Pteropus rufus 

Scope: Global
Language: English

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Taxonomy [top]

Kingdom Phylum Class Order Family
Animalia Chordata Mammalia Chiroptera Pteropodidae

Scientific Name: Pteropus rufus É. Geoffroy Saint-Hilaire, 1803
Common Name(s):
English Madagascan Flying Fox, Madagascar Flying-fox, Madagascar Fruit Bat
French Renard volant
Spanish Zorro Volador de Madagascar
Taxonomic Notes: Synonyms = Pteropus (edwardsi) rufus rufus Kaudern (1915); Pteropus phaiops Temminick (1825); Pteropus rufus Dorst (1948); Pteropus rufus princeps Grandidier & Petit (1932); Pteropus rufus princeps K. Andersen (1908); Pteropus rufus rufus Grandidier & Petit (1932); Pteropus rufus rufus K. Andersen (1912).

Assessment Information [top]

Red List Category & Criteria: Vulnerable A2acd ver 3.1
Year Published: 2016
Date Assessed: 2014-03-31
Assessor(s): Racey, P.A.
Reviewer(s): Mildenstein, T.
Contributor(s): Andriafidison , D., Goodman, S., Hutson, A.M., Jenkins, R.K.B., Razafimanahaka, J., Cardiff, S.G., Kofoky, A., Oleksy, R., Rabearivelo, A., Ranivo, J. & Ratrimomanarivo, F.H.
This species is listed as Vulnerable based on an estimated population reduction of over 30% in the last three generations (18 years; Pacifici et al. 2013). The species can be legally hunted and the reduction can be attributed to the increased level of hunting particularly with the introduction of shotguns and continued loss of its preferred roosting habitats.
Previously published Red List assessments:

Geographic Range [top]

Range Description:This species is endemic to Madagascar (Simmons 2005). It is one of the most widespread bat species on the island and appears to only be absent from the highly populated central highlands (MacKinnon et al. 2003). The highest density of roost sites is in coastal regions, especially from Morombe in the south-west to Antsiranana in the north (MacKinnon et al. 2003).
Countries occurrence:
Additional data:
Upper elevation limit (metres):1200
Range Map:Click here to open the map viewer and explore range.

Population [top]

Population:Roosts are conspicuous and noisy but the bats are often tightly clustered making abundance estimates at the larger sites difficult. MacKinnon et al. (2003) reported that colony size ranged from 10 to 5,000 animals with a median of 400. They found that nationally 17.5% of 154 roosts had been abandoned in the last ten years and desertion rates were much higher (70%) in the central highlands. The estimated national population size in 2000 was 300,000 (MacKinnon et al. 2003).
Current Population Trend:Decreasing
Additional data:
Number of mature individuals:300000
Population severely fragmented:No

Habitat and Ecology [top]

Habitat and Ecology:It roosts in a variety of different tree species where the bats hang on the outermost branches (MacKinnon et al. 2003, Jenkins et al. 2007a, Long and Racey 2007). Roosts are very rare inside intact forest and are usually found in forest fragments, small islands or mangroves (MacKinnon et al. 2003, Jenkins et al. 2007a, Rakotoarivelo and Randrianandriananina 2007). Occasionally, Eucalyptus plantations are also used (Jenkins et al. 2007a). The majority of roosts are in small areas of relatively degraded vegetation with a few large trees. The roosts are usually adjacent to water bodies (Jenkins et al. 2007a) and it seems that bats use them for navigation and orientation during foraging (Jenkins et al. 2007a, Oleksy 2014). Roosting bats are easily disturbed by people and cattle that venture near the roost and colonies readily take flight (MacKinnon et al. 2003). Once disturbed the bats may move to an alternative roost site in the vicinity and it appears that each colony requires more than one roost site as a response to disturbance but also maybe to shifting patterns of food availability (Jenkins et al. 2007a). This species can survive in heavily modified landscapes through feeding on a mixture of native and introduced plants (Raheriaisena 2005, Jenkins et al. 2007a, Long and Racey 2007) but it shows preferences towards native vegetation (e.g. forest fragments). It travels up to30 km a night between roosting and foraging areas (Long 2002, Oleksy 2014). The average flight speed of commuting flights is over 32 km/h and the highest recorded speed is 61 km/h (Oleksy 2014). In Berenty Reserve, south east Madagascar, home range of over 58,000 ha was recorded for 15 tracked individuals (Oleksy 2014). During the gestation period the females appear to fly significantly longer distances than males, probably to fulfil their increased energy requirements (Oleksy 2014).

The diet consists predominantly of fruit juices which are squeezed from fruit in the mouth. Large numbers of small seeds are accidentally ingested during feeding and are later dispersed at other foraging sites, during flight or at the roost. The average gut transit time of these bats is 12 min, however theyare able to retain the seeds in their gut for over 20h (Oleksy 2014). There is some evidence that seeds that have passed through the digestive tract of P. rufus incur a fitness advantage through increased germination success (Racey et al. 2009). This has been recently confirmed on two species of strangler figs (F. polita and F. grevei) in south east Madagascar. Both species showed significant increase in germination after bat ingestion and had higher survival rate than seedlings originating from unprocessed seeds (Oleksy 2014) Other plants parts are also consumed, including flowers, nectar and leaves. In the south, nectar from introduced plants is an important dietary component throughout the year (Long and Racey 2007). Some plants have evolved to attract nocturnal mammals like bats as pollinators and P. rufus feeds non-destructively on the nectar of two threatened baobab species (Baum 1995, Andriafidison et al. 2006). Because of its ability to travel long distances and its capability as a seed disperser and pollinator P. rufus is widely believed to be a key species in fragmented forest ecosystems (Bollen and Van Elsacker 2002, Bollen et al. 2004).
Generation Length (years):6

Use and Trade [top]

Use and Trade: It is an important subsistence food but also of commercial importance.

Threats [top]

Major Threat(s): There are numerous threats to this species. It is listed as a game species under Malagasy law and can be legally hunted between May and August (Durbin 2007). Only the few roosts that are located in protected areas therefore receive some protection (Racey et al. 2009). Pteropus rufus is hunted across Madagascar (MacKinnon et al. 2003, Rakotondravony 2006, Rakotonandrasana and Goodman 2007, Racey et al. 2009) where it is an important subsistence food but also of commercial importance. Investigations in eastern Madagascar revealed that people in urban areas are twice as likely to eat this species than those in rural areas: over 40% of the people interviewed in urban areas had eaten P. rufus compared with only 20% in rural areas (Jenkins et al. 2011). This proportion goes up to 60% in western Madagascar, with rural and urban areas combined (Razafimanahaka et al. 2012). Although quantitative data on the supply of P. rufus for food and the impact of this harvest on colonies/populations is scarce, there is some evidence that the offtake is locally unsustainable (Racey et al. 2009).
Hunting occurs at roost sites and where the bats forage, especially when they feed on trees in villages (Jenkins and Racey 2008). A study during the austral winter in the western Madagascar town of Mahabo revealed that seven active hunters can kill 381 bats over a period of 43 days, with average of almost 4 bats a night. Over 80% of their catch is sold to the public . Hunters usually operate for around 90 days a year. There are up to 30 active hunters in Mahabo alone, so they could potentially account for 9,000 bats during 90 days of active hunting a year (Oleksy 2010). There is also evidence that P. rufus from Ile Sainte Marie is hunted commercially and frozen shipments are sent to the mainland port of Toamasina (Rakotonandrasana and Goodman 2007). The conversion of the forest used by roosting bats into agriculture is another threat to P. rufus and results in the permanent loss of suitable trees which in many cases have been used for decades. Pteropus rufus feeds on a number of cultivated fruits that have a high economic value in rural Madagascar and is often subjected to persecution.

Conservation Actions [top]

Conservation Actions: It is listed on CITES Appendix II and is a game species under Malagasy law (Durbin 2007) but neither of these provide any practical in situ conservation measures (Racey et al. 2009). There are a few roosts in protected areas, notably Parc National Kirindy -Mité, Parc National de Masoala, Parc National de Mananara-nord (MacKinnon et al. 2003) and Berenty Private Reserve (Long 2002), but many of the existing parks and reserves appear to be without roosting colonies (e.g. Goodman 1996, 1999; Alonso et al. 2002; Goodman et al. 2005; Schmidand and Alonso 2005). The ongoing process to triple protected areas in Madagascar is providing an unprecedented opportunity to include traditional roosts in the new conservation sites. There is also significant scope for local institutions to conserve roosts and this is already occurring in some parts of Madagascar where the bats use sacred forests (Jenkins et al. 2007b) or where communities have created social contracts to protect the bats (Jenkins et al. 2007a).

Classifications [top]

1. Forest -> 1.5. Forest - Subtropical/Tropical Dry
1. Forest -> 1.6. Forest - Subtropical/Tropical Moist Lowland
1. Forest -> 1.7. Forest - Subtropical/Tropical Mangrove Vegetation Above High Tide Level
14. Artificial/Terrestrial -> 14.3. Artificial/Terrestrial - Plantations
14. Artificial/Terrestrial -> 14.6. Artificial/Terrestrial - Subtropical/Tropical Heavily Degraded Former Forest
1. Land/water protection -> 1.2. Resource & habitat protection
2. Land/water management -> 2.1. Site/area management
3. Species management -> 3.1. Species management -> 3.1.1. Harvest management
4. Education & awareness -> 4.3. Awareness & communications
5. Law & policy -> 5.1. Legislation -> 5.1.2. National level
5. Law & policy -> 5.1. Legislation -> 5.1.3. Sub-national level
5. Law & policy -> 5.4. Compliance and enforcement -> 5.4.1. International level
5. Law & policy -> 5.4. Compliance and enforcement -> 5.4.2. National level
5. Law & policy -> 5.4. Compliance and enforcement -> 5.4.3. Sub-national level
6. Livelihood, economic & other incentives -> 6.1. Linked enterprises & livelihood alternatives

In-Place Research, Monitoring and Planning
In-Place Land/Water Protection and Management
  Conservation sites identified:Yes, over entire range
In-Place Species Management
In-Place Education
  Subject to recent education and awareness programmes:Yes
  Included in international legislation:Yes
  Subject to any international management/trade controls:Yes
11. Climate change & severe weather -> 11.4. Storms & flooding
♦ timing:Ongoing    
→ Stresses
  • 1. Ecosystem stresses -> 1.2. Ecosystem degradation

2. Agriculture & aquaculture -> 2.1. Annual & perennial non-timber crops -> 2.1.1. Shifting agriculture
♦ timing:Ongoing    
→ Stresses
  • 1. Ecosystem stresses -> 1.1. Ecosystem conversion
  • 1. Ecosystem stresses -> 1.2. Ecosystem degradation

5. Biological resource use -> 5.1. Hunting & trapping terrestrial animals -> 5.1.1. Intentional use (species is the target)
♦ timing:Ongoing    
→ Stresses
  • 2. Species Stresses -> 2.1. Species mortality

5. Biological resource use -> 5.3. Logging & wood harvesting -> 5.3.5. Motivation Unknown/Unrecorded
♦ timing:Ongoing    
→ Stresses
  • 1. Ecosystem stresses -> 1.2. Ecosystem degradation

7. Natural system modifications -> 7.1. Fire & fire suppression -> 7.1.3. Trend Unknown/Unrecorded
♦ timing:Ongoing    
→ Stresses
  • 1. Ecosystem stresses -> 1.2. Ecosystem degradation

1. Research -> 1.2. Population size, distribution & trends
1. Research -> 1.3. Life history & ecology
3. Monitoring -> 3.1. Population trends

Bibliography [top]

Alonso, L. E., Schulenberg, T. S., Radilofe, S. and Missa, O. 2002. Evaluation rapide de la diversité biologique de reptiles et amphibians de la Reserve Naturelle Integrale d'Ankarafantsika. RAP Bulletin of Biological Assessment No. 23. Conservation International, Washington, DC, USA.

Andriafidison, D., Andrianaivoarivelo, R.A., Jenkins, R.K.B., Ramilijaona, O., Razanahoera, M., MacKinnon, J. and Racey, P.A. 2006. Nectarivory by endemic Malagasy fruit bats in the dry season. Biotropica 38: 85-90.

Baum, D. A. 1995. The comparative pollination and floral biology of Baobabs (Adansonia - Bombacaceae). Annals of the Missouri Botanical Gardens 82: 323-348.

Bollen, A. and van Elsacker, L. 2002. Feeding ecology of Pteropus rufus (Pteropodidae) in the littoral forest of Sainte Luce, SE Madagascar. Acta Chiropterologica 4: 33-47.

Bollen, A., van Elsacker, L. and Ganzhorn, J. U. 2004. Relations between fruits and disperser assemblages in a Malagasy littoral forest: a community-level approach. Journal of Tropical Ecology 20: 599-612.

Durbin, J. 2007. New legislation for the protection of Malagasy species. Lemur News 11: 4-6.

Goodman, S. M. 1996. Results of a bat survey of the eastern slopes of the Reserve Naturelle Integrale d'Andringitra, Madagascar. A floral and faunal inventory of the eastern slopes of the Reserve Naturelle Integrale d'Andringitra, Madagascar: With reference to elevational variation. Fieldiana: Zoology, new series 85: 284-288.

Goodman, S. M. 1999. Notes on the Bats of the Réserve Intégrale d'Andohahela and Surrounding Areas of Southeastern Madagascar. Fieldiana: Zoology 94: 251-257.

Goodman, S.M., Andriafidison, D., Andrianaivoarivelo, R., Cardiff, S.G., Ifticene, E., Jenkins, R.K.B., Kofoky, A., Mbohoahy, T., Rakotondravony, D., Ranivo, J., Ratrimomanarivo, F., Razafimanahaka, J. and Racey, P.A. 2005. The distribution and conservation of bats in the dry regions of Madagascar. Animal Conservation 8: 153-165.

IUCN. 2016. The IUCN Red List of Threatened Species. Version 2016-1. Available at: (Accessed: 30 June 2016).

Jenkins, R.K.B. and Racey, P.A. 2008. Bats as bushmeat in Madagascar. Madagascar Conservation and Development 3(1): 22-30.

Jenkins, R. K. B., Andriafidison, D., Razafimanahaka, J. H., Rabearivelo, A., Razafindrakoto, N., Andrianandrasana, R. H., Razafimahatratra, E. and Racey, P. A. 2007b. Not rare, but threatened : the Madagascar Flying Fox Pteropus rufus in a fragmented landscape. Oryx 41: 263-271.

Jenkins, R. K. B., Kofoky, A. F, Russ, J. M.Friafidison, A., Siemers, B. M., Randrianandrianina, F. H., Mbohoahy, T., Rahaingodrahety, V. N and Racey, P. A. 2007a. Ecology of Bats in the southern Anosy Region. In: J. U. Ganzhorn, S. M. Goodman, and M. Vincelette (eds), Biodiversity, ecology and conservation of littoral ecosystems in southeastern Madagascar, Tolagnaro (Fort Dauphin), Smithsonian Institution, Washington, DC, USA.

Jenkins, R., Keane, A., Rakotoarivelo, A., Rakotomboavonjy, V., Randrianandrianina, F., Razafimanahaka, H., Ralaiarimalala, S., Jones, J. 2011. Analysis of Patterns of Bushmeat Consumption Reveals Extensive Exploitation of Protected Species in Eastern Madagascar. PLoS ONE 6(12).

Long, E. 2002. The feeding ecology of Pteropus rufus in a remnant gallery forest surrounded by sisal plantations in South-east Madagascar. School of Biological Sciences, University of Aberdeen.

Long, E. and Racey, P. A. 2007. An exotic plantation crop as a keystone resource for an endemic megachiropteran, Pteropus rufus in Madagascar. Journal of Tropical Ecology 23: 1-11.

MacKinnon, J., Hawkins, C. E. and Racey, P. A. 2003. Pteropodidae. In: S. M. Goodman and J. P. Benstead (eds), The Natural History of Madagascar, pp. 1299–1302. The University of Chicago Press, Chicago, USA and London, UK.

Oleksy, R. 2010. Bats as bushmeat in western Madagascar. The role of the village kapok trees. BSc (Hons) Thesis, University of Aberdeen.

Oleksy, R. 2014. The contribution of fruit bats to forest regeneration in Madagascar. Ph.D. Thesis, University of Bristol.

Pacifici, M., Santini, L., Di Marco, M., Baisero, D., Francucci, L., Grottolo Marasini, G., Visconti, P. and Rondinini, C. 2013. Generation length for mammals. Nature Conservation 5: 87–94.

Racey, P.A., Goodman, S.M. and Jenkins, R.K.B. 2009. The ecology and conservation of Malagasy bats. In: T.H. Fleming and P.A. Racey (eds), Island Bats, Chicago University Press, Chicago, USA.

Raheriaisena, M. 2005. Régime alimentaire de Pteropus rufus (Chiroptera: Pteropodidae) dans le région sub-aride du sud de Madagascar. La Terre et la vie: Revue d'Ecologie 60: 255-264.

Rakotoarivelo, A. R. and Randrianandriananina, F. H. 2007. A chiropteran survey of the Lac Kinkony-Mahavavy area in western Madagascar. African Bat Conservation News 12: 2-4.

Rakotonandrasana, E. N. and Goodman, S. M. 2007. Bat inventories of the Madagascar offshore islands of Nosy Be, Nosy Komba and Ile Sainte Marie. African Bat Conservation News 12: 6-10.

Rakotondravony, H. 2006. Communautés locales et gibiers dans la région de Daraina, extrême Nord-Est de Madagascar. Madagascar Conservation and Development 1: 19-21.

Razafimanahaka, J.H., Jenkins, R.K.B., Andriafidison, D., Randrianandrianina, F.l., Rakotomboavonjy, V., Keane, A., Jones, J.P.G. 2012. Novel approach for quantifying illegal bushmeat consumption reveals high consumption of protected species in Madagascar. Oryx 46: 584-592.

Schmid, J. and Alonso, L. E. 2005. A rapid biological assessment of the Mantadia-Zahamena corridor, Madagascar. Conservation International, Washington, DC, USA.

Simmons, N.B. 2005. Order Chiroptera. In: D.E. Wilson and D.M. Reeder (eds), Mammal Species of the World, pp. 312-529. The Johns Hopkins University Press, Baltimore, MD, USA.

Citation: Racey, P.A. 2016. Pteropus rufus. In: The IUCN Red List of Threatened Species 2016: e.T18756A22087230. . Downloaded on 24 September 2017.
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