Madagascarophis fuchsi, Glaw, Frank, Kucharzewski, Christoph, Köhler, Jörn, Vences, Miguel & Nagy, Zoltán T., 2013

Glaw, Frank, Kucharzewski, Christoph, Köhler, Jörn, Vences, Miguel & Nagy, Zoltán T., 2013, Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar, Zootaxa 3630 (2), pp. 317-332 : 324-328

Madagascarophis fuchsi sp. nov.

Holotype. ZSM 2130/2007 (field number FGZC 1152; Figs. 4 and 5 ), adult female, collected close to the remains of the French Fort, Montagne des Français (12°19‘34“S, 49°20‘09“E, ca. 300 m above sea level), Antsiranana province, northern Madagascar, on 27 February 2007, by F. Glaw, J. Köhler, H. Enting, P. Bora, and A. Knoll.

Paratypes. ZSM 1620/2008 (MgF 0 61, FGZC 1737), adult female, collected at Montagne des Français, Antsiranana province, northern Madagascar (ca. 12°19‘S, 49°20‘E), at an uncertain date but most probably between 2005 and 2007, by the team of Frontiers Madagascar; ZMA 19623 (FGMV 2002.3093), adult female, collected at Montagne des Français, Antsiranana province, northern Madagascar, on 20 February 2003, by F. Glaw, R. D. Randrianiaina and A. Razafimanantsoa.

Diagnosis. The new species is characterized by the combination of the following characters: Relatively low number of midbody scale rows (25), relatively low number of ventral scales (171–172), a low ratio of ventrals/ subcaudals (2.38–2.61), a posterior part of the tail that is almost entirely black dorsally, and a relative high number of scales (13–16) bordering the parietals (temporals + occipital scales, without circumoculars and frontal). Madagascarophis fuchsi sp. nov. differs from all other species of the genus Madagascarophis by (1) the broad contact between the posterior inframaxillaries (genials) which in all other examined Madagascarophis specimens are separated by inserted gulars, and (2) by the lowest number of ventral scales within the genus (171–172 vs. 183–209 in M. colubrinus including M. c. citrinus , 205–224 in M. ocellatus , and 197–232 in M. meridionalis according to Domergue 1987). Furthermore the new species differs from M. meridionalis and M. ocellatus by having only 25 dorsal scale rows at midbody vs. 29–31 in M. ocellatus and 29–33 in M. meridionalis . Madagascarophis fuchsi differs from the syntopic subspecies M. c. septentrionalis (including the holotype and paratype of this taxon, see data in Table 1) by a lower number of midbody scale rows (25 vs. 27–29) and a distinctly lower number of ventral scales (171–172 vs. 186–200). It furthermore differs from M. colubrinus ssp. and M. meridionalis by substantial genetic differentiation (see Nagy et al. 2007, 2012 and Fig. 1 ).

Description of the holotype. Adult female in good state of preservation. Snout-vent length 417 mm, tail length 97 mm (18.9% of total length). Head length 23.1 mm, largest head width 13.3 mm. Body laterally depressed and head very distinct from neck. Pupil vertically elliptic. Eyes large, horizontal diameter 4.3 mm, larger than distance between anterior margin of eye and posterior edge of nostril (3.9 mm). Supralabials 9, not in contact with eye. Infralabials 12, 1st pair in contact behind mental and first 5 infralabials touching anterior inframaxillaries. Rostral broader than high (3.3 mm vs. 2.1 mm) and visible from above. Nasal divided below nostril, touching 1st and 2nd supralabial; upper margin of nostril almost touches internasal. Loreal 1, wider than high (1.8 mm vs. 1.3 mm) touching 2nd and 3rd supralabial. Circumoculars 9, including 1 large supraocular, 2 preoculars, 3 suboculars and 3 postoculars; upper preocular reaches top of head and touches frontal in one point at left side only. A small additional scale between 3rd and 4th supralabial and circumoculars on right side of head, this scale being not in contact with the loreal. Temporals according to formula 3+3. Upper surface of head with typical "colubrid" scalation consisting of 9 large shields. A pair of internasals (scale length 2.0 mm; suture 1.5 mm), a pair of prefrontals (scale length 2.6 mm; suture 2.5 mm); frontal longer than broad (6.4 mm vs. 4.0 mm), longer and broader than supraoculars (6.4 mm vs. 5.6 mm / 4.0 mm vs. 3.2 mm) and longer than distance between its anterior margin and posterior margin of rostral (6.4 mm vs. 4.2 mm); a pair of parietals (scale length 6.9 mm; suture 5.0 mm) which are bordered by 13 scales (temporals + occipital scales, without circumoculars and frontal). Two pairs of inframaxillaries of nearly same length but anterior ones broader. Posterior pair in broad contact with each other and separated by inserted gulars at posterior end only. 4 gulars separating posterior inframaxillaries from first preventral. 3 preventrals and 171 ventrals, cloacal scute divided, subcaudals 72 + terminal spine, mostly divided (exact formula: 1/1+4 [undivided] +24/24+2 [undivided] + 41/41+terminal spine). Ratio ventrals/subcaudals 2.38. Dorsal scales smooth, each with two apical pits along body, in 25-25-19 straight rows (counted at tenth ventral, midbody, and tenth ventral before vent) and homogeneous in size. Dorsocaudal scales smooth each with mostly 2 apical pits along the tail. For detailed dorsal scale row reduction and dorsocaudal scale row reduction see Appendix.

Dentition: 11 anterior maxillary teeth of nearly same size, followed after a diastema by two distinctly enlarged and grooved fangs. Maxillary bone reaching beyond palatine bone. Some palatine teeth enormously enlarged, larger than corresponding maxillary teeth. 6+13 mandibular teeth. Beside the small and indistinct anteriormost tooth the anterior 6 are distinctly enlarged (the 3rd – 5th largest) followed by 13 smaller teeth gradually decreasing in size posteriorly.

Coloration and pattern ( Fig. 5 a): Dorsal ground color of head, body and tail in life orange, which faded rapidly in alcohol to a pale cream ground color. Anterior part of head cape slightly darker, with a weakly developed dark postocular stripe. Color of iris in life orange (dark grey in preservative). Body pattern consists of irregular dark bands vertebrally and two rows of irregular shaped dark spots laterally, giving a reticulated appearance in general. Number of vertebral bands ca. 51. Upper row of lateral spots arranged alternating to bands and lower spots correspond to position of vertebral bands. Dorsal color of tail pale with irregular dark spots becoming almost entirely black posteriorly. Ventral surface of head, body and tail uniform pale cream. At posteriormost part of tail some subcaudals with black color.

Description of paratypes. ZSM 1620/2008: Adult female in good state of preservation. This paratype differs from the holotype in the following aspects: Snout-vent length 438 mm, tail length 99 mm (18.4% of total length). Tip of tail obtusely rounded, the terminal scale missing. Head length 24.5 mm, largest head width 15.2 mm. Horizontal diameter of eye 4.7 mm. Distance between anterior margin of eye and posterior edge of nostril 4.5 mm. Supralabials 8; infralabials 11, on right side 11th infralabial reaches slightly beyond posterior margin of last supralabial; first 4 infralabials in contact with anterior inframaxillaries. Rostral wider than high (3.7 mm vs. 2.0 mm); loreal wider than high (2.2 mm vs. 1.7 mm); upper preoculars reach top of head and touch the frontal on both sides; temporals according to formula 2+2 / 2+3. A pair of internasals (scale length 2.6 mm; suture 2.1 mm); a pair of prefrontals (scale length 2.8 mm; suture 2.5 mm); frontal longer than broad (6.9 mm vs. 4.5 mm), longer and broader than supraoculars (6.9 mm vs. 5.6 mm / 4.5 mm vs. 3.7 mm) and longer than the distance between its anterior margin and the posterior margin of rostral (6.9 mm vs. 4.9 mm); a pair of parietals (scale length 7.2 mm; suture 5.3 mm) which are bordered by 16 scales (temporals + occipital scales, without postoculars). 3–4 gulars separating posterior inframaxillaries from first preventral. 3 preventrals and 172 ventrals, cloacal scute divided, subcaudals 66 without terminal spine, mostly divided (exact formula: 2/2+3 [undivided] +21/21+2 [undivided] +11/11+9 [undivided] +1/1 +3 [undivided] +14/14). Ratio ventrals/subcaudals 2.61. Dorsal scales in 23-25-18 straight rows (counted at tenth ventral, midbody, and tenth ventral before vent). For detailed dorsal scale row reduction and dorsocaudal scale row reduction see Appendix.

Dentition similar to the holotype.

Coloration and pattern (in alcohol) rather different from the holotype: Dorsal ground color of body and head grey brown. Sides of head of same color with some light areas on supralabials. A slightly darker postocular stripe present. A dark stripe runs from suture of parietals onto neck, where it contacts a dark “∩” - shaped marking. Ventral surface of head strongly dark pale marbled. Body pattern consists of irregular black bands vertebrally and two rows of irregular shaped dark spots laterally. Number of vertebral bands ca. 44. Upper row of lateral spots arranged alternating to bands and lower spots corresponding to position of vertebral bands. Sometimes elements of pattern connected by thin dark lines giving a netlike appearance. Ventrolaterally dorsal ground color becomes marbled with yellowish ventral color. Ventral ground color yellowish irregularly powdered with dark pigment, the latter being more prominent on anterior and posterior part of venter. Outer edges of ventrals marbled with darker dorsal coloration. Dorsal color of the tail becomes entirely black at the tip. Subcaudals dark grey with only few pale markings.

Paratype ZMA 19623 was not available for study and the data provided here and in Table 1 are derived from pictures taken by E. Dondorp of the preserved specimen. According to these pictures, the dorsal coloration is yellowish with a moderately regular pattern of dark pigment which is concentrated on the edges of adjacent dorsal scales, forming a reticulated pattern similar to that found in the holotype ( Fig. 5 a). The ventral sides of head, body and tail are uniformly yellow without dark pigments. According to the field notes the dorsal coloration was orange in life.

Habitat, distribution and conservation status. The holotype was found active early at night during the rainy season in the karstic "tsingy" limestone area of Montagne des Français on the ground in deciduous dry forest. Although our observations are very limited it seems that M. fuchsi might be restricted to the tsingy formations whereas M. colubrinus is relatively abundant around and below the limestone karst. Our surveys in 2008 did not reveal any additional Madagascarophis fuchsi in Montagne des Français and surveys in nearby areas like Foret d'Orangea, Ampombofofo, Windsor Castle, Nosy Hara, Foret d'Ambre and Ankarana (D'Cruze et al. 2007, 2008; Megson et al. 2009; unpublished obs.) did not record this species. All these data indicate that M. fuchsi might be a microendemic species of Montagne des Français or a small region in northern Madagascar. Unfortunately, our surveys in 2007 and 2008 revealed that the forest of the Montagne de Français is under strong pressure and heavily degraded in almost all areas visited by us despite its recent inclusion in the network of Malagasy nature reserves. As in other parts of Madagascar the recent political instability seems to foster the degradation and the rate of forest clearing observed by us in 2007 and 2008 was clearly much higher than in the period of 2000–2007. Using the same rationale and IUCN criteria as applied during the assessment of Madagascan reptiles for presumably endemic species of Montagne des Français (e. g. Paroedura lohatsara ) we suggest to classify Madagascarophis fuchsi as Critically Endangered, because its Extent of Occurrence is less than 100 km 2, all individuals are from a single location, and there is continuing and serious decline in the extent and quality of its habitat at Montagne des Français.

Etymology. The senior author wishes to dedicate this new species to his former colleague and snake specialist Dieter Fuchs in recognition of his long-lasting work (1960–2007) as a technician in the herpetology section of the Zoologische Staatssammlung München, with congratulations and best wishes for his 70th birthday.

Figures

FIGURE 1. (A) Bayesian tree of representative Madagascarophis samples, based on 2197 nucleotide positions of two mitochondrial (16 S rRNA, cytochrome b) and one nuclear gene (c- mos). Bayesian posterior probabilities are given above branches, parsimony bootstrap values below branches. (B) Median-joining haplotype network of c- mos sequences. Circles represent haplotypes, short lines crossing indicate nucleotide substitutions (no heterozygous sequences were observed in the cmos data set).

FIGURE 4. Head drawings of Madagascarophis fuchsi sp. nov. (holotype, ZSM 2130 / 2007). (A) Lateral view; (B) dorsal view. Drawings by Ruth Kühbandner.

FIGURE 5. (A) Holotype of Madagascarophis fuchsi sp. nov. in life (ZSM 2130 / 2007). (B) Madagascarophis colubrinus septentrionalis from Windsor Castle (ZSM 2212 / 2007) in life, representing a typical color morph of the northernmost populations.