Liophidium sp.

Glaw, Frank, Kucharzewski, Christoph, Nagy, Zoltán T., Hawlitschek, Oliver & Vences, Miguel, 2014, New insights into the systematics and molecular phylogeny of the Malagasy snake genus Liopholidophis suggest at least one rapid reversal of extreme sexual dimorphism in tail length, Organisms Diversity & Evolution (New York, N. Y.) 14 (1), pp. 121-132 : 125-127

Liophidium sp. ( Domergue 1988)

"genus and species inquirenda" ( Cadle 1996a)

Holotype ZFMK 62235 , probably adult male (right hemipenis partially everted), collected dead on a trail near Hotel Feon'ny Ala (18° 56.845'S, 48° 25.078'E, ca. 940 m a. s. l.), at ca. 2.5 km distance from the village of Andasibe, central eastern Madagascar, on 31 January 1996 by F. Glaw.

Paratype MNHN 1988.331, probably adult male (hemipenes fully everted), collected at Perinet [=Andasibe], central eastern Madagascar, on 19 December 1966 by E. R. Brygoo (according to Domergue 1988), but see Cadle (1996a: 455).

Diagnosis A species attributed to the genus Liopholidophis because of its general morphology and its position in the molecular phylogenetic tree ( Fig. 1 ). The new species differs from all species of Liopholidophis , the superficially similar Liophidium , and all other Madagascan lamprophiid snakes by the presence of only 15 dorsal scale rows around the midbody (versus at least 17 scale rows at midbody in all other advanced Malagasy snakes). In addition it differs from all other species of Liopholidophis except L. rhadinaea by much smaller size of males (maximum total length 312.5 mm versus 889–1636 mm), lower number of subcaudals in males (71–77 versus 88– 221), short relative tail length (male tail length/total length 0.28–0.29 versus 0.30–0.55), presence of three light spots in the neck (versus absence) and a uniformly red belly in life that fades in preservative (never red in all other species). The new species is similar in general habitus and colouration (red belly in life, light spots in the neck) to Liopholidophis rhadinaea ( Fig. 5 ) but differs from this species by having fewer ventrals in males (149–158 versus 170–179), fewer subcaudals in males (71–77 versus 126–137), and smaller size (maximum total length of males 312.5 mm versus 749 mm, maximum snout-vent length of males 221.5 mm versus 429 mm). L. baderi is the sister species and most similar to L. oligolepis sp. nov., which will be described and diagnosed below. L. baderi differs furthermore from all other Liopholidophis species by substantial genetic differentiation.

Description of the holotype Adult male, in good state of preservation. Total length 283.0 mm, snout-vent-length 205.0 mm, tail length 78.0 mm (28% of total length). Greatest head width (temporal region) 5.1 mm, head length (tip of snout to end of mandibles) 8.4 mm. Distance between tip of snout and the posterior edge of the parietals 7.0 mm. Body dorsoventrally depressed (probably according to the finding situation dead on a road). Head only little wider than neck. Pupil round. Eye large, diameter horizontally 1.7 mm, larger than the distance between anterior margin of eye and posterior edge of nostril (1.1 mm). Dorsal scales smooth, without apical pits in 15–15–15 rows, but 17 scale rows directly behind the head. Scale row reduction from 17 to 15 rows at the level of ventral 5. Two preventrals, 158 rounded, not angulated ventrals. One gular scale between the posterior inframaxillaries and the first preventral. Anal scute divided. Subcaudals 71/72 all divided plus terminal spine. Reduction of dorsocaudal rows from 8 to 6 at the level of subcaudals 9– 12, from 6 to 4 at the level of subcaudals 33–36 and from 4 to 2 at the level of subcaudals 69–70.

Supralabials 8, with 4 th –5 th touching the eye. Infralabials 9, the first pair in contact behind the mental, 1 st –4 th touching the anterior inframaxillary, 4 th –5 th touching the posterior inframaxillary, the posteriormost infralabial reaches slightly beyond the posterior margin of the last supralabial. Two pairs of inframaxillaries both the same width, but the anterior (2.0 mm) pair slightly shorter than the posterior (2.4 mm). Rostral wider (1.8 mm) than high (0.9 mm) and visible from above. Nasal semi-divided (undivided above the nostril, divided below), touching first and second supralabial. One loreal present, higher (0.6 mm) than wide (0.4 mm), touching second and third supralabial. One preocular, scarely visible from above, not in contact with frontal, touching third and fourth supralabial. Two postoculars, lower one slightly larger than upper one, upper postocular in contact with parietal, lower postocular touching parietal, anterior temporal, fifth and sixth supralabial. Temporals 1+2+3. Dorsal surface of head shows the typical lamprophiid scalation consisting of nine shields. Frontal longer (2.8 mm) than wide (1.8 mm), longer than the distance to the posterior margin of rostral (1.5 mm). Parietals longer (3.3 mm) than the frontal.

Dentition: Maxillary teeth 25+2 without diastema, nearly same size, the last two teeth distinctly enlarged and ungrooved. Maxillary reaching beyond the palatine. Dentary teeth ca. 30, small, equal in size.

Colouration in preservative: General dorsal ground colouration brown. Dorsal rows 1–4 light brown become gradually darker vertebrally. Lower part of scale row five dark brown followed by a narrow pale stripe, situated in the middle of the scale and vertebrally bordered by very fine darker pigment. Upper part of row 5 and row 6 light brown. Rows 7+8 a little darker than rows before creating the appearance of a broad, indistinctly darker vertebral stripe. The pale lateral stripe starts six scales behind the posterior margin of the parietal and runs continuously on the fifth row on the body and on the second scale row on the tail nearly until the tip of tail. On the neck there are three distinct white spots. These ocelli occupy 1– 1.5 scales and are bordered by a thin dark ring. Ground colour of dorsal side of head reddish brown. Lower part of the supralabials pale. A white stripe runs from the rostral across the supralabials until the neck. This stripe is indistinctly lighter than the pale colour of the supralabials and irregularly bordered ventrally by dark pigment from the sixth supralabial until the neck and dorsally from a weakly developed dark postocular strip, which starts at the lower margin of the eye and runs until the level of the third ventral. Underside of head, body and tail uniform pale in preservative, except the extreme outer margins of ventrals and subcaudals, which show the same light brown colouration as the first dorsal scale row. Photographs of the preserved holotype will be made publically available via the Reptile Database (http://www.reptile-database.org/). Details of life colouration unknown, but still with reddish venter when found shortly after death, fading in preservative into whitish.

Description of the paratype A detailed description of the paratype was already provided by Domergue (1988) and Cadle (1996a).

Adult male. Total length 312.5 mm, snout-vent length 221.5 mm, tail length 91.0 mm (29% of total length). Greatest head width (temporal region) 4.8 mm, head length (tip of snout to end of mandibles) 9.1 mm. Distance between tip of snout and the posterior edge of the parietals 7.3 mm. Body slightly compressed laterally. Head only a little wider than neck. Pupil round. Eye large, diameter horizontally 1.6 mm, larger than the distance between anterior margin of eye and posterior edge of nostril (1.2 mm). Dorsal scales smooth, without apical pits in 15–15–15 rows, but 17 scale rows directly behind the head, scale row reduction from 17 to 15 rows at the level of ventral 4. Two preventrals, 149 rounded, not angulated ventrals. One gular scale between posterior inframaxillaries and the first preventral. Anal scute divided. Subcaudals 77/77 all divided plus terminal spine. Reduction of dorsocaudal rows from 8 to 6 at the level of subcaudal 11, from 6 to 4 at the level of subcaudals 31–33 and from 4 to 2 at the level of subcaudal 71.

Supralabials 8, with 4 th –5 th touching the eye. Infralabials 9, the first pair in contact behind the mental, 1 st –4 th touching the anterior inframaxillary, 4 th –5 th touching the posterior inframaxillary. Two pairs of inframaxillaries both the same width, but the anterior (2.3 mm) pair slightly shorter than the posterior (2.6 mm). Rostral wider (1.8 mm) than high (1.0 mm), visible from above. Nasal semi-divided (undivided above the nostril, divided below), touching first and second supralabial. One loreal present, higher (0.7 mm) than wide (0.4 mm), touching second and third supralabial. One preocular, scarely visible from above, not in contact with frontal, touching third and fourth supralabial. Two postoculars, lower one slightly larger than upper one, upper postocular in contact with parietal, lower postocular touching parietal, anterior temporal, fifth and sixth supralabial. Temporals 1 +1 +2/1 +1+ 3. Dorsal surface of head shows the typical lamprophiid scalation consisting of nine large shields. Frontal longer (2.7 mm) than wide (2.1 mm), longer than the distance to the posterior margin of rostral (1.5 mm). Parietals longer (3.8. mm) than the frontal.

Dentition: Maxillary teeth 26+2 [27+2 on the right side, Cadle 1996a] without diastema, the last two teeth distinctly enlarged and ungrooved. Maxillary reaching beyond the palatine. Dentary teeth ca. 31, small, equal in size.

Colouration in preservative: In comparison with Figure 42 in Cadle (1996a) the specimen has significantly faded, especially the three light spots on the neck. Nevertheless, the colouration and pattern of the paratype generally correspond with the holotype. The white lateral stripe starts seven scales behind the parietal and runs on the lower part of the fifth scale row.

Habitat, natural history, and conservation status The male holotype was collected dead on a trail near the hotel "Feon' ny Ala", at ca. 2.5 km distance from the village Andasibe. This locality was largely surrounded by secondary forest, but was very close to the Reserve Speciale Analamazoatra , which appears well protected. L. baderi is only known from a single location (Andasibe) in central eastern Madagascar. Although the area around this village has been extensively surveyed by herpetologists, local guides and tourists, only two specimens are known so far, one collected in 1966 and the other collected 30 years later in 1996, suggesting that this snake is rather secretive or rare at least around the type locality. Regular surveys of snake roadkills between the village of Andasibe and the Hotel Feon' ny Ala did not reveal the snake either ( R. Dolch, pers. comm.). Due to the very limited knowledge we suggest considering this species as "Data Deficient" according to the IUCN criteria used for the recent assessment of Malagasy reptiles .

Etymology The specific name is a patronym for Frank Bader ( Germany) in recognition of his support of research and nature conservation through the BIOPAT initiative.

Figures

Fig. 1 Phylogenetic tree of all Liopholidophis species based on sequences of three mitochondrial and one nuclear gene (16S rRNA, cytb, COI and c-mos). For L. baderi no 16S sequence was available. Bayesian posterior probabilities (10 million generations) are given above branches, parsimony bootstrap values (2,000 replicates) below branches. The relation of tail length/snout-vent length (red: female, blue: male) is given for all species of Liopholidophis, and for Liophidium torquatum, on the right of the species names. Reconstructed values for nodes are given at the branches, as results of parsimony/maximum likelihood (95% confidence interval). The lower figure shows the same tree with sexual dimorphism in relative tail length plotted on taxa (expressed as ratio of relative tail length of males/ females). Ancestral values calculated on ML-reconstructed values for males and females. Symbols show origin (>) and reversal (<) of extreme dimorphism (values 0.43-0.53, i.e. females having relative tail lengths roughly half that of males or lower)

Fig. 5 Liopholidophis rhadinaea in life, from Talatakely, Ranomafana National Park (ZSM 1602/2008)