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An Evaluation of the Nomina for Death Adders (Acanthophis Daudin, 1803) Proposed by Wells & Wellington (1985), and Confirmation of A

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An Evaluation of the Nomina for Death Adders (Acanthophis Daudin, 1803) Proposed by Wells & Wellington (1985), and Confirmation of A Zootaxa 4995 (1): 161–172 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2021 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4995.1.9 http://zoobank.org/urn:lsid:zoobank.org:pub:959FF3A5-63AD-496D-AB24-B704C998B8FF An evaluation of the nomina for death adders (Acanthophis Daudin, 1803) proposed by Wells & Wellington (1985), and confirmation of A. cryptamydros Maddock et al., 2015 as the valid name for the Kimberley death adder RYAN J. ELLIS1,2, HINRICH KAISER3, SIMON T. MADDOCK4,5,6, PAUL DOUGHTY1 & WOLFGANG WÜSTER7,* 1Collections & Research, Western Australian Museum, Welshpool WA 6016, Australia https://orcid.org/0000-0003-0631-0571 2Biologic Environmental Survey, 24–26 Wickham St, East Perth, Western Australia 6004, Australia. https://orcid.org/0000-0002-9272-8204 3Department of Vertebrate Zoology, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; and Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA https://orcid.org/0000-0002-0001-9428 4Faculty of Science and Engineering, University of Wolverhampton, Wolverhampton, WV1 1LY, United Kingdom https://orcid.org/0000-0002-5455-6990 5Department of Life Sciences, The Natural History Museum, London, SW7 5BD, United Kingdom 6Island Biodiversity and Conservation Centre, University of Seychelles, Mahé, Seychelles 7 Molecular Biology and Evolution at Bangor, School of Natural Sciences, Bangor University, Bangor, LL57 2UW, United Kingdom *Corresponding author. [email protected]; https://orcid.org/0000-0002-4890-4311 Abstract We assess the availability of four names proposed by Wells & Wellington (1985) for Australian death adders (Acanthophis). In agreement with previous literature, A. hawkei is an available name, whereas A. armstrongi, A. lancasteri, and A. schistos are not described in conformity with the requirements of Articles 13.1.1 or 13.1.2 of the International Code of Zoological Nomenclature and are therefore considered nomina nuda. Consequently, A. cryptamydros Maddock et al., 2015, is confirmed as the valid name for the Kimberley death adder of Western Australia. We comment on the need for greater clarity in the Code, and emphasise that the responsibility for establishing the availability of new nomina rests with their authors, not subsequent researchers. Key words: Nomenclature, taxonomy, nomen nudum, Australia, Reptilia, Serpentes, Elapidae Introduction Death adders (genus Acanthophis Daudin, 1803) are a group of uniquely specialised Australasian elapid snakes best known for their morphological and ecological similarity to vipers (Shine 1980; Greene 1997; Shine et al. 2014). The species-level taxonomy of the genus has remained poorly understood, in part as a consequence of contributions by authors operating outside the peer-reviewed scientific literature (see Aplin 1999; Aplin & Donnellan 1999). Among these, Wells & Wellington (1985) proposed four new names for putative species of Acanthophis: A. armstrongi, A. hawkei, A. lancasteri, and A. schistos. None of the four descriptions provided any morphological or other descrip- tive information associated with the designated holotype specimens, and largely referred to the published works of other authors for diagnoses and descriptions, including specimen images. While these descriptions were initially ignored in the subsequent literature, the confirmation of considerable cryptic diversity in death adders (Aplin & Donnellan 1999; Wüster et al. 2005; Maddock et al. 2015) brought these names back into contention as potentially available nomina. This has resulted in taxonomic confusion and nomenclatural instability, particularly in the case of northern Australian Acanthophis, which had long been considered part of A. praelongus Ramsay, 1877 (e.g., Storr 1981; Cogger 1983, 2000; Ehmann 1992; Storr et al. 2002; Wilson & Swan 2003, 2013). Accepted by V. Deepak: 8 Jun. 2021; published: 29 Jun. 2021 161 Licensed under Creative Commons Attribution-N.C. 4.0 International https://creativecommons.org/licenses/by-nc/4.0/ Wells & Wellington (1985) named A. lancasteri from the Kimberley region of Western Australia (WA) and the ‘Top End’ of the Northern Territory (NT). Several authors (Shea 1987; Aplin 1999; Aplin & Donnellan 1999; Maddock et al. 2015; Wüster, 2021) have explicitly considered A. lancasteri a nomen nudum: the diagnosis was outsourced to Storr (1981), but in these authors’ view, Storr did not provide diagnoses compliant with Article 13 of the International Code of Zoological Nomenclature (hereafter, the Code; ICZN 1999), making the species epithet lancasteri unavailable. Shea (1987: 258) identified A. lancasteri as one of the many species described by Wells & Wellington (1985), for which “the attempted diagnosis of new species on the basis of previously published photo- graphs or previously published descriptions [does] not provide a statement in words of characters that are purported to differentiate the taxon (Article 13a (i–ii),” and he further stated (p. 259) that “In a few of these cases, the publica- tion cited provides a diagnosis of the taxon in question as part of a redescription. However, in none of the instances cited above does such a diagnosis differentiate between the population described and the typical population.” Since the original publication of A. lancasteri, the name had not been used as valid in the scientific herpeto- logical literature (i.e., in peer-reviewed publications; Kaiser et al., 2013) until a plea for acceptance by Wellington (2016). The populations concerned were generally included in A. praelongus (e.g., Storr et al. 1986; Cogger 1986, 1992, 1996, 2000; Ehmann 1992; Storr et al. 2002; Wilson & Swan 2003, 2008, 2010, 2013) or, after the phylogeo- graphic study of Wüster et al. (2005), in A. rugosus by some (e.g., Eipper 2012; Cogger 2014). Maddock et al. (2015) found that death adders from the Kimberley region of WA constitute a separate species from the populations of the A. rugosus complex (sensu Wüster et al. 2005: 11) in the NT and elsewhere. Noting the status of A. lancasteri as a nomen nudum, they described this taxon as a new species, A. cryptamydros Maddock et al., 2015. However, Wellington (2016) argued that, contrary to Shea (1987) and Aplin (1999), Storr’s (1981) descrip- tion should be considered sufficient as the primary diagnosis to make the name A. lancasteri available, which would relegate A. cryptamydros to the status of its junior synonym. The result of this dispute is that the nomenclature of the Kimberley death adder is currently split in the scientific literature, with some sources, primarily field guides (Swan 2017; Wilson & Swan 2017, 2020; Cogger 2018; Eipper & Eipper 2019), regarding A. lancasteri as the valid name, whereas others (White 2016; Ellis 2017; Mirtschin et al. 2017; WHO 2017; Chapple et al. 2019; Altherr & Lame- ter 2020; Boundy 2020; de Solan et al. 2020; Wüster 2021) use the younger name A. cryptamydros. Databases are similarly divided: the Australian Reptile Online Database (Macdonald 2021), the Integrated Taxonomic Information System (ITIS 2020), the Reptile Database (Uetz et al. 2020), the International Union for the Conservation of Nature (Cogger et al. 2017), and the Western Australian Naturemap (DBCA 2020) are using A. cryptamydros, whereas the Australian Faunal Directory (ABRS 2020) uses A. lancasteri. Settling the status of contentious names requires careful analysis to assess their availability under the Code (e.g., Iverson et al. 2001). To alleviate the regrettable confusion over the scientific name of the Kimberley death adder, in the following paragraphs we analyse in detail the availability and validity of the nomen A. lancasteri under the Code. To further clarify the nomenclature of the genus, we also reanalyse the availability of the other three death adder names coined by Wells & Wellington (1985): A. armstrongi, A. hawkei, and A. schistos. The assessment of these names raises questions as to the applicable version of the Code. Wells & Wellington (1999) claimed that their 1985 paper may have predated the third edition of the Code. The issue of nomina coined by Wells & Wellington (1985) thus potentially extends to three editions of the Code, published in 1964, 1984, and 1999. However, as is clear from Article 86.3, older editions of the Code “have no force unless reaffirmed” in the latest edition, and only as specified in that edition. Moreover, in any case, a review of all three editions of the Code shows their equivalence with respect to the current Article 13 and its application to this case. Our discussion herein is therefore based on the text of the current, fourth edition of the Code (ICZN 1999). Article 13 of the Code The key article of the Code that governs all four names coined by Wells & Wellington (1985) is Article 13. This article is augmented by Recommendation 13A, which is not binding but serves to further explain the intentions of the article itself. The article states (words printed in bold, italic text indicate our emphasis): “Article 13. Names published after 1930. 13.1. Requirements. To be available, every new name published after 1930 must satisfy the provisions of Article 11 and must 162 · Zootaxa 4995 (1) © 2021 Magnolia Press ELLIS ET AL. 13.1.1. be accompanied by a description or definition that states in words characters that are purported to differentiate the taxon, or 13.1.2. be accompanied by a bibliographic reference to such a published
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