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Accepted by J. Padial: 28 Oct. 2014; published: 26 Nov. 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3887 (3): 459
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http://dx.doi.org/10.11646/zootaxa.3887.4.4
http://zoobank.org/urn:lsid:zoobank.org:pub:06CD5DE9-10BE-4DFF-8E5E-FD27808DA6E7
A new species of Psychrophrynella (Anura: Craugastoridae) from the Cordillera
Real, Department La Paz, Bolivia
IGNACIO DE LA RIVA
1
& PATRICIA A. BURROWES
2
1
Museo Nacional de Ciencias Naturales, CSIC, C/ José Gutiérrez Abascal, 28006 Madrid, Spain. E-mail: iriva@mncn.csic.es
2
Department of Biology, University of Puerto Rico, P.O. Box 23360, San Juan, PR 00931
Abstract
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
Introduction
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
(2009).
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 (www.fonozoo.com).
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®.
Systematics
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
mm).
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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
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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
9.9.
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-
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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 www.jampatu.org). 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
swab
), adult male MNCN 45703 had 130,400 Z
swab
. 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, bolivianamphibianinitiative.org and the website of the P.
illimani project, www.jampatu.org).
Acknowledgments
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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