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Boiga multomaculata (BOIE, 1827)

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Higher TaxaColubridae, Colubrinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesBoiga multomaculata multomaculata (BOIE 1827)
Boiga multomaculata ochracea (THEOBALD 1868)
Boiga multomaculata septentrionalis KÖHLER et al. 2023 
Common NamesE: Many-spotted Cat Snake, Ocellated Asian Cat snake
G: Gefleckte Katzennatter, Vielflecken-Nachtbaumnatter
Chinese: 繁花林蛇 
SynonymDipsas multomaculata BOIE 1827
Dipsas multomaculata REINWARDT 1827
Dipsas multimaculata [sic] — DUMÉRIL & BIBRON 1854: 1139
Dipsas multimaculata — THEOBALD 1868: 53
Dipsadomorphus multimaculatus — BOULENGER 1896
Boiga multimaculata SMITH 1923
Boiga multimaculata hainanensis MELL 1931
Boiga multimaculata sikiangensis MELL 1931
Boiga multimaculata — SMITH 1943
Boiga multomaculata — MANTHEY & GROSSMANN 1997: 323
Boiga multomaculata — COX et al. 1998: 76
Boiga multomaculata — GROEN 2008
Boiga multomaculata — COX et al. 2012: 122
Boiga multomaculata — WALLACH et al. 2014: 105
Boiga multomaculata — WANG et al. 2022
Boiga multomaculata multomaculata — KÖHLER et al. 2023: 170

Boiga multomaculata ochracea (THEOBALD 1868)
Dipsas ochraceus THEOBALD 1868: 53
Dipsas ochraceus — GÜNTHER 1868: 425
Dipsas hexogonatus STOLICZKA 1871 (non BLYTH)
Dipsadomorphus hexogonotus — BOULENGER 1896 (non BLYTH)
Dipsadomorphus hexagonotus — WALL 1909: 154
Boiga ochracea — SMITH 1943: 348
Boiga ochracea walli SMITH 1943
Boiga ochraceus — DAS 1996: 54
Boiga ochracea cf. walli — MAHONY & ALI REZA 2007
Boiga ochraceus — GROEN 2008
Boiga ochracea walli — MAHONY et al. 2009
Boiga walli — WALLACH et al. 2014: 106
Boiga ochracea — WALLACH et al. 2014: 106
Boiga multomaculata ochracea — KÖHLER et al. 2023

Boiga multomaculata septentrionalis KÖHLER et al. 2023: 177
Boiga multomaculata indica MELL 1931: 213 [partim]
Boiga ochracea walli SMITH 1943: 349 [partim] 
DistributionW Malaysia, Cambodia, Thailand, Vietnam (Hoa Binh),
Myanmar (= Burma),
India (Assam, Arunachal Pradesh, Nagaland),
S China (Fujian, incl. Hong Kong and Hainan)
Indonesia (Java, Sumatra, Bali);
Bangladesh; Hong Kong; Laos

Type locality: Java

septentrionalis: N Myanmar (Kachin state and Sagaing region), NE India (Assam and Nagaland), possibly S China (1 specimen, CAS 242550 from Yunnan Province).  
Reproductionoviparous 
TypesLectotype: RMNH.RENA 979b, an adult female from Java, Holotype unlocated (fide NGUYEN et al. 2009).
Holotype. CAS 241272, an adult male from Indawgyi lake wildlife Sanctuary (25.30347, 96.35417; 235 m a.s.l.), vicinity Kyang Kyar village, Moenyin township, Myitkyina District, Kachin State, Myanmar, collected on 20 july 2008 by j.A. wilkinson, j.v. vindum, S.l. Oo, K.t. Kyaw, and M. win. Original field tag MhS 25921. Paratypes (3). CAS 241150, an adult female from Indawgyi lake wildlife Sanctuary (25.30347, 96.35417; 235 m a.s.l.), E of Nat Mouk Kan village, Moenyin township, Myitkyina District, Kachin State, Myanmar, collected on 14 july 2008 by G.O.u. wogan, j.A. wilkinson, j.v. vindum, A.K. Shein. CAS 241550, an adult female from Indawgyi lake wildlife Sanctuary (25.17869, 96.29156; 270 m a.s.l.), vicinity of lwe Mon village, Moenyin township, Myitkyina District, Kachin State, Myanmar, collected on 26 july 2008 by j.A. wilkinson, K.t. Kyaw, and j.v. vindum. SMF 106288 (formerly CAS 239222), an adult male from linpha village (25,80489, 95,52667; 140 m a.s.l.), hkanti township, hkanti District, Sagaing region, Myanmar, collected on 10 September 2006 by K.S. lwin, S.l. Oo, and A.K. Shein. [septentrionalis]
Neotype. BMNH 1946.1.2.60 (formerly BMNH 68.4.3.16), an adult female from “Pegu” [Bago, Bago region, Myanmar, approx. 17.32°N, 96.47°E], collected by william theobald and donated to the BMNh collection in spring 1868, designated by Köhler et al. 2023. Original type lost. [ochracea]
Syntypes: BMNH 1946.1.2.60–61, collected by W. Theobald [walli] 
DiagnosisDiagnosis: Taylor 1965: 869

Description: Taylor 1965: 869

Description of the lectotype. Adult female, indicated by the absence of hemipenes; 1/1 loreal, wider than high; nasal scale completely divided; 1/1 preocular; 1/1 supraocular; 2/2 postoculars, upper not reaching onto top of head; 2 prefrontals; 2/1 anterior and 2/2 posterior temporals; supralabials 8/8, 3rd–5th supralabials entering eye; 11/11 infralabials, first four in contact with anterior chin shields; dorsal scales in 19-19-15 rows, smooth with single tiny apical pits on body and single or paired apical pits on dorsocaudal scales; vertebral scale row significantly enlarged; no preventral; 214 ventrals; cloacal plate entire; 89 paired subcaudal scales. Body slender; tail long (tl/Svl 0.243); Svl 592 mm; tl 144 mm, head length measured from tip of snout to posterior border of parietals 14.2 mm, head length measured from tip of snout to posterior edge of mandible 18.8 mm, head width 11.1 mm; diameter eye 3.3 mm; distance anterior border eye to tip of snout 5.2 mm. (Köhler et al. 2023)

Dentition. Maxillary bone with 11/11 prediastemal teeth, followed by a distinct diastema which is 65% longer than the socket of the last prediastemal tooth and followed by two distinctly enlarged, grooved and posteriorly bent postdiastemal teeth. Prediastemal teeth slightly decrease in size posteriorly, the anterior two distinctly posteriorly hooked, the following with less pronounced curvature. On the left side, prediastemal teeth number five, seven, nine, 11, and second postdiastemal loose. On the right side, prediastemal teeth two, four, five, and second postdiastemal tooth loose. Medial to each maxillary tooth is a single replacement tooth at different growth stages.
Palatine bone with 6/6 posteriorly curved teeth, shorter that the prediastemal teeth and slightly decreasing in size posteriorly. tooth one broken, teeth two, four and six loose on left side. tooth three broken, teeth two, four and six loose on the right side. lateral to each palatine tooth is a single replacement tooth at different growth stages. Pterygoid bone with 12/12 posteriorly curved teeth, shorter than the palatine tooth, gradually decreasing in size posteriorly. teeth one, three, four, five and 12 loose on left side. teeth three, four, five and nine loose on right side. the posterior 60% of the pterygoid bone are without teeth.
Mandibular bone with 17/17 posteriorly curved teeth, shorter than maxillary teeth, gradually decreasing in size posteriorly. Medial to each mandibular tooth is a single replacement tooth in different growth stages. teeth two, four and six loose on left side. teeth two, four, six, eight and 10 loose on right side. (Köhler et al. 2023)

Coloration: after approximately 200 years preservation in ca. 70% ethanol was recorded as follows: Dorsal ground color tawny Olive (Color 17) with Natal Brown (49) mottling on scales and with 62 dorsolateral pairs of Sepia (279) colored blotches on body that have a paler (Amber 51 suffused with Smoke Gray 266) center and a white to Pale Buff (1) border; dorsolateral body blotches separated by a Pale Buff (1) vertebral interspace; below the dorsolateral row of blotches an alternating row of smaller, irregularly shaped blotches with the same coloration; dorsal head with same ground color as dorsal body and a v-shaped Sepia (279) marking open towards the neck, starting from the inner posterior edges of the internasals and ending at the level of the last third of the last supralabial; an oval marking on the neck partly inside the bifurcation of the v-shaped element and with the same coloration as described for the body blotches; a Sepia (279) colored postocular stripe ending at the angle of the mouth; edges of posterior upper labials and edges of infralabials Sepia (279). venter Cream Color (12) with tawny Olive (17) colored alternating irregular shaped spots; ventral head Cream Color (12) without prominent markings. (Köhler et al. 2023)

Dorsal scale reduction formula: Köhler et al. 2023: 172.

Variation. Paralectotypes and additional examined material agree well with the lectotype in general appearance, morphometrics and scalation (table 1).
variation in dentition. Nine to 13 prediastemal teeth followed by a distinct diastema and two enlarged grooved postdiastemal teeth; six to seven palatine, eight to 12 pterygoid, and 17 to 19 dental teeth.
variation in live coloration and pattern. Only a blotched morph is known (Fig. 10). the dorsal ground color can vary from Pale Neutral Gray (color 296) to true Cinnamon (260), blotches on body Dark Grayish Brown (284) to Sepia (286), edged Smoky white (261) or Cream white (52). Iris may vary, in accordance with the dorsal ground color, from Pale Neutral Gray (color 296) to Medium Neutral Grey (299) or from Orange-rufous (56) to true Cinnamon (260), with black Sepia (286) pupil. ventral head and throat Smoky white (261), rest of venter can vary from Pale Neutral Gray (296) to Cream white (52) with small Pale Neutral Grey (296) or Cinnamon (255) spots. (Köhler et al. 2023)

Diagnosis (septentrionalis). A subspecies of Boiga multomaculata that differs from the other two subspecies (i.e., B. m. multomaculata and B. m. ochracea) by mean pairwise genetic distances of 3.3–5.8% (ND4) and 3.7–6.5% (CYtB), respectively. It further differs from B. m. multomaculata by having on average more ventral scales in both sexes (males 227.7±5.19, females 228.8±1.71 in B. m. septentrionalis versus males 214.4±8.06, females 217.1±9.50 in B. m. multomaculata), by having on average more subcaudal scales in both sexes (males 99.1±5.82, females 97.3±2.06 in B. m. septentrionalis versus males 94.5±5.33, females 89.8±4.13 in B. m. multomaculata), and by having both blotched and patternless individuals (versus only blotched specimens in Boiga m. multomaculata). It further differs from B. m. ochracea by having on average fewer ventral scales in females (228.8±1.71 in B. m. septentrionalis versus 238.9±6.30 in B. m. ochracea). (Köhler et al. 2023)

Description of the holotype (septentrionalis): Adult male, indicated by fully developed everted hemipenes; 1/1 loreal, wider than high; nasal scale partly divided below the naris; 1/1 preocular; 1/1 supraocular; 2/2 postoculars; 2 prefrontals; 2/2 anterior and 3/2 posterior temporals; supralabials 8/8, 3rd–5th supralabials entering eye; 11/11 infralabials, first five in contact with anterior chin shields; dorsal scales in 19-19-15 rows, smooth with single or paired apical pits on body and single to triple apical pits on dorsocaudal scales; vertebral scale row distinctly enlarged; two preventrals and 226 ventrals; cloacal plate entire; subcaudal scales 92, paired. Body slender; tail long (tl/Svl 0.235); Svl 578 mm; tl 136 mm; head length measured from tip of snout to posterior edge of mandible 18.1 mm, head width 10.5 mm; diameter eye 3.4 mm; distance anterior border eye to tip of snout 4.9 mm. Figs. 14 and 15,

Dentition. Maxillary bone with 11/11 prediastemal teeth, followed by a distinct diastema which is 40% longer than the socket of the last prediastemal tooth and followed by two distinctly enlarged, grooved and posteriorly bent postdiastemal teeth. Prediastemal teeth slightly decrease in size posteriorly, the anterior three distinctly posteriorly hooked, the following with less pronounced curvature. On the left side, prediastemal teeth number two, four, six, eight and ten, and first postdiastemal loose. On the right side, prediastemal teeth one, three, five, seven and nine, and first postdiastemal tooth loose. Medial to each maxillary tooth is a single replacement tooth at different growth stages.
Palatine bone with 6/6 posteriorly strongly curved teeth, shorter that the prediastemal teeth and slightly decreasing in size posteriorly. tooth three six loose on left side. teeth two, four and six loose on the right side. lateral to each palatine tooth is a single replacement tooth at different growth stages.
Pterygoid bone with 11/12 posteriorly curved teeth, shorter than the palatine tooth, gradually decreasing in size posteriorly. teeth two, five, seven and 10 loose on left side. teeth one, three, five, 10 and 12 loose on right side. the posterior 63% of the pterygoid bone are without teeth.
Mandibular bone with 18/17 posteriorly curved teeth, shorter than maxillary teeth, gradually decreasing in size posteriorly. Medial to each mandibular tooth is a single replacement tooth in different growth stages. teeth one, three, five, seven, nine, 11, 13 and 15 loose on left and right side. tooth 17 missing on right side.

Coloration: after about 13 years preservation in 70% ethanol was recorded as follows: Dorsal ground color Smoke Gray (Color 266) with Natal Brown (49) mottling on scales and with 62 Sepia (286) blotches that have a paler (Grayish Olive 274 suffused with Smoke Gray 266) center and a white to Pale Buff (1) border.

Dorsal scale reduction formula:

Variation. Paratypes agree well with the holotype in general appearance, morphometrics and scalation (table 1). Most paratypes are blotched as is the holotype. An exception is SMF 106288 (formerly CAS 239222) that has no blotches but exhibits 58 indistinct transverse dark bars composed of dark edges of adjacent dorsal scales.

Variation in dentition. ten to 12 prediastemal teeth followed by a distinct diastema and two enlarged grooved postdiastemal teeth; five to six palatine, seven to 12 pterygoid, and 16 to 19 dental teeth.
variation in live coloration and pattern. two color morphs, blotched and unicolored are known. the blotched morph resembles the color and pattern of brownish individuals of the nominate subspecies. the unicolored morph (Fig. 16) resembles the unpatterned morph of B. m. ochracea.
An intermediate color variation is known from Nagaland, NE India (wII-AD 856), which resembles the ground coloration of the unicolored morph but shows a raw umber (280) postocular stripe, faint dark dorsal had pattern similar to that of the nominate subspecies and an irregular strippled raw umber (280) pattern forming broken bands along dorsal body and tail.

 
CommentSynonymy: B. multomaculata and B. ochracea are closely related any may be conspecific (Weinell et al. 2020, 2021).

SMITH 1943 gave “1929” for MELL’s paper. Not listed by GRANDISON 1977.

Distribution: May have been erroneously reported from Borneo, but not listed for Borneo by MALKMUS et al. (2002). There are two records from Sulawesi which are from before 1900 and are thus questionable (DE LANG & VOGEL 2005, Koch 2012). Possibly in Bhutan (Lenz 2012). Not in Singapore or not established there (Figueroa et al. 2023).

Behavior: nocturnal.

Habitat: fully arboreal (Harrington et al. 2018). 
EtymologyNamed after Latin macula = spot, for its multitude of spots.

The subspecies epithet is formed from the latin words septentrio (= “north”) and the suffix -alis (to form an adjective) and refers to the geographic distribution of this taxon. Boiga m. septentrionalis has the most northern distribution of the subspecies of B. multomaculata. 
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