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Calamaria nebulosa LEE, 2021

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Higher TaxaColubridae, Calamariinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Clouded Reed Snake 
SynonymCalamaria nebulosa LEE 2021
Calamaria yunnanensis – STUART & HEATWOLE, 2008
Calamaria yunnanensis – QUAH et al., 2008 (in part)
Calamaria cf. yunnanensis – WEINELL et al. 2021 
DistributionLaos (Phongsaly Province)

Type locality: Laos, Phongsaly Province, Phongsaly District, on the mountainous road to Udomxai [= Oudomxai] about 25 km from Phongsaly City (21º29’N, 102º12’E; ~1000 meters above sea level)  
Reproductionoviparous (presumed, fide Lee, 2021) 
TypesHolotype: FMNH 258666 (field tag: HKV 64382), an adult female collected dead on road (DOR) by Mr. Bounthavy Phommachanh on 6 October 1999. 
DiagnosisDiagnosis: A species of Calamaria distinguished from all other members of the genus from mainland Southeast Asia by having the following combination of morphological characters: (1) rostral wider than high; (2) portion of rostral scale visible from above approximately half the length of the prefrontal suture; (3) six scales and shields surrounding the paraparietal scale; (4) absence of a preocular scale; (5) four supralabials, with the second and third touching the eye; (6) five infralabials, the first pair separating the mental scale from the anterior chin shields; (7) 179 ventrals and three gular scales; (8) 22 paired subcaudals; (9) dorsal color pattern in preservative bluish-gray with five indistinct dark-brown longitudinal stripes; (10) ventral surface of tail in preservative uniform yellow, immaculate; and (11) absence of a distinct white nuchal collar posterior to the head. (Lee, 2021)

Comparisons: Calamaria nebulosa sp. nov. was compared to all other members of the genus inhabiting mainland Southeast Asia. Since C. nebulosa sp. nov. is only known from a single female, morphological data from male Calamaria specimens were excluded from the comparisons. C. nebulosa sp. nov. is morphologically assigned to Calamaria by having an elongate, cylindrical-shaped body, absence of a loreal scale, four supralabials, five infralabials, and 13 rows of iridescent dorsal scales throughout the body.
The absence of a preocular scale in Calamaria nebulosa sp. nov. distinguishes it from all other congeners inhabiting mainland Southeast Asia (i.e. Calamaria abramovi Orlov, 2009, Calamaria albiventer (Gray, 1835), Calamaria andersoni Yang and Zhang. 2018, Calamaria buchi Marx and Inger, 1955, Calamaria concolor Orlov, Nguyen, Nguyen, Ananjeva and Ho, 2010, Calamaria gialaiensis Ziegler, Sang and Nguyen, 2009, Calamaria ingeri Grismer, Kaiser and Yaakob, 2004, C. lumbricoidea, C. pavimentata, C. sangi Nguyen, Sang and Ziegler, 2009, C. septentrionalis and C. strigiventris Poyarkov, Nguyen, Orlov and Vogel, 2019) except for Calamaria dominici Ziegler, Tran and Nguyen in Ziegler et al., 2019, Calamaria lovii gimletti Boulenger, 1905, C. lovii ingermarxorum Darevsky and Orlov, 1992, C. schlegeli schlegeli, C. thanhi Ziegler and Quyet, 2005 and C. yunnanensis, where the condition of the preocular is either absent or variable. From these taxa, C. nebulosa sp. nov. is distinguished from C. schlegeli schlegeli (the only species of Calamaria in the region where the preocular can be present or absent) by having the end of the tail tapering to an obtuse point (vs. blunt point), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs. reducing above the 3rd–25th subcaudal), and by having a bluish-gray dorsum with longitudinal stripes (vs. dorsum usually black with a reddish head or a yellow nuchal collar with no light vermiculations on dorsal scales); from C. dominici by having the mental separated from the anterior genials (vs. mental in narrow contact with anterior genials), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs. reducing above the 5th–6th subcaudal), 179 ventrals and 22 subcaudals in female (vs. 174 ventrals and 17–18 subcaudals in females), tail length 7.9% of total length in female (vs. 6.0% of total length in females) and by having a bluish-gray dorsum with longitudinal stripes (vs. dark-blue dorsum covered with bright yellow spots); from C. lovii gimletti by having the posterior genials meeting only anteriorly (vs. meeting at midline), the end of the tail tapering to an obtuse point (vs. tapering to a blunt point), 179 ventrals and 22 subcaudals in female (vs. 215–249 ventrals and 10–12 subcaudals in females) and by having a bluish-gray dorsum with longitudinal stripes (vs. dorsal color variable, but usually with a light nuchal collar, small yellow spots on the nape and tail and no light vermiculations on dorsal scales); from C. lovii ingermarxorum by having the portion of the rostral scale visible from above roughly equal to the half the length of the prefrontal suture (vs. greater than half the length of the prefrontal suture), the length of the prefrontal smaller than the frontal scale (vs. longer than frontal), mental separated from the anterior genials (vs. in contact with anterior genials), posterior genials meeting only anteriorly (vs. meeting at midline), the end of the tail tapering to an obtuse point (vs. tapering to a blunt point), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs. reducing above the 4th–5th subcaudal), 9 modified maxillary teeth (vs. 8), and by having a bluish-gray dorsum with longitudinal stripes (vs. dorsum blue-gray with a light nuchal collar and grayish venter); and from C. thanhi by having the end of tail tapering to an obtuse point (vs. tapering gradually to a point), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs. not reducing to four rows), 179 ventrals and 22 subcaudals in female (vs. 198 ventrals and 21 subcaudals in female) and by having a bluish-gray dorsum with longitudinal stripes (vs. dorsum dark blue or gray with several light body and tail rings).
Most specimens of C. yunnanensis can be distinguished from C. nebulosa sp. nov. by having a distinct nuchal color posterior to the head and a dark-brown midventral stripe under the surface of the tail (vs. nuchal collar and midventral stripe absent) and by having dark-brown margins on each of the ventral scales (vs. ventral scales immaculate). However, some specimens of C. yunnanensis do not share these color pattern traits. In specimens where the coloration is difficult to interpret in preservative or is variable, C. nebulosa sp. nov. can be differentiated from C. yunnanensis by having 179 ventrals in female (vs. 199), subcaudal/total body scale ratio 10.9% in female (vs. 8.7%), tail length approximately 7.9% of total length in female (vs. 5.0%), dorsal scales reducing to six rows above the 12th subcaudal (vs. 3rd to 5th subcaudal) and reducing to four rows above the last subcaudal (vs. third-to-last subcaudal). All specimens of C. yunnanensis have narrow dark-brown longitudinal stripes present on the margins of each dorsal scale row, occasionally restricted to the flanks in some specimens (vs. five, wider dark-brown longitudinal stripes, not present on every margin of each dorsal scale row), but these characters can be difficult to interpret on some specimens and should be treated with some wariness. (Lee, 2021)

Color in life: Not recorded (Lee, 2021) 
CommentSimilar Species: Calamaria yunnanensis CHERNOV, 1962. Most specimens of C. yunnanensis can be distinguished from C. nebulosa sp. nov. by having a distinct nuchal color posterior to the head and a dark-brown midventral stripe under the surface of the tail (vs. nuchal collar and midventral stripe absent) and by having dark-brown margins on each of the ventral scales (vs. ventral scales immaculate). However, some specimens of C. yunnanensis do not share these color pattern traits. In specimens where the coloration is difficult to interpret in preservative or is variable, C. nebulosa sp. nov. can be differentiated from C. yunnanensis by having 179 ventrals in female (vs. 199), subcaudal/total body scale ratio 10.9% in female (vs. 8.7%), tail length approximately 7.9% of total length in female (vs. 5.0%), dorsal scales reducing to six rows above the 12th subcaudal (vs. 3rd to 5th subcaudal) and reducing to four rows above the last subcaudal (vs. third-to-last subcaudal). All specimens of C. yunnanensis have narrow dark-brown longitudinal stripes present on the margins of each dorsal scale row, occasionally restricted to the flanks in some specimens (vs. five, wider dark-brown longitudinal stripes, not present on every margin of each dorsal scale row), but these characters can be difficult to interpret on some specimens and should be treated with some wariness. (Lee, 2021).

Distribution: Presumably has a wider distribution extending across northern Laos, northern Thailand, southern Yunnan Province, China and perhaps also Vietnam.

Conservation: Recommended IUCN status: Data Deficient fide. Lee, 2021. 
EtymologyNamed after the nominative form the word “nebulous”, meaning “misty, foggy or cloudy” in Latin language and is given in feminine form to match the female genitive declension of the genus name Calamaria. The species name is an allusion to both the type locality of this snake, situated in the mountainous regions of Laos, and the generally clouded appearance of its dorsal ground color in preservative. 
References
  • Inger, R. F. & H. MARX 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana: Zoology 49: 1-304. - get paper here
  • Lee, Justin L. 2021. Description of a new species of Southeast Asian reed snake from northern Laos (Squamata: Colubridae: Genus Calamaria F. Boie, 1827) with a revised diagnosis of Calamaria yunnanensis Chernov, 1962. Journal of Natural History, 55:9-10, 531-560, - get paper here
  • Marx, H. & R. F. INGER 1955. Notes on the snakes of the genus Calamaria. Fieldiana: Zoology 37: 167 - 209 - get paper here
  • Quah, Evan S H; Shahrul Anuar, Lee L Grismer, Perry L Wood, Siti Azizah Mohd Nor 2019. Systematics and natural history of mountain reed snakes (genus Macrocalamus; Calamariinae). Zoological Journal of the Linnean Society, 188: 1236 - 1276 - get paper here
  • Stuart, B.L. & Heatwole, H. 2008. Country records of snakes from Laos. Hamadryad 33: 97–106 - get paper here
  • Weinell, J. L., Leviton, A. E., & Brown, R. M. 2021. A new species of reed snake, genus Calamaria (Colubridae: Calamariinae), from Mindoro Island, Philippines. Philippine Journal of Systematic Biology, DOI 10.26757/pjsb2020b14006 - get paper here
 
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