Named after Stanley Kubrick, a new species of frog is a ‘clockwork orange’ of nature

Two new treefrog species were discovered in the Amazon Basin of Bolivia, Peru, and Brazil. Both had been previously misidentified as another superficially identical species.

Curiously, one of them received a name translating to ‘demon’ or ‘devil’. The second one was named in honor of famous American filmmaker Stanley Kubrick, because of his masterpiece A Clockwork Orange.

Having conducted analyses of genetic, morphologic, and bioacoustic data, scientists C. Daniel Rivadeneira, Dr. Pablo J. Venegas, and Dr. Santiago R. Ron concluded that the amphibians represented two previously unknown species that used to go by the name of the Sarayacu treefrog (Dendropsophus parviceps).

As a result of this research collaboration between Ecuador’s Catholic University (PUCE) and Peru’s Centro de Ornitología y Biodiversidad (CORBIDI), the treefrogs were recently described as new to science in the open access journal ZooKeys.

DSC06331The scientists remind that, back in 1972, when Anthony Burgess explained the title of his famous novel A Clockwork Orange, he said: “I’ve implied the junction of the organic, the lively, the sweet – in other words, life, the orange – and the mechanical, the cold, the disciplined (…)”

“Without knowing, he was also giving a good metaphor to describe ecosystems,” comment the researchers. “Nature works as the interplay between life and its cold, mechanical, and disciplined physical matrix.”

Furthermore, both new frogs, scientifically listed as D. kubricki and D. kamagarini, are characterized by having a bright orange blotch on the shanks, reminiscent of the ‘orange pieces of nature’.

On the other hand, the species name kamagarini refers to ‘demon’ or ‘devil’, as per its translation from the Matsigenka language spoken in southeastern Peru. One of the characteristic features of the new species D. kamagarini are namely its horn-like protuberances on the upper eyelids.Bolivia

Amphibians are important pieces in ecosystems as secondary consumers in food chains. They also play a significant role in decomposition and nutrient cycling.

Stanley Kubrick – arguably one of the most brilliant and influential film directors of all time – left an immemorial legacy in cinema. His masterpiece, A Clockwork Orange (1971), was based on Anthony Burgess’s 1962 novel of the same name.

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Original source:

Rivadeneira CD, Venegas PJ, Ron SR (2018) Species limits within the widespread Amazonian treefrog Dendropsophus parviceps with descriptions of two new species (Anura, Hylidae). ZooKeys 726: 25-77. https://doi.org/10.3897/zookeys.726.13864

New species of frog from the Neotropics carries its heart on its skin

In the Neotropics, there is a whole group of so-called glassfrogs that amaze with their transparent skin covering their bellies and showing their organs underneath. A recently discovered new species from Amazonian Ecuador, however, goes a step further to fully expose its heart thanks to the transparent skin stretching all over its chest as well as tummy.

The new amphibian is described by a team of scientists led by Dr. Juan M. Guayasamin, Universidad San Francisco de Quito, Ecuador, in the open access journal ZooKeys.

It can also be distinguished by the relatively large dark green spots at the back of its head and the foremost part of the body. Additionally, the species has a characteristic long call.

The new frog is named Hyalinobatrachium yaku, where the species name (yaku) translates to ‘water’ in the local language Kichwa. Water and, more specifically, slow-flowing streams are crucial for the reproduction of all known glassfrogs.

The reproductive behaviour is also quite unusual in this species. Males are often reported to call from the underside of leaves and look after the egg clutches.

Having identified individuals of the new species at three localities, the researchers note some behavioural differences between the populations. Two of them, spotted in the riverine vegetation of an intact forest in Kallana, have been calling from the underside of leaves a few metres above slow-flowing, relatively narrow and shallow streams. Another frog of the species has been observed in an area covered by secondary forests in the Ecuadorian village of Ahuano. Similarly, the amphibian was found on the underside of a leaf one metre above a slow-flowing, narrow and shallow stream.

oo_135330However, at the third locality – a disturbed secondary forest in San José de Payamino – the studied frogs have been perching on leaves of small shrubs, ferns, and grasses some 30 to 150 cm above the ground. Surprisingly, each of them has been at a distance greater than 30 metres from the nearest stream.

The researchers note that, given the geographic distance of approximately 110 km between the localities where the new species has been found, it is likely that the new species has a broader distribution, including areas in neighbouring Peru.

The uncertainty about its distributional range comes from a number of reasons. Firstly, the species’ tiny size of about 2 cm makes it tough to spot from underneath the leaves. Then, even if specimens of the species have been previously collected, they would be almost impossible to identify from museum collection, as many of the characteristic traits, such as the dark green marks, are getting lost after preservation. This is why the conservation status of the species has been listed as Data Deficient, according to the IUCN Red List criteria.

Nevertheless, the scientists identify the major threats to the species, including oil extraction in the region and the related water pollution, road development, habitat degradation and isolation.

“Glassfrogs presumably require continuous tracts of forest to interact with nearby populations, and roads potentially act as barriers to dispersal for transient individuals,” explain the authors.

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Original source:

Guayasamin JM, Cisneros-Heredia DF, Maynard RJ, Lynch RL, Culebras J, Hamilton PS (2017) A marvelous new glassfrog (Centrolenidae, Hyalinobatrachium) from Amazonian Ecuador. ZooKeys 673: 1-20. https://doi.org/10.3897/zookeys.673.12108

New frog from the Peruvian Andes is the first amphibian named after Sir David Attenborough

While there are already a number of species named after famous British broadcaster and naturalist Sir David Attenborough, including mammals, reptiles, invertebrates and plants, both extinct and extant, not until now has the host of the BBC Natural History’s Life series been honoured with an amphibian.

A new fleshbelly frog, recently discovered in the Peruvian Andes, is formally described as Pristimantis attenboroughi, while commonly it is to be referred to as the Attenborough’s Rubber Frog. The new species is published in the open-access journal ZooKeys.

Scientists Dr. Edgar Lehr, Illinois Wesleyan University, and Dr. Rudolf von May, University of Michigan, spent two years (2012-2014) surveying montane forests in central Peru, in order to document the local amphibians and reptiles, and evaluate their conservation statuses. Their efforts have been rewarded with several new species of frogs and a new spectacled lizard.

Each of these discoveries, including the Attenborough’s Rubber Frog, prove how beneficial it is to take into account both morphological and the genetic data, while looking for species new to science. For example, the authors report that when they spotted the P. attenboroughi frog for the first time, both of them were sure that they had found a species of another genus.

FIG. 2-female guarding eggs-not croppedThe Attenborough’s rubber frog is known to inhabit several localities across the Pui Pui Protected Forest, a nature reserve located at elevations between 3400 and 3936 m a.s.l. in central Peru. The adult males reach size of 14.6-19.2 mm in length, while the females are larger measuring between 19.2 and 23.0 mm. Their ground colour ranges from pale to dark gray, or reddish brown to brownish olive with dark gray scattered flecks. Meanwhile, the juveniles are paler (yellowish to reddish brown) with contrasting dark brown flecks and distinct stripes.

Due to the amphibian being known from fewer than ten localities, spread across less than 20,000 km2, the species should be deemed either Vulnerable or Endangered, according to the IUCN Red List Categories and Criteria. However, the authors suggest that the Attenborough’s Rubber frog should be listed as Near Threatened instead, since the Piu Piu forest is formally protected and still largely unknown, so it is likely that there are more additional populations of the new species. On the other hand, factors such as fungal infections, climate change, pollution, and man-made fires continue to be threats for many Andean amphibians even inside protected areas.

“We dedicate this species to Sir David Frederick Attenborough in honor for his educational documentaries on wildlife, especially on amphibians (e.g., Life in Cold Blood, Fabulous Frogs), and for raising awareness about the importance of wildlife conservation,” explain the authors.

In the present study, the scientists note that there are more terrestrial-breeding frogs from the surveyed montane forests that will be described in the near future.

Among the numerous namesakes of Sir David Attenborough to date, there are a rare genus of beautiful flowering plants, a rare butterfly species, commonly known as the Attenborough’s black-eyed satyr, a flightless weevil species, as well as a number of extinct species.

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Original source:

Lehr E, von May R (2017) A new species of terrestrial-breeding frog (Amphibia, Craugastoridae, Pristimantis) from high elevations of the Pui Pui Protected Forest in central Peru. ZooKeys 660: 17-42. https://doi.org/10.3897/zookeys.660.11394

Exceptional reproductive biology in extremely restricted critically endangered Nimba toad

The critically endangered Nimba toad is long known for its exceptional reproductive biology. The females of this unique species give live birth to fully developed juveniles, having for nine months continuously provided nutrition to the foetuses in the womb (matrotrophy). While live birth (viviparity) among frogs and toads is rather an exception than a common characteristic, matrotrophy, in place of alternatives such as the foetus being fed with yolk, unfertilized eggs, or smaller siblings, is what makes the Nimba toad one of a kind.

However, more than 40 years of research had not been comprehensively, accessibly and completely summarised. The gap has recently been filled with a new paper, published in the open access journal Zoosystematics and Evolution by German scientists Drs. Laura Sandberger-Loua and Mark-Oliver Rödel, both affiliated with Museum für Naturkunde, Berlin, and Dr. Hendrik Müller, Friedrich-Schiller-Universität Jena.

Studying the phenomenon, the scientists went through the literature published over four decades to gather the scattered details. They have also discussed the relationship between the toad’s reproductive biology and its specific habitat of merely 4 km² of high altitude grasslands located at a minimum of 1,200 m in the Nimba mountains, West Africa.

The climate of the area is characterised by a rainy season lasting from April to October and a dry season from November to February/March. These seasons are found to determine the activity of the Nimba toads. The amphibians are only active during the rainy season, when they give birth to their young, mate, and then find shelter underground, where they stay dormant during the dry season.

10489_ZSE_blogVisibly females can be distinguished from male Nimba toads by their differing cloaca and often larger size, compared to the males. Also, males show darker backs and, during most of their adult life, nuptial pads on their thumbs, which look like spiky swellings. This secondary sex characteristic, in its seasonal change linked to spermatogenesis, is used by the males to grasp tightly the female while mating.

In this species mating occurs without a copulatory organ. Instead, the sperm is transferred through connection of the cloacae, where the male’s swells and encloses the female’s cloaca. Furthermore, Nimba toads have a unique behavioural repertoire. Males crouch on their front legs and as soon as the female moves, follow her and grab her tightly in the groin. Due to the spiky nuptial pads, the males often injure their partner.

10489_ZSE_Image 3Giving birth in Nimba toads may take over two days, depending on the number of offspring, which can be up to 12 in older females – far fewer than the hundreds of eggs in most toad species. While giving birth, a female assumes a unique “birthing posture” to compensate for the lack of enough muscle power to expel juveniles. By the time the juveniles are ready to be born, they have already taken up nearly all the space in their mother’s body. The scientists conclude that the offspring play an active role in the process, as a juvenile toad’s death midway in the oviduct leads to the mother dying of sepsis.

Living exclusively in the Nimba mountains, and being listed as Critically Endangered, according to the IUCN, the studied toad is only one of the species restricted to the high altitude grasslands, which led to the declaration of the Nimba mountains as a World Heritage Site by UNESCO. Separated from other mountains, its inhabitants are isolated from external contacts, thus presumably leading to their evolutionary separation from related species. Furthermore, the toad’s unique reproductive biology is probably the result of this isolation, argue the authors.

In conclusion, the authors suggest that “it is likely that the harsh unpredictable environment and scarcity of open water promoted viviparity in Nimba toads, or supported the survival of this unique reproductive mode in these special and isolated conditions. Considering their complex life cycle, in which reproductive and seasonal cycles are tightly linked, understanding and protecting the Nimba toad’s threatened environment is of utmost importance.”

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Original source:

Sandberger-Loua L, Müller H, Rödel M-O (2017) A review of the reproductive biology of the only known matrotrophic viviparous anuran, the West African Nimba toad, Nimbaphrynoides occidentalis. Zoosystematics and Evolution 93(1): 105-133. https://doi.org/10.3897/zse.93.10489

Twenty-five frogs added to the amphibian fauna of Mount Oku, Cameroon

 

While amphibians all over the world are undergoing a continuous decline, their status in certain regions, such as Central Africa, remains unknown due to incomplete information. New paper, published by two scientists in the open access journal ZooKeys, addresses the knowledge gap by providing an updated list of already 50 amphibian species living on Mount Oku, Cameroon.

Scientists Dr Thomas M. Doherty-Bone, Royal Zoological Society of Scotland, and Dr Václav Gvoždík, affiliated with both the Czech Academy of Sciences and the National Museum in Prague, have spent more than 10 years studying the Cameroonian mountain. As a result of their thorough surveys, literature review, and re-examination of museum specimens, there are now 50 species known from the locality, which doubles previous numbers. In their newly published checklist the researchers have listed 49 species of frogs and toads, as well as one caecilian species – a limbless, snake-like amphibian.

However, the number of threatened species seems to increase quite proportionally. Many of the newly recorded frogs, for instance, appear to be extremely endangered, yet they have not been assessed by the International Union for Conservation of Nature (IUCN). Therefore, the authors have used the IUCN criteria to propose conservation assessments for them. If the suggested statuses are approved, together with the updated declines of previously abundant in the area species, the proportion of the threatened would rise to nearly half (48%) of the Mount Oku’s entire amphibian fauna. Meanwhile, it is 42% of amphibians at risk of extinction globally.

In their study, the scientists also review the research and conservation undertaken at the mountain so far, including the work they have initiated themselves. Although Mount Oku’s forest turned out to be the best managed among the rest in the region, threats such as forest loss, encroachment and degradation are still largely present and increasing. Additional threats, including use of agrochemicals, climate change and diseases, have also been identified. However, conservation actions for the amphibians of Mount Oku are on the rise, considering both the population and the ecosystem-level perspectives.

“Our paper provides a foundation for continuously improving amphibian conservation at Mount Oku, as well as other mountains in Cameroon,” conclude the authors.   

 

Original source:

Doherty-Bone TM, Gvoždík V (2017) The Amphibians of Mount Oku, Cameroon: an updated species inventory and conservation review. ZooKeys 643: 19-139. https://doi.org/10.3897/zookeys.643.9422

Disentangling the myth of the singing bushmaster viper with the help of tree frogs

Reaching over 3.5 m in length, the bushmaster (belonging in the Lachesis genus) is the largest viper in the western hemisphere. Legend spread among both colonists and natives from the Amazon region and Central America has it that it sings. Finding these numerous unrelated reports quite puzzling, since it is well known that snakes cannot sing, scientists took to finally disentangle the myth.photo-2

When the researchers recently conducted fieldwork in Amazonian Ecuador and Peru, they revealed it was not the snake singing. The ‘song’ was indeed the call of large tree frogs that live in hollow trunks in the forest.

While local guides in both countries attributed the songs to the bushmaster, the amphibians were almost completely unknown. To their surprise, instead of finding a snake, the field teams found two species of frogs of the genus Tepuihyla. The results are published in the open access journal ZooKeys in a collaborative effort by scientists from Catholic University of Ecuador, the Peruvian Institute of Research of the Amazon, Ecuadorian Museum of Natural Sciences, and Colorado State University, USA.

One of the tree frogs is a new species, Tepuihyla shushupe. The word shushupe is used by native people to refer to the bushmaster. The calls are highly unusual for frogs because they are a loud chuckle resembling the song of a bird. It is still unknown why locals associate the calls of the two species with the bushmaster.

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photo-3Original source:

Ron SR, Venegas PJ, Ortega-Andrade HM, Gagliardi-Urrutia G, Salerno P (2016) Systematics of Ecnomiohyla tuberculosa with the description of a new species and comments on the taxonomy of Trachycephalus typhonius (Anura, Hylidae). ZooKeys 630: 115-154. https://doi.org/10.3897/zookeys.630.9298

Scientists identify northeast Mindanao as major ‘bull’s-eye’ of biodiversity

Butuan City, Agusan del Norte. In a scientific report appearing Monday October 17th in the open access journal Zookeys, a team of researchers led by herpetologist Dr. Marites Sanguila of Father Saturnino Urios University announced that they have identified a new “epicenter” of southern Philippine biodiversity in amphibians and reptiles.

The international team of herpetologists (scientists who study frogs, lizards, snakes, turtles, and crocodiles) collaborating on the study is composed of scientists from Father Saturnino Urios University, the University of Kansas, the University of Oklahoma, the National Museum of the Philippines, Silliman University, and the Philippine Department of the Environment and Natural Resources. After an intensive five-year study, the team came to the ground-breaking conclusion that the Caraga Region of northeast Mindanao has the single highest herpetological species count of any similarly sized region in the country known to date.

Following a series of expeditions to four mountains in northern Mindanao, plus analyses of the distributions of species documented in museums around the world, the multi-institution effort culminated today with the announcement that the Caraga region is home total of 126 species of amphibians and reptiles.

According to Dr. Sanguila, this strikingly high diversity includes 40 species of frogs, one kind of caecilian (a secretive eel-like amphibian), 49 types of lizards, 35 varieties of snakes, plus one native freshwater turtle and, of course, one species of crocodile. According to the new study, the key to understanding the Caraga Region’s high biodiversity newly documented distributions of those 126 species, which overlap in northeast Mindanao. At a boundary between mainland Mindanao Island and the eastern Visayas (Samar, Leyte, Bohol, Dinagat, Siargao islands), “the Caraga region is an area where many species’ distributions come together and overlap, making this spot a kind of central hub of biodiversity,” said Dr. Sanguila. The new study brings more good news from the Philippines, a country internationally recognized as global biodiversity conservation hotspot of biodiversity.124836_web

“International collaborative biodiversity inventories are a great way to promote student training and faculty research development,” said Dr. Marites Sanguila, “In this research, we followed the example of the life-long collaboration between Dr. Angel Alcala, from the Silliman University, and Dr. Walter Brown, from California Academy of Sciences, and invited our U.S. counterparts to join in the effort to synthesize information on Caraga amphibian and reptile biodiversity. The results have unfolded in ways we could not have predicted, and generated opportunities for students on both sides of the Pacific.”

“Dr. Sanguila’s research tells us in a very special way something we have known intuitively for years, but have been unable to articulate: there is something very special about the unique biodiversity of the Caraga region! At the ‘center of the center’ of southern Philippine biodiversity, our small corner of Mindanao is undoubtedly unique, in need of conservation, and worthy of intensive scientific study,” said Rev. Fr. John Christian Young, president of Father Saturnino Urios University.

 

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Original Source:

Sanguila MB, Cobb KA, Siler CD, Diesmos AC, Alcala AC, Brown RM (2016) The amphibians and reptiles of Mindanao Island, southern Philippines, II: the herpetofauna of northeast Mindanao and adjacent islands. ZooKeys 624: 1-132. doi: 10.3897/zookeys.624.9814

In the belly of the Devil: New rare ant species found in the stomach of a poison frog

While new ant species are usually discovered in surveys involving researchers searching through leaf litter, it turns out that sifting through the stomach contents of insect-eating frogs might prove no less effective, especially when it comes to rare species. Such is the case of a new species of rarely collected long-toothed ant, discovered in the belly of a Little Devil poison frog in Ecuador.

The international team of Drs Christian Rabeling and Jeffrey Sosa-Calvo, both affiliated with University of Rochester, USA, Lauren A. O’Connell, Harvard University, USA, Luis A. Coloma, Fundación Otonga and Universidad Regional Amazónica Ikiam, Ecuador, and Fernando Fernández, Universidad Nacional de Colombia, have their study published in the open access journal ZooKeys.

The new ant species, named Lenomyrmex hoelldobleri after renowned myrmecologist Bert Hölldobler on the occasion of his 80th birthday, was described based on a single individual – a female worker, recovered from a Little Devil poison frog. It is the seventh known species in this rarely collected Neotropical genus.  

Similarly to its relatives within the group, this ant amazes with its slender and elongate mouthpart, yet it is larger than all of them. The remarkable jaws speak of specialised predatory habits, however, so far, nothing is known about these ants’ feeding behavior.in-full-face

The amphibian, whose diet majorly consists of ants, was collected from the Ecuadorian region Choco, which, unfortunately, despite being one of the most biologically diverse areas in the world with exceptionally high levels of endemism, is also one of Earth’s most threatened areas.

In conclusion, the authors point out that “studying vertebrate stomach contents is not only a way of studying the trophic ecology” (meaning the feeding relationships between organisms), “but also an interesting source of cryptic and new arthropod species, including ants.”

Furthermore, the scientists note that nowadays there is no need to kill a frog, in order to study its stomach. “Stomach flushing methods have been developed and successfully applied in numerous studies, which avoids killing individuals.”

 

Original source:

Rabeling C, Sosa-Calvo J, O’Connell LA, Coloma LA, Fernández F (2016) Lenomyrmex hoelldobleri: a new ant species discovered in the stomach of the dendrobatid poison frog, Oophaga sylvatica (Funkhouser). ZooKeys 618: 79-95. doi: 10.3897/zookeys.618.9692

Greek heroic deity Prometheus now has a namesake in a new tiny rain frog from Ecuador

Prometheus, the mythological Greek heroic deity, has been given a namesake in a new species of tiny rain frog, discovered in southwestern Ecuador. The name was chosen by the international team of scientists, led by Dr Paul Szekely, Ovidius University, Constanta, Romania, in acknowledgement of the Prometeo program, funded by the Ecuadorian government.

The description of this new species (Pristimantis prometeii) is the result of the cooperation between three Romanian Prometeo investigators affiliated with the Universidad Tecnica Particular de Loja and Universidad Nacional de Loja, and two Ecuadorian specialists from Pontificia Universidad Catolica del Ecuador. The full study is available from the open access ZooKeys.

During the day, frogs of the new species were found hiding in flowering plants, while at night — perching on leaves at low heights in well preserved cloud forests. They grow to 2-3 cm with the females being larger than the males.9121_Adult female in life img 2

The newly described species is part of a group of frogs called Terrarana (meaning ‘Land or terrestrial frogs’). This is a lineage of frogs that has evolved directly developing eggs, which are deposited in terrestrial habitats. Unlike other frogs, these ones do not have an aquatic tadpole stage and the embryos develop directly into froglets on land.

The newly described species is only known from Reserva Biologica Buenaventura, southwestern Ecuador, at elevations between 878 and 1082 m. This reserve is privately owned by the Jocotoco Conservation Foundation. The reserve has at least another four endemic species of amphibians, reptiles, and birds.

With more than 470 species, the directly developing rain frogs of the genus Pristimantiscontinue to surprise everyone.

“While new species are described every year, there are over a hundred discovered over the last decade only,” remind the authors.

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extraOriginal source:

Szekely P, Cogalniceanu D, Szekely D, Paez N, Ron SR (2016) A new species of Pristimantis from southern Ecuador (Anura, Craugastoridae). ZooKeys 606: 77-97. doi: 10.3897/zookeys.606.9121

Singing in the rain: A new species of rain frog from Manu National Park, Amazonian Peru

A new rain frog species has been described from Amazonian Peru and the Amazonian foothills of the Andes. The frog, given the name Pristimantis pluvialis, was found by researchers from Southern Illinois University Carbondale, the University of Michigan, and the National University of San Antonio Abad of Cusco in Peru. The discovery is published in the open access journal ZooKeys.

Several individuals of P. pluvialis were found during nocturnal surveys near Manu National Park, a region recognized as having the highest diversity of reptiles and amphibians of any protected area.

The species has also been collected within the private conservation area Bosque Nublado, owned by the Peruvian NGO Perú Verde, and within the Huachiperi Haramba Queros Conservation Concession, the first such type of concession granted to a native community in Peru.

The new species is likely found within the park as well, bringing the number of known amphibian species in this area to 156. Similarly to other species within its genus, which is among the largest vertebrate genera, the new rain frog exhibits direct development. This means that it is capable of undergoing its entire life cycle without a free-living tadpole stage.

It can be distinguished from other members of its genus by call, skin texture, and the presence of a rostral papilla. It was given the name “pluvialis”, translatable to “rainy” from Latin, to denote the incredibly rain-soaked habitat it lives in (>8 meters of rain yearly), and because it was found calling only after heavy rains.

Unfortunately, when a fungal disease, known as the amphibian chytrid fungus, arrived in the area back in the early 2000s, many frog species in and around the region began to decline. Out of the studied ten individuals of the presently described new species, four were found to be infected. However, the impact of the disease on these particular rain frogs is still unknown, and their numbers do not seem to have decreased.Image 3

“This discovery highlights the need for increased study throughout the tropics, for example Manu NP and its surrounding areas have been well studied, but despite these efforts, new species are being continuously discovered,” points out first author Alex Shepack, a PhD student in the laboratory of co-author Dr Alessandro Catenazzi at Southern Illinois University.

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Original source:

Shepack A, von May R, Ttito A, Catenazzi A (2016) A new species of Pristimantis (Amphibia, Anura, Craugastoridae) from the foothills of the Andes in Manu National Park, southeastern Peru. ZooKeys 594: 143-164. doi: 10.3897/zookeys.594.8295