Diversity and Conservation of the Amphibians of Bolivia

Author(s): Ignacio De la Riva and Steffen Reichle

Source: Herpetological Monographs, 28(1):46-65. 2014.
Published By: The Herpetologists' League
DOI: http://dx.doi.org/10.1655/HERPMONOGRAPHS-D-13-00009
URL: http://www.bioone.org/doi/full/10.1655/HERPMONOGRAPHS-D-13-00009

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Herpetological Monographs, 28 2014, 4665
E 2014 by The Herpetologists League, Inc.

DIVERSITY AND CONSERVATION OF THE AMPHIBIANS OF BOLIVIA

IGNACIO DE LA RIVA1,3 AND STEFFEN REICHLE2
1
Museo Nacional de Ciencias Naturales-CSIC, C/ Jose Gutierrez Abascal 2, 28006 Madrid, Spain
2
Quinta Tota, Santa Cruz de la Sierra, Bolivia

ABSTRACT: In the past decades, herpetologists have studied intensively the amphibians of Bolivia,
increasing dramatically the number of species known for the country. There are currently 266 species
recorded, but this number will increase with the addition of many new country records and the description of
species new to science, especially within Andean Craugastoridae. Deforestation, habitat destruction (mostly
due to agriculture), water pollution, and chytridiomycosis are the main causes of amphibian declines in
Bolivia. Andean frogs are much more affected than lowland species. Infection by the chytrid fungus
Batrachochytrium dendrobatidis is widespread. Forest species of the Andean genus Telmatobius have
disappeared from known sites and some other Andean taxa have declined severely. Here, we revise the
International Union for Conservation of Nature conservation status categories for some species of anurans.
Public awareness is increasing thanks to different local initiatives addressing projects to protect Bolivian
amphibians.
Key words: Anurans, chytridiomycosis; endemism; protected areas; Telmatobius

BOLIVIA is situated between latitudes 09u389S
and 22u539S and longitudes 57u269W and
69u389W and has an area of 1,098,581 km2.
The country is divided into nine administra-
tive departments (Beni, Chuquisaca, Cocha-
bamba, La Paz, Oruro, Pando, Potos, Santa
Cruz, and Tarija). The main physiographic
feature is the Andean Cordillera, which
crosses the country from northwest to south
and is the origin of rivers belonging to three
drainages: the Amazon Basin (66% of the
countrys extension), the Parana Basin (21%),
and the Altiplano Basin (13%). Roughly, half
of the country is in the Andean domain. The
Andes are divided here into two main ranges:
the volcanic Cordillera Occidental (which
delimits the Chilean border) and the Cordil-
lera Oriental (which contains most snow-
capped peaks and glaciers). Between these
two formations is the interior basin of the
Altiplano with an average elevation of 3700 m.
3
CORRESPONDENCE: e-mail, iriva@mncn.csic.es
Note: Volumes 9 and 11 in the series Amphibian
Biology, unlike other volumes in that series, are being
published in a number of parts, much like the separate
issues in the volume of a scientific journal. Published only
in Herpetological Monographs, this paper is Chapter 13,
Part 4 of Volume 9, Status of Decline of Amphibians:
Western Hemisphere, edited by H. Heatwole, C. Barrio-
Amoros, and J.W. Wilkinson. The previous parts (13) of
this volume covered the remaining countries of South
America, except Peru, and were published by Surrey
Beatty & Sons, Baulkham Hills, NSW, Australia; the rest
of the Western Hemisphere is pending.
The Altiplano contains large lakes and salty
flats. Lake Titicaca is 8300 km2 in extent
(3690 belonging to Bolivia and the rest to
Peru), whereas the Salar de Uyuni is the
largest salt-pan in the world, nearly 9000 km2.
The non-Andean regions of Bolivia are the
northern and eastern lowlands. Tropical
forests and savannahs predominate in the
center and north of the country, whereas the
drier Cerrado and Chaco formations occur in
the eastern and southeastern lowlands. The
main rivers in the Bolivian Amazon Basin are
Madre de Dios, Beni, Mamore, and Itenez,
and in the Parana Basin, the Pilcomayo
River; the largest river of the Altiplano Basin
is the Desaguadero, which connects Lake
Titicaca with Lake Poopo.
The Bolivian human population is slightly
over 10,000,000; roughly 62% is urban, with
most population (70%) concentrated in the
departments of La Paz, Santa Cruz, and
Cochabamba; the average human density is
8.49, being the lowest in Pando (0.8) and the
highest in Cochabamba (26.2; INE, 2002).
The demographic pressure is strong in some
areas, particularly around cities and in some
Andean valleys; in contrast, large parts of the
country are scarcely inhabited.
The main environmental concerns in Boli-
via are deforestation due to extensive clearing
of land for agricultural purposes (especially in
Santa Cruz Department, but lately also in the
Bolivian Yungas), slash-and-burn agriculture,

46
2014] HERPETOLOGICAL MONOGRAPHS
cattle ranching, timber exploitation, and min-
ing. The deforestation rate in Bolivia was
moderate a few decades ago, but the situation
changed dramatically by the end of the 20th
century. According to data provided by the
Ministerio de Desarrollo Sostenible y Medio
Ambiente, between 1975 and 1993, approxi-
mately 3.02 million hectares were deforested,
at a rate of 168,000 ha/yr; the average
deforestation rate then was 0.3% per year
(MDSMA, 1995), considerably lower than that
of most Latin American countries. According
to the Food and Agriculture Organization of
the United Nations, from 1990 to 1995 Bolivia
had the second highest annual rate of defores-
tation in South America, about 1.2% (FAO
1997). Cultivation of soybeans and other crops
to substitute for coca has contributed impor-
tantly to extensive deforestation since the 1990s
and has grown significantly in recent years.
Cuellar et al. (2012) show deforestation rates of
an average of 0.66% for Bolivia from 2000 to
2010, with peaks of 1.74% for the Cochabamba
Department, and 0.84% for the Santa Cruz
Department. Desertification and soil erosion
are rampant, especially in some Andean valleys,
as a consequence of overgrazing and poor
methods of cultivation. Pollution of water is
acute in some areas, mainly due to urban waste,
industry, and mining.
A substantial portion of the countrys
wilderness is protected by different adminis-
trative agencies such as national parks, natural
areas of integrated management, national
reserves, and biological stations. There are
probably over 100 protected areas, but only 22
are managed by the National Protected Areas
Service (SERNAP, Fig. 1); these collectively
account for roughly 17% of the national
territory (SERNAP 2000; Ibisch and Merida,
2003). These figures constantly change be-
cause the boundaries of some areas are
subject to reconsideration and, in recent
years, many municipal and private protected
areas were created. The degree of effective
protection, the budget for adequate manage-
ment, and the importance and intensity of
threats are extremely variable from site to site
but, in general, the situation is better for those
sites that are part of SERNAP.
The tropical location of Bolivia, together
with its elevational gradient and general
47
geographic heterogeneity, make it a country
of mega-diversity (Kohler, 2000). As habitat
loss is a major factor for declining amphibian
populations, it is worthwhile examining it on
an ecoregional scale in order to see how
strongly these ecoregions and their amphibian
populations and species are threatened by
changes in use of land in Bolivia.
ELEVATIONAL REGIONS AND ECOREGIONS
OF BOLIVIA
The World Wildlife Fund for Nature
recognizes 13 main ecoregions for Bolivia,
which are distributed in (1) the lowlands, (2)
the East Andean versants and the inter-Andean
dry valleys, and (3) the high mountain ranges
and highlands (Ibisch et al., 2003). The 22
National Protected Areas cover between 1.4%
to over 72.6% of each ecoregion (Table 1).
Figure 1 depicts a map of Bolivia showing the
ecoregions and the main protected areas.
The Lowlands
The lowlands include seven ecoregions: (1)
Southwest Amazonian Forests, (2) Madeira
Tapajos Moist Forest, (3) Cerrado, (4) Beni
Savannah, (5) Pantanal, (6) Chiquitano Dry
Forest, and (7) Dry Chaco. The Amazonian
rainforest covers most of Pando, northern
La Paz, some parts of northern Beni, and
northern Santa Cruz departments, as well as a
sub-Andean belt that reaches near the city of
Santa Cruz through the northern Cocha-
bamba Department. Deforestation has been
rampant in this last area due mainly to
agriculture (including coca), whereas in Pando
extensive cattle ranching is the main threat.
Large areas of rainforest are protected within
national parks and reserves (e.g., Madidi,
IsiboroSecure, Tahuamanu, ManuripiHeath,
Ros Blanco y Negro, Noel Kempff). Parts
of the Cerrado are threatened by soybean
cultivation and cattle ranching but, fortu-
nately, so far only on a relatively small
scale (compared to, for example, Brazil).
The seasonally inundated savannahs (Beni
Savannah and Pantanal) of Beni and Santa
Cruz are highly affected by cattle ranching
that is apparently shifting from extensive to
semiintensive. The most severe changes in
land use are occurring in the eastern parts of
the Chiquitano Dry Forest. The Chiquitania
48 HERPETOLOGICAL MONOGRAPHS [No. 28

FIG. 1.Map of Bolivia showing the main ecoregions and national protected areas (limits based on the World Wildlife
Fund for Nature and the National Protected Areas Service).
2014] HERPETOLOGICAL MONOGRAPHS
TABLE 1.Extent of protected areas in various Bolivian ecoregions.
Ecoregion Area protected in national reserves (ha)
Bolivian Yungas
Madeira- Tapajos Moist Forests
Southern Andean Yungas
Southwest Amazon Moist Forests
Bolivian Montane Dry Forests
Dry Chaco
Chiquitano Dry Forests
Beni Savannah
Cerrado
Pantanal
Central Andean Dry Puna
Central Andean Puna
Central Andean Wet Puna
Total
showed the highest deforestation rate in
the world during the late 1990s, reaching
1200 km2/yr (Steininger et al., 2001). Fortu-
nately, there are still large areas in the eastern
part of this ecoregion that are in very good
condition (Ibisch et al., 2002), although the
pressure upon this ecoregion is increasing in
recent years, as shown by Cuellar et al. (2012).
The Dry Chaco is still in good condition over
huge areas, with large portions protected (e.g.,
Kaa-Iya National Park); the main problem
outside these areas is extensive cattle ranching
and agriculture. Despite the existing pressures,
almost all amphibian species occurring in these
five ecoregions are relatively widespread and
viable populations are found in several national
parks and reserves.
The East Andean Versants and the
Inter-Andean Dry Valleys
This elevated region contains three eco-
regions: (1) Bolivian Yungas, (2) Southern
Andean Yungas, and (3) Bolivian Montane
Dry Forest.
This is the elevational region in Bolivia with
the most endemic amphibian species; several
of them have relatively small distributional
ranges and, consequently, this is the region
with the highest number of species suffering
conservation problems. Within its five eco-
regions, special attention should be paid to the
Bolivian Yungas and adjacent humid subpar-
amos, where roughly two-thirds of Bolivian
endemic amphibian species occur.
Parts of all these ecoregions suffer pres-
sures from marked changes in land use and
49
Percentage of ecoregion protected
3985.323 46.33
262.344 4.42
262.180 9.44
1676.704 9.91
416.599 5.72
3896.606 31.61
2763.828 16.82
180.476 1.44
432.835 72.68
1756.624 53,88
769.264 5.34
447.285 7.95
287.200 16.32
17,137.268 15.80
consequent erosion. In all these ecoregions,
untouched parts still remain, but also there
are valleys that have shifted almost completely
to agriculture and other human activities (e.g.,
Sorata, in La Paz Department). While large
parts of the Bolivian Yungas and the Southern
Andean Yungas are still in a good conserva-
tional state, the Bolivian Montane Dry Forests
especially have suffered severe impacts. Dur-
ing recent years, increasing cultivation of coca
in the Bolivian Yungas has caused further
deforestation and also encroachment into
protected areas. For example, the Carrasco
National Park currently has an even higher
deforestation rate (2.31%) than the national
one for 20002010 (0.66%; Cuellar et al.,
2012). Over 46% of the Bolivian Yungas are
under conservation in national protected areas
(see Table 1), but due to the high beta
diversity of amphibians in this ecoregion,
several species are not found in any of them.
The High Mountain Ranges and Highlands
The high mountains and highlands contain
three ecoregions: (1) Centran Andean Wet
Puna, (2) Central Andean Dry Puna, and (3)
Central Andean Puna. While vast regions are
still well preserved, mostly due to low human
population density, the Central Andean Wet
Puna has almost completely become an
anthropogenic area. Most of the amphibian
species found there are relatively capable of
dealing with human impacts. Nevertheless,
there are some aquatic species that are
threatened by direct human influences, as
for example, the Lake Titicaca Giant Frog.
50 HERPETOLOGICAL MONOGRAPHS
BOLIVIAN AMPHIBIAN DIVERSITY
In contrast with the scarcity of studies prior
to 1990, the amphibian fauna of Bolivia has
been the subject of considerable research over
the past two decades (De la Riva, 1990; De la
Riva et al., 2000; Kohler, 2000; Reichle, 2003,
2006). De la Riva et al. (2000) provided one of
the most comprehensive checklists of Bolivian
amphibians; they recognized 186 species, 44
genera, 11 families, and 3 orders. Almost
simultaneously, Kohler (2000) recognized 200
species. The differences between the two
studies stem mainly from the fact that Kohler
included in his checklist 10 species that were
at the time still undetermined. Since 2000,
many new records have been reported for the
country, and other species have been de-
scribed as new to science, dramatically raising
the number of Bolivian species (Reichle,
2003, 2006; Aguayo, 2009). Table 2 presents
an outline of amphibian diversity in Bolivia,
based on data currently published and updat-
ed through 30 April 2014. Table 2 is at the
same time an updated checklist of Bolivian
amphibians and constitutes the keystone upon
which the following discussion of the conser-
vation of Bolivian amphibians is based. Most
likely, the figure of 266 species (in 3 orders, 17
families, and 60 genera) presented herein will
continue changing rapidly. Published and
unpublished data indicate that several species
known hitherto from neighboring countries
most likely occur in Bolivia as well, and a
number of species, already in collections,
await description. Large areas of territory that
remain to be surveyed likely include new
species. Overall, considering all these factors,
a total of about 300350 species would seem
a reasonable prediction for the amphibian
diversity of Bolivia. This is about three times
the number of species (112) reported by De la
Riva (1990) in the first published checklist
of Bolivian amphibians. Whatever the final
figure, it will always be amenable to change
depending on taxonomic interpretations.
Table 2 presents the diversity and ende-
mism of the different taxa of Bolivian am-
phibians. The most diverse families are
Hylidae, Craugastoridae, and Leptodactylidae
with 78, 52, and 45 species, respectively.
Craugastoridae has by far the greatest ende-
mism: 32 of its species are endemic, in
[No. 28
contrast with only one endemic leptodactylid.
The families Craugastoridae and Telmatobi-
dae alone represent almost 63.6% of the total
amphibian endemism in Bolivia (66 species,
representing 24.8% of the Bolivian amphibian
fauna). These figures will undoubtedly in-
crease dramatically when the Bolivian fauna of
Psychrophrynella, a diverse craugastorid ge-
nus with highly endemic species (100%),
becomes better studied. Likewise, other
craugastorid genera, such as Pristimantis and
Noblella, will certainly add more species to the
list of endemics. It must be emphasized that
all these endemic members of Terrarana
(direct-developing frogs in the sense of
Hedges et al., 2008), and almost all non-
Terrarana endemics as well, occur in Andean
cloud forests or in humid subparamo forma-
tions.
At a spatial level, the richest Bolivian
regions in amphibian diversity are the north-
western part of the country (western Pando
and northern La Paz) and the Andean foothills
southwards, and the lowest diversity is found
in the cold Andean highlands (Embert et al.,
2011). The areas of the highest alpha diversity
and the highest endemic species richness do
not coincide; the maximum number of en-
demics is found in the Yungas of central
Bolivia (Cochabamba), where up to 15 sym-
patric endemic species can be potentially
found (Embert et al., 2011).
CONSERVATION STATUS OF BOLIVIAN AMPHIBIANS
The increase in knowledge of the Bolivian
amphibians during the past two decades has
only moderately been paralleled by a similar
increase in understanding their conservation
status. Data on possible threats or declines are
mostly anecdotal and programs of monitoring
populations and assessing putative declines
have only recently been implemented. Past
surveys provided useful information on local
diversity and gave some idea about possible
threats. However, these studies cover an
insufficient temporal scale for assessment of
demographic trends.
In the Red Book of Bolivian Vertebrates
(Ergueta and Morales, 1996), Ergueta and
Harvey (1996) made the first attempt to
categorize the conservation status of Bolivian
amphibians. These authors listed Telmatobius
2014] HERPETOLOGICAL MONOGRAPHS 51

TABLE 2.Checklist of described and reported amphibians of Bolivia, and their International Union for
Conservation of Nature Red List categories, as appearing in AmphibiaWeb (assigned by Stuart et al., 2008, in
Threatened Amphibians of the World [TAW]; these sources consider the whole distribution of the species), and Aguayo
(2009). If Aguayo changed TAWs category for Bolivia, we only show Aguayos designation. Categories proposed herein
refer to Bolivian populations only. Abbreviations for conservation status categories: LC 5 Least Concern; V 5
Vulnerable; NT 5 Near Threatened; EN 5 Endangered; CR 5 Critically Endangered; DD 5 Data Deficient.
Abbreviations for habitat mainly used by the species: D 5 dry; H 5 humid; F 5 forest; O 5 open; B 5 forest and open
areas and either dry or humid. Asterisks denote endemic species. Quotation marks indicate taxa of doubtful taxonomic
status. For some species there are no conclusive data on their presence in protected areas, although for most lowland
species it is highly likely. Nomenclature follows Frost (2014).

Conservation status

TAW/ Aguayo In protected

Taxon 2009 Proposed herein Lowland Andean area

Anura (14 families, 57 genera, 262 species, 66 endemics)

Aromobatidae (1 genus, 5 species, 1 endemic)
Allobatinae
Allobates brunneus LC DD D, O Yes
Allobates femoralis LC LC H, F Yes
Allobates flaviventris LC H, F Yes
Allobates mcdiarmidi* VU VU H, F Yes
Allobates trilineatus LC LC H, F Yes
Bufonidae (6 genera, 23 species, 5 endemics)
Amazophrynella minuta LC LC H, F, ?
Atelopus tricolor EN EN H, F Yes
Melanophryniscus rubriventris LC LC B, F Yes
Nannophryne apolobambica* DD DD H, F Yes
Rhaebo ecuadorensis LC H, F Yes
Rhaebo guttatus LC LC H, F Yes
Rhinella amboroensis* CR CR H, F Yes
Rhinella arenarum LC LC D, O Yes
Rhinella castaneotica LC LC H, F Yes
Rhinella justinianoi* EN EN H, F Yes
Rhinella leptoscelis VU H, F Yes
Rhinella major LC LC B, O Yes
Rhinella margaritifera LC LC H, F Yes
Rhinella marina LC LC H, O Yes
Rhinella mirandaribeiroi LC LC B, O Yes
Rhinella poeppigii LC LC H, B Yes
Rhinella quechua* EN EN H, F Yes
Rhinella rumbolli VU VU H, F Yes
Rhinella schneideri LC LC B, O Yes
Rhinella spinulosa LC NT D, O Yes
Rhinella stanlaii* LC DD H, F Yes
Rhinella tacana DD DD H, F Yes
Rhinella veraguensis LC VU H, F Yes
Centrolenidae (5 genera, 11 species, 2 endemics)
Centroleninae
Cochranella adenocheira DD DD H, F Yes
Cochranella nola NT NT H, F Yes
Cochranella phryxa* DD DD H, F Yes
Nymphargus bejaranoi* VU VU H, F Yes
Nymphargus pluvialis EN CR H, F ?
Rulyrana spiculata NT NT H, F ?
Vitreorana oyampiensis LC LC H, F Yes
Hyalinobatrachiinae
Hyalinobatrachium bergeri LC LC H, F Yes
Hyalinobatrachium carlesvilai LC H, F Yes
Hyalinobatrachium mondolfii LC H, F ?
Hyalinobatrachium munozorum LC H, F ?
52 HERPETOLOGICAL MONOGRAPHS [No. 28

TABLE 2.Continued.

Conservation status

TAW/ Aguayo In protected

Taxon 2009 Proposed herein Lowland Andean area

Ceratophryidae (3 genera, 5 species, 0 endemics)

Ceratophrys cornuta LC LC H, F Yes
Ceratophrys cranwelli LC LC D, O Yes
Chacophrys pierotti LC DD D, O Yes
Lepidobatrachus laevis LC LC D, O Yes
Lepidobatrachus llanensis LC DD D, O Yes
Craugastoridae (5 genera, 52 species, 32 endemics)
Holoadeninae
Noblella carrascoicola* LC VU H, F Yes
Noblella myrmecoides LC LC H, F Yes
Noblella ritarasquinae* VU VU H, F Yes
Psychrophrynella adenopleura* VU VU H, F Yes
Psychrophrynella ankohuma* VU VU H, O No
Psychrophrynella chacaltaya* VU VU H, O Yes
Psychrophrynella condoriri* EN EN H, O No
Psychrophrynella guillei* EN EN H, B Yes
Psychrophrynella harveyi* EN EN H, O No
Psychrophrynella iani* EN EN H, B No
Psychrophrynella iatamasi* VU VU H, F Yes
Psychrophrynella illampu* VU VU H, B No
Psychrophrynella illimani* EN EN H, B No
Psychrophrynella kallawaya* VU EN H, B Yes
Psychrophrynella katantika* VU VU H, F Yes
Psychrophrynella kempffi* VU VU H, F No
Psychrophrynella pinguis* EN EN H, F No
Psychrophrynella quimsacrucis * VU VU H, B No
Psychrophrynella saltator* VU VU H, F Yes
Psychrophrynella wettsteini* VU VU H, F Yes
Pristimantinae
Oreobates choristolemma* VU VU H, F Yes
Oreobates cruralis LC LC H, F Yes
Oreobates discoidalis LC LC H, F Yes
Oreobates heterodactylus DD DD D, O Yes
Oreobates ibischi* LC LC D, F ?
Oreobates madidi* LC LC H, F Yes
Oreobates quixensis LC LC H, F Yes
Oreobates sanctaecrucis* VU VU H, F Yes
Oreobates sanderi* VU VU H, F Yes
Oreobates zongoensis* CR DD H, F No
Pristimantis altamazonicus LC LC H, F Yes
Pristimantis carvalhoi LC LC H, F Yes
Pristimantis danae LC LC H, F Yes
Pristimantis dundeei DD DD H, O Yes
Pristimantis fenestratus LC LC H, F Yes
Pristimantis koehleri* LC H, F Yes
Pristimantis llojsintuta* LC LC H, F Yes
Pristimantis ockendeni LC LC H, F Yes
Pristimantis olivaceus DD LC H, F Yes
Pristimantis pharangobates LC LC H, F Yes
Pristimantis platydactylus LC LC H, F Yes
Pristimantis reichlei LC HF Yes
Pristimantis samaipatae* LC LC D, F ?
Pristimantis skydmainos LC LC H, F ?
Pristimantis toftae LC LC H, F Yes
Pristimantis ventrimarmoratus LC LC H, F Yes
Pristimantis zimmermanae LC LC H, F ?
2014] HERPETOLOGICAL MONOGRAPHS 53

TABLE 2.Continued.

Conservation status

TAW/ Aguayo In protected

Taxon 2009 Proposed herein Lowland Andean area

Strabomantinae
Yunganastes ashkapara* VU EN H, F Yes
Yunganastes bisignatus* EN EN H, F Yes
Yunganastes fraudator* VU VU H, F Yes
Yunganastes mercedesae DD DD H, F Yes
Yunganastes pluvicanorus* VU VU H, F Yes
Dendrobatidae (3 genera, 8 species, 3 endemics)
Colosthetinae
Ameerega boehmei* DD D, O Yes
Ameerega boliviana* LC LC H, F No
Ameerega hahneli LC LC H, F Yes
Ameerega picta LC LC H, B Yes
Ameerega trivittata LC LC H, F Yes
Ameerega yungicola* LC LC H, F Yes
Dendrobatinae
Adelphobates quinquevittatus LC LC H, F ?
Ranitomeya sirensis LC LC H, F ?
Hemiphractidae (2 genera, 6 species, 3 endemics)
Gastrotheca lauzuricae* CR CR H, F Yes
Gastrotheca marsupiata LC VU B, O Yes
Gastrotheca piperata* LC DD H, F Yes
Gastrotheca splendens* EN EN H, F Yes
Gastrotheca testudinea LC DD H, F Yes
Hemiphractus scutatus LC LC H, F ?
Hylidae (12 genera, 78 species, 9 endemics)
Hylinae
Dendropsophus acreanus LC LC H, F Yes
Dendropsophus brevifrons LC H, F ?
Dendropsophus coffea* LC LC H, F No
Dendropsophus delarivai LC LC H, F Yes
Dendropsophus joannae DD LC H, F ?
Dendropsophus juliani* LC LC D, O ?
Dendropsophus koechlini LC LC H, F Yes
Dendropsophus leali LC LC H, F Yes
Dendropsophus leucophyllatus LC LC H, F Yes
Dendropsophus marmoratus LC LC H, F Yes
Dendropsophus melanargyreus LC LC H, O Yes
Dendropsophus minutus LC LC B, B Yes
Dendropsophus nanus LC LC B, O Yes
Dendropsophus parviceps LC LC H, F Yes
Dendropsophus reichlei* DD DD H, F Yes
Dendropsophus rhodopeplus LC LC H, F Yes
Dendropsophus cf. riveroi LC LC H, F Yes
Dendropsophus rubicundulus LC LC D, O Yes
Dendropsophus salli LC LC H, F Yes
Dendropsophus sarayacuensis LC LC H, F Yes
Dendropsophus schubarti LC LC H, F ?
Dendropsophus triangulum LC LC H, F Yes
Dendropsophus tritaeniatus LC LC B, O Yes
Dendropsophus xapuriensis LC LC B, O ?
Dryaderces pearsoni LC LC H, F Yes
Hyloscirtus armatus VU VU H, F Yes
Hyloscirtus charazani* CR CR D, O Yes
Hyloscirtus chlorosteus* CR CR H, F Yes
Hypsiboas alboniger* NT NT D, O Yes
54 HERPETOLOGICAL MONOGRAPHS [No. 28

TABLE 2.Continued.

Conservation status

TAW/ Aguayo In protected

Taxon 2009 Proposed herein Lowland Andean area

Hypsiboas albopunctatus LC LC H, O Yes

Hypsiboas balzani* LC LC H, F Yes
Hypsiboas boans LC LC H, F Yes
Hypsiboas calcaratus LC LC H, F Yes
Hypsiboas callipleura* LC LC H, F Yes
Hypsiboas cinerascens LC LC H, F Yes
Hypsiboas geographicus LC LC H, F Yes
Hypsiboas lanciformis LC LC H, F Yes
Hypsiboas marianitae LC LC H, F Yes
Hypsiboas punctatus LC LC H, F Yes
Hypsiboas raniceps LC LC B, O Yes
Hypsiboas riojanus LC LC B, B Yes
Hypsiboas steinbachi* LC H, F Yes
Lysapsus boliviana DD LC H, O Yes
Lysapsus limellum LC LC B, O Yes
Osteocephalus castaneicola LC H, F Yes
Osteocephalus helenae LC LC H, F Yes
Osteocephalus cf. leprieurii LC LC H, F Yes
Osteocephalus mimeticus LC H, F Yes
Osteocephalus taurinus LC LC H, F Yes
Pseudis paradoxa LC LC B, O Yes
Pseudis platensis DD LC B, O Yes
Scarthyla goinorum LC LC H, F ?
Scinax acuminatus LC LC D, O Yes
Scinax castroviejoi DD LC D, F ?
Scinax chiquitanus LC LC H, F Yes
Scinax cf. fuscomarginatus LC LC B, O Yes
Scinax fuscovarius LC LC B, O Yes
Scinax garbei LC LC H, F Yes
Scinax ictericus LC LC H, F Yes
Scinax nasicus LC LC D, O Yes
Scinax nebulosus LC LC H, O Yes
Scinax parkeri LC LC B, O Yes
Scinax pedromedinae LC LC H, F ?
Scinax ruber LC LC H, F Yes
Scinax squalirostris LC DD D, O No
Sphaenorhynchus lacteus LC LC H, F Yes
Trachycephalus coriaceus LC LC H, F ?
Trachycephalus cunauaru LC LC H, F Yes
Trachycephalus typhonius LC LC B, B Yes
Phyllomedusinae
Phyllomedusa atelopoides LC LC H, F Yes
Phyllomedusa azurea LC LC B, O Yes
Phyllomedusa bicolor LC LC H, F Yes
Phyllomedusa boliviana LC LC B, B Yes
Phyllomedusa camba LC LC H, F Yes
Phyllomedusa palliata LC LC H, F Yes
Phyllomedusa sauvagii LC LC D, O Yes
Phyllomedusa tomopterna LC LC H, F Yes
Phyllomedusa vaillanti LC LC H, F Yes
Leptodactylidae (10 genera, 45 species, 1 endemic)
Leiuperinae
Edalorhina perezi LC LC H, F Yes
Engystomops freibergi LC LC H, F Yes
Eupemphix nattereri LC LC D, O Yes
Physalaemus albonotatus LC LC B, O Yes
2014] HERPETOLOGICAL MONOGRAPHS 55

TABLE 2.Continued.

Conservation status

TAW/ Aguayo In protected

Taxon 2009 Proposed herein Lowland Andean area

Physalaemus biligonigerus LC LC D, O Yes

Physalaemus centralis LC LC D, O Yes
Physalaemus cuqui LC LC D, O Yes
Physalaemus cuvieri LC LC D, O Yes
Pleurodema cinereum LC LC D, O Yes
Pleurodema guayapae VU VU D, O Yes
Pleurodema marmoratum LC LC D, O Yes
Pseudopaludicola boliviana LC LC B, O Yes
Pseudopaludicola mystacalis LC LC B, O Yes
Leptodactylinae
Adenomera andreae LC LC H, F Yes
Adenomera coca* DD LC H, B Yes
Adenomera diptyx LC LC B, B Yes
Adenomera hylaedactyla LC LC H, F Yes
Adenomera simonstuarti LC LC H, F ?
Hydrolaetare caparu DD LC H, B No
Hydrolaetare schmidti LC LC H, F ?
Leptodactylus bolivianus LC LC H, F Yes
Leptodactylus bufonius LC LC D, O Yes
Leptodactylus chaquensis LC LC B, O Yes
Leptodactylus didymus LC LC H, F Yes
Leptodactylus discodactylus LC LC H, F ?
Leptodactylus elenae LC LC B, O Yes
Leptodactylus fuscus LC LC B, O Yes
Leptodactylus gracilis LC LC H, O Yes
Leptodactylus griseigularis LC LC H, F Yes
Leptodactylus knudseni LC LC H, F Yes
Leptodactylus laticeps NT NT D, O Yes
Leptodactylus latinasus LC LC D, O Yes
Leptodactylus latrans LC LC D, O Yes
Leptodactylus leptodactyloides LC LC H, F Yes
Leptodactylus macrosternum LC LC H, O Yes
Leptodactylus mystaceus LC LC H, F Yes
Leptodactylus mystacinus LC LC D, O Yes
Leptodactylus pentadactylus LC LC H, F Yes
Leptodactylus petersi LC LC H, F Yes
Leptodactylus podicipinus LC LC H, F Yes
Leptodactylus rhodomystax LC LC H, F Yes
Leptodactylus rhodonotus LC LC H, F Yes
Leptodactylus syphax LC LC H, O Yes
Leptodactylus vastus LC LC D, O ?
Litodytes lineatus LC LC H, F Yes
Microhylidae (5 genera, 11 species, 0 endemics)
Gastrophryninae
Chiasmocleis albopunctata LC LC B, O Yes
Chiasmocleis bassleri LC LC H, F Yes
Chiasmocleis royi LC H, F Yes
Ctenophryne geayi LC LC H, F Yes
Dermatonotus muelleri LC LC D, O Yes
Elachistocleis bicolor LC LC B, O Yes
Elachistocleis haroi LC D, O Yes
Elachistocleis helianneae LC B, O ?
Elachistocleis ovalis LC LC H, O Yes
Hamptophryne alios DD DD H, F Yes
Hamptophryne boliviana LC LC H, F Yes
56 HERPETOLOGICAL MONOGRAPHS [No. 28

TABLE 2.Continued.

Conservation status

TAW/ Aguayo In protected

Taxon 2009 Proposed herein Lowland Andean area

Odontophrynidae (1 genus, 2 species, 0 endemics)

Odontophrynus cf. americanus LC LC D, O ?
Odontophrynus lavillai LC LC D, O ?
Pipidae (1 genus, 1 species, 0 endemics)
Pipa pipa LC LC H, F Yes
Ranidae (1 genus, 1 species, 0 endemics)
Lithobates palmipes LC LC H, F Yes
Telmatobiidae (1 genus, 14 species, 10 endemics)
Telmatobius bolivianus* EN CR H, F Yes
Telmatobius culeus CR CR D, O No
Telmatobius edaphonastes* EN CR H, F Yes
Telmatobius espadai* EN CR H, F Yes
Telmatobius gigas* CR CR D, O No
Telmatobius hintoni* VU EN D, O Yes
Telmatobius huayra* VU EN D, O Yes
Telmatobius marmoratus VU EN D, O Yes
Telmatobius sanborni VU CR H, F Yes
Telmatobius sibiricus* EN CR H, F Yes
Telmatobius simonsi* VU CR B, B Yes
Telmatobius timens VU CR H, F Yes
Telmatobius verrucosus* CR CR H, F Yes
Telmatobius yuracare* EN CR H, F Yes
CAUDATA (1 family, 1 genus, 1 species, 0 endemics)
Plethodontidae (1 genus, 1 species, 0 endemics)
Hemidactyliinae
Bolitoglossa cf. altamazonica LC LC H, F Yes
GYMNOPHIONA (2 families, 2 genera, 3 species, 0 endemics)
Caeciliidae (1 genus, 1 species, 0 endemics)
Caecilia marcusi LC LC H, F Yes
Siphonopidae (1 genus, 2 species, 0 endemics)
Siphonops annulatus LC LC H, F Yes
Siphonops paulensis LC LC B, O Yes

culeus as Vulnerable and T. albiventris (now
considered a synonym of T. culeus) as
Endangered due to overfishing and pollution
in Lake Titicaca. Furthermore, the workshop
for the preparation of the Red Book suggested
inclusion of Bolitoglossa altamazonica as
Vulnerable and recommended considering
some other species or subspecies, mostly
endemics, for inclusion in the book.
Until recently, the conclusions and recom-
mendations agreed upon in two workshops of
the Global Amphibian Assessment represent-
ed the only knowledge about the conservation
of Bolivian amphibians. In these two meet-
ings, held in Belo Horizonte (Brazil) and
Tandayapa (Ecuador) in 2003, attendants
addressed the status of Amazonian and
Andean species, respectively (Stuart et al.,
2004). Subsequently, Stuart et al. (2008)
evaluated the status of 209 species of Bolivian
amphibians following International Union for
Conservation of Natures (IUCNs) criteria.
Overall, they considered 21 species threat-
ened (5 Critically Endangered, 6 Endangered,
and 10 Vulnerable). Reichle (2006) recog-
nized 35 species as threatened (5 Critically
Endangered, 8 Endangered, and 22 Vulnera-
ble) and provided modelled potential distri-
bution maps for 206 species. The next
important step was the publication of the
Red Book of Wild Bolivian Vertebrates, in
which Aguayo (2009) made a complete
2014] HERPETOLOGICAL MONOGRAPHS
reevaluation of the IUCN categories for the
amphibians of Bolivia. Bolivian vertebrates
status was evaluated by means of a method
called MEGA, based on different variables
such as distribution, state of conservation of
habitats, population status, intrinsic biological
vulnerability (e.g., body size, reproductive
potential, vagility), and main threats to the
species (Aguirre et al., 2009). All these factors
were scored to estimate the degree of threat.
In his chapter on amphibians, Aguayo (2009)
refined this system for amphibian particular-
ities (e.g., low dispersal capability, small body
size, etc.). He included 254 species in his
analysis, of which 54 (21%) are in some
category of threat. Our Table 2 follows Stuart
et al. (2008) for the majority of species and
Aguayo (2009) for most of the species he
considered threatened (see below); for species
without a previously assigned category, this
chapter assigns one in Table 2, following
IUCN guidelines at regional and national
levels (IUCN, 2012). In Table 2, we provide
also a rough, generalized indication of habitat
use based on published records and collec-
tions, but it must be noted that compilations
like this are quite imperfect due to the
difficulty in properly allocating each species
within such broad habitat categories. For
example, several species from the lowlands
may also be present up to certain elevations in
the Andes (e.g., Rhinella marina, Dendropso-
phus parviceps), and some primarily Andean
species reach the adjacent lowlands as well
(e.g., Rhinella poeppigii). Likewise, some
species are able to colonize dry and humid
habitats and, within them, borders between
open habitats and forest are sometimes subtle
(e.g., Phyllomedusa boliviana). Accordingly,
these concepts are used here as general
vegetational domains, rather than as specific
kinds of habitats utilized by their resident
species. Thus, in Table 2, in general, we
indicate only the main kind of habitat used
by each species, unless the species is broadly
distributed throughout different kinds of
habitats. Overall, we treat 171 species as
lowland species and 95 as highland species
(64.3% and 35.7% of the total number of
species, respectively).
Some generalizations and conclusions stem
from the data presented in Table 2. First,
57
amphibians from the humid Andean montane
forests (Yungas) and wet puna are the most
threatened (70% of the total number of
threatened species; Aguayo, 2009), while
those from either the humid or dry lowlands
are not severely threatened. Embert et al.
(2011) stated that 95% of threatened Bolivian
amphibians occur in the Yungas and the
highlands. The Yungas of Cochabamba po-
tentially hold the highest number of sympatric
threatened species, with up to 13 species
(Embert et al., 2011); although most threat-
ened species in the Yungas occur in protected
areas, only 61% of the highland threatened
species occur in protected areas, mostly in
Madidi National Park and Cotapata National
Park (La Paz). These authors also stated that
habitat loss affects 100% of threatened
species, while chytridiomycosis affects 38%
of them (Embert et al., 2011). Most likely, the
main threat for montane species is chytridio-
mycosis, while for lowland species it is habitat
loss, which occurs at a fast pace in some
regions (e.g, Chapare, Cochabamba); howev-
er, large areas of suitable habitat are still
available in the lowlands, and the species as a
whole remain in good condition. In fact, no
species is considered threatened in the
lowland Pando and Beni departments, while
La Paz and Cochabamba have roughly 59%
and 48% of their species threatened, respec-
tively (Aguayo, 2009). Three exceptions are
the Chacoan species Ceratophrys cranwelli,
Chacophrys pierotti, and Leptodactylus lati-
ceps, which can be locally rare and subject to
some pet trade, at least in Argentina and
Brazil (Aguayo, 2009). In Bolivia, however, C.
cranwelli is locally common, albeit usually
difficult to find, and L. laticeps seems to be a
very rare species, perhaps because the Boli-
vian Chaco represents its distributional limit
and, accordingly, it is a suboptimal habitat for
this species. The status of many species may
change depending on the perspective from
which their distributions are viewed. When
the entire range of a species is considered, it
may be regarded as less threatened than when
it is considered on a narrower spatial scale.
For example, Bolivia holds a relatively small
area of the huge lowland rainforest that covers
most of the Amazon Basin, and some species
seem to occur only in an even smaller portion
58 HERPETOLOGICAL MONOGRAPHS [No. 28

FIG. 2.Some Bolivian amphibians. (A) Atelopus tricolor, the only species of this genus in the country, not seen since
2003; (B) Rhinella veraguensis, a formerly widespread and abundant species in montane Andean Bolivian forests, which
has experienced a drastic decline in the past two decades; (C) Hyalinobatrachium bergeri, a Glass Frog whose current
International Union for Conservation of Nature status of Low Concern can change if habitat degradation increases in
montane forests; (D) an undescribed species of Psychrophrynella from the Cordillera Real (La Paz), which, as with
many other species in this genus, might be threatened by habitat transformation; (E) Telmatobius culeus, the Titicaca
2014] HERPETOLOGICAL MONOGRAPHS 59

of this Bolivian segment (e.g., Ameerega
trivittata, Scarthyla goinorum, Phyllomedusa
atelopoides, Adelorhina perezi). Thus, wide-
spread Amazonian species may inhabit only a
small area in Bolivia, and the status at this
regional scale might not be considered the
same when seen more widely. In absolute
terms, however, the Bolivian lowlands still
retain large portions of well-preserved habi-
tats, big enough to safeguard viable amphibian
populations on a long-term basis.
Conservation threats are more important
for some Andean groups. Among the bufo-
nids, the dramatic population declines and
extinctions of members of the genus Atelopus
have been well documented throughout the
entire range of the genus (Young et al., 2001;
Lotters et al., 2004). So far, within this genus,
only A. tricolor (Fig. 2A) is known from
Bolivia, and its populations, considered En-
dangered, have not been properly monitored.
The species was rare in some areas (e.g.,
Yungas of Cochabamba), moderately common
in others (e.g., Serrana Eslabon and Serrana
Bellavista in La Paz Department), and seems
to be absent from apparently suitable areas
separating known populations. These facts
might indicate that population declines have
occurred in the past and/or are still taking
place. On the other hand, given the well-
documented propensity for all members of the
genus Atelopus to be severely affected by fatal
chytrid infections (Lotters et al., 2004), it
seems advisable to closely monitor the Boli-
vian populations of A. tricolor and of other
species occurring in the same habitat. Recent
efforts to find populations of A. tricolor in
Bolivia were undertaken in the Yungas of La
Paz and Cochabamba, but despite several
visits under suitable climatic conditions,
researchers failed to find any individual of
this species (I. De la Riva, personal observa-
tion; R. Aguayo, personal communication). As
far as we know, individuals were last seen in
December 2003 in Madidi National Park
(J.M. Padial, personal communication).
Cloud-forest species of Bufonidae in Bolivia
vary from quite rare species, such as Rhinella
justinianoi, to locally common ones, such as R.
veraguensis (Fig. 2B; however, this situation
changed a few years ago; now, R. veraguensis
has become remarkably rare). For some
species, e.g., R. tacana and Nannophryne
apolobambica, too few specimens and data
are available for assessment to be made.
Probably, the rarity of some species is natural,
but until monitoring programs are implement-
ed, it is difficult to ascertain the conservation
status and population trends of most species
of bufonids in the Bolivian Yungas. The first
record of chytridiomycosis in Bolivia was
reported for a bufonid, Rhinella quechua
(Barrionuevo et al., 2008).
A similar case is that of Glass Frogs
(Centrolenidae; Fig. 2C), which have been
victims of declines in some places. The
Bolivian endemic Nymphargus bejaranoi has
a rather extensive distributional area but, as in
the case of Atelopus tricolor, this species was
always rare in some areas, and perhaps has
real gaps in its distribution. Some attention
should be paid to N. pluvialis, a species only
known from the area of its Peruvian type
locality and one population close to Coroico in
the Yungas of La Paz. All visits to the Bolivian
locality since 2002 failed to find individuals
there, which might be due to nearby defores-
tation and major disturbance; consequently
Aguayos proposal that this species is Critically
Endangered is accepted here.
Among the aromobatids, Allobates mcdiar-
midi seems to be a rare species (based on the
scarcity of specimens in collections and our
own fieldwork experience), although with
relatively broad distribution in cloud forests
of Cochabamba and La Paz; the fragility of
this habitat makes it advisable to consider
the species as Vulnerable. By contrast, the

Giant Frog; (F) a poster aiming to encourage local people to protect the Titicaca Giant Frog from human consumption
(municipality of Sahuina, La Paz); (G) Telmatobius gigas, a species heavily infected by the chytrid fungus
Batrachochytrium dendrobatidis, but apparently not declining; (H) the last specimen known of Telmatobius yuracare,
a big male currently kept at the facilities of the Museo de Historia Natural Alcyde dOrbigny (Cochabamba). (Photos A,
B, C, D, F, and G by I. De la Riva; E and H by A. Munoz.)
60 HERPETOLOGICAL MONOGRAPHS
dendrobatid Ameerega boliviana has a more
restricted distribution in the Andean foothills
of La Paz, but is a species tolerant of
anthropogenic degradation of forests, and it
is rather common in the area. Thus, it is not
considered threatened, and a similar case is
that of Ameerega yungicola. The endemic
Ameerega boehmei deserves a special consid-
eration, for it has a restricted distribution in
an apparently unsuitable area for dendrobatid
frogs, the temperate forests of the Pre-
Cambrian shield in southeastern Santa Cruz.
Increasing frequency of fire and droughts at
these localities might threaten these popula-
tions and more data are needed to understand
how fire impacts them.
Among the marsupial frogs (Hemiphracti-
dae), Gastrotheca lauzuricae and G. splendens
seem to be extremely rare, and therefore are
considered threatened; they are known only
from one and two specimens respectively (in
the case of G. splendens, the provenance of
the holotype is unknown). Of course, this
apparent rarity might be due to inadequate
investigation of these particular species, and
their situation and taxonomic status may be
better understood when new studies and
material become available. Data are also scarce
for the three remaining species of Gastrotheca.
It seems that G. marsupiata was much more
common in the past (collection records show a
more or less continuous distribution in the
Bolivian humid highlands) and populations have
vanished from many areas (e.g., around La Paz
and some parts of the Altiplano). Gastrotheca
piperata is the most abundant species, although
its distribution is restricted to the cloud forests
of Cochabamba and Santa Cruz. Gastrotheca
testudinea, although widespread in the Amazo-
nian foothills of the Andes from Colombia to
Bolivia, is a rare species throughout its range.
In the diverse subfamily Hylinae (69 species)
only three species in the genus Hyloscirtus, all
of them endemic, are considered threatened.
Hyloscirtus armatus is considered Vulnerable;
it was formerly rather common but it seems
to be rarer now. Cortez (2009a) found dead
specimens in Zongo Valley, perhaps indicating
infection by chytrid fungus. Hyloscirtus char-
azani had apparently healthy populations, but
it is only known from a few streams in the area
of the dry Charazani Valley; this restricted
[No. 28
distribution represents a potential danger for
the species if environmental conditions change.
Finally, H. chlorosteus is known only from the
holotype, and its situation is similar to that
mentioned for Gastrotheca lauzuricae and G.
splendens. In addition, Hypsiboas alboniger is a
rare species with a relatively wide distribution
in the Andean dry valleys of the departments of
Cochabamba, Chuquisaca, and Potos, but only
a few specimens are known, and the species
might be affected by water pollution, droughts,
and erosion.
Among Andean craugastorids, Oreobates
zongoensis is known only from the holotype,
described in 1997; the type locality has been
severely disturbed, and further search by
different herpetologists and, specifically, by
C. Cortez recently, failed to find the species
(Cortez, 2009a,b). Thus, it was previously
considered as Critically Endangered. As a
species occurring at rather low elevation
(1200 m), O. zongoensis is not expected to
have a highly restricted distribution (as, for
example, highland species of the genus
Psychrophrynella). Probably, this is a very
rare species, as are other Andean Bolivian
craugastorids (e.g., Yunganastes bisignatus
and Y. mercedesae). Thus, it possibly occurs
in suitable areas nearby, so for the moment
this species should be considered as Data
Deficient. The situation in the genus Psychro-
phrynella is peculiar. Scant information is
available on the population dynamics of these
frogs, but two facts are evident: (1) some
species are restricted to a very small area (e.g.,
the surroundings of a stream at a given
elevation in a single valley) and in no case is
the distribution broad and (2) some species
(or populations) have a remarkably high
density of individuals, whereas other species
seem to be rather rare (De la Riva, 2007). The
conservation status of Psychrophrynella frogs
(Fig. 2D) is, in general, good, and no partic-
ular threats affect the extant species. Howev-
er, their extremely small distributions lead one
to think that, due to extensive habitat loss in
some regions, some species might have never
been discovered and are now extinct, and this
could happen in the future to others. For
example, most of the distribution of P. kempffi
likely falls outside the limits of the Carrasco
National Park, and developing agriculture
2014] HERPETOLOGICAL MONOGRAPHS
might affect this species severely. Climatic
change will, for sure, represent a threat in the
future for those species restricted to small
patches of suitable habitat, which might
disappear due to global warming and increas-
ing dryness. Thus, the status of each particular
species, many of them still undescribed,
should be carefully monitored, for the situa-
tion might change quite rapidly.
The status of the highly endemic, mostly
aquatic, frogs of the genus Telmatobius
deserves particular attention. Species of Tel-
matobius are affected by the same phenomena
that have led to the extinction of other
montane, stream-breeding amphibians in sev-
eral countries. The three species of Telmato-
bius from Ecuador seem to be already extinct
(Merino-Viteri et al., 2005), and cases of
infection by the chytrid fungus Batrachochy-
trium dendrobatidis (Bd) have been reported
in southern Peru in T. marmoratus (Seimon et
al., 2005; Catenazzi et al., 2010). Besides the
case of Rhinella quechua (Barrionuevo et al.,
2008), recent data indicate that Bd is broadly
present in several kinds of Andean habitats of
Bolivia, infecting many different species of
frogs, T. gigas among them (De la Riva and
Burrowes, 2011). Telmatobius gigas (Fig. 2G)
might have a quite restricted distribution and
lives in a habitat subject to perturbations from
natural and human causes (De la Riva, 2002);
apparently healthy, albeit infected, specimens
were found in 2008. The fact that some
Telmatobius still have healthy populations
should not make one trust that this situation
will continue. It has been suggested that even
persisting populations of frogs infected by Bd
can be at risk due to synergistic interactions
with climatic fluctuations and other stressors
(Longo and Burrowes, 2010). For this reason,
the category of Endangered is the one of least
threat that can be considered for any species
of this genus in Bolivia and the present
chapter makes some changes in relation to
Aguayos (2009) proposal. Two cases of severe
declines have been reported hitherto in the
country, in the department of Cochabamba
(De la Riva, 2005; De la Riva and Lavilla,
2008). A population of T. espadai and another
of T. hintoni, both in the same water
catchment system but at different elevations
close to the Colomi reservoir (department
61
of Cochabamba), were decimated in the
early 1990s. Only one other valley in the
department of La Paz is known to contain, or
have contained, T. espadai. The population in
Cochabamba is probably gone (De la Riva and
Lavilla, 2008) and nothing is known regarding
the fate of the one from La Paz, where the last
specimens were seen in 1999 (De la Riva, 2005).
Also, we found dead individuals of T. hintoni
near Aguirre (Cochabamba) in December 1999
and of T. bolivianus in Quime (La Paz) in
November 2002. Studies should be rapidly
carried out to assess the status of these and
other populations. The Titicaca Giant Frog, T.
culeus (Fig. 2E), is a special case because, in
addition to local problems of water pollution in
Lake Titicaca, the species is harvested for human
consumption. Unfortunately, research on its
captive breeding demonstrated it to be difficult
to rear and its slow life cycle makes com-
mercial exploitation probably neither feasible
nor profitable (Perez, 2005); however, the
Bolivian Amphibian Initiative (BAI, 2014)
has recently been successful at breeding both
T. culeus and T. hintoni in aquaria (Munoz,
2013). Telmatobius marmoratus is the most
widespread and common species in the
genus. Nevertheless, it is considered Endan-
gered, because it often lives in human-
modified landscapes and is affected by water
pollution and eutrophication caused by fer-
tilizers and by the excrement of domestic
camelids and sheep (De la Riva, 2005). The
species is absent from areas where it was
probably common in the past, and some
populations have declined severely. In gen-
eral, it seems that species of Telmatobius
from the dry puna or dry valleys (T. gigas, T.
hintoni, T. marmoratus, some populations of
T. simonsi), can persist locally, even infected
by Bd, whereas forest species are rapidly
decimated by the fungus (De la Riva and
Burrowes, 2011). For instance, besides the
case of T. espadai mentioned above, no
records of forest species of Telmatobius have
been obtained in recent years, in spite of, in
some cases, intensive search at type localities
and other areas where these species were
known to occur. As far as we know, T.
bolivianus was last seen in 2007 at Valle de
Zongo, La Paz (Cortez, 2009b); T. edapho-
nastes in 1998 at Carrasco National Park,
62 HERPETOLOGICAL MONOGRAPHS
Cochabamba (Aguayo, 2000); T. sanborni in
2001 at Pelechuco, La Paz (De la Riva, 2005);
T. sibiricus in 2003 at La Siberia, Cochabamba
(J. M. Padial, personal communication); T.
timens in 1999 at Tojoloque, La Paz (De la Riva
et al., 2005); T. verrucosus in 2004 at
Ancohuma, La Paz (J.M. Padial, personal
communication); and T. yuracare in 2007 at
Siberia, Cochabamba (A. Munoz, personal
communication). This last record corresponds
to an old male that is still kept alive at the
Museo de Historia Natural Alcyde dOrbigny,
in Cochabamba, as part of the captive breeding
program carried out by BAI; so far, no female
has been found to mate with him (Fig. 2H).
Finally, forest populations of T. simonsi seem
also to be gone (R. Aguayo and A. Munoz,
personal communication). We consider all
these forest species as Critically Endangered,
and perhaps some of them are already extinct.
The protected areas of Bolivia should be
able to preserve a great portion of the
countrys amphibian diversity, provided that
factors other than habitat loss and diseases are
not operative. Unfortunately, however, not
even an extensive network of national parks
and reserves covering the entire spectrum of
habitats and ecosystems seems to guarantee
the conservation of all amphibians. Emergent
infectious diseases, contamination, and the
negative effects of climatic change (higher
temperatures and lower humidity and precip-
itation) are factors acting across the boundar-
ies of protected areas. These adverse factors
affect mostly montane populations. On the
other hand, some endemic species with
greatly restricted distributions do not occur
in protected areas, as is the case of several
species of Psychrophrynella (e.g., P. anko-
huma, P. illampu, P. illimani, P. pinguis, P.
quimsacrucis). Small conservation areas in
municipalities might be able to cover and
protect some of these species, and plans for
conserving species should take into account
these political boundaries and conservation
tools (Embert et al., 2011).
There are no truly exhaustive inventories of
amphibians in any Bolivian national park. The
most comprehensive list is that of Madidi
National Park, where Domic et al. (2012)
reported 92 species. Considering the broad
elevational range of this national park (from
[No. 28
200 to 6000 m above sea level) and its many
types of ecosystems, the figure of 92 species is
probably low, but it indicates that at least
34.6% of the countrys 266 species have some
form of protection there. This number in-
creases when considering other large protect-
ed areas in other regions of the country (e.g.,
Kaa-Iya National Park or Noel Kempff
Mercado National Park), but they hold mostly
nonthreatened species. The perspective wors-
ens when only threatened amphibians are
considered. Aguayo (2009) stated that only
65% of threatened species are within protect-
ed areas. A large number of these species
occur in contiguous Carrasco National Park
(Cochabamba) and Amboro National Park
(Santa Cruz), with 39% and 21% of the
species, respectively (Aguayo, 2009). Unfortu-
nately, these areas suffer from heavy coloni-
zation pressure by farmers and illegal logging.
Embert et al. (2011) stated that the Yungas of
central Bolivia (mainly in Cochabamba) rep-
resent a well-defined conservation priority
area for amphibians, since this area combines
high species diversity with the highest number
of endemic species and threatened species.
Fortunately, most (80%) of this high-priority
area lies within protected areas, mostly in
Carrasco National Park. However, this is
still far from protecting all the endemic or
threatened species; for example, only two
species of Psychrophrynella (out of 17) and
four of Telmatobius (out of 14) occur in this
area (Embert et al., 2011). Overall, at least
222 species, representing a 83.4% of the total
amphibian fauna, are within protected areas,
with 144 (64.9%) being lowland species and
78 (35.1%) highland species (see Table 2).
Nevertheless, because of the scant geographic
records for many species (especially some
widespread lowland species), in many cases
we can only guess their presence within
protected areas; even those marked in Table 2
as not occurring within such areas might be
found there one day. For as many as 44
species (16.5%) there are no reliable records
within protected areas.
CONSERVATION PROCEDURES
Efforts specifically directed toward protect-
ing amphibians in Bolivia are still scarce. Two
notable exceptions are captive breeding of
2014] HERPETOLOGICAL MONOGRAPHS
the Titicaca Giant Frog, Telmatobius culeus
(Perez, 2005) and both T. culeus and T.
hintoni by BAI (Munoz, 2013). Conceivably,
if necessary, individuals from these attempts
at captive reproduction could be reintroduced
to restored habitats within the original range.
Furthermore, in the case of the Titicaca Giant
Frog, there is a growing concern about the
need to preserve this species and halt its
consumption by humans, with some interest-
ing local initiatives (Fig. 2F). Some Bolivian
herpetologists have already developed exper-
tise in monitoring populations. Several work-
shops on amphibian ecology and declines have
taken place in the country as, for example, the
Curso de Conservacion de Anfibios Altoan-
dinos, organized in October 2013 by BAI,
which has organized similar events in the past.
As this article was being written, BAI and
AmphibianArk were organizing a June 2014
workshop in Cochabamba on assessment of
needs for amphibian conservation. Likewise,
BAI intends to spread public awareness about
the need to protect amphibians, and runs
specific conservation projects. For example, a
project for evaluating the status of and ways to
protect Psychrophrynella illimani, threatened
by mining explotaition on the slopes of
Illimani Mountain (La Paz Department) was
recently approved and funded (A. Munoz,
personal communication). The Bolivian gov-
ernment is involved in developing a national
strategy for vertebrate conservation, which
should include specific strategies for amphib-
ians. By the end of 2012 the Direccion
General de la Biodiversidad submitted the
Action Plan for the Threatened Amphibians of
Bolivia, approved by the Bolivian government
but still pending funds (R. Aguayo, personal
communication). Additionally, the study of
past and current presence and the possible
effects of Bd in Bolivia is being addressed by
several researchers (e.g., R. Aguayo, J. Bosch,
P. Burrowes, C. Cortez, I. De la Riva, and A.
Munoz). The dramatic improvement in un-
derstanding the diversity and distribution of
Bolivian amphibians in the past few years has
made possible a more informed inclusion of
this group of vertebrates in inventories, tech-
nical reports, project proposals, area manage-
ment policies, and studies of environmental
impact. Besides continuing the necessary
63
studies of Bolivian amphibians from many
aspects, it is now time to initiate effective
conservation programs based on what is
already known. A nonexhaustive list of things
that should be done includes the following: (1)
creating microreserves protecting type locali-
ties of endemic species; often, this would
mean only a few square kilometers or even
less as, for example, for species of Psychro-
phrynella, which usually have extremely
restricted ranges; (2) searching for species
that are lost and, when feasible, the
implementation of ex-situ or in-situ captive-
breeding programs, such as the ones carried
out in places such as the El Valle Amphibian
Conservation Center, in Panama (e.g., Ga-
gliardo et al., 2008); (3) spreading public
awareness among local communities for whom
the knowledge that they hold species found
nowhere else in the world can be an incentive
to preserve their environment; (4) promoting
and facilitating taxonomic studies to improve
the knowledge of the real diversity and
distribution of Bolivian amphibians; (5) mon-
itoring extant populations of threatened
species; (6) limiting the uncontrolled use
of pesticides by farmers; (7) controlling the
introduction of invasive exotic fishes, such as
trout, which can be detrimental for native
amphibian species; and (8) implementing real
mechanisms to ensure effective preservation of
protected areas (especially those containing
high numbers of endemic or threatened
species); a first step in this direction would be
to abandon ambitious projects with unques-
tionable detrimental effects for the environ-
ment within such areas (e.g., oil, hydroelectric
power dams, roads, colonization programs).
Finally, a caution related to the fourth point
above: the Bolivian permitting authorities
should ensure that their policies do not stifle
scientific research. Collecting for scientific
purposes has never been the cause of
amphibian decline in Bolivia or almost any-
where else in the world, and new laws and
procedures should streamline permitting pro-
cedures. Arbitrary collection quotas should
not be imposed on scientists. Also, access to
necessary genetic data for taxonomic and
evolutionary studies should not be impeded.
Scientists pursuing these goals should not
be treated as commercial collectors or as
64 HERPETOLOGICAL MONOGRAPHS
commercially motivated individuals looking for
profitable, remunerative results (e.g., chemical
products, medicines). The scientific progress
that makes possible the basic knowledge of the
species, their biology, distribution, abundance,
and even their very existence, is often attained
despite, rather than thanks to, the legal
procedures. The last word on who, what,
where, how much, and when to collect should
rely on specialists and scientific, academic
authorities, and not on administrative staffs
that, unfortunately, often are in an almost
permanent state of flux regardless of how good
and efficient they are. Excessive bureaucracy
for scientists fieldwork, consisting, for exam-
ple, of having to get several mandatory permits
at different territorial and administrative levels,
should be kept at a minimum in order to
facilitate scientific progress, which for no good
reason often runs well behind the destruction
of our natural resources.
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