You are on page 1of 9

Graellsia, 58(1): 49-57 (2002

)

TAXONOMY AND DISTRIBUTION OF THE SOUTH AMERICAN TOAD BUFO POEPPIGII TSCHUDI, 1845 (AMPHIBIA, ANURA, BUFONIDAE)

I. De la Riva *

ABSTRACT The taxonomic status of the Andean toad Bufo poeppigii has been controversial since its description by Tschudi in the 19th Century, because of the similar appearance of the species with respect to Bufo marinus, and the fact that both species may occur together in some localities at the foot of the Andes. Bufo poeppigii is a valid species occurring on the Amazonian slopes of the Andes, at least from central Bolivia to northern Peru. It differs from B. marinus mainly by a less marked sexual dimorphism in size, smaller size, rugose skin always present in males, distribution of nuptial excrescences, and some features of the parotoid glands. The variability of B. marinus encompasses that of B. poeppigii; thus, in some cases it is difficult to identify some specimens. Key words: Anura, Bufonidae, Bufo poeppigii, B. marinus, B. arenarum, taxonomy, distribution, South America RESUMEN Taxonomía y distribución del sapo sudamericano Bufo poeppigii Tschudi (Amphibia: Anura: Bufonidae) La posición taxonómica del sapo andino Bufo poeppigii ha sido objeto de controversia desde su descripción por Tschudi en el siglo XIX. Ello se ha debido a su parecido con Bufo marinus y a que ambas especies pueden encontrarse juntas en algunas localidades del piedemonte andino. No obstante, Bufo poeppigii es una especie válida que habita en las laderas amazónicas andinas, al menos desde el centro de Bolivia hasta el norte de Perú. Difiere de B. marinus por tener dimorfismo sexual en tamaño menos marcado, menor talla, piel siempre rugosa en los machos, la distribución de las callosidades nupciales y algunos rasgos de las glándulas parotoides. La variabilidad de B. marinus abarca la de B. poeppigii, de modo que a veces es difícil identificar ciertos ejemplares. Palabras clave: Anura, Bufonidae, Bufo poeppigii, B. marinus, B. arenarum, taxonomía, distribución, Sudamérica

Introduction

Among the rich anuran fauna of the Andes are several species groups within the toad genus, Bufo. Whereas the Bufo veraguensis and Bufo spinulosus groups are endemic to the Andes, other, primarily

lowland groups have some representatives on the Andean slopes and/or at the foot of the mountains (Duellman & Schulte, 1992). Examples of these groups are the B. “margaritifer” complex (Hass et al., 1995) and the B. marinus group. Among members of the B. marinus group, several taxa have been

*

Departamento de Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales, CSIC. C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain

” and recognized the subspecies B. Calilegua. 1959. and 6) Bufo gallardoi Carrizo. Ruiz-Carranza et al.. Cei. 1972.g. Brazil). Bufo poeppigii has more frequently been considered as a valid species (e. Vellard. poeppigii “as well as different intermediate forms were completely mixed in Peruvian populations with no ecological differences between them. 1996. poeppigii has been changing continuously. Blair. either as a full species or as a subspecies of B. and Bolivia. inasmuch as these localities are in the mountains around Monobamba.. and topotypic material representing this taxon from the mountains of Monobamba. 1972. marinus (e. Rodríguez et al. 1996. B. 1979. Duellman & Schulte. the taxonomic status of B.. poeppigii is Moyobamba. Gorham. unanimously considered a synonym of B.. 1993) most continued recognizing B.g. MHNN 90. and reported from the eastern Andean slopes of Ecuador. was based. those from Tsioventini and Río Perene. 1979. a species occurring primarily in the lowlands of southern South America. 1990). marinus. a species closely related to B. that reaches the foot of the Andes in some parts of Bolivia (De la Riva.. The purpose of this paper is to discuss on the specific status of B. Furthermore.. reported or described explicitly from the Andean region of Peru and Bolivia. poeppigii are distinct species that are sympatric at some localities on the lower slopes. 1991.. Duellman & Toft.50 DE LA RIVA. 1999. Minas Gerais. 1925 (type locality: Belo Horizonte. 1845. This impression was reinforced by further examination of many Andean specimens of Bufo in museum collections (Appendix I). however. gallardoi in Bolivia are doubtful (Harvey. Peru and Bolivia (Frost. poeppigii. Glaw et al. poeppigii. Emmons. 1996.. 1867 (type locality: Río Grande do Sul. De la Riva. 2000 m)].. 1979. Cei. Henle. This assumption is unwarranted. Guttman. These are. 1972. marinus. poeppigii. 75º11’W. Paraguay. 1992. De la Riva et al. 1985). De la Riva et al. marinus and known from Argentina. The specimens from San Ramón and Yaupi (and. 1992. 1988. Bogart. poeppigii was made by Henle (1985). Lutz. Cadle & Patton. Mertens. arenarum and B. Since then. 1998. the taxonomic status of this species has been in a state of flux since its description. a widespread species in the Neotropics (it has also been introduced elsewhere) that reaches the Andean slopes and foothills from Venezuela to Bolivia (De la Riva et al. B. to a minor extent. Zug & Zug. Hass et al.g. but Nieden (1923) resurrected B. molitor in the synonymy of B. marinus Linnaeus. but also in the Andes of Central Bolivia up to 2600 m (Frost. 1993. 2) Bufo poeppigii Tschudi. Bertini & Cei. 1972. 1992. Frost. Taxonomic history of Bufo poeppigii de Monobamba” [a locality in the departament of Junín. Vellard. Jujuy. 1990.. 1995. Department of San Martín. described from Andean Argentina (type locality: Monolito. Köhler. 1961. Most authors have not provided justification on which their placement of B. 2000) and beyond the scope of this paper. 1997. 1700 m. ca. marinus poeppigii. poeppigii are superficially similar in appearance. marinus. marinus and B. 5) Bufo paracnemis A. poeppigii and provide remarks on the diagnosis. Brazil). Uruguay. 1941 (type locality: Roque. 2000). Duellman.. Almendariz. 1991. molitor to be a junior synonym. described from the Andes of Peru (type locality: “Montañas de Monobamba”). Duellman. Intensive field work in the Andean region of Bolivia led me to recognize that B. see Appendix I) may be considered as topotypic. marinus and B. 35 km SE Moyobamba. 1958. respectively from “Peru” and “Montañas I examined the holotype of B. 1983. distribution. Taxonomic status and diagnosis of Bufo poeppigii Tschudi (1845) described Bufo molitor and B. 2000. 2000). 1974). Although this criterion was followed by several authors (e. Roux (1907) considered both names as synonyms of B. 1758) [type locality: “America”. southeastern and central Brazil. poeppigii as a valid species and considered B. Departament of San Martín. Duellman. Peru. and ecology of the species. Peru). the species-group allocation and putative distribution of B. 1985). Morales. He stated that B. Peru. poeppigii as a full species (e. paracnemis does not present any problem regarding its identification or taxonomic status. 3) Bufo arenarum Hensel. all subsequent authors have placed B.. 4) Bufo marinus andinensis Melin. 1952.g. Argentina). restricted by Müller & Hellmich (1936) to Surinam]. being often considered as a subspecies of B. marinus. because there is no reason to suppose a misspelling of the type locality in Tschudi’s description. Harding. De la Riva et al. 1985) than as a synonym of B. 1995. 1955. by chronological order of description: 1) Bufo marinus (Linnaeus. I. The only comprehensive attempt to elucidate the taxonomic status of B.77. Peru (11º25’S. Henle (1985) assumed that the type locality of B. there are at least three different Peruvian localities called .

where the tubercles are more closely spaced (texture of the skin variable in amount. Fig. with many tubercles of similar size. II. marinus occurring in the eastern slopes of the Andes. and III (nuptial excrescences normally present on Fig. arenarum (KU 183186). 35 km SE Moyobamba. poeppigii. poeppigii (KU 183227). whereas B. often hypertrophied. skin texture. protuberant.9 mm). poeppigii is a juvenile or small subadult (SVL= 52. Peru. B. after comparing it with the syntypes of B. South America. marinus (KU 196665) y B. In many cases it is not easy to distinguish B. 1. marinus (Figs. decalcified. marinus from other regions (including Middle America. and dorsal and ventral patterns (personal observation. marinus. poeppigii shows little variation throughout its geographical range. marinus in parentheses): (1) males always with rugose skin. poeppigii. poeppigii. poeppigii is the name-bearing type of the species closely related to. marinus are conspecific. faded. poeppigii from B. Based on the poor condition of the holotype of B. marinus are (condition in B. marinus. I also examined the two syntypes of B.TAXONOMY AND DISTRIBUTION OF BUFO POEPPIGII 51 Moyobamba. West Indies. To the best of my knowledge. Fig. This is the only possible argument to assume that the holotype of B. B. the main differences between B. but different from B. Because the holotype of B. (2) parotoids subtriangular. poeppigii (KU 183227). a higher number of tubercles in the sacral region. each one situated in a different department (see Stephens & Traylor. marinus poeppigii) or is a juvenile of B. and Fiji). molitor and with a large sample of B.— Cabezas y glándulas parotoides de B. poeppigii and B. The latter is quite variable in size. poeppigii (or B. molitor (MHNN 90. arenarum (KU 183186) (de arriba a abajo). 1979). Australia. 1-3). and disposition of tubercles). poeppigii is the available name for this different taxon. In order to determine that no other described taxa correspond to what we recognize today as B. and B.— From top to bottom. development and shape of the parotoid glands. . their borders poorly defined. marinus (KU 196665). poeppigii and B. it is not surprising that Henle (1985). from Roque. Departamento San Martín. no specimens identified as B. 1. 2). 1983).87) and the two syntypes of B. 3). see also Zug & Zug. each covered by many keratinized spicules (Fig. marinus andinensis Melin (NHMG 457). marinus have been collected at high elevations in the area of the type locality of B. concluded that B. I agree with former authors in considering both nominal taxa as junior synonyms of B. flattened. never hypertrophied (parotoids subtriangular. and softened. heads and parotoid glands of B. I conclude that B. Thus. it is impossible to assess with certainty whether it actually represents what has been commonly recognized as B. (3) males bearing extensive nuptial excrescences on Fingers I. Keeping this in mind. size.

1972). One of the most consistent differences between the two species is overall size and degree of sexual dimorphism in size and skin texture. Perú (KU diapositiva # 10007). however. poeppigii there is no such marked sexual dimorphism in size. 2. well defined blotches. the largest female examined (AMNH 137401) is 132. poeppigii is a consistent character. . 1979)]. 1970. as pointed out by Vellard (1959). unless large samples are available. Peru (KU slide # 10007). if present on the other fingers. it becomes obvious that the presence of heavy skin rugosity in males of B. females of B. it seems that conclusive discrimination based on measurements is not possible. 1985. B. Duellman & Trueb. whereas females have a fairly smooth skin. when they are.— Bufo poeppigii (KU 183237) from Coroico. not sharply defined blotches. one female of B. The degree of difference in skin texture between males and females of B. In contrast. Males always have a highly rugose skin. 3. Fig.0 mm in SVL. poeppigii [although populations outside Neotropics may be of moderate size (Henle. and Köhler (2000). Bolivia (KU slide # 6440).3 mm in SVL. as was noted by Vellard (1959) for the region of Tingo María (Department of Huánuco. marinus. Peru). hybrid individuals can complicate even further the allocation of particular individuals to one or the other species. marinus across its entire range encompasses that of B. Finger I only and. poorly developed). 2. Fig.. spinulosus. marinus reach sexual maturity at a size of 90-100 mm. skin texture itself is not a reliable character to distinguish the two species. Cei (1958. especially in females and immature individuals). The range of variation in B. the calls of the two species are generally similar. Given the variation in B. Zug & Zug (1979) indicated that in Panama. marinus. attaining a snout-vent length (SVL) of 240 mm (Reed & Borowsky. Although distinguishing the two species should not be problematic when large samples are considered. Fig. females often having small dark.8 mm in SVL. 1979). As was demonstrated by Blair (1972) and Bogart (1972) by means of karyotype studies. and the largest male (one in the same lot as female USNM 193887) is 112. I tried unsuccessfully to find any ratio in body proportions to distinguish the two species properly.— Bufo poeppigii (KU 183237) de Coroico. 1996). at localities where both species occur it may be difficult to assign a particular specimen to one or the other species. and perhaps the call alone does not guarantee reproductive isolation (De la Riva et al. In B. (5) venter always uniformly pale (often marbled with gray). poeppigii. marinus are closely related species and may hybridize. Zug & Zug. poeppigii differs from that of B. immature individuals mostly gray with small dark blotches surrounding the dorsal tubercles (dorsum brown or tan. Obviously. (4) males with dorsum brown or tan. Bolivia (KU diapositiva # 6440). usually lacking pattern.— Bufo marinus (not collected) from Cuzco Amazónico. 1986). poeppigii with mature ovarian eggs (AMNH 72129) is only 63. I. and females are considerably larger than males (Zug & Zug. poeppigii is similar to that of the highland species B. Bufo marinus is by far a much larger species than B. The advertisement call of B.— Bufo marinus (ejemplar no colectado) del Cuzco Amazónico. Fig. poeppigii and B. usually with large black. 3.52 DE LA RIVA.

arenarum. In B. 800-1670 m.. 4. (1993) reported B. not to B.TAXONOMY AND DISTRIBUTION OF BUFO POEPPIGII 53 Fig. to the best of my knowledge. Nieden (1923) said that the species occurs in Colombia and Ecuador. poeppigii in the Peruvian departments of Huánuco. arenarum. Bolivia. Bufo poeppigii reaches the lowlands in areas adjacent to the foot of the mountains. Bufo arenarum and B. However. (1996). marinus poeppigii refer to B.. between elevations of 260-1829 m (Fig. spinulosus. are those published by Köhler (2000: Plate I-f) and De la Riva et al. Bolivia). poeppigii.. Peru. poeppigii provided by Vellard (1959: 36) with reservation. arenarum (Figs. B. Ecuador. 2000). Distribution and ecology The distribution of B. The report by Andersson (1938) of B. Bufo poeppigii occurs at least from the region of the upper Huallaga River (Department of San Martín. as well as . marinus. As was pointed out by Cei (1958. I misidentified as B. B. it is necessary to consider the list of localities for B. . Most Peruvian localities provided by Henle (1992) for B. and the posterior end of the parotoid is followed by a series of smaller glands. based on Frost’s information. Bolivia (KU diapositiva # 6432). Peru) to Amboró National Park and Buenavista area (Department of Santa Cruz. 1600 m) most probably corresponds to B. 5). Thus. poeppigii (De la Riva. poeppigii. Chacoan montane forests. 1990). 4. Loreto. 1 and 4). commonly they are in contact with the orbitotympanic crest or invest the postorbital crest. B. San Martín. In Bolivia. inter-Andean valleys and montane forests at least to 2600 m (De la Riva et al. Cochran & Goin (1970) did not report B. marinus poeppigii crosses the Huancabamba Depression in northern Peru and reaches the Pacific coast from Olmos (Departament of Lambayeque) to the Ecuadorian border. spinulosus at Mojos (La Paz. marinus) there is always a separate short postorbital crest (Fig. RuizCarranza et al. but it is not found far from the Andes. In summary. During the breeding season. and Bolivia. as stated above. poeppigii have a dorsal skin as rugose as that of the males of B. 1). Vellard (1959) stated that B. and more recently. New Grenada” (Colombia)] as a junior synonym of B. males of B. arenarum the first specimens of B. poeppigii that I collected at Villa Tunari and those from Caranda in the MNK [De la Riva. poeppigii is quite similar to B.— Bufo arenarum (KU 183186) from Aiquile. and the degree of sexual dimorphism in this character is similar in both species. poeppigii also is the subject of some controversy. a logical statement upon considering B. In B. poeppigii in her list of Ecuadorian amphibians. poeppigii. Junín. Following Henle’s results. From B.. poeppigii from Colombia and Miyata (1982) and Coloma (1991) did not report it in Ecuador. (2000: 80)]. the occurrence of the species in these two countries has not been documented. However. Fig. and Fig. the parotoids are long and enlarged anteriorly.— Bufo arenarum (KU 183186) de Aiquile. Bolivia (KU slide # 6432). marinus) in the Huancabamba Depression and I have examined no specimens from the Pacific versant of Peru and Ecuador that are referrable to B. On the other hand. in addition to that photograph and the one published herein. poeppigii do not occur sympatrically.”. 1990: 265. Duellman & Wild (1993) did not report the species (nor B. Madre de Dios. 7. included the species in the list of amphibians of Colombia but they explicitly stated that it is unknown in the country. Almendariz (1991) included B. Rodríguez et al. poeppigii the parotoids are subtriangular and (as in B. males congregate at ephemeral ponds and puddles (even in the middle of villages). although Frost (1985: 57) gave as distribution of the species “Cloud forests of the Andean slopes of Colombia. 1868 [type locality: “Valley of Quito and Carthagena. this is the first published photograph of the species. 1972). and Ucayali. arenarum occurs from Cochabamba to Tarija departments in Chacoan lowland forests. andianus Cope. poeppigii differs mainly by its slightly smaller size and by the shape of the parotoids.

1600 m in Venezuela (Rivero. Where B. comm.— Distribución de Bufo poeppigii en Perú y Bolivia.— Distribution of Bufo poeppigii in Peru and Bolivia.54 DE LA RIVA. poeppigii has a distribution mostly parapatric with that of B. marinus USNM 166768. why not consider it as a mere subspecies of B. their general pattern and morphology are also As has been explained. There is no known case of a widely distributed Neotropical anuran with a subspecies on the Andean slopes. marinus. coincident with B. Discussion 0 200 400 km 600 Area above 3000 m Fig. poeppigii as a valid species. there are some localities at which both species occur. marinus. . Bufo marinus typically is a lowland. marinus group occurring on the eastern Andean slopes of Peru and Bolivia. it could actually be a juvenile B. are herein tentatively considered as B. I adopt here the evolutionary species concept as explained by Wiley (1978) and expanded by Frost & Hillis (1990). Bolivia). Ecuador) (KU 147082). 5. marinus primarily inhabiting cloud forest in the central Andes. poeppigii is a poorly-preserved juvenile whose type locality is relatively imprecise. However. marinus seems to reach higher elevations [for example.). Because there is no unequivocal solution to this problem. We still know so little about tropical amphibians that the use of subspecies should be precluded. it is necessary to reiterate that because the holotype of B. namely Bulo-Bulo (Department of Cochabamba. From this point of view. Bolivia) and the region of Tingo María (Huánuco. 5. If so. Fig. poeppigii was no more than a regional race of B. there is no available name for the species considered herein as B. easier to diagnose than B. marinus) uniformly distributed across a defined area. the species seldom enters the closed rainforest. and that they might hybridize. poeppigii. marinus with a smaller size as a response to the environment at higher elevations (which might represent suboptimal conditions). In considering B. Further research in northern Peru and southern Ecuador is needed in order to ascertain whether or not B. However. its own evolutionary trend. 1961)]. anthropophyllous toad that colonizes new areas following roads and river banks and it is often found in villages and human settlements. however. Generally both species occur in open habitats and reproduce in similar places but usually they are not found together. marinus from a low locality in this area (B. in ponds isolated in river beds. Puerto Linares (La Paz. Peru) (McDiarmid. that they are sometimes barely distinguishable. Bolivia). marinus adapted for living at higher altitudes? Bufo poeppigii is not simply a set of populations of B. the binomial Bufo poeppigii is the name traditionally linked to the smaller species of the B. populations of B. poeppigii does not occur. so there is no clinal variation in size. I recommend that the name B. The two species might occur in the zone of the upper Tambopata River (Madre de Dios. a pattern that could be expected in other species if B. B. Given that B. pers. marinus. poeppigii actually occurs in these regions. among other reasons. poeppigii continue to be applied to the small species of Bufo closely related to B. poeppigii. one from Moyobamba (Department of San Martín. B. poeppigii from localities at 260 m are indistinguishable from those at 1800 m. I. Henle’s (1985) conclusion about the synonymy of the two nominal taxa was correct. poeppigii is a single lineage with reproductive cohesion. Rurrenabaque (Beni. and representing a homogeneous entity (in fact. Peru). Two specimens. Peru) (KU 211746) and another from Misión Bomboiza (Province of Morona Santiago. is a voucher for the occurrence of the species at low elevations in the region). If this is true.

Austin & London: 171-195. Amphibian species of the world: Additions and corrections. 35(1): 60-70. much field work is needed. 1990. L. I. 1972.). 1992(1): 162-172.. United States National Museum Bulletin. W. Kansas: 371-459. F. E. Bufo of South America. Batrachians from Burma collected by Dr. 1990). Duellman (ed. L. C. E. reviewed previous drafts of the manuscript. 1979. E. DUELLMAN. R. & WILD. & GOIN. Lawrence. D. E. F. 1972. R. particularly when no obvious morphological differences are present. F. M. Politécnica. Karyotypes. Coloma for their useful comments on the complicated subject of the taxonomic status and distribution of B.& MÁRQUEZ. LÖTTERS. Proceedings of a workshop on Neotropical Distribution Patterns. E. Mendelson III.. J. C. Evidence from hybridization. northern Peru: Systematics. R. and especially J. 1938.. and mammals of the eastern Andean slopes of southern Peru. I. E. distribution. In such a complex fauna as South American amphibians.).. The herpetofauna of Andes: patterns of distribution. and iconography. Anfibios y reptiles. 1988. DUELLMAN. CEI. 1999. BOSCH. 16(3): 89165. COLOMA.. & TRUEB. 13: 1-20. In: W. L. Ministerio de Ciencia y Tecnología. Distribution patterns of Amphibians in South America. Apart from some anonymous reviewers. Blair (ed. 1991. W. and biogeography. Biology of the Amphibians. ANDERSSON. 8(1): 261-319. Tapia and M. 1958. Copeia. R Heyer & P. In: W. W. References ALMENDARIZ. AKNOWLEDGMENTS I thank W. J. and D. 1993. 30: 1-24. COCHRAN. Anfibios del Ecuador: Lista de especies. Description of a new species of Bufo from northern Peru with comments on phenetic groups of South American toads (Anura: Bufonidae).). Anuran amphibians from the Cordillera de Huancabamba. University of Kansas Museum of Natural History Special Publications. literature. 1986. Hamnarland. Austin & London: 196232. E.. Dufour (MHNN) and G. F.. W. Blair (ed. C. R. The South American herpetofauna: Its origin. Reportes técnicos Ecociencia. Anurans from Serranía de Sira. Austin & London: 81-92. 6(2): 59-61.TAXONOMY AND DISTRIBUTION OF BUFO POEPPIGII 55 because we could be underestimating biodiversity. 14: 19-164. A. BERTINI. Baltimore: 255-328.. G. D. 1972. J. DUELLMAN. R. J. A. 1992. Patterns of Distribution of Amphibians. Arkiv för Zoologi. McDiarmid. E. Simmons helped me with the photos in the dark room. In: W. Evolution in the genus Bufo. S. J. Duellman and L. Nilson (NHMG) graciously sent to me the type material under their care. Shiel. Herpetologica. BLAIR. Blair (ed.. In: W. The advertisement calls of two Bolivian species of toads (Anura: Bufonidae: Bufo). and there are not objective reasons to do this (Frost & Hillis. Seroprotein patterns in the Bufo marinus Complex. In: W. Duellman (ed. S. evolution and dispersal.). C. & REICHLE. ecology. Remarks on the geographical distribution and phyletic trends of South American toads. The publication of this paper has been partially funded by project REN2001-1046. Revista Española de Herpetología. E. in many cases. New York. E. Heyer. 2000. E. Malaise and from Bolivia and Ecuador collected by Dr.). diferentiation and present communities. DUELLMAN. 1993. Collection of natural history data and direct experience with the species in the field prove to be invaluable in order to distinguish species. 1996. R. J. taxonomic problems. Evolution in the genus Bufo. KÖHLER. DE LA RIVA. & SCHULTE. E. C.. University of Texas Press. Kizirian provided space and help during my visits to the AMNH. Amazonian Perú: Taxonomy and biogeography. 288: 1-655. Reynolds helped me in many ways during my stay at the Smithsonian Institution. BOGART. 1961. McGraw-Hill Inc. E. Herpetologica. & PATTON. M. R. I.. W. E. CADLE. and R. DE LA RIVA. reptiles. Duellman. J. Ten years of research on Bolivian amphibians: updated checklist. Vanzolini (eds. 21: 1-372. University of Texas Press. University of Texas Press.. In: W. The Herpetological Journal. M. W. 1970. 1979. Bolletino del Museo regionale di Scienze naturali Torino. F. P. CEI. Lista preliminar comentada de los anfibios de Bolivia con datos sobre su distribución. The Pearce-Sellards series. ubicación altitudinal y referencias bibliográficas.. J. DUELLMAN. Montaño (MNK) sent me important data.). improving it considerably. 670 pp. and J. Ford. Frogs of Colombia.. DE LA RIVA. Rio de Janeiro: 225-244. W. J. M. Academia Brasileira de Ciências. DUELLMAN. conclusions obtained only from the study of preserved specimens can be misleading. Myers. University of Kansas Monographies 7. DUELLMAN. Lista de vertebrados del Ecuador. L. origin. The John Hopkins University Press. C. W. Distribution patterns of some amphibians. Spain (Principal Investigator: Ignacio De la Riva).. W.. 46 pp. Museum of Natural History. & TOFT. 1991.. . A Global Perspective. Evolution in the genus Bufo. E. poeppigii.. W. The Museo de Historia Natural “Noel Kempff Mercado” and the Colección Boliviana de Fauna facilitated my field work in Bolivia. & CEI. L. Frost. 17(4): 231-238.

Ornithological Gazetteer of Peru. M. C. 54: 1-70. & TRAYLOR. Recibido. D. 126: 1-207. B. In: T. 2000.. O.. L. A. 1959. R.. Bonner Zoologische Beiträge. MIYATA. 43(1): 79-129. 11: 150-170. Sectio 1..). W. 1990. A... Species concept and practice: Herpetological applications.. V. C. TSCHUDI. Blood proteins. L. 5: 141-161.. University of Texas Press. K. HARVEY. (A).. J. Jena: 257-266. Smithsonian Herpetological Information Service. M. & BOROWSKY. C. R. Gattungen und Arten. L. Amphibien und Reptilien. Studies on Neotropical Fauna and Environment. K. A. Salientia of Venezuela. L. HENLE. EMMONS. 46: 1-584. HENLE. Saint John. Pergamon Press.. Washington D. Amphibian species of the world. 8: 1-48. Stuttgart. J. I. Subordo Aglossa und Phaneroglossa. In: G. 46(1): 87-104. J. In: R. 271 pp. Biotropica. GORHAM. Herpetologica.. J. ARDILA-ROBAYO. B. 1996. New York. O. Observations on the herpetofauna.. J. Amphibien und Reptilien. Studies on Fauna of Surinam & Guyanas. 406 pp. Wissenschaftliche Ergebnisse der Deutschen Gran Chaco-Expedition. Smithsonian Herpetological Information Service. el 16-IV-2002 Aceptado. Evolution in the genus Bufo. F. E. 1936.. Bolivia. A taxonomic and geographical reference. Washington D. MAXSON. Amphibian diversity in Bolivia: a study with special reference to montane forest regions. HASS. R.56 DE LA RIVA. The evolutionary species concept reconsidered. & LYNCH. M.. M. DUNSKI. KÖHLER. 20(77): 365-415.). RUIZ-CARRANZA. Bolivia. 1983. F. Schulenberg & K. R. 1995. VELLARD. The “World’s largest toad” and other herpetological specimens from southern Surinam. Allen Press and the ASC. & ZUG. MÜLLER. & HILLIS. 157: 1-53. 12: 159-171.. GLAW. Revue Suisse de Zoologie. S. A rapid assessment of the humid forests of south central Chuquisaca. Vol. & HOOGMOED. J. 1974. Austin & London: 265-278. RAP Working Papers 1. Teil: Amphibia. RIVERO. Revista de la Academia Colombiana de Ciencias Exactas. J. New Brunswick. MERTENS. 27(1): 17-26.: 3336. Beiträge fur Fauna Perus. Bufo marinus: A natural history resumé of native populations. El género Bufo. The status of Bufo poeppigii Tschudi. 173 pp. & HELLMICH. STEPHENS. VON. 1982. FROST. 284: 158. MORALES. C. 120 pp. HOFRISCHTER. Systematik der Amphibien: Liste der rezenten Familien. III: Wissenschaftliche Bearbeitungen. 1845. Strecker und Schröder Verlag. RODRÍGUEZ. el 2-VII-2002 Publicado. 1993. R. REED. Check list and taxonomic bibliography of the amphibians from Perú. 1992. ZUG. A. Bulletin of the Museum of Comparative Zoology... Anura I. Répartition des batraciens dans les Andes au sud de l’équateur. Smithsonian Contributions to Zoology. Augsburg: 252-258. Lista preliminar de los anfibios del Perú. A. D. & DUBOIS. Hofrischter (ed. A. Kansas. 15: 293-303. & ICOCHEA.. Loricata.. 1961. S. WILEY. W. Reptilium conspectus quae in Republica Peruana reperiuntur et pleraque vel collecta sunt in itinere. H. 27(2): 238-249. Checklist of world amphibians. Publicaciones del Museo de Historia Natural UNMSM. 1923. Lista actualizada de la fauna de Amphibia de Colombia. J. 1998. A. In: W. 1972. Harvard University. Awbrey (eds... R. Físicas y Naturales. P... Travaux de l’Institute Française d’Études Andines.). GUTTMAN. S. J. Lawrence. Reptiles and amphibians from the vicinity of El Palmar in the Andes of Chuquisaca.). Systematic Zoology. V. Das Tierreich. 1995. FROST.. F. 107: 1-20. 1978. Zur Amphibienfauna Perus nebst Beschreibung eines neuen Eleutherodactylus (Leptodactylidae). Amphibien.: 74. The marine toad. Bailey (eds. 1907. I. Conservation International. J. 1985. NIEDEN. VELLARD. I. The University of Kansas. 1955. Archiv für Naturgeschichte. J. Catalogue of New World Amphibians. 45: 1-22. Naturbusch Verlag. A biological assessment of the Alto Madidi region and adjacent areas of northwest Bolivia. RAP Working Papers 8. 1991..). Cambridge. S. Divergent lineages within the Bufo margaritifera complex (Amphibia: Anura: Bufonidae) revealed by albumin immunology. 20: 167-173. HARDING. F. G. 1845 (Anura: Bufonidae). Révision de quelques espèces de reptiles et de batraciens du Pérou décrits par Tschudi. Conservation International. 732 pp. K. Blair (ed. L. Estudios sobre batracios andinos.. R. A check list of the amphibians and reptiles of Ecuador with a bibliography of Ecuadorian herpetology. 1970. Chelonia. K. Memorias del Museo de Historia Natural Javier Prado. Parker & B. J. 1983. Arcifera. M.. In: T. 1952. 83-84. H. 1997. D. 1979. Fischer (ed.. 1985. el 5-IX-2002 ... M. C. Bonner Zoologische Monographien. KÖHLER. ROUX. R. Occasional Papers of the Museum of Natural History. CÓRDOVA. 48: 1243. P.

MNK AM 239. Morona-Santiago: 5.Beni: Confluence Río Blanco and Iténez. 1995). Cochabamba: Carahuasi. 281021-2. Bufo arenarum: ARGENTINA. 2650 m. Río Chipiriri. poeppigii). ECUADOR. are listed followed by the authority of the report.No locality: MHNN 90. 5 miles above entrance of the Río Ipoki. USNM 332403. 3000 ft.. Huánuco: Aucuyacu. 12 km S Quillabamba. La Paz: Alto Madidi. Cuzco Amazónico. Amboró National Park. ca. 300 m. KU 147082.77 (holotype of B. MNK AM 314. 90 m. 5 km S Guayaramerín. BOLIVIA. KU 211746. NHMG 457 (two syntypes of Bufo marinus andinensis). Pasco: Nevati Mission. 139432-3. KU 194914. 1938).. Río Napo at Río Mangua. 281019. USNM 279869-88. AMNH 114843-4. Río Cheyo. Río Beni. Bufo poeppigii: BOLIVIA. KU 196660. Tsioventini. USNM 193902 (seven specimens). Zona Reservada Tambopata-Candamo. Roque. MNK AM 299.. Upper Ucayali River. 312. Yaupi. 200 m. Puno: Vicinity of San Juan.— Specimens examined and localities. 183807. Junín: Chancharia on Río Perene below Pampa Silva. 215. ca. USNM 193898. Amboró National Park. Bolivia). KU 154903. 300 m. 20 km (by river) N Puerto Linares. Misión Bomboiza. 700 m. KU 136191-6. Machu Picchu. 25 m. 840 m.Tucumán: El Cadillal.3 km NE Plan de Milagro. AMNH 78970. poeppigii not supported by voucher specimens. 193903. KU 202273. AMNH 5166-7. USNM: United States National Museum of Natural History.Cochabamba: Aiquile. Cliza. USNM 193890 (four specimens).5 km N Chipiriri. Estirón. 281032. PERU. Cochabamba: Bulo-Bulo (personal observation). 1979). MNK AM 109. KU 136197. Göteborg. AMNH: American Museum of Natural History. Villa Tunari. ca. Caranavi. Río Llullapichis..87 (two syntypes of Bufo molitor).Guayas: 15 km NNW Guayaquil. 15 km E Puerto Maldonado. Tumbes: Matapalo. KU 136048.5 km N Olmos. AMNH 78971. AMNH 108305. KU 183186. Huánuco: Finca Panguana. 600 m. Coroico. USNM 60802-3. KU 183191-261. AMNH 78972. but that represent plausible localities. 1260 m. Piura: Río Piura in Piura. Santa Cruz: Caranda. Lambayeque: 7. ca. 5000 ft. 1750. Pasco: Cacazú (Hass et al.. Valle de Sajta. Rurrenabaque. USNM 281016. 154906-8. MHNN 90. 497 m. 2 km above Costa Márquez. MNK AM 5. 1010 m. KU 136094. 300 m. 2300 m. 680 m. Tumi Chucua. 280448. 171907-13. 281037.. 600 m. Río Mamoré. Río Curanja. KU 219744. KU 219742-3. MHNN: Museé d’Histoire Naturelle de Neuchâtel. ca. Río Saguayo. 2650 m. USNM 166757. USNM 280441.TAXONOMY AND DISTRIBUTION OF BUFO POEPPIGII 57 Appendix I. 800 m. 12 km N Olmos. 670 m. Museum acronyms are: KU: Natural History Museum. KU 196655. USNM 193899-900. Cochabamba. 183803-6. CET 1627-8. NHMG: Naturhistoriska Museet. Balta. Ríos Pitasana y Surutú. Río Iténez. AMNH 72106-29. AMNH 78973. 183808. 193904. KU 154904-5. Loreto: Río Ampiyacu. Pastaza: Limoncocha. KU 160284-97. Mojos. KU 183262-4. Montañas de Monobamba.. KU 196662-7. Guayaramerín. MNK AM 10-11. PERU. Apéndice I. 205826. San Ramón. Río Manu. 2052356. Tingo María. 1829 m. The University of Kansas. Serranía de Sira (Duellman & Toft. 1880 m. MNK: Museo de Historia Natural Noel Kempff Mercado (Santa Cruz. 281034-5. CET A 1666-8. 11 km N Olmos. 187. KU 206086. Puerto Linares. 1550 m.Beni: Rurrenabaque. 1800 m. CET: Centro de Estudios Tropicales (Sevilla.Cuzco: Puente de San Miguel. 1600 m (Andersson. AMNH 137401-3. 154910-1. Bulo-Bulo (personal observation). Smithsonian Institution. USNM 336131-3. USNM 166768. 35 km SE Moyobamba. Amboró National Park. KU 173228. 860 m. USNM 281017. and those in which voucher specimens have not been examined personally by the author. Bufo marinus: BOLIVIA. ca. 500 m. USNM 280437-40. San Martín: Moyobamba. San Buenaventura.— Ejemplares examinados y localidades. and San Ignacio. KU 196661. 270 m. KU 160281-2. USNM 193887 (13 specimens). 281028-9. 1620 m. Madre de Dios: Cocha Cashu. 300 m. San Martín: Tocache Nuevo. KU 191988. 400 m. 670 m. 190. KU 120342-4. KU 139053-75. Spain). 6. 18 km SSE Tafí del Valle. 260 m. Tingo María. 178995. The distributional records of B. 13. .. La Paz: Puerto Linares. KU 98975-83. USNM 280443-4.