Phylogeography of the Asian lesser white-toothed shrew, Crocidura shantungensis, in East Asia: role of the Korean Peninsula as refugium for small mammals.

Many peninsulas in the temperate zone played an important role as refugia of various flora and fauna, and the southern Korean Peninsula also served as a refugium for many small mammals in East Asia during the Pleistocene. The Asian lesser white-toothed shrew, Crocidura shantungensis, is a widely distributed species in East Asia, and is an appropriate model organism for exploring the role of the Korean Peninsula as a refugium of small mammals. Here, we investigated phylogenetic relationships and genetic diversity based on the entire sequence of the mitochondrial cytochrome b gene (1140 bp). A Bayesian tree for 98 haplotypes detected in 228 C. shantungensis specimens from East Asia revealed the presence of three major groups with at least 5 subgroups. Most haplotypes were distributed according to their geographic proximity. Pairwise FST's and analysis of molecular variance (AMOVA) revealed a high degree of genetic differentiation and variance among regions as well as among populations within region, implying little gene flow among local populations. Genetic evidence from South Korean islands, Jeju-do Island of South Korea, and Taiwan leads us to reject the hypothesis of recent population expansion. We observed unique island-type genetic characteristics consistent with geographic isolation and resultant genetic drift. Phylogeographic inference, together with estimates of genetic differentiation and diversity, suggest that the southern most part the Korean Peninsula, including offshore islands, played an important role as a refugium for C. shantungensis during the Pleistocene. However, the presence of several refugia on the mainland of northeast Asia is also proposed.

Historical Relationships among Wild Boar Populations of the Ryukyu Archipelago and Other Eurasian regions, as Inferred from Mitochondrial Cytochrome b Gene Sequences.

The Ryukyu wild boar (Sus scrofa riukiuanus) is an endemic, morphologically defined subspecies of the Eurasian wild boar (S. scrofa) found on five islands of the Ryukyu Archipelago (a group of small islands stretching from mainland Japan to Taiwan). Two hypothetical scenarios have been proposed regarding the origin of the current Ryukyu wild boar populations: 1) natural dispersal and 2) transportation and subsequent release by prehistoric humans. To test these two hypotheses, we compared the mitochondrial cytochrome b gene sequence (1140 base pairs) in 352 individual wild boar samples that included representatives of all five insular populations of the Ryukyu wild boar and populations of other conspecific subspecies in insular East and Southeast Asia and the Eurasian Continent. A total of 68 haplotypes were recognized, of which 12 were unique to the Ryukyu wild boar populations. The results of Bayesian phylogenetic analyses supported monophyly of the five Ryukyu populations (posterior probability value of 92), confirming the validity of the subspecies as a natural group. Coalescent analysis estimated the divergence times between the Ryukyu wild boar and the other conspecific subspecies as 144-465 thousand years ago (Kya), with a 95% HPD (highest posterior density) range of 51-837 Kya, and with no significant migration. Taking the broadly accepted date of initial human migration to the Ryukyus (no earlier than 50 Kya) into consideration, our results strongly suggest that the ancestral form of the Ryukyu wild boar first entered the Ryukyu Archipelago by natural dispersal prior to the arrival of prehistoric humans.

Molecular phylogenetic reconstructions identify East Asia as the cradle for the evolution of the cosmopolitan genus Myotis (Mammalia, Chiroptera).

Sequences of the mitochondrial cytochrome b (1140 bp) and nuclear Rag 2 (1148 bp) genes were used to assess the evolutionary history of the cosmopolitan bat genus Myotis, based on a worldwide sampling of over 88 named species plus 7 species with uncertain nomenclature. Phylogenetic reconstructions of this comprehensive taxon sampling show that most radiation of species occurred independently within each biogeographic region. Our molecular study supports an early divergence of species from the New World, where all Nearctic and Neotropical species plus a lineage from the Palaearctic constitute a monophyletic clade, sister to the remaining Old World taxa. The major Old World clade includes all remaining Eurasian taxa, most Oriental species, one Oceanian, and all Ethiopian species. Another lineage, including M. latirostris from Taiwan, appears at the base of these two major biogeographic clades and, because it bears nyctalodont molars, could be considered as a distinct genus. However, this molar configuration is also found in crown-group species, indicating that these dental characters are variable in the genus Myotis and may confound interpretation of the fossil record. Molecular datings suggest an origin of all recent Myotis in the early Miocene (about 21MYA with 95% highest posterior density interval 23-20MYA). This period was characterized by a global climatic cooling that reduced the availability of tropical habitats and favoured the development of more temperate vegetation. This sharp climatic change might have triggered the evolution of Myotis in the Northern continents, because Myotis ancestors seem to have been well adapted and successful in such temperate habitats. Ancestral area reconstructions based on the molecular phylogeny suggest that the eastern portion of the Asian continent was an important center of origin for the early diversification of all Myotis lineages, and involved relatively few subsequent transcontinental range expansions.

Morphometric variation in the pusillus group of the genus Rhinolophus (Mammalia: Chiroptera: Rhinolophidae) in East Asia.

Based on 203 specimens belonging to the Rhinolophus "pusillus group" (Mammalia: Chiroptera: Rhinolophidae), univariate and multivariate morphometric analyses using 19 characters were performed to assess the confused species taxonomy. The results indicated that R. pusillus (including calidus, parcus, and szechuanus) in the continental region and Hainan Island of China and "R. cornutus" in Japan are morphologically divergent species. Rhinolophus cornutus should be further split into R. cornutus (including orii, pumilus, and miyakonis) in the main islands of Japan, the Amami and Okinawa Group of the central Ryukyu Archipelago, and Miyako Group of the southern Ryukyus; and R. perditus and R. imaizumii from the Yaeyama Group in the southern Ryukyus. Rhinolophus monoceros from Taiwan is morphologically more similar to species in Japan than to R. pusillus. In addition to R. pusillus, another form that is morphologically similar to species in Japan was recognized from Langzhong in Sichuan Province; this may represent an undescribed species, and further examination is necessary to determine its taxonomic status. Specimens from Guang'an in Sichuan Province, China, are also different from the others, and are characterized by the smallest skull size. Although further studies are required, these specimens were tentatively identified as R. subbadius.

A multi-locus phylogeny of Nectogalini shrews and influences of the paleoclimate on speciation and evolution.

Nectogaline shrews are a major component of the small mammalian fauna of Europe and Asia, and are notable for their diverse ecology, including utilization of aquatic habitats. So far, molecular phylogenetic analyses including nectogaline species have been unable to infer a well-resolved, well-supported phylogeny, thus limiting the power of comparative evolutionary and ecological analyses of the group. Here, we employ Bayesian phylogenetic analyses of eight mitochondrial and three nuclear genes to infer the phylogenetic relationships of nectogaline shrews. We subsequently use this phylogeny to assess the genetic diversity within the genus Episoriculus, and determine whether adaptation to aquatic habitats evolved independently multiple times. Moreover, we both analyze the fossil record and employ Bayesian relaxed clock divergence dating analyses of DNA to assess the impact of historical global climate change on the biogeography of Nectogalini. We infer strong support for the polyphyly of the genus Episoriculus. We also find strong evidence that the ability to heavily utilize aquatic habitats evolved independently in both Neomys and Chimarrogale+Nectogale lineages. Our Bayesian molecular divergence analysis suggests that the early history of Nectogalini is characterized by a rapid radiation at the Miocene/Pliocene boundary, thus potentially explaining the lack of resolution at the base of the tree. Finally, we find evidence that nectogalines once inhabited northern latitudes, but the global cooling and desiccating events at the Miocene/Pliocene and Pliocene/Pleistocene boundaries and Pleistocene glaciation resulted in the migration of most Nectogalini lineages to their present day southern distribution.

Origins and founder effects on the Japanese masked palm civet Paguma larvata (Viverridae, Carnivora), revealed from a comparison with its molecular phylogeography in Taiwan.

The source areas of the Japanese populations of the masked palm civet Paguma larvata (Viverridae, Carnivora), an alien species in Japan, have not been identified. In the present study, to reveal their origins and genetic features, we determined the full mitochondrial DNA cytochrome b sequences (1,140 base-pairs) of a total of 206 individuals of P. larvata from the Honshu and Shikoku islands of Japan (186 animals) and Taiwan (20 animals), and investigated their molecular phylogeography and the genetic relationships between populations in these countries. We found that each animal from Japan exhibited one of four haplotypes (JA1, JA2, JA4, and JA5), and that JA1 and JA4 were more frequent in eastern Honshu and Shikoku-central Honshu, respectively. By contrast, six haplotypes consisting of four new types (TW1, TW2, TW3, and TW4) and the previously reported two types (JA1 and JA4) were identified from 20 animals from native populations in Taiwan. Within Taiwan, one haplotype set (JA1, TW1, and TW2) was distributed in the western region, while a second (JA4, TW3, and TW4) was found in the eastern region; these regions are separated by high mountain ranges. Our comparison of haplotype distributions strongly demonstrated that the eastern Japanese populations originated from animals of western Taiwan, and that the western Japanese populations originated from those of eastern Taiwan. In addition, the lower genetic variability and particular distribution patterns of haplotypes in Japan showed founder effects, which may have resulted from multiple introductions of P. larvata to Japan from Taiwan.

Genome-Wide Microarray Analysis Suggests Transcriptomic Response May Not Play a Major Role in High- to Low-Altitude Acclimation in Harvest Mouse (Micromys minutus).

The harvest mouse (Micromys minutus) is a small rodent species with a wide range of vertical distribution in Taiwan, extending from the sea level to 3100 m altitude. This species has recently suffered from habitat loss in high-altitude areas due to orchard cultivation, which may have resulted in mouse migration from high to low altitude. To investigate whether there is any physiological mechanism involved in altitude acclimation, rat cDNA microarray was used to compare transcriptomic patterns of the skeletal muscle tissues taken from individuals native to the high-altitude environment and those transferred to the low-altitude captive site. Of the 23,188 genes being analyzed, 47 (33 up-regulated and 14 down-regulated) were found to have differential expression (fold change > 4 or < -4, ANOVA p < 0.05). However, after multiple testing correction with a false discovery rate (FDR), only the result for Tnfrsf12a was found to be statistically significant (fold change = 13, FDR p < 0.05). The result was confirmed by quantitative polymerase chain reaction (q-PCR). The expression of Tnfrsf12a possibly relates to the skeletal muscle biology and thus can be correlated with altitude acclimation. However, finding only one gene transcript with significant alteration suggests that transcriptomic response may not play a major role in high- to low-altitude acclimation in harvest mouse.

Karyotypes of three rat species (Mammalia: Rodentia: Muridae) from Hainan Island, China, and the valid specific status of Niviventer lotipes.

The karyotypes of three rat species from Hainan Island, China, were examined. Niviventer fulvescens (Gray, 1847) had 2n=46 and FN=64, similar to the karyotypes reported for N. fulvescens from Southeast Asia, while Niviventer lotipes () had 2n=52 and FN=66, which is distinct from the known karyotypes of other Niviventer species. Niviventer lotipes was recently considered conspecific with N. tenaster (), but the two were found to have extremely different karyotypes (2n=52 and FN=66 in N. lotipes; 2n=46 and FN=54 in N. tenaster). Therefore, in this paper N. lotipes is considered a valid species for the first time; it is distinct from N. tenaster and endemic to Hainan Island, where N. lotipes is differentiated from N. fulvescens by larger body and skull sizes, a shorter tail, and darker coloration. Rattus nitidus (Hodgson, 1845) from Hainan Island had 2n=42 and FN=62, which is similar to the reported karyotypes of conspecific populations in Southeast Asia.

Molecular diversity and phylogeography of the Asian leopard cat, Felis bengalensis, inferred from mitochondrial and Y-chromosomal DNA sequences.

To investigate genetic diversity and phylogeography of the Asian leopard cat (Felis bengalensis), mitochondrial DNA (mtDNA) sequences were determined for 39 individuals from various areas. Sequences combining the complete cytochrome b gene (1,140 bp) with the partial control region (646-810 bp) were classified into 24 haplotypes: 21 types from 21 animals, one from eight animals from Tsushima Islands, one from eight animals from Iriomote Island, and one from two animals from Southeast Asia. Phylogenetic trees of the 24 haplotypes clearly showed three clades: a Northern Lineage and Southern Lineages 1 and 2. The Northern Lineage consisted of animals from Tsushima Islands, the Korean Peninsula, the continental Far East, Taiwan, and Iriomote Island. Within the Northern Lineage, genetic contacts could have occurred between geographically neighboring populations before isolation by straits. Southern Lineage 1, comprising Southeast Asian animals, showed higher genetic diversity. Southern Lineage 2 had large genetic distances from other lineages. Within the control region, the Asian leopard cats shared two to four repetitive motifs, and the number of motifs and their constitution were highly variable among individuals. The motifs were polymorphic even within individuals and could be classified into 31 types. Finally, males of mtDNA Southern Lineage 1 had either of two types of the Y-chromosomal gene ZFY, whereas all males of Northern Lineage shared only one type. Our results indicate that the diversity of southern populations is higher and that genetic differentiation among northern local populations reflects past geographical isolation.

Accumulation of trace elements used in semiconductor industry in Formosan squirrel, as a bio-indicator of their exposure, living in Taiwan.

Concentrations of 17 trace elements were analyzed using inductively coupled plasma-mass spectrometry (ICP-MS) in Formosan squirrels (Callosciurus erythraeus) of Taiwan and Japan to document trace element pollution in Taiwan. High concentrations of elements used to produce semiconductors - Ga, As, Cd, In and Tl - were found in animals captured in Miaoli County, which is the nearest site to Hsinchu City, a chief city of Taiwan's semiconductor industry. Significant correlations between Ga, As, In and Tl were found in the kidney, liver, lung and muscle tissues of Taiwanese squirrels. Hierarchical cluster analysis indicated that Ga, As, In and Tl were of the same clade, indicating that Ga, As, In and Tl were discharged from an identical origin. Molar ratios of Ga/As concentration in lungs of animals captured in Miaoli resembled those of animals after intratracheal administration of particulate gallium arsenide (GaAs). This result might indicate that the higher concentrations of Ga and As in the specimens in Miaoli resulted from atmospheric exposure to GaAs.

Molecular phylogeny and morphological revision of Myotis bats (Chiroptera: Vespertilionidae) from Taiwan and adjacent China.

In taxonomic accounts, three species of Myotis have been traditionally reported to occur on the island of Taiwan: Watase's bat (M. formosus watasei Kishida), the Formosan broad-muzzled bat (M. muricola latirostris Kishida) and the Formosan mouse-eared bat (M. adversus taiwanensis Linde). The discovery in 1997 of an unknown taxon not fitting to the description of any of these species encouraged us to re-examine more thoroughly the systematics and phylogeny of Myotis bats inhabiting Taiwan. We used a combination of morphologic and molecular methods to aid the identification of the different taxa from this island and reconstruct their phylogenetic relationships. Multivariate analyses based on 17 craniodental characters of 105 specimens caught across Taiwan and further external characters allowed us to discriminate eight taxa of Myotinae co-occurring on this island. A subset of 80 specimens were further sequenced for the cytochrome b gene (1140 bp) and subjected to phylogenetic reconstructions including representative species from adjacent China and from all main lineages of the worldwide Myotis radiation. These molecular reconstructions showed that the Myotinae from Taiwan are phylogenetically diverse and are issued from several independent clades. The genetic results were completely congruent with the phenetic groupings based on craniodental and external morphology, as each of the eight Taiwanese taxa proved to be reciprocally monophyletic. Two unnamed taxa that did not fit into any of the known species were described as species new to science. Furthermore the taxon latirostris usually associated to the Asian M. muricola, was phylogenetically and morphologically distant from any other known Myotis and was assigned here to the fossil (Miocene) genus Submyotodon. Submyotodon latirostris, M. secundus sp. n. and M. soror sp. n. are endemic species from Taiwan, whereas the other five Myotis are more widespread and also found in the mainland. An identification key is provided to ease the discrimination of these Myotinae species in Taiwan and adjacent China.

The first archaic Homo from Taiwan.

Recent studies of an increasing number of hominin fossils highlight regional and chronological diversities of archaic Homo in the Pleistocene of eastern Asia. However, such a realization is still based on limited geographical occurrences mainly from Indonesia, China and Russian Altai. Here we describe a newly discovered archaic Homo mandible from Taiwan (Penghu 1), which further increases the diversity of Pleistocene Asian hominins. Penghu 1 revealed an unexpectedly late survival (younger than 450 but most likely 190-10 thousand years ago) of robust, apparently primitive dentognathic morphology in the periphery of the continent, which is unknown among the penecontemporaneous fossil records from other regions of Asia except for the mid-Middle Pleistocene Homo from Hexian, Eastern China. Such patterns of geographic trait distribution cannot be simply explained by clinal geographic variation of Homo erectus between northern China and Java, and suggests survival of multiple evolutionary lineages among archaic hominins before the arrival of modern humans in the region.

Morphometric geographic variation in the Asian lesser white-toothed shrew Crocidura shantungensis (Mammalia, Insectivora) in East Asia.

This study investigated geographic variation in the Asian lesser white-toothed shrew, Crocidura shantungensis Miller, 1901, among a number of localities in East Asia, including the Korean Peninsula, Tsushima Island, Cheju Island, and Taiwan, using univariate and multivariate analyses of cranial morphometric characters. Sexual differences were found in Taiwanese shrews, but were not evident in those from Korea or Tsushima. The samples from Taiwan had a larger overall cranial size than those from Korea and Tsushima, and were well separated from the latter two samples in the discriminant analyses. The overall size of the Cheju shrews was also greater, but lay within the variation of those from Taiwan. Crocidura dsinezumi quelpartis Kuroda, 1934, described from Cheju Island, is actually a junior synonym of C. shantungensis, and is not related to C. dsinezumi.

Insular variation of the craniodental morphology in the Siberian weasel Mustela sibirica.

We compared craniodental morphology among 5 populations of the Siberian weasel Mustela sibirica including 2 insular ones (Tsushima and Taiwan). Skulls of the insular individuals tended to be smaller than those of continental ones. Shape differences were also detected, but not so pronounced. Considering these results, the Taiwan population should be regarded as a distinct subspecies M. s. taivana from the mainland ones. The Tsushima population may also possibly be a distinct subspecies from the mainland ones, but more detailed studies using a larger number of specimens are needed for a conclusion. The introduced population in Honshu is also differentiated from the source population. This suggests a high morphological plasticity in M. sibirica.

A mitochondrial phylogeny and biogeographical scenario for Asiatic water shrews of the genus Chimarrogale: implications for taxonomy and low-latitude migration routes.

The six species and three subspecies in the genus Chimarrogale (Soricomorpha: Soricidae) are commonly referred to as Asiatic water shrews. The Chimarrogale are the most widely distributed group of Nectogaline shrews, extending throughout the Oriental region and Japan. Because of the limited numbers of specimens available for study, the phylogenetic relationships and biogeographical history of this genus have not been comprehensively discussed. We used mitochondrial cytochrome b gene sequences to estimate phylogenetic relationships and divergence times among four Chimarrogale species, including all three subspecies of Chimarrogale himalayica. We also conducted a species delimitation analysis and tested two alternative migration scenarios in Asia through species distribution modeling and a reconstruction of the ancestral distribution. Here, we present the first proposed hypothesis regarding the Asiatic water shrew phylogeny and reveal ten putative species within the four recognized species. Distinct phylogenetic statuses of Chimarrogale phaeura, Chimarrogale platycephala, and Chimarrogale styani were confirmed. Chimarrogale himalayica was strongly supported as paraphyletic. We suggest that three subspecies of Chimarrogale himalayica should be reconsidered as distinct species. However, these suggestions must be considered with caution because only a single locus of a mtDNA gene was used. Four additional putative species, possibly distributed in central southwestern China and Taiwan, are currently undescribed; therefore, comprehensive morphological analyses are warranted to test their taxonomic statuses. The estimated molecular divergence times indicated that rapid speciation occurred during the early Pliocene, and current distribution patterns may have been affected by global cooling during the Pliocene/Pleistocene boundary. Reconstruction of the ancestral distribution and species distribution modeling for Asiatic water shrews revealed a low-latitude migration route over which ancestral Chimarrogale migrated from Europe via Central Asia to their current distribution. Our results demonstrated that Asiatic water shrews could have evolved throughout the low-latitude migration route from Europe to East and Southeast Asia.

Genomic analyses of the Formosan harvest mouse (Micromys minutus) and comparisons to the brown Norway rat (Rattus norvegicus) and the house mouse (Mus musculus).

The harvest mouse, Micromys minutus (MMIN), has a very wide range of distribution (from the British Isles across the Euroasian continent to Japan and Taiwan). We studied an isolated population of MMIN in Taiwan, which is at the southeastern margin of the species' geographic distribution, and compared its genetic complement with those of the same species previously reported from other geographic locations and with two model rodent species, the house mouse (Mus musculus) and the brown Norway rat (Rattus norvegicus). The diploid number (2N) of MMIN was 68, consistent with that reported for other populations. However, variations were noted in the fundamental number (FN) and the shape and banding patterns of the individual chromosomes among populations. The FN of MMIN was estimated to be 72, including 2 bi-armed autosomes, 31 one-armed autosomes, and one pair of one-armed sex chromosomes. Here, we propose the first ideogram for MMIN. C-banding, Ag-NOR, and the locations of 18S rRNA gene sequences (MMIN chromosomes no. 10, 14, 19, 29, 31, 33, and X) mapped by fluorescence in situ hybridization (FISH) are also reported. Additionally, we compared the 18S rDNA sequences and performed cross-species X chromosome painting (FISH) for M. minutus, M. musculus, and R. norvegicus. The results indicate that both genetic elements are rather conserved across species. Thus, implications for the phylogenetic position of Micromys were limited.

Identification of novel Bartonella spp. in bats and evidence of Asian gray shrew as a new potential reservoir of Bartonella.

Many studies indicated that small mammals are important reservoirs for Bartonella species. Using molecular methods, several studies have documented that bats could harbor Bartonella. This study was conducted to investigate the relationship of Bartonella spp. identified in bats and small mammals living in the same ecological environment. During May 2009 and March 2010, a total of 102 blood specimens were collected. By whole blood culture and molecular identification, a total of 6 bats, 1 rodent and 9 shrews were shown to be infected by Bartonella species. After sequencing and phylogenetic analyses of the sequences of gltA, ftsZ, rpoB and ribC genes, these specific isolates from bats were not similar to the known Bartonella species (the similarity values were less than 91.2%, 90.5%, 88.8%, and 82.2%, respectively); these isolates formed an independent clade away from other known Bartonella type strains. The Bartonella spp. isolated from small mammals, which were closely related to Bartonella tribocorum, Bartonella elizabethae, Bartonella grahamii, Bartonella rattimassiliensis and Bartonella queenslandensis, were similar to the findings in previous studies worldwide. Therefore, the results implied that the species of Bartonella strains isolated from small mammals were different from those identified in bats. Our results strongly suggested that the bat isolate could be a new Bartonella species. This study is also the first one to isolate Bartonella organisms from Asian gray shrews, Crocidura attenuata tanakae.

Karyology of eight species of bats (Mammalia: Chiroptera) from Hainan Island, China.

Karyotypes and chromosomal data are presented for eight bat species representing two families (Rhinolophidae and Vespertilionidae) from Hainan Island, China. The species investigated were Rhinolophus lepidus (2n = 62, FN = 60), R. pusillus (2n = 62, FN = 60), R. affinis (2n = 62, FN = 60), R. sinicus (2n = 36, FN = 60), Myotis horsfieldi (2n = 44, FN = 52), Pipistrellus abramus (2n = 26, FN = 44), Miniopterus australis (2n = 46, FN = 50) and M. schreibersii (2n = 46, FN = 50). The karyotype of Rhinolophus lepidus is reported for the first time.

Phylogeography of the mole-shrew (Anourosorex yamashinai) in Taiwan: implications of interglacial refugia in a high-elevation small mammal.

To test the Pleistocene interglacial refugia hypothesis with a high-elevation mammal, we studied the phylogeography of the mole-shrew (Anourosorex yamashinai) using partial mitochondrial cytochrome b gene sequences (737 bases). This shrew is endemic to Taiwan. It is mainly distributed in the highlands from 1000 to 2500 m in elevation. We examined 103 specimens from 24 localities in three mountain ranges of Taiwan and found 36 haplotypes. These haplotypes separated into two major phylogroups (Northern and Southern) plus a minor phylogroup (Houhuan) of only one haplotype. This demonstrated strong association with geography. The formation of these three phylogroups may be the result of interglacial refugia during the middle Pleistocene. Distinct sublineages were not found within each major phylogroup, suggesting that the populations (phylogroups) explosively expanded from the interglacial refugia of ancestral founder haplotypes. The present distribution pattern of haplotypes suggests that Mount Houhuan is an effective refugium in central Taiwan. It was not possible to specify the refugia for the Northern and Southern phylogroups.

Intra-individual variation in the vocalized frequency of the Taiwanese leaf-nosed bat, Hipposideros terasensis, influenced by conspecific colony members.

We examined the intra-individual variation in resting frequency of the constant-frequency component of the second harmonic of the pulse (F (rest)) over 4 years in a laboratory colony of the Taiwanese leaf-nosed bat (Hipposideros terasensis). Patterns of change in F (rest) were observed when individuals were added to or removed from the colony so that we investigated whether F (rest) was affected by neighboring colony members. F (rest) of each bat continually showed a long-term gradual change throughout the year, and all bats in the colony increased or decreased their F (rest) in the same direction as a group non-seasonally. The greatest short-term changes were observed when new bats with a relatively low F (rest) joined the colony and F (rest) of new bats converged with those of the original colony members around 8 -16 days after their introduction. Conversely, a single individual showed sudden short-term decrease in F (rest) after its isolation from other colony members. These findings strongly indicate that F (rest) is flexible according to the presence of neighboring conspecific bats. We suggest that the audio-vocal feedback for conspecific pulses appears to be involved in the short- or long-term intra-individual variation in F (rest) other than factors previously thought such as age or season.