Cynopterus horsfieldiiHorsfield's fruit bat

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Geographic Range

Horsfield's fruit bats (Cynopterus horsfieldii) are found throughout peninsular Malaysia, Cambodia, southern Thailand and several islands in the Malay Archipelago including the Greater Sunda Islands (Sumatra, Java and Borneo) and the Lesser Sunda Islands (Lombok and Sumbawa). (Campbell and Kunz, 2006; Campbell, et al., 2006; Campbell, et al., 2007; Campbell, 2008; Funakoshi and Zubaid, 1997; Hodgkinson, et al., 2004; Lekagul and McNeely, 1977; Shmitt, et al., 1995; Tan, et al., 1999; Zubaid, et al., 2007)

Habitat

Horsfield's fruit bats can be found in several differing habitats ranging from primary and secondary forests, rainforests, agricultural lands, parks, mangroves, limestone caves and gardens. They are commonly found within ecotonal areas, the transitional areas between forests and cultivated land. A study in Malaysia looked at the vertical stratification of fruit bats based on wing morphology, Horsfield's fruit bats were commonly captured in mist nets from 1 to 28 m. Researchers in this study predicted that Horsfield's fruit bats are more prevalent in the mid-storey airspace, which is classified as 10 to 20 m from the forest floor. A comparative study of roosting ecology found that Horsfield's fruit bats have the second highest roosting diversity of their genus. Their roosting sites are less than 5 m above ground and within 0.25 km of the forest. These bats are commonly found utilizing both unmodified and modified roosting sites and show no preference between the two, with harems found in both. A study examining coexistence in Malaysian fruit bats found Horsfield's fruit bats have a 71.2% habitat overlap with greater short-nosed fruit bats and 35.7% habitat overlap with lesser short-nosed fruit bats. (Campbell and Kunz, 2006; Campbell, et al., 2006; Campbell, et al., 2007; Campbell, 2008; Funakoshi and Zubaid, 1997; Hodgkinson, et al., 2004; Tan, et al., 1999; Vijaya, et al., 2012; Zubaid, et al., 2007)

Physical Description

Horsfield’s fruit bats, also known as larger dog-faced fruit bats or peg-toothed short-nosed fruit bats, are moderately sized, with an average mass of 57.9 g. Their larger size distinguishes them from lesser short-nosed fruit bats and greater short-nosed fruit bats. Another distinguishing characteristic of this species is the presence of peg-like cusps in their lower 3rd premolar and their 1st molar. Horsfield's fruit bats are described as having the heaviest dentition in their genus; they have 30 teeth, with a dental formula of 2/2, 1/1, 3/3, 1/2. These bats have a muzzle that is both short and broad, with tubular nostrils. They have relatively large eyes and oval shaped ears that measure 17 to 22 mm and lack a tragus. Their ears, as well as their wings, have a distinctive white colored margin. Horsfield's fruit bats also have a short tail, with lengths ranging from 7 to 16 mm. Their wings have a low aspect (6.08) and high wing loading (15.98 Nm-2) ratio. Their wingspan is 465 mm and their wing area is 36,000 mm2. Their pelage is short and varies in color from dark or light, gray or brown. A study of Malaysian pteropodid hair morphology described Horsfield's fruits bats' hair as having entire coronal scales and a discontinuous medulla. Juveniles are gray, while breeding adults have an amber colored mantle, which is more distinguishable in males. Males also tend to have darker fur on the sides of their neck and chest and brighter fur on their shoulder and throat. In terms of their size, male forearms measure about 74.3 mm and female forearms measure about 75.1 mm. Their head and body length may range from 80 to 96 mm. (Campbell and Kunz, 2006; Campbell, et al., 2006; Campbell, et al., 2007; Campbell, 2008; Funakoshi and Zubaid, 1997; Hodgkinson, et al., 2004; Lekagul and McNeely, 1977; Vijaya, et al., 2012; Zubaid and Fatimah, 1990; Zubaid, et al., 2007)

  • Sexual Dimorphism
  • sexes colored or patterned differently
  • male more colorful
  • Average mass
    57.9 g
    2.04 oz
  • Average length
    80-96 mm
    in
  • Average wingspan
    465 mm
    18.31 in

Reproduction

Horsfield’s fruit bats exhibit polygyny with a harem-based social structure. Roosts typically consist of no more than one mature male, two to three mature females and their associated offspring, this is a relatively small harem compared to other members of genus Cynopterus. Adult males tend to roost separately from females, which tend to stay closer in proximity to each other. Although the sexes separate while roosting, there is no evidence of bachelor groups. In a study exploring the relationship between roosting ecology and the degree of polygyny, Horsfield's fruit bats, along with other members of Cynopterus, did not exhibit defense of females or food resources. In terms of their roosting sites, there was little evidence of defense by males. (Campbell and Kunz, 2006; Campbell, et al., 2006; Campbell, et al., 2007; Campbell, 2008; Funakoshi and Zubaid, 1997; Neuweiler, 2000)

Horsfield’s fruit bats are seasonally polyestrous and breed asynchronously, with pregnancy peaks occurring every 4 to 6 months. Reproductive females exhibit postpartum estrous and delayed embryonic development. There is limited data available specifically for this species, however, if they exhibit similar reproductive traits to greater and lesser short-nosed fruit bats their gestation period likely ranges from 4 to 6 months, males likely reach sexual maturity in 1 to 2 years, females likely reach sexual maturity in 6 to 8 months and weaning likely occurs after 40 days. Similar to greater short-nosed fruit bats, the average birth mass of Horsfield's fruit bats may be near 11 grams; this assumption is based on their comparable adult body masses. ("AnAge entry for Cynopterus brachyotis", 2009; "AnAge entry for Cynopterus sphinx", 2009; Campbell and Kunz, 2006; Crichton and Krutzsch, 2000; Mickleburgh, et al., 1992)

  • Key Reproductive Features
  • gonochoric/gonochoristic/dioecious (sexes separate)
  • sexual
  • viviparous
  • post-partum estrous
  • Breeding interval
    Asychronous
  • Breeding season
    Seasonally Polyestrous
  • Average number of offspring
    2
  • Average gestation period
    4-6 months
  • Range weaning age
    40 (low) days
  • Average age at sexual or reproductive maturity (female)
    6-8 months
  • Average age at sexual or reproductive maturity (male)
    1-2 years

There is limited species specific data for parental investment in Horsfield’s fruit bats. Most newborn bats are altricial at birth and in general, females provision and protect their young until independence. An important aspect of parental care during this period is the maintenance of the thermal environment. Female bats commonly carry their young while foraging until they are able to fly, which drastically increases their energy expenditure and reduces their foraging efficiency. (Crichton and Krutzsch, 2000; Mickleburgh, et al., 1992; Neuweiler, 2000)

  • Parental Investment
  • altricial
  • female parental care
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

There is limited data available on the longevity of this species, but a mark-recapture study of Horsfield’s fruit bats estimated an average lifespan of at least 31 months. (Campbell and Kunz, 2006)

  • Typical lifespan
    Status: wild
    31 (low) months

Behavior

Horsfield’s fruit bats are a gregarious species with a harem-based social structure. Roosts typically consist of no more than one mature harem male, two to three mature females and their associated offspring. Roost fidelity in this species is relatively low, with males and females frequently changing roosts. Roost modification is present in Horsfield’s fruit bats, with modified roosts described as inverted v-shape tents formed from severing a leaf halfway between its base and tip. A study examining the behavioral and reproductive ecology of Horsfield’s fruit bats found that they were most active 2 to 4 hours after sunset and 3 hours prior to sunrise. (Campbell and Kunz, 2006; Campbell, et al., 2006; Campbell, et al., 2007; Campbell, 2008; Funakoshi and Zubaid, 1997; Zubaid, et al., 2007)

Home Range

A study located in a secondary rainforest in Malaysia reported the average home range size of Horsfield’s fruit bats was 8.0 ha (n=2) in males and 5.8 ± 2.5 ha (n=4) in females. Their mean greatest distance traveled in this study was 475 ± 105 m. (Funakoshi and Zubaid, 1997)

Communication and Perception

Horsfield’s fruit bats rely on their eyesight to navigate through the forest at night, along with their sense of smell to detect fruit. (Crichton and Krutzsch, 2000; Mickleburgh, et al., 1992)

Food Habits

Horsfield’s fruit bats are predominately frugivorous, but also consume leaves, flowers and pollen. A study in peninsular Malaysia showed these bats feed on the following plant species: Elaeocarpus stipularis, Pternandra echinata, Artocarpus nitidus griffithii, Ficus variegata, Ficus fistulosa, Piper aduncum, along with unidentified leaves and pollen. Horsfield’s fruit bats also feed on Payena lucida and the flowers of Parkia speciosa. Studies have noted that their diet and food preference is based on seasonal availability. During the dry season, when fruit abundance is low, these bats shift their diet to an increased utilization of pollen. Horsfield’s fruit bats have also been observed feeding on fruits from orchards and gardens. These bats have been observed removing fruit from trees and moving to a feeding roost to consume them. They extract fruit juices by pushing them against their palate with their tongue, the pulp and seeds are spit out and fall to the ground. Smaller seeds are ingested and dispersed later in their feces. Body size plays an important role in resource partitioning within their genus. For instance, Horsfield’s fruit bats have a large body size and have been documented carrying heavy figs and producing heavy regurgitated pellets. There are also descriptions of these bats becoming intoxicated and lying at the base of trees after drinking palm juice collected in pots. (Campbell and Kunz, 2006; Campbell, et al., 2007; Fletcher, et al., 2012; Funakoshi and Zubaid, 1997; Lekagul and McNeely, 1977)

  • Plant Foods
  • leaves
  • fruit
  • pollen
  • flowers

Predation

There is limited data available on the predation of this species, but in general, fruit bats, especially those on islands, have few natural predators, limited to mammals, birds of prey and reptiles. (Mickleburgh, et al., 1992)

Ecosystem Roles

Horsfield’s fruit bats, along with other fruit bat species, play a crucial role in seed dispersal and pollination of a wide array of plant species. Their role in seed dispersal and pollination has several effects on the plant community including promoting increased biodiversity, increased germination, forest regeneration, protection from seed predators, forest gap development and alteration of successional development. (Funakoshi and Zubaid, 1997; Hodgkinson, et al., 2003)

  • Ecosystem Impact
  • disperses seeds
  • pollinates

Economic Importance for Humans: Positive

The economic importance of this species is centered on their role as a pollinator and seed disperser. Several species that are pollinated or have their seeds dispersed by these bats are used for medicine, timber or food sources, resulting in millions of dollars to local economies annually. (Campbell and Kunz, 2006; Fletcher, et al., 2012; Mickleburgh, et al., 1992)

  • Positive Impacts
  • pollinates crops

Economic Importance for Humans: Negative

Although there are no reported adverse effects of Horsfield’s fruit bats on humans, due to their frugivory, they are likely a pest to commercial fruit crops.

  • Negative Impacts
  • crop pest

Conservation Status

According to the International Union for Conservation of Nature (IUCN), Horsfield’s fruit bats are currently listed as a species of least concern because they are widespread, there are no major threats in their range and they are not declining at a rate that warrants an elevation of their status to a more threatened level. Horsfield’s fruit bats, along with other members of their genus, tolerate human disturbances and are able to roost in urban and agricultural areas. (Bates, et al., 2008; Campbell, et al., 2006)

Contributors

Erick Smogoleski (author), University of Wisconsin-Stevens Point, Christopher Yahnke (editor), University of Wisconsin-Stevens Point, Leila Siciliano Martina (editor), Animal Diversity Web Staff.

Glossary

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

folivore

an animal that mainly eats leaves.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

polygynous

having more than one female as a mate at one time

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

2009. "AnAge entry for Cynopterus brachyotis" (On-line). AnAge: The Animal Ageing and Longevity Database. Accessed May 10, 2013 at http://genomics.senescence.info/species/entry.php?species=Cynopterus_brachyotis.

2009. "AnAge entry for Cynopterus sphinx" (On-line). AnAge: The Animal Ageing and Longevity Database. Accessed May 10, 2013 at http://genomics.senescence.info/species/entry.php?species=Cynopterus_sphinx.

Bates, P., C. Francis, M. Gumal, S. Bumrunsri. 2008. "Cynopterus horsfieldii" (On-line). The IUCN Redlist of Threatened Species. Accessed May 09, 2013 at http://www.iucnredlist.org/details/6104/0.

Campbell, P. 2008. The Relationship between Roosting Ecology and Degree Polygyny in Harem-Forming Bats: Perspectives from Cynopterus. Journal of Mammalogy, 89/6: 1351-1360. Accessed February 27, 2013 at http://jstor.org/stable/30224403.

Campbell, P., T. Kunz. 2006. Cynopterus horsfieldii. Mammalian Species, 802: 1-5. Accessed February 24, 2013 at http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/.

Campbell, P., N. Reid, A. Zubaid, A. Adnan, T. Kunz. 2006. Comparative Roosting Ecology of Cynopterus (Chiroptera: Pteropodidae) Fruit Bats in Peninsular Malaysia. Biotropica, 38/6: 725-734. Accessed February 27, 2013 at http://jstor.org/stable/30044050.

Campbell, P., C. Schneider, A. Zubaid, A. Adnan, T. Kunz. 2007. Morphological and Ecological Correlates of Coexistence in Malaysian Fruit Bats (Chiroptera: Pteropodidae). Journal of Mammalogy, 88/1: 105-118.

Crichton, E., P. Krutzsch. 2000. Reproductive Biology of Bats. San Diego, CA: Academic Press.

Fletcher, C., Z. Akbar, T. Kunz. 2012. Fruit Diet of frugivorous bats (Cynopterus brachyotis and Cynopterus horsfieldii) in tropical hill forests of Peninsular Malaysia. Mammalia, 76/4: 389-397.

Funakoshi, K., A. Zubaid. 1997. Behavioural and reproductive ecology of the dog-faced bats, Cynopterus brachyotis and C. horsfieldii, in a Malaysian rainforest. Mammal Study, 22/1: 95-108.

Hodgkinson, R., S. Balding, A. Zubaid, T. Kunz. 2003. Fruit Bats (Chiroptera: Pteropodidae) as Seed Dispersers and Pollinators in a Lowland Malaysian Rainforest. Biotropica, 35/4: 491-502.

Hodgkinson, R., S. Balding, A. Zubaid, T. Kunz. 2004. Habitat Structure, Wing Morphology, and the Vertical Stratification of Malaysian Fruit Bats (Megachiroptera: Pteropodidae). Journal of Tropical Ecology, 20/6: 667-673. Accessed February 27, 2013 at http://jstor.org/stable/4092111.

Lekagul, B., J. McNeely. 1977. The Mammals of Thailand. Bangkok, Thailand: Association of Conservation and Wildlife.

Mickleburgh, S., A. Hutson, J. Racey. 1992. "Old World Fruit Bats: an action plan for their Conservation" (On-line pdf). Accessed April 25, 2013 at http://data.iucn.org/dbtw-wpd/edocs/1992-034.pdf.

Neuweiler, G. 2000. The Biology of Bats. New York: Oxford University Press.

Shmitt, L., D. Kitchener, R. How. 1995. A Genetic Perspective of Mammalian Variation and Evolution in the Indonesian Archipelago: Biogeographic Correlates in the Fruit Bat Genus Cynopterus. Evolution, 49/3: 399-412.

Tan, K., A. Zubaid, T. Kunz. 1999. Roost Selection and Social Organization in Cynopterus horsfieldii (Chiroptera: Pteropodidae). Malayan Nature Journal, 53/4: 295-298.

Vijaya, J., C. Laman, M. Abdullah. 2012. A Predictive Model to Differentiate the Fruit Bats Cynopterus brachyotis and C. cf. brachyotis Forest (Chiroptera: Pteropodidae) from Malaysia Using Multivariate Analysis. Zoological Studies, 51/2: 259-272.

Zubaid, A., A. Fatimah. 1990. Hair Morphology of Malausian Pteropodids. Mammalia, 54/4: 627-632.

Zubaid, A., T. Kunz, P. Campbell, C. Schneider, A. Adnan. 2007. Morphological and Ecological Correlates of Coexistence in Malaysian Fruit Bats (Chiroptera: Pteropodidae). Journal of Mammalogy, 88/1: 105-118. Accessed February 27, 2013 at http://jstor.org/stable/4126856.