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Urban expansion has caused major deforestation and forest fragmentation in the tropics. The impacts of habitat fragmentation on biodiversity are understudied in urban forest patches, especially in the tropics and little is known on the conservation value of the patches for maintaining mammalian biodiversity. In this study, camera trapping was used to determine the species composition and species richness of medium‐ and large‐sized mammals in three urban forest patches and a contiguous forest in Peninsular Malaysia. We identified the key vegetation attributes that predicted mammal species richness and occurrence of herbivores and omnivores in urban forest patches. A total number of 19 mammal species from 120 sampling points were recorded. Contiguous forest had the highest number of species compared to the urban forest patches. Sunda Pangolin and Asian Tapir were the only conservation priority species recorded in the urban forest patches and contiguous forest, respectively. Top predators such as Malayan Tiger and Melanistic Leopard were completely absent from the forest patches as well as the contiguous forest. This was reflected by the abundance of wild boars. We found that mammal species richness increased with the number of trees with DBH less than 5 cm, trees with DBH more than 50 cm, and dead standing trees. In the future, the remaining mammal species in the urban forest patches are expected to be locally extinct as connecting the urban forest patches may be infeasible due to land scarcity. Hence, to maintain the ecological integrity of urban forest patches, we recommend that stakeholders take intervention measures such as reintroduction of selected species and restocking of wild populations in the urban forest patches to regenerate the forest ecosystems.

Hipposideros kunzi sp. nov. Murray, Khan, Kingston, Akbar, and Campbell Kunz’s bicolored leaf-nosed bat Hipposideros bicolor (Temminck, 1834), part. Hipposideros atrox (Andersen, 1918), part. Hipposideros bicolor atrox (Kitchener et al., 1996), part. Hipposideros bicolor- 142 (Kingston et al., 2001) Hipposideros atrox (Douangboupha et al., 2010) Etymology 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. Holotype Texas Tech University TTU 108222 (tissue and karyotype TK 152065; 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. Type locality 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. Paratypes Texas Tech University TTU 108417 (tissue and karyotype number TK 152001), adult ♂ (4 August 2006), dry skin and skull with slight crack in brain case; TTU 108209 (tissue number TK 152051), 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 152992, adult ♀ (17 May 2008), dry skin and skull in Department of Wildlife and National Park (DWNP), Malaysia; specimen TK 153519, adult ♀ (20 May 2008), alcohol preserved specimen at Universiti Malaysia Sarawak. Both TK152992 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). Taxonomic notes 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. 4). 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. 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. 9). 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. 9). 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. 5 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. 6). 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. 9). 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. 9). 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. 7), 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. 2), 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. 9). Reproduction 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. 1, 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. 1), 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. 1) 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.

Lim, Lee-Sim, Csorba, Gábor, Wong, Cheng-Min, Zubaid, Akbar, Rahman, Sultana Parvin Habeebur, Kumaran, Jayaraj Vijaya, Khan, Faisal Ali Anwarali, Najimudin, Joe Chun- Chia Huang Nazalan, Görföl, Tamás (2016): The systematic position of Hypsugo macrotis (Chiroptera: Vespertilionidae) and a new record from Peninsular Malaysia. Zootaxa 4170 (1): 169-177, DOI: http://doi.org/10.11646/zootaxa.4170.1.10

Knowledge on home range sizes and movement patterns of animals through the environment is crucial for determining effects of habitat disturbance and fragmentation. To gauge the effects of land-use changes on Scotophilus kuhlii, a telemetric study was conducted between February 2014 and April 2014 in Tasik Chini, Pahang and Universiti Kebangsaan Malaysia (UKM), Bangi Campus. The home range sizes and movement patterns of S. kuhlii inhabiting agricultural landscape (Tasik Chini, Pahang) versus urban landscape (UKM) were compared. A total of ten individuals were successfully radio-tracked. Comparison of home range sizes of both sexes showed male S. kuhlii at Tasik Chini have larger mean home range sizes compared to UKM while female S.kuhlii in UKM have larger mean home range sizes compared to Tasik Chini. All individuals from both localities showed random movement. It is suggested that the home range and activity patterns might be influenced by food availability in the study area, food preferences and diet segregation and breeding behavior. This study provides baseline information on habitat utilization by S. kuhlii in relation to habitat perturbations.

The prediction that the spatial dispersion of resources that are important to females should dictate female dispersion and male mating tactics has not been previously tested in harem-forming bats. We investigated the relation between the estimated spatial distribution of available roosts, female group size, male roost fidelity and the strength of social associations in two species of harem-forming fruit bats in the genus Cynopterus. We evaluated the daily movements of individuals and groups among roosts using radiotelemetry and roost censuses at two sites in northern peninsular Malaysia. We found a high correspondence between the distribution of roost sites, female group size and male behaviour, supporting the prediction that clumped resources would promote female aggregation and high roost fidelity in males. However, there were significant interspecific differences in the strength of male–female associations, which suggested that, where roosts are abundant and similar in quality, regardless of their spatial distribution, the potential for males to monopolize mates depends on whether females move among roosts with, or independently of, males. We propose that interspecific differences in female behaviour may be related to the costs of moving between clumped versus randomly dispersed males.

The aims of this study were to investigate the roosting ecology and social organization of the spotted-winged fruit bat, Balionycteris maculata (Megachiroptera), within an old-growth Malaysian dipterocarp forest, and test the hypothesis that males spend a significantly greater proportion of the night in the immediate vicinity of their day roosts than females. Balionycteris maculata roosted in cavities and formed small harem groups that consisted of one adult male accompanied by up to nine adult females and their dependent young. Both male and female bats foraged locally, within approximately 1 km of their roost sites. However, in contrast to females, which remained away from the roost site throughout the night, both harem and solitary adult males divided their nightly activity between foraging and returning to their roost sites. Individual females were also found roosting sequentially with up to three different males. Hence, these results suggest that B. maculata has a resource-defence polygynous mating system, in which the roost cavity may represent a critical and defendable resource by which males recruit and gain improved reproductive access to females. Roost cavities occupied by B. maculata were found within a number of different forest structures, including ant nests, termite nests and the root masses of epiphytes. The consistent shape and position of the cavities, along with a single observation of cavity enlargement, provide the first evidence of roost-cavity excavation, by a megachiropteran bat, within the nests of social insects. Although the cues by which females select their roost sites are unclear, some aspect of roost cavity size, or condition, may provide a possible means by which females can assess the reproducÍtive fitness of potential mates.

We describe design of echolocation signals in 5 species of aerial-hawking insectivorous bats from Malaysia. These bats forage in open spaces above the forest or in large clearings and belong to 2 families: Molossidae (Chaerephon johorensis, Mops mops, Cheiromeles torquatus) and Vespertilionidae (Hesperoptenus blanfordi, Pipistrellus stenopterus). As is typical for aerial-hawking bats, all 5 species produced narrowband calls of long duration (6‐21 ms) and low peak frequency (16‐44 kHz). However, sequences recorded from bats flying at high altitude ( .10 m) were characterized by an alternation between calls that differed in frequency of maximum energy (peak frequency), switching between high- and low-frequency calls. In some species, the types of calls also differed in duration and sound pressure level. We consider possible implications of the alternation of types of calls for the detection of prey in open spaces.

Estimates of insectivorous bat diversity in the Palaeotropics have largely been hampered by the lack of long-term studies employing appropriate capture techniques. Using a variety of trapping methods, 45 insectivorous bat species were captured in approximately 3 km2 of primary dipterocarp rain forest in Malaysia over 8 y. The cumulative site list for Kuala Lompat Research Station, Krau Wildlife Reserve, now stands at 51 insectivorous species. Although this is likely still not a complete list, it is already one of the most species-rich in the world. We attribute much of our success in recording this diversity to the extensive use of harp traps. Of the 45 species, 38 were captured in an intensive harp-trapping programme (> 1030 harp-trap nights) of the forest interior (22 species exclusively so). Insectivorous bats of the forest interior are thus a key component of Old World bat diversity, particularly in South-East Asia, and are dominated by taxa capable of detecting and capturing prey in cluttered environments (Hipposideridae, Rhinolophidae, Kerivoulinae and Murininae).

Kerivoula pellucida is a small (4.5 g) vespertilionid bat, which forages in the understorey of tropical rainforests in Southeast Asia. Most echolocation signals are characteristic for the subfamily (very high-frequency, broadband FM calls (sweeping down from 178 kHz to 58 kHz) of low intensity and short duration (< 3.0 ms)), but this species also produces stereotypical calls of a markedly different structure. The arrangement of signal elements in these calls is unusual, and we present here a detailed analysis of calls from thirteen adult individuals (nine males and four females). These multiharmonic calls are of much lower frequency (fundamental peak frequency of 36 kHz), higher intensity and longer duration (12.0 ms) than the orientation calls, and are composites comprising a long quasi-constant frequency (QCF) syllable followed by an upward frequency modulated sweep (UFM). From considerations of signal design and preliminary field observations, we consider possible functions of these calls.

Echolocation signal design in nine syntopic vespertilionid bats from the Malaysian rain forest understorey was studied. Four species of Kerivoula, two species of Phoniscus (Kerivoulinae) and three species of Murina (Murininae) all emitted calls that are typical of species that glean insects from surfaces: broadband, frequency-modulated (FM) calls of low intensity and short duration. However, calls were highly distinctive in the use of very large bandwidths (range: 89–123 kHz) and extremely high frequencies (start frequency 152–180 kHz; end frequency 43–86 kHz). Furthermore, calls were produced in groups of 2–15 at very high pulse repetition rates (37–105 Hz). The functional significance of these characteristics with respect to foraging strategy is discussed. Large signal bandwidths facilitate highly accurate target localization in terms of both range and angle estimation and can thus be interpreted as an adaptation to foraging in the highly cluttered environment of the forest understorey. The use of high frequencies so far in excess of those seen in other FM gleaning bats of the same size is less easily explained, but may represent a mechanism by which these species distinguish prey items using echolocation alone, without recourse to visual or auditory cues. Species exhibited differences in echolocation parameters, particularly the end frequency and the number of calls per group of calls. The two subfamilies differed from each other in multivariate space derived from echolocation parameters; calls of the Murininae were of lower frequency than the Kerivoulinae and were typically produced in smaller groups or singly. Within the subfamilies there was considerable overlap between species of Murina, but the Kerivoula spp. were clearly distinct from one another but not from the two Phoniscus spp.

The wood mouse is primarily a woodland species but it also occurs on maritime sand-dunes. The mice living on the sand-dunes leave their nests earlier in the evening and spend more time out of the nest than those animals living in woodlands. Here we test the hypothesis that this difference is because woodland provides substantially more food than do sand-dunes. Our experimental approach was to provide supplementary food in the form of wheatgrain to a sand-dune population and compare activity patterns with a control population. The mice on the supplemented area were strictly nocturnal and spent less time above ground than the controls. Their activity pattern was essentially the same as that of wood mice living in deciduous woodland.

International audience; The impact of ecology and social organization on genetic structure at landscape spatial scales, where gene dynamics shape evolution as well as determine susceptibility to habitat fragmentation, is poorly understood. Attempts to assess these effects must take into account the potentially confounding effects of history. We used microsatellites to compare genetic structure in seven bat species with contrasting patterns of roosting ecology and social organization, all of which are codistributed in an ancient forest habitat that has been exceptionally buffered from radical habitat shifts. Over one thousand individuals were captured at foraging sites and genotyped at polymorphic microsatellite loci. Analyses of spatially explicit genotype data revealed interspecies differences in the extent of movement and gene flow and genetic structure across continuous intact forest. Highest positive genetic structure was observed in tree-roosting taxa that roost either alone or in small groups. By comparison, a complete absence of genetic autocorrelation was noted in the cave-roosting colonial species across the study area. Our results thus reveal measurable interspecies differences in the natural limits of gene flow in an unmodified habitat, which we attribute to contrasting roosting ecology and social organization. The consequences of ecology and behaviour for gene flow have important implications for conservation. In particular, tree-roosting species characterized by lower vagility and thus gene flow will be disproportionally impacted by landscape-scale forest clearance and habitat fragmentation, which are prevalent in the study region. Our method also highlights the usefulness of rapid sampling of foraging bats for assaying genetic structure, particularly where roosting sites are not always known.

International audience; The potential for parallel impacts of habitat change on multiple biodiversity levels has important conservation implications. We report on the first empirical test of the 'species-genetic diversity correlation' across co-distributed taxa with contrasting ecological traits in the context of habitat fragmentation. In a rainforest landscape undergoing conversion to oil palm, we show that depauperate species richness in fragments is mirrored by concomitant declines in population genetic diversity in the taxon predicted to be most susceptible to fragmentation. This association, not seen in the other species, relates to fragment area rather than isolation. While highlighting the over-simplification of extrapolating across taxa, we show that fragmentation presents a double jeopardy for some species. For these, conserving genetic diversity at levels of pristine forest could require sites 15-fold larger than those needed to safeguard species numbers. Importantly, however, each fragment contributes to regional species richness, with larger ones tending to contain more species.

The woodmouse is primarily a woodland species but it also occurs on maritime sand-dunes. The home ranges of mice living on the sand-dunes are much larger than those of animals living in woodlands. Here we test the hypothesis that this difference is due to the fact that woodland provides substantially more food than do sand-dunes. Our experimental approach was to provide supplementary food in the form of wheatgrain to a sand-dune population and to compare range sizes with a control population. Range sizes on the supplemented area were significantly smaller than the controls and essentially similar to those of woodmice living in deciduous woodland.

The woodmouse is a characteristic rodent of deciduous woodland but it also lives on maritime sand-dunes. The population density of woodmice on sand-dunes is much lower than in woodland. Hue we test the hypothesis that this difference is due to the fact that woodland provides substantially more food than do sand-dunes. Our experimental approach was to provide supplementary food (wheat grain) to a sand-dune population for a period of 22 months and to compare demographic responses with a control population. The supplemented population increased rapidly and reached a size 6–7 times that ofthe control population. The overall annual pattern of changes in density were similar in both populations with minimum numbers in late spring, maximum numbers in autumn and a loss of individuals over the winter. The woodmice receiving supplementary food bred earlier and for longer than did the control mice.