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Asthenodipsas stuebingi QUAH, GRISMER, LIM, ANUAR & IMBUN, 2019

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Higher TaxaPareidae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Common NamesE: Sabah Mountain Smooth Slug Snake 
SynonymAsthenodipsas stuebingi QUAH, GRISMER, LIM, ANUAR & IMBUN 2019
Amblycephalus laevis — BOULENGER 1896: 441 & 442 (in part)
Amblycephalus laevis — DE ROOIJ 1917: 276
Amblycephalus vertebralis — LOVERIDGE 1938: 43 (in part)
Asthenodipsas laevis — DAS 2006: 007 (in part)
Asthenodipsas laevis — DAS 2010: 344
Asthenodipsas laevis — GROSSMANN & TILLACK 2003: 188
Internatus laevis — MALKMUS et al. 2002: 342 & 343, Figs, 337 & 338
Internatus laevis — STUEBING et al. 2014: 85
Pareas laevis — MALKMUS 1996: 293
Pareas laevis — MANTHEY & GROSSMANN 1997: 377, Abb. 284
Pareas laevis — STUEBING 1991: 330 & 331 (in part)
Pareas laevis — STUEBING & INGER 1999: 87 (in part)
Pareas laevis — HAILE 1958: 766 (in part)
Asthenodipsas (Asthenodipsas) stuebingi — POYARKOV et al. 2022 
DistributionEast Malaysia (Borneo: Sabah)

Type locality: Minduk Sirung (Alab–Mahua trail), Crocker Range Park, Sabah, East Malaysia (estimated: N 5.823240, E 116.347238, 1859 m a.s.l.).  
TypesHolotype. SP 04679, adult female collected by Paul Yambun Imbun, Fred Tuh Yit Yu and Safrie on 25 March 2009. Paratypes. Adult male (SP 04806) collected by Siti Azizah Kusop, Junaidi Ais and R., Adzmie on 26 October 2014 from Kampung Desa Aman, Kundasang (estimated: N5.988087, E 116.570162, 1341 m a.s.l.). Adult female (ZMB 65429) collected by Rudolf Malkmus and Andreas Nӧllert in August 1994 on Kamborangoh Road, Mount Kinabalu, Sabah, West Malaysia (estimated: N6.0292, E116.5473, 1854 m a.s.l.). Adult male (MCZ R43591) col- lected by John A. Griswold Jr. in 1937 from Kadmayan R, near Kiau, Sabah, West Malaysia (estimated: N6.029788, E 116.489732, 971 m a.s.l.). 
DiagnosisDiagnosis. Morphological analyses of the four specimens from the highlands of Sabah confirms the placement of this population in the genus Asthenodipsas based on the combination of the following characters: smooth dorsal scales; absence of preoculars and suboculars; one or more supralabials in contact with the eye; a single anterior inframaxillary, followed by two or three pairs of inframaxillaries which are wider than long (Grossmann & Tillack 2003). Asthenodipsas stuebingi sp. nov. can be differentiated from its congeners by the following combination of characters: a maximum SVL of 557 mm; 15/15/15 dorsal scale rows; 165–175 ventrals; 35–47 subcaudal scales; one or two postoculars; 2(+0–1)+2 temporals; six supralabials, 3rd & 4th touching the eye; 4–6 infralabials, 2nd or 3rd pair in contact; a sharp vertebral keel; dorsum of adults beige to brown and overlain with 30–42 irregularly-shaped, rhomboidal dark brown bands beginning posterior to dark patch on the neck that and extends the length of body and tail and onto lateral edges of the ventral scales to form spots, but not encircling body; a narrow, light-coloured vertebral stripe; throat and ventrals beige to yellow and mottled with small, dark spots; head whitish to light-brown with dark speckling on the snout and crown; and iris and pupils black (Table 4, 5 & 6; Fig. 2 & 3 in Quah et al. 2019).

Comparison. Asthenodipsas stuebingi sp. nov. can be differentiated from Aplopeltura boa by its higher number of mid-dorsal scale rows (15 vs. 13) and divided subcaudals (de Rooij 1917; Grossmann & Tillack 2003; Stuebing et al. 2014). Asthenodipsas stuebingi sp. nov. can be differentiated from members of the genus Pareas by their possession of preocular and subocular scales (absent vs. present), supralabials in contact with orbit (3rd & 4th contact orbit vs. no supralabials in contact with orbit) and anterior single inframaxillary (present vs. absent) (Grossmann & Tillack 2003). Asthenodipsas lasgalenensis, A. tropidonotus and A. vertebralis can be differentiated from the new species by their higher number subcaudals (54–77 vs. 35–47), pairs of infralabials in contact (1st vs. 3rd) and more pairs of posterior inframaxillaries (three vs. two). In addition, A. vertebralis and A. tropidonotus can be differentiated from A. stuebingi sp. nov. by their higher number of ventrals (195–215 vs. 165–175). Adult A. lasgalensis can also be differentiated from A. stuebingi sp. nov. by their dorsal colour pattern that is solid dark brown to black and their whitish labials versus a beige to brown dorsum with dark-bands and a dark patch on the neck in the latter (Loredo et al. 2013). From Bornean populations of A. malaccana to which it is most similar in colour pattern, A. stuebingi sp. nov. can be differentiated by a lower number of supralabials (6 vs. 7–8) (Chan-ard et al. 2015; Das 2010; de Rooij 1917; Stuebing et al. 2014). From A. laevis to which it was previously confused, A. stuebingi sp. nov. can be differentiated by its larger adult length, (Max SVL 557mm vs. 373mm), dorsal scales rows (15/15/15 vs. 15/15/13) and sharp vertebral keel (present vs. absent) (Figs. 3F & 4F) (Tables 2–6). A. stuenbingi sp. nov. can be differentiated from A. jamilinaisi sp. nov. by its lower number of subcaudals (35–47 vs. 52–53), lower number of ventrals in males (165 vs. 173–175), size of vertebral scales (slightly enlarged vs. greatly enlarged), colour pattern (light-coloured head and dorsum with a dark neck patch and distinct bands vs. dark overall colouration of dorsum with muted banding) and body form (robust and stout vs. gracile and laterally compressed) (Tables 4–6). A key to the family Pareidae of Borneo is presented below. 
CommentHabitat: montane, only been found on the highlands of Sabah from 900–2000 m a.s.l.. This is in contrast to its congener A. laevis that is widely distributed in Borneo and recorded from many lowland localities up to 1150 m in elevation (Stuebing et al. 2014). At some locations such as the Crocker Range, both species occur (SP 04476, SP 04570 & SP 04679; see Table 5 in Quah et al. 2019). It is reportedly both terrestrial and arboreal (Malkmus et al. 2002). Malkmus et al. (2002) reported finding a specimen crossing the road at night in rainy weather and another specimen in a bush approximately 1m above the ground. The paratype ZMB 65429 was also collected at night fol- lowing light rains at approximately 22:00 h. A specimen was observed at 09:00 h close the power station of the Kinabalu park headquarters along Kamborangoh road (Andreas Nӧllert in litt. 2018).

Behavior: Similar to other species of slug snakes, A. stuebingi sp. nov. is nocturnal. When threatened, the snakes roll themselves up into tight coils and remain motionless — a behaviour observed in Aplopeltura boa (Jablonski & Hegner 2016).

Diet: Asthenodipsas stuebingi sp. nov. presumably feeds on snails or slugs like other members of the family. 
EtymologyThe specific epithet stuebingi is a patronym in honour of Robert B. Stuebing (born 1946) for his contributions to the field of herpetology in Borneo, especially on the study of snakes. 
  • Boulenger, G.A. 1896. Catalogue of the snakes in the British Museum, Vol. 3. London (Taylor & Francis), xiv + 727 pp. - get paper here
  • Das, I. 2010. A Field Guide to the Reptiles of Southeast Asia. New Holland, London. 376 pp. [book review in Sauria 33 (3): 51, and Russ. J. Herp. 18: 325] - get paper here
  • Das, Indraneil 2006. Crocker Range National Park, Sabah, as a refuge for Borneo's montane herpetofauna. Amphibian & Reptile Conservation 4 (1): 3-11 - get paper here
  • de Rooij, N. DE 1917. The Reptiles of the Indo-Australian Archipelago. Il. Ophidia. Leiden (E. J. Brill), xiv + 334 S. - get paper here
  • Grossmann, W. & Tillack, F. 2003. On the taxonomic status of Asthenodipsas tropidonotus (LIDTH DE JEUDE 1923) and Pareas vertebralis BOULENGER 1900) (Serpentes: Colubridae: Pareatinae). Russ. J. Herpetol. 10 (3): 175-190 - get paper here
  • Haile, N.S. 1958. The snakes of Borneo, with a key to the species. Sarawak Mus., Kuching, J. 8: 743-771.
  • Loveridge, A. 1938. New snakes of the genera Calamaria, Bungarus and Trimeresurus from Mt. Kinabalu, North Borneo. Proc. Biol. Soc. Washington 51: 43-46. - get paper here
  • Malkmus, R. 1996. Herpetologische Beobachtungen am Mount Kinabalu, Nord-Borneo. V. Mitteilungen aus dem Zoologischen Museum in Berlin, 72 (2): 277–295 - get paper here
  • Malkmus, R.; Manthey, U.; Vogel, G. Hoffmann, P. & Kosuch, J. 2002. Amphibians and reptiles of Mount Kinabalu (North Borneo). A.R.G. Ganther Verlag, Rugell, 404 pp.
  • Manthey, U. & Grossmann, W. 1997. Amphibien & Reptilien Südostasiens. Natur und Tier Verlag (Münster), 512 pp. - get paper here
  • Poyarkov NA, Nguyen TV, Pawangkhanant P, Yushchenko PV, Brakels P, Nguyen LH, Nguyen HN, Suwannapoom C, Orlov N, Vogel G. 2022. An integrative taxonomic revision of slug-eating snakes (Squamata: Pareidae: Pareineae) reveals unprecedented diversity in Indochina. PeerJ 10: e12713 - get paper here
  • Quah, Evan S H; Kelvin K P Lim, L Lee Grismer 2021. On the taxonomic status of Asthenodipsas vertebralis (Boulenger, 1900) (Squamata: Pareidae) in Borneo with the description of a new species. Zootaxa 4949 (1): 24–44
  • QUAH, EVAN S.H.; L. LEE GRISMER, KELVIN K.P. LIM, M.S. SHAHRUL ANUAR, & PAUL Y. IMBUN 2019. A taxonomic reappraisal of the Smooth Slug Snake Asthenodipsas laevis (Boie, 1827) (Squamata: Pareidae) in Borneo with the description of two new species. Zootaxa 4646 (3): 501–526 - get paper here
  • QUAH, EVAN S.H.; L. LEE GRISMER, KELVIN K.P. LIM, M.S. SHAHRUL ANUAR, KIN ONN CHAN 2020. A taxonomic revision of Asthenodipsas malaccana Peters, 1864 (Squamata: Pareidae) with a description of a new species from Borneo. Zootaxa 4729 (1): 1-24 - get paper here
  • Stuebing, R.B. 1991. A checklist of the snakes of Borneo. Raffles Bull. Zool. 39(2): 323-362. - get paper here
  • Stuebing, R.B. & INGER, R.F. 1999. A field guide to the snakes of Borneo. Natural history Publications (Borneo), Kota Kinabalu, 254 pp. [corrections in HR 31: 124]. - get paper here
  • Stuebing, R.B., Inger, R.F. & Lardner, B. 2014. A field guide to the snakes of Borneo, second edition. Natural history Publications (Borneo)
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