Human disturbance increases spatiotemporal associations among mountain forest terrestrial mammal species.

Spatial and temporal associations between sympatric species underpin biotic interactions, structure ecological assemblages, and sustain ecosystem functioning and stability. However, the resilience of interspecific spatiotemporal associations to human activity remains poorly understood, particularly in mountain forests where anthropogenic impacts are often pervasive. Here, we applied context-dependent Joint Species Distribution Models to a systematic camera-trap survey dataset from a global biodiversity hotspot in eastern Himalayas to understand how prominent human activities in mountain forests influence species associations within terrestrial mammal communities. We obtained 10,388 independent detections of 17 focal species (12 carnivores and five ungulates) from 322 stations over 43,163 camera days of effort. We identified a higher incidence of positive associations in habitats with higher levels of human modification (87%) and human presence (83%) compared to those located in habitats with lower human modification (64%) and human presence (65%) levels. We also detected a significant reduction of pairwise encounter time at increasing levels of human disturbance, corresponding to more frequent encounters between pairs of species. Our findings indicate that human activities can push mammals together into more frequent encounters and associations, which likely influences the coexistence and persistence of wildlife, with potential far-ranging ecological consequences.

Disassociation of social and sexual partner relationships in a gibbon population with stable one-male two-female groups.

Adult males living in a one-male multi-female social group are expected to try to monopolize copulations with resident females to increase reproductive fitness. Gibbons have traditionally been described as living in monogamous groups, with the sole resident adult male assumed to sire all of the group's offspring. Here, we used microsatellite analyses and behavioral observations to examine rates of extra-group paternity (EGP) over 16 years in a population of crested gibbons (Nomascus concolor) that form stable and long-term one-male two-female social units. Forty percent of offspring (N = 14) were sired by extra-group males. To understand this high level of EGP, we tested whether inbreeding avoidance was related to EGP. Females who engaged in EGP did not show larger pairwise relatedness with their resident male compared to females who did not engage in EGP. Nevertheless, the standardized heterozygosity of EGP offspring was significantly higher than for offspring sired by the group's resident male. These results provide partial support for the inbreeding avoidance hypothesis. It appears that resident male crested gibbons are unable to monopolize resident females' matings. Our results indicate that long-term social partners are often distinct from sexual partners in this population. Clearly, the breeding system of crested gibbons is more flexible than previously thought, indicating a need for integrating long-term behavioral data and genetic research to re-evaluate gibbon social and sexual relationships derived from concepts of monogamy and pair-bonding.

Multitrophic diversity and biotic associations influence subalpine forest ecosystem multifunctionality.

Biodiversity across multiple trophic levels is required to maintain multiple ecosystem functions. Yet it remains unclear how multitrophic diversity and species interactions regulate ecosystem multifunctionality. Here, combining data from 9 different trophic groups (including trees, shrubs, herbs, leaf mites, small mammals, bacteria, pathogenic fungi, saprophytic fungi, and symbiotic fungi) and 13 ecosystem functions related to supporting, provisioning, and regulating services, we used a multitrophic perspective to evaluate the effects of elevation, diversity, and network complexity on scale-dependent subalpine forest multifunctionality. Our results demonstrated that elevation and soil pH significantly modified species composition and richness across multitrophic groups and influenced multiple functions simultaneously. We present evidence that species richness across multiple trophic groups had stronger effects on multifunctionality than species richness at any single trophic level. Moreover, biotic associations, indicating the complexity of trophic networks, were positively associated with multifunctionality. The relative effects of diversity on multifunctionality increased at the scale of the larger community compared to a scale accounting for neighboring interactions. Our results highlight the paramount importance of scale- and context-dependent multitrophic diversity and interactions for a better understanding of mountain ecosystem multifunctionality in a changing world.

Echolocation in soft-furred tree mice.

Echolocation is the use of reflected sound to sense features of the environment. Here, we show that soft-furred tree mice (Typhlomys) echolocate based on multiple independent lines of evidence. Behavioral experiments show that these mice can locate and avoid obstacles in darkness using hearing and ultrasonic pulses. The proximal portion of their stylohyal bone fuses with the tympanic bone, a form previously only seen in laryngeally echolocating bats. Further, we found convergence of hearing-related genes across the genome and of the echolocation-related gene prestin between soft-furred tree mice and echolocating mammals. Together, our findings suggest that soft-furred tree mice are capable of echolocation, and thus are a new lineage of echolocating mammals.

Phylogenetic and morphological significance of an overlooked flying squirrel (Pteromyini, Rodentia) from the eastern Himalayas with the description of a new genus.

The flying squirrels (Pteromyini, Rodentia) are the most diverse and widely distributed group of gliding mammals. Taxonomic boundaries and relationships within flying squirrels remain an area of active research in mammalogy. The discovery of new specimens of Pteromys ( Hylopetes) leonardi Thomas, 1921 previously considered a synonym of Hylopetes alboniger, in Yunnan Province, China allowed a morphological and genetic reassessment of the status of this taxon. Phylogenetic reconstruction was implemented using sequences of two mitochondrial (12S ribosomal DNA and 16S ribosomal DNA) and one nuclear (interphotoreceptor retinoid-binding protein) gene fragments. Morphological assessments involved examinations of features preserved on skins, skulls, and penises of museum specimens, supplemented with principal component analysis of craniometric data. Together these assessments revealed that this taxon should be recognized not only as a distinct species, and should also be placed within a new genus, described here as Priapomys.

Historical biogeography, systematics, and integrative taxonomy of the non-Ethiopian speckled pelage brush-furred rats (Lophuromys flavopunctatus group).

BACKGROUND: The speckled-pelage brush-furred rats (Lophuromys flavopunctatus group) have been difficult to define given conflicting genetic, morphological, and distributional records that combine to obscure meaningful accounts of its taxonomic diversity and evolution. In this study, we inferred the systematics, phylogeography, and evolutionary history of the L. flavopunctatus group using maximum likelihood and Bayesian phylogenetic inference, divergence times, historical biogeographic reconstruction, and morphometric discriminant tests. We compiled comprehensive datasets of three loci (two mitochondrial [mtDNA] and one nuclear) and two morphometric datasets (linear and geometric) from across the known range of the genus Lophuromys. RESULTS: The mtDNA phylogeny supported the division of the genus Lophuromys into three primary groups with nearly equidistant pairwise differentiation: one group corresponding to the subgenus Kivumys (Kivumys group) and two groups corresponding to the subgenus Lophuromys (L. sikapusi group and L. flavopunctatus group). The L. flavopunctatus group comprised the speckled-pelage brush-furred Lophuromys endemic to Ethiopia (Ethiopian L. flavopunctatus members [ETHFLAVO]) and the non-Ethiopian ones (non-Ethiopian L. flavopunctatus members [NONETHFLAVO]) in deeply nested relationships. There were distinctly geographically structured mtDNA clades among the NONETHFLAVO, which were incongruous with the nuclear tree where several clades were unresolved. The morphometric datasets did not systematically assign samples to meaningful taxonomic units or agree with the mtDNA clades. The divergence dating and ancestral range reconstructions showed the NONETHFLAVO colonized the current ranges over two independent dispersal events out of Ethiopia in the early Pleistocene. CONCLUSION: The phylogenetic associations and divergence times of the L. flavopunctatus group support the hypothesis that paleoclimatic impacts and ecosystem refugia during the Pleistocene impacted the evolutionary radiation of these rodents. The overlap in craniodental variation between distinct mtDNA clades among the NONETHFLAVO suggests unraveling underlying ecomorphological drivers is key to reconciling taxonomically informative morphological characters. The genus Lophuromys requires a taxonomic reassessment based on extensive genomic evidence to elucidate the patterns and impacts of genetic isolation at clade contact zones.

Multilocus phylogeny suggests a distinct species status for the Nepal population of Assam macaques ( Macaca assamensis): implications for evolution and conservation.

Phylogenetic relationships within the sinica-group of macaques based on morphological, behavioral, and molecular characteristics have remained controversial. The Nepal population of Assam macaques ( Macaca assamensis) (NPAM), the westernmost population of the species, is morphologically distinct but has never been used in phylogenetic analyses. Here, the phylogenetic relationship of NPAM with other congeners was tested using multiple mitochondrial and Y-chromosomal loci. The divergence times and evolutionary genetic distances among macaques were also estimated. Results revealed two major mitochondrial DNA clades of macaques under the sinica-group: the first clade included M. thibetana, M. sinica, and eastern subspecies of Assam macaque ( M. assamensis assamensis); the second clade included M. radiata together with species from the eastern and central Himalaya, namely, M. leucogenys, M. munzala, and NPAM. Among the second-clade species, NPAM was the first to diverge from the other members of the clade around 1.9 million years ago. Our results revealed that NPAM is phylogenetically distinct from the eastern Assam macaques and closer to other species and hence may represent a separate species. Because of its phylogenetic distinctiveness, isolated distribution, and small population size, the Nepal population of sinica-group macaques warrants detailed taxonomic revision and high conservation priority.

Differences in Quaternary co-divergence reveals community-wide diversification in the mountains of southwest China varied among species.

The mountains of southwest China (MSWC) is a biodiversity hotspot with highly complex and unusual terrain. However, with the majority of studies focusing on the biogeographic consequences of massive mountain building, the Quaternary legacy of biodiversity for the MSWC has long been overlooked. Here, we took a statistical comparative phylogeography approach to examine factors that shaped community-wide diversification. With data from 30 vertebrate species, the results reveal spatially concordant genetic structure, and temporally clustered co-divergence events associated with river barriers during severe glacial cycles. This indicates the importance of riverine barriers in the phylogeographic history of the MSWC vertebrate community. We conclude that the repeated glacial cycles are associated with co-divergences that are themselves structured by the heterogeneity of the montane landscape of the MSWC. This orderly process of diversification has profound implications for conservation by highlighting the relative independence of different geographical areas in which some, but not all species in communities have responded similarly to climate change and calls for further comparative phylogeographic investigations to reveal the connection between biological traits and divergence pulses in this biodiversity hotspot.

The forty-year journey of Zoological Research: advancing with the times.

Since its establishment in 1980, Zoological Research ( ZR) has walked with many scientists during their academic careers (Yao & Zhang, 2015; Yao et al., 2019). During this joyful holiday season and hopeful new year, we are incredibly pleased to share our wonderful news with all authors, readers, editorial board members, and peer reviewers of ZR. Due to its growing academic influence over the last several years, ZR has attained its highest impact factor to date, reaching 2.638 by the end of 2019 (Q1, 12th/169 SCI journals). Furthermore, its current CiteScore has reached 4.3 (as of December 2020), a nearly 60% increase since 2019. Meanwhile, both the publishing capacity and efficiency of ZR have continued to expand. We are immensely proud of each of our efforts in promoting ZR. Of note, your enduring support and faith have helped drive the sustained growth and advancement of ZR, which continues to evolve as a prominent journal in the zoological field.

Isolated alpine habitats reveal disparate ecological drivers of taxonomic and functional beta-diversity of small mammal assemblages.

The interpretation of patterns of biodiversity requires the disentanglement of geographical and environmental variables. Disjunct alpine communities are geographically isolated from one another but experience similar environmental impacts. Isolated homogenous habitats may promote speciation but constrain functional trait variation. In this study, we examined the hypothesis that dispersal limitation promotes taxonomic divergence, whereas habitat similarity in alpine mountains leads to functional convergence. We performed standardized field investigation to sample non-volant small mammals from 18 prominent alpine sites in the Three Parallel Rivers area. We estimated indices quantifying taxonomic and functional alpha- and beta-diversity, as well as beta-diversity components. We then assessed the respective importance of geographical and environmental predictors in explaining taxonomic and functional compositions. No evidence was found to show that species were more functionally similar than expected in local assemblages. However, the taxonomic turnover components were higher than functional ones (0.471±0.230 vs. 0.243±0.215), with nestedness components showing the opposite pattern (0.063±0.054 vs. 0.269±0.225). This indicated that differences in taxonomic compositions between sites occurred from replacement of functionally similar species. Geographical barriers were the key factor influencing both taxonomic total dissimilarity and turnover components, whereas functional beta-diversity was primarily explained by climatic factors such as minimum temperature of the coldest month. Our findings provide empirical evidence that taxonomic and functional diversity patterns can be independently driven by different ecological processes. Our results point to the importance of clarifying different components of beta-diversity to understand the underlying mechanisms of community assembly. These results also shed light on the assembly rules and ecological processes of terrestrial mammal communities in extreme environments.

Influence of food availability and climate on behavior patterns of western black crested gibbons (Nomascus concolor) at Mt. Wuliang, Yunnan, China.

Food abundance and climatic factors can significantly affect the behavior of animals and constrain their activity budgets. The population of western black crested gibbons (Nomascus concolor) in Mt. Wuliang lives in montane forest and is close to the northern extreme of the distribution for gibbons (Hylobatidae). Their habitats show remarkable seasonal variation in terms of food availability, temperature, and rainfall. To understand behavioral adaptations of western black crested gibbons to different sets of ecological conditions, we examined relationships among food availability, mean temperature, rainfall, and behavior patterns by observing two groups for 1 year each. Our results revealed that activity budget was affected by food availability and mean temperature. The gibbons spent more time eating flowers when that resource was more available and spent less time moving when fruit was more available. The gibbons spent less time feeding and more time resting, and spent less feeding time on fruit and leaves when the mean temperature was lower. These results suggest that the gibbons displayed a pronounced preference for flowers as a food resource and adopted a time minimizer strategy when high-nutrient food items (i.e., fruit) were more available. In addition, the gibbons adopted an energy-conserving strategy during periods of low temperature. The flexibility of behavioral patterns in responding to food availability and temperature may potentially improve the gibbons' prospects of surviving and reproducing in a northern montane forest.

Discovery and description of a mysterious Asian flying squirrel (Rodentia, Sciuridae, Biswamoyopterus) from Mount Gaoligong, southwest China.

The flying squirrels of the tribe Pteromyini (Family Sciuridae) currently include 15 genera of which the genus Biswamoyopterus comprises two recognized species, B.biswasi Saha, 1981 and B.laoensis Sanamxay et al., 2013. These two species were each described from only one specimen that are separated from each other by 1,250 kilometres in southern Asia, where they occur in northeast India and central Lao PDR respectively. In 2017 and 2018, two specimens of Biswamoyopterus were discovered from Mount Gaoligong, west Yunnan province, southwest China (between the type locality of the two recognized species). This study aimed to evaluate the taxonomic status of these two newly acquired specimens of Biswamoyopterus by comparing their morphology with the two described species of the genus. The results of this study showed that the specimens from Yunnan province (China) differed from both B.laoensis and B.biswasi in both pelage colour and craniology, and should be recognised as a distinct species, B.gaoligongensis sp. nov., which is formally described here. This study contributes to the understanding of the flying squirrels of southern Asia and identifies an additional species that appears to be endemic to southwest China; however, more research is required to provide details of its ecology, distribution, and conservation status.

Small mammal diversity of Mt. Kenya based on carnivore fecal and surface bone remains.

Ecological dynamics and faunal diversity documentation is normally conducted by direct observation and trapping of live animals. However, surveys of carnivore scat prey and surface bone remains, which are relatively inexpensive, can provide complementary data that expand carnivore diet breadth and may improve accuracy regarding inferences of the ecological dynamics of a given ecosystem. We used this inexpensive method to document species diversity variation with elevation on the leeward (Sirimon) and windward (Chogoria) areas of Mt. Kenya. Bone and fecal specimens were opportunistically collected by walking 2 km in opposite directions from transect points selected at 200-m intervals along the elevational gradient of the study areas. We collected a total of 220 carnivore fecal and owl pellet specimens from both study sites, which were mainly deposited by the spotted hyena (Crocuta crocuta), leopard (Panthera pardus), serval (Leptailurus serval), genet (Genetta sp.), and Mackinder's Cape owl (Bubo capensis mackinderi). Serval scats were the most common, followed by those of the spotted hyena. Scats and bones were found at the lowest density at the lowest elevations, peaked at mid-higher elevations, and then declined at the highest elevations. Based on skeletal analysis only, there were more species in Sirimon (19) than in Chogoria (12). Small fauna (rodents to duiker size bovids) formed the bulk of the identified remains, representing 87.9% of the Sirimon fauna and 90.9% of the Chogoria fauna. The genus Otomys was the dominant prey of the owl and serval in both sites. Three giraffe teeth were found at 3 500 m a.s.l. in Chogoria on the edge of Lake Ellis, suggesting that it is an occasional visitor to such high elevations. This study underscores the value of fecal and bone surveys in understanding the diet and diversity of mammals in ecological ecosystems, but such surveys should be complemented with analysis of hairs found in scats to obtain a more complete list of carnivore prey at Mt. Kenya.

An annotated checklist of mammals of Kenya.

Kenya has a rich mammalian fauna. We reviewed recently published books and papers including the six volumes of Mammals of Africa to develop an up-to-date annotated checklist of all mammals recorded from Kenya. A total of 390 species have been identified in the country, including 106 species of rodents, 104 species of bats, 63 species of even-toed ungulates (including whales and dolphins), 36 species of insectivores and carnivores, 19 species of primates, five species of elephant shrews, four species of hyraxes and odd-toed ungulates, three species of afrosoricids, pangolins, and hares, and one species of aardvark, elephant, sirenian and hedgehog. The number of species in this checklist is expected to increase with additional surveys and as the taxonomic status of small mammals (e.g., bats, shrews and rodents) becomes better understood.

Diversity and distribution patterns of non-volant small mammals along different elevation gradients on Mt. Kenya, Kenya.

The distribution of small mammals in mountainous environments across different elevations can provide important information on the effects of climate change on the dispersal of species. However, few studies conducted on Afromontane ecosystems have compared the altitudinal patterns of small mammal diversity. We investigated the species diversity and abundance of non-volant small mammals (hereafter 'small mammals') on Mt. Kenya, the second tallest mountain in Africa, using a standard sampling scheme. Nine sampling transects were established at intervals of 200 m on the eastern (Chogoria) and western (Sirimon) slopes. A total of 1 905 individuals representing 25 species of small mammals were trapped after 12 240 trap-nights. Abundance was highest at mid-elevations on both slopes. However, species richness and distribution patterns differed between the two slopes. More species were recorded on Chogoria (24) than on Sirimon (17). On Chogoria, species richness was higher at mid-high elevations, with a peak at mid-elevation (2 800 m a.s.l.), whereas species richness showed little variation on the Sirimon slope. These results indicate that patterns of species diversity can differ between slopes on the same mountain. In addition, we extensively reviewed literature on Mt. Kenya's mammals and compiled a comprehensive checklist of 76 mammalian species. However, additional research is required to improve our understanding of small mammal diversity in mountain habitats in Africa.

A new genus of Asiatic short-tailed shrew (Soricidae, Eulipotyphla) based on molecular and morphological comparisons.

Blarinellini is a tribe of soricine shrews comprised of nine fossil genera and one extant genus. Blarinelline shrews were once widely distributed throughout Eurasia and North America, though only members of the Asiatic short-tailed shrew genus Blarinella currently persist (mostly in southwestern China and adjacent areas). Only three forms of Blarinella have been recognized as either species or subspecies. However, recent molecular studies indicated a strikingly deep divergence within the genus, implying the existence of a distinct genus-level lineage. We sequenced the complete mitochondrial genomes and one nuclear gene of three Asiatic short-tailed and two North American shrews and analyzed them morphometrically and morphologically. Our molecular analyses revealed that specimens ascribed to B. griselda formed two deeply diverged lineages, one a close relative to B. quadraticauda, whereas the other - comprised of topotype specimens from southern Gansu - diverged from other Blarinella in the middle Miocene (ca. 18.2 million years ago (Ma), 95% confidence interval=13.4-23.6 Ma). Although the skulls were similarly shaped in both lineages, we observed several diagnostic characteristics, including the shape of the upper P4. In consideration of the molecular and morphological evidence, we recognize B. griselda as the sole species of a new genus, namely, Pantherina gen. nov. Interestingly, some characteristics of Pantherina griselda are more similar to fossil genera, suggesting it represents an evolutionarily more primitive form than Blarinella. Recognition of this new genus sheds light on the systematics and evolutionary history of the tribe Blarinellini throughout Eurasia and North America.

Karyotypes of field mice of the genus Apodemus (Mammalia: Rodentia) from China.

Karyotypes of four Chinese species of field mice of the genus Apodemus were examined, including Apodemus chevrieri (diploid chromosome number, 2n=48, fundamental number of autosomal arms, FNa=56), A. draco (2n=48, FNa=48), A. ilex (2n=48, FNa=48), and A. latronum (2n=48, FNa=48). Karyotypes of A. chevrieri, A. draco, and A. ilex are reported here for the first time, providing useful information for their species taxonomy. Determining the karyotypes of all species of Apodemus in Asia, both in this and previous studies, provides a solid overview of the chromosome evolution and species differentiation of the genus in East Asia. In addition to allopatric speciation, chromosome rearrangements likely played an important role in the formation of the four Apodemus species groups as well as speciation within each group in East Asia. For example, increased centromeric heterochromatin in A. latronum may have contributed to the post-mating reproductive isolation from the A. draco-A. ilex-A. semotus clade.