New proposal for a subspecies definition triggered by a new longhorn beetle subspecies

The discovery of a new subspecies of longhorn beetle from Scandinavia triggered a discussion on the vague organism classification rank ‘subspecies’.

As a result, a newly proposed definition of subspecies has been published along with the description of the taxon in the open access journal ZooKeys by the research team of Henrik Wallin and Johannes Bergsten from the Swedish Museum of Natural History and Torstein Kvamme from the Norwegian Institute of Bioeconomy Research.

The northernmost populations of Saperda populnea display a number of divergent traits, including a shorter male antenna and reduced pubescens. In contrast with their most closely related subspecies (Saperda populnea populnea), whose favourite host plant, amongst others, is the European aspen, the new subspecies (Saperda populnea lapponica) specialised exclusively on downy willow (Salix lapponum).

Image 2According to the once no less disputed definition of species, regardless of their unique traits, populations cannot be considered as separate species until they are no longer able to produce fertile offspring according to the Biological Species concept, this being one of a number of proposed species concepts. A consensus is emerging around the unified species concept defining species as separately evolving (meta) population lineages.

However, differentiating between subspecies nowadays is a significantly tougher task, since there is no stable definition of the rank yet. Through the years, there have been various explanations of what a subspecies is and what criteria it needs to meet in order to be classified as one.

Compared to previous definitions, the researchers decode it quite simply. To them, the only necessary attributes a population needs to possess before being deemed a subspecies are that they are a potentially incipient new species; diagnosed by at least one heritable trait; and either partially or completely isolated geographically.

Furthermore, they refute a number of factors, including reciprocal monophyly in neutral markers, the “75% rule”, reproductive compatibility and the degree of gene flow.

The concept of subspecies has been so problematic that there have been even those scientists who have argued that taxonomy needs to discard it altogether.

However, the authors note that it is already formally recognised by the International Commission on Zoological Nomenclature (ICZN 1999), “albeit without giving any advice or criteria for its recognition.”

“The concept is more than a mere academic debate as subspecies are recognized in various Red Lists and conservation programs, and hence the recognition as a subspecies or not can have legal and monetary consequences.”

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Original source:

Wallin H, Kvamme T, Bergsten J (2017) To be or not to be a subspecies: description of Saperda populnea lapponica ssp. n. (Coleoptera, Cerambycidae) developing in downy willow (Salix lapponum L.). ZooKeys 691: 101-148. https://doi.org/10.3897/zookeys.691.12880

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

How bears bulk up ahead of the summer: A study into the Asiatic black bear’s spring diet

Much like gym enthusiasts, every year Asiatic black bears seem to be on the lookout for protein-rich food ahead of the summer, so that they can bulk up on lean muscle mass in place of the fat tissue formed last year prior to hibernation. This was concluded in a study by Dr. Shino Furusaka, Tokyo University of Agriculture and Technology and his team, based on direct observations on bears living across an area of about 60 km2 in Japan. The study is published in the open access journal ZooKeys.

In order to determine the bears’ food preferences and habits, the scientists followed a large number of animals in the Ashio area of the Ashio-Nikko Mountains in Japan from April to July in both 2013 and 2014. To avoid unnecessary intrusion, they stayed at a distance of at least 200 metres using video cameras with telescopic lenses to document the sightings. Having documented the plant species the bears consumed, the researchers studied their nutritional content and made conclusions about the nutrients needed for the species after hibernation.

While heavily dependent on food availability, generally the bears were noted to prefer food which is high in protein, but poor in fibre — likely because their stomachs and intestines were unable to efficiently digest the latter. Furthermore, the protein-rich diet ensures that the muscle mass is rebuilt to replace the lost winter fat.

Interestingly, the bears were observed to change their food preferences as spring progressed and that seemed to be linked to the shifts in the nutritional value of the available food.

Starting with their observations at the beginning of April, the scientists did not record any feeding behaviour until the end of the month. As leaf flush was yet to occur, the animals were active and feeding on overwintered grass. However, in early May, the bears began consuming newly emerged leaves, grass and, later in the month, they added flowers to their menu.

A shift in behaviour occurred in the following months. In June and July, the bears were seen to feed mainly on ants, with a small portion of their food intake consisting of grasses, sika deer carcasses and bees. Curiously, when the scientists looked into the nutritional content of the same plants which the animals sought only a few weeks ago, they found out that now they were significantly poorer in protein and richer in fibre.

Another finding showed that the calories in the different items were not related to the choice of food which likely proves that the key factor is none other than the amount of protein, provided that the fibre value is low enough for good digestibility.

Understanding the food preferences and habits of animals, as well as the reasons behind them, is essential for the development and revision of habitat management plans. However, previous knowledge of the feeding behaviour of Asiatic black bears has been based solely on faecal analyses which has not provided sufficient details on which nutritional factors influence the use of particular foods.

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Original source:

Furusaka S, Kozakai C, Nemoto Y, Umemura Y, Naganuma T, Yamazaki K, Koike S (2017) The selection by the Asiatic black bear (Ursus thibetanus) of spring plant food items according to their nutritional values. ZooKeys 672: 121-133. https://doi.org/10.3897/zookeys.672.10078

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.

CalinagaTogether, 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

Caught red-handed: The ‘Candy striped hermit crab’ is a new species from the Caribbean

Recent underwater photographs and video obtained using scuba equipment by underwater photographer Ellen Muller at dive sites in the National Marine Park of the southern Caribbean island of Bonaire revealed the presence of a small, secretive and brightly colored red-striped hermit crab that proved to represent a species new to science. The new few-millimeter species is described in the open access journal ZooKeys.

Extra ImageThe color pattern reminded author Dr. Rafael Lemaitre, Smithsonian Institution, USA, of traditional candy cane, and thus he assigned the common name “Candy striped hermit crab”. Meanwhile, the scientific name of the new species is Pylopaguropsis mollymullerae after Ellen Muller’s young granddaughter Molly Muller. The underwater photographer believes that the honor would “inspire her to continue the tradition of protecting the amazing and fragile diversity of marine life in Bonaire”.

The unusual hermit crab was first photographed inadvertently alongside a “flaming reef lobster”, while observing invertebrates that aggregate in crevices under a large coral ledge. Subsequently, more hermit crabs were photographed in various crevices shared with moray eels such as the “broad banded moray”, “spotted moray”, and “green moray”. When permits were obtained from the Government of the Island Territory of Bonaire, a few specimens were collected and brought for study to the Smithsonian Institution. The formal description was then prepared for publication and specimens were deposited in the collections of the National Museum of Natural History, as required by scientific rules when naming new species.

The shape of the right, or major, pincer of this new hermit crab species is remarkable and unique with its shape and massive size when compared to the body. The underside of the claw of this pincer is deeply excavated, scoop-like. The function of the pincer and claw, however, is at present unknown, although a video shows that it is used by the hermit crab to push itself while crawling along the bottom.

Image 1The behavior of this new hermit crab is intriguing. Is there an ecological association of this new species with moray eels? Could this new hermit crab species function as a “cleaner” or a “den commensal”? At least in one instance, an individual was observed crawling on the body of a “broad banded moray”, perhaps feeding on mucus or materials present on it. These observations could be interpreted as evidence of some kind of symbiotic association, or den commensalism, between the two animals. The brightly colored pattern of the hermit crab with red stripes and very long, hairy antennae are also typical for most crustaceans considered fish “cleaners”.

“Cleaning” parasites or fouling organisms from the body of many cooperating fish, or removal of undesired food particles by certain small and colorful shrimps has been known for nearly 60 years, but never has a hermit crab been documented to engage in such type of ecological association. Further studies are needed in order to confirm the true ecological role of this fascinating hermit crab.

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Original source:

Lemaitre R (2017) Discovery of a new species of hermit crab of the genus Pylopaguropsis Alcock, 1905 from the Caribbean: “den commensal” or “cleaner”? (Crustacea, Anomura, Paguridae). ZooKeys 646: 139-158, https://doi.org/10.3897/zookeys.646.11132

New crab species shares name with 2 ‘Harry Potter’ characters and a hero researcher

While not much is known about the animals living around coral reefs, ex-Marine turned researcher Harry Conley would often take to the island of Guam, western Pacific Ocean, and dig deep into the rubble to find fascinating critters as if by magic learnt at Hogwarts. Almost 20 years after his discoveries and his death, a secret is revealed on the pages of the open access journal ZooKeys – a new species and genus of crab, Harryplax severus.

Having dug as deep as 30 m into Guam’s coral reef rubble, Harry Conley collected many specimens which stayed in his personal collection until the early 2000’s when Dr. Gustav Paulay, currently affiliated with the University of Florida, handed the specimens to the second author of the present study, Dr. Peter Ng, National University of Singapore, which resulted in many discoveries and publications. Among the lot, however, were two unusual specimens which were not studied until much later. Only recently did Dr. Peter Ng and his colleague at the National University of Singapore and lead author of the paper, Dr. Jose Christopher E. Mendoza, discover that they represent not only a new species, but also a new genus.

Having chosen the name Harryplax for the new genus, the two authors pay tribute to the crab’s original collector Harry Conley, who they describe as a “soft-spoken ex-Marine with a steely determination and a heart of gold,” and whose endeavours “have substantially advanced the cause of marine science”. The name is also meant to allude to the main protagonist in J. K. Rowling’s famous fantasy novel series, whose magical abilities the scientists liken to Conley’s knack for finding rare or new species. Of the two authors, Dr. Mendoza is the self-confessed ‘Potterhead’, who was not about to pass up the chance of naming a new crab after his favourite fictional characters. In his turn, Dr. Ng, who knew Harry Conley personally, was quite amused and happy to agree.

Image1 Harryplax_severus_male paratype PRThe crab’s species name, severus, is inspired by another ‘Harry Potter’ character – Professor Severus Snape, who despite being a central character in the series, keeps his background and agenda mysterious until the very end, when he reveals a key secret. Showing his real identity, the character, to the authors, is “just like the present new species which has eluded discovery until now, nearly 20 years after it was first collected”.

The new species is a tiny crab measuring less than a centimeter in both length and width and can be found deep in coral rubble or under subtidal rocks, perhaps also in cavities. To survive in the dark depths, the species has evolved with reduced eyes, well developed antennae, and long, slender legs. For the time being it is known only from the island of Guam.

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Original source:

Mendoza JCE, Ng PKL (2017) Harryplax severus, a new genus and species of an unusual coral rubble-inhabiting crab from Guam (Crustacea, Brachyura, Christmaplacidae). ZooKeys 647: 23-35. https://doi.org/10.3897/zookeys.647.11455

A hair’s breadth away: New tarantula species and genus honors Gabriel García Márquez

With its extraordinary defensive hairs, a Colombian tarantula proved itself as not only a new species, but also a new genus. It is hypothesised that the new spider is the first in its subfamily to use its stinging hairs in direct attack instead of ‘kicking’ them into the enemy.

Described in the open access journal ZooKeys by an international research team, led by Carlos Perafán, University of the Republic, Uruguay, the name of the new spider genus honours an indigenous people from the Caribbean coast region, whose language and culture are, unfortunately, at serious risk of extinction. Meanwhile, its species’ name pays tribute to renowned Colombian author and Nobel laureate for his novel ‘One Hundred Years of Solitude’ Gabriel García Márquez.male kankuamo

The new tarantula, formally called Kankuamo marquezi, was discovered in Sierra Nevada de Santa Marta, Colombia. When examined, the arachnid showed something extraordinary about its defensive hairs and its genitalia. The hairs were noted to form a small oval patch of lance-shaped barbs, hypothesised by the scientists to have evolved to defend their owners by direct contact.

On the other hand, when defending against their aggressors, the rest of the tarantulas in this subfamily need to first face the offender and then vigorously rub their hind legs against their stomachs. Aimed and shot at the enemy, a ball of stinging hairs can cause fatal injuries to small mammals when landed into their mucous membrane (the layer that covers the cavities and shrouds the internal organs in the body). Once thrown, the hairs leave a bald spot on the tarantula’s belly.

“This new finding is a great contribution to the knowledge of the arachnids in Colombia and a sign of how much remains to be discovered,” point out he authors.

Figure 8“The morphological characteristics present on Kankuamo marquezi open the discussion about the phylogenetics relationship between subfamilies of Theraphosidae tarantulas and the evolutionary pressures that gave rise to the urticating hairs.”

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Original source:

Perafán C, Galvis W, Gutiérrez M, Pérez-Miles F (2016) Kankuamo, a new theraphosid genus from Colombia (Araneae, Mygalomorphae), with a new type of urticating setae and divergent male genitalia. ZooKeys 601: 89-109. doi: 10.3897/zookeys.601.7704

Celebration time: ZooKeys releases its 600th issue


zookeys 600 coverWith what already sounds like an annual tradition at this time of the year, we are delighted to announce yet another milestone that
ZooKeys just reached. Our 600th issue is now out and we are just as proud with it as we were exactly five years and a month ago, when we printed out our first three-digit issue number on a ZooKeys cover.

However, we feel nowhere near getting tired of counting pages, covers and issues, nor do we believe this will ever going to happen. Quite the contrary, every year we take more and more pleasure in adding new achievements next to the name of ZooKeys and Pensoft.   

Last year was no exception. During the past 13 months, we published a total of 673 articles, including research findings spectacular enough to reach out to not only the zoological fellowship, but to the wide audience from around the world. While our Impact Factor keeps on increasing, according to the figures Thomson Reuters released last week, we are gratified to observe our progressively growing impact on both the scholarly and the popular-science front.

Thanks to the discoveries, which found a suiting publication partner in ZooKeys, our authors and us made a lot of big headlines in outlets such as National Geographic, Science, CNN, BBC, Sky News, New York Times, Deutsche Welle, Der Standard, DR, Washington Post, Fox News, Huffington Post, The Guardian, NBC News, and a lot more. We had a bit of everything: record-breakers, species given mystic or splashy names and others bearing nerdy ones. Together, we also gave public voice to serious conservation issues, calling for immediate action.

Last June, we introduced you to the Hades centipede, known to be the world’s deepest-dwelling species of its kind. Who knew that the entrance to the Underworld is located in a Croatian cave?

Later on, in November, published with us snail species Acmella nana broke the World record for the tiniest land snail. Moreover, this happened only about a month after we published the previous ‘prizewinner’ Angustopila dominikae, and that one was already tiny enough to fit 10 of its shells within the eye of a needle at the very same time!  

Our pages, which have been and always will be openly available to read for anyone who is online, were also the first to let you know about the existence of the Johnny Cash tarantula, the (Edward) Snowden crayfish, the two daddy longlegs: Smeagol and the ‘Master-of-the-crypt’ Behemoth, the Chewbacca beetle and the Brad Pitt wasp, among many others.

About two months ago, graduate student Madhu Chetri spotted the ancient Himalayan woolly wolf in Nepal. The new knowledge about the beautiful and, sadly, Critically Endangered carnivore, which he acquired, will hopefully help in preventing its otherwise imminent extinction.

In the meantime, Deutsche Welle (DW) featured our Zorro fish along with the eight-legged Johnny Cash’s namesake in their rank list of the 7 “newcomer” species of the year.

While being in the spotlight is definitely a gratifying feeling, we also indulge in our successes achieved far from the eyes of the public, although we are certain that our authors will be just as excited to hear about. Such an accomplishment is our recently sealed partnership with open digital repository Zenodo, who are helping us, along with the rest of the journals, published by Pensoft, to keep our research findings safe and easily accessible by archiving all our articles in both PDF and XML format on the date of publication.

However, let’s not forget that nothing of all the above would be what it is without our authors, editors and reviewers, who have always done their best to keep ZooKeys at the World’s top open access academic journals. We’d especially like to thank our Most active authors, editors and reviewers for being substantial part of ZooKeys.

Critically Endangered and ancient Himalayan wolf needs global conservation attention

Although the Himalayan wolf is visibly distinct from its European cousin, its current distribution has mostly been a matter of assumption, rather than evident truth. The most ancient wolf lineage, known to science, has been listed as Critically Endangered in the National Red List.

Now, an international research team, led by Madhu Chetri, graduate student at the Hedmark University of Applied Sciences, Norway, report the wolf from Nepal’s largest protected area, thus confirming its existence in the country. Their findings are published in the open access journal ZooKeys.

When compared to the European wolf, this one stands out with its smaller size, unusually longer muzzle and stumpy legs. Another clearly distinctive feature is the white colouration around the throat, chest, belly and inner part of the limbs. On the other hand, its characteristic woolly body fur has given the subspecies the common name ‘woolly wolf’.

However, the distinctiveness of the Himalayan wolf is far more than skin-deep. The authors note that recent studies have already revealed that these wolves have split as a separate branch within the ‘tree of life’ so long ago that they are divergent from the whole globally distributed wolf-dog clade. Having undergone such an isolated evolution, the Himalayan wolf is considered of particular conservation concern.

However, the populations are still suffering heavy mortality. As a part of their research, the authors conducted both formal and informal interviews with about four hundred local herders, livestock owners, nomads and village elite to find out more about the status of the human-wolf conflict, as well as their attitudes and perceptions. As a result, they found out that the wolves are considered to pose a threat for the local livelihoods. They were persecuted and killed as a means of depredation.5966_Himalayan ancient wolf

“These genetically distinct Himalayan wolves deserve special conservation attention, at the same time that the conservation of this species in a context of human-wildlife conflict is challenging,” conclude the scientists. “A species action plan needs be formulated that develops mechanisms to minimize conflict, and strategies for motivating local communities towards wolf conservation.”

Original source:

Chetri M, Jhala YV, Jnawali SR, Subedi N, Dhakal M, Yumnam B (2016) Ancient Himalayan wolf (Canis lupus chanco) lineage in Upper Mustang of the Annapurna Conservation Area, Nepal. ZooKeys 582: 143-156. doi: 10.3897/zookeys.582.5966

The lizard of consistency: New iguana species which sticks to its colors found in Chile

During a field trip at 3000 metres above sea level, a group of scientists, led by Jaime Troncoso-Palacios, Universidad de Chile, discovered a new endemic iguana species, in the mountains of central Chile, scientists. Noticeably different in size and scalation, compared to the rest of the local lizards, what initially grabbed the biologists’ attention was its colouration. Not only was it unlike the already described ones, but also appeared surprisingly consistent within the collected individuals, even regardless of their sex. Eventually, it was this peculiar uniformity that determined the lizard’s name L. uniformis. The study is published in the open-access journal ZooKeys.

The researchers found the lizards quite abundant in the area, which facilitated their observations and estimations. Apart from a thorough description of the new iguana along with its comparisons to its related species, the present paper also provides an in-depth discussion about the placement of the new taxon, which had been confused with other species in the past.

While most of the other lizards from the area and its surroundings often vary greatly in colouration and pattern between populations and sexes, such thing is not present in the new species. Both males and females from the observed collection have their bodies’ upper side in brown, varying from dark on the head, through coppery on the back and light brown on the tail. The down side of the body is mainly yellowish, while the belly – whitish. The only variables the scientists have noticed in their specimens are slight differences in the shade with two females demonstrating unusual olive hues on their snouts. These differences in morphology were also strongly supported by the molecular phylogeny through the analysis of mitochondrial DNA, which was performed by Dr. Alvaro A. Elorza, from Universidad Andres Bello.

Accustomed to life in highland rocky habitats with scarce greenery, these lizards spend their active hours, estimated to take place between 09:00 h and 18:00 h hidden under stones. However, they might not be too hard to find due to their size of about 8.5 cm for the males and their abundance in the studied area. The females are more slender and measure 7 cm in length on average.

Having caught one of their specimens while holding a yellow flower in its mouth, the scientists conducted further examination of the stomach contents of the studied individuals and concluded that the species is omnivorous, feeding mainly on plants as well as insects and roundworms.

In conclusion, the researchers showed that there is still a huge gap in the knowledge of the close relatives of the newly described species and their “challenging taxonomy”.

 

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Original source:

Troncoso-Palacios J, Elorza AA, Puas GI, Alfaro-Pardo E (2016) A new species of Liolaemusrelated to L. nigroviridis from the Andean highlands of Central Chile (Iguania, Liolaemidae).ZooKeys 555: 91-114. doi: 10.3897/zookeys.555.6011