The story began in 2008, when surveyors captured a single distinctive vole in Lang County, on the right bank of the middle reaches of the Yarlung Zangbo River. The specimen was initially identified as A. stracheyi, a name later synonymized under A. stoliczkanus in a broad taxonomic revision of the genus. But the oddities never quite fit. That earlier revision had already flagged five different molar patterns within A. stoliczkanus and reported a surprisingly large genetic gap, roughly 4.3 percent at the cytochrome b gene, between Tibetan populations from Lang and Pulan counties. When eight more specimens turned up in 2023 from Lang and Longzi counties, closely resembling the 2008 animal and forming a single genetic lineage with it, the researchers decided the identification as A. stoliczkanus was untenable.

To test whether the vole represented something genuinely new, the team assembled an unusually broad evidence base. They sampled 93 specimens across much of the genus’s range, spanning Russia, Mongolia, Pakistan, Kazakhstan and northwestern China, and generated sequences from three mitochondrial genes, cytochrome b, cytochrome c oxidase subunit I and NADH dehydrogenase subunit 2, plus four nuclear genes, GHR, LCAT, BRCA1 and IRBP. Separate analyses of mitochondrial data, nuclear data, and the combined dataset all converged on the same result: the five sequenced individuals of the putative new species form a well-supported clade that is sister to A. stoliczkanus, meaning the two share an immediate common ancestor unique to them.

Three independent species delimitation methods, Bayesian assignment in BPP, automatic partitioning with ASAP, and the multi-rate PTP model, consistently treated the lineage as an independent species, recognizing 12, 11 and 14 candidate species within Alticola respectively. The genetic distances tell a striking story: Kimura-2-parameter distances at cytochrome b between the new vole and its eleven named relatives range from 4.0 percent against A. stoliczkanus to 10.3 percent against A. macrotis, averaging 8.1 percent. That 4.0 percent minimum is comparable to distances between other recognized species pairs in the genus, such as A. olchonensis and A. tuvinicus at 4.1 percent and A. argentatus and A. albicauda at 4.7 percent.

Dating the split with a relaxed molecular clock calibrated against the oldest fossils of Alticola and its sister genus Myodes revealed that the new species diverged from A. stoliczkanus roughly 490,000 years ago, with a 95 percent credible interval of 200,000 to 770,000 years. That makes it the youngest divergence node in the genus’s reconstructed tree, a product of recent Pleistocene speciation on a plateau whose complex topography and climatic history have repeatedly isolated small mammal populations in valleys and along river drainages.

Morphology backs up the molecules. The new vole carries the simplest third upper molar in the entire genus, marked by a distinctly truncated posterior lobe, a feature even simpler than that of A. stoliczkanus, which was previously considered the most reduced. It also has a slender, sparsely haired tail about a quarter of its head-body length, darker brown dorsal fur than any of its close relatives, and a body size larger than A. stoliczkanus but smaller than many other congeners. Factor analysis of sixteen external and cranial measurements from 67 intact specimens separated the new species from six of the seven other Alticola species occurring in China, with only A. macrotis overlapping heavily in overall morphospace, though post-hoc tests confirmed significant differences in head-body length, tail length and nasal length between the two.

Perhaps the most unexpected evidence came from reproductive anatomy, a classic but rarely deployed tool in microtine systematics. The glans penis and baculum of the new species differ markedly from both A. stoliczkanus and A. macrotis. Where A. stoliczkanus bears a single outer crater papilla on each side and A. macrotis has none, the new vole has four to five. Its proximal baculum has a semicircular base rather than a slightly enlarged one, its distal and lateral bacula are slender with a sharp tip on the lateral element, its urethral lappet forks into two long, deeply bifurcated branches, and its dorsal papilla carries a distinctive digitiform projection. In mammals, such differences in genital morphology often signal reproductive isolation, reinforcing the case that this is a genuinely separate evolutionary lineage rather than a geographic variant.

The holotype, an adult male weighing 29 grams with a head-body length of 104 millimeters, was trapped in August 2023 beside the Naijiaxiongqu stream in Longzi County at 4,022 meters elevation. All nine known specimens, four males and five females, came from riparian shrubland between 3,228 and 4,022 meters in Longzi and Lang counties, an habitat distinctly lower and lusher than the stony, glacial-moraine slopes above the timber line where A. stoliczkanus makes its home at 3,900 to 5,450 meters. The species epithet yarlungia honors the Yarlung Zangbo River, Tibet’s largest, whose basin drains the entire known range of the animal.

The discovery carries a broader message about how much life remains undocumented even in well-studied groups. Mountain voles are not obscure; they are widespread across Central Asian mountains and have been described and revised repeatedly since the nineteenth century. Yet the genus’s remote, high-elevation habitats have limited comprehensive sampling, particularly in China, where rugged terrain and poor transport infrastructure have left parts of the Yarlung Zangbo basin essentially unsurveyed. The researchers suggest that species diversity within Alticola remains underestimated and that the new vole likely also occupies the southern slopes of the eastern Himalayas, given the continuity of habitat across national borders.

For biodiversity science, the case of Alticola yarlungia is a textbook example of integrative taxonomy done right. No single line of evidence, geography, genetics, molar shape, pelage, or genital anatomy, would have been sufficient on its own to convince a skeptical reviewer. Together, they establish a species that diverged from its closest relative in the Middle Pleistocene and has since persisted, unnoticed, in the streamside shrubs of one of the planet’s most biologically productive and least explored highland regions. As surveying intensifies across the eastern Himalayan hotspot, more such hidden vertebrates are likely to come to light.

Subject of Research: A newly described species of mountain vole from the Yarlung Zangbo River basin in southern Tibet

Article Title: A new species of Alticola ( Mammalia , Rodentia , Arvicolinae ) from southern Tibet, China

Article References: Tang, M. K., Qiu, H., Peng, B. Q., Liao, R., Murphy, R. W., Wang, X. M., Pan, X., Wang, X., & Liu, S. Y. (2025). A new species of Alticola (Mammalia, Rodentia, Arvicolinae) from southern Tibet, China. Zoosystematics and Evolution, 101(4), 2387-2403. https://doi.org/10.3897/zse.101.170704

Image Credits: AI Generated

DOI: 10.3897/zse.101.170704

Keywords: Alticola yarlungia, mountain vole, Tibet, new species, integrative taxonomy, Qinghai-Tibet Plateau, rodents, phylogenetics, Yarlung Zangbo River, species delimitation, cytochrome b, Himalayas

The genus Glyptothorax is one of the most successful groups of small catfishes in Asia, with a range stretching from the Euphrates and Tigris river basins in Turkey in the west all the way to the Yangtze River in China in the east, and southwards to Borneo and Java. It is the most species-rich genus in the family Sisoridae, and its members share one of the most remarkable adaptations in freshwater fish biology: a thoracic adhesive apparatus, a patch of longitudinal or oblique skin folds arranged in an roughly elliptical area on the underside of the chest. This natural suction cup allows the fish to cling to rocks and gravel in fast-flowing hill streams, holding its position against currents that would sweep away less specialised fish.

China is a stronghold for the genus, with 27 documented species as of 2025. Most of these are concentrated in the south-west of the country, in the great river systems of the Brahmaputra, Irrawaddy, Nujiang-Salween, Lancangjiang-Mekong and Red River. Only four species were previously known from the vast area east of the Red River, encompassing the Pearl River, the Yangtze River, Hainan Island and the south-eastern coastal drainages. It was in this comparatively well-surveyed region that the new catfish turned up, a reminder that even familiar waters can still conceal undocumented biodiversity.

Between 2023 and 2024, researchers collected specimens from Yunnan and Guangxi, preserving them initially in 10 percent formalin before transferring them to 75 percent ethanol for long-term storage. Small clips of right pelvic fin tissue were kept in pure alcohol for genetic work. The team assembled a type series of 28 specimens, with the holotype, an 83.5 millimetre individual from Bamei Town in Guangnan County, and 17 paratypes deposited at the Kunming Zoological Museum of the Kunming Institute of Zoology, Chinese Academy of Sciences, and a further 10 paratypes held at the Fisheries Administration of Qiubei in Yunnan. The paratypes range from 47.1 to 106.4 millimetres in standard length and come from three localities: the Tuoniang River at Bamei, the Xiyang River at Banbang Township, and the Leli River in Tianlin County, Guangxi.

Morphologically, G. pulcher most closely resembles G. quadriocellatus, sharing a similar outline, skin tubercles and general colour pattern. Yet a careful combination of traits separates it from all of its relatives east of the Red River. The new species has a distinctive dark body marked with bright yellow spots and stripes, sparse soft tubercles on the skin that feel similar in texture to the surrounding tissue, and a smooth dorsal surface on the head. Its thoracic adhesive apparatus is unusually broad, with a length of just 1 to 1.2 times its width, and the apparatus spans 12.0 to 14.3 percent of standard length. The adipose fin base is short, measuring 11.7 to 15.4 percent of standard length, and the caudal peduncle, the narrow stalk preceding the tail fin, is notably deep and short, at 13.7 to 17.5 percent of standard length in depth terms and 10.3 to 12.3 percent in depth.

Compared specifically with G. quadriocellatus, the new species lacks the yellow spot behind the eye that its relative possesses, has smooth rather than tubercled skin on top of the head, a broader adhesive apparatus with a width of 12.0 to 14.3 percent of standard length versus 9.4 to 11.1 percent, no anteromedial striae on the adhesive disc, and a shorter adipose fin base. Against G. interspinalum, it again shows the broader adhesive disc and a shorter, deeper caudal peduncle. These measurements, taken with micrometer calipers to the nearest 0.1 millimetre and expressed as proportions of standard length and head length, follow established taxonomic conventions for the genus, and fin ray and vertebral counts were verified using X-ray imaging.

The colour pattern alone makes the fish unmistakable. In preserved specimens the head and body are brownish-black, with a light yellow ventral surface and adhesive apparatus. A yellow mid-dorsal stripe runs along the back, and yellow spots mark the dorsal fin origin, sometimes merging into a saddle patch. All of the fins are basically black with yellow margins: the dorsal fin carries a C-shaped yellow band, the paired and anal fins have irregular pincer-like yellow edges, and the caudal fin bears a large black dumbbell-shaped patch at its base with an M-shaped yellow posterior margin. In life, the brownish-black areas shift to a light chestnut colour and the adhesive apparatus turns pale pink, a living palette that justified the species name.

To test whether the morphology pointed to a genuinely distinct species, the team turned to DNA. They generated eleven new sequences of the mitochondrial cytochrome c oxidase subunit I gene, COI, and nine sequences of the cytochrome b gene from seven Glyptothorax species, adding eight previously published sequences to build a dataset of nineteen COI and seventeen Cyt b sequences. Two outgroup species, Bagarius protos and Pseudecheneis brachyura, were included to root the tree. Phylogenies were reconstructed with both Maximum Likelihood, using IQ-TREE with ultrafast bootstrap support over 1,000 iterations, and Bayesian Inference, using MrBayes with two independent runs of Markov Chain Monte Carlo for 10 million generations, with convergence confirmed when the average standard deviation of split frequencies fell below 0.01.

Both methods recovered the same topology and delivered a clear verdict. The new species forms a strongly supported monophyletic group, and within the broader tree it sits as the sister lineage to G. pallozonum, another Pearl River inhabitant. The two species fall within a clade of dark-bodied fish decorated with creamy or bright yellow spots and stripes, distinct from a second clade of species bearing black saddle-like patches. Kimura 2-parameter genetic distances between G. pulcher and G. pallozonum, based on the COI gene, range from 5.7 to 6.0 percent, a level of divergence consistent with separate species status in this group. The phylogeny also matched the patterns reported by earlier studies of the genus, reinforcing confidence in the result.

The study produced a second, equally consequential finding: a taxonomic revision that removes a name from the books. Glyptothorax zhujiangensis, described in 2003 from the Pearl River Delta, has long been treated as valid, but its type specimens have been lost. The holotype and nine paratypes once held at the Institute of Hydrobiology in Beijing cannot be found, the ten remaining paratypes were reportedly kept at an institution in Zhuhai that apparently never existed, and no photographs survive. The team collected fresh specimens matching the original description from the Pearl River Delta, roughly 50 kilometres from the type locality, and found that these fish nest firmly within G. pallozonum in the phylogenetic tree, with COI distances of just 0.3 to 1.0 percent. Given the overlapping morphology, the minimal genetic separation and the fact that G. pallozonum’s known range already includes the type locality of G. zhujiangensis, the authors conclude that the two are conspecific and that G. zhujiangensis should be regarded as a junior synonym of G. pallozonum, with the apparent differences likely reflecting populational variation, measurement error and the small size of the original type series.

For now, G. pulcher is known only from the Tuoniang River basin, a sub-basin of the upper Pearl River in Yunnan and Guangxi, and its discovery hints that the hill streams of southern China may hold more surprises. The combination of classical morphology, X-ray verified anatomy and mitochondrial genomics used here offers a template for untangling the region’s catfish diversity, and the loss of the G. zhujiangensis type material stands as a cautionary tale about the irreplaceable value of museum specimens. As researchers continue to survey the upper Pearl River’s rocky, fast-flowing tributaries, the beautiful new catfish serves as both a celebration of hidden biodiversity and an argument for protecting the clear hill streams that such specialised species depend on.

Subject of Research: Description of a new sisorid catfish species, Glyptothorax pulcher, from the upper Pearl River basin in China, with molecular phylogenetic analysis and taxonomic revision

Article Title:  Glyptothorax pulcher ( Siluriformes , Sisoridae ), a new species from the upper Pearl River of Yunnan and Guangxi, southwest China

Article References: Zeng, Y.-Y., Lin, F., Yang, H.-F., Pu, X.-R., Jiang, W.-S., & Chen, X.-Y. (2026). Glyptothorax pulcher (Siluriformes, Sisoridae), a new species from the upper Pearl River of Yunnan and Guangxi, southwest China. Zoosystematics and Evolution, 102(1), 17-26. https://doi.org/10.3897/zse.102.167142

Image Credits: AI Generated

DOI: 10.3897/zse.102.167142

Keywords: Glyptothorax pulcher, new species, catfish, Sisoridae, Pearl River, Yunnan, Guangxi, taxonomy, molecular phylogeny, COI, cytochrome b, freshwater biodiversity

The holotype, an adult male measuring 29.3 millimetres in standard length, was collected on 22 December 2016 in Yaw Stream, a tributary of the Yebya River near Pakokku City in Myanmar’s Magway Region. Over the following years, the team gathered additional specimens from several widely separated localities, including Myan Aww Stream inside Alaungdaw Kathapa National Park, the Salin Stream near Sinbyugyun, the lower Yemar River near Monywa, Nam Sa Pi Stream at Htamanthi, and even a stream flowing into Indawgyi Lake in Kachin State in the far north. That broad spread across the Chindwin–Irrawaddy drainage suggests the species may eventually be found throughout the tributaries of central Myanmar, particularly in the dry zones of the Sagaing, Mandalay, and Magway regions.

What makes Mustura arenaria recognisable is a distinctive combination of features rather than any single dramatic trait. The body carries 7 to 15 vertically elongated dark blotches along the flank, alternating with 9 to 17 dorsal saddles, and crucially the blotches are distinctly narrower than the pale spaces between them. Males and females differ in the shape of these markings, with the blotches more rounded in males and more elongated in females. The species also shows 8½ branched dorsal-fin rays, a forked tail with 8+8 branched caudal-fin rays, a complete lateral line, an axillary pelvic lobe, and a small notch in the upper lip. Two dark stripes running from the tip of the snout toward the front of the eyes, together with a pair of faint blotches between the nostrils, form an X-shaped pattern on the head, while a pair of blotches around the fontanelle creates a triangular mark extending to the nape.

Sexual dimorphism adds another layer of identification. Males possess a longer suborbital flap, a fleshy structure on the cheek studded with small tubercles, and their first branched pectoral-fin ray is broadened and angled slightly upward, with the branches pressed together and lacking a membrane except near the tip. Females show only a small flap, a groove, or nothing at all in that position. Males also carry more developed tubercles on the snout and head. These traits align the new species with the diagnostic profile of Mustura as defined by Swiss ichthyologist Maurice Kottelat when he established the genus in 2018, designating Mustura celata as its type species.

To test whether the morphology told the true evolutionary story, the researchers turned to genetics. They extracted DNA from fin tissue and sequenced two markers: the mitochondrial cytochrome b gene, a workhorse of fish phylogenetics, and the nuclear recombination-activating gene RAG1. The dataset ultimately included 57 Cyt b and 54 RAG1 sequences, with three species from other genera serving as outgroups. After alignment with MAFFT and trimming with Gblocks, the team built phylogenetic trees using both maximum likelihood methods in IQ-TREE, with 10,000 ultrafast bootstrap replicates, and Bayesian inference in MrBayes, running four Markov chain Monte Carlo chains for 10 million generations. An incongruence length difference test confirmed that the two genes did not significantly conflict, with a p-value of 0.335, allowing a combined analysis.

The genetic results were unambiguous about the new species itself. Mustura arenaria formed a well-supported, monophyletic lineage, and its pairwise Kimura two-parameter genetic distance from congeners, based on Cyt b, ranged from 13.49 to 16.16 percent, a deep divergence by any standard for closely related fish species. Within the species, sampled individuals differed by only 1.19 percent on average. For comparison, the smallest interspecific distance recorded in the study, 9.46 percent between Mustura sp. UF192977 and M. yangi, was still far above the new species’ internal variation, reinforcing its status as an independently evolving lineage.

But the wider picture was far messier. Across all analyses, Mustura as currently defined came out polyphyletic, meaning its member species do not all share a single recent common ancestor exclusive of other genera. The type species M. celata clustered with M. maepaiensis, M. arenaria, M. yangi, and two unassigned taxa, forming a clade that is sister to the genus Neonoemacheilus. The remaining species scattered into Schistura, Physoschistura, and other lineages. In other words, the genus is a morphological convenience rather than an evolutionary reality, a conclusion that echoes several recent phylogenetic studies of nemacheilid loaches.

The study also settled a long-running identity question for one particular fish. Mustura shanensis, first described in 1929 as Nemacheilus shanensis, has bounced between genera for decades, variously placed in Physoschistura, tentatively assigned to the Schistura rupecula group, and moved to Mustura on the basis of colour pattern and caudal-fin shape. Fresh examination of specimens from Inle Lake in Myanmar’s Shan State revealed that it lacks the sexual dimorphism typical of Mustura, has an air bladder divided into separate halves with a distinct oval posterior chamber, and possesses a lower lip broadly interrupted without the triangular cushion-like pads characteristic of true Mustura. Molecular data placed it firmly as a sister lineage to Schistura species, with maximal support values, and the authors formally transferred it to that genus as Schistura shanensis.

The deeper implication is that convergent evolution may be muddying loach classification across South and Southeast Asia. Species grouped in Mustura and related genera such as Pteronemacheilus, Physoschistura, and Petruichthys share features like mouth structure and male pectoral-fin modifications, yet the DNA indicates clear genetic divergence among these groups. Some of the characters long used to define genera may be less reliable than assumed. The authors note that air bladder variation has been linked to water velocity in the fish’s environment, and that preservation methods can make the small posterior chamber difficult to identify, while the rigid pectoral-fin rays of males appear only probabilistically across populations. Such traits may matter at the species level but must be applied cautiously when drawing genus boundaries.

For now, Mustura arenaria takes its place among the flowing, sandy-bottomed streams of the Irrawaddy basin, living in small aggregations alongside eels, loaches, and carps including Anguilla bengalensis, Pangio signicauda, Schistura callidora, and several species of Glyptothorax. Its description brings the count of recognised Mustura species, broadly defined, to 23, but the authors are clear that the number is a moving target. Several species provisionally assigned to the genus, including M. bella, M. geisleri, M. isostigma, and M. shuangjiangensis, appear genetically closer to other lineages, and one has already been moved to a newly erected genus. A comprehensive revision of Mustura and its relatives, combining broader molecular sampling with systematic morphological comparison, is now clearly needed. In the sandy shallows of Myanmar, a small fish has made a large case that the loach family tree still needs redrawing.

Subject of Research: Description of a new nemacheilid loach species from Myanmar and taxonomic revision of the genus Mustura

Article Title:  Mustura arenaria , a new species of genus Mustura ( Cypriniformes , Nemacheilidae ) from Myanmar, with taxonomic notes on Mustura shanensis

Article References: Lyu, J.-B., Lin, F., Oo, T. N., & Chen, X.-Y. (2026). Mustura arenaria, a new species of genus Mustura (Cypriniformes, Nemacheilidae) from Myanmar, with taxonomic notes on Mustura shanensis. Zoosystematics and Evolution, 102(1), 107-117. https://doi.org/10.3897/zse.102.170882

Image Credits: AI Generated

DOI: 10.3897/zse.102.170882

Keywords: Mustura arenaria, new species, Nemacheilidae, Myanmar, Irrawaddy basin, loach taxonomy, phylogenetics, cytochrome b, RAG1, Schistura shanensis, polyphyly, integrative taxonomy

The story begins with a taxonomic puzzle that has lingered since the nineteenth century. Schizopygopsis, a genus of highly specialized schizothoracine carps described by Steindachner in 1866, comprises scale-less, barbel-less fish with inferior mouths and sharp horny layers on their lower jaws, adaptations suited to scraping food from rocky riverbeds. Different authorities have recognized anywhere from nine to twelve species and subspecies, and the boundaries between them have been contested for decades. Schizopygopsis malacanthus, described by Herzenstein in 1891 from the Tongtian River headwaters of the Jinsha River, was thought to range widely across the upper Jinsha and Yalong Rivers, with populations in the two drainages treated merely as geographic variants of a single, broadly distributed species.

That assumption has now collapsed under the weight of integrative evidence. A research team from the Northwest Institute of Plateau Biology at the Chinese Academy of Sciences collected 41 specimens from the Yalong River and 20 from the Jinsha River system between 2021 and 2023, along with 29 samples of the coexisting species Herzensteinia microcephalus, which some classifications have historically placed within Schizopygopsis itself. The specimens were euthanized with MS222, preserved in 95 percent ethanol, and deposited as voucher specimens, providing a permanent physical record that other researchers can re-examine.

The morphological analysis went well beyond traditional external measurements. The team recorded seventeen body dimensions with digital callipers, counted the inner and outer gill rakers on the first gill arch, and used micro-computed tomography to count and image vertebrae non-destructively. Principal component analysis of the body proportions showed something intriguing: by overall body shape alone, the Yalong River fish and the Jinsha River fish were essentially indistinguishable, their measurements overlapping almost completely. If the researchers had stopped there, the case for a new species would have been weak. But the feeding apparatus told a very different story.

The Yalong fish carries 15 to 21 outer gill rakers and 25 to 39 inner gill rakers on its first gill arch, noticeably denser filters than those of its Jinsha River relative. More striking still are the pharyngeal teeth, the grinding structures that cyprinid fish use to process food. In the Yalong population, the pharyngeal bone is hooked and the teeth terminate in oblique-truncated tips with shovel-shaped grinding surfaces, whereas the Jinsha River fish has hooked teeth with concave grinding surfaces. These are not trivial differences. Pharyngeal tooth morphology is a cornerstone character in cyprinid taxonomy, and tooth shape tracks diet: hooked, grasping teeth typically belong to omnivores handling varied food items, while shovel-shaped teeth appear in fish that crush benthic invertebrates scraped from the river bottom.

The molecular evidence sealed the case. The team sequenced the mitochondrial cytochrome b gene, a 1,140-base-pair workhorse of fish phylogenetics, from all 75 newly collected specimens, generating 46 haplotypes, and combined these with GenBank sequences from seven other Schizopygopsis species plus five outgroup species. Genetic distances between the Yalong population and other Schizopygopsis species ranged from 2.03 to 6.16 percent, averaging 4.15 percent. The distance between the Yalong fish and S. malacanthus proper was 3.605 percent, which is larger than the distances separating several pairs of species that taxonomists already treat as fully valid, such as S. kessleri and S. pylzovi, which differ by a mere 0.132 percent. Genetic differentiation measured by Fst reached 0.5276 between the two populations, a level indicating profoundly restricted gene flow.

Perhaps the most surprising result came from the phylogenetic trees themselves. Both maximum likelihood and Bayesian inference analyses produced identical topologies, and they showed that the Yalong fish does not form the closest sister relationship with S. malacanthus at all. Instead, the strongly supported monophyletic Yalong lineage groups with S. kialingensis from the Jialing River, a distant tributary of the Yangtze, and then clusters with S. kessleri, S. pylzovi, and H. microcephalus. Earlier studies had concluded the Yalong and Jinsha populations were sister lineages, but only because those analyses omitted most of the genus. When all ten recognized species were finally included, the true picture emerged. Species delimitation methods, the Bayesian Poisson tree processes model and the automatic partitioning algorithm ASAP, both independently confirmed the Yalong fish as a distinct species, with bPTP assigning a Bayesian support value of 0.84.

The geographic context explains how such a split could occur. The dramatic uplift of the Qinghai-Tibetan Plateau in the late Pliocene reorganized the region’s drainage systems, isolating the Tongtian and Yalong Rivers from one another. Cut off from gene flow, the ancestral fish in each basin followed independent evolutionary trajectories over millions of years. Natural selection then sculpted their feeding machinery along divergent paths, likely in response to different food resources in the two rivers, a hypothesis supported by previous findings that the two populations harbor distinct gut microbial communities at the phylum, class, and genus levels. The result is two fish that look like twins in body plan but differ fundamentally in the tools they use to eat.

The formal description rests on a holotype collected in July 2023 at Zhenqin Village in Chenduo County, Qinghai Province, at an altitude of 4,172 meters, with 40 paratypes ranging from 65 to 196 millimeters in standard length. The species name yalongensis honors the Yalong River, its only known home. There, the fish inhabits cold streams with beds of sand, gravel, and pebbles, sharing the water with Gymnodiptychus pachycheilus and several species of stone loaches in the genus Triplophysa. In life, the fish displays a yellowish-brown back and head, a white belly, and blackish-grey spots scattered across the back, head, and dorsal fin.

The discovery carries weight beyond taxonomy. Fish diversity in the Yalong River is reportedly lower than in other Jinsha tributaries, so finding a species that exists nowhere else underscores how incomplete our inventories of plateau biodiversity remain, and it suggests that other cryptic species may be waiting in the Qinghai-Tibetan Plateau’s river systems. It also has conservation implications: S. malacanthus is considered vulnerable because of its slow growth, low reproduction rate, and restricted distribution, and the new species likely shares those biological traits. The authors argue that S. yalongensis warrants the same level of protection. A fish that survived Pliocene mountain-building and millions of years of isolation in glacial meltwater now faces a far more recent threat, and recognizing its existence is the first step toward safeguarding it.

Subject of Research: Description of a new fish species, Schizopygopsis yalongensis, from the Yalong River on the Qinghai-Tibetan Plateau using integrative taxonomy

Article Title: A new species of Schizopygopsis Steindachner, 1866 ( Cypriniformes , Cyprinidae ) from the Yalong River on the Qinghai-Tibetan Plateau, China

Article References: Zhao, K., Liu, S., Chen, S., & Tian, F. (2026). A new species of Schizopygopsis Steindachner, 1866 (Cypriniformes, Cyprinidae) from the Yalong River on the Qinghai-Tibetan Plateau, China. Zoosystematics and Evolution, 102(1), 131-144. https://doi.org/10.3897/zse.102.173298

Image Credits: AI Generated

DOI: 10.3897/zse.102.173298

Keywords: Schizopygopsis yalongensis, new species, Qinghai-Tibetan Plateau, integrative taxonomy, cytochrome b, phylogenetics, pharyngeal teeth, Yalong River, cryptic species, Cyprinidae, species delimitation, conservation

The keelback genus Hebius is the most species-rich lineage within the natricid snakes, a family of mostly semi-aquatic or terrestrial serpents spread across East and Southeast Asia. These are small to medium-sized animals with elongated bodies, a weak but visible boundary between head and neck, large eyes with round pupils, and rear maxillary teeth that grow progressively larger toward the back of the jaw. Their conservatism in outward appearance, combined with considerable variation within populations, has made species-level taxonomy notoriously difficult. Recent integrative analyses that merge morphology with molecular data have repeatedly suggested that the current count of 53 recognized species, 30 of them recorded from China and 11 from Vietnam, substantially underestimates the true diversity, with numerous cryptic lineages still awaiting formal recognition.

The Tonkin keelback itself has a tangled history. René Léon Bourret described it in 1934 as Natrix atemporalis, based on three specimens from Tonkin, now northern Vietnam, and its most striking feature was the absence of temporal scales on the sides of the head, a trait almost unique among its relatives. In 1969, Edmond Malnate and John Romer examined additional material from Hong Kong and confirmed that those snakes matched Bourret’s originals from what is now Tam Dao National Park in Phu Tho Province, though no formal restriction of the type locality was ever made. The species drifted through several genus names over the decades, from Natrix to Amphiesma, until molecular phylogenetic work in 2014 settled it firmly within Hebius. Throughout all of this shuffling, the snake was reported across northern Vietnam and southern China, from Guangdong and Guangxi to Guizhou, Yunnan, Hunan, Hubei, and Chongqing, giving it the appearance of a wide-ranging generalist.

Suspicion that this apparent range concealed more than one species had been building. Earlier molecular studies had already revealed two deeply divergent clades within what was then called Hebius atemporalis, hinting that the name might be papering over real biological boundaries. The challenge was that the morphological differences between populations were subtle, sampling was limited, and no comprehensive revision had ever been attempted. To resolve the problem, the research team assembled 27 preserved specimens of the species complex, including two syntypes held at the National Museum of Natural History in Paris, and supplemented them with 12 newly collected individuals from fieldwork conducted across Vietnam and China between 2020 and 2025.

On the molecular side, the team extracted genomic DNA from muscle or liver tissue and amplified a 1,098-base-pair fragment of the mitochondrial cytochrome b gene, a standard workhorse for snake phylogenetics. Sequences were aligned and uncorrected pairwise genetic distances calculated, and the resulting dataset was analyzed with both Maximum Likelihood and Bayesian Inference methods, using 36 additional Hebius sequences and two outgroup taxa from GenBank to place the findings in context. Both approaches produced largely congruent trees. The species complex emerged as a strongly supported monophyletic group, and within it, the population from the Yunnan-Guizhou Plateau stood out as a highly divergent lineage, separated from true Hebius atemporalis by uncorrected p-distances of 8.0 to 9.8 percent. For comparison, genetic divergence within the plateau population itself was less than 1 percent, while Chinese and Vietnamese populations of the true species differed by 6.4 to 7.0 percent, a level that itself approaches thresholds commonly used to infer species-level separation in natricine snakes.

The morphological evidence proved just as decisive. The plateau snakes are larger as adults, reaching a maximum snout-vent length of 416 millimeters against 294 millimeters for true Hebius atemporalis. They have proportionally shorter tails, with the tail making up 0.22 to 0.27 of total length compared with 0.28 to 0.33 in their Vietnamese and eastern Chinese relatives. They carry more ventral scales, 146 to 156 against 123 to 140, and correspondingly higher combined ventral and subcaudal counts. Most visually striking of all, they lack the white to orangish nuchal collar that gives the Tonkin keelback its characteristic neck band, and their dorsal coloration shifts from brown to dark olive rather than from reddish brown to blackish brown. Together with the shared absence of temporal scales, these characters set the plateau population apart not only from Hebius atemporalis but from all 37 other congeners recorded in China and the Indo-Burma region.

The new species has been named Hebius malnatei, honoring Edmond V. Malnate, the American herpetologist who first revised the taxonomic placement of the Tonkin keelback. The description is anchored by an adult female holotype collected in May 2024 at Pugu Village in Panzhou County, Guizhou Province, at an elevation of 1,750 meters, with five paratypes from Yunnan and Guizhou. The species is known from across the Yunnan-Guizhou Plateau at elevations of 1,750 to 1,950 meters, inhabiting forests, shrubland, and meadows near ponds and streams. It is active by day and night, peaking around sunset, and when captured it does not bite but instead releases a strong, fetid secretion from its cloacal glands. Earthworms are the only recorded prey, and the holotype contained five well-developed oviductal eggs, indicating oviparity and a late-spring breeding season. Based on IUCN Red List criteria, the authors recommend assigning it to Least Concern, though its restricted elevational range and specialized habitat association warrant targeted surveys and long-term monitoring.

True Hebius atemporalis, meanwhile, has received a long-overdue redescription based on its syntypes and additional material. It is now confirmed from northern Vietnam, including Phu Tho, Tuyen Quang, Dien Bien, and Nghe An provinces, and from southern and central China, including Hong Kong, Guangdong, Guangxi, Guizhou, Hunan, Hubei, and Chongqing, at elevations of 400 to 1,700 meters. It is a diurnal snake of subtropical montane forests that tolerates both primary and secondary habitats, from dense evergreen shrubland to comparatively dry hilltops without streams. When disturbed it moves rapidly and emits a strong, unpleasant odor, and like its newly recognized relative it is presumed to feed primarily on earthworms. Several historical records, including a report from Laos, remain unverified for lack of voucher specimens.

Questions remain. Three museum specimens from Yunnan collected decades ago cannot yet be assigned with confidence, and a short, potentially contaminated cytochrome b sequence attributed to the species from Yunnan was excluded from the analysis. The 6.4 to 7.0 percent genetic divergence between Chinese and Vietnamese populations of true Hebius atemporalis, together with roughly 4 percent divergence between Guizhou and Guangdong populations within China, suggests that further cryptic diversity may still be lurking within the complex. The authors hypothesize that the species’ relatively fossorial habits may restrict dispersal and promote such population structuring, a premise that broader geographic sampling and multi-locus genomic datasets will be needed to test.

Perhaps the most evocative finding is biogeographic. The estimated divergence between Hebius malnatei and Hebius atemporalis, dated to roughly 6.8 million years ago, coincides closely with major uplift phases of the Yunnan-Guizhou Plateau. This supports a scenario of allopatric speciation driven by geological and climatic processes that carved the region into a mosaic of topographic barriers and montane refugia. The plateau is increasingly recognized as a hotspot for cryptic amphibian and reptile diversity, with numerous lineages of frogs and snakes showing similar patterns of local endemism. The discovery of Malnate’s keelback reinforces the region’s status as both a cradle of herpetofaunal speciation and a priority area for biodiversity research and conservation, and it serves as a reminder that even familiar, wide-ranging species can conceal evolutionary stories written in stone.

Subject of Research: Taxonomic revision of the Hebius atemporalis snake complex and description of a new keelback species from China

Article Title: Taxonomic reassessment of Hebius atemporalis (Bourret, 1934) ( Squamata , Natricidae ), with the description of a new species from China

Article References: Li, M.-L., Nguyen, T. V., Huang, J.-J., Idiiatullina, S. S., Le, S. X., Poyarkov, N. A., David, P., Han, B.-Y., Wu, W., Li, J.-T., & Ren, J.-L. (2026). Taxonomic reassessment of Hebius atemporalis (Bourret, 1934) (Squamata, Natricidae), with the description of a new species from China. Zoosystematics and Evolution, 102(1), 235-251. https://doi.org/10.3897/zse.102.176342

Image Credits: AI Generated

DOI: 10.3897/zse.102.176342

Keywords: Hebius malnatei, Hebius atemporalis, keelback snakes, Natricidae, Yunnan-Guizhou Plateau, integrative taxonomy, cytochrome b, cryptic species, herpetology, China, Vietnam, species delimitation

The specimen was captured in July 2025 during a series of chiropteran surveys in southwestern Yunnan. Sweep netting at the eaves of the management station building within the Jinguang Temple Provincial Nature Reserve in Dali, at coordinates 25°11’55″N, 99°31’50″E and an elevation of roughly 2,500 metres, yielded one adult female, catalogued as GZHU 25209. Field protocols complied with Chinese law and live-animal handling followed American Society of Mammalogists guidelines. The voucher specimen, preserved in 75 percent ethanol, is now deposited at the South China Biodiversity Research Center at Guangzhou University. Adult status was confirmed by the degree of epiphyseal-diaphyseal fusion in the bones, a standard ageing method in bats.

Identifying the animal required an integrative approach combining morphology, morphometrics and genetics. The team took external measurements including body length, tail length, ear length, hind foot length, forearm length and tibia length, plus twelve craniodental dimensions measured with digital calipers accurate to 0.01 millimetres. The right wing was pinned flat, photographed perpendicular to the plane, and analysed with BAT-WINGAREA software to derive total wing area, wingspan and membrane areas, from which the three primary airfoil parameters were calculated. For the genetic component, total DNA was extracted and the mitochondrial cytochrome b gene, a 1,140-base-pair fragment widely used in bat systematics, was amplified and sequenced with specific primers.

The phylogenetic analysis aligned the new sequence with publicly available data from related genera including Pipistrellus, Vespertilio, Nyctalus and Falsistrellus, using Myotis chinensis as an outgroup. Maximum likelihood inference in IQ-TREE, with a TIM2+F+I substitution model and ultrafast bootstrap support estimated over 1,000 replicates, placed the Yunnan specimen firmly within a clade containing published sequences of Peters’s Pipistrelle from Luzon Island in the Philippines, with a bootstrap value of 100, the maximum possible. Pairwise genetic distances under the K2P model showed the new sample differed from Philippine H. petersi by only 0.10, whereas distances to other Hypsugo species ranged from 0.12 to 0.29. Notably, the Chinese sample formed the basal lineage relative to other conspecifics, hinting at an older, possibly ancestral population in the region.

Morphology told a consistent story. The bat is large for the genus, with a forearm length of 39.94 millimetres and a body mass of 7.5 grams. Its dorsal fur is long and woolly, dark chocolate-brown with lighter tips that give a frosted appearance, while the ventral fur is paler with brown tips. The wing membranes are blackish and hairless except for a small patch of pale-brown fur extending onto the ventral surface of the femur and the area around the anus, a hallmark of the species. The ears are dark, broad and rounded, the tragus short, broad and blunt, and the calcar bears a slight keel. The skull is long and narrow with slender zygomatic arches, well-developed occipital crests, and lower molars of the myotodont type, in which the talonid exceeds the trigonid in size, a diagnostic feature of Hypsugo that distinguishes it from Pipistrellus.

Multivariate statistics added a third line of evidence. Principal component analysis of six external measurements, supplemented with data from four other Chinese Hypsugo species, explained 69.6 percent of total variance in the first two components and separated five morphological groups, with the new specimen distinguished chiefly by its relatively long ears. A second PCA based on twelve craniodental measurements explained 72.1 percent of variance and clearly separated the Yunnan bat from all known Chinese Hypsugo, with condylobasal length and width across the upper canines driving the separation. Some measurements were slightly larger than previously published ranges, but the combined morphological, molecular and multivariate evidence left little doubt about the identification.

The taxonomic history of this bat is convoluted, which partly explains why it has been so easy to overlook. Described as Vespertilio petersi in 1899 from northern Sulawesi, the species was shuffled through the twentieth century between Pipistrellus and the genus Falsistrellus, erected in 1943 for Australian relatives. Hill and Harrison in 1987 even suggested that Asian Falsistrellus might be the eastern representative of Hypsugo. Only in 2018 did a comprehensive study integrating molecular, cranial, dental and multivariate evidence formally transfer the Asian members of Falsistrellus, including petersi, into Hypsugo, a revision now widely adopted. The genus Hypsugo itself comprises 22 species distributed across the southern Palaearctic, most of Africa and the Indomalayan region.

Because confirmed records of H. petersi remain so few, the International Union for Conservation of Nature classifies the species as Data Deficient, with critical gaps in knowledge about population trends, ecology and threats. The new record comes from southern Yunnan, adjacent to the Indo-China Peninsula biodiversity hotspot spanning Laos, Myanmar and Vietnam. The complete absence of records across that vast peninsula suggests a significant distributional gap and indicates that the species’ true geographic range is far less understood than its scattered island records imply. Targeted surveys and molecular screening of museum specimens and field samples across the region are now needed to map where this bat actually occurs.

Wing morphology offers clues to how the species lives. With high wing loading of about 16.77 newtons per square metre, an average aspect ratio of about 6.87 and a low tip-shape index, H. petersi is built for moderately fast flight with limited maneuverability, suited to foraging along semi-open corridors such as forest edges and gaps. This matches the field observation of the individual roosting in a building crevice surrounded by tall, dense coniferous forest at 2,539 metres elevation, and it aligns with earlier accounts of the species being caught in mossy and pine forests at high elevations in the Philippines and roosting under house eaves in Borneo and the Philippines. Remarkably, the Yunnan record extends the known altitudinal limit of the species upward by 400 metres.

To help future surveys, the researchers also published the first comprehensive identification key to all seven Hypsugo species now known from China, in both English and Chinese. Previous guides covered only three species, and the transfer of Asian Falsistrellus species to Hypsugo in 2018 had left the Chinese fauna without an up-to-date diagnostic tool. The key relies on forearm length, fur coloration on the femur and perianal region, dental characters such as the position of the first upper premolar, and tragus shape. The authors caution that the presence of two large-bodied relatives, H. affinis and H. mordax, remains unverified in China and flag them as requiring confirmation. The discovery underscores that bat diversity in China is still incompletely documented and that integrative taxonomy, combining genetics, morphology and morphometrics, remains essential for uncovering it.

Subject of Research: First record of the bat Hypsugo petersi in China based on morphological and genetic evidence

Article Title: First record of the Peters’s Pipistrelle Hypsugo petersi ( Chiroptera , Vespertilionidae ) from China and an updated identification key to Chinese Hypsugo

Article References: Guo, Y., Chen, K.-H., Fan, X.-D., Xue, L.-Y., Yang, H.-H., Xiao, Y.-J., Wang, X.-Y., Wu, Y., Yue, Y., & Yu, W.-H. (2026). First record of the Peters’s Pipistrelle Hypsugo petersi (Chiroptera, Vespertilionidae) from China and an updated identification key to Chinese Hypsugo. ZooKeys, 1290, 171-184. https://doi.org/10.3897/zookeys.1290.186733

Image Credits: AI Generated

DOI: 10.3897/zookeys.1290.186733

Keywords: Hypsugo petersi, bats, Chiroptera, Yunnan, China, new record, cytochrome b, phylogenetics, morphometrics, conservation, ZooKeys, taxonomy

The genus Heremites is a group of skinks in the family Scincidae that ranges from North Africa through the Middle East and Anatolia into Central Asia. Only three species were previously recognised: H. auratus, H. septemtaeniatus, and H. vittatus. Yet the taxonomy of these lizards has long been troubled by their morphological similarity and by persistent uncertainty about the status of named forms across the eastern part of the range. The easternmost populations, reported from eastern Afghanistan and northwestern Pakistan, were especially problematic. A single Afghan specimen collected in 1972 in Nangarhar Province sat in a museum collection as a biogeographic enigma, separated from the core range of the genus by the Hindu Kush mountains and intervening desert basins, with geographic gaps of roughly 700 kilometres to the nearest known occurrences.

The breakthrough came when researchers revisited a reproducing population discovered in Lower Dir District, Khyber Pakhtunkhwa Province, Pakistan, at around 1,110 metres elevation. Specimens collected there between 2013 and 2019 had been provisionally assigned to H. septemtaeniatus transcaucasicus based on morphology alone, but the same researchers noted that several traits deviated from what is typical for that subspecies and explicitly called for DNA data to test whether the isolated population represented a relict or an entirely unknown taxon. The new study answered that call with an integrative approach combining broad mitochondrial screening, a six-locus phylogeny, and comparative morphology.

The molecular work was substantial. The team first assembled a cytochrome b dataset of 209 terminals, including 195 Heremites sequences and 14 outgroups, to map the major mitochondrial lineages across the genus. They then built a concatenated matrix of 57 terminals and 238 locus sequences spanning six loci: two mitochondrial ribosomal genes (12S and 16S rRNA), two mitochondrial protein-coding genes (cytochrome b and ND2), and two nuclear genes (MC1R and RAG1). Maximum-likelihood analyses recovered four main well-supported clades corresponding to H. auratus, the H. septemtaeniatus complex, the H. vittatus complex, and the Hindu Kush lineage.

The genetic distances are remarkable. The Hindu Kush lineage differs from its western relatives by 15.2 to 15.8 percent uncorrected p-distance at cytochrome b and 17.0 to 21.0 percent at ND2, while showing lower but consistent nuclear differentiation of 1.17 to 2.59 percent at MC1R and 2.5 to 3.3 percent at RAG1. For context, the ND2 divergence between H. auratus and the H. septemtaeniatus complex is only 11.6 percent, and the maximum divergence within the H. vittatus complex is 6.8 percent. Allele networks for the nuclear markers revealed complete allelic differentiation among the sampled lineages, with the new species carrying private alleles at both RAG1 and MC1R, providing independent corroboration of the mitochondrial signal.

Divergence dating using a Bayesian framework with five secondary calibration points from a fossil-calibrated mabuyine phylogeny estimated that the lineage leading to H. afghanicus diverged from the common ancestor of H. auratus and the H. septemtaeniatus complex approximately 10.2 million years ago. This deep split predates much of the diversification within the western lineages of the genus, suggesting that the Hindu Kush population represents an old, independently evolving branch rather than a recent offshoot.

Morphological evidence reinforced the genetic picture. The researchers measured 37 characters across 27 museum specimens, including 14 individuals of the new lineage, six of H. auratus, and five representing the H. septemtaeniatus complex. Multivariate analyses using principal component analysis and multiple factor analysis separated the Hindu Kush lineage into a discrete cluster. The new species is diagnosable by a unique combination of traits: parietal shields consistently in contact (present in all 14 examined specimens, unlike the separated parietals of the H. septemtaeniatus complex), contact between the third supraocular and the frontal, 34 to 38 scales around midbody, 52 to 60 dorsal scale rows, 60 to 69 ventral scales, three-keeled dorsal scales, and a comparatively short eye-nostril distance of 2.7 to 5.1 millimetres.

The species is currently known from only two foothill localities along the southern margin of the Hindu Kush, approximately 140 kilometres apart. The type locality in Lower Dir District sits in a montane landscape of moist temperate forest, subtropical Chir pine, and broad-leaved woodland, where lizards were found on rocky slopes with boulders and crevices. The Afghan paratype locality near Dar-e-Nur in Nangarhar Province lies in a rugged, monsoon-influenced valley northeast of Jalalabad. The documented area of occupancy, calculated on a standard 2 by 2 kilometre grid, is just 8 square kilometres, corresponding to two occupied grid cells.

The name afghanicus honours not only Afghanistan in a geopolitical sense but explicitly the Pashtun-speaking people associated with the southern slopes of the Hindu Kush, reflecting the biogeographic and cultural-historical linkage of the region. The species has been suggested vernacular names in English, Pashto, and Urdu, including Afghan grass skink and Dir grass skink.

The discovery raises important questions about hidden biodiversity in one of the world’s most topographically complex regions. Whether H. afghanicus is truly restricted to these two known localities or represents a more broadly distributed species along the Hindu Kush foothills remains unknown. Substantial portions of the genus’ range, particularly in central and western Afghanistan, remain unsampled, and socio-political constraints make targeted fieldwork difficult. The authors emphasise that denser geographic sampling and higher genomic resolution will be necessary to fully delimit species boundaries and reconstruct the evolutionary history of the genus, but their findings represent a significant step toward understanding the poorly investigated biodiversity of the region between the Indus River and the Hindu Kush mountains.

Subject of Research: Discovery and integrative taxonomic description of a new skink species from the Hindu Kush foothills

Article Title: A new species of Heremites Gray, 1845 ( Squamata , Scincidae ) from the transboundary Hindu Kush foothills

Article References: Jablonski, D., Hofmann, S., Idrees, M., Khan, M. A., Jamal, Q., Böhme, W., & Masroor, R. (2026). A new species of Heremites Gray, 1845 (Squamata, Scincidae) from the transboundary Hindu Kush foothills. ZooKeys, 1291, 111-136. https://doi.org/10.3897/zookeys.1291.1863652

Image Credits: AI Generated

DOI: 10.3897/zookeys.1291.1863652

Keywords: Heremites afghanicus, skink, Hindu Kush, new species, phylogenetics, cytochrome b, Scincidae, Afghanistan, Pakistan, biogeography, mitochondrial DNA, taxonomy

The genus Garrulus currently comprises three forest-dwelling corvids. The Eurasian Jay Garrulus glandarius sprawls across an enormous range, from the Atlas Mountains of North Africa across northern Eurasia to Japan, and south into the Himalayas and continental Southeast Asia. Its two relatives are far more restricted: the Lanceolated Jay Garrulus lanceolatus lives in the western Himalayas, while Lidth’s Jay Garrulus lidthi is confined to three islands of Japan’s Ryukyu Arc. Across its vast distribution, the Eurasian Jay displays a bewildering array of plumage variation, and the world’s major bird checklists recognise between 34 and 40 subspecies, traditionally arranged into eight groups.

That traditional arrangement, first formalised by Erwin Stresemann in 1940, divides the species into the European glandarius group, the North African cervicalis group, the Middle Eastern and Caucasian atricapillus group, the Caspian hyrcanus group, the largely Siberian brandtii group, the Japanese japonicus group, the Himalayan and southern Chinese bispecularis group, and the Southeast Asian leucotis group. Some of these groups have repeatedly been proposed as full species, and the checklist authority del Hoyo and Collar recently elevated the two Southeast Asian groups to species rank, a move not adopted by other major checklists. The resulting taxonomic confusion prompted the research team to treat the aggregate as the Garrulus glandarius complex and to test whether plumage-based groupings correspond to real evolutionary lineages.

To do so, the researchers analysed sequences of two mitochondrial markers: the protein-coding cytochrome b gene and the mitochondrial control region, the latter generally more variable and well suited to resolving differences among populations. The study relied heavily on museum specimens, with DNA extracted from small foot pad samples, two to three millimetres in diameter, taken from 54 bird skins held in the Natural History Museum Vienna, alongside tissue samples from 38 additional individuals. In total, 92 jays were sampled, including 87 individuals of the glandarius complex covering nine of the ten subspecies groups the authors now recognise, plus specimens of the two other Garrulus species. Even specimens collected between 1831 and 1836 yielded usable DNA fragments of 343 base pairs, a testament to the value of historical collections for modern genomics.

Working with such old material required careful technique. Foot pad samples were digested for up to 71 hours, extractions were performed in a clean room with regular UV irradiation, and blank controls were run at a ratio of at least one per ten samples. Because a C-stretch near the start of the control region hampered sequencing, the team designed new primers and assembled complete sequences from overlapping fragments of roughly 300 base pairs each. The final analyses combined 2483 aligned positions and employed Bayesian inference with millions of Markov chain Monte Carlo generations, complemented by median-joining haplotype networks to visualise relationships among individual birds.

The results confirm that Garrulus is monophyletic and split into two major mitochondrial lineages: one containing the Lanceolated Jay and Lidth’s Jay, which differ from the glandarius complex by uncorrected genetic distances of roughly 10.5 to 12.2 percent, and one containing all Eurasian Jay populations. Within the complex, at least eight major clades emerged, generally matching the established subspecies groups. The most basal split separates the Japanese japonicus group from all remaining jays, a result consistent with earlier, more limited studies. A Southeast Asian clade unites the bispecularis and leucotis groups, while the brandtii and hyrcanus groups each form distinct lineages. The Western Palearctic taxa, comprising the glandarius, cervicalis, atricapillus and glaszneri groups, cluster together in a well-supported western clade, though the relationships among these four remain less certain.

Several findings stand out. The insular subspecies Garrulus g. taivanus from Taiwan proved to be, alongside the japonicus group, the most genetically distinct member of the entire complex, with distances of up to 3.4 percent from its closest relatives, a striking contrast to the morphological uniformity of the bispecularis group to which it belongs. The Cyprus Jay, Garrulus g. glaszneri, emerged as the most distinct lineage within the western clade, supporting its recognition as a separate subspecies group rather than a member of the nominate European group, and suggesting it is a relict of a formerly more widespread form. On Crete, the subspecies Garrulus g. cretorum showed a control-region haplotype separated by at least 15 substitutions from all other European jays, indicating long isolation. In Siberia, jays from Hokkaido were clearly differentiated from mainland and Sakhalin birds in the control-region data, hinting at an independent glacial refugium on the northern Japanese island.

Not everything fits the neat subspecies map, however. The study detected several cases of possible interbreeding between representatives of different groups. Two birds from Azerbaijan with intermediate plumage between the hyrcanus and atricapillus groups carried mitochondrial DNA of the atricapillus clade, strongly suggesting they are genuine intergrades. A specimen of the Turkish subspecies ferdinandi, normally assigned to the European group, clustered within the Caucasian atricapillus clade, complicating interpretations of intergradation around the Bosporus. Most puzzling, five Crimean jays of the subspecies iphigenia, traditionally placed in the black-capped atricapillus group, instead clustered within the European glandarius clade. The authors caution that a major invasion of migrating jays occurred in Europe during the winter when most of these birds were collected, so the samples might represent migrants from more northern breeding grounds rather than local birds, underscoring how migration can blur phylogeographic patterns.

The Southeast Asian picture is equally intriguing. The two sampled leucotis individuals from Myanmar did not form their own cluster but nested among bispecularis haplotypes, rendering the bispecularis group paraphyletic. The authors propose two main explanations: the sampled birds may be intergrades carrying mitochondrial DNA of the bispecularis group, or mitochondrial capture through large-scale introgression may have replaced the authentic leucotis lineage. Either scenario implies interbreeding in the contact zone of northern Myanmar, where intermediate birds have long been recorded and even described as named subspecies. Notably, the data do not support the recent species-level split of the bispecularis and leucotis groups, because such a move would render the remainder of Garrulus glandarius polyphyletic given the even stronger differentiation of the Japanese japonicus group.

Biogeographically, the results complicate the long-standing assumption that Garrulus originated in the tropical rainforests of Southeast Asia. Although the deepest mitochondrial splits occur in eastern Asia, the genus’s closest relatives appear to be arid-country corvids such as the magpie genus Pica, the Ethiopian Bush-Crow Zavattariornis, the Piapiac Ptilostomus and the ground jays Podoces, an assemblage adapted to open landscapes. The authors suggest that the radiation of this group may have occurred along the mid-latitude belt during the cooling and aridification of Eurasia that began in the early Miocene, and that Garrulus may have emerged in more northern regions of Eurasia, its current distribution reflecting ancient woodland refugia rather than a recent spread out of the tropics.

For now, the authors recommend treating the glandarius complex as a single species while regarding its subspecies groups, and the distinctive island forms of Cyprus and Taiwan, as Evolutionarily Significant Units for conservation purposes. Formal species status for the Cyprus Jay would likely lead to its classification as globally threatened, given forest fragmentation, corvid shooting and illegal trapping on the island. The study deliberately stops short of taxonomic changes, noting that several groups were represented by only one or two individuals and that the minor group of the Atlas Mountains was not sampled at all. Future work with larger sample sizes, nuclear genomic markers such as whole-genome or RAD sequencing, and detailed studies of contact zones and vocalisations will be needed to settle, once and for all, how many jays really share the name Garrulus glandarius.

Subject of Research: Mitochondrial phylogenetics and phylogeography of the jay genus Garrulus and the Eurasian Jay subspecies complex

Article Title: Phylogenetic relationships within the genus Garrulus Brisson, 1760 ( Corvidae , Aves ) with special emphasis on the Garrulus glandarius complex based on mitochondrial marker sequences

Article References: Wolf, P., Töpfer, T., Kryukov, A., Aoki, D., & Haring, E. (2026). Phylogenetic relationships within the genus Garrulus Brisson, 1760 (Corvidae, Aves) with special emphasis on the Garrulus glandarius complex based on mitochondrial marker sequences. Annals of the Natural History Museum Vienna, 127, 13-37. https://doi.org/10.3897/anhmw.168785

Image Credits: AI Generated

DOI: 10.3897/anhmw.168785

Keywords: Garrulus glandarius, Eurasian Jay, mitochondrial DNA, phylogeography, museum specimens, cytochrome b, control region, corvids, subspecies, taxonomy, biogeography, Cyprus Jay

Researchers analyzed Cytochrome-b sequences from 41 Myotis capaccinii samples collected from eight localities across Türkiye, comparing them with previously published sequences from European, Tunisian, and Iranian populations. The team identified nine haplotypes, three of which were recorded for the first time from two Mediterranean locations in southern Türkiye. The findings provide compelling evidence that Bulgarian and Turkish populations form a distinct mitochondrial clade separate from other European samples, while also revealing surprisingly low genetic differentiation among Turkish subpopulations.

The long-fingered bat is classified as Vulnerable by the IUCN due to population declines linked to habitat degradation, water pollution, and disturbance of cave roosts. Its distribution stretches from the Iberian Peninsula and North Africa through Southeastern Europe, Türkiye, and the Levant, reaching Western Iran. This cave-dwelling species is closely associated with aquatic habitats for foraging, making it particularly sensitive to environmental changes that affect freshwater ecosystems.

The taxonomic status of Myotis capaccinii has been controversial for decades. Previous researchers have proposed different subspecies assignments based on morphological differences between European and Anatolian subpopulations. Some studies included specimens from Turkish Thrace in the subspecies M. c. bureschi, while others placed Turkish specimens in the nominative subspecies. More recently, a new subspecies called M. c. abidinbudaki was proposed from southern Central Anatolia based on lighter dorsal fur color and biometric measurements. However, these morphologically defined races have not coincided with the lineages revealed by molecular phylogenetic analyses.

The research team generated new Cytochrome-b sequences from specimens collected in Antalya and Mersin provinces in Mediterranean Türkiye, combining them with sequences from GenBank representing localities across the species’ range. Using Bayesian Inference methods, the researchers constructed phylogenetic trees that revealed three main mitochondrial clusters. Cluster I consisted primarily of western European and Tunisian samples, Cluster II included Turkish, Bulgarian, and Greek samples together with part of the western European samples, and Cluster III comprised mainly Turkish and Iranian samples along with some southeastern European specimens.

Genetic distance analysis showed that the distance between Turkish and European samples was 3.0 percent, while distances among the six Turkish population groups ranged from just 0.10 to 0.76 percent. Haplotype diversity for Turkish samples was 0.786 with nucleotide diversity of 0.005, values quite similar to those found in European populations. The highest genetic distance, 4.0 percent, was observed between Turkish and Iranian samples, suggesting that Iranian populations represent the most divergent group in the study.

Divergence time estimates using Bayesian molecular clock analysis suggest that Cluster III likely separated from Clusters I and II approximately 1.64 million years ago during the mid-Pleistocene, while Clusters I and II diverged around 0.70 million years ago. These timelines align with previous estimates suggesting that Myotis capaccinii diverged from other Myotis species approximately 8 million years ago. The researchers propose that the species’ dispersal into southeastern Mediterranean refugia during the Last Glacial Maximum, followed by recolonization into Europe, facilitated gene exchange between Thracian, Anatolian, and European populations.

Tajima’s D neutrality test yielded a significantly negative value for the Turkish population, potentially indicating recent population expansion. However, Fu’s Fs was not statistically significant, so this signal should be interpreted cautiously. The Median-joining network analysis showed that although common haplotypes were shared among Tunisian, European, Turkish, and Iranian subpopulations, haplotype differentiation in the Bulgarian and Turkish samples was evident, with new haplotypes separated from the central haplotype group by varying mutational steps.

The findings have important implications for bat conservation and taxonomy. The low genetic diversity among Turkish subpopulations is thought to be related to the species’ dispersal ability, with movement among nearby caves and foraging habitats potentially facilitating gene flow. The researchers conclude that there is currently no clear mitochondrial evidence supporting additional subspecies differentiation within Turkish populations, as genetic distances are well below the subspecies level. However, they caution that mitochondrial markers reflect only maternal lineage history, and more comprehensive geographic sampling combined with nuclear markers and genomic data would be necessary to fully evaluate the taxonomic validity of proposed subspecies.

Anatolia’s location between the Middle East and Europe appears to have served as a genetic bridge for eastern and western lineages of this species. The study highlights the importance of the Thrace region, along the Greek-Bulgarian-Turkish border, as a potential contact zone where European and Anatolian lineages meet. Protecting this region is considered crucial for maintaining the species’ genetic diversity. As climate change and habitat destruction continue to threaten cave-dwelling bats across the Mediterranean, understanding the evolutionary history and population structure of vulnerable species like the long-fingered bat becomes increasingly urgent for developing effective conservation strategies.

Subject of Research: Phylogeny and phylogeography of the long-fingered bat Myotis capaccinii using Cytochrome-b mitochondrial DNA

Article Title: Contributions to the phylogeny and phylogeography of Myotis capaccinii (Bonaparte, 1837) ( Chordata , Mammalia , Chiroptera , Vespertilionidae ), inferred with Cytochrome-b

Article References: Barlas, E., Çetintürk, D., & Yiğit, N. (2026). Contributions to the phylogeny and phylogeography of Myotis capaccinii (Bonaparte, 1837) (Chordata, Mammalia, Chiroptera, Vespertilionidae), inferred with Cytochrome-b. Subterranean Biology, 56, 179-194. https://doi.org/10.3897/subtbiol.56.183693

Image Credits: AI Generated

DOI: 10.3897/subtbiol.56.183693

Keywords: Myotis capaccinii, long-fingered bat, phylogeography, Cytochrome-b, mitochondrial DNA, haplotypes, Anatolia, Bulgaria, Last Glacial Maximum, subspecies, cave-dwelling bats, Türkiye

Camarotoscena species live across the temperate Palaearctic, from the Mediterranean Basin to the Far East, with their greatest diversity in the arid mountain regions of Asia. All species known so far are narrowly oligophagous on poplars, genus Populus, and their immatures induce leaf roll galls on the leaves, sometimes on the very same tree, and occasionally even within the same gall, as two species share a host plant. Because poplars are widely planted for timber, fuel, windbreaks and fodder, and because some Camarotoscena species are considered pests, a reliable way to identify these insects matters far beyond the walls of taxonomy. For any pest control programme, a sound taxonomic base is crucial, and until now that base has been shaky for this particular genus.

The challenge stems from the fact that Camarotoscena is morphologically what the late Russian specialist Marina Loginova once described as a monolithic genus. The male and female terminalia, which in most psyllids provide dependable diagnostic characters, are unusually uniform across the group. Adults are small, typically between 2.0 and 2.5 millimetres long, ochreous or light brown with small dark dots, and bear whitish or yellowish forewings. Species are separated by subtle features: the arrangement of surface spinules on the forewing, the density and colour of the dot pattern, body size, and fine details of the genital structures. Loginova recognised 13 species in her 1975 review, three of which have since been transferred to other genera, leaving twelve recognised species going into the new study.

The revision, based on material from museums in Bulgaria, China, Switzerland, Czechia and Basel, confirms the ten species that remain after careful re-examination. The authors propose five new synonymies, collapsing four names into existing species. Camarotoscena ujenci, erected in 1982 for just two males from Mongolia, is now considered the same as C. subrubescens, a bright orange species from the Mediterranean region. Three Chinese species described in 2011 from single females, C. wulingshanica, C. wutaishanica and C. xinjiangica, are formally synonymised with C. bianchii, a species characterised by its square female subgenital plate. A doctoral thesis had already suggested these synonymies in 2016, but under the International Code of Zoological Nomenclature such acts are only valid when formally published, which the new paper now does.

The morphological work went beyond adults. Fifth instar immatures, known for only half of the species, turned out to carry their own diagnostic characters. The researchers examined the chaetotaxy, the arrangement of setae and pointed sectasetae on the caudal plate and forewing pads, as well as the degree of sclerotisation of the ventrite bearing the circumanal ring. These immature characters independently supported the species boundaries diagnosed from adults, a satisfying convergence for a genus where adult features are so subtle. The previously unknown female of C. lauta, the largest species in the genus at 2.8 to 3.2 millimetres, is now diagnosed and illustrated, and illustrated identification keys are provided for both adults and, where known, the immatures.

Then came the molecular surprise. The team sequenced two mitochondrial markers, cytochrome c oxidase subunit I and cytochrome b, from specimens of five species collected across India, Jordan, China, Czechia and Bulgaria, using both traditional Sanger sequencing and Oxford Nanopore technology. A commonly applied 3 percent COI threshold for species delimitation failed spectacularly in two cases. Between C. fulgidipennis and C. hoberlandti, two species that coexist on black poplar in the Middle East, Central Asia and the western Himalaya, there were zero differences in COI. Between C. speciosa and C. subrubescens, the Mediterranean species pair that sometimes shares the same trees, the difference was essentially nil in both genes. And cytb additionally failed to distinguish C. trjapitzini from the fulgidipennis-hoberlandti pair, even though COI separated that species by 2.6 percent.

The phylogenetic picture was not entirely bleak. A maximum likelihood tree built from concatenated COI and cytb sequences recovered two main groups, each also defined by morphology: one containing C. speciosa and C. subrubescens, the other containing C. fulgidipennis, C. hoberlandti and C. trjapitzini. Between these groups, genetic distances ranged from about 7.3 to 9.1 percent, clearly signalling separate lineages. Within groups, however, the barcodes went silent. The authors suggest that mitochondrial introgression or incomplete lineage sorting, phenomena well documented across the animal kingdom, may explain the mismatch, and they recommend that future work include nuclear genes or genomic-scale data to resolve the conflict.

Could the species pairs actually be one species each, perhaps seasonal forms of a single lineage as known from some pear psyllids? The authors consider this unlikely, because the two forms within each pair occur simultaneously at the same localities, independent of season. The presence of two species within a single leaf roll gall has a simpler explanation: females of both species lay eggs close together on the same leaf, and the gall is induced by the sucking of the first instar immatures, so all nearby individuals end up enclosed regardless of which species initiated it. Field observations from southern France add ecological texture, with C. subrubescens favouring black poplar along watercourses while C. speciosa also colonises trees at ditch edges.

The study also redraws the map. New country records include C. fulgidipennis and C. hoberlandti from Ladakh in India, the first record of the genus and of any psyllid from that cold desert region, C. lauta from Ankara in Turkey, and C. trjapitzini from Tehran and Yazd in Iran and from Spain. Perhaps most tellingly, a specimen from Jordan previously identified as C. hoberlandti was re-examined and found to be C. trjapitzini, a correction that matters because the two species are so similar that even published DNA sequences had been attached to the wrong name.

The broader lesson is one that molecular ecologists are hearing with increasing frequency. DNA barcoding is a fast and inexpensive tool that works well for many insect groups, and it has successfully delimited species across all seven recognised psyllid families, but it is not infallible. In other psyllid genera such as Aphalara, Cacopsylla and Melanastera, mitochondrial barcodes generally identify species correctly, yet even there exceptions abound, with intraspecific divergence reaching 14 percent in some Melanastera species while morphologically distinct Aphalara species differ by less than 1 percent. The Camarotoscena results highlight the limitations of relying solely on mitochondrial DNA from a few specimens of few localities for species delimitation. Rather than risk ill-founded taxonomic upheaval, the authors retain the traditional ten-species concept and call for targeted fieldwork, fresh material from more localities, nuclear gene sequencing, and descriptions of the still-unknown immatures. For now, the poplar galls keep their secrets, and the humble microscope remains the psyllid taxonomist’s most trustworthy instrument.

Subject of Research: Species delimitation and DNA barcoding of poplar-associated psyllids in the genus Camarotoscena

Article Title: Species concepts in the psyllid genus Camarotoscena ( Hemiptera , Psylloidea ) associated with Populus ( Salicaceae )

Article References: Burckhardt, D., Haseen, C., He, Z., Luo, X., Pramatarova, M., Grimaud, S., Hussain, B., Song, F., & Serbina, L. Š. (2025). Species concepts in the psyllid genus Camarotoscena (Hemiptera, Psylloidea) associated with Populus (Salicaceae). Alpine Entomology, 9, 153-172. https://doi.org/10.3897/alpento.9.172655

Image Credits: AI Generated

DOI: 10.3897/alpento.9.172655

Keywords: Camarotoscena, psyllids, DNA barcoding, Populus, taxonomy, COI, cytochrome b, leaf roll galls, Hemiptera, species delimitation, Palaearctic, morphology