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Madagascarophis lolo RUANE, BURBRINK, RANDRIAMAHATANTSOA & RAXWORTHY, 2016

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Higher TaxaPseudoxyrhophiidae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common Names 
SynonymMadagascarophis lolo RUANE, BURBRINK, RANDRIAMAHATANTSOA & RAXWORTHY 2016 
DistributionMadagascar (Antsiranana)

Type locality: Madagascar, Antsiranana Province, Diana Region, Ankarana National Park, ~4 km northwest of the village of Mahamasina, tsingy karst trail, 102 m elevation, 49.115078° E, 12.942108° S.  
Reproduction 
TypesHolotype: AMNH 176422 (RAX 12626), adult male (Fig. 1 in Ruane et al. 2016), Madagascar, Antsiranana Province, Diana Region, Ankarana National Park, ~4 km northwest of the village of Mahamasina, tsingy karst trail, 102 m elevation, 49.115078E, 12.942108S, 9 February 2014, 1930 hours, B. Randriamahatantsoa, C. Raxworthy, S. Ruane. 
DiagnosisDiagnosis: 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 in Ruane et al. 2016). 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. 189 M. lolo). However, this 15 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 syntopi- cally), 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 1⁄4 9.6% (Table 1).
 
CommentBehavior: crepuscular/nocturnal

Habitat: tsingy karst rocks, in an exposed area with low scrub habitat. Semi-arboreal (Harrington et al. 2018). 
EtymologyThe 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. 
References
  • Harrington, Sean M; Jordyn M de Haan, Lindsey Shapiro, Sara Ruane 2018. Habits and characteristics of arboreal snakes worldwide: arboreality constrains body size but does not affect lineage diversification. Biological Journal of the Linnean Society 125 (1): 61–71 - get paper here
  • Ruane, Sara; Frank T. Burbrink, Bernard Randriamahatantsoa, and 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: September 104 (3): 712-721 - get paper here
 
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