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Following extensive field work in Thailand (2006–2008), this paper reviews the taxonomic status of the three species of the Hipposideros bicolor group. Based on morphometric characters and acoustic data, the two phonic types, H. bicolor (131 kHz) and H. bicolor (142 kHz) are treated as distinct species. Hipposideros bicolor (131 kHz) is designated as H. bicolor; H. bicolor (142 kHz) is provisionally designated as H. atrox. The morphometric characters, acoustic data, and geographical distribution of H. pomona are also reviewed. The diagnostic characters of these frequently confused taxa are discussed, with a detailed study of the external, cranio-dental, and bacular morphology, and acoustic features. New data on the conservation status, distribution and ecology of these three species are included.
Since its discovery, the taxonomic status of the only species of Kerivoula (Chiroptera: Vespertilionidae: Kerivoulinae) to be found on Taiwan has been confused. Previous studies have assigned this species to either Kerivoula hardwickii or K. titania, both of which occur on continental SE Asia. This uncertainty supports repeated suggestions in the literature that specimens of K. hardwickii collected and/or sampled across SE Asia are likely to represent multiple cryptic taxa. To address these issues, we combined new and existing data from the genus Kerivoula on Taiwan and continental Asia, and performed diagnostic analyses in steps. First, phylogenetic reconstructions based on mitochondrial and nuclear DNA revealed a well-supported group comprising all taxa currently recognized as K. hardwickii, together with the Taiwanese Kerivoula and Kerivoula kachinensis to the exclusion of all other congeneric species. Second, focusing on all members of this monophyletic clade (i.e., K. hardwickii complex) together with K. titania, we used multivariate statistical methods to separate taxa based on morphometric data. Our results provide strong evidence that among these bats, the Taiwanese Kerivoula is a new species that also occurs on continental Asia, for which we provide a formal description and name. In addition, we show that the subspecies K. hardwickii depressa should be elevated to species status. We discuss our findings and the caveats of this and similar studies.
Variation in the acoustic structure of bat echolocation calls can often provide sufficient information for reliable and efficient species identification. The aim of this study was to investigate the use of echolocation call structure to identify a number of bats in the families Vespertilionidae, Emballonuridae, Nycteridae and Megadermatidae from Thailand. These species typically emit echolocation calls with a frequency-modulated (FM) sweep dominating part or all of their calls. A total of 510 echolocation calls from free-flying individuals were recorded throughout Thailand. According to the frequency-time spectra, these calls were categorized into four types: broadband FM (eight species), narrowband FM (seven species), long multiharmonic (four species) and short multiharmonic (three species). Discriminant function analysis was used to classify calls from individual bats to species. Correct classification levels were 85.9% for individuals emitting broadband FM calls (six species with adequate sample sizes), 70.4% for narrowband FM bats (seven species), 84.4% for species emitting long multiharmonic calls (four species) and 96.7% for species emitting short multiharmonic calls (two species with adequate sample sizes). However, classification rates were often low at the species level. Acoustic identification of bats emitting FM calls should be approached with caution in species-rich communities, in contrast with the identification of rhinolophoid bats where many species use distinctive constant frequencies that can facilitate identification, and provides a basis for rapid acoustic surveys of large areas in Thailand, and potentially other parts of Southeast Asia.
Kerivoula kachinensis is reported for the first time from Cambodia, Lao PDR and Thailand. In April, 2005 and January, 2006, three individuals were collected in deciduous dipterocarp forest, near bamboo, in the Seima Biodiversity Conservation Area, Mondul Kiri Province, Cambodia. In September, 2007, two individuals were collected in lower montane forest, which included some mixed deciduous forest, bamboo groves and banana trees, in the hills of Phu Suan Sai National Park, Loei Province, and a single individual was collected in mixed deciduous forest, near bamboo, in the Nam Nao National Park, Petchabun Province, Thailand. In 1996–1998, seven specimens were collected from five localities in north, central and southern Lao PDR; most were associated with evergreen forest at altitudes between 150–800 m a.s.l. The species appears to be relatively widespread in continental Southeast Asia. Locally common, it is probably not currently at risk.
Bats are an important component of biodiversity in Southeast Asia, and are key indicators of habitat quality. Acoustic analysis of echolocation calls not only allows the identification of bat species that are difficult to capture, but also allows for rapid and standardised ways to survey and monitor bats over large areas. However keys based on call parameters must also account for geographic variation in call parameters, and consider any effects of morphology and sex on call frequency. Presence-only modelling can predict likely geographic locations of specific taxa, and can used to refine decision making so that species unlikely to occur in a specific region can be omitted from more localised acoustic libraries. Here we develop an acoustic library for the echolocation calls of rhinolophoid bats in Thailand, and use presence-only modelling (Maxent) to explore how species with similar calls in a library developed at the national level can sometimes be largely allopatric, and hence identifiable, once geographic range is predicted. Presence-only modelling can also be used to explore whether species with similar calls adjust call frequency in likely areas of sympatry. We analysed calls from fourteen species of rhinolophid and twelve hipposiderid species from Thailand. Calls from a further three rhinolophid and one hipposiderid species are also described but not analysed statistically because of small sample sizes. Even without considering geographic variation, 69% of rhinolophid (14 species with a minimum of five individuals/ species) and 91% of hipposiderid calls (12 species) could be classified successfully to species using two call parameters (frequency of most energy (FMAXE) and duration) in a discriminant function analysis. Most of the discrimination was achieved because species often utilised different frequency bands. Morphology can also affect call frequency both across and within species. In both rhinolophids and hipposiderid families there was a negative relationship between FMAXE and forearm length. Within species, FMAXE was negatively related to forearm length in Rhinolophus microglobosus, R. pusillus and R. thomasi, and positively related to forearm length in R. affinis and R. pearsonii. Male R. pusillus called at higher frequencies than females, although there was no sexual size dimorphism in forearm length. Call frequency was negatively related to relative humidity in R. pusillus, suggesting that bats called at lower frequencies in humid environments to counter increases in atmospheric attenuation. Presence-only modelling was used to show that some species with similar call frequencies (e.g., R. lepidus and R. microglobosus; R. yunanensis and R. trifoliatus are predicted to occur largely in allopatry, and so could be identified reliably in particular parts of the country. Presenceonly modeling can assist in predicting areas of overlap between species with similar call frequencies. Other species (e.g., R. malayanus, R. coelophyllus) overlap in frequency when data from all of Thailand are combined, but seem to avoid call overlap when syntopic. Hence acoustic identification can be improved by taking into account local distribution patterns and patterns of species coexistence. The creation of call libraries at a local scale would have extensive potential as a resource to monitor changes in species distributions through time.
Until recently, the taxon Murina cyclotis was considered to be a widespread species, albeit one that exhibited considerable individual, sexual and geographical variation. Subsequently however, it was recognised that this taxon was in fact a complex of species. As such, in 2012, two larger forms were recognised as separate and distinct species, namely: M. peninsularis in the Sunda region and M. fionae in Laos and Vietnam. In the current paper, a new cryptic species of the cyclotis-complex is described from peninsular Thailand based on a combination of external, craniodental and genetic differences. In addition, the population previously referred to M. cyclotis from the Nicobar Islands is described as a new subspecies of this new species. Despite this work and the research of others, the taxonomy of M. cyclotis still requires further study. The description of M. peninsularis is emended and the extensive variation in its morphological characters is addressed. The diagnostic characters of each taxon, as well as the additional data on ecology, zoogeography, distribution, echolocation and genetics, where available, are summarised and discussed.
Based on recent field surveys in Thailand, Myanmar, and northern peninsular Malaysia, this paper reviews the taxonomy, morphometric and acoustic characters, distribution and ecology of the little known, globally endangered species Hipposideros halophyllus. It lists nine new localities records, including the first from northern and peninsular Thailand, which represent a substantial increase in the species' known range; it confirms the record from northern peninsular Malaysia. In addition, it provides further information on two other small species of the Hipposideros bicolor group, H. ater and H. cineraceus.
The taxon Hipposideros nicobarulae is elevated to a distinct species and an amended description is included. It is geographically restricted to the Nicobar Islands in the Andaman Sea where it has been found roosting in caves and deserted buildings. It is differentiated by size and external, cranial and dental morphology from Hipposideros ater from the Indian Subcontinent. Both taxa are distinguished from a new species, Hipposideros sp. nov., herein described from the Rakhine coast of western Myanmar, with referred material from Tanintharyi Division in southern Myanmar. In all cases, individuals were found roosting in human habitation. Information on the conservation status, distribution and ecology of the three species are included.
A new species of woolly horseshoe bat in the Rhinolophus trifoliatus species group is described from Sabah in Malaysian Borneo. Two specimens from Central and West Kalimantan, Indonesia are referred to this species. A fourth specimen from western Thailand is referable to this species but on the basis of ~10% genetic divergence at the cytochrome oxidase-I gene is described as a separate subspecies. Morphologically and acoustically the two subspecies are similar. With a forearm length of 52.90–54.70 mm, a skull length of 24.27–26.57 mm and a call frequency of 49.2–50.0 kHz, the new species overlaps in size and call frequency with the sympatric R. trifoliatus. However, it differs significantly in having a dark noseleaf and a uniformly dark brown pelage, resembling, but being intermediate in size between R. sedulus and R. luctus, which have a skull length of 18.99–20.17 and 26.35–32.07 mm, respectively. It also differs from R. trifoliatus in the shape and size of the rostral inflation. It can be distinguished from R. beddomei (forearm length 55.00–63.44 mm) and R. formosae (forearm length 53.85–62.40 mm), which are endemic to the Indian Subcontinent and Taiwan, respectively, by its relatively smaller body size. Acoustic and genetic data are included in the comparison between the species. Both character states support the conclusions based on morphology. Further surveys in intact evergreen forest together with a re-examination of museum specimens may reveal that this species is widespread in Southeast Asia.
A new species of Rhinolophus in the pusillus group is described from Ratchaburi, Kamphaeng Phet and Loei Provinces where it was found in evergreen forest at elevations ranging from 550 to 1,320 m a.s.l. It is distinguished from R. shortridgei and other similar species in the same group by its broad, parallel-sided sella, which is squared-off at the tip, relatively large body size with a forearm length of 42.2–44.1 mm, and bulbous rostral swellings. The echolocation frequency from hand-held individuals is 84.1–93.0 kHz. Bayesian analyses of a 654 bp of cytochrome oxidase subunit I (DNA barcode), and an 878 bp fragment of cytochrome b also support differences at the species level. Three specimens from Loaung Namtha, Lao PDR are referred to this new species based on DNA barcodes. Based on distinctive DNA barcodes and craniodental morphology, the taxon refulgens, is here regarded as a separate species from R. lepidus. Morphological comparisons between similar species are discussed and notes on ecology included.
Recent field studies have provided new data for a review of the taxonomy, acoustic characters, distribution, and ecology of two often confused rhinolophid species, which have essentially parapatric distributions in continental Southeast Asia. Rhinolophus coelophyllus is widespread ranging from northern Myanmar to northern Malaysia, eastern Thailand and provisionally western Lao PDR. R. shameli is restricted to eastern Thailand, Cambodia, Lao PDR, and central and southern Vietnam. There are well defined differences in skull morphology, size, and echolocation call frequency, which discriminate between the two taxa.
Rhinolophus affinis sensu lato is a widespread bat species in South and Southeast Asia which shows considerable geographical variation in its morphology, echolocation call frequencies and genetics. The taxonomic status of the taxon in the Sundaic subregion remains uncertain however as the limited studies to date have been largely based on morphology. The aim of the present study was to determine the taxonomic status of subspecific forms recognized in the subregion and to evaluate phylogeographic distinctiveness between those occurring in Borneo and the Malay Peninsula using genetic, morphological and acoustic datasets. Two forms were confirmed: R. a. nesites from Borneo and R. a. superans from the peninsula. The previous recognition of a population from southernmost Sumatra as R. a. superans was not supported, however, as this form is likely R. a. affinis. Genetic divergence between these three forms is rather deep and is estimated to have occurred during the arid climatic period of the Pleistocene when suitable habitats were reduced to isolated pockets. Our results support the phylogeographic distinctiveness hypothesis as R. affinis sensu lato shows discrete affinities between Borneo and the Malay Peninsula. Discovery of new forms of R. affinis is likely with greater sampling effort throughout the region. Our study also demonstrates the importance of employing multiple datasets in taxonomic evaluations, as the use of morphological and/or acoustic datasets alone could lead to erroneous conclusions.
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