The Swiss paper wasp, a new species of social Hymenoptera in Central Europe

Swiss scientists have discovered a new species of aculeate wasp, not hidden somewhere in a jungle on a remote continent, but in Central Europe, in a swampy area just a few kilometers from Zurich. The new species named “Polistes helveticus“, or the Swiss paper wasp, was described in the open access journal ZooKeys.

Paradoxically, this species has been long known in Central Europe but was confused for decades with a closely related species native to southern Europe. Only after the latter expanded its range to northern Switzerland, possibly following climate change, was the confusion noticed.

Six years ago, the Swiss entomologist Rainer Neumeyer discovered a population of paper wasps in wetlands close to Zurich. He thought that he had found P. bischoffi, a well-known species in Switzerland. But at the same location he found another species which, although quite similar to the first, was unusual and could not be identified with certainty.

Neumeyer contacted scientists Gaston-Denis Guex from the University of Zurich, Hannes Baur from the Natural History Museum of Bern, and Christophe Praz from the University of Neuchatel. Together, the scientists used genetic and morphometric analyses to confirm the coexistence of two distinct species at the locality close to Zurich: a northern taxon, already well known from Central Europe, and a little known taxon mostly distributed in southern Europe.

After reviewing the information on the type material, the scientists came to the conclusion that the southern species was P. bischoffi, described 80 years ago from Sardinia. The northern species was given the name P. helveticus.

The southern species was not present in northern Switzerland until recently. The confusion was only revealed after P. bischoffi expanded its distribution northwards, possibly following climate change.

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Original Source:

Neumeyer R, Baur H, Guex G-D, Praz C (2014) A new species of the paper wasp genus Polistes (Hymenoptera, Vespidae, Polistinae) in Europe revealed by morphometrics and molecular analyses. ZooKeys 400: 67–118. doi: 10.3897/zookeys.400.6611

Putting the endoparasitic plants Apodanthaceae on the map

The Apodanthaceae are small parasitic plants living almost entirely inside other plants. They occur in Africa, Iran, Australia, and the New World. Bellot and Renner propose the first revision of the species relationships in the family based on combined molecular and anatomical data. They show that Apodanthaceae comprise 10 species, which are specialized to parasitize either legumes or species in the willow family.

Few plants are obligate parasites, and fewer still are endo-parasites, meaning they live entirely within their host, emerging only to flower and fruit. Naturally, these plants are rarely collected, and their ecology, evolution, and taxonomy are therefore poorly understood. Perhaps the weirdest of these families is the Apodanthaceae, which Bellot and Renner now deal with in a paper in PhytoKeys.

Based on most material available of this family, they conclude that it has 10 species occurring in Australia, Africa, Iran, California, Central America and South America. Because the environment that matters to Apodanthaceae is the host, not anything outside it, these plants occur from the lowlands to 2500 m altitude and from deserts to Amazonian forest.

Bellot and Renner obtained DNA data to investigate species limits, and they also provide a key to all species, many illustrations, and a distribution map. The work will thus literally help putting Apodanthaceae on the map.

“I am currently assembling the plastid genome of the African species,” says Bellot, “and am already seeing that it is extremely reduced, fitting with the special lifestyle of Apodanthaceae.”

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Original source:

Bellot S, Renner SS (2014) The systematics of the worldwide endoparasite family Apodanthaceae (Cucurbitales), with a key, a map, and color photos of most species. PhytoKeys 36: 41. doi: 10.3897/phytokeys.36.7385

 

Additional information:

Funding came from the German Research Foundation (DFG RE 603/16-1).

 

Two new species of yellow-shouldered bats endemic to the Neotropics

Lying forgotten in museum collections two new species of yellow-shouldered bats have been unearthed by scientists at the American Museum of New York and The Field Museum of Natural History and described in the open access journal ZooKeys. These two new additions to the genus Sturnira are part of a recent discovery of three bats hidden away in collections around the world, the third one still waiting to be officially announced.

Up until recently the genus Sturnira was believed to contain only 14 species. In the last years closer morphological and molecular analysis have revealed an unexpected species richness in the genus. Sturnira now includes 22 described species, making it the most speciose genus in the Neotropical bat family Phyllostomidae.

Phyllostomidae, or the New World leaf-nosed bats are exclusively found in the biodiversity rich tropical areas of Central and South America. Both the scientific and common names of these bats refer to their often large, lance-shaped noseleaves. Because these bats use echolocation to orientate in the darkness the “nose-leaf” is thought to serve some role in fine-tuning their call.

All species in the yellow-shouldered genus Sturnira are frugivorous which means they feed largely on fruit. They are endemic to the Neotropics where they inhabit tropical lowland and montane forests. In fact the greatest diversity in the genus occurs on the elevated forested slopes of the Andes where at least 11 species occur.

The two newly described species, Sturnira bakeri and Sturnira burtonlimi occur in western Ecuador and in Costa Rica and Panama. The reason why they went unrecognized in collections is a superficial resemblance with other species in the genus, most of which were described without adequate illustrations to communicate identifying characteristics. Only after an in-depth molecular analysis that included over 100 samples from most of the species of the genus could the new species be identified. “Modern electronic publications likeZooKeys permit extensive and detailed color photography to accompany taxonomic descriptions. Any reader can easily and clearly appreciate the character states we use to distinguish these new taxa.” said co-author Bruce Patterson.

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Original Source:

Velazco PM, Patterson BD (2014) Two new species of yellow-shouldered bats, genus Sturnira Gray, 1842 (Chiroptera, Phyllostomidae) from Costa Rica, Panama and western Ecuador. ZooKeys 402: 43-66. doi:10.3897/zookeys.402.7228

Name of new weakly electric fish species reflects hope for peace in Central Africa

Two new species of weakly electric fishes from the Congo River basin are described in the open access journal ZooKeys. One of them, known from only a single specimen, is named “Petrocephalus boboto.” “Boboto” is the word for peace in the Lingala language, the lingua franca of the Congo River, reflecting the authors’ hope for peace in troubled Central Africa.

On a 2010 field trip to the Congo River of Democratic Republic of the Congo, in the riverside village of Yangambi-Lokélé, French ichthyologist Sébastien Lavoué of the Taiwan Institute of Oceanography and American ichthyologist John Sullivan of Cornell University, both specialists on mormyrid weakly electric fishes, captured a single individual of the genus Petrocephalus not quite like any they had seen before.

“Sébastien has the best eye of anyone in the world for Petrocephalus,” says Sullivan. “So when he wasn’t certain what species it belonged to, we flagged it as one to look at carefully once we got home.”

As they had for hundreds of other Petrocephalus specimens collected in Central Africa since the late 1990s, they placed the small, silvery fish in a small basin with water from the river and recorded its electric organ discharge (EOD) with an oscilloscope, humanely euthanized it, took its photo, procured a tissue sample for DNA analysis, tagged it and preserved it in formaldehyde.

Petrocephalus are African weakly fishes of the family Mormyridae that produce pulses of only a few hundred millivolts from an organ made of modified muscle cells in front of their tail. Receptor cells on the fishes’ skin detect distortions to the electric field created by nearby objects in the water. In this way, they are able to “electrolocate” through their complex aquatic environment at night. Their short electric pulses, too weak to be sensed by touch, are also used to communicate the sender’s species identity and gender to other electric fishes.

When Sullivan and other researchers collect live electric fishes in the field, they routinely make recordings of these EODs, one fish at a time, so that they can later study and compare the signals and the specimens’ anatomy together. Specimens are individually tagged so that after they are preserved and placed in a museum collection, they remain linked to their EOD recordings. “With these electric fishes it is often difficult to determine where the species boundaries are, and we need all the available evidence, from morphology, DNA and their EODs to figure it out.” Sullivan said. In this case, the EOD of this odd Petrocephalus was not so different from EODs of other species in this genus.

Back in his laboratory in Taipei, Lavoué sequenced the gene cytochrome b from the specimen. Comparing this sequence to those from other Petrocephalus specimens and careful examination of the fish’s morphology and its EOD helped him determine that it belonged to an undescribed species. “Describing a new species from a single specimen is far from ideal,” Lavoué said, “but in this case it seemed the best thing to do. In the places we’ve sampled, it’s obviously very rare. Since we haven’t yet found any locality where it’s common, it’s unlikely we’ll find such a locality anytime soon.”

Lavoué and Sullivan named this species “Petrocephalus boboto“: the word “boboto” means peace and fellowship in the Lingala language spoken along the Congo River.

“We named this hard-to-find Petrocephalus species “boboto” in the hopes that solutions for peace—though elusive like this fish—can be found in eastern D.R. Congo and the other troubled areas of Central Africa,” said Sullivan.

The other new Petrocephalus species Lavoué and Sullivan describe in the paper, also from the Congo River basin, is named for their colleague, Matthew Arnegard, another mormyrid researcher currently working at the Fred Hutchinson Cancer Research Center in Seattle, Washington.

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Funding:

The research that led to the discovery of the two new Petrocehpalus species was funded by the National Science Foundation of the United States and a Fulbright scholarship to J. Sullivan.

 

Original Source:

Lavoué S, Sullivan JP (2014) Petrocephalus boboto and Petrocephalus arnegardi, two new species of African electric fish (Osteoglossomorpha, Mormyridae) from the Congo River basin. ZooKeys 400: 43. doi: 10.3897/zookeys.400.6743

 

Contact:

John P. Sullivan,
PhD., Collection Archivist & Curatorial Affiliate,
Cornell University Museum of Vertebrates,
159 Sapsucker Woods Rd, Ithaca, NY 14850.
Email: jpsullivan@cornell.edu
Tel: 607.342.2234

Counting the invisible by sound — a new approach to estimate seabird populations

Seabirds nest in places that are inaccessible for most humans – vertical cliffs and remote islands surrounded by raging waves. Worse still, many seabirds lay their eggs in burrows or cavities where they are protected from inclement weather and invisible for researchers. Hidden under rocks or in burrows during the day, and flying around only during dark nights – counting these birds is a researcher’s nightmare.

Despite their cryptic behaviour, the seabirds are ill-prepared to fend off furry invaders. Humans have brought cats and rats to many islands around the world, where the cats and rats roam freely and kill seabirds. Especially those seabirds that nest in burrows are often unable to escape, and many species have disappeared from islands where cats or rats have been introduced.

Although researchers have known for decades that many seabirds are in trouble, it is surprisingly hard to put a number on how fast populations decline. “Those species that are most vulnerable to rats are often the ones that are the most difficult to count.” says Steffen Oppel, a Conservation Scientist with the RSPB who recently tested a new approach to count the invisible birds with colleagues from SPEA in Portugal.

Seabirds that nest underground may be all but invisible in their breeding colonies, but they are very noisy at night. And the more birds there are, the louder a colony is. Oppel and his colleagues set up sound recorders on a remote island in the North Atlantic for two years to ‘count’ the number of nesting birds by recording their calls at night. They painstakingly counted every nest near the recorders to test whether larger colonies do in fact make more noise. The study was published in the open access journal Nature Conservation.

“Recording seabird calls for a few months is the easy part – but making sense of 1000s of hours of sound recording is quite tricky.” says Oppel. Together with Matthew McKown, a seabird researcher who specialises on sound recordings, the team developed an algorithm that automatically counted the seabird calls in terabytes of recordings. The results conformed with expectations: places with the most nests did indeed register the highest number of calls. With that relationship established, the team then extrapolated the seabird population size for the entire island – a number that had so far been derived from wild guesses.

“Estimating exactly how many birds nest on a cliff is not very precise.” admits Oppel, but the sound recordings provide a very valuable index of how large seabird colonies are. “We can use this index over time to assess whether colonies are stable or decreasing – which is extremely important for many remote colonies about which we know very little.”

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Original source:

Oppel S, Hervías S, Oliveira N, Pipa T, Silva C, Geraldes P, Goh M, Immler E, McKown M (2014) Estimating population size of a nocturnal burrow-nesting seabird using acoustic monitoring and habitat mapping. Nature Conservation 7: 1–13. doi: 10.3897/natureconservation.7.6890

A new tiny species of crayfish from the swamps of coastal eastern Australia

Hidden in one of Australia’s most developed and fastest growing areas lives one of the world’s smallest freshwater crayfish species. Robert B McCormack the Team Leader for the Australian Crayfish Project described the new species belonging to the genus Gramastacus, after 8 years of research in the swamps and creeks of coastal New South Wales, Australia. The study was published in the open access journal ZooKeys.

Being a small crayfish species it has remained undescribed and undiscovered in one of the fastest developing regions of Australia. Only one other species of Gramastacus crayfish is known and it occurs some 900 km away in the Grampians region of Victoria.

This new species is found in lowland ephemeral habitats surrounding coastal lakes and lagoons from Wamberal Lagoon, north along the coastal strip to Wallis Lake. Being dependent on regular natural flooding and drying cycles, only lowland, swampy areas are suitable for this tiny crayfish.

Each crayfish digs a small rounded cross-section burrow up to one metre deep into the water table to survive the drying cycle. Some areas are riddled with these small burrows as they are a very prolific species and can occur in very high numbers in small habitat areas.

The newly described crayfish are found in one of Australia’s most developed regions. Unfortunately, this means that much of their habitat has been lost in the past as these ephemeral areas are the first to be drained or reclaimed to make way for agriculture, industry, housing developments, golf courses, infrastructure, etc. Now, being found and officially described, this crayfish must be considered in any further developments and hopefully future habitat loss will be reduced.

The scattered populations of Gramastacus seem highly fragmented and many are increasingly threatened by a range of risks other than human development. Invasive crayfish, pest fish species like plague minnows and swordtails, rising sea levels and falling water tables all are increasing dangers. Luckily, the large number of National Parks and Reserves along the coastal strip provides safe refuges for some populations.

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About the Australian Crayfish Project:

Started in 2005 The Australian Crayfish Project (ACP) is a privately funded project run entirely by volunteers with a thirst for knowledge on freshwater crayfish. We all freely give our time and resources to support the project. Australia has one of the most diverse ranges of freshwater crayfish species on the planet yet there is much confusion with the description and distribution of species and many species remain undiscovered & undescribed. The project is discovering dozens of new species or new populations that were never known to exist. Our aim is to find and identify all species of freshwater crayfish and their habitat, the bottom line is that if no one knows these species are there and they don’t even have a name then how can they be protected and conserved into the future? The results of this project will ensure that all management agencies take the crayfish and their habitat into consideration for future environmental, conservation, development and catchment management plans, etc.

Australia’s unique crayfish are important keystone species which have a disproportionately large effect on the whole catchment relative to their abundance. They play a critical role in maintaining the structure of the whole ecological community, their prosperity and abundance directly affecting the health and abundance of numerous other species (macroinvertebrates, fish, eels, turtles, lizards, snakes, water rats, platypus and birds, to name just a few) in the ecosystem. Please help us help the crayfish and all the other species that rely on them for their survival. http://www.aabio.com.au/the-australian-crayfish-project/

 

Original source:

McCormack RB (2014) The eastern swamp crayfish Gramastacus lacus sp. n. (Decapoda, Parastacidae) a new species of freshwater crayfish from coastal New South Wales, Australia. ZooKeys 398: 53–67. doi: 10.3897/zookeys.398.7544

 

Additional Information:

Article under conservation and research Australian Aquatic Biological.


For a full list of all Australian Freshwater Crayfish:

http://www.aabio.com.au/crayfish-list/

A new species of horseshoe worm discovered in Japan after a 62 year gap

The horseshoe worm is a worm-like marine invertebrate inhabiting both hard and soft substrates such as rock, bivalve shells, and sandy bottom. The name “horseshoe” refers to the U-shaped crown of tentacles which is called “lophophore.” Horseshoe worms comprise a small phylum Phoronida, which contains only ten species decorating the bottom of the oceans.

The new species Phoronis emigi, the eleventh member of the group described in the open access journal ZooKeys, comes after a long 62 year gap of new discoveries in the phylum. It is unique in the number and arrangement of body-wall muscle bundles and the position of the nephridia which is the excretory organ of some invertebrates. The new species is morphologically similar to sand-dwelling species Phoronis psammophila and it is also closely related to Phoronis hippocrepia, which inhabits hard substrate.

The morphology of the topotypes for Phoronis ijimai is also described in this study after 117 years since its original description. The combination of a detailed observation of the internal morphologies and the molecular phylogenetic analyses including the topotypes ensure a synonymy between P. ijimai and the northeastern pacific species Phoronis vancouverensis that has long been disputed.

“It is necessary to use both internal anatomy and molecular data for reveal the global diversity of horseshoe worm. The known phoronid diversity still remains low, with all specimens reported from limited habitats and the localities by the limited reports. Investigations at new localities or habitats may yield additional species in the future.”, explains Dr Masato Hirose, Atmosphere and Ocean Research Institute, The University of Tokyo, Japan.

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Original source:

Hirose M, Fukiage R, Katoh T, Kajihara H (2014) Description and molecular phylogeny of a new species of Phoronis (Phoronida) from Japan, with a redescription of topotypes of P. ijimai Oka, 1897. ZooKeys 398: 1. doi: 10.3897/zookeys.398.5176

Brand new old journal

Zoosystematics and Evolution moves to Pensoft Publishers to combine tradition and innovation in a new publishing format

Established in 1898 as Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische Reihe by the Museum für Naturkunde Berlin, Zoosystematics and Evolution is one of the oldest European zoological journals. The journal joined Pensoft Publishers Open Access family earlier this year and the first issue using the benefits of the new publishing platform is now a fact.

Marking the new collaboration and combining tradition and innovation, this first issue for 2014, is now available in a novel, attractive open access, semantically enriched format, while keeping at the same time the traditional PDF and print versions.

“We are excited to welcome Zoosystematics and Evolution in Pensoft. The attractive combination between historical legacy and tradition and the advanced publishing technologies of Pensoft ensures for this authoritative journal a jump right into the future!” said the Managing Director of Pensoft Publishers, Prof. Lyubomir Penev.

Being already a member of Pensoft’s family of advanced open access journals, Zoosystematics and Evolution now benefits from the innovative “platinum “open access model. For Pensoft, “platinum” open access means not just that the articles and all associated materials are free to download and that there are no author-side fees but even more so that novel approaches are used in the dissemination and reuse of published content. This publishing model includes:

  • Free to read, reuse, revise, remix, redistribute
  • Easy to discover and harvest by both humans and computers
  • Content automatically harvested by aggregators
  • Data and narrative (text) integrated to the widest extent possible
  • Community peer-review and rapid publication
  • Post-publication peer-review
  • Easy and efficient communication with authors and reviewers
  • Active promotion and dissemination of scientists’ work in the society
  • A “fremium” model assuming no author-side fees within an yearly limit of number of pages

“Making published data available online is good but not sufficient. The ultimate goal is to make the journal content more attractive for the readers and authors, so that to increase visibility and re-usability of the published information to generate new knowledge”, said Dr Matthias Glaubrecht from the Museum für Naturkunde, Editor-in-Chief of Zoosystematics and Evolution.

Dr. Gregor Hagedorn, Head of the Museum’s science division “Digital World and Information Science”, adds: “Original scientific articles and the underlying scientific data have to be published in new, connected ways if we want to be able to face the challenges to mitigate global biodiversity loss. By making this journal accessible under Open Access and by joining our efforts in developing the Museum’s digital infrastructures with the novel publishing technologies invented by Pensoft, we make a significant step towards these goals.”

“I am glad to see Zoosystematics and Evolution, originally established more than a Century ago as a journal to publish studies on Museum’s collections, now serving the great task to open the treasury of biodiversity data from these collections gathered by generations of naturalists with the strong support from society over 250 years of history of explorations!” concluded Prof. Johannes Vogel, Director General of the Museum für Naturkunde, Berlin.

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Additional Information:

Museum für Naturkunde – Leibniz Institute for Research on Evolution and Biodiversity is a research museum within the Leibniz Association. It is one of the most significant research institutions worldwide in biological and geo-scientific research on evolution and biodiversity. Established in 1810, its collections comprise more than 30 million items relating to zoology, palaeontology, geology and mineralogy, which are highly significant for science as well as for the history of science and at the centre of the research performed at the Museum.

Pensoft Publishers specialize in academic and professional book and journal publishing, mostly in the field of biodiversity science and natural history. In 2008, Pensoft launched its first open-access journals named ZooKeys and BioRisk, and since then, it has taken a leadership in introducing innovations in the field of open access. On 16th of September 2013, Pensoft launched the Biodiversity Data Journal (BDJ) and the associated Pensoft Writing Tool (PWT), as the first workflow ever to put together article authoring, community peer-review, publishing and dissemination within a single online collaborative platform.

Killing a name of an extinct sea cow species

Sirenians, or sea cows, are a particular group of mammals that superficially resembles whales in having, amongst other features, a streamlined-body and horizontal tail fluke. Though belonging to the so-called marine mammals, such as whales and seals, sea cows are members of a group having a single origin that includes their closest living relatives, the proboscideans (or elephants in the broader sense).

Today, sirenians are known by only four species, but their fossil record is much more diverse documenting the transition from land-dwelling animals to fully aquatic ones. This makes fossil representatives of this group not only very fascinating, but also crucial for the understanding of the transition from life on land to the sea and the past diversity of the order Sirenia.

Considering the oldest known member of this order, Prorastomus from the Eocene (~50 Ma – Millions of years before present) of Jamaica, first fully aquatic sea cows already occurred by approximately 42 Ma, which indicates a relatively fast transitional process. Later forms still retained a small femur, which however was no more visible outside. Amongst such sirenians, one species was distributed in a shallow marine area, now occupied by Germany and Belgium, about 30 Millions of years ago in the early Oligocene. This respective species was hitherto known as Halitherium schinzii and believed to represent the only sea cow that evolved in that region to that time.

In a recent publication of the open access journal Zoosystematics and Evolution, Manja Voss from the Museum für Naturkunde Berlin deals with a contrasting hypothesis of two distinct species in the early Oligocene of Central Europe. Considering the always incomplete fossil record, palaeontologists face the challenge to determine species on a material basis being as informative as possible. In the present study Manja Voss provides arguments why this is not the case with the so-called holotype, a single tooth, of H. schinzii and explains why this species name is not applicable to any currently known sirenian.

Additionally, Manja Voss emphasises that consequences are to be drawn including a morphological re-assessment of skeletal material originally assigned to this species and the requirement of new generic names for all species formerly grouped under Halitherium. Moreover, it is acknowledged that this genus does not only include unrelated species, but is also the name giving basis for a subfamily, the Halitheriinae, that is likewise far from forming a single entity. Therefore, the proposed rejection of these specific termini shall help to better handle the known diversity of the sirenian order.

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Original Source:

Voss M (2014) On the invalidity of Halitherium schinzii Kaup, 1838 (Mammalia, Sirenia), with comments on systematic consequences. Zoosystematics and Evolution 90(1): 87-93. DOI: 10.3897/zse.90.7421

One or 2? How to decide how many species you have got

It is often difficult to decide whether two animals belong to the same or two distinct species. This can be especially challenging for animals which externally look very similar. In a recent study, published in the open access journal Zoosystematics and Evolution, scientists from the Museum für Naturkunde Berlin use genetic data and sound analysis to test if treefrogs from West and Central Africa belong to different or the same species.

Due to the fact that, when external characters are used, only size is useful to distinguish these frogs the scientists employed additional characters to determine species affiliations. One the one hand, they used genetics, and, on the other, advertisement calls of different populations. Male frogs have to attract their females via species specific calls, so call characteristics (e.g. duration, frequency) reliably tell whether animals belong to one or several species.

The two sets of populations have been previously declared as belonging to the same species based on morphological, meaning external, similarities. With their work the researchers from Berlin proved their expectations right, and revealing they were in fact working with four and not two species: two different sets of one large and one small specie, which live in West and Central Africa respectively.

In order to facilitate other researchers the verification of these results, not only genetic but also acoustic data are publicly available and the latter can be downloaded from the animal sound archive of the Museum für Naturkunde.

Species are the basis for all biological questions. Therefore, they are not only important for taxonomists but also a reference parameter for physiologists, ecologists or conservationists. Thus it is of central importance to be able to distinguish whether animals of two different populations belong to one or two distinct species. A clear showcase is an example, which was examined in a recent publication by zoologist from Berlin and Geneva.

“A colleague grouped animals to two species which were formerly known as four species, each now with a distribution across West and Central Africa. However, this contradicted known zoogeographical facts which found that forest frogs live either in West or in Central Africa but very rarely simultaneously in both regions. The clarification of that question is for example important to understand how the landscapes in these regions have developed over the past millions of years.” explains Dr. Rödel, one of the authors if the recent study.

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Original Source:

Roedel M, Emmrich M, Penner J, Schmitz A, Barej M (2014) The taxonomic status of two West African Leptopelis species: L. macrotis Schiøtz, 1967 and L. spiritusnoctis Rödel, 2007 (Amphibia: Anura: Arthroleptidae). Zoosystematics and Evolution 90(1): 21-31. DOI: 10.3897/zse.90.7120