Madascincus pyrurus, Miralles, Aurelien, Koehler, Joern, Glaw, Frank & Vences, Miguel, 2016

Miralles, Aurelien, Koehler, Joern, Glaw, Frank & Vences, Miguel, 2016, Species delimitation methods put into taxonomic practice: two new Madascincus species formerly allocated to historical species names (Squamata, Scincidae), Zoosystematics and Evolution 92 (2), pp. 257-275 : 264-267

Madascincus pyrurus sp. n. Figs 3A, 4D, E

Holotype.

ZSM 520/2001 (MV 2001-445), adult male from Mont Ibity, approximately at 20°14'S, 47°03'E, 1700-1800 m above sea level, Fianarantsoa province, central Madagascar, collected on 10 March 2001 by M. Vences, D.R. Vieites, L. Raharivololoniaina and D. Rakotomalala.

Paratypes

(n=7). MNHN 1980-1217, Mont Ibity, Fianarantsoa province, central Madagascar, coll. by Y. Thérézien and R. Capuron; ZSM 518-519/2001 (MV 2001-441, 2001-444), UADBA uncatalogued (MV2001.442 and 443), same data as holotype; ZSM 521/2001 (MV 2001-611), UADBA uncatalogued (MV2001.610), Itremo (camp, 20°36 ’08’’ S, 46°34 ’16’’ E, 1648 m a.s.l.), Fianarantsoa province, coll. on 10 March 2001 by M. Vences, D.R. Vieites, L. Raharivololoniaina and D. Rakotomalala.

Chresonyms.

Scelotes igneocaudatus - Blanc (1967), Brygoo (1981, partim);

Amphiglossus igneocaudatus - Brygoo (1984, partim), Whiting et al. (2004), Schmitz et al. (2005);

Madascincus igneocaudatus - Glaw and Vences (2007, partim), Crottini et al. (2009; partim), Miralles et al. (2011a, c, partim; 2011b);

Madascincus igneocaudatus " igneocaudatus -C clade" - Miralles and Vences (2013);

Madascincus sp. " igneocaudatus " central clade - Miralles et al. (2015).

Diagnosis.

A member of the genus Madascincus based on its molecular phylogenetic relationships (see Fig. 1). Within the genus Madascincus , Madascincus pyrurus is distinguished from all its congeners by the following combination of characters: medium body size with a maximum snout-vent length (SVL) of 54.2 mm (versus, in smaller species, a maximum SVL of 33.6 mm in Madascincus nanus complex); 71-79 rows of paravertebral scales (versus 51-62 in Madascincus melanopleura , 57-65 in Madascincus minutus , 52-62 in Madascincus ankodabensis , 60-65 in Madascincus mouroundavae , and 50-57 in Madascincus nanus complex); 73-78 rows of ventral scales (versus 55-63 in Madascincus minutus , 56-61 in Madascincus melanopleura , 52-60 in Madascincus nanus complex, 59-63 in Madascincus ankodabensis , 65-73 in Madascincus miafina , 63-66 in Madascincus mouroundavae ); 15-18 subdigital lamellae under the fourth toes (versus 5-8 in Madascincus nanus complex, 9-13 in Madascincus minutus , 18-23 in Madascincus miafina and 12-15 in Madascincus ankodabensis ); 22-24 rows of scales around midbody (versus 18-20 in Madascincus nanus complex, 28-30 in Madascincus mouroundavae , 24-26 in Madascincus igneocaudatus and 30-32 in Madascincus stumpffi ); pentadactyl forelimbs (versus 3-5 digits in Madascincus nanus complex; the presence of postnasal scales (always absent in Madascincus arenicola ); the frontal is bell-shaped (versus hourglass shaped in Madascincus nanus , Madascincus minutus , Madascincus melanopleura , Madascincus ankodabensis , Madascincus mouroundavae , and in half (52.8%) of the specimen examined of Madascincus stumpffi ); the frontal is always separated from the interparietal (versus most often (87.5%) fused in Madascincus mouroundavae ); lower eyelid window is spectacled (versus scaly in Madascincus arenicola , Madascincus nanus , Madascincus mouroundavae , Madascincus polleni , Madascincus miafina and Madascincus stumpffi ), the presence of two (21.4%) or three (78.6%) rows of enlarged nuchal scales (versus absence or presence of a single row in Madascincus nanus , Madascincus mouroundavae , Madascincus miafina and Madascincus stumpffi ). More specifically, Madascincus pyrurus differs from its sister species Madascincus igneocaudatus in having a shorter and rounder snout (versus a relatively long and pointed snout usually characterizing semi-fossorial species found in sandy habitat) and in being oviparous (versus viviparous). More generally, Madascincus pyrurus can also be easily distinguished from all the other members of the genus Madascincus by its very characteristic pattern of coloration, being the only species with six well-defined very dark stripes (a pair of thin dorsal, a pair of wide upper lateral and a pair of thin lower lateral stripes) running along the body, and one of the few species (together with Madascincus miafina and Madascincus igneocaudatus ) in which the tail might be bright red in some specimens (see also Tables 1 and 2).

Description of the holotype.

ZSM 520/2001 (MV 2001-445, Fig. 3A). Adult male, with both hemipenes everted. Good state of preservation, with exception of a little incision on the right flank (ca. 10 mm). SVL (49.0 mm) 7.1 times head length (6.9 mm), shorter than the tail (58.0 mm, apparently regenerated). Limbs relatively short: SVL 6.2-6.5 times front limb length (7.5-7.9 mm) and 3.4 times hind limb length (14.4-15.0 mm). Snout relatively short and rounded on lateral aspect, with a rostral tip bluntly rounded in dorsal aspect. Rostral wider than high/long, contacting first supralabials, nasals, and supranasals. Paired supranasals in median contact, contacting loreals. Frontonasal roughly pentagonal, wider than long, contacting loreals, and first suproculars. Prefrontals absent. Frontal longer than wide, wider posteriorly, in contact with frontonasal, supraoculars, parietals and interparietal. Supraoculars four, all of them in contact with frontal; the second anteriormost pair larger and the posteriormost pair significantly smaller; the first pair not constricting frontal (bell-shaped). Frontoparietals absent. Interparietal longer than wide, well separated from supraoculars; parietal eyespot present but poorly marked. Parietals contact posterior to interparietal. Two pairs of primary nuchal scales (with two cycloids scales inserted between the parietal and the first nuchal, on the right side). Nasals slightly larger than nostrils; contacting rostral, first supralabials, postnasals and supranasals. Postnasals present, separating supranasals from first supralabials, and nasals from loreals. Loreal single, as high as long. Preocular higher than long, single. Presubocular single. Six supraciliaries on both sides, in continuous row; last pair slightly larger than the others, and projecting onto supraocular shelf. Pretemporals two, both contacted by parietal. Postsuboculars two; upper contacting lower pretemporal; both contacting penultimate supralabial. Lower eyelid moveable, with a transparent and undivided disc, and one row of small scales across its dorsal edge. Contact between upper palpebrals and supraciliaries seemingly direct but flexible, i.e. palpebral cleft narrow. Primary temporal single. Secondary temporals two; the upper one elongated, contacting lower pretemporal anteriorly and overlapping lower secondary temporal ventrally. Two tertiary temporals bordering lower secondary temporal. Supralabials six, with the fourth being the enlarged subocular contacting scales of the lower eyelid. Postsupralabial single. External ear opening roughly triangular, without lobules. Mental wider than long, posterior margin straight. Postmental wider than long, contacting first two pairs of infralabials. Infralabials six. Three pairs of large chin scales, both members of first pair in contact, both members of second pair separated by a single median scale, and members of third pair separated by three scale rows. No scales extending between infralabials and large chin scales; two asymmetrical postgenials posterolaterally in contact with the third pair of chin scales. Gulars similar in size and outline to ventrals. All scales, except head shields and scales on palms, soles, and digits, cycloid, smooth, and imbricate; longitudinal scale rows at midbody 22; paravertebrals 71, similar in size to adjacent scales; ventrals 73. Inner preanals overlapped by outer. Both pairs of limbs pentadactyl; fingers and toes relatively short, clawed; relative length of toes in the following order: I<II=V<III<IV. Subdigital lamellae smooth, single, 10 under fourth finger of both left and right manus, 17 under right fourth toe, and 15 under left fourth toe.

Coloration in preservative with upper side of the head, neck, back, limbs, and tail dark bronze. Venter, lower side of head, throat, lower side of limbs, and tail withish/cream. Lateral borders on the ventral side maculated by very small dark dots. Six very well defined very dark stripes run along the body, continuing along the first third of the tail, then abruptly ending where the tail is regenerated. Two thin blackish dorsal stripes formed by succession of contiguous dots start on the supranasal; at midbody, each dorsolateral stipe is less than one scale wide and both are separated by two rows of dorsal scales. Two wide dark brown upper lateral stripes; margins slightly darker and very sharp; about two scales wide at midbody and overlying three rows of scales; starting from the rostral, where the stripes all meet, extending on the upper half of each supralabial, the loreals, around the eyes, above ear opening, and above forelimbs and hindlimbs. Two thin dark lower lateral stripes, starting on the last infralabials, extending through the forelimb and hindlimb insertion. At midbody, each lower lateral stripe is less than one scale wide, with irregular margin. Four very light stripes run along the body, continuing along the first third of tail, then abruptly ending where the tail is regenerated. Two whitish dorsolateral stripes separate dark dorsal stripes from the upper lateral dark stripes; about one scale wide at midbody and overlying two rows of scales. Two whitish lateral stripes separating the dark upper lateral stripes from the dark lower lateral stripes; about one scale wide at midbody and overlying two rows of scales. Regenerated part of the tail cream, maculate with many small dark dots on the dorsal side. Palms and soles darker than the ventral side.

Life coloration for the holotype has not been documented, but it is apparently very similar to the coloration in preservative, with exception of the tail which is usually bright red, or pinkish brown in some specimens (cf. Fig. 4D, E).

Variation.

For variation in measurements and scale characters see Table 1. Some variation is evident with respect to tail coloration which may be bronze or reddish (cf. Fig. 4D, E) and ventral coloration which may be maculated by small black dots or uniformly whitish ( Blanc and Blanc 1967).

Etymology.

The specific epithet pyrurus is based on Greek roots pûr (πῦρ) and ourá (οὐρά), respectively meaning “fire” and “tail”. This word is here treated as invariable noun and has the same meaning as an other specific epithet in the genus, igneocaudatus , which is based on Latin roots. This epithet has been chosen to highlight the morphological similarity of Madascincus igneocaudatus and Madascincus pyrurus , both these sister species being characterized by a tail which may be red and reminding fire.

Distribution, habitat and habits.

The species is known from the dry environments on two massifs in the central highlands of Madagascar, in Mont Ibity and in Itremo (Fig. 5), dominated by rock outcrops and tapia woodlands (loose forests of Uacapa bojeri trees). This montane lizard has been found at altitudes ranging from 1648 to 1922 m a.s.l. in Itremo, and up to 2252 m a.s.l. in Ibity ( Brygoo 1984, present study). It can be commonly found under stones on usually somewhat sandy substrate (quartz sand especially on Mont Ibity), in open areas. On Mont Ibity, 6 eggs of Madascincus pyrurus have been collected at the end of December which measured 18x12 mm, the hatched juveniles measuring 25 mm SVL ( Blanc and Blanc 1967, Brygoo 1984). In contrast, in Madascincus igneocaudatus , developed juveniles have been found in a female dissected from Ifaty, indicating that the sister species of Madascincus pyrurus , endemic to the dry lowlands of the western coast, is viviparous ( Glaw and Vences 2007). Interestingly, the different modes of reproduction observed in these two sister species appear to be in contradiction with the trend usually observed in lizard taxa having both oviparous and viviparous species/populations: typically, evolution to viviparity in these groups is interpreted as local adaptation to cold climates, either at high altitude or latitude ( Pianka and Vitt 2003).