Eleven new species of theridiosomatid spiders from southern China (Araneae, Theridiosomatidae)

Abstract Two new genera of the spider family Theridiosomatidae, Alaria gen.n. with the type species Alaria chengguanensis sp. n., Menglunia gen.n. with the type species Menglunia inaffecta sp. n., are described from Guizhou and Yunnan, China. Nine more new species from Guangxi, Guizhou, Hainan and Yunnan Provinces of southern China are described: Baalzebub rastrarius sp. n.,Baalzebub youyiensis sp. n., Karstia nitidasp. n., Karstia prolata sp. n.,Ogulnius hapalus sp. n., Theridiosoma plumaria sp. n., Theridiosoma triumphalis sp. n., Theridiosoma vimineum sp. n., Zoma fascia sp. n. The type specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences in Beijing.


Introduction
Theridiosomatids are small (usually ≤ 3mm), widely distributed, and cryptozoic spiders, which can be found in damp, dark habitats (litter layer of forest or caves). They can be easily recognized by the presence of a pair of pits on the anterior margin of the sternum near the labial base (Fig. 9D), the disproportionately large globular pedipalps (except for genus Menglunia), and the long dorsal trichobothria on the third and fourth tibia (Coddington 1986).
The genera of family Theridiosomatidae were reviewed and revised by Coddington in 1986. Based on cladistic analysis, he recognized 4 subfamilies: Platoninae, Epeirotypinae, Ogulniinae and Theridiosomatinae. He also described/redescribed 9 genera and validated 28 species in his paper. Since then, 5 new genera and 25 new species have been discovered worldwide. In 2011, Wunderlich introduced three new theridiosomatid species from Laos and recognized one new subfamily: Luangnaminae. He also downgraded Platoninae, Epeirotypinae, and Ogulniinae to tribal rank based on rare or special characters. According to Platnick (2012), there are currently 16 genera and 89 species belonging to this family. Three widespread genera including Ogulnius, Theridiosoma, Wendilgarda, can be found in China and other Asian countries. The genus Coddingtonia was the first genus known only from China. As a monotypic genus, Coddingtonia is distinguished from other theridiosomatids by its distantly-separated spermathecae and long copulatory ducts. Miller et al. (2009) also discovered new species from other six genera, three of which were found in China for the first time. Zoma dibaiyin as the second speices and first known male from the genus Zoma, gives us new thoughts about the validation of Theridiosoma taiwanica (Zhang & Zhu, 2006). According to the illustrations given by that paper, we found that it resembles several critical features possessed by Zoma rather than Theridiosoma. We have no doubt that it will make more sense if it could be transferred to genus Zoma. The genus Karstia, including two species, was created by Chen (2010). As the second genus endemic to China, Karstia is placed in the Theridiosomatinae for three synapomorphies it shares with the other members of this subfamily: a row of short bristles on the cymbium at the junction with cymbial lamella, elongated median apophysis with a trough or groove along its upper surface, and singly attached egg sacs (Chen 2010: figs 12, 13, 30). Two new species of Kastia are described here in this paper, merely on the basis of the female characteristics, further diagnosis will be done when males are collected. To a certain extant, Karstia and Baalzebub are seemingly closely related, despite the fact Karstia has a few unique features: presence of cymbium apophysis, and stout, overlapped spermathecae, the accuracy or verity of it is still questionable to us, and more study efforts desperately need to be put into this extraordinary genus in the future.
Theridiosomatids in China are mainly found in the southern provinces: Guangxi, Guizhou, Hainan, Taiwan and Yunnan, which is consistent with the tropical and subtropical preferences typical for this family. Some theridiosomatids are associated with caves, which are characterized by stable humidity and temperature. In this paper, we provide descriptions and distribution data for eleven new species collected in Guangxi, Guizhou, Hainan and Yunnan.

Method
Specimens were examined using a LEICA M205 C stereomicroscope. Further details were studied under an Olympus BX51 compound microscope. All illustrations were made using a camera lucida attached to an Olympus BX51 compound microscope, and then inked on ink jet plotter paper. Male and female genitalia were examined and illustrated after being dissected from the spiders' bodies. Left pedipalps of male spiders were illustrated, except as otherwise indicated. Vulvae of female were removed and cleared in lactic acid or warm 10% potassium hydroxide (KOH) solution before illustration. All embolic divisions and vulvae were illustrated after being embedded in Arabic gum. Type specimens examined were preserved in 75% ethanol solution. Photos were taken with an Olympus c7070 wide zoom digital camera (7.1 megapixels) mounted on an Olympus SZX12 stereomicroscope. Images from multiple focal planes were combined using Helicon Focus (version 3.10.3) image stacking software. All measurements are given in millimeters. Leg measurements are shown as: total length (femur, patella, tibia, metatarsus, tarsus).
SEM images were taken using the HITACHI S-3000N at the Institute of Genetics and Developmental Biology, Chinese Academy of Sciences. Specimens for SEM examination were critical point dried and sputter coated with gold-palladium. Specimens were mounted on copper pedestal using double-sided adhesive tape.
All type specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences in Beijing.
Chaetotaxy. Macrosetae are marked for the dorsal (d), prolateral (p), retrolateral (r), and ventral (v) surfaces of the legs. Metatarsal trichobothrium (Tm) is given as the ratio of the distance between the proximal margin of the metatarsus and the root of the trichobothrium divided by the total length of the metatarsus (Locket and Millidge 1953) and Tm value for each leg is given as Tm I, Tm II, Tm III, Tm IV respectively.
Abbreviations and conventions. Abbreviations used in the text are given in Table  1. References to figures in cited papers are listed in lowercase type ( fig.); Figures of this paper are noted with an initial capital ( Fig.).
When extra materials are examined and recorded, and the paratype's collecting information is the same as holotype's, it will be implied in brackets as [same data as holotype]. The turning made by the copulatory ducts bends outwardly (Fig. 20B). Median apophysis mesally with a projection oriented distoventrally (Fig. 19A). Embolic apophysis long, whip-like (Fig. 19C)  Scape less sclerotized, smaller and partially exposed (Coddington 1986: fig. 206 The upper rim of the spermathecae is the same height as the copulatory ducts (Miller et al. 2009: fig. 11B The mesal bristle of the embolic apophysis protruding from beneath the conductor and lying long the mesal side of the conductor itself (Coddington 1986: fig. 198 Conductor with plumose branching (Fig. 22C). Median apophysis with a curved lobe attenuates distally ( Etymology. The generic epithet is derived from the Latin 'alarius', meaning 'of wings', which refers to the two projections of the copulatory ducts at each side, which resembles a pair of wings. Gender is feminine. Diagnosis. The unique structure of epigyne distinguishes Alaria from other theridiosomatids. Like in Wendilgarda and Chthonopes, the scape in female Alaria protrudes from beneath epigynal plate (Coddington 1986: figs 206, 213;Wunderlich 2011: figs 18d-e), but is utterly exposed, and more sclerotized, like a shield attached to the abdomen (Figs 2A-B, 4A-B). The conformation of the copulatory ducts is similar to that in Ogulnius obtectus (Coddington 1986: fig. 113), but copulatory ducts make one coil before the conjuncture with spermathecae instead a fold (Fig.  2B). The paracymbium in Alaria is neither a T-shaped lobe as in most thridiosomatids nor a broad apophysis as in Epeirotypinae, it is a thick, elongated structure with a small hooked projection (Fig. 3D). The long, whip-like embolus in Alaria resembles embolic apophysis in Ogulnius (Coddington 1986: figs 100-101, 116, 118), but proportionately much longer and mostly enveloped in conductor ( Fig. 1B-D). The median apophysis of Alaria is disproportionately large, stretching along the longitudinal axis of pedipalp with two curved, pointed distal ends (Figs 1A, 3A), which is never seen in any other theridiosomatid genus. Based on the combination of features mentioned above, Alaria should be recognized as a new genus, and is likely close to Wendilgarda and Chthonopes.
Species. Alaria chengguanensis sp. n. Vulva: Epigyne with long, tongue-shaped scape protruding from beneath its concaved margin. Scape slightly humped with a small transverse opening at its distal end. Spermathecae juxtaposed. Copulatory ducts rise and curl up to form a blunt-tipped projection at each side (     Etymology. The epithet comes from Latin word 'rastrarius' which means 'of a hoe', referring to the hoe-shaped conductor in prolateral view; adjective. Diagnosis. The presence of small, cleft median apophysis and blunt, spatulate processes of embolic apophysis in males, and tip-fused spermathecae in females indicates that this species belongs to the genus Baalzebub. Conductor in males envelopes the entire embolic apophysis (Fig 6A), which is similar to B. albinotatus, but the rectangular conductor and the small, pointed cymbium apophysis are different from other described Baalzebub species. Females distinguished by the triangular epigynal plate, similar to B. baubo (Coddington 1986: figs 183, 184), but distinguished by the narrower, arcade-shaped spermathecae (Fig. 7B).   Description. Carapace yellow tan. Sternum yellow with dark orange margins and setae. Legs yellow, dark brown distally at joints, especially tibiae. Abdomen tan with 2-4 rows of dark grey patches, the first two of which are connected by a perpendicular grey patch at midline, and two pairs of small dorsal brown spots (Figs 7C-F).
Vulva: Epigyne with a smooth triangular scape protruding from posterior margin of epigyne plate (Figs 7A-B). A deep pit lies on the epigynal mesal anterior edge. Spermathecae long, narrow, meeting at the tip to form an arcade-shaped structure. Copulatory ducts follow simple curve (Figs 7B, 8D Etymology. This specific name formed from the Chinese words for friendship yŏu yì ( ), which is the name of the county where this species was collected; adjective. Diagnosis. Females distinguished from other described Baalzebub by the shape of the semi-transparent scape, which is proportionately shorter and with a blunt tip (Fig. 9A). Spermathecae, contrasted with narrow, long spermathecae in other Baalzebub species, is relatively shorter and smaller compared to the ovoid loops made by copulatory ducts (Fig. 9B)  Description. Carapace broad, orange. Sternum yellow with dusky margins. Legs yellow, brown from patella to tarsus (except for leg I, which only brown distally at joints). Abdomen beige with irregular light, greenish-grey patches (Figs 9C-E).
Vulva: Epigyne with a short, pointed triangular scape with concave lateral margins protruding from posterior margin of epigyne plate, through which dark orange vulva is visible (Fig. 9A). Scape translucent. Epigyne plate with transverse grooves (Fig. 9A). Spermathecae small, elliptical, joining each other at the tip (Figs 9B, 10B) Diagnosis. Females distinguished by the following combination of characters: the structure of the scape, the stout, overlapped spermathecae (Fig. 11B), and the habitus of this species (Figs 11C-E). Spermathecae oval-shaped with vertically longer diameter, slightly detached from each other along their inner margin. The abdomen large, contrasted with distinctly small epigynal area. The scape structure is quite different from other Karstia or Baalzebub species: the tip of scape is swollen and shimmery, and the lateral margins of the plate extend toward the scape tip to form a armet-  shaped conformation (Fig. 11A). Generic placement tentative pending discovery and examination of the male.
Description. Carapace pale yellow with yellow ocular region. Sternum yellow with tan margins. Legs yellow, brown distally at joints. Abdomen dark grey mottled with white patches (Figs 11C-E).
Vulva: Epigyne small, with short, distally spherical scape protruding from posterior margin of epigyne plate. Epigyne plate extends posteriorly, together with scape to form a barbute-shaped conformation (Figs 11A, 12A). End of scape glossy with black purfle (Fig. 11A). Spermathecae peanut-shaped, juxtaposed, slightly detached from each other (Figs 11B, 12B). Copulatory ducts follow simple rout to form small loops and one turning before connected with spermathecae at the bottom (Figs 11B, 12B Diagnosis. Females distinguished by the protruding scape and the overlapped, stout spermathecae. The long acute-angled scape protrudes vertically from the posterior epigynal margin (Figs 13B, F), which is different from other known Karstia species and Baalzebub species.
Description. Relatively large in total body length, compared to other theridiosomatid species. Carapace greenish tan, with brown ocular region. Sternum yellow with dark margins. Femur yellow, brown from patella to tarsus. Abdomen beige with evenly distributed silver spots within dorsal area and 8 or more rows of mesally disrupted greenish grey patches, with a large, wedge-shaped greenish grey patch between epigyne and spinnerets (Figs 13D-E).
Vulva: Epigyne with a long, apiculate scape protruding perpendicularly (slightly tilted) from posterior margin of epigyne plate, flanked by a cluster of long sinuous setae on each side (Figs 13A, E-F). Two deep grooves occur at the posterior base of epigyne which are likely connected to the copulatory openings (Figs 13B, E). Tip of  scape glossy black (Fig. 13A). Spermathecae peanut-shaped, juxtaposed (Fig. 13C) Diagnosis. Distinguished from other theridiosomatids by the extremely simple, short embolus, and the round, separated spermathecae. The pedipalp in males is an elliptical (slightly rectangular in total), theca-textured, and obscurely circumscribed structure (Figs 15A, D). Conductor is less extensive, and fully covers the embolus (Fig.  18A). The embolus is beak-like, and enveloped in conductor (Fig. 15B). Unlike any other theridiosomatid genus, the median apophysis in Menglunia is merely a small projection, mildly curved without any sharp tip or trough (Fig. 15A). Spermathecae similar to Coddingtonia euryopoides (Miller et al. 2009) and Luangnam discobulbus (Wunderlich, 2011), but instead of being elliptical and separated by their diameter, they are more rounded and separated by less than half of their diameter. Copulatory duct is shorter and forms a simple loop which is about the same height as spermathecae's diameter, compared to the big loop and higher-positioned copulatory ducts in C. euryopoides and L. discobulbus (Fig. 16B).
Species. Menglunia inaffecta sp. n.    Description. Carapace yellow tan. Sternum yellow with greenish brown margins and sparse hairs. Legs thick, yellow. Abdomen beige with small, irregularly-distributed greenish brown spots.

Genus
Diagnosis. Though lack of one genetic feature, which is fourth legs are longer than the first legs (subequal in females), other generic characteristics of Ogulnius can be seen in this species: tapering whip-like embolic apophysis in males (Fig. 19C), transverse grooves on epigyne in females, separated and juxtaposed posterior median eyes (Coddington 1986). Males distinguished from other described Ogulnius species by the shape of median apophysis and the proportion of embolic apophysis. The median apophysis in O. hapalus is similar to that in O. gloriae (Coddington 1986: fig. 99): mesally wide, with a projection oriented distoventrally (Fig. 19A). The embolic apophysis is proportionately longer than that in O. barbandrewsi (Miller et al. 2009: fig. 5F). The Females distinguished by the routing of copulatory ducts and the peanut-shaped spermathecae. The fold made by the copulatory ducts is bending outwardly (Fig. 20B), instead of in-   wardly as in O. barbandrewsi (Miller et al. 1986: fig. 3D). The posterior lip of epigyne in O. hapalus concave in stead of convex as in O. pullus (Brignoli 1981: fig.1).

Genus
Diagnosis. Males, similar to T. gemmosum (L. Koch) (Coddington 1986: figs 134, 135), distinguished from any other known male Theridiosoma from Asia by the following characters: broader, slightly groovy median apophysis with an acuminated tip, and the exposed branchings of embolic apophysis (Fig. 22A).
Description. Carapace yellow tan. Sternum yellow with brown margins. Legs yellow. Abdomen tan with sparse silver specks and symmetric dark grey patches.
Diagnosis. Males distinguished from other described Asian Theridiosoma species by the shape of median apophysis: a small, curved projection with an attenuated tip extending distoventrally (Fig. 24B). Embolic apophysis fragmented, similar to T. caaguara (Rodrigues and Ott 2005: figs 4, 5), but different in details: four pieces of different shapes, the longest one with a whip-like tip, the longer one broad, with a blunt end (Fig. 24C).
Description. Carapace yellow tan. Sternum yellow with dark margins. Legs yellow. Due to poor preservation condition, abdomen too shriveled to make out its original color pattern.
Male pedipalp: Patella with strong macroseta. Paracymbium elongate with attenuated tip. Tegulum smooth with tuberculate ventral ridge. Median apophysis curved lobe with attenuated distal end. Conductor translucent theca covering most fragmented embolic apophysis, with a small rough ventral area (Fig. 24A). Two embolic apophysis fragment tips protruding out of the conductor tip to form a 'V'-shaped conformation (Fig. 24B) Etymology. The specific name is derived from Latin word 'vimineus' meaning 'pliant', refers to the long, pliant apophysis that protrudes from the conductor (Fig. 26B); adjective.
Diagnosis. Males similar to T. semiargentum (Simon) (Coddington 1986: figs 154,156), but different in the details of embolic apophysis fragments and the shape of median apophysis. The apophysis protruding from the conductor is longer and slimmer in T. vimineum, spur on the conductor is absent, and the median apophysis lacks a pointed hook at the tip.
Description. Carapace yellow. Sternum tan with dark margins. Legs yellow. Abdomen tan with light grey spots randomly distributed on the dorsal area.
Male pedipalp: Patella with macroseta. Tibia with one trichobothrium. Paracymbium elongate with sharp tip. Tegulum smooth with rough mesal lobe. Median apophysis slightly grooved, semi-transparent at the distal end (Figs 26B, 27B). Conductor translucent theca with a hooked end oriented proximally (Fig. 26A). A piece of long, flat embolic apophysis fragment with a attenuated tip protruding from the ridge of conductor and stretching toward median apophysis (Figs 26B, D). Embolic apophysis fragmented into long, slim bristles (Fig. 26C)    Etymology. The specific name comes from a Latin word 'fascia', which refers to the silver band on its abdomen; noun.

Diagnosis.
Males distinguished by the presence of a filiform embolic apophysis extending beyond the conductor tip and a 'Z'-shaped embolus (Figs 28B, D). Exposed portion of embolic apophysis is smaller compared to Z. didaiyin (Miller et al. 2009: fig. 10F).
Female distinguished from Z. didaiyin by the mildly convex posterior margin of the epigyne and the higher position of the spermathecae relative to the copulatory ducts (Figs 29A, B).
Description. Carapace brownish yellow in males, brown in females. Sternum yellow with dark brown margins. Legs yellow. Abdomen beige in males, dark grey in females with silver pecks forming a transverse belt.