42 Australasian Journal of Herpetology Australasian Journal of Herpetology 33:42-51. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online)

Not a monotypic genus! Aplopeltura boa (Boie, 1828) divided!

RAYMOND T. HOSER

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 5 January 2016 Accepted 24 July 2016, Published 1 August 2016.

ABSTRACT Until now the Blunt-headed Slug Eating from South-east Asia (Family: Pareidae), has been treated as being of a single species, namely Aplopeltura boa (Boie, 1828). However inspection of specimens from across the known range, shows that they are sufficiently divergent from one another in geographically separated populations to warrant being named as separate species. Available distributional and molecular evidence supports this contention. Hence the previously monotypic genus Aplopeltura Duméril, 1853 is herein divided into six well-defined species, five formally named for the first time according to the International Code of Zoological Nomenclature (Ride et al. 1999). Another species within the Pareidae, namely carinatus (Boie, 1828) is known to have divergent populations, which have been variously described as species and subspecies. Recognizing the three previously named forms (including the nominate one) herein as subspecies, two more obviously unnamed geographically separated subspecies are also formally named for the first time. Keywords: ; ; South-east Asia; Asia; Aplopeltura; boa; new species; shireenae; omarelhelou; lynnejohnstoneae; daranini; gibbonsi; new subspecies; sumatrensis; malayensis.

INTRODUCTION Until now the Blunt-headed Slug Eating Snake from south-east this sooner, rather than later, so as not to jeopardize their very Asia (Family: Pareidae), has been treated as being of a single existence. widespread species, namely Aplopeltura boa (Boie, 1828). I also note the abysmal environmental record of governments However inspection of specimens by myself over some years worldwide in the past 200 years as detailed by Hoser (1989, from across the known range, shows that they are sufficiently 1991, 1993 and 1996). divergent from one another in geographically separated Noting that publicly available distributional and molecular populations to warrant being named as separate species. evidence supports the contention that there is more than one Significant data was obtained from across the range of the species under the label Aplopeltura boa, I have now made the putative species, but this was stolen in an illegal armed raid on decision to formally name the most obviously divergent groups my facility on 17 August 2011 by corrupt wildlife officers seeking in accordance with the International Code of Zoological to permenantly disable my successful wildlife education Nomenclature (Ride et al. 1999). business, “Snakebusters, Australia’s best shows” (Court Hence the previously monotypic genus Aplopeltura Duméril, of Appeal Victoria 2014, VCAT 2015). 1853 is herein divided into six well-defined species, five formally None of this material was returned. named for the first time. Shortly after this illegal armed raid, Hoser (2012a) did the I also note that relevant texts (e.g. Das 2012) speak of putative obvious step of dividing the related genus Pareas Wagler, 1830 Aplopeltura boa as being widely distributed throughout the as then known, along obvious morphological and phylogenetic South-east Asian realm. However a survey of museum holdings lines. Hoser (2012a) also erected a subgenus for another records suggests that the distribution is somewhat disjunct and divergent lineage. largely confined to hilly areas and those immediately proximate As the extensive data gathered relevant to the species-level to them. See for example the distribution data in David and taxonomy was not returned, this material was not published in Vogel (1996). 2012. Hence the concept that currently divided populations were most However with ongoing environmental distruction in the South- likely similarly divided during recent glacial maxima is the one I east Asian realm coupled with the fact that this is likely to get am subscribing to and in the absence of evidence to the worse rather than better in the foreseeable future, it is important contrary. that regional populations warranting taxonomic recognition get While putative Aplopeltura boa vary significantly in both

Hoser 2016 - Australasian Journal of Herpetology 33:42-51. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 43

colouration and scalation within a single locality, there are NOTES ON THE DESCRIPTIONS FOR ANY POTENTIAL differences that are consistent between locations and these are REVISERS used as the basis to diagnose each species. Unless mandated by the rules of the International Code of Another species within the family Pareidae, namely Pareas Zoological Nomenclature, none of the spellings of the newly carinatus (Boie, 1828) is known to have divergent populations, proposed names should be altered in any way. Should one or which have been variously described as species and more newly named taxa be merged by later authors to be subspecies. treated as a single species, the order of prority of retention of The best known of these is P. nuchalis (Boulenger, 1900), names should be the order as listed in the keywords part of the treated by authors in the past as either a full species, or a abstract. synonym of P. carinatus. APLOPELTURA SHIREENAE SP. NOV. While Hoser (2012a) treated P. nuchalis as a full species, in Holotype: A preserved specimen held at the Carnegie Museum accordance with the views of de Rooij (1917), Malkmus (1996), of Natural History, Pittsburgh, Pennsylvania, USA, specimen Malkmus et al. (2002) and evidence of Guo et al. (2011), this number: CM Herps R2427, collected from Agusan Province, was contradicted by the results of Pyron et al. (2013), that Mindanao, , Asia. The Carnegie Museum of Natural implied P. nuchalis was conspecific with P. carinatus. History, Pittsburgh, Pennsylvania, USA, allows access to its In light of this new evidence of Pyron et al. (2013) and that of holdings. Guo et al. (2011), I herein conservatively treat both P. nuchalis Paratypes: Two preserved specimens held at the Carnegie and the previously described subspecies of P. carinatus Museum of Natural History, Pittsburgh, Pennsylvania, USA, described as Amblycephalus carinatus unicolor Bourret, 1934 as specimen numbers: CM Herps R2428 and CM Herps R2429, subspecies of P. carinatus. collected from Agusan Province, Mindanao, Philippines, Asia. Recognizing the three previously named forms, including the nominate one herein as subspecies, two more obviously Diagnosis: Aplopeltura shireenae sp. nov. from the main unnamed geographically separated subspecies are also formally Philippine Islands, including Mindanao, are readily separated named for the first time. from all other Aplopeltura species by the following suite of characters: A generally very indistinct dorsal pattern, being a As for A. boa above, I also note that relevant texts (e.g. Das light reddish-brown in colour, the lateral white blotches rising 2012) speak of putative P. carinatus as being widely distributed from the belly on the lower flanks are so heavily shaded that throughout the South-east Asian realm. However a survey of they are barely noticeable and this includes for the anterior museum records suggests that the distribution is somewhat section of the body, which is only similarly seen in Aplopeltura disjunct (but wider than for A. boa) and largely confined to hilly lynnejohnstoneae sp. nov. from the lower Malay Peninsula. A. areas and those immediately proximate to them. See for shireenae sp. nov. is separated from A. lynnejohnstoneae sp. example the distribution data in David and Vogel (1996). nov. by several characters including a lack of melanism on the I also note that the basis or material and methods underpinning dorsal surface of the head (see in A. lynnejohnstoneae sp. nov.) the taxonomy herein has been an inspection of live and dead and a lack of the profuse blackish specking seen on the dorsal specimens, photos with good locality data and the relevant surface of A. lynnejohnstoneae sp. nov.. available literature that summarizes relevant facts about the The top of the head of A. shireenae sp. nov., while also reddish relevant taxa. brown in colour has two distinct white patches somewhat Museum records were audited to ascertain the extent of known anterior to the eyes, pointing towards the middle of the head, but populations, via collection records and localities. not meeting. The iris is brownish in colour and characterised by The results are of course the formal taxonomic proposals within the presence of whitish specks. this paper as outlined both above and in the descriptions below. Aplopeltura boa (Boie, 1828) from Java is readily separated from Important references relevant to Aplopeltura and the taxonomic others in the genus (as defined herein) by significant somewhat decisions within this paper include the following: Boie (1828), irregular shaped white patches along the dorsal midline. The Boulenger (1894, 1896), Chan-ard et al. (2015), Cox et al. dark patch below the eye that commences on the jawline, does (1998), Das (2012), David and Vogel (1996), de Rooij (1917), not extend as far as the eye or if so, only just intersects it. Dowling and Jenner (1998), Duméril (1853), Grandison (1978), Aplopeltura omarelhelou sp. nov. from southern Thailand on the Grismer et al. (2010), Grossmann and Tillack (2001), Hofmann Isthumus of Kra in the region generally north of and including (2015), Hoser (2012a), Kopstein (1938), Manthey and Songkhla and Trang are readily separated from those on the Grossmann (1997), Pyron et al. (2013), Schlegel (1837), Smith lower Malay Peninsula (A. lynnejohnstoneae sp. nov.) and all (1943), Taylor (1965), Teynié et al. (2010), Tweedie (1950) and other species of Aplopeltura by the well defined whitish etchings sources cited therein. on the rear margins of otherwise darker labials. Important references relevant to Pareas carinatus (Boie, 1828), A. omarelhelou sp. nov. are also separated from the other as recognized to date including the taxon, P. nuchalis species by the obvious dark purplish red markings on the head (Boulenger, 1900), either as a full species, subspecies of P. and dorsal surface. carinatus or treated as synonymous to it, include: Boie (1828), White markings in the form of irregular shaped, but vaguely Boulenger (1900), Chan-ard and Nabhitabhata (2015), Chan-ard triangular blotches are very distinct and well defined in A. et al. (1999), Cox et al. (1998), Danaisawadi et al. (2016), Das omarelhelou sp. nov., as opposed to being either moderately (2012), de Rooij (1917), David and Vogel (1996), Dowling and well-defined or poorly defined in all other species. Jenner (1998), Geissler et al. (2011), Götz (2001, 2002), Aplopeltura lynnejohnstoneae sp. nov. from the lower Malay Grismer et al. (2008), Guo and Deng (2009), Hoser (2012a), Peninsula are readily separated from all other species of Kopstein (1936, 1938), Lang (2012), Malkmus and Sauer (1996), Aplopeltura by having significant melanism on the upper surface Malkmus et al. (2002), Manthey and Grossmann (1997), of the head, very white upper labials, the white lacking any Mertens (1930), Pauwels et al. (2003), Pyron et al. (2013), Sang darker pigment, sheen or flecks as seen in all other species, this et al. (2009), Savage (2015), Schlegel (1837), Schmidt and being except for a small triangular dark patch under the eye Kunz (2005), Sclater (1891), Smedley (1931), Smith (1943), (such a dark patch in one form or other is seen in all species), Stuart and Emmett (2006), Taylor (1965), Teynié et al. (2010), but which is far narrower than the eye itself in this species, and Wagler (1830), Zhao and Adler (1993), Ziegler et al. (2006, an indistinct dorsal pattern also defined by the presence of 2007) and sources cited therein. numerous irregular black or blackish flecks throughout. Aplopeltura daranini sp. nov. from Sumatra is readily separated from all other species of Aplopeltura by being similar in most

Hoser 2016 - Australasian Journal of Herpetology 33:42-51. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 44 Australasian Journal of Herpetology respects to A. lynnejohnstoneae sp. nov. as defined above, but attempted to unlawfully steal “name authority” for a species of by the following obvious differences. It has an orange sheen or monitor lizard formally named by Richard Wells and Cliff Ross peppering over the white upper labials so that they do not Wellington, some years earlier (Hoser 2007). appear to be an immaculate white in colour, as well as the APLOPELTURA OMARELHELOU SP. NOV. markings of the iris. In A. lynnejohnstoneae sp. nov. the iris is Holotype: A preserved specimen at the KU Biodiversity Institute more-or-less of one colour being a brownish colour, wheras in A. and Natural History Museum, University of Kansas, Lawrence. daranini sp. nov. the iris is generally whitish in colour, but with USA, specimen number: KU KUH 328493, collected at Mountain darker reddish veins or markings running more-or-less radially View Bungalos, near the entrance to the Khao Sok National out from the centre. On A. daranini sp. nov. the indistinct dorsal Park, Thailand, Latitude 8.91, Longitude 98.53. pattern is defined by the presence of irregular black spots as opposed to smaller blackish flecks as seen in A. This is a facility that allows access to its holdings. lynnejohnstoneae sp. nov.. Diagnosis: Aplopeltura omarelhelou sp. nov. from southern Aplopeltura gibbonsi sp. nov. from Borneo is separated from all Thailand on the Isthumus of Kra in the region generally north of other Aplopeltura by the following characters: Having a and including Songkhla and Trang are readily separated from reasonably distinct dorsal pattern consisting of irregular those on the lower Malay Peninsula (A. lynnejohnstoneae sp. markings or blotches on the dorsal surface, the most obvious nov.) and all other species of Aplopeltura by the well defined ones being an orangeish red in colour. The white markings on whitish etchings on the rear margins of otherwise darker labials. the lower flanks and white parts of the upper labials are heavily A. omarelhelou sp. nov. are also separated from the other peppered making them appear to be off-white and a shade of species by the obvious dark purplish red markings on the head the peppering, or alternatively fully shaded as another colour and dorsal surface. anyway (e.g. yellow). The dark patch under the eye is also well- White markings in the form of irregular shaped, but vaguely broken by one or more streaks of white pigment, this also being triangular blotches are very distinct and well defined in A. well peppered or shaded by a colour such as yellow, orange or omarelhelou sp. nov., as opposed to being either moderately red. well-defined or poorly defined in all other species. Members of the family Pareidae (alternatively referred to as the Aplopeltura lynnejohnstoneae sp. nov. from the lower Malay Pareatidae) are unique among Southeast Asian snakes and Peninsula are readily separated from all other species of diagnosed in having large scales overlapping on their chins Aplopeltura by having significant melanism on the upper surface rather than having them separated by a straight groove. These of the head, very white upper labials, the white lacking any snakes are known to feed mainly on terrestrial molluscs. darker pigment, sheen or flecks as seen in all other species, this being except for a small triangular dark patch under the eye They have short skulls; relatively large eyes; a large nasal gland; (such a dark patch in one form or other is seen in all species), have a relatively low number of scales at mid body (13-15 rows); but which is far narrower than the eye itself in this species, and the chin shields have no midline groove, and extend across the an indistinct dorsal pattern also defined by the presence of chin; they are nocturnal; they feed mostly upon gastropods and numerous irregular black or blackish flecks throughout. small vertebrates; and they lay eggs. Aplopeltura shireenae sp. nov. from the main Philippine Islands, The maxillary bone is very short, deep and with 5 or 6 subequal including Mindanao, are readily separated from all other mandibular teeth gradually decreasing in length. The head is Aplopeltura species by the following suite of characters: A distinct from the neck; eye moderate, with a vertical pupil; nasal generally very indistinct dorsal pattern, being a light reddish- single. Body more or less compressed; scales smooth or feebly brown in colour, the lateral white blotches rising from the belly on keeled, without pits, more or less oblique, in 13-15 rows, the lower flanks are so heavily shaded that they are barely vertebral row may be enlarged or not (depending on the genus). noticeable and this includes for the anterior section of the body, Ventrals rounded. Tail moderate or short; subcaudals divided. which is only similarly seen in Aplopeltura lynnejohnstoneae sp. Observations on captive and wild snakes eating snails suggests nov. from the lower Malay Peninsula. A. shireenae sp. nov. is that they rest the upper jaw on a snail (preferred food) and use separated from A. lynnejohnstoneae sp. nov. by several the mandibles to pull the snail’s body out of its shell. These are characters including a lack of melanism on the dorsal surface of thought to represent a basal lineage of the advanced snakes the head (see in A. lynnejohnstoneae sp. nov.) and a lack of the (Caenophidia). profuse blackish specking seen on the dorsal surface of A. There are 4 recognized genera and about 15 species currently lynnejohnstoneae sp. nov.. recognized (before the publication of this paper), all inhabiting The top of the head of A. shireenae sp. nov., while also reddish . brown in colour has two distinct white patches somewhat Genus Aplopeltura Duméril, 1838 has until now been treated as anterior to the eyes, pointing towards the middle of the head, but being monotypic for the species A. boa (Boie, 1828), but split six not meeting. The iris is brownish in colour and characterised by ways in this paper, with five new species formally named. the presence of whitish specks. Snakes in the genera Aplopeltura Duméril, 1838 and Aplopeltura boa (Boie, 1828) from Java is readily separated from Asthenodipsas Peters, 1864 are considerably thinner in build others in the genus (as defined herein) by significant somewhat and show significant vertical compression (also seen as a irregular shaped white patches along the dorsal midline. The distinct midline ridge along the back), that is not seen in Pareas dark patch below the eye that commences on the jawline, does Wagler, 1830 or Katrinahoserserpenea Hoser, 2012 as not extend as far as the eye or if so, only just intersects it. recognized to date. Aplopeltura daranini sp. nov. from Sumatra is readily separated Aplopeltura has 26-55 single subcaudals, versus 88-120 divided from all other species of Aplopeltura by being similar in most subcaudals in Asthenodipsas as well as numerous other respects to A. lynnejohnstoneae sp. nov. as defined above, but differences at outlined by Boulenger (1896) in his species-level by the following obvious differences. It has an orange sheen or descriptions. peppering over the white upper labials so that they do not Distribution: Philippine Islands. appear to be an immaculate white in colour, as well as the markings of the iris. In A. lynnejohnstoneae sp. nov. the iris is Etymology: Named in honour of my wife, Shireen Hoser in more-or-less of one colour being a brownish colour, wheras in A. recognition of many years services to herpetology and daranini sp. nov. the iris is generally whitish in colour, but with zoological taxonomy, including successfully assisting in darker reddish veins or markings running more-or-less radially petitioning the ICZN to stop taxonomic vandalism in the 1990’s, out from the centre. On A. daranini sp. nov. the indistinct dorsal in a case where Robert George Sprackland of the USA pattern is defined by the presence of irregular black spots as

Hoser 2016 - Australasian Journal of Herpetology 33:42-51. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 45

opposed to smaller blackish flecks as seen in A. number: 143540, collected at Selangor, Peninsular Malaysia. lynnejohnstoneae sp. nov.. This is a facility that allows access to its holdings. Aplopeltura gibbonsi sp. nov. from Borneo is separated from all Paratype: A preserved specimen at the Field Museum of other Aplopeltura by the following characters: Having a Natural History (FMNH) Chicago, Illinois, USA, specimen reasonably distinct dorsal pattern consisting of irregular number: 183769, collected at Selangor, Peninsular Malaysia. markings or blotches on the dorsal surface, the most obvious Diagnosis: Aplopeltura lynnejohnstoneae sp. nov. from the ones being an orangeish red in colour. The white markings on lower Malay Peninsula are readily separated from all other the lower flanks and white parts of the upper labials are heavily species of Aplopeltura by having significant melanism on the peppered making them appear to be off-white and a shade of upper surface of the head, very white upper labials, the white the peppering, or alternatively fully shaded as another colour lacking any darker pigment, sheen or flecks as seen in all other anyway (e.g. yellow). The dark patch under the eye is also well- species, this being except for a small triangular dark patch under broken by one or more streaks of white pigment, this also being the eye (such a dark patch in one form or other is seen in all well peppered or shaded by a colour such as yellow, orange or species), but which is far narrower than the eye itself in this red. species, and an indistinct dorsal pattern also defined by the Members of the family Pareidae (alternatively referred to as the presence of numerous irregular black or blackish flecks Pareatidae) are unique among Southeast Asian snakes and throughout. diagnosed in having large scales overlapping on their chins Aplopeltura shireenae sp. nov. from the main Philippine Islands, rather than having them separated by a straight groove. These including Mindanao, are readily separated from all other snakes are known to feed mainly on terrestrial molluscs. Aplopeltura species by the following suite of characters: A They have short skulls; relatively large eyes; a large nasal gland; generally very indistinct dorsal pattern, being a light reddish- have a relatively low number of scales at mid body (13-15 rows); brown in colour, the lateral white blotches rising from the belly on the chin shields have no midline groove, and extend across the the lower flanks are so heavily shaded that they are barely chin; they are nocturnal; they feed mostly upon gastropods and noticeable and this includes for the anterior section of the body, small vertebrates; and they lay eggs. which is only similarly seen in Aplopeltura lynnejohnstoneae sp. The maxillary bone is very short, deep and with 5 or 6 subequal nov. from the lower Malay Peninsula. A. shireenae sp. nov. is mandibular teeth gradually decreasing in length. Head is distinct separated from A. lynnejohnstoneae sp. nov. by several from the neck; eye moderate, with a vertical pupil; nasal single. characters including a lack of melanism on the dorsal surface of Body more or less compressed; scales smooth or feebly keeled, the head (see in A. lynnejohnstoneae sp. nov.) and a lack of the without pits, more or less oblique, in 13-15 rows, vertebral row profuse blackish specking seen on the dorsal surface of A. may be enlarged or not (depending on the genus). Ventrals lynnejohnstoneae sp. nov.. rounded. Tail moderate or short; subcaudals divided. The top of the head of A. shireenae sp. nov., while also reddish Observations on captive and wild snakes eating snails suggests brown in colour has two distinct white patches somewhat that they rest the upper jaw on a snail (preferred food) and use anterior to the eyes, pointing towards the middle of the head, but the mandibles to pull the snail’s body out of its shell. These are not meeting. The iris is brownish in colour and characterised by thought to represent a basal lineage of the advanced snakes the presence of whitish specks. (Caenophidia). Aplopeltura boa (Boie, 1828) from Java is readily separated from There are 4 recognized genera and about 15 species currently others in the genus (as defined herein) by significant somewhat recognized (before the publication of this paper), all inhabiting irregular shaped white patches along the dorsal midline. The Southeast Asia. dark patch below the eye that commences on the jawline, does Genus Aplopeltura Duméril, 1838 has until now been treated as not extend as far as the eye or if so, only just intersects it. being monotypic for the species A. boa (Boie, 1828), but split six Aplopeltura omarelhelou sp. nov. from southern Thailand on the ways in this paper, with five new species formally named. Isthumus of Kra in the region generally north of and including Songkhla and Trang are readily separated from those on the Snakes in the genera Aplopeltura Duméril, 1838 and lower Malay Peninsula (A. lynnejohnstoneae sp. nov.) and all Asthenodipsas Peters, 1864 are considerably thinner in build other species of Aplopeltura by the well defined whitish etchings and show significant vertical compression (also seen as a on the rear margins of otherwise darker labials. distinct midline ridge along the back), that is not seen in Pareas A. omarelhelou sp. nov. are also separated from the other Wagler, 1830 or Katrinahoserserpenea Hoser, 2012 as species by the obvious dark purplish red markings on the head recognized to date. and dorsal surface. Aplopeltura has 26-55 single subcaudals, versus 88-120 divided subcaudals in Asthenodipsas as well as numerous other White markings in the form of irregular shaped, but vaguely differences at outlined by Boulenger (1896) in his species-level triangular blotches are very distinct and well defined in A. descriptions. omarelhelou sp. nov., as opposed to being either moderately well-defined or poorly defined in all other species. Distribution: Southern Thailand on the Isthumus of Kra in the Aplopeltura daranini sp. nov. from Sumatra is readily separated region generally north of and including Songkhla and Trang. from all other species of Aplopeltura by being similar in most Etymology: Named in honour of Omar Elhelou of Melbourne, respects to A. lynnejohnstoneae sp. nov. as defined above, but Victoria, Australia, owner of Precision Security Australia who has by the following obvious differences. It has an orange sheen or made a significant contribution to wildlife conservation and peppering over the white upper labials so that they do not research in Australia by by installing security systems to appear to be an immaculate white in colour, as well as the safeguard the and property of Snakebusters: Australia’s markings of the iris. In A. lynnejohnstoneae sp. nov. the iris is best shows. more-or-less of one colour being a brownish colour, wheras in A. The patronym name would in the normal course of events be daranini sp. nov. the iris is generally whitish in colour, but with designated as “omarelhelouí” instead of the designated name darker reddish veins or markings running more-or-less radially “omarelhelou” but the spelling I have given it in this paper is out from the centre. On A. daranini sp. nov. the indistinct dorsal deliberate and should not be amended unless mandated by the pattern is defined by the presence of irregular black spots as International Code of Zoological Nomenclature. opposed to smaller blackish flecks as seen in A. APLOPELTURA LYNNEJOHNSTONEAE SP. NOV. lynnejohnstoneae sp. nov.. Holotype: A preserved specimen at the Field Museum of Aplopeltura gibbonsi sp. nov. from Borneo is separated from all Natural History (FMNH), Chicago, Illinois, USA, specimen other Aplopeltura by the following characters: Having a

Hoser 2016 - Australasian Journal of Herpetology 33:42-51. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 46 Australasian Journal of Herpetology reasonably distinct dorsal pattern consisting of irregular separated from all other species of Aplopeltura by being similar markings or blotches on the dorsal surface, the most obvious in most respects to A. lynnejohnstoneae sp. nov. (as defined in ones being an orangeish red in colour. The white markings on the description preceding or within this one, below), but by the the lower flanks and white parts of the upper labials are heavily following obvious differences. It has an orange sheen or peppered making them appear to be off-white and a shade of peppering over the white upper labials so that they do not the peppering, or alternatively fully shaded as another colour appear to be an immaculate white in colour, as well as the anyway (e.g. yellow). The dark patch under the eye is also well- markings of the iris. In A. lynnejohnstoneae sp. nov. the iris is broken by one or more streaks of white pigment, this also being more-or-less of one colour being a brownish colour, wheras in A. well peppered or shaded by a colour such as yellow, orange or daranini sp. nov. the iris is generally whitish in colour, but with red. darker reddish veins or markings running more-or-less radially Members of the family Pareidae (alternatively referred to as the out from the centre. On A. daranini sp. nov. the indistinct dorsal Pareatidae) are unique among Southeast Asian snakes and pattern is defined by the presence of irregular black spots as diagnosed in having large scales overlapping on their chins opposed to smaller blackish flecks as seen in A. rather than having them separated by a straight groove. These lynnejohnstoneae sp. nov.. snakes are known to feed mainly on terrestrial molluscs. Aplopeltura lynnejohnstoneae sp. nov. from the lower Malay They have short skulls; relatively large eyes; a large nasal gland; Peninsula are readily separated from all other species of have a relatively low number of scales at mid body (13-15 rows); Aplopeltura by having significant melanism on the upper surface the chin shields have no midline groove, and extend across the of the head, very white upper labials, the white lacking any chin; they are nocturnal; they feed mostly upon gastropods and darker pigment, sheen or flecks as seen in all other species, this small vertebrates; and they lay eggs. being except for a small triangular dark patch under the eye (such a dark patch in one form or other is seen in all species), The maxillary bone is very short, deep and with 5 or 6 subequal but which is far narrower than the eye itself in this species, and mandibular teeth gradually decreasing in length. The head is an indistinct dorsal pattern also defined by the presence of distinct from the neck; eye moderate, with a vertical pupil; nasal numerous irregular black or blackish flecks throughout. single. Body more or less compressed; scales smooth or feebly Aplopeltura shireenae sp. nov. from the main Philippine Islands, keeled, without pits, more or less oblique, in 13-15 rows, including Mindanao, are readily separated from all other vertebral row may be enlarged or not (depending on the genus). Aplopeltura species by the following suite of characters: A Ventrals rounded. Tail moderate or short; subcaudals divided. generally very indistinct dorsal pattern, being a light reddish- Observations on captive and wild snakes eating snails suggests brown in colour, the lateral white blotches rising from the belly on that they rest the upper jaw on a snail (preferred food) and use the lower flanks are so heavily shaded that they are barely the mandibles to pull the snail’s body out of its shell. These are noticeable and this includes for the anterior section of the body, thought to represent a basal lineage of the advanced snakes which is only similarly seen in Aplopeltura lynnejohnstoneae sp. (Caenophidia). nov. from the lower Malay Peninsula. A. shireenae sp. nov. is There are 4 recognized genera and about 15 species currently separated from A. lynnejohnstoneae sp. nov. by several recognized (before the publication of this paper), all inhabiting characters including a lack of melanism on the dorsal surface of Southeast Asia. the head (see in A. lynnejohnstoneae sp. nov.) and a lack of the Genus Aplopeltura Duméril, 1838 has until now been treated as profuse blackish specking seen on the dorsal surface of A. being monotypic for the species A. boa (Boie, 1828), but split six lynnejohnstoneae sp. nov.. ways in this paper, with five new species formally named. The top of the head of A. shireenae sp. nov., while also reddish Snakes in the genera Aplopeltura Duméril, 1838 and brown in colour has two distinct white patches somewhat Asthenodipsas Peters, 1864 are considerably thinner in build anterior to the eyes, pointing towards the middle of the head, but and show significant vertical compression (also seen as a not meeting. The iris is brownish in colour and characterised by distinct midline ridge along the back), that is not seen in Pareas the presence of whitish specks. Wagler, 1830 or Katrinahoserserpenea Hoser, 2012 as Aplopeltura boa (Boie, 1828) from Java is readily separated from recognized to date. others in the genus (as defined herein) by significant somewhat Aplopeltura has 26-55 single subcaudals, versus 88-120 divided irregular shaped white patches along the dorsal midline. The subcaudals in Asthenodipsas as well as numerous other dark patch below the eye that commences on the jawline, does differences at outlined by Boulenger (1896) in his species-level not extend as far as the eye or if so, only just intersects it. descriptions. Aplopeltura omarelhelou sp. nov. from southern Thailand on the Distribution: Lower Peninsula Malaysia in a region generally Isthumus of Kra in the region generally north of and including south of Songkhla and Trang, southern Thailand (not including Songkhla and Trang are readily separated from those on the those two named areas). lower Malay Peninsula (A. lynnejohnstoneae sp. nov.) and all Etymology: Named in honour of Lynne Johnstone in recognition other species of Aplopeltura by the well defined whitish etchings of her contributions to Australian culture via her long term on the rear margins of otherwise darker labials. relationship with lifetime partner Bruce Rogers, of Bend of A. omarelhelou sp. nov. are also separated from the other Islands, Kangaroo Ground, Victoria, Australia, one of the worlds species by the obvious dark purplish red markings on the head foremost Didgeridoo players and makers, who has been hailed and dorsal surface. as a hero for Indigenous Australians, even though he was not White markings in the form of irregular shaped, but vaguely born as one. In 2015, Bruce Rogers was diagnosed with an triangular blotches are very distinct and well defined in A. aggressive terminal cancer, (Leukemia), and died in July 2016. omarelhelou sp. nov., as opposed to being either moderately Brucerogersus Hoser, 2012 (type species Eutaenia well-defined or poorly defined in all other species. chrysocephala Cope, 1885) is a genus of North American Garter Aplopeltura gibbonsi sp. nov. from Borneo is separated from all Snakes named in honour of Bruce Rogers. other Aplopeltura by the following characters: Having a APLOPELTURA DARANINI SP. NOV. reasonably distinct dorsal pattern consisting of irregular markings or blotches on the dorsal surface, the most obvious Holotype: A preserved specimen at the American Museum of ones being an orangeish red in colour. The white markings on Natural History, New York, USA, specimen number: AMNH the lower flanks and white parts of the upper labials are heavily Herpetology R-2886 collected from Sumatra, . peppered making them appear to be off-white and a shade of This is a facility that allows access to its holdings. the peppering, or alternatively fully shaded as another colour Diagnosis: Aplopeltura daranini sp. nov. from Sumatra is readily anyway (e.g. yellow). The dark patch under the eye is also well-

Hoser 2016 - Australasian Journal of Herpetology 33:42-51. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 47

broken by one or more streaks of white pigment, this also being surface, the most obvious ones being an orangeish red in well peppered or shaded by a colour such as yellow, orange or colour. The white markings on the lower flanks and white parts red. of the upper labials are heavily peppered making them appear to Members of the family Pareidae (alternatively referred to as the be off-white and a shade of the peppering, or alternatively fully Pareatidae) are unique among Southeast Asian snakes and shaded as another colour anyway (e.g. yellow). The dark patch diagnosed in having large scales overlapping on their chins under the eye is also well-broken by one or more streaks of rather than having them separated by a straight groove. These white pigment, this also being well peppered or shaded by a snakes are known to feed mainly on terrestrial molluscs. colour such as yellow, orange or red. They have short skulls; relatively large eyes; a large nasal gland; Aplopeltura lynnejohnstoneae sp. nov. from the lower Malay have a relatively low number of scales at mid body (13-15 rows); Peninsula are readily separated from all other species of the chin shields have no midline groove, and extend across the Aplopeltura by having significant melanism on the upper surface chin; they are nocturnal; they feed mostly upon gastropods and of the head, very white upper labials, the white lacking any small vertebrates; and they lay eggs. darker pigment, sheen or flecks as seen in all other species, this being except for a small triangular dark patch under the eye The maxillary bone is very short, deep and with 5 or 6 subequal (such a dark patch in one form or other is seen in all species), mandibular teeth gradually decreasing in length. The head is but which is far narrower than the eye itself in this species, and distinct from the neck; eye moderate, with a vertical pupil; nasal an indistinct dorsal pattern also defined by the presence of single. Body more or less compressed; scales smooth or feebly numerous irregular black or blackish flecks throughout. keeled, without pits, more or less oblique, in 13-15 rows, Aplopeltura shireenae sp. nov. from the main Philippine Islands, vertebral row may be enlarged or not (depending on the genus). including Mindanao, are readily separated from all other Ventrals rounded. Tail moderate or short; subcaudals divided. Aplopeltura species by the following suite of characters: A Observations on captive and wild snakes eating snails suggests generally very indistinct dorsal pattern, being a light reddish- that they rest the upper jaw on a snail (preferred food) and use brown in colour, the lateral white blotches rising from the belly on the mandibles to pull the snail’s body out of its shell. These are the lower flanks are so heavily shaded that they are barely thought to represent a basal lineage of the advanced snakes noticeable and this includes for the anterior section of the body, (Caenophidia). which is only similarly seen in Aplopeltura lynnejohnstoneae sp. There are 4 recognized genera and about 15 species currently nov. from the lower Malay Peninsula. A. shireenae sp. nov. is recognized (before the publication of this paper), all inhabiting separated from A. lynnejohnstoneae sp. nov. by several Southeast Asia. characters including a lack of melanism on the dorsal surface of Genus Aplopeltura Duméril, 1838 has until now been treated as the head (see in A. lynnejohnstoneae sp. nov.) and a lack of the being monotypic for the species A. boa (Boie, 1828), but split six profuse blackish specking seen on the dorsal surface of A. ways in this paper, with five new species formally named. lynnejohnstoneae sp. nov.. Snakes in the genera Aplopeltura Duméril, 1838 and The top of the head of A. shireenae sp. nov., while also reddish Asthenodipsas Peters, 1864 are considerably thinner in build brown in colour has two distinct white patches somewhat and show significant vertical compression (also seen as a anterior to the eyes, pointing towards the middle of the head, but distinct midline ridge along the back), that is not seen in Pareas not meeting. The iris is brownish in colour and characterised by Wagler, 1830 or Katrinahoserserpenea Hoser, 2012 as the presence of whitish specks. recognized to date. Aplopeltura boa (Boie, 1828) from Java is readily separated from Aplopeltura has 26-55 single subcaudals, versus 88-120 divided others in the genus (as defined herein) by significant somewhat subcaudals in Asthenodipsas as well as numerous other irregular shaped white patches along the dorsal midline. The differences at outlined by Boulenger (1896) in his species-level dark patch below the eye that commences on the jawline, does descriptions. not extend as far as the eye or if so, only just intersects it. Distribution: Known only from the hillier sections of the island Aplopeltura omarelhelou sp. nov. from southern Thailand on the of Sumatra, Indonesia and mainly in the north. Isthumus of Kra in the region generally north of and including Etymology: Named in honour of Dara Nin of Ringwood, Victoria, Songkhla and Trang are readily separated from those on the Australia in recognition of a decade’s valuable wildlife lower Malay Peninsula (A. lynnejohnstoneae sp. nov.) and all conservation and education work with Snakebusters: Australia’s other species of Aplopeltura by the well defined whitish etchings best reptile displays, being the only wildlife display in Australia on the rear margins of otherwise darker labials. that lets people hold the animals, as opposed to boring static A. omarelhelou sp. nov. are also separated from the other displays done by less experienced imitators, this being the species by the obvious dark purplish red markings on the head sworn evidence of Ronald Leslie Waters, a former head of and dorsal surface. wildlife law enforcement at the Department of Sustainability and White markings in the form of irregular shaped, but vaguely Environment in Victoria (AKA DSE), in a court of law in 2015 triangular blotches are very distinct and well defined in A. (VCAT 2015). omarelhelou sp. nov., as opposed to being either moderately APLOPELTURA GIBBONSI SP. NOV. well-defined or poorly defined in all other species. Aplopeltura daranini sp. nov. from Sumatra is readily separated Holotype: A preserved specimen at the Field Museum of from all other species of Aplopeltura by being similar in most Natural History, Chicago, Illinois, USA, specimen number: respects to A. lynnejohnstoneae sp. nov. as defined above, but FMNH Amphibians and Reptiles 246150, collected at Sabah, by the following obvious differences. It has an orange sheen or Borneo, Malaysia. peppering over the white upper labials so that they do not The Field Museum of Natural History, Chicago, Illinois, USA, appear to be an immaculate white in colour, as well as the allows access to its holdings. markings of the iris. In A. lynnejohnstoneae sp. nov. the iris is Paratypes: Three preserved specimens at the Field Museum of more-or-less of one colour being a brownish colour, wheras in A. Natural History, Chicago, Illinois, USA, specimen numbers: daranini sp. nov. the iris is generally whitish in colour, but with FMNH Amphibians and Reptiles 246149, 246146 and 246151, darker reddish veins or markings running more-or-less radially collected at Sabah, Borneo, Malaysia. out from the centre. On A. daranini sp. nov. the indistinct dorsal Diagnosis: Aplopeltura gibbonsi sp. nov. from Borneo is pattern is defined by the presence of irregular black spots as separated from all other Aplopeltura species by the following opposed to smaller blackish flecks as seen in A. suite of characters: Having a reasonably distinct dorsal pattern lynnejohnstoneae sp. nov.. consisting of irregular markings or blotches on the dorsal Members of the family Pareidae (alternatively referred to as the Hoser 2016 - Australasian Journal of Herpetology 33:42-51. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 48 Australasian Journal of Herpetology

Pareatidae) are unique among Southeast Asian snakes and line with the other subspecies, the dorsal pattern beyond this diagnosed in having large scales overlapping on their chins point is one of alternating semi-destinct blackish crossbands rather than having them separated by a straight groove. These intersperced with wider areas of dark brown or greyish brown. snakes are known to feed mainly on terrestrial molluscs. On the forebody of P. carinatus sumatrensis subsp. nov. from They have short skulls; relatively large eyes; a large nasal gland; Sumatra, the lighter dorsal crossbands are significantly wider have a relatively low number of scales at mid body (13-15 rows); than the darker ones, versus the reverse situation (or roughly the chin shields have no midline groove, and extend across the equal size) as seen in nominate P. carinatus carinatus from chin; they are nocturnal; they feed mostly upon gastropods and Java, or those from Peninsula Malaysia, herein named as P. small vertebrates; and they lay eggs. carinatus malayensis subsp. nov.. The maxillary bone is very short, deep and with 5 or 6 subequal P. carinatus nuchalis (Boulenger, 1890) from Borneo, are readily mandibular teeth gradually decreasing in length. The head is distinguished from the other subspecies by the obvious distinct from the neck; eye moderate, with a vertical pupil; nasal melanism (blackish colouration) of the head and neck, meaning single. Body more or less compressed; scales smooth or feebly the temporal streak is either hidden or not obvious, the fact that keeled, without pits, more or less oblique, in 13-15 rows, the dark blackish bands are distinct and extend quite clearly vertebral row may be enlarged or not (depending on the genus). down the flanks to the ventral surface, and usually have Ventrals rounded. Tail moderate or short; subcaudals divided. individual white scales on the spine at the forebody being noticeably lighter in colour, forming a semi-distinct dotted Observations on captive and wild snakes eating snails suggests vertebral stripe. that they rest the upper jaw on a snail (preferred food) and use the mandibles to pull the snail’s body out of its shell. These are P. carinatus unicolor Bourret, 1934 from Cambodia are thought to represent a basal lineage of the advanced snakes separated from the other subspecies by the fact that the darker (Caenophidia). crossbands are significantly obscured on the mid flanks being There are 4 recognized genera and about 15 species currently near absent at this point. The mid and rear upper labials are recognized (before the publication of this paper), all inhabiting also characterised by dark etching (usually dark brown) on the Southeast Asia. scale margins. Genus Aplopeltura Duméril, 1838 has until now been treated as There is also an unsual unicolour phase in the range for P. being monotypic for the species A. boa (Boie, 1828), but split six carinatus unicolor Bourret, 1934, not seen elsewhere. ways in this paper, with five new species formally named. P. carinatus malayensis subsp. nov.. from Peninsula Malaysia and nearby places further north are similar in most respects to P. Snakes in the genera Aplopeltura Duméril, 1838 and carinatus carinatus but are separated from that subspecies by Asthenodipsas Peters, 1864 are considerably thinner in build the absence of the temporal streak on each side merging to and show significant vertical compression (also seen as a form a short thick black band on the back of the head and neck, distinct midline ridge along the back), that is not seen in Pareas which at the posterior end extends down the flanks to Wagler, 1830 or Katrinahoserserpenea Hoser, 2012 as immediately above the ventrals. recognized to date. P. carinatus malayensis subsp. nov.. are further separated from Aplopeltura has 26-55 single subcaudals, versus 88-120 divided the other subspecies by either the absence of a well defined subcaudals in Asthenodipsas as well as numerous other temporal streak, or alternatively one that is relatively indistinct. differences at outlined by Boulenger (1896) in his species-level descriptions. Snakes of the genus Pareas as presently recognized consists of (about) just one species as defined herein and are diagnosed by Distribution: The island of Borneo, Malaysia, Brunei and being small and slender with a blunt snout, no mental groove Indonesia, mainly in the hiller parts of the north and immediately and no teeth on the anterior part of the maxillary. adjacent areas. Snakes in the genera Aplobeltura Duméril, 1853 and Etymology: Named in honour of Dale Gibbons of Maiden Gully, Asthenodipsas Peters, 1864 are considerably thinner in build on the outskirts of Bendigo, Victoria, Australia in recognition of a and more vertically compressed than seen in Pareas and lifetime’s work with wildlife conservation and herpetology, Katrinahoserserpenea Hoser, 2013 as recognized to date. including captive breeding of rare and threatened species as well as a significant contribution to field work and surveys Pareas has since 2012 been restricted to the species taxa P. throughout the Australian state of Victoria. carinatus and putative P. nuchalis, treated here as a subspecies of one another, along with the other forms identified herein. PAREAS CARINATUS SUMATRENSIS SUBSP. NOV. Those taxa differ from taxa in the genus Katrinahoserserpenea Holotype: A preserved specimen at the Museum of by cephalic scalation and distribution pattern. Comparative Zoology, Harvard University in Cambridge, P. carinatus sensu lato (including putative P. nuchalis and other Massachusetts, USA, specimen number: MCZ:Herp:R-37764, forms identified above) share three anterior temporals in collected at Langhat, north-east coast of Sumatra, Indonesia, contrast to the one or two (rarely three) anterior temporals in Latitude 3.98,, Longitude 98.48. Katrinahoserserpenea species. The Museum of Comparative Zoology, Harvard University in Cambridge, Massachusetts, USA, allows access to its holdings. The frontal scale in P. carinatus sensu lato (including putative P. nuchalis and other forms identified above) is hexagonal with the Paratype: A preserved specimen at the National Museum of lateral sides parallel to the body axis; this scale in Natural History (USA), formerly known as the US National Katrinahoserserpenea is almost diamond-shaped or shield- Museum, administered by the Smithsonian Institution, shaped with the lateral sides converging posteriorly. Washington, D.C., USA, specimen number: USNM 103582.6076586, collected from Utara, Sumatra, Indonesia. The two anterior chin shields are longer than broad in Katrinahoserserpenea, whereas in Pareas they are broader than Diagnosis: Pareas carinatus sumatrensis sp. nov. from long; this is a consistent way to separate the two genera. Sumatra is readily separated from all other subspecies by the The snakes remaining in the genus Pareas occur mainly presence of extensive peppering on the light upper labials and throughout the Indochinese Peninsula and Sunda Islands. By head. The same peppering extends along the upper forebody. contrast most species of Katrinahoserserpenea occur in central Nominate P. carinatus carinatus from Java and islands east of and southern China and the northern Indochinese Peninsula, there, are separated from the other subspecies by a temporal with only two species, Katrinahoserserpenea margaritophorus streak on each side, merging to form a short thick black band on and K. hamptoni being found in the southern Indochinese the back of the head and neck, which at the posterior end Peninsula. extends down the flanks to immediately above the ventrals. In Hoser 2016 - Australasian Journal of Herpetology 33:42-51. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 49

Distribution: Hillier parts of Sumatra, Indonesia and nearby pattern. areas, including most of the northern quarter of the island and P. carinatus sensu lato (including putative P. nuchalis and other the hills and mountains running along the western side of the relevant taxa) share three anterior temporals in contrast to the island. one or two (rarely three) anterior temporals in Etymology: Named in reflection of where they naturally occur. Katrinahoserserpenea species. PAREAS CARINATUS MALAYENSIS SUBSP. NOV. The frontal scale in P. carinatus sensu lato (including putative P. Holotype: A preserved specimen at the California Academy of nuchalis and other relevant taxa) is hexagonal with the lateral Science, San Francisco, USA, specimen number: sides parallel to the body axis; this scale in CAS:HERP:73696, collected at Patani, southern Thailand. Katrinahoserserpenea is almost diamond-shaped or shield- shaped with the lateral sides converging posteriorly. The California Academy of Science, USA allows access to its holdings The two anterior chin shields are longer than broad in Diagnosis: P. carinatus malayensis subsp. nov.. from Peninsula Katrinahoserserpenea, whereas in Pareas they are broader than Malaysia are similar in most respects to P. carinatus carinatus long; this is a consistent way to separate the two genera. (see below) but are separated from that subspecies by the The snakes remaining in the genus Pareas occur mainly absence of the temporal streak on each side merging to form a throughout the Indochinese Peninsula and Sunda Islands. By short thick black band on the back of the head and neck, which contrast most species of Katrinahoserserpenea occur in central at the posterior end extends down the flanks to immediately and southern China and the northern Indochinese Peninsula, above the ventrals. with only two species Katrinahoserserpenea margaritophorus P. carinatus malayensis subsp. nov.. are further separated from and K. hamptoni being found in the southern Indochinese the other subspecies by either the absence of a well defined Peninsula. temporal streak, or alternatively one that is relatively indistinct. Finally I note that P. carinatus malayensis subsp. nov. is a legally available name under the rules of the the International Nominate P. carinatus carinatus from Java and islands east of Code of Zoological Nomenclature (Ride et al. 1999). The name there are separated from the other subspecies by a temporal Malayopython Reynolds et al. (2013a, b and 2014) is not! streak on each side, merging to form a short thick black band on the back of the head and neck, which at the posterior end See Hoser (2015) for the details. extends down the flanks to immediately above the ventrals. In Distribution: The lower Malay Peninsula, Asia, extending north line with the other subspecies, the dorsal pattern beyond this along most of the Isthumus of Kra. point is one of alternating semi-destinct blackish crossbands Etymology: Named in reflection of where they naturally occur. intersperced with wider areas of dark brown or greyish brown. REFERENCES CITED On the forebody of P. carinatus sumatrensis subsp. nov. from Boie, H. 1828. Auszüge aus Briefen von Heinr. Boie zu Java an Sumatra, the lighter dorsal crossbands are significantly wider Hn. Schlegel, Conservator anim. vertebr. am Königl. niederl. than the darker ones, versus the reverse situation (or roughly Museum. lsis von Oken, Jena, 21(10):1.025-1.035 equal size) as seen in nominate P. carinatus carinatus from Java, or those from Peninsula Malaysia, herein named as P. Boulenger, G. A. 1894. On the herpetological fauna of Palawan carinatus malayensis subsp. nov.. and Balabac. Ann. Mag. Nat. Hist. (6)14:81-90. Pareas carinatus sumatrensis sp. nov. is readily separated from Boulenger, G. A. 1896. Catalogue of the snakes in the British all other subspecies by the presence of extensive peppering on Museum (Natural History) Volume 3. London, UK: xiv+727 pp. the light upper labials and head. The same peppering extends Boulenger, G. A. 1900. Description of new reptiles and along the upper forebody. batrachians from Borneo. Proc. Zool. Soc. London:182-187. P. carinatus nuchalis (Boulenger, 1890) from Borneo are readily Chan-ard, T., Parr, J. W. K. and Nabhitabhata, J. 2015. A field distinguished from the other subspecies by the obvious guide to the reptiles of Thailand. Oxford University Press, NY, melanism (blackish colouration) of the head and neck, meaning USA:352 pp. the temporal streak is either hidden or not obvious, the fact that Chan-ard, T., Grossmann, W., Gumprecht, A. and Schulz, K. D. the dark blackish bands are distinct and extend quite clearly 1999. Amphibians and reptiles of peninsular Malaysia and down the flanks to the ventral surface, and usually have Thailand - an illustrated checklist [bilingual English and individual white scales on the spine at the forebody being German]. Bushmaster Publications, Würselen, Gemany:240 pp. noticeably lighter in colour, forming a semi-distinct dotted Cox, M. J., Van Dijk, P. P., Nabhitabhata, J. and Thirakhupt, K. vertebral stripe. 1998. A Photographic Guide to Snakes and Other Reptiles of P. carinatus unicolor Bourret, 1934 from Cambodia are Peninsular Malaysia, Singapore and Thailand. Ralph Curtis separated from the other subspecies by the fact that the darker Publishing:144 pp. crossbands are significantly obscured on the mid flanks being Court of Appeal Victoria 2014. Hoser v Department of near absent at this point. The mid and rear upper labials are Sustainability and Environment [2014] VSCA 206 (5 September also characterised by dark etching (usually dark brown) on the 2014). scale margins. Danaisawadi, P., Asami, T., Ota, H., Sutcharit, C. and Panha, S. There is also an unsual unicolour phase in the range for P. 2016. A snail-eating snake recognizes prey handedness. carinatus unicolor Bourret, 1934, not seen elsewhere. Scientific Reports 6(5 April): Article no. 23832. Snakes of the genus Pareas as presently recognized consists of Das, I. 2012. A Naturalist’s Guide to the Snakes of South-East about just one species as defined herein and are diagnosed by Asia: Malaysia, Singapore, Thailand, Myanmar, Borneo, being small and slender with a blunt snout, no mental groove Sumatra, Java and . Oxford John Beaufoy Publishing. and no teeth on the anterior part of the maxillary. David, P. and Vogel, G. 1996. The snakes of Sumatra. An Snakes in the genera Aplobeltura Duméril, 1853 and annotated checklist and key with natural history notes. Bücher Asthenodipsas Peters, 1864 are considerably thinner in build Kreth, Frankfurt/M. and more vertically compressed than seen in Pareas and Katrinahoserserpenea Hoser, 2013 as recognized to date. de Rooij, N. 1917. The Reptiles of the Indo-Australian Archipelago. Il. Ophidia. Leiden (E. J. Brill), xiv+334 S. Pareas has since 2012 been restricted to the species taxa P. carinatus and putative P. nuchalis, treated here as a subspecies. Dowling, H. G. and Jenner, J. V. 1988. Snakes of Burma: checklist of reported species and bibliography. Smithsonian Those two species differ from taxa in the genus Herp. Inf. Serv. (76):19 pp. Katrinahoserserpenea by cephalic scalation and distribution

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Duméril, A. M. C. 1853. Prodrome de la classifcation des Borneo). A.R.G. Ganther Verlag, Rugell:404 pp. reptiles ophidiens. Mém. Acad. Sci., Paris, 23:399-536. Manthey, U. and Grossmann, W. 1997. Amphibien and Reptilien Geissler, P., Nguyen, T. Q., Poyarkov, N. A. and Böhme, W. Südostasiens. Natur und Tier Verlag (Münster):512 pp. 2011. New records of snakes from Cat Tien National Park, Dong Mertens, R. 1930. Die Amphibien und Reptilien der Inseln Bali, Nai and Lam Dong provinces, southern Vietnam. Bonn Lombok, Sumbawa und Flores. Senck. Naturf. Gesell., Frankfurt zoological Bulletin 60(1):9-16. am Main, Abhandl. 42(3):117-344. Götz, M. 2001. Die Schneckennatter Pareas carinatus WAGLER Pauwels, O. S. G., David, P., Chimsunchart, C. and Thirakhupt, 1830 - Terrarienhaltung, Beutefangverhalten und Zucht. K. 2003. Reptiles of Phetchaburi Province, Western Thailand: a Salamandra 37(1):49-58. list of species, with natural history notes, and a discussion on Götz, M. 2002. The feeding behavior of the snail-eating snake the biogeography at the Isthmus of Kra. Natural History Journal Pareas carinatus Wagler 1830 (: ). of Chulalongkorn University 3(1):23-53. Amphibia-Reptilia 23(4):487-493. Peters, W. 1864. Über neue Amphibien (Typhloscincus, Grandison, A. G. C. 1978. Snakes of West Malaysia and Typhlops, Asthenodipsas, Ogmodon). Mber. k. preuss. Akad. Singapore. Annalen des Naturhistorischen Museums in Wien Wiss., Berlin:271-276. 81[1977]:283-303. Pyron, R. A., Burbrink, F. T. and Wiens, J. J. 2013. A phylogeny Grismer, L. L., Neang, T., Chav, T. and Grismer, J. L. 2008. and revised classification of Squamata, including 4161 species Checklist of the amphibians and reptiles of the Cardamom of lizards and snakes. BMC Evolutionary Biology 13:93. region of Southwestern Cambodia. Cambodian Journal of Reynolds, R. G., Niemiller, M. L. and Revella, L. J. 2013a. Natural History 2008(1):12-28. Toward a Tree-of-Life for the boas and pythons: Multilocus Grismer, L. 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with the description of a new species of the genus Bufo. Zootaxa Oxford/Ohio:522 pp. 2416:1-43. Ziegler, T., Ohler, A., Thanh, V. N., Quyet, L. K., Thuan, N. X., Tweedie, M. W. F. 1950. Notes on Malayan reptiles, No.2. Bull. Tri, D. H. and Thanh, B. N. 2006. Review of the amphibian and Raffl. Mus. 23:191-199. reptile diversity of Phong Nha - Ke Bang National Park and Victorian Civil and Administrative Tribunal (VCAT). 2015. Hoser adjacent areas, central Truong Son, Vietnam. Herpetologica v Department of Environment Land Water and Planning (Review Bonnensis II: 247-262. and Regulation) [2015] VCAT 1147 (30 July 2015). Ziegler, T., Hendrix, R., Thanh, V. N., Vogt, M., Forster, B. and Wagler, J. G. 1830. Natürliches System der Amphibien, mit Kien, D. N. 2007. The diversity of a snake community in a karst vorangehender Classification der Säugetiere und Vögel. Ein forest ecosystem in the central Truong Son, Vietnam, with an Beitrag zur vergleichenden Zoologie. 1.0. Cotta, München, identification key. Zootaxa 1493:1-40. Stuttgart, and Tübingen:354 pp. CONFLICT OF INTEREST Zhao, E. and Adler, K. 1993. Herpetology of China. SSAR, The author has no known conflicts of interest in terms of this paper and conclusions within. Snakebusters Australia’s best reptiles

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