Paroedura hordiesi, Glaw, Frank, Roesler, Herbert, Ineich, Ivan, Gehring, Philip-Sebastian, Koehler, Joern & Miguel Vences,, 2014

Glaw, Frank, Roesler, Herbert, Ineich, Ivan, Gehring, Philip-Sebastian, Koehler, Joern & Miguel Vences,, 2014, A new species of nocturnal gecko (Paroedura) from karstic limestone in northern Madagascar, Zoosystematics and Evolution 90 (2), pp. 249-259 : 249-255

Paroedura hordiesi sp. n. Figs 2, 3, 4, Table 1

Remarks.

This species has been treated or figured under the name Paroedura homalorhinus ( Henkel and Schmidt 1995), Paroedura karstophila ( Glaw et al. 2001), Paroedura sp. "Montagne des Français” ( Glaw and Vences 2007), Paroedura sp. ( D’Cruze et al. 2007), Paroedura aff. karstophila ( Schönecker 2008), Paroedura cf. karstophila ( Megson et al. 2009), Paroedura sp. n. / sp. n. 1 ( Jackman et al. 2008 and Nagy et al. 2012). Its karyotype (2n = 36) has been described under the names Paroedura sp. n. I 'Montagne des Français’ ( Aprea et al. 2013) and Paroedura karstophila ( Koubová et al. 2014). DNA-sequences of this species have been published in Jackman et al. (2008), Aprea et al. (2013), and Koubová et al. (2014).

Holotype.

ZSM 342/2004 (field number FGZC 639), adult male with (broken) original tail and everted hemipenes, collected at Montagne des Français (12°19 ‘34“ S, 49°20 ‘09“ E, 334 m above sea level), Antsiranana Province, north Madagascar, on 18 February 2004 by F. Glaw, M. Puente and R. Randrianiaina. GenBank accession numbers for sequences of the holotype ( Jackman et al. 2008): EF536213 (ND2), EF536239 (ND4), EF536165 (RAG1) and EF536189 (PDC).

Paratypes.

All paratypes were collected in the “tsingy” limestone massif at Montagne des Français, Antsiranana province, north Madagascar. Specimens were collected on the tsingy outcrops along the way between the Hotel "Kings Lodge" (12°18 ’44,8’’ S, 49°20 ’22,6’’ E, 10 m a.s.l.) and the remains of the French Fort (12°19 ’34’’ S, 49°20 ’09’’ E, 334 m), except where other locality information and coordinates are given in the following: UADBA uncatalogued (FG/MV 2000-317), sex unknown, and ZSM 531/2000 (FG/MV 2000-316), adult male with everted hemipenes, both collected on 14 March 2000 by F. Glaw, K. Glaw and M. Vences; ZSM 532/2000 (no field number), adult female, collected on 21 March 2000 by F. Glaw and K. Glaw; ZSM 1108/2003 (no field number), adult female, collected on ca. 20 February 2003 by F. Glaw and R. D. Randrianiaina; ZSM 337/2004 (FGZC 634), subadult, and ZSM 338/2004 (FGZC 635), subadult, both collected at 12°19 ‘34“ S, 49°20 ‘09“ E, 334 m a.s.l., on 23 February 2004 by F. Glaw, M. Puente and R. D. Randrianiaina; ZSM 339/2004 (FGZC 636), adult female, ZSM 340/2004 (FGZC 637), adult female, ZSM 341/2004 (FGZC 638), adult female [original tail broken], all three with same data as holotype; ZSM 343/2004 (FGZC 640), adult male without tail, and ZSM 350/2004 (FGZC 647), adult female, both without reliable locality and collection data, but most likely with same data as holotype; UADBA uncatalogued (FGZC 612), sex unknown, collected at 12°19 ‘34“ S, 49°20 ‘09“ E, 334 m a.s.l., on 20-28 February 2004 by F. Glaw, M. Puente and R. D. Randrianiaina; ZSM 352/2004 (FGZC 649), adult, and ZSM 353/2004 (FGZC 650), adult female, both collected at 12°19 ‘34“ S, 49°20 ‘09“ E, 334 m a.s.l., on 18-23 February 2004 by F. Glaw, M. Puente and R. D. Randrianiaina; UADBA-R 70183 (FGZC 1109), adult male with everted hemipenes, UADBA-R 70185 (FGZC 1112), male, UADBA-R 70184 (FGZC 1114), adult female, ZSM 2106/2007 (FGZC 1099), juvenile, ZSM 2107/2007 (FGZC 1100), juvenile, ZSM 2113/2007 (FGZC 1115), adult female, all six collected around the remains of the French Fort (12°19 ’33’’ S, 49°20 ’17’’ E), collected on 27 February 2007 by P. Bora, H. Enting, F. Glaw, A. Knoll and J. Köhler; UADBA-R 70281 (FGZC 1659), sex unknown, ZSM 1530/2008 (FGZC 1660), adult female, both collected in the cave between Andavakoera and remains of French Fort, on 16 February 2008 by M. Franzen, F. Glaw, J. Köhler and Z. T. Nagy.

Diagnosis.

Paroedura hordiesi sp. n. is a medium-sized species (SVL up to 58 mm, tail length up to 53 mm), having moderately prominent dorsal tubercles disposed into moderately distinct, and generally regular longitudinal rows and an original tail with no spines.

The new species can be easily attributed to the genus Paroedura based on its nested phylogenetic position within the genus ( Jackman et al. 2008) and its morphological similarity to other Paroedura species, especially concerning the ventral structure of their fingers and toes which comprise a pair of squarish terminal adhesive pads. Among Malagasy geckos this terminal toe structure is only found in Paroedura and the related genus Ebenavia (Glaw & Vences, 2007). The latter genus can be easily distinguished from Paroedura by its much narrower and strongly pointed head, its elongated body, and smaller size.

Comparisons.

The new species can be distinguished from the 17 other currently recognized Paroedura species (including the three available junior synonyms in the genus) as follows: From Paroedura androyensis , Paroedura bastardi , Paroedura ibityensis , Paroedura lohatsara , Paroedura maingoka , Paroedura picta , and Paroedura vahiny by having the nostril in contact with the rostral scale; from Paroedura gracilis by absence of a raised vertebral ridge on the body and shorter forelimbs which are not extending forward beyond tip of snout; from Paroedura masobe by much smaller size (SVL up to 58 mm versus 107 mm), much smaller eyes with a pigmented iris (versus black iris) and absence of a dorsal row of paired spines on the tail; from the two Comoroan species Paroedura sanctijohannis and Paroedura stellata by slightly smaller size (SVL up to 58 mm versus 68 mm and 62 mm, respectively) and absence of whorls with distinct spiny tubercles of the original tail; from the syntopically distributed Paroedura stumpffi by smaller size (SVL up to 58 mm versus 70 mm) and absence of whorls with distinct spiny tubercles of the original tail; from Paroedura tanjaka by much smaller size (SVL up to 58 mm versus 102 mm) and absence of whorls with distinct spiny tubercles of the original tail; from Paroedura vazimba by larger size (SVL up to 58 mm versus 49 mm) and absence of whorls with distinct spines of the original tail; from Paroedura oviceps from its type locality Nosy Be by smaller size (SVL up to 58 mm versus 69 mm) and rather regularly arranged tubercle rows on the back (versus rather irregular rows of tubercles); from Paroedura karstophila by the absence of whorls with distinct spiny tubercles of the original tail (and by a smoother regenerated tail, see Nussbaum and Raxworthy 2000) and by colouration (see Fig. 2 versus Fig. 5); and from its close relative Paroedura homalorhina ( Jackman et al. 2008) by shorter limbs (finger tips reach the anterior margin of eye versus snout tip when forelimbs are adpressed along the body), slightly smaller size (SVL up to 58 mm versus 65 mm, see Table 2), distinct and generally regularly arranged tubercles rows on the back (versus less distinct and less regular rows), and a more slender habitus (see Fig. 2 versus Fig. 5). In addition, Paroedura hordiesi can be easily distinguished from most other Paroedura species ( Paroedura androyensis , Paroedura bastardi , Paroedura gracilis , Paroedura ibityensis , Paroedura lohatsara , Paroedura maingoka , Paroedura masobe , Paroedura oviceps , Paroedura picta , Paroedura sanctijohannis , Paroedura stellata , Paroedura stumpffi , Paroedura tanjaka , Paroedura vahiny and Paroedura vazimba ) by adult colouration in life (see colour photographs in Glaw and Vences 2007, Schönecker 2008, Hawlitschek and Glaw 2013) and from Paroedura androyensis , Paroedura bastardi , Paroedura gracilis , Paroedura homalorhina , Paroedura ibityensis , Paroedura lohatsara , Paroedura maingoka , Paroedura masobe , Paroedura picta , Paroedura sanctijohannis , Paroedura stellata , Paroedura stumpffi , Paroedura tanjaka , and Paroedura vazimba by juvenile colouration ( Glaw and Vences 2007, Schönecker 2008, Hawlitschek and Glaw 2013, FG pers. obs.; juvenile colouration of the other species still unknown). Genetically, Paroedura hordiesi can be distinguished from all other species in the genus by its molecular differentiation in mitochondrial and nuclear genes ( Jackman et al. 2008, Nagy et al. 2012, Hawlitschek and Glaw 2013, Fig. 1) except for Paroedura vahiny for which DNA sequences are not yet available.

Description of the holotype.

SVL 52.6 mm, further measurements and counts are given in Table 1. Holotype in good condition, with complete but broken original tail and everted hemipenes. Head distinctly wider than neck and wider than body. Snout angled downward to tip, slight depression between poorly developed canthal ridges. Ear opening is a vertical slit. Original tail slightly shorter than snout-vent length, nearly round in cross section in its proximal part, laterally compressed in its distal half, with sharply pointed tip; ventral pygal section with pair of postcloacal sacs. Digits moderately expanded at tips. Rostral scale rectangular, much wider than tall and wider than mental. Nostril in contact with rostral, first supralabial, and five further scales. First supralabial largest, labials smooth. Snout and interorbital scales juxtaposed, some raised, some scales in front of orbits tuberculate, as are some larger lateral occipital scales. Dorsolateral neck and body scales very heterogeneous with about eight partly poorly recognizable longitudinal rows at midbody of enlarged tubercles; enlarged tubercles separated partly by small flat scales and smaller tubercles. Dorsal scales of forelimbs mostly flat. Dorsal scales of hindlimbs large and tuberculate, much smaller above knee. Ventral scales of forelimbs slightly smaller than surrounding ventral scales of the body. Dorsal pygal scales like dorsal body scales; lateroventral pygals tuberculate. Tail scales flat, tail segments without any transverse row of spiny tubercles. Mental triangular, bordered posteriorly by a pair of elongate, irregular hexagonal postmentals. Postmentals contact mental, first and second infralabial, one enlarged lateral gular, one smaller posterolateral gular, and one larger central gular. First three infralabials not significantly larger than others. Gulars small, granular. Ventrals of chest and abdomen flat. Proximal subdigitals in rows of 2-3. Pair of squarish, terminal pads. Claws curving downwards between terminal pads of digits.

Colour after 10 years in alcohol (Fig. 3): head dorsally beige with almost no recognizable pattern except a whitish dorsolateral spot above each ear opening and a well-defined blackish area above each eye which represents, however, no pigmentation of the skin, but is due to the blackish eyeball shining through the skin. Dorsum greyish with four poorly defined beige spots on the back which are the remains of dorsal crossbands: one distinct spot between the forelimbs, two poorly recognizable spots on the back between forelimbs and hindlimbs, and a fourth poorly recognizable spot between the hindlimbs. Dorsal surfaces of forelimbs and hindlimbs marbled with beige and grey. Flanks with similar colour as dorsum, but without any recognizable traces of light crossbands. Tail dorsally with whitish-grey and brown alternating transverse bands which are poorly delimited in the distal portion of the tail. Throat, chest, venter, ventral parts of forelimbs and hindlimbs and ventral side of tail whitish. Iris dark grey, pupil white. Colour of holotype in life unknown (no colour photographs available).

Variation.

Morphometric and meristic variation of ten specimens of the type series is summarized in Table 1, but there is also a remarkable individual variation in dorsal colouration and pattern (Fig. 2). Paratype ZSM 2106/2007, the smallest known juvenile (SVL 28.1 mm, tail length 25.2 mm), shows a moderately distinct juvenile colouration both in life (Fig. 2) and in preservative consisting of four light crossbands on the back between the insertion of forelimbs and the insertion of hindlimbs. A similarly distinct banding on body and original tail is still visible in the larger juvenile paratype ZSM 2107/2007 (Fig. 2, SVL 35.2 mm, tail length 34.7 mm). Although the juvenile colouration is distinct compared to most adults, it is less colourful and less contrasting in comparison with many other Paroedura species. The adult paratypes usually have a less distinct dorsal colour pattern than the juveniles, ranging from poorly recognizable (e. g. ZSM 532/2000) to distinct and well delimited (e. g. ZSM 531/2000) after 14 years in alcohol. Additional photographs of living individuals are provided in Henkel and Schmidt (1995), Glaw and Vences (2007), and Schönecker (2008). Most of the adult type specimens have a regenerated tail (without distinct transversal banding of dark and white), indicating a high pressure by predation or intraspecific aggression.

Hemipenis.

The following description is based on the everted right organ of ZSM 343/2004 (Fig. 4). Hemipenis medium-sized (total length 6.4 mm, width at apex 4.5 mm), apex divided in two lobes. Sulcus spermaticus forms an S-shaped, narrow and deep groove, bordered by moderately distinct lips reaching the apex. On the apex, the sulcus is becoming broader and divides in two branches. A horn-like apical cone is present in the center of each lobe. Calyces on the apex are only present at the border of the lobes. The hemipenes of the holotype (ZSM 342/2004) are virtually identical in every respect.

Habitat and habits.

Paroedura hordiesi was observed multiple times at night in karstic dry forest in the rainy season, mainly climbing on karstic rocks and the ruins of an old fort. It was found in close syntopy with Paroedura lohatsara in the karstic limestone areas, whereas Paroedura stumpffi was only encountered on the slope between the massif and the sea, in areas without karstic formations.

Etymology.

The specific name is dedicated to Freddy Hordies, in recognition of his support for biodiversity research and conservation through the BIOPAT initiative.

Distribution and conservation status.

Paroedura hordiesi is reliably known only from the recently established nature reserve of Montagne des Français. The species possibly also occurs at Ampombofofo, ca. 25 km north of this massif ( Megson et al. 2009: ZSM 1531/2008), but the identity of this population needs further study as molecular data are not available thus far. Surveys in other limestone areas of northern Madagascar revealed a superficially similar Paroedura at Nosy Hara ( Metcalf et al. 2007) which is, however, strongly differentiated by colouration (see photo in Glaw and Vences 2007) and mitochondrial DNA sequences (Fig. 1 and Nagy et al. 2012), and is therefore considered a further undescribed candidate species. The Ankarana massif is known to harbour Paroedura homalorhina , Paroedura karstophila , and Paroedura stumpffi ( Nussbaum and Raxworthy 2000, Raselimanana 2008). Our own surveys in Ankarana revealed only Paroedura homalorhina and Paroedura stumpffi and we consider the occurrence of Paroedura hordiesi unlikely in this massif. Rakotondravony (2006a) found Paroedura sp. at Binara in the Loky-Manambato region which potentially could refer to Paroedura hordiesi , but the identity of this record remains to be studied. No unidentified Paroedura species was found at Analamera ( Rakotondravony 2006b). Thus current evidence suggests that Paroedura hordiesi is microendemic of Montagne des Français and perhaps the adjacent Ampombofofo region.

For consistency with the IUCN Red List Assessment for Paroedura lohatsara ( Raxworthy et al. 2011) and other potential microendemic species of the Montagne des Français region, we suggest a classification as "Critically Endangered" on the basis that Paroedura hordiesi has an extent of occurrence of at most 50 km², it is known from a single location, and there is a continuing decline in the extent and quality of its habitat.