From Guizhou, China

Zootaxa 4701 (5): 434–442 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4701.5.3 http://zoobank.org/urn:lsid:zoobank.org:pub:5E5066BE-AD97-4D8D-82D2-0335311C28A5

Vappolotes, a new genus of coelotine spiders (Araneae, Agelenidae) from Guizhou, China

BING LI1, 2, ZHE ZHAO2, 3, YUANXUE CHEN2, HAIFENG CHEN1, ZHIYAN WU1, 3 & SHUQIANG LI2 1 Hebei Key Laboratory of Animal Diversity, College of Life Science, Langfang Normal University, Langfang 065000, China 2 Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China 3 Corresponding authors. Zhiyan Wu ([email protected]); Zhe Zhao ([email protected])

Abstract

A new genus of the subfamily Coelotinae F.O. Pickard-Cambridge, 1893, Vappolotes Zhao et S. Li gen. n., with two new species, V. ganlongensis Zhao et S. Li sp. n. (♂♀) and V. jianpingensis Zhao et S. Li sp. n. (♀), is described. The genus is restricted to southern China (Guizhou). Its relationship to other coelotine genera is discussed. A partial fragment of the mitochondrial cytochrome oxidase subunit I of both species were obtained to aid species identification at the molecular level.

Key words: Coelotinae, morphology, new species, taxonomy, biodiversity

Introduction

The spider family Agelenidae comprises 82 genera and 1323 valid species worldwide. The subfamily Coelotinae F.O. Pickard-Cambridge, 1893 comprises 32 genera and 58% of agelenid species. It is one of the best studied spider groups in East Asia, particularly in China, where more than half (~54%) of all coelotine species are found (World Spider Catalog, 2019). The group is well-studied on China because there have been major biodiversity programs in China in recent years (Li and Quan, 2017), and many new coelotine taxa have been reported from mountains and caves (include entrance and depth) (Chen et al., 2015a, 2015b; Chen et al., 2016a, 2016b; Li et al., 2018; Li et al.,2018a, 2018b; Li et al., 2019; Li et al., 2019a, 2019b; Zhang et al., 2016, 2017; Zhang and Zhao, 2017; Zhu et al., 2017). However, there are still many undescribed spider species in China, especially in the southern part. Herein, a new genus is proposed that includes two new species from caves in southern China (Guizhou). The genus Vappolotes gen. n. is quite different from other coelotines but has some morphological characters similar to Papiliocoelotes Zhao and Li (2016). Molecular data have indicated that Vappolotes gen. n. is the sister genus to Papiliocoelotes Zhao et Li, 2016 and close to the clade Longicoelotes Wang, 2002, Spiricoelotes Wang, 2002, Pla- tocoelotes Wang, 2002: ((((Vappolotes + Papiliocoelotes) Spiricoelotes) Longicoelotes) Platocoelotes) (Fig. 1).

Materials and methods

All specimens were collected from Guizhou Province. All type specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing. Specimens were examined with a LEICA M205 C stereomicroscope. Photos were captured at multiple focal ranges with an Olympus C7070 wide zoom digital camera (7.1 mega- pixels) mounted either on an Olympus SZX12 dissecting microscope or on an Olympus BX51 compound microscope, and the images were combined using Helicon Focus (Version 6.80) photo stacking software. The epigyne and male palps were dissected from the body for examination. The epigynum was treated in a warmed 10% potassium hydroxide (KOH) solution. Measurements were obtained with a LEICA M205 C stereomicroscope and are given in millimeters. Eye diameters were measured as the maximum diameter from either dorsal or frontal views. Leg measurements are given as: total length (femur, patella + tibia, metatarsus, tarsus). Photos of the left male palp are

434 Accepted by D. Harms: 16 Oct. 2019; published: 2 Dec. 2019 presented. The terminology of the structures follows Wang (2002) and Zhao and Li (2016). Figures from this paper are noted with an initial capital (Fig. or Figs) and figures from cited papers are listed in lowercase (fig. or figs). Abbreviations: A = atrium; ALE = anterior lateral eye; ALE–PLE = distance between ALE and PLE; AME = anterior median eye; AME–ALE = distance between AME and ALE; AME–AME = distance between AME and AME; AME–PME = distance between AME and PME; C = conductor; CD = copulatory duct; CDA = conductor’s dorsal apophysis; CF = cymbial furrow; E = embolus; EB = embolic base; FD = fertilization duct; H = hood; LC = lamella of conductor; PA = patellar apophysis; PLE = posterior lateral eye; PMA = posterior margin of atrium; PME = posterior median eye; PME–PLE = distance between PME and PLE; PME–PME = distance between PME and PME; RTA = retrolateral tibial apophysis; S = spermathecae; ST = subtegulum; T = tegulum; TA = tegular apophysis; TS = tegulum sclerite; VTA = ventrolateral tibial apophysis. In addition to morphological study, we reconstructed the phylogeny of coelotine genera based on molecular data from Zhao and Li (2017). DNA sequences were obtained for all specimens for delimiting and matching purposes. A partial fragment of the mitochondrial cytochrome oxidase subunit I (COI) gene was amplified and sequenced for Vappolotes gen. n., using the primers LCO1490-oono (5’-CWACAAAYCATARRGATA-TTGG-3’), HCO2198-zz (5’-TAAACTTCCAGGTGACCAAAAAATCA-3’) and C1-N-2776 (5’-GGATAATCAGAATANCGNCGAGG- 3’). See Zhao and Li (2017) for additional information on extraction, amplification, sequencing procedures, and the methods of tree construction. All sequences were checked using Basic Local Alignment Search Tool (BLAST) and are deposited in GenBank. The accession numbers are provided in Table 1.

Table 1. Voucher specimen information. Species Voucher code GenBank Sequence Collection localities accession number length V. ganlongensis sp. n. IZCAS-Ar39637 (ZZ550) KY778934 1194bp Miao Autonomous County of Songtao, Guizhou Province, China V. jianpingensis sp. n. IZCAS-Ar39645 (ZZ071) KY778933 562bp Zhengan County, Guizhou Prov- ince, China

Taxonomy

Family Agelenidae C.L. Koch, 1837

Subfamily Coelotinae F.O. Pickard-Cambridge, 1893

Genus Vappolotes Zhao et S. Li, gen. n.

Type species. Vappolotes ganlongensis Zhao et S. Li, sp. n. Etymology. The generic name is derived from the Latin “vappo”, meaning a variety of nocturnal moth. It refers both to the spiders being collected from the dark zone of a cave and to the copulatory ducts in the females which are shaped like the wings of a moth. The second part of the name “-loetes” comes from Coelotes Blackwall, 1841, referring to the similarity with the nominal genus of the subfamily. The gender is masculine. Diagnosis. The type species of the new genus is similar to Longicoelotes, Platocoelotes, Papiliocoelotes, and Spiricoelotes by the slender embolus (Fig. 2A, B), the small or absent median apophysis (Fig. 2C), and the lack of epigynal teeth (Figs 3A, 4A). The new genus is most similar to Papiliocoelotes with which it shares a bifurcated tegular apophysis (Fig. 2B, C). However, the male of the new genus can be easily distinguished from males of Papil- iocoelotes by having a dorsal apophysis and a lamella of the conductor (Fig. 2A–C), whereas in Papiliocoelotes the conductor has a slightly bifurcated distal process and lacks a lamella (fig.1A–C in Zhao and Li, 2016). Additionally, in Vappolotes, the tegular apophysis is bifurcated at the base (Fig. 2B, C), whereas in Papiliocoelotes it is bifurcated at the tip (fig. 2A, B in Zhao and Li, 2016). The female of the new genus can be easily distinguished from females of Papiliocoelotes by the bracketed lateral edges of the epigynum (Figs 3A, 4A); in Papiliocoelotes, the lateral edges are concave medially (fig. 2A in Zhao and Li, 2016). Also, in Vappolotes, the epigynal hoods are located anterolater- ally (Fig. 3A, 4A), whereas in Papiliocoelotes they are located mediolaterally (fig. 2A in Zhao and Li, 2016).

Vappolotes, New Coelotinae Genus From China Zootaxa 4701 (5) © 2019 Magnolia Press · 435 Description. Medium sized, with a total length of 5–6 mm. Body color yellow-brown generally, with black stripes (Fig. 3C–E); chelicerae with 3 promarginal and 2 retromarginal teeth in both sexes; carapace yellow-brown with radial pattern, pear-shaped (Fig. 3C, D); clypeus yellow-brown or brown (Fig. 3E); endites and labium brown (Fig. 3E); sternum yellow-brown (Fig. 3E); legs yellow-brown with the formula 4 > 1 > 2 > 3 (Fig. 3C–E); abdomen gray or yellow-brown, oval, with 3–4 chevrons posterodorsally (Fig. 3C, D); spinnerets yellow-brown (Fig. 3C–E). Male palp with one patellar apophysis, one retrolateral tibial apophysis (RTA), and a ventrolateral tibial apophysis (VTA) (Fig. 2C); median apophysis absent; cymbial furrow less than 1/2 the length of cymbium (Fig. 2C); conductor with a lamella (LC) and a dorsal apophysis (CDA) (Fig.2A, B); tegulum brown, sclerite of tegulum (ST) as long as embolic base (EB), and tegular apophysis bifurcated (Fig. 2A–C); subtegulum with many wrinkles (Fig. 2A); embolus filiform and long (Fig. 2A, B). Epigynum with deep anterolateral hoods (Figs 3A, 4A); atrium relatively large, with lateral and posterolateral margins conspicuous (Figs 3A, 4A); epigynal teeth and median septum absent; copulatory openings located posteromedially (Figs 3A, 4A); copulatory ducts broad and transparent, like the wings of a moth (Figs 3B, 4B); spermathecae nephroid, spermathecal bases close to each other (Figs 3B, 4B); fertilization ducts at least twice as long as wide (Figs 3B, 4B). Distribution. Guizhou, China (Fig. 5).

Figure 1. Simplified phylogenetic tree of coelotine spiders taken from Zhao and Li (2017). The numbers at the nodes show the Bayesian posterior probabilities and the Maximum Likelihood bootstrap values of correlated clades (with dots) respectively.

Vappolotes ganlongensis Zhao et S. Li, sp. n. Figs 2, 3, 5

Type material. Holotype: ♂ (IZCAS-Ar39637, ZZ550): Chenjia Cave (N28.38187º, E108.67368º, 792 m), Gan- long Town, Miao Autonomous County of Songtao, Guizhou Province, China, 9.III.2013, H.F. Zhao and J.C. Liu leg. Paratypes: 7♀♀ (IZCAS-Ar39638–Ar39644, ZZ550): same data as holotype. Etymology. The specific name refers to the type locality, Ganlong Town; adjective. Diagnosis. Vappolotes ganlongensis sp. n. can be distinguished from V. jianpingensis sp. n. by the following: the posterior margin of the atrium is concave medially (Fig. 3A), whereas it is arciform in V. jianpingensis sp. n. (Fig. 4A); the copulatory ducts are large, about 2 times larger as the spermathecae (Fig. 3B), whereas the copulatory ducts are nearly the same size as the spermathecae in V. jianpingensis sp. n. (Fig. 4B).

436 · Zootaxa 4701 (5) © 2019 Magnolia Press LI ET AL. Description. Male. Clypeus brown, abdomen gray, other parts are the same color as for the genus (Fig. 3C). Total length 5.76. Carapace 3.04 long, 2.19 wide. Abdomen 2.72 long, 1.75 wide. Eye sizes and interdistances: AME 0.11, ALE 0.16, PME 0.15, PLE 0.16; AME–AME 0.04, AME–ALE 0.02, AME–PME 0.04, ALE–PLE 0, PME–PME 0.06, PME–PLE 0.05. Leg measurements: I 12.24 (3.15, 4.06, 2.94, 2.09); II 11.52 (3.19, 3.52, 2.93, 1.88); III 11.11 (2.67, 3.46, 3.05, 1.93); IV 14.41 (3.71, 4.39, 4.12, 2.19). Palp: Femur 4.5 times longer than wide; patella with a long anterodorsal macroseta, almost 2.3 times longer than the patella, patellar apophysis about 1/2 the patella length, with blunt tip; tibia and base of cymbium with long dorsal and prolateral macrosetae; VTA and RTA spine-shaped, VTA about 3 times longer than RTA, extending beyond the distal margin of the tibia; cymbium 2.2 times longer than bulb, cymbial furrow short, less than 1/2 the cymbial length; dorsal apophysis of the conductor pointed, and the large lamella of conductor spoon-shaped; tegular apophysis shorter than 1/2 the cymbial furrow, with bifurcated tip; embolus originates around the 6 o’clock position (Fig. 2A–C). Female (IZCAS-Ar39638). Same color as male except abdomen yellow-brown (Fig. 3D, E). Total length 5.95. Carapace 2.56 long, 1.81 wide. Abdomen 3.39 long, 2.34 wide. Eye sizes and interdistances: AME 0.10, ALE 0.14, PME 0.12, PLE 0.13; AME–AME 0.05, AME–ALE 0.03, AME–PME 0.05, ALE–PLE 0, PME–PME 0.07, PME–PLE 0.06. Leg measurements: I 9.04 (2.59, 2.89, 2.11, 1.45); II 8.77 (2.54, 2.78, 2.06, 1.39); III 8.68 (2.41, 2.72, 2.24, 1.31); IV 11.98 (3.35, 3.67, 3.31, 1.65). Epigynum: about 1.7 times wider than long; atrium occupying 3/4 of epigynum, length equal to width, posterior lateral margin of atrium about 1/2 the width of the lateral margin, concave medially; copulatory ducts large, about 3 times larger than spermathecae; spermathecae small, occupying 1/4 of epigynum, over 2 times longer than wide; fertilization ducts 2 times longer than wide (Fig. 3A, B). Variation. Total length of females 5.72–5.96 (n=7). Distribution. Guizhou, China (Fig. 5).

Figure 2. Left male palp of Vappolotes ganlongensis sp. n. A, Prolateral view; B, Ventral view; C, Retrolateral view. Scale bar: equal for A, B and C.

Vappolotes, New Coelotinae Genus From China Zootaxa 4701 (5) © 2019 Magnolia Press · 437 Figure 3. Epigynum and habitus of Vappolotes ganlongensis sp. n. A, Epigynum, ventral; B, Internal genitalia, dorsal; C, Male habitus, dorsal; D, Female habitus, dorsal; E, Female habitus, ventral. Scale bars: equal for A and B, equal for C, D and E.

438 · Zootaxa 4701 (5) © 2019 Magnolia Press LI ET AL. Figure 4. Epigynum and habitus of Vappolotes jianpingensis sp. n. A, Epigynum, ventral; B, Internal genitalia, dorsal; C, Female habitus, dorsal; D, Female habitus, ventral; E, Female habitus, lateral. Scale bars: equal for A and B, equal for C, D and E.

Type material. Holotype: ♀ (IZCAS-Ar39645, ZZ071): Xinlong Cave (N28.51247º, E107.46134º, 521 m), Jian- ping Township, Zheng’an County, Zunyi City, Guizhou Province, China, 5.VIII.2012, H.F. Zhao leg. Paratypes: 5♀♀ (IZCAS-Ar39646–Ar39650, ZZ071): same data as holotype. Etymology. The specific name refers to the type locality, Jianping Township; adjective. Diagnosis. Vappolotes jianpingensis sp. n. can be distinguished from V. ganlongensis sp. n. by the following: the posterior and lateral atrium margins are the same width (Fig. 4A), whereas the posterior margin of the atrium is 1/2 as wide as the lateral margin in V. ganlongensis sp. n. (Fig. 3A); the size of the spermathecae is subequal to the size of the copulatory ducts (Fig. 4B), whereas the spermathecae are approximately 1/2 the size of the copulatory ducts in V. ganlongensis sp. n. (Fig. 3B). Description. Female (IZCAS-Ar39645). Clypeus yellow-brown, abdomen yellow-brown, other parts the same color as for the genus (Fig. 4C–E). Total length 5.44. Carapace 2.56 long, 1.64 wide. Abdomen 2.88 long, 2.24 wide. Eye sizes and interdistances: AME 0.10, ALE 0.13, PME 0.12, PLE 0.14; AME–AME 0.04, AME–ALE 0.04, AME–PME 0.08, ALE–PLE 0, PME–PME 0.08, PME–PLE 0.10. Leg measurements: I 9.16 (2.66, 2.87, 2.09, 1.54); II 8.25 (2.23, 2.45, 2.18, 1.39); III 7.84 (2.16, 2.54, 1.96, 1.18); IV 9.69 (2.54, 3.48, 2.14, 1.53). Epigynum: about 1.5 times wider than long; atrium large, occupying 2/3 of epigynum, length subequal to width, width of the posterolateral margin equal to the width of the lateral margin; copulatory ducts large, the same size as the spermathecae; spermathecae occupying 1/2 of the epigynum, about 2 times longer than wide; fertilization ducts 2.5 times longer than wide (Fig. 4A, B). Male. Unknown. Variation. Total length of females 5.34–5.53 (n=6). Distribution. Guizhou, China (Fig. 5).

Figure 5. Localities of new species of Vappolotes gen. n. from Guizhou, China. 1, V. ganlongensis sp. n.; 2, V. jianpingensis sp. n.

The manuscript benefitted greatly from comments by Danilo Harms, Nadine Dupérré, and another anonymous reviewer. Sarah Crews kindly checked English, and Francesco Ballarin checked the Latin names. This study was supported by the National Natural Sciences Foundation of China (NSFC–31772418, 31530067) and the program of Youth Innovation Promotion Association of Chinese Academy of Sciences (No. 2019087).

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