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ASG Chile Leads Update of the Extinction Risk of Chilean Amphibians for The IUCN Red List of Threatened Species

6 | FrogLog 23 (4), Number 116 (October 2015)
Twelve years have elapsed since the rst workshop aimed
at assessing the extinction risk of Chilean amphibians oc-
curred, with their resulting classication being published
in The IUCN Red List of Threatened SpeciesTM (Universidad de
Concepción, October 2003). Since then, new and dierent lines of
research have contributed additional knowledge for this taxonomic
group (1–4). Much of this work has comprised of updates on dis-
tribution ranges, identication of relevant threats, evaluation of
taxonomic identity and studies on ecological features for several
anuran species inhabiting Chile (salamanders and caecilians are not
present in Chile). There are also newly described endemic species
that have increased the number of species for the country. Given
this and the fact that IUCN Red List assessments have a shelf life
of ten years or less, it was imperative to make eorts to update the
conservation status of Chilean amphibians so that a valid list was
current and useful.
Organized by the Chilean branch of the IUCN SSC Amphibian
Specialist Group (ASG), an assessment process to update the con-
servation status of Chilean amphibians for The IUCN Red List of
Threatened SpeciesTM was initiated in May 2015. The assessment
process followed three steps: 1) compilation of existing published
information, 2) a public consultation period to collect any relevant
information needed for the extinction risk assessment, and 3) an
expert workshop facilitated by two experienced Specialist Group
Chairs and the participation of 19 local herpetologists belonging to
dierent institutions, including academia, NGOs and government
agencies (Fig. 1). The workshop was hosted by the Universidad An-
dres Bello (UNAB), Santiago, Chile, on 910 July 2015. The event
was funded by the Outreach Scheme of the Dirección de Extensión
Académica-UNAB, with additional funding kindly provided by the
IUCN Species Survival Commission.
At the time of writing this note, the assessments were being pre-
pared for the next step in the assessment process, an external re-
view on the application of the IUCN Red List methodology based
on existing documentation. Once this process is completed, species
proles, along with their range maps, will be submitted for publica-
tion on the IUCN Red List of Threatened SpeciesTM. Although there
is a possibility that changes could arise from the review process,
workshop results are summarized here.
Sixty-one species were assessed, encompassing 97% of the na-
tive anuran species previously identied for the country (5). One
assessment concluded that Telmatobius peruvianus is not present in
Chile (although it is still present in Peru), as previous records from
Putre in northern Chile, have been assigned to T. marmoratus based
on phylogenetic analyses (6). Also, the recently revalidated Telmato-
bius laevis (7), was not assessed at the workshop given that no new
data exist for this species since its original description in 1902 and
therefore to date no natural population can be assigned to this spe-
cies. From the species evaluated, 72% were identied as endemic
to Chile. If species having marginal distribution in Argentina are
included, this percentage increases to 90%. In addition, four species
were assessed for the rst time; three recently discovered species:
Alsodes cantillanensis (8), Eupsophus altor (9) and Telmatobufo ignotus
(10); and one taxonomic re-validation: Alsodes coppingeri (11).
Changes in the number of species within each conservation cat-
egory are shown in Fig. 2. The percentage of Threatened species
(i.e., Critically Endangered [CR], Endangered [EN] and Vulnerable
[VU]) increased from 38 to 47%. This increase is explained by the
addition into the Threatened categories of recently discovered or
previously described Data Decient (DD) species (Fig. 2C). It is
important to note that DD does not mean that the species is not
of conservation concern, but appropriate data on its distribution
and/or population is needed to make a consistent assessment of
its risk of extinction. Almost half of the assessed species (45%) ex-
perienced a change in category. Seven DD species changed their
category as follows: Least Concern (LC; one species), VU (2 spp.),
EN (1 sp.) and CR (3 spp.). The species that changed from DD to
CR were Telmatobius dankoi, T. fronteriensis and T. vilamensis, all
ASG Chile Leads Update of the Extinction Risk
of Chilean Amphibians for The IUCN Red List of
Threatened SpeciesTM
1Centro de Investigación para la Sustentabilidad, Universidad Andres Bello,
Santiago, Chile, 2ONG Ranita de Darwin, Santiago, Chile, 3Departamento
de Zoología, Universidad de Concepción, 4Departamento de Ciencias
Básicas, Campus Los Ángeles, Universidad de Concepción, 5Red Chilena
de Herpetología, 6Centro de Estudios Agrarios y Ambientales, Valdivia,
Chile, 7Instituto de Ciencias Marinas y Limnológicas, Universidad Austral
de Chile, Valdivia, Chile, 8Centro de Gestión Ambiental y Biodiversidad,
Facultad de Ciencias Veterinarias y Pecuarias, Universidad de Chile,
Santiago, Chile, 9Laboratorio de Genética y Evolución, Facultad de
Ciencias, Universidad de Chile, Santiago, Chile, 10Instituto de Ecología y
Biodiversidad, 11Programa de Fisiología y Biofísica, Facultad de Medicina,
Universidad de Chile, 12Instituto de Ciencias Ambientales y Evolutivas,
Universidad Austral de Chile, Valdivia, Chile, 13Programa de Bachillerato en
Ciencias, Universidad Santo Tomás, Santiago, Chile, 14Universidad del Bío-
Bío, Chillán, Chile 15IUCN SSC Amphibian Specialist Group.
By 1Claudio Soto-Azat, 1,2Andrés Valenzuela-Sánchez, 3Juan Carlos Ortiz, 4Helen Díaz-Páez, 3Camila Castro, 5Andrés Charrier, 3Claudio Correa,
6,7César Cuevas, 8Gabriel Lobos, 9,10Marco A. Mendez, 11Mario Penna, 1AlexandraPeñael-Ricaurte,12Felipe Rabanal, 13Claudia M. Vélez-R,
14Marcela A. Vidal & 15Ariadne Angulo.
Fig. 1: Herpetologists at the workshop, Universidad Andres Bello. From left to right in
upper row: Mario Penna, Edgardo Flores, Juan Carlos Ortiz, Claudia Vélez, Camila Castro,
Felipe Rabanal, Claudio Correa, Marco Mendez, Sandra Díaz, Mariella Superina, Reinaldo
Avilés, César Cuevas, Claudio Soto, Andrés Charrier and Ariadne Angulo. Lower row:
Andrés Valenzuela and Charif Tala. Absent from the picture: Marcela Vidal, Helen Diaz
and Alexandra Peñafiel. Photo: Claudio Soto-Azat.
FrogLog 23 (4), Number 116 (October 2015) | 7
from northern Chile (Fig. 3). When DD species are not considered,
changes in categories involve six up-listings and 12 down-listings.
Among the latter, only two species were removed from a Threat-
ened status. Two CR species were considered as Possibly Extinct
(a tag used in conjunction with the CR category to describe those
instances where there is a possibility a species may be extinct):
Rhinoderma rufum and Telmatobius pefauri. In fact, R. rufum has not
been recorded since 1980, despite numerous attempts to nd it
(12,13). The reasons for the sudden decline of this species are not
fully understood, but the extensive habitat loss across its historical
distribution and possibly chytridiomycosis could have played an
important role (13). On the other hand, Telmatobius pefauri is only
known from its holotype, collected in 1976 at the locality of Mur-
muntani in northern Chile (14). This species has not been observed
since, in spite of attempts to nd it.
The main threats identied for Chilean amphibians (3,4,15) are:
a) water scarcity due to anthropogenic modication of natural
systems, b) mining activities in northern and central Chile; c) im-
pacts related to agriculture, d) residential development in central
and southern Chile, e) exotic tree plantations, and f) anthropogenic
res in southern Chile, which cause loss of habitat and refuges. In
addition, livestock pressure and invasive species (particularly sal-
monids and the African Clawed Frog Xenopus laevis) were cited as
threats to amphibians across the country.
Following IUCN criteria for assessing species in one of the
Threatened categories, most (76%) of the amphibians were catego-
rized by their restricted geographic range (criteria B1 and B2). An-
other 10% were assessed according to a population size reduction
(criteria A) and 14% following very small or restricted populations
(criteria C). This overview highlights an imperative necessity for
local herpetologists to conduct studies and generate data on popu-
lation ecology, helping make assessment more informative in the
future, with the nal purpose of contributing to the conservation
of these species.
We are very grateful to all those herpetologists who made
possible this fruitful assessment process. Special thanks to Dr.
Mariella Superina, Chair of the IUCN SSC Anteater, Sloth and
Armadillo Specialist Group, who acted as expert facilitator at the
workshop. We would also wish to extend our acknowledgements
to all the people who supported the assessment process, providing
valuable information through the open consultation period, and to
the IUCN SSC Amphibian Red List Authority and IUCN Red List
Unit for processing these assessments. We believe the workshop
results will be an important contribution to the conservation of
these amazing animals.
1. F. E. Rabanal, J. J. Nuñez, Anbios de los bosques templados de Chile
(Universidad Austral de Chile, Valdivia, Chile, 2008).
2. M. A. Vidal, A. Labra, Eds, Herpetología de Chile (Science Verlag, Santiago,
Chile, 2008).
3. C. Soto-Azat, A. Valenzuela-Sánchez, Eds, Conservación de anbios de Chile
(Univ. Andrés Bello, Santiago, Chile, 2012).
4. G. Lobos et al., Anbios de Chile, un desafío para la conservación (Ministerio
del Medio Ambiente, Fundación Facultad de Ciencias Veterinarias y Pecuarias
de la Univ. de Chile y Red Chilena de Herpetología, Santiago de Chile, 2013).
5. D. R. Frost, Amphibian Species of the World: an Online Reference. Version 6.0. (2015).
6. P. A. Saéz et al., Zool. J. Lin. Soc., 171, 769 (2014).
7. C. C. Cuevas, Herpetol. J., 23, 145 (2013).
8. A. Charrier, C. Correa, C. Castro, M. A. Méndez, Zootaxa, 3,915, 540 (2015).
9. J. J. Nuñez, F. Rabanal, J. R. Formas, Zootaxa 3,305, 53 (2012).
10. C. C. Cuevas, Gayana 74, 102 (2010).
11. J. R. Formas, J. J. Nuñez, C. C. Cuevas, Rev. Chil. Hist. Nat., 81, 3 (2008).
12. J. Bourke, K. Busse, W Böhme, North-West. J. Zool. 8, 99 (2012).
13. C. Soto-Azat et al., PLOS One 8: e79862 (2013).
14. A. Veloso, L. Trueb, Occas. Pap. Mus. Nat. Hist. (Lawrence), 1 (1976).
 
n  
n  
   
 
 
 
   
Fig. 2: Percentage of Chilean amphibian species within each conservation category at A)
present (IUCN Red List assessments, accessed on 20 August 2015), and B) under the new
proposed assessments. The changes in these percentages per category are presented
in C), where negative values (blue) indicate a decrease in the percentage of species
within the category, while the positive values (red) indicates an increase. CR: Critically
Endangered; VU: Vulnerable; EN: Endangered; NT: Near Threatened; LC: Least Concern;
DD: Data Deficient.
Fig. 3: Telmatobius vilamensis from northern Chile. Previously assessed as Data Deficient
and now proposed as Critically Endangered as a result of the Chilean IUCN Red List
workshop. Photo: Felipe Rabanal.
... This included the finding of Bd infection in a Puerto Eden frog (Chaltenobatrachus grandisonae) near Villa O'Higgins ( Fig. 1A), extending the previously known southernmost global record of Bd by 588 km further 59,60 . From the total of 40 sampled anuran species in Chile (64% of total richness) 49 , we found that 24 species showed evidence of Bd infection (see Table 1). All 11 species which had a Bd infection prevalence > 30% were exclusively aquatic amphibians, likely due to a higher contact rate with the infective stage of Bd in the aquatic environment 53,61 . ...
... The highest observed Bd prevalence was in the Chilean Matorral ecoregion, an area considered as a priority for global biodiversity conservation 83 . This region harbours a high level of anuran endemism 49 , yet contains the highest human population density in the country (almost 90% of the Chilean population). Consequently, increasing urbanization is resulting in deforestation and habitat loss which is negatively impacting amphibian populations 84 . ...
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Amphibian chytridiomycosis, caused by the fungus Batrachochytrium dendrobatidis (Bd), has caused the greatest known loss of biodiversity due to an infectious disease. We used Bd infection data from quantitative real-time PCR (qPCR) assays of amphibian skin swabs collected across Chile during 2008–2018 to model Bd occurrence with the aim to determine bioclimatic and anthropogenic variables associated with Bd infection. Also, we used Bd presence/absence records to identify geographical Bd high-risk areas and compare Bd prevalence and infection loads between amphibian families, ecoregions, and host ecology. Data comprised 4155 Bd-specific qPCR assays from 162 locations across a latitudinal gradient of 3700 km (18º to 51ºS). Results showed a significant clustering of Bd associated with urban centres and anthropogenically highly disturbed ecosystems in central-south Chile. Both Bd prevalence and Bd infection loads were higher in aquatic than terrestrial amphibian species. Our model indicated positive associations of Bd prevalence with altitude, temperature, precipitation and human-modified landscapes. Also, we found that macroscale drivers, such as land use change and climate, shape the occurrence of Bd at the landscape level. Our study provides with new evidence that can improve the effectiveness of strategies to mitigate biodiversity loss due to amphibian chytridiomycosis.
... Exotic fish may affect native anurans by predating eggs, larvae and adults (Welsh et al. 2006;Jara and Perotti 2006). Although the number of replicates we used to evaluate predation of larvae was low due to bioethical (Russell and Burch 1959;Santos et al. 2009) and legal considerations (Soto-Azat et al. 2015), there was a clear tendency; A. facetus indistinctly predated larvae of all anuran species tested, with much less predatory activity by smaller fish (both native C. pisciculus and non-native G. holbrooki). It is probable that smaller fish had a physical limitation for eating larvae (mean length 30-40 mm, to predate on larvae with mean lengths of 26-31 mm), although it has been suggested that some species such as those of the genus Rhinella may have developed toxins that make them unpalatable (Kiesecker et al. 1996;Jara and Perotti 2006). ...
... The vulnerability of Chilean anurans to exotic fish is worrying, since all the anurans tested have been recognized as having some degree of threat (Díaz-Páez and Ortiz 2003;Soto-Azat et al. 2015). This study recognizes that the exotic fish are a high-risk factor for the Chilean Mediterranean zone (which may also occur in other Mediterranean areas), especially due the isolation in the evolution of its biota. ...
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Mediterranean biomes have been recognized as having high value due to their diversity; they are under threat due to anthropic pressures. Both freshwater fish and amphibians show high endemism and vulnerability to several threats; among the latter is the introduction of fishes. In Chile there is scarce documentation of the presence of the non-native Australoheros facetus (chameleon cichlid), where it cohabits with other non-native and native fish and anurans. In this study we first sampled for the presence/absence of A. facetus in 69 localities in the Chilean Mediterranean region. Secondly, to estimate their potential impact, we evaluated the predatory capacity of A. facetus and other two sympatric fish, Gambusia holbrooki (mosquitofish; non-native) and Cheirodon pisciculus (pocha; native), on larvae of four species of native anurans and on the eggs of a non-native anuran. We report that A. facetus has expanded in the entire Chilean Mediterranean region; its presence is related to low altitude areas. We confirmed that lentic environments and irrigation systems of streams increase the probability of successful establishment. The results for predatory capacity showed that A. facetus regularly predated larvae of all the tested anurans, being significantly more successful in predating larvae, but also significantly less successful in predating eggs, compared to other fish species. The control of non-native fish is a priority to avoid the invasion of new localities in these ecosystems.
... Regarding the conservation status of the genus, the Chilean species are classified according to the IUCN as data deficient (2), critical risk (5) and vulnerable (2). A recent report of the Chilean branch of the IUCN SSC Amphibian Specialist Group (ASG) indicated the urgent necessity for local herpetologists to conduct more studies to help make assessment more informative in the future, with the final purpose of contributing to the conservation of Chilean species [9]. ...
... In the case of T. marmoratus, T. fronteriensis, T. peruvianus and T. philippii, for the crossamplification, 6 microsatellites markers (Tchus_3, Tchus_7, Tchus_11, Tchus_21, Tchus_22 and Tchus_25) were polymorphic among all species tested. No data for Telmatobius halli was obtained in this study, because it has never been seen after it was first described [6,9]. The characterization of these 22 markers is useful for further population genetic studies, to provide a better understanding to undertake future : TAC ATA ATA TAA GCC AGG TC Tchus_2 F: CTG TAC CTC ATT ATG TAA ACT (TA)4 ...
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The genus Telmatobius Wiegmann, 1834 is composed of a wide variety of species and is one of the most diverse native com- ponents of the high-altitude Andean environments. The species of the genus present in Chile are considered as endangered, critically endangered or data deficient. We isolated and evaluated 44 microsatellites in 80 individuals of 8 species of Telma- tobius present in Chile, obtaining 22 polymorphic microsatellite loci for Telmatobius chusmisensis. The cross-amplification test was successful in all other species tested. For the first time, microsatellite markers are described for Telmatobius. The description of these primers will be useful for further genetic studies for T. chusmisensis and other species of the same genus; allowing further analyses of population structuring, dispersal patterns, recent demographic history and population effective size. This information is also significant to undertake conservation actions for the species of the genus Telmatobius, since most species have conservation issues.
... Currently, no ranavirus lineage is known to be endemic to Chile, and available data suggests that the virus may not be native to the country (Soto-Azat et al., 2016). Chile is known for its high rate of amphibian and fish endemism, with most endemic species being threatened and restricted to small areas (Soto-Azat et al., 2015), which makes them prone to the negative impacts of stochastic events such as the emergence of a disease such as ranavirosis. However, the epidemiology of ranavirus in Chile has not been studied; for instance, the origin, extent and impacts of ranavirus are unknown. ...
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Ranaviruses have been associated with amphibian, fish and reptile mortality events worldwide and with amphibian population declines in parts of Europe. Xenopus laevis is a widespread invasive amphibian species in Chile. Recently, Frog virus 3 (FV3), the type species of the Ranavirus genus, was detected in two wild populations of this frog near Santiago in Chile, however, the extent of ranavirus infection in this country remains unknown. To obtain more information about the origin of ranavirus in Chile, its distribution, species affected, and the role of invasive amphibians and freshwater fish in the epidemiology of ranavirus, a surveillance study comprising wild and farmed amphibians and wild fish over a large latitudinal gradient (2,500 km) was carried out in 2015–2017. In total, 1,752 amphibians and 496 fish were tested using a ranavirus-specific qPCR assay, and positive samples were analyzed for virus characterization through whole genome sequencing of viral DNA obtained from infected tissue. Ranavirus was detected at low viral loads in nine of 1,011 X. laevis from four populations in central Chile. No other amphibian or fish species tested were positive for ranavirus, suggesting ranavirus is not threatening native Chilean species yet. Phylogenetic analysis of partial ranavirus sequences showed 100% similarity with FV3. Our results show a restricted range of ranavirus infection in central Chile, coinciding with X. laevis presence, and suggest that FV3 may have entered the country through infected X. laevis , which appears to act as a competent reservoir host, and may contribute to the spread the virus locally as it invades new areas, and globally through the pet trade.
... Introduced salmonids are another threat for native amphibians in Chile (Soto-Azat et al. 2015). A recent study reports the presence of Oncorhynchus mykiss (Walbaum, 1792) in several locations of the upper Loa, being Sapunta the nearest sampling point to the source (approx. ...
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Telmatobius halli was the first representative of its genus to be described exclusively for Chile, yet for 85 years no new individuals could be located due to the vagueness with which its type locality was described. The type series was collected by one of the members of the International High Altitude Expedition to Chile (IHAEC) of 1935. Recently, three studies successively claimed to have located the type locality in different places. The third study proved, according to the chronicles of the IHAEC, that the actual locality is Miño, at the origin of the Loa River, where currently there are no published records of Telmatobius . In this study, additional documentary antecedents and graphic material are provided that corroborate that Miño is indeed the type locality of T. halli. Additionally, the recently rediscovered Telmatobius population from Miño and the environment it inhabits are described. The external characteristics of the frogs are consistent with the description of T. halli . Furthermore, molecular phylogenetic analyses were performed that showed that T. halli , T. dankoi , and T. vilamensis , all known only from their type localities in Chile, comprise a clade without internal resolution. A detailed comparison of the diagnoses of the three species revealed that the few phenotypic differences between these taxa were based on characteristics that vary widely within and between populations of the genus, hence their conspecificity is proposed. The implications of this synonymy for the taxonomy, biogeography, and conservation of the Telmatobius from the extreme south of its distribution in Chile are discussed.
... Rhinoderma darwinii está categorizada En Peligro de extinción (EN) por la Lista Roja de la UICN (IUCN SSC Amphibian Specialist Group 2018). El reciente cambio en la categoría de conservación de esta especie en la Lista Roja de la UICN de Vulnerable a En Peligro, surgió del taller de reevaluación de anfibios de Chile para la Lista Roja (ver Soto-Azat et al. 2015). La justificación de su actual categoría se basa en una limitada área de ocupación (estimada en 264 km 2 ), severa fragmentación de sus poblaciones y continua declinación de su hábitat. ...
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Binational Conservation Strategy fro Darwin's frogs
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Telmatobius halli Noble, 1938 was the first representative of its genus to be described for Chile, but for 80 years no new individuals could be located due to the vagueness with which its type locality was described. The type series was collected by one of the members of the International High Altitude Expedition to Chile (IHAEC) that took place in 1935. Recently, three studies successively claimed to have rediscovered the type locality in different places. The third study proved, considering the chronicles of the IHAEC, that the actual locality is Miño, at the origin of the Loa River. In the contemporary herpetological literature, there are no records of Telmatobius from this locality. In this study, we provide additional documentary antecedents and graphic material that corroborate that the mentioned location is indeed the historic type locality of T. halli . Additionally, we describe the recently discovered Telmatobius population from Miño, whose external characteristics are consistent with the description of T. halli , and the environment it inhabits. Furthermore, we performed a molecular phylogenetic analysis that strongly suggests that T. halli (from Miño), T. dankoi and T. vilamensis , all known only from their type locality in Chile, are conspecific. Neither of the populations from the previously proposed rediscoveries grouped with the one from the genuine type locality. We discuss the implications that these findings have for the taxonomy, biogeography and conservation of the populations from the extreme south of the distribution of the genus in Chile.
Full-text available
Darwin's frogs Rhinoderma darwinii and Rhinoderma rufum are the only known species of amphibians in which males brood their offspring in their vocal sacs. We propose these frogs as flagship species for the conservation of the Austral temperate forests of Chile and Argentina. This recommendation forms part of the vision of the Binational Conservation Strategy for Darwin's Frogs, which was launched in 2018. The strategy is a conservation initiative led by the IUCN SSC Amphibian Specialist Group, which in 2017 convened 30 governmental, non-profit and private organizations from Chile, Argentina and elsewhere. Darwin's frogs are iconic examples of the global amphibian conservation crisis: R. rufum is categorized as Critically Endangered (Possibly Extinct) on the IUCN Red List, and R. darwinii as Endangered. Here we articulate the conservation planning process that led to the development of the conservation strategy for these species and present its main findings and recommendations. Using an evidence-based approach, the Binational Conservation Strategy for Darwin's Frogs contains a comprehensive status review of Rhinoderma spp., including critical threat analyses, and proposes 39 prioritized conservation actions. Its goal is that by 2028, key information gaps on Rhinoderma spp. will be filled, the main threats to these species will be reduced, and financial, legal and societal support will have been achieved. The strategy is a multi-disciplinary, transnational endeavour aimed at ensuring the long-term viability of these unique frogs and their particular habitat.
Full-text available
Darwin’s frogs Rhinoderma spp. are the only known mouth-brooding frogs on Earth. The southern Darwin’s frog R. darwinii is found in the temperate forests of southern South America, is listed as endangered, and could be the only extant representative of this genus. Based on stomach content, invertebrate prey availability and stable isotope analysis, we determined for the first-time trophic ecological parameters for this species. Our results showed that R. darwinii is a generalist sit-and-wait predator and a secondary consumer with a trophic position of 2.9. Carbon and nitrogen isotope composition indicated that herbivore invertebrates were their main prey, detected in 68.1% of their assimilated food. The most consumed prey included mosquitoes, flies, crickets, grasshoppers and ants. Detritivore and carnivore invertebrates were also ingested, but in lower proportions. Our results contribute to a better understanding of the feeding habits of this fully terrestrial amphibian and provide the first insight on their role linking low forest trophic positions with intermediate predators. We provide valuable biological information for in situ and ex situ conservation, which can be used when developing habitat protection, reintroduction and captive breeding programmes. As revealed here, stable isotope analysis is a valuable tool to study the trophic ecology of highly endangered and cryptic species.
A new species, Telmatobufo ignotus, from the Reserva Nacional Los Queules, Cauquenes Province, central Chile is described herein. The new taxon is hypothesized based on a new combination of adult, larval, and chromosomal characters. The new species present the more septentrional distribution on the coastal range and a very restrictive distribution area compared to its congeners; because of this, taxonomic and conservation comments are discussed and advanced.
  • J R Formas
  • J J Nuñez
  • C C Cuevas
J. R. Formas, J. J. Nuñez, C. C. Cuevas, Rev. Chil. Hist. Nat., 81, 3 (2008).
  • A Charrier
  • C Correa
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A. Charrier, C. Correa, C. Castro, M. A. Méndez, Zootaxa, 3,915, 540 (2015).
  • J Bourke
  • K Busse
  • Böhme
J. Bourke, K. Busse, W Böhme, North-West. J. Zool. 8, 99 (2012).
  • A Veloso
  • L Trueb
A. Veloso, L. Trueb, Occas. Pap. Mus. Nat. Hist. (Lawrence), 1 (1976).
Amphibian Species of the World: an Online Reference
  • D R Frost
D. R. Frost, Amphibian Species of the World: an Online Reference. Version 6.0. (2015).
  • C Soto-Azat
C. Soto-Azat et al., PLOS One 8: e79862 (2013).
  • J J Nuñez
  • F Rabanal
  • J R Formas
J. J. Nuñez, F. Rabanal, J. R. Formas, Zootaxa 3,305, 53 (2012).
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P. A. Saéz et al., Zool. J. Lin. Soc., 171, 769 (2014).