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A new species of Psychrophrynella (Anura: Craugastoridae) from the Cordillera Real, Department La Paz, Bolivia


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We describe a new species of Psychrophrynella from a single locality in the Cordillera Real, in the Andes of Department of La Paz, Bolivia. Psychrophrynella teqta sp. nov. is highly variable in color pattern and differs from the species geographically closer mostly by presenting yellow or red blotches, usually absent in other species. In addition, the new species differs from all species in the genus because the males have a pulsed call. Males of this species show parental care, and the high number of eggs per nest suggest that perhaps more than one clutch might be attended by a single male. As other species in the genus, the new species is infected by Batrachochytrium dendrobatidis, and thus, a conservation threat is already present upon its discovery.
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Accepted by J. Padial: 28 Oct. 2014; published: 26 Nov. 2014
ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3887 (3): 459
A new species of Psychrophrynella (Anura: Craugastoridae) from the Cordillera
Real, Department La Paz, Bolivia
Museo Nacional de Ciencias Naturales, CSIC, C/ José Gutiérrez Abascal, 28006 Madrid, Spain. E-mail:
Department of Biology, University of Puerto Rico, P.O. Box 23360, San Juan, PR 00931
We describe a new species of Psychrophrynella from a single locality in the Cordillera Real, in the Andes of Department
of La Paz, Bolivia. Psychrophrynella teqta sp. nov. is highly variable in color pattern and differs from the species geo-
graphically closer mostly by presenting yellow or red blotches, usually absent in other species. In addition, the new species
differs from all species in the genus because the males have a pulsed call. Males of this species show parental care, and
the high number of eggs per nest suggest that perhaps more than one clutch might be attended by a single male. As other
species in the genus, the new species is infected by Batrachochytrium dendrobatidis, and thus, a conservation threat is
already present upon its discovery.
Key words: Amphibia, Andes, Bd, bioacoustics, parental care
Members of the genus Psychrophrynella represent a remarkable radiation of small direct-developing frogs
inhabiting the upper cloud forests, elfin forests, and wet subparamos of the Cordillera Oriental of the Andes from
southern Peru to central Bolivia (Frost 2014). Similar patterns of radiation are represented in central and southern
Peru by the genera Phrynopus and Bryophryne, respectively. The most striking feature of these groups is the high
beta-diversity and the limited distributions of species, which are usually known only from the type locality and, in
rare cases, some other places nearby (De la Riva 2007). All species described occur in the Amazonian versants of
the Andean Cordillera Oriental, and they are not found in the more xeric valleys facing the puna grasslands of the
Altiplano basin. The number of new species increases steadily as previously unexplored areas are surveyed, and the
actual number of species is still far from being known (De la Riva & Reichle 2014). While until now only three
species of Psychrophrynella have been described for Peru (but there are several others in collections awaiting
description; De la Riva et al. unpubl.), Bolivia holds a much richer Psychrophrynella fauna, with 17 species
described hitherto. Some areas, as the Cordillera de Apolobamba and the Cordillera Real, both in the Department
of La Paz, seem to have an especially high diversity of these frogs, often with at least one microendemic species
occurring in each major valley with suitable habitat.
During the early austral summer of 2012 we crossed the Cordillera Real from the Altiplano to the Amazonian
slopes, in search of Psychrophrynella frogs, and we found a beautiful new species in a valley not previously
surveyed. Herein we present the description of this species, describe and comment on particularities of its call,
report aspects of its reproductive biology, and assess conservation risk with respect to infection by the pathogenic
chytrid fungus, Batrachochytrium dendrobatidis Longcore, Pessier and Nichols.
Material and methods
Given the highly restricted distribution and microendemism of all species of Psychrophrynella, for practical
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purposes comparisons in the diagnosis are restricted to species geographically closest to the new species, or to
those that are superficially more similar. Specimens were fixed and preserved in ethanol 70%. They were examined
under a stereomicroscope and, when not possible to assess externally, the sexual and reproductive condition was
determined through dissection. Morphometric characters and format of diagnosis and description follow De la Riva
(2007); measurements were taken with a digital caliper to the nearest 0.01 mm and subsequently rounded to 0.1
mm. Abbreviations for morphometrics are as follows: SVL (snout–vent length), HL (head length, from rictus to tip
of snout), HW (head wide, at level of rictus), IND (internarial distance), END (eye–nostril distance, straight line
distance between anterior corner of orbital opening and posterior margin of external nares), ED (eye diameter,
horizontal), IOD (interorbital distance), TL (tibia length), and FL (foot length, distance from posterior margin of
inner metatarsal tubercle to tip of fourth toe). Terminology for skin texture and structures follow Duellman & Lehr
Call recordings (48 kHz, 24-bit, WAV format) were obtained by means of a Olympus LS-11 digital recorder,
and analyzed with Raven 1.4 software (Cornell University, Ithaca, NY) using both oscillogram and spectrogram.
After recording, we registered air temperature. For each vocalization, a total of 16 acoustic parameters were
measured, eleven in the temporal domain (call duration, inter-call interval, call rise duration, call fall duration, call
rise ratio, number of pulses, duration of first, intermediate and last pulses, pulse rise duration, and pulse fall
duration) and five in the spectral domain (call and pulse dominant frequency, frequency modulation, number of
harmonics, and first harmonic energy). Spectral parameters were calculated through fast Fourier transform (FFT)
set at a length of 512 points (Hann window, 50% overlap), providing a resolution of 94 Hz and 11 ms. Recordings
are deposited at the Fonoteca Zoológica Digital (
Samples of DNA from the skin of living frogs were taken as swabs, which were stored in individual airtight
plastic containers at ambient temperature during fieldwork and then at -20 °C in the laboratory until processed for
detection of the chytrid fungus Batrachochytrium dendrobatidis (Bd). DNA extraction was done using 50 ul of
PrepMan Ultra (Hyatt et al. 2007). We used a Taqman Real-Time quantitative PCR system for detection and
quantification of the level of infection by Bd on frog epidermal tissue, according to Boyle et al. (2004), and
infection intensity was calculated as the number of Bd zoospore genomic equivalents in each swab sample.
Coordinates were obtained with a Garmin 12XL GPS device and then adjusted with Google Earth®.
Psychrophrynella teqta, sp. nov.
(Figs. 1–2, 5)
Holotype. CBF 6725 (field tag 4421), an adult female from Pablo Amaya, Provincia Larecaja, Departament La
Paz, Bolivia (15º58’52.55” S, 68º12’19.6” W), 3700 m.a.s.l., collected on 6 November 2012 by Ignacio De la Riva
and Patricia A. Burrowes.
Paratypes. CBF 6726 (4415) and MNCN 45702–3 (4413, 4414), adult males; CBF 6727–8 (4417, 4422) and
MNCN 45704–5 (4416, 4418), adult females; and CBF 6729 (4420) and MNCN 45706 (4419), juveniles, same
data as the holotype.
Diagnosis. A member of Craugastoridae assigned to the genus Psychrophrynella based on biogeographical and
morphological grounds. Two other genera of Holoadeninae (sensu Padial et al. 2014) occur in Bolivia: Noblella
and Oreobates. Noblella has terminal phalanges narrowly T-shaped, discs and distal circumferential grooves
present distally, and tips of at least toes III and IV acuminate (terminal phalanges knob-shaped, circumferential
grooves absent, and tips of all digits rounded in Psychrophrynella); Oreobates has dentigerous processes of vomers
prominent (absent in Psychrophrynella) (Duellman & Lehr 2009). Psychrophrynella teqta is diagnosed by the
following combination of characters: (1) medium size (maximum SVL 27.8 mm), body robust, legs short (average
TL + FL between 69.8–75.7% SVL; n = 8); (2) tympanic membrane absent, tympanic annulus present, visible
under the skin; (3) first finger slightly shorter than second; (4) tips of digits slightly swollen, not expanded
laterally; (5) webbing of toes and lateral fringes absent; (6) two metatarsal tubercles, tarsal fold absent; (7) dorsal
skin and flanks shagreen to pustulate; dorsolateral folds irregular, reaching midbody; ventral skin areolate; (8)
snout rounded in dorsal view and in profile; (9) color on dorsum variable, from dark brown to beige, with or
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without irregular cream, reddish-brown blotches; (10) color on venter variable, from dark brown to cream, with a
pattern of irregular blotches cream, yellow, or gray.
FIGURE 1. Dorsolateral and ventral view of the living holotype of Psychrophrynella teqta sp. nov. (CBF 6725; SVL 26.1
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FIGURE 2. Variation in color pattern of Psychrophrynella teqta sp. nov. Dorsolateral and ventral views of A: MNCN 45704
(female; SVL 27.8 mm); B: CBF 6726 (male, 23.6 mm SVL); C: CBF 6727 (female, 27.2 mm SVL); D: CBF 6728 (female,
25.8 mm SVL).
Psychrophrynella teqta is distinguished from other species in the genus for which the call is known, by its
pulsed call. Regarding morphology and coloration, the new species is superficially similar to P. wettsteini (Parker)
and P. condoriri (De la Riva, Aguayo and Padial), which furthermore are species whose type localities are
geographically close to that of P. teqta. The new species is distinguished from P. wettsteini mostly by its smaller
size (maximum SVL in P. wettsteini 33.4 mm) and shorter legs (TL + FL > 80% SVL in P. wettsteini); the color
Zootaxa 3887 (3) © 2014 Magnolia Press
pattern in P. wettsteini is mostly gray or reddish-brown above with or without pale spots and ventrally cream with
reddish-brown spots or reticulations, while in P. te q t a coloration is extremely variable and often there are reddish
and yellow blotches, which are never present in P. wettsteini. From P. condoriri, the new species differs mostly by
having dorsal skin shagreen or rugose (mostly smooth in P. condoriri), and a variable color pattern both on dorsum
and venter, including reddish and yellow blotches (dorsum brown with diffuse darker areas, venter gray with brown
markings). The small species P. chacaltaya (De la Riva, Padial and Cortéz) is found in the upper Zongo Valley, not
far from the type locality of P. teqta, and it is easily distinguishable from the new species by being much smaller
(maximum SVL 20.4 mm), and having a different color pattern, consisting of dorsum dark grayish-brown with
darker marks and venter uniformly cream or brown with irregular brown blotches.
Description of the holotype. Body robust; dorsal skin covered with many small pustules, more abundant on
flanks, posterior part of body, and upper surface of thighs; a pair of dorsolateral folds reaching midbody (pustules
and dorsolateral folds barely visible in preservative for changes in skin texture); ventral skin areolate; no thoracic
or discoidal fold; a pelvic patch on lower surfaces of thighs, rugose, slightly swollen. Head wider than long, its
width 35.6% of SVL; head length 27.9% of SVL; snout moderately long, slightly rounded in dorsal view and in
profile; nostrils not protuberant, directed laterally, closer to snout than to eyes; canthus rostralis poorly marked,
straight, convex in dorsal view and in profile; eye-nostril distance 84.6% of eye length; loreal region barely
concave, interorbital region flat, lacking cranial crests; tubercles on upper eyelid absent; tympanic membrane
absent, two lower thirds of tympanic annulus slightly noticeable under the skin; supratympanic fold short, weak,
poorly marked; no postrictal tubercles or glands; tongue large, oval; choanae oval, small, widely spaced; vomerine
odontophores absent. Limbs moderately short; tips of digits slightly swollen, not expanded laterally; ulnar tubercle
and fold absent; inner palmar tubercle single, oval, poorly defined, smaller than round outer; fingers not fringed;
subarticular tubercles round, poorly defined; supernumerary tubercles, round, small; first finger slightly shorter
than second, relative length of fingers 1<2<4<3; tibia length 34.8% of SVL; tarsus lacking tubercle and fold; inner
metatarsal tubercle oval, larger than round, poorly defined outer; plantar surface smooth, no supernumerary
tubercles; subarticular tubercles round, slightly swollen; toes moderately long and slender, not webbed or fringed;
relative length of toes 1<2<3<5<4; foot length 37.9% of SVL.
Measurements (in mm) of the holotype. SVL 26.1; HL 7.3; HW 9.3; IND 2.3; END 2.2; ED 2.6; TL 9.1; FL
Color. In life, upper surfaces of body, head and limbs black; a few pinky-cream, irregular blotches on dorsum
and upper parts of flanks; creamy-yellow stripes on the upper edge of canthus rostralis, converging on the snout
and forming and inverted V from above; upper lip and post-commissural areas pale yellow; diffuse brown areas on
upper surfaces of forearms; venter and throat black with large, creamy-yellow blotches forming an irregular
pattern, becoming greenish-beige on belly; a blotch of this same color on the lower surface of each limb; lower
surface of thighs pale brown, with a reddish-brown, rugose pelvic patch; ventral surfaces of digits and inner part of
palms fleshy; iris greenish-brown, heavily reticulated in black, especially on the lower two thirds. In preservative,
dorsum, head, and upper parts of the extremities dark brown, with irregular areas slightly paler; gray, cream, and
pinky cream irregular blotches on dorsolateral areas and flanks; gray stripes bordering the canthus rostralis above;
upper lip and post-commissural area pale cream, with a fine dark brown line on the rim of the lip; diffuse brown
areas on upper surfaces of forearms; throat and venter dark brown, almost black, with large cream blotches forming
an irregular pattern; lower thighs pale brown; ventral surfaces of digits and inner part of palms pale cream.
Va ri at io n . Morphometric variation is given in Table 1. Males lack nuptial pads, and have small vocal slits and
a rugose, moderately developed, subgular vocal sac. Variation in color pattern is highly remarkable (Fig. 2; notes
based on coloration in life). The male MNCN 45702 has upper parts, throat and lower surfaces of limbs dark
brown; there is a greenish-beige line along the lower jaw; the venter is reddish-brown with large, irregular, bluish-
gray blotches. The male MNCN 45703 is pale brown above, with flanks reddish-beige with a reticulated pattern of
dark brown dots and blotches, all interconnected; there is a black supratympanic stripe; the throat is bluish-gray,
densely pigmented with dark brown; the venter is dark brown with small, scattered, irregular dark brown blotches.
The male CBF 6726 is pale brown above, with a black vertebral stripe and black irregular blotches forming an X-
shaped dorsal pattern; there is a black line from the tip of the snout to the shoulder, across the canthus rostralis, the
eye, and the supartympanic region; the flanks have an irregular pattern of black and golden-beige blotches; the
venter is bluish-gray with large black blotches, and the throat is dark brown with bluish-gray flecks. Two large
females—CBF 6727 and MNCN 45705—are, overall, similar to the holotype, but the former has a fine, greenish-
Zootaxa 3887 (3) © 2014 Magnolia Press
beige vertebral stripe, and the lower parts of head and body are black with irregular, greenish-beige blotches
(forming a longitudinal stripe on the throat); there are orange-beige blotches over the insertion of the forearms and
on the posterior part of the flanks; a reddish line is present along the posterior surfaces of thighs, meeting on the
cloacal region, where they form a heart-like shape. The female CBF 6728 is uniformly dark brown, almost black
above, with irregular, reddish-brown blotches on dorsum and flanks; the throat is reddish-beige and the venter is
greenish-beige with black. A specimen with a remarkable pattern is the female MNCN 45704, which is olive-
brown above, and has reddish flanks with a finely reticulated, brown pattern; there is a fine, beige vertebral stripe
starting on the tip of the snout; the throat, chest, and venter are greenish-beige with broad, reticulated, dark brown
blotches; the same reticulated pattern appears on the lower surface of limbs, encircling reddish blotches; there is a
reddish-brown line along the posterior surface of the thighs. The two juveniles collected are equally variable.
MNCN 45706 is pale reddish-brown above with diffuse areas olive-brown, and the flanks are dark brown with pale
red spots on the inguinal region; the toe tips are red. CBF 6729 is dark brown dorsally, with a fine, pale beige
vertebral stripe; there is a fine reddish line along the posterior surface of thighs, converging at the cloacal region,
forming a heart-like shape.
TABLE 1. Morphometrics of Psychrophrynella teqta sp. nov. Means followed by ranges in parentheses. For
abbreviations, see text.
Distribution and ecology. This species is known only from the type locality (Fig. 3). The locality of Pablo
Amaya is surrounded by agricultural fields and pastures for llamas, leaving very little suitable habitat for
Psychrophrynella frogs. The new species was on the more humid slopes passing the village, midway between the
old road and the small river that flows along the valley. Individuals were found under stones by day. One individual
(MNCN 45704) was on the bank of a narrow, small canyon, and the rest were under the scattered stones pertaining
to the ruins of an old construction (Fig. 4), which created a micro-environment with humidity and vegetation highly
suitable for the species. No other species of anurans were found in syntopy. The type locality of P. condoriri is 5.25
km to the northwest of Pablo Amaya, that of P. w et t st e i n i is 45 km to the southeast, and P. cf. chacaltaya occurs
25.5 km also to the southeast, in the upper Zongo Valley.
Natural history. Reproduction data on species of Psychrophrynella are not easy to gather. So far, only the
eggs of P. w et ts te i n i and P. illampu (De la Riva, Reichle and Padial) had been observed or photographed [Ergueta
1993; De la Riva 2007; recently, nests of P. illimani (De la Riva and Padial) were found and photographed by A.
Muñoz and colleagues (A. Muñoz, in litt.)]. We found two clutches of P. teqta under stones, each of them guarded
by a male occupying a small chamber. The first clutch contained 41 well-developed eggs (average diameter 5.11
mm, n=5; Fig. 5), containing embryos that moved frequently inside the egg capsule; the guarding male (MNCN
45703) was quite small (23.5 mm SVL). This clutch proved to be difficult to extract from the ground, for it was
deeply intermingled with small, very slender grass roots. The second clutch had 28, smaller, yellowish-white eggs
Character Adult Females (n=5) Adult Males (n=3)
SVL 26.8 (25.8–27.8) 23.2 (22.7–23.6)
HL 7.1 (6.7–7.5) 6.2 (6.1–6.4)
HW 8.9 (8.4–9.4) 8.0 (7.4–8.6)
IND 2.2 (1.9–2.3) 1.9 (1.8–2.2)
END 2.1 (2.1–2.2) 1.7 (1.5–1.9)
ED 2.4 (2.2–2.6) 2.1 (1.9–2.2)
TL 9.4 (9.1–9.7) 8.1 (7.7–8.6)
FL 10.2 (9.7–11.0) 9.0 (8.7–9.2)
HL/SVL 0.26 (0.25–0.27) 0.26 (0.26–0.27)
HW/SVL 0.33 (0.31–0.35) 0.34 (0.31–0.37)
END/ED 0.90 (0.84–0.95) 0.82 (0.68–0.94)
TL/SVL 0.34 (0.34–0.36) 0.34 (0.33–0.36)
FL/SVL 0.37 (0.35–0.39) 0.38 (0.38–0.39)
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(average diameter 4.74 mm, n=5). In both cases, the males remained on or beside the eggs, presumably offering
some type of parental care. They had the snout slightly protruding, of a translucent gray color, different from the
rest of the head; this probably indicates certain burrowing activity to construct and maintain the chambers where
they guard the clutch. The holotype had large, developed oviducts but no female among our sample showed
oviductal eggs.
FIGURE 3. Map of northern Bolivian Andes showing the distribution of Psychrophrynella teqta sp. nov. (star), and general
aspect of the type locality.
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FIGURE 4. Ruins of old local constructions, which create an optimal microhabitat for Psychrophrynella teqta sp. nov.
FIGURE 5. Male of Psychrophrynella teqta sp. nov. (MNCN 45073) guarding eggs.
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FIGURE 6. Advertisement call of Psychrophrynella teqta sp. nov. Air temperature at the time of recording, 10.8ºC. Above,
oscillogram and power spectra (power spectra analysis bandwidth: 15 Hz). Below, spectrogram and oscillogram (spectrogram
analysis bandwidth: 94 Hz).
Advertisement call. Calls of Psychrophrynella are remarkably similar across species, and most calls consist of
a simple, tonal, rather short, high-pitched note [so far, the most divergent call is that of P. saltator (De la Riva,
Reichle and Bosch), consisting of series of 7–36 short notes; see De la Riva 2007]. At the time of collecting P. teqta
(05–07 h pm) some frogs were calling. Surprisingly, calls were pulsed, not tonal. Thus, it was very important to
capture a voucher calling male in order to be sure that the recorded call indeed belonged to the new species. The
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voucher male (MNCN 45702) was in the middle of a thick mass of moss, over a large rock. At an air temperature of
10.8ºC and a relative humidity of 77%, this male emitted pulsed calls consisting of a simple note with an average
duration of 270 ms (n=8; Fig. 6), uttered in irregular series of several calls. Based on the total time of recording, the
overall call rate is approximately 6.5 calls/minute (average interval between calls, 2,365 seconds; n=6). Each call
has 16–19 pulses of variable duration, being the first pulses shorter. For example, the first pulse lasts, on average, 8
ms, while the last pulse has an average duration of 26 ms. The pulse amplitude is also progressively higher along
the call, with a maximum at approximately 221 ms from the onset of the call (i.e., call rise time 82% of call
duration). The spectral structure of this advertisement call is characterized by an average dominant frequency of
2.2 kHz, corresponding with the fundamental frequency of the signal and lacking frequency modulation. The call
shows 3–5 apparent harmonics, of which the first one accounts for an average of 95% of the total signal energy
(Fig. 6).
Etymology. The specific name is a mostly arbitrary combination of letters that the authors do like.
Remarks and discussion
Because no eggs were found inside the body of the collected females, we have no idea on the potential clutch size
in P. teqta. The number of eggs in the chambers, combined with their large egg diameter, seem to be excessive for
a small, direct-developing species. The scant information on reproductive biology of Psychrophrynella frogs do not
allow for extensive comparisons. Ergueta (1993) reported in P. wettsteini an average number of 13 ovaric eggs in
females (egg diameter, 5.0 mm) and the observation of two nests containing 8 and 20 froglets (average SVL, 6.3
mm in the first nest); in both cases, an adult, putatively a female, was also in the nest, but how the sex of this
individual was determined is not explained. De la Riva (2007) reported a nest of P. illampu containing 8 eggs with
a diameter of 5.5 mm, but no adult specimen was observed guarding the eggs. Catenazzi (2006) reported in a
Peruvian species of the closely related genus Bryophryne [B. cophites (Lynch)] a clutch of 20 eggs ca. 4 mm in
diameter, guarded by a female (again, it is not possible to know if sex was unequivocally determined or was
assumed). The clutches of P. illimani observed by a team from the University of Cochabamba were guarded by
males (see Thus, it is not clear if there is interspecific variation regarding the gender
responsible for parental care. In the case of the first guarding male of P. t eqt a observed with 41 eggs, it is plausible
to assume that these eggs may correspond to more than one female, indicating that a complex reproductive
behavior is involved, such as that reported for other members of Craugastoridae (Burrowes 2000). Evidently, there
are still many aspects of the reproductive biology of Psychrophrynella frogs to be investigated.
As far as it is known, the peculiar advertisement call of P. t eq t a is unique within the genus in being pulsed
instead of tonal. So far, the calls of eight Bolivian species have been analyzed and described (De la Riva 2007), and
in four of them they consist of a single, tonal, wistle-like vocalization, as are the calls recorded in Peru from P.
boettgeri (Lehr) and one undescribed species (De la Riva et al. unpubl.). This seems to be the most common call
structure in the genus. The Bolivian species P. illampu (De la Riva, Reichle and Padial) and P. ka t a n t i k a (De la
Riva and Martínez-Solano) emit calls consisting of a series of 2–4 notes, while P. wettsteini produces call groups
containing 7–18 short notes, and finally, in the most distinctive case of P. saltator, the notes are extremely short and
are emitted in groups of up to 36 notes per call (De la Riva 2007). These notes are nonetheless tonal, instead of
pulsed as in P. teqta.
Conservation of Psychroprynella frogs can be worrisome because they are sensitive to changes in land use, and
their geographic ranges are usually extremely restricted (De la Riva 2007). Climate warming can pose a threat too,
as temperature, humidity and –hence–, vegetation cover, might in turn compromise the future habitat quality for
these frogs. The fungal disease chytridiomycosis, caused by the pathogenic chytrid fungus Batrachochytrium
dendrobatidis (Bd), is considered the main threat to amphibians (Stuart et al. 2004; Mendelson et al. 2006), and it is
also a concern. Results of molecular diagnosis of Bd in P. t eq ta revealed that two out of six individuals tested were
infected. This is not surprising since Bd had already been reported for other members of this genus (Catenazzi et al.
2011; De la Riva & Burrowes 2011). None of the two individuals of P. t e q t a showed signs of being affected by
chytridiomycosis in spite of having moderate to high infection loads. While juvenile MNCN 45706 had 1088 Bd
zoospore genomic equivalent (Z
), adult male MNCN 45703 had 130,400 Z
. This level of infection is much
higher than that found by Catenazzi et al. (2011) for two Peruvian species of Psychrophynella and by De la Riva &
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Burrowes (2011) for the Bolivian species P. adenopleura (Aguayo and Harvey). Because population studies on
species of this genus are not available we cannot infer at this point the effect of Bd in Psychrophynella species that
are presently persisting with this pathogen. While their condition of terrestrial direct-developers may work to their
advantage in face of a mostly aquatic pathogen (Longcore et al. 1999), these frogs might be confronting a higher
risk of extinction if they are also subject to other environmental stressors presently affecting the Andes, like climate
warming and drought (Seimon et al. 2007). These factors can work in synergy with Bd to increase susceptibility of
frogs to chytridiomycosis as reported for other members of Brachycephaloidea (Longo & Burrowes 2010; Longo et
al. 2013; De la Riva & Burrowes 2011).
Beta-diversity of Psychrophrynella in Bolivia is astonishing. With 18 species described hitherto, all of them
endemic, this genus represents a 27.3% of the total endemic Bolivian anuran fauna—by far the greatest
contribution of any amphibian single group—and many more species await description (De la Riva 2007; De la
Riva & Reichle 2014; De la Riva, unpubl.). Indeed, no other vertebrate group reaches a similar degree of
endemicity; this alone stands as an important reason to pay especial attention to these little, otherwise scarcely
visible frogs, and justify the need to include them in conservation planning and environmental official agendas. A
recent project led by Bolivian Amphibian Initiative (BAI) is currently assessing the conservation status of P.
illimani and trying to create public awareness among locals about the necessity to preserve this and other frog
species (De la Riva & Reichle 2014; see BAI’s website, and the website of the P.
illimani project,
This research was possible thanks to the funding provided by project CLG2011-30393 of the Spanish Ministry of
Science and Innovation (PI, Ignacio De la Riva). We are grateful to James Aparicio for his help at the Colección
Boliviana de Fauna and to the DGB (Dirección General de la Biodiversidad) for providing exportation permits. We
are especially indebted to Diego Llusia for kindly helping with the analyses and figures of the advertisement calls,
to Jandrely López for assistance with qPCR detection, and to Noemí Goicoechea for help with Figure 3.
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... This supposition is supported by the discovery of 42 species of high Andean terrara- nas in the last 10 years during explorations in Bolivia and Peru (e.g. De la Riva, 2007;Lehr & Catenazzi, 2009;Lehr et al., 2012; De la Riva & Burrowes, 2014;Chaparro et al., 2007Chaparro et al., , 2015. Guided by the absence of species records for the dozens of unexplored glacial valleys in the cordilleras Apolobamba (Peruvian sec- tion) and Carabaya and intermediate areas in south- ern Peru -in strong contrast with our findings of many new species in Bolivia (De la Riva, 2007) -we surveyed several of those valleys and found multiple unknown species of terraranas. ...
... Mating calls are known for 11 spe- cies of Microkayla (including those of M. boettgeri and M. chapi, described herein). The call usually consists of a single, isolated, tonal, whistle-like, short note, but there are some exceptions: M. teqta has a pulsed call (De la Riva & Burrowes, 2014) while M. wettsteini and M. saltator emit several notes per call in a rapid series (De la Riva, 2007). Microkayla saltator is the most peculiar species in the genus; it lives at relatively low elevations (c. ...
... Also, several specimens pos- sess a protruding, translucent callosity on the tip of the snout, that covers the anterior area of the snout and part of the upper lip. So far, in Holoadeninae, this structure has been only described in males of the Bolivian species M. teqta (De la Riva & Burrowes, 2014). Those males were guarding egg clutches in subterranean chambers under stones; thus, the men- tioned peculiar rostral morphology is probably a struc- ture for digging (De la Riva & Burrowes, 2014). ...
Despite recent efforts to accelerate exploration and species description, the diversity of high Andean frogs remains highly underestimated. We report high levels of species diversity in direct-developing frogs or terraranas inhabiting the wet puna and adjacent cloud forests of the Amazonian versant of the Andes in Bolivia and Peru. Descriptive evidence of external morphology, distribution patterns and molecular phylogenetic analyses support the existence of nine unnamed species in two clades, which represents a 30% increase in species diversity for those clades. The relationships of these species and their relatives in Holoadeninae are tested using nuclear and mitochondrial genes for 159 terminals representing the 11 genera in this subfamily and 25 species of previously unknown relationships. Our results corroborate species monophyly in all but three cases and support the monophyly of all Holoadeninae genera, albeit the position of some differs between analyses. We propose a new genus (Microkayla gen. nov.) for the clade containing all Bolivian species formerly in Psychrophrynella plus five species from southern Peru. The new genus is monophyletic and supported by anatomical synapomorphies. Psychrophrynella is re-diagnosed and redefined to include three species from the Andes of southern Peru. We discuss the taxonomic instability associated with Noblella and Psychrophrynella due to the fact that the type species of both genera share a number of traits that support a close relationship. We also name and describe three new species of Bryophryne and two of Microkayla from Peru, provide baseline data for the future description of four Bolivian species of Microkayla, and describe the unknown mating calls of two species. Our results support that the grasslands of the Amazonian versant of the Andes harbour a large diversity of species with small altitudinal and horizontal distributions that replace each other along a latitudinal axis. These species belong to different lineages whose closest relatives are forest species, often from distant parts of the continent. These patterns suggest that high Andean environments were colonized several times independently by species with forest ancestors and which radiated into a multitude of species with remarkably similar ecomorphologies. The extent of these radiations remains obscured by a still rudimentary knowledge of species diversity due to insufficient fieldwork and taxonomic research.
... The frog genus Psychrophrynella currently includes 21 species distributed across the humid grasslands and forests from 1830 to 4190 m.a.s.l. in the Amazonian slopes of the Andes in southern Peru and Bolivia (De la Riva & Burrowes, 2014; Duellman & Lehr, 2009, Frost, 2015). The genus was placed within the Holoadeninae in the family Strabomantidae by Hedges, Duellman & Heinicke (2008), but Pyron & Wiens (2011) synonymized Strabomantidae with Craugastoridae. ...
... 1) lack a tubercle or fold on the inner edge of tarsus. Furthermore, among species whose advertisement calls is known, P. chirihampatu shares with P. bagrecito, P. saltator and P. usurpator the characteristic of having a call composed of multiple notes (Table 1; unpublished data for calls of P. bagrecito and P. usurpator), whereas the call is pulsed in P. teqta or composed of short, single notes in other congeneric species (De la Riva, 2007; De la Riva & Burrowes, 2014). Morphologically, the new species is most similar to P. usurpator (characters in parentheses;Table 1), from which it differs by having yellow ventral coloration with reddish-brown or grey flecks (dull brown, gray or black with cream flecks), Finger I shorter than Finger II (slightly shorter or same length), smaller SVL reaching 27.5 mm in females (SVL up to 30.5 mm), slender head (wider and shorter head), and inner metatarsal tubercle at least three times the size of outer metatarsal tubercle (about same size). ...
... The new species is yet another addition to the ever growing list of small craugastorid frogs (genera Bryophryne, Noblella and Psychrophrynella) from the eastern slopes of the Peruvian and Bolivian Andes (Catenazzi, Uscapi & Von May, 2015; De la Riva, 2007; De la Riva & Burrowes, 2014; De la Riva, Chaparro & Padial, 2008a; Harvey et al., 2013; Lehr & Catenazzi, 2009a; Lehr & Catenazzi, 2009b; Lehr & Catenazzi, 2010). Most if not all of these species have narrow distribution ranges often restricted to the type locality and surrounding mountaintop region, although large areas in between the type localities of these species remain unexplored. ...
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We describe a new species of Psychrophrynella from the humid montane forest of the Department Cusco in Peru. Specimens were collected at 2,670–3,165 m elevation in the Área de Conservación Privada Ukumari Llakta, Japumayo valley, near Comunidad Campesina de Japu, in the province of Paucartambo. The new species is readily distinguished from all other species of Psychrophrynella but P. bagrecito and P. usurpator by possessing a tubercle on the inner edge of the tarsus, and from these two species by its yellow ventral coloration on abdomen and limbs. Furthermore, the new species is like P. bagrecito and P. usurpator in having an advertisement call composed of multiple notes, whereas other species of Psychrophrynella whose calls are known have a pulsed call ( P. teqta ) or a short, tonal call composed of a single note. The new species has a snout-vent length of 16.1–24.1 mm in males and 23.3–27.7 mm in females. Like other recently described species in the genus, this new Psychrophrynella inhabits high-elevation forests in the tropical Andes and likely has a restricted geographic distribution.
... SVL); (2) tympanic membrane and tympanic annulus present, poorly visible; (3) first finger slightly shorter than second; (4) tips of digits slightly swollen, not expanded laterally, lacking circumferential grooves and, in males, nuptial excrescences; (5) webbing of toes and lateral fringes absent; (6) two metatarsal tubercles, tarsal fold absent; (7) dorsal skin and flanks shagreen with scattered small warts; ventral skin finely granular; (8) snout rounded in dorsal view and in profile; (9) dorsum uniformly reddish brown or with small yellowish blotches; (10) venter fleshy cream, chest and throat dark brown. Among the eight species of Microkayla so far described from the Cordillera Real, the new species is superficially similar to M. wettsteini (Parker, 1932) and M. condoriri (De la Riva, Aguayo & Padial, 2007), which furthermore are, together with M. chacaltaya (De la Riva, Padial & Cortez, 2007) and M. teqta (De la Riva & Burrowes, 2014), the species whose localities are geographically closer to that of M. huayna. From M. wettsteini the new species differs mostly by its smaller size (maximum male SVL 26.6 mm [range = 20.9-26.6; ...
... SVL, vs. 70% in M. condoriri [De la Riva 2007]). The new species is distinguished from M. teqta by males being larger (maximum male SVL 26.6 mm vs. 23.6 mm in males of M. teqta), having a large subgular vocal sac (less developed in M. teqta), and lacking yellow or red blotches on dorsum or venter; additionally, the call of the new species is tonal, while that of M. teqta is pulsed (De la Riva & Burrowes 2014). Microkayla huayna sp. ...
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We describe a new species of direct-developing frog of the genus Microkayla from the Cordillera Real of the Bolivian Andes, in the Department of La Paz. The new species, Microkayla huayna sp. nov., is closely related to M. teqta and can be distinguished from other species of the genus by its brown dorsal skin and the presence of a large dark brown vocal sac in males. This is the second species of Microkayla known from the Zongo Valley, and the ninth in the Cordillera Real, contributing to a total of 22 described species in Bolivia. Given its small distribution range, we recommend to considering it as Vulnerable according to IUCN criteria.
... Phrynopus is now restricted to central Peru, whereas all the species occurring in Bolivia were assigned to the genus Psychrophrynella Hedges, Duellman & Heinicke, 2008, where they represent an important proportion of the Andean anuran diversity. Eighteen species of Psychrophrynella, all of them endemic, are known thus far from Bolivia, and many more await description (De la Riva 2007, De la Riva & Burrowes 2014). The diversity of these frogs seems to be especially high in certain regions, among them the Cordillera de Apolobamba, which extends across de Bolivian-Peruvian border. ...
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Three new species of Psychrophrynella from the Bolivian section of the Cordillera de Apolobamba are described. The new species are distinguished from their closest relatives mainly by characters such as colour pattern, size, and skin texture. With the addition of these three new species, the diversity of the genus Psychrophrynella in Bolivia increases to 21 species. The protected Area Natural de Manejo Integrado Nacional (ANMIN) Apolobamba holds seven endemic species, and it is highly likely that more undescribed forms will be discovered when new surveys are conducted in this region, underscoring the need to preserve its rich endemic amphibian fauna.
... The structure of the advertisement call, when known, appears to be similar in both genera, i.e., a whistle-like call, but composed of a single note in Microkayla vs. 2-4 notes in Qosqophryne (except for Q. gymnotis). There is limited information on parental care, but it appears that females attend clutches in Q. gymnotis [39], whereas males attend clutches in M. illimani and M. teqta [50,51]. Similarly to Qosqophryne, females attend clutches in B. cophites [52], B. hanssaueri and B. nubilosus (Catenazzi, pers. ...
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We propose to erect a new genus of terrestrial-breeding frogs of the Terrarana clade to accommodate three species from the Province La Convención, Department of Cusco, Peru previously assigned to Bryophryne: B. flammiventris, B. gymnotis, and B. mancoinca. We examined types and specimens of most species, reviewed morphological and bioacoustic characteristics, and performed molecular analyses on the largest phylogeny of Bryophryne species to date. We performed phylogenetic analysis of a dataset of concatenated sequences from fragments of the 16S rRNA and 12S rRNA genes, the protein-coding gene cytochrome c oxidase subunit I (COI), the nuclear protein-coding gene recombination-activating protein 1 (RAG1), and the tyrosinase precursor (Tyr). The three species are immediately distinguishable from all other species of Bryophryne by the presence of a tympanic membrane and annulus, and by males having median subgular vocal sacs and emitting advertisement calls. Our molecular phylogeny confirms that the three species belong to a new, distinct clade, which we name Qosqophryne, and that they are reciprocally monophyletic with species of Microkayla. These two genera (Qosqophryne and Microkayla) are more closely related to species of Noblella and Psychrophrynella than to species of Bryophryne. Although there are no known morphological synapomorphies for either Microkayla or Qosqophryne, the high endemism of their species, and the disjoint geographic distribution of the two genera, with a gap region of ~310 km by airline where both genera are absent, provide further support for Qosqophryne having long diverged from Microkayla. The exploration of high elevation moss and leaf litter habitats in the tropical Andes will contribute to increase knowledge of the diversity and phylogenetic relationships within Terrarana.
... and Castro-H. 1999, Mendoza-Quijano et al. 2002, Ryan et al. 2010, De la Riva and Burrowes 2014. In Peru, seven species of Craugastoridae are known to exhibit parental care: Bryophryne cophites (Lynch, 1975) (Catenazzi 2006), Lynchius oblitus Motta, Chaparro, Pombal, Guayasamin, De la Riva, andPadial, 2016 (Motta et al. 2016), Phrynopus curator Lehr, Moravec, andCusi, 2012 (Lehr et al. 2012), Pristimantis attenboroughi von May, 2017 (Lehr andvon May 2017), P. buccinator (Rodrigues, 1994) (Carrillo and Catenazzi 2007), P. colodactylus (Lynch, 1979), and P. lymani (Barbour and Noble, 1920) (Duellman and Lehr 2009). ...
... Previous research described a single egg clutch containing 8 eggs in P. illampu, and for P. wettsteini clutches with an average of 18 eggs were found, with paternal clutch-tending being known from the latter (de la , Terceros 2010). In the newly described P. teqta, two egg clutches have been observed holding 28 and 41 eggs, both being tended to by the male (de la Riva & Burrowes 2014). We obtained temperature and relative humidity data for two egg clutches for 28 consecutive days between mid-October and mid-November of 2013. ...
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Amphibian populations have been declining worldwide for decades with a multitude of causes having been identified. Conservationists try to reverse the situation, but for many species, important information on distribution, habitat and ecology are missing, which makes the assessment of conservation priorities problematic. Although South America holds the largest number of extant amphibian species in the world, many of them are poorly studied. This is also the case for most species of Psychrophrynella, a genus of cold-adapted frogs occurring in the high Andes, the majority of which having only recently been described. We organized an extensive field survey to study the ecology of Psychrophrynella illimani, a critically endangered species endemic to a single valley in Bolivia that has not been reported on again since its discovery in 2002. We found P. illimani to be locally common and here report new localities, extending its known distribution. Furthermore, we provide new information on its morphology, ecology, and reproductive behaviour and describe for the first time its call. We also identify and discuss several threats that might affect this species’ survival.
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A new frog of the genus Psychrophrynella is described based on specimens from the Cordillera de Vilcabamba, in the department of Cusco in southeastern Peru. The new species inhabits the humid puna and is only known from its type locality in Challcha, near the road between Vilcabamba and Pampaconas, at 3,707 m asl. This new taxon is assigned to the genus Psychrophrynella based on a narrowest genetic distance of 16S rRNA with P. glauca (8.3%) and the presence of a fold-like tubercle on the inner edges of the tarsus. The description of Psychrophrynella vilcabambensis sp. nov. is based on three individuals. This new species can be differentiated from other members of the genera Psychrophrynella and Noblella by the combination of the following characters: light reddish-brown to tan coloration on the dorsum with dark brown markings, the presence of a thoracic fold, ulnar tubercles, a tubercle on the heel, three tubercles on outer edge of tarsus, and toes with lateral fringes. The SVL of male and female specimens are 16.5 and 16.6 mm, respectively. Resumen.-Describimos una nueva rana del género Psychrophrynella de la Cordillera de Vilcabamba, en el departamento de Cusco al sudeste del Perú. La nueva especie habita la puna húmeda y sólo se conoce de su localidad tipo en Challcha, cerca de la carretera entre Vilcabamba y Pampaconas, a 3,707 m snm. El nuevo taxón se asigna al género Psychrophrynella, basándose en la distancia genética de 16S ARNr más estrecha con P. glauca (8.3%), y la presencia de un tubérculo alargado similar a un pliegue en el borde interior del tarso. Psychrophrynella vilcabambensis sp. nov. fue descrita en base a tres individuos. Esta nueva especie se puede diferenciar de otros miembros de los géneros Psychrophrynella y Noblella por la siguiente combinación de caracteres: coloración marrón rojiza clara a marrón bronceada en el dorso con manchas marrón oscuras, presencia de pliegue torácico, tubérculos cubitales, un tubérculo en el talón, tres tubérculos en el borde exterior del tarso y dedos con rebordes laterales. El SVL del macho y de la hembra es 16.5 y 16.6 mm, respectivamente.
Several groups of direct-developing frogs of the Superfamily Brachycephaloidea (= Terraranae) are high Andean and occur in cloud forests, elfin forests, humid paramos, and grasslands, from Colombia to Bolivia. They belong to different genera and experienced independent radiations, in some cases leading to stunning beta-diversity and high level of endemism. Despite the remarkable accumulation of knowledge on the systematics and diversity of these frogs over the last decades, there has been no attempt to synthesize what is known about their morphology, life history, and evolutionary history. This is the goal of this contribution. Through this review, it becomes evident that similar life histories and ecological niches are associated with a particular, convergent morphology, here referred to as the “phrynopoid” ecomorph, which is also present in members of other groups that occupy those same habitats. It refers to small frogs with plump bodies, short legs, and simple digital tips, presumably adapted to a terrestrial life amidst the humid, mossed floor of paramos and upper cloud forests. Although it is possible to speculate on the mechanisms underlying the remarkable diversity of phrynopoid frogs, the geological and climatic processes that promoted the diversification and the present species diversity are still too poorly known to enable inferences about their evolutionary history. Much fieldwork and integrative studies are needed before we can attain a comprehensive knowledge of this important component of the Andean biota. Hopefully, this review provides a starting point.
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The reproductive biology of Eleutherodactylus cooki was studied in a cave system in San Lorenzo, Puerto Rico. I investigated frequency of multiple clutches in nests, clutch size, nest site locations, reproductive timing, embryonic development, and hatching success for 16 months from September 1995 to December 1996. Males guarded multiple clutches of eggs from up to four different females at a time. The mean clutch size was 17.35 eggs per clutch, and the difference in developmental timing among multiple clutches was between 4-14 days. Triple and quadruple clutches were less frequent than single and double clutches, and were associated with nest location. I examined experimentally the fitness benefit of parental care in the field. Compared to control clutches (males not removed), experimental clutches had significantly lower hatching success. Paternal care offers a significant fitness value through increased hatching success at virtually no cost to males in terms of energy expenditure or missed opportunities to mate. An evaluation of the differential reproductive effort, parental investment, and potential reproductive rate between sexes, considering the operational sex ratio, suggests that sexual selection acts on males in this species. I suggest that the mechanisms of sexual selection are female choice and intrasexual exploitative competition. Males with the greatest reproductive success had more yellow coloration than average in the ventral region and guarded at least one clutch of eggs in recessed surfaces of rock, thereby suggesting that visual cues are important in mate choice. Females were scarce and had higher parental investment, and a lower potential reproductive rate than males. Hence, they represent a resource for which males compete.
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Captive and wild frogs from North and Central America and Australia recently have died with epidermal infections by chytridiomycete fungi. We isolated a chytridiomycete into pure culture from a captive, blue poison dart frog that died at the National Zoological Park in Washington, D.C. Using this isolate, we photographed developmental stages on nutrient agar, examined zoospores with transmission electron microscopy, and inoculated test frogs. This inoperculate chytrid develops either monocentrically or colonially and has thread-like rhizoids that arise from single or multiple areas on the developing zoosporangium. The taxonomically important features of the kinetosomal region of the zoospore indicate that this chytrid is a member of the Chytridiales but differs from other chytrids studied with transmission electron microscopy. Its microtubule root, which begins at kinetosome triplets 9-1 and extends parallel to the kinetosome into the aggregation of ribosomes, is distinctive. Histologic examination of test frogs revealed that the pure culture infected the skin of test frogs, whereas the skin of control frogs remained free of infection. The fungus is described as Batrachochytrium dendrobatidis gen. et sp. nov.
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This paper contains a synopsis of the genus Phrynopus in Bolivia, including the description of twelve new species. With seventeen species currently known, the genus has a much higher diversity in Bolivia than previously expected. Whereas intraspecific variation is evident only in color pattern, interspecific variation is noticeable in size, robustness, shape of head, relative lengths of extremities and digits, and skin texture. However, some species are quite similar in appearance and would be difficult to identify in the field were it not for their extremely restricted distributions. The genus Phrynopus has an amazing radiation and its species diversity has been severely underestimated.
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Chytridiomycosis-induced mortalities often occur upon the emergence of Batrachochytrium dendrobatidis (Bd) in naïve amphibian populations. We report chytridiomycosis-associated mortalities in the wild of the coqui (Eleutherodactylus coqui), a declining direct-developing frog with persistent Bd infections. These findings provide additional evidence of decreased host defenses during cool-dry seasons in Puerto Rico.
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High-alpine life forms and ecosystems exist at the limits of habitable environments, and thus, are especially sensitive to environmental change. Here we report a recent increase in the elevational limit of anurans following glacial retreat in the tropical Peruvian Andes. Three species have colonized ponds in recently deglaciated terrain at new record elevations for amphibians worldwide (5244–5400 m). Two of these species were also found to be infected with Batrachochytrium dendrobatidis (Bd), an emerging fungal pathogen causally associated with global amphibian declines, including the disappearance of several Latin American species. The presence of this pathogen was associated with elevated mortality rates of at least one species. These results represent the first evidence of upward expansion of anurans to newly available habitat brought about by recent deglaciation. Furthermore, the large increase in the upper limit of known Bd infections, previously reported as 4112 m in Ecuador, to 5348 m in this study, also expands the spatial domain of potential Bd pathogenicity to encompass virtually all high elevation anuran habitats in the tropical Andes.
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Amphibians are declining worldwide, but these declines have been particularly dramatic in tropical mountains, where high endemism and vulnerability to an introduced fungal pathogen, Batrachochytrium dendrobatidis (Bd), is associated with amphibian extinctions. We surveyed frogs in the Peruvian Andes in montane forests along a steep elevational gradient (1200-3700 m). We used visual encounter surveys to sample stream-dwelling and arboreal species and leaf-litter plots to sample terrestrial-breeding species. We compared species richness and abundance among the wet seasons of 1999, 2008, and 2009. Despite similar sampling effort among years, the number of species (46 in 1999) declined by 47% between 1999 and 2008 and by 38% between 1999 and 2009. When we combined the number of species we found in 2008 and 2009, the decline from 1999 was 36%. Declines of stream-dwelling and arboreal species (a reduction in species richness of 55%) were much greater than declines of terrestrial-breeding species (reduction of 20% in 2008 and 24% in 2009). Similarly, abundances of stream-dwelling and arboreal frogs were lower in the combined 2008-2009 period than in 1999, whereas densities of frogs in leaf-litter plots did not differ among survey years. These declines may be associated with the infection of frogs with Bd. B. dendrobatidis prevalence correlated significantly with the proportion of species that were absent from the 2008 and 2009 surveys along the elevational gradient. Our results suggest Bd may have arrived at the site between 1999 and 2007, which is consistent with the hypothesis that this pathogen is spreading in epidemic waves along the Andean cordilleras. Our results also indicate a rapid decline of frog species richness and abundance in our study area, a national park that contains many endemic amphibian species and is high in amphibian species richness.
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.
Padial et al. (2014) applied the name Pristimantinae Ohler & Dubois, 2012 to a taxon including the genera Ceuthomantis, Dischidodactylus, Pristimantis, and Yunganastes. However, Ceuthomantidae Heinicke, Duellman, Trueb, Means, MacCulloch & Hedges, 2009, type genus Ceuthomantis Heinicke, Duellman, Trueb, Means, MacCulloch & Hedges, 2009, has priority over Pristimantinae Ohler & Dubois, 2012, a fact that we overlooked and correct herein. Ceuthomantinae is thus the correct subfamily name for the taxon including Ceuthomantis, Dischidodactylus, Pristimantis, and Yunganastes, and Pristimantinae Ohler & Dubois, 2012 is its junior synonym. We provide an amended Figure 22 (page 50) reflecting the current classification of Brachycephaloidea as now listed in Frost (2014) and provide the pertinent correction to page 125 of Appendix 2, which should read as follows: