Lygodactylus roavolana, Puente & Glaw & Vieites & Vences, 2009

Puente, Marta, Glaw, Frank, Vieites, David R. & Vences, Miguel, 2009, Competitividad del Sector Forestal de México Tendencias y Perspectivas, Zootaxa 2103, pp. 1-76 : 24-32

Lygodactylus pictus group

Contains: Lygodactylus pictus , L. tuberosus and L. roavolana sp. nov.

This is a poorly defined group of Malagasy Lygodactylus species that are mainly characterized by a semidivided mental scale (divided into three parts by incomplete sutures), and only two postmental scales. However, it needs to be stressed that like most meristic characters in Lygodactylus , there is variation in the number of postmentals, and some individuals assigned to species in this group indeed have three postmentals. Other characters shared by these species are: granular dorsal scales, 3–4 pairs of subdigital lamellae on the fourth toe, 6–8 preanal pores in males, no whorls on tail, absence of distinct dorsal pattern ( Fig. 12 ). The first finger is present, and it can bear a claw ( L. pictus , L. tuberosus ) or not ( L. raovolana ). The hemipenes are characterized by being bilobed with two rather short lobes, a long pedicel, and fields and serrated ridges with pointed papillae covering the asulcal side.

Lygodactylus pictus ( Peters, 1883)

( Fig. 13 )

Scalabotes pictus Peters, 1883 .— Name-bearing type: no data from original description; male lectotype (designated by Pasteur 1965: 94) ZMB 10453 according to Bauer et al. (1991), not examined by us.— Type locality: “Es fehlt leider die genaue Angabe des Fundorts, aber sie gehören unzweifelhaft dem Centrum dieser grossen Insel an”, thus central Madagascar, according to the original description.— Other types: no data from the original description; three males (ZMB 104454A–C), same data as lectotype and ZMB 15818 from " Madagascar ".— Etymology: probably based on the coloration, derived from Latin pictus (painted): “Unten schmutziggelb, am Unterlippenrande und in der Submentalgegend schwarz punktirt”, according to the original description.

Lygodactylus robustus Boettger, 1913 .— Name-bearing type: male lectotype SMF 8956 from Ankarimbela, designated by Mertens (1967).— Type locality: “Süd und Südost- Madagascar, die 25 vorliegenden Originale von Ankarimbela.”, according to the original description, but restricted to Ankarimbela by the lectotype designation.— Other types: According to Boettger (1913) the original description was apparently based on 30 syntypes (Senckenberg No. 4160,6a including 25 syntypes from Ankarimbela and Senckenberg No. 4160,4b with 5 syntypes (3 females, 2 juveniles) from Fianarantsoa.— Etymology: Derived from Latin robustus (robust).

Diagnosis. L. pictus is a relatively large species of Lygodactylus that is common on the central high plateau of Madagascar. It can be assigned to the L. pictus group based on the characters listed in the group definition above. It differs from L. tuberosus by usually having distinct dark spots on the throat (vs. no or only faint spots in L. tuberosus ), by a yellow ventral colour in life (usually whitish and not distinctly yellow in L. tuberosus ; the yellow pigment, however, quickly fades in preservative), and by the presence of only a dark stripe behind the eye (vs. presence in most individuals of a dark marking behind the eye which includes spiny scales). Typically, specimens of L. pictus are relatively large, but also some individuals classified as L. tuberosus can attain large sizes (up to 38 mm). Clearly, the definitions of these species require revision, and the specimens assigned here to either L. pictus or L. tuberosus do not form homogeneous groups. Distinction of L. pictus from L. roavolana is easy by the absence of distinct black markings above the forelimbs (vs. presence), by the number of 4–6 infralabial scales (vs. 7), and by the presence of a claw on the first finger (vs. apparent absence).

Description. (1, 2) A large Lygodactylus , adult SVL 20.4–37.7 mm (32.7± 3.4 mm, n=40; only a single young male being below 26 mm); (2) TAL 26.7–52.5 mm (37.0 ± 6.7 mm, n=15); (3) granular dorsal scales; (4, 5) first finger present, bearing a claw; (6) 3–4 pairs of subdigital lamellae on the fourth toe (3.38± 0.49 mm, n=45); (7, 8) mental scale semi-divided, its posterior projection can be in indistinct contact with the first infralabial; (9, 10) usually two (but sometimes three) symmetrical postmental scales (2.24±0.48, n=45); (11) usually five postpostmental scales (4.67±0.52, n=45); (12) 4–6 infralabial scales (5.44±0.58, n=45); (13) 5–8 supralabial scales (6.40±0.65, n=45); (14) 1–3 internasal scales, (1.67±0.56, n=45); (15) males with 6–8 preanal pores (7.06±0.44, n=16), with enlarged but unpigmented preanofemoral scales; (16) tail without whorls but with 7–11 rings of slightly contrasting colour giving the impression of whorls; (17, 18) 4–6 dorsolateral tubercles usually present, each composed of 1–6 scales; (19, 20) beige-brownish dorsal colour without a clearly defined pattern; often a black stripe anterior and posterior to the eye; (21) ventrally distinctly yellow in life (fades in preservative); throat always with dark spots, sometimes forming longitudinal lines; often a lateral dark line made up by dark markings on the infralabial scales; (22) 143–200 dorsal scales along the body (175.67±24.95, n=6); (23) 41–87 dorsal scales around the body (74.17±17.39, n=6) ( Figs. 12a–b , 13 ).

Variation: Two specimens, MNHN 1990.3561 and MNHN 1930.267, show somewhat unusual meristic characters; they have three internasal scales, one of them with the third scale being very small (MNHN 1930.267), and the other furthermore with only three postpostmental scales (MNHN 1990.3561).

The specimen ZMA 19595 from Analalava forest shares morphological characters typical for L. pictus (male, 31.4 mm SVL, granular dorsal scales, first finger present and bearing a claw, three divided lamellae under fourth toe, mental scale semidivided, two asymmetrical postmental scales, four postpostmental scales, six infralabial and seven supralabial scales, two internasal scales, seven preanal pores, beige dorsal colouration without a distinct pattern). However, based on molecular data it was genetically strongly divergent from the other L. pictus included in the analysis ( Puente et al. 2005b). Values from this specimen have not been included in the morphometric and meristic values given above for L. pictus .

Material examined. MNHN 1899.342 – 343 (Nov. 1898, Ikongo forest, Vinanitelo ) , MNHN 1930.267 – 268 ( Ivohibe ) , MNHN 1950.261 (Millot, Sep. 1949, Andringitra, Fivahona forest ) , MNHN 1965.282 (collector and locality unknown, May 1951) , MNHN 1990.105 – 106 (20 Jan., Andringitra, Andohabatomanara, 106: Ampanasana river, small tributary to the Zamandao in the Vohidray Massif ) , MNHN 1990.1889 – 1891 (C.P. Blanc, Mar. 1967, between Ambositra and Manakara ) , MNHN 1990.3550 – 3564 (C.P. Blanc, winter 1970–1971, forested areas of Andringitra, Ambalamarovandana ) , ZFMK 58224 (K. Liebel, Antananarivo) , ZFMK 17712 (H. Meier, Jan. 1976, Fianarantsoa) , ZFMK 20724 (Forsyth Major, 1903, Fianarantsoa) , ZFMK 47245 (H. Meier, Nov. 1987, Tolongoina ) , ZFMK 54516 (G. Trautmann, locality unknown) , ZFMK 59843 (F. Glaw & M. Vences, 15–20 Jan. 1994, near Sendrisoa ) , ZMA 19531–19532 (M. Vences, P. Bora & F. Rabemananjara, 17 Jan. 2003, Ambositra ) , ZMA 19595 ( Analalava Forest , Isalo; see remark below) , ZMA 19597 and 19621 (limit of Ranomafana National Park , parcel 2, road to Fianarantsoa) , SMF 8956 (lectotype of Lygodactylus robustus, A. Voeltzkow, Ankarimbela , S- Madagascar) , ZSM 7 /2004–8/2004, 10/2004–13/2004, 15/2004 (F. Glaw, M. Puente, M. Thomas & R. Randrianiaina, 19–20 Jan. 2004, Antsirabe ) .

Distribution. According to the specimens examined in this study, Lygodactylus pictus is known from the following precise localities:

(1) Ambositra , (2) Andringitra (various collection sites: Ambalamarovandana , Andohabatomanara , Ampanasana river , Fivahona forest ), (3) Ankarimbela , (4) Antsirabe , (5) Fianarantsoa, (6) Ikongo forest ( Vinanitelo ), (7) Ivohibe , (8) limit of Ranomafana National Park at road to Fiaranantsoa , (9) near Sendrisoa , (10) Tolongoina. Most of these sites are located in the Southern Central East of Madagascar. Most are at higher elevations (above 1000 m), but some (e.g., Tolongoina ) are at lower elevations, and the identity of the specimens from these sites warrants a more detailed revision. The locality Antananarivo (indicated by specimen ZFMK 58224 ) requires confirmation and is not considered here, because it is distinctly more northern than the other localities known. The locality Analalava forest probably refers to a distinct, undescribed species (see remark below) .

Habitat: Specimens were observed active during the day, on the trunks of large eucalypt trees and on agava leaves near Sendrisoa (F. Glaw, D. R. Vieites and M. Vences, personal observation), and on tree trunks, walls, and fences in the towns of Ambositra and Antsirabe (M. Puente and F. Glaw personal observation). At Antsirabe, ca. 10– 12 eggs (not glued) were found deposited in the bark of a tree, probably representing a communal egg deposition site (F. Glaw, M. Puente, M. Thomas, R. Randrianiaina, personal observation, January 2004). At Ambositra, pairs of eggs (not glued) were found between old wood in a hotel garden (M. Vences, personal observation, 2003).

Lygodactylus tuberosus Mertens, 1965

( Fig. 14 )

Lygodactylus tuberosus Mertens, 1965 .— Name-bearing type: holotype, SMF 8949, female; not examined by us.— Type locality: “Tsimanampetso, SW-Madagaskar.” (= Tsimanampetsotsa), according to the original description.— Other types: paratypes SMF 8950–8952, three females, same data as holotype.— Etymology: no data from the original description.— Remark: see synonymy account of Lygodactylus tolampyae and Mertens (1965) for the nomenclatural history of this species which was in part formerly known as Lygodactylus tuberifer Boettger, 1913 .

Diagnosis. Lygodactylus tuberosus is a relatively small species of Lygodactylus that is common in the arid South of Madagascar. It can be assigned to the L. pictus group based on the characters listed in the group definition above. It differs from L. pictus by usually having only few and indistinct dark spots on the throat (vs. usually at least some distinct spots), and by a white or only faint yellowish ventral colour in life (vs. usually distinctly yellow; but the yellow pigment quickly fades in preservative), furthermore by the presence in most specimens of a dark marking behind the eye which includes spiny scales (vs. presence of only a dark stripe). Typically, specimens of L. tuberosus are relatively small, but some can attain large sizes (up to 38 mm). Clearly, the definition of this species requires revision. Distinction of L. tuberosus from L. roavolana is easily possible by the presence of only small black markings on the neck (vs. presence of large and very distinct markings above the forelimbs), and by the presence of a claw on the first finger (vs. apparent absence).

Description. (1, 2) A small to moderately sized Lygodactylus . Adult SVL 18.9–38.2 mm (28.69± 3.56 mm, n=225) (see remark below); (2) TAL 8.6–43.0 mm (29.9 ± 5.05 mm, n=94); (3) granular dorsal scales, (4, 5) first finger present and bearing a claw; (6) usually three pairs of subdigital lamellae on the fourth toe (3.01±0.14, n=228); (7, 8) mental scale semidivided into three parts by sutures, contact between first infralabial scale and posterior projection of mental scale can exist but is not distinct; (9, 10) usually two postmental scales (2.02±0.21, n=230), asymmetrical or symmetrical; (11) usually four or five postpostmental scales (4.66±0.63, n=230); (12) 4–8 infralabial scales (5.64±0.58, n=230); (13) 5–9 supralabial scales (6.70±0.76, n=230); (14) 0–2 internasal scales (1.30±0.47, n=230); (15) males with 5–9, usually seven, preanal pores (6.85±1.14, n=126), with enlarged but unpigmented preanofemoral scales; (16) tail without whorls but with rings of slightly different colour giving the impression of whorls; (17, 18) often with 3–7 dorsolateral tubercles each composed of 1–7 scales; (19, 20) dorsally beige-greyish without a clearly defined pattern, often with a dark stripe anterior and posterior to the eye, usually a dark marking with spiny scales appears behind the eye (near the ear opening); (21) ventrally usually uniform, not distinctly yellow, without or with few dark spots on the throat; some stripes can be present on the throat; (22) 143 dorsal scales along the body (n=1); (23) 80 dorsal scales around the body (n=1); (24) 87–89 ventral scales (n=2) ( Figs. 12c–d , 14 ).

Hemipenial structure. Based on ZSM 583/2000, adult male from Ifaty. Hemipenis subcylindrical, total length ca. 3.5 mm, with relatively large pedicel of ca. 1.8–2 mm. The truncus and the lobes are covered with fields and serrated ridges of pointed papillae. The apex is divided in two lobes of 1.5 mm length each. Sulcus spermaticus formed by two channels finishing in each apex arm, no distinct sulcal lips and no papillae around the sulcus ( Fig. 14 ).

Material examined. MNHN 1901.239 (Ambovombe) , MNHN 1901.147 (Andrahomana) , MNHN 1920.420 (Pere Appert [?], Morombe ) , MNHN 1924.69 ( Onilahy valley ) , MNHN 1929.70 (near Ianzamaly ) , MNHN 1929.73 ( Toliara ) , MNHN 1930.265 (Antanimoza near Tolagnaro ) , MNHN 1930.266 (Andranomana) , MNHN 1935.123 – 124 (Amboasary, Mandrare river ) , MNHN 1938.193 – 196 (S Madagascar) , MNHN 1990.401 (8 Nov. 1961, Vohitomotsy , 80 km SE Toliara) , MNHN 1990.402 (11 Nov. 1961, Vohitomotsy, "sur tronc d'un Kily ") , MNHN 1990.403 – 406 (Nov. 1961, Ankazomanga, Plateau Mahafaly ) , MNHN 1990.407 (23 Nov. 1961, Ankazomanga, Plateau Mahafaly , "tronc d'arbre pourri") , MNHN 1990.408 (27 Nov., Andreboka, 35 km SSW Betioky ) , MNHN 1990.409 (23 Nov., Egogy, 17.5 km SW Ampanihy ) , MNHN 1990.410 (Jun. 1961, Aniriky , 3 km W Mahavelo) , MNHN 1990.411 (spiny forest between Marolinta and Ampokata ) , MNHN 1990.413 (19 Nov., Kaz (Mahafaly), "sur tronc Kily ") , MNHN 1990.414 (22 Nov., Kaz (Mahafaly, "sur tronc kily") , MNHN 1990.412 , 415 (3 Jul., Zampongotra ) , MNHN 1990.416 (15 Jul., Zampongotra ) , MNHN 1990.417 (Saraondry, "sur tronc de kily") , MNHN 1990.418 ( Sakaraha forest , on dead wood) , MNHN 1990.419 (C.P. Blanc, 2 Jan. 1964, Toliara, on dead tree) , MNHN 1990.421 (C.P. Blanc, 22 Dec. 1966, Toliara, on dead tree) , MNHN 1990.422 (3 Jul., around Zampongotra, 30 km W Beloha ) , MNHN 1990.423 – 425 (15 Jul. 1966?, Zampongotra ) , MNHN 1990.426 – 430 (C.A. Domergue, 25 Aug. 1971, Ihotry ) , MNHN 1990.431 – 433 (C.A. Domergue, Jan. 1966, Ampanihy ) , MNHN 1990.434 (C.A. Domergue, 12. Nov. 1965, Ihotry ) , MNHN 1990.438 (C.A. Domergue, 1–2. Nov. 1967, Ihotry ) , MNHN 1990.439 – 440 (C.A. Domergue, 27–28 Apr. 1969, Ihotry ) , MNHN 1990.441 (C.A. Domergue, 11 Jul. 1968, Ihotry ) , MNHN 1990.462 – 488 (10–11 Dec. 1972, Ihotry ) , MNHN 1990.501 – 511 (18 Mar. 1973, Ihotry ) , MNHN 1990.603 – 613 (6 Dec. 1975, Ihotry ) , MNHN 1990.624 – 645 (15 Jan. 1966, Ampanihy ) , MNHN 1990.646 – 659 (15 Jan. 1966, Ampanihy ) , MNHN 1990.669 – 672 (5–6 Apr. 1967, Belalanda ) , MNHN 1990.1860 – 1866 (C.A. Domergue, Ihotry ) , MNHN 1990.1840 – 1841 (C.A. Domergue, Toliara harbour) , MNHN 1990.1842 – 1843 (C.A. Domergue, Ihotry ) , MNHN 1990.1844 – 1846 (C.A. Domergue, 28 Mar. 1968, Ampanihy ) , MNHN 1990.3233 – 3240 (C.A. Domergue, 15 Jan. 1966, Ampanihy ) , MNHN 1990.3250 – 3263 (C.A. Domergue, 2 Mar. 1967, Belalanda ) , MNHN 1990.3264 – 3270 (C.A. Domergue, 5–6 Apr. 1967, Belalanda ) , MNHN 1994.1737 (Lamberton, southern Madagascar) , ZFMK 21814 (leg. H. Meier, Jan. 1978, Toliara) , ZFMK 59803 (M. Vences & F. Glaw, 23–27 Jan. 1994, Toliara) , ZFMK 70005 (N. Lutzmann, Toliara) , ZMA 19608 (M. Puente, 2003, Ifaty ) , ZMA 19600 (M. Puente, 2003, Arboretum Toliara) , ZSM 583 /2000 (F. Glaw, K. Schmidt, L. Rakotozafy & R. Razafindrasoa, 26 Mar. 2000, Ifaty ) , ZSM 584 /2000 (F. Glaw, K. Schmidt, L. Rakotozafy & R. Razafindrasoa, 1 Apr. 2000, Toliara) , ZSM 591 /2000 (F. Glaw, K. Schmidt, L. Rakotozafy & R. Razafindrasoa, 27 Mar. 2000, Ifaty ) , ZSM 947 /2003 (M. Puente & M. Thomas, 4 Feb. 2003, Arboretum Toliara) .

Distribution. According to the specimens examined in this study, Lygodactylus tuberosus is known from the following localities: (1) Amboasary, (2) Ambovombe, (3) Ampanihy, (4) Andrahomana, (5) Aniriky, (6) Ankazomanga, (7) Antanimoza, (8) Belalanda, (9) near Ianzamaly, (10) Ifaty, (11) Ihotry, (12) Kaz (Mahafaly), (13) Morombe, (14) Onilahy valley, (15) Saraondry, (16) Sakaraha forest, (17) Toliara, (18) Arboretum Toliara, (19) Tsimanampetsotsa (type locality), (20) Vohitomotsy, (21) Zampongotra. Furthermore, Glaw & Vences (1994, 2007) provide a record for the species from (22) Berenty. We are uncertain whether the locality information “Kaz (Mahafaly)” as given for MNHN 1990.413 and 1990.414 refers to an actual locality (as interpreted here), or “Kaz” may in fact be an abbreviation.

Habitat. L. tuberosus is a species inhabiting arid zones in the South. All localities are in this area. Specimens in the MNHN collection have been found on "Kily" trunks which refers to Tamarindus indicus trees. We observed the species active during the day, in open forests, on isolated trees or fallen tree trunks, and in dry bush areas (M. Puente & M. Thomas, personal observation, February 2003).

Remarks. Two specimens with SVL values of 12.7 and 15.3 mm have distinct preanal pores and thus could be classified as males. However, we consider these as too small to be sexually mature and thus have not included them in the calculation of average, minimum and maximum SVL values.

Four specimens with locality information outside the known range of L. tuberosus require further discussion:

The three specimens MNHN 1990.435 – 436 (10 Jan. 1970) and MNHN 1990.437 (20 Dec. 1966), two females and one male, from “Col des Tapias” have been catalogued as L. tuberosus and apparently belong to this species, but their locality is atypical for this species. Two other species, L. arnoulti and L. blanci , are known from the Col des Tapias / Mt. Ibity region, but the three specimens can be distinguished from these two species as follows: SVL 20.6 –31.0 mm ( L. blanci and L. arnoulti are bigger, the smallest specimen of each measuring> 21.4 mm); granular dorsal scales (partially keeled in L. blanci ); two symmetrical postmental scales ( L. blanci and L. arnoulti have three postmental scales); the male with seven preanal pores ( L. blanci and L. arnoulti have 9–11 preanal pores); intact tail with 12 faint coloured bands reminding whorls; dorsal colour beige-brown without a clear pattern; ventral colour light with spots. “ Tapia ” refers to a particular tree typical for some of the dryer areas of Madagascar's central highlands ( Uapaca bojeri ), and Col des Tapias usually refers to a locality along the National Road 7, between Antsirabe and Ambositra and next to Ibity mountain. However, according to a personal communication of C. P. Blanc in 2009, there is another site with this name in the South West between Ranohira and Sakaraha, and he has collected Lygodactylus from this site which may correspond to the MNHN specimens in question .

MNHN 1990.664 (Ambohitantely, "bambou au sol", collected by Seguier): This single specimen is catalogued as L. tuberosus , but the locality Ambohitantely is located in central Madagascar whereas L. tuberosus appears to be restricted to the South of Madagascar. The specimen is a female; 29.2 mm SVL; TAL (regenerated) 25.5 mm; dorsal scales granular; first finger present and with claw; five pairs of lamellae under fourth toe ( L. tuberosus with three pairs); mental scale semi-divided; no contact between posterior projection of mental and infralabial scales; two asymmetrical postmental scales; four postpostmental scales; six infralabial and seven supralabial scales; two internasal scales; dorsolateral tubercles absent; dorsal colour brown-beige, dorsal pattern tending to be striated ( L. tuberosus without a clear dorsal pattern, and ocular region striated with a dark spot); ventrally light with spots.

Figures

FIGURE 12. Photos of living specimens assigned to species in the Lygodactylus pictus group. (a) L. tuberosus, Berenty; (b) L. tuberosus, Ifaty; (c) and (d), L. pictus, near Sendrisoa, Andringitra; (e) and (f), dorsolateral and ventral view of L. roavolana, holotype specimen, from Manantantely. All photographs by F. Glaw & M. Vences.

FIGURE 13. Scalation characters of Lygodactylus pictus. (a) head in lateral view (scale bar = 1 mm); (b) lamellae of fourth toe in ventral view (scale bar = 1 mm); (c) head dorsal view (scale bar = 2 mm); (d) throat in ventral view (scale bar = 2 mm); all based on male specimen MNHN 1990.1859; (e) preanal pores in ventral view (scale bar = 2 mm), based on male specimen MNHN 1990.3550. All drawings by M. Puente.

FIGURE 14. Scalation and hemipenial characters of Lygodactylus tuberosus. (a) head in lateral view (scale bar = 1 mm), based on male specimen MNHN 1990.535; (b) head in dorsal view (scale bar = 2 mm), based on male specimen MNHN 1990.605; (c) throat in ventral view (scale bar = 2 mm), based on male specimen MNHN 1990.541; (d) preanal pores in ventral view (scale bar = 2 mm), based on male specimen MNHN 1990.524; (e) lamellae of fourth toe in ventral view (scale bar = 2 mm), based on male specimen MNHN 1990.535; (f) and (g), sulcal and asulcal view of the hemipenis based on male specimen ZSM 583/2000 (scale bar = 1 mm). All drawings by M. Puente.