Four new species of the trapdoor spider genus Conothele Thorell, 1878 from Mainland China and Laos (Araneae, Ctenizidae)

Abstract Here for the first time the presence of the trapdoor spider genus Conothele Thorell, 1878 (Araneae: Ctenizidae) is reported from mainland China and Laos. Four Conothele species collected from the regions are described as new to science, based on the female genital morphology: Conothele baiyunensis Xu, Xu & Liu, sp. n. (Guangdong Province), Conothele daxinensis Xu, Xu & Li, sp. n. (Guangxi Province), Conothele sidiechongensis Xu, Xu & Liu, sp. n. (Yunnan Province, China and Vietnam), Conothele yundingensis Xu, Xu & Li, sp. n. (Yunnan Province).


Introduction
The family Ctenizidae belongs to the suborder Opisthothelae and the infraorder Mygalomorphae. Ctenizids are widely distributed, and mainly found across Asia (China, India, Japan, Laos, Myanmar, Sumatra, Thailand, Vietnam), the Americas (North and South America), the Mediterranean, South Africa and Australia (World Spider Catalog 2016). Ctenizidae is a dispersal-limited, ground-dwelling lineage, members of which usually build underground silk lined burrows opening to the surface with a trapdoor. The trapdoor is sometimes covered with a layer of leaf litter or moss, which blends well with the surrounding environment, thus making it difficult to spot in the field (Gertsch and Wallace 1936;Hunt 1976;Platnick and Gertsch 1976;Bond and Coyle 1995).
Ctenizids were traditionally divided into two subfamilies based on morphological features, Ctenizinae and Ummidiinae (Raven 1985;Ortiz 2007). Ctenizinae includes six genera, Bothriocyrtum Simon, 1891, Cteniza Latreille, 1829, Cyclocosmia Ausserer, 1871, Cyrtocarenum Ausserer, 1871, Latouchia Pocock, 1901, and Stasimopus Simon, 1892 However, this grouping is not supported by any identified synapomorphies (Raven 1985). Ummidiinae contains three genera, Conothele Thorell, 1878, Hebestatis Simon, 1903, and Ummidia Thorell, 1875 and it is diagnosed by the presence of a saddle depression on the tibia III that may serve as an anchor in the burrow (Gertsch 1979;Coyle 1981Coyle , 1985. Recently Decae (2010) removed Hebestatis from this subfamily based on some distinct morphological characteristics, including the presence of lateral sternal sigilla, a less pronounced and glabrous dorsal saddle on the tibia III, an absence of curvy spines, a lack of tarsal clavate trichobothria, and the absence of centrally sclerotized spermathecae, which are not present in the other genera of Ummidiinae. Therefore, the taxonomic position of Hebestatis remains uncertain (but see Garrison et al. 2016). The phylogenetic structure of the family Ctenizidae is also contentious. Recent phylogenetic studies have recovered neither the monophyly of Ctenizidae nor of Ctenizinae (Hedin and Bond 2006;Ayoub et al. 2007;Bond et al. 2012;Opatavo et al. 2013), even though Garrison et al. (2016) recovered Cyclocosmia and Hebestatis as monophyletic. Most importantly, Cteniza, the type genus of the family, which is consistently supported as sister to the other Mediterranean genus Cyrtocarenum, is never recovered as closely related to any of the remaining genera of the family (Raven 1985;Decae 1996;Opatova 2013). Similarly, the genus Stasimopus never clusters with any other Ctenizidae genus (Hedin and Bond 2006).
In this study, four new Conothele species are diagnosed and described based on the morphology of female specimens collected from mainland China and Laos, where the genus had not been reported before. Although, ideally both male and female characters should be included in the description of new species, in trap-door spiders obtaining males is extremely difficult and indeed we were unable to obtain male Conothele specimens in this study. Direct collection by searching and digging burrows primarily results in either females or immature specimens. Males are short lived and leave the burrow immediately after the adult moult to search for females (Haupt and Shimojana 2001;Haupt 2003). Therefore, collecting males is only possible at certain times of the year, and thus is not feasible during all collection trips.

Materials and methods
Specimens were examined under an Olympus SZX16 stereomicroscope, and photographed using an Olympus BX51 compound microscope. Genitalia were cleaned in boiling KOH for a few minutes to dissolve soft tissues. All the specimens were deposited at the Centre for Behavioural Ecology and Evolution (CBEE), College of Life Sciences, Hubei University, Wuhan, China. All measurements are in millimetres. Leg and palp measurements are given in the following order: total length (femur + patella + tibia + metatarsus + tarsus).
Abbreviations used are:

Genus Conothele Thorell, 1878
Type. Conothele malayana (Doleschall 1859): 5, pl. 5, figure 8 (described female). Diagnosis. The genus Conothele can be distinguished from the genus Ummidia by their burrowing habits, the former constructs a short, parallel to the surface of ground, superficial burrow, whereas the latter digs a several centimeters long burrow in the soil (Haupt 2006). Moreover, Conothele distributes in the Oriental and Australian regions, whereas Ummidia is only found from New World and the Mediterranean region (Haupt 2006;Siliwal et al. 2009;World Spider Catalog 2016). Etymology. 'Baiyun' refers to the type locality of this species. Diagnosis. Female of C. baiyunensis sp. n. can be distinguished from other species of Conothele by the slightly globular lobes of spermathecae in the terminal part; stalks with sclerotized and inward-bent distal part; stalk terminal parts relatively short, simple and direct (Fig. 1E).
Legs black brown, light-coloured ventrally, with long and short brown sparse setae, short thorn-like and normal spines. Tibia III with a saddle-like depression dorsally on the basal part (Fig. 1C). Metatarsus III with three prolateral spines. Femur III thickest. Scopulae and claw tufts absent on trasi of all palp and legs. Palpal claw with a single branched tooth; legs each with three tarsal claws, paired claws with two denticles. Leg formula: 4123. Measurements of palp and legs: palp 7.35 (2.80 + 1.25 + 1.75 + 1.55),  Opisthosoma black, scattered with thick and slender black setae. Spinnerets brownish yellow, PMS one-segmented, 0.60 long, PMS-PMS 0.10; PLS three-segmented, 1.85 long, covered with brown spines, apical segment dome-shape (Fig. 1D). Genitalia with paired slightly globular lobes of spermathecae in the terminal part, each stalk slender, long, at the distal part is sclerotized and bent, yet the bending is relatively short and looks more simple and direct (Fig. 1E).
Legs light brown, light-colored ventrally, with long and short brown sparse setae. All tarsi with tadpole-shaped trichobothrial hairs besides the normal ones. Basal part of tibia III with a saddle-like depression dorsally (Fig. 2C). Tibia and tarsus of palp, distal three segments of legs I and II with bands of short thorn-like spines laterally; tibia III with 4 short spines distally (Fig. 2C). Femur III thickest. Scopulae and claw tufts absent. Palpal claw with a single branched tooth; legs each with 3 tarsal claws, paired claws with two denticles. Opisthosoma black, scattered with thick and slender black setae. Spinnerets brownish, PMS one-segmented, 0.50 long, PMS-PMS 0.20; PLS three-segmented, 0.60 long, thicker and shorter (Fig. 2D). Genitalia with a pair of spermathecae, each stalk slender, long, broader towards the base, distally gradually sclerotized and incurved around 110°, terminating with face-to-face bowl-shaped lobes (Fig. 2E).
Distribution. Guangxi Province (Chongzuo), China. Etymology. 'Sidiechong' refers to the type locality of the holotype specimen of this species.

Conothele sidiechongensis
Diagnosis. Female genitalia of C. sidiechongensis sp. n. resembles to C. taiwanensis (Tso, Haupt & Zhu, 2003), but can be distinguished from the latter by more or less upwards oriented bowl-shape lobes and stalk bent in zigzag pattern distally (Fig. 3E).
Opisthosoma black, scattered with thick and slender black setae. Spinnerets brownish, PMS one-segmented, 1.20 long, PMS-PMS 0.10; PLS three-segmented, 2.50 long, thicker, covered with brown spines, apical segment dome-shape (Fig. 3D). Genitalia with a pair of spermathecae, with bowl-shaped lobes facing up at the terminal part, long stalks, slightly broader at the basal part, strongly sclerotized and bent in zigzag pattern at the distal part (Fig. 3E).
Distribution. Yunnan Province (Mojiang, Mengla), China; Oudomxay Province, Laos. Etymology. 'Yunding' refers to the type locality of this species. Diagnosis. Female of C. yundingensis sp. n. can be distinguished from C. daxinensis sp. n. by the slightly upwards and globular lobes terminally (Fig. 4E), can be distinguished from C. baiyunensis sp. n. by the distal part of stalks bent towards inside about 90° (Fig. 4E).