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Rhinolophus beddomei sobrinus Rhinolophus beddomei sobrinus Andersen 1918 Kala Oya, North Central Province, Ceylon (= Sri Lanka). Sri Lankan Woolly Horseshoe Bat. Diagnosis: A medium-sized Rhinolophid belonging to trifoliatus group, with an average forearm length of 57.36 ± 1.11 mm. Horseshoe is broad, does not cover the whole of the muzzle, and possesses well-developed basal lappets. One mental groove is seen on the lower lip. Sella long (50–60% of the lancet) and distinctly shaped; superior connecting process rounded off to an anteriorly-projected inferior extremity; inferior connecting process slightly wavy. Lancet high, with a broad concave lower part (about 60% from the base) and a distinctly narrower tapering upper part. Skull robust with a CCL of 22.41 ± 0.26 mm; the maxillary tooth row (CM 3) is 9.58 ± 0.28 mm. Supraorbital depression deep and the supraorbital ridges are well-defined. First upper premolar (PM 2) small with a distinct cusp, and is in the tooth row. The second lower premolar (PM 3) is minute and is slightly extruded from the tooth row, leaving a small gap between PM 2 and PM 4. Body covered with dense and woolly fur, which is light brown to dark greyish brown. Baculum measures 4.94 mm long with a long shaft and a two-pronged broad base. External characters: This is a medium-sized Rhinolophid (FA 57.36 ± 1.11 mm; Table 3). Ears large (EL 29.50 ± 2.56 mm) and broad, with a pointed tip; antitragus well-developed and broadly triangular in shape, tragus absent; ridges 9 in number; the basal half of the ear covered with hair while the apical half scantly haired and the tip is totally devoid of hair (Fig. 9a); a concavity is observed on the outer border just below the tip. The lower lip has one mental groove (Fig. 9b). Horseshoe broad and flat, does not cover the muzzle entirely, with a well-developed, deep, and broad anterior median emargination (Fig. 9d). Internarial cup of the horseshoe broad, attached to the horseshoe by a narrow and short stalk, and flanked by two distinct lateral basal lappets when viewed frontally; nares teardrop-shaped and relatively large, located on either side of the base of the internarial cup. Nares large in size causing the lateral borders of the internarial cup to be upturned. Sella long (50–60% of the height of the lancet) and projects outwards (Fig. 9e), distinctly complex when viewed laterally, typical to species of the Rhinolophus trifoliatus group. Superior connecting process of the sella rounded off to an anteriorly-projected inferior extremity; inferior connecting process slightly wavy, connecting basally on both sides to the lateral lappets. Base of the sella broad, concave in the middle; it narrows halfway up its length towards the upper part, and the sides are parallel continuing towards a rounded-off tip (Fig. 9c). Lancet high, with a broad concave lower part (about 60% from the base) and a distinctly narrower tapering upper part. In the wing, the third metacarpal is much shorter than the fourth and fifth metacarpals (40.01 ± 1.5 mm vs. 45.57 ± 0.38 mm and 47.47 ± 1.62 mm respectively). The first and second phalanges of the third metacarpal are 52.26% and 83.41% of the third metacarpal respectively. The wing membrane is attached to the base of the first toe, and the interfemoral membrane is attached to the tibia (Fig. 9h). Body covered with dense fur. Face is hairy, but hair around eyes and the horseshoe less dense. On the ventral surface the fur extends along the forearm and a little below the anal region. The other parts of the membranes are devoid of hairs. On the dorsal surface the fur does not extend beyond the body. The penis is slender, parallel-sided, tapering to a narrowly rounded tip (Fig. 9f, g). Colouration (live): Fur woolly with fur colour in live condition being greyish brown. Individual hairs have grey bases followed by dark brown to fawn middle portion and pale to white hair tips (Fig. 6b). Craniodental characters: The GTL and CCL of the skull are 26.64 ± 0.87 mm and 22.41 ± 0.26 mm respectively and the skull is narrow and robust (Fig. 10a) (Table 3). Sagittal crest well-developed, connects to the supraorbital ridges (Fig. 10a), and extends up to the parietal region of the cranium (Fig. 10b). Rostrum robust, bulged, and relatively tall. The nasal inflations are well-developed and are located median on the rostrum, anterior to a deep supraorbital depression, which is flanked by well-defined supraorbital ridges. The sagittal crest is higher than the height of the rostrum. The skull is broadest at the widest point of the zygomatic arches. Zygomatic arches wide and strong (ZB 13.12 ± 0.40 mm), with a blunt triangular dorsal arch immediately anterior to the widest point (Fig. 10c). The second upper premolar (PM 2) is small with a distinct cusp, situated in the tooth row in contact with the canine and PM 4. The fourth upper premolar (PM 4) is roughly 58% the height of the upper canine (C 1; Fig. 10b). The upper tooth row (CM 3 9.58 ± 0.28 mm) is anteriorly convergent; C 1 –C 1 is about 67.86% of M 3 –M 3. The first and the second upper molars (M 1 & M 2) are equal in size. The third upper molar (M 3) is two-third the size of the first and the second molars; the metacone of M 3 is reduced and the metastyle is lacking (Fig. 10d). Two pairs of tricuspidate mandibular incisors (I 1 –I 3) are present (Fig. 10 e & g); the second lower premolar (PM 2) is small and half the height of the fourth lower premolar (PM 4); The third lower premolar (PM 3) is minute and slightly extruded from the tooth row, leaving a gap between PM 2 and PM 4 (Fig. 10e). The fourth lower premolar (PM 4) is about 65% the height of the lower canine, and roughly the same height as the first lower molar (M 1; Fig. 10f). The first and second lower molars M 1 and M 2 are of the same size while the third molar M 3 is slightly smaller than the other two. The talonoid of M 3 is broader than the trigonid, and the entoconid is lacking. Baculum: The baculum of R. b. sobrinus measures 4.94 mm long and is comprised of a thick shaft which expands into a two-pronged broad and robust base (1.76 mm) (Fig. 11 a,b). The base shows the presence of a deep groove on the ventral surface. The shaft is slender, ends with a broadly rounded tip and shows a concavity just below the tip which is visible in the lateral profile (Fig. 11c). Ecology: The Sri Lankan Woolly Horseshoe Bat is mostly found in caves, overhanging rock ledges, tree hollows, wells, old buildings, old abandoned plumbago mines, and tunnels, in or near dense dry and tropical moist forests, typically hanging by one foot, with the wings wrapped around the body. They live solitary, in pairs or in small parties of up to four individuals. They produce one to two pups per brood in Sri Lanka (Phillips 1980; Edirisinghe et al. 2016; Kusuminda et al. 2018). It has been recorded at elevations ranging from 43 m asl to 462 m asl (one specimen from Medamahanuwera, Central Province was found at 1077 m asl; Bates & Harrison 1997). Distribution: The Sri Lankan Woolly Horseshoe Bat is endemic to Sri Lanka. It has been recorded from many parts of the island except the northern and the eastern Provinces. The northernmost and southernmost records of the species are from Kala Oya (North Central Province) and Thalgasmankada (Southern Province) respectively., Published as part of Srinivasulu, Aditya, Srinivasulu, Bhargavi, Kusuminda, Tharaka, Amarasinghe, Chamara, Ukuwela, Kanishka D. B., Karunarathna, Mathisha, Mannakkara, Amani, Yapa, Wipula B. & Srinivasulu, Chelmala, 2023, Taxonomic status of the Lesser Woolly Horseshoe bats (Chiroptera, Rhinolophidae Rhinolophus beddomei) in peninsular India and Sri Lanka, pp. 199-218 in Zootaxa 5301 (2) on pages 211-215, DOI: 10.11646/zootaxa.5301.2.3, http://zenodo.org/record/8030308, {"references":["Andersen, K. (1918) Diagnoses of new bats of the families Rhinolophidae and Megadermatidae. Annals and Magazine of Natural History, 2, 374 - 384.","Phillips, W. W. A. (1980) Manual of the mammals of Sri Lanka. Part 1. 2 nd Revised Edition. Wildlife and Nature Protection Society of Sri Lanka, Colombo, 116 pp.","Edirisinghe, W. G. M., de Silva, I. M. C., Kusuminda, T. G. T., Thilina, M. H. D. K. & Gunawardana, K. D. S. D. (2016) New breeding data on Rhinolophus beddomei in Sri Lanka: First record of juveniles. Barbastella, 9 (1), 1 - 4. https: // doi. org / 10.14709 / BarbJ. 9.1.2016.03","Kusuminda, T., Mannakkara, A., Patterson, B. D. & Yapa, W. B. (2018) Bats in tea plantations in Sri Lanka: species richness and distribution. Journal of Bat Research & Conservation, 11 (1), 96 - 105.","Bates, P. J. J. & Harrison, D. L. (1997) The Bats of the Indian Subcontinent. Harrison Zoological Museum Publications, Sevenoaks, 258 pp."]}

Rhinolophus beddomei beddomei Rhinolophus beddomei Andersen 1905, Wynaad, Madras (= Kerala), India. Lesser Woolly Horseshoe Bat. External characters: A large-sized Rhinolophid belonging to trifoliatus group, with an average forearm length of 63.19 ± 2.63 mm. Fur dense and woolly, dark grey in colour with pale hair tips. The hair is very densely arranged on the body both dorsally and ventrally. The extent of fur is little beyond the body and extends sparsely on to the interfemoral on the dorsal surface. Hair is seen on either side of the legs just above the knee area to upper half of the tibia. Ears tall (28.98 ± 2.04 mm) and broad with a pointed tip. Antitragus broad and tall and is half the height of the pinna. Ear has 12 ridges. Long dense hair covers half the pinna, the distal half has scattered hair and the extreme tip is devoid of any hair. A concavity is observed on the outer border just below the tip (Fig. 5a). Only one mental groove on the lower lip is seen (Fig. 5b). The horseshoe is about 4 mm broad and covers the whole of the muzzle completely. The median emargination is about 3 mm deep but does not divide the horseshoe into two. The outer borders of the horseshoe are wavy. Nostrils are large and teardrop shaped and placed very close to each other. The posterior borders of the internarial cup are upturned due to large size and placement of the nostrils (Fig. 5d). The base of the sella extends on either side and forms the basal lappets which are large and circular and hang over the internarial cup and the distal half of the narial openings. The sella forms a complex structure when viewed laterally and overhangs the internarial cup (Fig. 5e). The inferior extremity of the sella is concave and slightly wavy. The sella is small about 40% the height of the lancet, has a broad base and is narrow distally (Fig. 5c). The superior connecting process of the sella broadly rounded off. The lancet is about 4 mm tall and ends with a narrowly rounded tip. In the wing the third metacarpal is shorter than the fourth metacarpal and the fifth metacarpals (42.71 ± 0.78 mm vs 50.38 ± 2.11 mm and 49.87 ± 1.99 mm). The first and the second phalanges are 57.57% and 76.75% of the third metacarpal respectively. Penis is thick, short and has a groove just below its tip, which is broadly rounded off (Fig. 5f). The wing membranes essentially naked with sparse and scattered hair. On the ventral surface the membranes have scattered sparse hair. Face and noseleaf very hairy. Feet hairy and large. Wings attached to the base of the first toe and the interfemoral membrane is attached to the ankle (Fig. 5g). Colouration (live): In the live condition the fur is woolly, long and the colour is dark grey to black with paler hair tips (Fig. 6a). Craniodental characters: Skull is long, narrow and robust (GTL: 28.12 ± 0.43 mm; CCL: 24.01 ± 0.47 mm) (Fig. 7a). Rostrum is elevated and has nasal swellings located medially on the rostrum. The sagittal crest is well developed and joins the supraorbital processes just behind the rostrum to form a deep supraorbital depression. The sagittal crest extends up to the parietal region of the skull. The lambdoid crests are not well developed. The sagittal crest and the rostrum are of the same height (Fig. 7b). Zygoma are robust, well developed and flared (ZB: 14.06 ± 0.56 mm); a triangle shaped dorsal arch seen on each zygoma (Fig. 7c). The upper tooth row (CM 3) averages 10.50 ± 0.38 mm. The second upper premolar (PM 2) is small, present in the tooth row, and located between the canine and the fourth upper premolar (PM 4). In comparison to PM 4, PM 2 seems minute and is situated between the canine and the PM 4. The PM 2 is in contact with the canine and is almost touching PM 4 (Fig. 7d). The fourth premolar is about three quarters the height of the robust canine. M 1 and M 2 are of the same size. M 3 is three quarters the size of M 2 but lacks the metastyle portion of the molar. Two pairs of tricuspidate mandibular incisors are present. The lower canines are slender than the upper. The canine shows a slight concavity on the posterior border (Fig. 7e, f). The second lower premolar (PM 2) is about 66% the height of the fourth premolar (PM 4). The third lower premolar (PM 3) is minute and is in the tooth row in some specimens resulting in a gap between PM 2 and PM 4 and in some slightly extruded from the tooth row resulting in PM 2 and PM 4 almost touching. M 1 and M 2 equal in size, M 3 slightly smaller than M 2. The talonoid of M 3 is broader than the trigonid, and the entoconid less developed (Fig. 7f). Baculum: The baculum of R. b. beddomei is long, measuring 5.8 mm, comprise a narrow distal shaft and a thick triangular base (2.2 mm wide) with a deep median groove (Fig. 8a). The shaft ends with a narrowly rounded tip. The base has a clear sulcus just above the median groove (Fig. 8b). The distal end of the shaft possesses a ridge-like protuberance towards the tip of the shaft on the ventral surface (Fig. 8c). Ecology: The Lesser Woolly Horseshoe Bat has been observed to roost solitary or in pairs, often mothers with pups have been observed to roost together. Roosting sites include old dilapidated buildings, temples, abandoned wells, tree hollows, old forts near the proximity of forested areas and caves. It has been observed with one pup and has been also observed to become inactive, tightly wrapping itself with its wings during winter season and sharing its roost with other Rhinolophid species. Distribution: The Lesser Woolly Horseshoe Bat is endemic to India, and is distributed in Maharashtra, Karnataka, Kerala, and Andhra Pradesh., Published as part of Srinivasulu, Aditya, Srinivasulu, Bhargavi, Kusuminda, Tharaka, Amarasinghe, Chamara, Ukuwela, Kanishka D. B., Karunarathna, Mathisha, Mannakkara, Amani, Yapa, Wipula B. & Srinivasulu, Chelmala, 2023, Taxonomic status of the Lesser Woolly Horseshoe bats (Chiroptera, Rhinolophidae Rhinolophus beddomei) in peninsular India and Sri Lanka, pp. 199-218 in Zootaxa 5301 (2) on pages 207-211, DOI: 10.11646/zootaxa.5301.2.3, http://zenodo.org/record/8030308, {"references":["Andersen, K. (1905) On the Rhinolophus philippinensis group and five new species. Annals and Magazine of Natural History, 16 (7), 243 - 257."]}

103. Beddome’s Woolly Horseshoe Bat Rhinolophus beddomei French: Rhinolophe de Beddome / German: Beddome-Wollhaar-Hufeisennase / Spanish: Herradura de Beddome Other common names: Bedomme's Horseshoe Bat Taxonomy. Rhinolophus beddomei K. Andersen, 1905, “ Wynaad [= Wayanad], Mysore [= Mysuru], S[outh]. India.” Rhinolophus beddomei is included in the trifoliatus species group and appears to be close to R. trifoliatus, R.sedulus, and R. luctus. Two subspecies are recognized. Subspecies and Distribution. R. b. beddomei K. Andersen, 1905 — W & E India (Maharashtra, Karnataka, Kerala, and Andhra Pradesh). R.b. sobrinus K Andersen, 1918 - Sri Lanka. Descriptive notes. Head-body 65-75 mm, tail 37-48 mm, ear 21-34 mm, hindfoot 12- 1-18 mm, forearm 54-9—64- 3 mm. Fur is long and woolly; dorsal and ventral pelage is dark brown or black (hairs with paler tips). Ears are blackish and of medium length. Noseleaf is dark brown or blackish, and has long and pointed or rounded lancet; connecting process is relatively low and broadly rounded; sella is broad and widely rounded off with large circular lateral lappets on either side of base; horseshoe is very wide (11 mm), covering muzzle, and has wide median emargination. Lower lip has one mental groove. Skull is robust (zygomatic breadth is larger than mastoid breadth); anterior median swellings are elongated but do not expand anteriorly to upper edge of nasal orifice, formed by a protruding bony rim; posterior swellings are more laterally situated; frontal depression is deep and long; supraorbital crests are sharp; sagittal crest is very well developed; canines are strong but relatively short. P2 is medium-sized and within tooth row or slightly displaced; P4 is small to medium-sized and extruded from the tooth row to a variable degree; P2 and P4 are almost in contact. Chromosomal complement has 2n = 32 and FN = 60. Habitat. Mainly found in dense dry and tropical moist forests. Elevational range is from sea level up to elevations of 1070 m. Food and Feeding. Beddome’s Woolly Horseshoe Bat feeds on a variety of insects especially beetles and termites. It flies low over the ground while foraging, and will forage close to vegetation. Breeding. A pregnant female with one embryo was collected in mid-January in Sri Lanka. Young have been reported with females in Kerala, India, in mid-May, and in Sri Lanka in mid-March and late April. Females may not breed until two or three years old. Activity patterns. Beddome’s Woolly Horseshoe Bat roosts by day in small caves, dilapidated buildings, tree hollows, wells, culverts, and old and unused tunnels, or clings from overhanging rocky outcrops. It emerges late in the evening from its roosts to forage through the night. Call shape is FM/CF/FM with a peak F recorded at 38-5—38-7 kHz and durations of 48-2—58 milliseconds in the Western Ghats of India; another study in the Western Ghats reported peak frequencies of 41-7-43-3 kHz (mean 42-8 kHz) and durations of 1-4-51-5 milliseconds (mean 25-2 milliseconds). A recording inside a cave in Sri Lanka produced peak frequencies of 41—62-5 kHz but this huge range must be an error, perhaps including more than one species. Movements, Home range and Social organization. Beddome’s Woolly Horseshoe Bat roosts alone or in pairs, or sometimes in groups of three or four individuals. Status and Conservation. Classified as Least Concern on TheIUCNRed List. Beddome’s Woolly Horseshoe Bat is relatively widespread and common throughout much of its distribution. However, in Sri Lanka the species is regionally listed as Vulnerable according to the National Red List for the country, and is mostly restricted to old-growth forests with small populations. The species appears to be threatened most by deforestation for logging and agricultural expansion. Bibliography. Bates & Harrison (1997), Csorba eta /. (2003), Edirisinghe et al. (2016), Koubfnovà et al. (2010), Kusuminda et al. (2013), Molur et al. (2002), Naidu & Gururaj (1984), Raghuram et al. (2014), Rao et al. (2004), Sinha (1973), Soisook, Niyomwan et al. (2010), Soisook, Struebig et al. (2015), Srinivasulu & Molur (2008b), Srinivasulu & Srinivasulu (2012),Topàl & Csorba (1992), Volleth et al. (2017), Wordley et al. (2014)., 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 330, 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.

Bats play crucial roles in ecosystems, are increasingly used as bio-indicators and are an important component of tropical diversity. Ecological studies and conservation-oriented monitoring of bats in the tropics benefit from published libraries of echolocation calls, which are not readily available for many tropical ecosystems. Here, we present the echolocation calls of 15 species from the Valparai plateau in the Anamalai Hills, southern Western Ghats of India: three rhinolophids (Rhinolophus beddomei, R. rouxii (indorouxii), R. lepidus), one hipposiderid (Hipposideros pomona), nine vespertilionids (Barbastella leucomelas darjelingensis, Hesperoptenus tickelli, Miniopterus fuliginosus, M. pusillus, Myotis horsfieldii, M. montivagus, Pipistrellus ceylonicus, Scotophilus heathii, S. kuhlii), one pteropodid (Rousettus leschenaultii) and one megadermatid (Megaderma spasma). Discriminant function analyses using leave-one-out cross validation classified bats producing calls with a strong constant frequency (CF) component with 100% success and bats producing frequency modulated (FM) calls with 90% success. For five species, we report their echolocation calls for the first time, and we present call frequencies for some species that differ from those published from other parts of the species' ranges. This exemplifies the need for more local call libraries from tropical regions to be collected and published in order to record endemic species and accurately identify species whose calls vary biogeographically.

R. beddomei ��� Sri Lanka : ABBM 439-05 (HZM 1.27943), 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 139, DOI: 10.3161/15081109ACC2016.18.1.005, http://zenodo.org/record/3756730

Shade coffee has shown great promise in providing crucial habitats for biodiversity outside formal protected areas. Insectivorous bats have been understudied in coffee, although they may provide pest control services. We investigated the influence of local and landscape-level features of coffee farms on aerial insectivorous bats in Chikmagalur district in the Western Ghats biodiversity hotspot, India. Bats were monitored in 20 farm sites using ultrasound detectors, and the response of bat species richness and activity to changes in tree density, proportion of built-up area in the neighborhood, and distance of farm from forest areas quantified. We examined if models built to explain the species richness and activity could also predict them in nine additional sites. We detected nine phonic types/species in the study area. The quantified predictors had no effect on assemblage-level species richness and activity of bats. Responses of edge-space and cluttered-space forager guilds mirrored those of the overall assemblage, but some species vulnerable to forest conversion like Rhinolophus beddomei were detected rarely. Best models explained up to 20% and 15% variation in assemblage-level species richness and activity respectively, and were poor predictors of both response variables. We conclude that coffee farms in our study area offer an important commuting space for insectivorous bats across a gradient of shade management. Further research should include species-specific responses to management decisions for at-risk species and quantification of ecosystem services like natural pest control to inform biodiversity conservation initiatives in the Western Ghats coffee landscapes.

The nominate subspecies of the lesser woolly horseshoe bat

1 Assistant Conservator of Forests (Biodiversity), 3 Field Director, Project Tiger, 4 Field Research Assistant, ERM Laboratory, Nagarjunasagar Srisailam Tiger Reserve, Sunnipenta, Kurnool District, Andhra Pradesh 518102, India. 2 (Corresponding author) Wildlife Biology Section, Department of Zoology, University College of Science, Osmania University, Hyderabad, Andhra Pradesh 500007, India. 2 Email: hyd2_masawa@sancharnet.in with web supplement