Tag: new species

Shadow-loving insect named after Tuomas Holopainen of Nightwish

Tuomas Holopainen, the multi-talented musician and founder of the symphonic metal band Nightwish, is also a full-blooded nature person. This gave conservation biologist Jukka Salmela of Metsähallitus Parks & Wildlife Finland an idea for the name of a new species he found in Finland. Discovered in eastern Lapland during an insect survey, the fungus gnat was given the scientific name Sciophila holopaineni after Tuomas. The new species is described in the open access Biodiversity Data Journal.

“I am very, very touched. This is the highest honour a nature nerd like me can receive,” Tuomas Holopainen replied after Jukka, who collected and described the fungus gnat, asked him for permission to name the species after him.

The idea for the name came up to Salmela while he was thinking about the habitat and appearance of the species. Then, he recalled Tuomas Holopainen’s interest in the natural sciences.

So far, the new species of fungus gnat has been only known from two locations: the Törmäoja Natura Area in Savukoski, eastern Lapland, and a meadow close to the White Sea, Russian Karelia.

The dark and beautiful gnat thrives in shadowy environments. In Törmäoja, it was caught in a river gulch next to the river source, while hiding under the shelter of the forest. Salmela proposes ‘tuomaanvarjokainen’ as the common Finnish name, inspired by the latest Nightwish album. After all, the themes of the album, Endless Forms Most Beautiful, are evolution and the diversity of nature.

Fungus gnats are flies, which feed on dead wood or fungi. Some of the larvae are predaceous. At current count, there are almost 800 species in Finland and about 1,000 in the Nordic countries. In fact, the Fennoscandia region is one of Europe’s biodiversity hotspots for this group of insects.

The Tuomas Holopainen’s species is only one of the eight new flies described in the study. Among them are the Boletina norokorpii fungus gnat, named after Docent Yrjö Norokorpi and known only from Ylitornio; Phronia reducta, which inhabits Salla and Siberia; and Orfelia boreoalpina found in Törmäoja and the German Alps.

The Parks & Wildlife Finland of Metsähallitus is responsible for the management and species surveys of the State’s nature reserves. The collected data is needed in activities such as assessing the status of biodiversity, the protection of species, and planning the management and use of the reserves. Insects are as good an indicator of the state of the natural environment as better-known vertebrates or plants. The diversity of insect species forms part of natural biodiversity and is necessary to human well-being.

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Original source:

Salmela J, Kolcsár L (2017) New and poorly known Palaearctic fungus gnats (Diptera, Sciaroidea). Biodiversity Data Journal 5: e11760. https://doi.org/10.3897/BDJ.5.e11760

Bird spiders detectives: The solution to a 200-year-old hairy mystery

Three species and three genera of birdeater spiders are described as new to science in a paper recently published in the open access journal ZooKeys. In their study, the Brazilian spider experts, Drs. Caroline Fukushima and Rogério Bertani, Laboratory of Ecology and Evolution, Instituto Butantan, report the diversity of the oldest tarantula genus (Avicularia), whose name derives from a famous 18th-century illustration depicting a bird caught by a spider.

Even though these harmless tarantulas have long been a favourite exotic pet around the world, their identity has remained problematic ever since the first species was described back in 1758 by the “father of modern taxonomy”, Carl Linnaeus.

“He described the species based on a hodgepodge of spiders,” explain the authors. “Over the next centuries, other species with completely different characteristics were called Avicularia creating a huge mess.” As a result, basic questions, such as the characteristic traits of the genus, the number of its species and their localities, have been left unanswered.

To address the confusion, the team studied both newly collected specimens and also specimens from around the world which had previously been deposited in museum collections. Thus, they concluded that, instead of the 49 species previously assigned to the problematic genus, there are in fact only 12, including three new to science.

One of the new species, Avicularia merianae, is named after Maria Sybilla Merian (1647-1717), a pioneering scientist and a remarkable artist who made the famous illustration of a spider eating a bird. In fact, it was her work that gave birth to the popular name used for a whole group of spiders, also known as birdeaters or bird spiders.

Meanwhile, the name of the new genus Ybyrapora, which occurs in the Brazilian Atlantic rainforest, translates to “those that live in trees” from the indigenous local language Tupi. It refers to the arboreal habitat of these species. The other two tarantula genera live exclusively on the Caribbean Islands.

“People think all biologists are like Indiana Jones, with their daily lives full of adventures in the wild. But most of the time, they are much more like Sherlock Holmes – sitting on a chair, collecting and analysing clues (specimens and scientific papers) and then using logical reasoning to solve Nature’s mysteries,” comments Dr. Caroline Fukushima.

“We, taxonomists, are ‘wildlife detectives’ who play an essential role not only in biology and conservation. Our work can also become the grounds for new technologies, medicines and ideas that could solve a variety of problems,” she adds.

“We are delighted to have finally closed one of the oldest unsolved cases,” the authors conclude.

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Original source:

Fukushima CS, Bertani R (2017) Taxonomic revision and cladistic analysis of Avicularia Lamarck, 1818 (Araneae, Theraphosidae, Aviculariinae) with description of three new aviculariine genera. ZooKeys 659: 1-185. https://doi.org/10.3897/zookeys.659.10717

New species of parasitic wasp discovered in the eggs of leaf-rolling weevils in Africa

A new species of parasitic wasp has been obtained from the eggs of weevils, associated with bushwillows, collected and identified by Dr. Silvano Biondi. Given the tiny insect from northeastern Gabon is the first record of its genus for West-Central Africa, the researchers Dr. Stefania Laudonia and Dr. Gennaro Viggiani, both affiliated with Italy’s University of Naples Federico II, decided to celebrate it by assigning the species a name that refers to the continent. Their team has published the findings in the open access journal ZooKeys.

Named Poropoea africana, the new species belongs to a large worldwide group of wasps well-known as egg parasitoids of leaf-rolling weevils. Using characteristically long ovipositors, they lay their own eggs in the eggs of the hosts, found in cigar-like rolls.

The new wasp measures less than 2 mm. It can be distinguished from related species by a number of characters, including the structure of the antennae, and the front and hind legs, which are more robust than the middle ones. The latter, which is a unique trait for the genus, seems to be an adaptation to host parasitisation, where the modified legs likely support the body and improve the propulsive efficiency of the ovipositor.

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Original source:

Laudonia S, Viggiani G, Biondi S (2017) A new species of Poropoea Foerster from Africa (Hymenoptera, Chalcidoidea, Trichogrammatidae). ZooKeys 658: 81-87. https://doi.org/10.3897/zookeys.658.11501

Moth gift: Winner of an eBay auction thanks his mother by naming a new species after her

The loving son presented the name to his mother on St. Valentine’s Day

Winner of an eBay auction Steve Mix received the opportunity to pick the name for a new species of satiny-white winged moth collected from the white gypsum dunes of the White Sands National Monument, New Mexico. A fan of butterflies and moths himself, he chose to honor his supportive and encouraging mother Delinda Mix, so the moth is now formally listed under the species name delindae. It is described in the open access journal ZooKeys.

Having spent 10 years studying the moth fauna at the White Sands National Monument, Eric H. Metzler, curator at the Michigan State University, but also research collaborator at the National Museum of Natural History, Smithsonian Institution, and research associate at the University of New Mexico and the University of Florida, discovered the moth during the first year of the study, in 2007. Back then, he spotted a curious small white moth with a satiny appearance, which immediately drew his attention.

Already assigned to the genus Givira to the family commonly known as carpenter millers, the moth was yet to be identified as a species. While most of its North American ‘relatives’ are either dark-colored, or have substantial dark smudges on the forewings, there are only four of them, including the new species, which are substantially white with few or no dark markings.

Further hindrance occurred when the researcher tried to study the specimens, as pinned moths turned out greased due to their abdomens being full of fatty tissue. However, the specialist managed to degrease them by carefully brushing their scales, and, having compared them to related species, confirmed them as representatives of a species new to science.

Then, Eric joined the fundraising event, organized by the Western National Parks Association (WNPA), a non-profit education partner of the US National Park Service. The highest bidder in the eBay auction would receive the chance to pick the scientific name for the satiny-looking moth, and thus, become part of history. Having won the opportunity, Steve Mix, who himself had once been interested in studying butterflies and moths, and has been maintaining his fondness of them ever since, decided to name the species after his mother Delinda Mix, in gratitude for “the support and encouragement she gave to her son”.

“Steve Mix submitted the winning bid, and he chose to have the moth named after his mother because of the lasting nature of this naming opportunity”, shares Eric. “I received no remuneration in this fundraising venture, and by volunteering my personal money, time, expertise, and experience I was able to help WNPA gain world-wide positive publicity while raising some much needed cash. The rewards to me were being able to help WNPA and Steve Mix honor his mother, which is just so very sentimental”.

“WNPA is so pleased that we were able to work with Eric and we are grateful to Steve. This project is a shining example of working together towards the common good of our parks with the added value of providing a priceless experience for everyone involved”, says Amy Reichgott, Development Manager at the Western National Parks Association.

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Original source:

Metzler EH (2017) The Lepidoptera of White Sands National Monument, Otero County, New Mexico, USA 9. A new species of Givira Walker (Cossidae, Hypoptinae) dedicated to Delinda Mix, including a list of species of Cossidae recorded from the Monument. ZooKeys 655: 141-156. https://doi.org/10.3897/zookeys.655.11339

Champions of biodiversity: A weevil genus beats records of explosive evolutive radiation

With as many as 120 recently discovered weevils placed in the genus Laparocerus, it now hosts a total of 237 known species and subspecies. They are all flightless beetles and most of them endemic (living exclusively in one geographic location) to a single island of the archipelagos of Madeira, Selvagens and the Canary Islands (17 islands in total). Only two species inhabit Morocco, the nearest continental land.

Independent Canarian entomologist Dr. Antonio Machado, who has been collecting and studying this genus of weevils for the last sixteen years and researched 46,500 specimens so far, was helped by geneticist Dr. Mariano Hernández, from the University of La Laguna, Tenerife, Canary Islands, Spain, to undertake a phylogenetic study using three mitochondrial genes and one nuclear gene. The resulting phylogenetic tree also allowed for estimating the whole evolutionary process along a timeframe of about 11.2 million years. Their study is published in the open access journal ZooKeys.

The molecular analysis confirms that all Laparocerus weevils have a common evolutionary ancestor (monophyly), but could not clarify whether that ancient founding species arrived from southern Europe or northwestern Africa. The two extant Moroccan species were found to be the result of a back-colonisation from the Canary Islands to Africa, and not the ancestral source lineage, which unfortunately is still unknown.

Colonisation of Macaronesia started in Porto Santo, Madeiran archipelago, which is the oldest island, and from there it ‘jumped’ to Madeira and the Desertas. The colonisation of the Canary Islands started shortly after, and it basically moved stepwise from the east to the west in line with the decreasing age of the volcanic islands. Yet, there have been several back-colonisations, as well (see map). Large islands, such as Tenerife (2034 km2), ended up with 65 species and subspecies. Globally, there is an outstanding ratio of one endemic Laparocerus for each 35.7 km2; a record not beaten by any other genus of plant or animal in Macaronesia.

The evolutionary process responsible for such richness comprises sequential radiation events in these archipelagoes, each generating several monophyletic groups. These groups, 20 in total, have been recognised as subgenera of Laparocerus, and five of them — Aridotrox, Belicarius, Bencomius, Canariotrox, and Purpuranius — are described as new to science in this study. Colonisation routes, habitat shifts, disruption of populations by volcanism, dispersal by massive landslides, and other relevant aspects for adaptive and non-adaptive radiation, are largely discussed and confronted with previously published data referring to other groups of beetles or to other biological organisms (spiders, bush crickets, plants, etc.).

“If oceanic islands have been traditionally considered as laboratories of evolution and species-producing machines, Laparocerus will become the ideal guinea-pig for broadening studies in dispersal and speciation processes of all kinds,” say the authors. “Working with such a group is like getting a picture of Nature with more pixels. Several intriguing cases highlighted in this contribution may turn into the inspiration for further phylogeographic research.”

The scientists hope that, in near future Laparocerus will merit sharing the podium with Darwin´s finches or Drosophila in the studies of island evolution”.

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Original source:

Machado A, Rodríguez-Expósito E, López M, Hernández M (2017) Phylogenetic analysis of the genus Laparocerus, with comments on colonisation and diversification in Macaronesia (Coleoptera, Curculionidae, Entiminae). Zookeys 651: 1-77 (02 Feb 2017) https://doi.org/10.3897/zookeys.651.10097

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.

The 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.

The 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

Chinese scientists discover a new species of catfish in Myanmar

During a survey of the freshwater fishes of the Mali Hka River drainage in the Hponkanrazi Wildlife Sanctuary, Myanmar, scientists Xiao-Yong Chen, Tao Qin and Zhi-Ying Chen, from the Chinese Academy of Sciences (CAS), identified a new catfish species among the collected specimens. It is distinct with a set of morphological features including its mouthparts and coloration. The discovery is published in the open access journal ZooKeys.

The new catfish belongs to a genus (Oreoglanis) of 22 currently recognised species. They are characterised with unusual teeth. While pointed in the upper and the back of the lower jaw, the teeth at the front of the lower jaw are shorter and broad. The latter are placed in a continuous dent. Out of the 22 species of the genus, there are only two known to live in Myanmar.

The new catfish, scientifically named Oreoglanis hponkanensis, has a moderately broad and strongly depressed head and body, and small eyes. The species is predominantly brown in colour, with light yellow belly and several yellowish patches across the body. Noticeable are also two round, bright orange patches in the middle of the fin.

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Original source:

Chen X-Y, Qin T, Chen Z-Y (2017) Oreoglanis hponkanensis, a new sisorid catfish from north Myanmar (Actinopterygii, Sisoridae). ZooKeys 646: 95-108. https://doi.org/10.3897/zookeys.646.11049

New species of moth named in honor of Donald Trump ahead of his swearing-in as president

Days before Donald J. Trump steps forward on the Inaugural platform in Washington to assume the role of the 45th President of the United States of America, evolutionary biologist and systematist Dr. Vazrick Nazari named a new species in his honour. The author, whose publication can be found in the open access journal ZooKeys, hopes that the fame around the new moth will successfully point to the critical need for further conservation efforts for fragile areas such as the habitat of the new species.

While going through material borrowed from the Bohart Museum of Entomology, University of California, Davis, Dr. Vazrick Nazari stumbled across a few specimens that did not match any previously known species. Following thorough analysis of these moths, as well as material from other institutions, the scientist confirmed he had discovered the second species of a genus of twirler moths.

While both species in the genus share a habitat, stretching across the states of California, USA, and Baja California, Mexico, one can easily tell them apart. The new moth, officially described as Neopalpa donaldtrumpi, stands out with yellowish-white scales present on the head in adults. In fact, it was in these scales that the author found an amusing reference to Mr. Trump’s hairstyle and turned it into an additional justification for its name.

Donald Trump’s flying namesake is announced only a month following the recently described species of basslet named after predecessor President Barack Obama, also published in ZooKeys. The fish is only known from coral reefs in the Papahānaumokuākea Marine National Monument, Northwestern Hawaii, a nature reserve which the 44th President of the United States of America expanded to become the largest protected marine area in the world.

Being a substantially urbanized and populated area, the habitat of N. donaldtrumpi is also under serious threat.

“The discovery of this distinct micro-moth in the densely populated and otherwise zoologically well-studied southern California underscores the importance of conservation of the fragile habitats that still contain undescribed and threatened species, and highlights the paucity of interest in species-level taxonomy of smaller faunal elements in North America,” says discoverer Dr. Vazrick Nazari. “By naming this species after the 45th President of the United States, I hope to bring some public attention to, and interest in, the importance of alpha-taxonomy in better understanding the neglected micro-fauna component of the North American biodiversity.”

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Original source:

Nazari V (2017) Review of Neopalpa Povolný, 1998 with description of a new species from California and Baja California, Mexico (Lepidoptera, Gelechiidae). ZooKeys 646: 79-94. https://doi.org/10.3897/zookeys.646.11411

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

Hawaii’s newest species named in honor of President Obama

The new species is the only coral-reef fish that lives exclusively within the marine protected area, which was recently expanded by the President

Scientists from the Bishop Museum, NOAA, and the Association for Marine Exploration published the description of a new species of coral-reef fish that they named in honor of President Barack Obama. The fish, which now bears the formal scientific name Tosanoides obama, was discovered during a June 2016 NOAA expedition to Papahānaumokuākea Marine National Monument in the remote Northwestern Hawaiian Islands. The study is published in the open-access scientific journal ZooKeys.

“We decided to name this fish after President Obama to recognize his efforts to protect and preserve the natural environment, including the expansion of Papahānaumokuākea,” said Richard Pyle, Bishop Museum scientist and lead author of the study. “This expansion adds a layer of protection to one of the last great wilderness areas on Earth.” The Museum is currently showcasing the exhibit Journeys: Heritage of the Northwestern Hawaiian Islands, featuring the Northwestern Hawaiian Islands and the Monument.

On August 26 of this year, at the urging of Sen. Brian Schatz (D-Hawaii), conservationists, and many marine scientists, President Obama expanded Papahānaumokuākea Marine National Monument. At 582,578 square miles, it is the largest permanent marine protected area on Earth. On September 1, during his trip to Midway Atoll within the Monument, legendary scientist, conservationist and deep ocean explorer Dr. Sylvia Earle gave the President a photograph of the fish that now bears his name. The exchange will be featured in the National Geographic global broadcast special, “Sea Of Hope” scheduled to be released on January 15, 2017.

The small pink and yellow fish is a kind of basslet, a group that includes many colorful reef fishes popular in the marine aquarium fish trade. There are two other species in the genus Tosanoides, both from the tropical northwestern Pacific Ocean. Males of the new species have a distinctive spot on the dorsal fin near the tail, which is blue around the edge and red with yellow stripes in the center. “The spot on the males is reminiscent of President Obama’s campaign logo,” said Pyle. “It seemed especially appropriate for a fish named in honor of the president.”

“The new fish is special because it is the only known species of coral-reef fish endemic to the Monument (meaning that the species is found nowhere else on Earth). Our research has documented the highest rate of fish endemism in the world — 100% — living on the deep reefs where we found this new species,” said NOAA scientist Randall Kosaki, chief scientist of the research cruise, and co-author on the paper. However, unlike all the other Hawaiian endemic species, which also occur in the main Hawaiian Islands, this new species is special because it is the only one that is limited to within the Monument itself. “Endemic species are unique contributions to global biodiversity,” Kosaki added. “With the onslaught of climate change, we are at risk of losing some of these undiscovered species before we even know they exist.”

The new fish was first discovered and collected on a dive to 300 feet at Kure Atoll, 1200 miles northwest of Honolulu. Kure is the northernmost of the Hawaiian Islands, and is the highest latitude coral atoll in the world. Deep coral reefs at depths of 150 to 500 feet, in the so-called “Twilight Zone” (also known as mesophotic coral ecosystems), are among the most poorly explored of all marine ecosystems. Located deeper than divers using conventional scuba gear can safely venture, these reefs represent a new frontier for coral-reef research. Pyle and co-authors Brian Greene and Randall Kosaki pioneered the use of advanced mixed-gas diving systems known as closed-circuit rebreathers for Twilight Zone research, and have been documenting the previously unexplored deep reefs throughout Hawai’i and the broader Pacific for the past three decades.

“These deep coral reefs are home to an incredible diversity of fishes, corals, and other marine invertebrates,” said Brian Greene, an experienced deep diver and researcher with the Association for Marine Exploration, and co-author of the paper. “There are many new species still waiting to be discovered down there.”

This is the second new species of fish from Papahānaumokuākea named this year. In August, Pyle and Kosaki published the description of a new species of butterflyfish (Prognathodes basabei) based on specimens collected on deep reefs at Pearl and Hermes Atoll earlier this year. President Obama also has several species from other locales named after him: a trapdoor spider, a speckled freshwater darter (fish), a parasitic hairworm, and an extinct lizard.

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Original source:

Citation: Pyle RL, Greene RD, Kosaki RK (2016) Tosanoides obama, a new basslet (Perciformes, Percoidei, Serranidae) from deep coral reefs in the Northwestern Hawaiian Islands. ZooKeys 641: 165-181. https://doi.org/10.3897/zookeys.641.11500