Contributed Papers Montane and Communities in Madagascar

CHRISTOPHER J. RAXWORTHY AND RONALD A. NUSSBAUM Museum of Zoology, University of Michigan, Ann Arbor, MI 48109-1079, U.S.A.

Abstract: Two habitats in Madagascar, secondary, montane heathland and high plateau prairie, are consid- ered artificial, having been created by human-set fires soon after the island was colonized less than 2000 years ago. These prairie and secondary heathland habitats are also thought to be degraded and faunistically depauperate. To test the depauperate fauna hypothesis we measured levels of herpetofaunal (amphibian and reptile) endemicity between primary forest, montane secondary heathland, and high plateau prairie. We found no endemic to high plateau prairie, but a significant percentage of the montane herpetofauna is restricted to post-fire secondary heathland. These results support the human-origin hypothesis for prairie, but secondary heathland appears to be a natural, post-fire successional stage that leads to a climax of sclero- phylloas forest. This suggests that careful management of montane heathland could establish new dispersal corridors between currently isolated montane forest blocks and, therefore, could offer new opportunities for conservation in Madagascar. Despite widespread burning, the montane heathlands of Madagascar have di- verse herpetofaunal communities, demonstrating that these montane communities are not as seriously de- graded as previously believed, and that they may be naturally resistant to fire.

Comunidades de Anfibios y de Montafia en Madagascar Resumen: En Madagascar los h~bitats de brezal secundarto de montana y los de la pradera alta (High Pla- teau) son considerados como artifictales debido a que se origtnaron por incendios provocados por humanos poco despu#s de la colonizaci6n de la tsla menos de 2000 a~os a.C. Tambidn se piensa que estos h~bitats es- t~n degradados y la fauna se encuentra en condiciones paup4rrimas. Para evaluar la htp6tesis de paupertza- ci6n de la fauna, medimos los niveles de endemismo de la herpetofauna (anfibtos y reptiles) entre bosque pri- mario, brezal secundarto y la pradera High Plateau. No encontramos especies enddmicas a la pradera High Plateau, pero un procentaje significativo de la herpetofauna de monta~a estd restringido a tierras de post-in- cendio secundario. Estos resultados soportan la hip6tesis de origen-humano para las regiones de pradera, pero los hd~bitats de brezal secundarios parecen presentar un estado seral natural post-incendio que con- ducen aun clfmax de bosque escler6filo. Esto sugiere que un manejo culdadoso de los brezales de montar~a puede permitir nuevos corredores de disperst6n entre bloques de bosque de montar~a actualmente aislados y de esta manera ofrecer nuevas oportunidades de conservaci6n en Madagascar. A pesar de los tan comunes incendios, los brezales de montana de Madagascar poseen comunidades diversas de herpetofauna que de- muestran que estas ~reas no se encuentran seriamente degradadas como previamente se habla pensado, ademds de que pueden presentar resistencia natural al fuego.

Introduction bert 1927), but it was actually a dynamic mosaic of heathland, grassland, and forest during the Holocene, The High Plateau of Madagascar (Fig. 1) was once as- 11,000-1240 years ago (MacPhee et al. 1985; Burney sumed to have been almost completely forested before 1987a, 1987b). The current prairies of the high plateau the arrival of humans (Perrier de la Bftthie 1921; Hum- are believed to be artificial habitats created by clearing, burning, and cattle grazing (Humbert 1927; Koechiin Paper submitted February 15, 1995; revised manuscript accepted 1972; Dewar 1984). In support of this interpretation, November 28, 1995. these prairies appear to have an impoverished fauna 750

Conservation Biology, Pages 750-756 Volmne 10, No. 3,June 1996 Raxworthy &Nussbaum Madagascan and Reptiles 751

quency of fires at Andringitra (Jenkins 1987) and our personal observations suggest that almost all extant heathland at this site is also secondary. The large areas of extant, secondary montane heath- land, typically dominated by Philippia and grasses, are usually interpreted as anthropogenic habitats (produced by human burning of sclerophyUous forest or primary heathland) that are faunistically depauperate (Humbert 1927, 1928; Perrier de la Bithie 1927). Therefore, these secondary heathlands are considered analogous to the pseudo-steppe prairies of the high plateau. In montane regions, however, there is evidence of natural wildfires occurring throughout the Holocene (Burney 1987a), which in addition to recent reports of lightning-caused fires (Jenkins 1987), suggests that secondary heathland is a natural, post-fire successional stage. The Madagascan herpetofauna, with about 550 total endemic species (Raxworthy & Nussbaum, unpublished data), represents about 65% of the endemic vertebrate species of Madagascar. This diverse group is well-suited for analyzing patterns of endemism within Madagascar because herpetofaunal species typically have restricted elevational and regional distributions, reflecting consid- erable habitat specialization (Raxworthy & Nussbaum 1994, 1995). None of these 550 island endemic species is restricted to obvious anthropogenic habitats (such as agricultural or urban areas), despite the widespread oc- currence of anthropogenic habitats throughout Mada- gascar. Herpetofaunal species that occur in anthropo- genic habitats appear to have invaded from adjacent natural habitats and probably have broad ecological tolerances. Figure 1. The high plateau of Madagascar (shaded > Based on this observation, we predict that if second- 800 m elevation) and associated major massifs (black ary montane heathiand and high plateau prairie are an- > 2000 m). In Madagascar montane habitats above thropogenic, with no history prior to human coloniza- 1500 m elevation are confined to the high plateau. tion, there should be no herpetofaunal species endemic to just these habitats. To test the working hypothesis that these habitats are of anthropogenic origin, we mea- (Perrier de la Bftthie 1936; Jenkins 1987), suggesting in- sured levels of herpetofaunal (amphibian and reptile) sufficient time since the creation of the prairies for the endemicity in primary forest, secondary heathland, and evolution of endemic species or subspecies. prairie habitats. The primary montane vegetation of the three highest massifs in Madagascar--Tsaratanana (2876 m), Ankaratra (2642 m), and Andringitra (2658 m)--is also thought to Methods be badly degraded or destroyed because of burning by humans (Jenkins 1987; Nicoll & Langrand 1989). Four Surveys were completed during the rainy seasons of primary montane (above 1500 m) vegetation types are 1993 (January-March, October-December) when most recognized currently: moist montane (moss) forest in species are breeding and activity is at its highest. Field less exposed areas; sclerophyftous (lichen) montane for- techniques used to sample were (1) opportunis- est; montane heathland (bushland) in isolated patches tic day and night searching; (2) refuge examination (un- on exposed ridges above 2000 m; and rupicolous shrub der and in fallen logs and rotten tree stumps, under land restricted to rocky outcrops (Perrier de la B~thie bark, under rocks, in leaf litter, in root-mat and soft, and 1927; White 1983). No primary heathland survives at in leaf axils of screw palms of the genus Pandanus); and Ankaratra and Tsaratanana; both massifs have been (3) pitfall trapping with drift fences. Opportunistic burned frequently during this century (Perrier de la searching and refuge examination were done through- B~thie 1927; Humbert 1928; Guillaumet 1984). The fre- out the full elevational range of habitats available. Night

Conservation Biology Volume 10, No. 3,June 1996 752 MadagascanAmphibians and Reptiles Ram,Jorthy &Nussbaum

Table 1. Habitat distribution of species of amphibians and reptiles endemic to the high plateau (800-1500 m elevation),a

High plateau habitat Species ° Prairie Forest Wetland Rupicolous Unknown Amphibia Hyperoliidae Heterixalus rutenbergi Boettger 1881 +c + Plethodontohyla A + gottlebei Busse & B6hme 1992 +a Mantellidae Mantella aurantlaca Mocquard 1900 + Mantella cowani Boulenger 1882 +e Mantidactylus ambohimitombi Boulenger 1919 + Rhacophoridae Boophis idae Steindachner 1867 +f Boophis goudoti Tschudi 1838 + Boophis mandraka Blommers-Schl6sser 1979 + Boophis rhodoscelis Boulenger 1882 +f Boophis erythrodactylus Guib~ 1953 + Boophis hillenii Blommers-Schl6sser 1979 +f Boophis A + Boophis B + Boophis C + Reptilia grandidieri Mocquard 1900 +g Chamaeleonidae Brookesia thieli Brygoo and Domergue 1969 + Brookesia therezteni Brygoo and Domergue 1970 + Calumma brevicornis tsarafldy Brygoo & Domergue 1970 +b Calummafallax Mocquard 1900 + Calumma globifer Giinther 1879 +t Calumma malthe Giinther 1879 + Calumma A + Cordyfidae Zonosaurus ornatus Gray 1845 + + Zonosaurus A + Typhlopidae Typhlops domerguei Roux-Est~ve 1980 + Colubridae Brygophis coulangesi Domergue 1988 +J Geodipsas vinckei Domergue 1988 +J Geodipsas A + Liopholidophis pinguts Parker 1925 + Liopholidophis sexlineatus Giinther 1882 + Pseudoxyrhopus ankafinaensts Raxworthy and Nussbaum 1994 + aData from Raxworthy and Nussbaum, unpublished unless otherwise indicated. OTaxa not assigned specific names were considered undescrtbed and are the subject of ongoing taxonomic studies to he published elsewhere. CBlommers~chl6sser (1982). aBusse & B6hrne (1992). eBoulenger (1882). f Blommers-Scbl6sser (1979). gBlanc (1977). bBrygoo (1971). tSeguier-Guis (1988). JDomergue (1988).

searches were made using headlights. Pitfall trapping lin and later transferred to alcohol. Liver and muscle proceeded as described by Raxworthy and Nussbaum were removed from representatives of almost all species (1994). Date, time, longitude and latitude, altitude, habi- and frozen in liquid nitrogen for genetic studies. Repre- tat, and microhabitat were recorded at the time of cap- sentative live individuals of most species were photo- ture of each individual. graphed to record color, which fades in preservative. Voucher specimens were fixed in 10% buffered forma- Voucher specimens were deposited either in the Museum

ConservationBiology Volume 10, No. 3,June 1996 Raxworthy &Nussbaum Madagascan Amphibians and Reph'les 753

Table 2. Habitat and elevattonal distribution of montane endemic (> 1500 m elevation) species of amphibians and reptiles.

Massifa Massif Elevation (m) Species ~ Tsaratanana Ankaratra Andringitra endemic min max Amphibia Microhylidae Anodonthyla montana Angel 1925 F,H + 1700 2300 Platypelts tsaratananaensis Guib~ 1974 F + 2350 2350 Platypelts A F + 2350 2350 Plethodontohyla guentherpeterst Guib~ 1974 F + 2300 2300 Plethodontohyla tuberata Millot and Guib~ 1954 F,H + 1700 2550 Plethodontohyla A H + 2700 2700 Plethodontohyla B F + 2050 2050 Plethodontohyla C F + 2050 2050 Scaphtophryne A F + 1900 19OO Stumpffia A F + 2050 2050 Mantellidae Mantidactylus aerumnali~ Peracca 1893 F H 1700 2150 Mantidactylus alutus Peracca 1893 F, H 1700 2500 Mantidactylus domerguei Guib~ 1974 F F, H c 1500 a 2150 Rhacophoridae Boophis microtympanum Boettger 1881 H H 1850 2550 Reptilia Gekkonidae Lygodactylus montanus Pasteur 1964 R + 2000 2250 Millottsaurus mirabilis Pasteur 1962 R + 2400 2640 Phelsuma barbourt Loveridge 1942 R R 16OO 2640 Phelsuma lineata punctulata Mertens 1970 R + 2700 2870 Chamaeleonidae Brookesia lolontany F + 1650 2050 Raxworthy and Nussbaum 1995 Calumma brevtcornis hillentusi F, H F, H 1550 2550 Brygoo, Blanc, and Domergue 1973 Calumma guibe Hillenius 1959 F + 16OO 2100 Calumma tsaratananaensis H + 2700 2800 Brygoo, Blanc, and Domergue 1969 Calumma A F + 2550 255O Calumma B F + 1630 2350 Furcifer campani Grandidier 1872 H H 1850 2500 Scincidae Amphiglossus tsaratananaenst$ Brygoo 1981 F, H + 2050 2870 e Amphiglossus A F, H H 1700 2640 Mabuya boettgeri Boulenger 1887 H H 1600 2400 Mabuya madagascartensis Mocquard 1908 H 1650 2400 Colubridae Geodipsas B F + 1620 2050 aF, montane moist and sclerophyllous forest; H, secondary heathland; R, ruptcolous habitats. bTaxa not assigned specific names were considered undescribed and are the subject of ongoing taxonomic studies to be published elsewhere. CMtllot & Guibd (1950). aBlommers.Schi6sser & Blanc (1991). eBrygoo (1981).

of Zoology, University of Michigan, or in the collections ordinates and further information of these sites are given of the Service de Zoologie, Universit6 d'Antananarivo. by the U.S. Department of the Interior (1955) and by A list of amphibians and reptiles endemic to the High Nicoll and Langrand (1989). Plateau and their habitat associations was compiled from The three montane systems surveyed were Ankaratra, our field work and from the literature (cited in Table 1). 10-24 February, 19°21'S, 47°18'E; 19°22'S, 47°19'E; Tsa- These data are based on studies made in the regions of ratanana, 18 March-4 April, 14°09'S, 48°58'E; 14°06'S, Ambatondrazaka, Ambatolampy, Ambositra, Ambohitan- 48°59'E; 14°ll'S, 48°57'E; and Andringitra, 8-17 De- tely, Andringitra, Antananarivo, Isalo, Mandraka, Man- cember, 22°12'S, 46°58'E; 22°10'S, 46°56'E. Montane tady, Perinet, Ranomafana, Tsarafidy, and Zahamena. Co- habitats were classified and identified as follows: (1)

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16 ~ ¢ Our surveys of the three major massifs (Andringitra, Ankaratra, Tsaratanana) yielded 30 amphibians and rep- 14 tiles restricted to 1500 m elevation or higher (Table 2). No additional species were found during the final 3-5 12 survey days at each site (Fig. 2), which indicates that

0 these species lists are nearly complete. Four geckos 10 (Gekkonidae) are restricted to montane rupicolous habi- tats. All other montane species (Microhylidae, Mantel- o~ 8 lidae, Rhacophoridae, Chamaeleonidae, Scincidae, and Colubridae) occur only in heathland and/or forest. "5 E= Ankaratra The herpetofaunal species diversity for secondary 6 0 Tsaratanana montane heathland is eight species at Andringitra, eight -=- Andringitra at Ankaratra, and three at Tsaratanana. Five species are 4 restricted to secondary montane heathland at Andringi- tra, four at Ankaratra, and two at Tsaratanana. At Tsara- 2 i tanana, 12 species are restricted to moist or sclerophyl- t / lous forest, compared to just two at Ankaratra and none 0 I i I I I I i I I I I i I , at Andringitra (Table 3). Secondary heathland is occu- 0 1 2 3 4 5 6 7 8 9 10 12 pied by 54% of the nonrupicolous montane herpeto- Sampling days fauna (all three massifs combined), and 23% of the spe- Figure 2. Accumulation curves illustrating the rate of cies are found only in this montane secondary habitat. discovery of new species of amphibians and reptiles at The significant percentage of montane endemic species each montane site. restricted to secondary heathland contrasts sharply with the results for the high plateau prairie, where no en- demic species were identified. We therefore reject the montane moist and sclerophyilous forest, trees forming hypothesis that montane secondary heathland is artifi- a continuous canopy; (2) secondary heathland, vegeta- cial and faunistically depauperate. tion dominated by ericoid shrubs and grasses with evi- dence of former burning (blackened tree trunks, stems, or roots); (3) rupicolous habitats, areas of rock surface Discussion and crevices. Montane moist forest was not distin- guished from sclerophyllous forest because the two The absence of endemic amphibians and reptiles in the high plateau prairie strongly suggests that the modem types proved to intergrade over broad areas. prairie is recent and artificial, and that the original vege- tation (and its endemic herpetofauna) has been lost. Claims that the original vegetation was extremely sus- Results ceptible to fire and that it was therefore destroyed by the very first human-set fires (Morat 1973) cannot be Of the herpetofauna surveyed, nine species of amphibi- supported because wildfires were common the high pla- ans and seven species of reptiles appear to be unde- teau throughout the Holocene (Burney 1987a, 1987b). scribed taxa, This reflects the general lack of herpetolog- It is also unlikely that such a large area of natural vegeta- ical survey work that has been conducted in the interior tion, 22,350-36,000 km 2 (Jenkins 1987), was destroyed of Madagascar. These new species are in the process of by clearing, because the original human population being described and in all cases are represented by must have been small (MacPhee et al. 1985). Grazing by voucher specimens. Because these new species repre- introduced domestic bovids also has been proposed to sent such a sizeable fraction of the surveyed herpeto- explain the destruction of the natural vegetation of the fauna, we have chosen to include them in this analysis. We identified 32 species of amphibians and reptiles as Table 3. Number of montane endemic (> 1500 m elevation) high plateau endemics (restricted to elevations between herpetological species in different habitats. 800 and 1500 m on the high plateau). These endemics and their habitat types are listed in Table 1. Of these, 25 Habitat ('78%) are restricted to forest habitat. Just two species Forest and (6%) occupy prairie, and neither is confined to this habi- Massif Ruptcoloas Forest Heathland heathland tat. The complete absence of endemism in the high pla- Tsaratanana 1 12 2 1 teau prairie supports the current belief that this habitat Ankaratra 1 2 4 4 is artificial and of recent origin. Andrin#tra 2 0 5 3

Conservation Biology Volume 10, No. 3, June 1996 Raxworthy &Nussbaum Madagascan Amphibians and Reptiles 755 high plateau (Dewar 1984). We suspect that the modem habitats have been destroyed or degraded has been largely practice of yearly burning for improved grazing (wide- responsible for the lack of research and conservation pro- spread throughout the high plateau) originated when grams in these regions. At Tsaratanana (the most diverse pastoralists first settled in Madagascar, and that regular, massif with probably the highest degree of local ende- repeated burning, rather than grazing per se, ultimately micity of any site in Madagascar) our survey was the first destroyed the original high plateau vegetation. Unlike scientific study made at the summit in almost 30 years, prairie, the montane habitats have not been frequently and there are no conservation programs currently active burned; therefore, their natural, post-fire successional within the Tsaratanana Reserve. Although conservation stages have not yet been destroyed. programs have recently been established at Andringitra We propose the following revised montane vegeta- and Ankaratra, the lack of research on the high-elevation tional model for Madagascar. Prior to human coloniza- habitats is preventing their effective management. tion, a mosaic of sclerophyllous forest and heathland Our results suggest that montane communities in was maintained by natural wildfires within each of the Madagascar may be more resistant to fire than currently major massifs of Madagascar. Secondary heathland devel- realized. Post-fire secondary heathland is a valuable habi- oped as a post-fire successional stage leading to a climax tat with a diverse endemic fauna. If protected from fur- of sclerophyllous forest, except on the most exposed ther burning or cattle grazing, it may develop into cli- ridges, plateaus, and summits, where heathland was the max sclerophyllous forest. This offers the possibility that climax vegetation. The rupicolous montane habitats, careful management of heathland could establish new largely protected from fire, remained stable. dispersal corridors between montane forest blocks. This At the northern massif, Tsaratanana, 75% of the mon- conclusion, if confirmed, offers new challenges and op- tane herpetofauna is restricted to moist or sclerophyl- portunities for conservation management in Madagascar. lous forest, a much higher percentage than on the two central massifs, Ankaratra and Andringitra, with just 17% and 0% respectively. This contrast may result from differ- Acknowledgments ences in the size of the montane forest refugia that ex- isted at each massif during glacial periods of the Pleis- We thank H. Wright and R. Burke for comments on an tocene. In East Africa, glacial periods of the Pleistocene earlier draft of this paper. We are greatly indebted to had much drier climates, which caused the montane for- Jean Baptiste Ramanamanjato, AchiUe Raselimanana, and ests to contract during these periods (Hamilton 1982). Anglien and Angeluc Razafimanantsoa for help in the Current rainfall is considerably higher at Tsaratanana field. This research was made possible through the co- (mean monthly rainfall 200-250 mm) than at the two operation of the Malagasy Minist6re de rEnseignement central massifs (mean monthly rainfall 100-150 mm) Sup6rieur, the Minist~re de la Production Animale et des (Donque 1972). If rainfall was also higher at Tsaratanana Eaux et For~t, and the Minist~re de la Recherche Scienti- during glacial periods than on the central massifs, then fique et Technologie pour le Developpement. This re- montane forest would have been much more extensive search was supported in part by a grant (BSR 90-24505) at Tsaratanana than Ankaratra or Andringitra. The larger from the National Science Foundation. Logistic support (and probably more stable) montane forest refuge at at Andringitra was provided by the World Wide Fund for Tsaratanana would, therefore, be expected to have a Nature. greater diversity of montane forest specialists than either Ankaratra or Andringitra. Literature Cited The secondary montane habitats in Madagascar are ob- viously not depauperate of endemic vertebrates, a result Blanc, C. P. 1977. Reptiles. Sauriens. Iguanidae. Faune de Madagascar 45:1-200. that has important implications for conservation. De- Blommers-SchlGsser, R. M, A. 1979. Biosystematics of the Malagasy spite frequent burning at Ankaratra and Andringitra, the , lI. The genus Boophts (Rhacophorid~e). Bijdragen Dierkunde secondary heathlands of both massifs still have diverse 49:261-312. herpetofaunas. The montane vegetation of Tsaratanana, Biommers-Schl6sser, R. M. A. 1982. Observations on the Malagasy ge- frequently reported to have been completely destroyed nus Hetertxalus Laurent, 1944 (Hyperoliidae). Beaufortia 32:1-11. by fire (Guillaumet 1984; Jenkins 1987; Nicoll & Lan- Blommers-Schl6sser, R. M. A., and C. P. Blanc. 1991. Amphibiens (premi6re partie). Faune de Madagascar 75:1-400. grand 1989), also has a secondary heathiand that sup- Boulenger, G. A. 1882. Catalogue of the Batrachia Salientia s. Caudata ports an endemic herpetofauna. The sclerophyllous for- in the collection of the British Museum. British Museum, London. est at Tsaratanana, claimed in 1924 to have been Brygoo, E. R. 1971. Reptiles Sauriens Chamaeleonidae, le genre completely burned above 2200 m elevation (Perrier de Chamaeleo. Faune de Madagascar 33:1-318. la Bfithie 1927), has actually regenerated or survived in Brygoo, E. R. 1981. Syst~matique des 16zards scincid~s de la r~gion malgache. VI. Deux scincid6s nouveaux. Bulletin de la Mus~e Na- some areas to an elevation of 2600 m (Raxworthy, per- tional de Histoire natureHe, Paris, 4th Series, SectionA 3:261-268. sonal observation). Burney, D. A. 1987a. Pre-settlement vegetation changes at Lake The widely held view that the Madagascan montane Tritrivakely, Madagascar. Palaeoecologyof Africa 18.-357-381.

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Burney, D. A. 1987b. Late Quatenmry stratigraphic charcoal records chronology and environment of Ampasambazimba, a Malagasy sub- from Madagascar. Quaternary Research 28:274-280. fossil lemur site. International Journal of Primatology 6:463-489. Busse, K, and W. B6hme. 1992. Two remarkable discoveries of Millot J., and J. Guib6. 1950. Les Batraciens du Nord de l'Andringitra the genera Mantella (Ranidae: Mantellinae) and Scaphiophryne (Madagascar). M~moires de l'Institute Scientifique de Madagascar (Microhylidae, Scaphiophryninae) from the west coast of Madagas- Series A 4:179-206. car. Revue Francaise d'Aquariologie, Herp6tologle 19:57-64. Morat, P. 1973. Les savanes de l'Ouest de Madagascar. M6moires de Dewar, R. E. 1984. Extinctions in Madagascar. Pages 574-593 in P. S. I'ORSTOM 68:191-193. Martin and R. G. Klein, editors. Quaternary extinctions: a prehis- Nicoll, N. E., and O. Langrand. 1989. Madagascar: Revue de la Conser- toric revolution. University of Arizona Press, Tucson. vation et des Aires Prot6g6es. Fonds Mondial pour la Nature, Gland, Domergue, C. A. 1988. Notes sur les serpents de la r~gion Malgache. Switzerland. VIII. Colubridae nouveaux. Bulletin de la Mus6e National de His- Perrier de la B~thie, H. 1921. La v6g6tation Malgache. Annales du toire naturelle, Paris, 4th Series, Section A 10:135-146. • Mus~e Colonial de Marseille, 3rd Series 9:1-29. Donque, G. 1972. The Climatology of Madagascar. Pages 87-144 in R. Perrier de la B$thie, H. 1927. Le Tsaratanana, l'Ankaratra et l'Andringi- Battistini and G. Richard-Vindard, editors. Biogeography and ecol- tra. M6moires de l'Acad~mie Malgache 3:1-52. ogy in Madagascar. Junk, The Hague. Perrier de la BSthie, H. 1936. Biog6ographie des Plantes de Madagas- car. Soci6t~ d'Editions G6ographiques, Maritimes, et Coloniales, Guillaumet, J. L. 1984. The vegetation: an extraordinary diversity. Paris. Pages 27-54 in A. Jolly, P. Oberld, and R. Albignac, editors. Key en- Raxworthy, C. J., and R. A. Nussbaum. 1994. A rainforest survey of am- vironments: Madagascar. Pergamon, Oxford, United Kingdom. phibians, reptiles and small mammals at Montagne d'Ambre, Mada- Hamilton, A. C. 1982. Environmental history of East Africa. Academic gascar. Biological Conservation 69:65-74. Press, London. Raxworthy, C. J., and R. A. Nussbaum. 1995. Systematics, speciation, Humbert, H. 1927. La destruction d'une flore Insulaire par le feu. Prin- and biogeography of the dwarf chameleons Brookesia Gray (Rep- cipaux aspects de la v~g~tation ~t Madagascar. M6moires. de l'Acad- tiUa; Sauria; Chamaeleontidae) of northern Madagascar. Journal of ~mie Malgache 5:1-78. Zoology, London 235:525-558. Humbert, H. 1928. V6g~tation des hautes montagnes de Madagascar. Seguier-Guis, M. 1988. Contribution ~ I'inventoire des vertebres de la M~moires de la Soci6t6 de Biogeographie 2:195-220. For6t d'Ambohitantely (Tampoketsa d'Ankazobe). Thesis. Univer- Jenkins, M. D. 1987. Madagascar, an environmental profile. World Con- sit~ de Madagascar, Antananarivo. servation Union, Gland, Switzerland. U.S. Department of the Interior. 1955. Gazetteer 2. Madagascar, Koechlin, J. 1972. Flora and vegetation of Madagascar. Pages 145-190 R~union and the Comoro Islands. Office of Geography (Depart- in R. Battistini and G. Richard-Vindard, editors. Biogeography and ment of the Interior), Washington, D. C. ecology in Madagascar. Junk, The Hague. White, F. 1983. The vegetation of Africa. United Nations Educational, MacPhee, R. D. E., D. A. Burney, and N. A. Wells. 1985. Early Holocene Social, and Cultural Organization, Paris.

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