Rev. Esp. Herp. (2000) 14:19-164 19

Ten years of research on Bolivian : updated checklist, distribution, taxonomic problems, literature and iconography

IGNACIO DE LA RIVA1, JÖRN KÖHLER2, STEFAN LÖTTERS2 & STEFFEN REICHLE2

1 Museo Nacional de Ciencias Naturales, CSIC, C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain 2 Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany.

Abstract: After ten years of intensive research, an updated checklist of the amphibians of was compi- led. For each species, information on its general distribution within the Neotropics, its occurrence within Boli- vian departments and ecoregions, and use is provided. Key literature and taxonomic remarks, if perti- nent, are added. The list contains 186 valid species recorded from Bolivian territory, 33 (17.7 %) of which are Bolivian endemisms. Two species reported recently (Hyla walfordi and Phrynopus peruvianus) are not included because those reports were based on wrong identifications. Nine species (Allobates femoralis, Epipedobates tri- vittatus, Hyla koechlini, H. sarayacuensis, Phrynohyas resinifictrix, Scinax squalirostris, Phyllomedusa tomop- terna, Eleutherodactylus ventrimarmoratus, and Ischnocnema quixensis) are recorded for the first time. More- over, 67 additional species are expected to occur in the country. Finally, we provide a comprehensive biblio- graphy on Bolivian amphibians and an iconography, figuring almost all Bolivian species in color. Keywords: Amphibia, Bolivia, checklist, , distribution, bibliography, iconography

Resumen: Diez años de investigación sobre anfibios de Bolivia: lista actualizada, distribución, problemas taxonómicos, bibliografía e iconografía.- Se presenta una lista actualizada de los anfibios de Bolivia, que re- copila los resultados de diez años de investigación. Para cada especie se proporciona información sobre distri- bución general en el Neotrópico y en los departamentos y ecoregiones bolivianas, uso del hábitat y bibliografía clave, así como comentarios taxonómicos, cuando proceden. La lista contiene 186 especies válidas registradas en territorio boliviano, de las cuales 33 (17.7 %) son endémicas. Dos especies citadas recientemente (Hyla wal- fordi y Phrynopus peruvianus) no se incluyen en la lista porque dichas citas se basaron en identificaciones erró- neas. Nueve especies (Allobates femoralis, Epipedobates trivittatus, Hyla koechlini, H. sarayacuensis, Phry- nohyas resinifictrix, Scinax squalirostris, Phyllomedusa tomopterna, Eleutherodactylus ventrimarmoratus, e Ischnocnema quixensis) se citan en este trabajo por primera vez. Además, 67 especies aún no citadas podrían existir en el país. Finalmente, se proporciona una bibliografía exhaustiva sobre los anfibios de Bolivia y foto- grafías en color de casi todas las especies registradas. Palabras clave: Amphibia, Bolivia, lista taxonómica, taxonomía, distribución, bibliografía, iconografía

INTRODUCTION distribution, included data from several col- lections, added some new findings, and pro- For many decades, the fauna of vided color photographs of 62 species, some Bolivia has been one of the least known in the of them never illustrated before. This list con- world. Until 1990, the information available tained 112 species accepted as valid, plus se- was scattered among several publications de- veral others of doubtful taxonomic status. In voted mainly to original descriptions of spe- addition, a total of 47 species from four differ- cies, reports on collecting trips, and revisions ent biogeographic areas were considered as of several taxa which included Bolivian mate- likely occurring in the country. rial. The first comprehensive list of Bolivian Throughout the 1990s, the Bolivian am- amphibians was provided by DE LA RIVA phibians have been the subject of intensive re- (1990a), who compiled most of the published search, which is obvious from the increased information, arranged the available data on number of recent publications (see biblio- 20 DE LA RIVA et al. graphic section below). As a result, the num- cludes papers in press about which we are ber of species known for the country by early aware. Basically we intend to reflect the cur- 2000 is much higher than the 1990 number. rent state of taxonomy of Bolivian amphib- This fact means not only a dramatic improve- ians. Thus, in general, in this paper we do not ment of the knowledge of the diversity of Bo- make taxonomic decisions or changes al- livian amphibians, but also of that of their though we are aware of some cases in which taxonomy, distribution, ecology, and natural such changes would be necessary and justi- history. Especially remarkable are the many fied. Several species are reported for the first publications dealing with vocalizations, which time in the present paper, based mostly on vou- have led to the description of the calls of near- cher specimens. Records based solely on per- ly 100 Bolivian species of anurans. sonal communications by other persons have We are convinced of the importance of pa- not been included, unless they have been cor- pers summarizing the state of the art of the roborated by us. Apart from published infor- distribution and diversity of regional faunas, mation, additional data from collections and especially those of poorly surveyed countries. the personal experience of the authors have The first step is “to know what is known”. been used to generate the patterns of geogra- Thus, these summaries, although best consi- phic and ecological distributions of the spe- dered works in-progress, are essential not only cies. Orders, families, subfamilies (when per- as keystones on which the advance of know- tinent), genera, and species are arranged in al- ledge, future research and conservation poli- phabetical order. Each species entry consists cies must be based, but also as an important of the scientific name, author, and year of de- stimulus and guidance for students and other scription, which is followed by an asterisk people interested in natural resources. Follo- when the type locality is in Bolivia, and two wing this philosophy and given the necessity asterisks when the species is endemic to the of a new compilation of the huge amount of country. Additional information is arranged new information generated in the last few into three sections: “Distribution”, “Key liter- years, we have prepared the present contribu- ature” and “Remarks”. tion, whose main goals are to: (1) provide an Distribution.- Under this heading, for each updated checklist of Bolivian amphibians; species, the following five kinds of informa- (2) give brief guidelines about their geograph- tion are provided: ic and ecological distributions; (3) point out (1) Patterns of distribution in the Neotro- particular cases in which taxonomic problems pics.- They are general geographic areas sha- preclude accurate or definitive statements; red by several species with similar ecological (4) include a comprehensive bibliography; requirements and/or biogeographical histo- (5) discuss the present results and predict ries. Some broad patterns encompass other, plausible, future findings; and (6) show color more restricted ones. The patterns are: photographs of almost every amphibian spe- Amazonia, which comprises the entire cies in Bolivia, many of them illustrated here- Amazon basin plus, usually, the Upper Ori- in for the first time. noco basin and the Guianan region. Andean Highlands, which includes the MATERIAL, METHODS AND GUIDELINES puna and Altiplano from, at its greatest exten- sion, S Peru to N Argentina and Chile. Unless stated otherwise, the information Bolivian-Argentinean Forests, compris- provided in the list mainly comes from pub- ing the semi-humid montane Andean forests lished papers cited in the literature, which has from Santa Cruz (Bolivia) to N Catamarca been updated until April 2000, and also in- (Argentina). TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 21

Cerrado domain, which comprises the open temperate and subtropical areas south of the formations whose core area lies in Central and Chaco and the Cerrado, in Argentina, Bolivia, SE Brazil and may also include open areas in SE Brazil, Paraguay and Uruguay. Bolivian the lower Amazon and NE Brazil (Caatingas), species with this general pattern usually occur reaching the Guianas, and/or the Plata Basin in only in the inter-Andean valleys. Argentina and adjacent countries. Temperate Inter-Andean Valleys of Bo- Chaco, which extends mainly throughout livia and adjacent areas of N Argentina. N Argentina, Paraguay, SW Brazil and SE Bo- Tropical Middle and South America, in- livia. The Chaco is often divided into dry and cluding Costa Rica and/or Panama, the Ama- humid Chaco. The later holds some species zon and Orinoco basins and the Guianas, and which are widespread in other areas or are may include the Brazilian Atlantic forest; it may otherwise linked to tropical environments, also include the Pacific lowlands of Colombia whereas the dry Chaco is inhabited by the typ- and Ecuador. ical Chacoan species adapted to the extreme Tropical South America, which is like the conditions of this ecosystem. previous one but excluding Middle America. Humid Montane Forests, and/or humid Upper Amazon Basin, which comprises subparamo in the from S Peru to Bo- roughly the western half of the Amazon Basin livia or only those of Bolivia. and may include or not the Guianan region. Lower Amazon Basin, which comprises All these patterns represent only a rough the eastern half of the Amazon Basin and may way to make species distributions fit in a include the Guianan region. scheme. Consequently, some species can be Middle and South America, which corre- assigned to more than one region or cannot be sponds to species with a broad distribution easily assigned to any of these regions. that may reach up to Mexico in the north and (2) Bolivian departments where the species SE Brazil and northern or central Argentina in occurs. The nine Bolivian departments are the south, albeit more commonly these distri- (Figure 1): Beni, Cochabamba, Chuquisaca, butions range from Panama or Costa Rica to La Paz, Oruro, Pando, Potosí, Santa Cruz, and Bolivia, comprising both tropical and subtro- Tarija. For each species, those departments for pical environments (often such broad distribu- which we are aware of published records, tions reflect a poor knowledge on the system- voucher specimens or personal observation by atics of a given species complex and, as this any of the authors, are marked in boldface. If knowledge improves, less species will fall the presence of the species is not cofirmed but within this pattern of distribution). only expected (for example, because it occurs Peri-Andean Forests, which is the pattern in neighboring areas of other departments or of those species occurring in the foothills and the type of habitat is the same), the depart- lower Andean slopes from Colombia or Ecuador ment appears in normal letters. to Bolivia, but more often from SE Peru to Bo- (3) Bolivian ecoregions (Figure 2), general- livia. ly following IBISCH (1996). Expectedly, some South America, which comprises tropical of them overlap partially or to a great extent and subtropical regions from Colombia to SE the major geographic regions listed in (1). Brazil and northern or central Argentina. These are: Southwestern Amazonia, comprising rou- Amazonian Rainforests: located in Beni, ghly the Madeira river basin in SE Peru, Ama- La Paz, Pando, as well as at the Andean foot- zonian Bolivia, and the Brazilian states of hills in Cochabamba and Santa Cruz; 100-600 Acre and Rondônia. m a.s.l.; mean annual temperature 29-27 °C; Subtropical South America, comprising mean annual precipitation 1800-2200 mm, 22 DE LA RIVA et al.

a.s.l.; mean annual temperature 25-26 °C; mean annual precipitation 400-900 mm, around 1000 mm at the Andean foothills and in the northern transition zone to the Pantanal; 6-8 arid months; low dry-forests with several suc- culent plants. Chaco Montane Forests: located in Chu- quisaca, Santa Cruz, and Tarija; 600-1500 m a.s.l.; mean annual temperatures 18-22 °C; mean annual precipitation 1000-2000 mm; 6- 7 arid months; deciduous forests of medium height; relationships to Caatinga formations. High Andean Vegetation: located in Chu- quisaca, Cochabamba, La Paz, Oruro, Potosí, and Tarija; 2500-4600 m a.s.l.; Polylepis for- est growth up to 5200 m a.s.l. around Volcán Sajama; mean annual temperature below 10 °C; Figure 1.- Map of Bolivian departments. mean annual precipitation 500-700 mm; 6-8 Figura 1.- Mapa de los departamentos Bolivianos. arid months; puna, low evergreen montane forests. more at the Andean foothills; 0-3 arid months; Humid Transition Lowland Forests: lo- tall evergreen rainforest. cated in Beni, Cochabamba, and Santa Cruz; Cerrado formations: located in Santa Cruz forest type unique to Bolivia; 150-300 m a.s.l.; and Beni; 200-1000 m a.s.l.; mean annual mean annual temperatures around 25 °C; mean temperatures 18-25 °C; mean annual precipi- annual precipitation 1200-1800 mm; 2-4 arid tation 1000-1500 mm; 3-5 arid months; main- months; evergreen rainforests of medium ly dry open savanna formations but includes height; azonal gallery forests along rivers. also low dry-forests; frequent natural fires. Inter-Andean Valleys: located in Chu- “Ceja de Montaña” - Cloud Forests: lo- quisaca, Cochabamba, La Paz, Santa Cruz, cated in La Paz, Cochabamba, and Santa and Tarija; 1300-3000 m a.s.l.; mean annual Cruz; 2500-3500 m a.s.l.; mean annual tem- temperature 12-16 °C, maximum above 30 °C, perature 10-14 °C; mean annual precipitation minimum below 0 °C; mean annual precipita- 2500-3500 mm; 0-2 arid months; low ever- tion 500-700 mm; 6-8 arid months; deciduous green cloud forest, subparamo. and semideciduous forests of medium height; Chiquitanía Forests: located in the pro- biogeographical relationships with the dry Cha- vincias Velasco, Ñuflo de Chávez, Sandoval, co forests. and Chiquitos in the Departamento de Santa Pre-Cambrian Shield Humid Forests: Cruz; forest type unique to Bolivia; transition located in N Santa Cruz and parts of Beni; zone between humid Amazonian forests and 200-1000 m a.s.l.; mean annual temperatures dry Chaco forests; 300-1200 m a.s.l.; mean 18-25 °C; mean annual precipitation 1500- annual temperatures 18-23 °C; mean annual 1800 mm; 2-4 arid months; evergreen forest precipitation 1000-1500 mm; 3-5 arid months; of medium height; azonal vegetation on rock semi-deciduous forests of medium height, large outcrops (inselbergs) and sandstone ridges. parts evergreen. Tucumanian-Bolivian Montane Forests: Chaco Lowland Forests: located in Chu- located in Chuquisaca, Santa Cruz, and Tarija; quisaca, Santa Cruz, and Tarija; 300-600 m 800-2600 m a.s.l.; mean annual temperature TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 23

Figure 2.- Map of Bolivian ecoregions. Figure 2.- Mapa de las ecoregiones de Bolivia. 24 DE LA RIVA et al.

13-23 °C; mean annual precipitation 1000- species with digital pads adapted to climb on 2000 mm; 3-5 arid months; montane forests of vegetation, although they can also move on the medium height. ground), Fossorial (when the species spends a Wet Savannas: located in Beni, Santa great part of its life underground, for which it Cruz, and N La Paz; 130-250 m a.s.l.; mean has some special adaptations), and Terrestrial annual temperature around 26 °C; mean an- [many terrestrial or arboreal species are usual- nual precipitation 1000-2000 mm; 2-6 arid ly near bodies of water, either ponds or streams, months; swamps and grass savannas with although they do not necessarily enter the wa- scattered forest islands. ter (e. g., centrolenids)]. Yungas-Montane Rainforests: located in 5) Type of habitat: Forest or Open for- Cochabamba, La Paz, and Santa Cruz; 600- mation. This is an important distinction. Of 2500 m a.s.l.; divided into upper montane course, most amphibians from savannas live rainforests (1500-2500 m a.s.l.) and lower in open formations, but many species from the montane rainforests (600-1500 m a.s.l.); mean forest domain do not occur in the forest itself, annual temperature 12-24 °C, minimum in up- but in the open areas within it (e. g., forest per regions below 0 °C; mean annual precipi- gaps, forest edges, river edges, human clea- tation 2500 to more than 6000 mm, maximum rings, etc.). Although trees in the Chaco can rainfall between 1500 and 1800 m a.s.l.; 0-2 be closely spaced, they are not too tall and the arid months; characterized by steep slopes and Chaco is thus considered an open formation. deep valleys; evergreen montane rainforest of Two facts must be taken into account: first, medium height. these two kinds of major are not clear- The transitional zones between two or more cut categories (there are open secondary of these ecoregions may contain species of forests, areas disturbed by human activity, different origin (for example, in Beni, where patchy environments, etc.); and second, sever- savannas and forest intermingle to form a al species are adapted to live in transitional patchy environment). Likewise, it must be zones or can occur in the two types of habitats taken into account that some species (e. g., (althoug often not being equally abundant in Bufo marinus, Hyla geographica, etc.) are both of them). linked to the rainforest domain, but they in- With these five kinds of information, the habit open areas within it (river shores, clea- putative area of distribution and habitat of a rings, etc.). Moreover, some Cerrado and/or particular species in the country can be easily Chacoan amphibians can enter open areas outlined, and it is possible to predict its occur- within the rainforest domain, either as a natu- rence in areas where it has not been found be- ral colonization or, quite often, taking advan- fore. tage of human perturbations to the landscape Key literature.-This section points out, by (roads, deforestation, etc.). As a result, a Cha- chronological order, the most relevant publi- coan or Cerrado species may be found, for ex- cations on each species in Bolivia, especially ample, in Pando (e. g., elenae). those that appeared after 1990. This section is Likewise, some Chacoan or Cerrado species not included in accounts of species reported may be found in temperate inter-Andean val- herein for the first time, a fact which will be leys (e. g., Scinax squalirostris). Finally, some indicated in the “Remarks” section. Addition- Amazonian species enter the northern Chaco al references, remarks, and data on the distri- (e. g., Sphaenorhynchus lacteus, Leptodacty- bution of many species are found in DE LA lus leptodactyloides). RIVA (1990a) but, in general, this publication (4) Habitat use: Aquatic (when the species will not be cited again as a key literature re- is almost always in the water), Arboreal (those ference for every species, unless no other rel- TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 25 evant publications exist. When the species amphibians presented herein includes every considered has its type locality in Bolivia but paper in which Bolivian amphibians are con- is not a Bolivian endemism, the original cita- sidered in any way. We provide references tion is provided only if it contains additional dealing with ecology, taxonomy, distribution, information (on taxonomy, ecology, distribu- etc. of Bolivian amphibian species as long as tion, natural history, etc.) other than the mere they deal with Bolivian specimens. Papers on description of the species. When a species has species of amphibians that occur in Bolivia been reported in the country only once, the but which are based on research carried out in corresponding reference is given. In some other countries, are not considered (for exam- cases this publication may be a short note on ple, papers dealing with descriptions of species distribution which, although it contains little that occur in Bolivia but whose type locality is information in general terms, represents all not in Bolivia). As an exception, a few refer- the information available for that particular ences cited in the main text do not contain species in the country. Papers and reports on specific information on Bolivian amphibians, faunal lists and surveys of particular localities but are relevant to understand some problems. or areas, inventories, etc., usually deal with no Besides published papers, we also list unpub- single species in particular. Thus, a paper of lished reports and dissertations. This “gray li- this kind is not mentioned in this section, un- terature” is usually difficult to obtain (and we less it contains the only known Bolivian are aware that we might have overlooked some record of the species, unless the species is of these papers), but we consider it is impor- treated more extensively, and/or unless sub- tant to take these papers into account because, stantial information other than geographical is in some instances, they deal with faunas of provided. Likewise, taxonomic reviews of poorly surveyed regions and very often are the genera or species groups which include Boli- only source of information about the amphib- vian specimens, but where they are not the ians occurring in such areas. Communications main subject, are not cited here unless they presented at congresses and meetings are ex- treat Bolivian endemic species. Instead, the cluded, unless they were published as books general bibliography should be consulted. or edited conference proceedings. Likewise, Remarks.-This section (which does not ap- references in which Bolivian amphibians are pear if there is no particular comment about treated in a vague or general context (for ex- the species) includes pertinent comments on ample, non technical books) are excluded, es- taxonomic problems, subspecies, distribution, pecially if there is more accurate information ongoing research on particular topics, etc., published on the same topic elsewhere. Two or with a special reference to the changes that more references by a given author in the same have taken place since DE LA RIVA’s (1990a) year are ordered as a, b, c, etc., only when they list. Recent synonymizations are indicated, es- are cited in this way in the main text. Refe- pecially if the synonymized species has its rences in the list are in alphabetical order. type locality in Bolivia, but older, well settled When there are more than two authors, the ci- taxonomic changes are not usually mentioned. tation in the main text is given with the name In the list we have not included doubtful of the first author followed by “et al.”. In this species [those known only from lost types, or case, if there are several references of the whose type locality is unknown, etc.; see DE same first auhor and year, the citations have LA RIVA (1990a)]. However, we consider them been marked as a, b, c, etc. in the main text, in a separate section because future research following the order with which they appear, may require us to take them into account. and irrespective of the alphabetical order fol- The comprehensive literature on Bolivian lowed in the list of references. 26 DE LA RIVA et al.

RESULTS Pando; 3) Amazonian Rainforests; 4) Ter- restrial; 5) Forest. Updated checklist of Amphibians of Bolivia. Key literature: KÖHLER & LÖTTERS, 1999a.

Anura Bufo fissipes Boulenger, 1903 Bufonidae Distribution: 1) Humid Montane Forests; 2) Cochabamba, La Paz; 3) Yungas-Montane Atelopus tricolor Boulenger, 1902 Rainforests; 4) Terrestrial; 5) Forest. Distribution: 1) Humid Montane Forests; 2) Key literature: DE LA RIVA, 1990a. Cochabamba, La Paz; 3) Yungas-Montane Remarks: FROST (1985) reported this Peru- Rainforests; 4) Terrestrial; 5) Forest. vian species (type locality: Santo Domingo, Key literature: REYNOLDS & FOSTER, 1992; Puno) in the Departamento of Cochabamba, LAVILLA et al., 1997; LÖTTERS & DE LA but we are unaware of voucher specimens RIVA, 1998; LÖTTERS et al., 1999a; KÖH- other than the holotype. It is surprising that LER, 2000a. several species of the B. veraguensis group Remarks: LÖTTERS & DE LA RIVA (1998) have been described in the last years from placed Atelopus willimani Donoso-Barros Peru and Bolivia (and some others still [type locality: Runerrabaque (sic) (= Rur- await description), but this species has yet to renabaque, Beni)] in the synonymy of A. tri- be collected in Bolivia.. color. Bufo granulosus Spix, 1824 Bufo amboroensis Harvey & Smith, 1993* Distribution: 1) Middle and South America; Distribution: 1) Humid Montane Forests; 2) 2) Beni, Cochabamba, Chuquisaca, La Paz, Cochabamba, La Paz Santa Cruz; 3) Yun- Pando, Santa Cruz, Tarija; 3) Amazonian gas-Montane Rainforests; 4) Aquatic, Ter- Rainforests, Cerrado, Chiquitanía Forests, restrial; 5) Forest. Chaco Lowland Forests, Humid Transition Key literature: HARVEY & SMITH, 1993. Lowland Forests, Pre-Cambrian Shield Hu- mid Forests, Wet Savannas; 4) Terrestrial; Bufo arenarum Hensel, 1867 5) Open. Distribution: 1) Subtropical South America; Key literature: REYNOLDS & FOSTER, 1992; 2) Cochabamba, Chuquisaca, Santa Cruz, KÖHLER, 1995a; REICHLE, 1996; 1997a; Tarija; 3) Chaco Lowland Forests, Chaco KÖHLER et al., 1997; HARVEY et al., 1998; Montane Forests, Inter-Andean Valleys, Tu- REICHLE & KÖHLER, 1998. cumanian-Bolivian Montane Forests; 4) Te- Remarks: Bufo granulosus is a polytypic rrestrial; 5) Open. species with 14 described subspecies (GAL- Key literature: DE LA RIVA, 1990a; KÖHLER, LARDO, 1965). Some of them have been 2000a. considered as full species by different au- Remarks: According to LAURENT (1969) the thors. We agree that some of these taxa de- subspecies present in Bolivia is B. arenarum serve specific status, but a revision of the arenarum. Figure 7 in DE LA RIVA (1990a; whole complex is desirable before any taxo- p. 305) actually corresponds to B. poeppigii, nomic decision is made. Four subspecies as well as the record from Caranda, Santa have been reported for Bolivia: B. g. goeldii Cruz (p. 265). Gallardo in the north of the Bolivian Ama- zon, B. g. major Müller & Hellmich in the Bufo castaneoticus Caldwell, 1991 Chacoan area (type locality: San José de Distribution: 1) Amazonia; 2) Beni, La Paz, Chiquitos, Santa Cruz), B. g. mini Gallardo in TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 27

the Beni area, (type locality: Upper Beni, tifer (or B. typhonius) complex is perhaps below mouth of Mapiri) and B. g. miran- one of the most chaotic of all Neotropical daribeiroi Gallardo in northern Santa Cruz. anuran groups (HOOGMOED, 1986; 1989; Bufo g. goeldii, B. g. mini, and B. g. major 1990; HASS et al., 1995). HOOGMOED (1986) are similar in morphology, although B. g. stated that the small toads of the Bolivian major is larger than the two other subspecies. lowlands are similar to B. acutirostris Spix The most distinctive form is B. g. miran- which otherwise occurs in E Colombia, S daribeiroi, which otherwise seems to be more Venezuela and adjacent Brazil. Bioacoustic closely related to the nominal subspecies studies carried out in N Santa Cruz, north- (from NE Brazil) than the other Bolivian ern Chaco of Santa Cruz, and Beni indicate subspecies. Thus, at least two different line- that all these populations are conspecific ages seem to be present in the country. In (DE LA RIVA et al., 1996a; KÖHLER et al., summary, the splitting of B. granulosus 1997). KÖHLER & LÖTTERS (1999a) stated might lead to the recognition of 2-4 species that B. castaneoticus is one of two forms that in Bolivia. occur in the Departament of Pando, and LÖTTERS & KÖHLER (2000a) are describing Bufo guttatus Schneider, 1799 a new species from the humid montane Distribution: 1) Amazonia; 2) Beni, La Paz, forests of Cochabamba and Santa Cruz. The Pando, Santa Cruz; 3) Amazonian Rain- type locality of Bufo pleuropterus Schmidt forests, Humid Transition Lowland Forests, (currently considered a synonym of B. mar- Pre-Cambrian Shield Humid Forests; 4) Ter- garitifer) is given as “Grenzgebiet von Bo- restrial; 5) Forest. livia gegen Peru, in etwa 3000 [feet] Höhe” Key literature: LÖTTERS et al., 2000. (at the border of Bolivia and Peru, approxi- mately 700 m) by SCHMIDT (1858). According Bufo justinianoi Harvey & Smith, 1994** to the loss of Bolivian territory in 1909 and Distribution: 1) CFP; 2) Cochabamba, La the travel route of the collector J. v. Warsze- Paz, Santa Cruz; 3) Yungas-Montane Rain- wiez, this locality is most probably situated forests; 4) Terrestrial; 5) Forest. in present-day Peru. The taxonomic status Key literature: HARVEY & SMITH, 1994; KÖH- of some populations of the B. margaritifer LER, 2000a. complex in the Bolivian lowlands and val- leys remains a topic to be investigated, but it Bufo “margaritifer” (Laurenti, 1768) is almost sure that not less than four differ- Distribution: 1) Tropical Middle and South ent species occur in the country. America; 2) Beni, Cochabamba, Chuqui- saca, La Paz, Pando, Santa Cruz; 3) Ama- Bufo marinus (Linnaeus, 1758) zonian Rainforests, Chiquitanía Forests, Distribution: 1) Middle and South America; Humid Transition Lowland Forests, Inter- 2) Beni, Cochabamba, La Paz, Pando, San- Andean Valleys, Pre-Cambrian Shield Hu- ta Cruz; 3) Amazonian Rainforests, Pre- mid Forests, Tucumanian-Bolivian Montane Cambrian Shield Humid Forests; 4) Terres- Forests, Wet Savannas, Yungas-Montane trial; 5) Open. Rainforests; 4) Terrestrial; 5) Forest, Open. Key literature: DE LA RIVA et al., 1996a. Key literature: REYNOLDS & FOSTER, 1992; KÖHLER, 1995a; DE LA RIVA et al., 1996a; Bufo paracnemis Lutz, 1925 KÖHLER et al., 1997; KÖHLER & LÖTTERS, Distribution: 1) Bolivian-Argentinean Forests, 1999a. Cerrado, Chaco, Subtropical South America; Remarks: The systematics of the B. margari- 2) Beni, Cochabamba, Chuquisaca, Santa 28 DE LA RIVA et al.

Cruz, Tarija; 3) Cerrado, Chiquitanía Fo- tinean species might be related either to the rests, Chaco Lowland Forests, Chaco Mon- spinulosus group (CARRIZO, 1992), or to tane Forests, Humid Transition Lowland the marinus group (HARVEY, 1997). Although Forests, Tucumanian-Bolivian Montane Fo- B. gallardoi or a similar form most probably rests, Wet Savannas; 4) Terrestrial; 5) Open. occurs in the country, we do not include it in Key literature: REYNOLDS & FOSTER, 1992; the list until a reliable record exists. DE LA RIVA, 1993a; KÖHLER, 1995a; 2000a; REICHLE, 1996; 1997a; KÖHLER Bufo veraguensis Schmidt, 1857 et al., 1997. Distribution: 1) Humid Montane Forests; 2) Co- chabamba, Chuquisaca, La Paz, Santa Bufo poeppigii Tschudi, 1845 Cruz, Tarija; 3) Inter-Andean Valleys, Tu- Distribution: 1) Humid Montane Forests, Peri- cumanian-Bolivian Montane Forests, Yun- Andean Forests; 2) Cochabamba, La Paz, gas-Montane Rainforests; 4) Terrestrial; Santa Cruz; 3) Amazonian Rainforests, 5) Forest. Yungas-Montane Rainforests; 4) Terrestrial. Key literature: REYNOLDS & FOSTER, 1992; 5) Forest, Open. HARVEY & SMITH, 1993; 1994; KÖHLER et Key literature: DE LA RIVA et al., 1996a; al., 1995b; KÖHLER, 2000a. KÖHLER, 2000a. Remarks: Some species in the B. veraguensis Remarks: See remarks under B. arenarum. group remain to be described, whereas the taxonomic status of others is doubtful. See Bufo quechua Gallardo, 1961** remarks under B. fissipes. Distribution: 1) Humid Montane Forests; 2) Cochabamba, La Paz, Santa Cruz; 3) Ceja Dendrophryniscus minutus (Melin, 1941) de Montaña-Cloud Forests, Yungas-Mon- Distribution: 1) Amazonia; 2) Beni, Cocha- tane Rainforests; 4) Terrestrial; 5) Forest. bamba, La Paz, Pando, Santa Cruz; 3) Ama- Key literature: GALLARDO, 1961a; HARVEY & zonian Rainforests; 4) Terrestrial; 5) Forest. SMITH, 1993; 1994; KÖHLER, 1995a; 2000a; Key literature: DE LA RIVA, 1999a. KÖHLER et al., 1995a. Remarks: HARVEY & SMITH (1993) placed Melanophryniscus rubriventris (Vellard, 1947) Bufo echinodes Reynolds & Foster [type lo- Distribution: 1) Temperate Inter-Andean cality: Parjacti (sic), Cochabamba (=Parac- Valleys; 2) Cochabamba, Chuquisaca, ti)] in the synonymy of B. quechua. Santa Cruz, Tarija; 3) Inter-Andean Val- leys; 4) Terrestrial. Bufo spinulosus Wiegmann, 1834 Key literature: DE LA RIVA, 1995a. Distribution: 1) Andean Highlands; 2) Cocha- Remarks: The Bolivian specimens known bamba, Chuquisaca, La Paz, Oruro, Po- seem to belong to the nominotypical form. tosí; 3) High Andean Vegetation, Inter-An- dean Valleys; 4) Terrestrial. Centrolenidae Key literature: ERGUETA, 1991; APARICIO, 1993; IBISCH & BÖHME, 1993; KÖHLER, 1995a. Cochranella bejaranoi (Cannatella, 1980)** Remarks: GALLARDO (1961a) described the Distribution: 1) Humid Montane Forests; 2) subspecies B. s. altiperuvianus from Bolivia Cochabamba, Chuquisaca, La Paz, Santa (type locality: Challapata, Oruro). HARVEY Cruz; 3) Yungas-Montane Rainforests; 4) (1997) reported a toad tentatively identified Arboreal; 5) Forest. as B. gallardoi Carrizo from El Palmar Key literature: CANNATELLA, 1980; CANNA- (Chuquisaca). This poorly known Argen- TELLA & DUELLMAN, 1982; SANCHÍZ & DE TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 29

LA RIVA, 1993; HARVEY, 1996; MÁRQUEZ Colostethus brunneus (Cope, 1887) et al., 1996; KÖHLER, 2000a. Distribution: 1) Amazonia, Cerrado; 2) Santa Remarks: HARVEY (1996) placed Centrolene- Cruz; 3) Amazonian Rainforests, Humid lla flavidigitata Reynolds & Foster (type lo- Transition Lowland Forests, Pre-Cambrian cality: Road to San Onofre, 3.3 km by road Shield Humid Forests; 4) Terrestrial; 5) For- N of the road from Cochabamba to Villa Tu- est. nari, Cochabamba) in the synonymy of C. Key literature: GONZÁLES et al., 1999. bejaranoi. Colostethus mcdiarmidi Reynolds & Foster, Cochranella nola Harvey, 1996** 1992** Distribution: 1) Humid Montane Forests, Peri- Distribution: 1) Humid Montane Forests; 2) Andean Forests; 2) Cochabamba, Chuqui- Cochabamba, La Paz; 3) Yungas-Montane saca, Santa Cruz; 3) Amazonian Rainfo- Rainforests; 4) Terrestrial; 5) Forest. rests, Tucumanian-Bolivian Montane Forests, Key literature: REYNOLDS & FOSTER, 1992; Yungas-Montane Rainforests; 4) Arboreal; GONZÁLES et al., 1999. 5) Forest. Key literature: HARVEY, 1996; KÖHLER, 2000a; Colostethus trilineatus (Boulenger, 1884 LÖTTERS & KÖHLER, 2000b. “1883”) Distribution: 1) Southwestern Amazonia; 2) Cochranella pluvialis (Cannatella & Duell- Beni, Cochabamba, La Paz, Pando, Santa man, 1982) Cruz; 3) Amazonian Rainforests, Humid Distribution: 1) Humid Montane Forests; 2) Transition Lowland Forests; 4) Terrestrial; La Paz, Cochabamba; 3) Yungas-Montane 5) Forest. Rainforests; 4) Arboreal; 5) Forest. Key literature: MORALES, 1994; DE LA RIVA Key literature: KÖHLER & REICHLE, 1998. et al., 1996b; REICHLE & KÖHLER, 1996a; GONZÁLES et al., 1999; KÖHLER & LÖT- Hyalinobatrachium bergeri (Cannatella, 1980)* TERS, 1999a. Distribution: 1) Humid Montane Forests, Peri- Remarks: MORALES (1994) stated that C. tri- Andean Forests; 2) Cochabamba, La Paz, lineatus is the proper name for southern Santa Cruz; 3) Amazonian Rainforests, populations of Amazonian Colostethus for- Yungas-Montane Rainforests; 4) Arboreal; merly considered as C. marchesianus (Melin). 5) Forest. Populations not formally described from Key literature: CANNATELLA, 1980; CANNA- Cochabamba (EDWARDS, 1974) are also C. TELLA & DUELLMAN, 1982; SANCHÍZ & DE trilineatus (DE LA RIVA et al., 1996a). An LA RIVA, 1993; MÁRQUEZ et al., 1996; unidentified, related species occurs in Pan- KÖHLER, 2000a. do (KÖHLER & LÖTTERS, 1999a).

Dendrobatidae Epipedobates bolivianus (Boulenger, 1902)** Distribution: 1) Humid Montane Forests; 2) Allobates femoralis (Boulenger, 1884 “1883”) La Paz; 3) Yungas-Montane Rainforests; 4) Distribution: 1) Amazonia; 2) Beni, La Paz, Terrestrial; 5) Forest. Pando; 3) Amazonian Rainforests; 4) Ter- Key literature: BOULENGER, 1902; SILVER- restrial; 5) Forest. STONE, 1976; RODRÍGUEZ & MYERS, 1993; Remarks: Found by J. E. Cadle and S. Reichle GONZÁLES et al., 1999. in November 1999 and reported herein for Remarks: This species has been only recent- the first time. ly rediscovered after almost one century 30 DE LA RIVA et al.

without new findings (GONZÁLES et al., ña-Cloud Forests, Yungas-Montane Rain- 1999). forests; 4) Arboreal; 5) Forest. Key literature: DE LA RIVA, 1992a. Epipedobates hahneli (Boulenger, 1884 Remarks: Known only from the holotype. “1883”) Distribution: 1) Upper Amazon Basin; 2) Be- Gastrotheca marsupiata (Duméril & Bibron, ni, La Paz, Pando; 3) Amazonian Rain- 1841) forests; 4) Terrestrial, Arboreal; 5) Forest. Distribution: 1) Andean Highlands, Humid Key literature: DE LA RIVA et al., 1996b; Montane Forests; 2) Cochabamba, Chuqui- GONZÁLES et al., 1999; KÖHLER & LÖT- saca, La Paz, Oruro, Potosí, Santa Cruz; TERS, 1999a. 3) Ceja de Montaña-Cloud Forests, High Andean Vegetation, Yungas-Montane Rain- Epipedobates pictus (Bibron in Tschudi, 1838)* forests; 4) Arboreal; 5) Forest, Open. Distribution: 1) Southwestern Amazonia, Cer- Key literature: ERGUETA, 1991; DE LA RIVA, rado, Humid Montane Forests; 2) Beni, 1992a; VELLARD, 1992; APARICIO, 1993; Cochabamba, La Paz, Santa Cruz; 3) Ama- IBISCH & BÖHME, 1993; DE LA RIVA et al., zonian Rainforests, Cerrado, Chiquitanía 1995; KÖHLER, 1995a; KÖHLER et al., 1995a. Forests, Humid Transition Lowland Forests, Remarks: The variation in populations from Pre-Cambrian Shield Humid Forests, Wet the highlands and those from the Yungas Savannas, Yungas-Montane Rainforests; was studied by DE LA RIVA (1992a), who 4) Terrestrial; 5) Forest, Open. suggested the possibility that more than one Key literature: REYNOLDS & FOSTER, 1992; species is hidden under the name G. marsu- DE LA RIVA, 1993a; HADDAD & MARTINS, piata in Bolivia. 1994; KÖHLER, 1995a; 2000a; DE LA RIVA et al., 1996b; KÖHLER & LÖTTERS, 1999a; Gastrotheca splendens (Schmidt, 1857) GONZÁLES et al., 1999. Distribution: 1) Humid Montane Forests; 2) Co- Remarks: An undescribed species from La chabamba, Santa Cruz; 3) Yungas-Montane Paz similar to E. pictus is being studied by Rainforests; 4) Arboreal; 5) Forest. S. Lötters and S. Reichle. Key literature: DE LA RIVA, 1992a; DUELLMAN & DE LA RIVA, 1999. Epipedobates trivittatus (Spix, 1824) Remarks: This species was “lost” for almost a Distribution: 1) Amazonia; 2) Beni, La Paz, century and a half. The type locality is un- Pando; 3) Amazonian Rainforests; 4) Ter- known, and the two only specimens known restrial; 5) Forest. at present are the holotype and a specimen Remarks: Found by J. E. Cadle and S. Reichle recently collected in cloud forests of Santa in November 1999 and reported herein for Cruz (DUELLMAN & DE LA RIVA, 1999). It the first time. is likely that G. splendens is a Bolivian en- demism. Hylidae Gastrotheca testudinea (Jiménez de la Espa- Hemiphractinae da, 1871) Distribution: 1) Humid Montane Forests, Peri- Gastrotheca lauzuricae De la Riva, 1992 Andean Forests; 2) Cochabamba, La Paz, “1991” ** Santa Cruz; 3) Amazonian Rainforests, Distribution: 1) Humid Montane Forests; 2) Co- Yungas-Montane Rainforests; 4) Arboreal; chabamba, Santa Cruz; 3) Ceja de Monta- 5) Forest. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 31

Key literature: DE LA RIVA, 1992a; KÖHLER, LER, 1995a; 2000a; KÖHLER et al., 1995b; 2000a. DUELLMAN et al., 1997. Remarks: DUELLMAN et al. (1997) elevated Hylinae Hyla pulchella andina to specific status. Hyla andina shows remarkable variation Hyla acreana Bokermann, 1964 and further studies might demonstrate that Distribution: 1) Southwestern Amazonia; 2) more than one species is involved. Figures 5 Beni, Cochabamba, La Paz, Pando, Santa and 6 in KÖHLER et al. (1995b; p. 17), pur- Cruz; 3) Amazonian Rainforests, Humid portedly showing specimens of H. pulchella Transition Lowland Forests, Pre-Cambrian andina, actually correspond to H. marianitae. Shield Humid Forests; 4) Arboreal; 5) For- HARVEY (1997) was unsure if specimens he est. assigned to H. pulchella (= H.andina) from Key literature: DE LA RIVA, 1993a; MÁRQUEZ El Palmar (Chuquisaca) could actually be et al., 1993. Hyla varelae Carrizo. He remarked that Ca- rrizo’s description was inadequate to make a Hyla albonigra Nieden, 1923** definite identification without examination Distribution: 1) Andean Highlands, Temperate of comparative material. Inter-Andean Valleys; 2) Cochabamba, Chuquisaca, Potosí, Santa Cruz; 3) High Hyla armata Boulenger, 1902* Andean Vegetation, Inter-Andean Valleys; Distribution: 1) Humid Montane Forests; 2) Co- 4) Arboreal; 5) Open. chabamba, La Paz, Santa Cruz; 3) Yun- Key literature: DUMÉRIL & BIBRON, 1841; gas-Montane Rainforests; 4) Arboreal; 5) Fo- D’ORBIGNY, 1847; NIEDEN, 1923; ANDERS- rest. SON, 1938; DUELLMAN et al., 1997. Key literature: REYNOLDS & FOSTER, 1992; Remarks: DUELLMAN et al. (1997) placed MÁRQUEZ et al., 1993; DUELLMAN et al., Hyla ocapia Andersson (type locality: Puka 1997; KÖHLER, 2000a. Khara, Potosí) in the synonymy of H. albo- Remarks: There is considerable variation in nigra. skin texture, color pattern, color of the iris and, apparently, advertisement calls, which Hyla albopunctata Spix, 1824 suggests that more than one species might Distribution: 1) Cerrado; 2) Santa Cruz; 3) be involved under the name H. armata (DU- Cerrado; 4) Arboreal; 5) Open. ELLMAN et al., 1997). Key literature: DE SÁ, 1995; DE LA RIVA et al., 1997. Hyla balzani Boulenger, 1898* Distribution: 1) Humid Montane Forests; 2) Co- Hyla andina Müller, 1924 chabamba, La Paz; 3) Yungas-Montane Rain- Distribution: 1) Andean Highlands, Humid forests; 4) Arboreal; 5) Forest. Montane Forests, Temperate Inter-Andean Key literature: REYNOLDS & FOSTER, 1992; Valleys; 2) Cochabamba, Chuquisaca, La DUELLMAN et al., 1997; KÖHLER, 2000a. Paz, Santa Cruz, Tarija; 3) Ceja de Mon- Remarks: DUELLMAN et al. (1997) placed Hyla taña-Cloud Forests, High Andean Vegeta- callipleura Boulenger [type locality: Charu- tion, Inter-Andean Valleys, Yungas-Mon- playa (Cochabamba) and San Ernesto (La tane Rainforests; 4) Terrestrial, Arboreal; Paz)] in the synonymy of H. balzani. How- 5) Forest, Open. ever, further evidence based mainly on vo- Key literature: ERGUETA, 1991; REYNOLDS & calizations seems to indicate that perhaps H. FOSTER, 1992; MÁRQUEZ et al., 1993; KÖH- callipleura should be resurrected (J. Köhler 32 DE LA RIVA et al.

and S. Reichle, pers. obs.). Additionally, Key literature: VELLARD, 1970; DUELLMAN et some populations of the H. pulchella al., 1997. group in the inter-Andean temperate va- Remarks: This poorly known species (see DU- lleys possibly represent an undescribed ELLMAN et al., 1997, p. 12) was recently re- species (see KÖHLER et al., 1995b; DUELL- discovered by J. Aparicio and S. Reichle at MAN et al., 1997). the type locality. It is similar to Hyla arma- ta, but seems to represent a different species Hyla bifurca Andersson, 1945 whose status must be studied together with Distribution: 1) Upper Amazon Basin; 2) Beni, that of certain populations currently consi- Cochabamba, La Paz, Pando, Santa Cruz; dered as H. armata (see remarks on H. ar- 3) Amazonian Rainforests, Humid Tran- mata). sition Lowland Forests, Pre-Cambrian Shield Humid Forests; 4) Arboreal; 5) For- Hyla chlorostea Reynolds & Foster, 1992** est, Open. Distribution: 1) Humid Montane Forests; 2) Key literature: DE LA RIVA, 1993a; MÁRQUEZ Cochabamba; 3) Yungas-Montane Rain- et al., 1993. forests; 4) Arboreal; 5) Forest. Remarks: There are considerable differences Key literature: REYNOLDS & FOSTER, 1992. in color pattern between Santa Cruz popula- Remarks: Known only from the holotype. Un- tions and those from Ecuador (where the certain taxonomic status. species has its type locality). In Pando there are specimens like those from Ecuador and Hyla fasciata Günther, 1859 those from Santa Cruz, as well as intermedi- Distribution: 1) Upper Amazon Basin; 2) Beni, ate ones (DE LA RIVA, 1993a). Cochabamba, La Paz, Pando, Santa Cruz; 3) Amazonian Rainforests, Humid Transi- Hyla boans (Linnaeus, 1758) tion Lowland Forests, Pre-Cambrian Shield Distribution: 1) Tropical Middle and South Humid Forests; 4) Arboreal; 5) Forest. America; 2) Beni, Cochabamba, La Paz, Key literature: DE LA RIVA, 1993a; MÁRQUEZ Pando, Santa Cruz; 3) Amazonian Rain- et al., 1993. forests, Humid Transition Lowland Forests, Remarks: DE LA RIVA (1990a) placed Hyla Pre-Cambrian Shield Humid Forests; 4) Ar- steinbachi Boulenger (type locality: Pro- boreal; 5) Forest, Open. vince Sarah, Santa Cruz) in the synonymy Key literature: DE LA RIVA, 1990a. of H. fasciata.

Hyla calcarata Troschel, 1848 Hyla geographica Spix, 1824 Distribution: 1) Upper Amazon Basin; 2) La Distribution: 1) Amazonia; 2) Beni, Cocha- Paz, Pando; 3) Amazonian Rainforests, bamba, La Paz, Pando, Santa Cruz; 3) Ama- Humid Transition Lowland Forests; 4) Ar- zonian Rainforests, Humid Transition Low- boreal; 5) Forest. land Forests, Pre-Cambrian Shield Humid Key literature: DE LA RIVA, 1990a; APARICIO, Forests, Wet Savannas; 4) Arboreal; 5) Open. 1992. Key literature: DE LA RIVA, 1993a; MÁRQUEZ et al., 1993; KÖHLER, 1995a; REICHLE, Hyla charazani Vellard, 1970** 1996; 1997b; REICHLE & KÖHLER, 1998. Distribution: 1) Temperate Inter-Andean Val- Remarks: DE LA RIVA’s (1990a) statement leys; 2) La Paz; 3) Ceja de Montaña-Cloud that the subspecies present in Bolivia is H. Forests, Inter-Andean Valleys; 4) Aquatic, g. punctatissima (Reinhardt & Lütken) is Arboreal; 5) Open. unfounded. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 33

Hyla granosa Boulenger, 1882 group (the H. leucophyllata group). Although Distribution: 1) Amazonia; 2) La Paz, Pando; H. rossalleni has not been found in Bolivia, 3) Amazonian Rainforests; 4) Arboreal; its occurrence in the country is expected. 5) Forest, Open. Key literature: KÖHLER & LÖTTERS, 1999a. Hyla leucophyllata (Beireis, 1783) Distribution: 1) Amazonia; 2) Beni, Cocha- Hyla koechlini Duellman & Trueb, 1989 bamba, La Paz, Pando, Santa Cruz; 3) Ama- Distribution: 1) Southwestern Amazonia; 2) zonian Rainforests, Humid Transition Low- Beni, La Paz, Pando, Santa Cruz; 3) Ama- land Forests, Pre-Cambrian Shield Humid zonian Rainforests; 4) Arboreal; 5) Forest. Forests, Wet Savannas; 4) Arboreal; 5) For- Remarks: This is the first record for the est, Open. species in the country. Key literature: DE LA RIVA, 1993a; MÁRQUEZ et al., 1993; KÖHLER, 1995a; REICHLE, Hyla lanciformis (Cope, 1870) 1996; 1997b; REICHLE & KÖHLER, 1998. Distribution: 1) Amazonia; 2) Beni, Cocha- bamba, La Paz, Pando, Santa Cruz; 3) Ama- Hyla marianitae Carrizo, 1992 zonian Rainforests, Humid Transition Low- Distribution: 1) Bolivian-Argentinean Forests, land Forests, Pre-Cambrian Shield Humid Humid Montane Forests, Temperate Inter- Forests, Wet Savannas; 4) Arboreal; 5) For- Andean Valleys; 2) Cochabamba, Chuqui- est, Open. saca, Santa Cruz, Tarija; 3) Amazonian Key literature: MÁRQUEZ et al., 1993. Rainforests, Inter-Andean Valleys, Tucum- anian-Bolivian Montane Forests, Yungas- Hyla leali Bokermann, 1964 Montane Rainforests; 4) Arboreal; 5) Forest, Distribution: 1) Southwestern Amazonia; 2) Be- Open. ni, Cochabamba, La Paz, Pando, Santa Key literature: MÁRQUEZ et al., 1993; DUELL- Cruz; 3) Amazonian Rainforests, Humid MAN et al., 1997; HARVEY, 1997; LÖTTERS Transition Lowland Forests, Pre-Cambrian et al., 1999b; KÖHLER, 2000a. Shield Humid Forests, Wet Savannas; 4) Ar- Remarks: The description of the call of H. cal- boreal; 5) Forest, Open. lipleura (= H. balzani) in MÁRQUEZ et al. Key literature: DE LA RIVA, 1993a; MÁRQUEZ (1993) actually corresponds to H. mariani- et al., 1993; REICHLE, 1996; 1997b; REICH- tae, as stated by DUELLMAN et al. (1997). LE & KÖHLER, 1998. See remarks under H. andina. Remarks: This species was erroneously con- sidered a junior synonym of Hyla rossalleni Hyla marmorata (Laurenti, 1768) Goin by DUELLMAN (1974). Although HE- Distribution: 1) Upper Amazon Basin; 2) Be- YER (1977) clearly stated that both species ni, La Paz, Pando, Santa Cruz; 3) Ama- were different and further publications treat- zonian Rainforests, Humid Transition Low- ed H. leali and H. rossalleni as separate spe- land Forests, Pre-Cambrian Shield Humid cies [for example, FROST (1985)], a certain Forests; 4) Arboreal; 5) Forest. degree of confusion persisted for many Key literature: KÖHLER, 1995a; 1995b; KÖH- years. This situation led DE LA RIVA (1990a) LER & BÖHME, 1996. to report H. rossalleni in Bolivia. Finally, DE LA RIVA & DUELLMAN (1997) clarified Hyla melanargyrea Cope, 1887 the taxonomic status of this last mentioned Distribution: 1) Cerrado; 2) Santa Cruz; 3) Ce- species, which is easily distinguishable from rrado, Chiquitanía Forests; 4) Arboreal; H. leali and belongs to a different species 5) Open. 34 DE LA RIVA et al.

Key literature: DE LA RIVA, 1993a; MÁRQUEZ Cruz; 3) Amazonian Rainforests, Chiqui- et al., 1993; KÖHLER, 1995a; KÖHLER & tanía Forests, Chaco Lowland Forests, Hu- BÖHME, 1996. mid Transition Lowland Forests, Pre-Cam- brian Shield Humid Forests, Wet Savannas; Hyla minuta Peters, 1872 4) Arboreal; 5) Open. Distribution: 1) South America; 2) Beni, Key literature: DE LA RIVA, 1993a; MÁRQUEZ Cochabamba, Chuquisaca, La Paz, Pando, et al., 1993; KÖHLER, 1995a; REICHLE, Santa Cruz, Tarija; 3) Amazonian Rain- 1996; 1997b; REICHLE & KÖHLER, 1998. forests, Cerrado, Chiquitanía Forests, Chaco Remarks: The validity of the Chacoan sub- Montane Forests, Humid Transition Low- species Hyla punctata rubrolineata Lutz (type land Forests, Inter-Andean Valleys, Pre- locality: Buenavista, Santa Cruz) is doubtful. Cambrian Shield Humid Forests, Tucuma- nian-Bolivian Montane Forests, Wet Savan- Hyla raniceps (Cope, 1862) nas; 4) Arboreal; 5) Forest, Open. Distribution: 1) Cerrado, Chaco; 2) Beni, Key literature: DE LA RIVA, 1993a; MÁRQUEZ Santa Cruz; 3) Cerrado, Chiquitanía Fo- et al., 1993; KÖHLER, 1995a; 2000a; KÖH- rests, Chaco Lowland Forests, Humid Tran- LER et al., 1995b; REICHLE, 1996; 1997b; sition Lowland Forests, Pre-Cambrian Shield REICHLE & KÖHLER, 1998. Humid Forests, Wet Savannas; 4) Arboreal; Remarks: A species related to H. minuta from 5) Open. the Chapare region of Cochabamba is being Key literature: DE LA RIVA, 1993a; MÁRQUEZ described by KÖHLER & LÖTTERS (2000). et al., 1993; KÖHLER, 1995a; REICHLE, 1996; 1997b; REICHLE & KÖHLER, 1998. Hyla nana Boulenger, 1889 Distribution: 1) Cerrado, Chaco, Subtropical Hyla rhodopepla Günther, 1859 South America; 2) Beni, Chuquisaca, Santa Distribution: 1) Upper Amazon Basin; 2) Beni, Cruz; 3) Cerrado, Chiquitanía Forests, Cha- Cochabamba, La Paz, Pando, Santa Cruz; co Lowland Forests, Wet Savannas; 4) Arbo- 3) Amazonian Rainforests, Yungas-Monta- real; 5) Open. ne Rainforests; 4) Arboreal; 5) Forest, Open. Key literature: LANGONE & BASSO, 1987; DE Key literature: MÁRQUEZ et al., 1993. LA RIVA, 1993a; MÁRQUEZ et al., 1993; KÖHLER, 1995a; REICHLE, 1996; 1997b; Hyla riveroi Cochran & Goin, 1970 REICHLE & KÖHLER, 1998. Distribution: 1) Upper Amazon Basin; 2) Beni, Cochabamba, La Paz, Pando, Santa Cruz; Hyla parviceps Boulenger, 1882 3) Amazonian Rainforests, Humid Transi- Distribution: 1) Upper Amazon Basin; 2) Be- tion Lowland Forests, Pre-Cambrian Shield ni, Cochabamba, La Paz, Pando, Santa Humid Forests, Wet Savannas; 4) Arboreal; Cruz; 3) Amazonian Rainforests, Humid 5) Open. Transition Lowland Forests, Pre-Cambrian Key literature: DE LA RIVA, 1993a; MÁRQUEZ Shield Humid Forests; 4) Arboreal; 5) Fo- et al., 1993; REICHLE, 1996; 1997b; REICH- rest, Open. LE & KÖHLER, 1998. Key literature: DE LA RIVA, 1993a; MÁRQUEZ et al., 1993. Hyla rubicundula Reinhardt & Lütken, 1862 Distribution: 1) Cerrado; 2) Santa Cruz; 3) Hyla punctata (Schneider, 1799) Cerrado, Chiquitanía Forests; 4) Arboreal; Distribution: 1) Tropical South America; 2) Be- 5) Open. ni, Cochabamba, La Paz, Pando, Santa Key literature: DE LA RIVA, 1990a. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 35

Hyla sarayacuensis Shreve, 1935 Osteocephalus pearsoni (Gaige, 1929)* Distribution: 1) Upper Amazon Basin; 2) Beni, Distribution: 1) Southwestern Amazonia; 2) Be- La Paz, Pando; 3) Amazonian Rainforests; ni, Cochabamba, La Paz, Pando; 3) Ama- 4) Arboreal; 5) Forest. zonian Rainforests; 4) Arboreal; 5) Forest. Remarks: This is the first record for the spe- Key literature: GAIGE, 1929; DE LA RIVA, cies in the country. 1990a.

Hyla schubarti Bokermann, 1963 Osteocephalus taurinus Steindachner, 1862 Distribution: 1) Southwestern Amazonia; 2) Be- Distribution: l) Amazonia; 2) Beni, Cocha- ni, La Paz, Pando; 3) Amazonian Rainfo- bamba, La Paz, Pando, Santa Cruz; 3) Ama- rests, Humid Transition Lowland Forests; zonian Rainforests, Humid Transition Low- 4) Arboreal; 5) Forest. land Forests, Pre-Cambrian Shield Humid Key literature: DE LA RIVA, 1999b. Forests; 4) Arboreal; 5) Forest. Key literature: DE LA RIVA, 1993a; DE LA Hyla tritaeniata Bokermann, 1965 RIVA et al.,1995; KÖHLER, 1995a; KÖHLER Distribution: 1) Cerrado; 2) Beni, Santa Cruz; & BÖHME, 1996. 3) Cerrado, Chiquitanía Forests, Wet Savan- nas; 4) Arboreal; 5) Open. Phrynohyas coriacea (Peters, 1867) Key literature: REICHLE, 1996; 1997b; REICH- Distribution: 1) Upper Amazon Basin; 2) Be- LE & KÖHLER, 1996a; 1998. ni, La Paz, Pando, Santa Cruz; 3) Amazon- Remarks: Based on this species’ remarkable ian Rainforests, Humid Transition Lowland intra- and interpopulational variation, NA- Forests, Pre-Cambrian Shield Humid Forests; POLI & CARAMASCHI (1999) suggested that 4) Arboreal; 5) Forest. H. tritaeniata is a complex of species. Key literature: DE LA RIVA, 1993a; 1994a; DE LA RIVA et al., 1995. Osteocephalus buckleyi (Boulenger, 1882) Distribution: 1) Upper Amazon Basin; 2) Be- Phrynohyas resinifictrix (Goeldi, 1907) ni, Cochabamba, La Paz, Pando, Santa Distribution: 1) Amazonia; 2) Beni, La Paz, Cruz; 3) Amazonian Rainforests, Yungas- Pando, Santa Cruz; 3) Amazonian Rain- Montane Rainforests; 4) Arboreal; 5) For- forests; 4) Arboreal; 5) Forest. est. Remarks: Found by J. E. Cadle and S. Reichle Key literature: DE LA RIVA, 1990a. in November 1999 and reported here for the first time. Osteocephalus leprieurii (Duméril & Bibron, 1841) Phrynohyas venulosa (Laurenti, 1768) Distribution: 1) Upper Amazon Basin; 2) Be- Distribution: 1) Middle and South America; ni, La Paz, Pando, Santa Cruz; 3) Amazo- 2) Beni, Cochabamba, Chuquisaca, La nian Rainforests, Humid Transition Low- Paz, Pando, Santa Cruz, Tarija; 3) Ama- land Forests, Pre-Cambrian Shield Humid zonian Rainforests, Cerrado, Chiquitanía Forests; 4) Arboreal; 5) Forest. Forests, Chaco Lowland Forests, Chaco Key literature: DE LA RIVA, 1993a; DE LA Montane Forests, Humid Transition Low- RIVA et al., 1995. land Forests, Inter-Andean Valleys, Pre- Remarks: SMITH et al. (2000) are describing a Cambrian Shield Humid Forests, Tucuman- new species of Bolivian Osteocephalus ian-Bolivian Montane Forests, Wet Savan- which corresponds to what is here consi- nas; 4) Arboreal; 5) Forest, Open. dered O. leprieurii. Key literature: DE LA RIVA, 1993a; DE LA 36 DE LA RIVA et al.

RIVA et al., 1995; KÖHLER, 1995a; 2000a; when other populations are considered. Ho- KÖHLER et al., 1995b; REICHLE, 1996; wever, many species have a distribution 1997b; REICHLE & KÖHLER, 1998. comprising the temperate and subtropical valleys of NW Argentina up to Santa Cruz Scarthyla ostinodactyla Duellman & de Sá, or Cochabamba (HARVEY, 1997). Thus, sin- 1988 ce the presence of one species does not ex- Distribution: 1) Upper Amazon Basin; 2) La clude the other (they are sympatric at some Paz, Pando; 3) Amazonian Rainforests; Bolivian localities), the occurrence of S. cas- 4) Aquatic, Arboreal; 5) Forest. troviejoi in Argentina would still be plausi- Key literature: DE LA RIVA, 1999c. ble even if the paratypes from Jujuy proved to be S. fuscovarius. Scinax acuminatus (Cope, 1862) Distribution: 1) Cerrado, Chaco; 2) Chuqui- Scinax chiquitanus (De la Riva, 1990)* saca, Santa Cruz, Tarija; 3) Cerrado, Chaco Distribution: 1) Southwestern Amazonia; 2) Be- Lowland Forests, Wet Savannas; 4) Arbo- ni, Cochabamba, La Paz, Pando, Santa real; 5) Open. Cruz; 3) Amazonian Rainforests, Humid Key literature: DE LA RIVA, 1990a. Transition Lowland Forests, Pre-Cambrian Shield Humid Forests; 4) Arboreal; 5) For- Scinax castroviejoi De la Riva, 1993* est. Distribution: 1) Bolivian-Argentinean Forests, Key literature: DE LA RIVA, 1990b; 1993a; DE Temperate Inter-Andean Valleys; 2) Cocha- LA RIVA et al., 1994; KÖHLER, 1995a. bamba, Chuquisaca, La Paz, Santa Cruz, Remarks: DUELLMAN & WIENS (1993) pro- Tarija; 3) Inter-Andean Valleys, Tucuma- vided data on this species from Peru. The nian-Bolivian Montane Forests, Yungas- male shown in their Fig. 2 (p. 8) has a color Montane Rainforests; 4) Arboreal; 5) Open. pattern which differs markedly from that of Key literature: DE LA RIVA, 1993b; DE LA Bolivian males, which otherwise present lit- RIVA et al., 1994; KÖHLER, 1995a; 2000a; tle variation. The taxonomic status of Peru- KÖHLER et al., 1995b. vian specimens identified as S. chiquitanus Remarks: This species was originally descri- should be reassessed. bed from temperate valleys of Santa Cruz and from a locality in Jujuy (Argentina). Scinax fuscovarius (Lutz, 1925) Subsequent surveys carried out at the Jujuy Distribution: 1) Cerrado, Chaco; 2) Beni, Chu- locality failed to find the species. Instead, quisaca, Santa Cruz, Tarija; 3) Cerrado, S. fuscovarius was common. These are sib- Chiquitanía Forests, Chaco Lowland Forests, ling species difficult to distinguish mor- Chaco Montane Forests, Humid Transition phologically, but they have very different Lowland Forests, Inter-Andean Valleys, Pre- advertisement calls. Argentinean paratypes Cambrian Shield Humid Forests, Tucuma- of S. castroviejoi were assigned to this spe- nian-Bolivian Montane Forests, Wet Savan- cies based on morphology only. Thus, it is nas; 4) Terrestrial, Arboreal; 5) Open. possible that these specimens actually are Key literature: DE LA RIVA, 1993a; 1993b; DE S. fuscovarius, a species whose geographic LA RIVA et al., 1994; KÖHLER, 1995a; variation could have enhanced the similari- 2000a; KÖHLER & BÖHME, 1996. ty between the Jujuy and Santa Cruz popu- Remarks: We follow LUTZ (1973) and CEI lations. Bolivian samples of these two (1980) in considering Scinax trachythorax species can be distinguished by meristic (Müller & Hellmich) as a synonym of S. characters, but perhaps this is not the case fuscovarius. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 37

Scinax garbei (Miranda-Ribeiro, 1926) ried out by DE LA RIVA et al. (1994; 1997) Distribution: 1) Upper Amazon Basin; 2) Be- suggested that they are not conspecific. ni, Cochabamba, La Paz, Pando, Santa However, this must be corroborated with Cruz; 3) Amazonian Rainforests, Cerrado, more data. As stated by DE LA RIVA et al. Humid Transition Lowland Forests, Pre- (1997), the taxonomic status of the Ama- Cambrian Shield Humid Forests; 4) Arbo- zonian species of the Scinax staufferi group real; 5) Forest, Open. needs urgent revision. In Bolivia, at least Key literature: DE LA RIVA, 1993a; DE LA another species (probably undescribed) in RIVA et al., 1994. the staufferi group occurs in the open areas Remarks: DUELLMAN & WIENS (1993) placed of Beni and Santa Cruz. S. epacrorhina (Duellman) [which was re- ported for Bolivia; see DE LA RIVA (1990a)] Scinax ruber (Laurenti, 1768) in the synonymy of S. garbei. Distribution: 1) Tropical Middle and South America; 2) Beni, Cochabamba, La Paz, Scinax nasicus (Cope, 1862) Pando, Santa Cruz; 3) Amazonian Rain- Distribution: 1) Cerrado, Chaco; 2) Beni, forests, Chiquitanía Forests, Humid Transi- Cochabamba, Chuquisaca, Santa Cruz, tion Lowland Forests, Pre-Cambrian Shield Tarija; 3) Cerrado, Chiquitanía Forests; Humid Forests, Wet Savannas; 4) Arboreal; 4) Arboreal, Terrestrial; 5) Open. 5) Forest, Open. Key literature: DE LA RIVA, 1993b; DE LA Key literature: DE LA RIVA, 1993a; DE LA RIVA et al., 1994; KÖHLER, 1995a. RIVA et al., 1994; KÖHLER, 1995a; KÖHLER & BÖHME, 1996; REICHLE, 1996; 1997b; Scinax nebulosus (Spix, 1824) REICHLE & KÖHLER, 1998. Distribution: 1) Lower Amazon Basin; 2) Be- Remarks: Given the broad distribution and ni, Santa Cruz; 3) Amazonian Rainforests, variation of S. ruber, probably there are Cerrado, Humid Transition Lowland Fo- more than one species hidden under this rests, Pre-Cambrian Shield Humid Forests, name (DUELLMAN & WIENS, 1993). As far Wet Savannas; 4) Arboreal; 5) Open. as we know, nobody is currently reviewing Key literature: DE LA RIVA, 1990a; HOOG- this species complex. Given that the type MOED, 1993; DE LA RIVA et al., 1994; KÖH- locality (Paramaribo, Surinam) is far away LER, 1995a. from Bolivia, it would be likely that Boli- vian populations would receive another na- Scinax parkeri (Gaige, 1929)* me if S. ruber were split into several Distribution: 1) Lower Amazon Basin, Cerra- species. do; 2) Beni, Santa Cruz; 3) Amazonian Rainforests, Chiquitanía Forests, Humid Scinax squalirostris (A. Lutz, 1925) Transition Lowland Forests, Pre-Cambrian Distribution: 1) Cerrado; 2) Beni, La Paz, Shield Humid Forests, Wet Savannas; 4) Ar- Santa Cruz; 3) Cerrado; 4) Arboreal; 5) Open. boreal; 5) Open. Remarks: This species is reported here for the Key literature: DE LA RIVA et al., 1994; 1997; first time, based on specimens collected by KÖHLER, 1995a; REICHLE, 1996; 1997b; J. Bosch, I. De la Riva and S. Reichle near REICHLE & KÖHLER, 1998. Apolo, La Paz, in November 1999. The oc- Remarks: Many authors have followed LUTZ currence of S. squalirostris in the country (1973) in considering this species a junior was expected in the Pantanal area of Santa synonym of S. fuscomarginatus (Lutz), but Cruz. Admittedly, its discovery for the first comparisons of the calls of both species car- time at a locality so far away from the Pan- 38 DE LA RIVA et al.

tanal is surprising. However, the temperate ported specimens of P. boliviana actually valleys around Apolo are suitable for this corresponded to a different species (see re- and other species typical of temperate low- marks under P. camba). lands and/or valleys (e. g., Scinax castro- viejoi and Phyllomedusa boliviana, which Phyllomedusa camba De la Riva, 2000 were also found in the area). We predict that “1999”* other populations of S. squalirostris will be Distribution: 1) Southwestern Amazonia; 2) Be- found in suitable habitats of La Paz, Beni ni, Cochabamba, La Paz, Pando, Santa and Santa Cruz. Cruz; 3) Amazonian Rainforests, Chiqui- tanía Forests, Humid Transition Lowland Sphaenorhynchus lacteus (Daudin, 1802) Forests, Pre-Cambrian Shield Humid Fo- Distribution: 1) Amazonia; 2) Beni, Cocha- rests; 4) Arboreal; 5) Forest. bamba, La Paz, Pando, Santa Cruz; 3) Ama- Key literature: DE LA RIVA, 1993a; DE LA zonian Rainforests, Chiquitanía Forests, Hu- RIVA et al., 1995; KÖHLER & LÖTTERS, mid Transition Lowland Forests, Pre-Cam- 1999a; DE LA RIVA, 2000. brian Shield Humid Forests, Wet Savannas; Remarks: The existence of this species, con- 4) Arboreal; 5) Forest, Open. fused with P. boliviana by several authors, Key literature: DE LA RIVA, 1993a; DE LA was announced long ago by CANNATELLA RIVA et al., 1995; KÖHLER, 1995a; REI- (1983) and it has been repeatedly reported CHLE, 1996; 1997c; REICHLE & KÖHLER, (usually as Phyllomedusa sp.) from several 1998. localities in SW Peru, Brazil, and Bolivia (see DE LA RIVA, 2000). Phyllomedusinae Phyllomedusa hypochondrialis (Daudin, Phyllomedusa bicolor (Boddaert, 1772) 1800) Distribution: 1) Upper Amazon Basin; 2) Be- Distribution: 1) Tropical South America; 2) ni, La Paz, Pando, Santa Cruz; 3) Amazo- Beni, Santa Cruz; 3) Chiquitanía Forests, nian Rainforests; 4) Arboreal; 5) Forest. Chaco Lowland Forests, Pre-Cambrian Key literature: REICHLE et al., 1997. Shield Humid Forests, Wet Savannas; 4) Ar- boreal; 5) Open. Phyllomedusa boliviana Boulenger, 1902* Key literature: DE LA RIVA et al., 1995; KÖH- Distribution: 1) Bolivian-Argentinean Fo- LER, 1995a; REICHLE, 1996; 1997c. rests, Cerrado, Humid Montane Forests, Remarks: LESCURE et al. (1995) stated that Temperate Inter-Andean Valleys; 2) Beni, the correct spelling of the specific name is Cochabamba, Chuquisaca, La Paz, Santa hypochondrialis instead hypocondrialis, a Cruz, Tarija; 3) Chiquitanía Forests, Inter- name frequently found in the literature. The Andean Valleys, Tucumanian-Bolivian Mon- record of P. rohdei Mertens in Beni by tane Forests, Yungas-Montane Rainforests; FUNKHOUSER (1957) must be in error and 4) Arboreal; 5) Open. correspond to P. hypochondrialis. After CEI Key literature: CANNATELLA, 1983; KÖHLER, (1980), the subspecies P. h. azurea Cope oc- 1995a; 2000a; KÖHLER et al., 1995b; DE LA curs in the Chaco, whereas the nominal sub- RIVA, 2000. species would occur in N South America. Remarks: CANNATELLA (1983) synonymized The distribution of both subspecies seems to Phyllomedusa pailona Shreve (type locali- be disjunct. The variation and taxonomic ty: El Pailón, Santa Cruz) with P. boliviana status of Bolivian populations remain to be and stated that FUNKHOUSER’s (1957) pur- studied. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 39

Phyllomedusa palliata Peters, 1872 Cruz, Tarija; 3) Chiquitanía Forests, Chaco Distribution: 1) Upper Amazon Basin; 2) Be- Lowland Forests; 4) Fossorial, Terrestrial; ni, Cochabamba, La Paz, Pando; 3) Ama- 5) Open. zonian Rainforests, Humid Transition Low- Key literature: DE LA RIVA, 1990a; GON- land Forests; 4) Arboreal; 5) Forest. ZÁLES, 1998. Key literature: REICHLE & KÖHLER, 1996a; KÖHLER & LÖTTERS, 1999a. Chacophrys pierottii (Vellard, 1948) Distribution: 1) Chaco; 2) Chuquisaca, Santa Phyllomedusa sauvagii Boulenger, 1882 Cruz, Tarija; 3) Chaco Lowland Forests; Distribution: 1) Chaco; 2) Chuquisaca, Santa 4) Fossorial, Terrestrial; 5) Open. Cruz, Tarija; 3) Chaco Lowland Forests; Key literature: KÖHLER, 1997a. 4) Arboreal; 5) Open. Key literature: DE LA RIVA, 1990a; GON- Lepidobatrachus laevis Budgett, 1899 ZÁLES, 1998. Distribution: 1) Chaco; 2) Chuquisaca, Santa Cruz, Tarija; 3) Chaco Lowland Forests; Phyllomedusa tomopterna (Cope, 1868) 4) Aquatic, Fossorial, Terrestrial; 5) Open. Distribution: 1) Upper Amazon Basin; 2) Beni, Key literature: KÖHLER, 1997b. Cochabamba, La Paz, Pando; 3) Amazon- ian Rainforests; 4) Arboreal; 5) Forest. Remarks: This is the first record for the species in the country. Adenomera andreae Müller, 1923 Distribution: 1) Amazonia; 2) Beni, Cocha- Phyllomedusa vaillanti Boulenger, 1882 bamba, La Paz, Pando, Santa Cruz; 3) Ama- Distribution: 1) Upper Amazon Basin; 2) Beni, zonian Rainforests, Humid Transition Low- Cochabamba, La Paz, Pando, Santa Cruz; land Forests, Pre-Cambrian Shield Humid 3) Amazonian Rainforests, Humid Transi- Forests; 4) Terrestrial; 5) Forest. tion Lowland Forests, Pre-Cambrian Shield Key literature: DE LA RIVA, 1995b; MÁRQUEZ Humid Forests; 4) Arboreal; 5) Forest. et al., 1995. Key literature: DE LA RIVA, 1993a; DE LA RIVA et al., 1995. Adenomera diptyx (Boettger, 1885) Distribution: 1) Cerrado; 2) Chuquisaca, San- ta Cruz; 3) Cerrado, Chiquitanía Forests, Chaco Lowland Forests, Wet Savannas; 4) Te- Ceratophryinae rrestrial; 5) Open. Key literature: DE LA RIVA, 1995b; MÁRQUEZ Ceratophrys cornuta (Linnaeus, 1758) et al., 1995; DE LA RIVA, 1996. Distribution: 1) Amazonia; 2) Beni, Cocha- Remarks: This species is very similar to A. hy- bamba, La Paz, Pando, Santa Cruz; 3) Ama- laedactyla but differs in its reproductive zonian Rainforests, Humid Transition Low- mode (DE LA RIVA, 1995b). It enters the hu- land Forests, Pre-Cambrian Shield Humid mid Chaco. Its advertisement call was des- Forests; 4) Fossorial, Terrestrial; 5) Forest. cribed by MÁRQUEZ et al. (1995) as that of Key literature: DE LA RIVA, 1993a; MÁRQUEZ A. andreae. et al., 1995. Adenomera hylaedactyla (Cope, 1868) Ceratophrys cranwelli Barrio, 1980 Distribution: 1) Amazonia; 2) Beni, Cocha- Distribution: 1) Chaco; 2) Chuquisaca, Santa bamba, La Paz, Pando, Santa Cruz; 3) Ama- 40 DE LA RIVA et al.

zonian Rainforests, Humid Transition Low- Remarks: As stated by DE LA RIVA & MAL- land Forests, Pre-Cambrian Shield Humid DONADO (1999), with the present state of Forests, Wet Savannas; 4) Terrestrial; 5) knowledge L. chaquensis is indistinguisha- Open. ble from L. macrosternum Miranda-Ribeiro, Key literature: DE LA RIVA, 1995b; KÖHLER, which is distributed from N South America 1995a; 2000a; MÁRQUEZ et al., 1995; KÖH- to Brazil and Bolivia (FROST, 1985). The LER & BÖHME, 1996; REICHLE, 1997c. only revision of the L. ocellatus group was Remarks: See remarks under A. diptyx. carried out by GALLARDO (1964a), who re- ported L. chaquensis and L. macrosternum Leptodactylus bolivianus Boulenger, 1898* in Bolivia, but it is impossible to assign Distribution: 1) Tropical Middle and South these species to localities with certainty. America; 2) Beni, Cochabamba, La Paz, Provisionally, it can be assumed that Ama- Pando, Santa Cruz; 3) Amazonian Rain- zonian records (Beni, Cochabamba, La Paz, forests, Humid Transition Lowland Forests, northern Santa Cruz) correspond to L. ma- Pre-Cambrian Shield Humid Forests; 4) Ter- crosternum, whereas L. chaquensis is rather restrial; 5) Forest, Open. restricted to the Chaco (Chuquisaca, S San- Key literature: DE LA RIVA, 1993a; MÁRQUEZ ta Cruz, Tarija). However, the Chacoan and et al., 1995. Cerrado affinities of the Beni and E Santa Remarks: Leptodactylus insularum Barbour Cruz herpetofauna make this assumption from northern South America and Middle too feeble. The taxonomy of the pair of spe- America has been often considered a syn- cies L. chaquensis-L. macrosternum is chao- onym of L. bolivianus. If it proved to be a tic and needs revision urgently. different species, then the corresponding pattern of distribution of L. bolivianus would Leptodactylus didymus Heyer, García-López be Amazonian, instead of Tropical Middle & Cardoso, 1996 and South America. Distribution: 1) Southwestern Amazonia; 2) Beni, La Paz, Pando; 3) Amazonian Rain- Leptodactylus bufonius Boulenger, 1894 forests, Humid Transition Lowland Forests, Distribution: 1) Cerrado, Chaco; 2) Chuqui- Pre-Cambrian Shield Humid Forests; 4) Ter- saca, Santa Cruz, Tarija; 3) Chaco Low- restrial; 5) Forest. land Forests; 4) Terrestrial; 5) Open. Key literature: KÖHLER & LÖTTERS, 1999a. Key literature: HEYER, 1978; DE LA RIVA, Remarks: Leptodactylus didymus and L. mys- 1990a. taceus are sibling species barely distinguis- hable based on morphology. However, the calls Leptodactylus chaquensis Cei, 1950 of L. didymus are more similar to those of L. Distribution: 1) Cerrado, Chaco; 2) Beni, Co- elenae than to those of L. mystaceus (MÁR- chabamba, Chuquisaca, La Paz, Pando, QUEZ et al., 1995; HEYER et al., 1996). Santa Cruz, Tarija; 3) Cerrado, Chiquita- nía Forests, Chaco Lowland Forests, Chaco Leptodactylus elenae Heyer, 1978 Montane Forests, Humid Transition Low- Distribution: 1) Cerrado, Chaco; 2) Beni, Co- land Forests, Pre-Cambrian Shield Humid Fo- chabamba, La Paz, Pando, Santa Cruz; rests, Wet Savannas; 4) Terrestrial; 5) Open. 3) Cerrado, Chiquitanía Forests, Chaco Key literature: DE LA RIVA, 1990a; KÖHLER, Lowland Forests, Wet Savannas; 4) Terres- 1995a; 2000a; REICHLE, 1996; 1997c; KÖH- trial; 5) Open. LER & LÖTTERS, 1999a; DE LA RIVA & Key literature: DE LA RIVA, 1993a; KÖHLER, MALDONADO, 1999. 1995a; MÁRQUEZ et al., 1995. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 41

Leptodactylus fuscus (Schneider, 1799) tion Lowland Forests; 4) Terrestrial; 5) Fo- Distribution: 1) Middle and South America; rest. 2) Beni, Cochabamba, Chuquisaca, La Key literature: DE LA RIVA, 1990a. Paz, Pando, Santa Cruz, Tarija; 3) Ama- zonian Rainforests, Cerrado, Chiquitanía Leptodactylus labyrinthicus (Spix, 1824) Forests, Humid Transition Lowland Forests, Distribution: 1) Cerrado; 2) Beni, Cochabam- Pre-Cambrian Shield Humid Forests, Wet ba, Chuquisaca, Santa Cruz; 3) Cerrado, Savannas; 4) Terrestrial; 5) Open. Chiquitanía Forests, Pre-Cambrian Shield Key literature: REYNOLDS & FOSTER, 1992; Humid Forests; 4) Terrestrial; 5) Open. KÖHLER, 1995a; MÁRQUEZ et al., 1995; Key literature: DE LA RIVA, 1993a; KÖHLER, KÖHLER & BÖHME, 1996; REICHLE, 1996; 1995a; 2000a; MÁRQUEZ et al., 1995; KÖH- 1997c; REICHLE & KÖHLER, 1998. LER & BÖHME, 1996.

Leptodactylus gracilis Duméril & Bibron, Leptodactylus laticeps Boulenger, 1918 1841 Distribution : 1) Chaco; 2) Chuquisaca, Santa Distribution: 1) Subtropical South America; Cruz, Tarija; 3) Chaco Lowland Forests; 2) Cochabamba, Chuquisaca, Santa Cruz, 4) Terrestrial; 5) Open. Tarija; 3) Inter-Andean Valleys, Tucuman- Key literature: DE LA RIVA, 1990a. ian-Bolivian Montane Forests; 4) Terrestri- Remarks: FROST (1985) reported the species al; 5) Open. in Bolivia, but there is no published locality Key literature: KÖHLER, 1995a; 2000a; KÖH- hitherto. LER et al., 1995b; KÖHLER & LÖTTERS, 1999b. Leptodactylus latinasus Jiménez de la Espa- da, 1875 Leptodactylus griseigularis (Henle, 1981) Distribution: 1) Chaco; 2) Chuquisaca, Santa Distribution: 1) Humid Montane Forests, Peri- Cruz, Tarija; 3) Chaco Lowland Forests; Andean Forests; 2) Cochabamba, La Paz, 4) Terrestrial; 5) Open. Santa Cruz; 3) Amazonian Rainforests, Key literature: DE LA RIVA, 1990a. Yungas-Montane Rainforests; 4) Terrestrial; Remarks: GALLARDO (1964b) described the 5) Forest. subspecies L. l. anceps, which would be the Key literature: HEYER, 1994; KÖHLER, 2000a. one present in Bolivia; this subspecies was Remarks: HEYER (1994) split the former, invalidated by KLAPPENBACH & LANGONE widespread L. wagneri (Peters) into several (1992). species. As a result, L. wagneri no longer oc- curs in Bolivia. Leptodactylus griseigularis Leptodactylus leptodactyloides (Andersson, is the name applied to the populations of the 1945) Andean slopes and adjacent foothills, whe- Distribution: 1) Amazonia; 2) Beni, Cocha- reas L. leptodactyloides is the species wi- bamba, La Paz, Pando, Santa Cruz; 3) Ama- despread in the lowlands. Both species are zonian Rainforests, Chiquitanía Forests, Hu- mostly parapatric, but come into contact at mid Transition Lowland Forests, Pre-Cambrian the foothills of the Andes. Shield Humid Forests, Yungas-Montane Rain- forests; 4) Terrestrial; 5) Forest, Open. Leptodactylus knudseni Heyer, 1972 Key literature: DE LA RIVA, 1993a; HEYER, Distribution: 1) Upper Amazon Basin; 2) Beni, 1994; KÖHLER, 1995a; 1995b; 2000a; MÁR- Cochabamba, La Paz, Pando, Santa Cruz; QUEZ et al., 1995; REICHLE, 1996; 1997c; 3) Amazonian Rainforests, Humid Transi- REICHLE & KÖHLER, 1998. 42 DE LA RIVA et al.

Remarks: Most records compiled by DE LA America; 2) Beni, Cochabamba, La Paz, RIVA (1990a) as L. wagneri correspond to L. Pando, Santa Cruz; 3) Amazonian Rain- leptodactyloides. See remarks under L. gri- forests; 4) Terrestrial; 5) Forest. seigularis. Key literature: APARICIO, 1992.

Leptodactylus macrosternum Miranda-Ribei- Leptodactylus petersii (Steindachner, 1864) ro, 1926 Distribution: 1) Amazonia, Cerrado; 2) Beni, Distribution: 1) Amazonia; 2) Beni, Cocha- Cochabamba, La Paz, Pando, Santa Cruz; bamba, La Paz, Pando, Santa Cruz; 3) Ama- 3) Amazonian Rainforests, Humid Transi- zonian Rainforests, Humid Transition Low- tion Lowland Forests, Pre-Cambrian Shield land Forests; 4) Terrestrial; 5) Open. Humid Forests; 4) Terrestrial; 5) Forest. Key literature: DE LA RIVA & MALDONADO, Key literature: HEYER, 1994; REICHLE, 1996; 1999. 1997c; REICHLE & KÖHLER, 1998. Remarks: See remarks under L. chaquensis. Remarks: This species was formerly consid- ered a subspecies of L. podicipinus. Some Leptodactylus mystaceus (Spix, 1824) of the locality records provided by DE LA Distribution: 1) Amazonia; 2) Beni, Cocha- RIVA (1990a) for L. podicipinus might cor- bamba, La Paz, Pando, Santa Cruz; 3) Ama- respond to L. petersii. zonian Rainforests, Humid Transition Low- land Forests, Pre-Cambrian Shield Humid Leptodactylus podicipinus (Cope, 1862) Forests; 4) Terrestrial; 5) Forest, Open. Distribution: 1) Amazonia, Cerrado; 2) Beni, Key literature: DE LA RIVA, 1993a; MÁRQUEZ Cochabamba, La Paz, Pando, Santa Cruz; et al., 1995. 3) Amazonian Rainforests, Cerrado, Chiq- uitanía Forests, Humid Transition Lowland Leptodactylus mystacinus (Burmeister, 1861) Forests, Pre-Cambrian Shield Humid Fo- Distribution: 1) Cerrado, Chaco, Subtropical rests; 4) Terrestrial; 5) Open. South America; 2) Chuquisaca, Santa Cruz, Key literature: HEYER, 1994; KÖHLER, 1995a; Tarija; 3) Cerrado, Chaco Lowland Forests; MÁRQUEZ et al., 1995. 4) Terrestrial; 5) Open. Remarks: See remarks under P. petersii. Key literature: DE LA RIVA, 1990a. Leptodactylus rhodomystax Boulenger, 1884 Leptodactylus ocellatus (Linnaeus, 1758) Distribution: 1) Upper Amazon Basin; 2) Beni, Distribution: 1) Subtropical South America; La Paz, Pando; 3) Amazonian Rainforests, 2) Santa Cruz; 3) Wet Savannas; 4) Terres- Humid Transition Lowland Forests; 4) Ter- trial; 5) Open. restrial; 5) Forest. Key literature: DE LA RIVA & MALDONADO, Key literature: FUGLER, 1983. 1999. Remarks: Many old Bolivian records actually Leptodactylus rhodonotus (Günther, 1869) corresponding to the pair L. chaquensis-L. Distribution: 1) Humid Montane Forests, Peri- macrosternum were assigned to this species Andean Forests; 2) Cochabamba, La Paz, in the past. Its distribution in the country is Santa Cruz; 3) Amazonian Rainforests, discussed by DE LA RIVA & MALDONADO Yungas-Montane Rainforests; 4) Terrestrial; (1999). 5) Forest. Key literature: REYNOLDS & FOSTER, 1992; Leptodactylus pentadactylus (Laurenti, 1768) KÖHLER, 1995a; 2000a; KÖHLER & LÖT- Distribution: 1) Tropical Middle and South TERS, 1999b. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 43

Leptodactylus syphax Bokermann, 1969 two names represent different species. PER- Distribution: 1) Cerrado; 2) Santa Cruz; ACCA (1897) reported specimens of both 3) Cerrado, Chiquitanía Forests, Pre-Cam- species (as Paludicola fuscomaculata Stein- brian Shield Humid Forests; 4) Terrestrial; dachner and Paludicola signifera Girard) at 5) Open. some localities in Tarija, but their assign- Key literature: KÖHLER, 1995a; 1995b; KÖH- ment to one or the other species was not LER & BÖHME, 1996. convincing (CEI, 1990). The locality of the rediscovered Paraguayan sample studied Lithodytes lineatus (Schneider, 1799) by CEI (1990) and, especially, the type lo- Distribution: 1) Amazonia; 2) Beni, Cocha- cality of P. fuscomaculatus (“Caissará”, bamba, La Paz, Pando, Santa Cruz; 3) Ama- Mato Grosso do Sul, Brazil) are very close zonian Rainforests, Humid Transition Low- to the Bolivian border. Thus, the presence land Forests, Pre-Cambrian Shield Humid of the species in the country is very plausi- Forests, Yungas-Montane Rainforests; 4) Te- ble but, according to CEI (1990), P. fusco- rrestrial; 5) Forest. maculatus is less abundant than P. biligo- Key literature: DE LA RIVA, 1993a; MÁRQUEZ nigerus and has a distribution restricted et al., 1995. to the lower Paraguay basin. For this reason, the records from Tarija are doubtful and we Physalaemus albonotatus (Steindachner, 1864) refrain from including P. fuscomaculatus in Distribution: 1) Cerrado, Chaco; 2) Beni, Chu- the Bolivian fauna until conclusive eviden- quisaca, Santa Cruz, Tarija; 3) Cerrado, ce of its occurrence in the country is avai- Chiquitanía Forests, Chaco Lowland Fo- lable. rests, Chaco Montane Forests; 4) Terrestrial; 5) Open. Physalaemus cuqui Lobo, 1993 Key literature: DE LA RIVA, 1993a; KÖHLER, Distribution: 1) Chaco; 2) Chuquisaca, San- 1995a; 2000a; MÁRQUEZ et al., 1995. ta Cruz, Tarija; 3) Chaco Lowland Fo- Remarks: Many old records of this species rests, Chaco Montane Forests; 4) Terres- were given as P. cuvieri, which is very simi- trial; 5) Open. lar (they are indistinguishable in alcohol). Key literature: LAVILLA & SCROCCHI (1999). See remarks under P. cuqui. Remarks: This species, very similar to P. al- bonotatus, has been reported in the country Physalaemus biligonigerus (Cope, 1861 only recently. Some Chacoan records of P. “1860”) albonotatus might actually correspond to Distribution: 1) Cerrado, Chaco; 2) Chu- P. cuqui. quisaca, Santa Cruz, Tarija; 3) Cerrado, Chiquitanía Forests, Chaco Lowland Fo- Physalaemus cuvieri Fitzinger, 1826 rests, Chaco Montane Forests; 4) Terrestrial; Distribution: 1) Cerrado; 2) Santa Cruz; 3) Ce- 5) Open. rrado; 4) Terrestrial; 5) Open. Key literature: MÁRQUEZ et al., 1995; KÖH- Key literature: DE LA RIVA, 1990a. LER, 2000a. Remarks: See remarks on P. albonotatus. Remarks: This species has been mistaken for P. fuscomaculatus (Steindachner) in the lit- Physalaemus nattereri (Steindachner, 1863) erature for a long period. As a result, the ge- Distribution: 1) Cerrado; 2) Santa Cruz; 3) Ce- ographic distribution of both species is con- rrado; 4) Terrestrial; 5) Open. fused. These problems were discussed by Key literature: DE LA RIVA, 1990a; KÖHLER, CEI (1990), who clearly stated that these 1995a; MÁRQUEZ et al., 1995. 44 DE LA RIVA et al.

Physalaemus petersi (Jiménez de la Espada, to P. borellii have been recently found far 1872) away from the Andean foothills (CÉSPEDEZ, Distribution: 1) Amazonia; 2) Beni, Cocha- 1997). Admittedly, less evidence is available bamba, La Paz, Santa Cruz, Pando; 3) Ama- in Bolivia to either recognize or reject P. zonian Rainforests, Humid Transition Low- borellii as a valid species. Since PERACCA land Forests; 4) Terrestrial; 5) Forest. (1897) reported P. borellii in Tarija, we ten- Key literature: CANNATELLA & DUELLMAN, tatively recognize it as valid and include it in 1984; MÁRQUEZ et al., 1995. the list. Remarks: CANNATELLA & DUELLMAN (1984) placed Eupemphix freibergi Donoso-Barros Pleurodema cinereum Cope, 1877 [type locality: Runerrabaque (sic), Beni] in Distribution: 1) Andean Highlands, Temperate the synonymy of Physalaemus petersi, and Inter-Andean Valleys; 2) Cochabamba, Chu- in this were followed by subsequent authors. quisaca, La Paz, Oruro, Potosí; 3) High Recently, CANNATELLA et al. (1998), in a Andean Vegetation, Inter-Andean Valleys; paper dealing with the phylogeny of the en- 4) Terrestrial; 5) Open. tire P. pustulosus species group, again used Key literature: ERGUETA, 1991; VELLARD, the name Physalaemus freibergi, the distri- 1992; APARICIO, 1993; CHRISTMANN, 1995; bution of which would be the SW Amazon DE LA RIVA et al., 1995; IBISCH & BÖHME, in Peru and Bolivia. We prefer to maintain 1995; KÖHLER, 1995a; 2000a; KÖHLER the name P. petersi for these populations un- et al., 1995b; MÁRQUEZ et al., 1995; DE LA til further studies confirm their taxonomic RIVA & GONZÁLES, 1998. position. Remarks: See remarks under P. borellii.

Pleurodema borellii (Peracca, 1895) Pleurodema guayapae Barrio, 1964 Distribution: 1) Chaco, Temperate Inter-An- Distribution: 1) Chaco; 2) Chuquisaca, Santa dean Valleys; 2) Chuquisaca, Tarija; 3) Cha- Cruz, Tarija; 3) Chaco Lowland Forests; 4) co Montane Forests, Inter-Andean Valleys, Terrestrial; 5) Open. Tucumanian-Bolivian Montane Forests; 4) Te- Key literature: DE LA RIVA & GONZÁLES, rrestrial; 5) Open. 1998; GONZÁLES, 1998. Key literature: PERACCA, 1897; DE LA RIVA & Remarks: Whereas the presence of this species GONZÁLES, 1998. in poorly surveyed areas of Chuquisaca and Remarks: This species has been considered Tarija is plausible, the Chacoan regions of either as a good species or as a synonym of northern Argentina are well studied and the P. cinereum, primarily on the basis of their occurrence of the species there is discarded. almost identical morphology. However, it The perplexing distribution of this species, seems odd that a species may range from the with a gap of 1000 km between the Bolivian hot, lowland Chaco to the high, cold puna and Argentinean populations (which live in (that is, from 400 to 4500 m or more), an ar- salty, arid environments), suggests that they gument sometimes utilized to justify the might represent different species, although recognition of the two species (DUELLMAN the morphological evidence does not sup- & VELOSO, 1977). However, the bioche- port such an assumption (DE LA RIVA & GON- mical and biacoustical evidence seems to ZÁLES, 1998). indicate that only one species might be in- volved (MCLISTER et al., 1991). Argenti- Pleurodema marmoratum (Duméril & Bi- nean herpetologists tend to recognize two bron, 1841)* separate species, and populations assigned Distribution: 1) Andean Highlands; 2) Co- TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 45

chabamba, La Paz, Oruro, Potosí; 3) High La Paz, Pando; 3) Amazonian Rainforests; Andean Vegetation; 4) Terrestrial; 5) Open. 4) Terrestrial; 5) Forest. Key literature: ERGUETA, 1991; REYNOLDS & Key literature: KÖHLER & LÖTTERS, 1999a. FOSTER, 1992; VELLARD, 1992; APARICIO, 1993. Telmatobiinae

Pseudopaludicola boliviana Parker, 1927* Eleutherodactylus ashkapara Köhler, 2000** Distribution: 1) Tropical South America; 2) Be- Distribution: 1) Humid Montane Forests; 2) Co- ni, Cochabamba, La Paz, Santa Cruz; 3) Ce- chabamba, La Paz, Santa Cruz; 3) Ceja de rrado, Chiquitanía Forests, Humid Tran- Montaña-Cloud Forests, Yungas-Montane sition Lowland Forests, Pre-Cambrian Shield Rainforests; 4) Arboreal; 5) Forest. Humid Forests, Wet Savannas; 4) Terrestri- Key literature: KÖHLER, 2000a; 2000b. al; 5) Open. Key literature: KÖHLER, 1995a; MÁRQUEZ Eleutherodactylus cruralis (Boulenger, 1902)* et al., 1995; REICHLE, 1996; 1997c; REICH- Distribution: 1) Bolivian-Argentinean Fo- LE & KÖHLER, 1998. rests, Humid Montane Forests, Peri-Andean Remarks: LYNCH (1989a) reviewed the Pseu- Forests; 2) Cochabamba, Chuquisaca, La dopaludicola of N South America, and he Paz, Santa Cruz; 3) Amazonian Rainfo- could not find differences between speci- rests, Tucumanian-Bolivian Montane Forests, mens from eastern Colombia, southern Ve- Yungas-Montane Rainforests; 4) Terrestrial, nezuela, and Bolivia. The currently accep- Arboreal; 5) Forest. ted distribution of P. boliviana comprises Key literature: LYNCH, 1989b; DE LA RIVA, eastern Colombia, S Venezuela, Guyana, 1993c; MÁRQUEZ et al., 1995; KÖHLER, Surinam, Roraima (N Brazil), E Bolivia, Pa- 1995a; 2000a; KÖHLER et al., 1995b; REICH- raguay, SW Brazil and N Argentina (LYNCH, LE & KÖHLER, 1997; REICHLE et al., 2001. 1989a; LOBO, 1992; 1994). Thus, there are Remarks: LYNCH (1989b) stated that there is two sets of populations separated by the Ama- remarkable variation in the size of males and zon Basin. This strange distribution roughly females across the range of the species, a parallels that of Phyllomedusa hypochon- fact that might indicate that more than one drialis. Although it is not necessarily un- species is involved under the name E. cru- tenable that both sets of populations of Pseu- ralis. LAVILLA & SCROCCHI (1999) report dopaludicola boliviana are conspecific, fur- two unidentified species of the E. discoidalis ther research is necessary to ascertain their group in Tarija. In our opinion, the morpho- phylogenetic relationships and taxonomic logical variation in samples identified as E. status, and perhaps the northern populations cruralis from lowland rainforests in SW might deserve to be considered a different Peru and central Bolivia, as well as in those species. from cloud forests and temperate valleys, are incompatible with the idea of a single Pseudopaludicola mystacalis (Cope, 1887) species. Bioacoustical evidence also sup- Distribution: 1) Cerrado; 2) Beni, Santa Cruz; ports the presence of more than one species. 3) Cerrado, Wet Savannas; 4) Terrestrial; 5) A new species is being described by REICH- Open. LE et al. (2001) and three others will be de- Key literature: DE LA RIVA, 1990a. scribed by M. Harvey and R. Aguayo.

Vanzolinius discodactylus (Boulenger, 1883) Eleutherodactylus danae Duellman, 1978 Distribution: 1) Upper Amazon Basin; 2) Beni, Distribution: 1) Humid Montane Forests, Peri- 46 DE LA RIVA et al.

Andean Forests; 2) Cochabamba, La Paz, Eleutherodactylus fraudator Lynch & McDi- Santa Cruz; 3) Amazonian Rainforests, armid, 1987** Yungas-Montane Rainforests; 4) Arboreal; Distribution: 1) Humid Montane Forests; 2) Co- 5) Forest. chabamba, La Paz, Santa Cruz; 3) Ceja de Key literature: KÖHLER, 1995a; 2000a; KÖH- Montaña-Cloud Forests, Yungas-Montane LER & JUNGFER, 1995. Rainforests; 4) Terrestrial; 5) Forest. Key literature: LYNCH & MCDIARMID, 1987; Eleutherodactylus discoidalis (Peracca, 1895) DE LA RIVA, 1993c; KÖHLER, 1995a; 2000a; Distribution: 1) Bolivian-Argentinean Forests, KÖHLER et al., 1995a; DE LA RIVA & Humid Montane Forests; 2) Cochabamba, LYNCH, 1997. Chuquisaca, Santa Cruz, Tarija; 3) Tucu- manian-Bolivian Montane Forests, Yungas- Eleutherodactylus llojsintuta Köhler & Löt- Montane Rainforests; 4) Terrestrial, Arbore- ters, 1999** al; 5) Forest. Distribution: 1) Humid Montane Forests; 2) Co- Key literature: DE LA RIVA, 1993c; HARVEY, chabamba, La Paz, Santa Cruz; 3) Yun- 1997. gas-Montane Rainforests; 4) Arboreal; 5) Fo- Remarks: DE LA RIVA (1990a; 1993c) over- rest. looked the old record of this species in Tari- Key literature: KÖHLER & LÖTTERS, 1999c; ja by PERACCA (1897). See remarks under KÖHLER, 2000a. E. cruralis. Remarks: See remarks under E. platydacty- lus. Eleutherodactylus fenestratus (Steindachner, 1864) Eleutherodactylus mercedesae Lynch & Mc- Distribution: 1) Amazonia; 2) Beni, Cocha- Diarmid, 1987** bamba, La Paz, Pando, Santa Cruz; 3) Ama- Distribution: 1) Humid Montane Forests; 2) Co- zonian Rainforests, Humid Transition Low- chabamba, La Paz, Santa Cruz; 3) Yungas- land Forests, Pre-Cambrian Shield Humid Montane Rainforests; 4) Terrestrial; 5) For- Forests, Yungas-Montane Rainforests; 4) Ter- est. restrial, Arboreal; 5) Forest. Key literature: LYNCH & MCDIARMID, 1987; Key literature: REYNOLDS & FOSTER, 1992; DE LA RIVA, 1993c; KÖHLER, 2000a. DE LA RIVA, 1993c; MÁRQUEZ et al., 1995; REICHLE, 1999; KÖHLER, 2000a. Eleutherodactylus olivaceus Köhler, Morales, Remarks: Many locality records assigned to Lötters, Reichle & Aparicio, 1998* this species are probably in error. Conside- Distribution: 1) Humid Montane Forests, Peri- rable confusion exists regarding the taxo- Andean Forests; 2) Cochabamba, La Paz, nomic status of some Peruvian and Bolivian Santa Cruz; 3) Amazonian Rainforests, populations of in the E. conspicillatus Yungas-Montane Rainforests; 4) Arboreal; group, which have similar morphologies, 5) Forest. moderate to broad distributions, and poorly Key literature: DE LA RIVA, 1993c; KÖHLER et known regional variation. After REICHLE al., 1998b; KÖHLER, 2000a. (1999) Hylodes gollmeri bisignatus Werner Remarks: This recently described species in- [type locality: Chaco, Bolivia (a cloud for- cludes the populations on which former Bo- est locality in Departamento La Paz)], cur- livian records of E. mendax Duellman were rently in the synonymy of E. fenestratus, is based (see DE LA RIVA, 1993c). However, a different species whose taxonomic posi- the presence of E. mendax in the country is tion is still under study (see KÖHLER, 2000a). still plausible. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 47

Eleutherodactylus peruvianus (Melin, 1941) Distribution: 1) Humid Montane Forests; Distribution: 1) Humid Montane Forests, Peri- 2) Cochabamba, La Paz, Santa Cruz; Andean Forests, Upper Amazon Basin; 2) Co- 3) Yungas-Montane Rainforests; 4) Terres- chabamba, La Paz, Santa Cruz; 3) Ama- trial; 5) Forest. zonian Rainforests, Yungas-Montane Rain- Key literature: KÖHLER, 1995a; 2000a; KÖH- forests; 4) Terrestrial, Arboreal; 5) Forest. LER et al., 1995a; DE LA RIVA & LYNCH, Key literature: DE LA RIVA, 1994b; KÖHLER, 1997. 1995a; 2000a. Remarks: Unlike most Peruvian specimens of Eleutherodactylus rhabdolaemus Duellman, E. peruvianus, Bolivian ones often lack or- 1978 ange spots on the posterior surfaces of the Distribution: 1) Humid Montane Forests; thighs. There is a considerable variation in 2) Cochabamba, La Paz, Santa Cruz; these frogs and, given the fact that the type 3) Yungas-Montane Rainforests; 4) Terres- locality of the species (Roque, Department trial, Arboreal; 5) Forest. of San Martín, Peru) is far away from Bo- Key literature: LYNCH & MCDIARMID, 1987; livia, it would not be surprising that some REYNOLDS & FOSTER, 1992; DE LA RIVA, Bolivian populations represent a different 1993c; KÖHLER, 1995a; 2000a. species (see KÖHLER, 2000a). Eleutherodactylus samaipatae Köhler & Eleutherodactylus platydactylus (Boulenger, Jungfer, 1995** 1903) Distribution: 1) Bolivian-Argentinean Forests, Distribution: 1) Humid Montane Forests; 2) Temperate Inter-Andean Valleys; 2) Co- Cochabamba, La Paz, Santa Cruz; 3) Ceja chabamba, Chuquisaca, Santa Cruz, Tari- de Montaña-Cloud Forests, Yungas-Mon- ja; 3) Inter-Andean Valleys, Tucumanian- tane Rainforests; 4) Arboreal; 5) Forest. Bolivian Montane Forests; 4) Terrestrial, Key literature: DE LA RIVA, 1993c; KÖHLER, Arboreal; 5) Forest, Open. 1995a; 2000a; KÖHLER et al., 1995a; MÁR- Key literature: KÖHLER, 1995a; 2000a; KÖH- QUEZ et al., 1995; DE LA RIVA, 1997; KÖHLER LER & JUNGFER, 1995; KÖHLER et al., & LÖTTERS, 1999c. 1995b; REICHLE, 1999. Remarks: Recently, DE LA RIVA (1997) placed E. bockermanni Donoso-Barros [type local- Eleutherodactylus toftae Duellman, 1978 ity: Runerrabaque (sic), Beni (probably in Distribution: 1) Peri-Andean Forests; 2) Co- error)] in the synonymy of E. platydactylus. chabamba, La Paz, Santa Cruz; 3) Ama- The striking variation of this species has zonian Rainforests; 4) Terrestrial, Arboreal; been pointed out and illustrated repeatedly, 5) Forest. and it has been suggested that more than Key literature: DE LA RIVA, 1993c; MÁRQUEZ one species might be hidden under the name et al., 1995. E. platydactylus. DE LA RIVA (1997) refrai- ned from naming new species but described Eleutherodactylus ventrimarmoratus (Bou- three morphs which might prove to be valid lenger, 1912) species. Eleutherodactylus llojsintuta is a Distribution: 1) Humid Montane Forests, Up- sibling species described by KÖHLER & per Amazon Basin; 2) Beni, La Paz, Pando; LÖTTERS (1999c) based mainly on differ- 3) Amazonian Rainforests, Yungas-Mon- ences in the advertisement calls. tane Rainforests; 4) Arboreal; 5) Forest. Eleutherodactylus pluvicanorus De la Riva Remarks: Reported herein for the first time & Lynch, 1997** for Bolivia. 48 DE LA RIVA et al.

Eleutherodactylus zongoensis Reichle & still unclear, and both forms seem to be Köhler, 1997** sympatric in some areas. Distribution: 1) Humid Montane Forests; 2) La Paz; 3) Yungas-Montane Rainforests; 4) Te- Phrynopus kempffi De la Riva, 1992** rrestrial; 5) Forest. Distribution: 1) Humid Montane Forests; 2) Co- Key literature: REICHLE & KÖHLER, 1997. chabamba, Santa Cruz; 3) Ceja de Mon- Remarks: Known only from the holotype. taña-Cloud Forests; 4) Terrestrial; 5) Forest. Key literature: DE LA RIVA, 1992b; MÁRQUEZ Ischnocnema quixensis (Jiménez de la Espa- et al., 1995. da, 1872) Distribution: 1) Amazonia; 2) Beni, La Paz, Phrynopus laplacai (Cei, 1968)** Pando; 3) Amazonian Rainforests; 4) Ter- Distribution: 1) Humid Montane Forests; 2) restrial; 5) Forest. La Paz; 3) Ceja de Montaña-Cloud Forests, Remarks: Found by J. E. Cadle and S. Reichle Yungas-Montane Rainforests; 4) Terrestrial; in November 1999 and reported herein for 5) Forest, Open. the first time. Key literature: CEI, 1968; LYNCH, 1975; ER- GUETA, 1993. Ischnocnema sanctaecrucis Harvey & Keck, 1995** Phrynopus pinguis Harvey & Ergueta, 1998** Distribution: 1) Humid Montane Forests; 2) Co- Distribution: 1) Humid Montane Forests; 2) chabamba, La Paz, Santa Cruz; 3) Yungas- La Paz; 3) Ceja de Montaña-Cloud Forests; Montane Rainforests; 4) Terrestrial; 5) For- 4) Terrestrial; 5) Forest. est. Key literature: HARVEY & ERGUETA, 1998. Key literature: HARVEY & KECK, 1995; REI- Remarks: The genus Phrynopus seems to be CHLE, 1999; KÖHLER, 2000a. much more speciose in Bolivia than previ- ously expected. Two new species are being Odontophrynus americanus (Duméril & Bi- described by R. Aguayo and M. Harvey, and bron, 1841) some more will be described by I. De la Distribution: 1) Subtropical South America; Riva and S. Reichle. 2) Cochabamba, Chuquisaca, Santa Cruz, Tarija; 3) Chaco Lowland Forests, Chaco Phyllonastes carrascoicola De la Riva & Montane Forests; 4) Fossorial, Terrestrial; Köhler, 1998** 5) Open. Distribution: 1) Humid Montane Forests; Key literature: DE LA RIVA, 1990a; MÁRQUEZ 2) Cochabamba, Santa Cruz; 3) Ceja de et al., 1995; KÖHLER, 2000a. Montaña-Cloud Forests, Yungas-Montane Rainforests; 4) Terrestrial; 5) Forest. Odontophrynus lavillai Cei, 1985 Key literature: REYNOLDS & FOSTER, 1992; Distribution: 1) Chaco, Temperate Inter-An- KÖHLER, 1995a; 2000a; 2000c; KÖHLER et dean Valleys; 2) Chuquisaca, Santa Cruz, al., 1995a; DE LA RIVA & KÖHLER, 1998. Tarija; 3) Chaco Lowland Forests, Inter- Andean Valleys; 4) Fossorial, Terrestrial; Phyllonastes ritarasquinae Köhler, 2000** 5) Open. Distribution: 1) Humid Montane Forests; 2) Co- Key literature: DE LA RIVA et al., 1996c; chabamba, La Paz, Santa Cruz; 3) Yungas- KÖHLER, 2000a. Montane Rainforests; 4) Terrestrial; 5) For- Remarks: The distribution and variation of O. est. americanus and O. lavillai in the country is Key literature: KÖHLER, 2000a; 2000c. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 49

Remarks: More species of the genus Phyl- Paz; 3) High Andean Vegetation; 4) Aquat- lonastes [or a closely related minute lepto- ic; 5) Open. dactylid genus (M. B. Harvey, in litt.)] are Key literature: BARBOUR & NOBLE, 1920; expected to be found in the country as clos- PARKER, 1940; VELLARD, 1951; 1953; er attention be paid to the forest litter am- 1955; 1960; DE MACEDO, 1960; CEI, 1986; phibian fauna. VELLARD, 1992; PÉREZ, 1998. Remarks: The description of subspecies on Telmatobius bolivianus Parker, 1940** the basis of morphological differences has Distribution: 1) Humid Montane Forests; 2) La created a taxonomic chaos in some species Paz; 3) Ceja de Montaña-Cloud Forests, groups of Telmatobius. Up to six subspecies Yungas-Montane Rainforests; 4) Aquatic; of T. culeus have been recognized. The nom- 5) Forest. inal subspecies has its type locality in Bo- Key literature: PARKER, 1940; VELLARD, 1951; livia [Achacache, Lago Titicaca (La Paz) 1970; LAVILLA & DE LA RIVA, 1993. (FROST, 1985) or “Proximidades de la Isla Remarks: The taxonomy of the Yungas Telma- del Sol” (La Paz) (VELLARD, 1953)]. Fur- tobius is obscured by historical problems re- thermore, VELLARD (1951; 1960) described lated to unclear type localities, state of flux three subspecies of T. albiventris Parker, a in relation to synonymizations and revalida- species that he later placed in the synonymy tions, the intrinsic difficulty posed by the of T. culeus, but he still recognized all the frogs themselves due to their intra- and in- former subspecies of T. albiventris (VEL- terpopulational variation, and the scarcity of LARD, 1992). One of these subspecies, T. a. museum specimens. Telmatobius bolivianus globulosus Vellard, has its type locality in was placed in the synonymy of T. verruco- Bolivia [Playa de Copani, Huiñamarca (La sus Werner by VELLARD (1951), who later Paz)]. As a result, the number of subspecies considered both forms as subspecies of T. of T. culeus rose to twelve. Fortunately, a marmoratus (VELLARD, 1970). DE LA RIVA thorough study carried out by SINSCH et al. (1990a) considered doubtful the validity of (1995) demonstrated that the morphological T. verrucosus (and, as a consequence, that of variation in Telmatobius does not match T. bolivianus). Later, LAVILLA & DE LA subspecific partitions like this. We follow RIVA (1993) described the putative tadpole VELLARD (1992) in that T. albiventris is a of T. bolivianus. The tadpole was assigned synonym of T. culeus. to this species on the basis of the compari- son of some adult frog specimens (for which Telmatobius edaphonastes De la Riva, 1995 the tadpole was known) with the holotype of “1994” ** T. bolivianus. Further observations led to Distribution: 1) Humid Montane Forests; 2) conclude that more than one species is in- Cochabamba, Santa Cruz; 3) Ceja de volved and the tadpoles were wrongly as- Montaña-Cloud Forests, Yungas-Montane signed (I. De la Riva, pers. obs.). Telmato- Rainforests; 4) Aquatic, Terrestrial; 5) For- bius bolivianus and T. verrucosus seem to est. be different species (but see remarks under Key literature: DE LA RIVA (1995c). T. verrucosus) and at least there is another related species, which is being described by Telmatobius huayra Lavilla & Ergueta, 1995** I. De la Riva. Distribution: 1) Andean Highlands; 2) Potosí; 3) High Andean Vegetation; 4) Aquatic; 5) Telmatobius culeus (Garman, 1875)* Open. Distribution: 1) Andean Highlands; 2) La Key literature: LAVILLA & ERGUETA, 1995a. 50 DE LA RIVA et al.

Telmatobius ifornoi Lavilla & Ergueta, 1999** Key literature: PARKER, 1940; VELLARD, Distribution: 1) Humid Montane Forests; 2) 1951. La Paz; 3) Ceja de Montaña-Cloud Forests, Remarks: Certain populations of cloud for- Yungas-Montane Rainforests; 4) Aquatic; est Telmatobius in Cochabamba and Santa 5) Forest. Cruz were tentatively identified as T. si- Key literature: LAVILLA & ERGUETA, 1999. monsi (DE LA RIVA, 1990a; 1994c). These populations actually represent an un- Telmatobius jahuira Lavilla & Ergueta, named species, which is being described 1995** by I. De la Riva and M. Harvey, who are Distribution: 1) Humid Montane Forests; 2) also reviewing the taxonomic status of La Paz; 3) Ceja de Montaña-Cloud Forests, some other populations allied to T. simon- Yungas-Montane Rainforests; 4) Aquatic, si from the temperate valleys of Santa Terrestrial; 5) Forest. Cruz. Key literature: LAVILLA & ERGUETA, 1995b. Telmatobius verrucosus Werner, 1899** Telmatobius marmoratus (Duméril & Bi- Distribution: 1) Humid Montane Forests; 2) bron, 1841) La Paz; 3) Ceja de Montaña-Cloud Forests, Distribution: 1) Andean Highlands; 2) Cocha- Yungas-Montane Rainforests; 4) Aquatic; bamba, Chuquisaca, La Paz, Oruro, Po- 5) Forest. tosí; 3) High Andean Vegetation, Inter-An- Key literature: WERNER, 1899; BARBOUR & dean Valleys; 4) Aquatic; 5) Open. NOBLE, 1920; VELLARD, 1951; 1970. Key literature: PARKER, 1940; VELLARD, Remarks: This species has not been found 1951; 1953; 1955; 1960a; 1969; 1970; again, and its uncertain taxonomic status is 1992; CEI, 1986; ERGUETA, 1991; APARI- being investigated by I. De la Riva. See re- CIO, 1993; IBISCH & BÖHME, 1993; KÖH- marks under T. bolivianus. LER, 1995a. Remarks: This is the most widespread species Telmatobius yuracare De la Riva, 1994** of Telmatobius and the one with the most Distribution: 1) Humid Montane Forests; 2) complicated taxonomy. Once T. bolivianus Cochabamba, Santa Cruz; 3) Ceja de is resurrected as a valid species, still nine Montaña-Cloud Forests, Yungas-Montane subspecies of T. marmoratus remain. Two of Rainforests; 4) Aquatic; 5) Forest. them have their type locality in Bolivia: T. Key literature: DE LA RIVA, 1994c; KÖHLER, m. hintoni Parker (Tolota, Cochabamba) and 1995a; 2000a; KÖHLER et al., 1995a. T. m. gigas Vellard (Huayllamarca, Oruro). Furthermore, the nominal subspecies and T. Microhylidae m. rugosus Vellard would also occur in the country. The validity of all these subspecies Microhylinae and the real specific diversity of the T. mar- moratus complex in Bolivia is being re- Chiasmocleis albopunctata (Boettger, 1885) viewed by I. De la Riva. Distribution: 1) Cerrado; 2) Santa Cruz; 3) Cerrado, Chiquitanía Forests, Chaco Low- Telmatobius simonsi Parker, 1940** land Forests, Chaco Montane Forests, Pre- Distribution: 1) Temperate Inter-Andean Val- Cambrian Shield Humid Forests; 4) Terres- leys; 2) Cochabamba, Chuquisaca, Potosí, trial; 5) Open. Santa Cruz, Tarija; 3) Inter-Andean Val- Key literature: DE LA RIVA, 1993a; DE LA leys; 4) Aquatic; 5) Open. RIVA et al., 1996c; KÖHLER, 2000a. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 51

Chiasmocleis ventrimaculata (Andersson, Santa Cruz; 3) Amazonian Rainforests, 1945) Cerrado, Chiquitanía Forests, Chaco Low- Distribution: 1) Upper Amazon Basin; 2) Beni, land Forests, Chaco Montane Forests, Hu- Cochabamba, La Paz, Pando, Santa Cruz; mid Transition Lowland Forests, Inter-An- 3) Amazonian Rainforests, Humid Transi- dean Valleys, Pre-Cambrian Shield Humid tion Lowland Forests; 4) Terrestrial; 5) For- Forests, Tucumanian-Bolivian Montane est. Forests, Wet Savannas; 4) Fossorial, Terres- Key literature: DE LA RIVA, 1995d. trial; 5) Open. Remarks: WALKER (1973) reported a speci- Key literature: DE LA RIVA, 1993a; DE LA men of C. bassleri Dunn from Bolivia. So RIVA et al., 1996c; REICHLE, 1997c; KÖH- far, C. bassleri is known from E Peru and LER, 2000a. Ecuador (FROST, 1985), but new findings in Remarks: See comments under E. bicolor. Brazil, close to the Bolivian border, make Walker’s report plausible (S. Reichle, pers. Hamptophryne boliviana (Parker, 1927)* obs.). Distribution: 1) Upper Amazon Basin; 2) Beni, Cochabamba, La Paz, Pando, Santa Dermatonotus muelleri (Boettger, 1855) Cruz; 3) Amazonian Rainforests, Humid Distribution: 1) Chaco, Cerrado; 2) Chuqui- Transition Lowland Forests, Pre-Cambrian saca, Santa Cruz, Tarija; 3) Chaco Low- Shield Humid Forests; 4) Terrestrial; 5) For- land Forests; 4) Fossorial, Terrestrial; 5) Open. est. Key literature: DE LA RIVA, 1990a. Key literature: DE LA RIVA, 1993a; DE LA RIVA et al., 1996c. Elachistocleis bicolor (Valenciennes, 1838) Distribution: 1) Cerrado, Chaco, South- Pipidae western Amazonia; 2) Beni, Cochabamba, La Paz, Pando, Santa Cruz; 3) Amazon- Pipinae ian Rainforests, Chiquitanía Forests, Cha- co Lowland Forests, Chaco Montane Fo- Pipa pipa (Linnaeus, 1758) rests, Humid Transition Lowland Forests, Distribution: 1) Amazonia; 2) Beni, Cocha- Pre-Cambrian Shield Humid Forests, Wet bamba, La Paz, Pando, Santa Cruz; 3) Savannas; 4) Fossorial, Terrestrial; 5) Amazonian Rainforests, Humid Transition Open. Lowland Forests, Pre-Cambrian Shield Hu- Key literature: KÖHLER, 1995a; 2000a; mid Forests; 4) Aquatic; 5) Forest, Open. 1995b; DE LA RIVA et al., 1996c; REICHLE, Key literature: REICHLE, 1997a; HARVEY et 1996; 1997c; REICHLE & KÖHLER, 1998. al., 1998. Remarks: Considerable confusion took place in the past regarding the identity of E. bicol- Pseudidae or vs. E. ovalis, whose main external differ- ence is the ventral coloration (see FROST, Lysapsus limellus Cope, 1862 1985). Today most authors apply the name Distribution: 1) Cerrado, Lower Amazon E. bicolor to the frogs with an immaculate Basin; 2) Beni, Cochabamba, La Paz, San- venter. ta Cruz, Pando; 3) Cerrado, Wet Savannas; 4) Aquatic; 5) Open. Elachistocleis ovalis (Schneider, 1799) Key literature: KÖHLER, 1995a; BOSCH et al., Distribution: 1) South America; 2) Beni, Co- 1996; REICHLE, 1996; 1997a; REICHLE & chabamba, Chuquisaca, La Paz, Pando, KÖHLER, 1998. 52 DE LA RIVA et al.

Remarks: The taxonomy of L. limellus was Ranidae reviewed by GALLARDO (1961b; 1964c) but the status of several populations re- Raninae mains problematic. In Bolivia the sub- species L. limellus bolivianus Gallardo Rana palmipes Spix,1824 (type locality: Reyes, Beni) occurs, and pro- Distribution: 1) Amazonia; 2) Beni, Cocha- bably L. l. limellus, whose type locality bamba, La Paz, Pando, Santa Cruz; 3) Ama- (Corumbá, Mato Grosso do Sul) lies on the zonian Rainforests, Humid Transition Low- Brazilian bank of the Paraguay river, which land Forests, Pre-Cambrian Shield Humid divides the Brazilian and Bolivian Pan- Forests; 4) Terrestrial; 5) Forest. tanal. KLAPPENBACH (1985) stated that L. Key literature: DE LA RIVA, 1990a. l. laevis Parker, from the lower Amazon, should be considered a good species, and Caudata suggested that L. l. bolivianus might be ei- Plethodontidae ther a valid species or a subspecies of L. laevis. At least two different morphs corre- Plethodontinae sponding to two kinds of advertisement calls are found in the Beni area (S. Reich- Bolitoglossa altamazonica (Cope, 1874) le, pers. obs.). Distribution: 1) Amazonia; 2) Beni, Cocha- bamba, La Paz, Santa Cruz, Pando; 3) Ama- Pseudis paradoxa (Linnaeus, 1758) zonian Rainforests, Yungas-Montane Rain- Distribution: 1) South America; 2) Beni, forests; 4) Arboreal, Terrestrial; 5) Forest. Cochabamba, Chuquisaca, La Paz, Pando, Key literature: REICHLE et al., 2000. Santa Cruz; 3) Cerrado, Chiquitanía Fo- Remarks: Bolitoglossa altamazonica is thought rests, Chaco Lowland Forests, Humid Tran- to include populations from the mouth of the sition Lowland Forests, Pre-Cambrian Shield Amazon to the Andean cloud forests from Humid Forests, Wet Savannas; 4) Aquatic; Colombia to Bolivia (there are scant records 5) Open. from the central Amazon). This is one of the Key literature: BOSCH et al., 1996; REICHLE, least known species complex of South 1996; 1997a; REICHLE & KÖHLER, 1998; American amphibians, and it is expected DE LA RIVA, 1999d. that, upon future research, the Bolivian pop- Remarks: Three subspecies have been report- ulations will be identified as one or more ed in the country: P. p. paradoxa (Linnaeus) different species (REICHLE et al., 2000). in the Beni region, P. p. platensis Gallardo in the Cerrado and P. p. occidentalis Gallar- Gymnophiona do (type locality: El Pailón, Santa Cruz) in the Chaco (GALLARDO, 1961b; 1964c). As Caeciliidae stated by BOSCH et al. (1996) and DE LA RIVA (1999d), the validity and distribution Caecilia marcusi Wake, 1984** of these forms in the country needs to be Distribution: 1) Peri-Andean Forests, South- studied, especially the presence of the nom- western Amazonia; 2) Beni, Cochabamba, inal subspecies, whose core area of distribu- La Paz, Santa Cruz; 3) Amazonian Rain- tion is the Guianan region and for which forests, Humid Transition Lowland Forests; there are no records in the intervening Ama- 4) Fossorial; 5) Forest. zonian region. Key literature: WAKE, 1984; REICHLE & KÖH- LER, 1996b. TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 53

Siphonops annulatus (Mikan, 1820) Species deletions.-There are some species Distribution: 1) Tropical South America; 2) deleted from the 1990 list but, due to taxo- Beni, Cochabamba, La Paz, Pando, Santa nomic or nomenclatural reasons, they have Cruz; 3) Amazonian Rainforests, Humid been replaced by another species’ name (e. g., Transition Lowland Forests, Pre-Cambrian Scinax parkeri vs. S. fuscomarginatus). In Shield Humid Forests; 4) Fossorial; 5) For- other cases, they have been identified as new est. species and described (e. g., Eleutherodacty- Key literature: DE LA RIVA, 1990a. lus olivaceus vs. E. mendax). These cases do not alter the number of species. However, Siphonops paulensis Boettger, 1892 there are two cases of species present in the Distribution: 1) Cerrado; 2) Beni, Santa 1990 list that have been completely removed Cruz; 3) Cerrado, Chiquitanía Forests, Cha- from the present one: the hylids Hemiphractus co Lowland Forests; 4) Fossorial; 5) Open. johnsoni (Noble) and Hyla rossalleni. The Key literature: DE LA RIVA, 1990a. case of H. rossalleni has been already com- mented in the remarks section of Hyla leali. CONCLUSIONS AND DISCUSSION There are two references suggesting the pres- ence of Hemiphractus johnstoni in Bolivia. Taxonomic diversity and changes since BOULENGER (1903a) treated the reproduction 1990. of this species (as Ceratohyla bubalus) based In total, up to date, there are three orders, on a specimen obtained by Mr. Ockenden at eleven families, 44 genera and 186 valid species Santo Domingo (Puno, Peru) and commented of amphibians recognized from Bolivia. Their that it is “...an inhabitant of the Andes of taxonomic distribution is summarized in Ecuador, Bolivia and Peru”. However, as far Table 1. These figures represent a dramatic in- as we know, no voucher specimens from Bo- crease with respect to DE LA RIVA‘s (1990a) livia are available. TRUEB (1974) listed a pur- list. Frogs belonging to nine genera not previ- ported Bolivian specimen of H. johnstoni ously reported in the country have been found from “Linimbane”, 1000 m. [housed at the (either as undescribed species or as already Natural History Museum, London (BM)]. The known ones): Dendrophryniscus, Melanophry- name of this locality is an obvious mispelling niscus, Allobates, Scarthyla, Chacophrys, Le- corresponding either to Limbani or, more pidobatrachus, Vanzolinius, Ischnocnema, and probably, to the Linimbare (=Inambari) river. Phyllonastes. As could be expected, the most Both sites, in the Peruvian department of significant change is that of the total number Puno, were visited in 1900 by P. O. Simons, of species, which has increased by 74, from collector of the BM. These localities, as well 112 to 186. This change, which overall repre- as Santo Domingo, are not far away from the sents an increase of 66 % in only ten years, is Bolivian border (ca.70 km by airline dis- due to several factors, namely: species over- tance). Thus, H. johnstoni is likely to occur in looked in the 1990 list, species deletions, the country, but its presence is still to be con- taxonomic changes, rediscovery of doubtful firmed. forms that proved to be valid species, species new to science, and new species records for Taxonomic changes.-Three kinds of taxonom- the country. ic changes have taken place: (1) synonymiza- tions (the synonymyzed “species” disappears Species overlooked in the 1990 list.-Lepto- from the list); (2) change from subspecific to dactylus macrosternum, Pleurodema borellii, specific status (a species is added if more than and Eleutherodactylus discoidalis. one subspecies were already cited); and (3) 54 DE LA RIVA et al.

Table 1: Taxonomic diversity and number of endemic species of Amphibians of Bolivia. Tabla 1: Diversidad taxonómica y número de especies endémicas de anfibios de Bolivia.

Order Family Genus Spp Endemics

Anura Bufonidae Atelopus 1 0 Bufo 14 2 Dendrophryniscus 1 0 Melanophryniscus 1 0 ______4 17 2

Centrolenidae Cochranella 3 2 Hyalinobatrachium 1 0 ______2 4 2

Dendrobatidae Allobates 1 0 Colostethus 3 1 Epipedobates 4 1 ______3 8 2

Hylidae Gastrotheca 4 2 Hyla 32 3 Osteocephalus 4 0 Phrynohyas 3 0 Scarthyla 1 0 Scinax 10 0 Sphaenorhynchus 1 0 Phyllomedusa 8 0 ______8 63 5

Leptodactylidae Ceratophrys 2 0 Chacophrys 1 0 Lepidobatrachus 1 0 Adenomera 3 0 Leptodactylus 23 0 Lithodytes 1 0 Physalaemus 6 0 Pleurodema 4 0 Pseudopaludicola 2 0 Vanzolinius 1 0 Eleutherodactylus 17 7 TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 55

Table 1: Continuation. Tabla 1: Continuación.

Order Family Genus Spp Endemics

Ischnocnema 2 1 Odontophrynus 2 0 Phrynopus 3 3 Phyllonastes 2 2 Telmatobius 10 8 ______16 80 21

Microhylidae Chiasmocleis 2 0 Dermatonotus 1 0 Elachistocleis 2 0 Hamptophryne 1 0 ______4 6 0

Pipidae Pipa 1 0

Pseudidae Lysapsus 1 0 Pseudis 1 0 ______2 2 0

Ranidae Rana 1 0

Caudata Plethodontidae Bolitoglossa 1 0

Gymnophiona Caeciliidae Caecilia 1 1 Siphonops 2 0 ______2 3 1

3 11 44 186 33 resurrection from synonymy (a species is steinbachi (it was listed in the 1990 list, but added if the senior synonym represents a synonymized with H. fasciata therein), Ololy- species also present in the country). gon (= Scinax) epacrorhina, Eleutherodacty- Synonymizations affected eight species: lus bockermanni, and Telmatobius albiventris. Atelopus willimani, Hyla callipleura, H. cari- Case (2) affects five species: Bufo poep- nata (DE LA RIVA, 1993d), H. ocapia, H. pigii, Epipedobates hahneli, Hyla andina, 56 DE LA RIVA et al.

Leptodactylus petersii, and Telmatobius boli- by HARVEY & SMITH (1993) and HARVEY vianus. (1996). Case (3) is represented by four species: Sci- Overall, of 50 valid species (at present-day) nax parkeri, Adenomera diptyx, Leptodactylus described from Bolivia over a period of 162 griseigularis, and L. leptodactyloides. years (in 1838 was described the first species whose type locality is in Bolivia, Epipedo- Rediscovery of doubtful forms.-Two species bates pictus), 25 (50%) have been described listed as doubtful in 1990 are included in the since 1990. This figure gives an idea about present list: Gastrotheca splendens and Tel- how much the research in Bolivia has lately matobius verrucosus. become dynamic and intense in comparison to past periods. New species are expected to be Species new to science.-A total of 28 Bolivian described in the next years, especially, but not species of anurans in 13 genera have been exclusively, in the genera Bufo, Hyla, Eleu- described as new since 1990: Bufo amboroen- therodactylus, Phrynopus, and Telmatobius. sis, B. castaneoticus (described from Brazil), B. justinianoi, Cochranella nola, Gastrotheca New species for the country.-They are 37 lauzuricae, Hyla chlorostea, H. marianitae species in 22 genera, accounting for the (described from Argentina), Scinax castrovie- largest number of additions to the 1990 list. joi, S. chiquitanus, Phyllomedusa camba, These species are: Bufo guttatus, Dendrophry- Leptodactylus didymus (described from Peru), niscus minutus, Melanophryniscus rubri- Physalaemus cuqui (described from Argen- ventris, Cochranella pluvialis, Allobates tina), Eleutherodactylus ashkapara, E. lloj- femoralis, Colostethus brunneus, C. trilinea- sintuta, E. olivaceus, E. pluvicanorus, E. tus, Epipedobates hahneli, E. trivittatus, Hyla samaipatae, E. zongoensis, Ischnocnema sanc- albopunctata, H. koechlini, H. marmorata, H. taecrucis, Phrynopus kempffi, P. pinguis, sarayacuensis, H. schubarti, H. tritaeniata, Phyllonastes carrascoicola, P. ritarasquinae, Phrynohyas resinifictrix, Scarthyla ostin- Telmatobius edaphonastes, T. huayra, T. ifornoi, odactyla, Scinax squalirostris, Phyllomedusa T. jahuira, and T. yuracare. bicolor, P. palliata, P. tomopterna, Chaco- All of them except those described from phrys pierottii, Lepidobatrachus laevis, Lep- other countries plus Bufo amboroensis (which todactylus gracilis, L. ocellatus, L. pentadac- occurs in Peru), S cinax castroviejoi (which tylus, L. syphax, Physalaemus cuqui, Pleu- presumably occurs in Argentina), Phyllome- rodema guayapae, Vanzolinius discodactylus, dusa camba (which occurs in Peru and Brazil), Eleutherodactylus danae, E. peruvianus, E. and Eleutherodactylus olivaceus (which oc- platydactylus, E. toftae, Ischnocnema quixen- curs in Peru) are Bolivian endemisms. The sis, Odontophrynus lavillai, and Chiasmocleis number of endemic species is now 33, which ventrimaculata. represents 17.7 % of the total amphibian fau- The occurrence of 19 of these species was na of the country. Most of these endemic expected by DE LA RIVA (1990a). The number species (21) belong to the family Leptodacty- of new records increases steadily with every lidae. herpetological field trip to poorly surveyed ar- Finally, two species described in the last eas of the country, especially those close to decade were considered as not valid in the the border of neighboring countries. same period: Bufo echinodes Reynolds & Fos- Two species of frogs reported recently in ter, and Centrolenella flavidigitata Reynolds Bolivia have not been included in the present & Foster were, respectively, synonymized list. Hyla walfordi Bokermann was reported in with B. quechua and Cochranella bejaranoi the country by DE LA RIVA et al. (1997), based TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 57 mainly on bioacoustical criteria. Further ex- Hyla molitor Schmidt, 1857. A case equal amination of the specimens and comparative to that of Gastrotheca splendens [see DUELL- studies of their advertisement calls revealed MAN (1970)]. that the reported population actually corre- Hyla roeschmanni De Grys, 1938. Type lost sponds to H. tritaeniata (I. De la Riva, pers. and type locality imprecise (“Beni, Bolivia”). obs.). Thus, H. walfordi has not been included Leptodactylus andicola Boettger, 1891. in the present list, but its occurrence in the Type lost and generic assignment doubtful country is highly plausible [see LANGONE & (see DE LA RIVA, 1990a). BASSO (1987)]. Phrynopus peruvianus (No- ble) was reported by KÖHLER (1995a; b), but Species expected to occur. further examination of the single specimen Many species not yet discovered are ex- collected demonstrated that it was a juvenile pected within the Bolivian boundaries when of Pleurodema marmoratum (I. De la Riva & further surveys are done. Some of the species S. Reichle, pers. obs.). listed below have been reported very close to the Bolivian border; others are known from Species of doubtful status. more distant localities, but substantial range For many species named long ago there extensions into Bolivia are plausible. In spite may be no more published information than of the fact that many species predicted by DE that provided in the original description. How- LA RIVA (1990a) have already been found in ever, these species may be common and/or Bolivia (see above), the number of potential have broad distributions. The redescription species (we propose 67) remains roughly as and publication of biological or ecological high as in 1990. This is due to two facts: the data on these species is always advisable. improvement of the knowledge on the distri- More remarkable are those cases in which a bution of many species and the description of species has remained unknown for a long pe- several new species from localities close to riod and the expectations to get new informa- Bolivia. In DE LA RIVA’s (1990a) arrange- tion are scant, due to the loss of the type ma- ment, there were four different geographical terial or because their type locality is un- areas in Bolivia to be considered in relation to known. We are unhappy with lists containing species potentially occurring in the country. species whose information is almost unre- Here we add a fifth zone, the Cordillera Occi- trievable. Such lists obscure the reality of dental. The areas and potential species are: what is well known about the composition and Amazonia: Allophryne ruthveni, Bufo gla- diversity of a given fauna. However, we are berrimus, Colostethus peruvianus, Dendro- aware of the importance of taking species in- bates biolat, D. quinquevittatus, D. vanzolinii, quirenda into account in order to carry out Epipedobates macero, E. simulans, Hyla al- further taxonomic research properly. The case lenorum, H. bokermanni, H. brevifrons, H. of Gastrotheca splendens commented above rossalleni, H. timbeba, H. walfordi, H. xa- is an illustrative example: being unaware of puriensis, Scinax ictericus, S. pedromedinae, the existence of such a “lost” taxon, probably Sphaenorhynchus carneus, Osteocephalus sub- would have led the taxonomist to describe a tilis, Agalychnis craspedopus, Phyllomedusa new species. Thus, a briefly commented list of atelopoides, perezi, Hydrolaetare Bolivian (or putatively Bolivian) doubtful schmidti, Leptodactylus dantasi, Eleuthero- species follows: dactylus altamazonicus, E. buccinator, E. imi- Bufo simus Schmidt, 1857. A case equal to tatrix, E. skydmainos, E. zimmermanae, Phy- that of Gastrotheca splendens [see SAVAGE llonastes myrmecoides, Phyzelaphryne miria- (1972)]. mae, Altigius alios, Chiasmocleis bassleri, C. 58 DE LA RIVA et al. shudikarensis, Ctenophryne geayi, Caecilia and mostly excluded from biodiversity inven- tentaculata, Oscaecilia bassleri, and Ty- tories, faunal lists, and regional surveys, pri- phlonectes compressicauda. marily due to lack of information and taxo- Chaco and Cerrado: Melanophryniscus nomic expertise. For example, The Red Data stelzneri, Epipedobates braccatus, Hyla vare- Book of Bolivian Vertebrates (ERGUETA & lae, Lepidobatrachus llanensis, Physalaemus MORALES, 1996) includes only two species of centralis, P. fuscomaculatus, Eleutherodacty- amphibians (Telmatobius albiventris and T. lus dundeei, and Chiasmocleis meheleyi. culeus, which are synonyms), and contains Northern Cordillera Oriental: Atelopus ery- recommendations about some others, but thropus, Cochranella phenax, C. spiculata, C. these recommendations are only tentative, for truebae, Hemiphractus johnsoni, Eleuthero- sound knowledge of the real problems affect- dactylus cosnipatae, E. lindae, E. mendax, E. ing Bolivian amphibians is lacking. Financial ockendeni, E. salaputium, Phrynopus peru- support and academic stimulus to carry out vianus, and Telmatobius sanborni. herpetological research are paramount if am- Southern Cordillera Oriental: Bufo gallar- phibians are to occupy the place they deserve doi, B. gnustae, B. rumbolli, Gastrotheca in conservation policies. Especially, the situa- chrysosticta, G. gracilis, and Telmatobius oxy- tion in Bolivia concerning the global phenom- cephalus. enon of declining amphibian populations is Cordillera Occidental: Telmatobius halli, T. completely unknown. Some observations seem pefauri, and T. zapahuirensis. to indicate that drastic declines might be af- fecting certain populations of montane species Conservation, recommendations and future [e. g., some Telmatobius; (ERGUETA & HAR- research. VEY, 1996; I. De la Riva, pers. obs.)]. Locally, The figure of 186 species of amphibians in rivers and lakes are being polluted and large Bolivia is still very conservative. Considering tracts of forest are being wiped out by agricul- the expected descriptions of new species plus ture, mining, logging, ranching, erosion, road the finding of species known from neighbor- construction, oil exploitation, etc. Informa- ing areas of other countries, a more realistic tion on amphibian diversity and distribution is figure for the Bolivian amphibian diversity is a first step to know what is being destroyed. around 250-300 species. The majority of the Nevertheless, overall, Bolivia is still well pre- endemic amphibians of Bolivia occur in the served and, if natural resources are wisely Cordillera Oriental of the Andes, and have re- managed, there are still great opportunities to stricted distributions in fragile ecosystems leave a rich natural heritage to future genera- (see KÖHLER et al., 1998b; KÖHLER 2000a). tions. We hope that further updated lists of The existence of this rich endemic fauna must Bolivian amphibians will not have to include a be necessarily taken into account by Bolivian section of “probably extinct” species. agencies involved in the management and pro- tection of natural resources, in order to make Acknowledgments sound and responsible environmental policy. This amphibian endemism often parallels Understandably, our years of work on Boli- equally interesting faunas and floras, but many vian amphibians which resulted in this list groups of and plants remain poorly in- would not have been possible without the kind vestigated. Thus, in general, preserving areas help of many friends and colleagues. To list all of amphibian endemism warrants the preser- of them would probably require a separate vation of other valuable biological resources. publication. Nevertheless, we want to thank However, the amphibians are often ignored all these persons and institutions who helped TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 59 with loaning specimens, providing miscella- APARICIO, J. (1993): Herpetofauna de Huara- neous information, or supporting our work in co, un ecosistema andino en el altiplano any other way. We specially thank the kind central de Bolivia. Ecol. Bol. Doc. ser. collaboration and support provided by J. Zool., 4: 1-38. Aparicio and the personnel at the the Colec- APARICIO, J. (1996): Herpetofauna. Pp 169- ción Boliviana de Fauna, La Paz, the Museo 172, in: Plan de Manejo del Area Natural de de Historia Natural “Noel Kempff Mercado”, Manejo Integral “Parque Nacional Sajama”. Santa Cruz, and the Fundación Amigos de la VAIPO-DGB-GTZ, La Paz (unpublished). Naturaleza, Santa Cruz. This paper greatly APARICIO, J. (1996): Herpetofauna de la car- benefitted from the kindness of J. Dorda, W. retera San Borja-Trinidad, Departamento Duellman, M. Göd, L. Gonzáles, P. Ibisch, K.- del Beni (Bolivia). Rev. Bol. de Ecol. 1 (1): H. Jungfer, A. Kwet, the late A. Mújica, H. 53-60. Nickel, D. Rumiz, and E. Wild, who loaned APARICIO, J. (1999): Herpetofauna. Pp. 66- some slides, and from the patience and effi- 70, in: Estudio de Biodiversidad de la cient management by the REH’s editors. The Reserva Nacional de Flora y Fauna Tariquía manuscript was improved by the wise com- (Tarija). IE-FUNDECO, La Paz (unpub- ments provided by M. B. Harvey, M. S. Hoog- lished). moed and J. A. Langone. Fieldwork of JK and APARICIO, J. (1999): Herpetofauna de la Re- SL was funded by grants of the German Aca- seva Nacional de Fauna Altoandina Ulla- demic Exchange Service (DAAD) and the Ulla, Departamento de La Paz (Bolivia). “Graduiertenförderung des Landes Nor- Rev. Ecol. Cons. Amb. Fund. Simón Patiño drhein-Westfalen”. 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Color Photographs.— Abbreviations of authors of the photographs are as follows: IR, I. De la Riva; JD, J. Dorda; WED, W. E. Duellman (courtesy of the Natural History Museum, The University of Kansas); MG, M. Göd; PI, P. Ibisch; KHJ, K.-H. Jungfer; JK, J. Köhler; AK, A. Kwet; JL, J. Lynch (courtesy of the Natural History Museum, The University of Kansas); AM, A. Mújica; SR, S. Reichle; and EW, E. Wild. Fotografías en color.— Las abreviaturas de los autores de las fotografías son: IR, I. De la Riva; JD, J. Dorda; WED, W. E. Duellman (courtesy of the Natural History Museum, The University of Kansas); MG, M. Göd; PI, P. Ibisch; KHJ, K.-H. Jungfer; JK, J. Köhler; AK, A. Kwet; JL, J. Lynch (courtesy of the Natural History Museum, The University of Kansas); AM, A. Mújica; SR, S. Reichle; y EW, E. Wild TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 75

Atelopus tricolor. Chapare, Cochabamba (JK)

Bufo amboroensis. Río Chua Khocha, Cochabamba (IR) 76 DE LA RIVA et al.

Bufo arenarum. Vaca Guzmán, Chuquisaca (JK)

Bufo castaneoticus. Cobija, Pando (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 77

Bufo granulosus. Rurrenabaque, Beni (SR)

Bufo guttatus. P. N. Noel Kempff Mercado, Santa Cruz (SR) 78 DE LA RIVA et al.

Bufo justinianoi. Chapare, Cochabamba (JK)

Bufo “margaritifer.” La Bola, Santa Cruz (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 79

Bufo marinus. Valle de Sajta, Cochabamba (IR)

Bufo paracnemis. Puerto Almacén, Santa Cruz (IR) 80 DE LA RIVA et al.

Bufo poeppigii. Carahuasi, Cochabamba (IR)

Bufo quechua. Incachaca, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 81

Bufo spinulosus. López Mendoza, Cochabamba (IR)

Bufo veraguensis. La Yunga, Santa Cruz (IR) 82 DE LA RIVA et al.

Dendrophryniscus minutus. Valle de Sajta, Cochabamba (IR)

Melanophryniscus rubriventris. López Mendoza, Cochabamba (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 83

Cochranella bejaranoi. Sehuencas, Cochabamba (JK)

Cochranella nola. El Fuerte de Samaipata, Santa Cruz (JK) 84 DE LA RIVA et al.

Cochranella pluvialis. Coroico, La Paz (SR)

Hyalinobatrachium bergeri. Chapare, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 85

Allobates femoralis. Chalalán, La Paz, (SR)

Colostethus brunneus. P. N. Noel Kempff, Santa Cruz (SR) 86 DE LA RIVA et al.

Colostethus mcdiarmidi. Río Yuyo, La Paz (IR)

Colostethus trilineatus. Estación Biológica Beni, Beni (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 87

Epipedobates bolivianus. Correo, La Paz (IR)

Epipedobates hahneli. Cobija, Pando (JK) 88 DE LA RIVA et al.

Epipedobates pictus. P. N. Noel Kempff Mercado, Santa Cruz (IR)

Epipedobates trivittatus. San Sebastián, Pando (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 89

Gastrotheca lauzuricae. Serranía Siberia, Cochabamba (IR)

Gastrotheca marsupiata. Sehuencas, Cochabamba (JK) 90 DE LA RIVA et al.

Gastrotheca splendens. Abra de la Cruz, Santa Cruz (IR)

Gastrotheca testudinea. Chapare, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 91

Hyla acreana. Mataracú, Santa Cruz (SR)

Hyla albonigra. Toro Toro, Potosí (PI) 92 DE LA RIVA et al.

Hyla albopunctata. P. N. Noel Kempff Mercado, Santa Cruz (IR)

Hyla andina. Aguirre, Cochabamba (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 93

Hyla armata. Chaco, La Paz (IR)

Hyla balzani. Coroico, La Paz (SR) 94 DE LA RIVA et al.

Hyla bifurca. Puerto Almacén, Santa Cruz (IR)

Hyla boans. P. N. Noel Kempff Mercado, Santa Cruz (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 95

Hyla calcarata. San Sebastián, Pando (SR)

Hyla charazani. Charazani, La Paz (IR) 96 DE LA RIVA et al.

Hyla chlorostea (holotype). Paracti, Cochabamba (IR)

Hyla fasciata. Cobija, Pando (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 97

Hyla geographica. Puerto Almacén, Santa Cruz (IR)

Hyla granosa. Cobija, Pando (JK) 98 DE LA RIVA et al.

Hyla koechlini. Rurrenabaque, Beni (SR)

Hyla lanciformis. Valle de Sajta, Cochabamba (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 99

Hyla leali. Estación Biológica Beni, Beni (SR)

Hyla leucophyllata. Estación Biológica Beni, Beni (SR) 100 DE LA RIVA et al.

Hyla marianitae. Carahuasi, Cochabamba (IR)

Hyla marmorata. “Inselberg” Santa Cruz (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 101

Hyla melanargyrea. P. N. Noel Kempff Mercado, Santa Cruz (SR)

Hyla minuta. Laguna Volcán, Santa Cruz (SR) 102 DE LA RIVA et al.

Hyla nana. Estación Biológica Beni, Beni (SR)

Hyla parviceps. Puerto Almacén, Santa Cruz (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 103

Hyla punctata. Puerto Almacén, Santa Cruz (IR)

Hyla raniceps. La Bola, Santa Cruz (IR) 104 DE LA RIVA et al.

Hyla rhodopepla. Mataracú, Santa Cruz (SR)

Hyla riveroi. Cobija, Pando (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 105

Hyla sarayacuensis San Sebastián, Pando (SR)

Hyla schubarti. Cuzco Amazónico, Peru (EW) 106 DE LA RIVA et al.

Hyla tritaeniata. Estación Biológica Beni, Beni (SR)

Osteocephalus buckleyi. Mataracú, Santa Cruz (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 107

Osteocephalus cf. leprieurii. Los Fierros, Santa Cruz (SR)

Osteocephalus pearsoni. Rurrenabaque, Beni (SR) 108 DE LA RIVA et al.

Osteocephalus taurinus. Estación Biológica Beni, Beni (SR)

Phrynohyas coriacea. Puerto Almacén, Santa Cruz (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 109

Phrynohyas resinifictrix. Centro Científico Yasuní, Napo, Ecuador (KHJ)

Phrynohyas venulosa. P. N. Noel Kempff Mercado, Santa Cruz (IR) 110 DE LA RIVA et al.

Scarthyla ostinodactyla. Iquitos, Peru (MG)

Scinax acuminatus. Lago Ipacaraí, Paraguay (KHJ) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 111

Scinax castroviejoi. Laguna Volcán, Santa Cruz (IR)

Scinax chiquitanus. Puerto Almacén, Santa Cruz (AM) 112 DE LA RIVA et al.

Scinax fuscovarius. Puerto Almacén, Santa Cruz (IR)

Scinax garbei. Cobija, Pando (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 113

Scinax nasicus. Estación Biológica Beni, Beni (SR)

Scinax nebulosus. P. N. Noel Kempff Mercado, Santa Cruz (SR) 114 DE LA RIVA et al.

Scinax parkeri. P. N. Noel Kempff Mercado, Santa Cruz (IR)

Scinax ruber. Estación Biológica Beni, Beni (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 115

Scinax squalirostris. Apolo, La Paz (IR)

Sphaenorhynchus lacteus. Puerto Almacén, Santa Cruz (IR) 116 DE LA RIVA et al.

Phyllomedusa bicolor. Pingo de Oro, Pando (SR)

Phyllomedusa boliviana. P. N. Amboró, Santa Cruz (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 117

Phyllomedusa camba. Puerto Almacén, Santa Cruz (IR)

Phyllomedusa hypochondrialis. Buenavista, Santa Cruz (IR) 118 DE LA RIVA et al.

Phyllomedusa palliata. Estación Biológica Beni, Beni (SR)

Phylomedusa sauvagii. Raco, Tucumán, Argentina (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 119

Phyllomedusa tomopterna. Cuzco Amazónico, Perú (IR)

Phyllomedusa vaillanti. Mataracú, Santa Cruz (JK) 120 DE LA RIVA et al.

Ceratophrys cornuta. Puerto Almacén, Santa Cruz (IR)

Ceratophrys cranwelli. , Santa Cruz (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 121

Chacophrys pierottii. Captive specimen from the pet trade (IR)

Lepidobatrachus laevis. Cerro Cortado, Santa Cruz (SR) 122 DE LA RIVA et al.

Adenomera andreae. Cobija, Pando (JK)

Adenomera diptyx. Santa Cruz de la Sierra, Santa Cruz (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 123

Adenomera hylaedactyla. Cobija, Pando (JK)

Leptodactylus bolivianus. Puerto Almacén, Santa Cruz (IR) 124 DE LA RIVA et al.

Leptodactylus bufonius. La Bola, Santa Cruz (IR)

Leptodactylus “chaquensis-macrosternum”. Vaca Guzmán, Chuquisaca (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 125

Leptodactylus didymus. Cobija, Pando (JK)

Leptodactylus elenae. Macuñucu, Santa Cruz (JK) 126 DE LA RIVA et al.

Leptodactylus fuscus. Buenavista, Santa Cruz (JK)

Leptodactylus gracilis. El Fuerte de Samaipata, Santa Cruz (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 127

Leptodactylus griseigularis. Chapare, Cochabamba (JK)

Leptodactylus knudseni. Rurrenabaque, Beni (SR) 128 DE LA RIVA et al.

Leptodactylus labyrinthicus. Puerto Almacén, Santa Cruz (IR)

Leptodactylus laticeps. Argentina (WED) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 129

Leptodactylus latinasus. Arroyo Los Naranjos, Jujuy, Argentina (WED)

Leptodactylus leptodactyloides. P. N. Noel Kempff Mercado, Santa Cruz (IR) 130 DE LA RIVA et al.

Leptodactylus mystaceus. Río Chevejecure, Beni (SR)

Leptodactylus mystacinus. Paraguay (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 131

Leptodactylus ocellatus. Lago Ipacaraí, Paraguay (KHJ)

Leptodactylus pentadactylus. Isla Coiba, Panama (IR) 132 DE LA RIVA et al.

Leptodactylus petersi. Buenavista, Santa Cruz (JK)

Leptodactylus podicipinus. San Ramón, Santa Cruz (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 133

Leptodactylus rhodomystax. Manaus, Brazil (KHJ)

Leptodactylus rhodonotus. La Hoyada, Santa Cruz (JK) 134 DE LA RIVA et al.

Leptodactylus syphax. “Inselberg”, Santa Cruz (JK)

Lithodytes lineatus. Mataracú, Santa Cruz (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 135

Physalaemus albonotatus. Santa Cruz de la Sierra, Santa Cruz (JK)

Physalaemus biligonigerus. Santa Cruz de la Sierra, Santa Cruz (IR) 136 DE LA RIVA et al.

Physalaemus cuvieri. Porto Alegre, Brazil (AK)

Physalaemus nattereri. Santa Rosa de la Roca, Santa Cruz (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 137

Physalaemus petersi. Mataracú, Santa Cruz (JK)

Pleurodema cinereum. Huayllamarca, Oruro (IR) 138 DE LA RIVA et al.

Pleurodema guayapae. Izozog, Santa Cruz (SR)

Pleurodema marmoratum. Comanche, La Paz (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 139

Pseudopaludicola boliviana. Estación Biológica Beni, Beni (SR)

Pseudopaludicola mystacalis. P. N. Noel Kempff Mercado, Santa Cruz (IR) 140 DE LA RIVA et al.

Vanzolinius discodactylus. Cobija, Pando (JK)

Eleutherodactylus ashkapara. Chapare, 2100 m, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 141

Eleutherodactylus cruralis. Mataracú, Santa Cruz (SR)

Eleutherodactylus danae. Chapare, Cochabamba (JK) 142 DE LA RIVA et al.

Eleutherodactylus discoidalis. La Yunga, Santa Cruz (IR)

Eleutherodactylus fenestratus. Santa Cruz del Valle Ameno, La Paz (IR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 143

Eleutherodactylus fraudator. Serranía Siberia, Cochabamba (IR)

Eleutherodactylus llojsintuta. Sehuencas, Cochabamba (JK) 144 DE LA RIVA et al.

Eleutherodactylus mercedesae. Road Caranavi-Yucumo, La Paz (SR)

Eleutherodactylus olivaceus. Chapare, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 145

Eleutherodactylus peruvianus. Santa Cruz del Valle Ameno, La Paz (IR)

Eleutherodactylus platydactylus. Sehuencas, Cochabamba (JK) 146 DE LA RIVA et al.

Eleutherodactylus pluvicanorus. Carahuasi, Cochabamba (SR)

Eleutherodactylus rhabdolaemus. Río Ronquito, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 147

Eleutherodactylus samaipatae. El Fuerte de Samaipata, Santa Cruz (JK)

Eleutherodactylus toftae. Paractito, Cochabamba (JD) 148 DE LA RIVA et al.

Eleutherodactylus ventrimarmoratus. Mera, Ecuador (JL)

Eleutherodactylus zongoensis. Zongo Valley, La Paz (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 149

Ischnocnema quixensis. P. N. Amacayacu, Colombia (IR)

Ischnocnema sanctaecrucis. Carahuasi, Cochabamba (JK) 150 DE LA RIVA et al.

Odontophrynus americanus. Pampa Grande, Santa Cruz (SR)

Odontophrynus lavillai. Santa Cruz de la Sierra, Santa Cruz (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 151

Phrynopus kempffi. Serranía Siberia, Cochabamba (IR)

Phrynopus laplacai. Unduavi, La Paz (SR) 152 DE LA RIVA et al.

Phrynopus pinguis (paratypes). Choquetanga Chico, La Paz (IR)

Phyllonastes carrascoicola. Chapare, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 153

Phyllonastes ritarasquinae. Chapare, 1250 m, Cochabamba (JK)

Telmatobius bolivianus. Chaco, La Paz (IR) 154 DE LA RIVA et al.

Telmatobius culeus. Lago Titicaca, La Paz (KHJ)

Telmatobius edaphonastes. Serranía Siberia, Santa Cruz (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 155

Telmatobius huayra (holotype). Campamento Khastor, Potosí (IR)

Telmatobius jahuira. Kkota Pata, La Paz (IR) 156 DE LA RIVA et al.

Telmatobius marmoratus. Comanche, La Paz (IR)

Telmatobius simonsi. , Chuquisaca (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 157

Telmatobius verrucosus (holotype). Chaco, La Paz (IR)

Telmatobius yuracare. Sehuencas, Cochabamba (JD) 158 DE LA RIVA et al.

Chiasmocleis albopunctata. Río Seco, Santa Cruz (JK)

Chiasmocleis ventrimaculata. Estación Biológica Beni, Beni (SR) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 159

Dermatonotus muelleri. Paraguay (JK)

Elachistocleis bicolor. Estación Biológica Beni, Beni (SR) 160 DE LA RIVA et al.

Elachistocleis ovalis. Río Seco, Santa Cruz (JK)

Hamptophryne boliviana. Puerto Almacén, Santa Cruz (JD) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 161

Pipa pipa. Captive specimen from the pet trade (IR)

Lysapsus limellus. P. N. Noel Kempff, Santa Cruz (JK) 162 DE LA RIVA et al.

Pseudis paradoxa. Montero, Santa Cruz (JK)

Rana palmipes. Paracti, Cochabamba (JK) TEN YEARS OF RESEARCH ON BOLIVIAN AMPHIBIANS 163

Bolitoglossa altamazonica. Villa Tunari, Cochabamba (SR)

Caecilia marcusi. Totaizal, Beni (SR) 164 DE LA RIVA et al.

Siphonops annulatus. Puerto Maldonado, Peru (WED)

Siphonops paulensis. Santa Cruz de la Sierra, Santa Cruz (JK)