Year: 2006
Type: article
Abstract: Cynopterus horsfieldiiGray, 1843:38. Type locality “Java,” Indonesia. Cynopterus marginatus: Blyth, 1852:345. Part, not Pteropus marginatusE. Geoffroy, 1810:97. Pteropus [Pachysoma] marginatus: Wagner, 1853, 1855:609. Part, not Pteropus marginatusE. Geoffroy, 1810:97. Pachysoma horsfieldii: Fitzinger, 1869:33. Name combination. Cynopterus marginatus var. horsfieldii: Gray, 1870:123. Name combination. Cynopterus princepsMiller, 1906:61. Type locality “Mojeia River, Nias Island, west Sumatra.” Niadias minorLyon, 1908:665. Type locality “confluence of Gasip and Siak rivers, eastern Sumatra.” Preoccupied by Cynopterus (Cynonycteris) minorTrouessart, 1878:206. Cynopterus (Niadius) harpaxThomas and Wroughton, 1909:439. Type locality “Semangko Pass, Pahang boundary,” Selangor– Pahang boundary, peninsular Malaysia, 3,000 feet (Andersen 1912a:633). Cynopterus horsfieldiAnderson, 1912a:827. Name combination and unjustified emendation. Cynopterus persimilisAndersen, 1912b:641. Type locality “Sarawak, Borneo.” Order Chiroptera, suborder Megachiroptera, family Pteropodidae, subfamily Pteropodinae, tribe Cynopterini, subtribe Cynopterina, genus Cynopterus. Niadius was synonymized with Cynopterus and the combined genus was divided in 2 sections (Anderson 1912a): the Niadius section with C. harpax and C. horsfieldii, and the Cynopterus section with C. angulatus (= C. s. angulatus), C. brachyotis, C. sphinx, and C. s. titthaecheilus (= C. titthaecheilus). C. harpaxThomas and Wroughton, 1909 was synonomized with C. h. minor by Hill (1961), who considered the type specimen to be morphologically indistinguishable from the Sumatran subspecies. C. h. lyoniAndersen, 1912a was considered the valid name and C. h. minor a homonym (Hill 1983). However, Koopman (1994) treated C. h. lyoni as a junior synonym of C. h. harpax. Four subspecies are recognized (Corbet and Hill 1992; Koopman 1994; Mickleburgh et al. 1992): C. h. harpaxThomas and Wroughton, 1909:439; see above; lyoni (Anderson) and minor (Lyon) are synonyms. C. h. horsfieldii (Gray, 1843:38); see above. C. h. persimilisAndersen, 1912b:641; see above. C. h. princepsMiller, 1906:61; see above. Cynopterus horsfieldii (Fig. 1) is readily distinguished from Pteropus by its smaller size. Aethalops alecto, Balionycteris maculata, Chironax melanocephalus, and Macroglossus are all considerably smaller than C. horsfieldii. Rousettus has 5 cheek teeth in upper toothrow and is generally larger than C. horsfieldii (Lekagul and McNeely 1977; Payne et al. 1985). Among medium-sized members of Pteropodidae, C. horsfieldii can be differentiated as follows: Megaerops and Penthetor lucasi lack white ear margins and have 1 pair of lower incisors whereas C. horsfieldii has 2; Megaerops lacks a tail. Eonycteris has a longer, narrower rostrum than C. horsfieldii and lacks claw on 2nd digit. Dyacopterus spadiceus is slightly larger than C. horsfieldii, with a broader rostrum, deeper jaw, massive squared cheek teeth, and 2 upper premolars as compared to 3 in C. horsfieldii (Lekagul and McNeely 1977; Medway 1983; Payne et al. 1985). Photograph of an adult male Cynopterus horsfieldii from Krau Wildlife Reserve in peninsular Malaysia. Used with permission of the photographer, T. Kingston Photograph of an adult male Cynopterus horsfieldii from Krau Wildlife Reserve in peninsular Malaysia. Used with permission of the photographer, T. Kingston Cynopterus horsfieldii is distinguished from all other congeners by having peglike cusps on the 3rd and 4th lower cheek teeth (p3, m1—Corbet and Hill 1992; Lekagul and McNeely 1977; Medway 1983; Payne et al. 1985). In peninsular Malaysia, C. horsfieldii is considerably larger than C. brachyotis and averages larger than C. sphinx in all characters except length of rostrum. Relative to C. sphinx, C. horsfieldii has a broader cranium, blunter rostrum, and more-robust cheek teeth (Campbell et al., in press). Cynopterus horsfieldii is a medium-sized fruit bat with large eyes and simple ears that lack a tragus. Skull (Fig. 2) is robust with a short, broad rostrum, and nostrils are subtubular. C. horsfieldii has 4 cheek teeth in each upper toothrow, 1 pair of upper and lower canines, and 2 pairs of upper and lower incisors. Well-developed surface cusps are on the 3rd lower premolar and 1st lower molar (Andersen 1912a; Lekagul and McNeely 1977; Medway 1983; Payne et al. 1985). Dorsal, ventral, and lateral views of cranium and lateral view of mandible of an adult male Cynopterus horsfieldii from Taman Negara, peninsular Malaysia (Department of Wildlife and National Parks, Kuala Lumpur, Malaysia, catalogue no. NP042). Greatest length of skull is 34.1 mm. Used with permission of the photographer, S. W. Murray Dorsal, ventral, and lateral views of cranium and lateral view of mandible of an adult male Cynopterus horsfieldii from Taman Negara, peninsular Malaysia (Department of Wildlife and National Parks, Kuala Lumpur, Malaysia, catalogue no. NP042). Greatest length of skull is 34.1 mm. Used with permission of the photographer, S. W. Murray Pelage is short and dense (Andersen 1912a). Color in adults ranges from light or yellowish gray to dark or bright brown; juveniles are dull buff or gray. Breeding adults have a reddish brown or orange mantle (Lekagul and McNeely 1977; Medway 1983; Payne et al. 1985). Pelage around throat and shoulders of adult males is stiffer, brighter, and richer in color than that of females. In males, mantle may extend ventrally across chest (Andersen 1912a; Medway 1983; Payne et al. 1985). Color and area of mantle are the only consistently sexually dimorphic characters; males and females do not differ significantly in size (Campbell et al., in press). Margins of ears and membranes overlying metacarpals and phalanges are edged in white. Mean external measurements (± SD, in g or mm, sexes combined unless noted) for C. horsfieldii in peninsular Malaysia are: body mass, 57.9 (4.8, n = 31—Campbell et al., in press); 55.6 (8.9, n = 7—Hodgkison et al. 2004a); 56.7 (6.1, n = 19 males), 59.7 (6.9, n = 26 females—Funakoshi and Zubaid 1997); length of forearm, 73.2 (1.79, n = 27—Campbell et al., in press); 74.2 (1.7, n = 14 males), 74.7 (1.9, n = 24 females—Hodgkison 2001); 74.3 (3.1, n = 19 males), 75.1 (2.3, n = 26 females—Funakoshi and Zubaid 1997); and length of tibia, 27.0 (1.2, n = 27—Campbell et al., in press). Lengths of ear and tail (range, in mm) are: 19–22 and 12–16, respectively (n = 8—Lim et al. 1999). Length of forearm (range, in mm, sexes combined) for C. horsfieldii in Indonesia is: 64.5–71 (n = 15, Java), 71.5–77.5 (n = 3, Sumatra), and 87.0–89.5 (n = 2, Nias Island—Andersen 1912a). Mean cranial measurements (± SD, in mm, sexes combined) for C. horsfieldii in peninsular Malaysia are: total length of skull, 32.4 (0.76); maximum zygomatic breadth, 22.6 (0.85); maximum posterior skull width at mastoid, 14.4 (0.38); length of rostrum, 7.0 (0.32); length of palate, 17.0 (0.53); length from canine to M1, 10.7 (0.31); width of palate at M1, 9.7 (0.32); width of palate at P2, 8.6 (0.33); width of palate at canines, 6.9 (0.27); length of dentary, 24.8 (0.58); condylocanine length, 24.2 (0.59); length from condyle to p3, 18.7 (0.53); length from condyle to m1, 16.2 (0.48); length from condyle to m2, 13.5 (0.45); and height of coronoid process, 13.2 (0.51, n = 27—Campbell et al., in press). Cranial measurements (ranges or individual measurements, in mm, sexes combined) for C. horsfieldii in Indonesia are: total length of skull, 30.5–33.3 (n = 16, Java), 33.5–35.8 (n = 3, Sumatra), 38.5, 39.3 (n = 2, Nias Island); condylobasal length, 29.5– 31.5 (n = 16, Java), 31.8–34.0 (n = 3, Sumatra), 36.2, 37.2 (n = 2, Nias Island); length of rostrum (orbit to nares), 6.2–7.3 (n = 16, Java), 7.0–7.8 (n = 3, Sumatra), 8.7, 9.2 (n = 2, Nias Island); length from canine to M1, 10.5–11.5 (n = 16, Java), 11.0–12.0 (n = 3, Sumatra), 13.2, 13.5 (n = 2, Nias Island); total length of dentary, 23.0–25.3 (n = 16, Java), 25.5–26.8 (n = 3, Sumatra), 29.0, 30.5 (n = 2, Nias Island); length from canine to m2, 11.7– 12.4 (n = 16, Java), 12.4–13.3 (n = 3, Sumatra), 14.4, 14.8 (n = 2, Nias Island); and height of coronoid process, 12–13.2 (n = 16, Java), 13.7–14.5 (n = 3, Sumatra), 16.8 (n = 2, Nias Island— Andersen 1912a). Dental measurements (ranges or individual measurements, in mm, sexes combined) at crowns are: length and width of P3, 2.2– 2.7 and 1.7–2.0 (n = 15, Java), 2.5–2.6 and 1.8–2.0 (n = 3, Sumatra), 2.8, 3.0 and 2.2, 2.3 (n = 2, Nias Island); length and width of M1, 2.3–2.7 and 1.6–1.9 (n = 15, Java), 2.6–2.8 and 1.8– 1.9 (n = 3, Sumatra), 3.0, 3.1 and 2.0, 2.2 (n = 2, Nias Island); length and width of p1, 0.7–1.1 and 1.1–1.7 (n = 15, Java), 1.0– 1.2 and 1.7 (n = 3, Sumatra), 1.2 and 1.7, 1.8 (n = 2, Nias Island); length and width of p3 (= p2), 2.0–2.5 and 1.7–2.0 (n = 15, Java), 2.2–2.5 and 1.8–2.0 (n = 3, Sumatra), 2.5, 2.8 and 2.3 (n = 2, Nias Island); length and width of p4 (= p3), 2.2–2.7 and 1.8–2.0 (n = 15, Java), 2.5–2.7 and 1.8–2.0 (n = 3, Sumatra), 2.8, 3.0 and 2.3 (n = 2, Nias Island); length and width of m1, 2.5–2.8 and 1.7–2.0 (n = 15, Java), 2.7–2.8 and 1.8 (n = 3, Sumatra), 2.9, 3.0 and 2.1, 2.2 (n = 2, Nias Island); and length and width of m2, 1.6–1.9 and 1.0–1.6 (n = 15, Java), 1.6–1.8 and 1.2–1.4 (n = 3, Sumatra), 1.8, 2.0 and 1.5, 1.7 (n = 2, Nias Island—Andersen 1912a). Cynopterus horsfieldii (Fig. 3) occurs from ca. 9°N in southern Thailand, throughout peninsular Malaysia, on the Indonesian islands of Sumatra, Java, Bali, Lombok, and Sumbawa, and on the island of Borneo in Kalimantan (Indonesia), Sabah and Sarawak (Malaysia), and Brunei (Campbell et al. 2006b; Corbet and Hill 1992; Lekagul and McNeely 1977; Medway 1983; Mickleburgh et al. 1992; Schmitt et al. 1995; Simmons 2005). No recent records are available from Singapore (Teo and Rajathurai 1997). Geographic distribution of Cynopterus horsfieldii, redrawn from Corbet and Hill (1992) with modifications according to Campbell et al. (in press) and Kitchener et al. (1990): 1, C. h. harpax, southern Thailand, peninsular Malaysia, and Sumatra; 2, C. h. horsfieldii, Java to Sumbawa; 3, C. h. persimilis, Borneo; 4, C. h. princeps, Nias Island. Inset map localizes the Malay peninsula and western Indonesia (black) in Asia Geographic distribution of Cynopterus horsfieldii, redrawn from Corbet and Hill (1992) with modifications according to Campbell et al. (in press) and Kitchener et al. (1990): 1, C. h. harpax, southern Thailand, peninsular Malaysia, and Sumatra; 2, C. h. horsfieldii, Java to Sumbawa; 3, C. h. persimilis, Borneo; 4, C. h. princeps, Nias Island. Inset map localizes the Malay peninsula and western Indonesia (black) in Asia Subspecies of C. horsfieldii have the following distributions: C. h. harpax, southern Thailand, peninsular Malaysia, and Sumatra; C. h. horsfieldii, Java to Sumbawa; C. h. persimilis, Borneo; C. h. princeps, Nias Island, Indonesia (Corbet and Hill 1992; Hill 1983; Koopman 1994; Lekagul and McNeely 1977; Medway 1983). No fossils are known. Dental formula of C. horsfieldii is i 2/2, c 1/1, p 3/3, m 1/2, total 30 (Andersen 1912a). Premolars and molars are broad and subrectangular, with well defined surface cusps on p3 and m1 (Andersen 1912a; Koopman 1994). Cynopterus horsfieldii has relatively short, broad wings (low aspect ratio) and a relatively high ratio of body mass to wing area (high wing loading—Campbell et al., in press; Hodgkison et al. 2004a; McKenzie et al. 1995). This unspecialized wing morphology indicates that C. horsfieldii is capable of slow flight in moderately cluttered airspace, but is ill-suited to foraging over long distances or maneuvering in dense vegetation (Campbell et al., in press; Norberg and Rayner 1987). Mean brain mass of C. horsfieldii is 1,370 mg (n = 3, sexes combined—Bhatnagar et al. 1990). Pineal gland is deeply recessed and covered by cerebral hemispheres. Pineal volume is 0.4853 mm3 and greatest width of pineal gland is 1,010 μm (Bhatnagar et al. 1986, 1990). In peninsular Malaysia, C. horsfieldii is seasonally polyestrous and reproductively asynchronous, with pregnant females recorded throughout the year (Medway 1983). Two loosely synchronized peaks in birth occur per annum; during a 30-month study, peak numbers of lactating females were observed in February–March and July–August (Funakoshi and Zubaid 1997). Concentrations (mg/g dry mass) of minerals and nitrogen in milk of C. horsfieldii are: iron, 0.173; calcium, 6.11; magnesium, 0.17; potassium, 7.9; sodium, 5.37; and nitrogen, 28.03 (n = 1— Studier et al. 1995). With the exception of sodium, concentrations of minerals in milk of C. horsfieldii are lower than those in milk of 2 other Old World fruit bats, D. spadiceus and Megaerops ecaudatus (Studier et al. 1995). Based on mark–recapture data from peninsular Malaysia, average longevity for C. horsfieldii is at least 31 months (Hodgkison 2001). Cynopterus horsfieldii is a habitat generalist, occurring in diverse lowland habitats, from primary rain forest to suburban parks (Hodgkison et al. 2004a; Lim 1966; Tan et al. 1999). Although relatively common, C. horsfieldii is typically less abundant than sympatric congeners, particularly C. sphinx and C. brachyotis (Campbell et al., in press; Francis 1994; Hodgkison et al. 2004b; Lim 1966; Zubaid 1993, 1994). In peninsular Malaysia, C. horsfieldii is captured in both primary and disturbed lowland dipteropcarp forest (Campbell et al., in press; Fletcher 2001; Francis 1994, 1995; Funakoshi and Zubaid 1997; Hodgkison et al. 2004a; Zubaid 1993), limestone hill forest (Campbell et al., in press), fruit orchards (Campbell et al., in press; Lim 1966; Lim et al. 1999), and suburban gardens and parks (Campbell et al., in press; Tan et al. 1999). On the Indonesian island of Lombok, C. horsfieldii occurs in agricultural areas and secondary and primary rain forest (Kitchener et al. 1990). Comparison of the relative abundance of Cynopterus species across adjacent habitat types in northern peninsula Malaysia indicates that estimated overlap in habitat use between C. horsfieldii and C. sphinx is relatively high (71.2%). However, C. horsfieldii is strongly associated with transitional habitat between forest and cultivated land, whereas C. sphinx is most abundant in highly disturbed habitats and suburban areas where C. horsfieldii is less common (Campbell et al., in press). Within the forest, C. horsfieldii is captured in both understory and subcanopy (Francis 1994; Zubaid 1994), usually below 20 m (Hodgkison et al. 2004a), with a significant negative association between capture rate and density of vegetation (Hodgkison 2001). In lowland rain forest in central peninsular Malaysia, C. horsfieldii is a transient member of the fruit bat assemblage. Aseasonal shifts in abundance of C. horsfieldii are associated with exploitation of “big bang” food resources such as fruit of strangler figs (Ficus), Elaeocarpus stipularils, and Payena lucida, and flowers of Parkia speciosa (Hodgkison et al. 2004b). Data from the same site indicate that C. horsfieldii relies year-round on food resources in gardens and secondary habitats near the forest edge (Hodgkison 2001). On Lombok Island, 11 pollen types were identified from stomachs and alimentary tracts of C. horsfieldii (X̄ ± SD, 2.5 ± 1.12 pollen types/individual, n = 16) from various habitats (Kitchener et al. 1990). The pollen dietary niche breadth of C. horsfieldii was low relative to predominantly nectarivorous species such as Eonycteris spelaea and Macroglossus minimus and was intermediate to that of C. brachyotis (= C. nusatenggara, sensu Kitchener and Maharadatunkamsi 1991) and C. titthaecheilus. Pollen intake by C. horsfieldii was strongly influenced by the temporal and spatial availability of fruit: larger quantities of pollen were ingested during the dry season when fruit was less abundant and pollen dietary niche breadth was highest at sites where fruit was scarce. Across seasons and sites, C. horsfieldii exhibited high pollen dietary niche overlap with C. brachyotis and C. titthaecheilus (Kitchener et al. 1990). When feeding in fruit trees, C. horsfieldii removes single fruits from parent tree to nearby day or feeding roosts (Funakoshi and Zubaid 1997; Hodgkison 2001). Germination rates for Ficus seeds ingested by C. horsfieldii are ca. twice that of controls (Fletcher 2001). In secondary forest in peninsular Malaysia, mean home range sizes of adult C. horsfieldii were 5.8 ha ± 2.5 SD (n = 4 females) and 8.0 ha (n = 2 males—Funakoshi and Zubaid 1997). However, at another peninsular Malaysian site, individuals of C. horsfieldii that were captured while foraging in fruit orchards commuted up to 1.6 km to roosts in adjacent forest fragments (Fletcher 2001). Cynopterus horsfieldii roosts in small groups in foliage, rock shelters, and limestone solution cavities on open cliff faces or near the mouths of caves (Campbell et al. 2006a; Funakoshi and Zubaid 1997; Lim 1966; Medway 1983; Payne et al. 1985; Tan et al. 1999). C. horsfieldii modifies foliage roosts in an opportunistic manner; individuals switch roosts frequently and do not exhibit detectable preferences for modified or unmodified roosts (Campbell et al. 2006a; Funakoshi and Zubaid 1997; Tan et al. 1999). In a comparative study of the roosting ecology of Cynopterus species in northern peninsular Malaysia, estimated overlap in roost plant choice between C. horsfieldii and sympatric congeners ranged from 0.00% to 6.73% (Campbell et al. 2006a). All roosts of C. horsfieldii were located either in ecotonal habitat between cultivated land and secondary forest or <0.25 km inside the forest (Campbell et al. 2006a). Seventy-three percent (n = 35) of roosts of this species were in oblong leaves of wild and cultivated bananas (Musa). Ca. half of banana leaf roosts occupied by C. horsfieldii were modified into an inverted V-shaped structure by partial severing of midrib of leaf midway between base and tip. Height of roosts (X̄ ± SD, in m) used by C. horsfieldii was significantly lower than that of roosts occupied by C. sphinx and C. brachyotis (2.94 ± 0.98, n = 26—Campbell et al. 2006a). Other plant species used as roosts by C. horsfieldii in peninsular Malaysia include palms in the genera Arenga, Cocos, Corypha, and Nypa and the epiphytic bird's nest fern Asplenium nidas (Campbell et al. 2006a; Funakoshi and Zubaid 1997; Lim 1966; Tan et al. 1999). Mating system of C. horsfieldii is polygynous with a harem-based social structure that is maintained year-round (Campbell et al. 2006a; Tan et al. 1999). Harem groups contain 1 adult male and up to 5 adult females and their offspring (Campbell et al. 2006a; Tan et al. 1999). Individual composition of harem groups is relatively labile and females may periodically roost singly without attendant males (Campbell et al. 2006a). Harem groups are found in both modified and unmodified roosts (Campbell et al. 2006a; Tan et al. 1999). Cynopterus horsfieldii has = and = et al. of 11 pairs of or 2 pairs of and 3 pairs of is a medium-sized and is a small of C. horsfieldii is to those of C. s. titthaecheilus (= C. C. brachyotis, and C. sphinx et al. et al. et al. Mean in C. horsfieldii from Java, Bali, Lombok, and was compared with for C. sphinx, for C. brachyotis, and for C. and C. titthaecheilus. C. horsfieldii between and et al. 1995). In a for C. is to C. horsfieldii et al. in the Cynopterus The of C. horsfieldii is the Malaysian C. brachyotis (= C. brachyotis between the 2 is base partial and et al. and of all for C. horsfieldii at occur in C. brachyotis (Campbell et al. of C. horsfieldii in peninsular Malaysia and southern are not (Campbell et al. Cynopterus is from the and wing Cynopterus horsfieldii was in of an and who of and while in Java for the 1990). The type specimen of C. horsfieldii was those to the by on of the for C. horsfieldii include the larger fruit bat (Medway and the fruit bat (Lekagul and McNeely This is of the
Source: Mammalian Species
DOI: 10.1644/802.1
Authors: Polly Campbell, Thomas H. Kunz
Language: English
Domain: Life Sciences
Open Access status: bronze
OpenAlex ID: W4246419875