Rhinolophus nippon, Temminck, 1835

Burgin, Connor, 2019, Rhinolophidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 280-332 : 297-298

33 . GreaterJapanese Horseshoe Bat

Rhinolophus nippon

French: Rhinolophe nippon / German: Grosse Japan-Hufeisennase / Spanish: Herradura de Japón

Other common names: Japanese Greater Horseshoe Bat

Taxonomy. Rhinolophus nippon Temminck, 1835 ,

Japan .

Rhinolophus nippon is in the Jèmmeçwtnutn species group and sister to a clade including A clivosus and R ferrumequinum . Rhinolophus nippon m previously included in R ferrumequinum , but genetic data have shown that the two are distinct species. Distributional limits between R nippon and R ferrumequinum are still uncertain because there have been no genetic assessments of specimens from central and southern Asia. Which species tragatus is best included under has yet to be determined, but it is included as a subspecies here. Taxon korai is here synonymized under nominate nippon, although genetic and morphometric testing is needed to support this view. Two subspecies recognized.

Subspecies and Distribution.

R n. nippon Temminck, 1835 — C & E China (from Jilin SW to Sichuan and S to N Guangxi, Hunan, Jiangxi, and Fujian), Korean Peninsula, andJapan (including some offshore islands).

R n. tragatus Hodgson , 1835 — N India (Himachal Pradesh, Uttarakhand, West Bengal, Sikkim, Arunachal Pradesh, and Nagaland), Nepal, Bhutan, N Bangladesh, and SW China (Yunnan and Guizhou).

Descriptive notes. Head—body 55—79 mm, tail 25-44 mm, ear 18—29 mm, hindfoot 10—14 mm, forearm 49—64 mm. The GreaterJapanese Horseshoe Bat is similar to the Greater Horseshoe Bat ( A ferrumequinum ), but it seems to have somewhat darker pelage. Nominate subspecies has forearm lengths of 49—59 mm and tibia lengths of less than 25 mm; tragatus has forearm lengths of 58—64 mm and tibia lengths of more than 25 mm, being significantly larger than nippon with little overlap. Dorsal pelage is smoky gray-brown, pale brown, or chestnut-brown, with variable levels of red tinge (some individuals apparently look dark orange-brown); venter is pale buff. There are possibly orange-morph individuals in Japan . Males lack axillary tufts. Ears are relatively short (c.46% of forearm length on average). Noseleaf has subtriangular lancet, becoming slightly concave near bluntly pointed tip; connecting process is rounded and much higher than sella tip; sella is naked, relatively small, and curved forward, making front surface strongly concave while sides are only slightly concave and tip is pointed; and horseshoe is narrow, does not cover muzzle, and has lateral leaflets (although sometimes inconspicuous) and deep median emargination. Lower lip has one or three grooves, although lateral grooves can be inconspicuous if present. Wings and uropatagium are grayish brown. Detailed morphological comparisons between the GreaterJapanese Horseshoe Bat and the Greater Horseshoe Bat are required to differentiate the two species based on skull differences. Chromosomal complement has 2n = 58 and FN = 62.

Habitat. Variety of temperate forested habitats in montane and lowland regions from sea level to elevations of c. 3500 m.

Food and Feeding. Greater Japanese Horseshoe Bats forage by fly-catching from a perch (most often), slow hawking, and gleaning prey from the ground and vegetation. They forage in open forests, woodland paths, and forest edges, generally preferring open areas to cluttered areas. They generally prey on species of Diptera , Lepidoptera, Coleoptera, Trichoptera, Plecoptera, Odonata, and Hemiptera , but Lepidoptera , o ­ leoptera, and Diptera make up the largest proportion of diets. During winter hibernation, they are known to awake and feed almost exclusively on troglophilic moths in hibernacula and occasionally outside when it is warm enough.

Breeding. GreaterJapanese Horseshoe Bats are seasonally monoestrous, with delayed fertilization. Copulation occurs in late summer and early autumn before hibernation, and sperm is stored in females’ reproductive tracts. Births of single young occur synchronously in early summer. Mothers recognize their young by vocal communication, and their supersonic calls are synchronized. Young stay attached to their mothers’ underside and suckle. They begin to forage by themselves at c.32 days old, and weaning occurs at 40 days old. Juvenile mortality has been reported at 3-6%. Female Greater Japanese Horseshoe Bats generally take 2 - 4 years before rearing their first offspring, with fertility increasing with each year of life: one year (13 -1% fertility), two years (49 * 5 %), three years (95 - 2 %), and four years (100 %) in Ishikawa Prefecture and one (0-0%), two (27-2%), and three (93-4%) in Yamaguchi Prefecture. Females less than a year old generally do not produce any offspring, but it is possible. Females possess a strong loyalty to their natal sites for giving birth and raising their young. GreaterJapanese Horseshoe Bats are very long-lived; the oldest individual recorded was a 23-yearold female fromJapan.

Activity patterns. GreaterJapanese Horseshoe Bats are nocturnal and forage throughout the night. During the day, they can enter torpor, and during winter, they hibernate in more temperate parts of their distribution. They might not hibernate in southern parts of the distribution (e.g. India). Most roosts are in caves and roofs of abandoned or unused buildings. Call shape is FM /CF/FM, with F component of 65-69-8 kHz throughoutJapan. In China, resting frequencies ( RF) vary considerably (c.68—76 kHz) and are correlated with geographical location and mean annual temperature (higher temperatures are correlated with larger RF values).

Movements, Home range and Social organization. Greater Japanese Horseshoe Bats roost alone, in small groups, or in large colonies. Colonies can have hundreds of individuals. During mating season, females form tightly knit maternity colonies that average 85 individuals on central Honshu and 132 individuals (range 10-200) in western Honshu. Non-breeding and male individuals also can be found in these colonies, albeit in low numbers because they generally create their own roosts separate from maternity colonies. One maternity roost in Ishikawa Prefecture had 75-2% adult females, 24-2% subadult females, and 0-6% males. During hibernation, Greater Japanese Horseshoe Bats form mixed roosts with dense clusters of individuals. Weights increase 25-8-28-2% in late autumn before hibernation on Kyushu, Japan. Individuals will travel fairly long distances between summer roosts and hibernacula; distances of up to 130 km have been reported on Kyushu. Home ranges of adult females in summer averaged 1-5 ha in one study.

Status and Conservation. Not assessed on TAe IUCN ed List. The Greater Japanese Horseshoe Bat was previously included in R ferrumequinum , which is classified as Least Concern. It has a wide distribution and is considered common throughout much of its distribution. It is probably not threatened overall but might be locally threatened by habitat destruction from logging and agricultural expansion and roost disturbance from cave tourism and human cohabitation in building roosts.

Bibliography. Benda &Vallo (2012), Funakoshi & Maeda (2003), Koh Hung-Sun eta/. (2014), Matsumura (1979, 1981), Matsuta eta /. (2013), Mori eta/. (1982), OhYung-Keun eta/. (1983, 1985), Ohdachi eta/. (2009), Sano (2000a, 2000b), Smith & XieYan (2008), Stoffberg eta/. (2010), Sun Keping eta/. (2013),Taniguchi (1985).

Figures

On following pages: 27. African Forest Horseshoe Bat (Rhinolophus silvestris); 28. Upland Horseshoe Bat (Rhinolophus hillorum); 29. Sakeji Horseshoe Bat (Rhinolophus sakejiensis); 30. Bokhara Horseshoe Bat (Rhinolophus bocharicus); 31. Greater Horseshoe Bat (Rhinolophus ferrumequinum); 32. Geoffrey's Horseshoe Bat (Rhinolophus clivosus); 33. Greater Japanese Horseshoe Bat (Rhinolophus nippon); 34. Horacek's Horseshoe Bat (Rhinolophus horaceki); 35. Maclaud's Horseshoe Bat (Rhinolophus maclaudi); 36. Ziama Horseshoe Bat (Rhinolophus ziama); 37. Hill's Horseshoe Bat (Rhinolophus hilli); 38. Kahuzi Horseshoe Bat (Rhinolophus kahuzi); 39. Willard's Horseshoe Bat (Rhinolophus willardi); 40. Ruwenzori Horseshoe Bat (Rhinolophus ruwenzorii); 41. Chinese Horseshoe Bat (Rhinolophus xinanzhongguoensis); 42. Big-eared Horseshoe Bat (Rhinolophus macrotis); 43. Osgood's Horseshoe Bat (Rhinolophus osgoodi); 44. Allen's Horseshoe Bat (Rhinolophus episcopus); 45. Thai Horseshoe Bat (Rhinolophus siamensis); 46, Schnitzler's Horseshoe Bat (Rhinolophus schnitzten); M. King Horseshoe Bat (Rhinolophus rex); 48. Marshall’s Horseshoe Bat (Rhinolophus marshalli).

Subspecies and Distribution. R n. nippon Temminck, 1835 — C & E China (from Jilin SW to Sichuan and S to N Guangxi, Hunan, Jiangxi, and Fujian), Korean Peninsula, andJapan (including some offshore islands). R n. tragatus Hodgson , 1835 — N India (Himachal Pradesh, Uttarakhand, West Bengal, Sikkim, Arunachal Pradesh, and Nagaland), Nepal , Bhutan , N Bangladesh , and SW China (Yunnan and Guizhou).