ISSUE 31, PUBLISHED 1 AUGUST 2016 ISSN 1836-5698 (Print) ISSN 1836-5779 (Online) AustralasianAustralasian JournalJournal ofof HerpetologyHerpetology

CONTENTS PAGE 2 2 Australasian Journal of Herpetology Australasian Journal of Herpetology Issue 31, 1 August 2016 Contents

Acanthophis lancasteri Wells and Wellington, 1985 gets hit with a dose of Crypto! … this is not the last word on Death Adder and nomenclature. ... Raymond T. Hoser, 3- 11. A division of the elapid Salomonelaps McDowell, 1970 from the , including the resurrection of two and formal description of four other forms (Serpentes: : Micropechiini: Loveridgelapina). ... Raymond T. Hoser, 21-21. A division of the genus elapid genus Loveridegelaps McDowell, 1970 from the Solomon Islands, including formal description of four new species (Serpentes: Elapidae: Micropechiini: Loveridgelapina). ... Raymond T. Hoser, 22-28. A review of the and the Dragon javanicus Reinhardt, 1836 species group, including the formal description of three new species, a division of Peters, 1869 into two genera and Jerdon, 1870 into subgenera (; Serpentes, , Xenodermidae). ... Raymond T. Hoser, 29-34. A second new Tropidechis Günther, 1863 from far north Queensland (Squamata: Serpentes: Elapidae). ... Raymond T. Hoser, 35-38. A review of the bibroni species complex (Squamata: Serpentes: Candoiidae: Candoia). ... Raymond T. Hoser, 39-61. A new species of Denisonia from North-west Queensland, Australia (Serpentes: Elapidae). ... Raymond T. Hoser, 62-63.

Cover images: Front: Mating Acanthophis cummingi bred by Gordon Plumridge of Bendigo, Victoria. Back: Amelansitic Acanthophis bottomi bred by Andrew Gedye of Cairns, North Queensland. Photos by Raymond T. Hoser.

Australasian Journal of Herpetology ® Publishes original research in printed form in relation to , other fauna and related matters, including classification, ecology, public interest, legal, captivity, “academic misconduct”, etc. It is a peer reviewed printed journal published in hard copy for permanent public scientific record in accordance with the International Code of Zoological Nomenclature (Ride et al. 1999), with sizeable print run and global audience. Full details of acquiring copies, either printed or in identical form online, editorial policies, publication procedure, author submission guidelines, peer review guidelines, legal matters, advantages of publication in this journal and the like can be found by following links from: http://www.herp.net Published by Kotabi Pty Ltd ISSNISSN 1836-5698 1836-5698 (Print) (Print) PO Box 599 ISSN 1836-5779 (Online) Doncaster, Victoria, 3108. ISSN 1836-5779 (Online) Australia. Copyright. All rights reserved. Australasian Journal of Herpetology is also a registered trademark ® in all relevant areas and jurisdictions (Australian trademark number: 1686575). All Intellectual Property (IP) rights are reserved, including in relation to all IP generated by the journal in terms of both authors, publisher and the like. Misuse of the IP by way of illegal over-writing of names formally proposed for taxa for the first timewithin this journal in accordance with relevant rules, or by any means, outside of fair and reasonable scientific discourse is strictly forbidden and legally actionable. Online journals (this issue) do not appear for a month after the actual and listed publication date of the printed journals. Minimum print run of first printings is always at least fifty hard copies.

Hoser 2015 - Australasian Journal of Herpetology 28:1-64 and 29:65-128. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 3 Australasian Journal of Herpetology 31:3-11. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online)

Acanthophis lancasteri Wells and Wellington, 1985 gets hit with a dose of Crypto! … this is not the last word on Death Adder taxonomy and nomenclature.

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 6 September 2015, Accepted 8 September 2015, Published 1 August 2016.

ABSTRACT On the evening of Friday 28 August 2015 (East Australian time), social media was hit with a SPAM attack in the form of wide cross-posting of a PRINO (peer reviewed in name only) Zootaxa paper by a group known as the Wüster gang. Their online paper alleged that the taxon name Acanthophis lancasteri Wells and Wellington, 1985 for the Kimberley Death Adder was a nomen nudem and therefore not available. The paper redescribed the same species as Acanthophis cryptamydros Maddock et al., 2015. This paper argues that Maddock et al. are incorrect and that the authors have engaged in an illegal and creative interpretation of the rules of the International Code of Zoological Nomenclature in order to market their illegal junior synonym. This they have reinforced by hijacking key journals and internet properties for the express purpose of peddling their warped world view on others and without allowing any dissenting views to be aired. In fact less than six days later, a “Google” search for the term “Acanthophis cryptamydros” showed that the group had cross-posted their new name on no less than 3,530 different websites to cement the perception that theirs was the only correct name for the taxon. Furthermore, other fraudulent practices by the same authors in terms of their alleged interpretations of the taxonomy and nomenclature of Death Adders (Genus Acanthophis: Serpentes: Elapidae) are detailed. It is shown that similar acts of taxonomic vandalism by the same group of people with respect to the genus Acanthophis are almost certain. This paper, formally accepts the division of the genus Acanthophis as first proposed by Wells (2002) and in turn names the third major as yet unnamed clade at the subgenus level. There is also a note herein affirming that the name Acanthophis groenveldi Hoser, 2002 is in fact a junior synonym for Acanthophis ceramensis Günther, 1863 and the latter name is the one that should be used. Also noted is that the spellings for the species Acanthophis cummingi Hoser, 1998 and Acanthophis wellsei as first proposed by Hoser in 1998 are correct and intentional name formations. Keywords: Taxonomy; ; nomenclature; taxonomic vandalism; nomen nudem; Death Adder; Elapid; Acanthophis; Aggressiserpens; lancasteri; ceramensis; groenveldi; cryptamydros; taxonomic vandalism; Wüster; Günther; Wells; Wellington; Hoser; Maddock; Gower; new subgenus; platyelapid.

INTRODUCTION Significant recent papers on the genus and the taxonomy include those listed by Hoser (2014) and sources cited therein As iconic Australian snakes, the Death Adders (Genus and they are not relisted herein. Acanthophis Daudin, 1803) are well known to herpetologists globally. The paper of Hoser (2014) effectively resolved the taxonomy and nomenclature of known extant species and subspecies of A detailed account of the genus of the snakes, including life Death Adders, including those forms described by Hoser (1998, history and the like can be found in Hoser (1995) and is not 2002 and 2014) as well as those of Wells and Wellington (1985) repeated here. and this paper makes no alteration to that. That paper (Hoser The taxonomic status of various forms have been scrutinized 2014) should be read before proceeding with this paper. intensely by herpetologists in Australia ever since the genus was However there are matters relevant to the taxonomy and first described. nomenclature of the group not covered in that paper that are Numerous species have been formally described, named, and at dealt with herein. times redescribed and renamed. Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 4 Australasian Journal of Herpetology

1/ The species name Acanthophis groenveldi Hoser, 2002 is in Herein I argue that Maddock et al. are incorrect and that the fact a junior synonym for Acanthophis ceramensis Günther, authors have engaged in an illegal and creative interpretation of 1863 (Günther 1863) and the latter name is the one that should the rules of the International Code of Zoological Nomenclature be used. It is most important that the correct nomenclature is (Ride et al. 1999) in order to market their junior synonym. used and not who is the “name authority”. Unlike members of a This recent publication and relevant issues are discussed after group of thieves known as the Wüster gang (see Hoser 2012a, the formal description of the subgenus Platyelapid subgen. nov. 2012b, 2013, 2015a-f and sources cited therein), I will not break for the smooth-scaled New Guinea Death Adders. the rules of the International Code of Zoological Nomenclature SUBGENUS PLATYELAPID SUBGEN. NOV. (Codes 2-4 as cited and referenced herein) to impose my name authority over a rightful one for personal self-gratification. Type species: Acanthophis laevis Macleay, 1877. I also note herein that the spellings for the species Acanthophis Diagnosis: The genus Acanthophis Daudin, 1803 are readily cummingi Hoser, 1998 and Acanthophis wellsei as first proposed separated from all other elapid snakes by the fact that the tail by Hoser in 1998 are correct and intentional name formations. ends in a soft terminal spine used for the purpose of caudal They should not be amended in any way by any author unless luring. absolutely mandatory under provisions of the relevant zoological The subgenus Platyelapid subgen. nov. are readily separated code. from all other Acanthophis by the following suite of characters: The two names were formed intentionally and to factor in Generally smooth scalation, including on the head and neck relevant issues such as to avoid potential formation of non- (except for some island forms which have some rugosity around homonym names. the head and neck), either an absence of markings on the labials, or if present, only as spots, blotches or peppering, or The name cummingi is in honour of a female person (Fia alternatively the labials are mainly black; ventrals are dark at the Cumming), but her courageous actions in exposing corruption in front (near black) and light (near white) at the rear giving a the NSW, Australian National Parks and Wildlife Service distinct banded appearance, versus immaculate, peppered or (NPWS) took what in Australian slang was “balls’ to do this, a only slightly (indistinctly banded in appearance) in the other two male-type attribute. With this in mind the suffix to the name was subgenera. masculinised. Most if not all Platyelapid subgen. nov. have a prominently For the species name wellsei, in honour of Richard Wells, the raised supraciliary scale, but this trait is also seen to a lesser choice of the strict form “wellsi” was considered, but rejected on degree in other subgenera. While a low subcaudal count (below the basis most people would say it as “wellseyi” and so a 115) appears to be the main character state for Platyelapid spelling broadly equating that was chosen. subgen. nov. species this is not always so. In any event, I hereby act as “first reviser” as per the Distribution: Most of island New Guinea and Islands to the International Code of Zoological Nomenclature (Ride et al. west of there to Ceram and Obi, but not including the Halmahera 1999), and affirm the correct spellings of those names. Island complex. Beyond these statements, nothing further needs to be done Content: Acanthophis (Platyelapid) laevis Macleay, 1877 (type within this paper in terms of these issues. for the subgenus); A. (Platyelapid) barnetti Hoser, 1998; A. 2/ Wells (2002), proposed a division of the genus Acanthophis (Platyelapid) ceramensis Günther, 1863; A. (Platyelapid) along obvious morphological lines, this being the removal of the crotalusei Hoser 1998; A. (Platyelapid) macgregori Hoser, 2002; Acanthophis pyrrhus group and placement within a new genus A. (Platyelapid) yuwoni Hoser, 2002. he erected called Aggressiserpens Wells, 2002. Based on the ACANTHOPHIS LANCASTERI WELLS AND WELLINGTON, deep phylogenetic divergence of the group from the other 1985 GETS HIT WITH A DOSE OF CRYPTO! Acanthophis, the judgement of Wells has merit. However it is my considered opinion that the relevant group would be better “Crypto” is shorthand or slang among keepers for the treated as a subgenus and so this is the case herein. Cryptosporidium, a genus of protozoans that cause gastrointestinal disease and often death in snakes. This conservative judgement is made noting that to date there has been no comprehensive molecular phylogeny of Hence the poetic license in the statement “Acanthophis Acanthophis as widely recognized with a comparison to other lancasteri Wells and Wellington, 1985 gets hit with a dose of elapid genera and subgenera in order to best escertain whether Crypto!” in view of the fact that the species name “Acanthophis or not Aggressiserpens should be treated as a subgenus or full lancasteri” has been attacked and perhaps fatally so. genus. That is at least the hope of the proponants of the new name. This is the same position and contention made in Hoser (2014), Coincidentally the newly proposed name to replace Acanthophis at page 24, where I further note that at one point in the lancasteri Wells and Wellington, 1985 is “Acanthophis discussion a typographical error led to Aggressiserpens being cryptamydros Maddock et al. 2015”. identified as a subspecies and not a subgenus, although Hence the “Crypto” line. elsewhere in the paper, the correct status of the name was The sequence of events relating to the taxon, Acanthophis given. lancasteri, better known as the species of Death Adder from the Within the ambit of a subgeneric break-up of Acanthophis as Kimberley Ranges of north-west Western Australia can be presently recognized, the third major lineage of Death Adders, summarised by the following dateline. this being the New Guinea / Indonesian group with smooth 1985 - Wells and Wellington published the name for the species scales and reduced ventral count need also to be placed within taxon in Australian Journal of Herpetology via what appeared to their own subgenus and so this is done within this paper. be a fairly standard, albeit brief species description. 3/ On the evening of Friday 28 August 2015 (East Australian 1987 - Richard Shine as “The President, Australian Society of time), social media was hit with a SPAM attack in the form of Herpetologists” attempted to suppress the entire contents of the wide cross-posting of a PRINO (peer reviewed in name only) relevant journal by a petition to the International Commission of Zootaxa paper by a group known as the Wüster gang. Their Zoological Nomenclature or ICZN. online paper alleged that the taxon name Acanthophis lancasteri 1991 - The ICZN rejected the petition by Shine and over a Wells and Wellington, 1985 for the Kimberley Death Adder was hundred other supporters and ruled in favour of the Wells and a nomen nudem and therefore not available. The paper Wellington journal stating that the names were nomenclaturally redescribed the same species as Acanthophis cryptamydros available. Maddock et al., 2015. 1998 - Hoser published a genus-wide revision of the Death

Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 5

Adders (Acanthophis) and used the name Acanthophis Acanthophis species.” lancasteri Wells and Wellington, 1985 for the relevant taxon. This leads one directly to the paper of Aplin and Donnellan 1999 - Without giving a proper reason, Ken Aplin (Aplin 1999) (1999), which clearly most readers of Maddock et al. (2015) and Aplin and Donnellan (1999) stated that Acanthophis would not do, but I in fact did. lancasteri Wells and Wellington, 1985 was a nomen nudem and The relevant passage in Aplin and Donnellan (1999) read as therefore not available to be used for the relevant taxon. This he follows: repeated, but at no stage explained his position. “The nomenclature of Acanthophis has been impacted by two 2001 - ICZN again ruled in favour of the Wells and Wellington works published by ‘amateur’ herpetologists in unrefereed journals following a second suppression attempt by Robert contexts. Wells and Wellington (1985) proposed four additional Sprackland, Pete Strimple and Hobart Smith. species of Acanthophis in their essentially self-published 2002 - Shea repeated the nomen nudem claim in an email and “Classification of the Amphibia and Reptilia of Australia”. Three this was published by Hoser (2002), who while publishing the of these proposed taxa (armstrongi, lancasteri, schistos) were comments in a further revision of Acanthophis, did not take a based solely on Storr’s (1981) figures and descriptions of each definitive position one way or other and chose not to rename the of the three Western Australian populations; these are nomina relevant taxon (or others similarly alleged to be nomen nudem). nuda because they do not include or point to previously 2012 - Wüster and associates circulated a petition globally published differential diagnoses. The fourth Wells and (Kaiser 2012, Kaiser et al. 2012) seeking that herpetologists Wellington name, A. hawkei, proposed for the ‘Barkly Adder’, ignore the rules of the International Code of Zoological minimally satisfies the conditions for ‘availability’ as set out by Nomenclature and rename taxa properly named by Hoser and the International Code of Zoological Nomenclature (1985). Wells. This claim has been amended in 2013 (Kaiser 2013), However, the taxon has not been adequately diagnosed and for Kaiser et al. 2013) and again in 2014 (Kaiser 2014a, 2014b) and the present is best treated as a junior synonym of A. most recently in 2015 (Rhodin et al. 2015), as detailed by Hoser antarcticus.” (2015). The relevant statement herein is: 2013 - Following the urging of Wüster and associates, their “these are nomina nuda because they do not include or point to friends start renaming dozens of taxa properly named by Hoser previously published differential diagnoses.” and Wells and Wells and Wellington in breach of the rules of the This brings us to the “International Code of Zoological International Code of Zoological Nomenclature, usually merely Nomenclature (1985)” as cited by Aplin for an explanation, which citing the Kaiser et al. (2013) “point of view” as a veto to enable I might add is highly unlikely to be consulted by a casual reader them to step outside the rules (e.g. Schleip 2014, who after of Maddock et al. (2015). recognizing Leiopython hoserae Hoser, 2014 as valid for some The terms nomen nudem, or the plural nomina nuda are defined six years chose to rename it L. meridionalis Schleip, 2014) in in the relevant codes, and in the current, fourth edition the one of the most blatant attempts to steal name authority in all following is written: the history of Zoology. “The provisions of this Code supersede those of the previous 2014 - Hoser published an updated revision of Acanthophis, editions with effect from 1 January 2000 … naming new taxa, but following cross referencing the Wells and nomen nudum (pl. nomina nuda), n. Wellington description/s of 1985 with the relevant parts and definitions in the International Code of Zoological Nomenclature A Latin term referring to a name that, if published before 1931, (as also done in this paper), this time took the position that fails to conform to Article 12; or, if published after 1930, fails to Acanthophis lancasteri Wells and Wellington, 1985 was not a conform to Article 13. A nomen nudum is not an available name, nomen nudem according to any of editions 2-4 of the and therefore the same name may be made available later for International Code of Zoological Nomenclature (cited herein). the same or a different concept; in such a case it would take authorship and date [Arts. 50, 21] from that act of establishment, 2015 - On the evening of 28 August, Wolfgang Wüster and co- not from any earlier publication as a nomen nudum.” authors launched an internet blitz promoting their new paper in PRINO (peer reviewed in name only) Journal Zootaxa that The preceding leads us to Article 13 of the Code, and here the alleged (without proper explanation) that Acanthophis lancasteri audit becomes more interesting as there are potentially three Wells and Wellington, 1985 was nomen nudem and that they issues of the Code to deal with. had renamed the same taxon as Acanthophis cryptamydros At the time of the publication of the Wells and Wellington paper, Maddock et al. 2015. the second edition of the code was in force. The third edition While the preceding timeline sets out the sequence of events carried a publication date of February 1985, but it was not relevant to the naming and use of the taxon name Acanthophis actually printed until 1988 based on date stamps on library lancasteri, and the associated issue of Wüster and associates copies, including that posted online by the Smithsonian in the seeking to steal name authority for other people’s taxa, the only USA. relevant issue in terms of Acanthophis lancasteri is whether or The fourth edition, published in 1999, has an explicit statement not the Wells and Wellington description of 1985 is valid that it supersedes the rest in any event. according to the rules of the International Code of Zoological However Article 13 is much the same in each code. Nomenclature. So there can be no doubt as to what is said in each edition, I The paper Maddock et al. (2015) in summary remanufactured copy them in full below: well known information about Acanthophis as “new” research Code 2nd Edition 1964 with the simple objective of renaming A. lancasteri. ”Article 13 The relevant passage in their paper read as follows: Article 13. Names published after 1930. “The consistent differences between the Kimberley death adders (a) Names in general. and all other Acanthophis across three independent genetic loci, morphology, and color pattern lead us to conclude that these In addition to satisfying the provisions of Article 11, a name populations represent a separate species from all other published after 1930 must either be: Australian Acanthophis. Since the only existing name applicable (i) accompanied by a statement that purports to give to this taxon, Acanthophis lancasteri Wells and Wellington, characters differentiating the taxon; or 1985, is a nomen nudum (Aplin and Donnellan 1999), we (ii) accompanied by a definite bibliographic reference to such a describe it as a new species below, diagnosing it from its statement; or congeners and all other currently recognized Australian Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 6 Australasian Journal of Herpetology

(iii) proposed expressly as a replacement for a pre-existing words characters that are purported to differentiate the available name. taxon” (b) Genus-group names. A genus-group name published after Code 4th Edition 1999 (2000) 1930 must, in addition to satisfying the provisions of Section (a), “accompanied by a description or definition that states in be accompanied by the definite fixation of a type-species [Art. words characters that are purported to differentiate the 68].” taxon” Code 3rd Edition 1985 (1988) The significant part of each section, never quoted by either Aplin and Donellan (1991) or of course Maddock et al. (2015), “Article 13. Names published after 1930. summarised in the table immediately above is the use of the (a) Requirements. word “purports” or “purported”. To be available, every new scientific name published after 1930 The word purport is not defined in any edition of the code, but it must satisfy the provisions of Article 11, and must be: is in most dictionaries and online as well, where on 3 September (i) accompanied by a description or definition that states in 2015 Google defined it as follows: words characters that are purported to differentiate the “appear to be or do something, especially falsely.” taxon, or In other words, even if the Wells and Wellington description for (ii) accompanied by a bibliographic reference to such a Acanthophis lancasteri pointed to a document that did not carry published statement even if contained in a work published a description or diagnosis, the mere fact their description before 1758 or that is not consistently binominal (for information purported this, means that it is valid under any of the three excluded for reasons of anonymity after 1950 see Article 14), or relevant editions of the code. (iii) proposed expressly as a new replacement name (nomen Now just to remove any doubt at all as to the nomenclatural novum) for an available name. availability of Acanthophis lancasteri and that the original Recommendation 13A. Comparisons.—In describing a new description did “purport” to “differentiate the taxon” (Code edition nominal taxon, an author should make his intention to 3), I copy the description in its entirety within this paper. differentiate clear to others by giving a summary of characters Elsewhere within a separate as yet unpublished paper, Ross that in the author’s opinion differentiate the taxon from other Wellington summed up the situation when he wrote: named taxa of the same rank as the new taxon. “By any reasonable objective interpretation of the Code Rule in (b) Genus-group names.—Every new genus-group name relation to the description of Acanthophis lancasteri Wells and published after 1930 (but not a name published at any time for a Wellington 1985, it is described. The ICZN (1991) has ruled that collective group or an ichnotaxon [Art. 66]) must, in addition to Wells and Wellington 1984 and 1985 are publications and are satisfying the provisions of Section a of this Article, be available for nomenclatural purposes. The description for accompanied by the fixation of a type species for that nominal Acanthophis lancasteri does provide a Holotype WAM R70690 genus-group taxon by original designation [Art. 68b] or by from a Type locality of 45 km NNE of Halls Creek, WA. Also in indication [Arts 67h, 68c-e]. accordance with the above Article 13 and in contradiction to Aplin (1999); Aplin and Donnellan (1999) and by implication also (i) If the name of a genus-group taxon established before 1931 Maddoock, Ellis, Doughty, Smith, and Wüster (2015) who relied is replaced after 1930, the type species of that nominal taxon upon Aplin’s (and Donnellan) incorrect assertions, the must then be designated, if that has not already been done.” description of Acanthophis lancasteri in fact does provide a Code 4th Edition 1999 (2000) statement that purports to show difference between the then “Article 13. Names published after 1930. new species and other species in the Acanthophis complex, it 13.1. Requirements. To be available, every new name published also provides further information, other references and to after 1930 must satisfy the provisions of Article 11 and must defined published source information in support of the purported 13.1.1. be accompanied by a description or definition that difference statement. For example it also provides further states in words characters that are purported to differentiate interpolative information in the description of Acanthophis the taxon, or hawkei (same paper) as well as in the references section. The W&W description demonstrates, unequivocally the entity to 13.1.2. be accompanied by a bibliographic reference to such a which the description applies. As with any description additional published statement, even if the statement is contained in a information could have been provided but the description, work published before 1758, or in one that is not consistently although brief, as it stands did conform to the minimum binominal, or in one that has been suppressed by the requirements (then required) of a valid description and hence is Commission (unless the Commission has ruled that the work is available.” to be treated as not having been published [Art. 8.7]), or Wells and Wellington (1999) also published a direct rebuttal to 13.1.3. be proposed expressly as a new replacement name Aplin (1999) and his claims against their taxon Acanthophis (nomen novum) for an available name, whether required by any lancasteri, but this was evidently deliberately ignored by Wüster provision of the Code or not. and his gang. Recommendation 13A. Intent to differentiate. When describing a In other words Acanthophis lancasteri is available for the taxon new nominal taxon, an author should make clear his or her and is the name that must be used. purpose to differentiate the taxon by including with it a diagnosis, that is to say, a summary of the characters that differentiate the Acanthophis cryptamydros Maddock et al. 2015 is merely a new nominal taxon from related or similar taxa. junior synonym of the former and should not be used. Recommendation 13B. Language. Authors should publish BAD MOTIVE ON THE PART OF MADDOCK AND WÜSTER. diagnoses of new taxa in languages widely used internationally Of course one needs evidence to assert such a thing. After all, in zoology. The diagnoses should also be given in languages one may assume for a moment that Maddock et al. published used in the regions relevant to the taxa diagnosed.” their incorrect assessment of the nomenclatural validity of ICZN Code Edition Acanthophis lancasteri due to a human error or inadvertent Proposed New Name Requirements failure to check the relevant parts of the International Code of Code 2nd Edition 1964 Zoological Nomenclature. “accompanied by a statement that purports to give That we know this is not the case and that they have acted at all characters differentiating the taxon” times with improper motive comes from the mouth of Maddock Code 3rd Edition 1985 (1988) himself (Proud 2015). “accompanied by a description or definition that states in On the website at: Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 7

http://www.nhm.ac.uk/our-science/science-news/2015/august/ (supervisor Dr David Gower)” new-highly-venomous-snake-species-discovered-in- Now who is David Gower? australia.html A quick search on “Google” shows he is listed as an editor at the with a posting date of 28 August 2015, is the statement: PRINO journal Zootaxa! “A team led by a Natural History Museum scientist has So there you have it! discovered a new species of highly venomous Australian death Wüster and an ex-student conspire to steal name rights for a adder in the Kimberley region of the country.” species and then have their paper published in a journal where a and co-worker Gower, who works with Maddock is able to bypass “The team, which included researchers from Bangor University any credible quality control. and the Western Australian Museum, identified the new species Nowhere in the relevant Maddock et al. paper is this critically while researching the genetics and ecological characteristics of important conflict of interest disclosed. snakes living in the Kimberley region.” Now this of course doesn’t explain the role of the other listed co- We know these statements to be false because even back in authors, but this is easily ascertained. 1985 when Wells and Wellington first formally named Ryan J. Ellis works with Wüster at Bangor University and plays a (discovered?) the same species, it was well known in Australia key role in creating the spiffy looking graphics you see in his and I had also caught and kept them for some years prior to that papers, so in gratitude, Wüster has him listed as a co-author. date! The other two authors, Paul Doughty and Laurie Smith, Now noting I had confirmed the existence of the species employed at the Western Australian Museum have long been at “discovered’ by Wells and Wellington in 1985 in my papers in loggerheads with Wells and Wellington, including in the failed 1998, 2002 and again in 2014, (not that Wells and Wellington attempt to have the relevant publication suppressed by the ICZN ever had the audacity to claim they were the ones who first in the 1980’s and 1990’s. discovered them), the claim by Maddock, Wüster and their gang to have discovered this species must be patently false! Smith also described a species of python from Western Australia calling it “Liasis stimsoni”. The problem for him was So not only have Maddock et al. misrepresented the provisions that it was a junior synonym for Antaresia saxacola Wells and of the International Code of Zoological Nomenclature to steal Wellington, 1985, named some months earlier. “name authority” for a species of snake, but then they have publicly lied about their claim to have “discovered a new species In order to discourage usage of the correct Wells and Wellington of highly venomous Australian death adder in the Kimberley name, Smith and others at the Western Australian Museum region of the country”. actively supported the push to suppress the Wells and Wellington publications of 1984 and 1985 while simultaneously HOW THEN DID THE PAPER OF MADDOCK ET AL. (2015) aggressively marketing his own name on the basis that the GET PUBLISHED? attempted suppression of Wells and Wellington’s would Zootaxa alleges it is a “peer reviewed” scientific journal. succeed. Hoser (2015d) and sources cited therein give numerous By the time this suppression attempt failed in 1991, Smith’s examples of evidence to show that Zootaxa has never had name was already in widespread usage, while the Wells and anything resembling a proper peer review quality control system Wellington one effectively unused. in place. After I corrected this anomaly in Hoser (2000), and expecting In fact Zootaxa is a holotype PRINO (peer reviewed in name others to follow the logical and correct course of using the only) journal. correct senior synonym, a new claim was erected to allege that However this paper deals specifically with Maddock et al. (2015) the Wells and Wellington name “saxacola” was “nomen nudem”, and it is here I point out the obvious failings. which from any cross referencing of the description with any Had there been proper quality control, the reviewers would have edition of the International Code of Zoological Nomenclature followed the simple intellectual exercise I have now done several (editions 2-4) is clearly not the case. times in order to ascertain the legal availability of the name I also note that it is obvious that few if any lay people would Acanthophis lancasteri. have the inclination or capacity to do this, noting that in year With the entire substantive basis of Maddock et al. (2015) being 2000, the Code was not available online and hard copies to rename the taxon (the rest of the paper’s text is effective relatively rare outside of natural history musems. padding for that), had a reviewer done the relevant exercise of Hence it would not come as a surprise to find that these men cross-checking they’d have found that the name Acanthophis would jump at the chance to be listed as coathors in a paper that lancasteri was nomenclaturally available and rejected the had them steal yet another west Australian species name from Maddock et al. paper. Wells and Wellington. So it becomes relevant as to who actually edited and reviewed Plus of course they supplied the holotype for the allegedly “new” the paper at Zootaxa. species. First we deal with the listed authors. WHY THIS IS NOT THE LAST WORD ON DEATH ADDER Maddock, until now effectively unknown in herpetology, turns out TAXONOMY AND NOMENCLATURE! to be a recently graduated student of University lecturer Wüster and his gang have access to modern molecular methods Wolfgang Wüster at Bangor University, Wales, UK. to assist in ascertaining relationships between taxa. Maddock et He now has a position at the Natural History Museum in London, al. (2015) has used molecular data to “validate” their taxonomic UK. conclusion that they are naming a new species (ignoring the fact We know all this from his website at: that buried in the text of their paper is an oblique statement to http://www.ucl.ac.uk/~ucbtjjd/Site/Simon.html the effect they are stealing “name authority” from Wells and On that webpage he states: Wellington. “Simon graduated from the University of Wales, Bangor with a However there are some key facts worth noting in all this. Master of Zoology in 2011.” Large charismatic vertebrates, including Death Adders do not That confirms Wüster was his teacher and a close associate. need the services of molecular biologists to work out which The webpage further states: species is which. “Currently Simon is working towards his PhD, which is joint They are easily delineated by simply looking at the snakes between UCL and the Natural History Museum, London themselves.

Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 8 Australasian Journal of Herpetology

The same is the case for other species such as White-lipped Rather than producing material and data that they had available Pythons (genus Leiopython), which is another group the Wüster to them, that confirms the validity of the species Acanthophis gang have tackled with their taxonomic vandalism and barnetti Hoser, 1998, and using the correct nomenclature, nomenclatural misconduct along with selective use and non-use Maddock et al. chose to withhold such information. of molecular data (see Hoser 2009). However it was posted on Facebook in the week following the In the molecular results published by Maddock et al., the authors publication of Maddock et al., that the same authors were have conveniently omitted to show any molecular data that plotting ways they could try to steal name authority for that taxon validated taxa named by myself, even though they had such as well. material available. This included specimen data for either Hence we know that Maddock et al. is not the last word from the described subspecies of Acanthophis wellsei, namely A. wellsei Wüster gang on Death Adder taxonomy. hoserae Hoser, 2014 and A. wellsei donnellani Hoser, 2002, or DEALING WITH THE CRYPTO INFECTION! the northern New Guinea taxa, Acanthophis barnetti Hoser, 1998 and A. crotalusei Hoser, 1998. Within hours of Wüster and his gang posting and promoting links to his co-authored paper on “Facebook” on Friday 28 Of course there has been published molecular data for species August 2015 (east Australian time), Wells had published an groups with parallel distributions to these taxa and all have extensive rebuttal of the claims in the 2015 paper on Facebook validated the obvious species divisions. which was read and answered by both Maddock and Wüster. Wüster of course has been playing mental gymnastics for years They ignored this and continued to peddle their new name and to avoid having to recognize any taxa formally named by myself paper, as if it were the unimpeachable gospel. and this involves some ridiculous propositions, including that A. barnetti and A. crotalusei are merely variants of A. laevis. However the reach of Wüster and his gang in peddling their lies As of 3 September 2015, on peter Uetz’s “The Reptile and falsehoods was best demonstrated when less than six days Database” which Wüster effectively controls in materially later in the morning of 3 September 2015 (east Australian time), relevant ways, one sees for the entry for “Acanthophis wellsi I did a “Google” search for Acanthophis cryptamydros. Hoser, 1998” (sic), the following text: It showed that the group had cross-posted their new name on no “Synonymy: Not listed by COGGER 2000. The name was less than 3,530 different websites to cement the perception that emended to wellsi as the species was described in honor of theirs was the only correct name for the taxon and that they had Richard Wells. Acanthophis wellsi donnellani HOSER 2002 may discovered the species themselves. be a synonym of A. wellsi (WÜSTER, pers. comm. 15 Dec By contrast the valid name (lancasteri) was shown on only half 2010).” that number of webpages, even though it had been around for Of course material Wüster and co-authors themselves had on 30 years! hand that they should have published in Maddock et al. could This is perhaps the most stark example of extremist taxonomic have easily refuted his bogus claim that “Acanthophis wellsi vandalism and nomenclatural misconduct and the power of donnellani HOSER 2002 may be a synonym of A. wellsi”. reach by those who engage in these activities, seen to date in It is also worth mentioning that Cogger (2014) did in fact the age of internet and rapid dissemination of information, both recognize A. wellsei (spelt properly I might add) a fact Wüster incorrect and correct. and sidekick Uetz have conveniently chosen to ignore. It shows how by use of social media including via the dark art of Significant however is that without so much as a statement that Search Engine Optimisation (SEO), hijacking sites like he had been lying about Acanthophis wellsei and it’s alleged Wikipedia (also attacked by Wüster’s gang within hours of their synymy with A. pyrrhus Boulenger, 1898 for the previous 17 paper coming out) altered to reflect their warped dreams of total years, the coauthor Wüster accepts and uses the name “A. hegemony in terms of reptile taxonomy and nomenclature, the wellsi” (sic) for a taxon he (now) regards as valid in Maddock et group can and does dupe people into believing that their illegal al. (2015). minority view is in fact correct and consensus in methods There is also the issue of the molecular data Maddock et al. accurately detailed by Dubois (2015). present for A. wellsei in their paper on page 306. Data from Contrary to various claims made, I have no vested interest in snakes from three different locations is shown, implying all are Wells, Wellington or their taxonomy and nomenclature. the same species and with minimal divergence between the My only concerns are with the science and the rules of samples. engagement, these being published in the International Code of The number of course matches the three forms I have described Zoological Nomenclature. (in 1998, 2002 and 2014). If and when Wells and Wellington get things right, I support What is not readily disclosed and only becomes clear when the them. If they get it wrong I condemn them, or I don’t agree with named locailites are plotted against a map is that all come from them, I deal with that appropriately. the main range of the nominate subspecies of A. wellsei and While it is easy to identify defects in the description Acanthophis that none of the samples include the more recently described lancasteri, (my view is it is lousy), the fact remains it complied subspecies. with the rules of the Code and also was typical of others of the Wüster assisted his mate Wulf Schleip in 2008 when he time (1980’s and earlier), and in fairness to the authors should published his paper on Leiopython which produced molecular be viewed in that context. evidence to confirm the obvious fact that those from south of the As seen many times past, Wüster and his gang have more time New Guinea central range were a different species to those from and internet savvy than their opponents. the north. They have hijacked control of key internet properties such as (Signicant that time was that they did not publish molecular data “Wikipedia” and “The Reptile Database’ to peddle their distorted they had obtained showing all the brown Leiopython from north world views. In the offline world, their group has hijacked of the central range were one and the same species, with editorial influence in several formerly well-regarded scientific Schleip claiming in the paper and ever since to have journals in order to bypass proper peer review to get their “discovered” several new species). material published as fast as they write it. Noting that the barrier affecting those snakes is the same as for As seen by the many examples published in Hoser (2012a, Death Adders, even before one inspects the very different 2012b, 2013, 2015a-f) and sources cited therein, Wüster and his snakes from north and south of the range, it is clear that the gang of thieves will not voluntarily stop their attack on the rules Acanthophis from each side are different species. of the International code of Zoological Nomenclature in order to

Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 9

Never letting the truth get in the way of their lie ... Within days, Wolfgang Wüster and his band of thieves had plastered their illegal name Acanthophis cryptamydros all over the internet to swamp the correct name Acanthophis lancasteri. This was in order to convince the world that they had indeed discovered a new species and that their name was the correct one. This is seen by the number of online uses via a Google search on 3 September 2015.

Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 10 Australasian Journal of Herpetology steal name authority for names from authors everywhere. and his gang of thieves. Then the ICZN commissioners Perhaps the best to make this potentially happen is via a themselves could spend more time dealing with their real strongly worded ruling by the ICZN against nomenclatural passion, that being taxonomy! misconduct by the Wüster gang or like-minded individuals. REFERENCES CITED Victims of the gang include myself, who have been falsely Aplin, K. P. 1999. Amateur Taxonomy in Australian Herpetology - accused of stealing the work of others (only they have done that) Help or hindrance? Monitor - Journal of the Victorian and this problem, real or perceived needs to be dealt with. Herpetological Society 10(2/3):104-109. In fact the actions of the Wüster gang and like-minded Aplin, K. P. and Donnellan S. C. 1999. An extended description individuals has created a ridiculous situation whereby scientists of the Pilbara Death Adder, Acanthophis wellsi Hoser are wasting an inordinate amount of time arguing over name (Serpentes: Elapidae), with notes on the Desert Death Adder, A. authority for taxa that was properly named long ago, while pyrrhus Boulenger, and identification of a possible hybrid zone. thieves try to creatively interpret the rules and steal yet more Records of the Western Australian Museum 19:277-298. validly named taxon “name authority” from others. This is all Cogger, H. G. 2014. Reptiles and Amphibians of Australia happening when scientists should be more properly dealing with (Seventh edition), CSIRO. Sydney, Australia:1064 pp. the science of taxonomy and describing biodiversity before it is killed off by the human population explosion. Daudin, F. M. 1802. Histoire Naturelle, Générale et Particulière des Reptiles, Vol. 4. F. Dufart, Paris. However the problem of name authority disputes can in fact be easily solved via a revamped system of establishing availability Dubois, A. 2015. Zoological nomina in the century of extinctions: of names in Zoological Nomenclature. new proposals. Bionomina, 8:11-53. New names could be registered in a similar manner to that used Günther, A. 1863. Contribution to the herpetology of Ceram. worldwide for trademark registration. Proc. Zool. Soc. London 1863:58-60. Via an automated online system, new names could be submitted Hoser, R. T. 1995. Australia’s Death Adders, Genus (at the time the scientist first seeks to potentially name taxa) Acanthophis, The Reptilian 3(4)7-21 and cover, 3 (5):27-34. with a time limit imposed to publish a paper formally describing Hoser, R. T. 1998. Death Adders (Genus Acanthophis): An the taxon or taxa and satisfy the relevant code requirements (the overview, including descriptions of Five new species and One code currently recommends a year and that could be made subspecies. Monitor: Journal of the Victorian Herpetological mandatory). The publication is then also submitted and checked Society 9(2):20-41. for form by an examiner against the rules of the Code as is done Hoser, R. T. 2002. Death Adders (Genus Acanthophis): An for trademarks. Updated overview, including descriptions of 3 New Island They check against a trademarks registration manual. species and 2 New Australian subspecies. Crocodilian:Journal In line with the current rules, the ICZN would restrict its ambit to of the Victorian Association of Amateur Herpetologists nomenclature and not taxonomy. September 2002:5-11,16-22,24-30, front and back covers. A fee could be imposed to cover the costs of the system, with Hoser, R. T. 2009. Creationism and contrived science: A review fee waiver provisions for those unable to pay. of recent python systematics papers and the resolution of issues of taxonomy and nomenclature. Australasian Journal of In fact the ICZN could even run the system at a profit to cover Herpetology 2:1-34. the ongoing administration costs of the entity. Hoser, R. T. 2012a. Exposing a Fraud! Afronaja Wallach, Wüster As with trademark registrations, there could be an “opposition” and Broadley 2009, is a junior synonym of Spracklandus Hoser period, whereby people opposing registration could lodge 2009! Australasian Journal of Herpetology 9:1-64. objections (subject to the rules) and argue their cases, with the ICZN making a decision one way or other and before the name Hoser, R. T. 2012b. Robust taxonomy and nomenclature based even becomes “legal”. on good science escapes harsh fact-based criticism, but remains unable to escape an attack of lies and deception. As with trademarks, the names can be used pending registration Australasian Journal of Herpetology 14:37-64. or non-registration, with registration back-dated to application or other specified date after registration takes place. Hoser, R. T. 2013. The science of herpetology is built on evidence, ethics, quality publications and strict compliance with In line with trademarks, non registration of a name would mean the rules of nomenclature. Australasian Journal of Herpetology it could not be used as intended. 18:2-79. All this would limit the ongoing instability created by the use or Hoser, R. T. 2014. Tidying up Death Adder taxonomy non-use of names some people assert are not code-compliant (Serpentes: Elapidae: Acanthophis): including descriptions of and would have prevented or resolved such issues like the new subspecies and the first ever key to identify all recognized validity of Acanthophis lancasteri, at the time it was proposed species and subspecies within the genus. Australasian Journal and not 30 years later and with the full-blown intervention of the of Herpetology 23:22-34. ICZN commissioners themselves. Hoser, R. T. 2015a. Dealing with the “truth haters” ... a If necessary a limit system could be used to prevent persons or summary! Introduction to Issues 25 and 26 of Australasian groups monopolizing taxa, making ambit claims or in any way Journal of Herpetology. Including “A timeline of relevant key unfairly preventing others from using the system. publishing and other events relevant to Wolfgang Wüster and Such a system would accurately identify who first publicly his gang of thieves.” and a “Synonyms list”. Australasian Journal identified themselves as working on given taxa, thereby enabling of Herpetology 25:3-13. accusations of theft of work or ideas to be easily checked and refuted or accepted. Hoser, R. T. 2015b. The Wüster gang and their proposed “Taxon Filter”: How they are knowingly publishing false information, A system of name registration similar to that employed by recklessly engaging in taxonomic vandalism and directly trademarks offices worldwide, including those nations signatory attacking the rules and stability of zoological nomenclature. to the Madrid protocol would not only significantly improve the Australasian Journal of Herpetology 25:14-38. nomenclature system for new names of taxa, but also reduce the unnecessary dispute resolving workload of scientists and Hoser, R. T. 2015c. Best Practices in herpetology: Hinrich ICZN Commissioners alike. Kaiser’s claims are unsubstantiated. Australasian Journal of Herpetology 25:39-52. In fact, the ICZN would no longer have to regularly deal with cases of thieves trying to steal the work of others as has Hoser, R. T, 2015d. PRINO (Peer reviewed in name only) become common in recent years due to the actions of Wüster journals: When quality control in scientific publications fails. Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 11

Australasian Journal of Herpetology 26:3-64. History Museum - Cromwell Road, London SW7 5BD, UK (also Hoser, R. T, 2015e. Comments on Spracklandus Hoser, 2009 commonly cited as “The Rules”, “Zoological Rules” or “ICZN (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the 1985”) (the document appears to have been first printed in 1988 availability of the generic name and for the nomenclatural based on a receipt from the Smithsonian Library stamped on validation of the journal in which it was published (Case 3601; their online pdf of the document). see BZN 70: 234-237; comments BZN 71:30-38, 133-135). Ride, W. D. L. (ed.) et al. (on behalf of the International (Unedited version) Australasian Journal of Herpetology 27:37- Commission on Zoological Nomenclature) 1999. International 42. code of Zoological Nomenclature (Fourth edition). The Natural Hoser, R. T. 2015f. Rhodin et al. 2015, Yet more lies, History Museum - Cromwell Road, London SW7 5BD, UK (also misrepresentations and falsehoods by a band of thieves intent commonly cited as “The Rules”, “Zoological Rules” or “ICZN on stealing credit for the scientific works of others. Australasian 1999”). Journal of Herpetology 27:3-36. Stoll, N. R. (ed.) et al. (on behalf of the International Commission on Zoological Nomenclature) 1964. International ICZN 1991. Three works by Richard W. Wells and C. Ross code of Zoological Nomenclature (Second edition). The Natural Wellington: proposed suppression for nomenclatorial purposes. History Museum - Cromwell Road, London SW7 5BD, UK (also Bulletin of Zoological Nomenclature 48(4): 337-338. commonly cited as “The Rules”, “Zoological Rules” or “ICZN Kaiser, H. 2012. SPAM email sent out to numerous recipients on 1964”). 5 June 2012. Wells, R. W. 2002. Taxonomy of the Genus Acanthophis Kaiser, H. et al. 2012. Point of view. Hate article sent as (Reptilia: Elapidae) in Australia. Australian Biodiversity Record attachment with SPAM email sent out on 5 June 2012. 2002(5)(March):18. Kaiser, H. 2013. The Taxon Filter, a novel mechanism designed Rhodin, A. et al. (70 listed authors) 2015. Comment on to facilitate the relationship between taxonomy and Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): nomenclature, vis-à-vis the utility of the Code’s Article 81 (the request for confirmation of the availability of the generic name Commission’s plenary power). Bulletin of Zoological and for the nomenclatural validation of the journal in which it was Nomenclature 70(4) December 2013:293-302. published (Case 3601; see BZN 70: 234-237; 71: 30-38, 133- Kaiser, H. 2014a. Comments on Spracklandus Hoser, 2009 135, 181-182, 252-253). Bulletin of Zoological Nomenclature (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the 72(1)65-78. availability of the generic name and for the nomenclatural Schleip, W. D. 2008. Revision of the Genus Leiopython validation of the journal in which it was published (Case 3601; Hubrecht 1879 (Serpentes: Pythonidae) with the Redescription see BZN 70: 234-237). Bulletin of Zoological Nomenclature of Taxa Recently Described by Hoser (2000) and the Description 7(1):30-35. of New Species. Journal of Herpetology 42(4):645-667. Kaiser, H. 2014b. Best Practices in Herpetological Taxonomy: Schleip, W. D. 2014b. Two new species of Leiopython Hubecht Errata and Addenda. Herpetological Review, 45(2):257-268. (sic), 1879 (Pythonidae: Serpentes): Non-compliance with the Kaiser, H., Crother, B. L., Kelly, C. M. R., Luiselli, L., O’Shea, M., International Code of Zoological Nomenclature leads to Ota, H., Passos, P., Schleip, W. D. and Wüster, W. 2013. Best unavailable names in zoological nomenclature. Journal of practices: In the 21st Century, Taxonomic Decisions in Herpetology 48(2):272-275. Herpetology are Acceptable Only When supported by a body of Wells, R. W. and Wellington, C. R. 1984. A synopsis of the class Evidence and Published via Peer-Review. Herpetological Reptilia in Australia. Aust. Journal of Herpetology 1(3-4):73-129. Review 44(1):8-23. Wells, R. W. and C. R. Wellington. 1985. A classification of the Maddock, S. T. 2015. Webpage blog at: http://www.ucl.ac.uk/ Amphibia and Reptilia of Australia. Australian Journal of ~ucbtjjd/Site/Simon.html Herpetology Supplementary Series 1:1-61. Maddock, S. T., Ellis, R. J., Doughty, P., Smith, L. A. and Wells, R. W. and Wellington, C. R. 1999. A response to Ken Wüster, W. 2015. A new species of death adder (Acanthophis: Aplin’s article on Herpetology in Australia. Monitor:Journal of the Serpentes: Elapidae) from north-western Australia. Zootaxa Victorian Herpetological Society 10(2/3):110-112. 4007(3):301-326 online. Wüster, W., Broadley, D. G. and Wallach, V. 2014. Comments Proud, L. 2015. New highly venomous snake species discovered on Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): in Australia. Post on internet at: request for confirmation of the availability of the generic name http://www.nhm.ac.uk/our-science/science-news/2015/august/ and for the nomenclatural validation of the journal in which it was new-highly-venomous-snake-species-discovered-in- published (Case 3601; see BZN 70: 234-237). Bulletin of australia.html Zoological Nomenclature 7(1):37-38. Ride, W. D. L. (ed.) et al. (on behalf of the International CONFLICT OF INTEREST Commission on Zoological Nomenclature) 1985. International code of Zoological Nomenclature (Third edition). The Natural The author has no relevant conflicts of interest.

Australasian Journal of Herpetology ® Publishes original research in printed form in relation to reptiles, other fauna and related matters, including the subjects of classification, ecology, public interest, captivity, “academic misconduct”, etc. It is a peer reviewed printed journal published in hard copy for permenant public scientific record in accordance with the International Code of Zoological Nomenclature (Ride et al. 1999), with a sizeable print run and has a global audience. Full details of acquiring copies, either printed or in identical form online, editorial policies, publication procedure, author submission guidelines, peer review guidelines, legal matters, advantages of publication in this journal and the like can be found by following links from: http://www.herp.net Published by Kotabi Pty Ltd PO Box 599 ISSN 1836-5698 (Print) Doncaster, Victoria, 3108. ISSN 1836-5779 (Online) Hoser 2016 - Australasian Journal of Herpetology 31:3-11. Australia. Copyright. All rights reserved. Available online at www.herp.net Online journals (thisCopyright- issue) do not Kotabiappear for Publishing a month after -the All actual rights and reservedlisted publication date of the printed journals. Minimum print run of first printings is always at least fifty hard copies. 12 Australasian Journal of Herpetology Australasian Journal of Herpetology 31:12-21. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online)

A division of the elapid genus Salomonelaps McDowell, 1970 from the Solomon Islands, including the resurrection of two species and formal description of four other forms (Serpentes: Elapidae: Micropechiini: Loveridgelapina).

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 29 April 2016, Accepted 30 May 2016, Published 1 August 2016.

ABSTRACT Since 1970, the Solomon Islands elapid genus Salomonelaps McDowell, 1970 has been viewed by virtually all herpetologists as being comprised of a single variable species. Salomonelaps par (Boulenger, 1884) has not been subjected to any serious taxonomic scrutiny, since being described. The two species Hoplocephalus woodfordii Boulenger, 1888 and H. melanurus Boulenger, 1888 have been ignored by all authors since, except perhaps for Kinghorn (1928), Schmidt (1932) and Williams and Parker (1964), who recognized the species “par” and ”woodfordii” in their accounts. More recently when erecting the genus Salomonelaps, McDowell (1970) treated ”woodfordii” and “melanurus” as synonyms of “par”. However, inspection of specimens from the majority of islands Salomonelaps occurs confirms the belief of McCoy (2006) that populations are significantly different on various islands. This includes consistent differences in scalation, colouration, dentition and hemipene morphology and can be reliably used to separate each form. As a result, of an assessment of the snakes and the relevant available genetic evidence involving species affected by the same geographical barriers, e.g. lizards of the genera Corucia Gray, 1855 and Tribolonotus Duméril and Bibron, 1839 as detailed by Austin et al. (2010) and Hagen et al. (2012), and the geological evidence of relevance, it is clear that the relevant forms are sufficiently divergent to warrant taxonomic recognition. Thus seven distinctive forms are herein given taxonomic recognition. All of S. par, S. woodfordi and S. melanurus are recognized as full species. Of the remaining four forms that are named for the first time, three are treated as subspecies of S. par, as it is likely that their populations were connected in the recent past during glacial maxima as outlined by Hagen et al. (2012), even though they are now clearly isolated and evolving independently of one another. Another, population from , believed to have been separated from the other populations the longest and thought not to be connected in recent ice age maxima, is herein treated as a full species. Rafting between islands is not viewed as a significant means of dispersal or ongoing gene flow, beyond times of initial colonisation for reasons given by Hagen et al. (2012) and Balsai (1995) and also due to the absence of the genus from nearby islands such as San Cristobal or beyond the Bougainville group. Keywords: Taxonomy; snakes; genus; Salomonelaps; species; par; woodfordii; melanurus; Boulenger; Solomon Islands; Solomons; ; Ngela; Nggela, Malaita; San Cristobal; ; ; Fauro; Santa Isabel; Choiseul; Florida Islands; Bougainville; new species; desburkei; new subspecies; ngelaensis; choiseulensis; shortlandensis.

INTRODUCTION being described, save for McDowell’s work in 1970. Since 1970, the Solomon Islands elapid genus Salomonelaps McDowell did an excellent job of summarizing physical McDowell, 1970 has been viewed by all herpetologists as being differences between populations from different islands, but failed comprised of a single variable species. This view has not until to see the significance of these differences, considering them now been questioned or challenged in spite of mounting mere variations in a wide-ranging species. evidence to the contrary. Salomonelaps par (Boulenger, 1884) However it should be noted that in 1970, the time McDowell’s has not been subjected to any serious taxonomic scrutiny, since study was published, he was isolated from molecular studies not Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 13

available at the time. glacial maxima as outlined by Hagen et al. (2012), even though He also was unable to merge this evidence with what is now well they are now clearly isolated, morphologically distinct and known about the recent geological past, in terms of ice-age evolving independently of one another. Another form, from maxima, changing sea levels and climates and the roles these Malaita, believed to have been separated from the other play in speciation, either in these relevant snakes or other reptile populations the longest and thought not to be connected in taxa affected by the same factors, all of which have led to recent ice age maxima, is herein treated as a full species. different conclusions made here and in the face of a lot of Rafting between islands is not viewed as a significant means of identical evidence. dispersal or ongoing gene flow, beyond times of initial The two species Hoplocephalus woodfordii Boulenger, 1888 and colonisation for reasons given by Hagen et al. (2012) and Balsai H. melanurus Boulenger, 1888 have been ignored by all authors (1995) and also due to the absence of the genus from nearby since being named, except perhaps for Kinghorn (1928) and islands such as San Cristobal or beyond the Bougainville group. Schmidt (1932), who recognized the species “par” and The islands Guadalcanal and Malaita are separated from one ”woodfordii” in their accounts, and ignoring the account of another and the others by a sea depth of more than 200 metres Williams and Parker (1964) who only accepted H. woodfordii on and hence do not appear to have been joined at any stage in the the grounds of precedent (McDowell, 1970). last 5 million years. More recently when erecting the genus Salomonelaps McDowell MATERIALS AND METHODS (1970) treated both ”woodfordii” and “melanurus” as synonyms These are not formally explained in a number of my recent of “par”. papers under the heading “Materials and Methods” or similar, on the basis they are self evident to any vaguely perceptive reader. This appears to have been the taxonomy followed by all However, the process by which the following taxonomy and herpetologists since then. nomenclature in this and other recent papers by myself of Inspection of a large number of specimens (several dozen) from similar form, has been arrived at, is explained herein for the the majority of islands Salomonelaps is known from and a benefit of people who have recently published so-called review of the literature, including the work of McDowell (1970), “criticisms” online of some of my recent papers. They have confirms the statement of McCoy (2006) that populations are alleged a serious “defect” by myself not formally explaining significantly different on various islands. “Materials and Methods” under such a heading. This includes consistent differences in scalation, colouration, The process involved in creating the final product for this and hemipene morphology in males and dentition as outlined by other relevant papers has been via a combination of the McDowell (1970) and combined, these can reliably be used to following: separate each form. Genera and component species are audited to see if their Notwithstanding McCoy’s two books on Solomon Islands classifications are correct on the basis on known type herpetofauna (McCoy 1980 and McCoy 2006), which included specimens, locations and the like when compared with known colour photos of a number of different forms of Salomonelaps, phylogenies and obvious morphological differences between like until now, no one has taken the matter further in terms of species. assessing these snakes to see if they represent just one highly Original descriptions and contemporary concepts of the species variable species, or in fact more than one. are matched with available specimens from across the ranges of Hence I engaged in such an assessment by reconciling the species to see if all conform to accepted norms. morphological differences with geographical evidence to ascertain divergences between local forms to determine which These may include those held in museums, private collections, were consistently different enough to be recognized as either collected in the field, photographed, posted on the internet or subspecies or species. held by individuals, and only when the location data is good and any other relevant data available. As a result, of an assessment of the snakes and the relevant Where specimens do not appear to comply with the described available genetic evidence involving species studied already that species (and accepted concept of the species), this non- have been affected by the same geographical barriers (e.g. conformation is looked at with a view to ascertaining if it is Corucia Gray, 1856 and Tribolonotus Duméril and Bibron, 1839) worthy of taxonomic recognition or other relevant considerations as detailed in the papers by Austin et al. (2010) and Hagen et al. on the basis of differences that can be tested for antiquity or (2012), and the geological evidence of relevance, it is clear that deduced from earlier studies. there are at least six relevant forms are sufficiently divergent to warrant taxonomic recognition. When this appears to be the case (non-conformation), the Divergences were ascertained on the basis of previous ice-age potential target taxon is inspected as closely as practicable with maxima connections between relevant islands as explained by a view to comparing with the nominate form or forms if other authors such as Bruns et al. (2009), Russell and Coupe (1984) similar taxa have been previously named. and recent molecular studies on both Corucia Gray, 1856 and Other relevant data is also inspected, including any available Tribolonotus Duméril and Bibron, 1839 as published and the molecular studies which may indicate likely divergence of relevant sources cited within. populations. Notwithstanding the theft of relevant materials from this author in Where molecular studies are unavailable for the relevant taxon an illegal armed raid on 17 August 2011, which were not or group, other studies involving species and groups constrained returned (Court of Appeal Victoria 2014 and VCAT 2015), I have by the same geographical or geological barriers, or with like made a decision to publish this paper in view of the conservation distribution patterns are inspected as they give reasonable significance attached to the formal recognition of unnamed indications of the likely divergences of the taxa being studied species and subspecies and on the basis that further delays herein. may in fact put these unnamed taxa at greater risk of extinction. Additionally other studies involving geological history, sea level Thus seven distinctive forms are herein given taxonomic and habitat changes associated with long-term climate change, recognition on the basis that likely divergences exceed the including recent ice age changes in sea levels, versus known timeline determined as significant by Keogh et al. (2003). sea depths are utilized to predict past movements of species All of S. par, S. woodfordi, S. melanurus are recognized as full and genus groups in order to further ascertain likely divergences species. Of the remaining four forms that are named for the first between extant populations (as done in this very paper). time, three are treated as subspecies of S. par, as it is likely that When all available information checks out to show taxonomically their populations were connected in the recent past during distinct populations worthy of recognition, they are then

Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 14 Australasian Journal of Herpetology recognized herein according to the rules of the International characters: there is no dark etching of the scales of the upper Code of Zoological Nomenclature (Ride et al. 1999). labials, including the rear ones, or the scales on the head in This means that if a name has been properly proposed in the front of the eye. There is no distinct nape at the rear of the head, past, it is used. This is exactly what happens in this paper for the although there is slight darkening in this region. The body has taxon described as Hoplocephalus woodfordii Boulenger, 1888. semi-distinct to distinct banding, slight darkening of tail and an Alternatively, if no name is available, one is proposed accoding immaculate creamish-white venter. If there are dark etched to the rules of the Code as is done four times in this paper. scales on the venter, they are only on the underside of the tail. There is none, or very little lightening towards the front of the As a matter of trite I mention that if a target taxon or group does snout. 158-167 ventrals. check out as being “in order” or properly classified, a paper is usually not published unless some other related taxon is named The subspecies S. par choiseulensis subsp. nov. described for the first time. herein, is separated from all other species and subspecies of Salomonelaps McDowell, 1970 by the following suite of The published literature relevant to the taxonomic judgements characters: semi-distinct to distinct banding. The darker bands, made within this paper includes papers relevant to Solomon described by McDowell (1970) as “broad banded” are 3-5 scale Islands species affected by the same physical barriers to lengths wide, alternating with lighter bands 1-3 scale lengths dispersion as well as those directly relevant to Salomonelaps. wide. The dark bands are essentially formed by prominent dark Combined, they include the following: edging of the scales and similarly dark interstitial skin, whereas Adler, et al. (1995), Austin et al. (2010), Balsai (1995), Barbour the interstitial skin of the pale bands is whiteish. In this taxon, (1921), Boseto and Pikacha (2016), Boulenger (1884, 1886, the venter is not immaculate. It has numerous dark etched 1887), Bruns et al. (1989), Cogger (1972), Dahl (1986), Duméril scales over a shiny whitish background along the main part of and Bibron (1839), Gray (1856), Greer (1982), Greer and Parker the mid body created by extensions of the dorsal bands running (1967), Greer and Simon (1982), Hagen et al. (2012), Hall across. These become prominent as one moves away from the (2002), Iskandar and Erdelen (2006), Keogh et al. (2003), anterior end of the snake. There is none, or very little lightening Kinghorn (1928, 1937), McCoy (1980, 2006), McDowell (1970), towards the front of the snout. Mys (1988), Ogilby (1890), Pianka and Vitt (2003), Pyron et al. The subspecies S. par ngelaensis subsp. nov. from the Florida (2013), Reeder (2003), Rittmeyer and Austin (2015), Russell and Islands group is separated from all other species and Coupe (1984), Schmidt (1932), Williams and Parker (1964), subspecies of Salomonelaps McDowell, 1970 by the following Zweifel (1966), and sources cited therein. suite of characters: bands invisible or rarely they are extremely GENUS SALOMONELAPS McDOWELL, 1970. indistinct (except in juveniles), scales of lower flanks have white Type species: Hoplocephalus par Boulenger, 1884. in the centers in a manner not seen in other forms, giving a Diagnosis: Because until now the elapid genus has been significantly different appearance to other forms. Most of the treated as monotypic, the diagnosis has been treated as the dorsal scales have a thick dark blackish etching, infilled with same for both genus and species. colour (most commonly reddish). The rostral is strongly whitish, All species and subspecies in the genus (as recognized herein) which spreads to nearby scales of the snout. There is also a can be defined as follows: significant amount of darkening below the eye, but otherwise not around the eye. The head is slightly depressed, distinct from the neck. The eye is moderate and nearly as long as its distance from the mouth. Salomonelaps par shortlandensis subsp. nov. from the Shortland Nasal may be single, but is usually divided and if single always Islands group of the Solomon Islands is separated from all other has a groove. One preocular, two postoculars. Internasals half species and subspecies of Salomonelaps McDowell, 1970 by as long as the prefrontals The rostral is broader than deep and the following suite of characters: The pattern is banded dorsally, just visible from above. The frontal is as wide as long or slightly to a width where the dark bands are about 2 scales wide and the longer, nearly twice as broad as the supraocular, as long as its lighter bands are about 2-3 scales wide, being slightly wider than distance from the rostral, much shorter than the parietals. the dark ones (in contrast to other banded Salomonelaps). The Posterior nasal is in contact with the preocular. 7 Supralabials, in colour of the dorsal scales, apart from the black edging of the which the third and fourth enter the eye. Four lower labials are in scales in the dark bands, is tawny orange, in contrast to the contact with the anterior chin-shields, which are shorter than the yellowish white and immaculate belly. posterior. Temporals 1+2. The rostral is wider than deep and In specimens of other subspecies of S. par the scales of the visible from above. Midbody scales in 15-17 rows, 158-180 dark bands may have reddish brown centres, but the scales of ventrals and the anal is divided. 38-59 subcaudals may be either the light bands are coloured like the belly. all single, all divided or a combination of these, often relevant to Female Salomonelaps par shortlandensis subsp. nov. are a given island population. unusual in having six to eight non-canaliculate maxillary teeth, McDowell (1970) gives a more detailed description of the genus versus five (rarely 4 ot 6) in all other forms of Salomonelaps. and also comparisons with the related genera of Ogmodon The taxon is also unusual in having 15 mid-body rows, versus Peters, 1864 and Loveridgelaps McDowell, 1970, as well as the the usual number of 17 (rarely 15 or 16) in the other populations distantly related genus Parapistocalamus Roux, 1934. of Salomonelaps. Of the other forms of Salomonelaps 15 mid Content: Salomonelaps par (Boulenger, 1884) (type species) body rows is only seen in S. melanurus and even then it is (including three subspecies); S. desburkei sp. nov.; S. relatively unusual. Of course S. melanurus is unique in terms of melanurus (Boulenger, 1888); S. woodfordii (Boulenger, 1888). Salomonelaps by the configuration of the banding. SALOMONELAPS PAR (BOULENGER, 1884). The species S. woodfordi (Boulenger, 1888) from the New Holotype: A specimen at the Museum of Natural History, UK, Georgia island group in the Western District is separated from specimen number: BMNH 1946.1.20.66 from , all other species and subspecies of Salomonelaps McDowell, Solomon Islands, situated immediately south-east of 1970 by the following suite of characters: The entire body, Bougainville and west of the Bougainville Strait. including the flanks is of a uniform colour, with each scale bordered by black, which results in a reticulated pattern as noted Diagnosis: Salomonelaps par is diagnosed as for the genus by Boulenger (1888), McDowell (1970) and McCoy (2006). The and separated from the other species and subspecies in the back of the head is slightly darker in colour, while the front of the genus, as follows: snout is lighter, but not white. All the dorsal head shields have The nominate subspecies of S. par from Bougainville and dark etching and there are whitish upper labials along the lip immediately offshore islands is separated from all other line, which darken higher up, near the eye. There is no subspecies and species by the following unique suite of darkening on the posterior dorsal surface. Subcaudals are white

Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 15

and patternless or rarely with darker margins. 165-179 ventrals. genus, as follows: Most or all subcaudals are divided (versus all or mainly single in The nominate subspecies of S. par from Bougainville and the other species). immediately offshore islands is separated from all other Rarely specimens of S. woodfordi may have a dorsal surface subspecies and species by the following unique suite of that is nearly a uniform dark grey colour, but all the scales of the characters: there is no dark etching of the scales of the upper first row, as well as some of those of the second row, have pale labials, including the rear ones, or the scales on the head in centres; ventrals and the subcaudals have dark etching on the front of the eye. There is no distinct nape at the rear of the head, margins. although there is slight darkening in this region. The body has The species S. melanurus (Boulenger, 1888) from Guadalcanal semi-distinct to distinct banding, slight darkening of tail and an in the central district is characterised and separated from all immaculate creamish-white venter. If there are dark etched other Salomonelaps species and subspecies by the following scales on the venter, they are only on the underside of the tail. characters: dark etching of the upper labials, but not those in There is none, or very little lightening towards the front of the front of the eyes. There is significant darkening around the eye, snout. especially posterior to it and darkening at the rear of the head The subspecies S. par choiseulensis subsp. nov. described forms a distinct nape. There is slight to significant darkening of herein, is separated from all other species and subspecies of the tail. Dorsal cross bands are noticeably more numerous and Salomonelaps McDowell, 1970 by the following suite of thinner than in other forms and were described by McDowell characters: semi-distinct to distinct banding. The darker bands, (1970) as “fine banded”. described by McDowell (1970) as “broad banded” are 3-5 scale McDowell’s description of the dorsal colouration was as follows: lengths wide, alternating with lighter bands 1-3 scale lengths “The dark bands are narrow (about one to one and a half scale wide. The dark bands are essentially formed by prominent dark lengths wide) and separated by pale bands only slightly, if at all, edging of the scales and similarly dark interstitial skin, whereas wider (one to two scale lengths wide). The general colour is the interstitial skin of the pale bands is whiteish. In this taxon, neutral or greyish brown, without orange tone, although the the venter is not immaculate. It has numerous dark etched scales over a shiny whitish background along the main part of scales may become almost uniform black, particularly the mid body created by extensions of the dorsal bands running posteriorly, so that the banding pattern is visible only on the across. These become prominent as one moves away from the interstitial skin. The bands do not extend onto the ventral anterior end of the snake. There is none, or very little lightening surface and there is no contrast between the colour of the belly towards the front of the snout. and that of the pale dorsal bands.” S. melanurus has 164-170 ventrals. The subspecies S. par ngelaensis subsp. nov. from the Florida Islands group is separated from all other species and The new species S. desburkei sp. nov. from Malaita in the subspecies of Salomonelaps McDowell, 1970 by the following Malaita district is characterised and separated from the other suite of characters: bands invisible or rarely they are extremely species of Salomonelaps McDowell, 1970 by the presence of indistinct (except in juveniles), scales of lower flanks have white upper labials that are lighter near the lip, darkening towards the in the centers in a manner not seen in other forms, giving a eye-line and a dorsal pattern with semidistinct bands in adults. significantly different appearance to other forms. Most of the The darker bands, described by McDowell (1970) as “broad dorsal scales have a thick dark blackish etching, infilled with banded” are 3-5 scale lengths wide, alternating with lighter colour (most commonly reddish). The rostral is strongly whitish, bands 1-3 scale lengths wide. The dark bands are essentially which spreads to nearby scales of the snout. There is also a formed by prominent dark edging of the scales and similarly significant amount of darkening below the eye, but otherwise not dark interstitial skin, whereas the interstitial skin of the pale around the eye. bands is whiteish. The dorsal pattern does not appear Salomonelaps par shortlandensis subsp. nov. from the Shortland reticulated as seen in S. woodfordi (described below). Islands group of the Solomon Islands is separated from all other In S. desburkei sp. nov. the dorsal pattern does not extend to species and subspecies of Salomonelaps McDowell, 1970 by the venter as seen in broad banded Salomonelaps or those with the following suite of characters: The pattern is banded dorsally, semi-distinct or indistinct bands from other islands, such as to a width where the dark bands are about 2 scales wide and the Ngella or Santa Isabella. lighter bands are about 2-3 scales wide, being slightly wider than Male S. desburkei sp. nov. are further separated from all other the dark ones (in contrast to other banded Salomonelaps). The species in the genus by the following suite of characters for the colour of the dorsal scales, apart from the black edging of the hemipenes, Organ to subcaudal (13), forked at subcaudal (11), scales in the dark bands, is tawny orange, in contrast to the Sulcus forked at subcaudal (9), Large spines begin at subcaudal yellowish white and immaculate belly. 7. In specimens of other subspecies of S. par the scales of the Distribution: The nominate subspecies of S. par par is believed dark bands may have reddish brown centres, but the scales of to be restricted to the Bougainville Group of islands only, as in the light bands are coloured like the belly. islands west of the Bougainville Strait including Fauro and Buka Female Salomonelaps par shortlandensis subsp. nov. are islands. See the following subspecies descriptions for the unusual in having six to eight non-canaliculate maxillary teeth, distribution of other forms of S. par in the Solomon Islands, to versus five (rarely 4 ot 6) in all other forms of Salomonelaps. complete the distribution for all S.par subspecies. The taxon is also unusual in having 15 mid-body rows, versus SALOMONELAPS PAR CHOISEULENSIS SUBSP. NOV. the usual number of 17 (rarely 15 or 16) in the other populations Holotype: A preserved specimen at the Museum of of Salomonelaps. Of the other forms of Salomonelaps 15 mid Comparative Zoology, Harvard University, USA, specimen body rows is only seen in S. melanurus and even then it is number: R-193030 collected from a trail running from Ghargara relatively unusual. Of course S. melanurus is unique in terms of Village to Sarelata Camp, , Solomon Islands. Salomonelaps by the configuration of the banding. The Museum of Comparative Zoology, Harvard University, USA The species S. woodfordi (Boulenger, 1888) from the New is a public facility that allows access to its specimen holdings. Georgia island group in the Western District is separated from Paratype: A preserved specimen at the Australian Museum, all other species and subspecies of Salomonelaps McDowell, Sydney, NSW, Australia, specimen number: R.127354 collected 1970 by the following suite of characters: The entire body, from Gnulahaghe Village, Ysabel Island (Santa Isabel), Solomon including the flanks is of a uniform colour, with each scale Islands (8°07' S, 159°32' E). bordered by black, which results in a reticulated pattern as noted Diagnosis: Salomonelaps par is diagnosed as for the genus by Boulenger (1888), McDowell (1970) and McCoy (2006). The back of the head is slightly darker in colour, while the front of the Hoser 2016 - Australasian Journal of Herpetology 31:12-21. and separated from the other species and subspecies in the Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 16 Australasian Journal of Herpetology snout is lighter, but not white. All the dorsal head shields have Diagnosis: Salomonelaps par is diagnosed as for the genus dark etching and there are whitish upper labials along the lip and separated from the other species and subspecies in the line, which darken higher up, near the eye. There is no genus, as follows: darkening on the posterior dorsal surface. Subcaudals are white The nominate subspecies of S. par from Bougainville and and patternless or rarely with darker margins. 165-179 ventrals. immediately offshore islands is separated from all other Most or all subcaudals are divided (versus all or mainly single in subspecies and species by the following unique suite of the other species). characters: there is no dark etching of the scales of the upper Rarely specimens of S. woodfordi may have a dorsal surface labials, including the rear ones, or the scales on the head in that is nearly a uniform dark grey colour, but all the scales of the front of the eye. There is no distinct nape at the rear of the head, first row, as well as some of those of the second row, have pale although there is slight darkening in this region. The body has centres; ventrals and the subcaudals have dark etching on the semi-distinct to distinct banding, slight darkening of tail and an margins. immaculate creamish-white venter. If there are dark etched The species S. melanurus (Boulenger, 1888) from Guadalcanal scales on the venter, they are only on the underside of the tail. in the central district is characterised and separated from all There is none, or very little lightening towards the front of the other Salomonelaps species and subspecies by the following snout. characters: dark etching of the upper labials, but not those in The subspecies S. par ngelaensis subsp. nov. from the Florida front of the eyes. There is significant darkening around the eye, Islands group described herein, is separated from all other especially posterior to it and darkening at the rear of the head species and subspecies of Salomonelaps McDowell, 1970 by forms a distinct nape. There is slight to significant darkening of the following suite of characters: bands invisible or rarely they the tail. Dorsal cross bands are noticeably more numerous and are extremely indistinct (except in juveniles), scales of lower thinner than in other forms as were described by McDowell flanks have white in the centers in a manner not seen in other (1970) as “fine banded”. forms, giving a significantly different appearance to other forms. McDowell’s description of the dorsal colouration was as follows: Most of the dorsal scales have a thick dark blackish etching, “The dark bands are narrow (about one to one and a half scale infilled with colour (most commonly reddish). The rostral is lengths wide) and separated by pale bands only slightly, if at all, strongly whitish, which spreads to nearby scales of the snout. wider (one to two scale lengths wide). The general colour is There is also a significant amount of darkening below the eye, neutral or greyish brown, without orange tone, although the but otherwise not around the eye. scales may become almost uniform black, particularly The subspecies S. par choiseulensis subsp. nov. described posteriorly, so that the banding pattern is visible only on the herein, is separated from all other species and subspecies of interstitial skin. The bands do not extend onto the ventral Salomonelaps McDowell, 1970 by the following suite of surface and there is no contrast between the colour of the belly characters: semi-distinct to distinct banding. The darker bands, and that of the pale dorsal bands.” described by McDowell (1970) as “broad banded” are 3-5 scale S. melanurus has 164-170 ventrals. lengths wide, alternating with lighter bands 1-3 scale lengths wide. The dark bands are essentially formed by prominent dark The new species S. desburkei sp. nov. from Malaita in the edging of the scales and similarly dark interstitial skin, whereas Malaita district is characterised and separated from the other the interstitial skin of the pale bands is whiteish. In this taxon, species of Salomonelaps McDowell, 1970 by the presence of the venter is not immaculate. It has numerous dark etched upper labials that are lighter near the lip, darkening towards the scales over a shiny whitish background along the main part of eye-line and a dorsal pattern with semidistinct bands in adults. the mid body created by extensions of the dorsal bands running The darker bands, described by McDowell (1970) as “broad across. These become prominent as one moves away from the banded” are 3-5 scale lengths wide, alternating with lighter anterior end of the snake. There is none, or very little lightening bands 1-3 scale lengths wide. The dark bands are essentially towards the front of the snout. formed by prominent dark edging of the scales and similarly Salomonelaps par shortlandensis subsp. nov. from the Shortland dark interstitial skin, whereas the interstitial skin of the pale Islands group of the Solomon Islands is separated from all other bands is whiteish. The dorsal pattern does not appear species and subspecies of Salomonelaps McDowell, 1970 by reticulated as seen in S. woodfordi (described below). the following suite of characters: The pattern is banded dorsally, In S. desburkei sp. nov. the dorsal pattern does not extend to to a width where the dark bands are about 2 scales wide and the the venter as seen in broad banded Salomonelaps or those with lighter bands are about 2-3 scales wide, being slightly wider than semi-distinct or indistinct bands from other islands, such as the dark ones (in contrast to other banded Salomonelaps). The Ngella or Santa Isabella. colour of the dorsal scales, apart from the black edging of the Male S. desburkei sp. nov. are further separated from all other scales in the dark bands, is tawny orange, in contrast to the species in the genus by the following suite of characters for the yellowish white and immaculate belly. hemipenes, Organ to subcaudal (13), forked at subcaudal (11), In specimens of other subspecies of S. par the scales of the Sulcus forked at subcaudal (9), Large spines begin at subcaudal dark bands may have reddish brown centres, but the scales of 7. the light bands are coloured like the belly. Distribution: Choiseul, Santa Isabel and immediately offshore Female Salomonelaps par shortlandensis subsp. nov. are smaller islands, Solomon Islands. unusual in having six to eight non-canaliculate maxillary teeth, Etymology: Named in relflection of where the taxon is found. versus five (rarely 4 ot 6) in all other forms of Salomonelaps. SALOMONELAPS PAR NGELAENSIS SUBSP. NOV. The taxon is also unusual in having 15 mid-body rows, versus Holotype: A preserved specimen at the Australian Museum, the usual number of 17 (rarely 15 or 16) in the other populations Sydney, NSW, Australia, specimen number: R.91262, collected of Salomonelaps. Of the other forms of Salomonelaps 15 mid from Mboromole, Nggela Island, Solomon Islands (9°03' S, body rows is only seen in S. melanurus and even then it is 160°18' E). relatively unusual. Of course S. melanurus is unique in terms of Salomonelaps by the configuration of the banding. The Australian Museum, Sydney, NSW, Australia is a public facility that allows access to its specimen holdings. The species S. woodfordi (Boulenger, 1888) from the New Georgia island group in the Western District is separated from Paratypes: Three preserved specimens at the Australian all other species and subspecies of Salomonelaps McDowell, Museum, Sydney, NSW, Australia, specimen numbers: R.91261, 1970 by the following suite of characters: The entire body, R.91227 and R.91263, collected from Boromole, Nggela Island, including the flanks is of a uniform colour, with each scale Solomon Islands (9°03' S, 160°18' E). bordered by black, which results in a reticulated pattern as noted Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 17

by Boulenger (1888), McDowell (1970) and McCoy (2006). The is a facility that allows access to its specimen holdings. back of the head is slightly darker in colour, while the front of the Paratypes: Preserved specimens at the Museum of snout is lighter, but not white. All the dorsal head shields have Comparative Zoology, Harvard University, USA, specimen dark etching and there are whitish upper labials along the lip numbers: R-88457, R-88458, R-89475 and R-89476 collected line, which darken higher up, near the eye. There is no from Nohu, Magusaiai Island, in the Group of darkening on the posterior dorsal surface. Subcaudals are white the Solomon Islands. and patternless or rarely with darker margins. 165-179 ventrals. The holotype and paratypes are all females. Most or all subcaudals are divided (versus all or mainly single in the other species). Diagnosis: Salomonelaps par shortlandensis subsp. nov. from the Shortland Islands group of the Solomon Islands is separated Rarely specimens of S. woodfordi may have a dorsal surface from all other species and subspecies of Salomonelaps that is nearly a uniform dark grey colour, but all the scales of the McDowell, 1970 by the following suite of characters: The pattern first row, as well as some of those of the second row, have pale is banded dorsally, to a width where the dark bands are about 2 centres; ventrals and the subcaudals have dark etching on the scales wide and the lighter bands are about 2-3 scales wide, margins. being slightly wider than the dark ones (in contrast to other The species S. melanurus (Boulenger, 1888) from Guadalcanal banded Salomonelaps). The colour of the dorsal scales, apart in the central district is characterised and separated from all from the black edging of the scales in the dark bands, is tawny other Salomonelaps species and subspecies by the following orange, in contrast to the yellowish white and immaculate belly. characters: dark etching of the upper labials, but not those in In specimens of other subspecies of S. par the scales of the front of the eyes. There is significant darkening around the eye, dark bands may have reddish brown centres, but the scales of especially posterior to it and darkening at the rear of the head the light bands are coloured like the belly. forms a distinct nape. There is slight to significant darkening of the tail. Dorsal cross bands are noticeably more numerous and Female Salomonelaps par shortlandensis subsp. nov. are thinner than in other forms as were described by McDowell unusual in having six to eight non-canaliculate maxillary teeth, (1970) as “fine banded”. versus five (rarely 4 ot 6) in all other forms of Salomonelaps. The taxon is also unusual in having 15 mid-body rows, versus McDowell’s description of the dorsal colouration was as follows: the usual number of 17 (rarely 15 or 16) in the other populations “The dark bands are narrow (about one to one and a half scale of Salomonelaps. Of the other forms of Salomonelaps 15 mid lengths wide) and separated by pale bands only slightly, if at all, body rows is only seen in S. melanurus and even then it is wider (one to two scale lengths wide). The general colour is relatively unusual. Of course S. melanurus is unique in terms of neutral or greyish brown, without orange tone, although the Salomonelaps by the configuration of the banding. scales may become almost uniform black, particularly Salomonelaps par is otherwise diagnosed as for the genus and posteriorly, so that the banding pattern is visible only on the separated from the other species and subspecies in the genus, interstitial skin. The bands do not extend onto the ventral as follows: surface and there is no contrast between the colour of the belly and that of the pale dorsal bands.” The nominate subspecies of S. par from Bougainville and immediately offshore islands is separated from all other S. melanurus has 164-170 ventrals. subspecies and species by the following unique suite of The new species S. desburkei sp. nov. from Malaita in the characters: there is no dark etching of the scales of the upper Malaita district is characterised and separated from the other labials, including the rear ones, or the scales on the head in species of Salomonelaps McDowell, 1970 by the presence of front of the eye. There is no distinct nape at the rear of the head, upper labials that are lighter near the lip, darkening towards the although there is slight darkening in this region. The body has eye-line and a dorsal pattern with semidistinct bands in adults. semi-distinct to distinct banding, slight darkening of tail and an The darker bands, described by McDowell (1970) as “broad immaculate creamish-white venter. If there are dark etched banded” are 3-5 scale lengths wide, alternating with lighter scales on the venter, they are only on the underside of the tail. bands 1-3 scale lengths wide. The dark bands are essentially There is none, or very little lightening towards the front of the formed by prominent dark edging of the scales and similarly snout. dark interstitial skin, whereas the interstitial skin of the pale The subspecies S. par ngelaensis subsp. nov. from the Florida bands is whiteish. The dorsal pattern does not appear Islands group described herein, is separated from all other reticulated as seen in S. woodfordi (described below). species and subspecies of Salomonelaps McDowell, 1970 by In S. desburkei sp. nov. the dorsal pattern does not extend to the following suite of characters: bands invisible or rarely they the venter as seen in broad banded Salomonelaps or those with are extremely indistinct (except in juveniles), scales of lower semi-distinct or indistinct bands from other islands, such as flanks have white in the centers in a manner not seen in other Ngella or Santa Isabella. forms, giving a significantly different appearance to other forms. Male S. desburkei sp. nov. are further separated from all other Most of the dorsal scales have a thick dark blackish etching, species in the genus by the following suite of characters for the infilled with colour (most commonly reddish). The rostral is hemipenes, Organ to subcaudal (13), forked at subcaudal (11), strongly whitish, which spreads to nearby scales of the snout. Sulcus forked at subcaudal (9), Large spines begin at subcaudal There is also a significant amount of darkening below the eye, 7. but otherwise not around the eye. Distribution: Belived to be restricted to the Florida islands in The subspecies S. par choiseulensis subsp. nov. described the Central District of the Solomon Islands (Ngela (AKA Nggela), herein, is separated from all other species and subspecies of including Nggela Sule, Nggela Pile, etc). Salomonelaps McDowell, 1970 by the following suite of Etymology: This snake is named in reflection of where the characters: semi-distinct to distinct banding. The darker bands, taxon is found. The spelling is intentional, even if treated in error described by McDowell (1970) as “broad banded” are 3-5 scale by a later reviser and should not be changed. lengths wide, alternating with lighter bands 1-3 scale lengths SALOMONELAPS PAR SHORTLANDENSIS SUBSP. NOV. wide. The dark bands are essentially formed by prominent dark edging of the scales and similarly dark interstitial skin, whereas Holotype: A preserved specimen at the Museum of the interstitial skin of the pale bands is whiteish. In this taxon, Comparative Zoology, Harvard University, USA, specimen the venter is not immaculate. It has numerous dark etched number: R-88455 collected from Maliai, Magusaiai Island, in the scales over a shiny whitish background along the main part of Shortland Islands Group of the Solomon Islands. the mid body created by extensions of the dorsal bands running The Museum of Comparative Zoology, Harvard University, USA across. These become prominent as one moves away from the

Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 18 Australasian Journal of Herpetology anterior end of the snake. There is none, or very little lightening SALOMONELAPS WOODFORDI (BOULENGER, 1888). towards the front of the snout. Holotype: A female specimen from Rubiana, New Georgia, The species S. woodfordi (Boulenger, 1888) from the New Solomon Islands. Georgia island group in the Western District is separated from Diagnosis: The species S. woodfordi (Boulenger, 1888) from all other species and subspecies of Salomonelaps McDowell, the New Georgia island group in the Western District is 1970 by the following suite of characters: The entire body, separated from all other species and subspecies of including the flanks is of a uniform colour, with each scale Salomonelaps McDowell, 1970 by the following suite of bordered by black, which results in a reticulated pattern as noted characters: The entire body, including the flanks is of a uniform by Boulenger (1888), McDowell (1970) and McCoy (2006). The colour, with each scale bordered by black, which results in a back of the head is slightly darker in colour, while the front of the reticulated pattern as noted by Boulenger (1888), McDowell snout is lighter, but not white. All the dorsal head shields have (1970) and McCoy (2006). The back of the head is slightly dark etching and there are whitish upper labials along the lip darker in colour, while the front of the snout is lighter, but not line, which darken higher up, near the eye. There is no white. All the dorsal head shields have dark etching and there darkening on the posterior dorsal surface. Subcaudals are white are whitish upper labials along the lip line, which darken higher and patternless or rarely with darker margins. 165-179 ventrals. up, near the eye. There is no darkening on the posterior dorsal Most or all subcaudals are divided (versus all or mainly single in surface. Subcaudals are white and patternless or rarely with the other species). darker margins. 165-179 ventrals. Most or all subcaudals are Rarely specimens of S. woodfordi may have a dorsal surface divided (versus all or mainly single in the other species). that is nearly a uniform dark grey colour, but all the scales of the Rarely specimens of S. woodfordi may have a dorsal surface first row, as well as some of those of the second row, have pale that is nearly a uniform dark grey colour, but all the scales of the centres; ventrals and the subcaudals have dark etching on the first row, as well as some of those of the second row, have pale margins. centres; ventrals and the subcaudals have dark etching on the The species S. melanurus (Boulenger, 1888) from Guadalcanal margins. in the central district is characterised and separated from all For diagnostic features of the other species and subspecies of other Salomonelaps species and subspecies by the following Salomonelaps, see the preceding description of S. par. characters: dark etching of the upper labials, but not those in Distribution: The New Georgia group of islands in the Western front of the eyes. There is significant darkening around the eye, District of the Solomon Islands. especially posterior to it and darkening at the rear of the head forms a distinct nape. There is slight to significant darkening of SALOMONELAPS MELANURUS BOULENGER,1888. the tail. Dorsal cross bands are noticeably more numerous and Holotype: Specimen number BM 1946.1.18.61 at the Museum thinner than in other forms as were described by McDowell of Natural History, UK, collected from Guadalcanal, as (1970) as “fine banded”. designated by McDowell (1970) as first reviser. McDowell’s description of the dorsal colouration was as follows: Diagnosis: The species S. melanurus (Boulenger, 1888) from “The dark bands are narrow (about one to one and a half scale Guadalcanal in the central district is characterised and lengths wide) and separated by pale bands only slightly, if at all, separated from all other Salomonelaps species and subspecies wider (one to two scale lengths wide). The general colour is by the following characters: dark etching of the upper labials, but neutral or greyish brown, without orange tone, although the not those in front of the eyes. There is significant darkening scales may become almost uniform black, particularly around the eye, especially posterior to it and darkening at the posteriorly, so that the banding pattern is visible only on the rear of the head forms a distinct nape. There is slight to interstitial skin. The bands do not extend onto the ventral significant darkening of the tail. Dorsal cross bands are surface and there is no contrast between the colour of the belly noticeably more numerous and thinner than in other forms as and that of the pale dorsal bands.” were described by McDowell (1970) as “fine banded”. S. melanurus has 164-170 ventrals. McDowell’s description of the dorsal colouration was as follows: “The dark bands are narrow (about one to one and a half scale The new species S. desburkei sp. nov. from Malaita in the lengths wide) and separated by pale bands only slightly, if at all, Malaita district is characterised and separated from the other wider (one to two scale lengths wide). The general colour is species of Salomonelaps McDowell, 1970 by the presence of neutral or greyish brown, without orange tone, although the upper labials that are lighter near the lip, darkening towards the eye-line and a dorsal pattern with semidistinct bands in adults. scales may become almost uniform black, particularly The darker bands, described by McDowell (1970) as “broad posteriorly, so that the banding pattern is visible only on the banded” are 3-5 scale lengths wide, alternating with lighter interstitial skin. The bands do not extend onto the ventral bands 1-3 scale lengths wide. The dark bands are essentially surface and there is no contrast between the colour of the belly formed by prominent dark edging of the scales and similarly and that of the pale dorsal bands.” dark interstitial skin, whereas the interstitial skin of the pale S. melanurus has 164-170 ventrals. bands is whiteish. The dorsal pattern does not appear Salomonelaps par is diagnosed as for the genus and separated reticulated as seen in S. woodfordi (described below). from the other species and subspecies in the genus, as follows: In S. desburkei sp. nov. the dorsal pattern does not extend to The nominate subspecies of S. par from Bougainville and the venter as seen in broad banded Salomonelaps or those with immediately offshore islands is separated from all other semi-distinct or indistinct bands from other islands, such as subspecies and species by the following unique suite of Ngella or Santa Isabella. characters: there is no dark etching of the scales of the upper Male S. desburkei sp. nov. are further separated from all other labials, including the rear ones, or the scales on the head in species in the genus by the following suite of characters for the front of the eye. There is no distinct nape at the rear of the head, hemipenes, Organ to subcaudal (13), forked at subcaudal (11), although there is slight darkening in this region. The body has Sulcus forked at subcaudal (9), Large spines begin at subcaudal semi-distinct to distinct banding, slight darkening of tail and an 7. immaculate creamish-white venter. If there are dark etched scales on the venter, they are only on the underside of the tail. Distribution: Shortland Island and the immediately adjacent There is none, or very little lightening towards the front of the Magusaiai Island, Solomon Islands. snout. Etymology: This snake is named in relflection of where the The subspecies S. par choiseulensis subsp. nov. described taxon is found. The spelling is intentional, even if treated in error herein, is separated from all other species and subspecies of by a later reviser, and should not be changed.

Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 19

Salomonelaps McDowell, 1970 by the following suite of upper labials that are lighter near the lip, darkening towards the characters: semi-distinct to distinct banding. The darker bands, eye-line and a dorsal pattern with semidistinct bands in adults. described by McDowell (1970) as “broad banded” are 3-5 scale The darker bands, described by McDowell (1970) as “broad lengths wide, alternating with lighter bands 1-3 scale lengths banded” are 3-5 scale lengths wide, alternating with lighter wide. The dark bands are essentially formed by prominent dark bands 1-3 scale lengths wide. The dark bands are essentially edging of the scales and similarly dark interstitial skin, whereas formed by prominent dark edging of the scales and similarly the interstitial skin of the pale bands is whiteish. In this taxon, dark interstitial skin, whereas the interstitial skin of the pale the venter is not immaculate. It has numerous dark etched bands is whiteish. The dorsal pattern does not appear scales over a shiny whitish background along the main part of reticulated as seen in S. woodfordi (described below). the mid body created by extensions of the dorsal bands running In S. desburkei sp. nov. the dorsal pattern does not extend to across. These become prominent as one moves away from the the venter as seen in broad banded Salomonelaps or those with anterior end of the snake. There is none, or very little lightening semi-distinct or indistinct bands from other islands, such as towards the front of the snout. Ngella or Santa Isabella. The subspecies S. par ngelaensis subsp. nov. from the Florida Male S. desburkei sp. nov. are further separated from all other Islands group is separated from all other species and species in the genus by the following suite of characters for the subspecies of Salomonelaps McDowell, 1970 by the following hemipenes, Organ to subcaudal (13), forked at subcaudal (11), suite of characters: bands invisible or rarely they are extremely Sulcus forked at subcaudal (9), Large spines begin at subcaudal indistinct (except in juveniles), scales of lower flanks have white 7. in the centers in a manner not seen in other forms, giving a Distribution: Guadalcanal in the central district of Solomon significantly different appearance to other forms. Most of the Islands, and immediately adjacent islets, but not including the dorsal scales have a thick dark blackish etching, infilled with Florida Islands (Ngela, AKA Nggela Sule, Nggela Pile, etc) to colour (most commonly reddish). The rostral is strongly whitish, the near north. which spreads to nearby scales of the snout. There is also a significant amount of darkening below the eye, but otherwise not SALOMONELAPS DESBURKEI SP. NOV. around the eye. Holotype: A preserved specimen at the Australian Museum, Salomonelaps par shortlandensis subsp. nov. from the Shortland Sydney, NSW, Australia, specimen number: R.87374, collected Islands group of the Solomon Islands is separated from all other within a 3 km radius of Bitaama, North Malaita, Solomon Islands species and subspecies of Salomonelaps McDowell, 1970 by (8°24' S, 160°36' E). the following suite of characters: The pattern is banded dorsally, The Australian Museum, Sydney, NSW, Australia is a public to a width where the dark bands are about 2 scales wide and the facility that allows access to its specimen holdings. lighter bands are about 2-3 scales wide, being slightly wider than Paratypes: A preserved specimen at the Australian Museum, the dark ones (in contrast to other banded Salomonelaps). The Sydney, NSW, Australia, specimen number: R.87375, collected colour of the dorsal scales, apart from the black edging of the within a 3 km radius of Bitaama, North Malaita, Solomon Islands scales in the dark bands, is tawny orange, in contrast to the (8°24' S, 160°36' E) and a preserved specimen at the Australian yellowish white and immaculate belly. Museum, Sydney, NSW, Australia, specimen number: R.137237 In specimens of other subspecies of S. par the scales of the collected at Bsurata Village, Malaita Island, Solomon Islands dark bands may have reddish brown centres, but the scales of (8°49' S, 160°49' E). the light bands are coloured like the belly. Diagnosis: The new species S. desburkei sp. nov. from Malaita Female Salomonelaps par shortlandensis subsp. nov. are in the Malaita district is characterised and separated from the unusual in having six to eight non-canaliculate maxillary teeth, other species of Salomonelaps McDowell, 1970 by the presence versus five (rarely 4 ot 6) in all other forms of Salomonelaps. of upper labials that are lighter near the lip, darkening towards The taxon is also unusual in having 15 mid-body rows, versus the eye-line and a dorsal pattern with semidistinct bands in the usual number of 17 (rarely 15 or 16) in the other populations adults. The darker bands, described by McDowell (1970) as of Salomonelaps. Of the other forms of Salomonelaps 15 mid “broad banded” are 3-5 scale lengths wide, alternating with body rows is only seen in S. melanurus and even then it is lighter bands 1-3 scale lengths wide. The dark bands are relatively unusual. Of course S. melanurus is unique in terms of essentially formed by prominent dark edging of the scales and Salomonelaps by the configuration of the dorsal banding. similarly dark interstitial skin, whereas the interstitial skin of the The species S. woodfordi (Boulenger, 1888) from the New pale bands is whiteish. The dorsal pattern does not appear Georgia island group in the Western District is separated from reticulated as seen in S. woodfordi (described below). all other species and subspecies of Salomonelaps McDowell, In S. desburkei sp. nov. the dorsal pattern does not extend to 1970 by the following suite of characters: The entire body, the venter as seen in broad banded Salomonelaps or those with including the flanks is of a uniform colour, with each scale semi-distinct or indistinct bands from other islands, such as bordered by black, which results in a reticulated pattern as noted Ngella or Santa Isabella. by Boulenger (1888), McDowell (1970) and McCoy (2006). The Male S. desburkei sp. nov. are further separated from all other back of the head is slightly darker in colour, while the front of the species in the genus by the following suite of characters for the snout is lighter, but not white. All the dorsal head shields have hemipenes, Organ to subcaudal (13), forked at subcaudal (11), dark etching and there are whitish upper labials along the lip Sulcus forked at subcaudal (9), Large spines begin at subcaudal line, which darken higher up, near the eye. There is no 7. darkening on the posterior dorsal surface. Subcaudals are white Salomonelaps par is diagnosed as for the genus and separated and patternless or rarely with darker margins. 165-179 ventrals. from the other species and subspecies in the genus, as follows: Most or all subcaudals are divided (versus all or mainly single in The nominate subspecies of S. par from Bougainville and the other species). immediately offshore islands is separated from all other Rarely specimens of S. woodfordi may have a dorsal surface subspecies and species by the following unique suite of that is nearly a uniform dark grey colour, but all the scales of the characters: there is no dark etching of the scales of the upper first row, as well as some of those of the second row, have pale labials, including the rear ones, or the scales on the head in centres; ventrals and the subcaudals have dark etching on the front of the eye. There is no distinct nape at the rear of the head, margins. although there is slight darkening in this region. The body has The new species S. desburkei sp. nov. from Malaita in the semi-distinct to distinct banding, slight darkening of tail and an Malaita district is characterised and separated from the other immaculate creamish-white venter. If there are dark etched species of Salomonelaps McDowell, 1970 by the presence of scales on the venter, they are only on the underside of the tail. Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 20 Australasian Journal of Herpetology

There is none, or very little lightening towards the front of the margins. snout. The species S. melanurus (Boulenger, 1888) from Guadalcanal The subspecies S. par choiseulensis subsp. nov. described in the central district is characterised and separated from all herein, is separated from all other species and subspecies of other Salomonelaps species and subspecies by the following Salomonelaps McDowell, 1970 by the following suite of characters: dark etching of the upper labials, but not those in characters: semi-distinct to distinct banding. The darker bands, front of the eyes. There is significant darkening around the eye, described by McDowell (1970) as “broad banded” are 3-5 scale especially posterior to it and darkening at the rear of the head lengths wide, alternating with lighter bands 1-3 scale lengths forms a distinct nape. There is slight to significant darkening of wide. The dark bands are essentially formed by prominent dark the tail. Dorsal cross bands are noticeably more numerous and edging of the scales and similarly dark interstitial skin, whereas thinner than in other forms as were described by McDowell the interstitial skin of the pale bands is whiteish. In this taxon, (1970) as “fine banded”. the venter is not immaculate. It has numerous dark etched McDowell’s description of the dorsal colouration was as follows: scales over a shiny whitish background along the main part of “The dark bands are narrow (about one to one and a half scale the mid body created by extensions of the dorsal bands running lengths wide) and separated by pale bands only slightly, if at all, across. These become prominent as one moves away from the wider (one to two scale lengths wide). The general colour is anterior end of the snake. There is none, or very little lightening neutral or greyish brown, without orange tone, although the towards the front of the snout. scales may become almost uniform black, particularly The subspecies S. par ngelaensis subsp. nov. from the Florida posteriorly, so that the banding pattern is visible only on the Islands group is separated from all other species and interstitial skin. The bands do not extend onto the ventral subspecies of Salomonelaps McDowell, 1970 by the following surface and there is no contrast between the colour of the belly suite of characters: bands invisible or rarely they are extremely and that of the pale dorsal bands.” indistinct (except in juveniles), scales of lower flanks have white S. melanurus has 164-170 ventrals. in the centers in a manner not seen in other forms, giving a significantly different appearance to other forms. Most of the Distribution: Malaita in the Malaita district of Solomon Islands, dorsal scales have a thick dark blackish etching, infilled with and immediately adjacent islets, but not including the Florida colour (most commonly reddish). The rostral is strongly whitish, Islands (Ngela, AKA Nggela Sule, Nggela Pile, etc) to the near which spreads to nearby scales of the snout. There is also a south-west. significant amount of darkening below the eye, but otherwise not Etymology: Named in honour of Desmond (Des) Burke, of around the eye. Joseph Burke Law, in recognition of his significant contributions Salomonelaps par shortlandensis subsp. nov. from the Shortland to the Criminal Justice system in Victoria as well as his Islands group of the Solomon Islands is separated from all other significant work in the past involving rodent breeding at this species and subspecies of Salomonelaps McDowell, 1970 by author’s research facility. the following suite of characters: The pattern is banded dorsally, REFERENCES CITED to a width where the dark bands are about 2 scales wide and the Adler, G. H., Austin, C. C. and Dudley, R. 1995. Dispersal and lighter bands are about 2-3 scales wide, being slightly wider than speciation of skinks among archipelagos in the tropical Pacific the dark ones (in contrast to other banded Salomonelaps). The Ocean. Evolutionary Ecology 9:529-541. colour of the dorsal scales, apart from the black edging of the Austin, C. C., Rittmeyer, E. N., Richards, S. J. and Zug, G. R. scales in the dark bands, is tawny orange, in contrast to the 2010. Phylogeny, historical biogeography and body size yellowish white and immaculate belly. evolution in Pacific Island Crocodile skinks Tribolonotus In specimens of other subspecies of S. par the scales of the (Squamata; Scincidae). Molecular Phylogenetics and Evolution dark bands may have reddish brown centres, but the scales of 57(1):227-236. the light bands are coloured like the belly. Balsai, M. J. 1995. Husbandry and Breeding of the Solomon Female Salomonelaps par shortlandensis subsp. nov. are Islands Prehensile-tailed Skink, Corucia zebrata. The Vivarium unusual in having six to eight non-canaliculate maxillary teeth, 7(1):4-11. versus five (rarely 4 ot 6) in all other forms of Salomonelaps. Barbour, T. 1921. Reptiles and amphibians from the British The taxon is also unusual in having 15 mid-body rows, versus Solomon Islands. Proc. New England zool. Club 7:91-112. the usual number of 17 (rarely 15 or 16) in the other populations Boseto, D. and Pikacha, P. (eds) 2016. A. report on baseline of Salomonelaps. Of the other forms of Salomonelaps 15 mid biodiversity inventory of Mount Maetambe to Kolobangara River body rows is only seen in S. melanurus and even then it is Corridor, Choiseul Island, Solomon Islands. Downloaded from relatively unusual. Of course S. melanurus is unique in terms of the website of Ecological Solutions Solomon Islands at: http:// Salomonelaps by the configuration of the dorsal banding. ecologicalsolutions-si.com/files/110494600.pdf The species S. woodfordi (Boulenger, 1888) from the New Boulenger, G. A. 1884. Diagnoses of new reptiles and Georgia island group in the Western District is separated from batrachians from the Solomon Islands, collected and presented all other species and subspecies of Salomonelaps McDowell, to the British Museum by H. B. Guppy, Esq., M. B., H. M. S. 1970 by the following suite of characters: The entire body, ‘Lark.’. Proc. Zool. Soc. London 1884:210-213. including the flanks is of a uniform colour, with each scale bordered by black, which results in a reticulated pattern as noted Boulenger, G. A. 1886. On the reptiles and batrachians of the by Boulenger (1888), McDowell (1970) and McCoy (2006). The Solomon Islands. Trans. Zool. Soc. London 12:35-62. back of the head is slightly darker in colour, while the front of the Boulenger, G. A. 1887. Catalogue of the Lizards in the British snout is lighter, but not white. All the dorsal head shields have Museum (Nat. Hist.) III. Lacertidae, Gerrhosauridae, Scincidae, dark etching and there are whitish upper labials along the lip Anelytropsidae, Dibamidae, Chamaeleontidae. London:575pp. line, which darken higher up, near the eye. There is no Bruns, T. R., Vedder, J. R. and Cooper, A. K. 1989. Geology of darkening on the posterior dorsal surface. Subcaudals are white the Shortland Basin Region, Central Solomons Trough, Solomon and patternless or rarely with darker margins. 165-179 ventrals. Islands - Review and New Findings. pp. 125-144 in Vedder, J.G., Most or all subcaudals are divided (versus all or mainly single in and Bruns, T. R., (editors), 1989. Geology and offshore the other species). resources of Pacific island arcs Solomon Islands and Rarely specimens of S. woodfordi may have a dorsal surface Bougainville, Papua New Guinea Regions: Houston, Texas, that is nearly a uniform dark grey colour, but all the scales of the Circum-Pacific Council for Energy and Mineral Resources, Earth first row, as well as some of those of the second row, have pale Science Series, v. 12. centres; ventrals and the subcaudals have dark etching on the Cogger, H. G. 1972. A new scincid lizard of the genus Hoser 2016 - Australasian Journal of Herpetology 31:12-21. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 21

Tribolonotus from Manus Island, New Guinea. Zool. 161:145-190. Mededelingen 47:202-210. Mys, B. 1988. The zoogeography of the scincid lizards from Court of Appeal Victoria. 2014. Hoser v Department of North Papua New Guinea (Reptilia: Scincidae). I. The Sustainability and Environment [2014] VSCA 206 (5 September distribution of the species. Bull. Inst. Roy. Sci. Nat. Belgique 2014). (Biologie) 58:127-183. Dahl, A. L. 1986. Review of the protected areas system in Ogilby, J. D. 1890. Report on a zoological collection from the Oceania. IUCN/UNEP, Gland, Switzerland. Solomon Islands. Part 2. Rec. Austr. Mus. 1:5-7. Duméril, A. M. C. and Bibron, G. 1839. Erpétologie Générale on Pianka, E. R. and Vitt, L. J. 2003. Lizards - Windows to the Histoire Naturelle Complète des Reptiles. Vol.5. Roret/Fain et Evolution of Diversity. University of California Press, Thunot, Paris:871 pp. Berkeley:347 pp. Gray, J. E. 1856. New genus of fish-scaled lizards (Scissosarae) Pyron, R. A., Burbrink, F. T. and Wiens, J. J. 2013. A phylogeny from New Guinea. Ann. Mag. Nat. Hist. (2)18:345-346. and revised classification of Squamata, including 4161 species Greer, A. E. 1982. A new species of Geomyersia (Scincidae) of lizards and snakes. BMC Evolutionary Biology 13:1-53. from the Admirality Islands, with a summary of the genus. Reeder, T. W. 2003. A phylogeny of the Australian Journal of Herpetology 16(1):61-66. Sphenomorphus group (Scincidae: Squamata) and the Greer, A. E. and Parker, F. 1967. A new scincid lizard from the phylogenetic placement of the crocodile skinks (Tribolonotus): northern Solomon Islands. Breviora (275):1-20. Bayesian approaches to assessing congruence and obtaining confidence in maximum likelihood inferred relationships. Greer, A. E. and Simon, M. 1982. Fojia bumui, an unusual new Molecular Phylogenetics and Evolution 27:384-397. genus and species of scincid lizard from New Guinea. Journal of Herpetology 16(2):131-139. Ride, W. D. L. (ed.) et al. (on behalf of the International Hagen, I. J., Donnellan, S. C. and Bull, M. 2012. Commission on Zoological Nomenclature) 1999. International Phylogeography of the prehensile-tailed skink Corucia zebrata code of Zoological Nomenclature (Fourth edition). The Natural on the Solomon Archipelago. Ecology and Evolution (2012), History Museum - Cromwell Road, London SW7 5BD, UK (also 2(6):1220-1234. commonly cited as “The Rules”, “Zoological Rules” or “ICZN 1999”). Hall, R., 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based Rittmeyer, E. N. and Austin, C. C. 2015. Combined next- reconstructions, model and animations. J. Asian Earth Sci. generation sequencing and morphology reveal fine-scale 20:353-431. speciation in Crocodile Skinks (Squamata: Scincidae: Tribolonotus). Mol Ecol. 2015 Jan, 24(2):466-83. doi: 10.1111/ Iskandar, D. T. and Erdelen, W. R. 2006. Conservation of mec.13030. Epub 2015 Jan 9. amphibians and reptiles in Indonesia: issues and problems. Russell, E. and Coupe, S. 1984. The Macquarie World Amphibian and Reptile Conservation 4(1):60-87. Illustrated Atlas. Kevin Weldon, Macquarie Library, Chatswood, Keogh, S. J., Scott, A. W., Fitzgerald, M. and Shine, R. 2003. NSW, Australia:511 pp. Molecular phylogeny of the Australian venomous snake genus Hoplocephalus (Serpentes, Elapidae) and conservation genetics Schmidt, K. P. 1932. Reptiles and Amphibians from the Solomon of the threatened H. stephensii. Conservation Genetics 4:57-65. Islands. Field Mus. Nat. Hist. Zool. Ser. 18(9):175-190. Victorian Civil and Administrative Tribunal (VCAT) 2015. Hoser v Kinghorn, J. R. 1928. Herpetology of the Solomon Islands. Rec. Department of Environment Land Water and Planning (Review Austral. Mus. 16:123-178. and Regulation) [2015] VCAT 1147 (30 July 2015). Kinghorn, J. R. 1937. A new species of skink from the Solomon Islands. Records of the Australian Museum 20(1):1-2. Williams, E. E. and Parker, F. 1964. The snake, genus Parapistocalamus on Bougainville, Solomon Islands. McCoy, M. 1980. Reptiles of the Solomon Islands. Wau Ecology Senckenberg. biol. 45:543-552. Institute Handbook 7. Wau Ecology Institute, Wau, Papua New Guinea. Zweifel, R. G. 1966. A New Lizard of the Genus Tribolonotus (Scincidae) from New Britain. American Museum Novitates McCoy, M. 2006. Reptiles of the Solomon Islands. Pensoft 2264:1-12. Series Faunistica 57:212 pp. CONFLICT OF INTEREST McDowell, S. B. 1970. On the status and relationships of the Solomon Island elapid snakes. Journal of Zoology, London The author has no known conflicts of interest in terms of this paper and conclusions within.

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Hoser 2016 - Australasian Journal of Herpetology 31:12-21. printed journals. Minimum print run of first printings is always at least fifty hard copies. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 22 Australasian Journal of Herpetology Australasian Journal of Herpetology 31:22-28. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online)

A division of the genus elapid genus Loveridegelaps McDowell, 1970 from the Solomon Islands, including formal description of four new species (Serpentes: Elapidae: Micropechiini: Loveridgelapina).

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 12 April 2016, Accepted 20 May 2016, Published 1 August 2016.

ABSTRACT The species described as Hoplocephalus elapoides Boulenger, 1890, from Florida Island in the Solomon Since the original description, widely divergent specimens have been found across the Solomon Islands. However, no herpetologist has considered whether or not there is more than one species currently under this umbrella. Inspection of specimens from the majority of islands Loveridegelaps have been found shows significant variation between specimens and of sufficient basis to warrant division into separate species. This includes consistent differences in scalation, colouration and hemipene morphology and can be reliably used to separate each form. As a result, of an assessment of the snakes and the relevant available genetic evidence involving species affected by the same geographical barriers, e.g. lizards of the genera Corucia Gray, 1855 and Tribolonotus Duméril and Bibron, 1839 as detailed by Austin et al. (2010) and Hagen et al. (2012), and the geological evidence of relevance, it is clear that the relevant forms are sufficiently divergent to warrant taxonomic recognition. Thus five distinctive forms are herein given taxonomic recognition as full species. Other than Loveridegelaps elapoides (Boulenger, 1890), none have available names and so four are named for the first time according to the provisions of the International Code of Zoological Nomenclature (Ride et al. 1999). These are: Loveridgelaps sloppi sp. nov. from the New Georgia Group of Islands. L. josephburkei sp. nov. from the Shortland Islands, L. yeomansi sp. nov. from Guadalcanal and L. fiacummingae sp. nov. from Malaita. Keywords: Taxonomy; snakes; genus; Loveridgelaps; species; elapoides; Boulenger; Solomon Islands; Solomons; Guadalcanal; Ngela; Nggela, Malaita; Shortland Island; New Georgia; Gizo; Santa Isabel; Florida Islands; Bougainville; new species; sloppi; josephburkei; yeomansi; fiacummingae.

INTRODUCTION The Solomons Black-banded Krait was originally described as between specimens and of sufficient basis to warrant division Hoplocephalus elapoides Boulenger, 1890, from a specimen into separate species. caught on Florida Island in the Solomon Islands. This includes consistent differences in scalation, colouration and It was transferred to a newly created monotypic genus hemipene morphology and can be reliably used to separate Loveridegelaps by McDowell in 1970 on the basis of significant each form, including the substantial body of evidence published morphological differences to all other elapid species. by McDowell (1970), who also inspected a number of specimens from across the Solomon Islands. Hoser (2012), assigned this and related species to a relevant tribe and subtibe, Micropechiini and Loveridgelapina As a result, of an assessment of the snakes and the relevant respectively. available genetic evidence involving species affected by the same geographical barriers, e.g. lizards of the genera Corucia Since the original description widely divergent specimens have Gray, 1855 and Tribolonotus Duméril and Bibron, 1839 as been found across most major island groups within the Solomon detailed by Austin et al. (2010) and Hagen et al. (2012), and the Islands. geological evidence of relevance, it is clear that the relevant However, until now no herpetologist has considered whether or forms are sufficiently divergent to warrant taxonomic recognition. not there is more than one species currently under this umbrella. They are clearly morphologically distinct, have significant Inspection of specimens from the majority of islands divergences with respect to very conservative characters, such Loveridegelaps have been found shows significant variation as hemipene morphology, indicating deep divergence and based Hoser 2016 - Australasian Journal of Herpetology 31:22-28. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 23

on parallel studies involving species affected by the same “Materials And Methods” under such a heading. barriers, clearly form genetically distinct, separately evolving The process involved in creating the final product for this and populations. other relevant papers has been via a combination of the Thus five distinctive forms are herein given taxonomic following: recognition as full species. Other than Loveridegelaps Genera and component species are audited to see if their elapoides (Boulenger, 1890), none have available names and so classifications are correct on the basis on known type four are named for the first time according to the provisions of specimens, locations and the like when compared with known the International Code of Zoological Nomenclature (Ride et al. phylogenies and obvious morphological differences between like 1999). species. These are: Loveridgelaps sloppi sp. nov. from the New Georgia Original descriptions and contemporary concepts of the species Group of Islands. L. josephburkei sp. nov. from the Shortland are matched with available specimens from across the ranges of Islands, L. yeomansi sp. nov. from Guadalcanal and L. the species to see if all conform to accepted norms. fiacummingae sp. nov. from Malaita. These may include those held in museums, private collections, It should also be noted that at the time of McDowell’s (1970) collected in the field, photographed, posted on the internet or study, he was isolated from molecular studies not available at held by individuals, and only when the location data is good and the time and therefore could only speculate as to the taxonomic any other relevant data available. significance of divergent traits he observed and documented. Where specimens do not appear to comply with the described However prior to the publication of this paper I was able to species (and accepted concept of the species), this non- match this evidence with what is now well known about the conformation is looked at with a view to ascertaining if it is recent geological past, in terms of ice-age maxima, changing worthy of taxonomic recognition or other relevant considerations sea levels and climates and the roles these play in speciation, on the basis of differences that can be tested for antiquity or either in these relevant snakes or other reptile taxa affected by deduced from earlier studies. the same factors. When this appears to be the case (non-conformation), the Divergences were ascertained on the basis of previous ice-age potential target taxon is inspected as closely as practicable with maxima connections between relevant islands as explained by a view to comparing with the nominate form or forms if other authors such as Bruns et al. (2009), Russell and Coupe (1984) similar taxa have been previously named. and recent molecular studies on both Corucia Gray, 1856 and Other relevant data is also inspected, including any available Tribolonotus Duméril and Bibron, 1839 as published by Austin et molecular studies which may indicate likely divergence of al. (2010) and Hagen et al. (2012), and the relevant sources populations. cited within. Where molecular studies are unavailable for the relevant taxon Notwithstanding the theft of relevant materials from this author in or group, other studies involving species and groups constrained an illegal armed raid on 17 August 2011, which were not by the same geographical or geological barriers, or with like returned (Court of Appeal Victoria 2014 and VCAT 2015) and not distribution patterns are inspected as they give reasonable returned in breach of various earlier court orders, I have made a indications of the likely divergences of the taxa being studied decision to publish this paper in view of the conservation herein. significance attached to the formal recognition of unnamed species and on the basis that further delays may in fact put Additionally other studies involving geological history, sea level these otherwise unnamed taxa at greater risk of extinction. and habitat changes associated with long-term climate change, including recent ice age changes in sea levels, versus known I also note that Boseto and Pikacha (2016), wrote of a serious sea depths are utilized to predict past movements of species alleged decline in abundance of Loveridgelaps in recent years, and genus groups in order to further ascertain likely divergences meaning the species in the genus are at heightened risk. between extant populations (as done in this very paper). They wrote: “Locals from Sasamugga also claimed that the rare When all available information checks out to show taxonomically and poorly known Loveridgelaps elapoides, one of the two distinct populations worthy of recognition, they are then terrestrial elapid snake species that has been previously recognized herein according to the rules of the International documented on Choiseul, was once common in the Sirebe Code of Zoological Nomenclature (Ride et al. 1999). Rainforest area, but that the arrival of R. marina caused it to decline dramatically.” This means that if a name has been properly proposed in the past, it is used. This is exactly what happens in this paper for the Thus five distinctive forms are herein given taxonomic taxon originally described as Hoplocephalus elapoides recognition on the basis that likely divergences exceed the Boulenger, 1890. timeline determined as significant by Keogh et al. (2003). Alternatively, if no name is available, one is proposed accoding Rafting between islands is not viewed as a significant means of to the rules of the Code as is done four times in this paper. dispersal or ongoing gene flow, beyond times of initial colonisation for reasons given by Hagen et al. (2012) and Balsai As a matter of trite I mention that if a target taxon or group does (1995) and also due to the absence of the genus from nearby check out as being “in order” or properly classified, a paper is island archipelagos beyond the Bougainville group. usually not published unless some other related taxon is named for the first time. Of relevance also is that the islands Guadalcanal and Malaita are separated from one another and the others by a sea depth The published literature relevant to the taxonomic judgements of more than 200 metres and hence do not appear to have been made within this paper includes papers relevant to Solomon joined at any stage in the last 5 million years. Islands species affected by the same physical barriers to MATERIALS AND METHODS dispersion as well as those directly relevant to Loveridgelaps and combined, they include the following: These are not formally explained in a number of my recent papers under the heading “Materials and methods” or similar, on Adler et al. (1995), Austin et al. (2010), Balsai (1995), Barbour the basis they are self evident to any vaguely perceptive reader. (1921), Boseto and Pikacha (2016), Boulenger (1884, 1886, However, the process by which the following taxonomy and 1890), Bruns et al. (1989), Cogger (1972), Dahl (1986), Duméril nomenclature in this and other recent papers by myself of and Bibron (1839), Gray (1856), Greer (1982), Greer and Parker similar form, has been arrived at, is explained herein for the (1967), Greer and Simon (1982), Hagen et al. (2012), Hall benefit of people who have recently published so-called (2002), Iskandar and Erdelen (2006), Keogh et al. (2003), “criticisms” online of some of my recent papers. They have Kinghorn (1928, 1937), McCoy (1980, 2006), McDowell (1970), alleged a serious “defect” by myself not formally explaining Mys (1988), Ogilby (1890), Pianka and Vitt (2003), Pyron et al. Hoser 2016 - Australasian Journal of Herpetology 31:22-28. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 24 Australasian Journal of Herpetology

(2013), Reeder (2003), Rittmeyer and Austin (2015), Russell and Malaita). In common with L. fiacummingae sp. nov. the Coupe (1984), Schmidt (1932), Williams and Parker (1964), hemipenis of L. elapoides is forked at subcaudal 7 or 8, versus 6 Zweifel (1966), and sources cited therein. in L. sloppi sp. nov. and 9 in L. yeomansi sp. nov.. The sulcus is Some material within descriptions below is repeated for different forked at subcaudals 5-8 in all species, but usually 7 in L. described taxa and this is in accordance with the provisions of elapoides. the International Code of Zoological Nomenclature and the legal Distribution: Restricted to the Florida Islands, Santa Isabel and requirements for each description. I make no apologies for this. Choiseul. GENUS LOVERIDGELAPS McDOWELL, 1970. LOVERIDGELAPS SLOPPI SP. NOV. Type species: Hoplocephalus elapoides Boulenger, 1890. Holotype: A male specimen at the Museum of Natural History, Diagnosis: Loveridgelaps McDowell, 1970 is defined in detail by London, UK, specimen number: 1933.3.4.2, from Gizo Island in McDowell (1970) and this diagnosis is adopted herein as correct the New Georgia group of islands in the Solomon islands. for the genus. The Museum of Natural History, London, UK is a facility that In summary the genus is defined as follows: Head slightly allows access to its holdings. flattened and barely distinct from the neck. Eyes very small and Diagnosis: Loveridgelaps sloppi sp. nov. from the New Georgia a diagnostic difference between this and other Solomon Islands group of islands is separated from all other Loveridgelaps elapids. Nasal is single or divided which contacts the preocular. McDowell, 1970, by the following suite of characters: The entire Rostral broad, frontal is as wide as long and wider than the head and anteriormost neck are yellowish white, except for a few supraoculars. 7 supralabials, with numbers 3 and 4 entering the dark flecks on the internasals and rostral and a narrow black eye. 1 or 2 postoculars. Temporals 1+2. Body is of moderate border around each eye and nostril; the black crossbands are shape and size is to about 1 meter in total length in adults. noticeably narrow laterally, but about five scale-lengths wide 17 Mid body rows, 193-218 ventrals, anal entire and 31-38 all vertebrally, where the separating orange-yellowish white zones divided subcaudals. are two or three scale-lengths wide. There are 42 dark crossbands on the body and tail, versus never more than 34 in The dorsal colouration is black with a regular series of bright any other species of Loveridgelaps. yellow bands along the vertebral line. Laterally the banding is white, usually separated from the yellow bands by one or two The pale zones and belly lack scattered black pigment, although rows of black scales. The head is usually white with irregular the black crossbands extend onto the tips of the ventrals and black markings on the rostral, labials, orbits and sometimes the completely traverse the subcaudals to form rings. occiput. Some melanotic forms are known. The hemipenis in male L. sloppi sp. nov. is unique for Distribution: Endemic to the Solomon Islands Archipelago, Loveridgelaps in the following properties: everted organ length to including: Shortland, Choiseul, Santa Isabel, Rob Roy, Vella subcaudal 6 (versus 7 or more in all other species), forked at Lavella, Gizo, Guadalcanal, Ngela (AKA Nggela) or Florida subcaudal 6 (versus 7 or more in all other species) and sulcus is Islands, Malaita. forked at subcaudal 5 (in common with L. fiacummingae sp. nov. Content: Loveridgelaps elapoides (Boulenger, 1890) (type from Malaita). species); L. sloppi sp. nov.; L. josephburkei sp. nov.; L. Loveridgelaps elapoides (Boulenger, 1890) from the Florida yeomansi sp. nov.; L. fiacummingae sp. nov.. Islands Group, Santa Isabel and Choiseul is separated from all LOVERIDGELAPS ELAPOIDES BOULENGER, 1890. other Loveridgelaps McDowell, 1970 (excluding L. josephburkei sp. nov.), by the following suite of characters: The snout and Holotype: A specimen at the Natural History Museum, London, ocular region are black, although the rest of the head and UK, specimen number, BM 1946. 1.18.98 (originally, anteriormost neck are yellowish white, with or without a pair of 89.3.29.191) collected at Florida Islands, Solomon Islands. small black spots on the occipital region of the head, behind the Diagnosis: Loveridgelaps elapoides (Boulenger, 1890) from the parietals. The black crossbands are of nearly equal width Florida Islands Group, Santa Isabel and Choiseul is separated laterally and vertebrally, about five or six scale-lengths wide, and from all other Loveridgelaps McDowell, 1970 (excluding L. separated by yellowish zones about three to four scales wide. josephburkei sp. nov.), by the following suite of characters: The Posteriorly, the pale zones contain black spots, and the black snout and ocular region are black, although the rest of the head crossbands extend onto the tips of the ventrals and encircle the and anteriormost neck are yellowish white, with or without a pair tail to form rings. of small black spots on the occipital region of the head, behind L. elapoides is separated from all other Loveridgelaps by having the parietals. The black crossbands are of nearly equal width a belly that is either unmarked and unspotted (Florida Islands laterally and vertebrally, about five or six scale-lengths wide and ) or with considerable black spotting except on the separated by yellowish zones about three to four scales wide. forebody (Choiseul and Santa Isabel). Posteriorly, the pale zones contain black spots, and the black For L. elapoides there are about 22 crossbands on the body and crossbands extend onto the tips of the ventrals and encircle the tail, (Florida Islands animals) or 34 (Choiseul and Santa Isabel). tail to form rings. L. elapoides is separated from all other Loveridgelaps by having The hemipenis in male L. elapoides is unique for Loveridgelaps a belly that is either unmarked and unspotted (Florida Islands in the following properties: The everted organ extends to animals) or with considerable black spotting except on the subcaudal nine, (versus 6 in Loveridgelaps sloppi sp. nov. from forebody (Choiseul and Santa Isabel). the New Georgia Group of Islands; 10 in L. yeomansi sp. nov. from Guadalcanal and 7-8 in L. fiacummingae sp. nov. from L. josephburkei sp. nov. known only from the Shortland Islands Malaita). In common with L. fiacummingae sp. nov. the is similar in most respects to L. elapoides which it would hemipenis of L. elapoides is forked at subcaudal 7 or 8, versus 6 otherwise be identified as, but differs from it by having small in L. sloppi sp. nov. and 9 in L. yeomansi sp. nov.. The sulcus is black spots, flecks and markings on the lower belly, but not on forked at subcaudals 5-8 in all species, but usually 7 in L. the mid-belly, and in not alternatively having an unmarked belly. elapoides. For L. elapoides there are about 22 crossbands on the body and L. josephburkei sp. nov. known only from the Shortland Islands tail, (Florida Islands animals) or 34 (Choiseul and Santa Isabel). is similar in most respects to L. elapoides which it would The hemipenis in male L. elapoides is unique for Loveridgelaps otherwise be identified as, but differs from it by having small in the following properties: The everted organ extends to black spots, flecks and markings on the lower belly, but not on subcaudal nine, (versus 6 in Loveridgelaps sloppi sp. nov. from the mid-belly, and in not alternatively having an unmarked belly. the New Georgia Group of Islands; 10 in L. yeomansi sp. nov. L. yeomansi sp. nov. from Guadalcanal is separated from all from Guadalcanal and 7-8 in L. fiacummingae sp. nov. from Hoser 2016 - Australasian Journal of Herpetology 31:22-28. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 25

other Loveridgelaps McDowell, 1970, by the following suite of animals) or with considerable black spotting except on the characters: Head as in L. elapoides, but black occipital spots forebody (Choiseul and Santa Isabel). There are anywhere from expanded into large blotches that extend nearly or to the edges 22 to 34 crossbands on the body and tail. of the parietals. The black crossbands are about four to six The hemipenis in male L. elapoides is unique for Loveridgelaps scale-lengths wide, and the light zones may or may not contain in the following properties: The everted organ extends to some black spotting, but not so much as to connect the black subcaudal nine, (versus 6 in Loveridgelaps sloppi sp. nov. from bands. The belly has a small amount of black spotting, and the the New Georgia Group of Islands; 10 in L. yeomansi sp. nov. black crossbands impinge extensively on the ventrals (so that from Guadalcanal and 7-8 in L. fiacummingae sp. nov. from the last one or two bands on the body may be complete rings, Malaita). In common with L. fiacummingae sp. nov. the like those of the tail). The crossbands are moderate in number hemipenis of L. elapoides is forked at subcaudal 7 or 8, versus 6 (28 to 33 on body and tail). in L. sloppi sp. nov. and 9 in L. yeomansi sp. nov.. The sulcus is The hemipenis in male L. yeomansi sp. nov. is unique for forked at subcaudals 5-8 in all species, but usually 7 in L. Loveridgelaps in the following properties: its length when everted elapoides. is 10 subcaudals (versus 9 or less for all other species), it is Loveridgelaps sloppi sp. nov. from the New Georgia group of forked at subcaudal number 9, versus 8 or less for all other islands is separated from all other Loveridgelaps McDowell, species, and the sulcus is forked at subcaudal number 7 or 8. 1970, by the following suite of characters: The entire head and L. fiacummingae sp. nov. from Malaita is separated from all anteriormost neck are yellowish white, except for a few dark other Loveridgelaps McDowell, 1970, by the following suite of flecks on the internasals and rostral and a narrow black border characters: The colouration noticeably tends towards being around each eye and nostril; the black crossbands are melanotic as described by both McCoy (2006) and McDowell noticeably narrow laterally, but about five scale-lengths wide (1970). In more detail, the black occipital blotches extend well vertebrally, where the separating orange-yellowish white zones onto the parietals and become confluent with the black ocular are two or three scale-lengths wide. There are 42 dark regions and with one another, thus isolating the white area on crossbands on the body and tail, versus never more than 34 in the frontal as an irregular pale crown patch. The dark any other species of Loveridgelaps. The pale zones and belly crossbands are very broad, but become narrower laterally, and lack scattered black pigment, although the black crossbands tend to fuse with one another through connection with the black extend onto the tips of the ventrals and completely traverse the pigment in the whitish zones, which makes the counting of subcaudals to form rings. blotches somewhat arbitrary; the pale zones are reduced in The hemipenis in male L. sloppi sp. nov. is unique for width to one scale-length vertebrally. The belly is white and Loveridgelaps in the following properties: everted organ length to without flecks or blotches, but the tail is encircled by black rings. subcaudal 6 (versus 7 or more in all other species), forked at Hemipene characteristics for L. fiacummingae sp. nov. appear subcaudal 6 (versus 7 or more in all other species) and sulcus is within the mid-range for the genus, reaching to subcaudal 7 or 8 forked at subcaudal 5 (in common with L. fiacummingae sp. nov. when fully everted, versus 6 in L. sloppi sp. nov., 9 in L. from Malaita). elapsoides and 10 in L. yeomansi sp. nov.. L. yeomansi sp. nov. from Guadalcanal is separated from all Distribution: L. sloppi sp. nov. is restricted to the New Georgia other Loveridgelaps McDowell, 1970, by the following suite of Group of Islands in the Solomon Islands. characters: Head as in L. elapoides, but black occipital spots Etymology: Named in honour of our living Great Dane (dog), expanded into large blotches that extend nearly or to the edges named “Slopp” for services to educating people about being nice of the parietals. The black crossbands are about four to six to animals, via our live shows and displays business. scale-lengths wide, and the light zones may or may not contain some black spotting, but not so much as to connect the black LOVERIDGELAPS JOSPEHBURKEI SP. NOV. bands. The belly has a small amount of black spotting, and the Holotype: A specimen at the Australian Museum, Sydney, black crossbands impinge extensively on the ventrals (so that NSW, Australia, specimen number: R.126267, from Near the last one or two bands on the body may be complete rings, Harehare Village, Shortland Island, Solomon Islands (7°03' S, like those of the tail). The crossbands are moderate in number 155°52' E). (28 to 33 on body and tail). The Australian Museum, Sydney, NSW, Australia is a facility that allows access to its holdings. The hemipenis in male L. yeomansi sp. nov. is unique for Diagnosis: L. josephburkei sp. nov. known only from the Loveridgelaps in the following properties: its length when everted Shortland Islands is similar in most respects to L. elapoides is 10 subcaudals (versus 9 or less for all other species), it is which it would otherwise be identified as, but differs from it by forked at subcaudal number 9, versus 8 or less for all other having small black spots, flecks and markings on the lower belly, species, and the sulcus is forked at subcaudal number 7 or 8. but not on the mid-belly, and in not alternatively having an L. fiacummingae sp. nov. from Malaita is separated from all unmarked belly. other Loveridgelaps McDowell, 1970, by the following suite of The hemipenes in male L. josephburkei sp. nov. are essentially characters: The colouration noticeably tends towards being similar to those of L. elapoides. melanotic as described by both McCoy (2006) and McDowell (1970). In more detail, the black occipital blotches extend well Loveridgelaps elapoides (Boulenger, 1890) from the Florida onto the parietals and become confluent with the black ocular Islands Group, Santa Isabel and Choiseul is separated from all regions and with one another, thus isolating the white area on other Loveridgelaps McDowell, 1970 (excluding L. josephburkei the frontal as an irregular pale crown patch. The dark sp. nov.), by the following suite of characters: The snout and crossbands are very broad, but become narrower laterally, and ocular region are black, although the rest of the head and tend to fuse with one another through connection with the black anteriormost neck are yellowish white, with or without a pair of pigment in the whitish zones, which makes the counting of small black spots on the occipital region of the head, behind the blotches somewhat arbitrary; the pale zones are reduced in parietals. The black crossbands are of nearly equal width width to one scale-length vertebrally. The belly is white and laterally and vertebrally, about five or six scale-lengths wide, and without flecks or blotches, but the tail is encircled by black rings. separated by yellowish zones about three to four scales wide. Hemipene characteristics for L. fiacummingae sp. nov. appear Posteriorly, the pale zones contain black spots, and the black within the mid-range for the genus, reaching to subcaudal 7 or 8 crossbands extend onto the tips of the ventrals and encircle the when fully everted, versus 6 in L. sloppi sp. nov., 9 in L. tail to form rings. elapsoides and 10 in L. yeomansi sp. nov.. L. elapoides is separated from all other Loveridgelaps by having a belly that is either unmarked and unspotted (Florida Islands Distribution: Known only from the Shortland Islands, Solomon

Hoser 2016 - Australasian Journal of Herpetology 31:22-28. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 26 Australasian Journal of Herpetology

Islands, but may also occur elsewhere in the Bougainville group in L. sloppi sp. nov. and 9 in L. yeomansi sp. nov.. The sulcus is of islands. forked at subcaudals 5-8 in all species, but usually 7 in L. Etymology: Named in honour of Joseph Burke of Joesph Burke elapoides. Law, Melbourne, Victoria in recognition of his services to the L. josephburkei sp. nov. known only from the Shortland Islands administration of justice in Melbourne, Australia, by defending is similar in most respects to L. elapoides which it would people against improper attacks from corrupt government otherwise be identified as, but differs from it by having small employees. black spots, flecks and markings on the lower belly, but not on LOVERIDGELAPS YEOMANSI SP. NOV. the mid-belly, and in not alternatively having an unmarked belly. Holotype: A specimen at the Australian Museum, Sydney, Loveridgelaps sloppi sp. nov. from the New Georgia group of NSW, Australia, specimen number: R.118881 from islands is separated from all other Loveridgelaps McDowell, Guadalcanal, Solomon Islands (9°32’S, 160°12’E). 1970, by the following suite of characters: The entire head and The Australian Museum, Sydney, NSW, Australia is a facility that anteriormost neck are yellowish white, except for a few dark allows access to its holdings. flecks on the internasals and rostral and a narrow black border around each eye and nostril; the black crossbands are Paratypes: A specimen at the Australian Museum, Sydney, noticeably narrow laterally, but about five scale-lengths wide NSW, Australia, specimen number: R.9301, from Guadalcanal, vertebrally, where the separating orange-yellowish white zones Solomon Islands (9°32’S, 160°12’E). are two or three scale-lengths wide. There are 42 dark A female specimen at the Museum of Comparative Zoology, crossbands on the body and tail, versus never more than 34 in Harvard University, USA, specimen number: MCZ 66899 from any other species of Loveridgelaps. Guadalcanal, Solomon Islands. The pale zones and belly lack scattered black pigment, although A male specimen at the Museum of Natural History, London, the black crossbands extend onto the tips of the ventrals and UK, specimen number: 1936.10.4.64 from Guadalcanal, completely traverse the subcaudals to form rings. Solomon Islands. The hemipenis in male L. sloppi sp. nov. is unique for A female specimen at the Museum of Natural History, London, Loveridgelaps in the following properties: everted organ length to UK, specimen number: 1967.834 from Guadalcanal, Solomon subcaudal 6 (versus 7 or more in all other species), forked at Islands. subcaudal 6 (versus 7 or more in all other species) and sulcus is Diagnosis: L. yeomansi sp. nov. from Guadalcanal is separated forked at subcaudal 5 (in common with L. fiacummingae sp. nov. from all other Loveridgelaps McDowell, 1970, by the following from Malaita). suite of characters: Head as in L. elapoides, but black occipital L. fiacummingae sp. nov. from Malaita is separated from all spots expanded into large blotches that extend nearly or to the other Loveridgelaps McDowell, 1970, by the following suite of edges of the parietals. The black crossbands are about four to characters: The colouration noticeably tends towards being six scale-lengths wide, and the light zones may or may not melanotic as described by both McCoy (2006) and McDowell contain some black spotting, but not so much as to connect the (1970). In more detail, the black occipital blotches extend well black bands. The belly has a small amount of black spotting and onto the parietals and become confluent with the black ocular the black crossbands impinge extensively on the ventrals (so regions and with one another, thus isolating the white area on that the last one or two bands on the body may be the frontal as an irregular pale crown patch. The dark complete rings, like those of the tail). The crossbands are crossbands are very broad, but become narrower laterally, and moderate in number (28 to 33 on body and tail). tend to fuse with one another through connection with the black The hemipenis in male L. yeomansi sp. nov. is unique for pigment in the whitish zones, which makes the counting of Loveridgelaps in the following properties: its length when everted blotches somewhat arbitrary; the pale zones are reduced in is 10 subcaudals (versus 9 or less for all other species), it is width to one scale-length vertebrally. The belly is white and forked at subcaudal number 9, versus 8 or less for all other without flecks or blotches, but the tail is encircled by black rings. species, and the sulcus is forked at subcaudal number 7 or 8. Hemipene characteristics for L. fiacummingae sp. nov. appear Loveridgelaps elapoides (Boulenger, 1890) from the Florida within the mid-range for the genus, reaching to subcaudal 7 or 8 Islands Group, Santa Isabel and Choiseul is separated from all when fully everted, versus 6 in L. sloppi sp. nov., 9 in L. other Loveridgelaps McDowell, 1970 (excluding L. josephburkei elapsoides and 10 in L. yeomansi sp. nov.. sp. nov.), by the following suite of characters: The snout and Distribution: Guadalcanal Island in the Solomon Islands. ocular region are black, although the rest of the head and Etymology: Named in honour of now deceased UK anteriormost neck are yellowish white, with or without a pair of herpetologist, Luke Yeomans. For details relating to the small black spots on the occipital region of the head, behind the etymology, see Hoser (2012). parietals. The black crossbands are of nearly equal width laterally and vertebrally, about five or six scale-lengths wide and LOVERIDGELAPS FIACUMMINGAE SP. NOV. separated by yellowish zones about three to four scales wide. Holotype: A male specimen at the American Museum of Natural Posteriorly, the pale zones contain black spots, and the black History (AMNH), New York, USA, specimen number: AMNH crossbands extend onto the tips of the ventrals and encircle the 43399, from Malaita, Solomon Islands. tail to form rings. The American Museum of Natural History (AMNH), New York, USA, is a facility that allows access to its holdings. L. elapoides is separated from all other Loveridgelaps by having a belly that is either unmarked and unspotted (Florida Islands Paratypes: 1/ A male specimen at the American Museum of animals) or with considerable black spotting except on the Natural History (AMNH), New York, USA, specimen number: forebody (Choiseul and Santa Isabel). AMNH 43400, from Malaita, Solomon Islands. For L. elapoides there are about 22 crossbands on the body and 2/ A specimen at the Australian Museum, Sydney, NSW, tail, (Florida Islands animals) or 34 (Choiseul and Santa Isabel). Australia, specimen number: R.2379 from Malaita, Solomon The hemipenis in male L. elapoides is unique for Loveridgelaps Islands (9°00’S, 161°00’E). in the following properties: The everted organ extends to 3/ A specimen at the Australian Museum, Sydney, NSW, subcaudal nine, (versus 6 in Loveridgelaps sloppi sp. nov. from Australia, specimen number: R.87382 from within a 3km radius the New Georgia Group of Islands; 10 in L. yeomansi sp. nov. of Bitaama, North Malaita, Solomon Islands (8°24’S, 160°36’E). from Guadalcanal and 7-8 in L. fiacummingae sp. nov. from Diagnosis: L. fiacummingae sp. nov. from Malaita is separated Malaita). In common with L. fiacummingae sp. nov. the from all other Loveridgelaps McDowell, 1970, by the following hemipenis of L. elapoides is forked at subcaudal 7 or 8, versus 6 suite of characters: The colouration noticeably tends towards

Hoser 2016 - Australasian Journal of Herpetology 31:22-28. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 27

being melanotic as described by both McCoy (2006) and L. yeomansi sp. nov. from Guadalcanal is separated from all McDowell (1970), separating this taxon from others in the genus. other Loveridgelaps McDowell, 1970, by the following suite of In more detail, the black occipital blotches extend well onto the characters: Head as in L. elapoides, but black occipital spots parietals and become confluent with the black ocular regions expanded into large blotches that extend nearly or to the edges and with one another, thus isolating the white area on the frontal of the parietals. The black crossbands are about four to six as an irregular pale crown patch. The dark crossbands are very scale-lengths wide, and the light zones may or may not contain broad, but become narrower laterally, and tend to fuse with one some black spotting, but not so much as to connect the black another through connection with the black pigment in the whitish bands. The belly has a small amount of black spotting, and the zones, which makes the counting of blotches somewhat black crossbands impinge extensively on the ventrals (so that arbitrary; the pale zones are reduced in width to one scale- the last one or two bands on the body may be complete rings, length vertebrally. The belly is white and without flecks or like those of the tail). The crossbands are moderate in number blotches, but the tail is encircled by black rings. (28 to 33 on body and tail). Hemipene characteristics for L. fiacummingae sp. nov. appear The hemipenis in male L. yeomansi sp. nov. is unique for within the mid-range for the genus, reaching to subcaudal 7 or 8 Loveridgelaps in the following properties: its length when everted when fully everted, versus 6 in L. sloppi sp. nov., 9 in L. is 10 subcaudals (versus 9 or less for all other species), it is elapsoides and 10 in L. yeomansi sp. nov.. forked at subcaudal number 9, versus 8 or less for all other Loveridgelaps elapoides (Boulenger, 1890) from the Florida species, and the sulcus is forked at subcaudal number 7 or 8. Islands Group, Santa Isabel and Choiseul is separated from all Distribution: Malaita Island in the Solomon Islands. other Loveridgelaps McDowell, 1970 (excluding L. josephburkei Etymology: Named in honour of Fia Cumming, former sp. nov.), by the following suite of characters: The snout and investigative journalist, of Lyons, ACT, Australia, formerly of ocular region are black, although the rest of the head and Chatswood, NSW, for her enormous contributions to wildlife anteriormost neck are yellowish white, with or without a pair of conservation in Australia as detailed in the book Smuggled-2: small black spots on the occipital region of the head, behind the wildlife Trafficking, Crime and Corruption in Australia (Hoser, parietals. The black crossbands are of nearly equal width 1996). laterally and vertebrally, about five or six scale-lengths wide, and The previous naming of one or more taxa in her honour as separated by yellowish zones about three to four scales wide. “cummingi” in the masculine, was deliberate as in Australian Posteriorly, the pale zones contain black spots, and the black slang language “it took balls’, an alleged male quality to take the crossbands extend onto the tips of the ventrals and encircle the enormous personal risks and costs she endured when tail to form rings. publishing her detailed expose’s of wildlife crime in Australia, L. elapoides is separated from all other Loveridgelaps by having and so the name “cummingi” as proposed by Hoser (1998) and/ a belly that is either unmarked and unspotted (Florida Islands or elsewhere, should not be amended unless mandatory animals) or with considerable black spotting except on the according to the rules of the International Code of Zoological forebody (Choiseul and Santa Isabel). Nomenclature (Ride et al. 1999). For L. elapoides there are about 22 crossbands on the body and NOTES ON THE DESCRIPTIONS FOR ANY POTENTIAL tail, (Florida Islands animals) or 34 (Choiseul and Santa Isabel). REVISORS The hemipenis in male L. elapoides is unique for Loveridgelaps Unless mandated by the rules of the International Code of in the following properties: The everted organ extends to Zoological Nomenclature, none of the spellings of the newly subcaudal nine, (versus 6 in Loveridgelaps sloppi sp. nov. from proposed names should be altered in any way. Should one or the New Georgia Group of Islands; 10 in L. yeomansi sp. nov. more newly named taxa be merged by later authors to be trated from Guadalcanal and 7-8 in L. fiacummingae sp. nov. from as single species, the order of prority of retention of names Malaita). In common with L. fiacummingae sp. nov. the should be as follows: sloppi; josephburkei; yeomansi; hemipenis of L. elapoides is forked at subcaudal 7 or 8, versus 6 fiacummingae, which is the order (page priority) of the in L. sloppi sp. nov. and 9 in L. yeomansi sp. nov.. The sulcus is descriptions within this text. forked at subcaudals 5-8 in all species, but usually 7 in L. REFERENCES CITED elapoides. Adler, G. H., Austin, C. C. and Dudley, R. 1995. Dispersal and L. josephburkei sp. nov. known only from the Shortland Islands speciation of skinks among archipelagos in the tropical Pacific is similar in most respects to L. elapoides which it would Ocean. Evolutionary Ecology 9:529-541. otherwise be identified as, but differs from it by having small Austin, C. C., Rittmeyer, E. N., Richards, S. J. and Zug, G. R. black spots, flecks and markings on the lower belly, but not on 2010. Phylogeny, historical biogeography and body size the mid-belly, and in not alternatively having an unmarked belly. evolution in Pacific Island Crocodile skinks Tribolonotus Loveridgelaps sloppi sp. nov. from the New Georgia group of (Squamata; Scincidae). Molecular Phylogenetics and Evolution islands is separated from all other Loveridgelaps McDowell, 57(1):227-236. 1970, by the following suite of characters: The entire head and Balsai, M. J. 1995. Husbandry and Breeding of the Solomon anteriormost neck are yellowish white, except for a few dark Islands Prehensile-tailed Skink, Corucia zebrata. The Vivarium flecks on the internasals and rostral and a narrow black border 7(1):4-11. around each eye and nostril; the black crossbands are noticeably narrow laterally, but about five scale-lengths wide Barbour, T. 1921. Reptiles and amphibians from the British vertebrally, where the separating orange-yellowish white zones Solomon Islands. Proc. New England zool. Club 7:91-112. are two or three scale-lengths wide. There are 42 dark Boseto, D. and Pikacha, P. (eds) 2016. A. report on baseline crossbands on the body and tail, versus never more than 34 in biodiversity inventory of Mount Maetambe to Kolobangara River any other species of Loveridgelaps. The pale zones and belly Corridor, Choiseul Island, Solomon Islands. Downloaded from lack scattered black pigment, although the black crossbands the website of Ecological Solutions Solomon Islands at: http:// extend onto the tips of the ventrals and completely traverse the ecologicalsolutions-si.com/files/110494600.pdf subcaudals to form rings. Boulenger, G. A. 1884. Diagnoses of new reptiles and The hemipenis in male L. sloppi sp. nov. is unique for batrachians from the Solomon Islands, collected and presented Loveridgelaps in the following properties: everted organ length to to the British Museum by H. B. Guppy, Esq., M. B., H. M. S. subcaudal 6 (versus 7 or more in all other species), forked at ‘Lark.’. Proc. Zool. Soc. London 1884:210-213. subcaudal 6 (versus 7 or more in all other species) and sulcus is Boulenger, G. A. 1886. On the reptiles and batrachians of the forked at subcaudal 5 (in common with L. fiacummingae sp. nov. Solomon Islands. Trans. Zool. Soc. London 12:35-62.

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Boulenger, G. A. 1890. Fourth contribution to the herpetology of 161:145-190. the Solomon Islands. Proc. Zool. Soc. London 1890:30-31. Mys, B. 1988. The zoogeography of the scincid lizards from Bruns, T. R., Vedder, J. R. and Cooper, A. K. 1989. Geology of North Papua New Guinea (Reptilia: Scincidae). I. The the Shortland Basin Region, Central Solomons Trough, Solomon distribution of the species. Bull. Inst. Roy. Sci. Nat. Belgique Islands - Review and New Findings. pp. 125-144 in Vedder, J.G., (Biologie) 58:127-183. and Bruns, T. R., (editors), 1989. Geology and offshore Ogilby, J. D. 1890. Report on a zoological collection from the resources of Pacific island arcs Solomon Islands and Solomon Islands. Part 2. Rec. Austr. Mus. 1:5-7. Bougainville, Papua New Guinea Regions: Houston, Texas, Pianka, E. R. and Vitt, L. J. 2003. Lizards - Windows to the Circum-Pacific Council for Energy and Mineral Resources, Earth Evolution of Diversity. University of California Press, Science Series, v. 12. Berkeley:347 pp. Cogger, H. G. 1972. A new scincid lizard of the genus Pyron, R. A., Burbrink, F. T. and Wiens, J. J. 2013. A phylogeny Tribolonotus from Manus Island, New Guinea. Zool. and revised classification of Squamata, including 4161 species Mededelingen 47:202-210. of lizards and snakes. BMC Evolutionary Biology 13:93. Court of Appeal Victoria. 2014. Hoser v Department of Reeder, T. W. 2003. A phylogeny of the Australian Sustainability and Environment [2014] VSCA 206 (5 September Sphenomorphus group (Scincidae: Squamata) and the 2014). phylogenetic placement of the crocodile skinks (Tribolonotus): Dahl, A. L. 1986. Review of the protected areas system in Bayesian approaches to assessing congruence and obtaining Oceania. IUCN/UNEP, Gland, Switzerland. confidence in maximum likelihood inferred relationships. Duméril, A. M. C. and Bibron, G. 1839. Erpétologie Générale on Molecular Phylogenetics and Evolution 27:384-397. Histoire Naturelle Complète des Reptiles. Vol.5. Roret/Fain et Ride, W. D. L. (ed.) et al. (on behalf of the International Thunot, Paris:871 pp. Commission on Zoological Nomenclature) 1999. International Gray, J. E. 1856. New genus of fish-scaled lizards (Scissosarae) code of Zoological Nomenclature (Fourth edition). The Natural from New Guinea. Ann. Mag. Nat. Hist. (2)18:345-346. History Museum - Cromwell Road, London SW7 5BD, UK (also Greer, A. E. 1982. A new species of Geomyersia (Scincidae) commonly cited as “The Rules”, “Zoological Rules” or “ICZN from the Admirality Islands, with a summary of the genus. 1999”). Journal of Herpetology 16(1):61-66. Rittmeyer, E. N. and Austin, C. C. 2015. Combined next- Greer, A. E. and Parker, F. 1967. A new scincid lizard from the generation sequencing and morphology reveal fine-scale northern Solomon Islands. Breviora (275):1-20. speciation in Crocodile Skinks (Squamata: Scincidae: Greer, A. E. and Simon, M. 1982. Fojia bumui, an unusual new Tribolonotus). Mol Ecol. 2015 Jan, 24(2):466-83. doi: 10.1111/ genus and species of scincid lizard from New Guinea. Journal of mec.13030. Epub 2015 Jan 9. Herpetology 16(2):131-139. Russell, E. and Coupe, S. 1984. The Macquarie World Illustrated Atlas. Kevin Weldon, Macquarie Library, Chatswood, Hagen, I. J., Donnellan, S. C. and Bull, M. 2012. NSW, Australia:511 pp. Phylogeography of the prehensile-tailed skink Corucia zebrata on the Solomon Archipelago. Ecology and Evolution (2012), Schmidt, K. P. 1932. Reptiles and Amphibians from the Solomon 2(6):1220-1234. Islands. Field Mus. Nat. Hist. Zool. Ser. 18(9):175-190. Hall, R., 2002. Cenozoic geological and plate tectonic evolution Victorian Civil and Administrative Tribunal (VCAT). 2015. Hoser of SE Asia and the SW Pacific: computer-based v Department of Environment Land Water and Planning (Review reconstructions, model and animations. J. Asian Earth Sci. and Regulation) [2015] VCAT 1147 (30 July 2015). 20:353-431. Williams, E. E. and Parker, F. 1964. The snake, genus Hoser, R. T. 1996. Smuggled-2: wildlife Trafficking, Crime and Parapistocalamus on Bougainville, Solomon Islands. Corruption in Australia. Kotabi P/L., Doncaster, Victoria, 3108, Senckenberg. biol. 45:543-552. Australia:260 pp. Zweifel, R. G. 1966. A New Lizard of the Genus Tribolonotus Hoser, R. T. 1998. Death Adders (Genus Acanthophis): An (Scincidae) from New Britain. American Museum Novitates overview, including descriptions of Five new species and One 2264:1-12. subspecies. Monitor - Journal of the Victorian Herpetological CONFLICT OF INTEREST Society 9(2):20-41. The author has no known conflicts of interest in terms of this Hoser, R. T. 2012. Yeomansus: A New Genus for the Slender paper and conclusions within. Racer (Serpentes:). Australasian Journal of Herpetology 14:3-5. Iskandar, D. T. and Erdelen, W. R. 2006. Conservation of amphibians and reptiles in Indonesia: issues and problems. Amphibian and Reptile Conservation 4(1):60-87. Australasian Journal of Herpetology ® Keogh, S. J., Scott, A. W., Fitzgerald, M. and Shine, R. 2003. ISSN 1836-5698 (Print) Molecular phylogeny of the Australian venomous snake genus ISSN 1836-5779 (Online) Hoplocephalus (Serpentes, Elapidae) and conservation genetics Publishes original research in printed form in of the threatened H. stephensii. Conservation Genetics 4:57-65. relation to reptiles, other fauna and related matters, Kinghorn, J. R. 1928. Herpetology of the Solomon Islands. Rec. including the subjects of classification, ecology, Austral. Mus. 16:123-178. public interest, legal, captivity, exposure of frauds, Kinghorn, J. R. 1937. A new species of skink from the Solomon “academic misconduct”, etc. Islands. Records of the Australian Museum 20(1):1-2. It is a peer reviewed printed journal published in McCoy, M. 1980. Reptiles of the Solomon Islands. Wau Ecology hard copy for permanent public scientific record, Institute Handbook 7. Wau Ecology Institute, Wau, Papua New with a sizeable print run and has a global audience. Guinea. Full details at: McCoy, M. 2006. Reptiles of the Solomon Islands. Pensoft http://www.herp.net Series Faunistica 57:212 pp. McDowell, S. B. 1970. On the status and relationships of the Solomon Island elapid snakes. Journal of Zoology, London Hoser 2016 - Australasian Journal of Herpetology 31:22-28. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 29 Australasian Journal of Herpetology 31:29-34. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online)

A review of the Xenodermidae and the Dragon Snake Xenodermus javanicus Reinhardt, 1836 species group, including the formal description of three new species, a division of Achalinus Peters, 1869 into two genera and Stoliczkia Jerdon, 1870 into subgenera (Squamata; Serpentes, Alethinophidia, Xenodermidae).

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 3 September 2015, Accepted 8 September 2015, Published 1 August 2016.

ABSTRACT Snakes in the genera Xenodermis Reinhardt, 1836 and Achalinus Peters, 1836 were reviewed. Regional variants of the putative species X. javanicus Reinhardt, 1836 were found to be sufficiently divergent to warrant being treated as full species. Other genera within the Xenodermidae were also reviewed. The species currently known as Achalinus meiguensis Hu and Zhao, 1966 was found to be sufficiently divergent both morphologically and by molecular analysis from other Achalinus Peters, 1869 species to warrant being placed in a separate genus. Stoliczkia Jerdon, 1870 currently contains two species that are divergent geographically and to a lesser extent morphologically and well separated by habitat. Therefore one is transferred to a new subgenus. As a result this paper formally names three new species of Xenodermus, namely X. oxyi sp. nov., X. crottyi sp. nov. and X. sloppi sp. nov., a new monotypic genus Fereachalinus gen. nov. and a new subgenus within Stoliczkia, namely Parastoliczkia subgen. nov.. Keywords: Taxonomy; snakes; nomenclature; Asia; Xenodermus; Achalinus; species; javanicus; meiguensis; new species; oxyi; crottyi; sloppi; new genus; Fereachalinus; new subgenus; Parastoliczkia.

INTRODUCTION In recent years, significant numbers of the distinctive Asian these snakes are found, indicating recent gene flow between the Dragon Snake Xenodermus javanicus Reinhardt, 1836, have populations is not likely to have happened. made their way into the private pet trade in Europe and the Based on both the genetic isolation of the known populations United States. The nominate form is allegedly common in parts and the morphological divergences of each, it is appropriate that of Java. The putative species is also known from elsewhere in they be recognized as distinct species. south-east Asia, including Peninsula Malaysia, Thailand, As there are no names available for the Peninsula Malaysia/ Sumatra and Borneo. Thailand population, that from northern Borneo or that from While this appears to be a widespread distribution, a close north-west Sumatra, each are named according to the rules of inspection of known specimens found that the taxon as currently the International Code of Zoological Nomenclature (Ride et al. recognized is largely confined to small areas of hilly forested 1999). habitat at higher elevations or immediately adjacent sites. By I note herein that the names Gonionotus plumbeus Gray, 1846 way of example, no specimens are known from flatter areas in and Xenoderma gonyonotus Lichtenstein and Martens, 1856 are Borneo, including the far south, or anywhere in Sumatra except unavailable for the two unnamed populations as both apply to for the far north-west which is relatively hilly. snakes from Java (Boulenger, 1893). Specimens from Thailand, possibly Burma and Peninsula At the same time this review took place, other taxa within the Malaysia are only known from the border areas of the Isthmus of Xenodermidae were inspected and reviewed in order to check Kra in the regions where the three countries (more-or-less) abut. that all were appropriately placed at the genus level and/or if Specimens from all known collection locations were inspected there were any obviously unrecognized taxa within any genus. and found to be morphologically divergent in characteristics that At the first level, it became clear that one species within the indicated long-term divergence of populations, notwithstanding genus Achalinus Peters, 1869 was significantly divergent from the evolutionarily conservative nature of the snakes. the others in the genus in terms of morphology and genetically. In spite of land bridges between the relevant parts of South-east This was the taxon formally described as Achalinus meiguensis Asia during ice-age minima, it is unlikely any gene flow would Hu and Zhao, 1966 which has noticeably larger body scales have occurred between populations in the recent past on the (evidenced by a lower mid body scale rows count), as well as basis of generally unsuitable habitat in the intervening spaces. significantly different head scalation to all the other species. Even now, the putative species Xenodermus javanicus remains Set apart morphologically, this taxon was also shown to be absent on low-lying islands between the higher land masses that significantly divergent in the molecular results of Pyron et al. Hoser 2016 - Australasian Journal of Herpetology 31:29-34. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 30 Australasian Journal of Herpetology

(2013) and so I have no hesitation in erecting a new genus for Chicago, Illinois, USA, specimen number: FMNH248958, this taxon. It is called Fereachalinus gen. nov.. collected from the Tawau District, Sabah, Borneo, Malaysia. The genus Stoliczkia Jerdon, 1870 currently contains two Diagnosis: Xenodermus oxyi sp. nov. is readily separated from species that are divergent geographically and to a lesser extent the other three species of Xenodermus by the presence of light morphologically and well separated by habitat. Therefore one is tubercles on the labial line. It is further distinguished from the transferred to a new subgenus named Parastoliczkia subgen. other species by the presence of a large number of red specks nov.. on the upper surface of the head and forebody. When I note further that the two relevant species are confined to specimens of Xenodermus from Java, Sumatra or the Isthmus highland areas and so have no obvious bridge between of Kra have red or orange specks, they are very few. populations. Xenodermus crottyi sp. nov. is readily separated from the other Publications relevant to the snakes currently referred to as three species of Xenodermus by the presence of a distinct Xenodermus javanicus Reinhardt, 1836 sensu lato, including the reddish tinge dorsally. taxonomic judgements made herein include the following: It is further separated from the other species by the ventral Boulenger (1893), Brongersma (1929), Chan-ard et al. (2015), colouration. In life, the individual scales are mainly black (in the Das (2012), David and Vogel (1996), de Rooij (1917), Flower centre), with thick white edges. In the other species, except for (1896), Gower et al. (2012), Grandison (1978), Gray (1849), X. sloppi sp. nov. this edging is bluish-black tinged. Haas (1950), Jan (1863), Kopstein (1938a, 1938b), Kudryavtsev X. sloppi sp. nov. is separated from the other three species in and Latyshev (2015), Lampe (1902), Lönnberg and Rendahl the genus by colouration. In this species the dorsal surface has (1925), Manthey and Grossmann (1997), Müller (1887, 1890), a dark brownish tinge (as opposed to being overwhelmingly Reinhardt (1836), Robinson and Kloss (1920), Rovatsos et al. bluish-grey in the others, excluding X. crottyi sp. nov. which is (2015), Savage (2015), Smith (1930, 1943), Stuebing and Inger bluish-grey above, but with a noticeable reddish tinge that (1999), Taylor (1965), Teynié et al. (2010), Tweedie (1983), Volz separates it from the rest). (1904), Welch (1988), Werner (1900, 1922) and sources cited therein. Ventrally, X. sloppi sp. nov. differs from the other species with each subcaudal being dark brown in colouration, with thick off- Key publications relevant to the snakes in the genus Achalinus white edges forming a well-defined boundary for each scale. Peters, 1869 as defined to date include the following: Barbour (1917), Boulenger (1893, 1888, 1908), Bourret (1937), Chen X. sloppi sp. nov. has 174 ventrals (similar to the other species), (2009), Fang and Wang (1983), Gao (1991), Goris and Maeda 128 subcaudals (versus 147 in X. javanicus) and 48 midbody (2004), Günther (1889), Guo et al. (1999), Hecht et al. (2013), rows, (versus 40 in X. javanicus, 48 in X. crottyi sp. nov. and 44 Hu and Zhao (1966), Hu et al. (1975), Inger et al. (1990), Kou in X. sloppi sp. nov.). and Wang (2003), Maki (1931), Mell (1931), Moriguchi and Naito X. javanicus is readily separated from the other species of (1979), Orlov, et al. (2000), Ota (2000), Ota and Toyama (1989a, Xenodermus by having (in life) a pinkish-white tongue, versus a 1989b), Ota et al. (1991), Peters (1869), Pyron et al. (2013), bluish-white tongue in the other three species. X. javanicus is Sang et al. (2009), Shie (2005), Smith (1943), Steindachner further separated by the very prominent blunt edged spines that (1913), Stejneger (1907, 1910), Toriba (1993), Van Denburgh form the longitunal rows running down the length of the body, (1912), Ziegler (2002), Zong and Ma (1983) and sources cited which are noticeably darker (as in black) than the bluish-grey therein. background colour of the upper body. While these raised spines Key publications relevant to the snakes in the genus Stoliczkia are also present in the other species, the individual spines are Jerdon, 1870 as defined to date include the following: Boulenger proportionately smaller and slightly more angular in appearance (1893, 1899), Das (1997, 2006, 2012), de Rooij (1917), Jerdon and only slightly darker in colour than the nearby greyish scales. (1870), Malkmus et al. (2002), Manthey (1983), Manthey and X. javanicus is also characterised by an average of 40 dorsal Grossmann (1997), Sharma (2004), Smith (1943) and sources mid body rows, versus 44 or more in the other three species. cited therein. The four species in the genus Xenodermus, namely X. oxyi sp. The materials and methods used as the basis for the following nov., X. crottyi sp. nov., X. sloppi sp. nov. and X. javanicus taxonomic results included a review of all available literature (as Reinhardt, 1836, and the genus itself are all defined and cited above) and specimens of all relevant taxa from all or most separated from all other snakes by the following unique suite of of where they are known to occur, when good locality information characters: was available for specimens. Teeth subequal, about 15 in each maxillary. Head distinct from I also note that, notwithstanding the theft of relevant materials neck, covered with granular juxtaposed scales; nostrils directed from this author in an illegal armed raid on 17 August 2011, forwards, in an undivided nasal; eye moderate, with a round which were not returned in breach of undertakings to the court pupil. Body slender, compressed, with small juxtaposed keeled (Court of Appeal Victoria 2014 and VCAT 2015), I have made a scales and longitudinal rows of large tubercles; ventrals well decision to publish this paper in view of the conservation developed. Tail long, with single subcaudals. Rostral small, significance attached to the formal recognition of unnamed taxa triangular, not visible from above; nasals meeting behind the and on the basis that further delays may in fact put these point of the rostral, followed by two pairs of small shields; labials unnamed taxa at greater risk of extinction, noting the extensive scarcely enlarged, about 20 on each side; no chinshields. About increase in human population in the area and associated habitat 40-48 scales across the body (mid body rows); the tubercles destruction occurring. disposed in pairs alternating with single ones on the vertebral XENODERMUS OXYI SP. NOV. line, and forming a single series along each side of the back. 173-185 ventrals; anal entire; 128-147 subcaudals. Colour is a Holotype: A preserved specimen at the Field Museum of dark greyish brown above and without markings, blotches or Natural History, Chicago, Illinois, USA, specimen number: stripes. The venter is light in colour, with or without darker FMNH 246192, collected from the Lahad Datu District in Sabah, markings in the center of each scale (adapted from Boulenger, Borneo, Malaysia. 1893). This facility allows access of its holdings to scientists. Distribution: Xenodermus oxyi sp. nov. is confined to the Paratypes: 1/ A preserved specimen at the Field Museum of northern two-thirds of Borneo, either in, or directly adjacent to Natural History, Chicago, Illinois, USA, specimen number: the northern and central mountain ranges. FMNH158613, collected from the Bintulu District, Sarawak, Etymology: Named in honour of my now deceased family pet Borneo, Malaysia. Great Dane named Oxyuranus (or “Oxy” for short), who spent 2/ A preserved specimen at the Field Museum of Natural History,

Hoser 2016 - Australasian Journal of Herpetology 31:29-34. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 31

some 8 years protecting the Hoser family children from thieves visible from above; nasals meeting behind the point of the and thugs and also played a valuable role in teaching people to rostral, followed by two pairs of small shields; labials scarcely be nice to animals via the Snakebusters educational wildlife enlarged, about 20 on each side; no chinshields. About 40-48 shows. scales across the body (mid body rows); the tubercles disposed I have no hesitation in naming a species in honour of a non- in pairs alternating with single ones on the vertebral line, and human inhabitant of this planet who has made a worthwhile forming a single series along each side of the back. 173-185 contribution to humanity and the welfare of other animals. ventrals; anal entire; 128-147 subcaudals. Colour is a dark By the way Oxyuranus Kinghorn, 1923 is the scientific name for greyish brown above and without markings, blotches or stripes. a genus of Australasian elapid snake. The venter is light in colour, with or without darker markings in the center of each scale, or as described for each species above XENODERMUS CROTTYI SP. NOV. (adapted from Boulenger, 1893). Holotype: A preserved specimen at the Field Museum of Distribution: Xenodermus oxyi sp. nov. is confined to the Natural History, Chicago, Illinois, USA, specimen number: northern two-thirds of Borneo, either in, or directly adjacent to FMNH178442, collected from Yala, southern Thailand. the northern and central mountain ranges. This facility allows access of its holdings to scientists. Etymology: Named in honour of my now deceased family pet Paratype: A specimen at the Museum of Natural History, UK, Great Dane/Rottweiler Cross named Crotalus (or “Crotty” for specimen number: 1939.1.4.5 (also stored as: short), who spent some 13 years protecting the Hoser family NHMUK:catalogue:1890625) from Victoria Point (now known as and research facility from thieves and thugs and also played a Kawthaung), Thailand. valuable role in teaching people to be nice to animals via the the Diagnosis: Xenodermus crottyi sp. nov. is readily separated educational displays we did at the time. from the other three species of Xenodermus by the presence of I have no hesitation in naming a species in honour of a non- a distinct reddish tinge dorsally. human inhabitant of this planet who has made a worthwhile It is further separated from the other species by the ventral contribution to humanity and the welfare of other animals. colouration. In life, the individual scales are mainly black (in the By the way Crotalus Linnaeus, 1758 is the scientific name for a centre), with thick white edges. In the other species, except for genus of mainly North American Pitvipers. X. sloppi sp. nov. this edging is bluish-black tinged. XENODERMUS SLOPPI SP. NOV. Xenodermus oxyi sp. nov. is readily separated from the other Holotype: A preserved specimen at the Museum Wiesbaden, three species of Xenodermus by the presence of light tubercles Wiesbaden, Germany, specimen number: 1119 collected at on labial line. It is further distinguished from the other species by Peiiak, Süd-Atjeh, Sumatra, Indonesia, in 1902 by. Dr. A. Fuchs the presence of a large number of red specks on the upper of Bornich, Germany. surface head and forebody. When specimens of Xenodermus from Java, Sumatra or the Isthmus of Kra have red or orange A detailed description of this specimen is in Lampe (1902). The specks, they are very few. Museum Wiesbaden allows access to its holdings. Diagnosis: Xenodermus sloppi sp. nov. is separated from the X. sloppi sp. nov. is separated from the other three species in other three species in the genus by colouration. In this species the genus by colouration. In this species the dorsal surface has a dark brownish tinge (as opposed to being overwhelmingly the dorsal surface has a dark brownish tinge (as opposed to bluish-grey in the others, excluding X. crottyi sp. nov. which is being overwhelmingly bluish-grey in the others, excluding X. bluish-grey above, but with a noticeable reddish tinge that crottyi sp. nov. which is bluish-grey above, but with a noticeable separates it from the rest). reddish tinge that separates it from the rest). Ventrally, X. sloppi sp. nov. differs from the other species with Ventrally, X. sloppi sp. nov. differs from the other species with each subcaudal being dark brown in colouration, with thick off- each subcaudal being dark brown in colouration, with thick off- white edges forming a well-defined boundary for each scale. white edges forming a well-defined boundary for each scale. X. sloppi sp. nov. has 174 ventrals (similar to the other species), X. sloppi sp. nov. has 174 ventrals (similar to the other species), 128 subcaudals (versus 147 in X. javanicus) and 48 midbody 128 subcaudals (versus 147 in X. javanicus) and 48 midbody rows, (versus 40 in X. javanicus, 48 in X. crottyi sp. nov. and 44 rows, (versus 40 in X. javanicus, 48 in X. crottyi sp. nov. and 44 in X. sloppi sp. nov.). in X. sloppi sp. nov.). X. javanicus is readily separated from the other species of Xenodermus oxyi sp. nov. is readily separated from the other Xenodermus by having (in life) a pinkish-white tongue, versus a three species of Xenodermus by the presence of light tubercles bluish-white tongue in the other three species. X. javanicus is on the labial line. It is further distinguished from the other further separated by the very prominent blunt edged spines that species by the presence of a large number of red specks on the form the longitunal rows running down the length of the body, upper surface head and forebody. When specimens of which are noticeably darker (as in black) than the bluish-grey Xenodermus from Java, Sumatra or the Isthmus of Kra have red background colour of the upper body. While these raised spines or orange specks, they are very few. are also present in the other species, the individual spines are Xenodermus crottyi sp. nov. is readily separated from the other proportionately smaller and slightly more angular in appearance three species of Xenodermus by the presence of a distinct and only slightly darker in colour than the nearby greyish scales. reddish tinge dorsally. X. javanicus is also characterised by an average of 40 dorsal It is further separated from the other species by the ventral mid body rows, versus 44 or more in the other three species. colouration. In life, the individual scales are mainly black (in the centre), with thick white edges. In the other species, except for The four species in the genus Xenodermus, namely X. oxyi sp. X. sloppi sp. nov. this edging is bluish-black tinged. nov., X. crottyi sp. nov., X. sloppi sp. nov. and X. javanicus Reinhardt, 1836, and the genus itself are all defined and X. javanicus is readily separated from the other species of separated from all other snakes by the following unique suite of Xenodermus by having (in life) a pinkish-white tongue, versus a characters: bluish-white tongue in the other three species. X. javanicus is Teeth subequal, about 15 in each maxillary. Head distinct from further separated by the very prominent blunt edged spines that neck, covered with granular juxtaposed scales; nostrils directed form the longitunal rows running down the length of the body, forwards, in an undivided nasal; eye moderate, with round pupil. which are noticeably darker (as in black) than the bluish-grey Body slender, compressed, with small juxtaposed keeled scales background colour of the upper body. While these raised spines and longitudinal rows of large tubercles; ventrals well developed. are also present in the other species, the individual spines are Tail long, with single subcaudals. Rostral small, triangular, not proportionately smaller and slightly more angular in appearance Hoser 2016 - Australasian Journal of Herpetology 31:29-34. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 32 Australasian Journal of Herpetology and only slightly darker in colour than the nearby greyish scales. long, subcaudals single. Hypapophyses developed throughout X. javanicus is also characterised by an average of 40 dorsal the vertebral column. mid body rows, versus 44 or more in the other three species. Distribution: Restricted to West and , China at The four species in the genus Xenodermus, namely X. oxyi sp. an elevation of 1200-1400 m. nov., X. crottyi sp. nov., X. sloppi sp. nov. and X. javanicus Etymology: Named in reflection of the fact that the relevant Reinhardt, 1836, and the genus itself are all defined and taxon is nearly, but not quite “Achalinus”. separated from all other snakes by the following unique suite of Content: Fereachalinus meiguensis (Hu and Zhao, 1966) characters: (monotypic). Teeth subequal, about 15 in each maxillary. Head distinct from PARASTOLICZKIA SUBGEN. NOV. the neck, covered with granular juxtaposed scales; nostrils Type species: Stoliczkia borneensis Boulenger, 1899. directed forwards, in an undivided nasal; eye moderate, with round pupil. Body slender, compressed, with small juxtaposed Diagnosis: Parastoliczkia subgen. nov. from the island of keeled scales and longitudinal rows of large tubercles; ventrals Borneo is physically similar to Stoliczkia Jerdon, 1870 from the well developed. Tail is long, with single subcaudals. Rostral Khazi Hills in north-east India. small, triangular, not visible from above; nasals meeting behind However, Parastoliczkia subgen. nov. differs from Stoliczkia by the point of the rostral, followed by two pairs of small shields; having a large triangular rostral (versus tiny in Stoliczkia), ten labials scarcely enlarged, about 20 on each side; no chinshields. supralabials (as opposed to eight in Stoliczkia), a large loreal About 40-48 scales across the body (mid body rows); the that is much longer than deeo (versus a tiny loreal in Stoliczkia), tubercles disposed in pairs alternating with single ones on the roughly 124 subcaudals in females (versus 115 in Stoliczkia); vertebral line, and forming a single series along each side of the colouration that is generally rufous, with large blackish spots, at back. 173-185 ventrals; anal entire; 128-147 subcaudals. Colour least as large as the space between them, disposed more or is a dark greyish brown above and without markings, blotches or less regularly in three longitudinal series, with a brownish venter, stripes. The venter is light in colour, with or without darker with each scale etched with yellowish (versus purplish brown markings in the center of each scale or as described for each above; three or four outer rows of scales and the ventrals are species above (adapted from Boulenger, 1893). white with brown edges, in Stoliczkia). Distribution: Known only from north-west Sumatra, in the Both Parastoliczkia subgen. nov. and Stoliczkia are separated vicinity of the locations of Perlak (AKA Peureulak) and from all other snakes by the following unique suite of characters: Sibolangit, north-west Sumatra, Indonesia. Teeth small, subequal, about 14 in each maxillary. Head distinct Etymology: Named in honour of the family pet Great Dane from neck, covered with large shields; nostrils directed forwards; named Slopp, who has spent some years protecting the Hoser nasal shield undivided; eye small, with round pupil. A pair of very family and research facility from thieves and thugs and also narrow internasals; a pair of large praefrontals, separated from played a valuable role in teaching people to be nice to animals the frontal and supraoculars by a series of small scales; frontal a via the the educational displays we do via Snakebusters: little broader than long, a little shorter than the parietals; Australia’s best reptiles, being the only hands on reptile shows in supraocular very small; narrow parietals nearly twice as long as Australia. the frontal; a large praeocular and two postoculars; eye very prominent, with vertically subelliptic pupil, nostril very large I have no hesitation in naming a species in honour of a non- temporals small, scale-like; a single pair of small chin-shields, in human inhabitant of this planet who has made a worthwhile contact with three lower labials. Scales in 30 rows, dorsals contribution to humanity and the welfare of other animals. separated by naked skin, laterals larger and juxtaposed. FEREACHALINUS GEN. NOV. Ventrals about 210 in females; anal single. Body slender, Type species: Achalinus meiguensis Hu and Zhao, 1966. compressed; scales elliptical, juxtaposed, strongly keeled, Diagnosis: Fereachalinus gen. nov. is readily separated from increasing in size towards the ventrals, which are well Achalinus Peters, 1869 by the following unique suite of developed. Tail long; subcaudals single. characters: The internasal is fused to the prefrontal (as opposed Distribution: Known only from hillier parts of the northern half of to being separated from the prefrontal by a suture in all species the island of Borneo, mainly, but not exclusively on the of Achalinus), the mental is in contact with the anterior Malaysian side. postmental (versus being separated from the anterior Etymology: Named Parastoliczkia as it isn’t exactly Stoliczkia postmental by the second infralabial in all species of Achalinus), Jerdon, 1870. 19 midbody scale rows (versus 21 or more in all species of Content: Stoliczkia (Parastoliczkia) borneensis Boulenger, 1899 Achalinus). (monotypic). The divergent species Achalinus formosanus Boulenger, 1908 NOTES ON THE DESCRIPTIONS FOR ANY POTENTIAL with 25 or 27 midbody rows has been placed in the genus REVISORS Achalinopsis Steindachner, 1913, which has been accepted by some authors (e.g. Zong and Ma, 1983), but not others (e.g. Unless mandated by the rules of the International Code of Shie, 2005). Zoological Nomenclature, none of the spellings of the newly proposed names should be altered in any way. Should one or In any event, this taxon is not as divergent from the nominate more newly named taxa be merged by later authors to be species for Achalinus, namely Peters, 1869 treated as a single species, the order of prority of retention of (23 midbody rows) as “Achalinus meiguensis Hu and Zhao, names should be the order (page priority) of the formal 1966”. descriptions within this text. Both the genera Fereachalinus gen. nov. and Achalinus are REFERENCES CITED separated from all other snakes by the following unique suite of characters: Maxillary teeth 22 to 25, small equal; mandibular Barbour, T. 1917. A most regretable tangle of names. teeth equal. Head not distinct from the neck; eye small, with Occasional Papers of the Museum of Zoology, University of round or vertically sub-elliptic pupil; nostril rather large, pierced Michigan (44):1-9. or not pierced in the anterior of two nasals; posterior nasal Boulenger, G. A. 1893. Catalogue of the snakes in the British concave; no praeocular, loreal extending from the nasals to the Museum (Nat. Hist.) I. London (Taylor and Francis):448 pp. eye; postoculars not distinct from anterior temporals. Body Boulenger, G. A. 1888. Description of two new snakes from cylindrical, slender; scales lanceolate, feebly imbricate, keeled, Hongkong, and note on the dentition of Hydrophis viperina. 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Hoser 2016 - Australasian Journal of Herpetology 31:29-34. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 33

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Achalinus formosanus Boulenger (Xenoderminae: Colubridae: Steindachner, F. 1913. Über zwei neue Schlangenarten aus Ophidia), with Description of a New Subspecies. Copeia 1989, Formosa. Anz. Kaiserl. Akad. Wiss. Wien, Math.-Nat. Ser. (3):597-602. 50:218-220. Ota, H., Kamezaki, N., Hikida, T., Matsui, M., Mori, A., Hayashi, Stejneger, L. H. 1907. Herpetology of Japan and adjacent T. and Tanabe, S. 1991. Karyotype of Achalinus spinalis from territory. Bull. US Natl. Mus. 58: xx+1-577. Japan: The first karyological description of a xenodermine snake Stejneger, L. H. 1910. The batrachians and reptiles of Formosa. (Colubridae). Japanese Journal of Herpetology 14:12-14. Proc. US Natl. Mus. 38:91-114. Peters, W. C. H. 1869. Über neue Gattungen und neue oder Stuebing, R. B. and Inger, R. F. 1999. A field guide to the weniger bekannte Arten von Amphibien (Eremias, Dicrodon, snakes of Borneo. Natural History Publications (Borneo), Kota Euprepes, Lygosoma, Typhlops, Eryx, Rhynchonyx, Kinabalu;254 pp. Elapomorphus, Achalinus, Coronella, Dromicus, Xenopholis, Taylor, E. H. 1965. The serpents of Thailand and adjacent Anoplodipsas, Spilotes, Tropidonotus). Monatsber. k. preuss. waters. Univ. Kansas Sci. Bull. 45(9):609-1096. Akad. Wiss. Berlin, 1869:432-447. Teynié, A., David, P. and Ohler, A. 2010. Note on a collection of Pyron, R. A., Burbrink, F. T. and Wiens, J. J. 2013. A phylogeny Amphibians and Reptiles from Western Sumatra (Indonesia), and revised classification of Squamata, including 4161 species with the description of a new species of the genus Bufo. Zootaxa of lizards and snakes. BMC Evolutionary Biology 13:1-53. 2416:1-43. Reinhardt, J. T. 1836. Afhandling om Xenodermus javanicus. Toriba, M. 1993. Present location of the type specimens Oversigt over det Kongelige Danske videnskabernes selskabs described by Maki. The Snake 25:71-72. forhandlinger. Kjobenhavn 3:6-7. Tweedie, M. W. F. 1983. The snakes of Malaya. (Third edition). Ride, W. D. L. (ed.) et al. (on behalf of the International Singapore National Printers, Singapore:1-167+12 plates. Commission on Zoological Nomenclature) 1999. International code of Zoological Nomenclature (Fourth edition). The Natural Van Denburgh, J. 1912. Advanced diagnoses of new reptiles History Museum - Cromwell Road, London SW7 5BD, UK (also and amphibians from the Loo Choo Islands and Formosa. commonly cited as “The Rules”, “Zoological Rules” or “ICZN Privately published by author, San Francisco:8 pp. 1999”). Victorian Civil and Administrative Tribunal (VCAT). 2015. Hoser Robinson, H. C. and Kloss, C. B. 1920. A nominal list of the v Department of Environment Land Water and Planning (Review species of reptiles and batrachians occurring in Sumatra. J. fed. and Regulation) [2015] VCAT 1147 (30 July 2015). Malay States Mus., 8(2):297-306. Volz, W. 1904. Schlangen von Palembang (Sumatra) (Reise von Rovatsos, P. M., Johnson, M. and Kratochviìl, L. 2015. Dr. Walter Volz). Zool. Jahrb. Jena, (Syst.), 20(5):491-508. Differentiation of Sex Chromosomes and Karyotype Welch, K. R. G. 1988. Snakes of the orient: a checklist. Robert Characterisation in the Dragonsnake Xenodermus javanicus F. Krieger Publishers, Malabar, Florida, USA:i-vii+183 pp. (Squamata: Xenodermatidae). Cytogenet Genome Res., DOI: Werner, F. 1900. Reptilienund Batrachier aus Sumatra, 10.1159/000441646 gesammelt von Herrn Gustav Schneider jr. im Jahre 1897-98. Sang, N. V., Cuc, H. T. and Nguyen, Q. T. 2009. Herpetofauna of Zool. Jb., (Abt. Sydt.), 13(6):479-508, pl.31-35. Vietnam. Chimaira, Frankfurt:768 pp. Werner, F. 1922. Synopsis der Schlangenfamilien der Savage, J. M. 2015. What are the correct family names for the Amblycephaliden und Viperiden nebst Übersicht über die taxa that include the snake genera Xenodermus, Pareas, and kleineren Familien und die Colubriden der Calamaria? Herpetological Review 46(4):664-665. Achrochordinengruppe. Auf Grund des Boulengerschen Sharma, R. C. 2004. Handbook Indian Snakes. Akhil Books, Schlangenkatalogs (1893-1896). Arch. Naturgesch. Berlin, (A), New Delhi:292 pp. 88(8):185-244. Shie, J. 2005. Phylogeography of Odd-scaled Snake Achalinus Ziegler, T. 2002. Die Amphibien und Reptilien eines in Taiwan. Masters thesis, National Sun Yat-sen University:53 Tieflandfeuchtwald-Schutzgebietes in Vietnam. Natur und Tier pp. Verlag (Münster):342 pp. Smith, M. A. 1930. The Reptilia and Amphibia of the Malay Zong, Y. and Ma, J. 1983. A new species of the genus Peninsula. Bull. Raffles Mus., 3:i-xviii+1-149. Achalinopsis from and the restoration of this genus. Acta Herpetologica Sinica 2(2):61-63. Smith, M. A. 1943. The Fauna of British India, Ceylon and Burma, Including the Whole of the Indo-Chinese Sub-Region. CONFLICT OF INTEREST Reptilia and Amphibia. 3 (Serpentes). Taylor and Francis, The author has no known conflicts of interest in terms of this London:583 pp. paper and conclusions within.

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A second new Tropidechis Günther, 1863 from far north Queensland (Squamata: Serpentes: Elapidae).

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 2 February 2016 Accepted 12 March 2016, Published 1 August 2016.

ABSTRACT The Australian Rough-scaled Snakes Tropidechis Günther, 1863, were formally divided by Hoser (2003) into two divergent species. These were the long-recognized T. carinatus (Krefft, 1863) from northern New South Wales and south-east Queensland, and the population from the wet tropics of far north-east Queensland, formally named as T. sadlieri Hoser, 2003. The two species are separated by a straight line distance of about 1,000 km at their closest points. Inspection of further specimens from the wet tropics has yielded two morphologically distinct and geographically isolated populations in the area. The nominate form of T. sadlieri is from the southern wet tropics, this area being Mount Spec in the south to Bellenden Ker, just south of Cairns in the north. The as yet unnamed population is found in the northern wet tropics in the region from Julatten/Mount Lewis in the south to at least the Windsor Tableland/Thornton Peak area in the north. It is herein formally named as T. jessejacksoni sp. nov. in honour of Jesse Louis Jackson, Sr. a well-known American civil rights activist, Baptist minister, and politician, in recognition of his lifetime’s struggle for the basic human rights for non-white citizens of the United States of America. The taxon is named in accordance with the provisions of the International Code of Zoological Nomenclature (Ride et al. 1999). Keywords: Taxonomy; Snakes; North Queensland; Queensland; Australia; wet tropics; northern wet tropics; Mount Lewis; Cairns; Julatten; Windsor Tableland; Tropidechis; carinatus; sadlieri; new species; jessejacksoni.

INTRODUCTION Following some years of field research, inspection of numerous By way of corroboration, I merely note that no other single live specimens in captivity and inspection of Tropidechis rainforest obligate species has a distribution that includes and is holdings in the Melbourne, Sydney and Brisbane Museums, restricted to south-east Queensland/Northern New South Wales Hoser (2003) divided the two widely disjunct populations of the rainforests and those of the north Queensland wet tropics. Rough-scaled Snake Tropidechis carinatus (Krefft, 1863) into Those species long thought of as inhabiting both regions (e.g. two species. Until then, Tropidechis Günther, 1863 had been Phyllurus cornutus Ogilby, 1892, a species more recently treated by all herpetologists as being monotypic for the one transferred to the genus Saltuarius Couper, Covacevich and species. Moritz, 1993), was split into several species some years back as detailed in Cogger (2014). The most obvious differences between the southern taxon from south-east Queensland and north-east New South Wales and An audit of rainforest obligate species in the wet tropics of the putative northern species from the wet tropics of north-east Australia, by myself in the period post-dating the 2003 Queensland, were spelt out by Hoser (2003) in the formal description of T. sadlieri, has found many putative species from description of T. sadlieri Hoser, 2003. the region being actually composite. This has usually been a situation of two species, one in the southern wet tropics and Notwithstanding the unscientific denials of the obvious by a band another in the northern wet tropics, separated by a gap zone of thieves known as the Wüster gang, who as of end 2015 were sited in the general area between Cairns in the south and still denying the existence of the taxon T. sadlieri as detailed on Julatten/Mount Lewis in the north. the website the gang control called the “Reptile Database” at http://reptile-database.reptarium.cz/ Some of these putative species as identified by Cogger et al. species?genus=Tropidechis&species=carinatus, which as of 12 (1983) that I have found to be actually two and divided by this December 2015 stated “Synonymy: Tropidechis sadlieri is barrier include Saltuarius cornutus Ogilby, 1892 (the southern probably a synonym of T. carinatus (Wilson & Swan 2008, population being the nominate form), Carphodactylus laevis Wüster, pers. comm., 15 Dec 2010).”; the specific status of T. Günther, 1897 (the southern population being the nominate sadlieri has been near universally recognized by Australian form) and Gnypetoscincus queenslandiae (De Vis, 1890) (the southern population being the nominate form) for all of which Hoser 2016 - Australasian Journal of Herpetology 31:35-38. herpetologists since the date of the original description. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 36 Australasian Journal of Herpetology molecular evidence is available and which confirms species gap between the two populations. level divergences. I am presuming it is more likely to be a predator in the lowlands With this in mind, putative T. sadlieri were revisited to see if A/ or drier habitats, as opposed to thermal or other properties of These snakes were divided by the same barrier and B/ If snakes the intervening habitat itself. However the inability of on either side of thre gap were the same or different. Tropidechis to breach dry zones between habitats is evident also These were the two questions asked (or hypotheses to be in the southern population from south-east Queensland and tested) and the answers were arrived at in the first instance by nearby New South Wales. simple checking of the Museum databases to see where Many rainforest obligate species and genera of reptiles and specimens had been found. frogs (e.g. Adelynhosersaur Hoser, 2013, Mixophyes Günther, Notwithstanding the ability of snakes such as Tropidechis 1864), were able to cross the dry zone of the Hunter Valley in species to hitch rides from place to place and so be caught and New South Wales, to be able to inhabit rainforests, south of recorded from locations that they do not originate from, the here, thus occupying both sides of this gap as indicated by the Museum databases for Australian holdings showed the well relevant distribution information in Hoser (1989) and/or Cogger defined gap from Cairns to Julatten/Mount Lewis having an (2014). However Tropidechis was unable to do so, only being absence of specimens. found north of this gap and not in areas of suitable habitat to the south (Hoser 1989, Cogger, 2014). Not one single animal had been either lodged in a museum, or reported on other specimen databases by way of “human Hence I have no doubt as to the effectiveness of the barrier observation” or the like. between the southern wet tropics population and the northern While this could be attributed to non-collection in this zone, this wet tropics populations of Tropidechis in the recent geological is thought unlikely due to the regular fieldwork in the area and past, including in the period preceding the most recent significant human presence in the region, but not of such a interglacial. degree as to exterminate such snakes if present. Also of relevance to this paper and the diagnosis of the new Of relevance also is the active capture and removal of all local species, I must mention a significant error in my book Australian species of snakes by government licensed snake handlers from Reptiles and Frogs (Hoser, 1989). properties in the area, which again has failed to yield a single On page 173 at the top right hand corner, I produced a photo Tropidechis specimen. captioned “Rough-scalled Snake Tropidechis carinatus (Jellaten, Due to the nature of the job of snake catchers in a given area QLD).” and that over time, all locally occurring species are found and The town “Jellaten” is in fact spelt “Julatten”, but that is not a caught by the snake catchers, the non-capture of Tropidechis in serious error as the mistake is obvious and so no confusion is the relevant gap zone, implies in the strongest possible terms of likely to have occurred. a genuine absence. The taxonomy used, reflected that at the time and as the book As for the snakes themselves, while the number of specimens was not about changing taxonomy or nomenclature, the from the northern wet tropics was small (I was only able to identification of the snake as “Tropidechis carinatus” was also in inspect less than ten in total), versus about 30 from the southern order. wet tropics (still a small number), the differences between both More significantly the snake depicted is almost certainly NOT groups was stark and consistent, as most readily evidenced in from Julatten as indicated. This I know as the head scalation the configuration of the labial scales. With these being a does not conform with those from Julatten, but rather it conservative character in terms of morphological evolution, it is conforms to the southern wet tropics form instead, defined clear that the two populations have been separated for an herein as T. sadlieri. The northern wet tropics form from extended geological period and not just the beginning of the Julatten has different scalation as outlined in the formal most recent interglacial about 12,500 years ago description of the new species below. Being aware of the fact that for most of the Pleistocene, the As to how the errors occurred, an explanation is required. climate in the region has been drier and that Tropidechis is in The snake photographed was in a cage at the private home of a the normal course of events a rainforest obligate species, it is Mr. Michael Cermak, who at the time when the photo was taken likely that the range of these snakes in north Queensland has in early 1983 lived at Manunda, Queensland. expanded in the recent past (last 12,500 years), rather than contracted and that the affected populations have been isolated He advised me of the alleged collection locality of the snake and over a time line similar to that indicated by the molecular wrote down the spelling of “Jellaten” on a piece of paper, both of evidence for the relevant Carphodactylidae species as outlined which were later transcribed to the slide mounts and then into by Couper, Covacevich and Moritz (2000) or for putative the book when it was published 6 years later. Gnypetoscincus queenslandiae as detailed by Moritz et al. It has since emerged that Mr. Cermak is at best described as (1993). unreliable, or perhaps better described as complete liar and For putative Gnypetoscincus queenslandiae Moritz et al. (1993) crook. To this end, he has established a reputation for making found the two populations diverged over 5 million years ago. false and unreliable statements for his own commercial purposes. In recent years (2010-2016) he has regularly made Significantly and in spite of presumed Holocene expansion of wild and ridiculous claims on social media (e.g. Facebook), the two Tropidechis populations in north Queensland, they have meaning that any information that may now be volunteered by not yet met and so remain isolated from one another and Cermac about the relevant snake would have to be dismissed as continue to evolve as separate species. unreliable in any event. On the basis of the obvious differences between the two While I could guess that the relevant snake came from a third populations of these snakes and continued isolation of each, I party and hence Cermak may not have even known where it have absolutely no hesitation in naming the northern population came from, this is speculation only. He may have caught it as a new species. himself! Of peripheral relevance and noting the ability of snakes in The only thing about the snake that is certain as of end 2015 is general to traverse substantial distances, including others within that it is 1/ A north Queensland Tropidechis and 2/ It is almost the broader so-called “Notechis clade” as outlined by Sanders et certainly NOT from Julatten in Queensland. al. (2008) and the four relevant sources cited therein, of which These facts need to be made known here as the relevant photo Tropidechis is part, the question I was not able to answer was published in Hoser (1989) has also been published on the why these snakes have been apparently unable to breach the internet since and reposted widely. People should be made

Hoser 2016 - Australasian Journal of Herpetology 31:35-38. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 37

aware of the error as they should not get the misguided idea that side, unless any are abnomally fused, this being obvious when it it is a typical example of the newly described species T. occurs by the scale being abnormally elongated (laterally) as jessejacksoni sp. nov. when it is not. compared to the others when viewed laterally. I also note that, notwithstanding the theft of relevant materials Labial number 6 (heading in a posterior direction) in T. from this author in an illegal armed raid on 17 August 2011, jessejacksoni sp. nov. either does not touch the jawline, being which were not returned in breach of undertakings to the court cut off by labials 5 and 7, or if it does, it does so only just and at (Court of Appeal Victoria 2014 and VCAT 2015), I have made a a very narrow point. By contrast in both T. sadlieri and T. decision to publish this paper. carinatus, labial 6 is more-or-less normal in that it is squarish This is even though significant data on specimens of all three and abuts the jawline by a normal wide boundary (as opposed to relevant species gathered over some decades were unlawfully being diamond-shaped in T. jessejacksoni sp. nov.). taken and never returned. Limited numbers of T. carinatus from the Mount Glorious area in This is in view of the conservation significance attached to the South East Queensland, do have labial 6 tending towards being formal recognition of unnamed taxa and on the basis that further diamond-shaped, but in these snakes, the lower boundary sits delays may in fact put the new previously unnamed species at on the jawline by a wide section and so cannot be confused with greater risk of extinction. T. jessejacksoni sp. nov.. This comment is made noting the extensive increase in human The front upper labials, those being between the eye and the population in Australia and the general environmental nasal are relatively short in T. carinatus. In that species, the destruction across the continent as documented by Hoser result is that the suture line running along the top of these (1991), including low density areas without a large permenant scales runs into the lower part of the nasal scale, running more- human population. or-less continuous with the lower line, as opposed to joining I also note the abysmal environmental record of Australian midway between the upper and lower line of the nasal. governments in the past 200 years as detailed by Hoser (1989, In both T. sadlieri and T. jessejacksoni sp. nov. the same upper 1991, 1993 and 1996). labials are higher and as a result the suture line running along TROPIDECHIS JESSEJACKSONI SP. NOV. the top of these scales runs into mid part of the nasal scale, running more-or-less continuous with the middle of the scale Holotype: A preserved specimen in the Queensland Museum, and not with either the upper or lower line. Brisbane, Queensland, Australia, specimen number: J71451, The shape of the preocular in T. carinatus is squarish, versus collected at Picaninny Ck, Windsor Tableland, North irregular in T. sadlieri and T. jessejacksoni sp. nov.. Queensland, Australia, Latitude -16.20, Longitude 144.97. The the Queensland Museum, Brisbane, Queensland, Australia, is a The nasal scale, past the nostril is heavily reduced in T. government-owned facility that allows access to its holdings. jessejacksoni sp. nov. being tiny in that section, versus being of Paratypes: Three preserved specimens in the Queensland similar size, both front and behind the nostril in both T. sadlieri Museum, Brisbane, Queensland, Australia, specimen numbers: and T. carinatus, although slightly smaller posteriorly than J41526 and J52857 collected at Mount Lewis, via Mount Molloy, anteriorly. North Queensland, Australia, Latitude -16.58, Longitude 145.28 The genus Tropidechis is defined as follows: A dangerously and the third preserved specimen collected 16km from Mount venomous genus of snakes (Gow 1983, Trinca, Craydon, Molloy on the Mount Lewis Road, North Queensland, Australia, Covacevich and Limpus 1971). The venom is not only strongly Latitude -16.58, Longitude 145.22. neurotoxic, but it also affects the blood and causes severe muscle damage (Gow 1983). Diagnosis: The three species of Tropidechis Günther, 1863 are It is generally separated from all other Australian elapids by it’s readily separated from one another on the basis of head strongly keeled scales along all or most of it’s body (Cogger scalation. 2000). Unlike Death Adders (Genus Acanthophis) which may In the absence of reliable locality data, the simplest way to sometimes have strongly keeled scales (particularly on the head separate the three species (carinatus Günther, 1863 versus and forebody) this species does not have a tail that terminates in sadlieri Hoser, 2003 and jessejacksoni sp. nov.) is by a cursory a well-defined spine. look at the frontal scale. In T. carinatus this scale is always Attaining an average adult length (total) of between .75 and 1 widest at the front point where it joins the supraoculars (on each metre, more than one death has been attributed to Tropidechis, side) at the point where the front border of the supraocular runs including the case of a 59 year old man dying within 5 minutes towards the sides of the head and borders the prefrontals. For T. after being bitten 3 times on the hand. sadlieri and T. jessejacksoni sp. nov. the frontal shield is widest The natural history of the species is discussed by Beard (1979). where it joins the supraoculars at the point where the Male combat has not been recorded in Tropidechis to date. supraoculars join the parietal shields. Rarely, in T. sadlieri and T. However based on the fact that adults are of similar size and jessejacksoni sp. nov., these two measurements are more-or- that Shine (1991) reported a sample of males being on average less the same, but still the front point is not distinctly wider as in a miniscule amount longer than a similar sample of females, the T. carinatus. possibility of male combat in Tropidechis should not be The best way to see this (in the first instance) is by comparative discounted (Hoser 2003). observation of the relevant head shields of specimens of both species, or by looking at photos of the same. Distribution: The Northern Wet tropics of Queensland, Australia T. carinatus is also separated from T. sadlieri and T. in an area bounded by Julatten/Mount Lewis in the south to at jessejacksoni sp. nov. by the nasal scale’s properties. In T. least the Windsor Tableland/Thornton Peak area in the north. sadlieri and T. jessejacksoni sp. nov. this scale is generally more Tropidechis sadlieri Hoser, 2003 is confined to the Southern Wet circular in form and lacks a distinct bulging back as in T. Tropics of Queensland, Australia in the area from Mount Spec in carinatus. The nasal in T. sadlieri and T. jessejacksoni sp. nov. the south to Bellenden Ker, just south of Cairns in the north and still has a raised surface posterior to the nostril. including the Atherton Tableland. Tropidechis carinatus is found Once again, the best way to see this (in the first instance) is by in a broad zone stretching from the wet northern parts of the comparative observation of the relevant head shields of Sunshine Coast in Queensland, south through the wetter ranges specimens of the three species, or by looking at photos of the and nearby areas to the dry zone of the Hunter Valley in the mid same. north coast of New South Wales and including Barrington Tops north-west of there. All three Tropidechis species can be readily separated from one another by the upper labial scales. Etymology: Named in honour of Jesse Louis Jackson, Sr. a well-known American civil rights activist, Baptist minister, and In all species in the normal situation there are seven on each Hoser 2016 - Australasian Journal of Herpetology 31:35-38. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 38 Australasian Journal of Herpetology politician, in recognition of his lifetime’s struggle for the basic Hoser, R. T. 1996. Smuggled-2: Wildlife Trafficking, Crime and human rights for non white citizens of the United States of Corruption in Australia. Kotabi Publishing, Doncaster, Victoria, America, a battle he has unfortunately not come even close to Australia:280 pp. winning, which is a fact that should make people of all racial Hoser, R. T. 2003. The Rough-scaled Snakes, Genus: backgrounds ashamed. Tropidechis (Serpentes:Elapidae), including the description of a REFERENCES CITED new species from far north Queensland, Australia. Crocodilian - Beard, D. J. 1979. Rough-scaled snake, Tropidechis carinatus. Journal of the Victorian Association of Amateur Herpetologists Herpetofauna 10:26-28. 4(2):11-14. (August 2003). Cogger, H. G. 2000. Reptiles and Amphibians of Australia (sixth Hoser, R. T. 2013. A seven way division of the Amphibolurinae edition), Reed/New Holland, Sydney, Australia:808 pp. (Squamata: Sauria: Agamidae). Australasian Journal of Cogger, H. G. 2014. Reptiles and Amphibians of Australia, 7th Herpetology 21:33-36. ed. CSIRO Publishing, Australia:xxx+1033 pp. Moritz, C., Joseph, L. and Adams, M. 1993. Cryptic diversity in Cogger, H. G., Cameron, E. E. and Cogger, H. M. 1983. an endemic rainforest skink (Gnypetoscincus queenslandiae). Zoological Catalogue of Australia (1): Amphibia and Reptilia. Biodiversity and Conservation 2:412-425. Australian Government Publishing Service, Canberra, ACT, Ogilby, J. D. 1892. Descriptions of three new Australian lizards. Australia:313 pp. Records of the Australian Museum 2:6-11. Couper, P. J., Schneider, C. J., Hoskin, C. J. and Covacevich, J. Ride, W. D. L. (ed.) et al. (on behalf of the International A. 2000. Australian leaf-tailed geckos: phylogeny, a new genus, Commission on Zoological Nomenclature) 1999. International two new species and other new data. Memoirs of the code of Zoological Nomenclature (Fourth edition). The Natural Queensland Museum 45(2):253-265. History Museum - Cromwell Road, London SW7 5BD, UK. Court of Appeal Victoria. 2014. Hoser v Department of Sanders, K, L., Lee, M. S. Y., Leys, R., Foster, R. and Scott Sustainability and Environment [2014] VSCA 206 (5 Sept). Keogh, J. 2008. Molecular phylogeny and divergence dates for de Vis, C. W. 1886. On certain geckos in the Queensland Australasian elapids and sea snakes (hydrophiinae): Evidence Museum. Proceedings of the Linnean Society of New South from seven genes for rapid evolutionary radiations. Journal of Wales (2)1(I):168-170. Evolutionary Biology 21:682-695. Gow, G. F. 1983. Snakes of Australia (revised edition), Angus Trinca, J. C., Craydon, J. J., Covacevich, J. and Limpus, C. and Robertson, Sydney, Australia:118 pp and 48 plates. 1971. ‘The rough-scaled snake (Tropidechis carinatus) a Günther, A. 1864. Third contribution to our knowledge of dangerously venomous Australian Snake’, Medical Journal of batrachians from Australia. Proceedings of the Zoological Australia:801-809. Society of London 1864:46-49 [46, pl. 7 fig. 1]. Uetz, P. 2015. Webpage at: http://reptile-database.reptarium.cz/ species?genus=Tropidechis&species=carinatus downloaded on Günther, A. 1897. Descriptions of new species of lizards and of 12 December 2015. a treefrog from north-eastern Queensland. Novitates Zoologicae, Zoological Museum, Tring, 4:403-406. Victorian Civil and Administrative Tribunal (VCAT). 2015. Hoser v Department of Environment Land Water and Planning (Review Hoser, R. T. 1989. Australian Reptiles and Frogs. Pierson and and Regulation) [2015] VCAT 1147 (30 July 2015, judgement Co., Mosman, NSW, 2088, Australia:238 pp. and transcript). Hoser, R. T. 1991. Endangered Animals of Australia. Pierson Wilson, S. and Swan, G. 2008. A complete guide to reptiles of Publishing, Mosman, NSW, 2088, Australia:240 pp. Australia. New Holland Publishers, Sydney, Australia:512 pp. Hoser, R. T. 1993. Smuggled: The Underground Trade in CONFLICT OF INTEREST Australia’s Wildlife. Apollo Publishing, Moss Vale, NSW, The author has no known relevant conflicts of interest. Australia:160 pp.

Australasian Journal of Herpetology ® Publishes original research in printed form in relation to reptiles, other fauna and related matters, including the subjects of classification, ecology, public interest, captivity, “academic misconduct”, etc. It is a peer reviewed printed journal published in hard copy for permenant public scientific record in accordance with the International Code of Zoological Nomenclature (Ride et al. 1999), with a sizeable print run and has a global audience. Full details of acquiring copies, either printed or in identical form online, editorial policies, publication procedure, author submission guidelines, peer review guidelines, legal matters, advantages of publication in this journal and the like can be found by following links from: http://www.herp.net Published by Kotabi Pty Ltd PO Box 599 ISSN 1836-5698 (Print) Doncaster, Victoria, 3108. ISSN 1836-5779 (Online) Australia. Copyright. All rights reserved. Online journals (this issue) do not appear for a month after the actual and listed publication date of the printed journals. Minimum print run of first printings is always at least fifty hard copies.

Hoser 2016 - Australasian Journal of Herpetology 31:35-38. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 39 Australasian Journal of Herpetology 31:39-61. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online)

A review of the Candoia bibroni species complex (Squamata: Serpentes: Candoiidae: Candoia).

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 12 April 2016, Accepted 20 May 2016, Published 1 August 2016.

ABSTRACT The Pacific Boas, genus Candoia Gray, 1842, have been subject of intense taxonomic scrutiny in recent years. This has included dissections of the three widely recognized putative species. Candoia carinata has been most recently split into three full species and a total of ten regionally distinct subspecies (Smith et al. 2001). C. aspera split into three subspecies (Hoser, 2013) and C.bibroni has long been recognized as consisting of two named subspecies, although one of the used names has been misapplied by various authors and is in fact probably a nomen nudem. Taxonomic treatments of the genus (e.g. McDowell 1979) and molecular treatments of the genus (e.g. Austin 2000) have tended to uphold these divisions and shown clearly that if anything, the taxonomic diversity of the group has been grossly understated. Hoser 2013, utilized these results and formally described Candoia aspera iansimpsoni, this being the most recent addition to the genus. Furthermore, by using existing available nomenclature Hoser (2013) placed each of the three well-known putative species into subgenera. Hoser (2013) also for the first time moved all species into the newly erected family Candoiidae Hoser, 2013 as distinct from the . This paper is a result of drawing on these facts, and available specimens of the putative species C. bibroni, to create a taxonomy that more properly reflects the morphology and phylogeny. The complex has been divided into eleven species and one subspecies; all except C. bibroni named for the first time. The division is along obvious morphological and geographical lines and each population is clearly genetically distinct and isolated. By simple analysis they are all significantly divergent. They are therefore evolving as species in the ordinary sense of the word and should be treated as such. The taxonomy is robust and conservative and in its entirety, in every case reflects recent “splits” of species across the exact same geographical barriers, likely to be affected by the same factors. This forms a necessary first step to preserving potentially threatened populations, which by virtue of their insular nature, must be treated as vulnerable. Furthermore it appears that the factor limiting the westward spread of the C. bibroni complex in the Solomon Islands has been the Brown Tree Snake, Boiga irregularis (Bechstein, 1802). Keywords: Taxonomy; Pacific Boa; snakes; genus; Candoia; bibroni; carinata; aspera; paulsoni; iansimpsoni; superciliosa; schmidti; australis; species; Fiji; Lau Islands; Loyalty Islands; Solomon Islands; Solomons; Rennell, Bellona; San Cristobal; ; Banks Islands; Tokelau; Samoa; Wallis; Futuna; Rotuma; Kandavu; Vanuatu; New Caledonia; new species; hoserae; woolfi; kimmooreae; malcolmmaclurei; boutrosi; Niraikanukiwai; Georgemacintyrei; louisemcgoldrickae; simonmcgoldricki; jamiekonstandinoui; new subspecies; georgekonstandinoui.

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 40 Australasian Journal of Herpetology

INTRODUCTION issues 1-32 of Australasian Journal of Herpetology has been via The Pacific Boas, genus Candoia Gray, 1842, have been subject a combination of the following: of intense taxonomic scrutiny in recent years (e.g. (Smith et al. Genera and component species are audited to see if their 2001, Hoser 2013). classifications are correct on the basis of known type This has included dissections of the three widely recognized specimens, locations and the like when compared with known putative species. Candoia carinata has been most recently split phylogenies and obvious morphological differences between like into three full species and a total of eleven regionally distinct species. subspecies (Smith et al. 2001). C. aspera into three subspecies Original descriptions and contemporary concepts of the species (Hoser, 2013) and C.bibroni has long been recognized as are matched with available specimens from across the ranges of consisting of two named subspecies, although one of the used the species to see if all conform to accepted norms. names has been misapplied and is in fact a nomen nudem. These may include those held in museums, private collections, Taxonomic treatments of the genus (e.g. McDowell 1979) and collected in the field, photographed, posted on the internet or molecular treatments of the genus (e.g. Austin 2000) have held by individuals. tended to uphold these divisions and shown clearly that if This is obviously only done when the location data is good and anything, the taxonomic diversity of the group has been when applicable, other relevant data that is available is also understated as compared to taxonomic treatments of other used. reptiles across the same biogepgraphical realm. Where specimens do not appear to comply with the described Hoser 2013, utilized these results and formally described species (and accepted concept of the species), this non- Candoia aspera iansimpsoni, this being the most recent addition conformation is looked at with a view to ascertaining if it is to the genus. Furthermore, by using existing available worthy of taxonomic recognition or other relevant considerations nomenclature Hoser (2013) placed each of the three well-known on the basis of differences that can be tested for antiquity or putative species into subgenera. deduced from earlier studies. Hoser (2013) also for the first time moved all species into the When this appears to be the case (non-conformation), the newly erected family Candoiidae Hoser, 2013. potential target taxon is inspected as closely as practicable with This paper goes further, by drawing on these facts, and available a view to comparing with the nominate form or forms if other specimens of the putative species C. bibroni, to marry what is similar taxa have been previously named. known of the morphology of local populations with what’s known Other relevant data is also inspected, including any available from relevant molecular studies and filling in the blanks with molecular studies, geological data and the like which may relevant geological evidence in terms of land bridges and the indicate likely divergence of populations. like which could aid shifts in populations and specimens to Where molecular studies are unavailable for the relevant taxon create a taxonomy that more properly reflects the morphology or group, other studies involving species and groups constrained and phylogeny. by the same geographical or geological barriers, or with like The complex has been divided into eleven species; all except for distribution patterns are inspected as they give reasonable C. bibroni are named for the first time. indications of the likely divergences of the taxa being studied They are separated along obvious morphological and herein. geographical lines, the latter shown to be major phylogeographic Additionally other studies involving geological history, sea level barriers for reptiles in other molecular studies such as Austin and habitat changes associated with long-term climate change, (2000), Keogh et al. (2008) and others. including recent ice age changes in sea levels, versus known It is self evident that they are therefore evolving as species in sea depths are utilized to predict past movements of species the ordinary sense of the word and should be treated as such. and genus groups in order to further ascertain likely divergences An additional subspecies is also named. between extant populations (as done in this very paper). The taxonomy presented here is robust and conservative and a When all available information checks out to show taxonomically necessary first step to preserving potentially threatened distinct populations worthy of recognition, they are then populations, which by virtue of their insular nature and ongoing recognized herein according to the rules of the International human pressures on the relevant islands, must be treated as Code of Zoological Nomenclature (Ride et al. 1999). vulnerable. This means that if a name has been properly proposed in the Notwithstanding the theft of relevant materials from this author in past, it is used. This is exactly what happens in this paper for the an illegal armed raid on 17 August 2011 (Court of Appeal taxon originally described as Enygrus bibroni Duméril and Victoria 2014 and VCAT 2015) and not returned in breach of Bibron, 1844. various earlier court orders, I have made a decision to publish Alternatively, if no name is available, one is proposed accoding this paper in view of the conservation significance attached to to the rules of the Code as is done numerous times in this the formal recognition of unnamed species. I also note that paper. further delays may in fact put these otherwise unnamed taxa at As a matter of trite I mention that if a target taxon or group does greater risk of extinction. check out as being “in order” or properly classified, a paper is MATERIALS AND METHODS usually not published unless some other related taxon is named These are not formally explained in a number of my recent for the first time. papers under the heading “Materials and methods” or similar, on The published literature relevant to the taxonomic judgements the basis they are self evident to any vaguely perceptive reader. made within this paper includes papers relevant to Solomon However, the process by which the following taxonomy and Islands and other south-west Pacific Island species affected by nomenclature in this and other recent papers by myself of the same physical barriers to dispersion. They detail phylogentic similar form, has been arrived at, is explained herein for the differences between similar taxa from nearby island groups, benefit of people who have recently published so-called dispersal dates and the like, geological factors and the like and “criticisms” online of some of my recent papers at the non-peer material directly relevant to Candoia. Combined, they include the reviewed website “Facebook” . They have alleged a serious following: “defect” by myself not formally explaining “Materials and Adler, et al. (1995), Austin (2000), Austin et al. (2010), Balsai methods” under such a heading. (1995), Barbour (1921), Bauer (1999), Bauer and Sadlier (2000), The process involved in creating the final product in terms of Bauer and Vindum (1990), Böhme et al. (2002), Boulenger taxonomy and nomenclature for this and other relevant papers in (1884, 1885, 1886, 1890, 1893, 1897), Brongniart (1800), Brown

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 41

(1991), Bruns et al. (1989), Cogger (1972), Colvée and Weffer Hamilton et al. (2010) also provide evidence contrary to the view (2004), Colvée and Martin (2005), Dahl (1986), Daza et al. that Candoia bibroni sensu lato originated from the Fiji region, (2015), de Rooij (1917), Duméril and Bibron (1839, 1844), dispersing outwards in most directions, indicating that the Duméril and Duméril (1851), Forcart (1951), Garman (1901), Gill founder stock may have originated in the Solomon Islands and (1993, 1995), Gray (1856), Greer (1982), Greer and Parker moved south-east over a substantial time frame by vicariance as (1967), Greer and Simon (1982), Hagen et al. (2012), Hall much as dispersal. (2002), Hamilton et al. (2010), Harvey et al. (2000), Higgins Sea currents and land mass evidence from glacial maxima also (1943), Hoser (2013), Ineich (2008, 2009, 2011), Iskandar and indicate a dispersal eastwards from the Solomons and then Erdelen (2006), Jacquinot and Guichenot (1853), Keith and south-west from the general vicinity of Samoa, Wallis and Marquet (2006, 2007), Keith et al. (2008, 2011), Keogh et al. Futuna towards Fiji and beyond. (2003, 2008), Kinghorn (1928, 1937), Koch et al. (2009), Mayr The name Candoia australis (Montrouzier, 1860), treated by (1931), McCoy (1980, 2006), McDiarmid et al. (1999), McDowell many authors as either a synonym for C. bibroni, (originally (1970, 1979), Medway (1974), Mertens (1928, 1931), named Boa australis Montrouzier, 1860) or a subspecies, is not Montrouzier (1860), Morrison (2003), Mys (1988), Ogilby (1890), available for any species or subspecies described herein. Oxley (2016), Ota et al. (1998), Parker (2012), Pianka and Vitt McDowell (1979) wrote of this alleged taxon: “Montrousier (2003), Pyron et al. (2013), Rahmstorf (2003), Reeder (2003), reported his Boa australis to come from New Caledonia, where Richmond et al. (2014), Rittmeyer and Austin (2015), Robinson there are no known terrestrial snakes and his description seems (1974), Roemmich (2007), Roux (1913), Russell and Coupe quite unidentifiable.” (1984), Sadlier and Bauer (1997), Schmidt (2010), Schmidt (1932), Schmidt and Burt (1930), Schweizer (1970), Sibley and I note also that the application of the name to the Solomon Monroe (1990), Wells and Wellington (1985), Williams and Islands population by authors in the past century following on Parker (1964), Zug (2012a, 2012, 2013) and sources cited from an error by Boulenger also has no tenable basis of fact or therein. availability under the rules of the various editions of Codes of Zoological Nomenclature (e.g. Ride et al. 1999). These papers combined with the findings of McDowell (1979) in terms of morphological divergences in populations, make the In terms of this nomenclatural problem, McDowell (1979) wrote: argument in favour of splitting the main populations within the C. “Just how Boulenger attached Montrousier’s name, based on a bibroni sensu lato group into full species as opposed to mere color description without scale counts and without known subspecies as would otherwise conservatively be the case (see preserved- types, to the form with high ventral count has never also below). been explained, for Montrousier reported his Boa australis to come from New Caledonia, where there are no known terrestrial Some material within descriptions below is repeated for different snakes and his description seems quite unidentifiable.” described taxa and this is in accordance with the provisions of the International Code of Zoological Nomenclature and the legal In other words Boa australis Montrouzier, 1860 is at this point in requirements for each description. I make no apologies for this. time clearly a nomen nudem. RESULTS In the alternative, if Boa australis Montrouzier, 1860 is not a nomen nudem, as first revisor, I hereby restrict the taxon to the A number of my recent taxonomic papers do not list any results type locality, New Caledonia, where no specimens occur. as such under any such heading. Also of note is the ultra conservative treatment of the various This is because the descriptions that follow the introduction are described taxa. in fact the results and the outcome arising from these. While two species groups are identified within the C. bibroni The relevant evidence is the animals themselves and from a complex herein (as identified by McDowell 1979 and Zug 2013), taxonomic standpoint, the relevant morphological features of I have taken the view that each of the divergent forms within them. each group should be treated as full species as done below. Evidence in the form of molecular evidence and geological The two main groups of species are what I call the C. bibroni history can only corroborate the evidence of the animals group from the east and north-east of the range, and the C. themselves. hoserae sp. nov. group from the north-west of the range. Part of the process of identifying and describing the relevant Diagnostic of the C. hoserae sp. nov. group of species is that species or subspecies in this and other papers involves a review the C. hoserae sp. nov. group is separated from the C. bibroni of the literature and conclusions drawn by these authors. group, by the form of the postorbital bone, which has its ventral I do not necessarily agree with all conclusions of authors in the (free) end abruptly flexed backward, as in C. carinata, rather cited literature and this is usually self-evident when my than forming a simple crescent arc as seen in C. bibroni. In the taxonomic judgements are cross-referenced with the cited C. bibroni group of species the premaxilla has the “typical boine” papers and the like. form described by Frazzetta (1959, 1975), with a large fenestra Some authors have clearly made similar taxonomic judgements between the narial vestibules, this fenestra bounded anteriorly to myself and others have not. by the premaxilla and posteriorly by the nasals; but C. hoserae In the case of my review of the available data, I do herein note a sp. nov. are like C. carinata (with which they are sometimes general concurrence in my conclusions from those inferred by sympatric) in having the intervestibular fenestra largely or McDowell (1979) and Austin (2000) in terms of the species entirely filled in by expansion of the premaxilla. Candoia bibroni (Duméril and Bibron, 1844) in that there should Of note also is that a reassessment of published records and be at least two species recognized (see also Zug 2013), one museum specimens indicates that the C. bibroni complex is not being primarily from the west of the range and the other from the as widespread as reported in a lot of the literature. east. There is currently no evidence the group occurs on any of The depth of the divergences is also corroborated by the Tokelau, Tonga, the lower New Hebrides or New Caledonia. findings of Hamilton et al. (2010), who significantly left Candoia These islands are also biogeographically separated from the out of their results, because they presumably had the belief that islands where the complex is known to occur. specimens on either side of their barrier (the so-called Cheesman’s line) were of the same species. AN EXPLANATION OF THE PRESENT DAY DISTRIBUTION OF THE CANDOIA BIBRONI SPECIES COMPLEX. It is in the light of the combined evidence available here, Previous authors such as McDowell, 1979 appeared to have previously unavailable to previous authors on which the current difficulty explaining the extant distribution of the complex. taxonomic and nomenclatural judgements published herein are made. Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 42 Australasian Journal of Herpetology

McDowell wrote: prevented rafting C. bibroni complex snakes from establishing. “the distribution of C. bibroni appears to centre around the Both species share the same ecological niche and nocturnal Pandora (or North Fiji) Basin, with a northeastward extension to habits and would clearly compete directly for food. B. irregularis Samoa, a southwestward extension to the Loyalty Islands, and would have a direct competitive advantage in that their northwestward extensions to Rennell and San Cristobal. cannibalistic and reptile eating habits are well known as is their However, it appears impossible to fit the distribution of the ability to live in high density and so overwhelm less numerous species to any known feature of submarine topography. Thus, competing species. the New Hebrides Trench has not prevented C. bibroni from Based on available evidence of distribution, these factors imply reaching the Loyalty Islands, yet the species does not extend to an overwhelming advantage over slower moving more heavy New Caledonia (not separated from the Loyalty Islands by any bodied snakes in the same ecospace. With sea currents in the submarine barrier). C. bibroni reaches San Cristobal at the Solomons running east to west as part of the well-established eastern end of the Solomons Ridge, but not the adjacent “Indonesian throughflow”, cross island colonisation of C. bibroni Guadalcanal or Malaita, on the same ridge; C. bibroni also complex snakes through the Solomon Islands should have been reaches , on the eastern end of the Rennell Ridge a mere formality, noting the cross sea distances to be rafted (essentially continuous with the Louisiade Rise) but does not were insignificant compared with those traversed in the south extend to the Louisiade Archipelago. In extending to Samoa, C. Pacific. The same applies for the islands of the Louisiade bibroni crosses the Tonga Trench and leaves the Melanesian Archipelago. Plateau.” This means that in the absence of some particularly effective To this extent, McDowell has identified the known range of the predator on the other islands, the C. bibroni complex snakes complex, but without identifying the means of dispersal. would have become established throughout the relevant A check of the geological history of the relevant island groups archipelagos. indicates at least 11 populations that have never been in land The only predatory species in the same ecospace which contact with one another meaning that the snakes must have appears to have a distribution that in total excludes C. bibroni either swam, or more likely “rafted” between the relevant land sensu lato is B. irregularis. masses. Hoser (1995), detailed how B. irregularis effectively caused the This is relatively simple over a geological time frame measured elimination of Hoplocephalus bungaroides (Schlegel, 1837) from in the millions of years when one looks at the prevailing sea areas north of Sydney harbour. The two species occupied currents in the region. The entire distribution of the C. bibroni identical habitats where they occur in the Sydney basin, but by complex clearly corresponds to the flow of the Equatorial being mutually exclusive. Countercurrent and the South Pacific Gyre as detailed by With B. irregularis the more recent immigrant to the region, we Rahmstorf (2003), Roemmich (2007) and D’Hondt et al. (2009). know emphatically that these snakes have successfully In fact it doesn’t seem necessary to identify where the snakes eliminated H. bungaroides from sandstone areas in Sydney’s originated, as the South Pacific Gyre would clearly transport north. rafting snakes to all known localities. The molecular evidence of Richmond et al. (2014) confirmed Having said this, based on the morphological evidence of that B. irregularis sensu lato migrated from west to east in the McDowell and the distributional evidence for Candoia as a recent geological past, this migration encompassing a region whole, centred on the islands north of New Guinea, including generally bound by Sulawesi in the West and Malaita and those accreted to it in the last few million years, it would appear Guadalcanal in the east of the Solomon Islands. Notably, it is that the Candoia bibroni group had its origins in the Solomon the next major island to the south-east, namely (AKA Islands rather than Fiji. San Cristobal) where Candoia bibroni complex makes its last Alternatively, if Fiji is the point of origin for the three species stand. In this general vicinity on other islets, uninhabited by B. groups, then Fijian animals most likely founded all the other irregularis, and islands to the east and south, also uninhabited putative species, including the C. hoserae sp. nov. group within by B. irregularis, C. bibroni complex snakes occur and in C. bibroni sensu lato. abundance. The molecular evidence of Austin (2000) also demonstrates that The division of the Solomon Islands into a western and central the relevant snakes had clearly been able to raft across sector with B. irregularis and a mutually exclusive eastern zone significant bodies of water in order to colonise the relevant island with C. bibroni complex snakes is seen in the distribution groups some millions of years ago, from where the populations information for B. irregularis and “C. bibroni” in McCoy 2006. have evolved more-or-less independently ever since. There he lists B. irregularis as being found in the following places: “Maluku (Moluccas), Sulawesi, New Guinea and The southern migration of the C. bibroni complex was almost adjacent archipelagos, Australia, Solomon Islands. There is also certainly limited by temperature, in that the climate further south an introduced population on Guam which has exterminated a was simply unsuitable for these particular snakes. number of species of native on that island. In the Hence these snakes are not found in places like New Zealand or Solomons: Bougainville, Shortland, Mono, Choiseul, , Norfolk Island. , New Georgia, Tetepare, , Isabel, The factor keeping the snakes from colonising New Caledonia is Guadalcanal, Ngela, Malaita.” not emphatically known, although it seems to apply to other For the species “C. bibroni” he lists the mutally exclusive species groups from the same distributional realm as C. bibroni distribution of “Solomon Islands, Vanuatu, Loyalty Islands, Fiji sensu lato, implying the local sea currents may exclude passge Islands, Tokelau Islands, Wallis and Futuna, the Samoas, to the island, even though the nearby Loyalty Islands group were Tonga. In the Solomons: Rennell, Makira, Ugi, Olu Malau, Santa reached. Ana, Santa Cruz, Reef Islands, , .” However the factor keeping the Candoia bibroni complex from No one, including McCoy, have until now picked up on this colonising any parts of tropical or eastern Australia, the western mutually exclusive distribution by two species that share a near parts of the Solomon Islands and the Louisiade Archipelago is in identical ecological niche. my view clearly obvious. That this has been overlooked until now is somewhat surprising, These areas are inhabited with a highly invasive, nocturnal especially considering the effect B. irregularis has had on Guam, predatorial tree snake in the form of the Brown Tree Snake where it was introduced inadvertently and has caused problems Boiga irregularis (Bechstein, 1802). ever since as noted by Rodda and Fritts (1992), Rodda and These snakes, which live in high densities, clearly must have Savidge (2010) and Rodda et al. (1999).

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Likewise in terms of the Louisiade Archipelago, where McDowell preponderance of females over males occurring in the new noted the absence of “C. bibroni”. de Rooij (1917), lists B. Georgia Group of the Solomons, where C. bibroni does not irregularis as occurring in the Louisiade Archipelago and other occur. nearby islands off the south-east coast of New Guinea, all of If one can judge from the collections of the Whitney South Sea which conspicuously lack “C. bibroni”. Expedition, the presence of one species of Candoia would seem McDowell (1979), spent considerable effort looking in the to have no depressing effect on the abundance of the other. The direction of another species of snake affecting the distributional largest collection of C. bibroni taken by that expedition was that limits of “C. bibroni” in the Solomon Islands and to this effect he from San Cristobal, where 18 were taken; the same expedition spent his efforts looking at the congeneric C. carinata took 19 C. carinata on San Cristobal, the largest collection of (Schneider, 1801) and not surprisingly he drew a blank in terms that species from a single island made by the Whitney South of any hard evidence implicating that species. Sea Expedition.” It is prudent for me to copy a sizeable chunk of his paper here to I also note here that McDowell also found that the largest see the basis of his analysis and why he drew a blank in the end numbers of both “C. bibroni” and C. carinata were found on San in terms of implicating the species C. carinata restricting the Cristobal which happens to be the only major island in the range or habits of “C. bibroni”. Solomons without Boiga irregularis, indicating that the tree McDowell (1979) wrote: snake species also has a depressing effect in numbers of C. “Over much of its range, C. bibroni is either the only snake in the carinata as well as “C. bibroni”. land fauna (Banks Islands, New Hebrides, Samoa, and probably As to why C. carinata sensu lato and the third species complex much of eastern Fiji) or is accompanied by a single form within Candoia, namely Candoia aspera (Günther, 1877) could obviously adapted to feeding on very small prey (Loyalty Islands, survive in areas inhabited by Boiga irregularis, including all where Typhlina willeyi occurs; larger Fiji Islands, where those Solomon Islands inhabited by B. irregularis for C. carinata Ogmodon vitianus occurs). It should be noted that of the four sensu lato (usually referred to as C. paulsoni (Stull, 1956) for cases of murid remains in the stomach of my material of C. Solomons specimens), the answer is self-evident. bibroni, two were from the New Hebrides (AMNH 81597, Aoba I.; Unlike C. bibroni sensu lato, the other two species groups are 81589, Maewo I.) and the others from Fiji (AMNH 40439, Viwa I.; relatively stout ground-dwelling snakes, not being obligatory tree 40451, Vomo I.) It seems possible, therefore, that C. bibroni dwellers, acting significantly outside the competitive niche of B. becomes more terrestrial in its foraging in regions where C. irregularis. carinata does not occur. As to why Boiga irregularis is not found on San Cristobal and It must be emphasised that my data are far too meagre to islands east and south of there, the reason is simple and again establish a character displacement in behavior, and in relates to the prevailing sea currents. morphology, where my data are more extensive, there is no The prevailing current in the region is the well-established evidence of a “character displacement” in either C. bibroni or C. “Indonesian throughflow”, which flows east to west and thereby carinata, at least no evidence that would fulfill the rigorous forms a so-called headwind against eastward migration of B. requirements of Grant (1972). It is true that western (“Long- irregularis by any form of rafting. tailed”) C. carinata are more like C. bibroni than are eastern All the islands colonised by B. irregularis in the Solomons fit one (“short-tailed”) C. carinata, particularly in such features as wide of the following profiles: separation of the postorbital from the frontal bone, high 1/ The islands were joined in ice-age minima into a single land subcaudal count, somewhat lower tooth counts (than in eastern bridge and so were easily crossed by the invasive snakes, which C. carinata, but still above those of C. bibroni), and frequent effectively traversed land bridges and/or proximal islands from occurrence of a striped color phase suggestive of some phases greater New Guinea, these land bridges detailed by Hagen et al. of C. bibroni. However, the geographic range of the eastern form (2012) and others, including a direct line of connection from of C. carinata far exceeds the overlap of that form with C. bibroni Bougainville to Ngela via joined islands in between. I note here (an overlap confined, so far as known, to San Cristobal that Ngela is directly proximal to the permanently separated and nearby Ugi, Bio, Santa Ana, and Three Sisters Islands, to Guadalcanal, the two islands split by a very narrow strait. the , and Rennell and Bellona). I can see no 2/ The other islands colonised, (e.g. the New Georgia group), indication, for example, that C. carinata from Guadalcanal, while never connected by land to the other Solomon Islands where C. bibroni does not occur, are any less-or any more- invaded by B. irregularis, effectively sat downcurrent from more different from C. bibroni than are C. carinata from San Cristobal. easterly islands that could be colonised via the above-referred to So far as variation in C. bibroni is concerned, there is nothing to land bridge/s (e.g. Ngela/Guadalcanal) via rafting. indicate that the populations sympatric with C. carinata are The significance of the exclusionary competitiveness of a morphologically more different from that species than are Colubrid (B. irregularis) to a Candoiid (C. bibroni sensu lato) populations from islands where C. carinata does not occur; if cannot be understated in terms of the study of modern anything, there may be some character convergence. Thus, in ecosystems and I suspect it is far more common than has been Fiji and New Hebrides C. bibroni the premaxilla has the “typical previously suspected across other regions. boine” form of Frazzetta (1959; 1975), with a large fenestra between the narial vestibules, this fenestra bounded anteriorly Hoser (2014) found the species Acanthophis pyrrhus by the premaxilla and posteriorly by the nasals; but eastern (Boulenger, 1898), to successfully exlude and/or eliminate Solomons C. bibroni are like C. carinata (with which they are Pilbara Death Adders Acanthophis wellsei Hoser, 1998 from sympatric) in having the intervestibular fenestra largely or areas they may otherwise cohabit. entirely filled in by expansion of the premaxilla. Again, the Hoser (2014) also confirmed that long term separation of A. highest tooth counts in C. bibroni are in specimens from the wellsei caused by intervening populations of A. pyrrhus also eastern Solomons and the small size of the vomerine foramen of caused a speciation process to occur in the former, as each eastern Solomons C. bibroni is a departure from Fiji and New population was isolated from one another and currently treated Hebrides specimens and a resemblance to the majority of C. as allopatric subspecies. carinata skulls. It may be added that the material available to me The situation of competitive species forming effective barriers to shows no significant (at 5 percent level) departure of C. bibroni gene flows in other taxon populations to allow speciation to from a 1:1 sex ratio or any significant differences in sex ratio occur, appears to have been largely unexplored in herpetology from island to island. It is noteworthy that on San Cristobal (and as a means to find new species. If pursued as a line of enquiry, also Guadalcanal) C. carinata shows at least a close there is no doubt that more cryptic species of reptile will be approximation to a 1:1 sex ratio, with the greatest found.

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THE FORMATION OF THE TAXONOMIC JUDGEMENTS ascertained by way of sea depth records and passages between WITHIN THIS PAPER island groups in excess of 120 metres. As noted by McDowell (1979) and further ascertained by myself This includes those populations identified herein as species from since, putative C. bibroni are morphologically distinct within the closely proximate locations such as the three from the Fiji various island groups they are found. Islands (excluding the distant Rotuma) and the three from the What McDowell (1979) and it seems all authors since him have eastern Solomon Islands. failed to do, is to try to assess these differences with a view to Furthermore as seen in the material cited earlier, each and every establishing a taxonomy for the complex. Or if changes in island group contains endemic species from genera that have taxonomy are indicated by the evidence, the authors have failed crossed the same ocean barriers and yet subsequently to take the logical next step. speciated in the relevant island groups. Zug (2013) used McDowell’s results to state “Subtle skeletal Hence my view that the putative Candoia bibroni from each differences hint that C. bibroni might consist of two species, a island group should also be treated as full species, rather than central Pacific one and a Vanuatu-Solomon Islands one”. subspecies. Austin (2000) wrote: “Both samples of C. bibroni in this study (n Evidence in the literature cited also suggests that the dispersal _ 3) come from the eastern portion of the range (Fiji and Samoa) for Candoia bibroni sensu lato across the existing range was and thus presumably represent only one form of McDowell’s rapid and in the order of 2-5 MYA, which again puts each postorbital bone division. Nevertheless, these samples show isolated population at the species level. over 6% sequence divergence. Sampling populations from Examples for each island group’s biogeographic isolation in the western portion of the range in the future may show levels of addition to isolation by sea-water barriers are given as follows: divergence indicative of specific differentiation within C. bibroni.” 1-3/ The distributions of the three species from Fiji (listed as 1-3 Of course, many species have been divided on far less than a above) effectively mirror in their entirety the co-evolving Iguana 6% sequence divergence (e.g. Scrub Python (Australiasis Wells species Brachylophus in their distributions, as outlined by Keogh and Wellington, 1985) species as defined by Harvey et al. 2000) et al. (2008). and so the case is already made by Austin (2000) for the Brachylophus fasciatus (Brongniart, 1800) appears to be Samoan snakes to be split off as a full species. endemic to the Lau group of islands. For the record, Harvey et al. erected the species Australiasis Robinson (1971) also confirms that the relevant island groups nauta (Harvey et al., 2000) on the basis of a maximum 1.4% have remained biogeographically separated for a significant divergence from nominate A. amethistina (Schneider, 1801) and period. other species were named on similarly low divergences. The Kadavu Rhipidura personata Ramsay, 1876 is a With the Solomons Islands C. bibroni complex snakes clearly species of in the fantail family . It is endemic to more divergent from the Fiji types, they too need to be redefined Kadavu and Ono in the Kadavu Islands, Fiji. On Vita Levu, Fiji it as a separate species. is replaced by the Rhipidura verreauxi Marie, However across the range of putative C. bibroni there is nothing 1870. The two similar taxa have speciated in nearby islands to suggest that either the Samoan, Solomons Islands or Fiji due to the same factors isolating the Candoia. (main island/Vita Levu) are any more divergent than those 4/ Endemisim in Pacific genera in the Loyalty Islands is seen in different forms from other island groups such as Wallis and the species Emoia loyaltiensis (Roux, 1913), including as Futuna, Rotuma, Loyalty Islands or Vanuatu, meaning that these referred to by Zug (2012). groups were also looked at to decide whether or not they should 5/ There are numerous endemic reptile species from the New be given taxonomic recognition and at what level. Hebrides, including the northern sector. Species include the McDowell (1979) when looking at morphological evidence, Toupeed Skink Emoia sanfordi Schmidt and Burt (1930), the stated that it implied parallel evolution of each population over Vanuatu Coppery Vine Skink Emoia nigromarginata (Roux, the recent geological past, adding to the case that all significant 1913), the Anatom Tree Skink Emoia aneityumensis Medway, island group populations should be equally recognized at the 1974, the Erronan Tree Skink Emoia erronan Brown, 1991, the same taxonomic level (i.e. as species or subspecies). Vanuatu Snake-eyed Skink Cryptoblepharus novohebridicus There appeared to be at least eleven main geographically Mertens, 1928, the Vanuatu Saw-tailed Gecko Perochirus isolated populations of putatuive C. bibroni which based on guentheri Boulenger, 1885, the Vanuatu Forest Gecko consistent criteria should be treated as full species in view of Lepidodactylus vanuatuensis Ota et al. 1998 and the Vanuatu both known divergences based on molecular and geological Ant-nest Gecko Lepidodactylus buleli Ineich, 2008. evidence and the accepted Darwinian concept of a species. Putative C. bibroni from the Banks Islands differ from those In no particular order, they are further south in the New Hebrides and are separated by deep water. But in the absence of a well-defined barrier, they are 1/ Nominate form from Fiji (main island/Vita Levu) and the only herein treated as a subspecies of the Vanuatu form. form with an available name, as in C. bibroni. 2/ Lau Group of islands, immediately east of Vita Levu in Fiji. 6-8/ The biogeographical barriers within the Solomon Islands in terms of terrestrial reptiles are well-known. The species Corucia 3/ Kandavu, immediately south of Vita Levu in Fiji. zebrata occurs in the San Christobal (AKA Makira) island group, 4/ Loyalty Islands, east of New Caledonia. but not in the Santa Cruz islands, east of San Christobal or the 5/ Vanuatu, and the Banks Islands. Rennell and Bellona islands, south of San Christobal (Hagen et 6/ Santa Cruz islands, east of the Solomon Islands. al. 2012), meaning each of these populations of putative C. bibroni are isolated from one another. 7/ Rennell and Bellona, south of the Solomon Islands. Furthermore the distribution of Euprepiosaurus juxtindicus 8/ San Christobal (AKA Makira) and small islands immediately (Böhme et al., 2002) is restricted to Rennell and Bellona islands north in the Solomon Islands. and nowhere else in the Solomon Islands (or anywhere else for 9/ Rotmuma Island. that matter), confirming the isolation of terrestrial reptiles here. 10/ Wallis and Futuna. 9/ Rotuma Island and its physical and biological isolation is 11/ Samoa. confirmed by unique species such as the Rotuma Forest Gecko All are clearly isolated island groups. Lepidodactylus gardineri Boulenger, 1897. Significantly, all were also isolated in the same way as at 10/ Futuna Island and its physical and biological isolation in present during Pleistocene Ice-Age sea level minima, as terms of terrestrial or non-marine vertebrates is confirmed by

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unique species such as the freshwater fish Stiphodon In C. bibroni the premaxilla has the “typical boine” form rubromaculatus Keith and Marquet, 2007, Akihito futuna Keith et described by Frazzetta (1959, 1975), with a large fenestra al., 2008, Sicyopus sasali Keith et al., 2011 and Stenogobius between the narial vestibules, this fenestra bounded anteriorly keletaona Keith and Marquet, 2006. by the premaxilla and posteriorly by the nasals; but C. hoserae 11/ Samoa and its physical and biological isolation in terms of sp. nov. and others in the species group are like C. carinata terrestrial or non-marine vertebrates is confirmed by unique (with which they are sometimes sympatric) in having the species such as Emoia samoensis (Duméril and Duméril, 1851). intervestibular fenestra largely or entirely filled in by expansion of GENUS CANDOIA GRAY, 1842. the premaxilla. Type species: Boa carinata Schneider, 1801. Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as defined in this paper, is the taxon from the main Fiji islands Diagnosis: All Candoia Gray, 1842 are separated from all other including Viti Levu and Vanua Levu and immediately adjacent Boas and Pythons and physically similar non-venomous islands, but not including Kadavu or the islands in the Lau constricting snakes with thick, muscular bodies, by the following group, each of which has a different species. suite of characters: The taxon C. bibroni is separated from all other species within Head distinct from the neck, Snout is obliquely truncate. The the C. bibroni species complex by the following suite of canthus rostralis is prominent. Body is slightly compressed. All characters: A blotched dorsal pattern with distinctive throat the dorsal scales are strongly keeled. Subcaudals are all single. stripes and a low ventral count ranging from 207-227 ventrals. There is a pair of longitudinal folds on each branch of hemipenis The species C. bibroni is further defined by the following suite of dividing each branch into a sulcal and an absulcal field; nasal characters: 18-22 maxillary teeth, (versus 17-19 in C. woolfi sp. bones overlapping dorsal surface of frontal bones, thus nov. from the Lau Group of islands), 53-59 subcaudals (versus separating prefrontals on the midline. 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- Distribution: The Pacific Islands from the Samoas in the east to 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the the islands north of New Guinea, including New Guinea and head at one head length behind the head (versus 33-34 in C. across to Halmahera. Not in Australia, New Caledonia, Tokelau, woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one Tonga or the lower New Hebrides. headlength anterior to the cloaca, hemipenis forks at subcaudal CANDOIA BIBRONI (HOMBRON AND JACQUINOT, 1853). 6-10. Type locality: Vita Levu, Fiji. The species Candoia woolfi sp. nov. from the Lau Group of Diagnosis: Candoia bibroni and other species described in this islands, east of the main Fiji islands is separated from all other paper that were formerly treated as populations of C. bibroni are species within the C. bibroni species complex by the following herein defined as being the C. bibroni species complex. suite of characters: A blotched pattern without stripes. The tail pattern is much more conspicuous than that of body, the pale The C. bibroni species group are those species not included in bordering of body blotches also replacing darker ground color on the C. hoserae sp. nov. species group, as defined below. the tail; 205-236 ventrals; there are 33-34 scale rows behind the All species within the C. bibroni species complex are separated head at one head length behind the head (versus 26-30 in C. from all other Candoia by having supralabials separated from bibroni). the eye by the subocular scales, versus C. carinata (Schneider, The species Candoia kimmooreae sp. nov. from the island of 1801) and C. paulsoni (Stull, 1956) which have them entering Kadavu, southern Fiji is is separated from all other species the eye; preocular differentiated; canthus rostralis rounded; a within the C. bibroni species complex by having a blotched relatively slender body with lengthwise ridges on the body scales pattern which includes a prominent median throat stripe and no that run parallel to the long axis of the body, versus a stout body lateral throat stripes. Otherwise this taxon is essentially similar with keels on the body scales forming oblique rows as in C. in form to nominate C. bibroni. aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. Furthermore the tail is more than twice as long as the head, The species Candoia malcolmmaclurei sp. nov. from the Loyalty being capable of more than one complete circle of coiling in the Islands, (territory controlled by New Caledonia) is separated vertical plane. from all other species within the C. bibroni species complex by having a blotched pattern without stripes and a tail similar in The species Candoia carinata (Schneider, 1801) including pattern to the body, or alternatively a blotched pattern with throat species and subspecies formerly treated as being within this stripes including a median row of dark spots on anterior ventrals; taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. dark perocular streak from angle of mouth (or angle of jaw) bibroni complex by having 2, or rarely 1 or 3 supralabials through eye to loreal region, dark spots on lips, and dark entering eye; an angulate canthus rostralis, 160-202 ventrals spotting on top of head and with a median ventral row of dark and 35-60 subcaudals. spots. Sometimes the colouration is dulled to appear unicolour. The species Candoia aspera (Günther, 1877) including species The species is further defined by having 13-16 infralabials, 27- and subspecies formerly treated as being within this taxon is 30 scale rows on the neck at one head length back, 20-22 scale separated from the C. bibroni complex by having a tail less than rows at one headlength anterior to the cloaca, 219-226 ventrals, twice as long as the head, usually being shorter than the head; 53-60 subcaudals and 19-22 maxillary teeth on either side. is incapable of forming a full circle of coiling in vertical plane; at The species Candoia boutrosi sp. nov. from the islands of least on the anterior two-thirds of body are keels of scales Samoa is separated from all other species within the C. bibroni forming curved diagonal ridges along the sides, which extend species complex by the following unique suite of characters: a backward and downward towards the belly; 127-153 ventrals; 11- distinctive blotched dorsal body pattern that is one or other of 22 subcaudals; no specially enlarged preocular; (supralabials being distinct or very indistinct, the large dark blotches being the excluded from the eye; canthus rostralis angular). main component of the dorsal midline for the anterior and mid- Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. body, with the lighter blotches dominating the midline posteriorly, species group are defined as the three species from the reversing on the tail to have the darker blotches dominate on the Solomon Islands, including outliers, as well as the single tail, which remains brightly marked for the whole length. The species from the Banks Islands and nearby New Hebrides. dorsal colouration is a combination of chocolate brown and These species are all separated from C. bibroni and others in beige in distinctly marked animals or a hybrid of this in the the C. bibroni species group (being all others in the C. bibroni indistinctly marked animals. On the head is a well-defined complex) by the form of the postorbital bone, which has its creamish bar on the upper labials above the jawline. ventral (free) end abruptly flexed backward, as in C. carinata, It is also defined as having 12 supralabials bilaterally (in rather than forming a simple crescent arc as seen in C. bibroni. Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 46 Australasian Journal of Herpetology common with the three species from the Fiji Islands), 15-16 Solomon Islands is separated from all other species within the infralabials (as for C. bibroni sensu stricto), and 31-32 scale C. hoserae sp. nov. species complex by the following unique rows on the neck, one head length back from the head (versus suite of characters: Similar in most respects to C. hoserae sp. 26-30 in C. bibroni sensu stricto). nov. and C. louisemcgoldrickae sp. nov from which it can be C. boutrosi sp. nov. is further defined and separated from the separated from it by a profusion of black pigment on the dorsal other species in the C. bibroni complex by the following suite of snout tip and including the rostral and all of first labial on either characters 37-38 mid-body rows, 22-24 scale rows at one head side. There is a particularly thick dark postorbital stripe (as length anterior to the cloaca, 237-252 ventrals, 51-61 opposed to medium in C. hoserae sp. nov. and C. subcaudals and 16-18 maxillary teeth on either side. louisemcgoldrickae sp. nov.). 38-41 midbody scale rows in C. The species Candoia niraikanukiwai sp. nov. from the islands of simonmcgoldricki sp. nov. versus 36-37 in C. hoserae sp. nov. Wallis and Futuna, including nearby islets is separated from all and C. louisemcgoldrickae sp. nov.. other species within the C. bibroni species complex by the The species Candoia simonmcgoldricki sp. nov. is further following unique suite of characters: It is essentially similar in diagnosed and defined by the following: 15-21 infralabials, 250- most respects to C. boutrosi sp. nov. from which it differs by 266 ventrals versus 235-257 for C. hoserae sp. nov. and C. having a distinctive well-defined stripe running through the eye louisemcgoldrickae sp. nov., 61-64 subcaudals versus 54-63 on either side of the head and the absence of a well-defined subcaudals for C. hoserae sp. nov. and Candoia creamish bar on the upper labials above the jawline. louisemcgoldrickae sp. nov., 18-22 maxillary teeth, hemipenis The species Candoia georgemacintyrei sp. nov. from the island fully everted extending from 9-20 subcaudals. of Rotuma, is separated from all other species within the C. The species Candoia jamiekonstandinoui sp. nov. from Vanuatu bibroni species complex by the following unique suite of and the nearby Banks Islands, including nearby islets is characters: 235-240 ventrals, 57-63 subcaudals, 37-39 mid separated from all other species within the C. hoserae sp. nov. body rows and significant lightening of the upper labials; the species group by the following unique suite of characters: A dorsal pattern is blotched and semi-distinct and there is distinctive dorsal colouration of pale reddish brown above with significant speckling and markings on the venter. semidistinct markings, with a dark brown vertebral stripe edged with yellow and a uniform yellow beneath. Males are unique in As mentioned already, Candoia hoserae sp. nov. and others in the C. bibroni species complex in that they have some the C. hoserae sp. nov. species group are defined as the three noticeable dark pigmentation on the hemipenes. 16-18 maxillary species from the Solomon Islands, including outliers, as well as teeth for the nominate subspecies, versus 19 in C. the single species from the Banks Islands and nearby New jamiekonstandinoui georgekonstandinoui subsp. nov.. Hebrides. These are C. hoserae sp. nov., C. louisemcgoldrickae sp. nov., The subspecies C. jamiekonstandinoui jamiekonstandinoui C. simonmcgoldricki sp. nov. and C. jamiekonstandinoui sp. subsp. nov. has 234-247 ventrals, versus 222-225 ventrals in C. nov., the latter of which includes the subspecies C. jamiekonstandinoui georgekonstandinoui subsp. nov.. jamiekonstandinoui georgekonstandinoui subsp. nov.. C. jamiekonstandinoui georgekonstandinoui subsp. nov. has the lowest ventral counts of any named taxon within the C. hoserae These species are all separated from C. bibroni and others in sp. nov. species group. the C. bibroni species group (being all others in the C. bibroni Distribution: The Candoia bibroni species group is found on the complex) as detailed in the precedning text, by the form of the Fiji Islands, Loyalty Islands (but not New Caledonia), Wallis and postorbital bone, which has its ventral (free) end abruptly flexed Futuna, Somoa Islands and Rotuma. backward, as in C. carinata, rather than forming a simple crescent arc as seen in the C. bibroni species group. In the C. Candoia bibroni is effectively restricted to the main islands of bibroni species group the premaxilla has the “typical boine” form Fiji, including Vita Levu and Ovalau and those immediately described by Frazzetta (1959, 1975), with a large fenestra north-west. between the narial vestibules, this fenestra bounded anteriorly CANDOIA HOSERAE SP. NOV. by the premaxilla and posteriorly by the nasals; but C. hoserae Holotype: A female specimen in the American Museum of sp. nov. and others in that species group are like C. carinata Natural History (AMNH), New York, USA, specimen number: (with which they are sometimes sympatric) in having the AMNH 40407, collected from San Cristobal Island, Solomon intervestibular fenestra largely or entirely filled in by expansion of Islands. the premaxilla (derived from McDowell, 1979). The American Museum of Natural History (AMNH), New York, Candoia hoserae sp. nov. from the island of San Cristobal (AKA USA, allows access to its holdings. Makira) and nearby islets is separated from all other species Paratypes: Specimens in the American Museum of Natural within the C. hoserae sp. nov. species complex by the following History (AMNH), New York, USA, specimen numbers: AMNH unique suite of characters: A striped dorsal body pattern, being 40412, 42092, 42093, 42161, 42165, 42166, 42168, 42198, light brown with a pair of lengthwise white dorsolateral stripes 42199, 42200, 42201, 42202, 42212, 44502 from San Cristobal from over the quadrate bone to near the tail-tip, the stripes Island, Solomon Islands. edged with brown and a brown streak from angle of mouth almost to eye; the belly is buffy white; 235-257 ventrals and 18- Diagnosis: Candoia hoserae sp. nov. and others in the C. 22 maxillary teeth on either side of the mouth. hoserae sp. nov. species group are defined as the three species from the Solomon Islands, including outliers, as well as the Candoia louisemcgoldrickae sp. nov. from the islands of Rennell single species from the Banks Islands and nearby New and Bellona in the Solomon Islands is separated from all other Hebrides, which is herein treated as two separate subspecies, species within the C. hoserae sp. nov. species complex by the separated by deep water. following unique suite of characters: Similar in most respects to C. hoserae sp. nov. from which it can be separated from by the These taxa are C. hoserae sp. nov., C. louisemcgoldrickae sp. fact that the upper surface of the head is generally dark in nov., C. simonmcgoldricki sp. nov. and C. jamiekonstandinoui colour, the generally dark head and the prominent thin, light sp. nov., the latter of which includes the subspecies C. temporal streak that runs to the neck is bounded by a distinctive jamiekonstandinoui georgekonstandinoui subsp. nov.. white border at the margins. This taxon is also separated from These species are all separated from C. bibroni and others in C. hoserae sp. nov. by being of a generally blotched or broken the C. bibroni species group (being all others in the C. bibroni blotched dorsal pattern. complex) as detailed in the preceding text, by the form of the Candoia simonmcgoldricki sp. nov. from the Santa Cruz island postorbital bone, which has its ventral (free) end abruptly flexed group, including the nearby Reef Islands within the eastern backward, as in C. carinata, rather than forming a simple

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 47

crescent arc as seen in the C. bibroni species group. In the C. The C. bibroni species group are those species not included in bibroni species group the premaxilla has the “typical boine” form the C. hoserae sp. nov. species group, as defined below. described by Frazzetta (1959, 1975), with a large fenestra Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. between the narial vestibules, this fenestra bounded anteriorly species group are defined as the three species from the by the premaxilla and posteriorly by the nasals; but C. hoserae Solomon Islands, including outliers, as well as the single sp. nov. and others in that species group are like C. carinata species from the Banks Islands and nearby New Hebrides. (with which they are sometimes sympatric) in having the These species are all separated from C. bibroni and others in intervestibular fenestra largely or entirely filled in by expansion of the C. bibroni species group (being all others in the C. bibroni the premaxilla (derived from McDowell, 1979). complex) by the form of the postorbital bone, which has its Candoia hoserae sp. nov. from the island of San Cristobal (AKA ventral (free) end abruptly flexed backward, as in C. carinata, Makira) and nearby islets is separated from all other species rather than forming a simple crescent arc as seen in C. bibroni. within the C. hoserae sp. nov. species complex by the following In C. bibroni the premaxilla has the “typical boine” form unique suite of characters: A striped dorsal body pattern, being described by Frazzetta (1959, 1975), with a large fenestra light brown with pair of lengthwise white dorsolateral stripes from between the narial vestibules, this fenestra bounded anteriorly over quadrate bone to near the tail-tip, the stripes edged with by the premaxilla and posteriorly by the nasals; but C. hoserae brown and a brown streak from angle of mouth almost to eye; sp. nov. and others in the species group are like C. carinata the belly is buffy white; 235-257 ventrals, 18-22 maxillary teeth (with which they are sometimes sympatric) in having the on either side of the mouth. intervestibular fenestra largely or entirely filled in by expansion of Candoia louisemcgoldrickae sp. nov. from the islands of Rennell the premaxilla. and Bellona in the Solomon Islands is separated from all other Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as species within the C. hoserae sp. nov. species complex by the defined in this paper, is the taxon from the main Fiji islands following unique suite of characters: Similar in most respects to including Viti Levu and Vanua Levu and immediately adjacent C. hoserae sp. nov. from which it can be separated from by the islands, but not including Kadavu or the islands in the Lau fact that the upper surface of the head is generally dark in group, each of which has a different species. colour, the generally dark head and the prominent thin, light The taxon C. bibroni is separated from all other species within temporal streak that runs to the neck is bounded by a distinctive the C. bibroni species complex by the following suite of white border at the margins. This taxon is also separated from characters: A blotched dorsal pattern with distinctive throat C. hoserae sp. nov. by being of a generally blotched or broken stripes and a low ventral count ranging from 207-227 ventrals. blotched dorsal pattern. The species C. bibroni is further defined by the following suite of Candoia simonmcgoldricki sp. nov. from the Santa Cruz island characters: 18-22 Maxillary teeth, (versus 17-19 in C. woolfi sp. group, including the nearby Reef Islands within the eastern nov. from the Lau Group of islands), 53-59 subcaudals (versus Solomon Islands is separated from all other species within the 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- C. hoserae sp. nov. species complex by the following unique 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the suite of characters: Similar in most respects to C. hoserae sp. head at one head length behind the head (versus 33-34 in C. nov. and C. louisemcgoldrickae sp. nov from which it can be woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one separated from it by a profusion of black pigment on the dorsal headlength anterior to the cloaca, hemipenis forks at subcaudal snout tip and including the rostral and all of first labial on either 6-10. side. There is a particularly thick dark postorbital stripe (as opposed to medium in C. hoserae sp. nov. and C. The species Candoia woolfi sp. nov. from the Lau Group of louisemcgoldrickae sp. nov.). 38-41 midbody scale rows in C. islands, east of the main Fiji islands is separated from all other simonmcgoldricki sp. nov. versus 36-37 in C. hoserae sp. nov. species within the C. bibroni species complex by the following and Candoia louisemcgoldrickae sp. nov.. suite of characters: A blotched pattern without stripes. The tail pattern is much more conspicuous than that of body, the pale The species Candoia simonmcgoldricki sp. nov. is further bordering of body blotches also replacing darker ground color on diagnosed and defined by the following: 15-21 infralabials, 250- the tail; 205-236 ventrals; there are 33-34 scale rows behind the 266 ventrals versus 235-257 for C. hoserae sp. nov. and head at one head length behind the head (versus 26-30 in C. Candoia louisemcgoldrickae sp. nov., 61-64 subcaudals versus bibroni). 54-63 subcaudals for C. hoserae sp. nov. and Candoia louisemcgoldrickae sp. nov., 18-22 maxillary teeth, hemipenis The species Candoia kimmooreae sp. nov. from the island of fully everted extending from 9-20 subcaudals. Kadavu, southern Fiji is is separated from all other species within the C. bibroni species complex by having a blotched The species Candoia jamiekonstandinoui sp. nov. from Vanuatu pattern which includes a prominent median throat stripe and no and the nearby Banks Islands, including nearby islets is lateral throat stripes. Otherwise this taxon is essentially similar separated from all other species within the C. hoserae sp. nov. in form to nominate C. bibroni. species group by the following unique suite of characters: A distinctive dorsal colouration of pale reddish brown above with The species Candoia malcolmmaclurei sp. nov. from the Loyalty semidistinct markings, with a dark brown vertebral stripe edged Islands, (territory controlled by New Caledonia) is separated with yellow; a uniform yellow beneath. Males are unique in the from all other species within the C. bibroni species complex by C. bibroni species complex in that they have some noticeable having a blotched pattern without stripes and a tail similar in dark pigmentation on the hemipenes. 16-18 maxillary teeth for pattern to the body, or alternatively a blotched pattern with throat the nominate subspecies, versus 19 in C. jamiekonstandinoui stripes including a median row of dark spots on anterior ventrals; georgekonstandinoui subsp. nov.. dark perocular streak from angle of mouth (or angle of jaw) through eye to loreal region, dark spots on lips, and dark The subspecies C. jamiekonstandinoui jamiekonstandinoui spotting on top of head and with a median ventral row of dark subsp. nov. has 234-247 ventrals, versus 222-225 ventrals in C. spots. Sometimes the colouration is dulled to appear unicolour. jamiekonstandinoui georgekonstandinoui subsp. nov.. The species is further defined by having 13-16 infralabials, 27- C. jamiekonstandinoui georgekonstandinoui subsp. nov. has the 30 scale rows on the neck at one head length back, 20-22 scale lowest ventral counts of any named taxon within the C. hoserae rows at one headlength anterior to the cloaca, 219-226 ventrals, sp. nov. species group. 53-60 subcaudals and 19-22 maxillary teeth on either side. Candoia bibroni and other species described in this paper that The species Candoia boutrosi sp. nov. from the islands of were formerly treated as populations of C. bibroni are herein Samoa is separated from all other species within the C. bibroni defined as being the C. bibroni species complex. species complex by the following unique suite of characters: a

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 48 Australasian Journal of Herpetology distinctive blotched dorsal body pattern that is one or other of so-called Cheesman’s Line as defined by Hamilton et al. (2010), being distinct or very indistinct, the large dark blotches being the which is a well-known faunal boundary for Pacific island fauna. main component of the dorsal midline for the anterior and mid- Candoia hoserae sp. nov. is restricted to San Cristobal (AKA body, with the lighter blotches dominating the midline posteriorly, Makira) and immediately adjacent islets including Ugi Island and reversing on the tail to have the darker blotches dominate on the Olu Malau. tail, which remains brightly marked for the whole length. The Etymology: Named in honour of Maxine Hoser of the United dorsal colouration is a combination of chocolate brown and Kingdom for services to herpetology, including logistical beige in distinctly marked animals or a hybrid of this in the assistance’s for various past projects. indistinctly marked animals. On the head is a well-defined creamish bar on the upper labials above the jawline. CANDOIA SIMONMCGOLDRICKI SP. NOV. It is also defined as having 12 supralabials bilaterally (in Holotype: A preserved male specimen at the American common with the three species from the Fiji Islands), 15-16 Museum of Natural History, New York, USA, specimen number: infralabials (as for C. bibroni sensu stricto), and 31-32 scale AMNH 40430 collected from Santa Cruz Island, Solomon rows on the neck, one head length back from the head (versus Islands. 26-30 in C. bibroni sensu stricto). The American Museum of Natural History, New York, USA C. boutrosi sp. nov. is further defined and separated from the allows access to its holdings. other species in the C. bibroni complex by the following suite of Paratpes: 1/ A preserved male specimen at the American characters 37-38 mid-body rows, 22-24 scale rows at one head Museum of Natural History, New York, USA, specimen number: length anterior to the cloaca, 237-252 ventrals, 51-61 AMNH 42215 collected from Santa Cruz Island, Solomon subcaudals and 16-18 maxillary teeth on either side. Islands. The species Candoia niraikanukiwai sp. nov. from the islands of 2/ A preserved male specimen at the American Museum of Wallis and Futuna, including nearby islets is separated from all Natural History, New York, USA, specimen number: AMNH other species within the C. bibroni species complex by the 42216 collected from Santa Cruz Island, Solomon Islands. following unique suite of characters: It is essentially similar in 3/ A preserved female specimen at the American Museum of most respects to C. boutrosi sp. nov. from which it differs by Natural History, New York, USA, specimen number: AMNH having a distinctive well-defined stripe running through the eye 42160 collected from Santa Cruz Island, Solomon Islands. on either side of the head and the absence of a well-defined Diagnosis: Candoia hoserae sp. nov. and others in the C. creamish bar on the upper labials above the jawline. hoserae sp. nov. species group are defined as the three species The species Candoia georgemacintyrei sp. nov. from the island from the Solomon Islands, including outliers, as well as the of Rotuma, is separated from all other species within the C. single species from the Banks Islands and nearby New bibroni species complex by the following unique suite of Hebrides, which is herein treated as two separate subspecies, characters: 235-240 ventrals, 57-63 subcaudals, 37-39 mid separated by deep water. body rows and significant lightening of the upper labials; the These taxa are C. hoserae sp. nov., C. louisemcgoldrickae sp. dorsal pattern is blotched and semi-distinct and there is nov., C. simonmcgoldricki sp. nov. and C. jamiekonstandinoui significant speckling and markings on the venter. sp. nov., the latter of which includes the subspecies C. All species within the C. bibroni species complex are separated jamiekonstandinoui georgekonstandinoui subsp. nov.. from all other Candoia by having supralabials separated from These species are all separated from C. bibroni and others in the eye by the subocular scales, versus C. carinata (Schneider, the C. bibroni species group (being all others in the C. bibroni 1801) and C. paulsoni (Stull, 1956) which have them entering complex) as detailed in the preceding text, by the form of the the eye; preocular differentiated; canthus rostralis rounded; a postorbital bone, which has its ventral (free) end abruptly flexed relatively slender body with lengthwise ridges on the body scales backward, as in C. carinata, rather than forming a simple that run parallel to the long axis of the body, versus a stout body crescent arc as seen in the C. bibroni species group. In the C. with keels on the body scales forming oblique rows as in C. bibroni species group the premaxilla has the “typical boine” form aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. described by Frazzetta (1959, 1975), with a large fenestra Furthermore the tail is more than twice as long as the head, between the narial vestibules, this fenestra bounded anteriorly being capable of more than one complete circle of coiling in the by the premaxilla and posteriorly by the nasals; but C. hoserae vertical plane. sp. nov. and others in that species group are like C. carinata The species Candoia carinata (Schneider, 1801) including (with which they are sometimes sympatric) in having the species and subspecies formerly treated as being within this intervestibular fenestra largely or entirely filled in by expansion of taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. the premaxilla (derived from McDowell, 1979). bibroni complex by having 2, or rarely 1 or 3 supralabials Candoia hoserae sp. nov. from the island of San Cristobal (AKA entering eye; an angulate canthus rostralis, 160-202 ventrals Makira) and nearby islets is separated from all other species and 35-60 subcaudals. within the C. hoserae sp. nov. species complex by the following The species Candoia aspera (Günther, 1877) including species unique suite of characters: A striped dorsal body pattern, being and subspecies formerly treated as being within this taxon is light brown with pair of lengthwise white dorsolateral stripes from separated from the C. bibroni complex by having a tail less than over quadrate bone to near the tail-tip, the stripes edged with twice as long as the head, usually being shorter than the head; brown and a brown streak from angle of mouth almost to eye; is incapable of forming a full circle of coiling in vertical plane; at the belly is buffy white; 235-257 ventrals, 18-22 maxillary teeth least on the anterior two-thirds of body are keels of scales on either side of the mouth. forming curved diagonal ridges along the sides, which extend Candoia louisemcgoldrickae sp. nov. from the islands of Rennell backward and downward towards the belly; 127-153 ventrals; 11- and Bellona in the Solomon Islands is separated from all other 22 subcaudals; no specially enlarged preocular; (supralabials species within the C. hoserae sp. nov. species complex by the excluded from the eye; canthus rostralis angular). following unique suite of characters: Similar in most respects to Distribution: The Candoia hoserae sp. nov. species group are C. hoserae sp. nov. from which it can be separated from by the found in the Solomon Islands, including San Cristobal (Makira), fact that the upper surface of the head is generally dark in colour Rennell, Bellona, Ugi, Olu Malau, Santa Ana, Santa Cruz, Reef and the generally dark head and the prominent thin, light Islands, Vanikuro, Utupua as well as the Banks Islands and temporal streak that runs to the neck is bounded by a distinctive Vanuatu. white border at the margins. This taxon is also separated from The southern boundary of the species group distribution is the C. hoserae sp. nov. by being of a generally blotched or broken

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 49

blotched dorsal pattern. (FMNH Amphibians and Reptiles) from , Solomon Candoia simonmcgoldricki sp. nov. from the Santa Cruz island Islands. group, including the nearby Reef Islands within the eastern Diagnosis: Candoia hoserae sp. nov. and others in the C. Solomon Islands is separated from all other species within the hoserae sp. nov. species group including C. louisemcgoldrickae C. hoserae sp. nov. species complex by the following unique sp. nov. are defined as the three species from the Solomon suite of characters: Similar in most respects to C. hoserae sp. Islands, including outliers, as well as the single species from the nov. and C. louisemcgoldrickae sp. nov from which it can be proximal Banks Islands and nearby New Hebrides, which is separated from it by a profusion of black pigment on the dorsal herein treated as two separate subspecies, being separated by snout tip and including the rostral and all of first labial on either deep water. side. There is a particularly thick dark postorbital stripe (as These taxa are C. hoserae sp. nov., C. louisemcgoldrickae sp. opposed to medium in C. hoserae sp. nov. and C. nov., C. simonmcgoldricki sp. nov. and C. jamiekonstandinoui louisemcgoldrickae sp. nov.). 38-41 midbody scale rows in C. sp. nov., the latter of which includes the subspecies C. simonmcgoldricki sp. nov. versus 36-37 in C. hoserae sp. nov. jamiekonstandinoui georgekonstandinoui subsp. nov.. and Candoia louisemcgoldrickae sp. nov.. These species are all separated from C. bibroni and others in The species Candoia simonmcgoldricki sp. nov. is further the C. bibroni species group (being all others in the C. bibroni diagnosed and defined by the following: 15-21 infralabials, 250- complex) as detailed in the preceding text, by the form of the 266 ventrals versus 235-257 for C. hoserae sp. nov. and postorbital bone, which has its ventral (free) end abruptly flexed Candoia louisemcgoldrickae sp. nov., 61-64 subcaudals versus backward, as in C. carinata, rather than forming a simple 54-63 subcaudals for C. hoserae sp. nov. and Candoia crescent arc as seen in the C. bibroni species group. In the C. louisemcgoldrickae sp. nov., 18-22 maxillary teeth, hemipenis bibroni species group the premaxilla has the “typical boine” form fully everted extending from 9-20 subcaudals. described by Frazzetta (1959, 1975), with a large fenestra The species Candoia jamiekonstandinoui sp. nov. from Vanuatu between the narial vestibules, this fenestra bounded anteriorly and the nearby Banks Islands, including nearby islets is by the premaxilla and posteriorly by the nasals; but C. hoserae separated from all other species within the C. hoserae sp. nov. sp. nov. and others in that species group are like C. carinata species group by the following unique suite of characters: A (with which they are sometimes sympatric) in having the distinctive dorsal colouration of pale reddish brown above with intervestibular fenestra largely or entirely filled in by expansion of semidistinct markings, with a dark brown vertebral stripe edged the premaxilla (derived from McDowell, 1979). with yellow; a uniform yellow beneath. Males are unique in the Candoia hoserae sp. nov. from the island of San Cristobal (AKA C. bibroni species complex in that they have some noticeable Makira) and nearby islets is separated from all other species dark pigmentation on the hemipenes. 16-18 maxillary teeth for within the C. hoserae sp. nov. species complex by the following the nominate subspecies, versus 19 in C. jamiekonstandinoui unique suite of characters: A striped dorsal body pattern, being georgekonstandinoui subsp. nov.. light brown with pair of lengthwise white dorsolateral stripes from The subspecies C. jamiekonstandinoui jamiekonstandinoui over quadrate bone to near the tail-tip, the stripes edged with subsp. nov. has 234-247 ventrals, versus 222-225 ventrals in C. brown and a brown streak from angle of mouth almost to eye; jamiekonstandinoui georgekonstandinoui subsp. nov.. the belly is buffy white; 235-257 ventrals, 18-22 maxillary teeth C. jamiekonstandinoui georgekonstandinoui subsp. nov. has the on either side of the mouth. lowest ventral counts of any named taxon within the C. hoserae Candoia louisemcgoldrickae sp. nov. from the islands of Rennell sp. nov. species group. and Bellona in the Solomon Islands is separated from all other Candoia bibroni and other species described in this paper that species within the C. hoserae sp. nov. species complex by the were formerly treated as populations of C. bibroni are herein following unique suite of characters: Similar in most respects to defined as being the C. bibroni species complex. C. hoserae sp. nov. from which it can be separated from by the The C. bibroni species group are those species not included in fact that the upper surface of the head is generally dark in the C. hoserae sp. nov. species group, as defined elsewhere in colour, the generally dark head and the prominent thin, light this paper. temporal streak that runs to the neck is bounded by a distinctive Distribution: The Candoia hoserae sp. nov. species group are white border at the margins. This taxon is also separated from found in the Solomon Islands, including San Cristobal (Makira), C. hoserae sp. nov. by being of a generally blotched or broken Rennell, Bellona, Ugi, Olu Malau, Santa Ana, Santa Cruz, Reef blotched dorsal pattern. Islands, Vanikuro, Utupua as well as the Banks Islands and Candoia simonmcgoldricki sp. nov. from the Santa Cruz island Vanuatu. group, including the nearby Reef Islands within the eastern The southern boundary of the species distribution is the so- Solomon Islands is separated from all other species within the called Cheesman’s Line as defined by Hamilton et al. (2010), C. hoserae sp. nov. species complex by the following unique which is a well-known faunal boundary for Pacific island fauna. suite of characters: Similar in most respects to C. hoserae sp. nov. and C. louisemcgoldrickae sp. nov from which it can be C. simonmcgoldricki sp. nov. is restricted to the Santa Cruz separated from it by a profusion of black pigment on the dorsal islands and nearby islands including the Reef Islands, Santa snout tip and including the rostral and all of first labial on either Ana, Vanikuro, Utupua and immediately adjacent islets. side. There is a particularly thick dark postorbital stripe (as Etymology: C. simonmcgoldricki sp. nov. is named in honour of opposed to medium in C. hoserae sp. nov. and C. Simon McGoldrick of Ringwood East, Melbourne, Victoria, louisemcgoldrickae sp. nov.). 38-41 midbody scale rows in C. Australia, for his contributions to wildlife conservation and simonmcgoldricki sp. nov. versus 36-37 in C. hoserae sp. nov. education including through his work with Snakebusters: and Candoia louisemcgoldrickae sp. nov.. Australia’s best reptiles shows. The species Candoia simonmcgoldricki sp. nov. is further CANDOIA LOUISEMCGOLDRICKAE SP. NOV. diagnosed and defined by the following: 15-21 infralabials, 250- Holotype: A specimen at the Field Museum of Natural History 266 ventrals versus 235-257 for C. hoserae sp. nov. and (FMNH), Chicago, Iliinois, USA, specimen number: 71741 Candoia louisemcgoldrickae sp. nov., 61-64 subcaudals versus (FMNH Amphibians and Reptiles) from Bellona Island, Solomon 54-63 subcaudals for C. hoserae sp. nov. and Candoia Islands. louisemcgoldrickae sp. nov., 18-22 maxillary teeth, hemipenis This facility allows access to its holdings by scientists. fully everted extending from 9-20 subcaudals. Paratype: A specimen at the Field Museum of Natural History The species Candoia jamiekonstandinoui sp. nov. from Vanuatu (FMNH), Chicago, Iliinois, USA, specimen number: 71742 and the nearby Banks Islands, including nearby islets is

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 50 Australasian Journal of Herpetology separated from all other species within the C. hoserae sp. nov. between the narial vestibules, this fenestra bounded anteriorly species group by the following unique suite of characters: A by the premaxilla and posteriorly by the nasals; but C. hoserae distinctive dorsal colouration of pale reddish brown above with sp. nov. and others in that species group are like C. carinata semidistinct markings, with a dark brown vertebral stripe edged (with which they are sometimes sympatric) in having the with yellow; a uniform yellow beneath. Males are unique in the intervestibular fenestra largely or entirely filled in by expansion of C. bibroni species complex in that they have some noticeable the premaxilla (derived from McDowell, 1979). dark pigmentation on the hemipenes. 16-18 maxillary teeth for Candoia hoserae sp. nov. from the island of San Cristobal (AKA the nominate subspecies, versus 19 in C. jamiekonstandinoui Makira) and nearby islets is separated from all other species georgekonstandinoui subsp. nov.. within the C. hoserae sp. nov. species complex by the following The subspecies C. jamiekonstandinoui jamiekonstandinoui unique suite of characters: A striped dorsal body pattern, being subsp. nov. has 234-247 ventrals, versus 222-225 ventrals in C. light brown with pair of lengthwise white dorsolateral stripes from jamiekonstandinoui georgekonstandinoui subsp. nov.. over quadrate bone to near the tail-tip, the stripes edged with C. jamiekonstandinoui georgekonstandinoui subsp. nov. has the brown and a brown streak from angle of mouth almost to eye; lowest ventral counts of any named taxon within the C. hoserae the belly is buffy white; 235-257 ventrals, 18-22 maxillary teeth sp. nov. species group. on either side of the mouth. Candoia bibroni and other species described in this paper that Candoia louisemcgoldrickae sp. nov. from the islands of Rennell were formerly treated as populations of C. bibroni are herein and Bellona in the Solomon Islands is separated from all other defined as being the C. bibroni species complex. species within the C. hoserae sp. nov. species complex by the The C. bibroni species group are those species not included in following unique suite of characters: Similar in most respects to the C. hoserae sp. nov. species group, as defined elsewhere in C. hoserae sp. nov. from which it can be separated from by the this paper. fact that the upper surface of the head is generally dark in colour and the generally dark head and the prominent thin, light Distribution: C. louisemcgoldrickae sp. nov. is restricted to temporal streak that runs to the neck is bounded by a distinctive Rennell and Bellona and immediately adjacent islets in the white border at the margins. This taxon is also separated from south-east Solomon Islands. C. hoserae sp. nov. by being of a generally blotched or broken The Candoia hoserae sp. nov. species group are found in the blotched dorsal pattern. Solomon Islands, including San Cristobal (Makira), Rennell, Bellona, Ugi, Olu Malau, Santa Ana, Santa Cruz, Reef Islands, Candoia simonmcgoldricki sp. nov. from the Santa Cruz island Vanikuro, Utupua as well as the Banks Islands and Vanuatu. group, including the nearby Reef Islands within the eastern Solomon Islands is separated from all other species within the The southern boundary of the species distribution is the so- C. hoserae sp. nov. species complex by the following unique called Cheesman’s Line as defined by Hamilton et al. (2010), suite of characters: Similar in most respects to C. hoserae sp. which is a well-known faunal boundary for Pacific island fauna. nov. and C. louisemcgoldrickae sp. nov from which it can be Etymology: C. louisemcgoldrickae sp. nov. is named in honour separated from it by a profusion of black pigment on the dorsal of Louise McGoldrick of Ringwood East, Melbourne, Victoria, snout tip and including the rostral and all of first labial on either Australia, for her contributions to wildlife conservation and side. There is a particularly thick dark postorbital stripe (as education including through her work with Snakebusters: opposed to medium in C. hoserae sp. nov. and C. Australia’s best reptiles shows over many years. louisemcgoldrickae sp. nov.). 38-41 midbody scale rows in C. CANDOIA JAMIEKONSTANDINOUI SP. NOV. simonmcgoldricki sp. nov. versus 36-37 in C. hoserae sp. nov. Holotype: A female specimen at the American Museum of and Candoia louisemcgoldrickae sp. nov.. Natural History (AMNH), New York, USA, specimen number: The species Candoia simonmcgoldricki sp. nov. is further 42004 collected at Malekula Island (Vanuatu), New Hebrides. diagnosed and defined by the following: 15-21 infralabials, 250- This is a facility that allows access to its holdings by scientists. 266 ventrals versus 235-257 for C. hoserae sp. nov. and Candoia louisemcgoldrickae sp. nov., 61-64 subcaudals versus Paratypes: 1/ A female specimen at the American Museum of 54-63 subcaudals for C. hoserae sp. nov. and Candoia Natural History (AMNH), New York, USA, specimen number: louisemcgoldrickae sp. nov., 18-22 maxillary teeth, hemipenis 81584 collected at Malekula Island (Vanuatu), New Hebrides. fully everted extending from 9-20 subcaudals. 2/ A female specimen at the American Museum of Natural The species Candoia jamiekonstandinoui sp. nov. from Vanuatu History (AMNH), New York, USA specimen number: 81583 and the nearby Banks Islands, including nearby islets is collected at Espiritu Santo (Vanuatu), New Hebrides. separated from all other species within the C. hoserae sp. nov. 3/ A male specimen at the American Museum of Natural History species group by the following unique suite of characters: A (AMNH), New York, USA specimen number: 42075 collected at distinctive dorsal colouration of pale reddish brown above with Espiritu Santo (Vanuatu), New Hebrides. semidistinct markings, with a dark brown vertebral stripe edged Diagnosis: Candoia hoserae sp. nov. and others in the C. with yellow; a uniform yellow beneath. Males are unique in the hoserae sp. nov. species group are defined as the three species C. bibroni species complex in that they have some noticeable from the Solomon Islands, including outliers, as well as the dark pigmentation on the hemipenes. 16-18 maxillary teeth for single species from the Banks Islands and nearby New the nominate subspecies, versus 19 in C. jamiekonstandinoui Hebrides, which is herein treated as two separate subspecies, georgekonstandinoui subsp. nov.. separated by deep water. The subspecies C. jamiekonstandinoui jamiekonstandinoui These taxa are C. hoserae sp. nov., C. louisemcgoldrickae sp. subsp. nov. has 234-247 ventrals, versus 222-225 ventrals in C. nov., C. simonmcgoldricki sp. nov. and C. jamiekonstandinoui jamiekonstandinoui georgekonstandinoui subsp. nov.. sp. nov., the latter of which includes the subspecies C. C. jamiekonstandinoui georgekonstandinoui subsp. nov. has the jamiekonstandinoui georgekonstandinoui subsp. nov.. lowest ventral counts of any named taxon within the C. hoserae These species are all separated from C. bibroni and others in sp. nov. species group. the C. bibroni species group (being all others in the C. bibroni Candoia bibroni and other species described in this paper that complex) as detailed in the preceding text, by the form of the were formerly treated as populations of C. bibroni are herein postorbital bone, which has its ventral (free) end abruptly flexed defined as being the C. bibroni species complex. backward, as in C. carinata, rather than forming a simple The C. bibroni species group are those species not included in crescent arc as seen in the C. bibroni species group. In the C. the C. hoserae sp. nov. species group, as defined elsewhere in bibroni species group the premaxilla has the “typical boine” form this paper. described by Frazzetta (1959, 1975), with a large fenestra Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 51

Distribution: The nominate form of C. jamiekonstandinoui C. hoserae sp. nov. species complex by the following unique jamiekonstandinoui subsp. nov. is found on the New Hebrides suite of characters: Similar in most respects to C. hoserae sp. islands of Malekula, Espiritu Santo, Ambae (AKA Aoba), nov. and C. louisemcgoldrickae sp. nov from which it can be and Maewo. The subspecies C. jamiekonstandinoui separated from it by a profusion of black pigment on the dorsal georgekonstandinoui subsp. nov. is found on the Banks Islands snout tip and including the rostral and all of first labial on either including Vanua Lava and presumably other major islands in the side. There is a particularly thick dark postorbital stripe (as group. opposed to medium in C. hoserae sp. nov. and C. Etymology: Named in honour of Jamie Christopher louisemcgoldrickae sp. nov.). 38-41 midbody scale rows in C. Konstandinoui of George’s Park Orchards Service Centre in simonmcgoldricki sp. nov. versus 36-37 in C. hoserae sp. nov. Ringwood, Victoria, Australia formerly of Park Orchards, and Candoia louisemcgoldrickae sp. nov.. Victoria, Australia, for vital logistical support to important The species Candoia simonmcgoldricki sp. nov. is further herpetological scientific research projects for more than a diagnosed and defined by the following: 15-21 infralabials, 250- decade. 266 ventrals versus 235-257 for C. hoserae sp. nov. and CANDOIA JAMIEKONSTANDINOUI Candoia louisemcgoldrickae sp. nov., 61-64 subcaudals versus GEORGEKONSTANDINOUI SUBSP. NOV. 54-63 subcaudals for C. hoserae sp. nov. and Candoia Holotype: A male specimen at the American Museum of Natural louisemcgoldrickae sp. nov., 18-22 maxillary teeth, hemipenis History (AMNH), New York, USA, specimen number: 81581 fully everted extending from 9-20 subcaudals. collected at Vanua Lava (Banks Islands), New Hebrides. The species Candoia jamiekonstandinoui sp. nov. from Vanuatu and the nearby Banks Islands, including nearby islets is This is a facility that allows access to its holdings by scientists. separated from all other species within the C. hoserae sp. nov. Paratype: A male specimen at the American Museum of Natural species group by the following unique suite of characters: A History (AMNH), New York, USA, specimen number: 81582 distinctive dorsal colouration of pale reddish brown above with collected at Vanua Lava (Banks Islands), New Hebrides. semidistinct markings, with a dark brown vertebral stripe edged Diagnosis: Candoia hoserae sp. nov. and others in the C. with yellow; a uniform yellow beneath. Males are unique in the hoserae sp. nov. species group are defined herein as the three C. bibroni species complex in that they have some noticeable species from the Solomon Islands, including outliers, as well as dark pigmentation on the hemipenes. 16-18 maxillary teeth for the single species from the Banks Islands and nearby northern the nominate subspecies, versus 19 in C. jamiekonstandinoui and central New Hebrides, which is herein treated as two georgekonstandinoui subsp. nov.. separate subspecies, separated by deep water. The subspecies C. jamiekonstandinoui jamiekonstandinoui These taxa are C. hoserae sp. nov., C. louisemcgoldrickae sp. subsp. nov. has 234-247 ventrals, versus 222-225 ventrals in C. nov., C. simonmcgoldricki sp. nov. and C. jamiekonstandinoui jamiekonstandinoui georgekonstandinoui subsp. nov.. sp. nov., the latter of which includes the subspecies C. C. jamiekonstandinoui georgekonstandinoui subsp. nov. has the jamiekonstandinoui georgekonstandinoui subsp. nov.. lowest ventral counts of any named taxon within the C. hoserae These species are all separated from C. bibroni and others in sp. nov. species group. the C. bibroni species group (being all others in the C. bibroni Candoia bibroni and other species described in this paper that complex) as detailed in the preceding text, by the form of the were formerly treated as populations of C. bibroni are herein postorbital bone, which has its ventral (free) end abruptly flexed defined as being the C. bibroni species complex. backward, as in C. carinata, rather than forming a simple The C. bibroni species group are those species not included in crescent arc as seen in the C. bibroni species group. In the C. the C. hoserae sp. nov. species group, as defined elsewhere in bibroni species group the premaxilla has the “typical boine” form this paper. described by Frazzetta (1959, 1975), with a large fenestra between the narial vestibules, this fenestra bounded anteriorly Distribution: The nominate form of C. jamiekonstandinoui by the premaxilla and posteriorly by the nasals; but C. hoserae jamiekonstandinoui subsp. nov. is found on the New Hebrides sp. nov. and others in that species group are like C. carinata islands of Malekula, Espiritu Santo, Ambae (AKA Aoba), Malo (with which they are sometimes sympatric) in having the and Maewo. The subspecies C. jamiekonstandinoui intervestibular fenestra largely or entirely filled in by expansion of georgekonstandinoui subsp. nov. as defined herein is found on the premaxilla (derived from McDowell, 1979). the Banks Islands including Vanua Lava and presumably other major islands in the group. Candoia hoserae sp. nov. from the island of San Cristobal (AKA Etymology: This Subspecies is named in honour of George Makira) and nearby islets is separated from all other species Konstandinoui of George’s Park Orchards Service Centre in within the C. hoserae sp. nov. species complex by the following Ringwood, Victoria, Australia formerly of Park Orchards, unique suite of characters: A striped dorsal body pattern, being Victoria, Australia, for vital logistical support to significant light brown with pair of lengthwise white dorsolateral stripes from herpetological scientific research projects and Snakebusters over quadrate bone to near the tail-tip, the stripes edged with wildlife conservation displays for more than a decade. brown and a brown streak from angle of mouth almost to eye; the belly is buffy white; 235-257 ventrals, 18-22 maxillary teeth CANDOIA WOOLFI SP. NOV. on either side of the mouth. Holotype: A male specimen at the Museum of Comparative Candoia louisemcgoldrickae sp. nov. from the islands of Rennell Zoology, Harvard University, USA, specimen number MCZ Herp and Bellona in the Solomon Islands is separated from all other R-15020, from Lakeba (AKA Lakemba) Island in the Lau group species within the C. hoserae sp. nov. species complex by the of islands, Fiji Islands. following unique suite of characters: Similar in most respects to The Museum of Comparative Zoology, Harvard University, USA, C. hoserae sp. nov. from which it can be separated from by the is a facility that allows access to its holdings by scientists. fact that the upper surface of the head is generally dark in colour Paratypes: Two female specimens at the Museum of and the generally dark head and the prominent thin, light Comparative Zoology, Harvard University, USA, specimen temporal streak that runs to the neck is bounded by a distinctive numbers MCZ Herp R-15019 and MCZ Herp R-15021, from white border at the margins. This taxon is also separated from Lakeba (AKA Lakemba) Island in the Lau group of islands, Fiji C. hoserae sp. nov. by being of a generally blotched or broken Islands. blotched dorsal pattern. Diagnosis: Candoia bibroni and other species described in this Candoia simonmcgoldricki sp. nov. from the Santa Cruz island paper that were formerly treated as populations of C. bibroni are group, including the nearby Reef Islands within the eastern herein defined as being the C. bibroni species complex. Solomon Islands is separated from all other species within the Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 52 Australasian Journal of Herpetology

The C. bibroni species group are those species not included in suite of characters: A blotched pattern without stripes. The tail the C. hoserae sp. nov. species group, as defined below or pattern is much more conspicuous than that of body, the pale elsewhere in this paper. bordering of body blotches also replacing darker ground color on All species within the C. bibroni species complex are separated the tail; 205-236 ventrals; there are 33-34 scale rows behind the from all other Candoia by having supralabials separated from head at one head length behind the head (versus 26-30 in C. the eye by the subocular scales, versus C. carinata (Schneider, bibroni). 1801) and C. paulsoni (Stull, 1956) which have them entering The species Candoia kimmooreae sp. nov. from the island of the eye; preocular differentiated; canthus rostralis rounded; a Kadavu, southern Fiji is is separated from all other species relatively slender body with lengthwise ridges on the body scales within the C. bibroni species complex by having a blotched that run parallel to the long axis of the body, versus a stout body pattern which includes a prominent median throat stripe and no with keels on the body scales forming oblique rows as in C. lateral throat stripes. Otherwise this taxon is essentially similar aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. in form to nominate C. bibroni. Furthermore the tail more than twice as long as the head, being The species Candoia malcolmmaclurei sp. nov. from the Loyalty capable of more than one complete circle of coiling in the Islands, (territory controlled by New Caledonia) is separated vertical plane. from all other species within the C. bibroni species complex by The species Candoia carinata (Schneider, 1801) including having a blotched pattern without stripes and a tail similar in species and subspecies formerly treated as being within this pattern to the body, or alternatively a blotched pattern with throat taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. stripes including a median row of dark spots on anterior ventrals; bibroni complex by having 2, or rarely 1 or 3 supralabials dark perocular streak from angle of mouth (or angle of jaw) entering eye; an angulate canthus rostralis, 160-202 ventrals through eye to loreal region, dark spots on lips, and dark and 35-60 subcaudals. spotting on top of head and with a median ventral row of dark The species Candoia aspera (Günther, 1877) including species spots. Sometimes the colouration is dulled to appear unicolour. and subspecies formerly treated as being within this taxon is The species is further defined by having 13-16 infralabials, 27- separated from the C. bibroni complex by having a tail less than 30 scale rows on the neck at one head length back, 20-22 scale twice as long as the head, usually being shorter than the head; rows at one headlength anterior to the cloaca, 219-226 ventrals, is incapable of forming a full circle of coiling in vertical plane; at 53-60 subcaudals and 19-22 maxillary teeth on either side. least on the anterior two-thirds of body are keels of scales The species Candoia boutrosi sp. nov. from the islands of forming curved diagonal ridges along the sides, which extend Samoa is separated from all other species within the C. bibroni backward and downward towards the belly; 127-153 ventrals; 11- species complex by the following unique suite of characters: a 22 subcaudals; no specially enlarged preocular; (supralabials distinctive blotched dorsal body pattern that is one or other of excluded from the eye; canthus rostralis angular). being distinct or very indistinct, the large dark blotches being the Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. main component of the dorsal midline for the anterior and mid- species group are defined as the three species from the body, with the lighter blotches dominating the midline posteriorly, Solomon Islands, including outliers, as well as the single reversing on the tail to have the darker blotches dominate on the species from the Banks Islands and nearby New Hebrides. tail, which remains brightly marked for the whole length. The These species which are defined in detail already in this paper, dorsal colouration is a combination of chocolate brown and are all separated from C. bibroni and others in the C. bibroni beige in distinctly marked animals or a hybrid of this in the species group (being all others in the C. bibroni complex) by the indistinctly marked animals. On the head is a well-defined form of the postorbital bone, which has its ventral (free) end creamish bar on the upper labials above the jawline. abruptly flexed backward, as in C. carinata, rather than forming It is also defined as having 12 supralabials bilaterally (in a simple crescent arc as seen in C. bibroni. In C. bibroni the common with the three species from the Fiji Islands), 15-16 premaxilla has the “typical boine” form described by Frazzetta infralabials (as for C. bibroni sensu stricto), and 31-32 scale (1959, 1975), with a large fenestra between the narial vestibules, rows on the neck, one head length back from the head (versus this fenestra bounded anteriorly by the premaxilla and posteriorly 26-30 in C. bibroni sensu stricto). by the nasals; but C. hoserae sp. nov. and others in the species C. boutrosi sp. nov. is further defined and separated from the group are like C. carinata (with which they are sometimes other species in the C. bibroni complex by the following suite of sympatric) in having the intervestibular fenestra largely or characters 37-38 mid-body rows, 22-24 scale rows at one head entirely filled in by expansion of the premaxilla. length anterior to the cloaca, 237-252 ventrals, 51-61 Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as subcaudals and 16-18 maxillary teeth on either side. defined in this paper, is the taxon from the main Fiji islands The species Candoia niraikanukiwai sp. nov. from the islands of including Viti Levu and Vanua Levu and immediately adjacent Wallis and Futuna, including nearby islets is separated from all islands, but not including Kadavu or the islands in the Lau other species within the C. bibroni species complex by the group, each of which has a different species. following unique suite of characters: It is essentially similar in The taxon C. bibroni is separated from all other species within most respects to C. boutrosi sp. nov. from which it differs by the C. bibroni species complex by the following suite of having a distinctive well-defined stripe running through the eye characters: A blotched dorsal pattern with distinctive throat on either side of the head and the absence of a well-defined stripes and a low ventral count ranging from 207-227 ventrals. creamish bar on the upper labials above the jawline. The species C. bibroni is further defined by the following suite of The species Candoia georgemacintyrei sp. nov. from the island characters: 18-22 Maxillary teeth, (versus 17-19 in C. woolfi sp. of Rotuma, is separated from all other species within the C. nov. from the Lau Group of islands), 53-59 subcaudals (versus bibroni species complex by the following unique suite of 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- characters: 235-240 ventrals, 57-63 subcaudals, 37-39 mid 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the body rows and significant lightening of the upper labials; the head at one head length behind the head (versus 33-34 in C. dorsal pattern is blotched and semi-distinct and there is woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one significant speckling and markings on the venter. headlength anterior to the cloaca, hemipenis forks at subcaudal Distribution: C. woolfi sp. nov. is found in the Lau Group of 6-10. islands, east of the main Fiji islands, in a region generally south The species Candoia woolfi sp. nov. from the Lau Group of of the deep water Nanuku Passage and east of the Koro Sea. islands, east of the main Fiji islands is separated from all other The taxon is found on most of the larger islands in the group species within the C. bibroni species complex by the following including Vanua Mbelavu, Lakemba, Ongea Levu and Fulanga.

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 53

Etymology: Named in honour of Paul Woolf, of Walloon, sympatric) in having the intervestibular fenestra largely or Queensland, Australia in recognition of contributions to entirely filled in by expansion of the premaxilla. herpetology spanning three decades, including in his role as Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as foundation president of the Herpetological Society of defined in this paper, is the taxon from the main Fiji islands Queensland Incorporated, HSQI, and management roles in the including Viti Levu and Vanua Levu and immediately adjacent society ever since, as well his defence of herpetology from islands, but not including Kadavu or the islands in the Lau illegal acts of taxonomic vandalism by the likes of convicted group, each of which has a different species. criminal David John Williams and associates in crime, Wolfgang The taxon C. bibroni is separated from all other species within Wüster, Mark O’Shea, Wulf Schleip, Scott Thomson, Van the C. bibroni species complex by the following suite of Wallach, George Zug, Anders Rhodin and Hinrich Kaiser. characters: A blotched dorsal pattern with distinctive throat CANDOIA KIMMOOREAE SP. NOV. stripes and a low ventral count ranging from 207-227 ventrals. Holotype: A preserved specimen at the Australian Museum, The species C. bibroni is further defined by the following suite of Sydney, NSW, Australia, specimen number: R.135213 collected characters: 18-22 Maxillary teeth, (versus 17-19 in C. woolfi sp. at Nakasaleka in the north of Kadavu Island, Fiji (18°57’S, nov. from the Lau Group of islands), 53-59 subcaudals (versus 178°23’E). 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- This is a government owned facility that allows access to its 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the holdings. head at one head length behind the head (versus 33-34 in C. Paratypes: Two preserved specimens at the Australian woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one Museum, Sydney, NSW, Australia, specimen numbers: headlength anterior to the cloaca, hemipenis forks at subcaudal R.135212 and R.135214 collected at Nakasaleka in the north of 6-10. Kadavu Island, Fiji (18°57’S, 178°23’E). The species Candoia woolfi sp. nov. from the Lau Group of Diagnosis: Candoia bibroni and other species described in this islands, east of the main Fiji islands is separated from all other paper that were formerly treated as populations of C. bibroni are species within the C. bibroni species complex by the following herein defined as being the C. bibroni species complex. suite of characters: A blotched pattern without stripes. The tail The C. bibroni species group are those species not included in pattern is much more conspicuous than that of body, the pale the C. hoserae sp. nov. species group, as defined below. bordering of body blotches also replacing darker ground color on the tail; 205-236 ventrals; there are 33-34 scale rows behind the All species within the C. bibroni species complex are separated head at one head length behind the head (versus 26-30 in C. from all other Candoia by having supralabials separated from bibroni). the eye by the subocular scales, versus C. carinata (Schneider, The species Candoia kimmooreae sp. nov. from the island of 1801) and C. paulsoni (Stull, 1956) which have them entering Kadavu, southern Fiji is is separated from all other species the eye; preocular differentiated; canthus rostralis rounded; a within the C. bibroni species complex by having a blotched relatively slender body with lengthwise ridges on the body scales pattern which includes a prominent median throat stripe and no that run parallel to the long axis of the body, versus a stout body lateral throat stripes. Otherwise this taxon is essentially similar with keels on the body scales forming oblique rows as in C. in form to nominate C. bibroni as described herein. aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. Furthermore the tail more than twice as long as the head, being The species Candoia malcolmmaclurei sp. nov. from the Loyalty capable of more than one complete circle of coiling in the Islands, (territory controlled by New Caledonia) is separated vertical plane. from all other species within the C. bibroni species complex by The species Candoia carinata (Schneider, 1801) including having a blotched pattern without stripes and a tail similar in species and subspecies formerly treated as being within this pattern to the body, or alternatively a blotched pattern with throat taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. stripes including a median row of dark spots on anterior ventrals; bibroni complex by having 2, or rarely 1 or 3 supralabials dark perocular streak from angle of mouth (or angle of jaw) entering eye; an angulate canthus rostralis, 160-202 ventrals through eye to loreal region, dark spots on lips, and dark and 35-60 subcaudals. spotting on top of head and with a median ventral row of dark spots. Sometimes the colouration is dulled to appear unicolour. The species Candoia aspera (Günther, 1877) including species The species is further defined by having 13-16 infralabials, 27- and subspecies formerly treated as being within this taxon is 30 scale rows on the neck at one head length back, 20-22 scale separated from the C. bibroni complex by having a tail less than rows at one headlength anterior to the cloaca, 219-226 ventrals, twice as long as the head, usually being shorter than the head; 53-60 subcaudals and 19-22 maxillary teeth on either side. is incapable of forming a full circle of coiling in vertical plane; at The species Candoia boutrosi sp. nov. from the islands of least on the anterior two-thirds of body are keels of scales Samoa is separated from all other species within the C. bibroni forming curved diagonal ridges along the sides, which extend species complex by the following unique suite of characters: a backward and downward towards the belly; 127-153 ventrals; 11- distinctive blotched dorsal body pattern that is one or other of 22 subcaudals; no specially enlarged preocular; supralabials being distinct or very indistinct, the large dark blotches being the excluded from the eye; canthus rostralis angular. main component of the dorsal midline for the anterior and mid- Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. body, with the lighter blotches dominating the midline posteriorly, species group are defined as the three species from the reversing on the tail to have the darker blotches dominate on the Solomon Islands, including outliers, as well as the single tail, which remains brightly marked for the whole length. The species from the Banks Islands and nearby New Hebrides. dorsal colouration is a combination of chocolate brown and These species which are defined in detail already in this paper, beige in distinctly marked animals or a hybrid of this in the are all separated from C. bibroni and others in the C. bibroni indistinctly marked animals. On the head is a well-defined species group (being all others in the C. bibroni complex) by the creamish bar on the upper labials above the jawline. form of the postorbital bone, which has its ventral (free) end It is also defined as having 12 supralabials bilaterally (in abruptly flexed backward, as in C. carinata, rather than forming common with the three species from the Fiji Islands), 15-16 a simple crescent arc as seen in C. bibroni. In C. bibroni the infralabials (as for C. bibroni sensu stricto), and 31-32 scale premaxilla has the “typical boine” form described by Frazzetta rows on the neck, one head length back from the head (versus (1959, 1975), with a large fenestra between the narial vestibules, 26-30 in C. bibroni sensu stricto). this fenestra bounded anteriorly by the premaxilla and posteriorly by the nasals; but C. hoserae sp. nov. and others in the species C. boutrosi sp. nov. is further defined and separated from the group are like C. carinata (with which they are sometimes other species in the C. bibroni complex by the following suite of Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 54 Australasian Journal of Herpetology characters 37-38 mid-body rows, 22-24 scale rows at one head 22 subcaudals; no specially enlarged preocular; (supralabials length anterior to the cloaca, 237-252 ventrals, 51-61 excluded from the eye; canthus rostralis angular). subcaudals and 16-18 maxillary teeth on either side. Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. The species Candoia niraikanukiwai sp. nov. from the islands of species group are defined as the three species from the Wallis and Futuna, including nearby islets is separated from all Solomon Islands, including outliers, as well as the single other species within the C. bibroni species complex by the species from the Banks Islands and nearby New Hebrides. following unique suite of characters: It is essentially similar in These species which are defined in detail already in this paper, most respects to C. boutrosi sp. nov. from which it differs by are all separated from C. bibroni and others in the C. bibroni having a distinctive well-defined stripe running through the eye species group (being all others in the C. bibroni complex) by the on either side of the head and the absence of a well-defined form of the postorbital bone, which has its ventral (free) end creamish bar on the upper labials above the jawline. abruptly flexed backward, as in C. carinata, rather than forming The species Candoia georgemacintyrei sp. nov. from the island a simple crescent arc as seen in C. bibroni. In C. bibroni the of Rotuma, is separated from all other species within the C. premaxilla has the “typical boine” form described by Frazzetta bibroni species complex by the following unique suite of (1959, 1975), with a large fenestra between the narial vestibules, characters: 235-240 ventrals, 57-63 subcaudals, 37-39 mid this fenestra bounded anteriorly by the premaxilla and posteriorly body rows and significant lightening of the upper labials; the by the nasals; but C. hoserae sp. nov. and others in the species dorsal pattern is blotched and semi-distinct and there is group are like C. carinata (with which they are sometimes significant speckling and markings on the venter. sympatric) in having the intervestibular fenestra largely or Distribution: The islands of Kadavu and Ono in southern Fiji, in entirely filled in by expansion of the premaxilla. a region generally bounded by the deep water Kadavu Passage Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as in the north and the Great Astrolabe Reef to the east. defined in this paper, is the taxon from the main Fiji islands Etymology: Named in honour of Kim Moore of Brisbane, including Viti Levu and Vanua Levu and immediately adjacent Queensland, Australia, widow of Michael Moore (Pike), for her islands, but not including Kadavu or the islands in the Lau services to herpetology, including in association with the group, each of which has a different species. Herpetological Society of Queensland Incorporated, over some The taxon C. bibroni is separated from all other species within decades. the C. bibroni species complex by the following suite of CANDOIA MALCOLMMCLUREI SP. NOV. characters: A blotched dorsal pattern with distinctive throat Holotype: A preserved female specimen at the American stripes and a low ventral count ranging from 207-227 ventrals. Museum of Natural History, New York, USA, specimen number: The species C. bibroni is further defined by the following suite of AMNH 60468 from Mare Island in the Loyalty Islands, (territory characters: 18-22 Maxillary teeth, (versus 17-19 in C. woolfi sp. of New Caledonia). nov. from the Lau Group of islands), 53-59 subcaudals (versus 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- This is a facility that allows access to its holdings. 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the Paratypes: Two preserved female specimens at the American head at one head length behind the head (versus 33-34 in C. Museum of Natural History, New York, USA, specimen numbers: woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one AMNH 61681 and AMNH 61682 from Ouvea Island in the headlength anterior to the cloaca, hemipenis forks at subcaudal Loyalty Islands, (territory of New Caledonia), as well as a male, 6-10. specimen number: AMNH 61711 from the same place. The species Candoia woolfi sp. nov. from the Lau Group of Diagnosis: Candoia bibroni and other species described in this islands, east of the main Fiji islands is separated from all other paper that were formerly treated as populations of C. bibroni are species within the C. bibroni species complex by the following herein defined as being the C. bibroni species complex. suite of characters: A blotched pattern without stripes. The tail The C. bibroni species group are those species not included in pattern is much more conspicuous than that of body, the pale the C. hoserae sp. nov. species group, as defined below. bordering of body blotches also replacing darker ground color on All species within the C. bibroni species complex are separated the tail; 205-236 ventrals; there are 33-34 scale rows behind the from all other Candoia by having supralabials separated from head at one head length behind the head (versus 26-30 in C. the eye by the subocular scales, versus C. carinata (Schneider, bibroni). 1801) and C. paulsoni (Stull, 1956) which have them entering The species Candoia kimmooreae sp. nov. from the island of the eye; preocular differentiated; canthus rostralis rounded; a Kadavu, southern Fiji is is separated from all other species relatively slender body with lengthwise ridges on the body scales within the C. bibroni species complex by having a blotched that run parallel to the long axis of the body, versus a stout body pattern which includes a prominent median throat stripe and no with keels on the body scales forming oblique rows as in C. lateral throat stripes. Otherwise this taxon is essentially similar aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. in form to nominate C. bibroni as described herein. Furthermore the tail more than twice as long as the head, being The species Candoia malcolmmaclurei sp. nov. from the Loyalty capable of more than one complete circle of coiling in the Islands, (territory controlled by New Caledonia) is separated vertical plane. from all other species within the C. bibroni species complex by The species Candoia carinata (Schneider, 1801) including having a blotched pattern without stripes and a tail similar in species and subspecies formerly treated as being within this pattern to the body, or alternatively a blotched pattern with throat taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. stripes including a median row of dark spots on anterior ventrals; bibroni complex by having 2, or rarely 1 or 3 supralabials dark perocular streak from angle of mouth (or angle of jaw) entering eye; an angulate canthus rostralis, 160-202 ventrals through eye to loreal region, dark spots on lips, and dark and 35-60 subcaudals. spotting on top of head and with a median ventral row of dark The species Candoia aspera (Günther, 1877) including species spots. Sometimes the colouration is dulled to appear unicolour. and subspecies formerly treated as being within this taxon is The species is further defined by having 13-16 infralabials, 27- separated from the C. bibroni complex by having a tail less than 30 scale rows on the neck at one head length back, 20-22 scale twice as long as the head, usually being shorter than the head; rows at one headlength anterior to the cloaca, 219-226 ventrals, is incapable of forming a full circle of coiling in vertical plane; at 53-60 subcaudals and 19-22 maxillary teeth on either side. least on the anterior two-thirds of body are keels of scales The species Candoia boutrosi sp. nov. from the islands of forming curved diagonal ridges along the sides, which extend Samoa is separated from all other species within the C. bibroni backward and downward towards the belly; 127-153 ventrals; 11- species complex by the following unique suite of characters: a

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 55

distinctive blotched dorsal body pattern that is one or other of capable of more than one complete circle of coiling in the being distinct or very indistinct, the large dark blotches being the vertical plane. main component of the dorsal midline for the anterior and mid- The species Candoia carinata (Schneider, 1801) including body, with the lighter blotches dominating the midline posteriorly, species and subspecies formerly treated as being within this reversing on the tail to have the darker blotches dominate on the taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. tail, which remains brightly marked for the whole length. The bibroni complex by having 2, or rarely 1 or 3 supralabials dorsal colouration is a combination of chocolate brown and entering eye; an angulate canthus rostralis, 160-202 ventrals beige in distinctly marked animals or a hybrid of this in the and 35-60 subcaudals. indistinctly marked animals. On the head is a well-defined The species Candoia aspera (Günther, 1877) including species creamish bar on the upper labials above the jawline. and subspecies formerly treated as being within this taxon is It is also defined as having 12 supralabials bilaterally (in separated from the C. bibroni complex by having a tail less than common with the three species from the Fiji Islands), 15-16 twice as long as the head, usually being shorter than the head; infralabials (as for C. bibroni sensu stricto), and 31-32 scale is incapable of forming a full circle of coiling in vertical plane; at rows on the neck, one head length back from the head (versus least on the anterior two-thirds of body are keels of scales 26-30 in C. bibroni sensu stricto). forming curved diagonal ridges along the sides, which extend C. boutrosi sp. nov. is further defined and separated from the backward and downward towards the belly; 127-153 ventrals; 11- other species in the C. bibroni complex by the following suite of 22 subcaudals; no specially enlarged preocular; (supralabials characters 37-38 mid-body rows, 22-24 scale rows at one head excluded from the eye; canthus rostralis angular). length anterior to the cloaca, 237-252 ventrals, 51-61 Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. subcaudals and 16-18 maxillary teeth on either side. species group are defined as the three species from the The species Candoia niraikanukiwai sp. nov. from the islands of Solomon Islands, including outliers, as well as the single Wallis and Futuna, including nearby islets is separated from all species from the Banks Islands and nearby New Hebrides. other species within the C. bibroni species complex by the These species which are defined in detail already in this paper, following unique suite of characters: It is essentially similar in are all separated from C. bibroni and others in the C. bibroni most respects to C. boutrosi sp. nov. from which it differs by species group (being all others in the C. bibroni complex) by the having a distinctive well-defined stripe running through the eye form of the postorbital bone, which has its ventral (free) end on either side of the head and the absence of a well-defined abruptly flexed backward, as in C. carinata, rather than forming creamish bar on the upper labials above the jawline. a simple crescent arc as seen in C. bibroni. In C. bibroni the The species Candoia georgemacintyrei sp. nov. from the island premaxilla has the “typical boine” form described by Frazzetta of Rotuma, is separated from all other species within the C. (1959, 1975), with a large fenestra between the narial vestibules, bibroni species complex by the following unique suite of this fenestra bounded anteriorly by the premaxilla and posteriorly characters: 235-240 ventrals, 57-63 subcaudals, 37-39 mid by the nasals; but C. hoserae sp. nov. and others in the species body rows and significant lightening of the upper labials; the group are like C. carinata (with which they are sometimes dorsal pattern is blotched and semi-distinct and there is sympatric) in having the intervestibular fenestra largely or significant speckling and markings on the venter. entirely filled in by expansion of the premaxilla. Distribution: C. malcolmmclurei sp. nov. is known only from the Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as Loyalty Islands of Ouvea, Lifou and Mare. defined in this paper, is the taxon from the main Fiji islands Etymology: Named in honour of Malcolm McLure of near Yea, including Viti Levu and Vanua Levu and immediately adjacent Victoria, Australia, formerly of Elwood, Victoria, Australia, in islands, but not including Kadavu or the islands in the Lau recognition of his many public interest contributions to Australia, group, each of which has a different species. including his battles against illegally imposed road tolls in The taxon C. bibroni is separated from all other species within Australia, via his organisation UPMART, political corruption in the C. bibroni species complex by the following suite of Australia, including by exposing tax-evading corporations characters: A blotched dorsal pattern with distinctive throat avoiding prosecution as a result of corrupt and illegal cash stripes and a low ventral count ranging from 207-227 ventrals. donations to politicians. The species C. bibroni is further defined by the following suite of CANDOIA BOUTROSI SP. NOV. characters: 18-22 Maxillary teeth, (versus 17-19 in C. woolfi sp. Holotype: A preserved male specimen at the American nov. from the Lau Group of islands), 53-59 subcaudals (versus Museum of Natural History, New York, USA, specimen number: 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- AMNH 41742 collected on the island of Savaii, Western Samoa. 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the head at one head length behind the head (versus 33-34 in C. This is a facility that allows access to its holdings. woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one Paratype: A preserved female specimen at the American headlength anterior to the cloaca, hemipenis forks at subcaudal Museum of Natural History, New York, USA, specimen number: 6-10. AMNH 41743 collected at the island of Savaii, Western Samoa. The species Candoia woolfi sp. nov. from the Lau Group of Diagnosis: Candoia bibroni and other species described in this islands, east of the main Fiji islands is separated from all other paper that were formerly treated as populations of C. bibroni are species within the C. bibroni species complex by the following herein defined as being the C. bibroni species complex. suite of characters: A blotched pattern without stripes. The tail The C. bibroni species group are those species not included in pattern is much more conspicuous than that of body, the pale the C. hoserae sp. nov. species group, as defined below. bordering of body blotches also replacing darker ground color on All species within the C. bibroni species complex are separated the tail; 205-236 ventrals; there are 33-34 scale rows behind the from all other Candoia by having supralabials separated from head at one head length behind the head (versus 26-30 in C. the eye by the subocular scales, versus C. carinata (Schneider, bibroni). 1801) and C. paulsoni (Stull, 1956) which have them entering The species Candoia kimmooreae sp. nov. from the island of the eye; preocular differentiated; canthus rostralis rounded; a Kadavu, southern Fiji is is separated from all other species relatively slender body with lengthwise ridges on the body scales within the C. bibroni species complex by having a blotched that run parallel to the long axis of the body, versus a stout body pattern which includes a prominent median throat stripe and no with keels on the body scales forming oblique rows as in C. lateral throat stripes. Otherwise this taxon is essentially similar aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. in form to nominate C. bibroni as described herein. Furthermore the tail more than twice as long as the head, being The species Candoia malcolmmaclurei sp. nov. from the Loyalty

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Islands, (territory controlled by New Caledonia) is separated Diagnosis: Candoia bibroni and other species described in this from all other species within the C. bibroni species complex by paper that were formerly treated as populations of C. bibroni are having a blotched pattern without stripes and a tail similar in herein defined as being the C. bibroni species complex. pattern to the body, or alternatively a blotched pattern with throat The C. bibroni species group are those species not included in stripes including a median row of dark spots on anterior ventrals; the C. hoserae sp. nov. species group, as defined below. dark perocular streak from angle of mouth (or angle of jaw) All species within the C. bibroni species complex are separated through eye to loreal region, dark spots on lips, and dark from all other Candoia by having supralabials separated from spotting on top of head and with a median ventral row of dark the eye by the subocular scales, versus C. carinata (Schneider, spots. Sometimes the colouration is dulled to appear unicolour. 1801) and C. paulsoni (Stull, 1956) which have them entering The species is further defined by having 13-16 infralabials, 27- the eye; preocular differentiated; canthus rostralis rounded; a 30 scale rows on the neck at one head length back, 20-22 scale relatively slender body with lengthwise ridges on the body scales rows at one headlength anterior to the cloaca, 219-226 ventrals, that run parallel to the long axis of the body, versus a stout body 53-60 subcaudals and 19-22 maxillary teeth on either side. with keels on the body scales forming oblique rows as in C. The species Candoia boutrosi sp. nov. from the islands of aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. Samoa is separated from all other species within the C. bibroni Furthermore the tail more than twice as long as the head, being species complex by the following unique suite of characters: a capable of more than one complete circle of coiling in the distinctive blotched dorsal body pattern that is one or other of vertical plane. being distinct or very indistinct, the large dark blotches being the The species Candoia carinata (Schneider, 1801) including main component of the dorsal midline for the anterior and mid- species and subspecies formerly treated as being within this body, with the lighter blotches dominating the midline posteriorly, taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. reversing on the tail to have the darker blotches dominate on the bibroni complex by having 2, or rarely 1 or 3 supralabials tail, which remains brightly marked for the whole length. The entering eye; an angulate canthus rostralis, 160-202 ventrals dorsal colouration is a combination of chocolate brown and and 35-60 subcaudals. beige in distinctly marked animals or a hybrid of this in the indistinctly marked animals. On the head is a well-defined The species Candoia aspera (Günther, 1877) including species creamish bar on the upper labials above the jawline. and subspecies formerly treated as being within this taxon is separated from the C. bibroni complex by having a tail less than It is also defined as having 12 supralabials bilaterally (in twice as long as the head, usually being shorter than the head; common with the three species from the Fiji Islands), 15-16 is incapable of forming a full circle of coiling in vertical plane; at infralabials (as for C. bibroni sensu stricto), and 31-32 scale least on the anterior two-thirds of body are keels of scales rows on the neck, one head length back from the head (versus forming curved diagonal ridges along the sides, which extend 26-30 in C. bibroni sensu stricto). backward and downward towards the belly; 127-153 ventrals; 11- C. boutrosi sp. nov. is further defined and separated from the 22 subcaudals; no specially enlarged preocular; (supralabials other species in the C. bibroni complex by the following suite of excluded from the eye; canthus rostralis angular). characters 37-38 mid-body rows, 22-24 scale rows at one head Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. length anterior to the cloaca, 237-252 ventrals, 51-61 species group are defined as the three species from the subcaudals and 16-18 maxillary teeth on either side. Solomon Islands, including outliers, as well as the single The species Candoia niraikanukiwai sp. nov. from the islands of species from the Banks Islands and nearby New Hebrides. Wallis and Futuna, including nearby islets is separated from all These species which are defined in detail already in this paper, other species within the C. bibroni species complex by the are all separated from C. bibroni and others in the C. bibroni following unique suite of characters: It is essentially similar in species group (being all others in the C. bibroni complex) by the most respects to C. boutrosi sp. nov. from which it differs by form of the postorbital bone, which has its ventral (free) end having a distinctive well-defined stripe running through the eye abruptly flexed backward, as in C. carinata, rather than forming on either side of the head and the absence of a well-defined a simple crescent arc as seen in C. bibroni. In C. bibroni the creamish bar on the upper labials above the jawline. premaxilla has the “typical boine” form described by Frazzetta The species Candoia georgemacintyrei sp. nov. from the island (1959, 1975), with a large fenestra between the narial vestibules, of Rotuma, is separated from all other species within the C. this fenestra bounded anteriorly by the premaxilla and posteriorly bibroni species complex by the following unique suite of by the nasals; but C. hoserae sp. nov. and others in the species characters: 235-240 ventrals, 57-63 subcaudals, 37-39 mid group are like C. carinata (with which they are sometimes body rows and significant lightening of the upper labials; the sympatric) in having the intervestibular fenestra largely or dorsal pattern is blotched and semi-distinct and there is entirely filled in by expansion of the premaxilla. significant speckling and markings on the venter. Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as Distribution: C. boutrosi sp. nov. is known only from the islands defined in this paper, is the taxon from the main Fiji islands of Samoa. including Viti Levu and Vanua Levu and immediately adjacent Etymology: Named in honour of Steve Boutros, from islands, but not including Kadavu or the islands in the Lau Templestowe, Victoria, Australia in recognition of his many group, each of which has a different species. contributions to the administration of justice via the legal system The taxon C. bibroni is separated from all other species within in Australia and other public benefit activities. the C. bibroni species complex by the following suite of CANDOIA NIRAIKANUKIWAI SP. NOV. characters: A blotched dorsal pattern with distinctive throat Holotype: A preserved adult specimen at the French National stripes and a low ventral count ranging from 207-227 ventrals. Museum of Natural History, known in French as the Muséum The species C. bibroni is further defined by the following suite of national d’histoire naturelle, in Paris, France, specimen number: characters: 18-22 Maxillary teeth, (versus 17-19 in C. woolfi sp. MNHN 1986.690, collected at Point Vele, Futuna, within the nov. from the Lau Group of islands), 53-59 subcaudals (versus territory of Wallis and Futuna. 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- This is a facility that allows access to its holdings. 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the Paratype: A preserved specimen at the French National head at one head length behind the head (versus 33-34 in C. Museum of Natural History, known in French as the Muséum woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one national d’histoire naturelle, in Paris, France, specimen number: headlength anterior to the cloaca, hemipenis forks at subcaudal MNHN RA 1993.228, collected on the island of Alafi, within the 6-10. territory of Wallis and Futuna. The species Candoia woolfi sp. nov. from the Lau Group of

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 57

islands, east of the main Fiji islands is separated from all other George’s Park Orchards Service Centre in Ringwood, Victoria, species within the C. bibroni species complex by the following Australia formerly of Park Orchards, Victoria, Australia, for vital suite of characters: A blotched pattern without stripes. The tail logistical support to significant herpetological scientific research pattern is much more conspicuous than that of body, the pale projects and Snakebusters wildlife conservation displays for the bordering of body blotches also replacing darker ground color on best part of a decade. It is also fitting that a man of Pacific the tail; 205-236 ventrals; there are 33-34 scale rows behind the Island descent should have a Pacific Island species named in head at one head length behind the head (versus 26-30 in C. his honour. The days of a gross over-representation of bibroni). Eurocentric patronyms in Zoology should have ended long ago! The species Candoia kimmooreae sp. nov. from the island of CANDOIA GEORGEMACINTYREI SP. NOV. Kadavu, southern Fiji is is separated from all other species Holotype: A preserved specimen at the Natural History within the C. bibroni species complex by having a blotched Museum, London, UK, specimen number: 1897.7.29.13 pattern which includes a prominent median throat stripe and no collected at the island of Rotuma, (territory controlled by Fiji). lateral throat stripes. Otherwise this taxon is essentially similar This is a facility that allows access to its holdings. in form to nominate C. bibroni as described herein. Diagnosis: The species Candoia georgemacintyrei sp. nov. The species Candoia malcolmmaclurei sp. nov. from the Loyalty from the island of Rotuma, is separated from all other species Islands, (territory controlled by New Caledonia) is separated within the C. bibroni species complex by the following unique from all other species within the C. bibroni species complex by suite of characters: 235-240 ventrals, 57-63 subcaudals, 37-39 having a blotched pattern without stripes and a tail similar in mid body rows and significant lightening of the upper labials; the pattern to the body, or alternatively a blotched pattern with throat dorsal pattern is blotched and semi-distinct and there is stripes including a median row of dark spots on anterior ventrals; significant speckling and markings on the venter. dark perocular streak from angle of mouth (or angle of jaw) Candoia bibroni and other species described in this paper that through eye to loreal region, dark spots on lips, and dark were formerly treated as populations of C. bibroni are herein spotting on top of head and with a median ventral row of dark defined as being the C. bibroni species complex. spots. Sometimes the colouration is dulled to appear unicolour. The species is further defined by having 13-16 infralabials, 27- The C. bibroni species group are those species not included in 30 scale rows on the neck at one head length back, 20-22 scale the C. hoserae sp. nov. species group, as defined below. rows at one headlength anterior to the cloaca, 219-226 ventrals, All species within the C. bibroni species complex are separated 53-60 subcaudals and 19-22 maxillary teeth on either side. from all other Candoia by having supralabials separated from The species Candoia boutrosi sp. nov. from the islands of the eye by the subocular scales, versus C. carinata (Schneider, Samoa is separated from all other species within the C. bibroni 1801) and C. paulsoni (Stull, 1956) which have them entering species complex by the following unique suite of characters: a the eye; preocular differentiated; canthus rostralis rounded; a distinctive blotched dorsal body pattern that is one or other of relatively slender body with lengthwise ridges on the body scales being distinct or very indistinct, the large dark blotches being the that run parallel to the long axis of the body, versus a stout body main component of the dorsal midline for the anterior and mid- with keels on the body scales forming oblique rows as in C. body, with the lighter blotches dominating the midline posteriorly, aspera (Günther, 1877); ventrals 203-266; subcaudals 44-67. reversing on the tail to have the darker blotches dominate on the Furthermore the tail more than twice as long as the head, being tail, which remains brightly marked for the whole length. The capable of more than one complete circle of coiling in the dorsal colouration is a combination of chocolate brown and vertical plane. beige in distinctly marked animals or a hybrid of this in the The species Candoia carinata (Schneider, 1801) including indistinctly marked animals. On the head is a well-defined species and subspecies formerly treated as being within this creamish bar on the upper labials above the jawline. taxon (e.g. C. paulsoni Stull, 1956) is separated from the C. It is also defined as having 12 supralabials bilaterally (in bibroni complex by having 2, or rarely 1 or 3 supralabials common with the three species from the Fiji Islands), 15-16 entering eye; an angulate canthus rostralis, 160-202 ventrals infralabials (as for C. bibroni sensu stricto), and 31-32 scale and 35-60 subcaudals. rows on the neck, one head length back from the head (versus The species Candoia aspera (Günther, 1877) including species 26-30 in C. bibroni sensu stricto). and subspecies formerly treated as being within this taxon is C. boutrosi sp. nov. is further defined and separated from the separated from the C. bibroni complex by having a tail less than other species in the C. bibroni complex by the following suite of twice as long as the head, usually being shorter than the head; characters 37-38 mid-body rows, 22-24 scale rows at one head is incapable of forming a full circle of coiling in vertical plane; at length anterior to the cloaca, 237-252 ventrals, 51-61 least on the anterior two-thirds of body are keels of scales subcaudals and 16-18 maxillary teeth on either side. forming curved diagonal ridges along the sides, which extend backward and downward towards the belly; 127-153 ventrals; 11- The species Candoia niraikanukiwai sp. nov. from the islands of 22 subcaudals; no specially enlarged preocular; (supralabials Wallis and Futuna, including nearby islets is separated from all excluded from the eye; canthus rostralis angular). other species within the C. bibroni species complex by the following unique suite of characters: It is essentially similar in Candoia hoserae sp. nov. and others in the C. hoserae sp. nov. most respects to C. boutrosi sp. nov. from which it differs by species group are defined as the three species from the having a distinctive well-defined stripe running through the eye Solomon Islands, including outliers, as well as the single on either side of the head and the absence of a well-defined species from the Banks Islands and nearby New Hebrides. creamish bar on the upper labials above the jawline. These species which are defined in detail already in this paper, The species Candoia georgemacintyrei sp. nov. from the island are all separated from C. bibroni and others in the C. bibroni of Rotuma, is separated from all other species within the C. species group (being all others in the C. bibroni complex) by the bibroni species complex by the following unique suite of form of the postorbital bone, which has its ventral (free) end characters: 235-240 ventrals, 57-63 subcaudals, 37-39 mid abruptly flexed backward, as in C. carinata, rather than forming body rows and significant lightening of the upper labials; the a simple crescent arc as seen in C. bibroni. In C. bibroni the dorsal pattern is blotched and semi-distinct and there is premaxilla has the “typical boine” form described by Frazzetta significant speckling and markings on the venter. (1959, 1975), with a large fenestra between the narial vestibules, this fenestra bounded anteriorly by the premaxilla and posteriorly Distribution: Candoia niraikanukiwai sp. nov. is known only by the nasals; but C. hoserae sp. nov. and others in the species from the islands of Wallis and Futuna. group are like C. carinata (with which they are sometimes Etymology: Named in honour of Nirai Karatia Kanukiwa of sympatric) in having the intervestibular fenestra largely or

Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 58 Australasian Journal of Herpetology entirely filled in by expansion of the premaxilla. length anterior to the cloaca, 237-252 ventrals, 51-61 Candoia bibroni (Hombron and Jacquinot, 1853) sensu stricto as subcaudals and 16-18 maxillary teeth on either side. defined in this paper, is the taxon from the main Fiji islands The species Candoia niraikanukiwai sp. nov. from the islands of including Viti Levu and Vanua Levu and immediately adjacent Wallis and Futuna, including nearby islets is separated from all islands, but not including Kadavu or the islands in the Lau other species within the C. bibroni species complex by the group, each of which has a different species. following unique suite of characters: It is essentially similar in The taxon C. bibroni is separated from all other species within most respects to C. boutrosi sp. nov. from which it differs by the C. bibroni species complex by the following suite of having a distinctive well-defined stripe running through the eye characters: A blotched dorsal pattern with distinctive throat on either side of the head and the absence of a well-defined stripes and a low ventral count ranging from 207-227 ventrals. creamish bar on the upper labials above the jawline. The species C. bibroni is further defined by the following suite of Distribution: Candoia georgemacintyrei sp. nov. is restricted to characters: 18-22 Maxillary teeth, (versus 17-19 in C. woolfi sp. the island of Rotuma, controlled politically by Fiji. nov. from the Lau Group of islands), 53-59 subcaudals (versus Etymology: Named in honour of George Hayden McIntyre of 45-56 in C. woolfi sp. nov. from the Lau Group of islands), 10- George’s Park Orchards Service Centre in Ringwood, Victoria, 16 supralabials, 12-17 infralabials, 26-30 scale rows behind the Australia formerly of Park Orchards, Victoria, Australia, for vital head at one head length behind the head (versus 33-34 in C. logistical support to significant herpetological scientific research woolfi sp. nov.), 30-37 midbody rows, 19-24 scale rows one projects and Snakebusters wildlife conservation displays for the headlength anterior to the cloaca, hemipenis forks at subcaudal best part of a decade. 6-10. NOTES ON THE DESCRIPTIONS FOR ANY POTENTIAL The species Candoia woolfi sp. nov. from the Lau Group of REVISORS islands, east of the main Fiji islands is separated from all other Unless mandated by the rules of the International Code of species within the C. bibroni species complex by the following Zoological Nomenclature, none of the spellings of the newly suite of characters: A blotched pattern without stripes. The tail proposed names should be altered in any way. Should one or pattern is much more conspicuous than that of body, the pale more newly named taxa be merged by later authors to be bordering of body blotches also replacing darker ground color on treated as a single species, the order of prority of retention of the tail; 205-236 ventrals; there are 33-34 scale rows behind the names should be the order (page priority) of the formal head at one head length behind the head (versus 26-30 in C. descriptions within this text. bibroni). REFERENCES CITED The species Candoia kimmooreae sp. nov. from the island of Adler, G. H., Austin, C. C. and Dudley, R. 1995. Dispersal and Kadavu, southern Fiji is is separated from all other species speciation of skinks among archipelagos in the tropical Pacific within the C. bibroni species complex by having a blotched Ocean. Evolutionary Ecology 9:529-541. pattern which includes a prominent median throat stripe and no lateral throat stripes. Otherwise this taxon is essentially similar Austin, C. C. 2000. Molecular phylogeny and historical in form to nominate C. bibroni as described herein. biogeography of Pacific Island boas (Candoia). Copeia 2000(2):341-352. The species Candoia malcolmmaclurei sp. nov. from the Loyalty Islands, (territory controlled by New Caledonia) is separated Austin, C. C., Rittmeyer, E. N., Richards, S. J. and Zug, G. R. from all other species within the C. bibroni species complex by 2010. 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Diagnoses of new reptiles and It is also defined as having 12 supralabials bilaterally (in batrachians from the Solomon Islands, collected and presented common with the three species from the Fiji Islands), 15-16 to the British Museum by H. B. Guppy, Esq., M. B., H. M. S. infralabials (as for C. bibroni sensu stricto), and 31-32 scale ‘Lark.’. Proc. Zool. Soc. London 1884:210-213. rows on the neck, one head length back from the head (versus Boulenger, G. A. 1885. Catalogue of the Lizards in the British 26-30 in C. bibroni sensu stricto). Museum (Nat. Hist.) I. Geckonidae, Eublepharidae, Uroplatidae, C. boutrosi sp. nov. is further defined and separated from the Pygopodidae, Agamidae. London:450 pp. other species in the C. bibroni complex by the following suite of Boulenger, G. A. 1886. On the reptiles and batrachians of the characters 37-38 mid-body rows, 22-24 scale rows at one head Solomon Islands. Trans. Zool. Soc. London 12:35-62.

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Hoser 2016 - Australasian Journal of Herpetology 31:39-61. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 62 Australasian Journal of Herpetology Australasian Journal of Herpetology 31:62-63. ISSN 1836-5698 (Print) Published 1 August 2016. ISSN 1836-5779 (Online)

A new species of Denisonia from North-west Queensland, Australia (Serpentes: Elapidae).

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 3 September 2015, Accepted 19 September 2015, Published 1 August 2016.

ABSTRACT A new species of elapid snake of the genus Denisonia Krefft, 1869 from North West Queensland is formally described according to the rules of the International Code of Zoological Nomenclature. The species has for many years been treated as a population of the species Denisonia devisi Waite and Longman, 1920 or occasionally as aberrant D. maculata (Steindachner, 1867). Distributionally and physically, these snakes sit more-or-less between the two. Denisonia devisi is primarily an inhabitant of black soil river flats associated with the Darling River and Northern Lake Eyre basins in inland Eastern Australia. D. maculatus is an inhabitant of Eastward flowing drainages in the dry zone between the south-east and North east Queensland wet zones, principally the Fitzroy and Burdekin River systems. This newly described taxon, Denisonia gedyei sp. nov. is primarily an inhabitant of the Flinders River system, draining into the Gulf of Carpentaria and may be more widely distributed than current museum records indicate. Keywords: Taxonomy; Denisonia; new species; gedyei; Queensland; Australia; elapid; snake; devisi maculatus; Flinders River; Gulf of Carpentaria.

INTRODUCTION the dry zone between the south-east and North east Queensland As part of a study of Australian snakes spanning more than 40 wet zones, principally the Fitzroy and Burdekin River systems. years, the genus Denisonia Krefft, 1869 was scrutinized in This newly described taxon, Denisonia gedyei sp. nov. described detail. In the 1970’s I became the first person known to have below is primarily an inhabitant of the Flinders River system, bred them in captivity, this being the breeding of a pair of D. draining into the Gulf of Carpentaria and may be more widely devisi originally caught from north-west of Nevertire in western distributed than current museum records indicate. New South Wales. Names applied to the other two species of Denisonia as listed in Capture of specimens of both recognized species of Denisonia, Cogger et al. (1983) are not available for this newly described namely D. devisi Waite and Longman, 1920 and D. maculatus taxon. These unavailable names are, “Hoplocephalus ornatus” (Steindachner, 1867) by myself and others indicated regional De Vis (1884), “Denisonia ornata” Krefft (1869) and variation. This appeared to be mainly of a clinal nature. “Hoplocephalus muelleri” Fischer (1885). However specimens from the region between Mount Isa and Were it not for the distributional differences between the known Charters Towers in Queensland appeared to be intermediate populations (in separated drainage basins) and the knowledge between the two recognized species and no clines between that Denisonia is a uniquely distinctive genus of snake, both these and the other two recognized taxa were known. morphologically, as seen by the account in Cogger (2014), or by D. devisi is found south of this area, while D. maculatus is found phylogeny e.g. Pyron et al. (2013), a strong case could be to the east and south-east. mounted to treat all Denisonia as belonging to a single and As recently as 2014, Cogger (2014) stated that the only obvious variable species, with the three relevant populations being difference between the two species was the presence or treated as geographical subspecies. absence of bands on the body of either taxon, although the While it is possible that a molecular study of population samples bands on aged specimens of D. devisi are sometimes hard to from across the range of the genus may yield cryptic species, detect, but appear to always be present. Scalation and other these being most likely in populations of D. devisi, it is clear that characteristics were identical for both species, although Cogger a third form from North-west Queensland is sufficiently distinct (2014) reported average maximum size of the species being as to warrant being treated as its own taxonomic entity. marginally different, but of no diagnostic help in separating the Being distributionally and reproductively isolated from the other taxa. two recognized species and easily diagnosed and separated As mentioned in the abstract, Denisonia devisi is primarily an from the others on the basis of colouration and patterning, with inhabitant of black soil river flats associated with the Darling no known intermediates, in spite of inspection of specimens River and Northern Lake Eyre basins in inland Eastern Australia. from close localities on the relevant boundaries for each form, I D. maculatus is an inhabitant of Eastward flowing drainages in have no hesitation in formally naming this taxon as a new Hoser 2016 - Australasian Journal of Herpetology 31:62-63. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 63

species, namely Denisonia gedyei sp. nov.. devisi, while those from the Burdekin system appear to conform The formal description follows. with typical D. maculata. As a passing note, I should add that further survey work will ETYMOLOGY AND RELEVANT NOMENCLATURAL almost certainly yield further populations of D. devisi in parts of COMMENTS the Lake Eyre drainage basin, in areas where to date none have Named in honour of Andrew Gedye, in recognition of his been seen or collected, as well as parts of the Murray Darling excellent work with reptiles spanning many decades. His main basin, that have yet to see specimens collected. activity has been in the captive breeding of many rare and DENISONIA GEDYEI SP. NOV. potentially threatened species as well as many months of extensive fieldwork in all parts of mainland Australia. Holotype: A preserved specimen at the Museum and Art Gallery of the Northern Territory, Reptile Collection, catalogue number He currently lives in a suburb of Cairns, Queensland. R36392, from Julia Creek in north-west Queensland, Latitude - The subgenus Geddykukrius Hoser, 2012 was also named after 20.667, Longitude 141.633. Andrew Gedye, and herein as first reviser and original author, I The Museum and Art Gallery of the Northern Territory is a note the following: The spelling of Andrew’s surname Gedye was publicly owned facility that allows access to specimens. incorrect in that paper. Notwithstanding this and in order to Diagnosis: Denisonia gedyei sp. nov. is most similar to the two maintain stability of nomenclature, the nomen Geddykukrius currently recognized species of Denisonia as defined by Cogger Hoser, 2012 should retain the original spelling as in the original (2014) and matches the genus diagnosis in that text. paper. Denisonia gedyei sp. nov. is separated from both other species Similarly in 2014, I named a taxon Broghammerus reticulatus in that genus (Denisonia) by the following characters: mandella subsp. nov., repeatedly mis-spelling the name Nelson There is banding or similar on the neck and/or forebody, and well Mandela and/or as a result the patronym mandella. past the nape, but not on the majority of the body or tail which is I thank the taxonomic vandal, thief and law-breaker Mark clearly unmarked. In D. maculata there is no banding or similar O’Shea for pointing out this error on numerous online forums at all on the neck or body beyond the nape. In D. devisi, there is and while much of what Mr. OShea has done is beneath banding or similar along the length of the entire body, which may contempt and most of what he has written and said about me or may not include the tail, noting that the tail will invariably have has been totally untrue, he was within his rights to point out my at least a few patches of dark pigment on the dorsal surface, error in this case and I thank him for this. which is not seen in D. gedyei sp. nov. or D. maculata. Post publication peer (and non-peer) review is as important as Within the constraints of the above, the three relevant species that which occurs prior to publication and accuracy of are characterised and separated from one another as follows: publications and noting of any errors are both paramount goals which should not be subsumed or ignored on the basis of ego, D. gedyei sp. nov. has from 2 to 12 distinctive bands on the or an author foolishly refusing to admit error when one has been upper neck region, which fade to merge into a dorsal body committed. pattern of one colour, characterised by being a brownish colour, Notwithstanding the above, and while acknowledging the with each scale containing a distinctive dark brown or black spelling errors in the first instance, for the same reasons as patch in the anterior section. The tail is distinctly lighter. given for Geddykukrius Hoser, 2012, and invoking the same D. maculata does not have 2-12 distinctive bands on the upper neck and while scales may be darker anteriorly than posteriorly, rights as per the International Code of Zoological Nomenclature, this is neither obvious or distinctive, in that the body appears at the nomen spelling Broghammerus reticulatus mandella Hoser, a glance to be of one colour only. 2014 should not have the original spelling changed. D. devisi has a dorsal body pattern consisting of fairly distinct This paper may be cited as the basis for retention of spelling in bands or broken bands along the entire length of the body. In both cases. some specimens, the bands become skewed to give another REFERENCES CITED form of pattern, but the distinctive body pattern, as opposed to Cogger, H. G., Cameron, E. E. and Cogger, H. M. 1983. the one colour body scheme of the other two species remains. Zoological Catalogue of Australia (1): Amphibia and Reptilia. This remains the case even in faded and older specimens (e.g. Aust. Gov. Publishing Service, Canberra, ACT, Australia:313 pp. as seen on page 85 of Emmott and Wilson 2009). Cogger, H. G. 2014. Reptiles and Amphibians of Australia D. gedyei sp. nov. has distinctive white barring of the upper (Seventh edition), CSIRO. Sydney, Australia:1064 pp. labials, a trait sometimes, but not always seen in the other two Emmott, A. and Wilson, S. G. 2009. Snakes of Western species. Queensland: A field guide. Desert Channels Queensland, D. gedyei sp. nov. and D. maculata both posess a tail that Australia:136 pp. dorsally is distinctly lighter in colour than the adjacent body. Hoser, R. T. 1989. Australian Reptiles and Frogs, Pierson and While this sometimes occurs in D. devisi, the latter taxon is Co., Mosman, NSW, Australia:239 pp. separated from the other two by patches of dark pigment on a Pyron, R. A., Burbrink, F. T. and Wiens, J. J. 2013. A phylogeny lighter background, versus a one colour tail in the other two (see and revised classification of Squamata, including 4161 species typical D. devisi at top of page 145 of Hoser 1989). of lizards and snakes. BMC Evolutionary Biology 13:93. Distribution: Known only from the vicinity of Julia Creek, Ride, W. D. L. (ed.) et al. (on behalf of the International Richmond and Hughenden in mid-north-west Queensland, Commission on Zoological Nomenclature) 1999. International Australia in the upper reaches of the Flinders River system code of Zoological Nomenclature (Fourth edition). The Natural where it is found immediately in or adjacent to watercourses History Museum - Cromwell Road, London SW7 5BD, UK. either under cover by day or active at night. This system drains Waite, E. R. and Longman, H. A. 1920. Description of little- into the Gulf of Carpentaria and there is a lot of potentially known Australian snakes. Records of the South Australian suiable habitat for this taxon north of where it is currently known Museum 1:173-180. from. I recommend that fieldwork be conducted in the relevant areas to determine the presence or absence of the taxon here. Steindachner, F. 1867. in: Reise der Österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858,1859 unter den That Denisonia gedyei sp. nov. appears to be confined to Befehlen des Commodore B. von Wüllerstorf-Urbair (Zoologie), drainages associated with the Flinders River system was Vol. 1, part 3 (Reptilien p.1-98). K. Gerold’s Sohn/Kaiserlich- confirmed via inspection of specimens from nearby major Königl. Hof- und Staatsdruckerei, Wien [1869 on title page] drainages, these being the upper Darling system, upper lake Eyre drainages and the Burdekin/Fitzroy River systems. CONFLICT OF INTEREST Specimens from near Winton appear to conform with typical D. The author has no relevant conflicts of interest. Hoser 2016 - Australasian Journal of Herpetology 31:62-63. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 64 Australasian Journal of Herpetology ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

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