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Peninsular India is an important region for mammalian diversification and harbors major biogeographic barriers. However, little is known about the role of this region in the diversification of bats though it harbors high chiropteran diversity. In this study, we used phenotypic, acoustic, and genetic markers to assess the diversification of Rhinolophus lepidus bats in South Asia. We first investigated if peninsular India is associated with speciation of R. lepidus. Further, we tested if the Palghat Gap acts as a biogeographic barrier to gene flow in this species. Our results revealed cryptic genetic diversity in peninsular India suggesting that this region holds at least one endemic species level lineage of the R. lepidus species complex. Analyses of populations of R. lepidus across the Palghat Gap in the Western Ghats revealed clinal variation in phenotype, with bats south of this barrier being bigger and emitting echolocation calls of higher frequency. We also observed that populations on either side of the Palghat Gap have remained genetically isolated since the mid-Holocene.

Species richness, diversity and abundance of bat fauna in Similipal Biosphere Reserve were studied during 2013–2015. Bats were surveyed by exploring the roosting sites and mist netting for 288 nights in primary forest, degraded forest, agricultural land and human habitation. A total of 21 species representing three species of frugivore bats and 18 species of insectivorous bats in seven families were recorded during the study, of which 152 individuals in 13 species were captured in mist nets. Species diversity and capture rate were recorded higher in primary forest. Vespertilionidae was reported as the most diverse family with five species, and Rhinolophus lepidus was reported to be the most abundant species with maximum capture rate during the study. As Similipal Biosphere Reserve is experiencing severe biotic pressure, it can be predicted that the structure and composition of bat fauna may be affected. The authors expect that their findings will be useful for future comparative studies to monitor the change over time and develop conservation plans.

Horseshoe bats (Family Rhinolophidae) show an impressive array of morphological traits associated with use of high duty cycle echolocation calls that they emit via their nostrils (nasophonation). Delicate maxilloturbinal bones inside the nasal fossa of horseshoe bats have a unique elongated strand-like shape unknown in other mammals. Maxilloturbinal strands also vary considerably in length and cross-sectional shape. In other mammals, maxilloturbinals help direct respired air and prevent respiratory heat and water loss. We investigated whether strand-shaped maxilloturbinals in horseshoe bats perform a similar function to those of other mammals, or whether they were shaped for a role in nasophonation. Using histology, we studied the mucosa of the nasal fossa in Rhinolophus lepidus, which we compared with Hipposideros lankadiva (Hipposideridae) and Megaderma lyra (Megadermatidae). Using micro-CT scans of 30 horseshoe bat species, we quantified maxilloturbinal surface area and skull shape within a phylogenetic context. Histological results showed horseshoe bat maxilloturbinals are covered in a thin, poorly vascularized, sparsely ciliated mucosa poorly suited for preventing respiratory heat and water loss. Maxilloturbinal surface area was correlated with basicranial width, but exceptionally long and dorsoventrally flat maxilloturbinals did not show enhanced surface area for heat and moisture exchange. Skull shape variation appears to be driven by structures linked to nasophonation, including maxilloturbinals. Resting echolocation call frequency better predicted skull shape than did skull size, and was specifically correlated with dimensions of the rostral inflations, palate, and maxilloturbinals. These traits appear to form a morphological complex, indicating a nasophonatory role for the strand-shaped rhinolophid maxilloturbinals. Anat Rec, 2018. © 2018 American Association for Anatomy.

Diurnal flight and foraging activity in insectivorous bats are atypical behaviours that have been recorded from islands with few avian predators and from locations with extended daylight hours. We present the first known observations of diurnal activity of Rhinolophus lepidus in forests on Tioman Island, Malaysia, recorded using visual surveys and acoustic monitoring. The bats were flying during the day and at night, and feeding buzzes detected suggest that they were actively foraging during the day. This appears to be a regular phenomenon on Tioman Island. The absence of resident diurnal avian predators that hunt below the forest canopy may account for the diurnal activity of R. lepidus in forests there.

Rhinolophus lepidus, Blyth, 1844, (family Rhinolophidae) is one of the 124 bat species found in India. Till date, 13 species have been reported from Delhi, with one of the identified roosting site present in a natural cave in Aravalli Biodiversity Park. Roost population with direct roost count estimation and morphometric measurements of 65 individuals (49h, 16f) were studied at the said location. Roosting behavior, and emergence time for Blyth’s horseshoe bat, R. lepidus were recorded from June 2015 to September 2016. Echolocation calls were recorded during roosting and release calls during foraging. The recordings revealed a frequency with the maximum energy of 108 kHz. This study also reports the phenomenon of torpor in bats from North India in the winter season for the first time.

Bats are unique mammals that are reservoirs of high levels of virus diversity. Although several of these viruses are zoonotic, the majority are not. Astroviruses, transmitted fecal-orally, are commonly detected in a wide diversity of bat species, are prevalent at high rates and are not thought to directly infect humans. These features make astroviruses useful in examining virus evolutionary history, epidemiology in the host, and temporal shedding trends. Our study screened for the presence of astroviruses in bats in Singapore, reconstructed the phylogenetic relations of the polymerase genes and tested for population characteristics associated with infection. Of the seven species screened, astroviruses were detected in Rhinolophus lepidus and Eonycteris spelaea. The R. lepidus sequences grouped with other Rhinolophus astrovirus sequences from China and Laos, while the Eoncyteris sequences formed a distinct clade with astroviruses from Rousettus spp. in Laos and Pteropus giganteus in Bangladesh, but not with other E. spelaea sequences. Longitudinal collections of Eonycteris feces demonstrated variable shedding. Juvenile status of bats was a risk factor for astroviruses. This study highlights the diversity of astroviruses in nectivorous and insectivorous bats in Singapore and provides a predictive framework for understanding astrovirus infection in these bats. It also suggests that in addition to host phylogenetic relatedness, host ecology, such as roosting behavior, may drive co-infections, virus maintenance and spillover.

74. Blyth’s Horseshoe Bat Rhinolophus lepidus French: Rhinolophe de Blyth / German: Blyth-Hufeisennase / Spanish: Herradura de Blyth Taxonomy. Rhinolophus lepidus Blyth, 1844, “ probably in the vicinity of Calcutta,” West Bengal, India. Rhinolophus lepidus is included in the pusillus species group, and appears to be sister to a clade including R. comutus, R. pumilus, R.perditus, R. monticolus, R. shortridgei, R. pusillus, R. refulgens, and R. cf. lepidus from South-east Asia, which makes the species basal to much of the pusillus species group. As currendy defined, R. lepidus probably represents a species complex. The Central Asian specimens originally misidentified as R. hipposideros in Uzbekistan and Kyrgyzstan were mentioned under the new name R. kirgisorum (or variously as R. aff. lepidus) by I. Horaòcek and colleagues in 2000, and were thought probably to represent a new species. There was no description given to the name, so it is a nomen nudum. Subsequent comparison by P. Benda and J. Gaisler in 2015 indicated that the specimens from these regions are similar to specimens in Afghanistan, suggesting that they should be included within the race monticola . Race monticola may represent a species distinct from the South Asian and South-east Asian taxa, pending further genetic and morphological studies. Distributional limits between the subspecies are not well defined. The juvenile type specimen of monticola may have a mismatched skull, as the specimen’s external measurements are within the range of R.pusillus whereas the skull is typical of lepidus. Three subspecies are currently recognized. Subspecies and Distribution. . l. lepidus Blyth, 1844 - most of India and Bangladesh; possibly in Bhutan. R..l. feae K. Andersen, 1907 — Myanmar, S China (Yunnan), N Thailand, SW Cambodia, and S Vietnam (possibly more widespread across Vietnam) including Con Son and Phu Quoc Is. . l. monticola K Andersen, 1905 - SE Uzbekistan, SW Kyrgyzstan, Afghanistan (Faryab, Kabul, Nangarhar, Parwan, and Zabul provinces), N Pakistan (Khyber Pakhtunkhwa and Punjab provinces), NW India (Himachal Pradesh and Uttarakhand), and Nepal. Descriptive notes. Head—body 33- 5—54 mm, tail 13- 3—26 mm, ear 11-20- 6 mm, hindfoot 5-5-10- 5 mm, forearm 37-42- 9 mm; weight 3-7-8- 2 g. Dorsal pelage varies between wood brown and cinnamon (hairs with drab bases with paler tip, giving fur glossy appearance), whereas ventral pelage is wood brown or drab. Ears are small or medium-sized. Noseleaf has strongly concave-sided lancet (may be almost parallel-sided in some individuals) and has either a rounded or pointed tip; connecting process is pointed triangle; sella is narrow and usually parallel-sided; horseshoe is relatively wide (6-8 mm) and does not cover the whole muzzle, has wide median emargination, and secondary leaflets are present. Lower lip has three mental grooves. Baculum is elongate, with dorsal bend near the basal cone and ventral bend near base, which is deeply bifurcated with depression on the ventral margin; tip is knobbed and wider in dorsal view, and shaft is cylindrical. Skull is small and robust (zygomatic width is usually subequal to, or sometimes slightiy larger than, mastoid width); anterior median swellings are small and subcircular; posterior swellings are relatively inflated; rostral profile is slightly sloped backwards or occasionally nearly straight; sagittal crest is moderately strong but weakened posteriorly; frontal depression is shallow to nearly straight; supraorbital ridges are low. C1 is well developed, usually greatly exceeding height of P4; P2 has distinct and generally well-developed cusp and is within tooth row; P3 is variably within or extruded from tooth row; P2 and P4 are in contact ifP3 is external. Chromosomal complement has 2n = 62 and FNa = 60 (southern India and Kyrgyzstan). Habitat. Both dry and moist forest and fringe areas, as well as some desert regions in Central Asia. Recorded at elevations of 100-2338 m. Food and Feeding. Blyth’s Horseshoe Bat appears to forage among vegetation and has been observed both aerial-hawking and gleaning prey off offoliage. It is a low, maneuverable flier. Known to feed on Lepidoptera, Coleoptera, Diptera, and Hymenoptera. Breeding. Females give birth to a single young in early May, in Maharashtra. Activity patterns. Blyth’s Horseshoe Bat roosts by day in caves, unused tunnels, old and ruined buildings, and old temples, and forages during the night. Search call shape is FM/CF/FM with a peak F recorded at 93-2-96-8 kHz and durations of 16-8-42-4 milliseconds in southern India. Peak frequency was reported as 104—105 kHz in another study in southern India and 103 kHz in South-east Asia. Movements, Home range and Social organization. Blyth’s Horseshoe Bat generally roosts in colonies ranging from a few individuals scattered within the roost or in compact clusters of a dozen or so, to several hundred individuals, some ofwhich may roost solitarily. Males and females roost together in mixed colonies. They reportedly urinate on intruders to deter them from disturbing their roosts. Status and Conservation. Classified as Least Concern on The IUCN ed List. Blyth’s Horseshoe Bat is generally widespread and does not seem to have any major threats affecting it as a whole. However, in parts of its Indian range, it is threatened by roost disturbance due to conversion of old forts into hotels as part of tourist-related development activities. Bibliography. Abramov eta/. (2007), Bates & Harrison (1997), Benda & Gaisler (2015), Benda, Hanak (2011), Bumrungsri, Francis & Csorba (2008), Csorba eta/. (2003), Das (1986a), Dejtaradol (2009), rancis (2008a), Horâcek eta/. (2000), Koublnova eta/. (2010), Kruskop (2013a), Raghuram eta /. (2014), Salim et al. (2017), Shahbaz et al. (2014), Sinha (1973), Smith &XieYan (2008), Soisook eta/. (2016)., Published as part of Burgin, Connor, 2019, Rhinolophidae, pp. 280-332 in Handbook of the Mammals of the World – Volume 9 Bats, Barcelona :Lynx Edicions on page 315, DOI: 10.5281/zenodo.3748525

To conserve biodiversity it is imperative that we understand how different species respond to land use change, and determine the scales at which habitat changes affect species' persistence. We used habitat suitability models (HSMs) at spatial scales from 100–4000 m to address these concerns for bats in the Western Ghats of India, a biodiversity hotspot of global importance where the habitat requirements of bats are poorly understood. We used acoustic and capture data to build fine scale HSMs for ten species (Hesperoptenus tickelli, Miniopterus fuliginosus, Miniopterus pusillus, Myotis horsfieldii, Pipistrellus ceylonicus, Megaderma spasma, Hipposideros pomona, Rhinolophus beddomei, Rhinolophus indorouxii and Rhinolophus lepidus) in a tea-dominated landscape. Small (100–500 m) scale habitat variables (e.g. percentage tea plantation cover) and distances to habitat features (e.g. distance to water) were the strongest predictors of bat occurrence, likely due to their high mobility, which enables them to exploit even small or isolated foraging areas. Most species showed a positive response to coffee plantations grown under native shade and to forest fragments, but a negative response to more heavily modified tea plantations. Two species were never recorded in tea plantations. This is the first study of bats in tea plantations globally, and the first ecological Old World bat study to combine acoustic and capture data. Our results suggest that although bats respond negatively to tea plantations, tea-dominated landscapes that also contain forest fragments and shade coffee can nevertheless support many bat species.

Rhinolophus lepidus Blyth, 1844 Rhinolophus lepidus Blyth, 1844: 486; Calcutta, INDIA (Collector unknown; Type unknown) [160]. Rhinolophus refulgens Andersen, 1905: 124, pl. 4; Gunung Igari, Perak, MALAYSIA, 2000 ft. (A. L. Butlerm Esq., presenter; BM(NH) 1898.11.29.2) [161]. Rhinolophus lepidus refulgens [9]. Common English name: Blyth’s Horseshoe Bat Barcode Index Number: BOLD:AAB9127 (5 DNA barcodes from Peninsular Malaysia; Fig 5) Remarks: DNA barcodes of R. lepidus are associated with three BINs (BOLD:AAB9127, BOLD:ABZ1016 and BOLD:ABZ2266; Fig 5). The BIN, BOLD:ABZ1016 contains DNA barcodes recorded as R. lepidus and R. pusillus. Some authors considered R. refulgens as a subspecies of R. lepidus [9, 11, 12] while some considered them to be distinct [50, 67]. Our NJ analysis (Fig 5) suggested that the DNA barcodes recorded as R. lepidus from Peninsular Malaysia may be distinct from DNA barcodes recorded as R. lepidus from Indochina. Similarly, Bumrungsri et al. [162] commented that R. lepidus from Peninsular Malaysia may represent a distinct taxon and the appropriate name would be R. refulgens based on the type locality. Due to the lack of DNA barcodes from the type locality of R. lepidus, India for comparison, we could not determine if DNA barcodes from Peninsular Malaysia represent the nominate R. lepidus or R. refulgens. Consequently, we tentatively retained only the name R. lepidus in this checklist (Fig 5). IUCN status: Least Concern Recorded at: Pahang: Krau Wildlife Reserve [11, 42], Kuala Atok, National Park [44], Lata Bujang Forest Reserve [56], Cameron Highland [60], Pulau Tioman [79], Bukit Ibam, Kemasul, Jengka, Klau Besar, Kenong and Gunung Aais [100]; Terengganu: Pasir Raja, Dungun [15], Tasik Kenyir [69], Gunung Tebu Forest Reserve [101]; Perak: Temengor Forest Reserve [46, 47, 111], Temenggor Lake [69]; Perlis: Wang Kelian State Park, [50], Gunung Stong State Park [100]; Selangor: Bukit Kutu Wildlife Reserve [51], Semangkok Forest Reserve [101]; Johor: Endau-Kota Tinggi Forest Reserve [56], Labis Forest Reserve [100]; Kedah: Ulu Muda Forest Reserve [57, 100], Bukit Hijau [100]; Kelantan: Air Panas-Gua Musang [61], Gunung Reng and Gua Musang [62]; Melaka: Unspecified [68]; Pulau Pinang: Bukit Panchor [100]; Negeri Sembilan: Gunung Angsi Forest Reserve [100, 101], Berembun Forest Reserve [101]. As R. refulgens: Perak: Maxwell Hill [23]; Pahang: Pulau Tioman [23], Krau Wildlife Reserve [41]; Johor: Pulau Pemanggil and Pulau Aur [23]; Negeri Sembilan: Pasoh Forest Reserve [45]; Kelantan: Gunung Stong State Park [67]. R. lepidus inhabits mature lowland and hill forests and has been reported roosting in caves and rock crevices, often with R. stheno [11, 14], Published as part of Voon-Ching Lim, Rosli Ramli, Subha Bhassu & John-James Wilson, 2017, A checklist of the bats of Peninsular Malaysia and progress towards a DNA barcode reference library, pp. 1-65 in PLoS ONE 12 (7) on pages 31-32, DOI: 10.1371/journal.pone.0179555, http://zenodo.org/record/4251352, {"references":["160. Blyth E. Notices of various Mammalia with descriptions of many new species. Journal Asiat Soc Bengal. 1844; 13: 463 - 494.","161. Andersen K. On the bats of the Rhinolophus philippinensis group, with descriptions of five new species. Annals Mag Nat Hist. 1905; 16: 546 - 550.","9. Corbet GB, Hill JE. The mammals of the Indomalayan region: a systematic review. Oxford: Oxford University Press; 1992.","11. Kingston T, Lim BL, Akbar Z. Bats of Krau wildlife reserve. Penerbit Universiti Kebangsaan Malaysia; 2006.","12. Csorba G. Description of a new species of Rhinolophus (Chiroptera: Rhinolophidae) from Malaysia. J Mammal. 1997; 78 (2): 342 - 347.","50. Jayaraj VK, Daud SH, Azhar MI, Sah SA, Mokhtar SI, Abdullah MT. Diversity and conservation status of mammals in Wang Kelian State Park, Perlis, Malaysia. Check List. 2013; 9 (6): 1439 - 48.","67. Jayaraj VK, Tahir NF, Udin NA, Baharin NF, Ismail SK, Zakaria SN. Species diversity of small mammals at Gunung Stong State Park, Kelantan, Malaysia. J Threat Taxa. 2012; 4 (6): 2617 - 2628.","162. Bumrungsri S, Francis CM, Csorba G. Rhinolophus lepidus; 2008 [cited 2016 May 22]. Database: The IUCN Red List of Threatened Species 2008: e. T 19547 A 8971569 [Internet]. Available from: http: // dx. doi. org / 10.2305 / IUCN. UK. 2008. RLTS. T 19547 A 8971569. en.","42. Anan S, Rashdi AM, Abdul J, Lim BL. A survey of small mammals at Sungai Chenderoh, Perlok, Krau Wildlife Reserve, Pahang. Journal of Wildlife and Parks. 1998; 16: 104 - 109.","44. Tingga RCT, Khan FA, Ridwan AM, Senawi J, Abdullah MT. Small Mammals from Kuala Atok, Taman Negara Pahang, Malaysia. Sains Malays. 2012; 41 (6): 659 - 669.","56. Mohd-Hanif RM, Nur-Aida MT, Zahirunisa AR, Mohd-Ridwan AR, Abdullah MT. Contribution of regenerated forest in conservation of bats in Peninsular Malaysia. J Trop For Sci. 2015; 27 (4): 506 - 16.","60. Shahfiz MA, Sahrul-Anuar MS, Shukor MN, Zalipah MN, Pan KA, Muin MA, et al. Survey of small mammals at two forest reserves in Cameron Highlands, Pahang, Peninsular Malaysia. In: Rahman A, Koh HL, Mohd-Paiz K, Muhamad A, Latiff A, editors. Proceedings for Forestry Department of Peninsular Malaysia, 3 - 5 August 2008. Department of Wildlife and National Parks, Malaysia; 2008. pp. 293 - 298.","79. Csorba G, Fuisz T, Kelen B. New birds and bats from Pulau Tioman, Malaysia. Mal Nat J. 1997; 50: 197 - 200.","100. Lim LS, Mohd-Adnan A, Zubaid A, Struebig MJ, Rossiter SJ. Diversity of Malaysian insectivorous bat assemblages revisited. J Trop Ecol. 2014; 30 (02): 111 - 121.","15. Wilson JJ, Sing KW, Halim MR, Ramli R, Hashim R, Sofian-Azirun M. Utility of DNA barcoding for rapid and accurate assessment of bat diversity in Malaysia in the absence of formally described species. Genet Mol Res 2014; 13 (1): 920 - 5. https: // doi. org / 10.4238 / 2014. February. 19.2 PMID: 24634112","69. Syaripuddin K, Kumar A, Sing KW, Halim MR, Nursyereen MN, Wilson JJ. Mercury accumulation in bats near hydroelectric reservoirs in Peninsular Malaysia. Ecotoxicology. 2014; 23 (7): 1164 - 1171. https: // doi. org / 10.1007 / s 10646 - 014 - 1258 - y PMID: 24840106","101. Joann CL, Fletcher C, Rahman KA. Spatial effects of virgin jungle reserves (VJR) on the community of insectivorous bats in Peninsular Malaysia. J Trop For Sci. 2013; 25 (1): 118 - 130.","46. Francis CM. The diversity of bats in Temengor Forest Reserve, Hulu Perak, Malaysia. Mal Nat J. 1995; 48: 403 - 408.","47. Shahfiz MA, Shahrul-Anuar MA, Kaviarasu M, Fauzi MZ, Rizal MR. Preliminary checklist of mammals at Sungai Enam in Temengor Forest Reserve, Perak, Peninsular Malaysia In: Mohamed AL, Krishnapillay DB, Kasim Z, Khushairi ZA, Ismail NI, Rezan EFN, Sadali A, Mohammad MS, Hassan MN, Aznan ARY, editors. Proceedings of the 2 nd Temengor Scientific Expedition 2012, 22 - 25 August 2013. Pulau Banding Foundation, Malaysia; 2013.","111. Joann CL, Fletcher C, Salim HMW, Abdul-Rahman K, Harrison RD, Potts MD. Insectivorous bat assemblage in the hill dipterocarp forest of Temengor Forest Reserve, Peninsular Malaysia. Mal Nat J. 2011; 63 (3): 569 - 576.","51. Lim BL, Majid RA, Norsham Y. Studies on the mammal fauna of Bukit Kutu Wildlife reserve, Hulu Selangor. Journal of Wildlife and Parks. 1999; 17: 1 - 6.","57. Norsham Y, Shariff F, Norhayati A, Nordin M, Lim BL. Pre-logging survey of mammal fauna at Sungai Weng sub-catchment, Ulu Muda Forest Reserve Forest Reserve, Kedah. Journal of Wildlife and Parks. 1999; 17: 28 - 43.","61. Hasan NH, Khan K, Senawi J, Ketol B, Sait I, Abdullah MT. A report on bats survey at Air Panas-Gua Musang, Kelantan, Malaysia. J Trop Biol Conserv. 2012; 9 (2): 156 - 162.","62. Jayaraj VK, Khan FA, Azhar I, WeeChen E, Ali MR, Ahmad A, et al. Diversity and conservation status of small mammals in Kelantan, Malaysia. Warasan Songkhla Nakharin. 2016; 38 (2): 213 - 20.","68. Shahfiz MA, Shahrul-Anuar MA, Nor-Zalipah M, Kaviarasu M, Pan KA, Yusof MO, et al. Surveys of bats at selected sites in Melaka Forest Reserves, Peninsular Malaysia. In: Rahman A, Koh HL, Muhamad A, Yusof WKW, Latiff A, editors. Proceedings for Forestry Department of Peninsular Malaysia, 1 - 2 April 2009 2008. Department of Wildlife and National Parks, Malaysia; 2009. pp. 265 - 270.","23. Medway L. The wild mammals of Malaya. London: Oxford University Press. 1969.","41. Zubaid A. A comparison of the bat fauna between a primary and fragmented secondary forest in Peninsular Malaysia. Mammalia. 1993; 57 (2): 201 - 206.","45. Francis CM. Trophic structure of bat communities in the understorey of lowland dipterocarp rain forest in Malaysia. J Trop Ecol. 1990; 6: 421 - 431.","14. Francis CM. A field guide to the mammals of South-East Asia. New Holland Publishers; 2008."]}

Mutual recognition is the product of species coexistence, and has direct effects on survival and reproduction of animals. Bats are able to discriminate between sympatric different heterospecifics based on their echolocation calls, which has been shown both in free-flying and captive bats. To date, however, the factors that may determine the behavioral responses of bats to echolocation calls from sympatric heterospecifics have rarely been tested, especially under well-controlled conditions in captive bats. Hence, we aimed at tackling this question by performing playback experiments (habituation–dishabituation) with three horseshoe bat species within the constant-frequency bat guild, which included big-eared horseshoe bats (Rhinolophus macrotis), Blyth’s horseshoe bats (Rhinolophus lepidus), and Chinese horseshoe bats (Rhinolophus sinicus). We studied the behavioral responses of these three species to echolocation calls of conspecifics, to other two species, and to another heterospecifics bat, Stoliczka’s trident bat (Asellisus stoliczkanus), which also belongs to this guild. We found that the three rhinolophid species displayed a series of distinct behaviors to heterospecific echolocation but few to conspecific calls after habituation, suggesting that they may have been able to discriminate sympatric heterospecific echolocation calls from those of conspecifics. Interestingly, the behavioral responses to heterospecific calls were positively correlated with the interspecific overlap index in trophic niche, whereas call design had only a minor effect. This implies that the behavioral responses of these bats to heterospecific echolocation calls may be related to the degree of interspecific food competition.

Rhinolophus lepidus ��� India: ♂ HZM.19.28161, ♀ HZM. 20.28162, Tughlakabad fort, New Delhi; ♂ HZM.31. 36209, ♀ HZM.30.36208, Shenkaltheri Kalakad Hill, Tamil Nadu; ♂ HZM.26.35089, Ayiraperi Cave, Courtallum Hills, Tamil Nadu; ♂ HZM.16.25684, Lohani caves, Mamdu, Madhya Pradesh; ♂ field no. 120429.1 (released individual), ♀ field no. 120428.1 (released individual), Enckikuki-Kukaran Podabu, Tamil Nadu; ♀ HZM.17.25685, HZM.18.25686, Dungeon, Hindola Palace, Dando, Madhya Prabesh; ♀ HZM.29.35005, HZM.27.35090, Kuraven Kuli Cave, near Inchikuli, Agesthiyar Hill, Tamil Nadu; ♂ BMNH.79.11.21.151 (type of monticola), Masuri, Kumaon; ��� Myanmar: ♂ BMNH.2003.63 (syntype of feae), ♀ BMNH.1907.1.9.16 (syntype of feae), Biapo, Karen Hills, NE of Tounshow (=Toungou?), S. Burma; ��� Vietnam: ♂ HZM.4. 32309, ♂ HZM.22.32310, ♀ HZM.23.32311, Kon Ka Kinh Nature Reserve, Gai Lai Province; ♂ HZM.21.32305, Kon Cha Rang Nature Reserve, Gai Lai Province; ��� Cambodia: ♂ HZM. 31.36495, Tatai Lieu, Koh Kong Province; ��� Thailand: ♀ PSUZC-MM.2007.257, Phu Suan Sai NP., Loei Province; ♂ PSUZC-MM2007.105, ♂ PSUZC-MM2007.106, Khao Plu, Lamae, Chumphon; ♀ PSUZC-MM2007.107, Silawan Cave, Chumphon; ♂ PSUZC-MM2007.88, Khao Chong, Trang; ♂ PSUZC-MM2006.101, ♂ PSUZC-MM2006.102, ♀ PSUZC- MM2006.100, ♀ PSUZC-MM2006.103, ♀ PSUZC-MM2006. 104, Boripatr Waterfall, Ton Nga-Chang WS, Rattaphum, Songkhla; ♀ PSUZC-MM2006.11, Ton Nga-Chang WS, Hat Yai, Songkhla; ♂ PSUZC-MM2006.105, ♀ PSUZC-MM2006. 106, ♂ PSUZC-MM2006.107, ♂ PSUZC-MM2006.108, ♂ PSUZC-MM2006.109, ♂ PSUZC-MM2006.145; ♀ PSUZC- MM2006.151, Khao Rak Kiet, Ratthaphum, Songkhla; ♂ PSUZC-MM2007.89, ♂ PSUZC-MM2007.91, ♀ PSUZC- MM2005.48, ♀ PSUZC-MM2007.90, Tarutao Island, Satun; ��� Malaysia: ♂ BMNH.67.1571, ♀ BMNH.67.1578, Base camp, Gunung Benom, Pahang, Malaysia., Published as part of Soisook, Pipat, Karapan, Sunate, Srikrachang, Mattana, Dejtaradol, Ariya, Nualcharoen, Kwan, Bumrungsri, Sara, Lin, Sai Sein, Aung, Moe Moe, Bates, Paul J. J., Harutyunyan, Margarita, Bu��, Magdalena M. & Bogdanowicz, Ies��aw, 2016, Hill forest dweller: a new cryptic species of Rhinolophus in the ' pusillus group' (Chiroptera: Rhinolophidae) from Thailand and Lao PDR, pp. 117-139 in Acta Chiropterologica 18 (1) on page 137, DOI: 10.3161/15081109ACC2016.18.1.005, http://zenodo.org/record/3756730

There is an apparent allometric relationship between peak frequency of echolocation and body size in rhinophilids. However, some rhinolophids deviate from this rule. To date this variation has been explained as a result of partitioning of communication channels. An alternative hypothesis that food resource partitioning results in this divergence in expected frequencies was tested by comparing prey selection between Rhinolophus macrotis Blyth, 1844 and Rhinolophus lepidus Blyth, 1844 in Yunnan Province, China. These two sympatric species are morphologically similar but acoustically divergent: R. macrotis has an echolocation frequency significantly lower than that predicted by the allometric relationship, whereas that of R. lepidus agreed with expectations. Prey selection experiments, conducted in a flight tent, indicated that the dominant prey taxa of R. macrotis were Lasiocampidae, Arctiidae and Noctuidae, whilst that of R. lepidus were Arctiidae, Noctuidae and Ichneumonidae. R. macrotis ate more earless moths and fewer eared moths than R. lepidus, and R. macrotis fed on larger prey in general and captured a wider size range than that captured by R. lepidus. These results confirmed the existence of finely tuned trophic niche differentiation and suggested that food resource partitioning is one of the factors leading to lower peak frequency of calls in R. macrotis.

Karyotypes and chromosomal data are presented for eight bat species representing two families (Rhinolophidae and Vespertilionidae) from Hainan Island, China. The species investigated were Rhinolophus lepidus (2n = 62, FN = 60), R. pusillus (2n = 62, FN = 60), R. affinis (2n = 62, FN = 60), R. sinicus (2n = 36, FN = 60), Myotis horsfieldi (2n = 44, FN = 52), Pipistrellus abramus (2n = 26, FN = 44), Miniopterus australis (2n = 46, FN = 50) and M. schreibersii (2n = 46, FN = 50). The karyotype of Rhinolophus lepidus is reported for the first time.

Rhinolophus lepidus Blyth, 1844 / R. pusillusTemminck, 1834 Blyth’s horseshoe bat/Least horseshoe bat New records Lampung Province: Lombok Village, Pemerihan Village, Sukaraja Forest, Sukaraja Village, Way Canguk Forest. New material Four individuals were collected as specimens. Lampung Province: Pemerihan Village, 1♀ (MZB 35051); Sukaraja Forest, 1♂, 1♀ (MZB 35795, 35796); Way Canguk Forest, 1♀ (MZB 35798). Previous records from Sumatra North Sumatra Province: Deli for R. lepidus (van Strien, 1996); Medan for R. pusillus (Csorba et al. 2003). Remarks This small Rhinolophus can be distinguished from other Rhinolophus species either by its smaller body size or by the shape of the connecting process, which is triangular and pointed at the tip. The posterior noseleaf (lancet) is triangular and pointed at tip. The coloration is light brown, grayish brown, or orange tipped with buffy brown. Individuals are generally characterized by yellowish brown and yellow skin of the face. Based on the characters above, this bat is most similar to R. lepidus and R. pusillus in the pusillus species group. The two Rhinolophus are hard to be distinguished externally and the taxonomic relationship between the two species is not clear (Csorba et al., 2003). Some studies suggest that peak frequency (F MAXE) is a diagnostic trait to distinguish R. lepidus and R. pusillus (Kingston et al., 2000; Zhang et al., 2009; Hughes et al., 2010). However, the discrepancy in echolocation calls reported from those studies indicates there is no clear relationship between peak frequency and species (R. lepidus versus R. pusillus: 100 kHz and 92.5 kHz in Kingston et al., 2000; 92–95 kHz and 100–111 kHz in Zhang et al., 2009; 100.1 kHz and 112.5 kHz in Hughes et al., 2010). Thus, we are not able to assign our samples to either of the two species. Individuals were captured with harp traps with an elevation from 50–507 m a.s.l. in lowland rainforest as well as plantations near forest in our study area. It was recorded roosting in the Gimbar 2 cave at Way Canguk Forest, and using houses as night roosts and feeding sites in Pemerihan Village., Published as part of Huang, Joe Chun-Chia, Jazdzyk, Elly Lestari, Nusalawo, Meyner, Maryanto, Ibnu, Maharadatunkamsi, Wiantoro, Sigit & Kingston, Tigga, 2014, A recent bat survey reveals Bukit Barisan Selatan Landscape as a chiropteran diversity hotspot in Sumatra, pp. 413-449 in Acta Chiropterologica 16 (2) on page 429, DOI: 10.3161/150811014X687369, http://zenodo.org/record/3943617, {"references":["VAN STRIEN, N. J. 1996. The checklist mammal fauna of the Gunung Leuser National Park. Pp. 133 - 203, in Leuser: a Sumatran sanctuary (C. P. VAN SCHAIK and J. SUPRIATNA, eds.). Yayasan Bina Sains Hayati Indonesia, Depok, iv + 348 pp.","CSORBA, G., P. UJHELYI, and N. THOMAS. 2003. Horseshoe bats of the world (Chiroptera: Rhinolophidae). Alana Ecology, Shropshire, xxxii + 160 pp.","KINGSTON, T., G. JONES, A. ZUBAID, and T. H. KUNZ. 2000. Resource partitioning in rhinolophoid bats revisited. Oecologia, 124: 332 - 342.","ZHANG, L., G. JONES, J. ZHANG, G. ZHU, S. PARSONS, S. J. ROS- SITER, and S. ZHANG. 2009. Recent surveys of bats (Mammalia: Chiroptera) from China. I. Rhinolophidae and Hipposideridae. Acta Chiropterologica, 11: 71 - 88.","HUGHES, A. C., C. SATASOOK, P. J. J. BATES, P. SOISOOK, T. SRITONGCHUAY, G. JONES, and S. BUMRUNGSRI. 2010. Echolocation call analysis and presence-only modeling as conservation monitoring tools for rhinolophoid bats in Thailand. Acta Chiropterologica, 12: 311 - 327."]}

The historical mammalian literature and recent surveys document a total of 24 microchiropteran species for the small equatorial island nation of Singapore. Intensive surveys carried out in the 1990s and subsequent records indicate that, of these 24 taxa, 15 are still present, another eight species, including all three hipposiderids, have become locally extinct and one other is indeterminate. Almost half (6) of the surviving species have very low abundances. Three new records of bat species in Singapore, namely Nycteris tragata, Rhinolophus lepidus and Murina suilla are assumed to have been present in the past but not detected in earlier surveys. Nycteris tragata and M. suilla are categorized as locally endangered. The vespertilionid, Myotis oreias, recorded previously only in Singapore, was not found in this survey and may be globally extinct.

Rhinolophus lepidus Blyth, 1844 Blyth’s horseshoe bat Rhinolophus lepidus Blyth, 1844: 486; Calcutta, India (see Das 1986). Distribution Afghanistan, Pakistan, N India, Nepal, Myanmar, Thailand, Szechwan and Yunnan (China), Peninsular Malaysia, Sumatra (Indonesia). Distribution in Lao PDR (Fig. 25): Louangnamtha Province: Ban Namya (ROM collection). Taxonomic notes Morphologically, in size and noseleaf structure, these specimens appear to match R. lepidus, but a detailed taxonomic study has not yet been undertaken to verify that this is the appropriate name for the species., Published as part of Thomas, Nikky M., Duckworth, J. W., Douangboubpha, Bounsavane, Williams, Meredith & Francis, Charles M., 2013, A checklist of bats (Mammalia: Chiroptera) from Lao PDR, pp. 193-260 in Acta Chiropterologica 15 (1) on page 208, DOI: 10.3161/150811013X667993, http://zenodo.org/record/3943341, {"references":["BLYTH, E. 1844. Notices of various Mammalia with descriptions of many new species. Journal of the Asiatic Society of Bengal, 13: 463 - 494.","DAS, P. K. 1986. Studies on the taxonomy of bats obtained by the Silent Valley (Kerala, India) Expedition, 1980. Records of the Zoological Survey of India, 8: 259 - 276."]}

This is the first karyotype description for Rhinolophus lepidus, which is widely distributed in Asia from Afghanistan to China, as well as in Sumatra [26]. The karyotype of R. lepidus from Hainan Island was 2n = 62 and FN = 60 (Fig. 1), consisting of 30 medium-sized to small acrocentric autosomal pairs, a large subtelocentric X chromosome, and a small acrocentric Y chromosome. Secondary constrictions were detected adjacent to one pair of medium-sized acrocentric chromosomes., Published as part of Wu, Yi, Motokawa, Masaharu, Li, Yu-Chun, Harada, Masashi, Chen, Zhong & Lin, Liang-Kong, 2009, Karyology of eight species of bats (Mammalia: Chiroptera) from Hainan Island, China, pp. 659-666 in International Journal of Biological Sciences 5 (7) on page 660, DOI: 10.7150/ijbs.5.659, http://zenodo.org/record/4266204, {"references":["26. Simmons NB. Order Chiroptera. In: Wilson DE, Reeder DM, eds. Mammal Species of the World: A Taxonomic and Geographic Reference, 3 rd ed. Baltimore: Johns Hopkins University Press; 2005: 312 - 529."]}

Rhinolophus lepidus Blyth, 1844 Blyth’s horseshoe bat Rhinolophus osgoodi Sanborn, 1939 Osgood’s horseshoe bat We consider these taxa together because it is not clear whether or not they are in fact conspecific (Corbet and Hill, 1992; Simmons, 2005). Csorba et al. (2003) consider R. osgoodi as a distinct species on the basis of lancet shape, its short upper canine and long palatal bridge, but recognise that specific distinctness of this taxon has not been accepted widely in the past. Rhinolophus osgoodi has only previously been recorded from one site (Nguluku, north of Likiang) in Yunnan. Osgood (1932) initially considered this taxon to be a subspecies of R. lepidus, and its specific status was introduced by Sanborn (1939) on the basis of its relatively long forearm and small cranial measurements. One male was captured in Fumin County, Yunnan on 4 November 2005, FA 43 mm, FMAXE 92 kHz. One female (FA 45.4 mm, FMAXE 95.2 kHz) was netted in Dali County, Yunnan Province on 4 September 2003. The Dali bat had a relatively long upper canine more typical of R. lepidus than R. osgoodi (Csorba et al., 2003), despite being captured close to the type locality for R. osgoodi. The similarities in forearm length and call frequencies of these bats suggest they may belong to the same taxon. Pottie et al. (2005) reported that R. lepidus in Singapore calls at 97.8 kHz, and had a FA of 39.9 mm, whereas this species in Malaysia calls at around 100 kHz and has a FA that ranges from 37.0– 43.9 mm (Kingston et al., 2000). More work is warranted to determine if R. osgoodi is distinct from R. lepidus. Previous records from China (R. lepidus): Anhui, Fujian, Guangxi, Guangdong, Guizhou, Hainan, Hunan, Hubei, Jiangxi, Sichuan, Yunnan and Zhejiang (Zhang, 1997; Wang, 2003). Rhinolophus osgoodi: Yunnan. Ecological Notes Seldom encountered and appeared to be rare and not widespread. Records were from caves., Published as part of Zhang, Libiao, Jones, Gareth, Zhang, Jinshuo, Zhu, Guangjian & Parsons, Stuart, 2009, Recent surveys of bats (Mammalia: Chiroptera) from China. I. Rhinolophidae and Hipposideridae, pp. 71-88 in Acta Chiropterologica 11 (1) on pages 76-77, DOI: 10.3161/150811009X465703, http://zenodo.org/record/3944511, {"references":["SANBORN, C. C. 1939. Eight new bats of the genus Rhinolophus. Publications of the Field Museum of Natural History Zoology, 24: 37 - 43.","SIMMONS, N. B. 2005. Order Chiroptera. Pp. 312 - 525, in Mammal species of the World: a taxonomic and geographic reference, 3 rd edition (D. E. WILSON and D. M. REEDER, eds.). The Johns Hopkins University Press, Baltimore, 2142 pp.","OSGOOD, S. I. 1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic expeditions. Publications of the Field Museum of Natural History Zoology, 18: 193 - 339.","POTTIE, S. A., D. J. W. LANE, T. KINGSTON, and B. P. Y. - H. LEE. 2005. The microchiropteran bat fauna of Singapore. Acta Chiropterologica, 7: 237 - 247.","KINGSTON, T., G. JONES, A. ZUBAID, and T. H. KUNZ. 2000. Resource partitioning in rhinolophoid bats revisited. Oecologia, 124: 332 - 342.","ZHANG, Y. Z. (ed.). 1997. Distribution of mammalian species in China. China Forestry Publishing House, Beijing, 280 pp.","WANG, Y. X. (ed.). 2003. A complete checklist of mammal species and subspecies in China: a taxonomic and geographic reference. China Forestry Publishing House, Beijing, 394 pp."]}

Extensive surveys were carried out from October 2001–September 2004 to assess the eco-status of chiropterans in and around the districts of Jodhpur (26°17′59N, 73°02′02E), Jaisalmer (26°54′47N, 70°54′52E) and Bikaner (28°01′22N, 73° 19′ 13E) in the Great Indian Desert (Thar Desert), India. The population dynamics and distributions of bats were examined at selected sites, revealing that the tremendous increase in human population, urbanization and construction of new buildings as well as the renovation of old heritage buildings have adversely affected chiropteran biodiversity in the region. Of the 11 micro- and one megachiropteran bat species recorded in Jodhpur in the early 1960s to 1980s, only eight species (seven micro- and one megachiropteran) have been found during the present study period, and dramatic changes have occurred at the roosting sites. Of the three microchiropteran species (Rhinopoma microphyllum kinneari, Rhinopoma hardwickii and Taphozous perforatus) recorded in Jaisalmer during the early 1960s, T. perforatus has vanished and has been replaced by Taphozous nudiventris at various sites. In Bikaner, four microchiropteran bat species were found at a number of roosting sites, whereas no megachiropteran has yet been documented. Of these, two species, namely Rhinopoma microphyllum kinneari and Pipistrellus tenuis, are here reported for the first time for this region, whereas the other two, namely Rhinopoma hardwickii and Rhinolophus lepidus, had been recorded already in the early 1960s.

An inventory of the mammal fauna of Phu Quoc Island, the largest offshore island of Vietnam lying 20 km off the Cambodian coast, has been conducted for the first time. A total of 17 species of small mammals were recorded: Cynopterus brachyotis, Cynopterus sphinx, Hipposideros ater, Rhinolophus shameli, Rhinolophus lepidus, Rhinolophus malayanus, Hesperoptenus blanfordi, Crocidura sp., Menetes berdmorei, Hylopetes alboniger, Berylmys berdmorei, Maxomys surifer, Rattus tanezumi, Rattus exulans, Niviventer fulvescens, Tupaia belangeri, Paradoxurus hermaphroditus. In addition, Pteropus vampirus, Callosciurus finlaysonii, Trachypithecus germani, Macaca fascicularis and Muntiacus muntjak were registered on the basis of field observations. Records for Sus scrofa and Cervus unicolor are based on interviews conducted. Most of the findings represent new distributional records for Vietnamese mammal fauna.

The description of this taxon as a subspecies of R. lepidus from Upper Burma (Myanmar) was published by O L D F I E L D THOMAS on behalf of ANDERSEN (1918), based on the short notes of the latter. The diagnostic characters of shortridgei ("skull and teeth averaging larger") appeared only in the key given for the species and subspecies of the pusillus-group but even without comparison of the measurements with the other named forms. According to SlNHA (1973) shortridgei differs from R. lepidus lepidus in having a longer hind foot (55-63% of the tibia, against 45.8-47.5%) and longer mandible. However, investigation of the type skull (BM(NH) 18.8.3.1) and other specimens (housed in the collection of USNM, FMNH, HNHM) revealed well-defined differences as compared with the other subspecies of R. lepidus; upper canines are strong, wide-based; sagittal crest extending posteriorly to the lambda and skull length is over 17 mm. Consequently, the taxon shortridgei is considered as a full species., Published as part of Csorba, G., 2002, Remarks on some types of the genus Rhinolophus (Mammalia, Chiroptera), pp. 217-226 in Annales Historico-Naturales Musei Nationalis Hungarici 94 on pages 219-220, DOI: 10.5281/zenodo.3839717, {"references":["SlNHA, Y. P. (1973): Taxonomic studies on the Indian horseshoe bats of the genus Rhinolophus Lacepede. - Mammalia 37 (4): 603 - 630."]}

Rhinolophus lepidus Blyth, 1844. J. Asiat. Soc. Bengal, 13:486. TYPE LOCALITY: India, Bengal, Calcutta (uncertain). DISTRIBUTION: Afghanistan, N India, Burma, Thailand, Szechwan and Yunnan (China), W Malaysia, Sumatra. SYNONYMS: cuneatus, feae, monticola, refulgens, shortridgei. COMMENTS: Includes feae, monticola, and refulgens; see Hill and Yoshiyuki (1980:180, 186); but also see Sinha (1973:620-621) who considered monticola a distinct species., Published as part of Karl F. Koopman, 1993, Order Chiroptera, pp. 137-241 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on pages 166-167, DOI: 10.5281/zenodo.7353060, {"references":["Hill, J. E., and M. Yoshiyuki. 1980. A new species of Rhinolophus (Chiroptera: Rhinolophidae) from Iriomote Island, Ryukyu Islands, with notes on the Asiatic members of the Rhinolophus pusillus group. Bulletin of the National Science Museum (Tokyo), ser. A (Zoology), 6: 179 - 189.","Sinha, Y. P. 1973. Taxonomic studies on the Indian horseshoe bats of the genus Rhinolophus Lacepede. Mammalia, 37: 603 - 630."]}

Rhinolophus lepidus Blyth, 1844. J. Asiat. Soc. Bengal, 13:486. TYPE LOCALITY: India, Bengal, Calcutta (uncertain). DISTRIBUTION: Afghanistan; N. India; Burma; Thailand; Szechwan, Yunnan (China); Malaya; Sumatra. COMMENT: Includes monticola and reftilgens; see Hill and Yoshiyuki, 1980, Bull. Nat. Sei. Mus. Tokyo, ser. A (Zool.), 6: 186; but also see Sinha, 1973, Mammalia, 37:620-621, who considered monticola a distinct species. ISIS NUMBER: 5301405007003035001 as R. lepidus. 5301405007003050001 as R. refulgens., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Chiroptera, pp. 111-215 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 145, DOI: 10.5281/zenodo.7352990, {"references":["Hill, J. E., and M. Yoshiyuki. 1980. A new species of Rhinolophus (Chiroptera: Rhinolophidae) from Iriomote Island, Ryukyu Islands, with notes on the Asiatic members of the Rhinolophus pusillus group. Bulletin of the National Science Museum (Tokyo), ser. A (Zoology), 6: 179 - 189.","Sinha, Y. P. 1973. Taxonomic studies on the Indian horseshoe bats of the genus Rhinolophus Lacepede. Mammalia, 37: 603 - 630."]}

on Rhinolophus lepidus Group. The ancestral species.��� The ancestors of the simpleX and lepidus groups were very closely related. The latter had a projectingconnecting process, a slightly smaller skull and teeth. But the general shape of the skull, the dentition, the nose-leaves, apart from the process and a very slight difference in the shape of the sella, the ears, the wing-structure, the length of the tail, and, we might even say, probably the size, were either identical or extremely similar in both of these extinct Bats. The place of origin. ���There can scarcely be any doubt that the lepidus group originated much farther westwards than the simpleX group. If we regard Japan as a continental group of islands, and put aside Java, on account of its peculiar geological history, we still find, not only the most primitive, but in fact all the species of the lepidus section on the Continent. It is only the section which has spread over the adjacent larger islands, one of which (Sumatra) has comparatively recently been continental, while another (Java), probably in a more remote period, seems to have been connected with some part or other of Indo�� China; and only one form, still so closely related to the Java species as hardly to be specifically different, has found its way so far eastwards as Lombok. The hypothesis, therefore, cannot be called unfounded, that of the two ancestral species, the ancient 1 simplex ��� and the ancient 11 lepidus, ��� the former was Eastern in range (Austro-Indo-Malayan), the latter Western (Oriental). Differentiation*.��� From a systematic point of view I found it convenient to divide the lepidus section into three ���types ���; I think that, phylogenetically speaking, there are two only: the lepidus and the minor type. The former, as coming nearest to simpleX in the proportionate size of the skull and teeth, is, probably, the more primitive; it is now distributed over the Indian Peninsula (lepidus), the Himalayas (monticola), and Malacca (refulgens). The latter, the minor -type, has spread from the Himalayas (minor) eastwards through S. China to Japan (cornutus); it is represented on the now quite isolated Anambas Islands ( ��� minutus ���); its occurrence in Java is not surprising, considering * the diagram on n. 138. the faunistic affinities of that island; and it has established itself on the western coast of the Indian Peninsula (gracilis). I have but very little doubt that now, when attention has been called to the differences of all these forms of the minor-type, it will be found also in other parts of the Indian Peninsula. If any inference can be drawn from fragments of a skull and the external characters, the subbadius-type would appear to be an offshoot of the minor-type: already in minor and cornutus the process is a little sharper-pointed than in lepidus; in subbadius and monoceros this tendency is carried much further. The skull of the species of the acuminatus section (Java- Lombok, Sumatra-Engano) is of the lepidus-ty pe; the process too; the colour remarkably like that of refulgens. This leads me to suppose that acuminalus and its allies (sumatranus, calypso) are scarcely more than giant representatives of the lepidus-type. It is the subbadius- type which, from a zoogeographical point of view, is by far the most interesting: it has spread southwestwards over a vast part of the Ethiopian Region, and westwards over the Mediterranean countries:��� (1) The empusa-type.��� Rh. empusa * and blasii have progressed further on the way already indicated by Rh. subbadius. They have the small skull and the small teeth characteristic of minor- subbadius; in the shape of the skull there is no essential difference; the dentition is identically the same; the process is that of a sub�� badius; the sella is deltoid, that is: the tendency, in the subbadius- sella (as emphasised above), towards assuming a subacute summit has been further developed; and we still see the constriction at the middle of the sella. But empusa and blasii are (as always the Ethiopian and W. Palaearctic species) in several points more highly developed: III.2 is lengthened (about, or more than, 1 the length of III1.); also IV.2 is very much longer (not far from twice the length of IV 1.). Rh. empusa is, however, an inhabitant of Nyasa- land, far S. of the Equator, Rh. blasii of the Mediterranean Subregion; thus, the two extremely closely allied species are now separated by an enormous tract, where no relative appears to occur. As we now know that they are descendants of the Oriental subbadius-type, the explanation seems to be quite clear: one branch spread south west wards, into the Ethiopian Region, and developed into Rh. empusa (slightly more primitive dentition; shorter ears, broader horse-shoe); another westwards into the Mediterranean countries, Rh. blasii. There is an instructive fact connected with these two Bats: I believe them to be comparatively recent intruders into their areas; Rh. empusa is known from one specimen only, from the very East of Tropical Africa; Rh. blasii is much more common in the Eastern Mediterranean tract, and still it does not seem to have reached Spain t. * Andersen, Ann. & Mag. Nat. Hist. (7) xiv. (1904) p. 378 (there is a p. 380: the length of the mandible is 2 ���, not 13*1 mm.). f Not recorded in Cabrera Latorre���s ���Quir��pteros de Espana, ��� Mem. Hist. Nat. ii. (1904). I am also not satisfied that there is any reliable the African coast of the Mediterranean. misprint on Soc. Espan. record from (2) The landeri-euryale type. ��� The Ethiopian Rh. landeri (Fernando Po, Gaboon), h. lobatus (Lower Zambesi to Mombasa), and Rh. dobsoni * (Kordofan) have the small skull and the small teeth characteristic of minor-subbadius; the same shape of the skull; the same dentition (no vacillation in the position of p); the process is that of a subbadius. In so far there is no difference at all between this section and the former (empusa-blasii). But in the shape of the sella and in a certain peculiarity in the wingstructure they have taken a course of their own:���We have seen, in the simpleX group, a progressive development from a sella constricted at the middle, through a parallel-margined stage, to a pandurate sella; we have seen in the lepidus group, too, the constricted sella {minor) modified into the parallel-margined (gracilis); the Ethiopian species here under consideration represent the third and final stage, the pandtvrate sella. In addition to this: in all of them IV.1 is peculiarly shortened���, less than (extremely rarely, as a slight individual atavism, equal to) half the length of IV2. As in Rh. empusa and blasii, III.2 is lengthened. Rh. euryale, from the Mediterranean Subregion, is so extremely closely allied to the above-named Ethiopian species that it shares with them all essential characters (even the highly peculiar shortening of TV. 1), with one exception: it has retained the parallelmargined sella. Summary.��� When discussing the affinities of the Ethiopian species of the Rh. simpleX group (above, pp. 117-20), I arrived at the conclusion that they are undoubtedly derived from Oriental types, and that, most probably, the ancestral species have spread directly from South Asia into the Ethiopian Region. As will be observed from this, a study of the Ethiopian representatives of the Rh. lepidus group leads to quite the same result: they have their closest known allies in the Oriental Region, but they are, without exception, considerably more highly developed than any of their Oriental relatives. Bats of the subbadius- type have evidently spread from some part of South Asia south westwards into the Ethiopian Region { empusa; landeri, lobatus, dobsoni ), and westwards over the Mediterranean countries (blasii; euryale). Of all the species of the Rh. lepidus group only one has found its way to Lower Egypt, Rh. euryale. It is a species exclusively Mediterranean in range, and unusually liable to differentiation into slightly differing local forms I. Its presence in Lower Egypt is easily explained by invasion from the adjacent Asiatic coast of the Mediterranean, where it is very common (specimens from Lower Egypt are indistinguishable from the Palestine form, Rh. e. judaicus) J. * Thomas, Ann. & Mag. Nat. Hist. (7) xiv. (1904) p. 156. f Andersen andMatschie, ���Ueber einige geographische Formender Untergattung Euryalis��� (SB. Ges. naturi'. Fr. Berlin, 1904, pp. 71-83). I Although it is beyond the strict limits of the present paper, I propose to insert a few words on the remaining Ethiopian species of the genus:���The aethiops section (Rh. aethiops, hildebrandti, and fumigatus) are very closely related to the Himalayan Rh. maorotis, but much more highly developed in the dentition, the wing- The probable affinities and phylogeny of the principal forms of the Rh. lepidus group are expressed in the subjoined diagram (Ethiopian types marked with an asterisk), Published as part of Andersen, Knud, 1905, On some Bats of the Genus Rhinolophus, with Remarks on their Mutual Affinities, and Descriptions of Twenty-six new Forms., pp. 75-145 in Proceedings of the Zoological Society of London 2 on pages 135-138, DOI: 10.5281/zenodo.3757451