Fauna of Madagascar

The fauna of Madagascar is a part of the wildlife of Madagascar.

The history of the fauna of Madagascar in the context of plate tectonics and paleoclimate over the last 200 million years (Aepyornithidae arrived later than is indicated).
A good example of Malagasy convergent evolution is the fossa, a Malagasy carnivore that has evolved in appearance and behaviour to be so like a large cat that it was originally classified in Felidae, when it is in fact more closely related to the mongoose

Madagascar has been an isolated island for about 70 million years, breaking away from Africa around 165 million years ago, then from India nearly 100 million years later. This isolation led to the development of a unique endemic fauna.

Before humans arrived about 2,000 years ago, there were many large and unusual animals living there, descended from species that were originally present when Madagascar became an island, or from species that later crossed the sea to Madagascar. Ecological niches were often filled by animals with quite different histories from those on the African mainland, often leading to convergent evolution. A large proportion of these endemic Malagasy animals have died out since the arrival of humans, most particularly the megafauna.

Despite this, and massive deforestation, Madagascar is still home to an incredible array of wildlife, the vast majority of which is unique in the world.[1] Madagascar is a primary spot for ecotourism,[2] with more than fifty national parks and other protected reserves.

There are believed to have been only four colonization events of terrestrial mammals from mainland Africa. They brought to Madagascar the ancestors of its tenrecs, lemurs, carnivorans and nesomyine rodents. The other mammalian colonizations were those of the amphibious hippopotamuses (now extinct) and bats.

Tailless tenrec (Tenrec ecaudatus), the largest of the tenrecs.

Mammals
Main articles: List of mammals of Madagascar and List of lemur species

The lemurs are the best known of Madagascar's mammals.[3] They can only be found on Madagascar. In the absence of monkeys and other competitors, these primates have adapted to a wide range of habitats and diversified into numerous species. The tenrecs are another group of mammals characteristic of Madagascar. Most of the world's species of these small to medium-sized insectivores are found on the island. Rodents are poorly represented on the island with only a handful of native species, all belonging to the subfamily Nesomyinae. They include the large, endangered Malagasy giant rat. Due to their ability to fly, a greater variety of bats have reached the island. However, only about a third of these are endemic, a lower proportion than in other groups of land mammals. There are eight members of the Carnivora on the island, all now classified in the endemic family Eupleridae. The cat-like fossa is the largest of these.

Malagasy living mammals

Lemurs: As of 2008, there are officially 99 species and subspecies of lemur, 39 of which have been described by zoologists between 2000 and 2008.[4] Of the 51 species that were evaluated by the IUCN that year, 43 were categorized as threatened to some degree, with six species being classified as "Critically Endangered", the designation of highest risk.[5] The evaluations became considerably more pessimistic in 2012.[6]

Eupleridae: Primary among these malagasy carnivores is the fossa (Cryptoprocta ferox), an animal similar in appearance to a feline. Other Malagasy carnivores include the fanaloka (Fossa fossana), which, despite its scientific name, should not be confused with the fossa. Nor should it be confused with the very similarly named falanouc (Eupleres goudotii), also known as the Malagasy small-toothed civet. Five species of mongoose are also found in Madagascar, as is the small Indian civet, one of the carnivores believed to have been introduced to the island.

Tenrecidae: Three species of tenrec (the otter shrews) are found on the African mainland. The majority of tenrecs, around 30 species, are found in Madagascar. They have radiated into many different niches. For example, web-footed tenrecs (Limnogale mergulus) resemble river otters in appearance and behaviour. The lesser hedgehog tenrec (Echinops telfairi) resembles its namesake the hedgehog.

Rodents: About 30 species of indigenous rodent are known from Madagascar, three of which are now extinct. They are all members of the muroid subfamily Nesomyinae. These have also radiated into various niches, with vole-like forms, arboreal mice, fossorial varieties, and rabbit-like forms.

Bats: Approximately 69 species of bat are known from Madagascar, more than half of which are endemic to the island (see List of bats of Madagascar).

Malagasy extinct mammals

Plesiorycteropus (P. germainepetterae, P. madagascariensis): One or two species of mysterious aardvark-like tenrec relatives that may have gone extinct about 1000 years ago.

Giant lemurs: At least 17 species of lemur have become extinct since man arrived on Madagascar, all of which were larger than the surviving lemur species. They include the giant aye-aye, three to five times as heavy as the extant species. Megaladapis, which had three species, reached the size of orangutans. The eight sloth lemurs show evolutionary convergence with the South American sloths; Palaeopropithecus was chimpanzee-sized, while Archaeoindris grew to over 200 kg,[7] more massive than a silverback gorilla.

Giant fossa (Cryptoprocta spelea): The fossil record of Madagascar has yielded the remains of a recently extinct fossa which was about a quarter larger than the living species,[8] making it close to the size of an ocelot. This species was believed to have preyed upon the larger lemurs that inhabited Madagascar until the island was settled by man.

Malagasy hippopotamuses: There are thought to have been three species of dwarf or pygmy hippopotamus in Madagascar,[9] the last of which died out no earlier than 1,000 years ago, probably as a result of human settlement of the island.

Cuckoo-roller (Leptosomus discolor), the only member of a family unique to Madagascar and the nearby Comoros.

Birds
Main articles: List of birds of Madagascar and Endemic birds of Madagascar and western Indian Ocean islands

About 280 species of bird have been recorded on Madagascar and about 200 of these breed. Although these are relatively low numbers for a large tropical island, there is a high degree of endemism. Over 100 bird species are endemic and 49 of these are restricted-range endemics with a range of less than 50,000 km2. There are five bird families unique to Madagascar or shared only with the Comoro Islands: the mesites, the ground-rollers, the cuckoo-roller, the asities and the vangas. In addition, recent studies suggest that several songbirds should be grouped together in a new endemic family: the Malagasy warblers.

Elephant birds (e.g. Aepyornis maximus) were flightless ratites up to over three metres tall and half a tonne in weight. Genetic studies have revealed that their closest living relatives are the kiwi of New Zealand, rather than the nearby ostriches of Africa; i.e., their presence on the island appears to be due to long-range dispersal rather than vicariance.[10] They are thought to have become extinct within the last millennium, as a result of human activity. Other endemic birds that have disappeared since human settlement of the island include the Malagasy lapwing[11] and Malagasy shelduck.[12]

Panther chameleon (Furcifer pardalis), a large chameleon which has adapted to man-made habitats.

Reptiles
See also: List of reptiles in Madagascar

Relatively few families and genera of reptile have reached Madagascar but they have diversified into more than 260 species, with over 90% of these being endemic. The chameleons are very well represented with two-thirds of the world's species found there. The other lizard groups on Madagascar are the geckoes, skinks, girdle-tailed lizards and iguanids. Over 60 different snakes occur on the island; none of these are harmful to man. There are six land tortoises species, five endemics and one introduced species (Bell's hinge-back tortoise). Land based turtles include the African helmeted turtle, yellow-bellied mud turtle and East African black mud turtle. The shores are used for nesting by the loggerhead sea turtle, green sea turtle and hawksbill sea turtle while the olive ridley sea turtle forages here.[13] Madagascar's largest reptile is the Nile crocodile which has become very rare due to hunting for its skin. Endemic voay (V. robustus) became extinct.

Madagascar is a stronghold for a wide diversity of endemic species of chameleons and is considered the radiation point for day geckos. The two genera of iguanas present have a basal relationship to other iguanas and are believed to have been present on the island since it separated from Africa.[14] The Madagascar big-headed turtle, spider tortoise, flat-backed spider tortoise, radiated tortoise and ploughshare tortoise are endangered species.[13]

Amphibians
Mantella frogs have bright warning colours and are sometimes confused with the unrelated poison dart frogs of the Americas.[15]

There are more than 290 amphibian species in Madagascar with new species being found regularly.[16] Nearly all of these are endemic and the majority are restricted to primary forest. Examples of well-known amphibian species found in Madagascar include tomato frog and golden mantella frog.

Freshwater fish
The cichlid Ptychochromis insolitus was feared extinct in the wild (with only males in captivity) until rediscovered in 2013.[17]

Madagascar has a rich freshwater fish fauna with a very high rate of endemics. The full diversity is unclear, as new species are being described regularly and species may have disappeared before even being discovered; estimates suggest the island has between 135 and 150 native fish species that are restricted to freshwater (this number grows significantly if including widespread euryhaline species).[18] Among these are two families entirely restricted to Madagascar, the anchariid catfish and the Madagascar rainbowfish.[19][20] Additionally, several genera are endemic to Madagascar, including cichlids (Katria, Oxylapia, Paratilapia, Paretroplus, Ptychochromis and Ptychochromoides), round herrings (Sauvagella and Spratellomorpha), Old World silverside (Teramulus), sleeper gobies (Ratsirakia and the troglobitic Typhleotris), and aplocheilid killifish (Pachypanchax – strictly speaking near-endemic, as there is a single species from the Seychelles).[18][21] There are endemic species from other genera (notably Ambassis, Arius, Eleotris, Glossogobius, Kuhlia, Mesopristes, Ophiocara, Pantanodon and Sicyopterus),[21] but these also have other species found elsewhere.

The geographic history of Madagascar (long isolated, but part of Gondwana) has resulted in unusual biogeographic patterns, comparable to those seen in other animal and plant groups of the island. For example, the closest relatives of the Madagascar rainbowfish are the "true" rainbowfish of New Guinea and Australia,[22] while the killifish Pachypanchax and the cichlids Paretroplus have their closest relatives in South Asia.[23]

The freshwater fish fauna in Madagascar has declined drastically due to habitat loss (pollution, siltation following deforestation, damming and alike), overfishing and introduced species (tilapia, Nile perch, African arowana, blotched snakehead, green swordtail and many other), with some suggesting that only a remnant of the natives can be saved.[18] Among species already believed to be extinct are Pantanodon madagascariensis and Ptychochromis onilahy.[24][25]

Invertebrates

Oligochaeta

Thirty-eight species of earthworms are recorded in seven families, i.e. the Megascolecidae, Kynotidae (giant earthworms; endemic family with a single genus), Acanthodrilidae, Eudrilidae, Ocneodrilidae, Octochaetidae and Glossoscolecidae. Among the species reported 59% are endemic.[26]

Insects
Comet moth (Argema mittrei), one of the world's largest moths.
Main article: List of butterflies of Madagascar

Madagascar is home to huge variety of insects, the majority of which are endemic. Thousands of species are present in some groups such as the beetles and moths. There are approximately 100,000 species of insects and counting in Madagascar. Distinctive species include the long-necked giraffe weevil, the huge comet moth and the butterfly-like Madagascan sunset moth. About 80 species of stick insect occur; the Achrioptera species are large and colourful while others are small and very well-camouflaged. Many of the island's praying mantises are also well-camouflaged, mimicking dead leaves or bark. There are over 100 cockroaches, including the large Madagascar hissing cockroach.

The mosquitoes of Madagascar include 235 species, among which 138 (59%) are endemic and 64 (27%) have a known medical or veterinary interest because they can transmit diseases.[27]

Beetles: Various species of beetles. For example, various tiger beetles (Cicindelidae): 109 species from genus Pogonostoma,[28] 65 species from genus Physodeutera.[29]

Madagascar is home to the smallest bee in the World, Liotrigona bitika (Brooks & Michener 1988) with a body length of less than 2 mm, and to several other species of Meliponini.[30]

The freshwater crayfish Astacoides appears to be a Gondwanan relict.[31]

Spiders

Darwin's bark spider was discovered in 2009. It is famous for making the strongest and largest spider webs (ranging from 900 - 28000 square centimetres). The silk produced by this spider is twice as strong as other common spider silk.[32]

Molluscs
Main article: List of non-marine molluscs of Madagascar

Freshwater crustaceans

Madagascar is home to several endemic freshwater crustaceans, including 17 species of potamonautid crabs (Boreas, Foza, Hydrothelphusa, Madagapotamon, Malagasya, Marojejy and Skelosophusa), 7 species of parastacid crayfish (Astacoides), and atyid shrimp.[31][33][34][35]

Marine life
Robust ghostpipefish (Solenostomus cyanopterus) off Nosy Be.
Omura's whale off Nosy Be.

The seas around Madagascar host a high diversity of wildlife, including invertebrates.[36] There is a still a large area of mangrove swamp around the coast, particularly in the west, and there are extensive coral reefs around the island. Thousands of fish species occur, including the rare coelacanth. Four species of sea turtle breed on the beaches, but many are taken for food by local people. For whales, humpback whales breed off the south-western coasts such as at Île Sainte-Marie, Omura's whales notably around Nosy Be,[37] pygmy blue whales on the Madagascar Plateau,[38] and southern right whales slowly increasing along Malagasy coasts.[39]

Malagasy faunal names

Fish

Selected Malagasy fish names:[40][41][42][43]

  • gogo: Arius spp.
  • mahalogy: Caesio spp.
  • tsenaby: Cheilio inermis
  • lemeleme: Coris formosa
  • lanora: Elagatis bipinnulata
  • alovo: Epinephelus fasciatus, Epinephelus coeruleopunctatus
  • angoa, vohy: Euthynnus affinis
  • tserakantsiva: Fistularia petimba
  • ambariake: Gerres oyena
  • soavahindriaka: Hippocampus cf. borboniensis
  • ndwaro: Istiophorus platypterus
  • fiambondis (?): Labroides dimidiatus
  • tapaporoha: Lethrinus harak
  • ambitsy: Lethrinus nebulosus
  • romanjia: Lethrinus variegatus
  • tsivaravana: Lutjanus bohar
  • amposama: Lutjanus fulviflamma
  • fiamasika: Lutjanus kasmira
  • ndwaro: Makaira indica
  • kipela, talatala: Monodactylus argenteus
  • amgelika: Monotaxis grandoculis
  • dangiara, filao papango: Platax orbicularis
  • fiandolo, lafalovo: Plotosus lineatus
  • bodoloha: Pseudanthias squamipinnis
  • lemilemy: Pseudochromis dutoiti
  • sabonto: Rastrelliger kanagurta
  • soroboa: Rhynchobatus djiddensis
  • maro taobana: Sardinella albella
  • lamatra, talafeta: Scomberomorus commerson
  • lamatra: Scomberomorus plurilineatus
  • akihoviko: Sphyrna spp.
  • drihy, tsarabora: Terapon jarbua
  • lemeleme: Thalassoma hebraicum
  • sampramale, vivano: Variola louti
  • fiam'akoho: Zanclus cornutus

Mammals
Main article: List of Malagasy mammal common names

Extinct megafauna
Birds
Lemurs
Further information: Subfossil lemur
Others[44]

Habitat loss

Most of Madagascar's habitats are in danger; they are being threatened by today's demands and growing needs. Environmental damage has been especially severe for humid forests.[45] Fauna and Flora International, abbreviated as FFI is one of the organizations that supports Madagascar's habitats.[46]

  • As a part of conservation efforts, the Wildlife Conservation Society has recently opened a Madagascar! exhibit at the Bronx Zoo.
  • The 2005 animated film Madagascar featured a number of animals of Madagascar as cartoon characters.
gollark: No, if it is not GTech-recognized.
gollark: We do not accept those.
gollark: In any case, Θ-3 is different from Γ-3 thanks to GTech's "extreme management" structure.
gollark: Well, you need GTech-recognized clearance, obviously...
gollark: Where did you get Θ-3 clearance from‽

See also

References
  1. Benstead, Jonathan P.; Goodman, Steven D. (2003). The natural history of Madagascar. Chicago: University of Chicago Press. ISBN 0-226-30306-3.
  2. Schuurman, Derek; Nick Garbutt; Bradt, Hilary (2001). Madagascar Wildlife: A Visitor's Guide (2nd ed.). Bradt Travel Guides. ISBN 1-84162-029-7.
  3. Harris, Bruce. "Nature of Madagascar". mada.moreoless.net.au. Retrieved May 2004. Check date values in: |accessdate= (help)
  4. Mittermeier, R., Ganzhorn, J., Konstant, W., Glander, K., Tattersall, I., Groves, C., Rylands, A., Hapke, A., Ratsimbazafy, J., Mayor, M., Louis, E., Rumpler, Y., Schwitzer, C. & Rasoloarison, R. (December 2008). "Lemur Diversity in Madagascar". International Journal of Primatology. 29 (6): 1607–1656. doi:10.1007/s10764-008-9317-y. hdl:10161/6237.CS1 maint: multiple names: authors list (link)
  5. Jane Thornback; Harcourt, Caroline (1990). Lemurs of Madagascar and the Comoros: The IUCN Red Data Book. World Conservation Union. ISBN 2-88032-957-4.
  6. Black, Richard (13 July 2012). "Lemurs sliding toward extinction". BBC News. Archived from the original on 27 August 2012. Retrieved 26 August 2012.
  7. Jungers WL, Godfrey LR, Simons EL, Chatrath PS (October 1997). "Phalangeal curvature and positional behavior in extinct sloth lemurs (Primates, Palaeopropithecidae)". Proc. Natl. Acad. Sci. U.S.A. 94 (22): 11998–2001. Bibcode:1997PNAS...9411998J. doi:10.1073/pnas.94.22.11998. PMC 23681. PMID 11038588.
  8. Goodman SM, Rasoloarison RM, Ganzhorn JU (2004). "On the specific identification of subfossil Cryptoprocta (Mammalia, Carnivora) from Madagascar" (PDF). Zoosystema. 26 (1): 129–143. Archived from the original (PDF) on 2006-11-27.
  9. Nick Garbutt (1999). Mammals of Madagascar. New Haven, Conn: Yale University Press. ISBN 0-300-07751-3.
  10. Mitchell, K. J.; Llamas, B.; Soubrier, J.; Rawlence, N. J.; Worthy, T. H.; Wood, J.; Lee, M. S. Y.; Cooper, A. (2014-05-23). "Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution" (PDF). Science. 344 (6186): 898–900. Bibcode:2014Sci...344..898M. doi:10.1126/science.1251981. hdl:2328/35953. PMID 24855267.CS1 maint: ref=harv (link)
  11. Hume, Julian P; Walters, Michael (2012). Extinct Birds. Poyser Monographs. A & C Black. p. 125. ISBN 9781408158623.
  12. Goodman, SM (1999). In: Adams, N.J.; Slotow, R.H. Proc. 22 Int. Ornithol. Congr; Durban (eds.). Holocene bird subfossils from the sites of Ampasambazimba, Antsirabe and Ampoza, Madagascar: Changes in the avifauna of south central Madagascar over the past few millennia. Johannesburg: BirdLife South Africa. pp. 3071–3083. Archived from the original on 2014-05-19.
  13. Rhodin, Anders G.J.; van Dijk, Peter Paul; Inverson, John B.; Shaffer, H. Bradley (2010-12-14). "Turtles of the World 2010 Update: Annotated Checklist of Taxonomy, Synonymy, Distribution and Conservation Status" (PDF). Archived from the original (PDF) on 2010-12-15. Retrieved 2010-12-15.
  14. Okajima, Y.; Kumazawa, Y. (2009-07-15). "Mitogenomic perspectives into iguanid phylogeny and biogeography: Gondwanan vicariance for the origin of Madagascan oplurines". Gene. Elsevier. 441 (1–2): 28–35. doi:10.1016/j.gene.2008.06.011. PMID 18598742.
  15. Staniszewski, M. S: The Mantella FAQ. Retrieved 29 June 2014
  16. http://amphibiaweb.org/. Retrieved 28 June 2014. Missing or empty |title= (help)
  17. Mongabay (20 December 2013). Conservation Hail Mary works: Mate for near-extinct fish found!. Retrieved 29 June 2014.
  18. IUCN Freshwater Fish Specialist Group: Madagascar. Archived 2014-11-29 at the Wayback Machine Retrieved 29 June 2014.
  19. Ng, and Sparks (2005). Revision of the endemic Malagasy catfish family Anchariidae (Teleostei: Siluriformes), with descriptions of a new genus and three new species. Ichthyol. Explor. Freshwaters 16(4): 303–323.
  20. Sparks; and Smith (2004). Phylogeny and biogeography of the Malagasy and Australasian rainbowfishes (Teleostei: Melanotaenioidei): Gondwanan vicariance and evolution in freshwater. Molecular Phylogenetics and Evolution 33(3): 719–734
  21. FishBase: List of Freshwater Fishes for Madagascar. Retrieved 29 June 2014.
  22. Sparks; and Smith (2004). 'Phylogeny and biogeography of the Malagasy and Australasian rainbowfishes (Teleostei: Melanotaenioidei): Gondwanan vicariance and evolution in freshwater. Molecular Phylogenetics and Evolution 33(3): 719–734.
  23. Sparks (2004). Molecular phylogeny and biogeography of the Malagasy and South Asian cichlids (Teleostei: Perciformes: Cichlidae). Molecular Phylogenetics and Evolution 30(3): 599–614.
  24. Loiselle, P.; Participants of the CBSG; ANGAP CAMP "Faune de Madagascar" workshop (2004). "Pantanodon madagascariensis". IUCN Red List of Threatened Species. Mantasoa, Madagascar. 2004.CS1 maint: ref=harv (link)
  25. Loiselle, P. & participants of the CBSG/ANGAP CAMP "Faune de Madagascar" workshop, Mantasoa, Madagascar 2001 (2004). "Ptychochromis onilahy". IUCN Red List of Threatened Species. 2004.CS1 maint: multiple names: authors list (link) CS1 maint: ref=harv (link)
  26. Razafindrakoto, M., Csuzdi, Cs. & Blanchart, E. 2011. New and little known giant earthworms from Madagascar (Oligochaeta: Kynotidae). African Invertebrates 52 (2): 285-294. Archived 2011-10-21 at the Wayback Machine
  27. Tantely, Michaël Luciano; Le Goff, Gilbert; Boyer, Sébastien; Fontenille, Didier (2016). "An updated checklist of mosquito species (Diptera: Culicidae) from Madagascar". Parasite. 23: 20. doi:10.1051/parasite/2016018. ISSN 1776-1042. PMC 4840257. PMID 27101839.
  28. Moravec Jiří (2007). "A monograph of the genus Pogonostoma" (30 colour plates, 499 pp.). Tiger beetles of Madagascar. 1. Zlín: Nakladatelství Kabourek. ISBN 978-80-86447-13-1.
  29. Moravec Jiří (2007). "A monograph of the genus Physodeutera" (21 colour plates, 290 pp.). Tiger beetles of Madagascar. 2. Zlín: Nakladatelství Kabourek. ISBN 978-80-86447-13-1.
  30. Koch, H (2010). "Combining morphology and DNA barcoding resolves the taxonomy of Western Malagasy Liotrigona Moure, 1961" (PDF). African Invertebrates. 51 (2): 413–421. doi:10.5733/afin.051.0210. Archived from the original (PDF) on 2011-07-06.
  31. Jones; Andriahajaina; Hockley; Crandall; and Roavohangimalala (2007). The ecology and conservation status of Madagascar’s endemic freshwater crayfish (Parastacidae; Astacoides). Freshwater Biology 52(9): 1820–1833.
  32. "TOP 10 RARE MADAGASCAR ANIMALS". Listverse Info. Retrieved 2016-03-19.
  33. Cumberlidge (2008). Insular species of Afrotropical freshwater crabs (Crustacea: Decapoda: Brachyura: Potamonautidae and Potamidae) with special reference to Madagascar and the Seychelles. Contributions to Zoology 77(2): 71–81.
  34. Hogan (26 August 2008). Biological diversity in Madagascar and the Indian Ocean Islands. Conservation International, EoEarth. Retrieved 29 June 2014.
  35. Cumberlidge, Klaus, Meyer & Koppin (2015). New collections of freshwater crabs from northern Madagascar, with the description of a new species of Foza Reed & Cumberlidge, 2006 (Brachyura, Potamonautidae), and comments on their conservation status. European Journal of Taxonomy 109: 1-15.
  36. Dell'Angelo, B., Prelle, G., Sosso, M. & Bonfitto, A. 2011. Intertidal chitons (Mollusca: Polyplacophora) from southern Madagascar. African Invertebrates 52 (1): 21-37. Archived 2011-09-03 at the Wayback Machine
  37. Jefferson, Thomas; Marc A. Webber & Robert L. Pitman (2015). Marine Mammals of the World: A Comprehensive Guide to their Identification. London: Academic.
  38. WCS Madagascar - Blue Whale
  39. "Des baleines franches australes dans la Baie de Diego Suarez" [Southern right whales in the Diego-Suarez Bay]. La Tribune de Diego (in French). 13 October 2011.
  40. Blench, Roger. Notes on Malagasy fish names.
  41. Bauchot, M-L. & G. Bianchi 1984. Guide de poisons commerciaux de Madagascar (espèces marines et d’eaux saumâtres). Rome : FAO.
  42. Pellegrin, M. Jacques 1933. Les poissons des eaux douces de Madagascar et des iles voisines (Comores, Seychelles, Mascareignes). Tananarive : G. Pitot et Cie.
  43. Richmond, M.D. 2002. A field guide to the seashores of Eastern Africa and the Western Indian Ocean islands (2nd ed). Milan: SIDA/SAREC - University of Dar es Salaam.
  44. More Madagascan mystery mammals
  45. "MADAGASCAR". 1250 24th Street, N.W. Washington, DC 20037. World Wildlife Fund. Retrieved 2013. Check date values in: |accessdate= (help)
  46. "Madagascar". Fauna & Flora International. Retrieved 2013. Check date values in: |accessdate= (help)
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