ARTÍCULO: DESCRIPTION OF A NEW SPECIES OF GROSPHUS SIMON (SCORPIONES, BUTHIDAE) FROM THE ANKARANA MASSIF, MADAGASCAR Wilson R. Lourenço & Steven M. Goodman Abstract Some morphological characteristics of Grosphus flavopiceus Kraepelin, 1900 are revised. A new species, Grosphus ankarana sp. n., related to G. flavopiceus, is described from the Ankarana Massif in the province of Antsiranana (Diégo Suarez), Madagascar. The morphology of the basal middle lamellae of the pectines is illustrated by using scanning electron microscopy. A revised key to the 12 known species of this genus is presented. Key words: Scorpions, Grosphus, new species, speciation, Madagascar, Ankarana Massif. ARTÍCULO: Taxonomy: Grosphus ankarana sp. n. Description of a new species of Descripción de una nueva especie de Grosphus Simon (Scorpiones, Buthidae) Grosphus Simon (Scorpiones, del Macizo Ankarana, Madagascar Buthidae) from the Ankarana Massif, Madagascar Resumen Wilson R. Lourenço Se revisan los caracteres morfológicos de Grosphus flavopiceus Kraepelin, 1900. Se Département de Systématique describe una nueva especie, Grosphus ankarana sp. n., próxima a G. flavopiceus, del et Evolution, Section Macizo de Ankarana, en la provincia de Antsiranana (Diégo Suarez), Madagascar. La Arthropodes (Arachnologie), morfología de la lamela basal media de las pectinas es ilustrada usando microscopio Muséum National d’Histoire electrónico. Se presenta una clave revisada para las 12 especies conocidas del género. Naturelle, Palabras clave: Escorpiones, Grosphus, nueva especie, especiación, Madagascar, Ankarana 61 rue de Buffon Massif. 75005 Paris, France Taxonomía: Grosphus ankarana sp. n. [email protected] Steven M. Goodman Field Museum of Natural History, Introduction Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605, As discussed in recent papers (Lourenço, 1999, 2001, 2003), the first Grosphus USA, species to be described was Scorpio (Androctonus) madagascariensis Gervais and WWF, BP 738, (1843/1844) [=Grosphus madagascariensis] (Gervais, 1843/1844). Kraepelin Antananarivo (101), Madagascar (1900) contributed to the study of the genus Grosphus, and described several new [email protected] species. In his monograph on the scorpions of Madagascar, Fage (1929) described a new variety of Grosphus limbatus, which he named annulata and was subse- Revista Ibérica de Aracnología ISSN: 1576 - 9518. quently raised to specific rank as G. annulatus (Lourenço, 1996). After Fage’s Dep. Legal: Z-2656-2000. (1929) monograph only four new species of this genus have been described Vol. 7, 30-VI-2003 (Lourenço, 1996, 1999, 2001, 2003). Sección: Artículos y Notas. The original descriptions of these Grosphus species were in most cases quite Pp: 19–28 clear and the associated proposed distinguishing characters have been confirmed in Edita: subsequent diagnoses (Fage, 1929; Lourenço, 1996), although some features may Grupo Ibérico de Aracnología require re-examination. The taxonomy of Grosphus is based mainly on two (GIA) principal characters: the pattern of coloration, and the morphology of the basal Grupo de trabajo en Aracnología de la Sociedad Entomológica middle lamellae of the female pectines. For some species, such as G. flavopiceus, Aragonesa (SEA) recent studies (see Lourenço, 1996) were based on insufficient material. Moreover, Avda. Radio Juventud, 37 the precise distributional range of this species was not properly understood. For the 50012 Zaragoza (ESPAÑA) Tef. 976 324415 present study, freshly collected material was examined and the morphology of the Fax. 976 535697 basal middle lamella of the pectines studied using scanning electron microscopy. As C-elect.: [email protected] already suspected by Fage (1929; see discussion), two species are in fact present Director: A. Melic over what was previously considered the distributional range of G. flavopiceus. A Información sobre suscripción, new species of Grosphus, related to G. flavopiceus is herein described from the índices, resúmenes de artículos on Ankarana Massif. line, normas de publicación, etc. en: Página web GIA: http://entomologia.rediris.es/gia Ecology and Biogeography of the Ankarana Massif Página web SEA: The Ankarana Massif is an outcrop of middle Jurassic limestone, oriented NE-SW http://entomologia.rediris.es/sea with about 25 km long and 8 km large (Fig. 1). Quaternary earth movements (circa 1.5 MYA) resulted in the splitting of the massif and elevating its western wall 20 Wilson R. Lourenço & Steven M. Goodman Fig. 1. Map showing the location of Ankarana Massif and Montagne des Français (Modified from Bardot-Vaucoulon, 1997). (Wilson, 1987), which is now marked by vertical cliffs with an average daily temperature of 27.5°C (Bardot- of 100 to 150 m high. A series of volcanic eruptions Vaucoulon, 1997). over the course of recent geological time have produced The narrow and often deep canyons within the a series of lava flows in to the area, including to the east massif regularly have tall canopy semi-evergreen forests along the Besaboba River (2 MYA), to the west along resting on slightly acidic soils of basaltic rock. These the Analatelo Plain (1 MYA), and directly into the canyons provide a buffer against the relatively extreme canyons of Ankarana (0.5 MYA). The vast majority of dry conditions of this general region of Madagascar, in the massif is contained with the Réserve Spéciale (RS) particularly shelter against the sun and often-strong d’Ankarana and the highest point in the reserve is at winds (Hawkins et al., 1990). Further, an extensive about 500 m. subterranean aquifer passing under the Ankarana The Ankarana Massif lies at the extreme northern resurges in these canyons or is close to the surface level end of Madagascar about 75 km south of Antsiranana and helps to maintain mesic conditions even during the (Diégo Suarez, 12°55’S, 49°06’E) and to the southwest heart of the dry season. The forested habitat within the of Montagne d’Ambre – this latter site is an isolated deep canyons of the Ankarana form an ecological volcano and associated with orographic factors the island, isolated from the surrounding areas by dry upper slopes of which received considerable rainfall deciduous forests or anthropogenic savannas. The biota and hold humid forests. The Ankarana region is slightly of these canyons contain many endemic plant and drier and with slightly less than 2000 m of rainfall each animal species (Cardiff and Befourouack, in press), year (Hawkins et al., 1990). Even with this relatively some with affinities of modern eastern humid forests high annual precipitation the site has dry deciduous and others to the western deciduous forests. On the vegetation and this is due to an extended dry season basis of subfossil evidence it is clear that the massif (Lowry et al., 1997), which lasts from May to Novem- supported more mesic conditions during the Quaternary ber. The average daily temperature during this season is (Godfrey et al., 1996, 1999; Jungers et al., 1995) and 26°C. The rainy season lasts from December to April, the remaining canyon forests hold some relict taxa of and accounts for 93% of the annual precipitation, and this more mesic period. Description of a new species of Grosphus Simon from the Ankarana Massif 21 Fig. 2-5. Grosphus ankarana sp. n. 2-3: Male holotype, dorsal and ventral aspects. 4-5: Idem, Female paratype. Taxonomic Treatment 12° 54.5’ S – 49° 06.6’ E. (S. Goodman, leg.), 6 males Description of a new species and 8 females (paratypes). Holotype and paratypes deposited inthe Field Museum of Natural History, Grosphus ankarana sp. n. Chicago, with the exception of one paratype deposited (Figs. 2-11, 16-24) in the Muséum National d’Histoire Naturelle, Paris, and Grosphus flavopiceus; Fage, 1929: 649 (in part) two paratypes deposited in the Natural History Mu- seum, Geneva. MATERIAL EXAMINED: Madagascar, Province d’Antsi- The new species was collected together with ranana, Réserve Spéciale d’Ankarana, 2.6 km E Andra- several specimens of Grosphus madagascariensis, also fiabe, in forest near Andrafiabe Cave, 12° 55.9’ S – 49° in the collections of the Field Museum of Natural 03.4’ E, 20621/I/2001, about 50 m, dry deciduous forest History. at base of tsingy, pitfall-traps (S. Goodman, leg), FMHD # 01-41. 1 male (holotype), 10 males and 3 ETYMOLOGY: The name of the type locality (Ankarana females (paratypes). Campement des Anglais (Anilotra), Massif) is placed in apposition to the generic name. 22 Wilson R. Lourenço & Steven M. Goodman Fig. 6-11. Grosphus ankarana sp. n. (female paratype). 6: Pecten showing the basal middle lamella. 7: Basal middle lamellae, in detail. 8: Microstructure of peg sensilla on one tooth. 9-11: Peg sensilla in detail. DIAGNOSIS: The morphology of the new species shows DESCRIPTION BASED ON MALE HOLOTYPE AND ONE that it is close to Grosphus flavopiceus Kraepelin, but FEMALE PARATYPE. it can be readily distinguished from that species by the Measurements in Table I. following characters: (i) overall larger size – see Table I; (ii) greater number of pectinial teeth in both males Coloration. Basically reddish-yellow to gray-yellowish and females – see Table II; (iii) 14 to 15 rows of or gray-reddish. Prosoma: carapace reddish-yellow; granules on the pedipalp tibia fingers; and (iv) more eyes surrounded by black pigment. Mesosoma: gray- elongated and curved basal middle lamellae of the yellowish. Metasoma: segment I gray-yellowish; female pectines, covering the first 3 proximal teeth. segments II-IV reddish-yellow; segment V reddish. Description of a new species of Grosphus Simon from the Ankarana Massif 23 Fig. 12-15. Grosphus flavopiceus. 12: Pecten showing the basal middle lamella. 13: Microstructure of peg sensilla on one tooth. 14-15: Peg sensilla in detail. Vesicle dark reddish; aculeus with reddish base and constant in wide from the base to the apex, covering the blackish tip. Venter: coxapophysis, sternum and genital 3 proximal teeth. Sternites smooth, with elongated operculum gray-yellowish; pectines yellowish with stigmata; III-VI with two longitudinal furrows; VII with some diffuse gray spots; sternites gray-yellowish. four vestigial carinae. Metasoma: segments I and II with Chelicerae reddish-yellow without any dark variegated 10 carinae, moderately crenulate. Segments III and IV pigmentation along its entire surface; fingers reddish with 8 carinae, weakly crenulate. Segment V with 5 with teeth blackish. Pedipalps: femur yellowish, patella carinae. Dorsal carinae on segments I-IV with one or and tibia reddish with vestigial dark zones on patella two posterior spinoid granules. Intercarinal spaces and tibia; rows of granules on fingers blackish. Legs weakly granular to smooth. Telson smooth; aculeus yellowish without spots. strongly curved and slightly shorter than the vesicle; Morphology. Carapace moderately to strongly granu- subaculear tooth absent. Cheliceral dentition characte- lar; anterior margin almost straight, with a weak median ristic of the family Buthidae (Vachon, 1963); two concavity. All carinae weak; furrows moderately to distinct basal teeth present on the movable finger, but strongly developed. Median ocular tubercle anterior to very reduced; ventral aspect of both fingers and of the center of carapace; median eyes separated by more manus with dense, long setae. Pedipalps: femur pentaca- than one ocular diameter. Three pairs of lateral eyes. rinate; patella with a dorsointernal carina and with Sternum between sub-triangular and sub-pentagonal in several spinoid granules on the internal face; tibia shape. Mesosoma: tergites with a weak to moderate smooth, without carinae, all faces weakly granular to granulation. Median carina moderately developed in all smooth. Movable and fixed fingers with 14/15 oblique tergites. Tergite VII pentacarinate. Venter: genital rows of granules in males and females. Trichobothrio- operculum consisting of two subtriangular plates. taxy; orthobothriotaxy A-α (Vachon, 1974, 1975). Pectines: pectinal teeth count 40-41 (for variation see Legs: tarsus with numerous short thin setae ventrally. Table II); basal middle lamellae of each pecten not Tibial spurs present on legs III and IV; pedal spurs dilated in males; elongated and curved in females; present on legs I to IV; all spurs strong. 24 Wilson R. Lourenço & Steven M. Goodman Table I Morphometric values (in mm) of the male holotype and female paratype of Grosphus ankarana sp. n., and of male and female of Grosphus flavopiceus from the Montagne des Français area. Grosphus ankarana sp. n. Grosphus flavopiceus holotype paratype male female Total length 114.5 105.2 68.6 69.1 Carapace: length 10.9 11.4 7.4 8.2 anterior width 8.1 8.7 5.5 6.1 posterior width 11.8 12.9 8.0 9.3 Metasomal segment I: length 11.2 9.8 5.8 5.9 width 5.8 6.8 4.4 5.5 Metasomal segment V: length 15.3 13.2 9.4 8.8 width 5.2 5.5 4.2 5.0 depth 5.1 5.2 3.8 4.6 Vesicle: width 5.1 5.6 3.6 4.5 depth 5.1 5.6 3.6 4.3 Pedipalp: Femur length 12.2 11.3 7.5 7.5 Femur width 2.8 3.3 2.1 2.4 Patella length 12.9 12.7 8.4 8.9 Patella width 3.7 4.5 3.0 3.4 Chela length 20.7 20.9 13.9 14.7 Chela width 4.6 5.4 4.0 4.2 Chela depth 4.5 5.2 3.7 3.8 Movable finger: length 11.2 12.1 8.1 8.8 Table II Discussion Variation in pectinal tooth count in Grophus Fage (1929) in his monograph on the scorpions of ankarana sp. n., and in G. flavopiceus. Madagascar, noted the existence of two different forms Number G. ankarana G. flavopiceus within animals he referred to Grosphus flavopiceus. He of teeth Males Females Males Females remarked (pp. 648-650, translated from the French), 24 0 0 0 6 “The description proposed here applies to the specimens 25 0 0 0 14 deposited in the Muséum (Paris), and used by Kraepelin 26 0 0 0 6 (1900) for the original description. These are from 27 0 0 0 0 Diégo, Antsirana [sic] and Tamatave. There are, howe- 28 0 0 2 0 ver, in the collections specimens from Ambilobe, 29 0 0 4 1 collected by M. Waterlot which differ from the type 30 0 0 1 1 material in several points. The females range from 50 to 31 0 2 0 0 107 mm, and the males from 68 to 75 mm. They show 32 0 8 0 0 a greater number of pectinial teeth: 32-33 for females 33 0 5 0 0 and 33-37 for males and the basal middle lamella is 34 0 2 0 0 longer, not conic and rounded in its extremity (see also 35 0 3 0 0 fig. 4, a-b in page 648).” [Some of the specimen mate- 36 2 0 0 0 rial studied by Fage was not located in the Paris collec- 37 6 0 0 0 tions.] 38 7 0 0 0 Almost without a doubt, Fage had before him 39 11 0 0 0 material of Grosphus ankarana and, probably given the 40 4 0 0 0 size variation he presented, some of the individuals 41 4 0 0 0 were juveniles. Although there appear to be characters to differentiate G. flavopiceus from the “Ambilobe” Remarks: Coloration in females is slightly paler. The animal, Fage did not name the latter as a new species to body is bulkier but the total length is smaller than in science. This hesitation might be related to two different males (see Table I for morphometric values). points concerning members of the genus Grosphus: a Description of a new species of Grosphus Simon from the Ankarana Massif 25 Fig. 16-24. Grosphus ankarana sp. n. 16-23: Male holotype. 16: Movable finger of tibia pedipalp. 17: Detail of distal extremity of the same. 18: Metasomal segment V and telson, lateral aspect. 19-22: Trichobothrial pattern. 19: Chela, dorso-external aspect. 20: Chela, ventral aspect. 21: Patella, dorsal aspect. 22: Femur, dorsal aspect. 23: Chelicera. 24: Idem female paratype. Fig. 25-30. Grosphus flavopiceus (Montagne des Français). 25: Chelicera (female). 26: Metasomal segment V and telson, lateral aspect. 27-30: Trichobothrial pattern. 27: Chela, dorso-external aspect. 28: Chela, ventral aspect. 29: Patella, dorsal aspect. 30: Femur, dorsal aspect. 26 Wilson R. Lourenço & Steven M. Goodman lack of information on the intraspecific variability of 30 cm tall buckets. Clearly based on the placement of certain characters and an unclear view of the biogeo- these devices, this scorpion moves on the forest floor. graphic patterns of the different names species (see also Further, a number of individuals were observed and discussion in Lourenço, 2003). Given that at present collected at night climbing on exposed limestone rock this species is only known from the Ankarana Massif, forming the edge of the canyon walls or isolated out- the question remains if his specimens were indeed from crops within the canyon forest or on vertical tree trunks. Ambilobe, or this was simply the nearest town to the During a night walk in the Andrafiabe Forest a very collection site. hard rain fell and an individual of G. ankarana sp. n. The morphology of the basal middle lamellae of dropped from the tree canopy on to the head of one the female pectines proved to be a diagnostic character researcher. Thus, on the basis of current information for Grosphus species (Lourenço, 1996, 2003). In fact, this species is both terrestrial and arboreal. Fage (1946) himself emphasized this characteristic in At our campsites this scorpion was often found regard to the species of the Sambirano and Ankarana. under personnel or research gear left in the open during Moreover, Grosphus species show, almost without the night. On one occasion a member of our research exception, very clear and discrete distributional patterns team was stung on the hand by an individual of G. (Lourenço, 1996; Lourenço & Goodman, 1999). We ankarana sp. n. while removing it with a large forceps suggest here that on the basis of current information the from a pitfall bucket. She experienced a painful swe- new species G. ankarana sp. n. is endemic to the lling of the pain and stomach nausea that lasted for Ankarana Massif. several hours. It is not certain that she received a full dose of this scorpion’s venom. The number of tourists visiting the Réserve Spéciale d’Ankarana has been Ecology of Grosphus ankarana increasing in recent years (Cardiff and Befourouack, in Grosphus ankarana is known to occur in several press) and many of these visitors camp at several sites different habitats in the Ankarana. A considerable within the reserve. Given that G. ankarana sp. n. is proportion of specimens were obtained in 121 pitfall notably common at some of these campsites they pose buckets placed in lines on the forest floor. These lines a potential threat to these visitors. However, on the were in intact forest and were located in areas of canyon basis of current information the sting of this animal is forest with sufficient soil and humus to dig in the nearly not particularly poisonous. Key to the species of the genus Grosphus 1 Pectines with a maximum of 21 teeth .................................. ................. 2 – Pectines with more than 22 teeth ...................................... ................. 4 2 Coloration yellowish to reddish yellow, with variegated brownish pigmentation; body length about 30 mm ........................................... .......... G. garciai – Coloration dark, from reddish brown to dark brown; body length about 50 mm . ................. 3 3 Coloration reddish brown to dark brown, without light spots; metasomal segment I longer than wide; basal middle lamellae of female pectines oval in shape ...... .. G. madagascarienis – Coloration reddish brown with lighter spots; metasomal segment I wider than long; basal middle lamellae of female pectines subquadrangular in shape .......... ........... G. hirtus 4 Coloration blackish or reddish brown to yellowish; pectines with 30 to 40 teeth; body length more than 90 mm ........................................ ................. 5 – Coloration reddish brown to yellowish, never blackish; body length less than 90 mm................. 6 5 Coloration blackish throughout ....................................... ....... G. grandidieri – Coloration reddish brown to yellowish ................................. ... G. ankarana sp. n. 6 Mesosoma with homogenous coloration, reddish brown or yellowish ......... ................. 7 – Mesosoma with a blackish median longitudinal band, or with two blackish lateral longitudinal bands ................................................. ................ 10 7 Total length more than 65 mm; mesosoma reddish brown; basal middle lamellae of female pectines two times longer than wide at their base ................... ....... G. flavopiceus – Total length less than 60 mm; mesosoma yellowish; basal middle lamellae of female pectines three times longer than wide at the base ......................... ................. 8 Description of a new species of Grosphus Simon from the Ankarana Massif 27 8 Metasomal segment V and telson pale yellowish ......................... ....... G. intertidalis – Metasomal segment V and telson with blackish spots or blackish throughout ... ................. 9 9 Metasomal segment V and telson with blackish spots ..................... ........ G. annulatus – Metasomal segment V and telson blackish .............................. ............. G. feti 10 Mesosoma with a wide blackish median longitudinal band; basal middle lamellae of female pectines three times longer than wide at their base and covering the 4 proximal teeth .................................................... ......... G. limbatus – Mesosoma with two narrow blackish lateral longitudinal bands.............. ................ 11 11 Carapace without a blackish triangular spot; basal middle lamellae of female pectines weakly curved, widening in proximal half and covering the two proximal teeth ............................................................ ........ G. bistriatus – Carapace with a blackish triangular spot; basal middle lamellae of the female pectines curved and constantly narrowing from the base to apex covering the four proximal teeth .................................................... .... G. ankarafantsika Acknowledgements GODFREY, L. R., W. L. JUNGERS, E. L. SIMONS, P. S. CHA- TRATH & B. RAKOTOSAMIMANANA 1999. Past and We are very grateful to Laurent Albenga and Régis present distributions of lemurs in Madagascar. In New Cleva (Muséum National d’Histoire Naturelle, Paris) directions in lemur studies, B. Rakotosamimanana, H. for technical help with the preparation of SEM and Rasamimanana, J. U. Ganzhorn, and S. M. Goodman digital photos. For permission to work in the Ankarana (eds.), pp. 19-53. New York: Kluwer Academic / we would like to thank the Association National pour la Plenum Publishers. Gestion des Aires Protégées (ANGAP) based in Anta- HAWKINS, F. A., P. CHAPMAN, J. U. GANZHORN, Q. M. C. nanarivo and Ambilobe. Funds to conduct this field- BLOXAM, S. C. BARLOW & S. J. TONGE 1990. Vertebrate conservation in Ankarana Special Reserve, northern work were provided by the National Geographic Society Madagascar. Biological Conservation, 54: 83-110. and the Volkswagen Foundation. JUNGERS, W. L., L. R. GODFREY, E. L. SIMONS & P. S. CHA- TRATH 1995. Subfossil Indri indri from the Ankarana massif of northern Madagascar. American Journal of References Physical Anthropology, 97: 357-366. KRAEPELIN, K. 1900. Üeber einige neue Gliederspinnen. BARDOT-VAUCOULON, M. 1997. Observations sur le milieu et Abhandlungen aus dem Gebiete der Naturwissenschaf- la végétation du Massif de l’Ankarana (Nord de Mada- ten, 16: 3-17. gascar) et description de trois nouvelles espèces de Chlorophytum (Liliaceae), Tacca (Taccaceae) et Adenia LOURENÇO, W. R. 1995. Description de trois nouveaux genres et quatre nouvelles espèces de Scorpions Buthi- (Passifloraceae). Adansonia, sér. 3, 19(1): 139-163. dae de Madagascar. Bulletin du Muséum National CARDIFF, S. & J. BEFOUROUACK (in press). The Réserve d’Histoire Naturelle, Paris, 4e sér. 17(1-2): 95-106. Spéciale d’Ankarana, S. M. Goodman and J. P. Bens- tead (eds.): The Natural History of Madagascar, LOURENÇO, W. R. 1996. Scorpions (Chelicerata, Scorpiones). In: Faune de Madagascar N° 87. Muséum National Chicago: University of Chicago Press. d’Histoire Naturelle, Paris: 102p. FAGE, L. 1929. Les Scorpions de Madagascar. Faune des Colonies françaises 3. Société d’Editions Géographi- LOURENÇO, W. R. 1999. A new species of Grosphus Simon ques, Maritimes et Coloniales, Paris, pp. 637-694. (Scorpiones, Buthidae), the first record of an intertidal FAGE, L. 1946. Complément à la Faune des Arachnides de scorpion from Madagascar. Entomologische Mitteilun- Madagascar. Bulletin du Muséum National d’Histoire gen aus dem Zoologischen Museum Hamburg, 12(158): Naturelle, 2e sér., 18(3): 256-267. 297-307. GERVAIS, P. 1843. (Les principaux résultats d’un travail sur la LOURENÇO, W. R. 2001. Another new species of Grosphus famille des Scorpions). Société Philomatique de Paris. (Scorpiones, Buthidae) for Madagascar. Revue suisse de Extraits des Procès-Verbaux des Séances, 5(7): 129- Zoologie, 108(3): 455-461. 131. LOURENÇO, W. R. 2003. New taxonomic considerations on GERVAIS, P. 1844. Remarques sur la famille des Scorpions et some species of the genus Grosphus Simon, with description de description de plusieurs espèces nouve- description of a new species (Scorpiones, Buthidae). lles de la collection du Muséum. Archives du Muséum Revue suisse de Zoologie, 110(1). d’Histoire Naturelle, Paris 4: 201-240. LOURENÇO, W. R. & S. M. GOODMAN 1999. Taxonomic and GODFREY, L. R., J. M. WILSON, E. L. SIMONS, P. D. STEWART ecological observations on the scorpions collected in & M. VUILLAUME-RANDRIAMANANTENA 1996. Ankara- the Réserve Naturelle Intégrale d’Andohahela, Mada- na: Window to Madagascar's past. Lemur News, 2: 16-17. gascar. Fieldiana : Zoology, new series, 94: 149-153. 28 Wilson R. Lourenço & Steven M. Goodman LOWRY, P. P. II, G. E. SCHATZ & P. B. PHILLIPSON 1997. The pions. Bulletin du Muséum National d’Histoire Nature- classification of natural and anthropogenic vegetation lle, Paris, 3è sér., n° 140, Zool. 104: 857-958. in Madagascar. 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