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Lycodryas maculatus (GÜNTHER, 1858)

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Higher TaxaPseudoxyrhophiidae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesLycodryas maculatus maculatus GÜNTHER 1858
Lycodryas maculatus comorensis (PETERS 1874) 
Common NamesE: Spotted Tree Snake [maculatus]
Comoro Island, Johanna Island, Mayotte [sanctijohannis] 
SynonymDipsadoboa maculata GÜNTHER 1858: 183
Lycodryas sancti johannis GÜNTHER 1879: 218
Stenophis maculatus — BOULENGER 1896: 43
Lycodryas maculatus — MEIRTE 1999
Lycodryas sanctijohannis — MEIRTE 1999
Lycodryas sanctijohannis — NAGY et al. 2010
Lycodryas maculatus — NAGY et al. 2010
Lycodryas sanctijohannis — HAWLITSCHEK et al. 2011
Lycodryas maculatus — WALLACH et al. 2014: 399
Lycodryas sanctijohannis — EL-YAMINE et al. 2016

Lycodryas maculatus comorensis (PETERS 1874)
Dipsas (Heterurus) Gaimardii var. comorensis PETERS 1874
Lycodryas sanctijohannis var. mayottensis BOETTGER 1913
Lycodryas maculatus comorensis — HAWLITSCHEK et al. 2012 
DistributionComoro Islands (Mayotte)

Type locality: see comment, changed to ‘‘unknown’’ by Boulenger 1896.

sanctijohannis: Comoro Island, Johanna Island, Mayotte; Type locality: “Anjouan island’’  
TypesHolotype: BMNH 1946.1.21.47
Holotype: BMNH 1946.1.5.20 [sanctijohannis]
Holotype: Not traced and considered lost, juvenile [comorensis]
Neotype: ZSM 42/2010, adult male; collected 7 February 2010; Comoros Archipelago, MAYOTTE, Petite-Terre, near Moya, under bark on a tree; by O. Hawlitschek, J. Berger, B. Brückmann. [comorensis] 
DiagnosisDiagnosis (maculatus): Largest subspecies of Comoran tree snakes, snout-vent length max. 867 mm; max. snout-vent length for L. cococola cococola sp. n. 650 mm, for L. maculatus comorensis 757 mm and for L. cococola innocens ssp. n. 835 mm. 19 middorsal scale rows, no specimens with 17 known (see Table 1 and Fig. 3 for other taxa of Comoran Lycodryas). 233 to 259 ventral scales, 85 to 126 subcaudal scales, posterior ones divided. Loreal in contact with supralabials 2 and 3, as in L. maculatus comorensis, but unlike the other two subspecies. Anal shield divided. BMNH 1946.1.21.47 is the only specimen studied with 8/8 infralabial scales, while all other Comoran Lycodryas specimens have at least 9/9. Males dorsally grey, head with marbled pattern of darker spots and dots. Lower part of supralabials in males white, upper part brown or black. Body with regular middorsal band of dark brown blotches, scales between these blotches sometimes appearing brighter. Dark ventral stripe always absent. Females with typical pattern of reddish, brownish or yellowish dorsal and yellowish ventral side [from HAWLITSCHEK et al. 2012].

Diagnosis (comorensis): Subspecies of Comoran Lycodryas of intermediate snout-vent length (max. 757 mm); longer than L. cococola cococola sp. n. (max. 650 mm), shorter than L. maculatus maculatus (max. 867 mm) and L. cococola innocens ssp. n. (max. 835 mm). 19 middorsal scale rows, 17 in MNHN 1884-518 (see Table 1 for comparison with other taxa of Comoran Lycodryas). 227 to 261 ventral scales, 84 to 117 subcaudal scales, posterior ones divided. Loreal in contact with supralabials 2 and 3, like in L. maculatus maculatus, but unlike other taxa of Comoran Lycodryas. Anal shield divided. Males can be distinguished from other taxa of Comoran Lycodryas by a mostly clearly defined, sometimes diffuse dark stripe extending from the gular region to the tail tip along the row of ventral scales. Dorsal ground color of males grey or olive-grey, the latter being recorded in no other taxa of Comoran Lycodryas. Lower part of supralabials in males white, upper part brown or black; pattern of more or less diffuse dark spots extending over the dorsal side. Dark dorsal crossbands were never recorded in any adult specimen. Females show the typical pattern of reddish, brownish or yellowish dorsal and yellowish ventral side [from HAWLITSCHEK et al. 2012]. 
CommentDistribution: Erroneously reported from Madagascar. The original type locality given was “Central America”. The taxon Dipsas (Heterurus) gaimardii Schlegel var. comorensis Peters, 1874 [107], has been considered a record of L. gaimardii from Mayotte. According to HAWLITSCHEK et al. 2012, all Lycodryas from Mayotte belong to a single taxon that is not conspecific with L. gaimardii. Therefore, the designation of a neotype is necessary to stabilize the nomenclatural identity of Lycodryas populations from Mayotte.

Synonymy: after BOULENGER 1896 and HAWLITSCHEK et al. 2012 (who synonymized Lycodryas sanctijohannis with L. maculatus). However, Zaher et al. 2019 found that sanctijohannis is more closely related to cococola than maculatus and thus warrants recognition. However, the GenBank sequences Zaher used were from different individuals, some of which were maculatus, some cococola. They are listed as sanctijohannis on GenBank because they were uploaded before L. maculatus was revalidated and L. cococola described (O. Hawlitschek, pers. comm., 8 Jan. 2020).

NAGY et al. (2010) write “Lycodryas maculatus is here assigned to Lycodryas only tentatively. This taxon was (perhaps erroneously) reported from the region: It was originally described by Günther (1858) as Dipsadoboa maculata from Central America. The type of D. maculata has dorsal scales in 19 rows, has 243 ventrals and 126 subcaudals and an undivided anal scale. Furthermore, it has a single praeocular scale and three postoculars at both sides, and it is characterized by a total length of 655 mm (tail length: 165 mm) according to Boulenger (1896). According to Domergue (unpublished data), however, the specimen was not available for re-examination any more [in his unpublished notes he writes: ‘pas... reconnu (retrouve ́ )’.

Type species: Lycodryas sanctijohannis is the type species of the genus Lycodryas GÜNTHER 1879.

Definition and description (Lycodryas): Typically 8 (range: 7–9) upper labials (usually 4th and 5th touching eye); 6–10 lower labials; 1 loreal scale; 17–19 rows of dorsal scales at mid-body; 13 or 15 rows of dorsal scales at the last ventral; 185–284 ven- tral scales; anal plate usually divided; 70–130 subcaudals (divided, undivided or combination of both); pupil is a small vertical ellipse when contracted; hemipenis as far as known deeply divided. Further characters are given in Table 2 in NAGY et al. 2010.

Habitat: fully arboreal (Harrington et al. 2018). 
EtymologyNamed after Johanna (Anjouan) Island (Comoro Islands). 
  • Boulenger, G.A. 1896. Catalogue of the snakes in the British Museum, Vol. 3. London (Taylor & Francis), xiv + 727 pp. - get paper here
  • Domergue, C.A. 1970. Notes sur les serpents de la région malgache. Lycodryas maculatus (Günther, 1858), espèce des Comores. Description de deux femelles. Bull. Mus. natn. Hist. nat., Paris 3: 449-451. - get paper here
  • El-Yamine, Ali Mohamed; Hamada Chakira, Nadjim Ahmed Mohamed, Ouledi Ahmed, Anllaoudine Abou, Cheng Xiao Hu, Hantanirina Rasamimanana 2016. Distribution of the forest nocturnal reptiles of Anjouan Island, their main habitats significances and conservation status. Journal of Biodiversity and Environmental Sciences 9 (1): 159-176 - get paper here
  • Günther, A. 1858. Catalogue of Colubrine snakes of the British Museum. London, I - XVI, 1 - 281
  • 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
  • Hawlitschek, Oliver; Boris Brückmann, Johannes Berger, Katie Green, Frank Glaw 2011. Integrating field surveys and remote sensing data to study distribution, habitat use and conservation status of the herpetofauna of the Comoro Islands. ZooKeys 144: 21-79 - get paper here
  • Nagy, Z. T., Glaw, F. & Vences, M. 2010. Systematics of the snake genera Stenophis and Lycodryas from Madagascar and the Comoros. Zoologica Scripta 39, 426–435 - get paper here
  • Peters, Wilhem Carl Hartwig 1874. Über eine von Hrn. F. Pollen und van Dam auf Madagascar und anderen ostafrikanischen Inseln gemachte Sammlung von Amphibien. Monatsber. königl. Akad. Wiss. Berlin. 1873 (December): 792-795 - get paper here
  • Wallach, Van; Kenneth L. Williams , Jeff Boundy 2014. Snakes of the World: A Catalogue of Living and Extinct Species. [type catalogue] Taylor and Francis, CRC Press, 1237 pp.
  • Zaher H, Murphy RW, Arredondo JC, Graboski R, Machado-Filho PR, Mahlow K, et al. 2019. Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes). PLoS ONE 14(5): e0216148 - get paper here
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