Rhinolophus affinis, Horsfield, 1823

Burgin, Connor, 2019, Rhinolophidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 280-332 : 320-321

86 . Intermediate Horseshoe Bat

Rhinolophus affinis

French: Rhinolophe de Horsfield I German: Mittelgrosse Asiatische Hufeisennase / Spanish: Herradura de Horsfield

Taxonomy. Rhinolophus affinis Horsfield, 1823 , Java, Indonesia .

Rhinolophus affinis is here included in the euryotis species group, based on genetic data, although this species and R andamanensis are typically included in the megaphyllus species group. Genetic data seems to indicate that R. affinis is related to at least R. creaghi , R.arcuatus , R.subrufus , R.sitameli , and R stheno , although studies including more species and multiple genes are needed to determine this species’ phylogenetic relationship. R andamanensis wa& recendy recognized by. Srinivasulu and colleagues in 2019 as a distinct species from R affnis based on genetic and morphological data. The proposed subspecies tener is here synonymized with macrurus, and superans is synonymized with the nominate, based on morphological and genetic data. Distributional borders in the three Chinese subspecies hainanus, macrurus, and himalayanus are uncertain, and macrums seems to have originated from a hybrid himalayanus x hainanus, which evolved on Hainan but later recolonized mainland China. Subspecies princeps is currendy the only recognized subspecies that has not been validated using genetic and morphological studies. Six subspecies recognized.

Subspecies and Distribution.

R. a. afinis Horsfield, 1823 - Malay Peninsula (including Langkawi and Tioman Is), Sumatra, North Pagai I in Mentawai Is, and Java.

R. a. hainanus J. A. Allen, 1906 - Hainan I, China.

R. a. himalayanus K Andersen, 1905 — N India (Uttarakhand, Uttar Pradesh, Sikkim, West Bengal, Assam, Meghalaya, Arunachal Pradesh, and Nagaland), Nepal, Bhutan, NE Bangladesh, N Myanmar, and C & S China (Sichuan, Yunnan, Shaanxi, Hubei, Hunan, and Guizhou).

R.a. macrurus K. Andersen, 1905 - SE China (Jiangsu, Anhui, Zhejiang,Jiangxi, Fujian, Guangdong, Hong Kong, and Guangxi), S Myanmar, Thailand, Laos, and Vietnam (including Dao Tra Ban and Phu Quoc Is); possibly also Cambodia.

R. a. nesites K.Andersen, 1905 - Borneo (including Laut, Sebuku, and Laut Kecil Is) and Anamba and Natuna Is.

R. a. princeps K. Andersen, 1905 — Kangean (Kangean and Sepanjang Is), Lombok, Sumbawa, Flores, and Sumba Is.

A morphologically distinct subspecies (still unnamed) is known from E Myanmar and N Vietnam.

Descriptive notes. Head-body 46-68 mm, tail 174- 35 mm, ear 14—25 mm, hindfoot 9- 8-13 mm, forearm 46-57 mm; weight 9- 9-19 g. Dorsal pelage varies from dark to lighter brown, occasionally with an ocherous buff tinge; ventral pelage ranges from brown to creamy buff. Ears are comparatively small. Noseleaf has parallel-sided lancet with pointed tip; connecting process is rounded and sparsely haired; sella is slighdy concave and pandurate in shape; horseshoe is relatively broad (8-11- 4 mm) but does not cover whole muzzle and has well-defined median emargination. Lower lip has three mental grooves. Baculum has deeply notched ventral margin of the basal cone but is less notched on dorsal margin; it has smaller notches on lateral sides; shaft is roughly circular in cross section, slightly bending toward ventral side (in himalayanus), or is considerably shorter and more bent (in macrurus). Skull is robust (but zygomatic width is smaller than mastoid width) with a moderately long rostrum; anterior median swellings are comparatively less inflated; posterior swellings are well defined; sagittal crest is moderately to strongly developed; frontal depression is moderately developed; supraorbital crests are conspicuous. P2 is large and only slightly displaced, separating C1 from P 4; Ps is small to very small and generally fully (rarely partially) displaced from tooth row; P 2 and P4 are either touching or almost in contact. Chromosomal complement has 2n = 62 and FNa = 60 ( China and Vietnam).

Habitat. The Intermediate Horseshoe Bat is found in a wide variety ofhabitats and is very adaptable; it occurs in both primary and secondary forests as well as disturbed habitats, orchards, and other agricultural areas. In China, it can be found both in the wet and more temperate western highlands and in the more tropical eastern lowlands. In Vietnam, the species has been reported from both primary and secondary tropical forests but appears to avoid heavily disturbed landscapes, although it has been observed flying into houses and under tents in campsites. Recorded at elevations of 200-2000 m.

Food and Feeding. Intermediate Horseshoe Bats forage by aerial-hawking and perchhunting. They are frequently observed flying along streams and roads (c. 1-5- 2 m aboveground). They feed on a variety of insects but seem to prefer Coleoptera and Lepidoptera . In Jiangxi, China, this species fed mainly on Coleoptera (61-4% by volume) and Lepidoptera (28-9%), and less on Hymenoptera , Orthoptera, Homoptera, Megaloptera, Neuroptera , Diptera, Hemiptera, and Trichoptera ; this was a much wider variety than in the sympatric Pearson’s Horseshoe Bat (.pearsoni), with which has an extensive trophic niche overlap.

Breeding. Females give birth to a single young. In Peninsular Malaysia, there appear to be two breeding seasons per year, with pregnant females being collected in April-May and in October. Lactating females have also been captured in June.

Activity patterns. Intermediate Horseshoe Bats emerge from their roosts at dusk to forage; by day they roost mainly in caves, although they may also use rock crevices or hollow trees. Calls are FM/CF/FM shaped with a peak F recorded at c.90 kHz in Vu Quang and c.78 kHz on the Langbian Plateau, Vietnam; and 71-3 kHz in Thailand (duration of 29-2 milliseconds and interpulse interval of 59-4 milliseconds). InJiangxi, China, the peak F of the first harmonic averaged 42-9 kHz and the second harmonic averaged 85-9 kHz, with a duration of 46-5 milliseconds and interpulse interval of 45-2 milliseconds.

Movements, Home range and Social organization. Intermediate Horseshoe Bats roost in large colonies with up to thousands of individuals.

Status and Conservation. Classified as Least Concern on The IUCN Red List. The Intermediate Horseshoe Bat is widespread and common throughout much of its distribution. Although there do not seem to be any major threats currently affecting this species, limestone extraction in some regions of South Asia may threaten roosting sites.

Bibliography. Bates & Harrison (1997), Bates, Thi Mar-Mar et al. (2004), rancis (2008a), Harada, Yenbutra, Yosida &Takada (1985), Ith, Bumrungsri, Furey et al. (2015), Ith, Bumrungsri, Thomas et al. (2016), Jiang Tinglei, Feng Jiang et al. (2008), Jiang Tinglei, Lu Guanjun et al. (2013), Kingsada et al. (2011), Kruskop (2013a), Maharadatunkamsi et al. (2000), Mao Xiuguang, NieWenhui et al. (2007), Mao Xiuguang, Zhu Guangjian, Zhang Libiao et al. (2014), Mao Xiuguang, Zhu Guangjin, Zhang Shuyi & Rossiter (2010), Niu Huiling etal. (2007), Phommexay (2009), Sia et al. (2015), Sinha (1973), Smith & XieYan (2008), Srinivasulu & Srinivasulu (2012), Srinivasulu et al. (2019), Stoffberg et al. (2010), Tingga et al. (2012), Walston et al. (2008a), Wu Yi & Harada (2005),YuYan et al. (2006), Zhang Weidao (1985), Zhou Zhaomin et al. (2005).

Figures

On following pages: 78. New Guinea Broad-eared Horseshoe Bat (Rhinolophus euryotis); 79. Sulawesi Broad-eared Horseshoe Bat (Rhinolophus tatah; 80. Poso Horseshoe Bat (Rhinolophus belligerator); 81. Bornean Woolly Horseshoe Bat (Rhinolophus proconsulis); 82. Arcuate Horseshoe Bat (Rhinolophus arcuatus); 83. Philippine Forest Horseshoe Bat (Rhinolophus inops); 84. Small Rufous Horseshoe Bat (Rhinolophus subrufus); 85. Large Rufous Horseshoe Bat (Rhinolophus rufus); 86. Intermediate Horseshoe Bat (Rhinolophus affinis); 87. Homfray's Horseshoe Bat (Rhinolophus andamanensis); 88. Croslet Horseshoe Bat (Rhinolophus coelophyllus); 89. Shamel's Horseshoe Bat (Rhinolophus shameli); 90. Vietnamese Brown Horseshoe Bat (Rhinolophus microglobosus); 91. Lesser Brown Horseshoe Bat (Rhinolophus stheno).

Subspecies and Distribution. R. a. afinis Horsfield, 1823 - Malay Peninsula (including Langkawi and Tioman Is), Sumatra, North Pagai I in Mentawai Is, and Java. R. a. hainanus J. A. Allen, 1906 - Hainan I, China . R. a. himalayanus K Andersen, 1905 — N India (Uttarakhand, Uttar Pradesh, Sikkim, West Bengal, Assam, Meghalaya, Arunachal Pradesh, and Nagaland), Nepal , Bhutan , NE Bangladesh , N Myanmar , and C & S China (Sichuan, Yunnan, Shaanxi, Hubei, Hunan, and Guizhou). R.a. macrurus K. Andersen, 1905 - SE China (Jiangsu, Anhui, Zhejiang,Jiangxi, Fujian, Guangdong, Hong Kong , and Guangxi), S Myanmar, Thailand , Laos , and Vietnam (including Dao Tra Ban and Phu Quoc Is); possibly also Cambodia . R. a. nesites K.Andersen, 1905 - Borneo (including Laut, Sebuku, and Laut Kecil Is) and Anamba and Natuna Is. R. a. princeps K. Andersen, 1905 — Kangean (Kangean and Sepanjang Is), Lombok, Sumbawa, Flores , and Sumba Is. A morphologically distinct subspecies (still unnamed) is known from E Myanmar and N Vietnam .