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-Karyotypes of 2 species of Kerivoula. A) Karyotype of K. intermedia (TK 152019; female; 2n 5 28, FN 5 52). This karyotype was identical to that of K. minuta. B) Karyotypes of K. hardwickii (top: TK 152097; female; 2n 5 26, FN 5 48; and bottom: TK 152115; male; 2n 5 26, FN 5 46).
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Examining species boundaries using data from multiple independent sources is an appropriate and robust method to identify genetically isolated evolutionary units. We used 5 data sets—cytochrome b (Cytb), cytochrome c oxidase (COI), amplified fragment length polymorphisms (AFLPs), karyotypes, and morphology—to estimate phylogenetic relationships and...
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... based on comparison with the COI gene tree (deposited by researchers in the Barcode of Life Database) of Francis et al. (2007) and was further reinforced in our chromosomal analysis. Speci- mens of K. hardwickii (TK 152097, TK 152115) from the same locality in Borneo have a diploid number of 26 but have different fundamental numbers, 48 and 46 (Fig. 5B), respec- tively. In the karyotype TK 152097 (Fig. 5B) chromosome pair 12 is distinctly biarmed, whereas in TK 152115 ( Fig. 5B) this pair is acrocentric with a small 2nd arm. Individuals with different karyotypes had similar mtDNA haplotypes, thus indicating a chromosomal polymorphism but probably not cryptic ...
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... by researchers in the Barcode of Life Database) of Francis et al. (2007) and was further reinforced in our chromosomal analysis. Speci- mens of K. hardwickii (TK 152097, TK 152115) from the same locality in Borneo have a diploid number of 26 but have different fundamental numbers, 48 and 46 (Fig. 5B), respec- tively. In the karyotype TK 152097 (Fig. 5B) chromosome pair 12 is distinctly biarmed, whereas in TK 152115 ( Fig. 5B) this pair is acrocentric with a small 2nd arm. Individuals with different karyotypes had similar mtDNA haplotypes, thus indicating a chromosomal polymorphism but probably not cryptic ...
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... and was further reinforced in our chromosomal analysis. Speci- mens of K. hardwickii (TK 152097, TK 152115) from the same locality in Borneo have a diploid number of 26 but have different fundamental numbers, 48 and 46 (Fig. 5B), respec- tively. In the karyotype TK 152097 (Fig. 5B) chromosome pair 12 is distinctly biarmed, whereas in TK 152115 ( Fig. 5B) this pair is acrocentric with a small 2nd arm. Individuals with different karyotypes had similar mtDNA haplotypes, thus indicating a chromosomal polymorphism but probably not cryptic ...
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Species identification through genetic barcoding can augment traditional taxonomic methods, which rely on morphological features of the specimen. Such approaches are especially valuable when specimens are in poor condition or comprise very limited material, a situation that often applies to chiropteran (bat) specimens submitted to the Canadian Food...
The identification of two cryptic bat species of the genus Pipistrellus using a non-destructive and quick method of multiplex PCR and agarose gel electrophoresis is described. Two primer combinations were able to produce species-specific bands that identified reliably individuals that were previously identified by mtDNA sequencing. Robustness of th...
Citations
... Subsequently, Bates et al. (2007) divided the species into two morphological types, K. hardwickii (with domed skull) and K. depressa (Miller, 1906) (with flattened skull). However, molecular phylogenies indicate that the taxonomy and systematics of the K. hardwickii complex are still ambiguous because of the occurrence of multiple divergent lineages (Francis et al. , 2010Khan et al. 2010;Douangboubpha et al. 2016;Nguyen et al. 2016). Recent genetic analyses revealed that K. depressa can be divided into two distinct clades (Kuo et al. 2017). ...
... Recent genetic analyses revealed that K. depressa can be divided into two distinct clades (Kuo et al. 2017). Tu et al. (2018) described K. depressa and K. dongduongana (Tu et al. 2018) based on phylogenetic analyses using COI, Cytb, and RAG2 sequences from newly obtained specimens and those from previous studies across the Indo-China Peninsula and the Philippines (Hoofer et al. 2003;Stadelmann et al. 2004;Francis et al. 2010;Khan et al. 2010;Wu et al. 2012;Kruskop 2013;Kuo et al. 2014;Kuo et al. 2017). Currently, the K. hardwickii complex contains five species: K. hardwickii (sensu stricto), K. kachinensis (Bates et al., 2004), K. furva (Kuo et al., 2017), K. depressa, and K. dongduongana (Kuo et al. 2017;Tu et al. 2018). ...
... Three gene sequences from all 24 voucher Kerivoula specimens were obtained (GenBank accession numbers: COI: OM716930-OM716952; Cytb: OM735691-OM735714; RAG2: OM735715-OM735736). These sequences were compared with 87 COI, 38 Cytb, and 28 RAG2 sequences of the subfamily Kerivoulinae from NCBI nucleotide databases and with the three outgroup species, including Myotis muricola (Gray, 1846), Harpiocephalus harpia (Temminck, 1840), and Murina cyclotis (Dobson, 1872) (Hoofer et al. 2003;Khan et al. 2010;Ruedi et al. 2012). Newly generated and downloaded sequences are detailed in Appendix 1. ...
Recent studies have revealed that the Kerivoula depressa complex should be divided into two species, K. depressa distributed mainly in Myanmar, Vietnam, Laos and Cambodia, and K. dongduongana found only in the Annamite Mountains of Vietnam, Laos and Cambodia. In November 2018 and April 2019, 24 woolly bats were collected by two-band harp traps in Xishuangbanna, Yunnan, China. Based on morphological, morphometric, and phylogenetic ( COI , Cytb , and RAG2 gene sequences) analyses, these bats were identified as K. depressa and K. dongduongana , representing two new species records for the country. Including the new records, six Kerivoula species have been recorded in China, namely K. depressa , K. dongduongana , K. furva , K. kachinensis , K. picta and K. titania . To facilitate their identification and biological research in the future, we have provided an up-to-date key to all Kerivoula species occurring in China.
... Total genomic DNA was isolated from approximately 20 mg of muscle tissue using the traditional phenol chloroform extraction method. We further sequenced the partial segments of CoI, Cytb and Rag2 following protocol and primers from Francis et al. (2010), Khan et al. (2010) and Yu et al. (2018). Gene sequences of Kerivoula were downloaded from the NCBI-nt database and used for reconstruction phylogenetic trees with the newly generated sequences (GenBank accession numbers of K. kachinensis: CoI: MZ438737-MZ438743; Cytb: MZ438728-MZ438735; Rag2: MZ438744-MZ438750; GenBank accession numbers of K. titania: CoI: MZ417537-MZ417543; Cytb: MZ417524-MZ417530; Rag2: MZ417531-MZ417536). ...
In April 2019, 15 (10♂, 5♀) Kerivoula bats were collected by harp traps from Xishuangbanna, Yunnan Province, China. External and craniodental examination, multivariate statistical analyses and molecular phylogenetic inference ( CoI , Cytb and Rag2 gene markers) indicated they are Kerivoula kachinensis and Kerivoula titania , respectively. Former represents a new chiropteran record from China, while the latter is a valid occurrence of K. titania in this region because recent study indicate a misidentification of “ K. titania ” in Guangdong, Guangxi and Hainan, China. All specimens are presently preserved at Key Laboratory of Conservation and Application in Biodiversity of South China in Guangzhou University, Guangzhou, China. Nowadays, four woolly bats occur in China including, Kerivoula furva , K. kachinensis , Kerivoula picta and K. titania , whilst there is a risk of underestimation the actual species diversity in China region when comparing those of neighboring region such as Vietnam. Supports for field survey need to be continued in future.
... E. alecto/monticola was only identified to a species pair from a photo of four individuals roosting at a small cave entrance, following the description by Payne et al. (1985) and a description of the unique roosting posture by Kingston et al. (2006). In this study, we assigned individuals of Kerivoula papillosa to either the large or small form of the species (Khan et al. 2010, Hasan & Abdullah 2011. We considered individuals with forearm length 43.7 mm and above to be the large form and those with forearm 42.5 mm and below to be the small form. ...
... However, K. papillosa (both large and small form) had higher classification success (86% and 82% respectively). The small form, with forearm length <44 mm, is generally accepted as the true K. papillosa, while the larger form, with forearm length >44 mm, may represent an undescribed species endemic to Borneo (Khan et al. 2010, Hasan & Abdullah 2011. In this study, there was only 7% overlap in call parameters between the two forms, which supports the hypothesis that these are two separate species. ...
Acoustic monitoring with ultrasonic detectors has emerged, in recent years, as an essential tool to quantify the activity of echolocating, insectivorous bats and identify critical commuting and foraging habitats. Comprehensive reference call libraries are critical for the identification of species from their calls. This is especially important in species diverse areas like Gunung Mulu National Park (Sarawak). This study aims to (1) develop a call library for all known echolocating bat species found in Gunung Mulu National Park, (2) determine if calls of different species can be automatically classified using discriminant function analysis, and (3) examine intraspecific variation in relation to sex and geographical location, for calls in species of the families Rhinolophidae and Hipposideridae. Between 2012 and 2017, insectivorous bats were trapped within and outside the park. Echolocation calls were recorded from a total of 508 individuals, representing 31 species from 8 families. Results from discriminant function analysis indicated that the majority of cave roosting bats, which included Chaerephon plicatus, Miniopterus australis, Myotis horsfieldii and 13 species from the families Rhinolophidae, Hipposideridae, and Emballonuridae, could be readily distinguished from their calls, when manually separated into groups according to call structure. However, classification success was much lower for the remaining 15 species that consisted mainly of forest roosting bats from the family Vespertilionidae. This reference call library is expected to contribute to a regional online open-access database. It can be used to survey and monitor selected species in Gunung Mulu National Park as well as highlighting the importance of threatened habitats outside the boundary for these species.
... Many studies use karyological evidence as an element of integrated approach, in which case it becomes an extra argument to support a taxonomic hypothesis. Examples include the confirmation of generic rank of Neoromicia (Kearney et al., 2002), the description of cryptic diversity in Carollia castanea species complex (Solari et al., 2006), the description of the genus Hsunycteris and the tribe Hsunycterini (Parlos et al., 2014), and the study of phylogenetic relationships and cryptic diversity in the genus Kerivoula (Khan et al., 2010). ...
Bats are the second largest mammalian order with an almost worldwide distribution. Bat taxonomy remained almost unchanged for decades, and the diversity of the order was underestimated. The advent of molecular methods brought change to chiropteran taxonomy. The number of families increased from 17–18 to 21, and the relationships between them were revised, as were the composition of suborders and superfamilies. The number of recognized species and genera went up by almost a third. As a discipline, bat taxonomy has changed much methodologically and conceptually. After its long reign, comparative morphology has faded into the background. It has become clear that characters can diverge and converge in related species, masking true phylogenetic relationships. Not writing morphology off entirely, it does necessitate resorting to finer structures or multivariate data analysis. Karyology is of limited use in bat taxonomy, but methods such as FISH add to the understanding of relationships between suprageneric taxa. Mitochondrial DNA sequences are easy to obtain, and their analysis yields well-supported phylogenetic trees, but reticular processes and other factors may mask taxon boundaries. To resolve the uncertainty, nuclear markers are used, and their number and choice depends on taxon characteristics. Overall, building a consistent chiropteran system calls for an integration of all mentioned approaches.
... DNA microsatellite markers are highly versatile and useful as genetic markers for ecological and evolutionary studies, i.e. from species identification to the detection of the biological history of populations (Khan et al. 2010;Wilson et al. 2014). However, the lack and unavailability of these microsatellite markers for many non-model organisms, including bats from the Old World populations, has hampered efforts in the genetic conservation of the species. ...
Microsatellite markers are remarkably useful as genetic markers in ecological and evolutionary studies. However, the lack of microsatellite markers for many non-model organisms, including bats, especially those from the Old World populations, has hampered genetic conservation efforts. The Bornean bats-Hipposideros cervinus (Fawn leaf bat) and Myotis muricola (Whiskered myotis) are two examples of the Old World bats. In this study, genomic DNA from H. cervinus and M. muricola were isolated and sheared to approximately 6 kb fragments for SMRTbell template library generation. Genome shotgun sequencing was then performed for both the genomic DNA samples on a PacBio's RSII sequencer using the P6v2C4 chemistry platform. PacBio's High-throughput sequencing resulted in a total of 125,740 high-quality DNA circular consensus reads (CCS) for H. cervinus and 117,009 for M. muricola, respectively. The resulting high-quality CCS reads were mined for four types of microsatellites, i.e. di-, tri, tetra-and penta-nucleotide repeats. Subsequently, specific primers were developed for the microsatellite motifs with large tandem repeats to capture polymorphisms in the bat species. The information obtained from this work will assist in the genetic conservation of the two species.
... whereas Tylonycteris, Kerivoula, and Rhinolophus spp. have a much weaker dispersal ability because they are mostly forest-dwelling taxa (Khan et al., 2010;Mao et al., 2010;Tu et al., 2017Tu et al., , 2018. ...
... Thus, unlike forest-dependent species, the dispersal and associated gene flow among geographic populations of Asian Scotophilus might be less restricted by physical and ecological barriers (Hisheh et al., 2004;Yu et al., 2012). In addition, while the population structure and dynamics of species with low dispersal capacities were strongly influenced by the past compression and expansion of forests during glacial and interglacial periods in the Pleistocene (Khan et al., 2010;Mao et al., 2010;Tu et al., 2017Tu et al., , 2018 and by current patterns of deforestation (Kingston, 2010), Asian Scotophilus spp. ...
Yellow house bats (Scotophilus) have been known for centuries as a widespread genus of vesper bats in the Indomalayan Region. Despite this, their taxonomic status and phylogeographical patterns remain unclear due to differing criteria employed by early taxonomists and inconsistencies between morphological and molecular assessments. To address these issues, we undertook a comparative phylogeographic analysis of Asian Scotophilus spp. using integrated genetic and morphological analyses of samples collected across the region. These demonstrate that yellow house bats in Asia can be classified into just two widespread species, namely the smaller S. kuhlii (e.g., FA ≤ 53.1 mm, GLS ≤ 20.18 mm) and the larger S. heathii (e.g., FA ≥ 53.4 mm, GLS ≥ 20.85 mm), which occur in sympatry in different parts of the Indomalayan Region. Although these two sympatric species share similar eco‐ethological preferences, they differ considerably in their geographic distributions and intraspecific variation in mtDNA sequences and morphological traits. These disparities were likely misinterpreted as indicating potential cryptic diversity in previous studies, whereas we suggest they are related to interspecific differences in sex‐biased gene flow and phenotypic plasticity to adapt to varying environments. Our study highlights the importance of using multiple datasets to resolve taxonomic uncertainties and reconstruct demographic and phylogeographic histories of sympatric species.
... Additionally, we included several species that occur near the Philippines, including K. titania, K. intermedia, and K. minuta. We did not include several members of the K. hardwickii group, a complex that requires much detailed study (e.g., K. depressa, K. krauensis, K. furva or K. kachinensis; see Bates et al. 2007;Francis et al. 2007;Khan et al. 2010;Douangboubpha et al. 2015;Son et al., et al. 2015;Struebig et al. 2016;Kuo et al. 2017;Tu et al. 2018). ...
... In addition to the sequences of Cyt b we generated for this paper, we downloaded from the GenBank and aligned a set of representative Kerivoula lineages from Southeast Asia published by Khan et al. (2010), 2 sequences (JN112244 and JN112246) assigned to K. titania by Wu et al. (2012) and subsequently reassigned to K. furva by Tu et al. (2018), and one unpublished sequence of K. pellucida (AB444717; Bastian Jr, S.T., Malcampo, R.A., Yamagata, T. & Namikawa, T.) from the Philippines (Appendix B). We noticed that a series of sequences of K. intermedia (EU188789-188791) are wrongly labelled as K. whiteheadi in the GenBank, while a series of K. minuta (EU188774-188778) are wrongly labelled as K. intermedia. ...
... We noticed that a series of sequences of K. intermedia (EU188789-188791) are wrongly labelled as K. whiteheadi in the GenBank, while a series of K. minuta (EU188774-188778) are wrongly labelled as K. intermedia. As both series are correctly referenced in the Appendix I of Khan et al. (2010), we follow this original publication to label representatives of these lineages (see Appendix B). ...
Bats of the genus Kerivoula (Mammalia, Chiroptera) are widespread in the Philippines with four reported species, but have been poorly known due to a paucity of specimens. We provide the first molecular phylogeny for Philippine Kerivoula, which supports the existence of four distinct clades that we treat as species (K. hardwickii, K. papillosa, K. pellucida, and K. whiteheadi); these four overlap broadly geographically. Each of these may be recognized on the basis of cytochrome b sequences and external and craniodental morphology. Detailed examination of K. pellucida shows little geographic differentiation within the Philippines, but they differ subtly from those on the Sunda Shelf. We consider K. whiteheadi to be composed of four recognizable clades, each restricted to a geographic region within the Philippines. We consider K. bicolor, from peninsular Thailand, and K. pusilla, from Borneo, to be distinct from K. whiteheadi. Our data indicate the presence of two species within the Philippines currently lumped as K. hardwickii; further study of these is needed. A calibrated phylogeny suggests that Kerivoula began arriving in the Philippines about 10 MYA, with each of the four current lineages arriving independently.
... Evidently, the principal application of integrative taxonomy is in species delimitation, which includes the use of a species concept and tests for species boundaries (see Sites 2007). Publications dealing with species delimitation have used a combination of characters, including molecular, morphological, morphometric, echolocation calls, karyotypes, and virtually any dataset that can reinforce discrimination among taxa (Solari and Baker 2006;Khan et al. 2010;Velazco et al. 2010;Galimberti et al. 2012;Clare et al. 2013;Volleth et al. 2015;Pavan and Marroig 2016). To reiterate, the idea behind this approach is to avoid the possible subjectivity of a phylogenetic analysis based on a single dataset to make taxonomic decisions. ...
... Although independent molecular datasets (e.g., mitochondrial, nuclear, etc.) can be analyzed under similar conditions (rendering their independent topologies fully comparable) or even concatenated for a single analysis, an equivalent integration cannot be applied to morphological or karyological traits. An example of corroboration by different datasets, including mitochondrial and nuclear sequences, karyotypes, and morphological traits, was presented by Khan et al. (2010) in a study of Malaysian species of Kerivoula (Vespertilionidae). Most datasets converged on a similar result to provide a robust framework for species taxonomy and their phylogenetic relationships, comparable in large part to that elucidated by previous studies (Francis et al. 2007). ...
The order Chiroptera comprises all bat species and is the second-most diverse order of mammals. Organization of this diversity into well-delimited taxa embodying their evolutionary affinities remains challenging, particularly because the relationships among various groups at most taxonomic levels are not yet fully resolved. Historically, systematic questions were approached independently using tools and methodologies within researchers' individual expertise. With technological advances, this trend has shifted to more comprehensive analyses, merging independent datasets from various lines of research. Nevertheless, disagreements among phylogenetic relationships inferred from independent datasets, and derived from the complexity of modes of diversification and speciation, still preclude the achievement of a single, comprehensive, and stable classification framework for bats. In this context, integrative taxonomy uses complementarity among various disciplines to improve the outcome of hypothesis testing, which can be achieved by congruence or cumulation of datasets. Here, we provide a review of bat systematics, with emphasis on the instances where integrative taxonomy has proven to be crucial in delimitation of taxa. We focus on examples of true integration or integration using discrete data to support delimitation at different taxonomic levels. In addition, we discuss the difficulties associated with the integration of incongruent phylogenies, as well as how to incorporate information crucial to phylogenetic decisions from data not easily comparable to most common datasets. Our goal is to provide exemplar guidelines that can be useful to taxonomists for the creation of stable classifications without subjectivity and limitations of single datasets. Accordingly, we present a review of recent issues in chiropteran systematics, including comments on how the use of integrative taxonomy helped to solve them. El orden Chiroptera incluye todos los murciélagos (actuales y extintos) y es el segundo orden más diverso de los mamíferos. La organización de esta diversidad en taxones bien delimitados que incorporen sus afinidades evolutivas es un reto permanente, especialmente porque las relaciones a distintos niveles taxonómicos no han sido resueltas en su totalidad. Históricamente, las preguntas sistemáticas fueron abordadas independientemente, usando herramientas y metodologías asociadas a la experiencia individual de cada investigador. Los avances tecnológicos han cambiado esta tendencia hacia un análisis más comprensivo, combinando conjuntos de datos independientes desde las más diversas áreas de investigación. Sin embargo, desacuerdos entre hipótesis filogenéticas resultantes de datos independientes, derivados de la complejidad de los diversos modos de diversificación y especiación, siguen evitando la consolidación de una clasificación comprensiva y estable. En este contexto, la taxonomía integrativa favorece la complementariedad entre varias disciplinas para mejorar la robustez de las hipótesis sistemáticas, la cual puede lograrse por congruencia entre o acumulación de datos. En este estudio, presentamos una revisión de la sistemática actual de los murciélagos, con énfasis en casos en los que una aproximación integrativa de la taxonomía resultó crucial para delimitar taxones. Para ello, revisamos ejemplos de verdadera integración, o integración empleando datos discretos, para soportar la delimitación de entidades a distintos niveles taxonómicos.
... Using mtDNA barcode sequences (COI), Francis et al. (2007Francis et al. ( , 2010 found five divergent mitochondrial haplogroups in bats tentatively identified as K. hardwickii. Khan et al. (2010) also detected high levels of divergence (5.7%) in Cytb sequences between populations of K. hardwickii s.l. in Borneo. In Thailand, K. hardwickii specimens were divided into two morphotypes (domed-skulls vs. flattened skulls) and COI analyses indicated that specimens in the domed-skull group formed a monophyletic clade, whereas those with flattened skulls resided in two divergent monophyletic clades (Douangboubpha et al., 2015). ...
... S2). The three outgroup species were chosen on the basis of previous molecular studies (Hoofer et al., 2003;Khan et al., 2010;Ruedi et al., 2012) and represent three vespertilionid genera from two subfamilies: Myotis muricola belongs to Myotinae, Harpiocephalus harpia, and Murina cyclotis to Murininae. ...
... Within Kerivoula, K. picta and K. pellucida occured outside of the clade uniting all other studied species (PP=1 in COI/Cytb/mtDNA). In this large clade, two major groups were recovered in all analyses, namely: one group named "papillosa" uniting K. papillosa, K. lenis, and a potentially undescribed species of Kerivoula from Borneo (see Khan et al., 2010 for details) (PP=1), and another group named "hardwickii" including K. intermedia, K. kachinensis, K. krauensis, K. minuta, K. titania, and K. hardwickii sensu lato (PP COI/Cytb/mtDNA =0.4/0.8/0.98). ...
We undertook a comparative phylogeographic study using molecular, morphological and morphometric approaches to address systematic issues in bats of the Kerivoula hardwickii complex in Asia. Our phylogenetic reconstructions using DNA sequences of two mitochondrial and seven nuclear genes reveal a distinct clade containing four small-sized species (K. hardwickii sensu stricto, K. depressa, K. furva and Kerivoula sp. nov.) previously assigned to K. hardwickii sensu lato, as well as K. kachinensis, a distinctly larger taxon. Among the four small species, morphological analyses showed that K. hardwickii s.str. differs significantly from the other three species in skul shape, whereas K. depressa, K. furva and the new species appear to be morphologically cryptic species. Molecular dating estimates suggest that species within the hardwickii-complex diversified during the Late Pliocene/Early Pleistocene period, most probably in different glacial refugia in Asia. Available evidence indicates that allopatric speciation within the complex included morphological, acoustic, and cytogenetic divergence, although these were not always mutually exclusive. Such adaptive changes would explain how different taxa with overlapping morphological features can share ecological niches and maintain their gene flow in sympatry. Accordingly, we also suggest that two subspecies of K. hardwickii sensu lato (K. h. crypta and K. h. malpasi) originally described as distinct species (from Southern India and Sri Lanka, respectively) can be re-elevated to species rank. Should these be found to be conspecific however, the name crypta would have priority and malpasi should be treated as its subspecies.
... for 5 min for Co1 or 10 min for Cytb; and a final hold at 10 ? (Hoffmann & Baker 2001;Hoffmann et al. 2008;Hulva et al. 2004;Khan et al. 2010). DNA sequencing was performed using an ABI PRISM 3730 DNA Analyzer (Applied Biosystems). ...
Although increased survey efforts using improved capture methods (particularly harp traps) have greatly expanded the quantity of Kerivoula specimens available in China, the understanding of the genus has been long constrained. After the recently published revision of the hardwickii-complex with the description of K. furva and re-evaluation of occurrence of K. titania in Taiwan, the critical overview of the previous data of Chinese Kerivoula (with the exception of K. picta, a strikingly colored and unmistakable species) is imperative. To clarify the taxonomy and distribution of the hardwickii-complex in China, 40 additional specimens collected from South China were studied through detailed morphological comparisons, multivariate statistical methods and phylogenetic inference. Our results evidently classified these specimens as K. furva instead of K. titania or K. hardwickii sensu stricto and together with the critical review of literature data indicate that all previous Chinese records of the two latter species were based on either misidentifications or outdated taxonomy. K. furva have so far been recorded in the Chinese provinces of Chongqing, Fujian, Guangdong, Guangxi, Hainan, Hunan, Jiangxi, Yunnan and Taiwan, but more field surveys are needed to confirm whether it could be found in other nearby provinces.
