Ten years of research on Bolivian amphibians: updated checklist, distribution, taxonomic problems

146 Pages·2000·4.96 MB·English

Rev. Esp. Herp. (2000) 14:19-164 19
Ten years of research on Bolivian amphibians:updated checklist,
distribution,taxonomic problems,literature and iconography
IGNACIODELARIVA1, JÖRNKÖHLER2, STEFANLÖTTERS2& STEFFENREICHLE2
1Museo Nacional de Ciencias Naturales, CSIC,
C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain
2Zoologisches Forschungsinstitut und Museum Alexander Koenig,
Adenauerallee 160, 53113 Bonn, Germany.
Abstract:After ten years of intensive research, an updated checklist of the amphibians of Bolivia was compi-
led. For each species, information on its general distribution within the Neotropics, its occurrence within Boli-
vian departments and ecoregions, and habitat 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 walfordiand 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, taxonomy, 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-
fordiy 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 amphibian 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 DELARIVAet 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,METHODSANDGUIDELINES 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).
TENYEARSOFRESEARCHONBOLIVIANAMPHIBIANS 21
Cerradodomain, 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 Andes 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 DELARIVAet 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;
mean annual precipitation 500-700 mm; 6-8
Figure 1.-Map of Bolivian departments.
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;
Cerradoformations: 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
TENYEARSOFRESEARCHONBOLIVIANAMPHIBIANS 23
Figure 2.-Map of Bolivian ecoregions.
Figure 2.-Mapa de las ecoregiones de Bolivia.
24 DELARIVAet 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 habitats 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., Leptodactylus 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-
TENYEARSOFRESEARCHONBOLIVIANAMPHIBIANS 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
LARIVA(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 DELARIVAet 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: DELARIVA, 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 amboroensisHarvey & 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 castaneoticusCaldwell, 1991 Chacoan area (type locality: San José de
Distribution: 1) Amazonia; 2) Beni, La Paz, Chiquitos, Santa Cruz), B. g. mini Gallardo in
TENYEARSOFRESEARCHONBOLIVIANAMPHIBIANS 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; HASSet 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 isone 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: LÖTTERSet 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: DELARIVAet 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 DELARIVAet 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 quechuaGallardo, 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: DELARIVA, 1999a.
KÖHLERet 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 spinulosusWiegmann, 1834 Key literature: DELARIVA, 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. altiperuvianusfrom 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