List of bats of Madagascar
Bats are one of the major components of the indigenous mammalian fauna of Madagascar, in addition to tenrecs, lemurs, euplerid carnivores, and nesomyine rodents. Forty-six bat species have so far been recorded on Madagascar, of which thirty-six occur only on the island. However, new species continue to be discovered, causing the number of species to rise rapidly; for example, Nick Garbutt's Mammals of Madagascar listed only 36 species. Most Malagasy bats have their origins in nearby mainland Africa, but on at least three occasions—Pipistrellus raceyi, Pteropus rufus, and the species pair Emballonura atrata–E. tiavato—bats have colonized Madagascar from Asia.
Taxonomic classification
The following bat genera and families include species found on Madagascar :- Family Pteropodidae
- *Genus Eidolon
- *Genus Pteropus
- *Genus Rousettus
- Family Hipposideridae
- *Genus Hipposideros
- *Genus Macronycteris
- *Genus Paratriaenops
- *Genus Triaenops
- Family Emballonuridae
- *Genus Coleura
- *Genus Emballonura
- *Genus Taphozous
- Family Nycteridae
- *Genus Nycteris
- Family Myzopodidae
- *Genus Myzopoda
- Family Molossidae
- *Genus Chaerephon
- *Genus Mops
- *Genus Mormopterus
- *Genus Otomops
- *Genus Tadarida
- Family Miniopteridae
- *Genus Miniopterus
- Family Vespertilionidae
- *Genus Hypsugo
- *Genus Myotis
- *Genus Neoromicia
- *Genus Pipistrellus
- *Genus ''Scotophilus''
Family Pteropodidae
Pteropodidae are a diverse family, with 186 species recognized in 2005, which occurs across the tropical regions of the Old World. They include the largest bats, but also some smaller species, and are mostly diurnal and frugivorous. Three species are known from Madagascar; each is classified in its own genus and is most closely related to species from outside Madagascar.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Eidolon dupreanum | 1866, Pollen | Madagascar only | 115 to 130 mm | Vulnerable | |
| Pteropus rufus | 1803, E. Geoffroy | Madagascar only | 155 to 175 mm | Vulnerable | |
| Rousettus madagascariensis | 1928, G. Grandidier | Madagascar only; absent in SW | 65 to 75 mm | Near threatened |
Family Hipposideridae
Hipposideridae are a moderately diverse family—81 species were listed in 2005—and occur across the Old World tropics. Insectivorous, cave-roosting, and characterized by an elaborate noseleaf, they have often been united with the horseshoe bats into a single family Rhinolophidae, but are currently classified separately. Six species, all endemic, are known from Madagascar, of which four are extant. Macronycteris commersoni is the largest non-pteropodid bat of Madagascar and the extinct Hipposideros besaoka was even larger. The other species belong to the closely related genera Triaenops and Paratriaenops; the latter was split from Triaenops in 2009 and is restricted to Madagascar and the western Seychelles.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Hipposideros besaoka | 2007, Samonds | Anjohibe, NW Madagascar only | – | Extinct | |
| Macronycteris commersoni | 1813, E. Geoffroy | Madagascar only | 83 to 97 mm | Near threatened | |
| Macronycteris cryptovalorona | 2016, Goodman et al. | Madagascar only | 80 to 81 mm | Not evaluated | |
| Paratriaenops auritus | 1912, G. Grandidier | N and NW Madagascar only | 44 to 51 mm | Vulnerable | |
| Paratriaenops furcula | 1906, Trouessart | W and SW Madagascar only | 42 to 49 mm | Least concern | |
| Triaenops goodmani | 2007, Samonds | Anjohibe, NW Madagascar only | – | Extinct | |
| Triaenops menamena | 2009, Goodman and Ranivo | N, W, and S Madagascar only | 46 to 56 mm | Least concern |
Family Emballonuridae
With 51 species, Emballonuridae are a moderately diverse family. Found in tropical and subtropical regions across the world, they are characterized by a tail that extends beyond the uropatagium, but may be retracted into a sheath. Four species are known from Madagascar, of which two are endemic and two others are shared with mainland Africa.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Coleura afra | 1852, Peters | Ankarana, N Madagascar; Sub-Saharan Africa; Yemen | 45 to 55 mm | Least concern | |
| Emballonura atrata | 1836, Eydoux and Gervais | E Madagascar only | c. 37 to 40 mm | Least concern | |
| Emballonura tiavato | 2006, Goodman et al. | N and W Madagascar only | 35 to 41 mm | Least concern | |
| Taphozous mauritianus | 1818, E. Geoffroy | Madagascar and other W Indian Ocean islands; Sub-Saharan Africa | 58 to 64 mm | Least concern |
Family Nycteridae
Nycteridae is a small family of 16 species in a single genus found in Africa and east to the Sunda Islands. They are characterized by a groove on their face and are insectivorous. A single, poorly known species has been recorded from Madagascar.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Nycteris madagascariensis | 1937, G. Grandidier | N Madagascar only | 50 to 52 mm | Data deficient |
Family Myzopodidae
This family, characterized by suction disks on the hand and feet, is unique to Madagascar. A single species has historically been recognized, but eastern and western populations were classified as separate species in 2007.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Myzopoda aurita | 1878, Milne-Edwards and Grandidier | E Madagascar only | 46 to 49 mm | Least concern | |
| Myzopoda schliemanni | 2007, Goodman et al. | W Madagascar only | 45 to 49 mm | Least concern |
Family Molossidae
This diverse family of 100 species occurs across the world in tropical regions. The tail conspicuously projects from the uropatagium and the wings are long. Eight species are known from Madagascar, four of which are endemic.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Chaerephon atsinanana | 2010, Goodman et al. | E Madagascar | 37 to 42 mm | - | |
| Chaerephon jobimena | 2004, Goodman and Cardiff | Madagascar only | 45 to 48 mm | Least concern | |
| Chaerephon leucogaster | 1870, A. Grandidier | W Madagascar; Sub-Saharan Africa; Mayotte | 33 to 38 mm | Not evaluated | |
| Mops leucostigma | 1918, G.M. Allen | Madagascar only | 42 to 45 mm | Least concern | |
| Mops midas | 1843, Sundevall | Madagascar; Sub-Saharan Africa; Saudi Arabia | 62 to 63 mm | Least concern | |
| Mormopterus jugularis | 1865, Peters | Madagascar only | 37 to 40 mm | Least concern | |
| Otomops madagascariensis | 1953, Dorst | Madagascar only | 59 to 65 mm | Least concern | |
| Tadarida fulminans | 1903, Thomas | Madagascar; Sub-Saharan Africa north to Kenya | 57 to 60 mm | Least concern |
Family Miniopteridae
This family contains a single genus, Miniopterus, with 19 species recognized in 2005, which was classified in Vespertilionidae until recently. Insectivorous and characterized by long fingers, the species are all quite similar, leading to a confused classification. On Madagascar, four species were recognized as recently as 2007, but systematic research has led the number to increase to eleven, of which nine are restricted to Madagascar and two shared with the Comoros.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Miniopterus aelleni | 2009, Goodman et al. | N and W Madagascar; Anjouan, Comoros | 35 to 41 mm | Not evaluated | |
| Miniopterus brachytragos | 2009, Goodman et al. | N and W Madagascar only | 35 to 38 mm | Not evaluated | |
| Miniopterus egeri | 2011, Goodman et al. | E Madagascar only | 37 to 40 mm | Not evaluated | |
| Miniopterus gleni | 1995, Peterson et al. | Madagascar only, except S | 47 to 50 mm | Least concern | |
| Miniopterus griffithsi | 2009, Goodman et al. | S Madagascar only | 48 to 50 mm | Not evaluated | |
| Miniopterus griveaudi | 1959, Harrison | N and W Madagascar; Comoros | 35 to 38 mm | Data deficient | |
| Miniopterus mahafaliensis | 2009, Goodman et al. | S Madagascar only | 35 to 40 mm | Not evaluated | |
| Miniopterus majori | 1906, Thomas | Madagascar only | 43 to 47 mm | Least concern | |
| Miniopterus manavi | 1906, Thomas | Central Highlands, Madagascar only | 38 to 39 mm | Least concern | |
| Miniopterus petersoni | 2008, Goodman et al. | SE Madagascar only | 38 to 43 mm | Data deficient | |
| Miniopterus sororculus | 2007, Goodman et al. | Central Highlands, Madagascar only | 42 to 45 mm | Least concern |
Family Vespertilionidae
With 407 species, Vespertilionidae is the largest bat family. Characterized by a tail contained in the uropatagium, they occur around the world in many habitats and are insectivorous. Madagascar hosts an endemic species of the extremely widespread genus Myotis, four species of the house bat Scotophilus, and at least six of small vespertilionids in the genera Hypsugo, Eptesicus, Neoromicia, and Pipistrellus. The classification of the "pipistrelles" is confused, leading to many changing identifications. In addition to the six "pipistrelles" listed here, the African Neoromicia nanus has also been recorded from Madagascar, but the identification of the Madagascar records needs to be confirmed.| Scientific name | Classified | Distribution | Forearm | Conservation status | References |
| Hypsugo anchietae | 1900, Seabra | SW Madagascar; southern Africa | 28 to 31 mm | Least concern | |
| Myotis goudoti | 1834, A. Smith | Madagascar only | 32 to 41 mm | Least concern | |
| Neoromicia malagasyensis | 1995, Peterson et al. | Isalo, SC Madagascar only | 30 to 32 mm | Endangered | |
| Neoromicia matroka | 1905, Thomas and Schwann | E Madagascar only | 31 to 33 mm | Least concern | |
| Neoromicia robertsi | 2012, Goodman et al. | E Madagascar only | 34 to 38 mm | Not evaluated | |
| Pipistrellus hesperidus | 1840, Temminck | W Madagascar; Sub-Saharan Africa | 29 to 31 mm | Least concern | |
| Pipistrellus raceyi | 2006, Bates et al. | W and E Madagascar only | 28 to 31 mm | Least concern | |
| Scotophilus cf. borbonicus | 1803, E. Geoffroy | Sarodrano, SW Madagascar; Réunion | 51 to 52 mm | Data deficient | |
| Scotophilus marovaza | 2006, Goodman et al. | W Madagascar only | 41 to 45 mm | Least concern | |
| Scotophilus robustus | 1881, Milne-Edwards | Madagascar only | 62 to 65 mm | Least concern | |
| Scotophilus tandrefana | 2005, Goodman et al. | W Madagascar only | 44 to 47 mm | Data deficient |
Literature cited
General
- Garbutt, N. 2007. Mammals of Madagascar: A Complete Guide. London: A & C Black, 304 pp.
- Goodman, S.M., Weyeneth, N., Ibrahim, Y., Saïd, I. and Ruedi, M. 2010b. . Acta Chiropterologica 12:117–141.
- Nowak, R.M. 1994. Walker's Bats of the World. Baltimore: The Johns Hopkins University Press, 287 pp.
- Simmons, N.B. 2005. Order Chiroptera. Pp. 312–529 in Wilson, D.E. and Reeder, D.M.. . 3rd ed. Baltimore: The Johns Hopkins University Press, 2 vols., 2142 pp.
- Wilson, D.E. and Reeder, D.M.. 2005. . 3rd ed. Baltimore: The Johns Hopkins University Press, 2 vols., 2142 pp.
Pteropodidae
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008a.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 26, 2010.
- Andriafidison, D, Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Rabearivelo, A., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008b.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 26, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008c.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 26, 2010.
Hipposideridae
- Andriafidison, D, Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008f.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008o.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008p.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008q.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Benda, P. and Vallo, P. 2009. . Folia Zoologica 58 :1–45.
- Goodman, S.M. and Ranivo, J. 2009. . Mammalia 73:47–55.
- Ranivo, J. and Goodman S.M. 2006. Révision taxinomique des Triaenops malgaches. Zoosystema 28:963–985.
- Samonds, K.E. 2007. . Acta Chiropterologica 9:39–65.
Emballonuridae
- Goodman, S.M., Cardiff, S.G., Ranivo, J., Russell, A.L. and Yoder, A.D. 2006a. . American Museum Novitates 3538:1–24.
- Goodman, S.M., Cardiff, S.G. and Ratrimomanarivo, F.H. 2008a. . Systematics and Biodiversity 6:283–292.
- Hutson, A.M., Racey, P., Ravino, J., Mickleburgh, S., Bergmans, W. and Fahr, J. 2008c.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008a.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008b.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Mickleburgh, S., Hutson, A.M., Racey, P.A., Cardiff, S. and Bergmans, W. 2008c.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
Nycteridae
- Hutson, T., Racey, P.A. and Ravino, J. 2008a.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
Myzopodidae
- Goodman, S.M., Rakotondraparany, F. and Kofoky, A. 2007. . Mammalian Biology 72:65–81.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008c.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008d.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
Molossidae
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008i.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008j.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008n.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2014.. IUCN Red List of Threatened Species. Version 2014.3. Downloaded May 28, 2015.
- Cotterill, F.P.D, Hutson, A.M., Racey, P.A. and Ravino, J. 2008.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Goodman, S.M. and Cardiff, S.G. 2004. A new species of Chaerephon from Madagascar with notes on other members of the family. Acta Chiropterologica 6:227–248.
- Goodman, S.M., Buccas, W., Naidoo, T., Ratrimomanarivo, F., Taylor, P.J. and Lamb, J. 2010c. . Zootaxa 2551:1–36.
- Jenkins, R.K.B., Racey, P.A., Ranivo, J., Ratimomanarivo, F., Mickleburgh, S., Hutson, A.M., Bergmans, W., Cotterill, F.P.D. and Fahr, J. 2008e.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Mickleburgh, S., Hutson, A.M., Racey, P.A., Ravino, J., Bergmans, W., Cotterill, F.P.D. and Gerlach, J. 2008b.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Ratrimomanarivo, F.H., Goodman, S.M., Stanley, W.T., Naidoo, T., Taylor, P.J. and Lamb, J. 2009. . Acta Chiropterologica 11:25–52.
Miniopteridae
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008g.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008h.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Goodman, S.M., Bradman, H.M., Maminirina, C.P., Ryan, K.E., Christidis, L. and Appleton, B. 2008b. A new species of Miniopterus from lowland southeastern Madagascar. Mammalian Biology 73:199–213.
- Goodman, S.M., Maminirina, C.P., Bradman, H.M., Christidis, L. and Appleton, B. 2009. The use of molecular phylogenetic and morphological tools to identify cryptic and paraphyletic species: Examples from the diminutive long-fingered bats on Madagascar. American Museum Novitates 3669:1–34.
- Goodman, S.M., Maminirina, C.P., Bradman, H.M., Christidis, L. and Appleton, B.R. 2010a. . Journal of Zoological Systematics and Evolutionary Research 48:75–86.
- Goodman, S.M., Ramasindrazana, B., Maminirina, C.P., Schoeman, M.C. and Appleton, B. 2011. . Zootaxa 2880:1–19.
- Jenkins, R.K.B. and Rakotoarivelo, A.R. 2008a.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B. and Rakotoarivelo, A.R. 2008b.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008f.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
Vespertilionidae
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008d.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008e.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008i.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D, Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008k.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Andriafidison, D., Cardiff, S.G., Goodman, S.M., Hutson, A.M., Jenkins, R.K.B., Kofoky, A.F., Racey, P.A., Ranivo, J., Ratrimomanarivo, F.H. and Razafimanahaka, H.J. 2008l.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Bates, P.J.J., Ratrimomanarivo, F.H., Harrison, D.L. and Goodman, S.M. 2006. . Acta Chiropterologica 8:299–324.
- Goodman S.M., Jenkins R.K.B. and Ratrimomanarivo F.H. 2005. A review of the genus Scotophilus on Madagascar, with the description of a new species. Zoosystema 27:867–882.
- Goodman, S.M., Ratrimomanarivo, F.H. and Randrianandrianina, F.H. 2006b. A new species of Scotophilus from western Madagascar. Acta Chiropterologica 8:21–37.
- Goodman, S.M., Taylor, P.J., Ratrimomanarivo, F. and Hoofer, S.R. 2012. . Zootaxa 3250:1–25.
- Hutson, A.M., Racey, P.A., Goodman, S. and Jacobs, D. 2008b.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jacobs, D., Cotterill, F.P.D., Taylor, P.J., Monadjem, A. and Griffin, M. 2008.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008g.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008h.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Jenkins, R.K.B., Rakotoarivelo, A.R., Ratrimomanarivo, F.H. and Cardiff, S.G. 2008i.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.
- Mickleburgh, S., Hutson, A.M. and Racey, P.A. 2008a.. IUCN Red List of Threatened Species. Version 2009.2. Downloaded May 27, 2010.