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The Systematics and Biogeography of the Mite Harvestman Family

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The Systematics and Biogeography of the Mite Harvestman Family The systematics and biogeography of the mite harvestman family Sironidae (Arachnida : Opiliones : Cyphophthalmi) with the description of five new species Author(s): Gonzalo Giribet, Ligia R. Benavides and Izaskun Merino-Sáinz Source: Invertebrate Systematics, 31(4):456-491. Published By: CSIRO Publishing URL: http://www.bioone.org/doi/full/10.1071/IS16086 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. CSIRO PUBLISHING Invertebrate Systematics, 2017, 31, 456–491 http://dx.doi.org/10.1071/IS16086 The systematics and biogeography of the mite harvestman family Sironidae (Arachnida : Opiliones : Cyphophthalmi) with the description of five new species Gonzalo Giribet A, Ligia R. Benavides A,C and Izaskun Merino-Sáinz B AMuseum of Comparative Zoology & Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA. BDepartamento de Biología de Organismos y Sistemas, Universidad de Oviedo, Calle Catedrático Rodrigo Uría s/n, 33071 Oviedo, Asturias, Spain. CCorresponding author. Email: [email protected] Abstract. Sironidae, the first described family of Cyphophthalmi, is among the least understood phylogenetically. After examining recent collections across their distribution range, we provide the first comprehensive treatment of Sironidae by including molecular data from most of the known species, and all genera except for the monotypic Odontosiro Juberthie, 1961. We also revisit the male genitalic morphology for most genera by using confocal laser scanning microscopy and provide descriptions of five new species belonging to Iberosiro de Bivort & Giribet, 2004 (monotypic until now), Paramiopsalis Juberthie, 1962 and Siro Latreille, 1802. While the monophyly of Sironidae remains poorly supported using traditional Sanger-based markers, with the Mediterranean Parasiro Hansen & Sørensen, 1904 and the Japanese Suzukielus Juberthie, 1970b sometimes branching basally with respect to the other sironids, the remaining genera form a well- supported Laurentian/Laurasian clade. This group divides into a Western European/North American clade of Siro and the remaining genera, Iberosiro, Paramiopsalis and Cyphophthalmus Joseph, 1868. Iberosiro and Paramiopsalis form a well- supported clade from the NW corner of the Iberian Peninsula, while Cyphophthalmus is widespread in the Balkan region and Eastern Mediterranean. Finally, the following new taxa are described: Iberosiro rosae Giribet, Merino-Sáinz & Benavides, sp. nov., Paramiopsalis anadonae Giribet, Merino-Sáinz & Benavides, sp. nov., Paramiopsalis ramblae Benavides & Giribet, sp. nov., Siro ligiae Giribet, sp. nov., and Siro richarti Benavides & Giribet, sp. nov. Additional keywords: diversity, Iberian Peninsula, Iberosiro, Paramiopsalis, Siro. Received 19 December 2016, accepted 20 February 2017, published online 7 July 2017 Introduction exception of a morphological cladistic analysis (de Bivort and The Opiliones suborder Cyphophthalmi has received ample Giribet 2004). More recently, Dreszer et al.(2015) published attention from phylogenetic and biogeographic points of view an updated phylogenetic analysis of the family, but focused (Juberthie and Massoud 1976; Shear 1980; Boyer et al. 2007; mostly on the genus Cyphophthalmus and on previously Giribet et al. 2012). From its six currently recognised families, described taxa, and the outgroup sampling did not allow for Sironidae is restricted to the former Laurentian/Laurasian terranes testing of the monophyly of Sironidae. A study by Giribet et al. (Fig. 1), while the five remaining families have a Gondwanan (2012), which combined morphological and molecular data, was origin (Giribet and Prieto 2003; Sharma and Giribet 2009; Clouse ambiguous with respect to the monophyly of Sironidae, but found and Giribet 2010; Benavides and Giribet 2013; Giribet et al. four main clades: Parasiro Hansen & Sørensen, 1904 (with three 2016) (but see discussion about Marwe coarctata Shear, 1985 described species; endemic to the Western Mediterranean); below, a putative sironid found in Kenya). Recent work on sironid Suzukielus Juberthie, 1970b (monotypic; Honshu, Japan); Siro phylogeny has focussed on particular clades – the genera Siro (with 12 described extant species plus two amber fossils; Latreille, 1802 (Giribet and Shear 2010), Cyphophthalmus North America and Europe); and the clade formed by Joseph, 1868 (Boyer et al. 2005; Karaman 2009; Murienne Paramiopsalis (two described species; NW Iberian Peninsula) et al. 2010; Dreszer et al. 2015) and Paramiopsalis Juberthie, and Cyphophthalmus (33 species; Balkan region to Asia Minor). 1962 (Murienne and Giribet 2009) – but few studies have Siro, Paramiopsalis and Cyphophthalmus constituted the core addressed the global phylogeny of the family (Giribet and of Sironidae, sharing many of the characters typically assigned to Boyer 2002; Boyer et al. 2007; Giribet et al. 2012), with the the family and forming a clade in virtually all analyses. However, Journal compilation Ó CSIRO 2017 www.publish.csiro.au/journals/is Systematics and biogeography of Sironidae Invertebrate Systematics 457 (A) (B)(C) (D) (E) (F)(G) (H) Fig. 1. Geographical distribution of the sironid taxa included in our molecular study. (A) Distribution of the family worldwide. B–H, Detailed maps showing the generic sampling. (B) Cyphophthalmus depicted in Cyan; (C) Iberosiro represented in red (I. rosae, sp. nov. represented by a triangle); (D) Paramiopsalis in orange (P. anadonae, sp. nov. represented by a triangle; P. ramblae, sp. nov. by a square); (E) Parasiro in green; (F) Suzukielus in magenta; (G) European Siro and (H) North American Siro both in blue (S. ligiae, sp. nov. represented by a triangle; S. richarti, sp. nov. by a square). 458 Invertebrate Systematics G. Giribet et al. the membership of Parasiro and Suzukielus in the main sironid sputter-coated with gold using an EMS 300T D Dual Head clade is poorly supported, if at all, in Sanger-based molecular Sputter Coater (Electron Microscope Sciences, Hatfield, PA). analyses and in morphological analyses (e.g. Giribet and Boyer Images were taken using a Zeiss Supra 55VP Field Emission 2002; de Bivort and Giribet 2004; Boyer et al. 2007; Giribet Scanning Electron Microscope at the Harvard Center of et al. 2012). Iberosiro de Bivort & Giribet, 2004, a monotypic Nanoscale Systems (CNS). genus described from Portugal (de Bivort and Giribet 2004), was found to form a clade with the sympatric genus Paramiopsalis Auto-fluorescence imaging (de Bivort and Giribet 2004; Dreszer et al. 2015). We used an LSM 880 with Airyscan (Carl Zeiss, Jena, Here we provide a comprehensive phylogeny of the family Germany) system from the Harvard Center for Biological fi Sironidae based on ve genetic markers, including molecular Imaging (HCBI) to image the spermatopositor of the new data from multiple individuals of the genus Iberosiro, for which species. Spermatopositors were dissected from the ventral side we describe a second species from Asturias, Spain. This new of the opisthosoma and placed on slides containing Rapiclear species lives in a rather unorthodox habitat for Cyphophthalmi, (SunJin Lab Co., Hsinchu City, Taiwan). The master pinhole was and occurs in sympatry with a species of Paramiopsalis, set to 1–1.25 Airy unit (AU), and the 20Â Plan Apochromat long which we describe here as well. In addition, we name a second working distance objective was used. Laser wavelengths of 481 Paramiopsalis from Asturias, to bring the genus to four species and 565 nm were used to detect the auto-fluorescence of the (Juberthie 1962; Rambla and Fontarnau 1984; Murienne and chitin. A series of images (100–270) were taken in the z direction Giribet 2009), and two new species of Siro from North America, to generate a Z-stack. Zen 2 blue edition modular image- bringing the number of living Siro species to fourteen (Ewing processing software (Carl Zeiss, Jena, Germany) was used 1923; Hoffman 1963; Shear 1980; Giribet and Shear 2010). to assemble, edit and generate a 3D reconstruction of the Species descriptions are accompanied with detailed scanning spermatopositor for each species. All videos were deposited in electron micrographs and with novel imaging of their genitalia the MCZ database MCZbase (http://mczbase.mcz.harvard.edu) with confocal laser microscopy, as in previous recent studies associated with the specimen records. (Murienne and Giribet 2009; Dreszer et al. 2015). Molecules Methods Laboratory protocols Taxon sampling Foreach specimen, DNAwas extracted and isolated from up to All specimens used for this study were preserved in ~96% ethanol two legs using the Qiagen DNeasy
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