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2010. The Journal of Arachnology 38:57–72 The genus Taira, with notes on tibial apophyses and descriptions of three new species (Araneae: Amaurobiidae) Xin-Ping Wang: College of Life Sciences, Hebei University, Baoding, Hebei, 071002, China. E-mail: [email protected] Peter Ja¨ger: Arachnology, Research Institute Senckenberg, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany Zhi-Sheng Zhang: College of Life Sciences, Southwest University, Chongqing, 400715, China Abstract. Homologies of the tibial apophyses of Taira with those of other members of the subfamily Amaurobiinae are evaluated. The monophyly of the genus Taira and the phylogenetic relationships of its species are analyzed using parsimony. The genus Taira is supported by two putative synapomorphies: the presence of broad epigynal teeth and the distally originating tegular sclerite apophysis. A diagnosis and description of Taira and a key to its species are provided. Three new species are described from China: Taira qiuae new species (LK), T. sichuanensis new species (LK), and T. zhui new species (LK). Keywords: Identification key, taxonomy, homology, phylogenetic relationships, China Lehtinen (1967) created the new genus Taira for the METHODS species Amaurobius flavidorsalis Yaginuma 1964, described All measurements are in mm. Scale lines are 0.2 mm long from Japan, compared it with other genera of Amaurobiidae, except where indicated otherwise. Eye diameters are taken at and placed Taira with the genus Tamgrinia Lehtinen 1967 in the widest point. The length of body, prosoma, and the tribe Tairini within the Holarctic subfamily Amaurobii- opisthosoma do not include the length of the chelicerae or nae Thorell 1870. Only four papers have been published on spinnerets. The distribution map was generated using GIS Taira since its establishment in 1967. Wang (2000) ques- ArcView software, and the text files of the studied species are tioned the sister group relationship between Taira and downloadable from Wang (2009). More type specimen photos Tamgrinia by comparing spinnerets, tracheae, and tricho- of the species included in this paper can be viewed in Li & bothria of the family Amaurobiidae. Zhu et al. (2004) and Wang (2009). Wang & Ran (2004) described two new species based on Anatomical abbreviations used in the text and figures: Eyes: specimens from two closely situated localities in Libo AME–anterior median eyes; ALE–anterior lateral eyes; PLE– County, Guizhou, China. A study by Zhang et al. (2008) posterior lateral eyes; PME–posterior median eyes. Female concluded that T. lunaris Wang & Ran 2004 is a junior Genitalia: CD–copulatory duct; EL–epigynal median lobe; synonym of T. liboensis Zhu, Chen & Zhang 2004. In recent ET–epigynal tooth; FD–fertilization duct; S–spermatheca. years, field work by Zhang et al. (2008) in southern China Male palp: C–conductor hyaline apophysis; C1–conductor has yielded five more new species: T. cangshan Zhang, Zhu & apophysis I (from which hyaline apophysis arises); C2– Song 2008, T. latilabiata Zhang, Zhu & Song 2008, T. obtusa conductor apophysis II, which is an extension of the Zhang, Zhu & Song 2008, T. concava Zhang, Zhu & Song conductor apophysis I; C3–conductor apophysis III; CYM– 2008, and T. sulciformis Zhang, Zhu & Song 2008, and a new cymbium; dDTA–distal/subdistal tooth of DTA; DTA–dorsal combination, T. decorata (Yin & Bao 2001). As a result, a tibial apophysis; E–embolus; eDTA–ectal branch of DTA; total of eleven species from Japan and southern China are ITA–Intermediate tibial apophysis; MA–median apophysis; included in Taira, including the three new species described mDTA–mesal branch of DTA; RTA–retrolateral tibial here. In all previous studies, the diagnosis of the genus Taira, apophysis; ST–subtegulum; T–tegulum; TB– tibia; TS–tegular and therefore the distinction between Taira and other sclerite; TSA–tegular sclerite apophysis. amaurobiids, was only vaguely defined. Zhang et al. (2008) Specimens studied in the current paper are deposited in the provided a diagnosis for Taira that we argue was based on a California Academy of Sciences, San Francisco (CAS); Hunan misinterpretation of non-homologous features (i.e., tibial Normal University, Changsha, Hunan (HNU); Institute of apophyses, see below). This study is focused on diagnosing Zoology, Chinese Academy of Sciences, Beijing (IZCAS); and describing the genus Taira, discussing its monophyly, Museum of Hebei University, Baoding, Hebei (MHBU); estimating the species relationships, and providing a key to Southwest University, Chongqing (SWUC); and Senckenberg the Taira species described so far. The three new species Museum, Frankfurt (SMF). described here belong to a distinct group, which differs from other Taira by the widely separated, elongated spermathecae AMAUROBIINAE TIBIAL APOPHYSES in females and the long RTA (except T. sichuanensis), Tibial apophyses of the subfamily Amaurobiinae are presence of an intermediate apophysis between the RTA and important characters in species identification, generic limita- dorsal tibial apophysis, and the uniquely modified conductor tion, and, therefore, valuable tools for phylogenetic analyses. in males. In previous publications, however, different arachnologists 57 58 THE JOURNAL OF ARACHNOLOGY Figures 1, 2.—Schematic drawings of Taira tibia (TB), showing the retrolateral tibial apophysis (RTA), dorsal tibial apophysis (DTA), intermediate tibial apophysis (ITA), external branch (eDTA) and mesal branch (mDTA) of DTA, and teeth (dDTA) on DTA. CYM 5 cymbium. 1. Prolateral view. 2. Retrolateral view. have perceived and defined those apophyses in different ways have an RTA distinctly separated from the DTA. The RTA and given them different names. In this study, we have was labeled as the ectal process by Leech (1972: fig. 10) and conducted initial homology assessments of Amaurobiinae Yaginuma (1987: figs. C, D, G, H). tibial apophyses based on similarities in their position and morphology. There are at least three apophyses on amaur- DORSAL TIBIAL APOPHYSIS (DTA) obiine tibiae: a retrolateral tibial apophysis (RTA), a dorsal The presence of a DTA, in combination with a hyaline tibial apophysis (DTA) and an intermediate tibial apophysis conductor, is synapomorphic for the family Amaurobiidae (ITA), which originates between the RTA and DTA in most (Griswold et al. 2005). The DTA is usually long with a slender Amaurobiinae. The Taira DTA usually has branches on its distal end in Amaurobius and Callobius (Leech 1972: figs. 10, external and mesal side, or teeth on its distal half; here we 11). In Taira the DTA is broad, modified with grooves, an differentiate the external branch of the DTA (eDTA), mesal external branch (eDTA) and mesal branch (mDTA), or with branch of the DTA (mDTA), and distal teeth of the DTA one or more distal teeth (dDTA) (Figs. 1, 2). eDTA is a (dDTA) (Figs. 1, 2). branch protruding externally on the DTA toward the lateral side of the body, while mDTA is a branch protruding mesally RETROLATERAL TIBIAL APOPHYSIS (RTA) on the DTA toward the middle of the body in natural The presence of the RTA, plus the presence of tarsal condition (dorsal view of palp). The DTA was termed a trichobothria and three or more metatarsal trichobothria, ‘‘mesal process’’ by Leech (1972: fig. 10) and Yaginuma (1987. supports the monophyly of RTA-clade spiders (Coddington & figs. G, H), an ‘‘innere apophyse’’ by Thaler (1990: figs. 7–10), Levi 1991; Griswold et al. 2005). The morphology of the RTA a ‘‘prolateral-dorsal apophysis’’ by Thaler & Knoflach (1993: varies among amaurobiids. Rather than originating proximal- figs. 7–12), and a ‘‘prolateral apophysis’’ by Thaler & ly or medially on tibia, extending along the tibial length, Knoflach (1991: figs. 3, 4). Zhang et al. (2008) treated the protruding distally as in coelotines (Wang 2002: figs. 44–46; DTA as having two branches, iDTA (interior branch of DTA) Wang 2003; Wang & Ja¨ger 2007; Xu & Li 2008), the RTA of and eDTA (exterior branch of DTA). The iDTA of Zhang et most amaurobiines originates distally or medially on the tibia, al. is in fact the DTA (rather than a branch on DTA), and the and does not extend along the tibial length (Leech 1972; eDTA of Zhang et al. is the mDTA, which protrudes mesally Yaginuma 1987; Thaler & Knoflach 1993). In Taira, the RTA toward the middle of the body (rather than on the external varies from absent, as in T. liboensis (Zhang et al. 2008: figs. side of DTA). All Taira species have a long, deeply grooved 31–33), long, sharply pointed, originating on the distal half of DTA and a broad mDTA except T. liboensis, which has a less the tibia, as in T. qiuae new species (Figs. 19–21) and T. zhui deeply grooved DTA and a broad eDTA, but no mDTA new species (Figs. 33–35), to a small lobe, as in T. sichuanensis (Zhang et al. 2008: figs. 31–33). DTA of T. cangshan is less new species (Figs. 26–28) and five other remaining Taira grooved and slightly larger than mDTA (Zhang et al. 2008: species (i.e., T. cangshan, T. concava, T. decorata, T. figs. 14–16). flavidorsalis, and T. sulciformis) (Zhang et al. 2008: figs. 11, 16, 21, 26, 40). We follow Zhang et al. (2008) and treat this INTERMEDIATE TIBIAL APOPHYSIS (ITA) small apophysis as the RTA. Among those with a small RTA, The ITA is small, lobe-shaped, and present in most T. cangshan has a RTA originating on the proximal half of the amaurobiines between the RTA and DTA. The ITA was tibia; in the remaining species, it originates either medially or referred to as a ‘‘dorsal process’’ by Leech (1972: fig. 10) and distally on the tibia. Except for T. convava and T. sulciformis, Yaginuma (1987: figs. G, H), as a ‘‘mittlere apophyse’’ by which have an RTA situated close to the DTA, other species Thaler (1990: figs. 7–10), and as an ‘‘intermediate apophysis’’ WANG ET AL.—GENUS TAIRA (AMAUROBIINAE) 59 by Thaler & Knoflach (1993: figs. 7–12).