An additional species of the genus Nemaspela Silhavý, 1966 from a cave in Georgia, with a note on Nemaspela melouri (Opiliones: Nemastomatidae).

A cavernicolous harvestman species, Nemaspela kotia sp. nov., is described from the Kotia Cave (=Kotiasklde Cave) in Zemo Imereti plateau, Imereti Region, West Georgia. Its males lack a cheliceral apophysis, a feature shared with the Georgian cave-dwelling Nemaspela femorecurvata Martens, 2006 and N. prometheus Martens, Maghradze & Barjadze, 2021. Besides, Nemaspela melouri Martens, Maghradze & Barjadze, 2021, known only from its type locality, is found in Solkota Cave (Tskaltubo Municipality, Sataplia-Tskaltubo karst massif, Imereti region, West Georgia). A key for the identification of the known species of Nemaspela is provided.

Limited Song Mixing Without Genomic Gene Flow in a Contact Zone Between Two Songbird Species.

Song is considered to play an important role in the maintenance of prezygotic reproductive isolation between closely related songbird species. Therefore, song mixing in a contact zone between closely related species is often considered as evidence of hybridization. The Sichuan Leaf Warbler Phylloscopus forresti and the Gansu Leaf Warbler Phylloscopus kansuensis, which diverged 2 million years ago, have formed a contact zone in the south of the Gansu Province of China, where mixed songs have been observed. In this study, we investigated the potential causes and consequences of song mixing by integrating bioacoustic, morphological, mitochondrial, and genomic data with field ecological observations. We found that the two species display no apparent morphological differences, whereas their songs differ dramatically. We demonstrated that ∼11% of the males in the contact zone sang mixed songs. Two males singing mixed song were genotyped, and both were found to be P. kansuensis. Despite the presence of mixed singers, population genomic analyses detected no signs of recent gene flow between the two species, although two possible cases of mitochondrial introgression were identified. We conclude that the rather limited song mixing does not lead to, or result from, hybridization, and hence does not result in the breakdown of reproductive barriers between these cryptic species.

On the genus Tegestria Roewer, 1936, new senior synonym of Gintingius Roewer, 1938 with the description of a new species (Opiliones: Epedanidae: Sarasinicinae).

Tegestria altmannae sp. nov. is described and illustrated based on male and female specimens collected in Malaysia. It is characterized by the unarmed dorsal scutum, and basal segment of chelicerae dorsally with three seta-tipped tubercles. The type species of the monotypic genus Gintingius Roewer, 1938, G. robustus Roewer, 1938, is treated as a new synonym of Tegestria coniata Roewer, 1938 and the genus Gintingius as a new synonym of Tegestria Roewer, 1936, accordingly.

From the Ethiopian Bale Mountains hotspotFilopalpinae subfam. nov., a new taxon of Laniatorean harvestmen based on external and genital morphology (Arachnida, Opiliones, Assamiidae).

Filopalpinae subfam. nov. is proposed as a new subfamily within family Assamiidae based on a new genus, Filopalpus gen. nov. and on five new species: F. joschmidti sp. nov., F. bale sp. nov., F. kakaensis sp. nov., F. altomontanus sp. nov. and F. niger sp. nov. All originate from the highlands of Ethiopia, namely the Bale Mountains and adjacent volcanoes in Oromia State. Species of the new subfamily are characterized by dense body cover with pointed tubercles causing a hedgehog appearance of the body (dorsal side, partly ventral side), extreme sexual dimorphism in pedipalps (in males thread-like long and thin surpassing body length by about four to six times, devoid of apophyses and thorns; in females short and stout about body length, with a distal grasping hand formed by apophyses and strong setae of tibia and tarsus, and tarsal claw). Male genital morphology is characterized by a hemolymph-pressure system of truncus penis and glans, a dorsal sub-distal tube-like glans with an inner prickly funnel, which is everted by hemolymph pressure during courtship. These male genital characters place the present species in Assamiidae, but external morphology points to a strongly different separate evolutionary unit within this family. The five species known to date were secured at altitudes between 2830 and 4100 m. At least two of the collecting areas are now devoid of forest due to human impact. To secure the present specimens at high-altitude localities was possible only by searching in remote microhabitats unavailable for cattle and man-made fire. A perspective on current subfamilies of Assamiidae and on pedipalpal morphology and evolution is presented.

Speciation and population divergence in a mutualistic seed dispersing bird.

Bird-mediated seed dispersal is crucial for the regeneration and viability of ecosystems, often resulting in complex mutualistic species networks. Yet, how this mutualism drives the evolution of seed dispersing birds is still poorly understood. In the present study we combine whole genome re-sequencing analyses and morphometric data to assess the evolutionary processes that shaped the diversification of the Eurasian nutcracker (Nucifraga), a seed disperser known for its mutualism with pines (Pinus). Our results show that the divergence and phylogeographic patterns of nutcrackers resemble those of other non-mutualistic passerine birds and suggest that their early diversification was shaped by similar biogeographic and climatic processes. The limited variation in foraging traits indicates that local adaptation to pines likely played a minor role. Our study shows that close mutualistic relationships between bird and plant species might not necessarily act as a primary driver of evolution and diversification in resource-specialized birds.

Asymmetric allelic introgression across a hybrid zone of the coal tit (Periparus ater) in the central Himalayas.

In the Himalayas, a number of secondary contact zones have been described for vicariant vertebrate taxa. However, analyses of genetic divergence and admixture are missing for most of these examples. In this study, we provide a population genetic analysis for the coal tit (Periparus ater) hybrid zone in Nepal. Intermediate phenotypes between the distinctive western "spot-winged tit" (P. a. melanolophus) and Eastern Himalayan coal tits (P. a. aemodius) occur across a narrow range of <100 km in western Nepal. As a peculiarity, another distinctive cinnamon-bellied form is known from a single population so far. Genetic admixture of western and eastern mitochondrial lineages was restricted to the narrow zone of phenotypically intermediate populations. The cline width was estimated 46 km only with a center close to the population of the cinnamon-bellied phenotype. In contrast, allelic introgression of microsatellite loci was asymmetrical from eastern P. a. aemodius into far western populations of phenotypic P. a. melanolophus but not vice versa. Accordingly, the microsatellite cline was about 3.7 times wider than the mitochondrial one.

A new genus and two new species of Galumnidae (Acari, Oribatida) from Nepal.

One new genus and two new species of oribatid mites (Oribatida) of the family Galumnidae are described from soil in Nepal. Anomalogalumna gen. nov. (with type species Anomalogalumna dungeri sp. nov.) differs from Flagellozetes Balogh, 1970 by the absence of octotaxic system, and the presence of short, notogastral setae and areolate-reticulate surface of the notogaster and anogenital region. Allogalumna beateae sp. nov. differs from all other species of the genus by the presence of thickened rostral, lamellar and interlamellar setae, and the position of rostral seta (close to lamellar seta and distant from the rostrum).

Two new species of the genus emNemaspela/em Silhavý from caves in Georgia (Opiliones: Nemastomatidae).

Two highly specialized endemic troglobiotic harvestman species of the genus Nemaspela Silhavý, 1966 are described. N. melouri sp. nov. from Melouri Cave and N. prometheus sp. nov. from Prometheus Cave (Sataplia-Tskaltubo karst massif, Imereti region, western Georgia), respectively. Despite the fact that the entrances of the caves are positioned only 2.5 km apart, the new taxa differ from each other distinctly by presence vs. absence of male cheliceral apophysis, which is lacking in the second species. A key to the Caucasian species of the genus is provided. Relationships of Nemaspela species within the genus and with hypothetical epigean ancestors are discussed.

"Into and Out of" the Qinghai-Tibet Plateau and the Himalayas: Centers of origin and diversification across five clades of Eurasian montane and alpine passerine birds.

Encompassing some of the major hotspots of biodiversity on Earth, large mountain systems have long held the attention of evolutionary biologists. The region of the Qinghai-Tibet Plateau (QTP) is considered a biogeographic source for multiple colonization events into adjacent areas including the northern Palearctic. The faunal exchange between the QTP and adjacent regions could thus represent a one-way street ("out of" the QTP). However, immigration into the QTP region has so far received only little attention, despite its potential to shape faunal and floral communities of the QTP. In this study, we investigated centers of origin and dispersal routes between the QTP, its forested margins and adjacent regions for five clades of alpine and montane birds of the passerine superfamily Passeroidea. We performed an ancestral area reconstruction using BioGeoBEARS and inferred a time-calibrated backbone phylogeny for 279 taxa of Passeroidea. The oldest endemic species of the QTP was dated to the early Miocene (ca. 20 Ma). Several additional QTP endemics evolved in the mid to late Miocene (12-7 Ma). The inferred centers of origin and diversification for some of our target clades matched the "out of Tibet hypothesis' or the "out of Himalayas hypothesis" for others they matched the "into Tibet hypothesis." Three radiations included multiple independent Pleistocene colonization events to regions as distant as the Western Palearctic and the Nearctic. We conclude that faunal exchange between the QTP and adjacent regions was bidirectional through time, and the QTP region has thus harbored both centers of diversification and centers of immigration.

Phylogeny of the Eurasian Wren Nannus troglodytes (Aves: Passeriformes: Troglodytidae) reveals deep and complex diversification patterns of Ibero-Maghrebian and Cyrenaican populations.

The Mediterranean Basin represents a Global Biodiversity Hotspot where many organisms show high inter- and intraspecific differentiation. Extant phylogeographic patterns of terrestrial circum-Mediterranean faunas were mainly shaped through Pleistocene range shifts and range fragmentations due to retreat into different glacial refugia. Thus, several extant Mediterranean bird species have diversified by surviving glaciations in different hospitable refugia and subsequently expanded their distribution ranges during the Holocene. Such a scenario was also suggested for the Eurasian Wren (Nannus troglodytes) despite the lack of genetic data for most Mediterranean subspecies. Our phylogenetic multi-locus analysis comprised 18 out of 28 currently accepted subspecies of N. troglodytes, including all but one subspecies which are present in the Mediterranean Basin. The resulting phylogenetic reconstruction dated the onset of the entire Holarctic radiation of three Nannus species to the early Pleistocene. In the Eurasian Wren, two North African subspecies represented separate basal lineages from the Maghreb (N. t. kabylorum) and from the Libyan Cyrenaica (N. t. juniperi), being only distantly related to other Mediterranean populations. Although N. troglodytes appeared to be paraphyletic with respect to the Nearctic Winter Wren (N. hiemalis), respective nodes did not receive strong statistical support. In contrast, paraphyly of the Ibero-Maghrebian taxon N. t. kabylorum was strongly supported. Southern Iberian populations of N. t. kabylorum did not clade with Maghrebian populations of the same subspecies but formed a sister clade to a highly diverse European clade (including nominate N. t. troglodytes and eight further taxa). In accordance with a pattern also found in other birds, Eurasian populations were split into a western clade (Europe, Caucasus) and an eastern clade (Central Asia, Sino-Himalayas, East Asia). This complex phylogeographic pattern revealed cryptic diversification in N. troglodytes, especially in the Iberio-Maghrebian region.

A taxonomic study on Epedanidae from Thailand including functional aspects of male genital morphology (Opiliones, Laniatores).

The South-East Asian opilionid family Epedanidae Sørensen, 1886 has one of its strongholds in Thailand from where a multitude of genera and species have been described but the epedanid fauna of the country is still poorly known. This paper records four species from this country, three of which are new: Euepedanus dashdamirovi sp. nov. (male and female), Plistobunus jaegeri sp. nov. (male and female), and Toccolus kuryi sp. nov. (male and female). Toccolus globitarsis Suzuki, 1969 was previously known only from the type locality in Thailand and is redescribed here. Functional aspects of epedanid penial morphology are highlighted.

Genetic admixture despite ecological segregation in a North African sparrow hybrid zone (Aves, Passeriformes, Passer domesticus × Passer hispaniolensis).

Under different environmental conditions, hybridization between the same species might result in different patterns of genetic admixture. Particularly, species pairs with large distribution ranges and long evolutionary history may have experienced several independent hybridization events over time in different zones of overlap. In birds, the diverse hybrid populations of the house sparrow (Passer domesticus) and the Spanish sparrow (Passer hispaniolensis) provide a striking example. Throughout their range of sympatry, these two species do not regularly interbreed; however, a stabilized hybrid form (Passer italiae) exists on the Italian Peninsula and on several Mediterranean islands. The spatial distribution pattern on the Eurasian continent strongly contrasts the situation in North Africa, where house sparrows and Spanish sparrows occur in close vicinity of phenotypically intermediate populations across a broad mosaic hybrid zone. In this study, we investigate patterns of divergence and admixture among the two parental species, stabilized and nonstabilized hybrid populations in Italy and Algeria based on a mitochondrial marker, a sex chromosomal marker, and 12 microsatellite loci. In Algeria, despite strong spatial and temporal separation of urban early-breeding house sparrows and hybrids and rural late-breeding Spanish sparrows, we found strong genetic admixture of mitochondrial and nuclear markers across all study populations and phenotypes. That pattern of admixture in the North African hybrid zone is strikingly different from i) the Iberian area of sympatry where we observed only weak asymmetrical introgression of Spanish sparrow nuclear alleles into local house sparrow populations and ii) the very homogenous Italian sparrow population where the mitogenome of one parent (P. domesticus) and the Z-chromosomal marker of the other parent (P. hispaniolensis) are fixed. The North African sparrow hybrids provide a further example of enhanced hybridization along with recent urbanization and anthropogenic land-use changes in a mosaic landscape.

Ecotypic changes of alpine birds to climate change.

In endotherm animals, several traits are related to climate. For example, Bergmann's rule predicts a decrease in body size within species and across closely related species with increasing temperature, whereas Gloger's rule states that birds and mammals should be darker in humid and warm environments compared to colder and drier areas. However, it is still not clear whether ecotypic responses to variation in the local environment can also apply to morphological and colouration changes through time in response to climate change. We present a 100-year-long time series on morphological and melanin-based colours of snowfinch (325 Montifringilla, 92 Pyrgilauda and 30 Onychostruthus) museum specimens. Here we show that the tarsus length of the species has decreased and the saturation of the melanin-based colour has increased, which was correlated with the increase of temperature and precipitations. As ecotypic variations are tightly linked to individual behavioural and physiological responses to environmental variations, differently sized and coloured individuals are expected to be differently penalized by global changes. This study opens the pertinent question about whether ecotypic responses can enhance population persistence in the context of global change.

Rediscovery of Lobonychiumpalpiplus Roewer, 1938 (Opiliones, Laniatores, Epedanidae) in Sabah, Malaysia.

Lobonychiumpalpiplus Roewer, 1938, originally reported from Indonesian Borneo, is redescribed based on the specimens from Malaysia. The genitalia of this species are described for the first time and a new genital terminology is proposed. The rediscovery expands the known distribution of the species to Malaysian Borneo.

Two new species of Biantes from China and Malaysia (Opiliones: Laniatores: Biantidae).

Two new species of Biantes Simon, 1885 (Biantidae: Biantinae) are described: Biantes calyptroideus sp. nov. (male and female) from China and Biantes spatulatus sp. nov. (male and female) from Malaysia. Biantidae is recorded from China for the first time.

Ancient home or in exile? The easternmost species of genus Starengovia Snegovaya, 2010 found in China (Opiliones, Nemastomatidae, Nemastomatinae).

Starengovia quadrituberculatasp. n. is described and illustrated based on male and female specimens collected in Yunnan Province, China. The new species is distinct from the two other congeners, S. kirgizica Snegovaya, 2010 and S. ivanloebli Martens, 2017, in having two pairs of low submedian tubercles on abdominal areae III and IV; distal margin of the lateral foliate wing-like structures of the penis situated close to the glans base, the short rod-like stylus, the form and position of spines on the stylus of the penis, anvil-shaped tubercles mainly on front margin of prosoma. The occurrence of Starengovia in Yunnan, the second nemastomatine species in China, creates a huge distributional gap of roughly 2700 km distance to its closest neighbor S. ivanloebli in Northwest Pakistan. The historical relations of Chinese nemastomatines are discussed.

Distribution, diversity patterns and faunogenesis of the millipedes (Diplopoda) of the Himalayas.

The Himalayas support a highly rich, diverse, multi-layered, mostly endemic diplopod fauna which presently contains >270 species, 53 genera, 23 families and 13 orders. This is the result of mixing the ancient, apparently Tertiary and younger, Plio-Pleistocene elements of various origins, as well as the most recent anthropochore (= man-mediated) introductions. At the species and, partly, generic levels, the fauna is largely autochthonous and sylvicolous, formed through abounding in situ radiation and vicariance events. In general, the species from large genera and families tend to occupy a wide range of altitudes, but nearly each of the constituent species shows a distribution highly localized both horizontally and altitudinally, yet quite often with sympatry or even syntopy involved. The bulk of the fauna is Indo-Malayan in origin, with individual genera or families shared with those of SE Asia (mostly) and/or S India (few). Sino-Himalayan and, especially, Palaearctic components are subordinate, but also clearly distinguishable.

Two new species of Sabacon Simon, 1879 (Opiliones: Dyspnoi: Sabaconidae) from Yunnan Province, China.

Two new species of the harvestman genus Sabacon Simon, 1879 are recorded from Yunnan Province, southern China, Sabacon monacanthus sp. n. (male and female; relationships unclear) and S. suzukii sp. n. (male and female; closely related to S. kangding Martens, 2015 from Sichuan Province, southwestern China).

Improved sampling at the subspecies level solves a taxonomic dilemma - A case study of two enigmatic Chinese tit species (Aves, Passeriformes, Paridae, Poecile).

A recent full species-level phylogeny of tits, titmice and chickadees (Paridae) has placed the Chinese endemic black-bibbed tit (Poecile hypermelaenus) as the sister to the Palearctic willow tit (P. montanus). Because this sister-group relationship is in striking disagreement with the traditional affiliation of P. hypermelaenus close to the marsh tit (P. palustris) we tested this phylogenetic hypothesis in a multi-locus analysis with an extended taxon sampling including sixteen subspecies of willow tits and marsh tits. As a taxonomic reference we included type specimens in our analysis. The molecular genetic study was complemented with an analysis of biometric data obtained from museum specimens. Our phylogenetic reconstructions, including a comparison of all GenBank data available for our target species, clearly show that the genetic lineage previously identified as P. hypermelaenus actually refers to P. weigoldicus because sequences were identical to that of a syntype of this taxon. The close relationship of P. weigoldicus and P. montanus - despite large genetic distances between the two taxa - is in accordance with current taxonomy and systematics. In disagreement with the previous phylogenetic hypothesis but in accordance with most taxonomic authorities, all our P. hypermelaenus specimens fell in the sister clade of all western and eastern Palearctic P. palustris. Though shared haplotypes among the Chinese populations of the two latter species might indicate mitochondrial introgression in this part of the breeding range, further research is needed here due to the limitations of our own sampling.

Sinostoma yunnanicum, the first nemastomatine harvestman in China (Arachnida: Opiliones: Nemastomatidae).

The easternmost Nemastomatinae species, Sinostoma yunnanicum n. gen., n. sp., from northern Yunnan, China is described. It extends the geographic distribution of Nemastomatinae by roughly 3000 km southeastwards. Within Nemastomatinae Sinostoma displays plesiomorphic characters, including the long, basic bulb of the truncus shaft and the extremely short glans of penis, armed with short robust spines. Sinostoma may represent a relict line in the early evolution of nemastomatine harvestmen.