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(Opiliones, Cyphophthalmi, Sironidae), in North America with a Phylogenetic Analysis Based on Molecular Data and the Description of Four New Species

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(Opiliones, Cyphophthalmi, Sironidae), in North America with a Phylogenetic Analysis Based on Molecular Data and the Description of Four New Species THE GENUS SIRO LATREILLE, 1796 (OPILIONES, CYPHOPHTHALMI, SIRONIDAE), IN NORTH AMERICA WITH A PHYLOGENETIC ANALYSIS BASED ON MOLECULAR DATA AND THE DESCRIPTION OF FOUR NEW SPECIES GONZALO GIRIBET1 AND WILLIAM A. SHEAR2 ABSTRACT. The North American fauna of the Laur- unrelated to Siro rubens Latreille, 1804, asian family Sironidae is examined phylogenetically and and therefore considered a different genus compared with species from Europe and Japan. The North American clade is not resolved as monophyletic. (Boyer et al., 2005; Karaman, 2008; Muri- The phylogenetic analyses and detailed morphological enne et al., 2010). In this article, we restrict study identified four cryptic species of sironids in the the concept of Siro to a clade of recent western United States, formerly considered within the Western European species composed of S. geographical and morphological range of Siro acaroides (Ewing, 1923). These four species are described as Siro rubens; Siro carpaticus Rafalski, 1956; Siro boyerae sp. nov., Siro calaveras sp. nov., Siro clousi sp. crassus Novak & Giribet, 2006; and Siro nov., and Siro shasta sp. nov. We also provide new valleorum Chemini, 1990, and to a clade of localities for the previously known species in the several North American species: Siro acar- western United States. Siro boyerae sp. nov. forms a oides (Ewing, 1923); Siro exilis Hofman, clade with Siro kamiakensis (Newell, 1943) and with the East Coast species Siro exilis Hoffman, 1963, 1963; Siro kamiakensis (Newell, 1943); and characterized by the presence of narrow coxae III that Siro sonoma Shear, 1980. The four previ- do not meet along the midline. The affinities of S. ously known North American species were calaveras sp. nov., S. clousi sp. nov., and S. shasta sp. revised by Shear (1980) and profusely nov. remain largely unresolved, but S. clousi sp. nov., is illustrated by de Bivort and Giribet (2004: not related to S. acaroides despite being found sympatrically. figs. 10–39). Siro acaroides was described in 1923 as the INTRODUCTION type of the new genus Holosiro Ewing, 1923, this species being the first cyphophthalmid The cyphophthalmid genus Siro currently discovered in the New World (Ewing, 1923). includes a series of species found in North Later, it was recognized that the species could America and continental Western Europe not be easily distinguished from the European (Giribet, 2000; Juberthie, 1970; Novak and Siro at the generic level, and Holosiro was Giribet, 2006; Shear, 1980). The status of considered a junior synonym of Siro (Newell, the European members of the genus Siro 1943). In the same article, Newell described a has been recently revised, and the radiation new species of American sironid in the new of species related to Cyphophthalmus dur- genus Neosiro Newell, 1943, for the species icorius Joseph, 1868, in the Balkans and Neosiro kamiakensis.Thenewgenuswas adjacent geographic areas seems to be based on the divided fourth tarsus of the male. Both species inhabit western North America, 1 Museum of Comparative Zoology, Department of each originally described from single locali- Organismic and Evolutionary Biology, Harvard Uni- ties: S. acaroides from Benton County, versity, 26 Oxford Street, Cambridge, Massachusetts 02138. southwestern Oregon, and N. kamiakensis 2 Department of Biology, Hampden-Sydney College, from Whitman County in western Washing- Hampden-Sydney, Virginia 23943. ton. An eastern North American species, Siro Bull. Mus. Comp. Zool., 160(1): 1–33, October, 2010 1 2 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 1 exilis,foundintheAppalachianMountains been recently collected by G. Giribet, S. along the boundary between Virginia and Boyer, and R. M. Clouse in Calaveras Big West Virginia, was subsequently added to the Trees State Park, which was suitable for list (Hoffman, 1963). molecular work. Meanwhile, Davis (1933) had described The map Shear published in 1980 did not Siro americanus from northwestern Florida; correlate well with the list of localities given; after an unwarranted sojourn in the genus a location for S. acaroides is shown signif- Parasiro Hansen and Sørensen, 1904 (Hin- icantly south of the California/Oregon ton, 1938), this species was made the type of border, but only Del Norte County records the new genus Metasiro Juberthie, 1960 are listed in the text. This map symbol was (Juberthie, 1960), within the family Sironi- added late in the preparation of the paper dae (or Sironinae of Juberthie, 1970) and referred to Shasta County specimens (Giribet, 2000; Juberthie, 1970; Shear, that were then considered S. acaroides. 1980). Later on, Hoffman (1963) proposed They are of a yet another new species. the synonym genus Floridogovea Hoffman, A field trip through Idaho, Washington, 1963, for Metasiro. Based on ample mor- Oregon, and Northern California by G. phological and molecular evidence, Meta- Giribet, S. Boyer, and R. M. Clouse in June siro is now considered a member of 2005 yielded numerous collections of Cy- Neogoveidae (Boyer et al., 2007b; Giribet, phophthalmi, including all known species 2007). for the western United States, with the In 1980, Shear had access to a wide range exception of the elusive S. sonoma. The aim of collections that had been assembled since of this trip was to obtain more specimens of 1947 (Shear, 1980). Contrary to the asser- the new species from Calaveras County and tions of Ewing (1923) and Newell (1947), Shasta County, as well as to revisit other Shear (1980) proposed that S. acaroides was cyphophthalmid localities to obtain speci- widely distributed in the Coast Ranges from mens suitable for molecular work for all the northern California to Puget Sound and that NW U.S. species. Two specimens of S. N. kamiakensis occurred in at least one sonoma were collected by G. Giribet, T. more locality in western Washington (Mt. Briggs, and D. Ubick in Monte Rio, Spokane) and at three places in Kootenai December 2001. Phylogenetic analysis of and Idaho Counties, in northern Idaho. the new specimens further revealed the Furthermore, Shear argued that the pre- presence of multiple cryptic lineages in ponderance of characters of N. kamiakensis the previously considered widespread were consistent with a placement in the species S. acaroides. Two of these species genus Siro and so synonymized Newell’s that could be characterized morpho- genus Neosiro. Shear also added a distinc- logically are described here. The new tive fourth species of Siro, S. sonoma Shear, species double the number of known 1980, from Sonoma County in Northern American sironids but also indicate that California. our knowledge of the American sironid Some loose ends were mentioned in fauna is still in its infancy. Shear’s 1980 paper. In particular, a single California has not been intensively ex- female specimen from Calaveras County, plored for cyphophthalmids. They are most California, in the Sierra Nevada Mountains, easily collected from Berlese samples; the seemed clearly to be a new species, but success of this method was demonstrated by Shear was reluctant to describe it from a the many specimens and new records of S. single female example. Now additional acaroides obtained by Ellen Benedict material from that same collection has (Shear, 1980). We have also been successful become available, and it is clear that this collecting many live specimens by sifting population represents a new, fifth species of with a 4-mm mesh size or via extraction with American Siro. Additional material has also Winkler apparatus. But other than these PHYLOGENETIC ANALYSIS OF SIRO IN NORTH AMERICA N Giribet and Shear 3 examples, most specimens have been Morphological Methods obtained after occasional direct collecting. For each species, the male holotype and a We predict that a thorough search of proper female paratype were photographed using a habitats in the Sierra Nevada, and both JVC KY-F70B digital camera mounted on a northern and southern Coast Ranges in Leica MZ 12.5 stereomicroscope. A series California, will yield more new species of of images (ca. 10) were taken at different sironids. The distribution pattern of soil- focal planes and assembled with the dedi- dwelling organisms with species in the cated software package Auto-Montage Pro Appalachians in the east and the Coast Version 5.00.0271 (Syncroscopy, Freder- Ranges and northern Idaho in the west ick, Maryland, USA). Each specimen was often includes the central Rocky Mountains photographed in dorsal, ventral, and as well; Siro might be expected to turn up in lateral views, and when available, the Utah, Colorado, or New Mexico. With 43 holotype was always photographed. Full extant species of sironids in Europe, it body measurements of the holotype and a seems reasonable to expect that North America eventually could be shown to have female paratype were then taken from more species than the 10 we know now. these photographs in Adobe Photoshop CS3 with the ‘‘Analysis’’ menu and were MATERIALS AND METHODS recorded in a spreadsheet. Total body length refers to the distance between Abbreviations for Repository Institutions midpoint of anterior and midpoint of AMNH American Museum of Natural posterior margin of the dorsal scutum. History, New York, New York, Body width refers to the maximum width, USA whether recorded in the prosomal or in the BMNH The Natural History Museum, opisthosomal region. London, United Kingdom One male and one female specimen of CAS California Academy of each species were examined with a FEI Sciences, San Francisco, Quanta 200 SEM (Peabody, Massachusetts, California, USA USA). Appendage and body part measure- CNHM Field Museum of Natural ments were taken from the digital micro- History, Chicago, Illinois, USA graphs in Adobe Photoshop CS3 with the (usually, FMNH for Field ‘‘Analysis’’ menu and were recorded in a Museum of Natural History) spreadsheet. Measurements of the chelic- EME Essig Museum of Entomology, era, palp, and leg articles were mostly taken U.C. Berkeley, Berkeley, on their dorsal side, from the midpoint of California, USA the anterior margin to the midpoint of the FMNH Field Museum of Natural posterior margin.
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