Lygodactylus mirabilis

Puente, Marta, Glaw, Frank, Vieites, David R. & Vences, Miguel, 2009, Competitividad del Sector Forestal de México Tendencias y Perspectivas, Zootaxa 2103, pp. 1-76 : 34-44

Lygodactylus mirabilis group

Contains: Lygodactylus blanci , L. intermedius , L. mirabilis , and L. montanus

This group, for which the subgenus name Millotisaurus is available, contains four rock- and ground-dwelling species endemic to high elevations on mountain massifs of Central and South-East Madagascar. They can be distinguished from all other Malagasy Lygodactylus by their partially or completely keeled dorsal scales (granular and not keeled in all other species). The four species in this group furthermore all are characterized by having, within one population, specimens with an ocellated dorsal pattern and others with a striated pattern ( Fig. 16 ). This polymorphism is most extreme in L. mirabilis where the pattern differences between the two morphs are very distinct and constant, and less expressed in L. blanci where patterns intermediate between striated and ocellated occur. All species have a mental scale semi-divided by sutures and typically three postmentals (although in L. montanus , a relatively high prevalence of specimens with two postmentals is observed), and the tail is without whorls. The group contains small species such as L. mirabilis , and large species such as L. montanus . The SVL range of adults is 20.4–39.2 mm.

Two subgroups can be distinguished. L. mirabilis and L. intermedius have keeled dorsal scales, are relatively small-sized (SVL 20.4–30.9 mm), the first finger is completely absent or rudimentary without a claw, and males have seven preanal pores (rarely eight). In contrast, L. blanci and L. montanus are larger (SVL 22.8–39.2 mm), the dorsal scales are partially keeled, the first finger is present, without or with a claw, and males have 8–12 preanal pores.

The L. mirabilis group may be a monophyletic unit, and one more species may belong to it ( L. pauliani , distinguished by unkeeled granular scales; here not assigned to any species group). If hypothetically including L. pauliani , then these species would show a transition towards smaller-sized species with more strongly keeled scales, a more strongly expressed ocellated/striated polymorphism, and a stronger reduction of the first finger, in the direction pauliani-blanci-montanus-intermedius-mirabilis (see also Pasteur 1965, 1995). In fact, this order largely corresponds to the habitat of the species towards colder and more humid climates at higher elevations.

minimum-maximum, modal, and number of measured individuals (n).

Lygodactylus blanci Pasteur, 1967

( Fig. 17 )

Lygodactylus blanci Pasteur, 1967a .— Name-bearing type: male holotype, MNHN 1966.1003 .— Type locality: “Mont Ibity, en cohabitation avec L. arnoulti ”, according to the original description.— Other types: paratypes, six specimens, “dans la série de Ch. Blanc, capturés le 16/v/1964 (n° 69.64 de la collection de l’auteur) et, avec le type, à Noël, 1965 (3.66, 6.66, 9.66, 13.66 et 15.66).”, according to the original description.— Etymology: dedicated to Charles P. Blanc.

Diagnosis. A large species, endemic to high elevations on Ibity mountain. Assigned to the L. mirabilis group based on its partially keeled dorsal scales and slight ocellated/striated polymorphism of the dorsal pattern. L. blanci differs from L. mirabilis by being larger (maximum SVL 39.2 mm vs. 29.3 mm), having more preanal pores in males (10–12 vs. 7), a well developed first finger with claw (vs. first finger absent), and partially keeled dorsal scales (vs. fully keeled). It differs from L. intermedius by being larger (maximum SVL 39.2 mm vs. 30.9 mm), by having 10–12 preanal pores in males (vs. seven), a claw on the first finger (vs. claw absent), and partially keeled dorsal scales (versus fully keeled). It differs from L. montanus by presence of a claw on the first finger (vs. absence).

Lygodactylus blanci occurs on Mont Ibity in sympatry with L. arnoulti which differs by a grey (vs. brown) ground colour, granular dorsal scales (vs. partially keeled), and a strongly whorled tail (vs. without whorls).

Description. (1, 2) A large species, adult SVL 22.8–39.2 mm (33.3± 4.87 mm, n=16); (2) TAL 25.6–47.6 mm (38.4± 7.91 mm, n=7); (3) dorsal scales partially keeled; (4, 5) first finger present, with a strong claw; (6) 3–4 pairs of lamellae under fourth toe (3.06±0.24, n=17); (7, 8) mental scale semidivided by sutures, its posterior projection sometimes in contact with first infralabial scale; (9, 10) three usually bisymmetrical postmental scales (2.88±0.33, n=17); (11) 4–5 postpostmental scales (4.94±0.24, n=17); (12) five infralabial scales (5.06±0.55, n=17); (13) 5–7 supralabial scales (6.06±0.65, n=17); (14) one internasal scale (n=17); (15) males with 10–12 preanal pores (11.00±0.63, n=6); (16) tail without whorls, but sometimes with 11–20 rings of faintly different colour; (17, 18) sometimes with 4–5 dorsolateral tubercles, each composed of 1–4 scales; (19) grey-greenish dorsal colour; (20) dorsal pattern ocellated or, sometimes, striated; (21) light ventral colour with some dark spots on throat area, like an irregular striation following the body line; (22) 150–188 dorsal scales along the body (169±26.87, n=2); (23) 53–59 dorsal scales around the body (56±4.24, n=2) ( Figs. 16a–b , 17 ).

Material examined. MNHN 1990.1892 – 1900 (C.P. Blanc, Dec. 1965, Mont Ibity ) , MNHN 1966.1003 (holotype, C.P. Blanc, Dec. 1965, Mont Ibity ) , MNHN 1990.3565 (C.P. Blanc & F. Blanc, Dec. 1965, Mont Ibity ) , MNHN 1990.8 (C.P. Blanc, 16 May 1964, Mont Ibity ) , MNHN 1990.9 – 13 (C.P. Blanc, Dec. 1965, Mont Ibity ) .

Distribution. According to the specimens examined in this study, Lygodactylus blanci is only known from the type locality, the summital area of Mont Ibity. No further localities are mentioned in the literature.

Habitat. Briefly described by Blanc & Blanc (1967). During various expeditions to Mont Ibity and surrounding areas (Col des Tapias) we did not detect this species but instead collected the sympatric Lygodactylus arnoulti . Following indications of C.P. Blanc (personal communication in 2007), we could reach the original collecting locality by ascending the Ibity from Ihasy village in the north-west of the massif (D. R. Vieites, M. Vences & F. Ratsoavina, unpublished data from 2008). We observed numerous L. arnoulti on rocks close to the summit (2000–2100 m elevation), whereas L. blanci was only found on the summital area itself, in an area characterized by large rocks, very little vegetation, and a quartzitic sand substrate. Here, adults ( Figs. 16a–b ) and juvenile specimens were active on rocks and on stones between grass tufts, or found under large stones, in close syntopy with L. arnoulti . The colour in life was brownish, whereas adults as well as small juveniles of L. arnoulti had a very typical and invariable grey colour with dark grey pattern. Neither our own surveys nor those of C.P. Blanc (pers. comm. in 2008) in the southern Amborompotsy part of the massif, ascending from the Col des Tapias, have yielded records of this species.

Lygodactylus intermedius Pasteur, 1995

( Fig. 18 )

Lygodactylus (Millotisaurus) intermedius Pasteur, 1995 .— Name-bearing type: holotype MNHN 1990.50, female, 29 mm SVL (according to the original description)— Type locality: “Prairie altimontaine sèche de Manjarivolo à 1800 m, chaîne de l’Andrianony (au sud-est du Pic Boby)”, in the Andringitra Massif, according to the original description.— Other types: paratypes MNHN 1990.51–56, MNHN 1990.57–59 (females), MNHN 1990.60 (male), MNHN 1990.61–92, MNHN 1990.92–103, MNHN 1990.104 (young male), MHNG 2541.30 (male), MHNG 2541.31 (immature).— Etymology: from the Latin adjective intermedius ("intermediate"), because of the intermediate morphology between L. montanus and L. mirabilis .

Diagnosis. A relatively small species, endemic to high elevations on the Andringitra Massif. Assigned to the L. mirabilis group based on its keeled dorsal scales and clear ocellated/striated polymorphism of the dorsal pattern. Lygodactylus intermedius differs from L. mirabilis and L. blanci by the presence of a rudimentary first finger without a claw (vs. first finger absent in L. mirabilis and first finger present with claw in L. blanci ). Diagnosis from L. montanus is less clear; L. intermedius differs from this species by being smaller (maximum SVL 30.9 mm vs. 37.5 mm), having seven preanal pores in males (vs. 8–11), and having keeled dorsal scales (versus partially keeled), but this latter difference is clearly a gradual one.

Description. (1, 2) relatively small-sized, adult SVL 21.2–30.9 mm (27.52± 2.61 mm, n=40); (2) TAL 26.5–35.6 mm (30.7± 4.95 mm, n=4); (3) dorsal scales keeled; (4, 5) first finger present in most of the cases, but very small, and without a claw; (6) 3–4 pairs of lamellae under fourth toe (3.73±0.45, n=52); (7, 8) mental scale semi-divided by sutures, usually with broad contact between posterior projection of mental scale and first infralabial scale; (9, 10) 2–3 bisymmetrical postmental scales (2.98±0.13, n=56); (11) 5–6 postpostmental scales (5.13±0.33, n=54); (12) 3–6 infralabial scales (4.91±0.63, n=56); (13) 4–7 supralabial scales (5.22±0.66, n=55); (14) 1–3 internasal scales (1.30±0.50, n=56); (15) males with seven preanal pores (n=4) and with enlarged but unpigmented preano-femoral scales; (16) tail without whorls; (17, 18) mostly without dorsolateral tubercles, but two male specimens have 4–5 such tubercles composed of 2–5 scales each (MNHN 1990.53, and MNHN 1990.60); (19) dorsal colour usually grey-greenish, sometimes blackish (melanic); (20) dorsally with a distinct pattern, either striated or ocellated; (21) ventrally light coloured, with irregular dark spots, especially on throat, and sometimes with faint dark longitudinal lines along ventral side ( Figs. 16c–d , 18 ).

Material examined. MNHN 1990.50 (holotype, C.P. Blanc, 1970, Andringitra ) , MNHN 1990.51 – 56 (paratypes, C.P. Blanc, 29–30 Oct. 1970, Andringitra , Andrianony chain) , MNHN 1990.57 (paratype, C.P. Blanc, 23 Nov. 1970, Andringitra , Andrianony chain, "prairie altimontaine seche du Manjarivolo ", 1800 m) , MNHN 1990.58 (paratype, C.P. Blanc, 23 Nov. 1970, Andringitra , Cuvette Boby, 2500–2550 m) MNHN 1990.59 (paratype, C.P. Blanc, 12 Dec. 1970, Andringitra , Andrianony chain, 1800 m, "cours superieur de la Sahatena ") , MNHN 1990.60 (paratype, C.P. Blanc, 17 Dec. 1970, Andringitra , Pic Boby, 2550 m) , MNHN 1990.61 (paratype, C.P. Blanc, 19 Nov. 1970, Andringitra , Andohariana) , MNHN 1990.62 – 64 (paratypes, C.P. Blanc, 9 Dec. 1970, Andringitra , Varavarana) , MNHN 1990.65 – 66 (paratypes, C.P. Blanc, 9 Dec. 1970, Andringitra ) , MNHN 1990.67 – 73 (paratypes, C.P. Blanc, 5–12 Dec. 1970, Andringitra , Andohariana plateau, 2000–2100 m) , MNHN 1990.74 – 75 (paratypes, C.P. Blanc, Nov. 1970, Andringitra , Cirque Boby, 2550–2600 m) , MNHN 1990.76 – 91 (paratypes, C.P. Blanc, Nov. 1970, Andringitra , Pic Boby, 2550–2630 m) , MNHN 1990.92 (paratype, C.P. Blanc, 16 Dec. 1970, Andringitra , southern versant of the Ibory, 2500–2600 m) , MNHN 1990.93 – 102 (paratypes, C.P. Blanc, 16 Dec. 1970, Andringitra , southern versant of the Ibory, 2500–2600 m) , MNHN 1990.103 (paratype, C.P. Blanc, 16 Dec. 1970, Andringitra , southern versant of the Ibory) , MNHN 1990.104 (paratype, C.P. Blanc, 18 Jan.1971, Andringitra , Andohabatomanara) , MNHN 1990.1876 (C.P. Blanc, 17 Dec. 1970, Andringitra , Camp Mont Ibory) .

Distribution. According to the original description, the species is known exclusively from the Andringitra Massif. Here, specimens have been collected from several sites: Manjarivolo (type locality), Andohabatomanara, Andohariana plateau, Cirque Boby, Cuvette Boby, Ibory, Pic Boby, Varavarana.

Habitat. According to Pasteur (1995) the species lives in rocky environments at high altitudes of 2550–2600 m. At least during the rainy season, the species is not easy to observe, given that various expeditions to the Andringitra Massif (F. Glaw & M. Vences in 1994; D. R. Vieites & M. Vences in 2001; D. R. Vieites and M. Puente in 2003) failed to yield any records. Recently, P. Bora has been able to find specimens of L. intermedius at various localities on the Andringitra Massif (personal communication; expeditions in 2006/2007). The individuals collected were not examined for the present study, but it is relevant that at least one of these specimens showed a clearly melanic coloration ( Fig. 16d ; also pictured in Glaw & Vences 2007).

Lygodactylus mirabilis ( Pasteur, 1962)

( Fig. 19 )

Millotisaurus mirabilis Pasteur, 1962 .— Name-bearing type: male holotype (SVL 24 mm), no. 152.59P from Bons-Girot-Pasteur (BGP) collection, according to the original description); catalogued as MNHN 1966.1000 (collected by J. Millot, 8–9 Sep. 1959)— Type locality: “Mont Tsiafajavona, entre 2300 et 2500 mètres.”, according to the original description.— Other types: 47 paratypes all collected by Millot on the Tsiafajavona mountain at 2300 m: "un en juillet 1946 (MNHN 47.1), deux en juin 1947 (MNHN 48.1 et 48.9 d), les autres avec le type ou nés à Rabat d'oeufs récoltés en même temps (BGP 58–9.59, 61.59, 116–51.59 P et 153–57.59 P)." according to the original description.— Etymology: from the Latin adjective mirabilis ("remarkable"): “Je l’ai apellé ... Millotisaurus mirabilis , moins dans le sens de <admirable>, bien que sa pigmentation soit assez peu banale pour un gecko, que dans celui de <remarquable>: il est en effet quelque chose de nouveau non seulement parmi les Lygodactyles, mais même pour la famille des Gekkonidés, tout entière” ( Pasteur 1964).

Lygodactylus (Millotisaurus) mirabilis — Pasteur (1995).

Diagnosis. A small species with an apparently rather short snout, endemic to high elevations on the Ankaratra Massif. Assigned to the L. mirabilis group based on its keeled dorsal scales and clear ocellated/striated polymorphism of the dorsal pattern. It differs from the other three species in the Lygodactylus mirabilis group by the absence of the first finger (present but without claw in L. intermedius and L. montanus , and present with claw in L. blanci ). It further differs from L. intermedius by the absence of distinct dark spots on the throat (versus presence).

Description. (1, 2) A rather small species, adult SVL 20.8–29.3 mm (25.2± 1.7 mm, n=39); (2) TAL 19.9–39.0 mm (28.1 ± 7.10 mm, n=6); (3) dorsal scales keeled; (4, 5) first finger absent, therefore obviously claw on first finger absent as well; (6) 3–4 pairs of lamellae under fourth toe (3.07±0.25, n=73); (7, 8) mental scale semi-divided by sutures, its posterior projection in faint contact with first infralabial scale; (9, 10) usually three bisymmetrical postmental scales (range 2–4; 2.97±0.37, n=73); (11) 5–8 postpostmental scales (5.42±0.70, n=73); (12) 4–7 infralabial scales (4.83±0.68, n=73); (13) 4–7 supralabial scales (5.44±0.62, n=72); (14) 1–2 internasal scales (1.15±0.36, n=72); (15) males with usually seven, rarely eight preanal pores and enlarged but unpigmented preano-femoral scales; (16) tail without whorls; (17, 18) dorsolateral tubercles absent; (19) dorsally brown-grey to greenish; (20) distinct dorsal pattern present, specimens are either striated or ocellated; (21) ventrally with dark spots, especially on throat, which sometimes are arranged as faint ventral or ventrolateral lines; (22) 99–125 dorsal scales along the body (113.67±13.31, n=3); (23) 40–56 dorsal scales around the body (46.33±8.50, n=3); (24) 70–75 ventral scales (72.67±2.51, n=3) ( Figs. 16e–f , 19 ).

Hemipenial structure. Based on the adult male ZSM 389/2000. Hemipenis subcylindrical, total length ca. 3 mm, with a long pedicel, ca. 1.8–2.0 mm in lenth. Apex divided into two short lobes, each ca. 0.5–1 mm in length. Truncus and lobes are covered with fields and serrated ridges of pointed papillae. Sulcus spermaticus formed by a single channel on the pedicel, dividing into two channels, each running to a terminal position on one of the two lobes. No distinct sulcal lips. Sulcal area with some pointed papillae, especially along the lobes ( Fig. 19 ).

Material examined. BMNH 1962.269 (paratype, J. Millot, 8–9 Sept. 1959, Tsiafajavona , Ankaratra, 2300–2500 m) , MNHN 1990.3586 – 3593 (paratypes, J. Millot, 8 Sep. 1959, Tsiafajavona , Ankaratra, 2300–2500 m) , MNHN 1990.3594 – 3598 (paratypes, J. Millot, 8 Sep. 1959, Tsiafajavona , Ankaratra, 2300–2500 m) , MNHN 1990.3606 (C.P. Blanc, 7 Jan. 1972, Tsiafajavona , Ankaratra) , MNHN 1990.3572 – 3585 (paratypes, J. Millot, 8 Sep. 1959, Tsiafajavona , Ankaratra, 2300–2500 m) , MNHN 1966.999 (paratype, J. Millot, 8–9 Sep. 1959, locality not given in catalogue) , MNHN 1900.3601 – 3630 (C.P. Blanc, 7 Jan. 1972, Tsiafajavona , Ankaratra, 2300–2500 m) , MNHN 1990.3631 – 3636 (Llinares, 21 Feb. 1973, Tsiafajavona , Ankaratra, 2500 m) , MNHN 1990.3637 (Llinares, 13 Aug. 1973, Tsiafajavona , Ankaratra, 2500 m) , ZFMK 51146–51147 (F. Glaw & M. Vences, Ankaratra ) , ZSM 388 /2000 (F. Glaw & M. Vences, 12 Feb. 2000, Ankaratra ) , ZSM 389 /2000 (M. Vences, 3 Feb. 2000, Ankaratra ) , ZSM 796 /2003 ( Ankaratra ) , ZSM 1144 /2003 (G. Aprea, Jan. 2003, Tsiafajavona , Ankaratra) , ZSM 1143 /2003 (G. Aprea, 27. Jan. 2003, Tsiafajavona , Ankaratra) .

Distribution. All known specimens so far have been collected on or near Tsiafajavona mountain in the Ankaratra Massif, but during an extensive survey in 2006, we could find the species on additional neighbouring summits of Ankaratra (D. R. Vieites & M. Vences, personal observation). The precise distribution will be subject of a forthcoming paper.

Habitat. Basic ecological and biological data are given and discussed by Pasteur (1962) and Vences et al. (2002). The species is found only in high elevations of the Ankaratra Massif, an area of harsh and very variable weather conditions, with strong day-night temperature differences (2ºC in the night and more than 20ºC during the day). The species lives in montane grassland with some ericoid bushes, especially in areas with groups of stones or rocks. It appears to reproduce both in the dry and wet seasons: the small eggs are laid into the substrate and can be found under stones; they are not glued to each other or to the substrate. Specimens can be found during cloudy weather under stones above 2200 m elevation, but with sunny weather, they become active on the stones, rocks and grass tufts (M. Vences & D. R. Vieites, personal observations in 2006). Additional data on natural history and reproduction are summarized in Vences et al. (2002).

Lygodactylus montanus Pasteur, 1964

( Fig. 20 )

Lygodactylus montanus Pasteur, 1964 .— Name-bearing type: holotype, MNHN 1956.71, female (32.5 + 38 mm)— Type locality: “sommet du mont Ivohibe (Sud-Est central)”, according to the original description.— Other types: paratypes MNHN 1956.72–73.— Etymology: named after its occurrence on mountains.

Diagnosis. A large species, endemic to high elevations on Ivohibe mountain and on the Andohahela/Anosy chains. Assigned to the L. mirabilis group based on its partially keeled dorsal scales and ocellated/striated polymorphism of the dorsal pattern. L. montanus differs from L. mirabilis by being larger (maximum SVL 37.5 mm vs. 29.3 mm), having more preanal pores in males (8–11 vs. 7), having a small but well developed first finger with claw (vs. first finger absent), and by having partially keeled dorsal scales (vs. fully keeled). It differs from L. intermedius by being larger (maximum SVL 37.5 mm vs. 30.9 mm), by having 8–11 preanal pores in males (vs. seven), and having partially keeled dorsal scales (versus fully keeled). It differs from L. blanci by absence of a claw on the first finger (vs. presence).

Description. (1, 2) A large species, adult SVL 23.9–37.5 mm (33.8± 2.3 mm, n=110); (2) TAL 29.2–49.1 mm (38.4± 4.4 mm, n=35); (3) dorsal scales partially keeled; (4, 5) first finger present, small, without claw; (6) 3–4 pairs of lamellae under fourth toe (3.93±0.24, n=125); (7, 8) mental scale semi–divided by sutures, an indistinct contact of posterior projection of mental scale with first infralabial can be present; (9, 10) 2–3 usually bisymmetrical postmental scales (2.86±0.35, n=125); (11) 3–7 postpostmental scales (4.94±0.44, n=125); (12) 3–7 infralabial scales (5.46±0.68, n=125); (13) 4–8 supralabial scales (6.03±0.74, n=125); (14) 1–2 internasal scales (1.31±0.46, n=125); (15) males with 8–11 preanal pores (8.68±0.65, n=39) and distinct but unpigmented enlarged preano-femoral scales; (16) tail without whorls; (17, 18) usually with 3–9 dorsolateral tubercles, each composed of 1–8 scales; (19) grey-greenish dorsal colour; (20) dorsal pattern alternatively ocellated or striated; (21) light ventral colour with some dark spots in throat area, forming an irregular longitudinal striation along the venter; (22) 138–158 dorsal scales along the body (145.50±6.90, n=8); (23) 61–72 dorsal scales around the body (66.38±3.54, n=8) ( Figs. 16g –f , 20 ).

Hemipenial structure. Based on ZSM 5120/2005, adult male from Andohahela. Hemipenis subcylindrical, total length ca. 3 mm, with a large pedicel of ca. 2 mm in length. The apex is divided in two short lobes measuring ca. 1 mm each. The truncus and lobes are covered with fields and serrated ridges of pointed papillae. Sulcus spermaticus formed by one channel, which divides such that along each lobe there is one subchannels. No distinct sulcal lips, and with some fields of pointed papillae in the sulcal area ( Fig. 20 ).

Material examined. MNHN 1956.71 (holotype, J. Millot, Ivohibe , 2100 m) , MNHN 1956.72 – 73 (paratypes, J. Millot, Ivohibe 2100 m) , MNHN 1990.1877 – 1880 (C.P. Blanc, 20 Nov. 1971, Anosy chain) , MNHN 1990.1881 – 1886 (C.P. Blanc, 25 Nov. 1971, Anosy chain, "sommet E") , MNHN 1990.3452 .3486 (Betsch, Guillaumet & Blanc, 1971, Anosy chain, "cuvette") , MNHN 1990.3487 – 3496 (Betsch, Guillaumet & Blanc, 15–17 Nov. 1971, Anosy chain, "cuvette et premiers sommets") , MNHN 1990.3497 – 3511 (Betsch, Guillaumet & Blanc, 21 Nov. 1971, Anosy chain, "cuvette") , MNHN 1990.3512 – 3525 (Betsch, Guillaumet & Blanc, 23 Nov. 1971, Anosy chain, "sommet F") , MNHN 1990.3526 – 3531 (Betsch, Guillaumet & Blanc, 1971, Anosy chain, "sommet F") , MNHN 1990.3532 – 3538 (Betsch, Guillaumet & Blanc, 25 Nov. 1971, Anosy chain, "sommet D") , MNHN 1990.3539 – 3549 (Betsch, Guillaumet & Blanc, Nov. 1971, Anosy chain, "camp du Sommet ") , MNHN 1993.907 – 913 (J. Arnoult, Jan. 1954, Andohahela , 2000 m) , ZSM 5120 / 2005–5126/2005 (F. Glaw, M. Vences & P. Bora , 26–27 Jan. 2005, Andohahela) .

Distribution. According to the specimens examined in this study, Lygodactylus montanus is known from the following localities: (1) Ivohibe (type locality), (3) Andohahela, and (3) Anosy chain (various localities on the massif). No further localities are mentioned in the literature.

Habitat. According to Pasteur (1964) this species lives on rocks. At Andohahela, we found specimens during cloudy weather under stones which were laying on large and rather smooth outcrops of granitic rock.

Remark. In the catalogue of the Paris museum, the whole series MNHN 1990.3452–3549 is marked as originating from "Chaines Anosyennes du Massif l'Andringitra" or "Chaines Anosyennes. Massif de l'Andringitra", collected by C.P. Blanc or by the "Mission Betsch-Guillaumet-Blanc". However, we believe that all these specimens originate from the Anosy chain (=Chaines Anosyennes) and not from Andringitra for the following reasons: (1) they are marked as having been collected in November 1971 (some of them more precisely on 21–25 November 1971), which agrees with the dates at which C.P. Blanc had been collecting on the Chaines Anosyennes, as can be seen in the series of L. montanus MNHN 1990.1881–1886, or in numerous series of frogs collected from this massif (e.g., Gephyromantis spinifer ; Vences & Glaw 2001); (2) C.P. Blanc collected at Andringitra in the period between October 1970 and January 1971, as indicated by specimens of Lygodactylus pictus (MNHN 1990.3550–3564) and L. intermedius (MNHN 1990.60, MNHN 1990.61, MNHN 1990.62–64, MNHN 1990.65–66, MNHN 1990.67–73, MNHN 1990.74–75, MNHN 1990.76–91, MNHN 1990.92–102, MNHN 1990.103, MNHN 1990.104, MNHN 1990.50).

Variation. It is biogeographically interesting that this species occurs on Ivohibe and the more southern massifs of Anosy and Andohahela, whereas Andringitra (located rather close to Ivohibe) harbours a different species, L. intermedius . We could not detect any clear difference between the specimens from Ivohibe (the type material) and those from the Anosy chain. The Ivohibe specimens have two asymmetrical postmental scales, versus mostly three postmental scales in Anosy specimens, but also among the Anosy sample we observed specimens with two postmentals (11 cases).

One specimen (MNHN 1990.3536) catalogued as originating from "Sommet D" (referring to the Anosy Chain, collected on 25 Nov. 1971) is identified as L. montanus in the MNHN catalogue but differs morphologically: the specimen is a male of 34.2 mm SVL; granular dorsal scales (not partially keeled like in L. montanus ); first finger with claw ( L. montanus without claw). Values from this specimen have not been included in the description of morphological characters of L. montanus given above.

Figures

FIGURE 16. Photographs of living specimens attributed to species in the Lygodactylus mirabilis group. (a) and (b) male of L. blanci from Mt. Ibity; (c) and (d) specimens of L. intermedius from Andringitra; note melanic colouration in (d); (e) and (f), striped and ocellated specimens of L. mirabilis, Ankaratra Massif; (g) and (h), striped and ocellated specimens of L. montanus from Andohahela massif. All photographs by F. Glaw & M. Vences, except (c) which was taken by P. Bora.

FIGURE 17. Scalation characters of Lygodactylus blanci. (a) head in lateral view (scale bar=2 mm), based on of female specimen MNHN 1990.1897; (b) lamellae of fourth toe in ventral view (scale bar=2 mm), based on female specimen MNHN 1990.1895; (c) head in dorsal view (scale bar=2 mm), based on female specimen MNHN 1990.1897; (d) throat in ventral view (scale bar=2 mm), based on female specimen MNHN 1990.1897; (e) preanal pores in ventral view (scale bar=2 mm), based on male specimen MNHN 1990.10. All drawings by M. Puente.

FIGURE 18. Scalation characters of Lygodactylus intermedius. (a) head in lateral view (scale bar = 2 mm); (b) lamellae of fourth toe in ventral view (scale bar = 2 mm); (c) head dorsal view (scale bar = 2 mm); (d) throat in ventral view (scale bar = 2 mm); (e) preanal pores in ventral view (scale bar = 2 mm). Drawings based on male specimen MNHN 1990.52, except for (b) that is based on female specimen MNHN 1990.54. All drawings by M. Puente.

FIGURE 19. Scalation and hemipenial characters of Lygodactylus mirabilis. (a) head in lateral view (scale bar = 2 mm); (b) head dorsal view (scale bar = 2 mm); (c) throat in ventral view (scale bar = 2 mm); (d) preanal pores in ventral view (scale bar = 2 mm); (e) B, lamellae of fourth toe in ventral view (scale bar = 2 mm); (f) and (g), sulcal and asulcal view of the hemipenis (scale bar = 1 mm). Drawings based on male specimen MNHN 1990.3605 except for (b) that is based on male specimen MNHN 1990.3582 and (f) and (g) that are based on male specimen ZSM 389/2000. All drawings by M. Puente.

FIGURE 20. Scalation and hemipenial characters of Lygodactylus montanus. (a) head in lateral view (scale bar = 2 mm); (b) head in dorsal view (scale bar = 1 mm); (c) throat in ventral view (scale bar = 1 mm); (d) preanal pores in ventral view (scale bar = 2 mm); (e) lamellae of fourth toe in ventral view (scale bar = 1 mm); all based on based on male specimen MNHN 1990.3499; (f) and (g), sulcal and asulcal view of the hemipenis, based on male specimen ZSM 5120/ 2005 (scale bar = 1 mm). All drawings by M. Puente.