Murray, Susan W., Khan, Faisal A. A., Kingston, Tigga, Zubaid, Akbar & Campbell, Polly, 2018, A new species in the Hipposideros bicolor group (Chiroptera: Hipposideridae) from Peninsular Malaysia, Acta Chiropterologica 20 (1), pp. 1-29: 21-23
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Hipposideros kunzi sp. nov. Murray, Khan, Kingston, Akbar, and Campbell
Kunz’s bicolored leaf-nosed bat
Hipposideros bicolor- 142 ( Kingston et al., 2001)
The species is named after Thomas H. Kunz in recognition of his many contributions to the ecology and conservation of bats, and his dedication to the promotion of bat research in Malaysia.
Texas Tech University TTU 108222 (tissue and karyotype TK 152065View Materials; field number VJS 155), adult ♂, body in alcohol, skull extracted, collected and photographed by Robert J. Baker on 6 August 2006 during TTU-UNIMAS Sowell Expedition ( Khan et al., 2008). Although the echolocation calls were not recorded for the holotype and the paratypes described here, all of the type specimens had mtDNA haplotypes consistent with the 142 kHz phonic group. This was further supported through comparisons of the noseleaf morphology with that of individuals for which the echolocation call frequency was known.
Measurements (in mm) — forearm length: 43.31; fifth, fourth, and third metacarpals lengths, respectively: 32.20, 33.87, 32.88. Length of first and second phalanges of third digits, respectively: 17.47, 16.46; tail length: 25.0; hind-foot length: 7.0; tibia length: 19.70; ear height: 17; body mass: 6.5 g; anterior noseleaf width: 4.66. Skull measurements are provided in Table 3.
Bukit Rengit, Krau Wildlife Reserve, Pahang, Peninsular Malaysia ( WGS 84 03°35’45.6”N, 102°10’ 59.0”E — approximate elevation 72 m). The specimen was collected using a harp trap set across a trail near the Institute of Biological Diversity at Bukit Rengit.
Texas Tech University TTU 108417 (tissue and karyotype number TK 152001View Materials), adult ♂ (4 August 2006), dry skin and skull with slight crack in brain case ; TTU 108209 (tissue number TK 152051View Materials), adult ♀ (6 August 2006), dry skin (housed at the Universiti Malaysia Sarawak, but missing) and skull (housed at the Texas Tech University). Both TTU 108417 and TTU 108209 were captured in Krau Wildlife Reserve (03°35’45.6”N, 102°10’59.0”E — elevation 72 m). Specimen TK 152992View Materials, adult ♀ (17 May 2008), dry skin and skull in Department of Wildlife and National Park ( DWNP), MalaysiaGoogleMaps ; specimen TK 153519View Materials, adult ♀ (20 May 2008), alcohol preserved specimen at Universiti Malaysia Sarawak. Both TK152992View Materials and 153519 were collected by FAAK during DWNP biodiversity inventory at Kuala Atok , Pahang, peninsular Malaysia (04°16.281’N 102°22.316’E — approximate elevation 85 m)GoogleMaps .
All specimens previously referred to H. atrox ( Douangboubpha et al., 2010) and H. bicolor -142 are here referred to H. kunzi sp. nov. Based on length of forearm, Hill (1963) likely included both H. bicolor and H. kunzi as H. bicolor atrox , although the majority of these individuals are probably H. kunzi based on length of forearm (p. 29, Fig. 4View FIG). We cautiously assign the individuals of H. bicolor atrox from both Hill et al. (1986) and Zubaid and Davison (1987) to H. kunzi . It is unclear where the bats were collected, but it is suggested they were captured in Northern peninsular Malaysia, which would suggest that they are indeed H. kunzi . In his description of the new species Hipposideros gentilis, Andersen (1918) described the new subspecies H. g. atrox as having a wide range of forearm lengths that span both H. bicolor and H. kunzi : 42–46.2 mm ( Andersen, 1918: 380). Thus he likely measured both individuals of H. bicolor and H. kunzi for the subspecies description.
This is a small to medium-sized hipposiderid bat in the H. bicolor group with a forearm length ranging from 38.8 to 45.6 mm (mean = 42.9 mm ± 0.9), tibia length of 17.1 to 20.6 mm (mean = 18.8 mm ± 0.5), and mass varying from 6.0 to 12.0 g (mean = 8.5 g ± 0.9 — Table 2). The dorsal pelage varies from medium or dark brown to bright orange, but is always bicolored with a white base. The ventral pelage ranges from buff or golden, to bright orange ( Fig. 9View FIG). The wing and tail membranes are dark brown, as are the ears. The ears are large (mean = 17.6 mm ± 0.6) and rounded with a bluntly pointed tip. The noseleaf lacks supplementary lateral leaflets and has an internarial septum that is generally triangular in shape (wider at the base — Fig. 9View FIG). The posterior and anterior portions of the nose are dark brown-grey in color, while the central part of the noseleaf is more flesh colored. The tail is long (mean = 28.7 mm ± 1.8), extending the full length of the uropatagium. The fifth metacarpal is about 74% of forearm length and the first phalanx of the third digit is about 53% of third metacarpal. Echolocation call frequency of the CF component ranges from 133.2 to 147.5 kHz, with a mean call frequency of 143.1 ± 2.0 kHz ( Fig. 5View FIG and Table 2).
Hipposideros kunzi has a small and elongate skull with the greatest length of skull ( GSL) ranging from 17.69 to 19.13 mm (mean = 18.31 ± 0.33 mm). The skull is slightly wider across the zygomata (mean = 9.2 ± 0.2 mm) compared to across the mastoids (mean = 9.2 ± 0.2 mm — Table 3). The distal process of the jugal bone is low and not well defined ( Fig. 6View FIG). The rostrum is well developed with six nasal inflations. The sagittal crest is well developed and is taller more anteriorly. The constriction behind the orbits is well defined and narrower than the rostrum. The upper toothrow is shorter ( CM 3 mean = 6.3 ± 0.1 mm) than the lower ( CM 3 mean = 6.8 ± 0.1 mm). The upper incisor is small and both the upper and lower canines are of moderate size. The upper premolar (P 2) is minute and extruded from the toothrow, while the lower premolar (P 2) is about half the height of the second premolar (P 4). The species is sexually dimorphic with respect to magnitude of certain skull measurements: despite being smaller than females, males have longer and taller skulls and longer canines.
Comparisons with similar species
Hipposideros kunzi is one of several Hipposideros species described from the Indo-Malayan region, which superficially resemble H. bicolor and lack supplementary leaflets adjacent to the noseleaf. In peninsular Malaysia and southern Thailand, H. kunzi most closely resembles, and is easily confused with, both H. bicolor and H. pomona . Compared to H. bicolor , H. kunzi has a higher echolocation call frequency ( Table 2), is generally smaller in body size ( Table 2), and has a shorter but wider skull ( Table 3 and Fig. 9View FIG). In addition, H. kunzi has a narrower anterior noseleaf (Holotype: 4.66 mm) that is slightly curved upwards compared to H. bicolor , which has a wider anterior noseleaf (4.94−5.46 mm, n = 5) that is flattened and square in appearance ( Kingston et al., 2006), lighter in color, and has rudimentary supplementary lateral leaflets ( Fig. 9View FIG). The noseleaf characters, however, are only useful if both species are available for comparison in the field.
Based on appearance ( Murray et al., 2012: figure S1), echolocation call frequency ( H. pomona : 136.4 –139.4 kHz, n = 3), overall size ( H. pomona length of forearm: 42.7–44.8 mm, n = 3), and skull size and shape ( Fig. 7View FIG), it is very difficult to distinguish H. kunzi from H. pomona . The main morphological difference between these species is ear height, with H. pomona having a much larger ear compared to H. kunzi : 20.0– 21.5 mm (n = 3) versus 15.0– 19.5 mm (mean = 17.6 mm — Table 2), respectively. Hipposideros pomona and H. kunzi , however, are not closely related based on both mitochondrial and nuclear DNA ( Murray et al., 2012; this study).
Despite being sister taxa ( Fig. 2View FIG), having similar appearance, and overlapping in echolocation call frequencies ( Kingston et al., 2000), individuals of H. kunzi and H. cineraceus -B are easily distinguished using body size ( H. kunzi being larger; Table 2) and nose morphology: H. cineraceus -B has a small swelling in its internarial septum ( Fig. 9View FIG).
In both 2003 and 2004 in peninsular Malaysia, palpably pregnant females were captured in February and March, and lactating individuals were captured from April through September. Similarly, Nurul-Ain et al. (2017) found females from Krau Wildlife Reserve and Samad Cave (ca. 10 Km from Krau) to be seasonally monestrous, with a peak in pregnancy in March, and lactation in June, although lactating females were captured from April through October.
Distribution, ecological notes, and conservation status
Currently, H. kunzi has only been documented on the Malay Peninsula, between 3°12’N in peninsular Malaysia ( Fig. 1View FIG, site 12) and the Isthmus of Kra at 10°41’N in Southern Thailand (this study; Douangboubpha et al., 2010). Despite extensive sampling, Douangboubpha and colleagues did not capture H. kunzi in Central or Northern Thailand, suggesting that the northern limit of this species’ range is restricted to the Sundaic biogeographical region, as delimited by the Isthmus of Kra ( Douangboubpha et al., 2010). While we did not sample bats in the southern tip of peninsular Malaysia, we expect that H. kunzi should occur throughout the peninsula where suitable habitat exists. Lim et al. (2014) reported a positive correlation between the abundance of H. kunzi (as H. bicolor -142) and latitude across 15 forest sites in peninsular Malaysia, with few or no captures at sites in the southern third of the Peninsula (which may be attributable to the lack of karst). In Singapore, H. bicolor (= H. kunzi ) is considered locally extinct due to habitat loss ( Pottie et al., 2005). Douangboubpha et al. (2010) included Sumatra in the distribution of H. atrox (= H. kunzi ), but because of the high level of cryptic diversity within this group it is impossible to determine whether individuals from Sumatra are conspecific with H. kunzi without genetic data. Based on limited sampling in Borneo ( Fig. 1View FIG), there is currently no evidence that H. kunzi occurs in Borneo.
In peninsular Malaysia, individuals of H. kunzi were captured at all sampling sites ( Fig. 1View FIG) and were relatively common and widespread in karst regions, but were also common in some non-karst areas (e.g., Krau Wildlife Reserve). Colonies ranged in size from a few individuals to several hundred and were found in caves, mines, and rock crevices. Colonies of H. kunzi were almost always found in caves housing other bat species; these included H. cervinus , H. larvatus , H. armiger , Rhinolophus malayanus , R. stheno , Myotis siligorensis , M. ater , Miniopterus medius , and Taphozous melanopogon . Based on captures and wing morphology, H. kunzi is believed to forage in forested habitats; Douangboupha et al. (2010) suggested that H. kunzi forages in diverse forest types and may be somewhat tolerant of anthropogenically modified landscapes that retain vegetative structure (e.g., secondary forest, rubber and orchard plantations). Given the species’ distribution across the Malay peninsula into Southern Thailand, widespread occurrence and local abundance, we currently recommend H. kunzi be evaluated as a species of Least Concern, following IUCN Red List Categories and Criteria v. 3.1 ( IUCN, 2012). Loss and disturbance of caves and foraging habitats would support a higher category of risk.
We and others have noted the high levels of cryptic diversity in Hipposideros (e.g., Esselstyn et al., 2012; Murray et al., 2012; Foley et al., 2017). We hope that our taxonomic delineation of a new member of the bicolor species group, H. kunzi , will motivate further efforts to resolve the taxonomy of remaining cryptic lineages. Such efforts are essential to the conservation of the remarkable diversity that exists within this already speciose genus.
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|Murray, Susan W., Khan, Faisal A. A., Kingston, Tigga, Zubaid, Akbar & Campbell, Polly 2018|
Hipposideros bicolor ( Temminck, 1834 )
|Hipposideros bicolor ( Temminck, 1834|
Hipposideros atrox ( Andersen, 1918 )
|Hipposideros atrox ( Andersen, 1918|
Hipposideros bicolor atrox ( Kitchener et al., 1996 )
|Hipposideros bicolor atrox ( Kitchener et al., 1996|
Hipposideros bicolor- 142
|Hipposideros bicolor- 142 ( Kingston et al., 2001|
|Hipposideros atrox (Douangboupha et al., 2010|