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Crustacea: Isopoda: Oniscidea) from Neotropical Region Canadian Journal of Zoology Integrative taxonomy reveals a new genus and new species of Philosciidae (Crustacea: Isopoda: Oniscidea) from Neotropical region Journal: Canadian Journal of Zoology Manuscript ID cjz-2017-0289.R1 Manuscript Type: Article Date Submitted by the Author: 22-Jan-2018 Complete List of Authors: Zimmermann, Bianca; Universidade Federal de Santa Maria Soares Campos Filho, Ivanklin; Universidade Federal de Campina Grande Araujo, Paula;Draft Universidade Federal do Rio Grande do Sul Atlantoscia, Brazilian Atlantic Forest, isopods phylogeny, Neotropics, Keyword: Paratlantoscia gen. nov., Paratlantoscia robusta sp. nov. https://mc06.manuscriptcentral.com/cjz-pubs Page 1 of 38 Canadian Journal of Zoology 1 Integrative taxonomy reveals a new genus and new species of Philosciidae (Crustacea: Isopoda: Oniscidea) from Neotropical region Bianca L. Zimmermann1, Ivanklin S. Campos-Filho2, and Paula B. Araujo3 1 Centro de Ciências Naturais e Exatas, Departamento de Ecologia e Evolução, Universidade Federal de Santa Maria, Av. Roraima, 1000, Cidade Universitária, Bairro Camobi, 97105-900, Santa Maria, Brazil. Fone/Fax: +55 55 3220 8465. 2 Programa de Pós-Graduação em Recursos Naturais, Universidade Federal de Campina Grande, Av. Aprígio Veloso, 882, Bairro universitário, 58429-140, Campina Grande, Brazil. Email: [email protected] Draft 3 Instituto de Biociências, Departamento de Zoologia, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves 9500, Bairro Agronomia, 91501-970, Porto Alegre, Brazil. Email: [email protected] Corresponding author. B.L. Zimmermann (email: [email protected]). https://mc06.manuscriptcentral.com/cjz-pubs Canadian Journal of Zoology Page 2 of 38 2 Integrative taxonomy reveals a new genus and new species of Philosciidae (Crustacea: Isopoda: Oniscidea) from Neotropical region. B.L. Zimmermann, I.S. Campos-Filho, and P.B. Araujo. Abstract: Although new methods and data are conquering space in the field of taxonomy, such as integrative taxonomy, most of the terrestrial isopod species are still described based only on morphology. Species of the genus Atlantoscia Ferrara & Taiti, 1981 were the first and are the unique terrestrial isopods from the Neotropics for which a molecular phylogeny was already conducted. Previous results indicated thatDraft this genus could be paraphyletic, and a more detailed analysis would be required. Our aim was to reconstruct the phylogeny of Atlantoscia using mitochondrial and nuclear markers and test its monophyly by integrating molecular and morphological data. We observed that, indeed, Atlantoscia is paraphyletic. Atlantoscia ituberasensis Campos-Filho, Lisboa & Araujo, 2013 and Atlantoscia rubromarginata Araujo & Leistikow, 1999 were placed in a new genus of terrestrial isopods, Paratlantoscia gen. nov., together with a new species described in the present study, Paratlantoscia robusta sp. nov. The new genus is defined by the presence of specialized respiratory areas in the pleopod exopods and its validity is highly corroborated by molecular analyses and by biogeographic information. This study highlights the importance of multiple and complementary perspectives as a way to improve the quality of species hypothesis and associated descriptions. Keywords: Atlantoscia, Brazilian Atlantic Forest, isopods phylogeny, Neotropics, Paratlantoscia gen. nov., Paratlantoscia robusta sp. nov., https://mc06.manuscriptcentral.com/cjz-pubs Page 3 of 38 Canadian Journal of Zoology 3 Introduction Species are a cornerstone of biology, ecology, and conservation (Petit and Excoffier 2009). Hence, identifying the boundaries between a set of species has been a largely discussed issue in the last years in the field of systematic biology (e.g., De Queiroz 2007; Knowles and Carstens 2007; Aguilar et al. 2013; Grummer et al. 2014). Due to practical and historical reasons most species were primarily described based on morphology for most part of our history (Padial et al. 2010). Nonetheless, new methods and data are conquering space in taxonomy. For example, assigning groups of organisms to species is one of the most challenging aspects of phylogenetic studies (Coyne and Orr 2004). One of these new approaches is integrativeDraft taxonomy. It integrates information from different sources (usually from morphological and molecular data) to delimit and describe taxa. The result is improved rigor in species delimitation, which is its ultimate goal as well (Marshall et al. 2006; Schilick-Steiner et al. 2010; Castalanelli et al. 2017). Integrative taxonomy has gained many supporters in recent years, especially because this approach has been proved to be effective in solving systematic issues in a variety of groups (e.g., Castro-Romero et al. 2016; Decker et al. 2016; Ekimova et al. 2016; Hernandez-Orts et al. 2017; Mazancourt et al. 2017), thus improving the quality of species hypothesis and associated descriptions (Pante et al. 2015). In this context, terrestrial isopods (Oniscidea) are an interesting group, since the majority of the species are still described based only on morphology, even though in many cases morphological characters do not provide a clear taxonomic resolution (Schmalfuss 2003). Terrestrial isopods comprise the most successful colonizers of terrestrial environments among crustaceans and show morphological, physiological, and behavioral adaptations to this lifestyle (Richardson and Araujo 2015). For example, the development of the respiratory structures on the pleopod exopods was a key factor in https://mc06.manuscriptcentral.com/cjz-pubs Canadian Journal of Zoology Page 4 of 38 4 the success of terrestrial colonization in woodlice. Variations in these structures represent both convergence and functional constraints. The complexity of respiratory structures is generally linked to different levels of environment adaptation (Richardson and Araujo 2015) and could potentially be used in the species delimitation.. Among the main representatives of terrestrial isopods we can highlight the family Philosciidae Kinahan, 1857, one of the most important groups in tropical and wetlands habitats (Leistikow 2001; Schmalfuss 2003; Campos-Filho et al. 2014). Within Philosciidae, the genus Atlantoscia Ferrara & Taiti, 1981 consists of seven described species, mainly recorded in Brazil, except for the widely distributed species A. floridana Van Name, 1940 (Campos-Filho et al. 2013). Members of this genus differ in few morphological characteristics and are defined based mainly on the shape of the male pleopods (Campos-Filho et al. 2013).Draft Recently, Zimmermann et al. (2015) tried to overcome these taxonomic problems within Atlantoscia. The authors demonstrated that using integrative taxonomy by coupling morphology and DNA sequences is an efficient strategy for delimitation of Atlantoscia species and to investigate their relationships. However, some issues still persist For instance, Atlantoscia rubromarginata Araujo & Leistikow, 1999 and Atlantoscia ituberasensis Campos-Filho, Lisboa & Araujo, 2013 are quite different from the other congeneric species and thus might not belong to this genus (Zimmermann et al. 2015). Specifically, they are genetically and geographically distant and possess unique and specialized morphologies, with well developed respiratory areas in the pleopods (Campos-Filho et al. 2013). Taking into account that populations that are considered to be candidate species will typically exhibit some evidence of genetic isolation, either in the form of phenotypic/genetic differences or geographic separation (Jackson et al. 2017), our aim was to reconstruct the phylogeny of the species in Atlantoscia using mitochondrial and nuclear markers and test its monophyly by integrating molecular, morphological and biogeographical data. We observed, in this study, that Atlantoscia is paraphyletic. Atlantoscia ituberasensis and A. rubromarginata were grouped with a new species https://mc06.manuscriptcentral.com/cjz-pubs Page 5 of 38 Canadian Journal of Zoology 5 described in the present study and all of them should be placed in a new genus of terrestrial isopods. Information from morphology, genetic and biogeography corroborate these findings. Material and Methods Material studied Terrestrial isopods used in this study come from samples deposited in scientific collections of Museu de Zoologia (MZUSP), Universidade de São Paulo, São Paulo and Coleção de Carcinologia (UFRGS), Universidade Federal do Rio Grande do Sul, Porto Alegre. Individuals from an undescribed species sampled in the Brazilian state of Bahia and possibly related to the Atlantoscia genus (due to morphological similarities)Draft were also examined. Morphological Analysis: description of the new genus and new species The specimens were dissected and the appendages and pereonites were mounted on slides. Drawings were prepared using a camera lucida. The noduli laterales were measured and illustrated as in Vandel (1962). The terminology for the setae follows Campos-Filho et al. (2013). The descriptions are based on the paratypes mounted on slides. The material was deposited in the same museums mentioned in the previous session. Molecular Analysis: DNA extraction and PCR amplification Dissections were conducted as described by Bouchon et al. (1998). Total DNA was extracted using a PureLink Genomic DNA kit (Invitrogen/K1820-01). PCR amplifications were performed using https://mc06.manuscriptcentral.com/cjz-pubs Canadian Journal of Zoology Page 6 of 38 6 the LCO / HCO primer set (Folmer et al. 1994), which amplifies
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