Madagascar's Reptiles and Amphibians

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Madagascar is a herpetological treasure trove. A plump, blood-red tomato frog, a turquoise chameleon the size of a small cat, a jewel-eyed gecko edged with frills, and other bizarre creatures such as golden frogs and flat-tailed kapidolo tortoises, are only a few of the delights which await the avid herpetologist.

Madagascar is unique; an island continent whose plants and animals have evolved in isolation for more than 150 million years. Due to its proximity to Africa, the fauna and flora have an obvious kinship with the continent. However, some Malagasy reptiles have their closest relatives not in Africa, but in South America. There are also curious gaps and exciting excesses that make the island a very special region. Bird diversity is strangely reduced (some 250 species), only a third of the level found on the adjacent mainland. Adders and cobras are absent, as are ungulates, wild cats and canids. In glorious profusion, however, are orchids (more than in all of Africa), half of all chameleons, and the richest primate fauna in the world (albeit restricted to lemurs).

Humans arrived late, only some 1 500 years ago, sailing across the Indian Ocean from the East Indies. In a short time they had hunted to extinction all the large animals, including the world's heaviest bird (the flightless elephant bird), two species of giant tortoise, the tiniest hippopotamus, and a giant lemur. During the last few centuries, in the face of a population explosion, the eastern rainforests have been decimated, only remants of their former glory remaining. Most of the original habitats have been extensively altered, a process which continues unabated.

Although such habitat changes sometimes benefit certain animals, the majority are usually less tolerant, or entirely incapable, of adjusting to a modified environment. Specialist species with very particular habitat requirements often demand pristine conditions to ensure their long-term survival. In response to this environmental crisis, scientists are actively searching the remaining forest fragments for the survivors of this unique radiation.

Amphibians and reptiles usually proliferate in tropical areas, and Madagascar is no exception. The island's diverse climatic, geological and botanical elements ultimately contribute to its faunal richness and diversity: currently, more than 300 reptile species and nearly 170 frog species have been recorded. In spite of these already impressive figures, new species are being discovered and described at a rate higher than ever before, and within this decade 49 new species of amphibians and reptiles have been described while other discoveries await formal description.

One would expect the fauna of at least the nature reserves to be well documented, but this is not the case. For instance, much of the Montagne d'Ambre mountain range in northern Madagascar currently receives protection by means of a Special Reserve and a National Park which were created in 1958. Despite a history of collecting in the area that dates back over a century, a two-month herpetofaunal survey in 1991-92 more than doubled the amphibian and reptile checklist. Out of a total of 70 species found during the survey, 37 were new records for the area. Incredibly, seven of these proved to be new species.

Some newly discovered reptiles are so restricted that they are obvious conservation concerns. In fact, one new leaf gecko, Uroplatus malama, is known from a single specimen from a remnant of low montane rainforest in south-eastern Madagascar. A similar species, Uroplatus malahelo, is known from only two specimens collected in 1992 from a minute patch (1 500 x 800 metres) of rainforest in southern Madagascar. It is aptly named after a Malagasy word (pronounced 'mawlahayloo') meaning forlorn or sad. When collected, its habitat, the southernmost rainforests on the island, was already being selectively logged and was not expected to last more than 3-4 years. The species may already be extinct.

The large, attractive new species Paroedura masobe is considered by many to be the most beautiful gecko on the island. It is known from a single locality, fortunately within the boundaries of the Zahamena Strict Nature Reserve. But its size and beauty almost guarantee this gecko's entrance into the international pet trade within the next few years.

Of the approximately 170 frog species currently recognized from Madagascar, a stag-gering 99 per cent are endemic. Of the only two non-endemic frogs, one species, Hoplo-batrachus tigerinus, was introduced from Asia to satisfy the culinary cravings of the French community. The mosaic of eastern rainforests abound with frogs. More than 90 species have been recorded from moist montane forest in the region of Andasibe (Périnet), making this among the highest frog richness in the world. The impressive endemic amphibian species record, however, is contained in only a few families, and some frog families are notably absent. These include the almost cosmopolitan Bufonidae (toads) and limbless amphibians, also known as caecilians, which are present in much of Africa and on the Seychelles, but do not occur on Madagascar.

A photographer would be hard pushed to find a more photogenic group of animals than the Malagasy tree frogs of the genus Boophis. But beautiful pictures don't come easily! To find these exquisite creatures you must spend many hours at night in the rainforest, trying to home in on their calls. Rainy nights are the most productive, when the frogs engage in a frenzy of calling and spawning. Thirty-six species are currently known in the genus, but this is definitely not the sum total: since 1990, 13 new species and two subspecies have been described, while a few other new taxa still await scientific names.

Some of the most popular frogs in captivity are the mantellas. These small, diurnal frogs are strikingly beautiful, displaying a variety of vivid patterns. Their brilliant colours may serve to warn predators of toxic elements present in their skin, a characteristic (known as aposematic coloration) that they share with the South American poison arrow frogs (Dendrobatidae). This represents an extraordinary case of convergence (parallel evolution). Particularly prized is the golden mantella Mantella aurantiaca and thousands of specimens are exported annually for the exotic pet trade in America and Europe. Some other mantellas are also exported in large numbers.

Malagasy microhylids include a multitude of shapes, colours and sizes, and are represented by 10 genera. It is in this family that most new species have yet to be described. Probably the most famous species is the tomato frog Dyscophus antongili, an obese red frog which can secrete a toxic substance from its skin to deter potential predators such as the giant hog-nosed snake Leioheterodon madagascariensis.

The majority of Malagasy frogs do not follow the typical amphibian life-cycle in which eggs laid in water hatch into tadpoles that eventually metamorphose into small frogs. The numerous frogs of the genus Mantidactylus lay their eggs on the forest floor near water, or on to vegetation overhanging a stream or pool. After a while the tadpoles emerge and wiggle their way to the water, to complete their life-cycle in the usual way. But some frogs have skipped the free-swimming tadpole stage altogether and are terrestrial breeders.

One genus that has adopted this mode of reproduction is Stumpffia. These minute frogs lay their eggs in foam nests hidden among leaves on the forest floor, and the non-feeding tadpoles develop directly into tiny froglets. Those of Stumpffia pygmaea measure less than three millimetres in body length, making them the smallest frog, if not vertebrate, in the world.
The most striking feature of the more than 80 species of snakes which occur on Madagascar is the absence of deadly venomous species - all are clinically harmless. The larger, non-venomous species may inflict a painful bite, but the back-fanged species possess venoms that are either too weak or produced in too low a quantity to be of danger to humans. With such an absence of dangerous snakes, one would expect the local people to have no fear of snakes. However, the giant hog-nosed snake does scare some people, and the fandrefiala (Ithcyphus miniatus) is apparently feared throughout Madagascar. This slender-bodied, arboreal snake has a bright red tail tip, and is reputed to drop tail first from trees, stiff as an arrow, to stab any person or zebu unfortunate enough to be passing below!

The largest snakes include three boa species whose closest relatives are found not in Asia, but in the Amazon Basin in South America. Pythons, which abound in Africa and Australasia, are absent from Madagascar. All three boas are protected from international trade, although all remain relatively common and widespread on the island. There is evidence, however, that their flesh and skins are increasingly being exploited, and belts and other articles made from boa skin can be found in markets in the towns and cities.

New snake species are still being discovered. One of the island's most unusual serpents is the twig-mimicking snake Langaha madagascariensis. This species exhibits sexual dimorphism, the two sexes differing in coloration and nasal appendages. Another dimorphic species is Mimophis mahfalensis, in which the male is intricately patterned with a zigzag vertebral stripe, whereas the female is uniformly coloured.

Lizards are abundant on Madagascar, with more than 210 species in five families, four of which are common in Africa. Absent, however, are common African and Asian lizards such as lacertids, agamas and monitors. Repeating the theme found in snakes, the island is home to a few iguanid lizards, a group otherwise restricted to the Americas and a few Pacific islands.
Madagascar is justifiably famous for its chameleon diversity for it contains approximately half of the world's species, among them the biggest and smallest known chameleons. Perhaps the most peculiar are the small, stump-tailed chameleons, Brookesia spp. They clamber about the leaf litter on the forest floor, where their drab colour and twig-like body shape make them well-nigh invisible to the casual observer. Of the more typical chameleons, Oustalet's (Furcifer oustaleti) and Parson's (Calumma parsonii) chameleons are immense, both exceeding 600 millimetres in length.

The appearance of many chameleons differs so markedly between the sexes that, to the uninformed, they appear to be different species. This radical sexual dimorphism is best illustrated by Furcifer minor and F. labordi, in which only the males have extravagant nasal protuberances. The females, however, are brightly coloured, and early herpetologists, working with discoloured and shrivelled preserved specimens, can be forgiven for mistaking them as separate species.

Visitors are often fascinated by the beauty of chameleons and their ability to change colour. In contrast, the local people of Madagascar, living in a land teeming with these bizarre reptiles, often regard them with revulsion. Many of the local proverbs reflect this attitude: the Tsimihety saying 'Ratsy karaha Kandrondro' means 'ugly as a chameleon', while 'Tsatsaka no i Taha ka mifaninjy fa samy raty' of the Mahafaly tribe translates as 'between Phelsuma (day geckos) and chameleon, don't quarrel, you are equally ugly'. Even the small, stump-tailed chameleons (Brookesia spp.) are feared, and another Tsimihety proverb, 'Mahatsidia vokon' Anjava kely izy fa mafoaka' means 'step on a knot of Brookesia - a small animal but expect a great misfortune'.

Like chameleons, day geckos (Phelsuma spp.) and leaf-tailed geckos (Uroplatus spp.) elicit great excitement among visitors. The combination of brilliant neon coloration, general abundance, approachability, and their diurnal habits make day geckos ideal photographic subjects. While the intense green coloration of many of these geckos is important for intraspecific recognition and communication, in sharp contrast, the leaf-tailed geckos are masters of disguise. In addition to being cryptically coloured, ragged fringes of skin extend along the throat, body and limbs of some species, particularly Uroplatus fimbriatus and U. henkeli. When pressed against a tree, these fringes break the outline of the body and prevent tell-tale shadows. Achieving up to 330 milli-metres in total length, Uroplatus fimbriatus is one of the world's largest geckos; it also sports in excess of 300 teeth - more than any other reptile or mammal on earth.

The iguanids and gerrhosaurids are conspicuous, medium to large sized diurnal lizards. They inhabit a wide variety of habitats, including forests, deserts, and rocky outcrops. The gerrhosaurid Zonosaurus maximus is the largest Malagasy lizard and is locally known as the 'petit caiman', as a result of its large size and aquatic habits.

Not to be outdone in the speciality stakes, four of the five terrestrial tortoises present on Madagascar are endemic. The other, Bell's hingeback tortoise Kinixys belliana, is common in coastal regions of East Africa and was probably imported to the island by man. The angonoka Geochelone yniphora of Baly bay in the north-western coastal region, is considered the world's rarest tortoise and, in an effort to ensure its survival, the Jersey Wildlife Preservation Trust successfully established a captive-breeding programme at Ampijoroa. The programme received a setback in mid-1996, however, as two adults and 73 juveniles were stolen from the centre - smuggled to Europe for the illegal pet trade.

Also threatened is the attractively patterned radiated tortoise Geochelone radiata, which is restricted to the arid south and south-west of Madagascar. Although the local Antandroy and Mahafaly tribes regard it as fady (taboo) to eat these tortoises, people from other parts of Madagascar consider the meat a delicacy. Many tortoises are therefore collected illegally for food, while others are kept as pets or their shells sold as curios. Another very rare tortoise is the kapidolo or Malagasy flat-tailed tortoise Pyxis planicauda. During the dry season this small tortoise aestivates, and only emerges in response to heavy rains.

Madagascar's smallest chelonian is the spider tortoise Pyxis arachnoides, also known as tsakafy or kapila. It inhabits semi-arid thornbush scrub along a southern and south-western coastal belt, extending only from 10-50 kilometres inland.

Of the four freshwater terrapins on the island, the Madagascan big-headed terrapin Erymnochelys madagascariensis is the sole endemic - the other species are all widespread in Africa. The big-headed terrapin is the only Old World representative of a subfamily otherwise restricted to South America. It is fully protected by Malagasy law but enforcement is difficult, and terrapins accidentally caught in seine fishing nets are almost invariably slaughtered for local consumption.

This terrapin appears to be very sensitive to over-exploitation: local fishermen at Lac Marofilao were surprised when they suddenly disappeared from the lake in 1987 as, up until the previous year, they had been regularly caught. Various factors have contributed to the rapid decline of this species, but the most important appears to be its prolonged juvenile stage and large size at maturity: specimens become large enough to eat long before they reach sexual maturity. Marine turtles have also been heavily exploited for many years. A survey in 1971 estimated that each year 13 000 turtles are killed along the west coast.

It is painfully obvious that the island's herpetofauna is in great demand in inter-national trade. Madagascar's herpetofauna was poorly represented in the exotic pet market until the mid-1980s, when a rapid increase heralded their arrival. By far the most popular are the colourful day geckos (Phelsuma spp.): between 1986 and 1991, just under 145 000 specimens of 17 different species were exported from the island. Another popular group is the chameleons: CITES figures for the period 1986-1991 record the export of 38 325 specimens from 21 species. But the most apparent trade boom of Malagasy herpetofauna is in frogs; for instance, legally exported mantellas skyrocketed from 230 specimens in 1980 to 11 058 in 1989.

BIODEV, a local Malagasy task group working in conjunction with the IUCN, has investigated the economic potential of amphibian and reptile exports from Madagascar. They calculate an economic potential of US$1 million (including $150 000 for chameleons and $240 000 for day geckos) per year. This takes into account that most of the economic benefit from this animal trade will still accrue to the importing countries, where sales for chameleons alone will probably top $1 million. In terms of foreign revenue to the Malagasy people, this trade will be equivalent to clearing 16 000 hectares of forest every year. Such a reversal in conservation priorities from protection to utilization will only work, however, if the profits of sustainable harvesting are equitably distributed to the people whose survival currently depends on cutting and burning the forests.

Currently, the enormous profit margins obtained from exploiting Malagasy reptiles benefit just a few individual importers and exporters. A villager collecting a day gecko in the forest earns only FMG50 (US$0.03), whereas the local exporter sells it for US$9-13. By the time it retails in the USA it may cost US$75, a 2 500-fold increase in the price. Can Malagasy villagers be expected to value sustainable harvesting when they make only US$3 for every 100 lizards they catch?
A possible solution could be for the Madagascar authorities to place greater emphasis on ecotourism, to encourage more people to visit the island, and to place greater value on protecting the herpetofauna.

There is no doubt that Madagascar has the potential to become a prime ecotourism hot-spot - the unique and diverse wildlife of the island must impress any naturalist. But can amphibians and reptiles really serve as a drawcard for tourists when many people regard these creatures with abhorrence?
If one considers the Komodo dragon, the world's largest monitor lizard and a major drawcard for Indonesian ecotourism, the giant tortoises and marine iguanas of the Galapagos islands which annually attract thousands of visitors, as well as the numerous organized excursions to observe nesting marine turtles at sites worldwide, Madagascar must surely rate as one of the world's best countries in which to appreciate amphibians and reptiles in their natural habitat.

ALL CREATURES GREAT AND SMALL
The best way to find chameleons is to search for them by torchlight at night. But most people will be quite surprised - startled, even - when the light reveals a 'monster' such as the gigantic Parson's chameleon - it is a truly a magnificent beast. Locating a specimen of the pygmy stump-tailed chameleon is, however, no easy feat. This specimen perched on a human finger is an adult male which was found during the day among leaf litter on Nosy Mangabe.

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