Madagascarophis lolo, Sara Ruane, Frank T. Burbrink, Bernard Randriamahatantsoa & Christopher J. Raxworthy, 2016

Sara Ruane, Frank T. Burbrink, Bernard Randriamahatantsoa & Christopher J. Raxworthy, 2016, The Cat-eyed Snakes of Madagascar: Phylogeny and Description of a New Species of Madagascarophis (Serpentes: Lamprophiidae) from the Tsingy of Ankarana, Copeia 104 (3), pp. 712-721 : 716-718

Madagascarophis lolo , new species

urn:lsid:zoobank.org:act: E94496 B3-AC04-4FE2-AFA6- 8CC7DBE3A63A

Figures 1, 2, 3, 4, 5; Table 1

Holotype.— AMNH 176422 ( RAX 12626 ), adult male (Fig. 1), Madagascar, Antsiranana Province, Diana Region, Ankarana National Park, ~ 4 km northwest of the village of Mahamasina, tsingy karst trail , 1 0 2 m elevation, 4 9.1 1 5 0 7 ° E, 12.94210 ° S, 9 February 2014, 1930 hours, B. Randriamahatantsoa, C. Raxworthy, S. Ruane.

Diagnosis.— A new species of Madagascarophis than can be diagnosed from its congeners by the following combination of characters: an overall gray body color with a black vertebral stripe and alternating light gray blotches down the dorsum, 25 scale rows at midbody, 189 ventral scales and 56 divided subcaudal scales, with extended contact of the posterior inframaxillary scales. Madagascarophis lolo , new species, differs from all other species of Madagascarophis having a gray body color combined with an alternating pattern of pale gray blotches along the vertebral column and the presence of black scales on the vertebral row scales, giving the appearance of a thin black dorsal stripe (Figs. 1, 3, 4). This overall combination of color and pattern is unique among species of Madagascarophis (see Domergue, 1987:fig. 4 for comparison). Madagascarophis lolo , new species, differs from all other species with the exception of M. fuchsi by having extended contact of the posterior inframaxillary scales ( Fig. 5 ). We note the specimen of M. lolo , new species, has a single gular scale that infringes on the posterior end of the posterior inframaxillaries. However, with the exception of M. fuchsi , the posterior inframaxillary contact of M. lolo , new species, is still much greater than for the other species of Madagascarophis ( Fig. 5 ; see Glaw et al., 2013a for additional examples).

Madagascarophis lolo , new species, differs from M. fuchsi by having a lower number of infralabial scales (10 M. lolo vs. 12– 13 M. fuchsi) and a higher ventral scale count (171– 174 M. fuchsi vs. 1 8 9 M. lolo). However, this 1 5 ventral scale difference falls within the intraspecific range of other species (e.g., 35 ventral scales in M. meridionalis ). It differs from the other species of Madagascarophis except M. colubrinus by having a lower ventral scale count (189 M. lolo): 183–209 in M. colubrinus , 205–224 in M. ocellatus , and 197–232 in M. meridionalis . A general difference between M. lolo , new species, and most other Madagascarophis is the dorsal scale count at midbody. Madagascarophis lolo , new species, has 25 dorsal scale rows as does M. fuchsi , in contrast to the 27–29 typically found in M. colubrinus (rarely 25, and not syntopically), 29–33 in M. meridionalis , and 29–31 in M. ocellatus ( Glaw and Vences, 2007; Glaw et al., 2013a). It also differs genetically from all other species in the genus, e.g., M. lolo vs.

M. fuchsi , COI uncorrected pairwise distance =9.6% ( Table 1 ). For specimens not examined here (Appendix 1), additional data were used from Domergue (1987) and Glaw et al. (2013a) for the diagnosis.

Description.— Adult male in excellent state of preservation, tail complete, short ventral slit midbody for DNA tissue sample, lower body slit for assessing gonad development (fully formed mature testes; 10 mm length, 2 mm width). Snout–vent length 426 mm, tail length 65 mm, tail short (13% of total body length). Head length 20 mm, width 12 mm. Head distinct from neck. Eyes large, 3 mm horizontal diameter, pupil vertically elliptical. Supralabials 8, not in contact with the eye. Infralabials 10, first pair in contact behind mental, infralabials 1–5 in contact with inframaxillaries. Rostral broader than high, 3 mm wide/ 1.5 mm high, visible from above. Nasal divided below nostril, in contact with 1st and 2nd supralabials. Single loreal, in contact with nasal, preoculars, prefrontal, and supralabials 2 and 3. Circumoculars 9, 1 supraocular, 2 preoculars, 3 suboculars, and 3 postoculars. Temporals 4 + 4/4 + 5. Dorsal surface of head includes pair of internasals (width 1.7 mm /length of suture 2 mm), pair of prefrontals (width 2 mm /length of suture 2.1 mm), pair of supraoculars (width 2.8 mm /length 4.8 mm), frontal longer than wide (length 5.7 mm /anterior width 2.9 mm), pair of parietals (length of suture 4.6 mm). Two pairs of inframaxillaries (anterior inframaxillary length 4.9 mm, posterior inframaxillary length 3.1 mm), posterior inframaxillaries substantially in contact with each other excepting small gular scale at posterior end ( Fig. 5 ). Dorsal scale rows 23-25-19 at 10th ventral from anterior, midbody, and 10th ventral anterior to cloaca.

Coloration and pattern.— Dorsal ground color gray in life, alternating lighter gray blotches/squares alongside vertebral column, many vertebral row scales black in coloration, giving general appearance of black dorsal line interrupted occasionally by gray scales (Fig. 1). At roughly the posterior 1/3 of the body, dorsal scale rows 7 and 8 occasionally black, giving spotted appearance in lateral view. Overall coloration pale gray in preservation. Color of iris in life gray/silver with gold flecking (Fig. 1), opaque gray in preservation. Dorsum of head, including rostral, internasals, prefrontals, frontal, supraoculars, and parietals gray. Supralabials 4–8 with gray and white mottling, infralabials with gray and white mottling. Slightly darker diffuse brown line runs from the posterior of eye to posterior margin of mouth. Tail gray with black mottling, darker and more contrasting compared to body, with slight flush of pale orange towards tail tip. Ventral scales cream with no pattern anteriorly, small amounts of gray flecking on ventral scales beginning at the posterior 1/3 of the body, continuing and increasing in intensity onto the subcaudal scales.

Natural history.— Similar to other species of Madagascarophis , M. lolo appears to be crepuscular/nocturnal; the specimen was found active on the ground at 1930 hours on tsingy karst rocks, in an exposed area with low scrub habitat. This is very similar to what has been described for M. fuchsi ( Glaw et al., 2013a) , and our own observation of the M. fuchsi sample included here, which we found outside a small cave in the karst system of the Montagne des Français massif, approximately 7 0 km away. By contrast, the other species of Madagascarophis found at Ankarana, M. colubrinus , was common in canyon forests and surrounding relict forests, as well as in anthropogenically disturbed habitat. We suspect the reason that M. lolo has gone undetected for so long at Ankarana is that the exposed tsingy plateau has been poorly surveyed at night in previous expeditions due to problems of gaining safe access to these areas. Madagascarophis lolo may be endemic to the karst habitats of Ankarana, and possibly Analamerana, which is the closest karst system to the east.

Etymology.— The species name, lolo , is taken from the Malagasy word for ghost; it is a noun in apposition to the genus name. This name refers to 1) the ghostly pale gray color of the holotype, and 2) that M. lolo has eluded discovery for so long at Ankarana, arguably one of the better surveyed sites in Madagascar.

Figures

Fig. 5. Ventral head view, showing the posterior infralabial scale contact of Madagascarophis lolo (A; RAX 12626), M. fuchsi (B; RAX 12424), M. colubrinus (C; RAX 10540); a scale bar is provided for each species. Photos by SR (A, C) and FTB (B).