Sierra Jua´rez, Oaxaca, Me´xico. Herpetologica 38:355–361. CAMPBELL, J. A., AND D. R. FROST. 1993. Anguid lizards of the genus Abronia: revisionary notes, descriptions of four new species, a phylogenetic analysis, and key. Bulletin of the American Museum of Natural History (216):1–121. CAMPBELL, J. A., M. SASSA, M. ACEVEDO, AND J. R. MENDELSON III. 1998. A new species of Abronia (Squamata: Anguidae) from the high Cuchumatanes of Guatemala. Herpetologica 54:221–234. CASTRO-FRANCO, R. 1987. New records of reptiles from the Mexican state of Morelos. Bulletin of the Chicago Herpetological Society 22:69–70. CHIPPINDALE, P. T., L. K. AMMERMAN, AND J. A. CAMPBELL. 1998. Molecular approaches to the phylogeny of Abronia (Anguidae: Gerrhonotinae), with emphasis on relationships in subgenus Auriculabronia. Copeia 1998:883–892. DAVIS, W. B., AND J. R. DIXON. 1961. Reptiles (exclusive of snakes) of the Chilpancingo region, Mexico. Proceedings of the Biological Society of Washington 74:37–56. FLORES-VILLELA, O. 1993. Herpetofauna Mexicana. Special Publication Carnegie Museum of Natural History (17):1–73. FLORES-VILLELA, O., AND E. HERNANDEZ-GARCIA. 1989. New state records from northern Guerrero, Mexico. Herpetological Review 20:15–16. FLORES-VILLELA, O., AND A. MUN˜OZ-ALONSO. 1993. Anfibios y reptiles. Pp. 411–442. In I. Luna and J. Llorente (Eds.), Historia Natural del Parque Ecolo´gico Estatal Omiltemi, Chilpancingo, Guerrero, Me´xico. Comisio´n Nacional para el Conocimiento y Uso de la BiodiversidadUniversidad Nacional Auto´noma de Me´xico, Me´xico. GOOD, D. A. 1988. Phylogenetic relationships among gerrhonotine lizards: an analysis of external morphology. University of California Publications in Zoology 121:1– 139. GOOD, D. A., AND K. SCHWENK. 1985. A new species of Abronia (Lacertilia: Anguidae) from Oaxaca, Mexico. Copeia 1985:135–141. GRAY, J. E. 1838. Catalogue of the slender-tongued saurians, with descriptions of many new genera and species. Annals and Magazine of Natural History 2:287– 293. GU¨NTHER, A. C. L. G. 1902. Biologia Centrali Americana, Reptilia and Batrachia. Porter, London, England. SA´NCHEZ-HERRERA, O., AND W. LO´PEZ-FORMENT, C. 1980. The lizard Abronia deppei (Sauria: Anguidae) in the state

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of Mexico, with the restriction of its type locality. Bulletin of the Maryland Herpetological Society 16:83–87. SIBLEY, C. G., AND J. DAVIS. 1946. Real de Arriba, Me´xico, as a Deppe locality. Condor 48:279. SMITH, H. M., AND E. H. TAYLOR. 1950. An annotated checklist and key to the reptiles of Mexico, exclusive of the snakes. United States National Museum Bulletin 199:1–253. STRESSEMAN, E. 1954. Ferdinand Deppe’s travels in Mexico, 1824–1829. Condor 56:86–92. TAYLOR, E. H. 1969. Wiegmann and the herpetology of Mexico. Pp. iii–vi. In K. Adler (Ed.), Herpetologia Mexicana by A. F. A. Wiegmann. Facsimile Reprints in Herpetology, No. 23, Society for the Study of Amphibians and Reptiles, Notre Dame, Indiana, U.S.A. TIHEN, J. A. 1949. The genera of gerrhonotine lizards. American Midland Naturalist 41:580–601. ———. 1954. Gerrhonotine lizards recently added to the American Museum Collection, with further revisions of the genus Abronia. American Museum Novitates (1687):1–26. WIEGMANN, A. F. A. 1828. Beitra¨ge zur Amphibien-Kunde. Isis von Oken 21:364–383. ———. 1834. Herpetologia Mexicana seu descriptio amphibiorum Novae Hispaniae. Pars prima. Saurorum species. Facsimile Reprints in Herpetology No. 23, Society for the Study of Amphibians and Reptiles, Notre Dame, Indiana, U.S.A. Accepted: 14 March 2003 Associate Editor: Stephen Tilley

APPENDIX I Specimens Examined Abronia deppii.—MEXICO: MEXICO 1 km SSW from Valle de Bravo, 2000 m (UTA-R 31634, formerly MZFC 764); Valle de Bravo (MZFC 4307); Avandaro, approxi-


mately 2050 m (MZFC 2015); Ixtapan de la Sal, Km 9 on the road to Zacualpan-Mamatla, 1900 m (ENEPI 963); Km 9 on road from Zacualpan to Mamatla, 40 km from Ixtapan de la Sal (MZFC 6294, formerly ENEPI 1001); GUERRERO: Ixcateopan de Cuahute´moc, Km 28 on road TaxcoIxcateopan, 2100 m (MZFC 3992); Pedro Ascencio de Alquiciras, Cruz Alta, 2560 m (MZFC 3991); Tetipac, Arroyo las Damas, 1850 m (MZFC 3993); MORELOS: Derrame del Chichinautzin, 2250 m (MZFC 2015); Huitzilac, N of CEBETIS, 2600 m (EBUM 0425). Photographic material.—MEXICO: no specific locality (ZMB 1149, type); no specific locality (ZMB 1150, paratype). Abronia martindelcampoi.—MEXICO: GUERRERO: Amula (BMNH 1913.7.19.102); Atoyac, Carrizal de Bravos, 2407.9 m (UC-MVZ 164922); Carrizal de Bravos, 60.3 km SW by road (LACM 109262); Carrizal de Bravos, 0.96 km SW, 2200 m (UC-MVZ 134109); Chilpancingo Omiltemi, 2250 m (MZFC 2781, 2784, 2875; ENEPI 964; UC-MVZ 57173; UTA-R 4451, 5653); Omiltemi, 2436 m pine-oak forest (UTA-R 5645–46); Omiltemi, 2 km SE, 2250 m (MZFC 2774, 2775, 2777); Omiltemi, N of town, 2250 m (MZFC 2776, 2778 holotype); Omiltemi, 500 m E of town, 2250 m (MZFC 2779); Omiltemi, 2 km E, Barranca de Potrerillos, 2200 m (MZFC 2780); Omiltemi NW of town, 2250 m (MZFC 2782); Omiltemi SW side of town, 2250 m (MZFC 2783); Omiltemi (BMNH 1913.7.19.98–101); San Miguel Totolapan, 1 km W of Toro Muerto, 2600 m (MZFC 765); Filo de Caballo?, 2000 m (MZFC 766); Vicinity of Chilpancingo (AMNH 72543; UC-MVZ 45005); Puerto del Gallo (UTA-R 4151); 0.8–1.6 km NE Puerto del Gallo, 2560–2804 m (UTA-R 12136). Other materials examined.—Abronia martindelcampoi MEXICO: GUERRERO: Atoyac, Carrizal de Bravos, 1 km W by road (LACM 127415); Carrizal de Bravos, 0.96 km SW, 2200 m (MVZ 110941); Chilpancingo, Omiltemi 2286 m elevation (MVZ 57163).