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Insectivorous bats are major predators of nocturnal insects and have the potential to act as biological pest control agents in farmlands. The objective of the present study was to establish the diet of the guano bat, Tadarida plicata. The study was carried out at the Khao Chong Pran Cave, which houses 2.6 million bats, and is surrounded by rice fields. A total of 1,925 faecal pellets were collected from 385 bats during their morning return from January to December 2002. Faecal analysis indicated that T. plicata fed on at least nine insect orders: Homoptera (28.4%), Lepidoptera (20.8), Hemiptera (16.4), Coleoptera (14.4), Diptera (7.0), Hymenoptera (6.6), Odonata (6.0), Orthoptera (0.5) and Psocoptera (0.1). Light traps indicated that Coleoptera (41.2%), Homoptera (25.3), Hemiptera (18.8) and Diptera (12.7) were the most abundant insects in the study area. Homopterans, most of which were white-backed planthopper (Sogatella sp., Delphacidae) had the highest percentage frequency of occurrence in the bats' diet indicating that T. plicata potentially plays an important role in controlling this major crop pest. The presence of macropterous planthoppers and a large proportion of moths in its diet suggests that T. plicata feeds on windborne migrant insects at high altitude. Female bats fed significantly more on lepidopterans and coleopterans and less on damselflies than males. The diet diversity index of lactating females was higher than pregnant females. Diet did not differ significantly between the dry and rainy seasons for either sex.
We studied the foraging behaviour of the dawn bat Eonycteris spelaea, a cave roosting nectarivore widespread in SE Asia, and principal pollinator of economically important crops. We radio-tracked 17 individuals for five to 19 nights over a three month period. The bats were from three cave colonies in agricultural habitats in southern Thailand. They traveled between one and 17.9 km (fi01_307.gif ± SD: 4.4 km ± 5.07, median = 2.34) from their roosting cave to food sources. The mean home-range size of the individuals varied with the method used in its calculation from 518.4 ha (100% Minimum Convex Polygon, MCP) to 564.5 ha (100% Local Convex Hull method, LoCoH) and 460.8 ha (95% Kernel density estimation, KDE). The mean size of foraging areas used by the bats also varied according to the method of calculation from 14.26 ha (100% MCP), 13.25 ha (100% LoCoH) and 38.52 ha (95% KDE) and accounted for 21.9%, 20.08% and 40.5% of the respective home-range size. The bats foraged in one to three foraging areas each night. The greatest distance between feeding trees varied between 0.25 and 8 km (mean 1.25 km ± 2.19). Those bats with multiple foraging areas moved from patch to patch of Durio zibethinus and did not return to a previously visited patch, whereas those feeding on Parkia repeated their visits to several patches in a single night. Ninety percent of foraging areas used by the radio-tagged individuals were in managed habitat such as fruit orchards and yards of houses to which the bats maintained strong site fidelity.
Sympatric bats engage in various strategies for dietary niche partitioning such as different microhabitat use; however, no previous study has yet looked at potential dietary niche partitioning in mammals foraging in a space void of any physical structure. Here, we used stable isotope ratios of carbon and nitrogen to investigate if three insectivorous bats of central Thailand, Chaerephon plicatus, Taphozous melanopogon and T. theobaldi, partition food resources when foraging in the open space of the lower boundaries of the troposphere. We quantified the isotopic dietary niches of these species and compared niche dimensions within the guild of openspace foraging bats and between this guild and the edge-foraging bat Hipposideros larvatus. Our results showed that stable isotope ratios of bats differed between wet and dry seasons. Consistently, open-space foraging bat species shared a similar isotopic composition in both seasons, which contrasted that of the edge-space foraging H. larvatus. Isotopic niche dimensions of open-space foraging bats were smaller than those of the edge-space foraging bat. Based on isotopic data, we inferred that open-space foraging bats foraged mostly on dipterans which may fly or drift to higher altitudes where these bats hunt. In contrast, H. larvatus included mostly beetles from C4food webs in their diet, highlighting that this species is an important predator of pest insects of C4crops, namely cane sugar and corn. Our study emphasizes that the unstructured aerosphere in which open-space foraging bats hunt insects may promote a large overlap in the diet of these species. We conclude that mechanisms other than trophic niche differentiation, such as the motion capacity of bat species, both in terms of covered distances and accessed altitudes may facilitate the coexistence of high-altitude foraging bats.
The brown planthopper (Nilaparvata lugens) is one of the major insect pests of rice fields in Southeast Asia. They have been widely acknowledged for causing significant rice yield losses. However, the wrinkle-lipped free-tailed bat (Chaerephon plicatus Buchannan, 1800) is a known agent of pest suppression for white-backed planthoppers (Sogatella furcifera), and may also suppress brown planthopper populations. Hence, it is important to investigate the diet of C. plicatus in areas where brown planthoppers are common to determine whether these bats feed on these insects. To accomplish this objective, we analyzed the diet of C. plicatus from two caves that differed in the percentage of surrounding land area occupied by rice fields (70% versus 22%). Bat fecal pellets were collected monthly for a year. A total of 720 fecal pellets were analyzed, and the results revealed that C. plicatus feeds on at least eight insect orders, including Coleoptera, Homoptera, Hemiptera, Diptera, Lepidoptera, Odonata, Hymenoptera and Orthoptera. Specifically, homopterans comprised the greatest diet volume in the rice growing season, whereas coleopterans were most abundant in the diet when rice fields were fallow. Moreover, most homopterans were identified as brown planthoppers. To estimate the relative numbers of brown planthoppers consumed during each month, the number of genitalia of male brown planthoppers was counted. We recorded the greatest numbers of genitalia during the rice planting period, with an average of four genitalia per fecal pellet. Examining both the percent volume and percent frequency of each insect order in the diet of C. plicatus revealed that the two study caves were no significantly different, even though the proportion of surrounding active rice fields was different. Our results suggest that tens of millions of brown planthoppers are consumed by this bat species each night. The similar diets of the two study colonies may be due to their high altitude foraging and preference for migratory insects. Our results indicate that the wrinkle-lipped freetailed bat is an important biological suppression agent of brown planthoppers in rice fields.
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.
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.
The taxon Rhinolophus microglobosus is elevated to specific rank on the basis of clearly defined morphometric and acoustic characters which differentiate it from Rhinolophus stheno. It is recorded from Cambodia for the first time. Rhinolophus malayanus exhibits considerable geographical variation in echolocation calls, with apparently two phonic types: a northern population with lower frequency calls and a predominantly southern population with higher frequencies. However, this acoustic divergence is not reflected in any morphometric divergence, and the taxonomic status of the two phonic populations remains unclear. Discriminating characters of all three species are given, together with distribution data and short ecological summaries. The value of echolocation as an indicator of cryptic species and the zoogeographical implications of the study are briefly discussed.
The diet of E. spelaea was determined for bats captured monthly between June 2002-June 2003 at a cave entrance in Songkhla Province, Southern Thailand. Faecal analysis and pollen collected from the bats’ fur were used to identify the plant species ingested. From 1,155 diet records from 506 samples, at least eleven plant taxa were identified. Individual bats feed on flowers of up to six plant species each night. Parkia spp. (34%) and Musa spp. (28%) have the highest percentage frequency followed by Eugenia spp. (9.4%), Oroxylum indicum (6.4%), Durio zibethinus (6.2%), Ceiba pentandra (5.5%), Sonneratia spp. (5.2%), while Cocos nucifera and an unknown plant species, made up a minor proportion (<2.5%). Parkia and Musa were the main dietary items of E. spelaea in nearly every month, while the remaining components of the diet varied seasonally. Durio spp. is an important contributor to the diet during March–April (39–42%). The results from pollen collected from fur generally corresponded with those from faecal analysis, but Musa spp. had a higher percentage frequency on the fur (34%) than Parkia spp. (23%). The dawn return patterns of the bats to their roosts differ significantly between sexes. Most mature males return early in the night while most females return at dawn. Earlier returning males were significantly heavier than those returning later. This return pattern is similar to that reported during the breeding period for those polygynous fruit bats that maintain a harem. Thus, these results suggest that E. spelaea may exhibit a resource-defence polygynous mating system.
A review of the literature relating to the history of bat research in Thailand (1821–2006) is included, together with lists of the 119 bat species currently recorded from the country and the 16 that are omitted for lack of supporting data. The geographical distribution within Thailand of the some of the more significant bat field studies (1896–2004) is mapped and briefly discussed. Based on field work conducted in peninsular Thailand in 1993 and 2003–2004, eight bat species (Hipposideros ridleyi, Myotis hermani, Pipistrellus stenopterus, Hesperoptenus tomesi, Murina suilla, Murina aenea, Kerivoula pellucida, and Mops mops) are recorded from the country for the first time; information is provided on their taxonomy, distribution, and ecology. Recommendations are made for further bat studies in Thailand, with emphasis placed on selecting less well known species groups, such as forest bats, in under-researched habitats in neglected geographical areas (for example, the deciduous dipterocarp forests of eastern Thailand and the semi-evergreen forests of peninsular Thailand). A need to develop in-country skills in bat acoustics and taxonomy is also highlighted.
The diversity of Rhinolophidae in Thailand and Vietnam is briefly discussed and the taxonomy of Asian Kerivoulinae, with particular reference to the genus Phoniscus, is reviewed. Four new country records are included: Rhinolophus shameli and Kerivoula kachinensis from Vietnam and Phoniscus jagorii from Vietnam and Thailand. A second record of Phoniscus atrox from Thailand is also discussed.
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.
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 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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