Tag: biodiversity

The ‘Star dust’ wasp is a new extinct species named after David Bowie’s alter ego

During her study on fossil insects of the order Hymenoptera at China’s Capitol Normal University, student Longfeng Li visited the Smithsonian National Museum of Natural History, Washington, carrying two unidentified wasp specimens that were exceptionally well-preserved in Burmese amber. This type of fossilized tree resin is known for the quality of the fossil specimens which can be preserved inside it. Being 100 million years old, they provide an incredible view into the past.

The subsequent analysis of the specimens revealed that both represent species new to science. Furthermore, one of the wasps showed such amazing similarities to a modern group of wasps that it was placed in a currently existing genus, Archaeoteleiawhich has long been considered as an ancient lineage. The species are described in a study published in the open access Journal of Hymenoptera Research.

However, Archaeoteleia has changed since the times when the ancient wasp got stuck on fresh tree resin. The authors note that “a novice might not recognize the characters that unite the fossil with extant species”. For instance, the modern wasp species of the genus show visibly longer antennal segments and a different number of teeth on the mandible when compared to the fossil. In turn, the description of the new extinct species enhances the knowledge about living species by highlighting anatomical structures shared by all species within the genus.

This fossil wasp with living relatives received quite a curious name, Archaeoteleia astropulvis. The species name, astropulvis, translates from Latin to ‘star dust’. The discoverers chose the name to refer to both “the ancient source of the atoms that form our planet and its inhabitants”, as well as to commemorate the late David Bowie’s alter ego – Ziggy Stardust.

Unlike the Star dust wasp, the second new species belongs to a genus (Proteroscelio) known exclusively from Cretaceous fossils. Likewise, it is a tiny insect, measuring less than 2mm in length. It also plays an important role in taxonomy by expanding the anatomical diversity known from this extinct genus.

The authors conclude that their discovery, especially the Star dust wasp and its placement in an extant genus, where it is the only fossil species, “exemplifies the importance of understanding the extant fauna of a taxon to interpret fossils”.

“Such union of fossil and extant morphologies is especially illuminating and requires examination of both kinds of specimens,” they add.

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Original source:

Talamas EJ, Johnson NF, Buffington ML, Dong R (2016) Archaeoteleia Masner in the Cretaceous and a new species of Proteroscelio Brues (Hymenoptera, Platygastroidea). In: Talamas EJ, Buffington ML (Eds) Advances in the Systematics of Platygastroidea. Journal of Hymenoptera Research 56: 241-261. https://doi.org/10.3897/jhr.56.10388

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.

However, 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

Three new mini thorn snails described from Georgia (USA), Belize and Panama

Although computer tomography (CT) is widely used in medicine, its application in micro snail identification is still at the pioneering stage.

However, Dr Adrienne Jochum from the Naturhistorisches Museum der Burgergemeinde Bern (NMBE), Switzerland and her interdisciplinary team of German and Swiss scientists (Dr. Alexander M. Weigand, University of Duisburg-Essen, Estee Bochud and Thomas Inäbnit, NMBE and the University of Bern, Dorian D. Dörge, Goethe University, Frankfurt, Dr. Bernhard Ruthensteiner, Zoologische Staatssammlung Muenchen, Dr. Adrien Favre, Leipzig University, Gunhild Martels and Dr. Marian Kampschulte, Justus-Liebig University Giessen) have recently applied it in their research, now published in the journal ZooKeys.

As a result of their revolutionary approach, the scientists report three new thorn snail species – tiny, colourless and highly fragile creatures that measure less than 2 mm and belong to the genus Carychium.

Much like X-rays showing the degree of damage in broken bones, CT scans provide access to snail shells. Differences, such as the degree of sinuosity of the potato chip-like wedge (lamella), elegantly gliding along the spindle-like columella, become visible. These structures provide stability and surface area to exert muscular traction while manoeuvring the unwieldy shell into tight cavities. The alignment and degree of undulation of the lamella on the columella is also used by malacologists (mollusc specialists) to identify different thorn snail species.

Conventionally, examination of this signatory character requires cutting a hole in the shell with a fine needle under the microscope. This tedious method requires a much patience and dexterity and, all too often, the shell cracks open or disintegrates into dust under pressure. By exposing the delicate lamella to non-manipulative CT scans, Dr. Jochum and her team have found the best method to differentiate not only thorn snails but also many other micro creatures.

Together with G. Martels and Dr. M. Kampschulte, Dr. Jochum described new micro snails for the first time using CT in East Asian hypselostomatid snails in 2014. The first subterranean Asian relative of the thorn snails (Koreozospeum nodongense), was also described by Dr. Jochum thanks to CT scans in 2015.

The scientists studied and compared thorn snails collected from Mexico, Florida (USA) and Costa Rica.

Curiously, the new species Carychium hardiei was discovered by accident by Dr. Jochum en route to the Atlanta Airport during a rest stop in Georgia (USA). The snail is named after the American naturalist and field biologist Frank Hardie. Another species, Carychium belizeense, was found in the Bladen Nature Reserve in Belize and is named after its country of origin. The third, Carychium zarzaae from Panama, is named after Eugenia Zarza, collector of material for this study, including this species.

In total, there are fourteen species of thorn snails known in North and Central America. Their distribution ranges from as far north as northern Ontario, Canada through North America (including Bermuda and Jamaica) and south through Central America to Costa Rica. Thorn snails also live as far north as northern Sweden and as far south as sub-equatorial Java. Worldwide, this genus spans the Nearctic, Palearctic and Indomalayan biogeographic realms.

Thorn snails live in tropical and temperate forests, meadows and riparian zones, where they comprise the decomposer community in leaf litter of ecologically stable environments.

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Original Source:

Jochum A, Weigand AM, Bochud E, Inäbnit T, Dörge DD, Ruthensteiner B, Favre A, Martels G, Kampschulte M (2017) Three new species of Carychium O.F. Müller, 1773 from the Southeastern USA, Belize and Panama are described using computer tomography (CT) (Eupulmonata, Ellobioidea, Carychiidae). ZooKeys 675: 97-127. https://doi.org/10.3897/zookeys.675.12453

Conservation and nameless earthworms: Assessors in the dark?

Species that live exclusively in a single region are at a particular risk of extinction. However, for them to be protected, thorough assessments of the environmental impacts need to be performed.

There are more than 100 earthworm species living in the soil and dead wood of KwaZulu-Natal Province, South Africa. Most of them live exclusively in small regions in the province, which makes them extremely vulnerable.

To scientists Dr Adrian J. Armstrong, Ezemvelo KZN Wildlife, and Ms Thembeka Nxele, KwaZulu-Natal Museum, the problem is twofold. Firstly, they note that the expression “out of sight, out of mind” is very suitable for the case of the endemic earthworms in South Africa. Secondly, they point out that the lack of common names for these species is a stumbling block that hinders their inclusion in conservation assessments.

As a result, the researchers try to rectify this situation by assigning standardised English names to the endemic earthworms in KwaZulu-Natal. Their article is published in the open access journal African Invertebrates.

Scientific names are often intractable to non-specialists, and the lack of common names leaves environmental assessors in the dark when they need to figure out which earthworms may occur at a development site. In the meantime, it has been found that about 50% of the native vegetation in KwaZulu-Natal has already been removed as a result of infrastructure construction and the figure is rising.

“The indigenous earthworms generally don’t survive in developed areas,” say the authors.

For instance, the informal use of an English name (green giant wrinkled earthworm) for the species Microchaetus papillatus, has facilitated the inclusion of this species in environmental impact assessments in KwaZulu-Natal.

While the green giant wrinkled earthworm does occur in a relatively large and rapidly developing area in KwaZulu-Natal, other species live in smaller areas that have been urbanised even more.

The extinction of these earthworms is not only undesirable from the point of view of biodiversity advocates – the role of this group of soil organisms is impossible to replace fully with non-native earthworms. For example, some of the large indigenous earthworms (more than 1 m in length) burrow much deeper than the non-native species, thereby enriching and aerating the soil at greater depth.

The authors are hopeful that by giving the indigenous earthworms in KwaZulu-Natal common names, the threatened and endemic species will be conserved through inclusion in environmental impact assessments. Furthermore, they believe that earthworms could draw attention to the areas where they occur whenever a choice for new protected areas is to be made.

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Original source:

Armstrong AJ, Nxele TC (2017) English names of the megadrile earthworms (Oligochaeta) of KwaZulu-Natal. African Invertebrates 58(2): 11-20. https://doi.org/10.3897/AfrInvertebr.58.13226

Dig it! Two new shrimp species found in burrows at the bottom of the Gulf of California

Although the Santa María-La Reforma lagoon complex in the Gulf of California is one of the most important areas for shrimp fishery, little is known about the crustacean species that live in the burrows dug in the bottom.

In addition to presenting two species new to science, researchers Drs. José Salgado-Barragán, Universidad Nacional Autónoma de México, Manuel Ayón-Parente and Pilar Zamora-Tavares, both affiliated with Universidad de Guadalajara, México collaborated to build on the knowledge of small shrimp species living there. The study is published in the open access journal ZooKeys.

Over the span of about two years – between 2013 and 2015, the scientists conducted series of surveys of the bottom-dwelling crustaceans in Bahía Santa María-La Reforma lagoon, located in the southwest Gulf of California. Following a thorough examination of the collected specimens, they recorded five shrimp species of three genera, inhabiting burrows dug into either mud, sand, or sandy-mud. Two of these species turned out to be previously unknown.

One of the new species is named Alpheus margaritae after Dr. Margarita Hermoso-Salazar, a caridean shrimp expert who helped the authors with the identification of the species. This new crustacean lives in the intertidal zone, where it hides in soft mud and gravel of shells and rocks. So far, it is known exclusively from the coastal lagoon Bahía Santa María-La Reforma, Sinaloa, Mexico. Among its characteristic traits are creamy-white colouration splashed with sparse olive green to light brown patches.

The second new species, Leptalpheus melendezensis, is reported to live in the fine sand of the beach. It is named after the Melendez island – the only locality the species has been identified from. Unlike the rest seven members of its genus (Leptalpheus), its major cheliped lacks adhesive disks.

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Original source:

Salgado-Barragán J, Ayón-Parente M, Zamora-Tavares P (2017) New records and description of two new species of carideans shrimps from Bahía Santa María-La Reforma lagoon, Gulf of California, Mexico (Crustacea, Caridea, Alpheidae and Processidae). ZooKeys 671: 131-153. https://doi.org/10.3897/zookeys.671.9081

New butterfly species discovered in Israel for the first time in 109 years

Vladimir Lukhtanov, entomologist and evolutionary biologist at the Zoological Institute in St. Petersburg, Russia, made a startling discovery: what people had thought was a population of a common species, turned out to be a whole new organism and, moreover – one with an interesting evolutionary history. This new species is named Acentria’s fritillary (Melitaea acentria) and was found flying right over the slopes of the popular Mount Hermon ski resort in northern Israel. It is described in the open access journal Comparative Cytogenetics.

“To me, it was a surprise that no one had already discovered it,” says Vladimir Lukhtanov.

“Thousands of people had observed and many had even photographed this beautifully coloured butterfly, yet no one recognised it as a separate species. The lepidopterists (experts in butterflies and moths) had been sure that the Hermon samples belonged to the common species called Persian fritillary (Melitaea persea), because of their similar appearance, but nobody made the effort to study their internal anatomy and DNA”.

In 2012, Vladimir Lukhtanov, together with his students, initiated an exhaustive study of Israeli butterflies using an array of modern and traditional research techniques. In 2013, Asya Novikova (until 2012, a master’s student at St. Petersburg University and, from 2013, a PhD student at the Hebrew University, Jerusalem) sampled a few fritillaries from Mt. Hermon.

It was at that time when the researchers noticed that the specimens “didn’t look right” – their genitalia appeared different from those of the typical Persian fritillary. Over the next few years, Lukhtanov and his students studied this population in-depth. They carried out sequencing DNA from the specimens and found that they had a unique molecular signature – very different from the DNA of any other fritillary.

The Acentria’s fritillary seems to be endemic in northern Israel and the neighbouring territories of Syria and Lebanon. Its evolutionary history is likely to prove interesting.

“The species is probably one of a handful of butterflies known to have arisen through hybridisation between two other species in the past,” says Lukhtanov. “This process is known to be common in plants, but scientists have only recently realised it might also be present in butterflies.”

This is the first new butterfly species discovered and described from the territory of Israel in 109 years.

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Original source:

Lukhtanov VA (2017) A new species of Melitaea from Israel, with notes on taxonomy, cytogenetics, phylogeography and interspecific hybridization in the Melitaea persea complex (Lepidoptera, Nymphalidae). Comparative Cytogenetics 11(2): 325-357. https://doi.org/10.3897/CompCytogen.v11i2.12370

The Radiohead ant: A new species of ‘silky’ ant grows fungus gardens for food

The ants of the genus Sericomyrmex – literally translated as ‘silky ants’ – belong to the fungus-farming ants, a group of ants that have figured out how to farm their own food. The silky ants are the less well-known relatives of the famous leaf-cutter ants – well-studied, photogenic model organisms that you simply cannot avoid if you take a trip to the Neotropics.

For their study, now published in ZooKeys, Ana Ješovnik and Ted R. Schultz from the Smithsonian Institution‘s Ant Lab in Washington, D.C., collected silky ants from across their entire range in Central and South America, and revised the genus based on DNA sequence data and morphology. In the end, they turned out to have discovered three new species.

One of those species, Sericomyrmex radioheadi, collected in the Venezuelan Amazon, was named after the famous British music band Radiohead.

“We wanted to honor their music” one of the authors, Ana Ješovnik, says. “But more importantly, we wanted to acknowledge the conservation efforts of the band members, especially in raising climate-change awareness. ”

Using a scanning electron microscope, the authors found that the bodies of the ants are covered with a white, crystal-like layer. Curiously, this previously unknown layer is present in female ants (both workers and queens), but is entirely absent in males. Both the chemical composition and the function of this layer are unclear.

One possibility is that the layer is microbial in origin and that it has a role in protecting the ants and their gardens from parasites. This is interesting, because most of the fungus-farming ants cultivate antibiotic-producing bacteria on their bodies to protect their gardens from microbial weeds. In the meantime, in Sericomyrmex these bacteria are absent, yet their gardens are also parasite-free. Figuring out if this crystal-like layer has a role in protecting these ants’ fungus gardens might provide clues for managing diseases in human agriculture and medicine.

At only four million years, Sericomyrmex is an evolutionary youngster, the most recently evolved genus of fungus-farming ants, and an example of rapid radiation – comparable to other fast-evolving groups, such as the freshwater fishes in Africa, or the Hawaiian fruit flies.

Rapid radiation is a process in which organisms diversify quickly into a multitude of forms, making these ants good candidates for studies into speciation and evolution. In the present article, the authors acknowledge that some of the species they describe might, in fact, be multiple species that look alike, but because the ants are in the early stages of speciation, this is hard to detect.

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Original source:

Ješovnik A, Schultz TR (2017) Revision of the fungus-farming ant genus Sericomyrmex Mayr(Hymenoptera, Formicidae, Myrmicinae). ZooKeys 670: 1-109. https://doi.org/10.3897/zookeys.670.11839

Origins of an enigmatic genus of Asian butterflies carrying mythological names decoded

A group of rare Asian butterflies which have once inspired an association with Hindu mythological creatures have been quite a chaos for the experts. In fact, their systematics turned out so confusing that in order to decode their taxonomic placement, scientists had to dig up their roots some 43 million years back.

Now, having shed new light on their ancestors, a team of researchers from the Biodiversity Institute of Ontario at University of Guelph, Agriculture and Agri-Food Canada and University of Vienna, published their findings in the open access journal Zoosystematics and Evolution.

Together, Drs. Valentina Todisco, Vazrick Nazari and Paul Hebert arrived at the conclusion that the enigmatic genus (Calinaga) originated in southeast Tibet in the Eocene as a result of the immense geological and environmental impact caused by the collision between the Indian and Asian subcontinents. However, the diversification within the lineage was far from over at that point. In the following epochs, the butterflies had to adapt to major changes when Indochina drifted away, leading to the isolation of numerous populations; and then again, when the Pleistocene climatic changes took their own toll.

To make their conclusions, the scientists studied 51 specimens collected from a wide range of localities spanning across India, South China, Laos, Vietnam, Myanmar and Thailand. For the first time for the genus, the authors conducted molecular data and combined it with an examination of both genitalia and wing patterns – distinct morphological characters in butterflies. While previous estimates had reported existence of anywhere between one and eleven species in the genus, the present study identified only four, while confirming how easy it is to mislabel samples based on earlier descriptions.

However, the researchers note that they have not sampled specimens from all species listed throughout the years under the name of the genus, so they need additional data to confirm the actual number of valid Calinaga species. The authors are to enrich this preliminary study in the near future, analysing both a larger dataset and type specimens in collaboration with the Natural History Museum of London that holds the largest Calinaga collection.

Despite being beautiful butterflies, the examined species belong to a genus whose name derives from the Hindu mythical reptilian creatures Nāga and a particular one of them – Kaliya, which is believed to live in Yamuna river, Uttar Pradesh, and is notorious for its poison. According to the Hindu myths, no sooner than Kaliya was confronted by the major deity Krishna, did it surrender.

“It seems that the modern taxonomy of Calinaga is in need of a Krishna to conquer these superfluous names and cleanse its taxonomy albeit after careful examination of the types and sequencing of additional material,” comment the authors.

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Original source:

Todisco V, Nazari V, Hebert PDN (2017) Preliminary molecular phylogeny and biogeography of the monobasic subfamily Calinaginae (Lepidoptera, Nymphalidae). Zoosystematics and Evolution 93(2): 255-264. https://doi.org/10.3897/zse.93.10744

Of Star Trek, Mark Twain and helmets: 15 new species of wasps with curious names

A total of fifteen new species of parasitic wasps have been described from across the Neotropical region. Apart from belonging to a peculiar group of wasps distinct with large and elongated bodies, the new insects also draw attention with the curious names they have been formally assigned with.

Among them, there are species named after characters from the television series Star Trek and Mark Twain’s The Prince and the Pauper, and five wasps bearing species names all translating to ‘helmet’. The study, conducted by graduate student Katherine C. Nesheim and Dr. Norman F. Johnson, both affiliated with the Ohio State University, USA, and Dr. Lubomír Masner, Agriculture and Agri-Food Canada, is published in the open access journal ZooKeys.

The larvae of the studied wasps parasitise the eggs of lanternflies and planthoppers. These species inhabit exclusively the Neotropical region, with their range stretching from the Isthmus of Tehuantepec in the north to Misiones in southern Paraguay. Despite being quite abundant in the region, these insects have remained under-researched until recently.

One of the newly discovered wasp is named Phanuromyia odo, where the species name odo refers to the Star Trek: Deep Space Nine fictional character of the same name. In the popular sci-fi television series, Odo belongs to a species of shapeshifters called Changelings. The reason for the scientists to associate the parasitoid with the character is the spectacular variability observed within the insect species. In fact, it was this peculiarity that, at some point, led the entomologists believe they were dealing with two separate species.

The authors do not make a clear statement that the new species P. pauper has a name inspired by the famous novel The Prince and the Pauper by Mark Twain. Instead, they justify their choice with the fact that the species lacks a specific morphological feature – thus making it ‘poor’. On the other hand, the authors confirm that the new species called P. princeps is of ‘blue blood’ indeed, having its name derive from the other main character of the same book. Furthermore, both species are reported to look a lot like each other.

Among the curious names in the list of new species, there are also five wasps whose scientific names all translate to ‘helmet’ in three different languages – Greek, Latin and Old Norse. The reason behind is that they have unusually large heads, which reminded the scientists of a “knight wearing a helmet”. Likewise, a related species received a name that in Latin means ‘wearing a hood’.

There is also a species, whose name means ‘having long hair’, and another called ‘constellation’ in Latin.

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Original source:

Nesheim KC, Masner L, Johnson NF (2017) The Phanuromyia galeata species group (Hymenoptera, Platygastridae, Telenominae): shining a lantern into an unexplored corner of Neotropical diversity. ZooKeys 663: 71-105. https://doi.org/10.3897/zookeys.663.11554

The first crowdfunded study in Japan: Micro X-ray observation of a fleshy brittle star

Not only have scientists from Japan performed the first non-destructive morphological observations on the Fleshy brittle star, Asteronyx loveni, using micro X-ray tomography, but they also published their research as the first study supported via crowdfunding in the Asian country.

The team leader, Dr. Masanori Okanishi, Ibaraki University, managed to raise part of the funds via Japan’s pioneering crowd-funding platform academist. The study by Dr. Masanori Okanishi, Dr. Toshihiko Fujita, National Museum of Nature and Science, Tsukuba, Yu Maekawa and Dr. Takenori Sasaki, University of Tokyo, is now openly available in the open access journal ZooKeys.

While taxonomy is generally considered as “minor” and “basic” discipline within biology, it could be extremely strenuous for taxonomists to apply for and receive funding. Thus, Dr. Okanishi jumped to the conclusion that his planned study might have a go via crowdfunding instead.

Dr. Okanishi approached academist in April 2014, when he was a Postdoctoral researcher at Kyoto University. Titled “Taxonomy of bathyal euryalid ophiuroids”, it was not long before his research project successfully raised 634,500 JPY (ca. 5,600$).

Having already stumbled across several undescribed species of the brittle star genus Asteronyx, the scientists directed the raised funds towards the genetic and morphological analysis of Asteronyx loveni.

As suggested by its common name, the fleshy brittle star (Asteronyx loveni) is covered by thick skin, making it particularly difficult for scientists to observe the body in detail without dissolving the skin. However, modern computed tomography with micrometer resolution allowed for the 3D images of skeletal ossicles and soft tissues to be constructed with no physical intervention whatsoever.

“The present case indicates that crowdfunding will increase the chances to finance the funds for researchers in those disciplines and activate the research area,” conclude the authors.

The newly discovered brittle star species are yet to be published.

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Original source:

Okanishi M, Fujita T, Maekawa Y, Sasaki T (2017) Non-destructive morphological observations of the fleshy brittle star, Asteronyx loveni using micro-computed tomography (Echinodermata, Ophiuroidea, Euryalida). ZooKeys 663: 1-19. https://doi.org/10.3897/zookeys.663.11413