Plagiarism Checker X Originality Report Similarity Found: 5% Date: Saturday, July 21, 2018 Statistics: 1626 words Plagiarized / 35967 Total words Remarks: Low Plagiarism Detected - Your Document needs Optional Improvement. ------------------------------------------------------------------------------------------- 1 Integrative taxonomy of a new and highly-diverse genus of onchidiid slugs from the Coral Triangle (Gastropoda, Pulmonata, Onchidiidae) Tricia C. Goulding1,2, Munawar Khalil3, Shau Hwai Tan4, Benoît Dayrat1 1 Department of Biology, Pennsylvania State University, University Park, PA 16802, USA 2 Current address: Bernice Pauahi Bishop Museum, 1525 Bernice St, Honolulu, HI 96817 3 Department of Marine Science, Universitas Malikussaleh. Reuleut Main Campus, Kecamatan Muara Batu, North Aceh, Aceh, 24355, Indo - nesia 4 Marine Science Laboratory, School of Biological Sciences, and Centre for Marine and Coastal Studies, Universiti Sains Malaysia, 11800 Penang, Malaysia Corresponding author: Benoît Dayrat ([email protected]) Academic editor: N. Yonow | Received 27 September 2017 | Accepted 8 May 2018 | Published 4 June 2018 http://zoobank.org/90B77255-4C5E-436C-A793-D924892B5B14 Citation: Goulding TC, Khalil M, Tan SH, Dayrat B (2018) Integrative taxonomy of a new and highly-diverse genus of onchidiid slugs from the Coral Triangle (Gastropoda, Pulmonata, Onchidiidae). ZooKeys 763: 1–111. https://doi. org/10.3897/zookeys.763.21252 Abstract A new genus of onchidiid slugs, Wallaconchis Goulding & Dayrat, gen. n., is described, including ten species. Five species were previously described but known only from the type material: Wallaconchis ater (Lesson, 1830), W. graniferum (Semper, 1880), W. nangkauriense (Plate, 1893), W. buetschlii (Stantschinsky, 1907), and W. gracile (Stantschinsky, 1907), all of which were originally classified in Onchidium Buchannan, 1800. Many new records are provided for these five species, which greatly expand their known geographic distributions. Five species are new: Wallaconchis achleitneri Goulding, sp. n., W. comendadori Goulding & Dayrat, sp. n., W. melanesiensis Goulding & Dayrat, sp. n., W. sinanui Goulding & Dayrat, sp. n., and W. uncinus Goulding & Dayrat, sp. n. Nine of the ten Wallaconchis species are found in the Coral Triangle (eastern Indonesia and the Philippines). Sympatry is high, with up to six species found on the island of Bohol (Philippines) and eight species overlapping in northern Sulawesi (Indonesia). Wallaconchis is distin- guished from other onchidiids by its bright dorsal colors (red, yellow, orange) but those are extremely vari - able and not useful for specific identification. Internally, the reproductive system can be used to identify all Wallaconchis species. The copulatory organs of Wallaconchis species are especially diverse compared to other onchidiid genera, and the possible role of reproductive incompatibility in species diversification is discussed. All specimens examined were freshly collected for the purpose of a worldwide revision of the Onchidiidae Rafinesque, 1815. The species are well delineated using DNA sequences and comparative anatomy. Mito - chondrial DNA analysis yields thirteen molecular units separated by a large barcode gap, while nuclear DNA ZooKeys 763: 1–111 (2018) doi: 10.3897/zookeys.763.21252 http://zookeys.pensoft.net Copyright Tricia C. Goulding et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. MONOGRAPH Launched to accelerate biodiversity research A peer-reviewed open-access journal 2 yields nine units. By integrating nuclear DNA and mitochondrial DNA with morphology, ten species are recognized. The natural history of each species (e.g., the microhabitat where they are found) is also docu- mented. Nomenclature is addressed thoroughly (the types of all onchidiid species were examined, lectotypes were designated when needed, nomina dubia are discussed). Morphological characters, transitions to new microhabitats, and diversification processes are discussed in the context of a robust molecular phylogeny. Keywords Euthyneura, Indo-West Pacific, marine biogeography, phylogeography, systematics Table of contents Introduction .............................................................................................................3 Materials and methods ............................................................................................. 4 Collecting ............................................................................................................ 4 Specimens ............................................................................................................ 5 Anatomical preparations and descriptions ............................................................ 5 DNA extraction and PCR amplification ............................................................... 7 Phylogenetic analyses ........................................................................................... 7 Phylogenetic results ................................................................................................13 Molecular phylogenetic analyses .........................................................................13 Systematics and anatomical descriptions .................................................................22 Genus Wallaconchis Goulding & Dayrat, gen. n. ..............................................22 Wallaconchis sinanui Goulding & Dayrat, sp. n. .........................................25 Wallaconchis uncinus Goulding & Dayrat, sp. n.......................................... 33 Wallaconchis buetschlii (Stantschinsky, 1907), comb. n. .............................. 42 Wallaconchis gracile (Stantschinsky, 1907), comb. n. ...................................49 Wallaconchis nangkauriense (Plate, 1893), comb. n. .................................... 56 Wallaconchis ater (Lesson, 1830), comb. n. .................................................63 Wallaconchis graniferum (Semper, 1880), comb. n. ..................................... 76 Wallaconchis achleitneri Goulding, sp. n. .................................................... 84 Wallaconchis comendadori Goulding & Dayrat, sp. n. .................................88 Wallaconchis melanesiensis Goulding & Dayrat, sp. n. .................................93 Identification key ........................................................................................... 100 Discussion ............................................................................................................100 Higher relationships .......................................................................................100 Species delineation ......................................................................................... 101 Species diversity ............................................................................................. 102 Nomenclature ................................................................................................ 104 Diversification: transitions to vastly different microhabitats ............................104 Diversification: the role of reproductive isolation ........................................... 106 Endemism and conservation .......................................................................... 107 Acknowledgements ...............................................................................................108 References ............................................................................................................ 108 3 Introduction The Onchidiidae is a group of shell-less, marine, air-breathing slugs that live in in - tertidal habitats worldwide. Most of the diversity is found in mangroves of South- East Asia, where onchidiids have diversified into several genera, such as Onchidium Buchannan, 1800, Peronina Plate, 1893, Platevindex Baker, 1938, and Melayonchis Dayrat & Goulding, 2017. The genus Peronia Fleming, 1822, which can easily be identified externally by branched gills on the dorsal notum, has diversified in the rocky intertidal across the Indo-West Pacific. However, for decades the diversity of onchidiid slugs has remained poorly known for a variety of reasons: mangroves have rarely been explored; preserved slugs can hardly be distinguished externally; and tax - onomists have avoided the overwhelming nomenclature (Dayrat 2009). As a result, most species could not be reliably identified even though many species are geographi- cally widespread and common. The Dayrat lab is currently engaged in a global revision of the Onchidiidae, based on an integrative approach involving extensive fieldwork (sampling thousands of individuals at more than 300 stations worldwide), natural history observations, comparative anatomy, and DNA sequencing. An important component of integra - tive taxonomy is sound nomenclature; this revision has required the examination of the type material of all onchidiid species, the detailed analysis of all species descrip - tions, and the re-evaluation of the nomenclatural status of every genus- and species- group name. First, monophyletic genera had to be delineated, and now each genus is being revised separately (Dayrat et al. 2016, 2017; Dayrat and Goulding 2017; Goulding et al. in press). Here, a new genus, Wallaconchis Goulding & Dayrat, gen. n., is described from the Indo-West Pacific, with a geographic