The Madagascan Realm

The Madagascan Realm includes Madagascar along with the Mascarene, Comoros, and Seychelles island groups of the south-western Indian Ocean and a few smaller islands. Biogeographically speaking it has long been included with Africa, although the growing weight of evidence argues for separation. Most are essentially old pieces of the ancient supercontinent of Gondwana that broke away from Africa millions of years ago, although some (such as the Comoros and Mascarenes) are volcanic islands that formed much more recently.

 

Species and subspecies

The lesser yellow bat (Scotophilus borbonicus) was considered common on the island of Réunion in the Mascarenes during the early nineteenth century but soon disappeared. A single specimen was collected from southern Madagascar in 1868 (former Toliara province), but the species has not been recorded since. It is most likely extinct, a victim of habitat destruction.

The Madagascar harrier (Circus macrosceles) is a bird of prey found across the whole of Madagascar and on Grande Comore, Mohéli, and Anjouan in the Comoro Islands, where it is primarily associated with wetlands. Despite its wide distribution the species occurs at extremely low densities, with the total population estimated at less than 250.

The Madagascar sacred ibis (Threskiornis bernieri) is confined to coastal areas along the length of western Madagascar as well as in the Seychelles, where the total population is thought to be between 2000 and 3000.

Humblot’s heron (Ardea humbloti) is largely restricted to western coastal Madagascar but is also recorded as a vagrant from areas further east (i.e. the Central Highlands, especially Lake Alaotra), the Anorontany Archipelago, and the Comoros Islands. The total population is thought to be no more than 1500. The Madagascar pond heron (A. idae) is widespread in freshwater wetland areas across east-central Africa, Madagascar, and the Seychelles, Comoros, and Mascarene Islands. It is everywhere rare, however, with a total population estimated at between 2000 and 6000.

The Madagascar pratincole (Glareola ocularis) is a type of small, migratory wading bird found widely over centraleastern Africa, Madagascar, and the islands of the western Indian Ocean. Nevertheless, the total population is believed to be small and threatened by the loss and degradation of wetlands, particularly its breeding areas on the eastern coast of Madagascar.

Madagascar

The island of Madagascar is the world’s fourth largest and is, in many ways, a mini-continent. The seas around it are, in general, deep, with the wide Mozambique Channel which separates it from the East African coast reaching a depth of 3292 m. The prehistoric breakup of the supercontinent Gondwana separated the Madagascar–Antarctic–India landmass from the Africa–South America landmass around 135 million years ago. Madagascar later split from India about 88 million years ago, allowing plants and animals on the island to henceforth evolve in relative isolation. Biogeographically it is almost as peculiar as Australia, which has been isolated from Asia for more than 50 million years. In both cases evolution has followed a unique pattern, with prosimians in many ways taking over the role that marsupials serve in Australia. While much has been lost, much remains. Madagascar continues to be a major biodiversity hotspot, with over 90 per cent of its fauna and a similar percentage of its flora found nowhere else on Earth.

Madagascar is tropical and chiefly mountainous, with a diverse range of habitats. A narrow and steep escarpment runs along the length of the eastern coast, which contains much of the island’s remaining lowland rainforest. To the west of this ridge lies a plateau in the centre of the island ranging in altitude from 750 to 1500 m. These central highlands are the most densely populated part of the island and are characterized by terraced, rice-growing valleys lying between grassy hills and patches of the subhumid forests that formerly covered the entire region. To the west of the highlands, the increasingly arid terrain gradually slopes down to the Mozambique Channel and to mangrove swamps along the coast. The western and southern sides are home to dry deciduous forests, spiny thickets, and xeric shrublands. Madagascar’s highest peaks rise from three prominent highland massifs, with Maromokotro (2876 m) in the Tsaratanana Massif the island’s highest point. Many of its endemic species are very specialized in terms of certain bioregions.

Madagascar’s earliest human settlers found a rich prehistoric world very different from the one that we see today, one populated by an extraordinary megafauna including at least 17 species of lemur larger than any living species, some the size of a gorilla. Other highlights were at least two types of hippopotamus, gigantic crocodilians and tortoises, and an extraordinary burrowing creature that must have looked like a cross between an aardvark and a cow. However, it is the various species of enormous ratites known as elephant birds that were the most spectacular. The largest birds that ever lived, one of them (Vorombe titan) stood fully 3 m high and weighed up to 730 kg. How much responsibility these early people had in wiping out these creatures is still open to debate, although it is clear that at least some may have survived up into historic times. Madagascar’s megafaunal extinctions were among the most severe for any continent or large island, with all endemic wildlife over 10 kg (at least two dozen species) disappearing even before the arrival of Europeans.

The aye-aye (Daubentonia madagascariensis) is perhaps the most remarkable of all surviving lemurs. The world’s largest nocturnal primate, it resembles a witch’s cat in general appearance and is notable for its enormous ears, rodent-like, perpetually growing teeth and narrow middle fingers. All of these features are special adaptations for its unusual method of finding tree-boring insect larvae. Individuals tap on tree limbs while listening intently for the grubs burrowing within, then gnaw at the bark with their forward-slanting incisors in order to create a small hole into which they then insert their wire-like finger in order to extract it. A cryptic species, the aye-aye was rarely encountered after its discovery in the late eighteenth century, and by the 1930s was widely thought to be on the verge of extinction. It was rediscovered in 1957, however, and in 1966 nine individuals were transported to the island of Nosy Mangabe, in the Bay of Antongil off the north-eastern coast of Madagascar, in what was then considered a last-ditch effort to save it. By the 1980s sightings on the mainland became more frequent, and in the following decade a successful captive breeding programme was undertaken by a number of zoos around the world. While still rare and highly localized, the ayeaye is now known to be one of the most widespread of all lemurs, being found in both humid and dry forests across eastern, western, and northern Madagascar. Sadly, in many areas the species is shot on sight by local people in the erroneous belief that it is a threat to their crops, or because it is considered to be a harbinger of evil.

The indri (Indri indri), the largest living lemur, was at the beginning of the nineteenth century still so common in eastern and north-eastern Madagascar that a traveller reported that no one could journey from Tamatave to Antanarivo without often hearing their eerie, whale-like calls as they passed through the great forests. The species declined along with the latter, but is still found widely, if very locally, from Anjanaharibe-Sud and Antohaka Lava (former Antsiranana province) south to the Anosibe An-ala Classified Forest (former Toamasina province). It is everywhere under threat, however, from habitat destruction and illegal hunting, and has never been kept in captivity apart from a brief period in Paris in 1939.

Several species of sifaka (Propithecus) are found throughout the forested areas of Madagascar, where they are everywhere threatened by habitat destruction and, increasingly, subsistence hunting. Perrier’s sifaka (P. perrieri) is found in both dry and humid forest fragments within a very small area in northernmost Madagascar (former Antsiranana province), where the total population is thought to number less than 500. The silky sifaka (P. candidus), one of the rarest and most threatened of all sifakas, has a very restricted range in northeastern Madagascar (former Antsiranana province), where it lives in both lower- and upper-elevation rainforest. The total population is thought to be less than 250. The diademed sifaka (P. diadema), second only to the indri (Indri indri) in size, occurs widely but patchily throughout the rainforests of eastern and north-eastern Madagascar (former Toamasina province) from the Mangoro and Onive rivers north to the Mananara River. Milne-Edwards’ sifaka (P. edwardsi) is confined to a small area of middle- and upper-elevation rainforest in central-eastern Madagascar (former Fianarantsoa province). Tattersall’s sifaka (P. tattersalli) is confined to a small area of north-eastern coastal Madagascar (former Antsiranana province) between the Loky River in the north and the Manambato in the south. The range centres on the town of Daraina, and consists of human-altered savanna, dry scrub, agricultural land, gallery forests, and forest fragments. In recent years gold-mining activities in the region have posed an additional threat. Coquerel’s sifaka (P. coquereli) is found patchily in the lowland dry forests as well as coastal mangroves of north-western Madagascar (former Mahajanga province) to the north and east of the Betsiboka River. The crowned sifaka (P. coronatus) is found over a relatively wide area of northwestern and central-western Madagascar (former Mahajanga and Antananarivo provinces) from the Mahavavy River to the Betsiboka River. Von der Decken’s sifaka (P. deckenii) occurs patchily in north-western and central-western Madagascar (former Mahajanga and Toliara provinces) between the Mahavavy and Manambolo rivers. Verreaux’s sifaka (P. verreauxi) occurs in a variety of habitats in southern and southwestern Madagascar (former Toliara province) including lowland and montane dry deciduous, spiny thicket, and even some lowland rainforest. It is, however, most associated with spiny woodlands, where they can be seen leaping recklessly between thorny trunks and bounding along the ground. Some hunting continues, but the main threat is habitat loss due to slash-andburn agriculture and charcoal and fuelwood production.

The Sambirano woolly lemur (Avahi unicolor) is confined to two small, disjunct areas of the Sambirano region of northwestern Madagascar (former Antsiranana and Mahajanga provinces). Peyrieras’ woolly lemur (A. peyrierasi) is confined to a small area of lowland and montane rainforest in southeastern Madagascar (former Fianarantsoa province). Gmelin’s woolly lemur (A. laniger) remains widespread in eastern and north-eastern Madagascar, although all species are threatened by loss of habitat and subsistence hunting.

The ruffed lemurs (Varecia) are large and garishly coloured. Three subspecies of pied ruffed lemur (V. variegata) are found discontinuously throughout the eastern rainforests. The black and white ruffed lemur (V. v. variegata) occurs in eastern Madagascar (former Toamasina and Fianarantsoa provinces) south of the Anove River from about Ambatovaky south to Betampona and Zahamena National Park. The whitebelted ruffed lemur (V. v. subcincta) occurs in north-eastern Madagascar (former Toamasina province) from the Antainambalana River south to the Anove River, including parts of Makira, Mananara-Nord, Atialanankorendrina, and Marotandrano. This subspecies was additionally introduced to the island of Nosy Mangabe in the Bay of Antongil in the 1930s and still occurs there. Hill’s black and white ruffed lemur (V. v. editorum) is only known with certainty from southeastern Madagascar (former Fianarantsoa province) from Mantadia southwards to the Manombo Special Reserve. The red ruffed lemur (V. rubra) is confined to the Masoala Peninsula and the region immediately north of the Bay of Antongil in north-eastern Madagascar (former Antsiranana province). All of these forms are seriously threatened by habitat destruction and hunting.

The ring-tailed lemur (Lemur catta), an iconic symbol of Madagascar, remains relatively widespread in the dry forests and bush of the south and south-west (former Toliara and Fianarantsoa provinces). Nevertheless, its range is highly fragmented and population densities are low. Habitat destruction and hunting are the chief threats.

Several species of so-called true lemur (Eulemur) are found throughout Madagascar, where they are threatened by habitat destruction, hunting, and capture for use as pets. The crowned lemur (E. coronatus) inhabits extreme northern Madagascar (former Antsaranana province), where it prefers semideciduous dry lowland and mid-altitude forest but may be found in practically all forest types, including high-altitude moist forest, wooded savanna, and even agricultural areas. The mongoose lemur (E. mongoz) is found in north-western Madagascar (former Mahajanga province) in the region of Ambato-Boéni and Ankarafantsika, with introduced populations also occurring on the islands of Mohéli, Anjouan, and Grande Comore in the Comoros Islands. Sanford’s lemur (E. sanfordi) has a restricted range in northern Madagascar (former Antsiranana province), with the southerly limit being the Manambato River. The range is centred on Ankarana, Anamalerana, and Montagne d’Ambre, with a disjunct population in Daraina to the south-east. It is found from sea level to 1400 m and thus inhabits both dry and subhumid forest, but seems to be more common and to occur at higher densities in the latter. The red-bellied lemur (E. rubriventer) is found over a wide area from the Tsaratanana Massif in northern Madagascar south along the eastern rainforests (excluding the Masoala Peninsula) to the Pic d’Ivobe and the Manampatrana River, although at one time it ranged further south. The white-fronted lemur (E. albifrons) is found throughout most of the remaining rainforest in north-eastern Madagascar (former Antsiranana and Toamasina provinces), from the Bemarivo River near Sambava south to the region of Mananara Nord, including the Masoala Peninsula. There is an isolated population in the Betampona Nature Reserve further south, and the species has also been introduced to Nosy Mangabe. Its distribution south of Mananara remains to be clarified as there is significant hybridization with the brown lemur (E. fulvus) over a wide area. The white-collared lemur (E. cinereiceps) is largely confined to a thin strip of lowland rainforest running from the Andringitra Massif south to the Mananara River in south-eastern Madagascar (former Fianarantsoa province). Isolated populations also occur in the Manombo Special Reserve and in the Mahabo Forest south of Farafangana, and possibly at Vohipaho, south of the Mananara River. The rufous lemur (E. rufus) occurs in north-western Madagascar (former Mahajanga province) from the Betsiboka River south to the Tsiribihina. The red-fronted lemur (E. rufifrons) occurs disjunctly in south-eastern and south-western Madagascar (former Toliara, Fianarantsoa, and Toamasina provinces). The red-collared lemur (E. collaris) is found in south-eastern Madagascar (former Toliara and Fianarantsoa provinces) roughly from Tolagnaro north to the Mananara River, with the western limits of the range being the forests of the Kalambatritra region. The black lemur (E. macaco) occurs in moist forests of the Sambirano region of north-eastern Madagascar (former Antsiranana and Mahajanga provinces) and on the islands of Nosy Be, Nosy Komba, and Nosy Tanikely. Sclater’s lemur (E. flavifrons) is confined to a small area in north-eastern Madagascar (former Mahajanga province) consisting of the Sahamalaza Peninsula along with a narrow stretch of forest on the adjacent mainland. The brown lemur (E. fulvus) is found widely but disjunctly in northern and eastern Madagascar, with an introduced population in the Comoros Islands.

The greater bamboo lemur (Prolemur simus) is one of the world’s most critically endangered primates. Subfossil remains confirm that it was once widespread across northern, central and eastern Madagascar, but by the mid-twentieth century it was thought to be extinct, a victim of habitat destruction and extreme dietary specialization. A remnant population was discovered in 1986, however, and since then surveys in southern and central-eastern Madagascar have discovered around 500 individuals living in 11 subpopulations. Unfortunately, all live in areas of severely degraded and unprotected habitat.

Several bamboo lemur species and subspecies of the genus Hapalemur are threatened by habitat destruction, subsistence hunting, and capture for use as pets. The golden bamboo lemur (H. aureus) was first discovered in 1986 in what is now Ranomafana National Park, in central-east Madagascar (former Fianarantsoa province). It is now known to also occur in Andringitra National Park and in a forest corridor connecting the two, as well as in the region of Betsakafandrika. The northern bamboo lemur (H. occidentalis) is found in a number of discontinuous pockets in northern Madagascar (former Antsiranana, Mahajanga, and Toamasina provinces). It is known (at least formerly) from the forests of Ankarana and Analamerana in the far north and possibly on the Ankarana Massif as well, as well as from the Ampasindava Peninsula, and the Sambirano and Sahamalaza regions in the north-west; the Tsiombikibo, Baie de Baly; Tsingy de Namaroka and Bongolava regions in the central-west; Masoala, Maroansetra, and Ile Roger (Aye aye Island) in the north-east; and as far south as Zahamena and Marovohangy near Lake Alaotra. Three subspecies of grey bamboo lemur (H. griseus) are found disjunctly in eastern and western Madagascar. The Mantadia grey bamboo lemur (H. g. griseus) is restricted to east-central Madagascar (former Toamasina province), roughly from the Onibe River south to the Onive River. The Beanamalao bamboo lemur (H. g. gilberti) is known from a small area of central-eastern Madagascar (former Toamasina province) between the Onive and Nosivolo rivers. The Ranomafana bamboo lemur (H. g. ranomafanensis) occurs in two widely separated populations in east-central and west-central Madagascar. In the east it is known from the forests to the south of the Mangoro and Onive rivers in former Fianarantsoa province, while in the west it has been reported from the forests of Tsingy de Bemaraha, probably as far as the Betsiboka River in former Mahajanga province. The southern bamboo lemur (H. meridionalis) is found in south-eastern Madagascar (former Toliara and Fianarantsoa provinces), roughly from Andohahela north to the Mananara River.

The Sahamalaza sportive lemur (Lepilemur sahamalazensis) is restricted to the three remaining humid and dry forest blocks on the Sahamalaza Peninsula in north-eastern Madagascar (former Mahajanga province). Only some of its range lies within protected areas. The Daraina sportive lemur (L. milanoii) and the Ankarana sportive lemur (L. ankaranensis) are both confined to forest fragments in far northern Madagascar (former Antsiranana province). Seal’s sportive lemur (L. seali) is confined to a small area of north-eastern Madagascar (former Antsiranana province). The weasel sportive lemur (L. mustelinus) is found disjunctly in north-eastern coastal Madagascar (former Toamasina province). All are threatened by habitat destruction and hunting.

The Montagne d’Ambre fork-marked lemur (Phaner electromontis) is found sporadically in the regions of Montagne d’Ambre and Ankarana in far northern Madagascar (former Antsaranana province), with a further population at Daraina likely representing a distinct species. The Masoala forkmarked lemur (P. furcifer) is found over a relatively wide area of north-eastern Madagascar (former Antsiranana and Toamasina provinces). Both are threatened by loss of habitat.

Three species of dwarf lemur (Cheirogaleus) are threatened by habitat destruction and hunting. The Ankarana dwarf lemur (C. shethi) is found patchily in forest and shrubby areas of far northern Madagascar (former Antsiranana province). Crossley’s dwarf lemur (C. crossleyi) is found disjunctly in eastern and north-eastern Madagascar (former Antsiranana and Toamasina provinces). The Lavasoa dwarf lemur (C. lavasoensis) is known disjunctly from forest patches in the Lavasoa–Ambatotsirongorongo Mountains and from the Kalambatritra (Sahalava) Forest in south-eastern Madagascar (former Toliara and Fianarantsoa provinces).

The hairy-eared dwarf lemur (Allocebus trichotis) is a tiny, enigmatic species. Long known from only a few museum specimens collected during the late nineteenth century, it was not seen again until 1966. It was thereafter feared to be extinct until rediscovered in 1989 in lowland rainforest in northeastern Madagascar. Individuals have since been found living in highland rainforest at Andasibe in eastern Madagascar and in other localities as well, indicating a more widespread, if patchy, distribution.

The northern giant mouse lemur (Mirza zaza) is confined to an area of north-eastern Madagascar centred on the Ampasindava Peninsula (former Antsiranana province).

Several species of mouse lemur (Microcebus) are threatened by loss of habitat. The Tavaratra mouse lemur (M. tavaratra) and Arnhold’s mouse lemur (M. arnholdi) are both found patchily in far northern Madagascar (former Antsiranana province). Mittermeier’s mouse lemur (M. mittermeieri) is found patchily in both lowland and highland rainforest in northeastern Madagascar (former Antsiranana province). The Sambirano mouse lemur (M. sambiranensis) is known only from the few localities in north-eastern Madagascar (former Antsiratana and Mahajanga provinces). Margot Marsh’s mouse lemur (M. margotmarshae) is found mainly in the Antafondro Classified Forest of north-eastern Madagascar (former Mahajunga province), and perhaps also in highelevation areas of the Tsaratanana Special Reserve. The Anosy mouse lemur (M. tanosi) is confined to lowland and montane rainforests in the south-eastern corner of Madagascar (former Toliara province). Peters’ mouse lemur (M. myoxinus) has a disjunct distribution in central-western Madagascar (former Mahajanga and Toliara provinces) between the northern banks of the Tsiribihina River north to Baie de Baly, the forests of Belo sur Tsiribihina and Aboalimena, and in the Tsingy de Bemaraha and Tsingy de Namoroka national parks.

The fossa (Cryptoprocta ferox), the largest carnivore of Madagascar, is found throughout the country except for the Central Plateau, but is everywhere threatened by habitat destruction, hunting, and human persecution.

The eastern falanouc (Eupleres goudotii) is a mongooselike carnivore that is widespread throughout the length of eastern Madagascar, but rare and extensively hunted.

The spotted fanaloka (Fossa fossana) is a small, civet-like species found over much of eastern and north-eastern Madagascar, where it is vulnerable to hunting and habitat destruction.

The broad-striped vontsira (Galidictis fasciata) is a type of mongoose found from Marojejy National Park throughout the eastern rainforest as far south as Andohahela, although always at very low densities.

The web-footed tenrec (Limnogale mergulus) is a semiaquatic insectivore known only from a small number of stream and river habitats across a wide area of east-central Madagascar.

The northern shrew tenrec (Microgale jobihely) is known from two highly disjunct populations in eastern and northeastern Madagascar (former Mahajanga and Toamasina provinces). Nasolo’s shrew tenrec (M. nasoloi) is known only from a few specimens collected from three disjunct localities in south-western Madagascar (former Toliara province).

Petter’s tuft-tailed rat (Eliurus petteri) was, until 2003, known only from three specimens, but has since been discovered in a few localities in central-eastern Madagascar (former Toamasina province). Ellerman’s tuft-tailed rat (E. ellermani) is known only from two specimens, one collected some 40 km north-west of Maroantsetra in north-eastern Madagascar and the other in the vicinity of Lohariandava in the central-east (former Toamasina province). The whitetipped tuft-tailed rat (E. penicillatus) was originally described in the early twentieth century from Ampitambe Forest near Ambositra in east-central Madagascar (former Fianarantsoa province). It was not recorded again until 2000 when it was collected at Anerana, about 35 km northeast of Fandriana.

The Madagascan flying fox (Pteropus rufus) is widespread on Madagascar, with the highest density of roost sites being in the coastal regions, especially from Morombe in the south-west to Antsiranana in the north. It is intensively hunted for food.

The Madagascan fruit bat (Eidolon dupreanum) is found over much of Madagascar with the exception of the eastern rainforests. It is everywhere rare, however, and dependent upon suitable cave for roosting sites, where it too is frequently hunted by local people for food.

The Comoros bent-winged bat (Miniopterus griveaudi) is known from Grande Comore and Anjouan as well as from a few localities in northern and western Madagascar.

The western yellow bat (Scotophilus tandrefana) is similarly known only from two specimens, one from near Parc National Tsingy de Bemaraha and the second at Sarodrano (former Mahajanga and Toliara provinces).

The Madagascar slit-faced bat (Nycteris madagascariensis) is known only from two specimens collected in 1910, in the Irodo River Valley of northern Madagascar near Analamera (former Antsiranana province). It has not been found since, despite surveys.

Racey’s pipistrelle bat (Pipistrellus raceyi) is known only from a few specimens collected at two lowland rainforest localities in the central-east and two more, lowland dry forest sites in the central-west.

The Madagascar serpent eagle (Eutriorchis astur) is a rare inhabitant of the eastern rainforests from Marojejy in the north to Zahamena in the south, as well as the Central Highlands at Anjozorobe and Bemanevika. The total population likely numbers less than 1000.

The Madagascar fish eagle (Haliaeetus vociferoides) survives in low numbers within wooded areas adjacent to mangroves and shallow bays along the western coast of Madagascar (former Antsiranana, Mahajanga, and Toliara provinces). Its numbers have declined dramatically in recent years, and it is now one of the world’s rarest birds of prey. Only about 120 breeding pairs survive.

The Madagascar red owl (Tyto soumagnei) was long feared to be extinct but is now known to be fairly widespread throughout the eastern rainforests. However, the total population is small and severely fragmented.

Bernier’s teal (Anas bernieri) is found in a narrow strip along almost the whole of the west coast and extreme northeast of Madagascar (former Antsiranana, Mahajanga, and Toliara provinces), in both mangrove and freshwater wetlands. It is threatened by habitat destruction and disturbance, and is frequently hunted.

The Madagascar rail (Rallus madagascariensis) occupies suitable wetland habitat in the east up to 1800 m, where it is generally rare if locally not uncommon.

The white-breasted mesite (Mesitornis variegatus) is a raillike ground-dweller found at five localities in northern and central-western Madagascar (Menabe Forest, the Ankarafantsika complex, the Ankarana Special Reserve, Analamera Special Reserve, and the Daraina forests), as well as in the Ambatovaky Special Reserve in the east. Surveys have demonstrated its genuine absence frommany intervening areas, including somewith apparently suitable habitat.

The helmeted vanga (Euryceros prevostii) is a shrike-like bird found patchily in the rainforests of eastern and northeastern Madagascar (former Antsiranana and Toamasina provinces), where it is threatened by loss of habitat.

The short-legged ground-roller (Brachypteracias leptosomus) is still fairly common in eastern Madagascar from Daraina Forest in the north to Andohahela in the south, where it is restricted to undisturbed primary forest.

Appert’s greenbul (Xanthomixis apperti) is a small, terrestrial bird known only from four tiny forest patches in southwestern Madagascar (former Toliara province).

The spider tortoise (Pyxis arachnoides) is confined to the arid region of south-western Madagascar (former Toliara province) from the coast up to 10–50 km inland going as far north as Morombe. There are three subspecies. The nominate form (P. a. arachnoides) occurs in the region of the Onilahy River near Toliara, Brygoo’s spider tortoise (P. a. brygooi) occurs south of the Mangoky River, and the oblong spider tortoise (P. a. oblonga) along the southern coast. All are threatened by habitat destruction and fragmentation, as well as by hunting for food.

Petter’s chameleon (Furcifer petteri) is known from a few localities over a relatively wide area of northern Madagascar (former Antsiranana and Mahajanga provinces). The twobanded chameleon (F. balteatus) is known from two widely disjunct areas in the central south-east and far south-east (former Fianarantsoa and Toliara provinces), where it is rarely encountered. Laborde’s chameleon (F. labordi) is found in both dry deciduous and spiny succulent forest in western and south-western Madagascar (former Toliara province). All are threatened by habitat destruction and overcollection for the exotic pet trade.

Two subspecies of elongated leaf chameleon (Palleon nasus) inhabit areas of higher-elevation humid forest in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). Boulenger’s elongated leaf chameleon (P. n. nasus) is known only from its type locality of Manjarivolo, while Paulian’s elongated leaf chameleon (P. n. pauliani) is fairly widespread. Both are threatened by habitat destruction.

The leaf chameleons (Brookesia) are a group of dullcoloured, mostly terrestrial species that include some of the world’s smallest reptiles. The northern leaf chameleon (B. ebenaui) is known from a few small areas of both dry and humid lowland forest in far northern Madagascar (former Antsiranana and Mahajanga provinces). The mossy leaf chameleon (B. vadoni) is confined to a small area of north-eastern Madagascar (former Antsiranana and Toamasina provinces). Ramanantsoa’s leaf chameleon (B. ramanantsoai) is known only from a few localities in east-central Madagascar (former Antananarivo and Toamasina provinces). Peyrieras’ leaf chameleon (B. peyrierasi) is confined to the Masoala Peninsula and the island of Nosy Mangabe in north-eastern Madagascar (former Antsiranana and Toamasina provinces). The Nosy Be leaf chameleon (B. minima) is confined to lowland forest fragments on Nosy Be and Nosy Komba and a few other localities in the north-west including Manongarivo and, most probably, Sahamalaza. All are threatened by loss of habitat.

Seipp’s day gecko (Phelsuma seippi) is known from a few localities in the Sambirano region of northern Madagascar (former Antsiranana and Mahajanga provinces), and from the islands of Nosy Be and Nosy Komba. It is threatened by loss of habitat.

Boivin’s velvet gecko (Blaesodactylus boivini) is known from two disjunct areas of northern Madagascar (former Antsiranana and Mahajanga provinces) along with some offshore islands.

The leaf-tailed geckos (Uroplatus) are nocturnal and arboreal, with highly cryptic colouration used as camouflage. The Malahelo leaf-tailed gecko (U. malahelo) has been recorded from widely scattered localities in southern Madagascar including Ambatotsirongorongo, Andohahela, Kalambatritra, and Analavelona. The Malama leaf-tailed gecko (U. malama) has been recorded from a few localities in south-eastern Madagascar (former Fianarantsoa and Toliara provinces), including Andohahela, Ampamakiesiny, Kalambatritra, Ranomafana-Sud, Ivorona, Farafara, and Pic Ivohibe. Ebenau’s leaf-tailed gecko (U. ebenaui) is found in far northern Madagascar (former Antsiranana province), including the island of Nosy Be. The giant leaf-tailed gecko (U. giganteus), the world’s second largest living species of gecko, is found in north-eastern Madagascar, where it has been reported from Montagne d’Ambre and Marojejy south into the Masoala Peninsula (former Antsiranana province). All are highly sought after for the international pet trade.

The four-striped plated lizard (Zonosaurus quadrilineatus) is confined to a small area of south-western coastal Madagascar (former Toliara provinces), where it lives in dry forest patches and sand dunes. Boettger’s plated lizard (Z. boettgeri) is found widely but patchily in north-eastern Madagascar (former Antsiranana province), including the island of Nosy Be.

Peters’ keeled plated lizard (Tracheloptychus petersi) is found in a few mostly arid and sandy localities in southwestern coastal Madagascar (former Toliara province).

Millot’s skink (Paracontias milloti) is known only from museum specimens collected from the small islands of Nosy Mitsio and Nosy Mamoko off the north-western coast of Madagascar, and possibly from Marojejy in the north-east. It is possible that the species was confined to areas of now-lost lowland humid forest and has disappeared.

O’Shaughnessy’s skink (Brachyseps gastrostictus) has only been recorded from two widely separated localities in eastern Madagascar (Anjanaharibe-Sud in the north-east and Andrangoloaka further south), which suggests that the species is likely to occur between them as well. The Anosy skink (B. anosyensis) is known from a few scattered localities across a relatively wide area of south-eastern Madagascar (former Fianarantsoa and Toliara provinces). The splendid skink (B. splendidus) is a rare species known only from two extremely disjunct areas of central-western and southeastern Madagascar.

Alluaud’s skink (Flexiseps alluaudi) is only known from Montagne d’Ambre and a few localities within the Sambirano region of northern Madagascar (former Antsiranana province), as well as from the island of Nosy Be. The Mandokava skink (F. mandokava) is confined to a few localities in northern Madagascar (former Antsiranana and Mahajanga provinces).

Bocage’s skink (Pseudoacontias madagascariensis) is a large, fossorial species with severely reduced limbs known only from a single specimen collected from an undefined locality in northern Madagascar during the late nineteenth century. Angel’s skink (P. angelorum) has been recorded at two localities in north-eastern Madagascar (former Antsiranana province).

The Betsileo mabuya skink (Trachylepis betsileana) is known only from a single specimen said to have been collected during the early nineteenth century from east-central Madagascar, but which may have possibly originated from somewhere in mainland Africa.

Günther’s tree snake (Lycodryas guentheri) is known only from a few specimens collected in disjunct areas of the Central Highlands and south-eastern Madagascar. The lemon tree snake (L. citrinus) is known only from a few localities within a small area of western coastal Madagascar (former Mahajanga and Toliara provinces). Both are threatened by loss of habitat and collection for the international pet trade.

The yellow-striped water snake (Thamnosophis stumpffi) is confined to the island of Nosy Be and adjacent mainland areas of suitable habitat in north-western Madagascar (former Antsiranana province).

The Madagascar burrowing snake (Pararhadinaea melanogaster) is divided into two subspecies. The nominate form (P. m. melanogaster) is known only from a few localities across a relatively wide area of northern Madagascar (former Antsiranana province) and the island of Nosy Be.

The small-headed blind snake (Madatyphlops microcephalus) is a burrowing species known from the lowland humid and dry forests of Montagne d’Ambre, Ampombofofo, and Montagne des Français in north-eastern Madagascar (former Antsiranana province).

Trueb’s Madagascar tree frog (Madecassophryne truebae) is known only from a few localities in south-eastern Madagascar (former Toliara province).

The Maharipeo cophyline tree frog (Cophyla maharipeo) is known only from a small population living in an abbey garden at Joffreville, and in the nearby Fontenay Private Nature Park in far northern Madagascar (former Antsiranana province). The four-spotted tree cophyline tree frog (C. tetra) is known only from a few localities in northeastern Madagascar (former Antsiranana province). Millot’s cophyline tree frog (C. milloti) is confined to a small area of northern Madagascar (former Antsiranana province), including the island of Nosy Be. All are threatened by loss of habitat.

The Ranomafana climbing frog (Anodonthyla moramora) is known only from a small area of south-eastern Madagascar (former Fianarantsoa province), centred on Ranomafana National Park.

A large and unique group endemic to the Madagascan Realm are the so-called skeleton frogs (Boophis), so named for their almost translucent skin that allows their bones and internal organs to be observed. They are arboreal and notably ‘tree frog-like’ in terms of their appearance and mode of life. Many are highly threatened by loss of habitat. Sandra’s skeleton frog (B. sandrae) is known only from a few localities in the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). Andreone’s skeleton frog (B. andreonei) is known from a few localities within a small area of northern Madagascar (former Mahajanga and Antsiranana provinces). Jaeger’s skeleton frog (B. jaegeri) occurs on Nosy Be and the adjacent Sahamalaza Peninsula in north-eastern Madagascar (former Antsiranana and Mahajanga provinces). The Sambirano skeleton frog (B. sambirano) is known from the Manongarivo Special Reserve and in a few localities in and around the Tsaratanana Reserve in northern Madagascar (former Antsiranana province). Blommers’ skeleton frog (B. blommersae) is confined to two disjunct areas separated by unsuitable habitat in northern Madagascar (former Antsiranana province). Ulftunn’s skeleton frog (B. ulftunni) is known from a few localities in northern Madagascar (former Antsiranana province). The banded skeleton frog (B. vittatus) is found widely but disjunctly in northern Madagascar (former Anstiranana province). The Haingana skeleton frog (B. haingana) is known from two localities in the region of the Andohahela National Park in south-eastern Madagascar (former Toliara province), but likely occurs a little more widely. The golden-flanked skeleton frog (B. andohahela) is found widely but patchily in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). Böhme’s skeleton frog (B. boehmei) is found widely but patchily in eastern and south-eastern Madagascar (former Toamasina, Fianarantsoa and Toliara provinces). Boettger’s skeleton frog (B. brachychir) is found patchily over a wide area of northern and western Madagascar. The green mantella (Mantella viridis) is confined to a small area of far northern Madagascar (former Antsiranana province) and the island of Nosy Hara. It is threatened by loss of habitat and collection for the international pet trade.

Angel’s Madagascar frog (Boehmantis microtympanum) is confined to forest fragments in far south-eastern Madagascar (former Toliara province).

Boettger’s stump-toed frog (Stumpffia psologlossa) is known only from a small area of north-eastern Madagascar (former Antsiranana province), including the island of Nosy Be.

A number of grainy frogs (Gephyromantis) are threatened by loss of habitat. Klemmer’s grainy frog (G. klemmeri) is known only from Marojejy, Anjanaharibe-Sud, and Betaolana in north-eastern Madagascar (former Antsiranana province). Webb’s grainy frog (G. webbi) and the Silvanus grainy frog (G. silvanus) are both known only from a few localities in the region of Antongil Bay and on Nosy Mangabe in north-eastern Madagascar (former Fianarantsoa province). Boettger’s grainy frog (G. horridus) is known only from three localities in northern Madagascar (Montagne d’Ambre, Tsaratanana and, at least historically, the island of Nosy Be, all in former Antsiranana province). The Isalo grainy frog (G. corvus) is known only from the Isalo and Makay massifs of southwestern Madagascar (former Toliara province). The Ambohitra grainy frog (G. ambohitra) is known from Montagne d’Ambre, Tsaratanana, and Manongarivo in northern Madagascar (former Antsiranana province). The Zavona grainy frog (G. zavona) and the Tandroka grainy frog (G. tandroka) are both confined to a few localities in northern Madagascar (former Antsiranana and Mahajanga provinces). Enk’s grainy frog (G. enki) is confined to the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). The Salegy grainy frog (G. salegy) and the striated grainy frog (G. striatus) are both found relatively widely in northern Madagascar (former Antsiranana and Toamasina provinces). The greater spiny grainy frog (G. spiniferus) is known from a few localities in south-eastern Madagascar (former Fianarantsoa and Toliara provinces).

The marbled rain frog (Scaphiophryne marmorata) is known from a few lowland and highland localities in centraleastern Madagascar (former Toamasina province), where it is threatened by loss of habitat. The obscure rain frog (S. obscura) is known only from an undefined locality somewhere on the north-western coast of Madagascar. The warty rain frog (S. verrucosa) is known only from its original collection in the early twentieth century from coastal south-western Madagascar (former Toliara province).

Mass’ stream frog (Spinomantis massi) is known only from a few localities in northern Madagascar (former Mahajanga and Antsiranana provinces). The Tavaratra stream frog (S. tavaratra) is confined to a small area of northern Madagascar centred on the Marojejy Massif (former Antsiranana and Mahajanga provinces).

Mountains and Highlands

The most important highland regions of Madagascar include a narrow and steep escarpment running along the eastern coast, containing much of the island’s remaining tropical rainforest. To the west of this ridge lies a plateau in the centre of the island known as the Central Highlands and, north of that, the Northern Highlands. Madagascar’s highest peaks rise from three prominent massifs (Tsaratanana, Andringitra, and Ankaratra). They provide a range of habitats from subhumid forest, grassland and woodland mosaics, and montane heathlands.

Sibree’s dwarf lemur (Cheirogaleus sibreei) is known only from a few small but widely separated pockets of high-altitude rainforest in eastern Madagascar.

The yellow-bellied asity (Neodrepanis hypoxantha) is a type of sunbird that was long known only from specimens collected prior to 1933 and thought to be extinct. It has since been found in scattered higher-elevation forests throughout the length of eastern Madagascar.

O’Shaughnessy’s chameleon (Calumma oshaughnessyi) is found widely but patchily along the eastern and southern highlands of Madagascar (former Toamasina, Antananarivo, Fianarantsoa, and Toliara provinces), where it is threatened by loss of habitat.

The Makira skink (Paracontias vermisaurus) is known only from two localities within the Makira Reserve in north-eastern Madagascar (former Toamasina province). The Kankana skink (P. kankana) is known only from a single locality near Lake Alaotra, but likely occurs in a few other areas in northeastern Madagascar.

Coulanges’ tree snake (Brygophis coulangesi) was long known only from a single specimen collected in 1968 at Fierenana in central-eastern Madagascar (former Toamasina province). In the mid 1990s a second was discovered at Andasaribe-Sud in north-eastern Madagascar (former Antsiranana province), suggesting that the species is widespread but very rare and localized.

Grandidier’s water snake (Liopholidophis grandidieri) is a rare inhabitant of high-altitude rainforest and arid shrubland in east-central Madagascar (former Fianarantsoa province). Known localities include Ranomafana National Park and surroundings, as well as the Tsinjoarivo region.

The rounded rain frog (Scaphiophryne boribory) is known from two disjunct localities in north-eastern and northern Madagascar (former Toamasina province).

Guibe’s stream frog (Spinomantis guibei) is known only from a few localities in south-eastern Madagascar (former Toliara and Fianarantsoa provinces).

The Northern Highlands

The Northern Highlands are a mountainous region in northeastern Madagascar (former Antsiranana province) that includes the Tsaratanana Massif and smaller nearby massifs such as Marojejy, Anjanaharibe-Sud, and Manongarivo. It is separated from the Central Highlands by a valley known as the Mandritsara Window, which acts as a barrier to dispersal between the two. They are noted for their extremely high level of species endemism.

The montane shrew tenrec (Microgale monticola) is known from only four localities within a very narrow elevation range (1500–1950 m) on two massifs in north-eastern Madagascar (former Antsiranana province), the latter encompassing Marojejy National Park and the Anjanaharibe-Sud Special Reserve.

The hairy-tailed tree rat (Brachytarsomys villosa) is a poorly known nocturnal and arboreal species known only from five localities in the Northern Highlands (former Antsiranana, Mahajanga, and Toamasina provinces).

The bizarre-nosed chameleon (Calumma hafahafa) was originally known from a single male collected in 2003 from a forest at the Bemanevika Lakes in north-western Madagascar (former Mahajanga province). A handful of additional specimens have subsequently been taken from the same general area, as well as from a forest fragment near the Tsaratanana Massif (former Antsiranana provinces).

The lined leaf chameleon (Brookesia lineata) is known only from two localities at Manongarivo in north-eastern Madagascar (former Mahajanga province) along with a third at Tsaratanana (former Antsiranana provinces). The Bekolosy leaf chameleon (B. bekolosy) is known only from a single specimen collected in 1992 from the Bekolosy Plateau, within the Manongarivo Special Reserve in north-eastern Madagascar (former Antsiranana province).

The Tsaratanana cophyline tree frog (Cophyla tsaratananaensis) was long known only from three specimens collected in 1949 from the Tsaratanana Massif in north-eastern Madagascar, but has since been found at two additional montane localities. The yellowish cophyline tree frog (C. mavomavo) is known only from a small area of the Northern Highlands, but may occur more widely.

The Anjanaharibe skeleton frog (Boophis anjanaharibeensis) is known only from the Anjanahribe-Sud Special Reserve, Ambolokopatrika, and Tsararano, and Marojejy National Park (former Antsiranana province).

The Tahotra grainy frog (Gephyromantis tahotra) and Schilf’s grainy frog (G. schilfi) are both known from a few localities in northern Madagascar (former Antsiranana and Mahajanga provinces). Both are threatened by loss of habitat.

The chubby diamond frog (Rhombophryne botabota) is known only from a few localities within the Northern Highlands, including Marojejy National Park and the Makira Plateau.

Günther’s digging frog (Plethodontohyla guentheri) is known only from Marojejy National Park and the Anjanharibe-Sud Special Reserve.

Montagne d’Ambre (Amber Mountain) is an isolated volcanic massif located near the northern tip of Madagascar (former Antsiranana province). It contains a significant pocket of subhumid forest surrounded at lower elevations by dry deciduous forest. Known for its high levels of endemism, it has long been protected within Montagne d’Ambre National Park.

The Montagne d’Ambre dwarf lemur (Cheirogaleus andysabini) is confined to Montagne d’Ambre National Park and a few areas near the town of Joffreville.

The Montagne d’Ambre rock thrush (Monticola erythronotus) is confined to the subhumid forests of Montagne d’Ambre, where the total population is thought to be less than 5000.

The Montagne d’Ambre leaf chameleon (Brookesia tuberculata) is confined to Montagne d’Ambre. The Forêt d’Ambre leaf chameleon (B. desperata) is known only from the Forêt d’Ambre Special Reserve.

Puellars’ cophyline tree frog (Cophyla puellarum) and Noromalala’s cophyline tree frog (C. noromalalae) are both confined to high-elevation rainforest in Montagne d’Ambre National Park.

The Matavy diamond frog (Rhombophryne matavy) is known only from a single lowland forest locality within the Forêt d’Ambre Special Reserve.

The Montagne d’Ambre stump-toed frog (Stumpffia madagascariensis) is confined to Montagne d’Ambre.

The Marojejy Massif is a mountain chain located in northeastern Madagascar (former Antsiranana province). Access to the area around the massif was restricted to research scientists when it was set aside as a strict nature reserve in 1952. In 1998 it was opened to the public when it was converted into a national park. Originally seen as a transition zone between the eastern rainforests and the central highlands, Marojejy is now recognized as having its own unique features, with some of the richest biodiversity on the island. The wide range of elevations and rugged topography create diverse habitats that transition quickly with changes in altitude. Warm, dense rainforest can be found at lower elevations, followed by subhumid and cloud forest, and topped near the peaks with the only remaining, undisturbed montane shrubland in Madagascar. Unfortunately poaching, selective logging, and wood collection are still persistent problems within the park, particularly since the start of the 2009 political crisis. Moreover, a single outbreak of fire could result in the loss of large parts of the montane eroicoid habitat, as it already has at other sites.

Two chameleons of the genus Calumna from Marojejy are threatened by habitat destruction. The Marojejy peak chameleon (C. jejy) is confined to the higher elevations between 1800 and 2130 m. Peyrieras’ chameleon (C. peyrierasi) is known only from a few specimens.

The Marojejy leaf chameleon (Brookesia karchei) is confined to the Marojejy Massif.

The Marojejy burrowing snake (Pararhadinaea melanogaster marojejyensis) is known only from a single specimen collected within Marojejy National Park.

The Marojejy cophyline tree frog (Cophyla rava) is known only from high elevations within Marojejy National Park.

The Ranjomavo grainy frog (Gephyromantis ranjomavo) is known only from the Marojejy Massif.

The Vaventy diamond frog (Rhombophryne vaventy), tiny diamond frog (R. minuta), and Guibé’s diamond frog (R. serratopalpebrosa) are all known only from the Marojejy Massif, where they are threatened by loss of habitat.

The Marojejy stump-toed frog (Stumpffia roseifemoralis) is known only from the Marojejy Massif.

The Sorata Massif is located in north-eastern Madagascar (former Antsiranana province).

The Sorata diamond frog (Rhombophryne longicrus) is confined to the Sorata Massif.

The Manongarivo Massif is located in north-eastern Madagascar (former Antsiranana province).

The Manongarivo leaf chameleon (Brookesia valerieae) is known only from two isolated, low-elevation localities at Manongarivo.

The Tsaratanana Massif is located in north-central Madagascar (former Antsiranana province). It features the highest mountain in Madagascar, Maromokotro.

The Tsaratanana chameleon (Calumma tsaratananense) is confined to the high-elevation heathland of Tsaratanana.

The Tsaratanana skink (Paracontias manify) is confined to the Tsaratanana Massif.

Olga’s cophyline tree frog (Cophyla olgae) and Guibé’s cophyline tree frog (C. alticola) are both known only from a few specimens collected from the highest parts of the Tsaratanana Massif.

Nussbaum’s stream frog (Spinomantis nussbaumi) is known only from its type locality on the Befosa River, Antetikalambazaha, Tsaratanana Massif.

The ornate diamond frog (Rhombophryne ornata), Guenther Peters’ diamond frog (R. guentherpetersi), and the Tany diamond frog (R. tany) are all confined to the Tsaratnana Massif, where they are threatened by loss of habitat.

The Central Highlands

The Central Highlands or Central High Plateau is comprised of the contiguous part of the island’s interior above 800 m in the east and 600 m in the west. It is separated from the Northern Highlands by a low-lying valley known as the Mandritsara Window, which has apparently acted as a barrier to dispersal between the two. It is thought that subhumid forest originally extended over most of it, although this has long since been replaced by grassland and agriculture, with actual forest reduced to fragmented patches (most notably at Anjozorobe and Ambohitantely). A number of massifs rise out of the highlands and represent some of Madagascar’s highest mountains.

Wright’s sportive lemur (Lepilemur wrightae) is known only from the Kalambatritra Special Reserve and the unprotected Beakora Forest in south-eastern Madagascar (former Fianarantsoa province).

Goodman’s mouse lemur (Microcebus lehilahytsara) is confined to forest fragments in east-central Madagascar (former Toamasina province).

The eastern voalavo (Voalavo antsahabensis) is a type of rodent known only from montane forest surrounding Anjozorobe in east-central Madagascar (former Antananarivo province).

The four-toed rice tenrec (Oryzorictes tetradactylus) is a semi-fossorial species that appears to be restricted to the south-central highlands and the flank of the Eastern Escarpment (former Fianarantsoa province).

Several chameleons of the genus Calumma are threatened by loss of habitat. Hillenius’ chameleon (C. hilleniusi) is known from three disjunct localities in central Madagascar (Ankaratra, Andringitra, and Ivohibe in the central-west, with an isolated record as well from near Ambohijanahary in the south-east). The globe-horned chameleon (C. globifer) appears to be restricted to a relatively small area in the central-east highlands, having been reported from Ambohitantely and the Angavo-Anjozorobe corridor (former Antananarivo province). The Andringitra chameleon (C. andringitraense) is known from a number of localities around the Andringitra Massif and at Kalambatritra in south-eastern Madagascar, and may be present as well at Andohahela (former Fianarantsoa and Toliara provinces). The blunt-nosed chameleon (C. tsycorne) is found disjunctly in a few highland areas of southeastern Madagascar (former Fianarantsoa and Toliara provinces).

The jewelled chameleon (Furcifer campani) is found patchily in montane savanna areas from Andringitra National Park in the south to Ankaratra in the north (former Antananarivo and Fianarantsoa provinces). The species was formerly collected in large numbers for the international pet trade, but in recent years exports have fallen off considerably. The whitelipped chameleon (F. minor) is confined to a small area of the Central Highlands (former Antananarivo and Fianarantsoa provinces), where it is threatened by loss of its Tapia forest habitat.

Pronk’s day gecko (Phelsuma pronki) and the yellowthroated day gecko (P. flavigularis) are both confined to isolated rainforest fragments bordering the Central Highlands in central-eastern Madagascar (former Antananarivo and Toamasina provinces), where they are highly threatened by habitat destruction and overcollection for the international pet trade. Gould’s day gecko (P. gouldi) is known only from a type specimen collected from the Anja Reserve, 13 km south of Ambalavao in south-central Madagascar (former Fianarantoa province), along with a few additional photographs from the same locality.

Three species of dwarf gecko (Lygodactylus) are threatened by loss of habitat. Blanc’s dwarf gecko (L. blanci) is confined to Mount Ibity in central Madagascar (former Antananarivo province). The ornate dwarf gecko (L. ornatus) is known only from a single locality in north-eastern Madagascar (former Mahajanga province). Paulian’s dwarf gecko (L. pauliani) is known only from a handful of specimens collected from a small area of central Madagascar (former Fianarantsoa province).

The rusty skink (Madascincus macrolepis) is a leaf litterdependent species known from two disjunct localities in eastcentral Madagascar (former Toamasina and Fianarantsoa provinces).

The Ankafina brook snake (Pseudoxyrhopus ankafinaensis) is known only from a single specimen collected during the nineteenth century in eastern Madagascar (former Fianarantsoa province). Numerous dedicated searches have failed to find it, and the species is most likely extinct.

The Andrangoloaka skeleton frog (Boophis andrangoloaka) is known only from two small, disjunct localities in north-central Madagascar (former Antananarivo province). The spinophis skeleton frog (B. spinophis) is known only from two specimens collected from Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province) and from the Ambohitantely Special Reserve in north-central Madagascar (former Antananarivo province). It likely occurs patchily between these two disjunct localities. The bright-eyed skeleton frog (B. rhodoscelis) is found widely but patchily in central-eastern Madagascar (former Mahajanga, Toamasina, and Fianarantsoa provinces). All are threatened by loss of habitat.

Cowan’s mantella (Mantella cowanii) is only known for certain from three small, disjunct localities in east-central Madagascar (former Antananarivo and Fianarantsoa provinces), all of which are embedded within a heavily modified agricultural landscape. The species may possibly occur more widely, but this has yet to be confirmed.

Vallan’s climbing frog (Anodonthyla vallani) is only reliably known from a single high-elevation locality within the Ambohitantely Special Reserve, north-central Madagascar (former Antananarivo province).

The dotted Madagascar frog (Guibemantis punctatus) is confined to the vicinity of the Ambohitantely Special Reserve in north-central Madagascar (former Antananarivo province).

The Ambohimitombi Madagascar frog (Mantidactylus ambohimitombi) is known only from a small area of montane grassland in east-central Madagascar (former Antananarivo province). Delorme’s Madagascar frog (M. delormei) is known only from the Andringitra Massif and from the Maharira Forest in Ranomafana National Park in southeastern Madagascar (former Fianarantsoa province).

The Alaotran grainy frog (Gephyromantis mafy) is confined to a single forest fragment near Lake Alaotra in centraleastern Madagascar (former Toamasina province).

Helen’s diamond frog (Anilany helenae) is known only from two small forest fragments in the vicinity of Ambohitantely, in central Madagascar (former Antananarivo province).

The Eastern Escarpment is located along the eastern edge of the Central Highlands in central-eastern and south-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces).

The Betsileo woolly lemur (Avahi betsileo) is known only from the Bemosary Classified Forest in central-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces).

The Betsileo sportive lemur (Lepilemur betsileo) is known only from a small area of central-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces). The small-toothed sportive lemur (L. microdon) is known for certain only from a small area of south-eastern Madagascar (former Fianarantsoa province). Both are threatened by habitat destruction and hunting.

Groves’ dwarf lemur (Cheirogaleus grovesi) is known only from a few specimens collected within Ranomafana and Andringitra national parks (former Toamasina, Antananarivo, and Fianarantsoa provinces).

The rufous mouse lemur (Microcebus rufus) is confined to central-eastern and south-eastern Madagascar (former Toamasina, Antananarivo, and Fianarantsoa provinces), roughly from Andringitra National Park to Ranamafana National Park.

The Tarzan chameleon (Calumma tarzan) is known only from two small patches of degraded forest in central-eastern Madagascar (former Antananarivo province). Glaw’s chameleon (C. glawi) is known from a few localities in south-eastern Madagascar (former Toamasina and Antananarivo provinces).

Lamberton’s leaf chameleon (Brookesia lambertoni) is known only from a small area of central-eastern Madagascar (former Toamasina province).

The cork bark leaf-tailed gecko (Uroplatus pietschmanni) has been reported from a small area of central-eastern Madagascar (former Toamasaina province) that includes Ambatovy and Analamay.

The delightful brook snake (Pseudoxyrhopus oblectator) is known only from the forest of Ranomafana, Anjozorobe, and Ambatovy in central-eastern Madagascar (former Toamasina and Fianarantsoa provinces).

The Betsileo cophyline tree frog (Cophyla cowanii) is known only from a few specimens collected during the nineteenth century from an undefined locality in east-central Madagascar (former Toamasina and Fianarantsoa provinces).

Emile’s climbing frog (Anodonthyla emilei) is known only from two localities within Ranomafana National Park in south-eastern Madagascar (former Toamasina province). The Anamalazoatra climbing frog (A. pollicaris) is known only from a few localities in central-eastern Madagascar (former Toamasina province).

Several species of skeleton frog (Boophis) endemic to the Eastern Escarpment are threatened by loss of habitat. The Mandraka skeleton frog (B. mandraka) is known for certain only from a single locality in central-eastern Madagascar (former Toamasina province). The peppered skeleton frog (B. piperatus), Schuboe’s skeleton frog (B. schuboeae), Boppa’s skeleton frog (B. boppa), and Narin’s skeleton frog (B. narinsi) are all known only from the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). Liam’s skeleton frog (B. liami) is known only from Vohidrazana and near Andasibe in central-eastern Madagascar (former Toamasina province), although it may occur more widely. Burger’s skeleton frog (B. burgeri) is known only from a few specimens collected from a small area around Andasibe in east-central Madagascar (former Toamasina province). Lilian’s skeleton frog (B. lilianae) is known only with certainty from a few adults and tadpoles collected in southern east-central Madagascar (former Toamasina province). Pop’s skeleton frog (B. popi) and Major’s skeleton frog (B. majori) are both found widely but patchily in eastern-central Madagascar (former Toamasina and Fianarantsoa provinces).

Several species of mantella (Mantella) endemic to the Eastern Escarpment are threatened by loss of habitat and collection for the international pet trade. The black-eared mantella (M. milotympanum) was previously known only from a number of fragmented localities in east-central Madagascar covering a small area south of Fierenana (former Toamasina province). It has since been discovered in a few areas to the south-east along the Ivohitra River and around Ranomainty. The golden mantella (M. aurantiaca) has a very restricted distribution in east-central Madagascar (former Toamasina province) centred on the Torotorofotsy area and the Andromena Forest, with populations occurring in several other small forest fragments north and south of Moramanga. The eastern yellow mantella (M. crocea) is known only from a few scattered localities in east-central Madagascar (former Toamasina province). The painted mantella (M. madagascariensis) is known only from a few localities within a relatively wide area of east-central Madagascar (former Toamasina and Fianarantsoa provinces).

The white-moustached Madagascar frog (Mantidactylus albofrenatus) and the Andasibe Madagascar frog (M. zolitschka) are each known only from a small area of eastcentral Madagascar (former Toamasina province). The Ranomafana Madagascar frog (M. paidroa) is known only from the region of Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). All are threatened by loss of habitat.

The Kibomena stump-toed frog (Stumpffia kibomena) is known only from three localities within a small area of centraleastern Madagascar (former Toamasina province).

Runewsweek’s grainy frog (Gephyromantis runewsweeki) is only reliably known from near the summit of Mount Maharira, within Ranomafana National Park in south-eastern Madagascar (former Fianarantsoa province). The Thelen grainy frog (G. thelenae) and Eiselt’s grainy frog (G. eiselti) are each known only from a small area in central-eastern Madagascar (former Toamasina province). The horned grainy frog (G. cornutus) is known from two small, disjunct localities in central-eastern Madagascar (former Toamasina province).

The Betsileo digging frog (Plethodontohyla brevipes) is known only from a few individuals collected at Betsileo and Ranomafana in east-central Madagascar (former Toamasina and Fianarantsoa provinces).

The Ankaratra Range (also known as the Ankaratra Massif ) is an extinct/dormant volcanic range located in eastcentral Madagascar (former Antananarivo province). The highest peak in Ankaratra is Tsiafajavona, at 2644 m. Some trace of activity remains in the southernmost part of the range, creating hot springs in the area around Antsirabe.

The Ankaratra dwarf gecko (Lygodactylus mirabilis) is confined to montane grassland within the Ankaratra Range.

Williams’ skeleton frog (Boophis williamsi) is known from streams at between 2100 and 2381 m on Mount Tsiafajavona, an isolated peak in the Ankaratra Massif and the second highest in Madagascar. One of the most threatened amphibians in Madagascar, it lives in a severely degraded area where its habitat is under constant threat from fire, illegal logging, overgrazing by livestock, and expanding potato farming.

Paulian’s Madagascar frog (Mantidactylus pauliani) is known only from a small area of the Ankaratra Range. The Andringitra Massif is located in south-eastern Madagascar (former Fianarantsoa province). It is protected within Andringitra National Park.

The Andringitra dwarf gecko (Lygodactylus intermedius) is confined to rocky heathland on the Andringitra Massif.

The Andringitra skeleton frog (Boophis laurenti) is confined to montane heathland and some higher-elevation forest within Andringitra National Park.

The Andringitra Madagascar frog (Mantidactylus madecassus) and Bourgat’s Madagascar frog (M. bourgati) are both confined to the higher elevations of Andringitra.

The mountain climbing frog (Anodonthyla montana) is known only from a single locality in Andringitra, between 1900 and 2650 m.

The Anosy Chain (Chaînes Anosyennes in French) is located in south-eastern Madagascar (former Fianarantsoa and Toliara provinces).

Capuron’s chameleon (Calumma capuroni) is known only from Andohahela National Park.

The striped brook snake (Pseudoxyrhopus sokosoko) is found only in far south-eastern Madagascar (former Toliara province), with all known specimens having been found in the lower-elevation forests of the Anosy and Vohimena Mountains.

The Andohahela skeleton frog (Boophis miadana) is known only from a single locality within Andohahela National Park (former Toliara province).

The Andohahela climbing frog (Anodonthyla jeanbai) is known only from a single locality within Andohahela National Park. The Anosy climbing frog (A. rouxae) has a very small range close to the summit in the Anosy Mountains in south-eastern Madagascar, as well as in Andohahela National Park.

The Andohahela stream frog (Spinomantis microtis) is confined to the Anosy Chain.

The Isalo Massif is located in south-central Madagascar (former Fianarantsoa and Toliara provinces). It is protected within Isalo National Park.

The Isalo serotine bat (Neoromicia malagasyensis) is confined to the Isalo Massif region.

Nora Lott’s Madagascar frog (Mantidactylus noralottae) is currently known only from the vicinity of Isalo National Park, where it lives on canyon walls and beside rock pools.

The blue-legged mantella (Mantella expectata) is confined to a small area centred on Isalo National Park.

The Isalo grainy frog (Gephyromantis azzurrae) is known only from three localities on the Isalo Massif.

Gottlebe’s rain frog (Scaphiophryne gottlebei) is known only from a few localities in the Isalo Massif area.

Lowland Rainforests

Lowland rainforests occupy a relatively narrow strip between the eastern coast and the mountainous Central Highlands, from sea level to 800 m elevation. They extend from Marojejy in the north-east to the south-east corner of the island. At the northern edge around Vohemar, these moist forests transition to the dry deciduous forest characteristic of the western and far northern coast, and to higher-elevation subhumid forests to the east. The southern end lies at the crest of the Anosy Mountains, where a narrow belt of dry forest marks the transition to the spiny thickets beyond. While the lowland rainforests have been preserved generally better than have the original woodlands of the high Central Plateau, there has still been considerable loss. Although it has nearly been destroyed, the moist coastal forest of Madagascar may also be included in the lowland rainforest category. Taken together, the lowland rainforests represent a great reservoir of diversity and endemism. Nearly all of Madagascar’s endemic mammal genera are represented here, including all five families of lemurs.

Three species of woolly lemur (Avahi) from lowland rainforest areas are threatened by loss of habitat and subsistence hunting. The Masoala woolly lemur (A. mooreorum) is known only from Masoala National Park in north-eastern Madagascar (former Antsiranana province). The Manombo woolly lemur (A. ramanantsoavanai) is confined to a few areas of southeastern Madagascar (former Toliara and Fianarantsoa provinces). The southern woolly lemur (A. meridionalis) is confined to a small area of lowland rainforest and littoral forest in the south-eastern corner of Madagascar (former Toliara province).

Several species of sportive lemur (Lepilemur) are found in lowland rainforest areas, where they are threatened by habitat destruction and subsistence hunting. The Masoala sportive lemur (L. scottorum) is known only from Masoala National Park in north-eastern Madagascar (former Antsiranana province). The Mananara-Nord sportive lemur (L. hollandorum) is confined to a small area of coastal north-eastern Madagascar (former Toamasina province) centred on the Mananara-Nord Biosphere Reserve. The Manombo sportive lemur (L. jamesorum) is confined to the region of the Manombo Special Reserve in south-eastern Madagascar (former Fianarantsoa province) where the total population in 2010 was estimated at around 1400 and declining. Madame Fleurette’s sportive lemur (L. fleuretae) is confined to three small rainforest fragments in the south-eastern corner of Madagascar (former Toliara province).

In recent years a great many species of mouse lemur (Microcebus) have been described throughout Madagascar, each confined to a relatively small area where they are often threatened by habitat destruction and hunting. MacArthur’s mouse lemur (M. macarthurii) is known only from Makira National Park and the island of Nosy Mangabe in northeastern Madagascar (former Toamasina province). Simmons’ mouse lemur (M. simmonsi) is known only from the Betampona, Zahamena, and Tampolo regions of north-eastern Madagascar (former Toamasina province). The Sahafina mouse lemur (M. gerpi) is known only from the Sahafina Forest, near Mantadia National Park in central-eastern coastal Madagascar (former Toamasina province). The Marohita mouse lemur (M. marohita) is currently known only from the Marohita Forest in central-eastern coastal Madagascar (former Toamasina province). Jolly’s mouse lemur (M. jollyae) is confined to the Mananjary and Kianjavato regions of south-eastern Madagascar (former Fianarantsoa province).

The brown-tailed vontsira (Salanoia concolor) is a rare mongoose-like species confined to north-eastern Madagascar (former Antsiranana and Toamasina provinces).

The dryad shrew tenrec (Microgale dryas) is only known from a few specimens collected in north-eastern Madagascar (former Toamasina province).

Peterson’s long-fingered bat (Miniopterus petersoni) appears to be restricted to the lowland rainforests of the south-east corner of Madagascar (former Toliara province).

Bernier’s vanga (Oriolia bernieri) is a shrike-like bird that is rare and patchily distributed throughout its fairly wide range in eastern Madagascar.

The red-tailed newtonia (Newtonia fanovanae) is a type of passerine bird that was long known only from a single specimen collected in 1931 near Fanoyana, in east-central Madagascar. It was almost simultaneously rediscovered in Andohahela National Park in 1989 and in the Ambatovaky Special Reserve in 1990 and is now known from a number of low-elevation rainforest blocks, but is everywhere rare.

The dusky tetraka (Crossleyia tenebrosa) is a rare type of warbler confined to eastern and north-eastern Madagascar (former Antsiranana and Toamasina provinces). The scaly ground-roller (Geobiastes squamiger) is found throughout the lowland rainforests of eastern Madagascar, but is everywhere declining due to loss of habitat.

The brown mesite (Mesitornis unicolor) is a grounddwelling bird with a thin and patchy distribution down the length of the eastern rainforests from Marojejy and the Masoala Peninsula as far south as Taolañaro (former Fort Dauphin). It is everywhere uncommon and rarely seen.

Several chameleons of the genus Calumma are threatened by habitat destruction. The Vatosoa chameleon (C. vatosoa) and Vences’ chameleon (C. vencesi) are both known only from a small area of north-eastern Madagascar (former Antsiranana province). The Zahamena chameleon (C. furcifer) is known only from a few localities in central-eastern Madagascar (former Toamasina province). The lance-nosed chameleon (C. gallus) is found patchily in central-eastern Madagascar (former Toamasina province). The hooded chameleon (C. cucullatum) is found widely but patchily over a relatively wide area of north-eastern and eastern Madagascar.

The Zahamena ground gecko (Paroedura masobe) is restricted to low-elevation primary rainforest in the north of the Zahamena–Ankeniheny corridor, in central-eastern Madagascar (former Toamasina province). It is rarely encountered and difficult to find.

Spannring’s leaf-toed gecko (Matoatoa spannringi) was long known only from a few microhabitats at Fiadanana in south-eastern Madagascar (former Fianarantsoa province), although more recently a single specimen was recorded from Sainte Luce in the far south-east (former Toliara province).

Boettger’s dwarf gecko (Lygodactylus madagascariensis) is found patchily in north-eastern Madagascar (former Antsiranana province). The tiny-scaled dwarf gecko (L. bivittis) is found in eastern Madagascar (former Antsiranana and Toamasina provinces). The Roavolana dwarf gecko (L. roavolana) is found patchily in the south-eastern corner ofMadagascar (former Toliara province). All are threatened by loss of habitat.

Gabriellà’s gecko (Paragehyra gabriellae) is known from a few localities in the south-eastern corner of Madagascar (former Toliara province).

The Tsararano Forest skink (Paracontias tsararano) is known only from a single locality in north-eastern Madagascar (former Antsiranana province), but may be more widespread.

The banded skink (Brachyseps spilostichus) is known only from the Tsararano Forest in north-eastern Madagascar (former Antsiranana province).

The Masoala skink (Flexiseps stylus) is known only from the Masoala Peninsula in north-eastern Madagascar (former Antsiranana province), where it is threatened by loss of habitat.

The Mandady skink (Brachyseps mandady) is known only from the Masoala Peninsula in north-eastern Madagascar (former Antsiranana province), where it is threatened by loss of habitat.

The dwarf skink (Madascincus nanus) is known only from north-eastern Madagascar (former Mahajanga province).

Three species of plated lizard (Zonasaurus) are threatened by loss of habitat. Boettger’s plated lizard (Zonosaurus subunicolor) is known from two disjunct mainland localities in north-eastern Madagascar along with a few offshore islands (former Antsiranana province). The giant plated lizard (Z. maximus) and the Anelanelany plated lizard (Z. anelanelany) are both known only from a few localities in south-eastern Madagascar (former Fianarantsoa and Toliara provinces).

The Iaraka tree snake (Phisalixella iarakaensis) is known only from a single specimen collected at Iaraka in northeastern Madagascar (former Antsiranana province).

The Zeny forest snake (Compsophis zeny) is known only from a few specimens collected over a relatively wide area of south-eastern Madagascar (former Fianarantsoa and Toliara provinces). Vincke’s forest snake (C. vinckei) is known from two specimens, the first collected at Andasibe and the second at An’ala, in east-central Madagascar (former Toamasina province).

Blanc’s night snake (Ithycyphus blanci) is a venomous species known only from a single specimen collected from lowland rainforest within Marojejy National Park, northeastern Madagascar (former Antsiranana province). Most of this habitat has since been destroyed as a result of rosewood logging.

The small brook snake (Pseudoxyrhopus kely) is a naturally rare species known with certainty only from five localities in the south-eastern corner of Madagascar (former Toliara province). Specimens have also been recorded from the dry forests of Bemaraha in western Madagascar, but will almost certainly prove to be a separate taxon.

The Andasibe blind snake (Madatyphlops andasibensis) is known only from Andasibe in east-central Madagascar (former Toamasina province). Domergue’s blind snake (M. domerguei) and Rajery’s blind snake (M. rajeryi) are both known only from Ranomafana National Park in east-central Madagascar (former Fianarantsoa province). Given their cryptic lifestyle some or all of these species may prove to be more widespread than is currently understood.

Theo’s climbing frog (Anodonthyla theoi) is known only from a two small patches of lowland rainforest in the Manombo Special Reserve of coastal south-eastern Madagascar (former Fianarantsoa province). Hutchison’s climbing frog (A. hutchisoni) is known only from two localities within the Masoala Peninsula of northern Madagascar (former Antsiranana and Toamasina provinces). The blackthroated climbing frog (A. nigrigularis) is confined to a small area of extreme south-eastern Madagascar (former Toliara province). All are threatened by loss of habitat.

Several species of skeleton frog (Boophis) are threatened by loss of their lowland forest habitat. Fay’s skeleton frog (B. fayi) is known only from a few localities in northern Madagascar (former Toamasina province). Englaender’s skeleton frog (B. englaenderi) is known only from a few localities in northern Madagascar (former Antsiranana and Toamasina provinces). The Andasibe skeleton frog (B. feonnyala) is known only from a small area of central-eastern Madagascar (former Toamasina province). The secret skeleton frog (B. arcanus) is known only from a few localities within a small area of south-eastern Madagascar (former Fianarantsoa province). The Vohidrazana skeleton frog (B. solomaso) is known for certain only from the Vohidrazana Forest and nearby Analambalo in central-eastern Madagascar (former Toamasina province), but may also occur within the Betampona Reserve and in other localities between. The tiny skeleton frog (B. miniatus) is known only from a few localities within a relatively wide area of south-eastern Madagascar (former Fianarantsoa and Toliara provinces). The Toliara skeleton frog (B. haematopu) is known only from a few localities within a small area of extreme south-eastern Madagascar (former Toliara province).

The Tiarano web-footed frog (Paradoxophyla tiarano) is a semi-aquatic species that to date has only been collected from one area on the Masoala Peninsula in north-eastern Madagascar (former Antsiranana province), but is suspected to occur more widely.

The Savaka diamond frog (Rhombophryne savaka) is confined to lowland rainforest within Marojejy National Park in north-eastern Madagascar (former Antsiranana province).

The Ranomafana stump-toed frog (Stumpffia miery) is confined to a small area of south-eastern Madagascar (former Fianarantsoa province).

Bernhard’s mantella (Mantella bernhardi) is known only from a few localities within a relatively wide area of southeastern Madagascar (former Fianarantsoa province). The Antsiranana mantella (M. manery) is known from a few scattered localities in northern Madagascar (former Antsiranana province). Harald Meier’s mantella (M. haraldmeieri) is known from a few localities in extreme south-eastern Madagascar (former Toliara province). All are threatened by loss of habitat and collection for the international pet trade.

The Anosy stream frog (Spinomantis brunae) is known from Andohahela National Park and Manantantely in far south-eastern Madagascar (former Toliara province).

The stream grainy frog (Gephyromantis rivicola) is known from a few localities over a relatively wide area of northern Madagascar (former Antsiranana and Toamasina provinces). Hintelmann’s grainy frog (G. hintelmannae) is confined to a small area of coastal south-eastern Madagascar (former Toamasina province). Both are threatened by loss of habitat.

Watterson’s Madagascar frog (Guibemantis wattersoni) is confined to a few localities in extreme south-eastern Madagascar (former Toliara province). The white-flanked Madagascar frog (G. tasifotsy) is known from a few localities over a relatively wide area of south-eastern Madagascar (former Fianarantsoa province). Kathrin’s Madagascar frog (G. kathrinae) is known only from two small, highly disjunct localities in northern and central-eastern Madagascar. It is not known if it occurs in the intervening areas or if the two populations instead represent distinct species. All are threatened by loss of habitat.

The three-banded Madagascar frog (Mantidactylus tricinctus) is known only from a few localities in south-eastern Madagascar (former Toliara and Fianarantsoa provinces). Littoral Forests and Scrub

The moist coastal forests of eastern Madagascar are among the most threatened ecoregions in the country.

Thomas’ dwarf lemur (Cheirogaleus thomasi) is confined to littoral forest fragments along the south-eastern corner of Madagascar (former Toliara province).

The Vohibola chameleon (Calumma vohibola) is confined to littoral forest fragments between Ivoloina and Vohibola in eastern Madagascar (former Toamasina province).

Several species of day gecko (Phelsuma) are threatened by loss of habitat. The flat-tailed day gecko (P. serraticauda) is known from a few localities in coastal eastern and northeastern Madagascar (former Antsiranana and Toamasina provinces). The Masoala day gecko (P. masohoala) is known only from three specimens collected at Cap Est on the Masoala Peninsula of north-eastern Madagascar (former Antsiranana province). Extensive searches have failed to record it at any other sites. The Ampitambe day gecko (P. kely) is known only from the area around Lake Ampitambe in east-central coastal Madagascar (former Toamasina province). Hoesch’s day gecko (P. hoeschi) has so far only been recorded from Brickaville (Ampasimanaolotra) and Anosibe An’Ala in eastcentral coastal Madagascar (former Toamasina province). Additional subpopulations are likely to occur between Ampasimanolotra and Vatomandry. The southern day gecko (P. antanosy) is known from a few coastal forest fragments in south-eastern coastal Madagascar (former Toliara province), specifically Ambatotsirongorongo and Sainte Luce (former Toliara province). A third population, at Petriky, was extirpated before 1994 as a result of habitat destruction.

The rock skink (Flexiseps decaryi) was long known only from a single locality, but has since been reported from a few other areas on the south-eastern coast of Madagascar (former Toliara province).

The Manombo Madagascar frog (Guibemantis diphonus) is confined to the Manombo Special Reserve in coastal southeastern Madagascar (former Fianarantsoa province). The ringwearing Madagascar frog (G. annulatus) is confined to three isolated pockets of low-elevation littoral forest (Mandena, Saint Luce, and the Agnalaro Forest) in south-eastern Madagascar (former Toliara province), where it is highly dependent upon stands of Pandanus plants.

The southern jumping frog (Aglyptodactylus australis) is known only from a small area of extreme south-eastern Madagascar (former Toliara province), where it is threatened by loss of habitat.

Lowland Dry Deciduous Forests

The tropical dry deciduous forests that are to be found in Madagascar are among the richest and most distinctive of their kind in the world, with high numbers of plant and animal species. There are two separate areas within this bioregion: the western side of Madagascar from the Ampasindava Peninsula in the north-east to Belo-sur-Tsiribihina and Maromandia in the central-west; and the far northern tip of the island (apart from the high elevations of Montagne d’Ambre). Together, these dry deciduous forests span the coastal plain with its limestone plateaus (known as tsingy) emanating virtually at sea level to altitudes of roughly 600 m. The area also includes wetlands and grasslands (the latter mostly created by forest clearance for agriculture). The dry forests themselves are characterized by a deciduous canopy extending to a height of 10–15 m. Rainfall is more abundant than in the succulent woodlands and spiny thickets to the south, but lower by far than in the eastern lowland rainforests. The trees have adapted to the dry climate by shedding leaves in the winter months to limit fluid loss. Some, such as the baobabs, have adapted by evolving the ability to store copious amounts of water in their large, bulbous trunks. While human population densities are lower in these regions, the dry deciduous forests have nevertheless been largely cleared for slash-and-burn agriculture, pasture, firewood, and logging.

Lorenz Von Liburnau’s woolly lemur (Avahi occidentalis) is found in two disjunct areas of north-western Madagascar (former Mahajanga province) where it is threatened by habitat destruction and hunting.

Several dry forest species of sportive lemur (Lepilemur) are threatened by habitat destruction and hunting. The Sahafary sportive lemur (L. septentrionalis) is arguably the most endangered lemur species of them all. Only about 50 individuals survive in a few small patches of forest north of the Irodo River in far north-eastern Madagascar (former Antsiranana province), near the villages of Madirobe and Ankarongana. Gray’s sportive lemur (L. dorsalis) is confined to the Sambirano region of north-eastern Madagascar (former Mahajanga province). Mittermeier’s sportive lemur (L. mittermeieri) is confined to the Ampasindava Peninsula in northeastern Madagascar (former Antsiranana province). Milne- Edwards’ sportive lemur (L. edwardsi), Antafia sportive lemur (L. aeeclis), Grewcock’s sportive lemur (L. grewcockorum), Ahmanson’s sportive lemur (L. ahmansonorum), and Otto’s sportive lemur (L. otto) are all confined to small areas of coastal north-western Madagascar (former Mahajanga province). The Bemaraha sportive lemur (L. randrianasoloi) is confined to a small area of central-western coastal Madagascar (former Mahajanga province). The red-tailed sportive lemur (L. ruficaudatus) is found patchily in central-western coastal Madagascar (former Toliara province).

The pale fork-marked lemur (Phaner pallescens) is patchily but widely distributed along a narrow strip of western Madagascar (former Mahajunga and Toliara provinces). The northerly populations appear isolated and may possibly represent distinct taxa. The Sambirano fork-marked lemur (P. parienti) is confined to the Sambirano region of north-eastern Madagascar (former Antiranana and Mahajanga provinces), specifically the Ampasindava Peninsula south to the Andranomalaza River. Both are threatened by loss of habitat.

Coquerel’s giant mouse lemur (Mirza coquereli) is found patchily throughout western Madagascar (former Mahajunga and Toliara provinces), where it is threatened by habitat destruction.

Owing to their small size the mouse lemurs (Microcebus) of the western dry forests are less vulnerable to hunting pressure, although habitat destruction is a serious threat. The Bongolava mouse lemur (M. bongolavensis) is known only from three small dry forest fragments in the area around Port- Bergé in north-western Madagascar (former Mahajanga province). The Ambarijeby mouse lemur (M. danfossi) is only known from a few dry forest fragments between the Sofia and Maevarano rivers in north-western Madagascar (former Mahajanga province). The golden-brown mouse lemur (M. ravelobensis) occurs in the forests of Ankarafantsika as well as in the Bongolava and Mariarano Classified Forests in northwestern Madagascar (former Mahajanga province). Madame Berthe’s mouse lemur (M. berthae), the world’s smallest primate, is confined to the Menabe region south of the Tsiribihina River in western coastal Madagascar (former Toliara province). The total range is quite small and highly fragmented.

The western falanouc (Eupleres major) is a mongoose-like mammal that occurs patchily in north-western Madagascar (former Antsiranana and Mahajanga provinces) from near Baly Bay National Park north through the Ankarafantsika and Analalava forests to the Sambirano, east to the foothills of Tsaratanana, the Sahamalaza Peninsula, and possibly as far as the lower elevations of Montagne d’Ambre. It is rarely seen and presumably rare.

Jenkin’s shrew tenrec (Microgale jenkinsae) is known only from two specimens collected in the Mikea Forest of southwestern Madagascar (former Toliara province).

The giant jumping rat (Hypogeomys antimena) is a rabbitsized, kangaroo-like species restricted to a small area of coastal western Madagascar (former Toliara province) from the Tomitsy River north to the Tsiribihina River. Habitat destruction, hunting and introduced species threaten its survival.

The lowland red forest rat (Nesomys lambertoni) is a littleknown species confined to a small area of dry forest in western Madagascar (former Mahajanga province).

The greater big-footed mouse (Macrotarsomys ingens) is a nocturnal species known only from the vicinity of Ankarafantsika National Park in north-western Madagascar (former Mahajanga province), where it is threatened by feral cats and dogs as well as by wildfires. Petter’s big-footed mouse (M. petteri) is known only from a single specimen collected in 2003 in the Mikea Forest of south-western coastal Madagascar (former Toliara province).

Grandidier’s trident bat (Paratriaenops auritus) is an obligate cave-roosting species with a restricted range in the northeastern tip of the island (former Antsiranana province).

Van Dam’s vanga (Xenopirostris damii) is a rare shrikelike bird currently known only from two small, disjunct areas in north-western and far northern Madagascar (Ankarafantsika and Analamera), where it is under threat by fire and other pressures. However, there is still much suitable habitat between the two sites that have yet to be surveyed.

The flat-tailed tortoise or kapidolo (Pyxis planicauda) is restricted to dry forest fragments within a small area of western coastal Madagascar (former Toliara province).

The ploughshare tortoise or angonoka (Astrochelys yniphora), one of the world’s rarest tortoises, is confined to two localities around Baly Bay in north-western Madagascar (former Mahajanga province) which have recently been declared a national park. Nevertheless, the species may soon be extinct in the wild due to illegal collection for use in the international pet trade. It is well established in captivity.

The rhinoceros chameleon (Furcifer rhinoceratus) is known only from a few localities in north-western coastal Madagascar (former Mahajanga province). The Antimena chameleon (F. antimena) is known only from a small area of south-western coastal Madagascar (former Toliara province), but may be more widespread. Both are threatened by loss of habitat.

Decary’s leaf chameleon (Brookesia decaryi) and the toothed leaf chameleon (B. dentata) are both known only from Ankarafantsika National Park in north-western Madagascar (former Mahajanga province).

The Bemaraha day gecko (Phelsuma borai) is known for certain only from a single specimen collected from the Bemaraha region of central-western Madagascar (former Mahajanga province). However, published photos indicate that the species may occur to the north at Ankarafantsika, suggesting that it may occur in intervening areas as well. Van Heygen’s day gecko (P. vanheygeni) is known only from three localities on the Ampasindava Peninsula of northern Madagascar (former Antsiranana province). The yellowheaded day gecko (P. klemmeri) is known only from the Ampasindava Peninsula in northern Madagascar and from an area of dry forest near Mandrozo Lake some 600 km to the south. Surveys between these two localities as well as on coastal islands have failed to detect this highly conspicuous species, and the two populations are likely to be genuinely isolated from one another.

Henkel’s leaf-tailed gecko (Uroplatus henkeli) and Günther’s leaf-tailed gecko (U. guentheri) are both found widely but patchily in northern and western Madagascar (former Antsiranana, Mahajanga, and Toliara provinces).

The Vazimba Madagascar ground gecko (Paroedura vazimba) is known from Ankarafantsika, Anjiamangirana, and Bora in north-westernMadagascar (former Mahajanga province).

The Ambonihazo velvet gecko (Blaesodactylus ambonihazo) is known only from a few specimens collected from Ankarafantsika National Park in north-western Madagascar (former Mahajanga province).

Boettger’s skink (Voeltzkowia mira) is confined to a few localities in north-western Madagascar (former Mahajanga province).

The Tavaratra mabuya skink (Trachylepis tavaratra) is confined to far northern Madagascar (former Antsiranana province).

Yamagishi’s mermaid skink (Sirenoscincus yamagishi), sonamed for its lack of hindlimbs, is known from Belambo and Ankarafantsika in north-western Madagascar (former Mahajanga province).

The Menamainty skink (Pseudoacontias menamainty) is known only from a single specimen collected in north-western coastal Madagascar (former Mahajanga province).

The yellow skink (Flexiseps ardouini) is confined to far northern Madagascar (former Antsiranana province).

Petter’s short skink (Pygomeles petteri) is a burrowing species confined to the lowland dry forests of Ankarafantsika National Park in north-western Madagascar (former Mahajanga province).

The variable tree snake (Phisalixella variabilis) appears to be restricted to a few localities in far north-eastern Madagascar (former Antsiranana province), where it is relatively rare.

Marta’s water snake (Thamnosophis martae) is confined to lowland dry forests in the vicinity of Montagne des Français and Forêt d’Ambre in far north-eastern Madagascar (former Antsiranana province).

The Maintikibo snake (Liophidium maintikibo) is known only from two specimens collected from the Kirindy Forest and the Andranomena Special Reserve in western coastal Madagascar (former Toliara province). Therezien’s snake (L. therezieni) is known only from a few specimens collected from widely spaced localities in north-eastern and north-western Madagascar (former Antsiranana and Mahajanga provinces).

The Berara cophyline tree frog (Cophyla berara) is known only from a few areas of the Sahamalaza Peninsula in northeastern Madagascar (former Antsiranana province).

The Madinika Madagascar frog (Wakea madinika) is known only from specimens collected in 2001 from a temporary pond on a cacao plantation in north-eastern Madagascar (former Antsiranana province). It may occur more widely in surrounding areas, but extensive surveys have failed to locate it.

The Fôret d’Ambre skeleton frog (Boophis baetkei) is confined to a single locality on the eastern edge of the Fôret d’Ambre Special Reserve in far northern Madagascar (former Antsiranana province). The Berara skeleton frog (B. tsilomaro) appears to be restricted to in and around Berara Forest in north-eastern Madagascar (former Anstiranana province). The Ankarafa skeleton frog (B. ankarafensis) is confined to a single forest fragment on the Sahamalaza Peninsula, north-western Madagascar (former Antsiranana province).

The Antsohihy stump-toed frog (Stumpffia analamaina) is known only from a small fragment of dry deciduous forest along the road between Antosohihy and Mandritsara in northwestern Madagascar (former Mahajanga province). It may be more widespread within a small range of suitable habitat north and north-east of Antsohihy.

The dry forest jumping frog (Aglyptodactylus laticeps) is well known from a few widely separated localities in western Madagascar (former Mahajanga province), where it can be locally abundant but is thought to be declining overall by an increasing lack of suitable habitat.

Tsingy Forest

Tsingy forests are dry deciduous forest areas located on karstic limestone massifs and plateaus rising from sea level upwards to around 600 m.

The tsingy tuft-tailed rat (Eliurus antsingy) is known only from a few specimens collected in disjunct areas of limestone tsingy forests and other formations in western Madagascar (former Mahajanga province).

The Maramaintso plated lizard (Zonosaurus maramaintso) is known only from a single undefined locality in western Madagascar (though most likely the Tsingy de Bemaraha region).

The Tanjaka Madagascar ground gecko (Paroedura tanjaka) is found in a few rocky forested localities in western Madagascar (former Mahajnga province) at Namoroka and near Bemaraha.

The unexpected tree snake (Lycodryas inopinae) is known only from a few specimens collected from far northern Madagascar (former Antsiranana province).

Montagne des Français is located in far northern Madagascar (former Antsiranana province).

The Montagne des Français leaf chameleon (Brookesia tristis) is known only from the Montagne des Français Massif in north-eastern Madagascar (former Antsiranana province).

The Lohatsara Madagascar ground gecko (Paroedura lohatsara) is confined to the lower elevations of Montagne des Français in far northern Madagascar (former Anstiranana province).

The stout dwarf snake (Heteroliodon fohy) is known only from four specimens collected or observed at Montagne des Français and near the village of Ampombofofo in far northeastern Madagascar (former Antsiranana province).

Stafford’s stump-toed frog (Stumpffia staffordi) is confined to a karstic cave and its environs on Montagne des Français.

The Ankarana Massif is located in far northern Madagascar (former Antsiranana province). The important Ankarana Special Reserve was created in 1956.

Rösler’s day gecko (Phelsuma roesleri) is known only from a few specimens collected from lower elevation dry forests on the Ankarana Massif.

Mocquard’s keeled snake (Alluaudina mocquardi) is known only from karst outcrop caves at Ankarana, where it is thought to be threatened by sapphire mining operations.

The Ankarana tsingy frog (Tsingymantis antitra) is known only from the Ankarana Special Reserve where it lives in lowland limestone karst formations.

The Ankarana stump-toed frog (Stumpffia be) is known only from the eastern side of the Ankarana Special Reserve in far northern Madagascar (former Antsiranana province).

Tsingy de Namoroko is located in north-western Madagascar (former Mahajanga province).

The Namoroka leaf chameleon (Brookesia bonsi) is confined to an area of Tsingy de Namoroka Strict Nature Reserve.

Tsingy de Bemaraha is located in central-western coastal Madagascar (former Mahajanga province). It is protected within Tsingy de Bemaraha National Park.

The Bemaraha woolly lemur (Avahi cleesei) is known only from the Tsingy de Bemaraha region.

Nicosia’s chameleon (Furcifer nicosiai) is known only from the Tsingy de Bemaraha region.

The Antsingy leaf chameleon (Brookesia perarmata) and the dwarf leaf chameleon (B. exarmata) are both known only from Tsingy de Bemaraha National Park.

The Bemaraha digging frog (Plethodontohyla fonetana) is endemic to the Tsingy de Bemaraha National Park and adjacent Beanka Classified Forest.

The Antsingy grainy frog (Gephyromantis atsingy) is confined to Tsingy de Bemaraha.

Semi-deserts and Arid Shrublands

Semi-deserts and shrublands predominate in south-western and southern Madagascar as a result of rainshadow from the Central Highlands.

Petter’s sportive lemur (Lepilemur petteri) is relatively widespread in south-western and southern Madagascar (former Toliara province), but is everywhere threatened by loss of habitat.

Standing’s day gecko (Phelsuma standingi) is known only from a few localities in south-western Madagascar (former Toliara province). It is extensively collected for the international pet trade. The Toliara day gecko (P. breviceps) is known only from a few localities in south-western coastal Madagascar (former Toliara province).

Mocquard’s leaf-toed gecko (Matoatoa brevipes) is a rare species confined to areas of intact spiny thicket in coastal south-western Madagascar (former Toliara province), where it is known from Ambolisaka near Morombe, Tsivanoa near Belalanda, Ranobe, Mikea, Tsimanampetsotsa, and Cap St. Marie.

Decary’s dwarf gecko (Lygodactylus decaryi) is known only from specimens collected from Andrahamana, Angavo, and Behara in southern Madagascar (former Toliara province) between the 1930s and 1950s.

Dumas’ mabuya skink (Trachylepis dumasi) is found widely but very patchily throughout south-western and southern coastal Madagascar (former Mahajanga and Toliara provinces).

The Andranovaho skink (Flexiseps andranovahensis) is known only from three widely separated localities in coastal south-western Madagascar (former Toliara province).

Appert’s snake (Liophidium apperti) is known only from a single specimen collected during the 1980s from south-western Madagascar (former Toliara province). The forest where it was discovered has since been destroyed, and the species may be extinct.

Succulent Spiny Woodlands

The succulent spiny woodlands are xeric shrubland region found in south-central and south-western Madagascar (former Toliara and Fianarantsoa provinces). The climate is tropical and dry, with a marked dry season from May to October. To the north this bioregion transitions into dry deciduous forest, while to the south it borders the even drier spiny thickets. Subhumid forests lie to the east. The vegetation is similar to the dry deciduous forest but includes more dry-adapted, xerophytic species. Forests reaching 15 m in height contain a mixture of trees and shrubs, among them two species of baobab and succulents of the genus Pachypodium. This bioregion is threatened by burning, logging, and cattle and goat farming. Hunting is also a problem. Protected areas include the Zombitse-Vohibasia National Park, Kirindy Mitea National Park and Kirindy Forest, and the Andranomena Reserve.

Hubbard’s sportive lemur (Lepilemur hubbardi) are both confined to small areas of south-western Madagascar (former Toliara province).

The narrow-striped mongoose or bokiboky (Mungotictis decemlineata) is divided into two subspecies. The northern narrow-striped mongoose (M. d. decemlineata) inhabits the succulent spiny woodlands of south-western coastal Madagascar (former Toliara province), where it is threatened by loss of habitat, hunting, and predation by feral dogs.

The subdesert mesite (Monias benschi) is a rail-like terrestrial bird confined to a narrow coastal strip in south-western Madagascar (former Toliara province). Although still common in suitable habitat, it is threatened by habitat destruction.

The long-tailed ground-roller (Uratelornis chimaera) is a largely terrestrial bird restricted to a narrow coastal strip in south-western Madagascar (former Toliara province) between the Fiherenana and Mangoky rivers. Owing to its weak flight it is particularly vulnerable to hunting and predation by feral animals.

The Belalanda chameleon (Furcifer belalandaensis) is confined to two small localities near Toliara in south-western Madagascar (former Toliara province). Tuzet’s chameleon (F. tuzetae) is known only from a single specimen collected in 1972 at Andrenalamivola in south-western Madagascar (former Toliara province). It is possibly confined to gallery forest along the Mangoky River.

Hielsher’s day gecko (Phelsuma hielscheri) is known only from a few localities in the south-western and south-central Madagascar (former Toliara province).

The unadorned tree snake (Lycodryas inornatus) occurs in south-western coastal Madagascar (former Toliara province). Arid Spiny Bush

The arid spiny bush (also known as southern spiny thickets) is a bioregion in coastal southern and south-western Madagascar (former Toliara province). Around 95 per cent of the original flora is endemic, making it one of the most unique places on Earth, and many of these plants show extreme adaptation to the arid environment. The fauna is equally remarkable. As with other areas of Madagascar this habitat is under pressure from human activities such as conversion to grazing land, harvesting for charcoal and firewood, and logging. At present only about 3 per cent is protected in national parks and reserves.

The white-footed sportive lemur (Lepilemur leucopus) is confined to a few localities within a small area of southeastern Madagascar.

Grandidier’s vontsira (Galidictis grandidieri) is a type of mongoose that was only described to science in 1986 and has a restricted range around Lake Tsimanampetsotsa in southwestern Madagascar.

The southern narrow-striped mongoose (Mungotictis decemlineata lineata) is confined to arid spiny bush areas of south-western Madagascar, where it is threatened by loss of habitat, hunting, and predation by feral dogs.

The red-shouldered vanga (Calicalicus rufocarpalis) is a shrike-like bird that was originally described from two specimens collected in 1947 near Toliara in south-western Madagascar. It was not seen again until being photographed in 1992 and then in 1997, when a number of individuals were seen in the same area. It has since been located in a few other localities, but the total population is thought to be under 1000.

The radiated tortoise (Astrochelys radiata) is found primarily in the dry spiny bush of southern and south-western Madagascar, from the area of Amboasary in the south across the Karimbola and Mahafaly plateaus north of Toliara to Morombe. It is usually found in a narrow band within 50–100 km of the coast. Historically this species was quite abundant, often being found along roadways, although this is no longer the case. However, they may still be locally abundant in certain areas. The main threats area habitat destruction and collection, both for the international pet trade and for local consumption.

The Tsimanampetsotsa clawless gecko (Ebenavia maintimainty) is restricted to areas of dry limestone spiny bush in south-western Madagascar. Most of the known population is protected within Tsimanampetsotsa National Park.

Grandidier’s Madagascar ground gecko (Paroedura androyensis) is found in scattered localities throughout southern coastal Madagascar, including Ambatotsirongorongo, Andohahela, Antafoky, Cap Sainte Marie, Petriky, Sarodrano, Sept Lacs, Zombitse, and Tsivanoa.

The Lavenambato mabuya skink (Trachylepis vezo) is currently known only from a few specimens collected at a single locality near the mouth of the Onilahy River in south-western Madagascar. However, it likely also occurs on the Mahafaly Plateau and elsewhere.

The Madagascar three-lined snake (Liophidium trilineatum) is known from very few records from the region of Lake Tsimanampetsotsa, as well as Andranobe and Benonoka in south-western and southern Madagascar.

Lakes, Rivers, and Marshes

Madagascar has one of the most distinctive freshwater ecosystems in the world, with many endemic species. From the island’s Central Plateau a number of short, swift rivers flow over a narrow strip of land as they descend to the Indian Ocean to the east. In the west, longer rivers flow down into the Mozambique Channel. Due to climate variability throughout the island, the freshwater systems of the eastern side contrast greatly with those of the west, although they all experience seasonal flooding. Madagascar’s freshwater fishes are ‘living fossils’ and belong to the most primitive of catfish, herrings, cichlids, killifish, silversides, and the like.

The Madagascar grebe (Tachybaptus pelzelnii), an inhabitant of small lakes and forested wetlands, ranges across Madagascar and is still reasonably common in a few areas of the Central Highlands and west, although in small numbers. In 2007 the total population was estimated at between 1500 and 2500.

Meller’s duck (Anas melleri) is found mainly in the wetlands of the eastern and northern Central Highlands. There are additional, isolated populations on isolated massifs on the western edge of the plateau, and (at least formerly) an introduced population on the island of Mauritius that is now most likely extinct. The species has been in decline since the beginning of human colonization and is now rare throughout its range, with the exception of forested areas of the north-west and in the wetlands around Lake Alaotra.

The Madagascar pochard (Aythya innotata) is a type of diving duck that was considered to be relatively common at Lake Alaotra in central-eastern Madagascar (former Toamasina province) during the 1930s, but declined dramatically over the subsequent two decades. There were a few unconfirmed sightings thereafter, with the last certain record taking place in 1960. The species was presumed extinct until 1991, when a single male was captured alive, although subsequent searches failed to discover others. The species was once more considered to be extinct until 2006, when it was rediscovered in very small numbers in a small volcanic lake some 330 km north of Lake Alaotra in north-eastern Madagascar (former Mahajanga province).

The Madagascar snipe (Gallinago macrodactyla) is found throughout the eastern half of the island but is much more common over 700 m elevation. It is threatened by the increasing conversion of wetland-edge habitat for rice cultivation.

The Sakalava rail (Zapornia olivieri) is known from several widely separated wetland areas in lowland western Madagascar, where it is rare and localized. Specimens were taken in 1930 and 1962, and all records since have been field sightings. A single bird was seen in 1995 at Lake Bemamba and another in 1999; two were reported at Amboropotsy Marsh near Bekopaka in 2001; and a small group were observed over the course of several days at Lake Kinkony south-west of Mahajanga in 2003. Other sightings have occurred since at Lake Kinkony, Lake Ampandra, Lake Amparihy, Lake Sahapy, and Lake Mandrozo, all between the Betsiboka and Mangoky rivers. The total population may be no more than 250.

The slender-billed flufftail (Sarothrura watersi) is a small, secretive rail of isolated mid-altitude rainforest wetland areas across much of eastern Madagascar. It has been recorded from near Analamazaotra in 1928, near Andapa in 1930, and, in more recent years, from Anjozorobe, Torotorofotsy Marsh, Ranomafana/Vohiparara, Andringitra National Park, and from localities between the latter two areas. The total population is thought to be less than 1000.

The Madagascar big-headed turtle (Erymnochelys madagascariensis) inhabits slow-moving rivers, lakes, and swamps in western Madagascar (former Antsiranana, Mahajanga, and Toliara provinces), where it is known to be in serious decline. Its large size makes it an attractive food source for local people.

The black diamond cichlid (Paratilapia polleni) is a popular aquarium fish with a restricted range in north-eastern Madagascar (former Antsiranana and Mahajanga provinces), where it is threatened by deforestation and irrigation projects. It is still relatively common on the island of Nosy Be and in a few lakes on the mainland.

The Itasy cichlid (Ptychochromoides itasy) was last seen in Lake Itasy, central Madagascar (former Antananarivo province) in the 1970s, having been extirpated by a combination of habitat degradation and introduced species. It was rediscovered in 2010 in the Sakay River, a tributary of the Tsiribihina River.

The Fort Dauphin cichlid (Ptychochromis mainty) is known only from forested streams in extreme south-eastern Madagascar (former Toliara province). The Diana cichlid (P. oligacanthus) is found in northern Madagascar (former Antsiranana and Mahajanga provinces) and in crater lakes on the island of Nosy Be. The Antsirabe Nord cichlid (P. loisellei) is known from the Mahanara River and its tributaries in north-eastern coastal Madagascar (former Antsiranana province), with further subpopulations in Lake Nosiarina in the Bemariovo River drainage and from Lake Mahatsara near Vohemar. All are threatened by overfishing, loss of habitat, and invasive species.

Cichlids of the genus Paretroplus are largely restricted to lakes and rivers in north-western Madagascar, where they have declined drastically due to habitat destruction, overfishing, and invasive species. The damba cichlid (P. maculatus) lives in shallow floodplain lakes and small rivers in the Betsiboka and Mahajamba drainages, where it has suffered a severe decline in recent years. The Maromandia cichlid (P. maromandia) is known only from the Andanomalaza and Maevarano River drainages. It was present in Lake Andrapongy until the mid- 1990s, but has since been extirpated there. The Calico cichlid (P. kieneri) is still relatively widespread in north-western Madagascar but is everywhere rare. Van Dam’s cichlid (P. damii) is also found over a large area of western and far northern Madagascar, but has disappeared from areas where it formerly occurred, as for instance the island of Nosy Be. Petit’s cichlid (P. petiti) is known only from a juvenile type specimen of unknown origin. The Tsimoly cichlid (P. tsimoly) is known from two north-bank tributaries of the Betsiboka (Ankalimilotrabe and Boinakely rivers), with an additional population located in an unnamed lake east of the Kalamilotra River. Similar-looking specimens have also been collected from the upper reaches of the Kamoro River near the village of Tsimoly. Loiselle’s cichlid (P. loisellei) is so far known from the Mahanara River in northern coastal Madagascar (former Antsiranana province), but may be more widespread. Nourissat’s cichlid (P. nourissati) is known from the Amboaboa and Mangarahara rivers in northern Madagascar (former Mahajanga and Toamasina provinces). The Lamenbe cichlid (P. lamenabe) is confined to the lower reaches of the Mahajamba River near the town of Androka and to Lake Tseny in the Sofia River drainage (former Mahajanga province).

The elongated grunter (Mesopristes elongata) is known from several rivers along the eastern coast of Madagascar, where it is threatened by overfishing and habitat destruction.

Sauvage’s flagtail (Kuhlia sauvagii) is a perch-like species that occurs in a number of rivers draining eastern coastal Madagascar (former Toamasina, Fianarantsoa, and Toliara provinces). Although relatively widespread and common, it is vulnerable to deforestation as it prefers relatively pristine forested rivers and streams.

The Amboaboa round herring (Sauvagella robusta) is a small fish known from the middle and upper reaches of the Amboaboa, Mangarahara, and Anjombony rivers in northern Madagascar (former Mahajanga province), with additional populations in Lake Tseny and, at least formerly, Lake Sarodrono.

The tricolour rainbowfish (Bedotia tricolor) is only known from a single tributary of the Faraony River in south-eastern Madagascar (former Fianarantsoa province), whose watershed has been almost entirely stripped of forest cover. The species must also cope with an introduced predator, the spotted snakehead (Channa punctata). Geay’s rainbowfish (B. geayi) is known only from small forest streams associated with the Mananjary River drainage in south-central Madagascar (former Fianarantsoa province). The white-margined rainbowfish (B. albomarginata) is known only from small forested to moderately degraded tributaries in the upper to middle reaches of the Mananara and Rienana rivers in south-eastern Madagascar (former Fianarantsoa and Toliara provinces). The Rianila rainbowfish (B. leucopteron) is known from the middle reaches of the Iaroka–Rianila system in central-eastern Madagascar (former Toamasina province), but may be present in adjacent drainages. The long-finned rainbowfish (B. longianalis) is confined to a few rivers in north-eastern coastal Madagascar (former Toamasina province) and the island of Nosy Boraha.

Pellegrin’s rainbowfish (Rheocles pellegrini) was long known only from a single specimen collected in 1931 from an undefined area of northern Madagascar (former Antsiranana and/or Toamasina provinces). Further specimens have been collected from the same general region within the past couple of decades. Wright’s rainbowfish (R. wrightae) is known from a few rivers in central-eastern coastal Madagascar (former Toamasina province). The Alaotran rainbowfish (R. alaotrensis) appears to be confined to Lake Alaotra, a few smaller surrounding lakes and to the Maningory River in central-eastern Madagascar (former Toamasina province).

Waterlot’s silverside (Teramulus waterloti) is known from the Mananjeba and Anjingo rivers in north-western Madagascar (former Antsiranana and Mahajanga rivers). The range is notably fragmented, with numerous localities having been sampled without finding the species.

The Masoala goby (Sicyopterus punctissimus) is restricted to a few isolated localities on the Masoala Peninsula of northeastern Madagascar (former Antsiranana province), where it is uncommon.

The Pangalana sleeper goby (Eleotris vomerodentata) is known only from a single specimen collected during the 1980s from Pangalana Lagoon in eastern coastal Madagascar (former Toamasina province).

The Malagasy mountain mullet (Acentrogobius therezieni) is a small goby found in the Andranomavokely River and the associated tsingy pools and springs in the region of the Namoroka Strict Nature Reserve, north-western Madagascar (former Mahajanga province).

Blind and pigment-free gobies of the genus Typhleotris are endemic to subterranean habitats (caves, sinkholes, and wells) of south-western coastal Madagascar (former Toliara province), where they are seriously threatened by human disturbance. Petit’s cavefish (T. madagascariensis) is found in the Mahafaly Plateau karst formation, as well as in isolated localities along the coastal plain below and to the west. The Mararybe cavefish (T. mararybe) is confined to a single cave system near the town of Itampolo. Paulian’s cavefish (T. pauliani) inhabits coastal regions to the south of Morombe in the vicinity of Andalambezo.

Several killifish of the genus Pachypanchax are threatened by loss of habitat and invasive species. The powder-blue killifish (P. omalonota) is found in the Sambirano River drainage of northern Madagascar (former Antsiranana and Mahajanga provinces) and on the island of Nosy Be. Patricia Yazgi’s killifish (P. patriciae) occurs in the Mananjeba, Mahavavy du Nord, Ifasy, Manehoko, and Ampandra rivers of northern Madagascar (former Antsiranana province). Holly’s killifish (P. sakaramyi) is known from the Sakaramy and Antongombato rivers and possibly a few crater lakes as well in far northern Madagascar (former Antsiranana province). The Varatraza killifish (P. varatraza) is known from the Menambery, Fanambana, and Ampanobe rivers of northern Madagascar (former Antsiranana province), but may be more widespread. Arnoult’s killifish (P. arnoulti) is found patchily in the lakes, rivers, and streams of north-western Madagascar (former Mahajanga province).

The Madagascar lampeye (Pantanodon madagascariensis) was originally known from forest streams on the east-central slopes between Mahavelona and Fenoarivo in eastern coastal Madagascar (former Toamasina province). No specimens have been collected since the early 1960s, and the species is now believed to be extinct, a victim of conversion of its preferred swamp habitat into rice fields and competition from introduced Gambusia species.

Lake Alaotra

Lake Alaotra (Lac Alaotra in French) is located on the northern central plateau (former Toamasina province). The largest lake in Madagascar, it serves as an important habitat for waterfowl but is increasingly threatened owing to the destruction of the surrounding marshes by burning and conversion to rice fields. A 42,478-ha protected area was created around the lake in 2007.

The Alaotran bamboo lemur (Hapalemur alaotrensis) is confined to the papyrus and reed beds in and around the lake. Its unique, semi-aquatic habitat long provided it with protection, but during the mid-twentieth century the population began to decline dramatically due to habitat destruction and hunting by local fishermen (who employ a variety of cruel methods including harpoons, dogs or simply shake the reeds until the animals fall helplessly into their boats). It is also frequently captured for use as pets. By the early 1990s, when an international conservation programme began, the species was already on the verge of extinction. Numbers continued to drop in any case, reaching a low of just 2500 in 2013.

Durrell’s vontsira (Salanoia durrelli) is a mongoose-like species known only from the marshlands of Lake Alaotra.

The Alaotran grebe (Tachybaptus rufolavatus) was a nearly flightless waterbird known mainly from Lake Alaotra, where the species was last recorded in 1985. Thorough surveys since have failed to find any evidence of its existence, and it is now considered to be extinct.

The Mananara du Nord River

The Mananara du Nord River is located in north-eastern Madagascar (former Toamasina province).

Ernest Magnus’ cichlid (Ptychochromis ernestmagnusi) is known only from a few specimens collected from the Mananara du Nord River.

The Mananara du Nord catfish (Gogo atratus) is confined to the Mananara du Nord River.

The Lokoho River

The Lokoho River is located in north-eastern Madagascar (former Antsiranana province).

The Lokoho rainbowfish (Rheocles vatosoa) is confined to the upper reaches of the Lokoho River.

The Manentenina River is an effluent of the Lokoho River within Marojejy National Park.

The Manentenina rainbowfish (Bedotia marojejy) is confined to the Manantenina River.

The Sofia River

The Sofia River is located in north-western Madagascar (former Mahajanga province), where it rises on the Tsaratanana Massif.

The Mangarahara River is a major tributary of the Sofia River.

The Mangarahara cichlid (Ptychochromis insolitus) is known only from the Amboaboa River, (a tributary of the Mangarahara River), near the town of Marotandrano. It was previously also found in the Mangarahara River itself, but seems to have been extirpated there. The species was feared extinct, with only three males surviving in captivity (London Zoo and Berlin Aquarium), until a small number were rediscovered in the Amboaboa in 2013. The latter were taken into captivity in order to establish a breeding programme.

The Mandritsara cichlid (Paretroplus gymnopreopercularis) is known only from the Mangarahara River and its tributary, the Amboaboa River.

Derham’s rainbowfish (Rheocles derhami) is confined to the Amboaboa and Mangarahara rivers within the Sofia River drainage.

The Amboaboa catfish (Arius festinus) is known only from the Amboaboa River near its confluence with the Mangarahara River, and possibly Lake Tseny.

The Bemarivo River is a major tributary of the Sofia River.

The pinstripe cichlid (Paretroplus menarambo) was originally found in the floodplain lakes of the Bemarivo River, where it was at one time an important food fish. It was last reported from Lake Sarodrano in the late 1990s, but not seen again in the wild for many years. A remnant population was later discovered in Lake Tseny. Captive breeding populations exist in Europe, North America, and Madagascar.

The Ankofia River is a tributary of the Sofia River.

The Ankofia cichlid (Ptychochromis inornatus) is confined to the Ankofia River drainage, including Lake Andrapongy.

The Andrapongy catfish (Arius uncinatus) is confined to Lake Andrapongy within the Ankofia River drainage.

Sparks’ killifish (Pachypanchax sparksorum) is known only from streams flowing into the Ankofia River and its principal tributary, the Anjingo River.

The Betsiboka River

The Betsiboka River is located in west-central Madagascar (former Mahajanga and Antananarivo provinces).

The Ikopa River is located in the former Antananarivo province.

Bleeker’s cichlid (Paratilapia bleekeri) is known only from specimens collected during the nineteenth century from the Ikopa River.

The Mangoro River

The Mangoro River is located on the central-eastern coast of Madagascar (former Toamasina province).

The ornate catfish (Gogo ornatus) is confined to the Mangoro and Nosivolo rivers, where it is threatened by loss of habitat and introduced fish species.

The Nosivolo River is the primary tributary of the Mangoro River.

The songatana cichlid (Oxylapia polli) is known only from the Marolambo Rapids of the Nosivolo River, where it is threatened by habitat destruction and sedimentation caused by deforestation.

The Katria cichlid (Katria katria) is confined to the Nosivolo River.

The Mandolotra rainbowfish (Rheocles lateralis) is confined to the headwaters of the Mandolotra River, a tributary stream of the Nosivolo River.

The Onilahy River

The Onilahy is located in south-western Madagascar (former Toliara province).

The Onilahy cichlid (Ptychochromis onilahy) was first described in 2006 based on five preserved specimens in the Museum of Natural History, Paris that were originally collected from the Onilahy River in 1962. It is most likely extinct, although the possibility exists of small remnant populations in remote areas.

The trondo cichlid (Ptychochromoides betsileanus) was originally described from specimens collected in the Betsileo region of east-central Madagascar. It was subsequently recorded from the Ilanana River (a tributary of the Onilahy River), the Manantanana River (a tributary of the Mangoky), and the upper reaches of the Matsiatra, especially its tributary the Mandranofotsy, as well as in some waterfalls in streams near Ampamaherana and in the Manantanana and Zomandao rivers of the Ambalavao region. However, no more extant populations have been reported since the discovery of the one at Ilanana in 1997 despite numerous field surveys. It is now thought to be extinct owing to a combination of overfishing, declining water quality and competition with introduced Tilapia.

The Onilahy catfish (Ancharius griseus) is known only from the Ihazofotsy and Ilanana rivers within the Onilahy River drainage.

Miscellaneous Lakes, Rivers, and Marshes

Lake Kinkony (Lac Kinkony in French) is located in northwestern Madagascar (former Mahajanga province). It is the country’s second largest.

The Kinkony cichlid (Paretroplus dambabe) was formerly very common in the Mahavavy du Sud drainage but now appears to be mostly (or entirely) confined to Lake Kinkony, where it has been reduced to low numbers.

The Ankavanana River is located in north-eastern Madagascar (former Antsiranana province), on the Masoala Peninsula.

The Ankavanana rainbowfish (Bedotia masoala) is known only from a short stretch of the Ankananana River. The Antainambalana River is located in north-eastern Madagascar (former Toamasina province).

The Makira cichlid (Ptychochromis makira) is known only from two specimens collected from the Antainambalana River. However, the river has not yet been extensively surveyed for freshwater fish, and it is possible that the species is much more widespread.

The Sambirano River is located in north-western Madagascar (former Antsiranana province).

The Sambirano frillfin goby (Bathygobius samberanoensis) is a little-known species of questionable taxonomic validity, apparently confined to the Sambirano River.

The Manambolo River is located in eastern Madagascar (former Toamasina province).

Sikora’s rainbowfish (Rheocles sikorae) is known only from the Manambolo River, where it was long thought to be extinct until rediscovered in 1996. It is now known from a few forested streams near the town of Marolambo.

The Ivoloina River is located in eastern Madagascar (former Toamasina province).

The Ivoloina rainbowfish (Bedotia madagascariensis) is confined to the lower Ivoloina River drainage, where it is threatened by habitat destruction.

The Mananjary River is located in south-eastern Madagascar (former Fianarantsoa province).

The Mananjary catfish (Gogo brevibarbis) is known only from a single specimen collected from the Mananjary River.

The Mananara du Sud River is located in south-eastern Madagascar (former Fianarantsoa province).

The Vondrozo cichlid (Ptychochromoides vondrozo) is known only from the Mananara du Sud River and its tributaries.

The Andriambondro River is located in south-eastern Madagascar (former Fianarantsoa province).

The Sandrananta catfish (Gogo arcuatus) is known only from the Sandrananta River, within the Andriambondro River drainage.

Coasts and Satellite Islands

Madagascar features still extensive areas of coastal mangroves, wetlands, and littoral forests, although in general they receive little legal protection and are everywhere under threat by development.

The black-banded plover (Charadrius thoracicus) is found discontinuously in western and southern coastal Madagascar (former Mahajanga and Toliara provinces).

Petit’s gecko (Paragehyra petiti) is confined to cliffs along the Onilahy River near Lavenombato, in south-western coastal Madagascar (former Toliara province). The small area where this species occurs is potentially at risk from a proposed limestone mining operation which, if it goes ahead, may result in the complete destruction of its habitat.

The Fasika skink (Paracontias fasika) is known only from a single specimen collected from a small patch of coastal forest just 10 m above sea level in far northern Madagascar (former Antsiranana province). The minimal skink (P. minimus) is similarly known only from a small area of coastal shrub in far northern Madagascar (former Antsiranana province). Rothschild’s skink (P. rothschildi) is a rare sand-dweller from coastal dunes and degraded dry forests near Sakalava Bay in far northern Madagascar (former Antsiranana province).

The Antsiranana skink (Madascincus arenicola) is a fossorial or semi-fossorial skink known only from a small area of far northern coastal Madagascar (former Antsiranana province), where it has been recorded from Baie des Sakalava and Baie des Dunes in the Forêt d’Orangea region, and at an unnamed research site in the Ampombofofo region.

Grandidier’s blind snake (Xenotyphlops grandidieri) is known only from coastal forest and shrubby dunes of Sakalava Bay, in far northern Madagascar (former Antsiranana province). Mocquard’s blind snake (X. mocquardi) is known only from a single locality in far northern coastal Madagascar (former Antsiranana province).

The Maromandia diamond frog (Rhombophryne tetradactyla) is known only from the islands of Nosy Mangabe and Nosy Boraha (Ile St. Marie) off the north-eastern coast of Madagascar.

Megson’s stump-toed frog (Stumpffia megsoni) is known only from a few small caves in far northern coastal Madagascar (former Antsiranana province).

Nosy Hara

Nosy Hara is located off the far northern coast of Madagascar (former Antsiranana province).

The Nosy Hara leaf chameleon (Brookesia micra), the world’s smallest lizard, is confined to Nosy Hara.

The Nosy Hara diamond frog (Rhombophryne hara) is confined to Nosy Hara.

Nosy Mangabe

Nosy Mangabe is located in Antongil Bay off the north-east coast of Madagascar (former Toamasina province), near the town of Maroantsetra. Covered by dense rainforest, it has been designated a special reserve and there are no permanent settlements on the island.

The Nosy Mangabe diamond frog (Rhombophryne mangabensis) is known only from 13 specimens collected in 1991 and 2005.

Nosy Boraha

Nosy Boraha (Île Sainte-Marie in French) is located off the north-eastern coast of Madagascar (former Toamasina province).

Delalande’s coua (Coua delalandei) was a type of cuckoo restricted to Nosy Boraha. There it was often trapped for food and feathers, and the introduction of rats may have also contributed to its demise. Ultimately, however, it was the complete destruction of the island’s forests that was the cause of the extinction. It was last collected in 1834 (a report of a sighting from around 1930 appears to be unfounded).

Nosy Be and Nosy Komba

Nosy Be is a small volcanic island located off the north-eastern coast of Madagascar (former Antsiranana province). One of Madagascar’s largest and busiest tourist resorts, it has been heavily deforested. One of the only remaining natural areas for the island’s many endemic species is the small Lokobe Strict Nature Reserve. The smaller islet of Nosy Komba is located to the south-east.

The Nosy Be sportive lemur (Lepilemur tymerlachsoni) is confined to Nosy Be.

The Mamiratra mouse lemur (Microcebus mamiratra) is confined to Nosy Be.

The Nosy Be skink (Pseudoacontias unicolor) appears to be confined to the Lokobe Strict Nature Reserve on Nosy Be.

The Nosy Be mabuya skink (Trachylepis lavarambo) is confined to Nosy Be.

Reuter’s blind snake (Madatyphlops reuteri) and Boettger’s blind snake (M. madagascariensis) are both known only from Nosy Be.

The mystery cophyline tree frog (Cophyla occultans) appears to be confined to Nosy Be, with populations in north-eastern Madagascar formerly included within its range now thought to represent an as-yet undescribed species.

The Andoany diamond frog (Rhombophryne pygmaea) and Boettger’s diamond frog (R. testudo) are both known only from Nosy Be and Nosy Komba.

The Glorioso Islands

The Glorioso Islands (Îles Glorieuses in French) are a group of two small, uninhabited islands and three rocky islets located about 160 km north-west of Madagascar. The climate is lush and tropical, and the terrain low and flat. In 2012, France founded the Parc Naturel Marin des Glorieuses, a marine protected area that covers all flora and fauna.

The Glorioso snake-eyed skink (Cryptoblepharus gloriosus) is thought to be at risk, as its restricted range makes it vulnerable to stochastic events and sea level rise due to climate change.

The Comoros Islands

The Comoros Islands (Les Comores in French) are an archipelago of four main volcanic islands situated at the north end of the Mozambique Channel, north-west of Madagascar. Several smaller islands and submerged reefs are also included in the chain.

Livingstone’s flying fox (Pteropus livingstonii) is one of the world’s largest bats. Considered common at the time it was first discovered by the explorer David Livingstone in 1863, it is today restricted to the islands of Anjouan and Mohéli where the population stands at about 1200. In 1992 the Durrell Wildlife Conservation Trust sent an expedition to the Comoros in order to bring some of the bats back to Britain to establish a captive breeding population. This has proved successful and the species is now established in several other zoos. Its status in the wild, however, remains perilous due to habitat destruction, hunting, and the threat of stochastic events.

The Comoros rousette fruit bat (Rousettus obliviosus) is confined to the islands of Grande Comore, Anjouan, and Mohéli, where it is dependent upon caves for roosting.

The Comoros green pigeon (Treron griveaudi) is currently confined to Mohéli, although it is thought to have been historically present on Grand Comoro and Anjouan.

The Comoros cuckoo-shrike (Ceblepyris cucullatus) is found on the islands of Grande Comore and Mohéli, where the total population is thought to be less than 2500.

The Comoros ground gecko (Paroedura sanctijohannis) occurs on all four main islands, where with the exception of Mayotte it is confined to small, high-elevation areas.

The Comoros blind snake (Madatyphlops comorensis) is a burrowing species known only from a few specimens collected on Grande Comore and, in one instance, Anjouan.

Grande Comore

Grand Comore is the largest of the Comoro Islands and is comprised of two shield volcanoes.

The Comoro blue vanga (Cyanolanius comorensis) is divided into two subspecies. The Grande Comore blue vanga (C. c. bensoni) is a shrike-like bird known only from sightings in 1974 and 1981, and is possibly extinct.

The Grande Comore bulbul (Hypsipetes parvirostris) is confined to Grande Comore, where it remains reasonably common.

Mount Karthala

Mount Karthala, a highly active volcano, is the highest point in the Comoros. Nominally protected within Mount Karthala National Park, it provides a last refuge for several species of bird but continues to be threatened by deforestation.

The Grande Comore scops owl (Otus pauliani) is confined to Mount Karthala.

The Grande Comore drongo (Dicrurus fuscipennis) is confined to Mount Karthala, where in 1985 the total population was estimated at around 100.

Humblot’s flycatcher (Humblotia flavirostris) is confined to Mount Karthala.

The Karthala white-eye (Zosterops mouroniensis) is confined to Mount Karthala.

Anjouan

Anjouan is the easternmost of the Comoros Islands.

The Anjouan mouse-eared bat (Myotis anjouanensis) is a little-known species confined to Anjouan.

The Anjouan scops owl (Otus capnodes) was long feared to be extinct. Between 1884 and 1897 it was collected in large numbers for museums, but not seen after that. In 1992, however, it was rediscovered on Anjouan. The species remains highly threatened, with an estimated population of 50–100 pairs.

The Anjouan sparrowhawk (Accipiter francesiae pusillus) is confined to Anjouan, where it is seriously threatened. It was thought to be extinct until searches in the 1980s and in 2005 confirmed that it is still extant.

Mayotte

Mayotte consists of a large main island (Grande-Terre), a smaller one (Petite-Terre), and several islets.

The Mayotte drongo (Dicrurus waldenii) is a type of passerine bird confined to Mayotte, where it occurs widely but sparsely.

The black-striped day gecko (Phelsuma nigristriata) and Robert Mertens’ day gecko (P. robertmertensi) are both confined to the island.

Peters’ bright snake (Liophidium mayottensis) is found in very low numbers throughout the island.

Mohéli

Mohéli is the smallest of the four major Comoros Islands.

The Mohéli scops owl (Otus moheliensis) is confined to Mohéli.

The Mohéli bulbul (Hypsipetes moheliensis) is a type of passerine bird confined to Mohéli.

The Mohéli blue vanga (Cyanolanius comorensis comorensis) is confined to Mohéli.

The Seychelles

The Seychelles comprise four main islands and hundreds of smaller islands and islets about located 950 km north of Madagascar. There are two main divisions: the larger granitic Seychelles and the smaller, outlying coralline Aldabras. The natural forest has been largely destroyed throughout, resulting in a steppe-like vegetation, serious erosion, and a sinking water table. Habitats are deteriorating as well due to invasion by introduced plants, principally cinnamon (Cinnamomum verum), as well as the drying effects of climate change. This, together with introduced cats and dogs and uninvited rats, has become a great threat to many endemic species.

The Seychelles sheath-tailed bat (Coleura seychellensis) was historically found throughout the Seychelles, but declined rapidly with the destruction of its lowland forests in the late nineteenth and early twentieth centuries. Introduced predators and disturbance of its cave roosting sites have since made it one of the world’s rarest species, with the total population in 2016 estimated at less than 100. Two subspecies are recognized. The nominate form (C. s. seychellensis) is now confined to Mahé, having been extirpated from Praslin in the 1980s. The subspecies C. s. silhouettae is now confined to Silhouette, having been extirpated from La Digue in the 1980s.

The Seychelles parakeet (Psittacula wardi) was endemic to Mahé and Silhouette islands, and possibly Praslin as well. It became extinct as a result of hunting pressure. The last known individuals were shot in 1893, and none were found during a survey in 1906.

The Seychelles kestrel (Falco araeus) once occurred on almost all of the granitic islands but began to decline sharply after 1940, likely due to pesticide use and loss of habitat due to cinnamon cultivation and logging. By 1959 it was found only on Mahé, where there were less than 30 birds in 1964–65. Since then, it has made a remarkable recovery and now numbers around 800. The majority still live on Mahé (along with a few of its satellite islands Sainte Anne, Cerf, Conception, and Thérèse), but the species is once again to be found as well on Silhouette, Ile du Nord, Praslin, and occasionally La Digue and Félicité.

The Seychelles turtledove (Nesoenas picturatus rostrata) was formerly distributed over all of the main islands but was largely absorbed by the introduced Madagascar turtledove (N. p. picturatus). Pure populations of the Seychelles turtledove now survive only on Cousin and Cousine Islands.

The Seychelles paradise flycatcher (Terpsiphone corvina), which formerly occurred on several islands, has long been largely confined to the Veuve Nature Reserve on the western side of La Digue. A few birds have been found on neighbouring Marianne and Praslin but are not established there. Occasional sightings on Félicité have also been reported. On La Digue the population has been increasing steadily from an estimated 28 birds in 1965 to a current, stabilized 150–200. A translocation of 23 birds to Denis Island was conducted in 2008, with the aim of establishing a permanent population.

The Seychelles magpie-robin (Copsychus sechellarum) once ranged throughout the islands but had been reduced to just 12–15 individuals on Frégate by the mid 1960s. A small, introduced population on Alphonse survived until about 1960, when it was wiped out by feral cats. By the late 1970s the Frégate population numbered around 40 birds but had been reduced to half that by 1981. Numbers remained around this level until the 1990s. In 1994, with the advent of a recovery programme, the population had increased to 48, with two birds translocated to Aride. In 2000, following further translocations, the population reached 86 (comprising 46 on Frégate, 23 on Cousin, 15 on Cousine, and 2 on Aride). Since then the population has continued to climb, reaching 283 by 2015, with the species now established on Denis Island as well.

The Seychelles warbler (Acrocephalus sechellensis) was historically present on several of the inner granitic islands, until human disturbance in the twentieth century reduced the species to a single population of a hundred birds or less on Cousin Island by the 1960s. Conservation efforts have since helped it to recover on Cousin Island, and the species has been translocated to the islands of Aride, Cousine, Denis, and Frégate.

The Seychelles swiftlet (Aerodramus elaphrus) occurs on the islands of Mahé, Praslin and La Digue, with one known breeding site on each island. A colony on Félicité has disappeared, as has a second one on Mahé.

The Seychelles fody (Foudia sechellarum) is a small songbird that was reduced, by the early 1970s, to between 400 and 500 birds living on the islands of Frégate, Cousin, and Cousine, due mainly to a combination of human persecution (it was erroneously thought to consume crops) and habitat destruction. It has since recovered there and has been introduced to the predator-free islands of Aride, D’Arros, and Denis. The total population now stands at around 3500.

The Seychelles white-eye (Zosterops modestus) was long thought to survive only in three tiny areas on Mahé, with only 25–35 individuals known in 1996. That year, however, a previously unknown population of at least 250 was discovered on Conception. Since that time the species has been translocated to Frégate Island, North Island and Cousine.

The Seychelles giant tortoise (Aldabrachelys gigantea), one of the world’s largest, was formerly common throughout the Seychelles and Aldabra islands. Four subspecies have been identified, among them the Aldabra giant tortoise (A. g. gigantea), which is discussed separately in this volume. Günther’s giant tortoise (A. g. hololissa) historically lived on Cerf, Cousine, Frégate, Mahe, Praslin, Round, and Silhouette. Thought to have been exterminated by 1840, a few captive and semi-wild specimens of unknown provenance still survive.

The Seychelles black mud turtle (Pelusios subniger parietalis) occurs in the lowland marshes of Mahé, Cerf, Praslin, La Digue, Fregate, and Silhouette islands, where it has already lost most of its available habitat due to drainage. The total population is thought to be around 250.

The Seychelles tiger chameleon (Archaius tigris) is an arboreal species from the forests of Mahé, Silhouette, and Praslin. It is threatened by habitat degradation and overcollection for the international pet trade.

The giant bronze gecko (Ailuronyx trachygaster) is a canopy-dwelling species endemic to Silhouette and Praslin.

Brauer’s burrowing skink (Janetaescinus braueri) is confined to Mahé and Silhouette. Vesey-Fitzgerald’s burrowing skink (J. veseyfitzgeraldi) is known from Mahé, Silhouette, Curieuse, Félicité, La Digue and Frégate. Both are considered vulnerable to habitat destruction and degradation, as well as to predation by introduced tailless tenrecs (Tenrec ecaudatus).

Wright’s mabuya skink (Trachylepis wrightii) is confined to islands with seabird colonies (i.e. Aride, Cousin, Cousine, St. Pierre, Mammelles, Recifs, and Frégate).

The Seychelles wolf snake (Lycognathophis seychellensis) is confined to Mahé, Silhouette, Praslin, Aride, La Digue, and Frégate.

The Seychelles house snake (Lamprophis geometricus) is confined to Mahé, Silhouette, Praslin, and Frégate.

The Seychelles pygmy frog (Sooglossus sechellensis) is found on the islands of Mahé, Silhouette, and Praslin, where it is still relatively common. Thomasset’s frog (S. thomasseti) is confined to Mahé and Silhouette, where it lives in wet rocky areas.

Gardiner’s Seychelles frog (Sechellophryne gardineri), one of the world’s smallest frogs, is confined to high- and midaltitude rainforests on Mahé and Silhouette.

Cooper’s black caecilian (Praslinia cooperi) is a burrowing amphibian known from a handful of higher-elevation localities on Mahé and Silhouette, with a possible old record from Praslin as well. It has not been collected since the early 1990s.

The golden killifish (Pachypanchax playfairii) is a freshand brackish-water species endemic to streams and ponds in the granitic Seychelles. Little is known of its current status, although an introduced population on the island of Zanzibar is now believed to have been extirpated. Popularly kept as an aquarium fish, little is known of its status in the wild.

Mahé

Mahé is the largest island in the Seychelles. Considerable development and habitat alteration have taken place during the twenty-first century. Fortunately, several protected areas have been set aside, the largest of which is Morne Seychellois National Park, which covers almost a quarter of the island.

Arnold’s giant tortoise (Aldabrachelys gigantea arnoldi) and Daudin’s giant tortoise (A. g. daudinii) were both historically confined to Mahé, where they were thought to have been exterminated by 1840. Captive and semi-wild specimens of the former subspecies still survive; however, of unknown provenance.

The Seychelles scops owl (Otus insularis) is confined to Mahé, where the population appears to be stable.

The Mahé caecilian (Grandisonia brevis) is found in only a few scattered localities on Mahé and perhaps Silhouette. Rare and hard to detect, it has not been observed in many years.

Praslin

Lying 44 km north-east of Mahé, Praslin still has substantial tracks of tropical forest, with a large area in the south of the island having been designated as a national park.

The Seychelles black parrot (Coracopsis barklyi) is largely confined to Praslin Island, with occasional reports from Curieuse 1 km to the north. Less than 50 were estimated there in 1965, although today that number is nearer to 100000.

Silhouette

Silhouette Island is located 20 km north-west of Mahé and is the third largest in the Seychelles. Lush and mountainous, almost 95 per cent of it is protected within Silhouette National Park.

The Seychelles palm frog (Sechellophryne pipilodryas) is confined to the forests of Silhouette Island, where it is closely associated with the thief palm (Phoenicophorium borsigianum).

Marianne

Marianne is a small, nowadays uninhabited granitic islet that formerly served as a coconut plantation.

The Marianne white-eye (Zosterops semiflavus) was a type of passerine bird only known with certainty from this island, although it may have occurred as well on Praslin, La Digue, Silhouette, and Mahé as well. It became extinct around 1888.

The Aldabra Islands

The remote Aldabra Islands are a group of four raised coral atolls along with a number of islets located south of the Seychelles and north-west of Madagascar.

Aldabra Atoll

Aldabra is the world’s second largest coral atoll, and consists of four main islands along with some 40 small islets. During the 1960s there were plans to transform it into a military airbase, which would probably have doomed most of the native species. Fortunately, the project was eventually cancelled after strong opposition from conservationists.

The Aldabra flying fox (Pteropus aldabrensis) is confined to Aldabra Atoll, where the total population is reported to consist of only a few hundred animals.

The Aldabra trident bat (Paratriaenops pauliani) is littleknown and apparently confined to Picard Island.

The Aldabra brush warbler (Nesillas aldabrana) was discovered only in 1967. Confined to Ile Malabar, its population was already extremely small and vulnerable to predation and habitat alteration by invasive species. It was last recorded in 1983, and searches in 1986 confirmed its extinction.

The Aldabra white-throated rail (Dryolimnas cuvieri aldabranus) is the last surviving flightless rail of the Indian Ocean region. While limited in its range and vulnerable to feral cats, it does not appear to be threatened at present.

The Aldabra giant tortoise (Aldabrachelys gigantea gigantea) is endemic to the islands of the Aldabra Atoll, where for many centuries it was heavily exploited by European sailors but ultimately saved by some early established protected areas. Today over 100,000 there, with additional populations in the Sainte Anne Marine National Park on Moyenne Island in the Seychelles and on the island of Changuu, near Zanzibar. Populations have also been introduced to protected areas on Mauritius and Réunion.

Assumption Island

Located about 27 km south of Aldabra Atoll, Assumption is a coral island with extensive sandy dunes. Due to the disruptive effect of guano mining, which lasted until 1983, it is dominated by expanses of bare rock and caves and sparsely covered with low-growing vegetation.

The Assumption white-throated rail (Dryolimnas cuvieri abbotti) became extinct sometime during the early twentieth century.

The Assumption day gecko (Phelsuma abbotti sumptio) is confined to the island, where it appears to be fairly common.

The Amirante Islands

The Amirante Islands (Les Amirantes in French) are a group of 29 coral islands and atolls that belong to the outer islands of the Seychelles.

The endemic Amirante turtledove (Nesoenas picturatus aldabrana) is now extinct due to hybridization with the introduced Madagascar turtledove (N. p. picturatus).

The Mascarene Islands

The Mascarene Islands (Les Mascareignes in French) are located east of Madagascar, and consist of three large volcanic islands (Mauritius, Réunion, and Rodrigues) along with a number of remnants. The islands share a common geological origin and are notable for a unique, if sadly devastated, flora and fauna. Indeed, since the settlement of these islands by Europeans in the seventeenth and eighteenth centuries, no less than 45 endemic species and subspecies have become extinct. Many of these were birds that, living unmolested, evolved to flightlessness and were doomed when man and his introduced animals ravaged these islands. The explosion in human population, partly explained by labour imported from India after 1848, has also been fatal to wildlife. Cultivation spread up the volcanic slopes, destroying vegetation and leading to the most serious erosion. What little wilderness remains, however, has been well protected in recent decades.

Three species of flying fox (Pteropus) were historically found throughout the Mascarenes. The lesser Mascarene flying fox (P. subniger) is believed to have died out on Réunion in the 1860s and on Mauritius before 1873. The greater Mascarene flying fox (P. niger) was extirpated from Réunion in the early eighteenth century but has survived on Mauritius, where it is fairly common. The Rodrigues flying fox (P. rodricensis) was formerly found on Mauritius and Round Island but is today confined to Rodrigues. The small population is vulnerable to cyclones.

The Mascarene petrel (Pseudobulweria aterrima) is known to breed only on Réunion, where it is very rare, although subfossil evidence has been discovered on Rodrigues and a dead bird found on Mauritius.

The Mascarene grey parrot (Psittacula bensoni) was found on both Mauritius and Réunion. Easily hunted, the capture of one specimen would result in its calling out to summon the entire flock. It was exterminated on Réunion by the 1730s and by the 1760s on Mauritius.

The Mascarene coot (Fulica newtonii) is known only from early traveller’s reports and bones collected on Mauritius and Réunion. It was extinct before 1700.

Mauritius

Mauritius (Île Maurice in French) is a relatively large island surrounded by the world’s third largest coral reef and a ring of about 50 satellite islets, many of which have been set aside as protected areas. When early navigators first visited in the sixteenth century it was still uninhabited. Dense evergreen forests descended from the mountain peaks to the sea, and in drier areas there were palm savannas. Its position today is best described by mentioning that the population density is one of the highest on Earth. As a result, only about 17 per cent of the island remains forested, although of this only about 2 per cent can accurately be described as ‘natural forest’. Indeed, the remaining forests are nowhere entirely natural, because exotic, introduced plants have invaded the native vegetation. Severe loss and degradation of habitat has been compounded by introduced long-tailed macaques (Macaca fascicularis), sambar deer (Rusa unicolor), small Indian mongoose (Herpestes auropunctatus), rats, and feral cats. Frequent cyclones are another problem. All of these factors have served to threaten not only the indigenous flora but also the animals, of which Mauritius has already lost 11 species of endemic birds, and 3 reptiles.

The Mauritius free-tailed bat (Mormopterus acetabulosus) is confined to a few lava tube caves where it is vulnerable to disturbance.

The Mauritius owl (Mascarenotus sauzieri) was last recorded in 1837, and was certainly extinct by 1859.

The Mauritius kestrel (Falco punctatus) was historically widespread but had been reduced, by 1974, to just four wild birds (including a single breeding pair) due to loss of habitat, pesticide use, and introduced predators. It has since made a spectacular comeback with the help of a recovery programme, and by the early twenty-first century numbered between 600 and 800.

The Mauritius night-heron (Nycticorax mauritianus) is known only from subfossil bones. Last recorded in 1693, it was probably exterminated by hunters before 1700.

The Mauritius sheldgoose (Alopochen mauritiana) was last recorded in 1693, when it was said to be rare, and could not be found in 1698. The cause of its extinction is thought to have been hunting.

Theodor’s duck (Anas theodori) is known only from explorer’s accounts and bones, and was last recorded in 1696. It or a similar species may have also lived on Réunion.

The Mauritius red rail (Aphanapteryx bonasia) is known from traveller’s accounts, illustrations, and bones. It went extinct around 1693 due to hunting and perhaps predation by feral cats.

The broad-billed parrot (Lophopsittacus mauritianus) was probably also flightless to judge from traveller’s accounts as well as from a 1638 sketch and some subfossil bones. Heavily hunted, the last records date from 1673 to 1675. It was absent in 1693.

The Mauritius echo parakeet (Psittacula eques echo) is the only extant parrot of the Mascarene Islands, all others having become extinct due to human activity. It is also one of themost remarkable success stories in the history of conservation. By the early 1980s it appeared to be doomed. The roughly 10 individuals still surviving had hardly ever bred successfully due to a lack of suitable trees for nesting, nest predation, and competition from the introduced ring-necked parakeet (P. krameri). However, a dedicated research and captive-breeding programme was launched by the Durrell Wildlife Conservation Trust and Mauritian Wildlife Foundation to save the species. By the mid-1990s the numbers had risen to around 50–60. Since then the recovery has continued, with the total wild population sitting at 280–300.

Perhaps the most famous extinct species of them all is the dodo (Raphus cucullatus), which has become a symbol for the meaningless and short-sighted extermination of animals by man. A flightless, heavy pigeon weighing somewhere between 13 and 23 kg, it was driven to extinction as a result of hunting by settlers and nest predation by introduced pigs. The last individuals are thought to have been killed on the offshore islet of Ile d’Ambre in 1662. All that remains today are numerous bones and specimen fragments, along with contemporary reports and paintings.

The Mauritius wood pigeon (Columba thiriouxi) is known only from subfossil remains, and was extinct around 1730.

The Mauritius blue pigeon (Alectroenas nitidissimus) was last reported in 1832, and is thought to have become extinct a few years later.

The Mauritius pink pigeon (Nesoenas mayeri) was at one time one of the world’s rarest birds, having been reduced to just 12 wild individuals in 1986. A successful captive breeding and reintroduction programme has since raised the population to around 400 but the total range remains small, consisting as it does of the Black River Gorges of south-west Mauritius and the Ile aux Aigrettes just off the eastern coast. The Mauritius turtledove (N. cicur) is known only from subfossil remains. It was extinct around 1730.

The Mauritius cuckoo-shrike (Lalage typica) is a rare endemic species with a total population of around 1000.

The Mauritius black bulbul (Hypsipetes olivaceus) had been reduced to around 200 pairs in the mid- 1970s. Today the population remains dangerously small but stable.

The Mauritius paradise flycatcher (Terpsiphone bourbonnensis desolata) is endemic to the island, where the population is estimated at around 250.

The Mauritius olive white-eye (Zosterops chloronothos) had been reduced to around 200 pairs in the early 1990s. It has since recovered somewhat, but appears to be confined to the wettest native upland forests.

The Mauritius fody (Foudia rubra) is restricted to southwestern Mauritius, where it declined from around 250 pairs in the mid-1970s to 108–122 pairs by 2001. In 2005, 45 handreared chicks were released onto Ile aux Aigrettes (on which the species apparently occurred historically), where by 2014 the population rose to between 180 and 200.

Mauritius had at least two endemic species of giant land tortoise (Cylindraspis). Both were historically common, but by the beginning of the eighteenth century ships on their way to and from India were slaughtering or capturing thousands of them for food and export, particularly to hospitals. In addition, introduced species such as rats, cats, and pigs ate the eggs and hatchlings. As a result, both the domed Mauritius giant tortoise (C. triserrata) and the saddle-backed Mauritius giant tortoise (C. inepta) were likely extinct on the main island by about 1700, and on most of the surrounding islets by 1735. At least one of the species may have survived on Round Island until much later. An expedition there in 1844 found several very large specimens, although by that point the island was already overrun with introduced rabbits and all the tortoises appear to have been gone by 1893.

The Mauritius giant skink (Leiolopisma mauritiana) is known only from subfossil bones. It is thought to have gone extinct around 1600.

Bojer’s skink (Gongylomorphus bojerii) was historically widespread in the Mascarenes, where it was divided into two subspecies. The Mauritius Bojer’s skink (G. b. bojerii) was found on Mauritius and surrounding islands, but was extirpated from the former due to the introduction of predatory snakes and shrews and now survives only a handful of predator-free islands.

Hoffstetter’s blind snake (Madatyphlops cariei) was last recorded during the seventeenth century, and is thought to have been exterminated by introduced species.

Round Island

Round Island (Île Ronde in French) is located about 22 km off the north-eastern coast of Mauritius. The largest of a group of volcanic islets that serve as the last refuge of a remarkable reptilian fauna, it has never been inhabited and is almost inaccessible, rising steeply out of the sea. Only about 150 years ago it was covered by forest, but owing to monumental stupidity goats and rabbits were introduced there about 1850. The result was that, by the 1970s, there was no forest left and indeed very little vegetation at all, with only some small groups of endemic bottle palms and most of the soil eroded away. Insect life had all but disappeared and the endemic lizards that feed on them vanished too, along with the snakes that preyed upon the lizards. Fortunately, since the mid-1970s the island has been the subject of a concerted conservation programme. Goats and rabbits have been eliminated, the forest has begun to regenerate and the surviving reptiles, all highly threatened, have been safeguarded by a successful captive-breeding programme. More recently, Aldabra giant tortoises (Aldabrachelys gigantea gigantea) have been introduced.

The Round Island day gecko (Phelsuma guentheri), which historically occurred on Mauritius itself, was reduced by the 1970s to a few hundred individuals on Round Island. The population is now fairly stable.

Telfair’s skink (Leiolopisma telfairii) was near extinction by the late 1960s, but has made a remarkable recovery in recent years thanks to conservation efforts.

The Round Island burrowing boa (Bolyeria multocarinata) was last reported in 1975, and is now certainly extinct.

The Round Island keel-scaled boa (Casarea dussumieri) was thought to be extinct in the 1970s when it was unexpectedly rediscovered. Today the species is established on Round Island as well as on Gunner’s Quoin, Flat Island, and Ile de la Pas. Subfossil remains have shown that it was historically present on Mauritius itself.

Serpent Island

Serpent Island (Île Serpent in French) is located close to Round Island.

The Serpent Island night gecko (Nactus serpensinsula) and Durrells’ night gecko (N. durrellorum) are both confined to Serpent Island.

Gunner’s Quoin, Pigeon Rock, Ile aux Vacoas, and Flat Island

Gunner’s Quoin, Pigeon Rock, Ile aux Vacoas, and Flat Island are small islets located off the northern coast of Mauritius.

The lesser night gecko (Nactus coindemirensis) is confined to Gunner’s Quoin, Pigeon Rock, Ile aux Vacoas, and Flat Island.

Réunion

Réunion (La Réunion in French, and formerly Île Bourbon), located about 175 km south-west of Mauritius, is the westernmost and largest of the Mascarene Islands. Although the evergreen forests that originally covered the island have been destroyed and altered by humans, it has more undisturbed vegetation than the other Mascarene Islands. Nevertheless, at least 1 mammal, 11 birds, and 2 reptiles have become extinct since the arrival of Europeans, mainly due to hunting pressure and/or the predations of introduced rats and cats.

Barau’s petrel (Pterodroma baraui) is effectively endemic to Réunion for breeding purposes, but disperses widely throughout the tropical Indian Ocean at other times.

The Réunion owl (Mascarenotus grucheti) is known only from subfossil bones. It was exterminated in the early seventeenth century.

The Réunion kestrel (Falco duboisi) went extinct after 1672.

The Réunion harrier (Circus maillardi) is currently estimated to number between 200 and 560 mature individuals.

The Réunion sacred ibis (Threskiornis solitarius) is known from bones collected on Réunion, and is likely synonymous with the ‘solitaire’ known from numerous early accounts. If so, the last report dates from 1763.

The Réunion night heron (Nycticorax duboisi) was last recorded in 1674. It was probably driven to extinction by hunters before 1700.

The Réunion purple gallinule (Porphyrio caerulescens) was an inhabitant of montane forest, where it was described by early settlers. It was likely hunted to extinction around 1730.

The Réunion sheldgoose (Alopochen kervazoi) is known only from subfossil bones collected in 1974. It was exterminated after 1672.

The Réunion rail (Dryolimnas augusti) was described from fossils and thought to correspond to a rail species mentioned by Dubois in 1674. Likely flightless, it was exterminated in the late seventeenth century.

The Réunion parrot (Mascarinus mascarin) was a unique species with no near relatives. First mentioned in 1674, several live specimens were brought back to France in the late eighteenth century. The last reports of wild birds date from the 1770s, and the species may have been extinct by 1804 (although tradition has it that a single individual survived in the King of Bavaria’s menagerie until 1834). Two museum specimens are all that remain today.

The Réunion echo parakeet (Psittacula eques eques) has been extinct since about 1770.

The Réunion pink pigeon (Nesoenas duboisi) was last recorded in 1674. It is thought to have been extinct by the early eighteenth century.

The Réunion crested starling (Fregilupus varius) was apparently common as late as the 1830s, but was thereafter greatly affected by introduced diseases and hunting. The last known specimen was shot in 1837, and the species became extinct some time during the 1850s.

The Réunion cuckoo-shrike (Lalage newtoni) is restricted to two very small areas in the north-west of the island, where the total population is estimated at fewer than 30 pairs.

The Réunion fody (Foudia delloni) was a type of passerine bird that was once so abundant it was considered a crop pest. Last seen shortly after 1672, it was likely driven extinct by introduced rats.

The Réunion giant tortoise (Cylindraspis indica) was common up until the early eighteenth century, but was easy prey for European sailors and killed or collected in vast numbers. In addition, introduced pigs, cats, and rats destroyed the eggs and hatchlings. By the beginning of the nineteenth century only a few still survived in highland areas, where they were finally driven extinct during the 1840s.

The Réunion Bojer’s skink (Gongylomorphus bojerii borbonica) was observed and collected many times during the nineteenth century, but has not been recorded since.

Rodrigues

Rodrigues (Île Rodrigues in French) is located about 560 km east of Mauritius. The smallest of the Mascarene Islands, it is notably hilly with a high central ridge cut with deep valleys. A large fringing reef surrounds it, forming a lagoon within which lie 18 small islets. In 1691, when Europeans first settled here, it had been covered with forests, of which only patches remain on the reef. There is hardly any native vegetation left at all on Rodrigues and a large percentage of the island’s vertebrates have vanished.

The Rodrigues owl (Mascarenotus murivorus) is known only from bones and early reports. It was last recorded in 1726.

The Rodrigues night-heron (Nycticorax megacephalus) is known only from bones and early reports. It was last recorded in 1726, and mentioned as absent in 1761.

The Rodrigues flightless rail (Erythromachus leguati) was driven to extinction by heavy hunting. It was last recorded in 1726, with its absence being noted by Alexandre Guy Pingré in 1761.

The Rodrigues parrot (Necropsittacus rodricanus) was last seen about 1761, and presumably became extinct soon after.

The Rodrigues parakeet (Psittacula exsul), long reduced by habitat destruction and hunting, is thought to have ultimately gone extinct as a result of vicious cyclones in 1875.

The Rodrigues solitaire (Pezophaps solitaria), a flightless relative of the dodo, is known only from bones and the accounts of travellers. Reported in 1761, it had become extinct by 1778.

The Rodrigues blue pigeon (Alectroenas payandeei) is known only from a single subfossil bone. It was not reported by early travellers to the island and likely disappeared by the 1690s.

The Rodrigues turtledove (Nesoenas rodericanus) is known from subfossil bones. It went extinct between 1726 and 1761 as a result of predation by rats.

The Rodrigues starling (Necropsar rodericanus) was last recorded in 1726, and not found during a visit to the island in 1761.

The Rodrigues warbler (Acrocephalus rodericanus) was historically quite common but dwindled to very low numbers by the 1970s. Fortunately, it proved highly adaptable to introduced vegetation and has made a considerable comeback in recent years.

The Rodrigues fody (Foudia flavicans) was reduced to just five or six pairs in 1968. It has made a remarkable recovery in the decades since owing to the expansion of native and exotic woodland, and is no longer considered to be immediately threatened.

Two species of giant land tortoise (Cylindraspis) once roamed this island in huge numbers. When the French traveller François Leguat visited there in 1691, he reported ‘such a plenty of land turtles in this isle that sometimes you see two or three hundred of them in a flock, so that you may go above a hundred paces on their backs’. In 1759 one ship alone slaughtered 6000 tortoises and within 18 months 30,000 tortoises were captured. By the end of the century both the domed Rodrigues giant tortoise (C. peltastes) and the saddle-backed Rodrigues giant tortoise (C. vosmaeri) were on the point of extinction. One or the other species appears to have survived at least until 1802, when a few were reported killed in large fires used to clear the island’s vegetation for agriculture.

Two species of day gecko (Phelsuma) historically endemic to Rodrigues are now thought to be extinct. The Rodrigues giant day gecko (P. gigas) was common on Rodrigues in 1691–93, but seems then to have disappeared rapidly. In any case it was gone before 1874. The Rodrigues blue-dotted day gecko (P. edwardnewtoni) was probably extinct on Rodrigues before 1874, but subsisted on Ile aux Frégates at least until 1917. In 1963 no individuals were found there in spite of an intensive search.

 

Anthropogenic effects on the flora and fauna

While the precise timeline is still debated Madagascar was certainly one of the last major landmasses on Earth to be settled by humans, predating only that of Iceland and New Zealand. The earliest archaeological evidence for human presence there may be as much as 10,000 years old, although actual settlement by Austronesian peoples first began between 550 and 350 bc. The latter are believed to have arrived in successive waves by outrigger canoe from Borneo. These early settlers practiced slash-and-burn agriculture to clear the coastal rainforests for cultivation and hunted the island’s megafauna, wiping out most of them in the process. By ad 600 groups had begun to clear the forests of the central highlands as well. Arab traders first reached the island between the seventh and ninth centuries, and a wave of Bantu-speaking migrants crossed the Mozambique Channel from southeastern Africa around ad 1000. Others continued to reach on the island over time, including South Indian Tamil merchants who introduced the zebu (Bos primigenius indicus), a type of long-horned humped cattle, which they kept in large herds. Irrigated paddy fields were developed in the highlands and were later extended with terraced paddies. By the seventeenth century the rising intensity of land cultivation and the everincreasing demand for zebu pasturage had largely transformed the central highlands from a forest ecosystem to grassland. European contact with Madagascar began in 1500–01 with the Portuguese sea captain Diogo Dias first sighting the island. In 1528 Diogo Rodrigues explored what are now the Mascarene Islands, naming what are now Réunion, Mauritius, and Rodrigues. The first small Portuguese settlements began to appear along the coasts of Madagascar a few years later for the purposes of trade and missionary work. By the late seventeenth century the French, too, were establishing trading posts along the eastern coast, and the island began to gain prominence among European pirates and slave traders as well. Indeed, the small island of Nosy Boroha, off the northeastern coast, has been proposed by some historians as the site of the legendary pirate utopia of Libertalia. Many European sailors were shipwrecked along the coasts, among them the 17- year-old Englishman Robert Drury, whose journal is one of the few written depictions of life in southern Madagascar during the early eighteenth century. The wealth generated by maritime trade spurred the rise of organized native kingdoms, some of which grew quite powerful.

Unlike other African states, Madagascar had a relatively short period of colonization, lasting from the late nineteenth century until the early 1960s. The Mascarenes, by contrast, had already been settled by the Portuguese, Dutch, and French as early as the seventeenth century, in the process unleashing an unbelievably wanton destruction of animal life. In 1883 the French invaded Madagascar itself in what became known as the first Frano-Hova War. At its conclusion, the island ceded the northern port town of Diego Suarez (modern Antsiranana) to France. In 1890 the British accepted the full formal imposition of a French protectorate on the island, although this authority was not recognized by the native government. To force its capitulation, in 1894 the French bombarded and occupied the harbour of Tamatave (modern Toamasina) on the eastern coast, and the following year Majunga (modern Mahajanga) on the west coast. A French military expedition then marched to the capital Tana (modern Antananarivo), where the royal palace was bombed with heavy artillery leading Queen Ranavalona III to surrender and the royal family to go into exile. France formally annexed Madagascar in 1896 and declared the island a colony the following year. Slavery was abolished in 1896, and approximately 500,000 slaves were freed. Under colonial rule wide paved boulevards and buildings were constructed in Antananarivo, and the royal palace was turned into a museum. Schools were built throughout the island and a French-language education became mandatory. Taxation enabled the construction of a railway and roads linking key coastal cities to the capital. Plantations were established for the production of a variety of export crops, along with ambitious and often highly destructive development schemes. Even worse was the introduction of livestock and exotic species. Zebu cattle were brought in by the millions, causing enormous environmental damage. Many of the herds roamed in a semi-wild state, feeding, and reproducing freely but without being controlled by predators. Furthermore, between 1930 and 1940 red deer (Cervus elaphus) and fallow deer (Dama dama) were also introduced. They too were detrimental to the native vegetation, which had evolved without pressure from ungulate browsing. With independence and an ever-exploding human population, however, the destruction of the remaining forests accelerated, mainly due to slash-and-burn agriculture. It was only in relatively recent decades that an environmental awareness began to take hold and a system of protected areas put in place, with the first national parks established by the late 1950s. By then it was almost too late. The occupation of Madagascar during World War II had tarnished the prestige of the colonial administration and galvanized a growing independence movement, and the Malagasy Republic was ultimately proclaimed in 1958 as an autonomous state within the French Community. Since then, Madagascar has transitioned through four republics with a rather neo-colonial arrangement, all noted for their corruption. Today, what little remains of the island’s ecosystems and unique wildlife continue to be under threat by the encroachment of humans.

In recent historical time (i.e. since ad 1500), the Madagascan Realm has lost at least 49 species/3 subspecies of vertebrates. Among the extinct forms 2 species are mammals, 37 species/2 subspecies are birds, 7 species/2 subspecies are reptiles, and 3 species are freshwater fishes. One other species of mammal and 2 species of reptile are possibly extinct, and one species of fish is currently extinct in the wild.

In addition, there are 636 species/20 subspecies currently threatened with extinction (that is to say, either Critically Endangered, Endangered, or Vulnerable according to the IUCN Red List, as well as certain forms either listed as Data Deficient or Not Assessed but which are clearly at some risk of extinction). Of these, 143 species/5 subspecies are mammals, 65 species/6 subspecies are birds, 192 species/9 subspecies are reptiles, 163 species are amphibians, and 73 species are freshwater fishes.