Rhinolophus philippinensis, Waterhouse, 1843

Burgin, Connor, 2019, Rhinolophidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 280-332 : 305

50 . Large-eared Horseshoe Bat

Rhinolophus philippinensis

French: Rhinolophe des Philippines I German: Philippinen-Hufeisennase / Spanish: Herradura de ilipinas

Other common names: Enormous-eared Horseshoe Bat; Greater Large-eared Horseshoe Bat (large morph), Lesser Large-eared Horseshoe Bat (small morph)

Taxonomy. Rhinolophus philippinensis Waterhouse, 1843 ,

Luzon , Philippines .

Included in the philippinensis species group along with R montanus and R achilles , both of which have generally been regarded as subspecies of R philippinensis . The philippinensis group may be best included within the megaphyllus species group, based on genetic data which show that if not, the megaphyllus group is paraphyletic with respect to R philippinensis .

This has led to considerable taxonomic confusion, as populations from Australia (both small and large forms discussed under R achiUes) are closely related to R megaphyllus , while the Sabah (and probably also the Sarawak) and New Guinea populations form their own clades. Populations in the Philippines and Lesser Sundas have not been genetically tested and neither has R montanus , so at present it is impossible to make firm taxonomic conclusions. The Sabah and New Guinea populations are included here but Australian and Kai Island populations are recognized as a distinct species, based on genetic and morphological data (see R achiUes). Proposed races alleni and sanbomi are considered synonyms of mams, based on morphological similarities. Throughout the distribution of R philippinensis , there are large and small forms, and even within populations there can be several size morphs, such as three in Sabah and Sulawesi (large, intermediate, and small). The three morphs in Sulawesi cluster together genetically in an mtDNA study presented by T. Kingston and S. J. Rossiter in 2004, despite having distinct call frequencies and markedly different forearm lengths. In the same study, Buton Island (offsouth-east Sulawesi) specimens clustered separately from Australian R achiUes, although the authors only included specimens from these two populations. Continued research is desperately needed within the species complex. Two subspecies are recognized.

Subspecies and Distribution.

R p. philippinensis Waterhouse, 1843 - Philippines (Luzon, Polillo, Mindoro , Samar , Negros , Bohol, Siquijor, and Mindanao Is ).

R p. maros Tate & Archbold, 1939 - N Borneo (Sabah and Sarawak), SW Sulawesi, Kabaena, Buton, Flores, and E Timor Is, and C Papua New Guinea ( Mt Karimui and Waro).

Descriptive notes. Head—body c. 56-61 mm, tail 29—37 mm, ear 30-38 mm, hindfoot 10-11 mm, forearm 47-58 mm; weight 6-7- 12 g. The Large-eared Horseshoe Bat has distinct size morphs throughout much of its distribution. In one study, three size morphs were found in sympatry on Buton Island, off Sulawesi, average forearm lengths being large (56- 1 mm), intermediate (50- 6 mm), and small (47 mm). Dorsal pelage is light cinnamon buff or dark brown, while ventral pelage is paler and more grayish; there is no orange morph. Ears are enormous and antitragus is well developed and marked off by deep notch. Noseleaf has prominent lancet with a rounded tip connecting process is low, has convex outline, and joins lancet near base; sella is very large, long, and tongue-shaped, being narrowest at base with longitudinal median depression extending nearly to top; intemarial cup is very broad and extends laterally over much of horseshoe’s width; horseshoe is exceptionally broad, covers and extends well past muzzle, has deep median emargination, and lateral leaflets. Skull is narrow and long (zygomatic width is much less than mastoid width); median nasal swellings are conspicuous and elliptical and protrude anteriorly well beyond rostral wall; lateral compartments are elongate; sagittal crest is very low to low; frontal depression is deep or moderately deep; supraorbital crests are moderately developed. P2 is small but within tooth row completely, separating C 1 and P4; tiny to small P3 is also within tooth row separating P2 and P4.

Habitat. Primary and secondary lowland and montane rainforest, riparian forest, open forest, and woodland, as well as disturbed forest in Sulawesi and Flores. Large-eared Horseshoe Bats are typically found near caves or other roosting sites but have been recorded in forest where no caves are known to exist. They have been recorded from sea level up to 1500 m, typically occurring at higher densities at lower elevations.

Food and Feeding. The Large-eared Horseshoe Bat is insectivorous, foraging among dense vegetation.

Breeding. Litter size is a single young.

Activity patterns. The Large-eared Horseshoe Bat is nocturnal, roosting during the day in warm, humid small caves and abandoned mine shafts; also large, deep caves in the Philippines. Call shape is FM/CF/FM with a peak F recorded at 31-32 kHz on Bohol Island, Philippines, 36-6 kHz in Sabah, Borneo, 27 kHz in East Timor, 27-2 kHz (large morph), 39 kHz (intermediate morph) and 53-6 kHz (small morph) on Buton Island, Indonesia.

Movements, Home range and Social organization. Large-eared Horseshoe Bats appear to roost in relatively small numbers. When roosting, individuals hang separately rather than in a cluster.

Status and Conservation. Classified as Least Concern on The IUCNed List. Although widespread, the Large-eared Horseshoe Bat is not particularly common throughout its range. It may be threatened by general habitat loss and fragmentation, as well as roost disturbance.

Bibliography. Armstrong & Konishi (2012), Bonaccorso (1998), Cooper eta/. (1998), Csorba et al. (2003), Duya et al. (2007), Flannery (1995a, 1995b), Heaney et al. (2016), Kingston & Rossiter (2004), Tu Vuong Tan, Hassanin et al. (2017), Zhang Lin et al. (2018).

Figures

On following pages: 52. Eastern Horseshoe Bat (Rhinolophus megaphyllus}; 53. Insular Horseshoe Bat (Rhinolophus keyensis}; 54. Yellow-faced Horseshoe Bat (Rhinolophus virgo}; 55. Madura Horseshoe Bat (Rhinolophus madurensis}; 56. Sulawesi Horseshoe Bat (Rhinolophus celebensis}; 57. Robinson's Horseshoe Bat (Rhinolophus robinsoni); 58. Indochinese Horseshoe Bat (Rhinolophus chasem); 59. Anamban Horseshoe Bat (Rhinolophus nereis}; 60. Bornean Horseshoe Bat (Rhinolophus borneensis); 61. Malayan Horseshoe Bat (Rhinolophus malayanus}; 62. Acuminate Horseshoe Bat (Rhinolophus acuminatus}; 63. Acuminate Horseshoe Bat (Rhinolophus acuminatus}; 64. Little Japanese Horseshoe Bat (Rhinolophus cornutus}; 65. Yaeyama Horseshoe Bat (Rhinolophus perditus}; 66. Mountain Horseshoe Bat (Rhinolophus monticolus}; 67. Shortridge's Horseshoe Bat (Rhinolophus shortridgei}; 68. Formosan Horseshoe Bat (Rhinolophus monoceros}; 69. Least Horseshoe Bat (Rhinolophus pusillus}; 70. Little Nepalese Horseshoe Bat (Rhinolophus subbadius}; 71. Convex Horseshoe Bat (Rhinolophus convexus}; 72. Andaman Horseshoe Bat (Rhinolophus cognatus}; 73. Glossy Horseshoe Bat (Rhinolophus refulgens}; 74. Blyth's Horseshoe Bat (Rhinolophus lepidus};

Subspecies and Distribution. R p. philippinensis Waterhouse, 1843 - Philippines (Luzon, Polillo, Mindoro , Samar , Negros , Bohol, Siquijor, and Mindanao Is ). R p. maros Tate & Archbold, 1939 - N Borneo (Sabah and Sarawak ), SW Sulawesi, Kabaena, Buton, Flores, and E Timor Is, and C Papua New Guinea ( Mt Karimui and Waro).