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Rhinolophus luctuswoolly horseshoe bat

By Codey Dueck

Geographic Range

Great woolly horseshoe bats (Rhinolophus luctus) are distributed throughout South and South-East Asia. Their range extends north into southern China, west into India, and as far south as the island of Java. (Molur, et al., 2002; Walston, et al., 2008)

Habitat

Woolly horseshoe bats primarily live in caves and in tropical to subtropical forests among foliage or in tree cavities. Less commonly, R. luctus may be found roosting between rocks in mountain ranges up to 1600 m, in tunnels, and under tree bark. Additionally, while they generally roost away from human activity, they have been found in fragmented forests and abandoned human structures such as mine shafts. Agricultural land is an unsuitable habitat for R. luctus. (Lim, et al., 2014; Molur, et al., 2002; Walston, et al., 2008; Wordley, et al., 2015)

The differences between the R. luctus subspecies lie primarily in their ranges. R. l. foetidus is found on the island of Borneo; R. l. lanosus is found in the Fukien province of China; R. l. luctus is found on the island of Java, Singapore, and Tenasserim; R. l. morio is found throughout the Malay Peninsula; R. l. perniger is found in Burma and Nepal; and R. l. spurcus is found on the island of Hainan. (Andersen, 1918; Ellerman and Morrison-Scott, 1951)

  • Range elevation
    0 to 1600 m
    0.00 to 5249.34 ft

Physical Description

Like all horseshoe bats, Rhinolophus luctus has a complex nose-leaf that covers the upper lip and surrounds the nostrils. Specifically, woolly horseshoe bats have a broad sella and trifoliate nose-leaf. Their large ears (between 38 and 42 mm) and black-to-brown woolly fur set them apart from other Rhinolophus species. Additionally, their dental formula is I1/2 C1/1 P2/3 M3/3. (Andersen, 1918; Bates, et al., 2004; Topal and Csorba, 1992; Walker, 1975)

Woolly horseshoe bats are the largest species of horseshoe bat, with adults weighing between 22.3 and 33.9 g. Their bodies are 80 to 99 mm long and their tail length ranges from 46 to 59 mm. The wingspan of these animals has not been evaluated. Female-biased sexual dimorphism is present in the body mass and forearm length. On average, females weigh 27.81 g and their forearms are 67.77 mm in length whereas males weigh 22.29 g and have 63.27 mm long forearms. (Bates, et al., 2004; Wu, et al., 2018)

The six subspecies of R. luctus vary slightly in relative skull width (zygomatic width:mandible length) and relative length of lower tooth row (lower CM3 length:mandible length). In terms of the relative skull width, R. l. lanosus (0.64) is the smallest followed by R. l. perniger (0.68-0.70), R. l. foetidus (0.72), R. l. spurcus (O.72), R. l. morio (0.74), and R. l. luctus (0.78). In terms of the relative length of the lower tooth row, R. l. lanosus (0.54) is the smallest followed by R. l. spurcus (0.56), R. l. morio (0.57), R. l. perniger (0.57-0.58), R. l. foetidus (0.58), and R. l. luctus (0.63). (Volleth, et al., 2015)

  • Sexual Dimorphism
  • female larger
  • Range mass
    22.3 to 33.9 g
    0.79 to 1.19 oz
  • Range length
    80 to 99 mm
    3.15 to 3.90 in

Reproduction

Unlike the polygynous mating system of other horseshoe bats, woolly horseshoe bats are monogamous. Additionally, females show strong sexual selection on male echolocation frequency and weak sexual selection on male body size. (Wu, et al., 2018)

While there is little available information on the breeding cycles of R. luctus specifically, it is unlikely that they differ from the non-synchronous monoestry that is present in other horseshoe bats. However, one species description found a fetus in the advanced stage (10 g in weight) in the uterine horn of an individual woolly horseshoe bat that indicated a June parturition. (Krutzsch and Crichton, 2000; Saikia, et al., 2004)

Due to a lack of information, the following is an extrapolation to woolly horseshoe bats based on shared characteristics within their family. Horseshoe bats typically have delayed fertilization or implantation of only one embryo once a year in spring with a gestation period of approximately 7 weeks. In the similar-sized Rhinolophus ferrumequinum, lactation lasts for up to 45 days. Finally, horseshoe bats reach sexual maturity at approximately 2 years of age. (Krutzsch and Crichton, 2000; MacDonald, 2006; Walker, 1975)

  • Breeding interval
    Not reported, but likely once yearly
  • Breeding season
    Not reported, but likely spring or winter
  • Average number of offspring
    1
  • Average gestation period
    7 weeks
  • Average weaning age
    45 days
  • Average age at sexual or reproductive maturity (female)
    2 years
  • Average age at sexual or reproductive maturity (male)
    2 years

Woolly horseshoe bats are born in an altricial state and although there are no documented cases of parental care, they provide maternal care in the form of lactation (up to 45 days in the similar-sized Rhinolophus ferrumequinum). In addition to the two functional teats on the abdomen, woolly horseshoe bats have two additional dummy teats that are not connected to mammary glands. Juveniles wrap their legs around their mother's neck and attach to one of these dummy teats with their mouths during flight. (Krutzsch and Crichton, 2000; MacDonald, 2006; Walker, 1975)

Lifespan/Longevity

There is no data on the specific lifespan of woolly horseshoe bats.

Behavior

Great woolly horseshoe bats roost in monogamous pairs or solitarily. Additionally, in these monogamous pairs, individuals do not roost touching the other and do not share their roosts with any other species. Finally, these bats are found at very low densities which makes it difficult to accurately assess population sizes. (Lim, et al., 2014; Saikia, et al., 2004; Topal and Csorba, 1992; Walker, 1975; Walston, et al., 2008)

There is insufficient information about the specific activity of woolly horseshoe bats, but horseshoe bats generally are all nocturnal and forage solitarily later in the night than other bats. Additionally, Rhinolophus luctus can enter a deep torpor, but hibernation has not been assessed. (Baniya, et al., 2019; Walker, 1975)

Home Range

There is no data on the specific home range of woolly horseshoe bats.

Communication and Perception

The woolly horseshoe bat's small and partially obscured eyes combined with frequent cave roosts suggests that vision may not be an important sense. Echolocation has been directly observed in R. luctus, ranging from 32 to 43 Hz and which exhibits a positive relationship with the body size among the subspecies. (Klaus-Gerhard and Volleth, 1995; Molur, et al., 2002; Volleth, et al., 2015; Walston, et al., 2008)

Food Habits

Great woolly horseshoe bats are insectivorous, feeding primarily on beetles and termites, and secondarily on other insects. While feeding habits of R. luctus is understudied, Rhinolophid bats typically use slow maneuverable flight in combination with their echolocation to locate and capture prey in a gleaning (hunting grounded prey) or aerial hawking (catching flying prey) fashion. It is possible that R. luctus hunts in the same way. (Klaus-Gerhard and Volleth, 1995; MacDonald, 2006; Walker, 1975; Walston, et al., 2008)

  • Animal Foods
  • insects

Predation

No specific information on the predation of R. luctus is available. However, they likely are depredated by the same predators of bats in general, like owls, hawks, and snakes. Furthermore, it is possible that woolly horseshoe bats' thick black fur could function as a form of crypsis, especially in their roosts. (Bates, et al., 2004; Hill and Smith, 1984)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Great woolly horseshoe bats are insectivorous and consume beetles, termites, and other insects and thus negatively impact these populations. There is little information on these bats, but generally bats are not a primary prey group for many predators, but owls and snakes are known to depredate them. Additionally, bats also carry parasites such as mites, trematodes, and fleas, and there are no plausible reasons that woolly horseshoe bats would differ from this trend. Furthermore, Rhinolophus luctus has been shown to be a possible reservoir host for some economically important diseases such as Nipah virus and SARS-like coronavirus. (Hill and Smith, 1984; Plowright, et al., 2019; Walston, et al., 2008)

Commensal/Parasitic Species

Economic Importance for Humans: Positive

There are no positive human economic roles for R. luctus aside from being one of the species hunted for usage in Indian and Chinese Traditional Medicine. A drawback of these bats' rarity and solitary or paired roosting behavior is that they are not likely to significantly contribute to insect control. (Molur, et al., 2002; Riccucci, 2016)

Economic Importance for Humans: Negative

There are no major negative economic impacts on humans documented for R. luctus. A benefit of these bats' rarity and solitary or paired roosting behavior is that they are not likely to become a pest species in human developments.

It is worth noting, however, that R. luctus has been identified as a potential reservoir host for Nipah virus and SARS-like coronavirus, although their solitary nature, insectivorous diet, and usual avoidance of agricultural developments make the risk of zoonotic transmission low. (Plowright, et al., 2019; Tang, et al., 2006)

Conservation Status

Woolly horseshoe bat populations might be declining but not at a rate sufficient to put them into a more protected category. Deforestation and hunting are not a significant threat due to their tolerance of anthropological disturbance and wide range. Additionally, R. luctus resides in many protected regions such as Satpura National Park (India) and Vu Quang National Park (Vietnam). However, despite listing woolly horseshoe bats as Least Concern, the IUCN Red List recommends an increase in population studies and ecological monitoring. (Borissenko and Kruskop, 2003; Molur, et al., 2002; Walston, et al., 2008)

Other Comments

Subspecies of the great woolly horseshoe bat include Rhinolophus luctus foetidus, R. l. lanosus, R. l. luctus, R. l. morio, R. l. perniger (most common), and R. l. spurcus. Older sources may refer to Rhinolopus luctus as Rhinolophus perniger and Rhinolophus lanosus (now both are classified as subspecies), or Rhinolophus geminus. However, the taxonomy remains unresolved and in need of study. For example, R. l. morio has been called to be elevated to the species rank due to a Y-autosomal translocation and skull variations. (Andersen, 1918; Ellerman and Morrison-Scott, 1951; Simmons, 2005; Topal and Csorba, 1992; Volleth, et al., 2015; Walston, et al., 2008)

Contributors

Codey Dueck (author), University of Manitoba, Annemarie van der Marel (editor), University of Manitoba, Genevieve Barnett (editor), Colorado State University.

Glossary

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

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.

carnivore

an animal that mainly eats meat

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chemical

uses smells or other chemicals to communicate

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

delayed fertilization

a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.

delayed implantation

in mammals, a condition in which a fertilized egg reaches the uterus but delays its implantation in the uterine lining, sometimes for several months.

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

echolocation

The process by which an animal locates itself with respect to other animals and objects by emitting sound waves and sensing the pattern of the reflected sound waves.

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

fertilization

union of egg and spermatozoan

forest

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

insectivore

An animal that eats mainly insects or spiders.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

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

seasonal breeding

breeding is confined to a particular season

sexual

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

solitary

lives alone

suburban

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

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

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.

ultrasound

uses sound above the range of human hearing for either navigation or communication or both

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

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Topal, G., G. Csorba. 1992. The subspecific division of Rhinolophus luctus Temminck, 1835, and the taxonomic status of Rhinolophus beddomei Anderson, 1905 (Mammalia, Chiroptera). Miscellanea Zoologica Hungarica, 7: 101-116.

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