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Asthenodipsas ingeri (QUAH, LIM & GRISMER, 2021)

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Higher TaxaPareidae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Common NamesE: Inger’s Slug Snake 
SynonymAsthenodipsas ingeri QUAH, LIM & GRISMER 2021
Amblycephalus vertebralis – LOVERIDGE 1938: 43
Asthenodipsas vertebralis – STUEBING et al. 2014: 82 & 86 (part)
Pareas vertebralis – COX et al. 1998: 79 (part)
Pareas vertebralis – DE HAAS 1950: 529 (part)
Pareas vertebralis – HAILE 1958: 759, 766 (part)
Pareas vertebralis – ISKANDAR & COLIJN “2001” 2002: 113 (part)
Pareas vertebralis – MANTHEY & GROSSMANN 1997: 308, 378 (part)
Pareas vertebralis – STUEBING 1991: 331 (part)
Pareas vertebralis – STUEBING & INGER 1999: 87 (part)
Pareas vertebralis – INGER & VORIS 2001: 890 (part)
Amblycephalus laevis – SMITH 1925: 20 (part)
Asthenodipsas laevis – GROSSMANN & TILLACK 2003: 180 (part)
Asthenodipsas laevis – STUEBING et al. 2014: 82 (part)
Internatus laevis – MALKMUS et al. 2002: 342 (part)
Pareas laevis – STUEBING & INGER 1999: 87 (part)
Pareas carinatus – DAS 2012: 132 (part)
Pareas carinatus – DAS 2018: 132 (part) 
DistributionEast Malaysia (Borneo: Sabah and possibly Sarawak)

Type locality: Lumu Lumu, Mount Kinabalu, Sabah, East Malaysia (estimated: N 6.01966, E 116.539501, 1728 m a.s.l.).  
ReproductionOviparous (oresumably, Quah et al. 2021) 
TypesHolotype: Adult female MCZ R43592, collected by John A. Griswold Jr. in July 1937 
DiagnosisDiagnosis: Asthenodipsas ingeri sp. nov. can be differentiated from its congeners by the following combination of characters: a maximum SVL of 715 mm; 15/15/15 dorsal scale rows; 201 ventrals; 53 subcaudal scales; one or two postoculars; 2+2+2 temporals; seven supralabials, 3rd,4th & 5th (sometimes 3rd & 4th) touching the eye; six to seven infralabials, 1st and 3rd pair in contact; single anterior inframaxillary present; two pairs of posterior inframaxillaries; a sharp vertebral keel; dorsum of adults brown to grey and overlain with up to 51 irregularly-shaped, rhomboidal dark-brown bands beginning on the nape and extending the length of body and tail and onto the lateral edges of the ventral scales to form spots, but not encircling body; a narrow, light-coloured vertebral stripe; throat and ventrals white to cream-coloured with very fine speckling; top of head and snout darker than ground colouration of body while the supralabials and temporal region are lighter in colour; and iris and pupils black (Table 2 & 3; fig. 2, 4 & 5 in Quah et al. 2021).

Comparisons: Asthenodipsas ingeri sp. nov. can be differentiated from A. laevis, A. borneensis, A. malaccana, A. jamilinaisi and A. stuebingi by its higher number of ventrals (185–201 vs. 148–179) and the pairs of infralabials in contact ( 1st & 3rd vs. 2nd or 3rd). It can be further separated from A. laevis by its number of dorsal scale rows (15/15/15 vs. 15/15/13) and the sharp vertebral keel (present vs. absent) (Quah et al. 2019, 2020). From A. vertebralis, A. tropidonotus and A. lasgalenensis, the new species can be distinguished by the pairs of infralabials in contact (1st & 3rd vs. 1st) and fewer pairs of posterior inframaxillaries (two vs three) (fig. 5). Asthenodipsas ingeri sp. nov. can be further distinguished from A. vertebralis, A. tropidonotus and A. lasgalenensis by the number of supralabials in contact with the orbit (3–5 [rarely 3 & 4] vs. 3 & 4 [rarely 3–5 or only 3 or 4]) (fig. 4 & 6B), and having fewer subcaudals (53–57 vs. 57–79). In addition, A. ingeri sp. nov. can be further distinguished from A. tropidonotus by its higher number of postoculars (2 vs. 1), and from A. lasgalenensis by its dorsal colouration in adults (light-brown with dark bands and prominent vertebral stripe vs. solid dark-brown to black and no vertebral stripe) (fig. 6). In life A. ingeri sp. nov. can also be distinguished from A. vertebralis, A. lasgalenensis and A. tropidonotus by the colouration of its iris (dark brown vs. orange to reddish-brown) (fig. 2C–f, 4B–D & 6) (Table 3; Loredo et al. 2013; iNaturalist) (Table 4 in Quah et al. 2021).

Color in life: Based on photographs of other specimens from Mount Kinabalu, Sabah (fig. 2C–E) and Payeh Maga Highlands, Sarawak (fig. 2f) that are tentatively identified here as A. ingeri sp. nov. the dorsal ground colour in life ranges from beige to light-grey with more prominent dark banding on the body and the venter ranges from white to cream-coloured. The thin vertebral stripe is cream-coloured to light orange (Quah et al. 2021). 
CommentDiet: Presumably feeds on snails like other congeners (Quah et al. 2021)

Presumably lays eggs (Quah et al. 2021) 
EtymologyThe specific epithet ingeri is in honour of Robert f. Inger (10 September 1920–12 April 2019) for his extensive work on the reptiles and amphibians of Borneo and his overall contributions to the field of herpetology throughout Asia for nearly 80 years. 
  • Cox, Merel J.; Van Dijk, Peter Paul; Jarujin Nabhitabhata & Thirakhupt,Kumthorn 1998. A Photographic Guide to Snakes and Other Reptiles of Peninsular Malaysia, Singapore and Thailand. Ralph Curtis Publishing, 144 pp.
  • Das, I. 2012. A Naturalist's Guide to the Snakes of South-East Asia: Malaysia, Singapore, Thailand, Myanmar, Borneo, Sumatra, Java and Bali. Oxford J, ohn Beaufoy Publishing - get paper here
  • Das, I. 2018. A Naturalist’s Guide to the Snakes of Thailand and South-east Asia. 2nd edition. John Beaufoy Publishing, Oxford, 176 pp
  • 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
  • HAAS, C.P.J. De 1950. Checklist of the snakes of the Indo-Australian archipelago. Treubia 20 (3): 511-625 - get paper here
  • Haile, N.S. 1958. The snakes of Borneo, with a key to the species. Sarawak Mus., Kuching, J. 8: 743-771.
  • Inger, R.F. & Voris, H. K. 2001. The biogeographical relations of the frogs and snakes of Sundaland. Journal of Biogeography 28: 863-89 1
  • Iskandar, D.T. & E. Colijn. 2001. Checklist of Southeast Asian and New Guinea Herpetofauna II Reptilia, Serpentes. Treubia 31:135–313 - get paper here
  • 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.; 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
  • 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
  • Smith, M. A. 1925. Contribution to the herpetology of Borneo. Sarawak Mus. J. 3 (8): 15-34. - 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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