Cynopterus brachyotis (S. Müller, 1838)

Don E. Wilson & Russell A. Mittermeier, 2019, Pteropodidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 16-162 : 65-66

6.

Lesser Short-nosed Fruit Bat

Cynopterus brachyotis

French: Cynoptéere de Miller / German: Kleiner-Kurznasenflughund / Spanish: Cynéptero de Miller

Other common names: Common Short-nosed Fruit Bat, Lesser Dog-faced Fruit Bat, Sunda Short-nosed Fruit Bat

Taxonomy. Pachysoma brachyotis [sic] S. Miller, 1838 ,

Dewei (= Dewai) River, Borneo.

Taxonomy is challenging and in need of revision. Six, may be more, distinct mitochondrial lineages could be raised to species level. Cynopterus brachyotis is often confused with C. sphinx and other species with which it overlaps in many physical dimensions. Nominate subspecies has two sympatric morphs in Peninsular Malaysia: smaller “Forest” lineageis a forest specialist and larger “Sunda” lineage is found in more open and disturbed areas. These forms are probably distinct species. Records from Philippines and Sulawesi are now assigned to C. luzoniensis and C. minutus , respectively. Records of subspecies brachysoma might represent C. s. sphinx . Eight subspecies recognized.

Subspecies and Distribution.

C.b.brachyotisS.Miller,1838—SW&NEIndia,Bhutan,andBangladeshandmainlandSEAsiafromNMyanmartoSPeninsularMalaysia,Sumatra,andBorneo(Sundalineage).

C.b.altitudinisHill,1961—CameronHighlands,WPeninsularMalaysia.

C.b.brachysomaDobson,1871—AndamanIs.

C.b.ceylonensisJ.E.Gray,1870—C&SSriLanka.

C.b.concolorSody,1940—EngganoI.

C.b.hoffetiBourret,1944—Vietnam.SChinarecordsquestionable.

C.b.insularumK.Andersen,1910—KangeanIs.

C. b. javanicus K. Andersen, 1910 — Java, including Madura I, and Bali I.

Descriptive notes. Head-body 72-96 mm, tail 10-16 mm, ear 15-20 mm, hindfoot 11-15 mm, forearm 55-66 mm; weight 27-45 g (mean 33-1 g). The Lesser Shortnosed Fruit Bat is small, with white ear rims and wing digits. Muzzle is short, almost naked; skin is dark brown; nostrils are shortly tubular; philtrum ends in two upper lip pads; and two triangular pads occur on lowerlip. Forehead slopesslightly, top of head is rounded, and crown is distinctly brown in some specimens. Eyesare large;iris is chestnut-brown to olive-brown. Ears are moderately short, oval, attenuated at tip, and pale brown, edged with white. Head pelageis short and brown to orange/yellowish brown; pelage is longer on nape and dorsum, brown suffused with warm russet to tawny olive. Females are generally paler. Pelage reaches dorsal bases of forearms and center of uropatagium. Tail is short and dorsally attached to uropatagium; calcaris short. Ruff is intensely orange on males, more so on sides of neck and paler on chest; it is generally paler on females. Chest and belly are grayish brown, occasionally with orange or yellowish tinge. Wing membranes are dark gray and attach to first toe; index claw is present. Skull lacks basicranial deflection, rostrum is short, forehead is relatively low and sloping, orbit is large, and braincase is rounded. Zygomatic rootis only slightly above level of alveolar line, and zygoma is thin and gently arched posteriorly. Dorsally, paranasal recesses are inflated, reaching small postorbital foramina; postorbital processes are thin and directed posterolaterally. There is almost no postorbital constriction; braincase is almost perfectly rounded, with only traces of temporal lines; and nuchal crest is modest. Palate is flat, upper tooth row is almost parallel, end of post-dental palate is rounded, and mid-sphenoidalridge is low. Ectotympanic is small and wide, more so anteriorly; entotympanic is variously reduced. Mandible is straight, with sloping tall coronoid; condyle is slightly above alveolar line; and angle is rounded off. Upper incisors are small but rather long (I? longer); C' is almoststraight; anterior surface is convex; and P' is long and minute; posterior cheekteeth decrease in height, are rounded to elongate posteriorly, and have no additional cusps. Lower incisors are small; I, is shorter; C, is small and short; Pis low and peg-like; posterior premolars and molars decreasing in size, with oval outline and no additional cusps; and M,is peg-like. There are eight interdental palatal ridges, undivided, arched, and rather thick, followed by one thin medially divided ridge, spaced from two divided denticulate ridges on posterior one-half of post-dental palate. Chromosomal complement of subspecies Javanicus has 2n = 34 and FN = 58, with eleven pairs of variable-sized metacentric to submetacentric (pair with a secondary constriction), two pairs of subtelocentric, and three pairs of small to minute acrocentric chromosomes. X-chromosome is subtelocentric, and Y-chromosome is minute and acrocentric.

Habitat. Lowland to montane forests, including mangrove forests, in all degrees of modification, including deforested areas such urban environments near forests, rural areas, and orchards, from sea level up to elevations of ¢. 1600 m. Nominate brachyotis tends to diverge ecologically: Forest lineage is a forest specialist and Sunda lineage is found in secondary and disturbed forests and rural/agricultural areas.

Food and Feeding. The Lesser Short-nosed Fruit Bat is mainly frugivorous (95% of diet) but also eats leaves, pollen, and nectar mainly in dry season. It forages from ground level to subcanopy. Most fruits consumed are small (most less than 2 g but a few heavier than 8 g) and taken from early successional forests (two-thirds of diet) Juveniles concentrate on large-crop fruiting species and introduced steady-state fruiting plants toward adulthood. Lesser Short-nosed Fruit Bats feed at night roosts where they carry fruits and leaves; flower products are eaten on the plant. In Peninsular Malaysia, diet containsfruits of about 78 plant species from at least 24 families, leaves of 14 species, and nectar and pollen offive species (native plus exotic plants). Ficus fruits are heavily used (e.g. E fistulosa). Leaves from Fabaceae (e.g. Erythrina ) are eaten. Flowers include native Cassia , Peltophorum , Senna , and exotic Bauhinia (all Fabaceae ). In Borneo, fruits of at least 24 plant species in 16 genera and 15 families are eaten.

Breeding. Reproductive cycle of the Lesser Short-nosed Fruit Bat is seasonal bimodal polyestry, with postpartum estrus. Mating is polygynous. Females are sexually mature at 6-7 months of age and give birth to one young within theirfirst year and up to two offspring per year thereafter. Mating in male-defended roosts occurs from December onward, gestation lasts c¢.4 months,births occur in March-April and July-August, and lactation lasts ¢.2 months. Lactation occurs during onset and mid-rainy season when food availability is highest. Males are sexually mature at c.12 months of age but mate for the first time after they are two years old. Cycle of testis size is correlated with photoperiod and seasonality. Greater breeding success was reported for years after El Nino—Southern Oscillation (ENSO) events (e.g. 1999).

Activity patterns. The Lesser Short-nosed Fruit Bat becomes active shortly after sunset and moves directly to fruiting trees. Activity at feeding roosts had two peaks (19:00 20:00 h and 22:00-23:00 h), and activity declined as night progressed. Activity is reduced when intensity of moonlight is high, and individuals spend more time in the roost, especially during breeding season. Day roosts are located in dense foliage higher than 5 m aboveground in trees (e.g. Durio zibethinus, Malvaceae ) or bamboos (e.g. Gigantochloa scortechinii, Poaceae ). Many individuals make only transient use (1-5 days) of these roosts. Many ofthese roosts are tents constructed in foliage; males bite twigs and large leaves to create small (15-50 cm wide) spaces in vegetation that are occupied by females and defended as a harem territory by the male. The Lesser Short-nosed Fruit Bat builds stem tents using root masses of Phylodendron ( Araceae ) and Cymbidium (Orchidaceae) ; palmate umbrella tents are constructed by applying circular bites to base of palm fronds (e.g. Borassus , Cocos , Elaeis , Licuala , Sabal , and Roystonea , all Arecaceae ); conical tents are built by severing petioles of leaves in sapling stems (e.g. Dracaena fragrans, Asparagaceae ); and apical tents are made by chewing main veins of leaves (e.g. Scindapsus , Araceae ). Day roost tents are 3-10 m aboveground; tents used as feeding roost are 1-3 m aboveground. Adults spend c.18% of night actively defending day roost/tents.

Movements, Home range and Social organization. Each night, individuals commute 7-1-14-5 km between day roosts and feeding areas; both sexes commute double the distance during the breeding season. The Lesser Short-nosed Fruit Bat visits 1-6 feeding areas/night spending an average of one hour and 38 minutes in each feeding area during which individuals undertake 2-10 feeding bouts between fruiting trees and chosen night roosts where fruits are eaten. Total home ranges are 30-365 ha, marginally greater for females. Roost area is ¢.5% of total home range. Roost area is non-overlapping among males, which exhibit high roost and home range fidelity. Tents were used by a single solitary male or a harem group consisting of a defending male and 1-4 females with their young. The male roosts alone on the side of the tent, and harem females and their young cluster together. Tents can last up to years. Mating and lactation take place in tents. Activity of solitary vs. harem-holding malesis similar.

Status and Conservation. Classified as Least Concern on The IUCN Red List. The Lesser Short-nosed Fruit Bat faces no immediate conservation threats given its wide distribution, presumably large population, and environmental tolerance. It is considered vermin under Schedule V of the Indian Wildlife (Protection) Act. Resolution of exceedingly complex taxonomy of the C. brachyotis complex is required to correctly assess conservation status of these forms.

Bibliography. Ando et al. (1980a), Bumrungsri (2002), Bumrungsri et al. (2007), Campbell (2008), Campbell, Reid et al. (2006), Campbell, Schneider et al. (2006), Csorba, Bumrungsri, Francis, Bates, Gumal, Kingston, Molur & Srinivasulu (2008a), Fletcher et al. (2012), Francis (1994), Funakoshi & Zubaid (1997), Hodgkison (2001), Jones, Bielby et al. (2009), Kitchener & Maharadatunkamsi (1991), Magalhaes & Costa (2009), Mohd-Azlan et al. (2010), Tan et al. (1997, 1998, 2000), Tingga et al. (2012).

Figures

1. Temminck’s Tailless Fruit Bat (Megaerops ecaudatus), 2. Javan Tailless Fruit Bat (Megaerops kusnotoi), 3. Ratanaworabhan’s Tailless Fruit Bat (Megaerops niphanae), 4. White-collared Fruit Bat (Megaerops albicollis), 5. Greater Short-nosed Fruit Bat (Cynopterus sphinx), 6. Lesser Short-nosed Fruit Bat (Cynopterus brachyotis), 7. Horsfield’s Short-nosed Fruit Bat (Cynopterus horsfieldui), 8. Peters’s Short-nosed Fruit Bat (Cynoplerus luzoniensis), 9. Minute Short-nosed Fruit Bat (Cynopterus minutus), 10. Nusatenggara Short-nosed Fruit Bat (Cynopterus nusatenggara), 11. Indonesian Short-nosed Fruit Bat (Cynoplerus titthaecheilus), 12. Lesser Musky Fruit Bat (Ptenochirus minor), 13. Greater Musky Fruit Bat (Plenochirus jagorn), 14. Mindanao Fruit Bat (Ptenochirus wetmorei)

Subspecies and Distribution. C.b.brachyotisS.Miller,1838—SW&NEIndia,Bhutan,andBangladeshandmainlandSEAsiafromNMyanmartoSPeninsularMalaysia,Sumatra,andBorneo(Sundalineage). C.b.altitudinisHill,1961—CameronHighlands,WPeninsularMalaysia. C.b.brachysomaDobson,1871—AndamanIs. C.b.ceylonensisJ.E.Gray,1870—C&SSriLanka. C.b.concolorSody,1940—EngganoI. C.b.hoffetiBourret,1944—Vietnam.SChinarecordsquestionable. C.b.insularumK.Andersen,1910—KangeanIs. C. b. javanicus K. Andersen, 1910 — Java, including Madura I, and Bali I.