Acoustic divergence is a fundamental component of signal-based communication in animals, including humans, and has major implications for individual recognition, mate choice, and speciation. Bats have recently become an important model system for investigating acoustic communication due to their structurally complex vocal repertoires. This study explored communication calls of greater horseshoe bats ( Rhinolophus nippon) to determine whether geographically structured dialects occur and to identify factors associated with dialect formation. Communication calls from nine populations distributed across three regions were analyzed, and 12 syllable types shared among populations were identified. The associations of dialect differentiation with acoustic, climatic, morphological, and genetic distances, together with the influence of geographic barriers, were investigated. Results demonstrated significant acoustic variation among populations and regions for each syllable type, and this divergence was driven by diverse factors. As bats typically emit syllables in combination, all shared syllable types were subsequently analyzed as an integrated dataset. At this broader level, acoustic distance showed significant correlations with both geographic and genetic distance. Causal modeling further revealed that geographic distance and geographic barriers exerted direct effects on dialect differentiation. The sampled populations were classified into three dialect regions: Northeast, Central-East, and Southwest. The Qinling Mountains formed the boundary between the Central-East and Southwest regions. These findings provide clear evidence for the existence of dialect structure in communication calls of greater horseshoe bats and identify geographic isolation as a major force in the formation and preservation of vocal divergence. This study advances current understanding of animal acoustic evolution and offers insight into mechanisms linking vocal diversification with the generation of biological diversity.

Bats play important roles in ecosystems, and are thus considered bioindicators. Libraries of echolocation calls provide huge potential resources for bat species identifications, ecological studies and conservation surveys. Here, the echolocation calls of two morphologically similar bat species (Miniopterus magnater and Miniopterus fuliginosus) were recorded and described in order to characterize vocal signatures for field identification in China. Both M. magnater and M. fuliginosus emitted short frequency modulated echolocation calls with narrow bandwidths. Each call of the former species included two harmonics, with the first harmonic being the strongest, whereas calls of the latter species normally contained one harmonic. Although call durations were similar between the two species, there were significant differences in start, end and peak frequencies between M. magnater and M. fuliginous. The results showed that 92.3% of all calls recorded in China were attributed to the correct species based on spectral features of echolocation calls. We concluded that echolocation calls are valuable characters for the identification of morphologically similar bat species.

Evolutionary biologists had a long-standing interest in the evolutionary forces underlying geographical variation in the acoustic signals of animals. However, the evolutionary forces driving acoustic variation are still unclear. In this study, we quantified the geographical variation in the peak frequencies of echolocation calls in eight Miniopterus fuliginosus bat colonies, and assessed the forces that drive acoustic divergence. Our results demonstrated that seven of the colonies had very similar peak frequencies, while only one colony was significantly higher than the others. This similarity in echolocation call frequency among the seven colonies was likely due to frequent dispersal and migration, leading to male-mediated infiltration of nuclear genes. This infiltration enhances gene flow and weakens ecological selection, and also increases interactions in the presence of conspecifics. Significant correlations were not observed between acoustic distances and morphological distances, climatic differences, geographic distances or mtDNA genetic distances. However, variation in acoustic distances was significantly positive correlated with nDNA genetic distance, even after controlling for geographic distance. Interestingly, the relationship between call divergence and genetic distance was no longer significant after excluding the colony with the highest call frequency, which may be due to the minimal genetic distance among the other seven colonies. The highest frequencies of echolocation calls observed in the one colony may be shaped by selection pressure due to loud background noise in the area. Taken together, these results suggest that geographic divergence of echolocation calls may not be subject to genetic drift, but rather, that the strong selective pressure induced by background noise may lead to acoustic and genetic differentiation between JXT and the other colonies.

Acoustic signals mediate important life history events in a variety of species, providing new vistas for understanding speciation. It has been proposed that animal acoustic signals undergo complex interactions among morphology, ecology, social pressure, and phylogenetic history. Yet, the relative importance of these factors in shaping acoustic divergence is rarely assessed within a comparative framework. Herein, we aim to investigate the key determinants of social call divergence across 31 bat species from 5 families. We compiled a wide data set on bat aggressive calls, body size, foraging habitats, foraging modes, climatic conditions, colony size, and phylogenetic components. We identified remarkable interspecific divergence versus within-species variation in aggressive vocalizations. Despite weak effects of ecological factors, colony size, body size, and phylogenetic components accounted for the majority of variation in call parameters among species. The colony size and body size played a major role in influencing spectral parameters, whereas phylogenetic relationships determined call duration and minimum frequency. Together, our findings constitute convincing evidence that sociality, morphological constraint, and phylogenetic constraint mold social call divergence in bats. This study expands our limited knowledge of the evolution of bat social calls, and highlights the importance of sociality in driving acoustic phenotype diversity.

Drift, selection, or their combined effects commonly drive geographic variation in traits. Clarifying the relative roles of each process is a long-standing research goal in evolutionary biology. Acoustic signals of bats are a phenotypic characteristic that plays an important role in social organization and species recognition. We extensively sampled the Chinese rufous horseshoe bat (Rhinolophus sinicus) throughout China and Vietnam and reconstructed a species phylogeny to better understand the patterns and causes of the geographic variation of acoustic signals. Our results showed that the resting frequency (RF) of calls varied with latitude, sex, and distance among different colony locations. RF differences were not correlated with genetic distance (based on only one nuclear locus and the mitochondrial locus), climatic factors (mean annual temperature and mean annual precipitable water), or body size, although differences in calls increased with distance among various populations. This suggests that cultural drift may play more important roles than genetic drift and acoustic adaptation in shaping acoustic differences within regions in R. sinicus.

Why a variety of social animals vocalize during agonistic foraging interactions remains obscure. One possibility is that these signals advertise the signaller's competitive ability, playing a role in the defence of food resources, yet there is limited evidence in support of this idea. Here, we used adult female Asian particoloured bats, Vespertilio sinensis, to test whether individual variation in competitive ability when foraging can be explained by social calls. Using synchronized infrared video and audio recording, we monitored bat social vocalizations, dominance rank and weight gain in triads under controlled food conditions. Additionally, subsequent playbacks, consisting of experimental stimuli, echolocation pulses and silence, were presented to feeding bats in a laboratory. Analyses showed that females uttered low-requency social calls composed of one to five syllable types during feeding competition. The rates of social vocalizations increased with reduced food availability. Interestingly, dominance score and weight gain correlated weakly with body size, but positively with call rate and associated parameters. Playback of social calls inhibited the visits of bats to the focal food dish compared to playback of silence and echolocation pulses. The amount of food consumed was greatly reduced in the presence of experimental stimuli versus controls. Collectively, these results highlight that acoustic signals serve as an honest indicator of bat competitive ability. (C) 2017 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

To the Editor: Emerging infectious diseases have caused catastrophic declines in wildlife populations, and the introductions of many pathogen have been linked to increases in global trade and travel (1). Mapping the distribution of pathogens is necessary to identify species and populations at risk and identify sources of pathogen spillover and introduction. Once pathogen distributions are known, management actions can be taken to reduce the risk for future global spread (2). Bats with symptoms of white-nose syndrome (WNS) were first detected in the United States in 2006, and the disease has subsequently caused precipitous declines in temperate bat populations across eastern North America (3,4). Pseudogymnoascus destructans, the causative agent of WNS, is a cold-growing fungus that infects bats’ skin during hibernation, leading to more frequent arousals from torpor and death (3). P. destructans is widespread throughout Europe (5), but, to our knowledge, its presence in Asia has not been documented. We sampled bats and hibernacula surfaces (cave walls and ceilings) across northeastern China during 2 visits (June–July 2014 and March 2015) using a previously described swab-sampling technique (6). Bats were captured inside caves and at their entrances. DNA was extracted from samples by using a modified QIAGEN DNeasy blood and tissue kit (QIAGEN, Valencia, CA, USA) and tested in duplicate for the presence of P. destructans with a quantitative real-time PCR (qPCR) (6,7). In the summer of 2014 and winter of 2015, we collected 385 samples from hibernacula surfaces at 12 sites in 3 provinces and 1 municipality (Figure, panel A) and 215 samples from 9 species of bats at 10 sites (summer: Rhinolophus ferrumequinum, Rhinolophus pusillus, Myotis adversus, Myotis macrodactylus, Myotis pilosus, Myotis chinensis, Murina usseriensis; winter: R. ferrumequinum, Murina leucogaster, Myotis petax). During the summer, P. destructans was widely distributed across the study region with positive samples (determined on the basis of qPCR results) obtained from cave surfaces at 9 of 12 sites and from bats at 2 of the 9 sites where bats were sampled (Figure, panel A). Figure A) Distribution of Pseudogymnoascus destructans in cave environments during summer at 9 sites in northeastern China. Pie charts show the prevalence (red indicates fraction of positive samples) of P. destructans, and the size of pie graphs indicates the ... Prevalence of P. destructans was low during summer in the environment (mean prevalence across sites 0.06 ± 0.03) and in bats. Bats of 3 species tested positive for P. destructans in the summer: M. macrodactylus (1/10), M. chinensis (1/1), and M. ussuriensis (1/1). P. destructans was not detected in bats of 4 other species, of which >20 individual animals of each species were sampled (R. ferrumequinum, R. pusillus, M. pilosus, and M. adversus). The low prevalence of P. destructans in bats and on hibernacula surfaces in China during the summer was similar to comparable results from studies in North America (6). In winter, prevalence at the 2 sites we revisited was much higher; 75% of 85 samples from 3 species tested positive, including samples from 16/17 M. petax bats. We also detected P. destructans in bats from 2 additional species (R. ferrumequinum [11/19 bats] and M. leucogaster [11/16 bats]). In addition, during March 2015, we observed visual evidence of P. destructans in bats (M. petax; Figure, panel C) and obtained 2 fungal cultures from swab specimens taken from these bats. To isolate P. destructans from these samples, we plated swab specimens from visibly infected bats on Sabouraud dextrose agar at 10°C. We identified potential P. destructans isolates on the basis of morphologic characteristics. DNA was then extracted from 2 suspected fungal cultures and tested for P. destructans by qPCR, as previously described. To further confirm the presence of P. destructans, we prepared the fungal isolates for Sanger sequencing (Technical Appendix). The 600-nt amplification products from these 2 isolates were sequenced and found to be 100% identical to the P. destructans rRNA gene region targeted for amplification. In addition, using BLAST (http://www.ncbi.nlm.nih.gov/blast.cgi), we found that sequences were a 100% match with isolates from Europe (GenBank accession no. {type:entrez-nucleotide,attrs:{text:GQ489024,term_id:284944064,term_text:GQ489024}}GQ489024) and North America (GenBank accession no. {type:entrez-nucleotide,attrs:{text:EU884924,term_id:211905147,term_text:EU884924}}EU884924). This result confirms that the same species of fungus occurs on all 3 continents. We also obtained wing biopsy punches from these bats and found lesions characteristic of WNS by histopathologic examination (Figure, panel B; Technical Appendix). The occurrence of P. destructans at most sites sampled indicates that this pathogen is widespread in eastern Asia (Figure, panel A). The presence of P. destructans in bats from 6 species in China and on bats in 13 species in Europe (8) confirms the generalist nature of this fungus and suggests that it may occur throughout Eurasia (Figure, panel D). Decontamination and restrictions on the use of equipment that has been used in caves in Asia would help reduce the probability of introducing P. destructans to uninfected bat populations (e.g., western North America, New Zealand, southern Australia, and temperate areas of South America). These measures would also reduce the risk of introducing new strains of P. destructans to regions where bats are already infected (e.g., eastern North America and Europe). These measures are necessary to prevent the devastating effects this pathogen has had on bats in North America and would help maintain the ecosystem services that bats provide (9,10). Technical Appendix: The Technical Appendix describes the DNA sequencing of fungal isolates to confirm the presence of Pseudogymnoascus destructans and histologic examination of bat fungal lesions. Click here to view.(238K, pdf)

The geographical variation in animals’ social acoustical signals can reduce recognition across populations and may thus promote assortative mating, reproductive isolation and speciation. However, the social consequences of geographical variation in acoustical signals serving an ecological function are poorly known. Bat echolocation calls are considered to have a dual function; they are used not only for orientation and foraging but also for communication. In this study, we studied the behavioural response of Rhinolophus ferrumequinum to geographical variation in echolocation calls. Using habituation-dishabituation playback experiments, we found that all tested bats from northeast China exhibited obvious responses after switching playback from their own population to those from central east China and southwest China. Using two-choice playback experiments, we showed that the bats from northeast China responded more strongly to echolocation calls from their own population than to those from central east China. Our results demonstrate that Rhinolophus ferrumequinum is able to discriminate between echolocation calls of its local population and a foreign one and that local echolocation calls are preferred over foreign ones in a two-choice context. This study supports the communicative potential of bat echolocation calls and provides insight into the discrimination ability and behavioural preference of bats with respect to geographical variation in echolocation calls.

Bats have been implicated as important reservoir hosts of alpha- and betacoronaviruses. In this study, diverse coronaviruses (CoVs) were detected in 50 of 951 (positive rate 5.3%) intestinal specimens of eight bat species collected in four provinces and the Tibet Autonomous Region of China by pan-coronavirus RT-PCR screening. Based on 400-nt RNA-dependent RNA polymerase (RdRP) sequence analysis, eight belonged to genus Alphacoronavirus and 42 to Betacoronavirus. Among the 50 positive specimens, thirteen gave rise to CoV full-length RdRP gene amplification for further sequence comparison, of which three divergent sequences (two from a unreported province) were subjected to full genome sequencing. Two complete genomes of betacoronaviruses (JTMC15 and JPDB144) and one nearly-complete genome of alphacoronavirus (JTAC2) were sequenced and their genomic organization predicted. The present study has identified additional numbers of genetically diverse bat-borne coronaviruses with a wide distribution in China. Two new species of bat CoV, identified through sequence comparison and phylogenetic analysis, are proposed.

Predicting species' fates following the introduction of a novel pathogen is a significant and growing problem in conservation. Comparing disease dynamics between introduced and endemic regions can offer insight into which naive hosts will persist or go extinct, with disease acting as a filter on host communities. We examined four hypothesized mechanisms for host–pathogen persistence by comparing host infection patterns and environmental reservoirs for Pseudogymnoascus destructans (the causative agent of white-nose syndrome) in Asia, an endemic region, and North America, where the pathogen has recently invaded. Although colony sizes of bats and hibernacula temperatures were very similar, both infection prevalence and fungal loads were much lower on bats and in the environment in Asia than North America. These results indicate that transmission intensity and pathogen growth are lower in Asia, likely due to higher host resistance to pathogen growth in this endemic region, and not due to host tolerance, lower transmission due to smaller populations, or lower environmentally driven pathogen growth rate. Disease filtering also appears to be favouring initially resistant species in North America. More broadly, determining the mechanisms allowing species persistence in endemic regions can help identify species at greater risk of extinction in introduced regions, and determine the consequences for disease dynamics and host–pathogen coevolution.

Because of its complicated systematics, the bent-winged bat is one of the most frequently studied bat species groups. In China, two morphologically similar bent-winged bat species, Miniopterus fuliginosus and Miniopterus magnater were identified, but their distribution range and genetic differentiation are largely unexplored. In this study, we applied DNA bar codes and two other mitochondrial DNA genes including morphological parameters to determine the phylogeny, genetic differentiation, spatial distribution, and morphological difference of the M. fuliginosus and M. magnater sampled from China and one site in Vietnam. Mitochondrial DNA gene genealogies revealed two monophyletic lineages throughout the Tropic of Cancer. According to DNA bar code divergences, one is M. fuliginosus corresponding to the Chinese mainland and the other is M. magnater corresponding to tropical regions including Hainan and Guangdong provinces of China and Vietnam. Their most recent common ancestor was dated to the early stage of the Quaternary glacial period (ca. 2.26 million years ago [Ma] on the basis of D-loop data, and ca. 1.69-2.37 Ma according to ND2). A population expansion event was inferred for populations of M. fuliginosus at 0.14 Ma. The two species probably arose in separate Pleistocene refugia under different climate zones. They significantly differed in forearm length, maxillary third molar width, and greatest length of the skull.

Aim The goals of our study were to assess the population history and genetic structure of the widespread bat Hipposideros armiger, and to evaluate the effect of palaeoclimatic changes and dispersal patterns on this species.Location South China, mainland Southeast Asia and the South Himalayas.Methods We amplified two mitochondrial DNA (mtDNA) regions (cyt b and D-loop) and seven nuclear microsatellite loci (nSSRs) from 216 individuals of H. armiger. To examine the evolutionary history of this species, we constructed maximum likelihood and Bayesian phylogenetic trees based on the two mtDNA regions. From the mtDNA and/or nSSR data, we assessed population genetic structure using analysis of molecular variance (AMOVA) and median-joining network and structure analyses. We also estimated demographic history and gene flow using a Bayesian skyline plot and the program IMA2.Results Phylogenetic and median-joining network analyses revealed that H. armiger comprises two distinct mtDNA clades divided into seven subclades. The results of AMOVA suggested strong population genetic structure based on mtDNA, but weak structure based on nSSRs. structure analysis identified three population clusters and also showed weak genetic structure at the nuclear level. Demographic analyses revealed two population expansion events c. 0.62 Ma and c. 0.25 Ma. The basic phylogeographical structure of H. armiger was established by 0.24 Ma. IMA2 analysis demonstrated that substantial gene flow has occurred between different regions since then. Additionally, non-significant population structure and significant gene flow were detected between Taiwan and Hainan island populations and those from mainland China.Main conclusions Our results suggest that divergence and population expansion of H. armiger occurred in association with Pleistocene climatic changes and that multiple refugia may have existed for this species. Post-glacial male-biased dispersal was likely to be the primary contributor to the contemporary genetic structure of H. armiger populations. Gene flow may have contributed greatly to the genetic structure of insular populations and populations from mainland China.

Ecologists and evolutionary biologists have a long‐standing interest in the patterns and causes of geographical variation in animals’ acoustic signals. Nonetheless, the processes driving acoustic divergence are still poorly understood. Here, we studied the geographical variation in echolocation vocalizations (commonly referred to as echolocation ‘pulses’ given their short duration and relatively stereotypic nature, and to contrast them from the communicative vocalizations or ‘calls’) of a widespread bat species Hipposideros armiger in south China, and assessed whether the acoustic divergence was driven by either ecological selection, or cultural or genetic drift. Our results revealed that the peak frequency of echolocation pulses varied significantly across populations sampled, with the maximum variation of about 6 kHz. The peak frequency clustered into three groups: eastern and western China, Hainan and southern Yunnan. The population differences in echolocation pulses were not significantly related to the variation in climatic (mean annual temperature, mean annual relative humidity, and mean annual precipitable water) or genetic (genetic distance) factors, but significantly related to morphological (forearm length) variation which was correlated with mean annual temperature. Moreover, the acoustic differences were significantly correlated with geographical and latitudinal distance after controlling for ‘morphological distance’. Thus, neither direct ecological selection nor genetic drift contributed to the acoustic divergence observed in H. armiger. Instead, we propose that the action of both indirect ecological selection (i.e. selection on body size) as well as cultural drift promote, in part, divergence in echolocation vocalizations of individuals within geographically distributed populations.

The investigation of mechanism of species coexistence promotes understanding of the mechanistic processes behind community ecology and ecosystem functions. Niche theory declares that species coexistence within a community must partition the resources of their environment. Two sympatric and morphologically similar bat species, Rhinolophus affinis and Rhinolophus pearsoni , provided a unique opportunity to test the causal mechanism of coexistence. Previous study showed that their coexistence was promoted not by the trophic and spatial niche differentiation but the relatively high abundance of prey resources, which was not in accord with the prediction of niche theory. Here, therefore, we reanalyzed the dietary composition by fecal analysis and surveyed the feeding time of both species. Our results showed that R. affinis and R. pearsoni hunt mainly mostly on Coleoptera and Lepidoptera, and there was a very high overlap (0.84) of trophic niche between the two species. However, significant difference in the duration of the activity period between both species was detected, which illustrated that temporal partitioning of prey resource use facilitated their coexistence. Additionally, our work highlighted the importance of integration of the traditional methods and next-generation sequencing methods for identifying dietary composition of carnivores, and suggested that ongoing studies of species coexistence must consider simultaneously multiple niche axes.

The sensory drive hypothesis of speciation predicts that divergence in communication systems will occur when environments differ and that this sensory divergence can ultimately promote speciation. The factors affecting geographic evolution in acoustic signals remain poorly understood, especially in the contexts of high gene flow. This study investigated variation patterns in peak frequency emitted by the Chinese endemic Myotis davidii on a broad geographic scale by evaluating the relative importance of morphological, environmental, geographic, and genetic variables. Significant variation in peak frequency was observed among regions, but peak frequencies among populations within region had some percentage of similarity. Differences in peak frequency were not associated with morphological difference, genetic structure, and geographic distance among regions, which suggested that peak frequency divergences in M. davidii were not the primary driver of regions' isolation in a context of weak gene flow. Within the Middle East Plain (MEP), one of the regions delineated in this study, peak frequency differences of M. davidii were not significantly correlated with genetic distance and geographic distance among populations, suggesting that peak frequency was not be subject to cultural drift within MEP. Our results provide evidence that geographic variation in echolocation call design may evolve as a consequence of local adaptation to climate conditions.

Subterranean environments are essential for the survival of many bat species and other cave fauna but these places are subject to increasing human disturbance. To examine the significance of subterranean habitats for the conservation of bats in China we surveyed bat species in 225 underground sites during 2003-2011. Our results show that 77% of bat species in China, including 30 nationally Endangered or Vulnerable species and nine endemic species, roost in caves and other subterranean habitats. The number of species in occupied roosts was 1-15. Almost 90% of the roosts surveyed contained signs of human disturbance, most of which was from recreational activities. One hundred and twenty-one roosts merit special concern because they harbour >= 6 species or > 1,000 individuals, or species of special concern (threatened or endemic species). Generally, larger roosts support more species and a greater abundance of bats than smaller roosts but there is no direct correlation between the presence of species of special concern and roost size. Disused tourist caves have significantly more bat species than other types of roosts. Our data demonstrate that roost disturbance by recreational activities has pronounced detrimental effects on the number of bat species and the presence of species of special concern. We discuss the social, economic and political issues that could adversely affect bat conservation in caves in China, and we recommend that protection of subterranean habitats should be a high priority for bat conservation.

Background: China is characterized by complex topographic structure and dramatic palaeoclimatic changes, making species biogeography studies particularly interesting. Previous researchers have also demonstrated multiple species experienced complex population histories, meanwhile multiple shelters existed in Chinese mainland. Despite this, species phylogeography is still largely unexplored. In the present study, we used a combination of microsatellites and mitochondrial DNA (mtDNA) to investigate the phylogeography of the east Asian fish-eating bat (Myotis pilosus).Results: Phylogenetic analyses showed that M. pilosus comprised three main lineages: A, B and C, which corresponded to distinct geographic populations of the Yangtze Plain (YTP), Sichuan Basin (SCB) and North and South of China (NSC), respectively. The most recent common ancestor of M. pilosus was dated as 0.25 million years before present (BP). Population expansion events were inferred for populations of Clade C, North China Plain region, Clade B and YunGui Plateau region at 38,700, 15,900, 4,520 and 4,520 years BP, respectively. Conflicting results were obtained from mtDNA and microsatellite analyses; strong population genetic structure was obtained from mtDNA data but not microsatellite data. The microsatellite data indicated that genetic subdivision fits an isolation-by-distance (IBD) model, but the mtDNA data failed to support this model.Conclusions: Our results suggested that Pleistocene climatic oscillations might have had a profound influence on the demographic history of M. pilosus. Spatial genetic structures of maternal lineages that are different from those observed in other sympatric bats species may be as a result of interactions among special population history and local environmental factors. There are at least three possible refugia for M. pilosus during glacial episodes. Apparently contradictory genetic structure patterns of mtDNA and microsatellite could be explained by male-mediated gene flow among populations. This study also provides insights on the necessity of conservation of M. pilosus populations to conserve this genetic biodiversity, especially in the areas of YTP, SCB and NSC regions.

Although it is conjectured that diurnal capture may reduce the colony size of bats, little evidence is available. We monitored a maternity colony of cave-dwelling bats (Hipposideros armiger) both before and after casual diurnal disturbance and capture. Results from more than 40 days of monitoring after disturbance suggested that diurnal capture significantly reduced the colony size from more than 90 individuals to a few. Moreover, the trend for colony size after disturbance indicated that most bats may not return to the roost for the whole season. This study has implications for both bat researchers and wildlife managers.

Fourteen bats were captured at the Tongan Cave in Yongzhou of Hunan Province and identified as Myotis ricketti which is a new record of the Chiroptera in Hunan Province.The morphological measurements and skull characteristics were reported and compared to the same specimens in Zhejiang and Anhui Province.The species produces short echolocation calls with downward frequency-modulated(FM) components and low peak frequency.The special echolocation calls and morphology such as strong hind foot and sharp claws contribute M.ricketti to trawl fish over the water surface.