Three new species of the genus Coddingtonia from Asia (Araneae, Theridiosomatidae)

Abstract The current paper expands knowledge of the genus Coddingtonia Miller, Griswold & Yin, 2009. Based on morphological characters and molecular data, three species are documented as new to science: C. erhuan Feng & Lin, sp. nov. (♀) from China, C. lizu Feng & Lin, sp. nov. (♀) from China, and C. huifengi Feng & Lin, sp. nov. (♂♀) from Indonesia. The type of C. euryopoides Miller, Griswold & Yin, 2009 is also reexamined. DNA sequences (COI), detailed illustrations of habitus, male palp and epigyne are provided for these four species, as well as a key and a distribution map for Coddingtonia species.


Introduction
Coddingtonia was originally established by Miller et al. (2009) as a monotypic genus based on C. euryopoides Miller et al., 2009 from the Gaoligong Mountains in Southwest China. Labarque and Griswold (2014) reported two Coddingtonia species from Laos and Malaysia. Currently the genus Coddingtonia contains three valid species distributed in China, Laos, Thailand, and Malaysia (Lopardo and Hormiga 2015;World Spider Catalog 2019).
In a recent collection of theridiosomatids from China and Indonesia, we found three members of Coddingtonia and propose them as species new to science. Detailed diagnoses, descriptions, and identifying illustrations are provided for each. This work also represents the first record of this genus from Indonesia.

Materials and methods
All specimens were preserved in 95% ethanol. Specimens were examined and measured with a Leica M205 C stereomicroscope. Further details were studied using an Olympus BX53 compound microscope mounted with a Canon EOS 60D wide zoom digital camera (8.5 megapixels). Male and female copulatory organs were examined and photographed after they were dissected and detached from the bodies. Vulvae were treated with lactic acid before being photographed. The digital images were montaged using Helicon Focus 3.10 image stacking software (Khmelik et al. 2006). All measurements in the paper are in millimeters. Leg measurements are given in the following sequence: total length (femur, patella, tibia, metatarsus, and tarsus A partial fragment (636 bp) of the mitochondrial gene cytochrome c oxidase subunit I (COI) was amplified and sequenced in order to check the genetic distance between morphologically close related species and confirm identifications and the sex pairing accuracy. For the same reasons, sequences of Coddingtonia euryopoides Miller et al., 2009 were also included.
The primers used are as following: LCO1490 (5'-GGTCAACAAATCATCAT-AAAGATATTGG-3') and HCO2198 (5'-TAAACTTCAGGGTGACCAAAAAA TCA-3'). Raw sequences were edited and assembled using BioEdit v.7.2.5 (Hall 1999) and the uncorrected pairwise distance between the species was calculated using MEGA7.0.14 (Kumar et al. 2016). All sequences were incorporated in GenBank and the accession numbers are provided in Table 1. Results of the comparison between the genetic distances are shown in Table 2.
All examined materials are deposited in the Natural History Museum of Sichuan University in Chengdu (NHMSU), China, except the holotype of C. euryopoides, which is deposited in the School of Life Sciences, Hunan Normal University in Changsha (HNU), China.  (Coddington, 1986), long and coiled copulatory ducts surrounding the spermathecae, but lacking that in other theridiosomatids ( Distribution. Southern China (Yunnan, Hainan), Laos, Thailand, Malaysia, and Indonesia (Fig. 6). Diagnosis. The male of C. euryopoides differs from the males of other species by the mesal bristle of the embolic apophysis describing a semi-loop very close to the embolus base and a semi-loop around the bulb, and the straight median apophysis having a tapering tip (Lopardo and Hormiga 2015: fig. 124B, E, F). The female of C. euryopoides can be distinguished from the other five species by having 9 coils of the copulatory ducts (Fig. 1E), whereas other species have fewer coils. Moreover, C. euryopoides differs from C. anaktakun, C. discobulbus, and C. huifengi sp. nov. by having a posterior tubercle on the abdomen (Fig. 1A-C), whereas this tubercle is absent in the latter three species.
Description. See Fig. 1A-F and Miller et al. (2009: 30). Male of this species remains unknown.
* Only referring to characters of the vulva  Diagnosis. The male of this new species differs from the male of C. euryopoides by the median apophysis with a distal flexible hook, and the narrower, shorter conductor (Fig. 3A, D); in other similar species the tip is straight and wider and conductor is longer (see Labarque and Griswold 2014: figs 1C, 5D-F). The female can be distinguished from the other five species by having 3 coils (one thick, two thin) of copulatory ducts (Fig. 2F), whereas they are fewer or more in other species. Moreover, C. huifengi differs by the lack of a posterior tubercle on the abdomen ( Fig. 2A-D) vs. present in C. euryopoides, C. erhuan sp. nov., and C. lizu sp. nov. (Figs 1A-C, 4A-C, 5A, B).
Description. Females (holotype). Carapace nearly pentagonal, dim yellowish, cephalic area moderately raised. Anterior eye row precurved, posterior eye row straight. Sternum heart-shaped, grey yellow, with sparse setae. Mouthparts brown. Femora and patellae dim yellow, other segments brown. Abdomen round, dorsally grey, ventrally deeper, bears sparse long hairs, weakly ossified at hair base ( Fig. 2A, B) Epigyne (Fig. 2E-G): epigyne covered with sparse black setae in the central region; with deep central pit and 2 longitudinal grooves close to lateral margins of the plate. Spermathecae barely visible through the integument; LW well developed, like a pair of boxing gloves, swollen sacks with dorso-median glandular ducts; spermathecae globular, separated by one radius; copulatory ducts form an expanded posterolateral loop, and coiled into 2 slender posteromedian loops, finally connecting ventrally on the spermathecae; fertilization ducts arise from the dorsomesal the spermathecae.
Male (one paratype): Somatic features as in Fig. 2A   Palp (Fig. 3A-D): tibia small, cymbium narrow, about 2 times longer than width, with long setae; paracymbium short and small, about of 1/5 cymbial length; tegulum capacious; median apophysis lamellar, subrectangular; conductor disk shaped with a needle-like distal process; mesal bristle of the embolic apophysis describes a semi-loop above the tegulum and cymbium; embolus long, whip-like, extending far beyond the mesial embolic apophysis and coiling into one loop.
Male. unknown. Distribution. Known only from the type locality (Fig. 6). Diagnosis. This new species can be distinguished from the congeners by having 5 loops of unilateral copulatory duct ( Fig. 5D; Note: the broken first and fourth loops on the right side of copulatory duct in vulva are due to careless dissection). Moreover, it has a posterior tubercle on the abdomen (Fig. 5A, B), which is absent in C. anaktakun, C. discobulbus, and C. huifengi sp. nov. Description. Female (holotype): Carapace pear-shaped, black. Sternum dim, posteriorly contracted. Femora and tibiae of legs dark, other segments yellow to brown. Abdomen dark black, dorsal color lighter than venter, with posterior tubercle, covers sparse long, stiff setae (Fig. 5A, B) Epigyne (Fig. 5C-E): weakly sclerotized, nearly rectangular, black pigmentation in the central region; central pit and lateral pockets indistinct. lateral wings well developed, reniform and translucent; spermathecae small and round, separated by approximately one radius; copulatory ducts form a posterolateral auricular loop on the both sides of the lateral wings, followed by 5 loops, and finally connecting ventrally on the spermathecae (Fig. 5D); fertilization ducts short, arise from the dorsal-inner base of spermathecae (Fig. 5E).