Trigonopterus corona, the new species of tiny beetle named after the coronavirus

… and 27 other new species of beetles discovered on Sulawesi Island

Many curious animals can be found on the Indonesian Island of Sulawesi – such as the deer-hog and the midget buffalo. But the island’s tropical forests hide a diversity of tiny insects that still remains largely unexplored. Museum scientists from Indonesia and Germany have just discovered 28 new species of beetles, all belonging to the weevil genus Trigonopterus.

Twenty-four newly discovered species of the genus Trigonopterus from Sulawesi. Image by Alexander Riedel

Most of the new species were collected by Raden Pramesa Narakusumo, curator of beetles at the Museum Zoologicum Bogoriense, from two localities of Central Sulawesi Province: Mt. Dako and Mt. Pompangeo. In fact, the forests on their slopes had never been searched for small weevils before.

A view from a ridge over the cloudy slopes of Mt. Pompangeo. Photo by Raden Pramesa Narakusumo

His research partner, Alexander Riedel of the Natural History Museum Karlsruhe, had been studying this genus for the past 15 years and was planning for a research trip to Papua New Guinea, when the COVID-19 pandemic hit. Finding himself grounded, he decided to work on the specimens from Sulawesi together with Narakusumo instead.

After diagnosing the new species, it was a challenge to find suitable names for them. One obvious choice was Trigonopterus corona, which reflects the large impact of the COVID-19 pandemic on this project. However, T. corona is by far not the first insect species with a pandemic-inspired name. In the last year, we’ve seen the species descriptions of the caddisfly Potamophylax coronavirus and the wasps Stethantyx covida and Allorhogas quarentenus.

Trigonopterus corona.
Trigonopterus ewok.

While some of the newly described species go by rather ‘standard’ names that derive from either the localities they have been collected from or their distinct characters, others were given a free pass to the Hall of Fame. Two of them were named after Indonesian movie characters (T. gundala and T. unyil), while T. ewok is another addition based on the Star Wars universe – perfectly in line with T. chewbacca, T. yoda and T.porg, all described between 2016 and 2019 by teams involving Riedel. The two-millimeter-long, rust-coloured Trigonopterus ewok was found at 1900–2000 m on Mt Pompangeo, hiding among the leaf litter in the forest.

But how come the critters have remained overlooked for so long? Almost all of these beetles measure only 2-3 millimeters, while most entomologists have a preference for the larger and strikingly looking stag beetles or jewel beetles. 

A second factor is the superficial resemblance of many species: they are most easily diagnosed by their DNA sequences. Besides the publication in the open-access journal ZooKeys, high-resolution photographs of each species were uploaded to the Species ID website, along with a short scientific description. This provides a face to the species name, an important prerequisite for future studies.

R.P. Narakusumo during fieldwork at the top of Mt. Dako. Photo by Raden Pramesa Narakusumo

This is the duo’s second published paper on Trigonopterus weevils from Sulawesi – the first one describing the whopping 103 new species from the area. Currently, the known Trigonopterus species on the island amount to 132, which is likely a mere fraction of the real diversity. The numerous mountains of Sulawesi have a distinct fauna of endemics that have evolved over the past millions of years, and these wingless, flightless weevils, highly isolated in their habitats, are a good example of this diversification. Their evolution is interwoven with the island´s geological history. Riedel wants to increase the number of sampled localities: 

“Once we have enough locality coverage and understand the weevils’ evolution, we can draw conclusions on the geological processes that formed the island of Sulawesi. This is a fascinating subject, because this island was formed by the fusion of different fragments millions of years ago.” The new species thus fill an important gap required for solving the island´s geological puzzle.

For the Indonesian side, it is equally important to obtain an inventory of species: “A large percentage of Indonesian biodiversity is yet unknown and we need names and diagnoses of species, so we can use these in further studies on conservation and bioprospecting,” says R. Pramesa Narakusumo. “Two of the newly described species came from our museum collection, and this underlines the importance of museums as a source for biological discoveries,” he added.

With many more new species of this genus to be expected, it is a lucky coincidence that the number of Star Wars characters is equally long. May the Force be with these researchers!

Research article:

Narakusumo RP, Riedel A (2021) Twenty-eight new species of Trigonopterus Fauvel (Coleoptera, Curculionidae) from Central Sulawesi. ZooKeys 1065: 29-79. https://doi.org/10.3897/zookeys.1065.71680

The first case of a Portuguese beetle living exclusively in groundwater

New to science, the species was named after Pluto, the ruler of the underworld in Greek mythology

A diving beetle demonstrating various adaptations to the life underground, including depigmentation and evolutionary loss of eyes, was discovered at the bottom of a clay pound in the cave Soprador do Carvalho, Portugal. The species turned out to be the very first in the whole order of beetles (Coleoptera) to be known exclusively from the underground waters of the country.

The Soprador do Carvalho cave (Portugal) is the type locality of the newly described species Iberoporus pluto. Photo by Ignacio Ribera.

Despite not being able to find any other specimens during their study – save for the single female, the team of Dr Ignacio Ribera, Institute of Evolutionary Biology (Spain) and Prof Ana Sofia P. S. Reboleira, University of Copenhagen (Denmark) identified the beetle as new to science, thanks to its unambiguous morphology in combination with molecular data.

Profile view of the newly described species Iberoporus pluto. Photo by Ignacio Ribera.

Aptly named Iberoporus pluto in reference to the ruler of the underworld in Greek mythology Pluto, the species was recently described in the open-access journal ZooKeys.

With a uniformly pale orange body measuring 2.8 mm in length and 1.1 mm in its widest part, the beetle is larger than the rest species known in its genus, and its appendages are longer and more slender. While blindness and depigmentation are clear adaptation to life away from sunlight, the elongated limbs and antennae reflect poor swimming abilities needed in a subterranean habitat. Going for 4 km in horizontal direction, Soprador do Carvalho is the largest in the Dueça cave system, located in the north-eastern part of the Sicó karst area in central Portugal. In recent years, the cave is being explored for tourism.

“The knowledge of the subterranean fauna from Portugal has significantly increased over the last decade, with the description of a high number of obligate subterranean species (tripling their number) and the establishment of new biogeographic patterns,” explain the authors of the study. “A high number of these species are stygobiont (i.e. confined to groundwater), mostly from wells in the north of the country, where evapotranspiration is higher.”

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Original source:

Ribera I, Reboleira ASPS (2019) The first stygobiont species of Coleoptera from Portugal, with a molecular phylogeny of the Siettitia group of genera (Dytiscidae, Hydroporinae, Hydroporini, Siettitiina). ZooKeys 813: 21-38. https://doi.org/10.3897/zookeys.813.29765

Beetle named after actress & biologist Isabella Rossellini for her series about animals

A new species of beetle with remarkably long genitalia that hint at a curious evolutionary “sexual arms race” has been described from Malaysian Borneo.

The new insect was named after actress and biologist Isabella Rossellini in honour of her stage shows and Webby Award-winning series of films about animal reproduction, featured on SundanceTV.

The species is described by scientists Menno Schilthuizen and Iva Njunjic of Naturalis Biodiversity Center and Universiti Malaysia Sabah, and Michel Perreau, Sorbonne Université, Paris. Their paper is published in the open access journal ZooKeys.

The new species, Ptomaphaginus isabellarossellini, finds a place among the 30 known species of round fungus beetles (subfamily Cholevinae) recorded by the authors from the island of Borneo. Of these, there are a total of 14 which had remained unknown to science until now.

The reason why the scientists named this particular species after the famous actress is its genitalia. The beetle’s penis carries a long, whip-like thread, called flagellum, whereas the female has a similarly long tube leading up to a sperm storage organ.

Lead author Menno Schilthuizen, who himself has previously released a simultaneously educative and entertaining book about the evolution of genitals, says that such exaggerated male and female genitalia often betray an “evolutionary sexual arms race”.

On the male side, natural selection favours longer genitalia because of the ability to directly reach the female’s sperm storage organ. However, female beetles would rather retain their right to favour the DNA of a certain mate over the rest. The upshot is that, over long periods of evolution, penises get longer and vaginas get deeper. Similar evolutionary genital exaggeration is also known in rove beetles and ducks.

“This is better than winning the Oscar,” says honoured Isabella Rossellini.

The Italian-born actress, filmmaker, author, philanthropist, and model has even featured the new beetle in her new Link Link Circus stage shows. Dealing with animal behaviour, the series will be touring theaters in Europe this fall.

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Original source:

Schilthuizen M, Perreau M, Njunjic I (2018) A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae). ZooKeys 777: 57-108. https://doi.org/10.3897/zookeys.777.23212

New species of ground beetle described from a 147-year-old specimen

While new species are most commonly described based on recent field collections, undertaken at poorly explored places, some are identified in museum collections, where they have spent decades before being recognised as new to science. Such is the case of an unusually large and likely extinct ground beetle found at the Muséum national d’Histoire naturelle, Paris, whose story began in the distant 1860s with Dr. Eduard Graeffe’s trip to Samoa. Now, a century and a half later, the beetle is finally described by Dr. James K. Liebherr, Cornell University, USA, in the open access journal Zoosystematics and Evolution.

Much like the rest of the species within the genus, the beetle now going under the name Bryanites graeffi showed vestigial flight wings and other traits associated with flight-wing loss. However, at length of 16.2 mm it is the largest for the taxonomic group it is now assigned to. Although this may seem way too obvious for taxonomists to overlook, the beetle’s relatives are just as obscure. The Bryanites genus was previously known from two species represented by two specimens only, collected in 1924 from Savai?i Island by Edwin H. Bryan, Jr., Bernice P. Bishop Museum in Honolulu, during the Bishop Museum’s Whitney South Seas Expedition.

As a result, we now have three species representing an evolutionary radiation in Samoa, all known from single specimens collected long ago. The phylogenetics of these three species link them to other groups from Fiji and New Zealand.

What is the advantage of knowledge about species that existed some 90-150 years ago, but no longer? It might actually point us to the actual level of impact mankind has on natural ecosystems. The cause of the likely extermination of Bryanites graeffi might never be known with certainty, however, the colonization of many Pacific islands by the Polynesian rat has always been followed by the diminution or elimination of native insect species. Thus, we can add another likely victim to the list of species that have been adversely impacted by mankind’s commensal voyagers.

The species bears the name of its original collector to pay tribute to Dr. Graeffe and his hard work while collecting insects in the rain forest of Samoa well over a century ago .

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Original source:

Liebherr JK (2017) Bryanites graeffii sp. n. (Coleoptera, Carabidae): museum rediscovery of a relict species from Samoa. Zoosystematics and Evolution 93(1): 1-11. https://doi.org/10.3897/zse.93.10802

New Chinese leaf-roller weevil does not know how to roll leaves

A long-term project on insect-seed interactions, currently being carried out by researchers of the Institute of Zoology (Chinese Academy of Sciences) in a subtropical forest near Dujiangyan City, Sichuan, China, revealed the presence of larvae of an unknown weevil species eating the seeds in the pods of a shrubby legume.

Scientists from the Institute of Zoology, China, Xiangyang Lv, Zhishu Xiao, Zhiliang Wang, Runzhi Zhang, and Miguel A. Alonso-Zarazaga, also affiliated with the Museo Nacional de Ciencias Naturales (CSIC), Spain, published the description of the new genus and species, named Evemphyron sinense, and added data on its biology in the open access journal ZooKeys.

Because of its peculiar features, it was difficult to locate the closest relatives of this new species. However, a few characteristic traits of the body and genitalia, strongly pointed to its placement within the tribe Deporaini.

The closest, although seemingly rather far-related to the new species, beetles are considered to be a genus with scattered distribution, stretching from the Russian Far East to the Indian Himalaya.

However, they are smaller weevils whose females cut shoots to lay their eggs. On the other hand, the males in both genera share a peculiar patch of hairs, probably related to pheromone dispersal. Likewise, each species is associated with legumes.

The curious feature of this weevil group (Deporaini) is that the vast majority of its species are leaf-rollers. The females cut a hardwood leaf in a peculiar and mathematical way and roll it, laying one egg inside each one. This behaviour, which is known in other far-related weevils of the same family, seems to have appeared independently in different evolutionary branches. In the case of Deporaini, this behavioural trait evolved after the new genus became a distinct one.Figure 5 large

As a result, the new genus is considered to be one of the two most primitive within the tribe. In fact, it might be the most primitive one, taking into account a number of morphological traits as well.

It could be that the new beetle never knew how to roll a leaf to make nests and shelter its offspring.

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Original source:

Lv X, Alonso-Zarazaga MA, Xiao Z, Wang Z, Zhang R (2016) Evemphyron sinense, a new genus and species infesting legume seedpods in China (Coleoptera, Attelabidae, Rhynchitinae).ZooKeys 600: 89-101. doi: 10.3897/zookeys.600.6709

More than just hippos and crocs: The hidden biodiversity of the iSimangaliso Wetland Park

iSimangaliso Wetland Park, a UNESCO World Heritage Site in the sub-tropical north-eastern corner of South Africa has become famous for its birdlife, crocodiles and hippopotamuses that frolic in the warm estuarine waters of Lake St Lucia. However, there’s more to the park than the “big and hairy”, according to aquatic ecologist Prof Renzo Perissinotto at Nelson Mandela Metropolitan University (NMMU) in Port Elizabeth, whose research is published in the open access journal ZooKeys.

“Although we have spent several decades focusing on life in the estuary, we only recently came to realise that much of the wealth of biodiversity in the park exists in the small freshwater ponds that are adjacent to, but disconnected from, the main lake,” he says.Image 1

The St Lucia lake itself is generally brackish and is located on a large sandy expanse known as the Maputaland coastal plain. Dotted across the landscape of this coastal foreland are numerous temporary freshwater ponds, seeps and small streams that are disconnected from the brackish lake body.

A team of self-proclaimed “beetle nerds”, led by Prof Perissinotto, got together from NMMU and Plymouth University (UK) and uncovered more species of water beetles in these tiny water bodies than is known for any other similar-sized region in southern Africa.

The beetle collection trips were done over a 16-month period and revealed 68 species of predaceous water beetles alone, termed more formally as the “Hydradephaga”. The iSimangaliso Wetland Park houses approximately 20% of the total number of known species for this beetle group in the whole of southern Africa. Of the species collected during their expeditions, five have never been recorded in South Africa before, highlighting our poor understanding of aquatic insect distributions in this part of the world.

Most of the species collected (almost 80%) belonged to the family Dytiscidae, more commonly known as “diving beetles” due to their lifestyle that involves coming up for air and immediately diving back down to the depths to carry on hunting unsuspecting prey, which can be as large as small fish and amphibians.Image 2

Prof Perissinotto and his NMMU colleague Dr Matthew Bird, together with water beetle specialist Prof David Bilton (Plymouth University), collected specimens ranging from 1 mm to almost 5 cm in length (the tadpole eaters). According to Prof Bilton, “Irrespective of size, these water beetles are a crucial component of the iSimangaliso ecosystem in that they are the primary predators in these temporary wetlands, which generally lack fish. Their abundance and diversity can be used to gauge the overall health of wetland ecosystems as they are sensitive to pollution, for instance”.

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Original source:

Perissinotto R, Bird MS, Bilton DT (2016) Predaceous water beetles (Coleoptera, Hydradephaga) of the Lake St Lucia system, South Africa: biodiversity, community ecology and conservation implications. ZooKeys 595: 85-135. doi: 10.3897/zookeys.595.8614

New curiously scaled beetle species from New Britain named after ‘Star Wars’ Chewbacca

Chewbacca, the fictional ‘Star Wars’ character, has given his name to a new species of flightless beetle, discovered in New Britain, Papua New Guinea. Although Trigonopterus chewbacca was only one of the four black new weevil beetles found during the expedition, it stood out with its curious scales, which made the authors think of Han Solo’s loyal companion.

Scientists Dr Matthew H. Van Dam, SNSB-Zoological State Collection, Germany, Raymond Laufa, The University of Papua New Guinea and Dr Alexander Riedel, Natural History Museum Karlsruhe have their paper, where they describe the new species, published in the open access journal ZooKeys.

Failing to understand how was it possible that the hyperdiverse beetle genus Trigonopterus has never been spotted in New Britain, two of the researchers travelled to the island to double-check the foliage and leaf litter. Interestingly, the genus thrives best in Melanesia, the Oceania subregion, where Papua New Guinea is located, yet there had been only a single Trigonopterus species known from Bismarck Archipelago prior to the present study.

Eventually, having spent ten days sifting leaf litter and beating foliage, the authors discovered eighteen individuals in primary forests growing on limestone karst, and later assigned them to four separate species. However, these few findings are still striking, given the abundance of the beetles in similar localities in the New-Guinean mainland.

Unlike its sci-fi namesake, the Chewbacca beetle cannot rely on its measurements to scare other possibly malevolent species off. It only measures between 2.78 and 3.13 mm. Dissimilar again, is its body, which is black and rhomboid-shaped, while its legs and antenna appear rusty. What likens the beetle to “Chewie”, however, is its distinctively dense scales, covering its head and legs.

In conclusion, the authors note that the beetle genus must have colonised New Britain at least four times in the past. “Given the size, mountainous topography and tropical vegetation of New Britain, it is likely that Trigonopterus has undergone some local speciation on the island, but this possibility requires further investigation,” they say.

 

Original source:

 

Van Dam MH, Laufa R, Riedel A (2016) Four new species of Trigonopterus Fauvel from the island of New Britain (Coleoptera, Curculionidae). ZooKeys 582: 129-141. doi: 10.3897/zookeys.582.7709

Over 300 new beetle records for New Brunswick, Canada, in a special issue of ZooKeys

Beetles diversity in New Brunswick, Canada, has elicited the interest of biologists for over a century and continues to do so. In 1991, 1,365 species were known from New Brunswick. That number had increased to 2,703 by 2013, as a result of a series of publications in three previous special ZooKeys issues and other publications. In spite of that work, there were still gaps in the knowledge of the Coleopteran fauna.

Now, a group of insect specialists have joined forces in the name of their love for beetles, and compiled their findings from the last three years, reporting another 303 species for New Brunswick, including thirty-two species new to science. All of these records are published in a special issue, titled “The Coleoptera of New Brunswick and Canada: Providing baseline biodiversity and natural history data” of the open access journal ZooKeys.

It might have been only three years, but the authors of the present issue have expanded the beetle fauna of New Brunswick by 13%. On a longer timeline since 1991, the increase rises to an impressive 124%.

These figures come as a result of the 303 new records for New Brunswick that included 32 species, which the team have found to be new to science, 4 new North American records, 21 new Canadian records, 270 new provincial records, and 45 adventive species that have somehow arrived in the region from elsewhere. As a result, the beetle fauna of New Brunswick currently comprises 3,062 species.

“This information constitutes a baseline of biological knowledge that is critical to support other branches of science,” point out the authors.

“It is important to remind ourselves that the understanding of biological diversity is not possible without taxonomic research, which is thought by many to be the foundation of biological science,” they explain. “Data on the mega-diversity of life and knowledge on species identity and distribution require discovery, description, cataloguing, and organization in order to be made accessible to a wide audience.”

“This work would not have been possible to complete without the enthusiasm, determination, and professionalism of a small number of dedicated individuals who are acknowledged in the papers in this special issue,” the researchers conclude. “We hope that this special issue will generate a positive response and further interest in the Coleoptera fauna of New Brunswick and Canada, as many new discoveries await.”

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Original source:

Webster RP, Bouchard P, Klimaszewski J, Sweeney JD (2016) History of Coleoptera collecting in New Brunswick, Canada: advancing our knowledge of the Coleoptera fauna in the early 21st century. In: Webster RP, Bouchard P, Klimaszewski J (Eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys 573: 1-18. doi: 10.3897/zookeys.573.8123

Twenty-four new beetle species discovered in Australian rain forests

As many as twenty-four new species from Australian rainforests are added to the weevil genus Trigonopterus. Museum scientists Dr. Alexander Riedel, State Museum of Natural History Karlsruhe, Germany, and Rene Tanzler, Zoological State Collection Munich, Germany, have first discovered them among unidentified specimens in different beetle collections. The study is published in the open-access journal ZooKeys.

Australia is well known for its extensive deserts and savanna habitats. However, a great number of native Australian species are restricted to the wet tropical forests along the east coast of northern Queensland. These forests are also the home of the recent discoveries.

Most of the weevil species now recognised as new have already been collected in the 80s and 90s of the past century. Since then they had been resting in museum collections until German researcher Alexander Riedel had the opportunity to study them.

“Usually a delay of decades or even centuries occurs between the encounter of a new species in the field and its thorough scientific study and formal naming,” he explains. “This is due to the small number of experts who focus on species discovery,” he elaborates. “There are millions of unidentified insect specimens stored in collections around the world but only few people have the training necessary to identify those of special interest.”

However, old museum specimens alone are not enough either. Nowadays, researchers try to include DNA data in their descriptions, and the necessary sequencing techniques work more efficiently with freshly collected material. Therefore, the scientists set off to the field after they have studied the collections of others. Nevertheless, the German team were led to the discovery of one additional new species, which had never been seen before. They called itTrigonopterus garradungensis after the place where it was found.

All of the newly described weevils are restricted to small areas. Some are found only in a single locality. Presumably, this is a consequence of their winglessness, which has prevented them from spreading around. Furthermore, most of them dwell in the leaf litter where they are easily overlooked. Usually, they come to light during specific surveys of the litter fauna.

This is what Geoff Monteith from the Queensland Museum in Brisbane, for instance, has done in the past. As a result, his work is now relevant to conservation because highly localised species are extremely vulnerable to changes of their habitat such as climate change or the arrival of invasive species.

It is likely that Trigonopterus weevils have originated in Australia, the oldest landmass in the region. The island of New Guinea is geologically much younger, but there the genus has quickly enough diversified into hundreds of species. Studies investigating such evolutionary processes depend on names and clear diagnoses of the species. As a result of the present research, for the Australian fauna these are now available.

Besides the publication in the open-access journal ZooKeys, high-resolution photographs of each species are uploaded to the Species ID website, along with the scientific description. All this puts a face to the species name, and therefore is an important prerequisite for future studies on their evolution.

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Original Source:

Riedel A, Tanzler R (2016) Revision of the Australian species of the weevil genus Trigonopterus Fauvel. ZooKeys 556: 97-162. doi: 10.3897/zookeys.556.6126

Lava attraction: 74 new beetle species found hiding in plain sight on a Hawaiian volcano

Confined to the limits of Haleakala volcano, Maui Island, Hawaii, the beetle fauna there turns out to be not only extremely diverse, but very abundant as well. When Prof. James Liebherr of the Cornell University Insect Collection thoroughly sampled beetle populations on the volcano, he identified 116 species of round-waisted predatory beetles, including 74 new to science. The taxonomic revision, complete with descriptions of the new species, is now published in the open-access journal ZooKeys.

The present discoveries and observations are certainly surprising due to their scale, even though it has been long known that the Hawaiian Islands support disproportionately high levels of biodiversity. For this group of native round-waisted beetles, called Mecyclothorax in the zoological naming system, there are 239 species across the Hawaiian Islands, all of them descended from a single colonizing species.

The 116 species known from Haleakala make that volcano the center of biodiversity for this group within Hawaii. These beetles’ evolution during the 1.2-million-year lifespan of Haleakala volcano means they have speciated faster than most organisms on Earth, including the Hawaiian Drosophila and the cichlid fishes of eastern Africa.

No less striking is the fact that the 74 newly described beetle species previously evaded discovery within the limits of Haleakala’s 1,440 km2 of surface area. Reasons for this include the restricted distributions of many of the beetle species, and the previous lack of comprehensive field sampling. During his research, Prof. Liebherr examined all quarters of the mountain to eventually find many places of 1′ latitude × 1′ longitude where more than 20Mecyclothorax species lived closely together within a limited area of forest.

Most of these diverse microhabitats were discovered in windward rainforests. Moreover, different forest areas, geographically isolated from each other by volcanic lava flows, steep valleys, or extensive mudflows, supported different sets of species. “Haleakala volcano is a large pie with different sets of beetle species living in the different slices,” comments Prof. Liebherr. “Actually the different pie slices are just like the original Hawaiian land divisions called ahu pua’a, showing that the Hawaiian people had a keen sense for how their island home was organized.”

Additionally, the round-waisted beetle species seem to thrive across a wide range of altitudes, with their populations covering the major part of the mountain’s height. Historical as well as modern records have identified representatives of these insects from 450-metre elevation up to the volcano’s summit at 3000 m. However, given land conversion and the influx of alien invasive plants, habitats below about 1000 m have been seriously disrupted, and these elevations support few native beetles.

Looking to the future, Liebherr points out that “the substantial level of sympatry, associated with occupation of diverse microhabitats by these beetles, provides ample information useful for monitoring biodiversity of the natural areas of Haleakala.”

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Original source:

Liebherr JK (2015) The Mecyclothorax beetles (Coleoptera, Carabidae, Moriomorphini) of Haleakala-, Maui: Keystone of a hyperdiverse Hawaiian radiation. ZooKeys 544: 1-407. doi: 10.3897/zookeys.544.6074