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Research Article
A review of Crassignatha (Araneae, Symphytognathidae)
expand article infoYa Li, Yucheng Lin, Shuqiang Li§
‡ Sichuan University, Chengdu, China
§ Institute of Zoology, Chinese Academy of Sciences, Beijing, China
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Abstract

Crassignatha Wunderlich, 1995 is redefined to include species with six eyes in three diads, chelicerae fused only near the base, sculpturing on the carapace, one or two clasping spurs on tibia II, a bilateral scutum of the male abdomen, and globular spermathecae and adjacent copulatory openings in the female. A key and distribution map are provided for 24 Crassignatha species in this paper. Diagnoses and illustrated photographs are provided for 22 species from China, Malaysia, Thailand, and Vietnam. Thirteen species are described and documented as new to science: C. baihua Y. Lin & S. Li, sp. nov. (♂♀), C. bangbie Y. Lin & S. Li, sp. nov. (♀), C. changyan Y. Lin & S. Li, sp. nov. (♀), C. dongnai Y. Lin & S. Li, sp. nov. (♀), C. gucheng Y. Lin & S. Li, sp. nov. (♂♀), C. mengla Y. Lin & S. Li, sp. nov. (♂♀), C. nantou Y. Lin & S. Li, sp. nov. (♂♀), C. nasalis Y. Lin & S. Li, sp. nov. (♂♀), C. rostriformis Y. Lin & S. Li, sp. nov. (♂♀), C. shunani Y. Lin & S. Li, sp. nov. (♂♀), C. si Y. Lin & S. Li, sp. nov. (♂♀), C. thamphra Y. Lin & S. Li, sp. nov. (♀), and C. xichou Y. Lin & S. Li, sp. nov. (♀). Three new combinations are proposed: C. bicorniventris (Lin & Li, 2009), comb. nov., C. quadriventris (Lin & Li, 2009), comb. nov., and C. shiluensis (Lin & Li, 2009), comb. nov. are transferred from Patu Marples, 1951. DNA barcodes and genetic distances of seventeen species are obtained to confirm correct identification. Types of seven known Chinese Crassignatha species are re-examined, and the taxonomic placement of C. longtou Miller, Griswold & Yin, 2009 may be incorrect based on morphological and molecular data.

Keywords

Asia, barcode, new combination, new species, redescription, symphytognathids, taxonomy

Introduction

Symphytognathidae Hickman, 1931 is a category of super miniature (body size ca. 1 mm or less), poorly known, araneoid spiders that contains eight genera and 74 documented extant species (WSC 2020; Li 2020), but no fossil species have been recorded to date (Dunlop et al. 2020). With the exception of Antarctica, they are found on all continents and some oceanic islands, distributed in the tropical and subtropical regions. They are cryptozoic, commonly found in the leaf litter layer of forests, and some species inhabit dark caves (Cardoso and Scharff 2009; Lin and Li 2009; Miller et al. 2014).

Crassignatha Wunderlich, 1995 was erected as a monotypic genus and originally placed in Synaphridae Wunderlich, 1986. The phylogenetic analysis of Lopardo and Hormiga (2015) corroborated its placement within Symphytognathidae. Since its inception, the genus has included nine described species, which are only found in the Gaoligong Mountains of southwest China (Miller et al. 2009), Malaysia (Wunderlich, 1995; Miller et al. 2014). The distribution of Crassignatha is currently confined to Southern China and Southeast Asia. There has been no worldwide or regional taxonomic revision of the genus.

From 2006 to 2018, we have accumulated a considerable number of symphytognathid specimens during many field collecting trips in China, Vietnam, Thailand, Myanmar, and Indonesia. Some of these specimens were described as new species (Lin and Li 2009; Lin et al. 2009; Lin et al. 2013; Lin 2019). However, further morphological and molecular studies of this material have revealed an extraordinary species diversity in the aforementioned geographic areas.

The main aim of this paper is to provide a comprehensive overview of Crassignatha and to report 22 species which are found in Southern China and the Indo-China Peninsula. The circumscription and diagnosis of Crassignatha are reconfirmed. Thirteen species are described and designated as new members of Crassignatha, as well as three new combinations that are transferred from Patu Marples, 1951. The types of seven known Crassignatha species from the Gaoligong Mountains in Southwest China are re-examined. A key and distribution records are provided for all Crassignatha species.

Materials and methods

Most of the specimens from this study were collected by hand or sifting leaf litter and immediately preserved in a 95% ethanol solution. Type specimens of other known Chinese Crassignatha species were borrowed from the Life College of Sciences, Hunan Normal University in Changsha (HNU), China and the Institute of Zoology, Chinese Academy of Sciences in Beijing (IZCAS), China. All materials were examined using a Leica M205 C stereomicroscope and photographed with a Canon EOS 60D wide zoom digital camera (8.5 megapixels) mounted on an Olympus BX 43 compound microscope. Male palps and epigynes were examined and photographed after dissection. The left palp was photographed and described unless it is missing, then the right was selected. Epigynes were treated with lactic acid before being embedded in Hoyer’s gum to take photos of the vulva. The images were montaged using Helicon Focus 3.10 (Khmelik et al. 2006) image stacking software. All measurements are in millimeters. Leg measurements are given as follows: total length (femur, patella, tibia, metatarsus, and tarsus).

Tissue samples were taken from twenty-eight individuals of Crassignatha, including eleven new and six known species. Molecular data were obtained from specimens collected at the type locality, although not from the type specimens themselves. A partial fragment (636 bp) of the mitochondrial gene cytochrome c oxidase subunit I (COI) was amplified and sequenced to calculate the genetic distances between morphologically similar species and to confirm identifications and sex pairing accuracy.

The primers used are: LCO1490 (5'-GGTCAACAAATCATCATAAAGATATTGG-3') and HCO2198 (5'-TAAACTTCAGGGTGACCAAAAAATCA-3'). Raw sequences were edited and assembled using BioEdit v.7.2.5 (Hall 1999), and the uncorrected pairwise distances between species were calculated using MEGA7.0.14 (Kumar et al. 2016). Results of the genetic distance analysis are shown in Appendix 1.

Abbreviations used in the text or figures are given in Table 1. References to figures in the cited papers are in lowercase (fig. or figs), figures in this paper are noted with an initial capital (Fig. or Figs). New sequences generated for this study are available in GenBank, and the accession numbers are reported in Table 2. With the exception of the types of previously described species kept in HNU and IZCAS, all molecular vouchers are tentatively deposited in NHMSU in Chengdu, China, and examined morphological material is deposited in NHMSU and IZCAS.

Table 1.

List of abbreviations used in the text or figures.

Male palp Epigyne
C conductor CD copulatory duct
CB cymbium CO copulatory opening
CT cymbial tooth FD fertilization ducts
E embolus S spermathecae
EM embolic membrane Sp scape
Fe femur Ocular area
MA median apophysis ALE anterior lateral eye
Pa patella PLE posterior lateral eye
T tegulum PME posterior median eye
Ti tibia PER posterior eye row
TS tibial spur on leg II
Institutions
HNU College of Life Sciences, Hunan Normal University, Changsha, China
IZCAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China
NHMSU Natural History Museum of Sichuan University, Chengdu, China
Table 2.

GenBank accession numbers for new DNA sequence data from seventeen Crassignatha species.

Species Identifier Sex/Stage COI Collection localities
Crassignatha baihua sp. nov. HA109 ♂/adult MT992007 China, Yunnan, Longling Co., Baihualing Village
HA109 ♀/adult MT992006
Crassignatha bangbie sp. nov. HA137 ♀/adult MT992015 China, Yunnan, Longling Co., Bangbie Village
Crassignatha dongnai sp. nov. HA092 ♀/adult MT992004 Vietnam, Dong Nai Pro., Cat Tien National Park
Crassignatha ertou HA108 ♀/adult MT992005 China, Yunnan, Baoshan, Mangkuan Town, Baihualing
Crassignatha gucheng sp. nov. HA132 ♀/adult MT992014 China, Yunnan, Longling Co., Mt. Xiaohei Nature Reserve
Crassignatha mengla sp. nov. HA080 ♂/juvenile MT992000 China, Yunnan, Mengla Co., Baca Nature Reserve
HA080 ♀/adult MT991999
Crassignatha nantou sp. nov. HA055 ♂/adult MT991996 China, Taiwan, Nantou Co., Hehuan Hill
HA055 ♀/adult MT991995
Crassignatha nasalis sp. nov. HA041 ♂/adult MT991992 China, Sichuan, Guling Co., Yuhua Town, Taoyuan Cave
HA041 ♀/adult MT991991
Crassignatha pianma HA113 ♂/adult MT992009 China, Yunnan, Lushui Co., Pianma Town, broad-leaf forest
HA113 ♀/adult MT992008
Crassignatha quadriventris HA025 ♀/adult MT991990 China, Hainan, Dongfang City, Nanlang Village, E’xianling
Crassignatha rostriformis sp. nov. HA079 ♂/adult MT991998 China, Yunnan, Xichou Co., Xianren Cave
HA079 ♀/adult MT991997
Crassignatha shiluensis HA081 ♂/juvenile MT992002 China, Yunnan, Mengla Co., Xishuangbanna Botanical Garden
HA081 ♀/adult MT992001
Crassignatha shunani sp. nov. HA046 ♂/juvenile MT991994 China, Sichuan, Guling Co., Dahei Cave
HA046 ♀/adult MT991993
Crassignatha si sp. nov. HA141 ♂/juvenile MT992017 China, Yunnan, Yiliang Co., Dazhezong Village, Baiyan Cave
HA141 ♀/juvenile MT992016
Crassignatha thamphra sp. nov. HA089 ♀/adult MT992003 Thailand, Khon Kaen Pro., Phu Pha Man Distr., Tham Phra Cave
Crassignatha yamu HA115 ♂/adult MT992011 China, Yunnan, Fugong Co., Shilajia Village, Yamu River
HA115 ♀/adult MT992010
Crassignatha yinzhi HA117 ♂/adult MT992013 China, Yunnan, Longling Co., Mt. Xiaohei Nature Reserve
HA117 ♀/adult MT992012

Taxonomy

Family Symphytognathidae Hickman, 1931

Crassignatha Wunderlich, 1995

Crassignatha Wunderlich, 1995: 546

Crassignatha Miller et al., 2009: 68

Type species

Crassignatha haeneli Wunderlich, 1995 by original designation, from Malaysia.

Diagnosis

Crassignatha can be distinguished from Anapistula Gertsch, 1941 by having six eyes vs. four or absent in the latter; and from Anapogonia Simon, 1905 by the chelicerae fused near the base vs. unfused. (The latter is tentatively placed in Symphytognathidae (Plantnick and Forster 1989: 76)). Crassignatha differs from Globignatha Balogh & Loksa, 1968 and Symphytognatha Hickman, 1931 by the chelicerae fused only near the base vs. almost fully fused in the latter two (Balogh and Loksa 1968: fig. 10; Forster and Platnick 1977: fig. 41; Lin 2019: fig. 1H). Crassignatha is most similar to Curimagua Forster & Platnick, 1977 and Patu Marples, 1951 in habitus features and body size but differs from Curimagua by having six eyes in diads and lacking female palps rather than eyes in triads and female palps reduced to remnants but not absent (Fig. 1A, D vs. Forster and Platnick 1977: figs 40, 63); and from Patu by the sculptured carapace (Fig. 16A, D; Wunderlich 1995: fig. 15; smooth in a few species) and the male abdomen usually with a lateral scutum (Fig. 1C; absent in a few species).

Description

Body length 0.50–0.90 in male, 0.60–1.30 in female; six eyes in three diads. Ocular area in male raised more than in female. Carapace sub-rounded or pear shaped, brown or yellow-brown, usually sculptured on surface, but smooth in a few species. Cervical groove distinct. Clypeus concave. Chelicerae usually fused near base, with one or two retromarginal teeth. Labium triangular or semilunar, fused to sternum. Sternum scutellate or heart shaped, slightly plump, surface mostly sculptured, rarely smooth, truncated posteriorly. Legs pale yellow to brown-yellow. Leg formula: I-II-IV-III or I-VI-II-III. Male tibia II usually with two long clasping spurs on ventral-subdistal part (but only one spur in a few species). Abdomen globular or quadrate posteriorly in both sexes, male usually with weakly sclerotized abdominal scutum laterally and posteriorly (absent in few species), with an annular plate around spinnerets. Colulus absent.

Male palps oblate. Cymbium wraps around bulb on the prolateral-ventral surface, with a distal cymbial tooth. Median apophysis present, conductor absent. Embolus sclerotized, usually attached to a transparent embolic membrane at base.

Female genital area weakly sclerotized, internal structure faintly visible through tegument. Majority of species with protruded scape, copulatory opening located at apex of scape. Paired spermathecae globular, separated. Copulatory ducts tortile, usually connected to the posterolateral or dorsal surface of spermathecae. Fertilization ducts usually starting at the posterior or lower inner surface of spermathecae.

Composition

Crassignatha baihua sp. nov., C. bangbie sp. nov., C. bicorniventris (Lin & Li, 2009), C. changyan sp. nov., C. danaugirangensis Miller et al., 2014, C. dongnai sp. nov., C. ertou Miller, Griswold & Yin, 2009, C. gucheng sp. nov., C. gudu Miller, Griswold & Yin, 2009, C. haeneli Wunderlich, 1995, C. mengla sp. nov., C. nantou sp. nov., C. nasalis sp. nov., C. pianma Miller, Griswold & Yin, 2009, C. quadriventris (Lin & Li, 2009), C. quanqu Miller, Griswold & Yin, 2009, C. rostriformis sp. nov., C. shiluensis (Lin & Li, 2009), C. shunani sp. nov., C. si sp. nov., C. thamphra sp. nov., C. xichou sp. nov., C. yamu Miller, Griswold & Yin, 2009, and C. yinzhi Miller, Griswold & Yin, 2009. Patu bispina Lin, Pham & Li, 2009, and P. kishidai Shinkai, 2009 may also belong in this genus.

Distribution

Southern China (Guizhou, Yunnan, Hainan, and Taiwan), Central Japan (Honshu, Shikoku), Vietnam, Thailand, Malaysia.

Key to species of Crassignatha Wunderlich, 1995

1 Males 2
Females 18
2 Embolus long, extending beyond anterior edge of median apophysis (Figs 8B, 15B, 27A, 31B, 35B) 3
Embolus short, not extending beyond anterior edge of median apophysis (Figs 10B, 21B, 23B) 13
3 Embolus filiform, flexible (Figs 27A, 35B, 37A, B) 4
Embolus spiraled, stiff (Figs 8B, 15B, 31B) 8
4 Embolus coiled (Figs 27A, 35B) 5
Embolus not coiled (Fig. 37A; Wunderlich 1995: fig. 19; Miller et al. 2014: fig. 4) 6
5 Embolus coiled into more than two loops (Fig. 27A) C. shiluensis
Embolus coiled into fewer than two loops (Fig. 35B) C. yamu
6 Embolus filiform, without basal nodule 7
Embolus straight, pointed, with a basal nodule (Fig. 37A, B) C. yinzhi
7 Median apophysis bilobate (Wunderlich 1995: fig 18) C. haeneli
Median apophysis trilobate (Miller et al. 2014: fig. 4) C. danaugirangensis
8 Embolus twisted anticlockwise (Figs 8B, 15B, 25B) 9
Embolus twisted clockwise (Fig. 31B) C. si sp. nov.
9 Cymbial tooth large, hook shaped (Figs 8B, 25B) 10
Cymbial tooth small, tooth-like (Figs 15B, 17A, 29A) 11
10 Embolic base narrow (Fig. 8B)…. C. ertou
Embolic base wide (Fig. 25B) C. rostriformis sp. nov.
11 Embolic tip blunt, stiff (Fig. 17B) C. nasalis sp. nov.
Embolic tip sharp, narrow (Figs 15A, B, 29A, B) 12
12 Cymbial tooth sharp, median apophysis lacks a hook (Fig. 29A) C. shunani sp. nov.
Cymbial tooth blunt, median apophysis with a hook (Fig. 15A) C. nantou sp. nov.
13 Embolic tip blunt (Figs 10B, 19B, 23B) 14
Embolic tip pointed (Figs 2B, 13B, 21B) 16
14 Embolic base wide; cymbial tooth spur-like (Figs 19A, B, 23A, B) 15
Embolic base narrow; cymbial tooth spine-like (Fig. 10A, B) C. gucheng sp. nov.
15 Median apophysis subquadrate, with a hooked process (Fig. 19A, B) C. pianma
Median apophysis subtriangular, with a truncated process (Fig. 23A, B) C. quanqu
16 Embolic tip narrow; median apophysis with a process (Figs 2B, 21B) 17
Embolic tip wide; median apophysis lacks a process (Fig. 13A, B) C. mengla sp. nov.
17 Embolic apex flat (Fig. 2B) C. baihua sp. nov.
Embolic apex sloped (Fig. 21A, B) C. quadriventris
18 Scape long, distinctly protrudes from epigastric furrow (Figs 2D, 6D, 10D) 19
Scape short or absent (Figs 4D, 5D, 27C, 33D) 27
19 Copulatory ducts merged into a tubular atrium medially (Figs 11F, 35G) 20
Copulatory duct junction near copulatory opening (Figs 2F, 8F, 10F) 22
20 Copulatory atrium short, does not extend beyond spermathecal anterior margin 21
Copulatory atrium long, extends beyond spermathecal anterior margin (Fig. 31F) C. si sp. nov.
21 Copulatory ducts do not overlap atrium (Fig. 11F, G) C. gudu
Copulatory ducts overlap part of atrium (Fig. 35F, G) C. yamu
22 Copulatory ducts form a V-shape before junction (Figs 8F, 37F) 23
Copulatory ducts nearly parallel at center before junction (Figs 2F, 21F) 24
23 Copulatory ducts make two sharp turns (Fig. 37F) C. yinzhi
Copulatory ducts make four sharp turns (Fig. 8F) C. ertou
24 Copulatory duct connects to spermathecal dorsum (Figs 6G, 10G) 25
Copulatory duct connects to posterior margin of spermathecae (Fig. 2G, 21G) 26
25 Copulatory ducts twisted twice in center of vulva (Fig. 10F, G) C. gucheng sp. nov.
Copulatory ducts twisted once in center of vulva (Fig. 6F, G) C. dongnai sp. nov.
26 Copulatory ducts make four turns (Fig. 2F, G) C. baihua sp. nov.
Copulatory ducts make six turns (Fig. 21F, G) C. quadriventris
27 Scape absent (Figs 4D, 17D, 27C, 33D) 28
Scape present (Figs 3E, 19E, 29E, 32E, Miller et al. 2014: fig. 2) 33
28 Copulatory duct loops 3 ×, connects anterolaterally to spermathecae (Fig. 27D, E) C. shiluensis
Vulva not as above 29
29 Spermathecae separated by at least their diameter (Figs 5E, 33E) 30
Spermathecae separated by less than their diameter (Figs 4F, 15F, 17E) 31
30 Copulatory ducts vertically linked to copulatory opening (Fig. 33E) C. xichou sp. nov.
Copulatory ducts diagonally linked to copulatory opening (Fig. 5E) C. changyan sp. nov.
31 Copulatory ducts vertical and parallel in center of vulva (Fig. 15F) C. nantou sp. nov.
Copulatory ducts not as above 32
32 Copulatory ducts twisted 2× (Fig. 17E, F) C. nasalis sp. nov.
Copulatory ducts twisted 4× (Fig. 4F, G) C. bicorniventris
33 Spermathecae separated by less than 1.5× their diameter 34
Spermathecae separated by more than 3× their diameter (Miller et al. 2014: fig. 2) C. danaugirangensis
34 Copulatory ducts nearly vertically linked to copulatory opening (Figs 13F, 29F) 35
Copulatory ducts diagonally or horizontally linked to copulatory opening 37
35 Proximal part of copulatory ducts not confluent (Fig. 29F) C. shunani sp. nov.
Copulatory ducts confluent before reaching copulatory opening (Figs 13F, 25F) 36
36 Spermathecae separated by their diameter (Fig. 25F) C. mengla sp. nov.
Spermathecae spacing does not exceed their diameter (Fig. 32G) C. rostriformis sp. nov.
37 Copulatory ducts horizontally linked to copulatory opening 38
Copulatory ducts diagonally linked to copulatory opening 39
38 Copulatory duct has two inflection points in middle of vulva (Fig. 19F) C. pianma
Copulatory duct has one inflection point in middle of vulva (Fig. 3F) C. bangbie sp. nov.
39 Proximal copulatory ducts curved (Fig. 23D) C. quanqu
Proximal copulatory ducts straight (Fig. 32F) C. thamphra sp. nov.

Crassignatha baihua Y. Lin & S. Li, sp. nov.

Figs 1, 2, 38

Type material

Holotype ♂ (NHMSU Ar 001) and paratypes 3♂ 8♀ (NHMSU Ar 002–012), China: Yunnan Province, Longling County, Mangkuan Township, Zaotanghe at Baihua Ling (Flowers Ridge) Village, in leaf litter under undisturbed subtropical broadleaf forest (25.30450°N, 98.80059°E; 1635 m), 21.VIII.2018, Y. Lin et al. leg. 1♂ (NHMSU-HA109) and 1♀ (NHMSU-HA109) used for sequencing, GenBank: MT992007 and MT992006, same data as for preceding.

Figure 1. 

Crassignatha baihua sp. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Other material examined

1♂ 13♀ (NHMSU-HA110), same data as holotype; 1♂ 2♀ (NHMSU-HA104), China: Yunnan Province, Nujiang Prefecture, Fugong County, Shiyueliang Township (27.27012°N, 98.89803°E; 1647 m), 5.VII.2016, Y. Li leg.

Diagnosis

Crassignatha baihua sp. nov. is similar to C. quadriventris but can be distinguished by the short, rigid, distally flat embolus (Fig. 2A) and the long copulatory ducts that make four turns before reaching the copulatory opening (Fig. 2F, G).

Description

Male (holotype). Total length 0.64. Carapace 0.32 long, 0.32 wide, 0.36 high. Clypeus 0.10 high. Sternum 0.24 long, 0.20 wide. Abdomen 0.44 long, 0.44 wide, 0.48 high. Length of legs: I 0.94 (0.20, 0.10, 0.28, 0.16, 0.20); II 0.84 (0.16, 0.10, 0.24, 0.14, 0.20); III 0.60 (0.10, 0.06, 0.14, 0.12, 0.18); IV 0.68 (0.14, 0.06, 0.18, 0.10, 0.20).

Somatic characters (Fig. 1A–C). Coloration: carapace, sternum, chelicerae, endites, and labium dark brown. Abdomen light black, with numerous small, sclerotized patches, with single orange scutum laterally and posteriorly. Prosoma: carapace nearly pear shaped, surface granular, with two setae medially. Cephalic region elevated. PER strongly recurved. Chelicerae covered with setae anteriorly. Sternum almost heart shaped, rough, slightly swollen, truncated posteriorly, surface rough. Legs: light brown, covered with setae and bristles. Tibia II with two clasping spurs. Abdomen: anteriorly round, posteriorly square. Dorsally with pale yellow speckles, lateral scutum present, bears sparse, long setae. Spinnerets light brown, with circular plate.

Palp (Fig. 2A–C): bulb relatively large, ~½ size of carapace. Cymbium bears apical setae, cymbial tooth hooked. Tegulum smooth, globular. Median apophysis nearly square, with a prolateral-distal process. Embolic membrane arises between median apophysis and embolus. Embolus short, rigid, distally blunt, forming a torsion.

Figure 2. 

Crassignatha baihua sp. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Female (one of the paratypes). Total length 0.96. Carapace 0.36 long, 0.36 wide, 0.36 high. Clypeus 0.10 high. Sternum 0.24 long, 0.24 wide. Abdomen 0.64 long, 0.64 wide, 0.72 high. Length of legs: I 1.30 (0.42, 0.14, 0.30, 0.20, 0.24); II 1.06 (0.30, 0.14, 0.24, 0.16, 0.22); III 0.70 (0.18, 0.08, 0.20, 0.08, 0.16); IV 1.02 (0.30, 0.14, 0.24, 0.14, 0.20).

Somatic characters (Fig. 1D–F). Coloration: carapace, sternum, chelicerae, endites, and labium dark brown. Abdomen black, with numerous small sclerotized patches. Prosoma: carapace nearly pear shaped with small tubercles, with two strong setae medially. Cephalic region elevated. PER straight. Mouthparts and sternum as in male. Abdomen: anteriorly rounded, posteriorly square, with paired posterolateral small tubercles. Spinnerets as in male, circular plate absent.

Epigyne (Fig. 2D–G): epigynal area weakly sclerotized, with some setae. Scape distinctly extends beyond the epigastric furrow. Internal structures more or less visible via translucent tegument. Spermathecae separated by 1.2× their diameter. Fertilization ducts starting at posterior of spermathecae. Copulatory ducts long, connected to dorsal surface of spermathecae, forming four turns before curving downward. Subproximal copulatory ducts parallel, throughout the entire scape. Copulatory opening located at end of scape.

Etymology

The specific epithet is derived from the type locality; noun in apposition.

Distribution

China (Yunnan) (Fig. 38).

Crassignatha bangbie Y. Lin & S. Li, sp. nov.

Figs 3, 38

Type material

Holotype ♀ (NHMSU Ar 013), China: Yunnan Province, Longling County, Zhen’an Township, Bangbie Village, at stream at km 6.8 on Road S317, shady embankments along stream, dusting webs in understory (24.81333°N, 98.83280°E; 1560 m), 22.VIII.2018, Y. Lin et al. leg.; 1♀ (NHMSU-HA137) used for sequencing, GenBank: MT992015, same data as for preceding.

Diagnosis

Crassignatha bangbie sp. nov. is similar to C. pianma but can be distinguished by the copulatory duct having one inflection point in middle of vulva rather than two inflection points as in the latter (Fig. 3F, G).

Figure 3. 

Female of Crassignatha bangbie sp. nov. A habitus, dorsal B habitus, ventral C habitus, lateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.50 mm (A–C); 0.10 mm (D–G).

Description

Female (holotype). Total length 0.80. Carapace 0.32 long, 0.36 wide, 0.32 high. Clypeus 0.14 high. Sternum 0.20 long, 0.20 wide. Abdomen 0.52 long, 0.56 wide, 0.60 high. Length of legs: I 1.16 (0.40, 0.12, 0.28, 0.16, 0.20); II 0.92 (0.28, 0.12, 0.24, 0.12, 0.16); III 0.84 (0.24, 0.12, 0.16, 0.12, 0.20); IV 0.96 (0.32, 0.08, 0.24, 0.16, 0.16).

Somatic characters (Fig. 3A–C). Coloration: carapace, sternum, chelicerae, endites, and labium brown. Abdomen dark orange with some light patches. Prosoma: carapace nearly pear shaped. PER straight. Chelicerae covered with setae anteriorly. Labium nearly semicircular. Sternum smooth, bears sparse setae and small light patches, subcordate, truncated posteriorly. Legs: brown with a little black, covered with setae and bristles. Abdomen: anteriorly round, posteriorly relatively pointed, with light patches. Spinnerets gray, anterior spinnerets larger than posterior spinnerets.

Epigyne (Fig. 3D–G): epigynal area slightly sclerotized, with few setae. Scape stubby, protruded. Internal structures faintly visible via translucent tegument. Spermathecae separated by approximately their diameter. Fertilization ducts starting at posterior margin of spermathecae. Copulatory ducts connected to the dorsal-subcentral surface of spermathecae, bent toward the central area of vulva to form an inflection point, then retracing under the spermathecae, fused at the copulatory opening. Copulatory openings large, rounded in ventral view, located at end of scape.

Male. Unknown.

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

China (Yunnan) (Fig. 38).

Crassignatha bicorniventris (Lin & Li, 2009), comb. nov.

Figs 4, 38

Patu bicorniventris Lin & Li, 2009: 50, figs 1, 2A–D (♀).

Type material

Holotype ♀ and paratype 1♀ (IZCAS), China: Hainan Province, Changjiang Lizu Autonomous County, Qicha Town, Bawangling Nature Reserve (19.03333°N, 109.10000°E; 698 m), 29.VII.2007, S. Li and C. Wang leg. Examined.

Other material examined

1♀ (IZCAS-Ar 40996), China: Hainan Province, Dongfang City, Donghe Town, Yalong Village, outside of Yalong Huangxian Cave (18.97920°N, 108.88967°E; 264 m), 15.XII.2014, Q. Zhao and L. Shao leg.

Diagnosis

This species resembles C. nasalis sp. nov. but can be distinguished by the copulatory ducts twisting 4× but only 2× in the latter (Fig. 4D–G).

Description

Female (IZCAS-Ar 40996). Total length 0.80. Carapace 0.36 long, 0.32 wide, 0.32 high. Clypeus 0.12 high. Sternum 0.28 long, 0.20 wide. Abdomen 0.56 long, 0.60 wide, 0.56 high. Length of legs: I 1.16 (0.40, 0.14, 0.26, 0.18, 0.18); II 1.00 (0.32, 0.10, 0.24, 0.14, 0.20); III 0.68 (0.20, 0.10, 0.12, 0.12, 0.14); IV 0.98 (0.34, 0.12, 0.20, 0.14, 0.18).

Figure 4. 

Female of Crassignatha bicorniventris A habitus, dorsal B habitus, ventral C habitus, lateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.50 mm (A–C); 0.10 mm (D–G).

Somatic characters (Fig. 4A–C). Coloration: carapace light brown, centrally smooth, marginally darkish, without modified pattern. Chelicerae, endites, labium, and sternum light brown. Abdomen charcoal gray, with irregular light patches. Prosoma: carapace nearly pear shaped, with two strong setae medially. Cephalic region elevated. PER straight. Chelicerae covered with setae anteriorly. Sternum smooth, slightly plump, truncated posteriorly, fused to labium. Legs: light brown, each segment with distal black ring, covered with setae and bristles. Abdomen: anteriorly rounded, posteriorly quadrate, bears sparse, long setae, with a pair of posterolateral abdominal tubercles. Spinnerets light brown.

Epigyne (Fig. 4D–G): epigynal area lightly sclerotized, with some setae. Internal structures faintly visible via translucent tegument. Scape small, unobvious. Paired spermathecae separated by ~ ½ their diameter. Fertilization ducts originating inside middle margin of spermathecae, bent downward, reverse course under the spermathecae. Copulatory opening small, located at end of scape. Copulatory ducts long, connected to posteriorly ventral surface of spermathecae, twisted 4× under spermathecae. Bilateral copulatory ducts merged into a Y-shaped atrium before copulatory opening.

Male. Unknown.

Taxonomic justification

Although we were unable to obtain molecular data for this species, the configuration of the vulva and the modified habitus leave little doubt that it is a member of the genus Crassignatha and not Patu. Therefore, we propose a new combination, Crassignatha bicorniventris (Lin & Li, 2009) comb. nov., transferring it from Patu.

Distribution

China (Hainan) (Fig. 38).

Crassignatha changyan Y. Lin & S. Li, sp. nov.

Figs 5, 38

Type material

Holotype ♀ (NHMSU Ar 014) and paratypes 3♀ (NHMSU Ar 015–017), China: Yunnan Province, Lushui County, Pianma Township, Changyan River, 9.3 km ESE Pianma, mixed broadleaf deciduous and evergreen forest, dusting small webs near ground in forest understory (25.99363°N, 98.66651°E; 2470 m), 10.VIII.2018, Y. Lin et al. leg.

Diagnosis

Crassignatha changyan sp. nov. is similar to C. xichou sp. nov. but differs by the copulatory ducts diagonally linked to the copulatory opening but vertically linked in the latter (Fig. 5E, F).

Description

Female (holotype). Total length 0.88. Carapace 0.36 long, 0.32 wide, 0.36 high. Clypeus 0.14 high. Sternum 0.20 long, 0.20 wide. Abdomen 0.56 long, 0.56 wide, 0.60 high. Length of legs: I 1.12 (0.34, 0.20, 0.20, 0.14, 0.24); II 0.96 (0.32, 0.12, 0.16, 0.12, 0.24); III 0.78 (0.26, 0.10, 0.14, 0.08, 0.20); IV 0.98 (0.32, 0.12, 0.18, 0.12, 0.24).

Somatic characters (Fig. 5A–C). Coloration: prosoma and abdomen dark, genital area and spinnerets slightly pale. Legs pale brown, with dark pigmentation. Prosoma: carapace nearly pear shaped. PER recurved. Chelicerae bears setae anteriorly. Labium semilunar. Sternum heart shaped, smooth, slightly swollen, truncated posteriorly. Legs: a distal seta on patella dorsally, two on tibia dorsally. Abdomen: globose, modified by sparse, long setae and faint dots. Spinnerets distally pale.

Epigyne (Fig. 5D–F): epigynal area with a few setae. Scape knobbed. Paired spermathecae globose, widely separated by at least 1.5× their diameter. Copulatory ducts connected to ventral center of spermathecae, twisting into an S-shaped under spermathecae, and merging anteriorly with copulatory openings. Copulatory openings located below scape. Fertilization ducts starting at inside margin of spermathecae, then curving posteriorly.

Figure 5. 

Female of Crassignatha changyan sp. nov. A habitus, dorsal B habitus, ventral C habitus, lateral D epigyne, ventral E vulva, ventral F vulva dorsal. Scale bars: 0.50 mm (A–C); 0.10 mm (D–F).

Male. Unknown.

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

China (Yunnan) (Fig. 38).

Crassignatha dongnai Y. Lin & S. Li, sp. nov.

Figs 6, 38

Type material

Holotype ♀ (IZCAS-Ar 40997) and paratype 1♀ (IZCAS-Ar 40998), Vietnam: Dong Nai Province, Cat Tien National Park, Natural Forest (11.45008°N, 107.36438°E; 173 m), 5.IX.2015, Q. Zhao, Y. Li and Z. Chen leg.; 1♀ (NHMSU-HA092) used for sequencing, GenBank: MT992004, same data as for preceding.

Diagnosis

This species is similar to C. gucheng sp. nov. but can be distinguished by the copulatory ducts twisted once in the center of vulva vs. twisted twice in the latter (Fig. 6F, G).

Description

Female (holotype). Total length 1.28. Carapace 0.48 long, 0.40 wide, 0.40 high. Clypeus 0.14 high. Sternum 0.28 long, 0.28 wide. Abdomen 0.84 long, 0.76 wide, 0.80 high. Length of legs: I 1.60 (0.54, 0.18, 0.40, 0.24, 0.24); II 1.30 (0.40, 0.14, 0.32, 0.20, 0.24); III 1.00 (0.26, 0.14, 0.20, 0.16, 0.24); IV 1.20 (0.38, 0.14, 0.28, 0.18, 0.22).

Somatic characters (Fig. 6A–C). Coloration: carapace brown, marginally darker, mouthparts and sternum brown. Legs pale brown. Abdomen pale dorsally and ventrally, black laterally and posteriorly. Prosoma: carapace nearly pear shaped, granular with sulci, with two midline setae on cephalic area. Cephalic area elevated. PER recurved. Clypeus slightly concave. Labium tongue shaped, unfused to sternum. Sternum scutiform, flat, sculptured, truncated posteriorly. Legs: patellae with dorsal seta distally, tibia with two dorsal setae. Abdomen: anteriorly rounded, posteriorly subquadrate, with a pair of posterodorsal tubercles. Spinnerets slightly sclerotized.

Figure 6. 

Female of Crassignatha dongnai sp. nov. A habitus, dorsal B habitus, ventral C habitus, lateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.50 mm (A–C); 0.10 mm (D–G).

Epigyne (Fig. 6D–G): epigynal area distinctly sclerotized, bears a few setae. Scape large, finger-like, extended beyond the epigastric furrow. Internal structures faintly visible via the translucent tegument. Spermathecae separated by their diameter. Copulatory ducts long, connected to the dorsal center of spermathecae, curving downwards and going up to the center of vulva, then turning back vertically downward, merged before copulatory opening. Copulatory opening located at scape end. Fertilization ducts starting at inside lower margin of spermathecae, then curved posteriorly and laterally.

Male. Unknown.

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

Vietnam (Fig. 38).

Crassignatha ertou Miller, Griswold & Yin, 2009

Figs 7, 8, 38

Crassignatha ertou Miller et al., 2009: 74, figs 86D–F, 88A, B, 89A, B (♂♀).

Type material

Holotype ♂ (HNU-CASENT 9029324) and paratypes 3♀ (HNU-CASENT 9022397), China: Yunnan Province, Longling County, Mangkuan Township, Zaotanghe at Baihualing Village, undisturbed subtropical broadleaf forest, dusting webs in understory (25.30450°N, 98.80059°E; 1635 m), 2.VI.2005, C. Griswold leg. Examined.

Figure 7. 

Crassignatha ertou Miller, Griswold & Yin, 2009 A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Other material examined

1♂ 7♀ (NHMSU-HA108), China: Yunnan Province, Baoshan City, Longyang District, Mangkuan Township, Zaotanghe at Baihualing Village, good subtropical broadleaf forest, in leaf litter (25.30118°N, 98.79397°E; 1802 m), 18.VI.2016, Y. Li leg.; 1♀ (NHMSU-HA108) used for sequencing, GenBank: MT992005, same data as for preceding.

Diagnosis

The male of C. ertou is similar to that of C. rostriformis sp. nov. but can be distinguished by the narrower embolic base, wider in the latter (Fig. 8A). The female of C. ertou is most similar to C. yinzhi but differs by the copulatory ducts having four sharp turns, but only two in the latter (Fig. 8D–G).

Figure 8. 

Crassignatha ertou Miller, Griswold & Yin, 2009 A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Description

See Miller et al. (2009).

Distribution

China (Yunnan) (Fig. 38).

Crassignatha gucheng Y. Lin & S. Li, sp. nov.

Figs 9, 10, 38

Type material

Holotype ♂ (NHMSU Ar 018) and paratypes 2♂ 5♀ (NHMSU Ar 019–025), China: Yunnan Province, Longling County, Longjiang Township, Xiaoheishan Nature Reserve, Gucheng Mountain, in good forest (24.82886°N, 98.75917°E; 2010 m), 22.VIII.2018, Y. Lin et al. leg.; 1♀ (NHMSU-HA132) used for sequencing, GenBank: MT992014, same data as for preceding.

Figure 9. 

Crassignatha gucheng sp. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Diagnosis

The male of C. gucheng sp. nov. is similar to that of C. pianma and C. quanqu but can be distinguished by the narrower embolic base and spine-like cymbial tooth vs. a wider embolic base and spur-like cymbial tooth in the latter two (Fig. 10A). The female of C. gucheng sp. nov. differs from that of C. dongnai sp. nov. by the copulatory ducts twisted twice at the center of vulva, only once in the latter (Fig. 10F, G).

Description

Male (holotype). Total length 0.72. Carapace 0.32 long, 0.36 wide, 0.40 high. Clypeus 0.12 high. Sternum 0.24 long, 0.24 wide. Abdomen 0.44 long, 0.44 wide, 0.44 high. Length of legs: I 1.28 (0.38, 0.14, 0.32, 0.20, 0.24); II 0.98 (0.26, 0.14, 0.26, 0.14, 0.18); III 0.78 (0.20, 0.10, 0.16, 0.12, 0.20); IV 0.94 (0.30, 0.12, 0.18, 0.14, 0.20).

Somatic characters (Fig. 9A–C). Coloration: carapace pale brown, marginally darker. Mouthparts and sternum pale brown. Legs pale yellow, with faint pigmentation on each segment distally. Abdomen pale dorsally, brown-yellow laterally and ventrally, dark posteriorly. Prosoma: carapace nearly rounded, cephalic area elevated, cervical groove distinct, thoracic area and clypeus sculptured, clypeus concave. ALE protruded, PER recurved. Chelicerae bears setae anteriorly. Labium semicircular. Sternum heart shaped, slightly swollen, surface rough, truncated posteriorly. Legs: with dorsal seta on each patella distally, with two setae on each tibia subproximally. Tibia II with two clasping spurs. Abdomen: anteriorly rounded, posteriorly subquadrate, lateral scutum inconspicuous. Spinnerets slightly sclerotized, lack circular plate.

Palp (Fig. 10A–C): relatively large, no less than 1/3 carapace size. A few setae on cymbium distally. Apical cymbial tooth caniniform. Tegulum smooth, broad, globular. Plate-like median apophysis with a distal sclerotized process. Embolic membrane slender, laminar, translucent, arises from base of embolus. Embolus short and rigid, terminus blunt.

Figure 10. 

Crassignatha gucheng sp. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Female (one of the paratypes). Total length 1.08. Carapace 0.44 long, 0.40 wide, 0.40 high. Clypeus 0.12 high. Sternum 0.28 long, 0.24 wide. Abdomen 0.72 long, 0.72 wide, 0.84 high. Length of legs: I 1.48 (0.56, 0.16, 0.40, 0.14, 0.22); II 1.20 (0.38, 0.16, 0.34, 0.12, 0.20); III 0.84 (0.28, 0.14, 0.18, 0.10, 0.14); IV 1.1 (0.38, 0.14, 0.28, 0.18, 0.12).

Somatic characters (Fig. 9D–F). Coloration: carapace roughly the same as in male. Legs pale brown. Abdomen with black pigmentation laterally, posteriorly, and ventrally. Prosoma: carapace pear shaped, sculptured, cephalic area slightly lower than in male. PER recurved. Mouthparts and sternum as in male. Abdomen: anteriorly rounded and posteriorly subquadrate, surface with weakly sclerotized hairy patches. Spinnerets as in male.

Epigyne (Fig. 10D–G): epigynal area distinctly sclerotized, bears a few setae. Scape large, finger-like, protruded, copulatory opening located at its terminus. Internal structures faintly visible via translucent tegument. Spermathecae globose, strongly sclerotized, widely separated by 1.5× their diameter. Fertilization duct originates at medial posterior margin of spermathecae, extends to posterior margin of epigyne. Copulatory ducts long, connected to dorsal surface of spermathecae, extends below spermathecae toward the vulval center, and forms two return paths, then turns downward to copulatory opening.

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

China (Yunnan), Vietnam (Fig. 38).

Crassignatha gudu Miller, Griswold & Yin, 2009

Figs 11, 38

Crassignatha gudu Miller et al., 2009: 75, fig. 89C, D (♀).

Type material

Holotype ♀ (HNU-CASENT 9029318), China: Yunnan Province, Longling County, Mangkuan Township, Zaotanghe at Baihualing Village, good subtropical broadleaf forest, dusting webs in understory (25.30450°N, 98.80059°E; 1635 m), 2.VI.2005, C. Griswold leg. Examined.

Diagnosis

Crassignatha gudu can be easily distinguished from other congeners, except C. yamu and C. si sp. nov., by having a columnar atrium formed by the fusion of proximal copulatory ducts (Figs 11F, 31F, 35G). Crassignatha gudu is most similar to C. yamu and C. si sp. nov. and has a similar vulva configuration but differs by the longer copulatory atrium (Fig. 11E, F vs. Figs 31E, F, 35F, G).

Description

See Miller et al. (2009).

Distribution

China (Yunnan) (Fig. 38).

Figure 11. 

Female of Crassignatha gudu A prosoma, dorsal B prosoma, ventral C abdomen, dorsal D abdomen, ventral E vulva, ventral F vulva, dorsal. Scale bars: 0.50 mm (A–D); 0.10 mm (E, F).

Crassignatha mengla Y. Lin & S. Li, sp. nov.

Figs 12, 13, 38

Type material

Holotype ♂ (IZCAS-Ar 40999) and paratypes 4♂ 8♀ (IZCAS-Ar 41000–41011), China: Yunnan Province, Xishuangbanna Prefecture, Mengla County, Menglun Town, Entrance to Shenmi Cave, in good forest (21.97332°N, 101.24336°E; 776 m), 3.X.2017, Y. Lin and Y. Li leg.; 1♂ juvenile (NHMSU-HA080) and 1♀ (NHMSU-HA080) used for sequencing, GenBank: MT992000 and MT991999, same data as for preceding.

Diagnosis

The male of Crassignatha mengla sp. nov. is similar to that of C. baihua sp. nov. and C. quadriventris (Lin & Li, 2009) comb. nov. but differs by the wider and longer embolic tip and the median apophysis lacks a process, rather than a narrower (Fig. 13A, B), shorter embolic tip and a process on the median apophysis in the latter two (Figs 2A, B, 21A, B). The female of C. mengla sp. nov. is most similar to C. rostriformis sp. nov. in the vulva configuration but can be easily distinguished by the spermathecae separated by more than one diameter vs. less than one diameter (Figs 13F, G, 25E, F).

Figure 12. 

Crassignatha mengla sp. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Description

Male (holotype). Total length 0.72. Carapace 0.36 long, 0.32 wide, 0.36 high. Clypeus 0.16 high. Sternum 0.24 long, 0.20 wide. Abdomen 0.44 long, 0.40 wide, 0.44 high. Length of legs: I 1.24 (0.38, 0.14, 0.30, 0.18, 0.24); II 1.04 (0.28, 0.14, 0.24, 0.14, 0.24); III 0.74 (0.18, 0.10, 0.14, 0.12, 0.20); IV 0.88 (0.24, 0.12, 0.20, 0.12, 0.20).

Somatic characters (Fig. 12A–C). Coloration: prosoma dark brown. Legs brown-yellow. Abdomen charcoal gray, darker ventrally than dorsally. Prosoma: carapace sub-rounded, surface rough, sculptured. Cephalic area elevated. ALE protruded, PER strongly recurved. Clypeus concave. Mouthparts strongly sclerotized. Labium semicircular. Sternum heart shaped, slightly plump, surface rough, truncated posteriorly. Legs: tibia II with two clasping spurs subdistal-ventrally. Abdomen: nearly rounded dorsally, with lightly sclerotized dots dorsally. Lateral scutum weakly sclerotized, dark brown. Spinnerets brown, with a circular plate.

Palp (Fig. 13A–C): weakly sclerotized. Cymbium with few setae distally; cymbial tooth thin, located subapically. Tegulum globular and plump. Median apophysis lamellar, with sclerotized margin. Embolic membrane arises behind median apophysis, near embolic base. Embolus short, rigid, basolateral protrusion tapering distally, with a single bend.

Female (paratypes). Total length 1.00. Carapace 0.40 long, 0.36 wide, 0.36 high. Clypeus 0.14 high. Sternum 0.24 long, 0.24 wide. Abdomen 0.64 long, 0.64 wide, 0.60 high. Length of legs: I 1.38 (0.42, 0.14, 0.36, 0.20, 0.26); II 1.14 (0.30, 0.12, 0.30, 0.18, 0.24); III 0.88 (0.26, 0.10, 0.18, 0.14, 0.20); IV 1.10 (0.38, 0.12, 0.24, 0.14, 0.22).

Somatic characters (Fig. 12D–F). Coloration: prosoma as in male. Abdomen lighter than in male. Prosoma: carapace pear shaped. Cephalic region elevated, slightly lower than in male. PER slightly recurved. Labium triangular, unfused to sternum. Sternum heart shaped, surface rough, slightly plump, truncated posteriorly. Abdomen: anteriorly round, posteriorly nearly square. Spinnerets brown, weakly sclerotized.

Epigyne (Fig. 13D–G): epigynal area with few setae. Scape short, slightly protruded. Copulatory opening located at terminal part of scape. Internal structures faintly visible via translucent tegument. Spermathecae globose, widely separated by ~1.2× their diameter. Fertilization ducts short, thin, starting at the inside margin of spermathecae, deflexed, bifurcated distally. Copulatory ducts long, thick, connected to posterior margin of spermathecae, bent upward to center of vulva, then downward, fusing before reaching copulatory opening.

Figure 13. 

Crassignatha mengla sp. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

China (Yunnan) (Fig. 38).

Crassignatha nantou Y. Lin & S. Li, sp. nov.

Figs 14, 15, 38

Type material

Holotype ♂ (IZCAS-Ar 41012) and paratypes 17♀ (IZCAS-Ar 41013–41029), China: Taiwan Province, Nantou County, Ren’an Township, Hehuan Mountain, Yuanfeng Peak (24.11780°N, 121.23731°E; 2757 m), 2.VII.2013, S. Li and G. Zheng leg.; 1♂ (NHMSU-HA055) and 1♀ (NHMSU-HA055) used for sequencing, GenBank: MT991996 and MT991995, same data as for preceding.

Diagnosis

Crassignatha nantou sp. nov. differs from other congeners, except C. shunani sp. nov., by the long, spiral embolus with a sharp, narrow tip and the separate bases of copulatory ducts (Fig. 15A, B, F, G). It is similar to C. shunani sp. nov. by the shape of the male palp and vulva, but the male can be distinguished by having a blunt cymbial tooth and a hook on the median apophysis vs. a sharp cymbial tooth and lack of a hook in the latter (Figs 15A, 29A); the female differs by the closer spermathecae and larger copulatory opening (Figs 15F, G, 29F, G).

Figure 14. 

Crassignatha nantou sp. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Description

Male (holotype). Total length 0.80. Carapace 0.36 long, 0.36 wide, 0.40 high. Clypeus 0.20 high. Sternum 0.24 long, 0.24 wide. Abdomen 0.52 long, 0.40 wide, 0.60 high. Length of legs: I 1.34 (0.42, 0.14, 0.32, 0.20, 0.26); II 1.10 (0.32, 0.12, 0.26, 0.16, 0.24); III 0.80 (0.22, 0.10, 0.16, 0.12, 0.20); IV 1.02 (0.32, 0.12, 0.22, 0.12, 0.24).

Somatic characters (Fig. 14A–C). Coloration: carapace, sternum, chelicerae, endites, and labium dark brown. Abdomen blue-green with irregular sclerotized patches. Prosoma: carapace nearly rounded, surface granular, with small sulci. Cephalic region strongly elevated. ALE protruded, PER recurved. Clypeus concave. Chelicerae covered with setae anteriorly. Labium semilunar, fused to sternum. Sternum heart shaped, flat, surface rough, truncated posteriorly. Legs: tibia II with one clasping spur. Abdomen: sub-elliptic dorsally, with lateral scuta. Spinnerets weakly sclerotized, with a circular plate.

Palp (Fig. 15A–C): pale, weakly sclerotized. Cymbium large, with a few setae distally, cymbial tooth near distal margin. Tegulum large, smooth, and plump. Median apophysis disciform, with a hooked process on margin. Embolic membrane slender, laminar, translucent, arises near anterior part of median apophysis. Embolus long, flexible, spiraled, and wide basally, narrow distally.

Female (one of paratypes). Total length 1.08. Carapace 0.44 long, 0.40 wide, 0.40 high. Clypeus 0.16 high. Sternum 0.28 long, 0.28 wide. Abdomen 0.68 long, 0.60 wide, 0.80 high. Length of legs: I 1.68 (0.58, 0.16, 0.42, 0.24, 0.28); II 1.38 (0.46, 0.14, 0.34, 0.22, 0.22); III 1.00 (0.28, 0.12, 0.22, 0.16, 0.22); IV 1.24 (0.42, 0.14, 0.26, 0.18, 0.24).

Somatic characters (Fig. 14D–F). Coloration: prosoma as in male. Abdominal color lighter than in male, dorsum lighter than venter. Prosoma: carapace nearly pear shaped, sculptured, and granular. Cephalic region lower than in male. PER slightly recurved. Chelicerae fused near base, covered with setae anteriorly. Labium subtriangular, fused to sternum. Sternum as in male. Abdomen: nearly globular dorsally, surface modified by sclerotized dots. Spinnerets weakly sclerotized, with a circular plate.

Epigyne (Fig. 15D–G): epigynal area lightly sclerotized, with setae on lateral margins. Scape short, wide, copulatory opening located at its terminus, split into two labella. Internal structures more or less visible via translucent tegument. Paired spermathecae globose, separated by half their diameter. Fertilization ducts thin, starting at inside central margin of spermathecae, bent downward, twisted, and furcate at end. Copulatory ducts thick, connected to posterior margin of spermathecae, passing under the spermathecae, up into the center of vulva, deflexed to copulatory opening, their proximal base unfused.

Figure 15. 

Crassignatha nantou sp. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

China (Taiwan) (Fig. 38).

Crassignatha nasalis Y. Lin & S. Li, sp. nov.

Figs 16, 17, 38

Type material

Holotype ♂ (NHMSU Ar 026) and paratypes 9♀ (NHMSU Ar 027–035), China: Sichuan Province, Luzhou City, Gulin County, Yuhua Township, Taoyuan Cave (27.98293°N, 105.99833°E; 910 m), 21.IV.2014, Y. Lin, H. Zhao, and Y. Li leg. 1♂ (NHMSU-HA041) and 1♀ (NHMSU-HA041) used for sequencing, GenBank: MT991992 and MT991991, same data as for preceding.

Diagnosis

The male of C. nasalis sp. nov. is similar to that of C. rostriformis sp. nov. but can be distinguished by the sharp hook of the median apophysis and a straight cymbial tooth vs. a blunt hook of the median apophysis and a hook-like cymbial tooth (Figs 17A, B, 25A, B). The female is similar to C. quanqu in epigyne shape but differs from the latter by the indistinct scape and the copulatory ducts nearly forming a closed rhombic area at center of vulva vs. obvious scape and copulatory ducts not forming a closed area at center of vulva (Fig. 17E, F vs. Fig. 23D, E).

Figure 16. 

Crassignatha nasalis sp. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Description

Male (holotype). Total length 0.80. Carapace 0.40 long, 0.32 wide, 0.44 high. Clypeus 0.12 high. Sternum 0.24 long, 0.20 wide. Abdomen 0.52 long, 0.44 wide, 0.56 high. Length of legs: I 1.10 (0.30, 0.14, 0.30, 0.16, 0.20); II 0.92 (0.24, 0.12, 0.24, 0.14, 0.18); III 0.72 (0.20, 0.10, 0.14, 0.10, 0.18); IV 0.82 (0.22, 0.10, 0.20, 0.12, 0.18).

Somatic characters (Fig. 16A–C). Coloration: carapace, sternum, chelicerae, endites, and labium brown. Legs yellow-brown. Abdomen pale gray, ventrally darker than dorsally, with sclerotized dots and sparse setae; posterolateral scutum light brown. Prosoma: carapace nearly rounded, thoracic center smooth, cephalic area and margins granulated and pitted, with two strong setae medially. Cephalic area strongly elevated. ALE protruded, PER strongly recurved. Clypeus concave. Chelicerae covered with setae anteriorly. Sternum almost heart shaped, slightly plump, truncated posteriorly, surface rough. Legs: tibia II with two large clasping spurs. Abdomen: sub-rounded dorsally. Spinnerets weakly sclerotized, with a circular plate.

Palp (Fig. 17A–C): pale, weakly sclerotized. Cymbium with a few setae apically, cymbial tooth spur-like, located at distal terminus. Tegulum globose, smooth. Plate-like median apophysis with a spike-shaped distal process. Embolic membrane arises behind the median apophysis. Embolus short, rigid, basally constricted and mesally widened, distally forming an inverted Z-shape.

Figure 17. 

Crassignatha nasalis sp. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E vulva, ventral F vulva, dorsal. Scale bars: 0.10 mm (A–F).

Female (one of paratypes). Total length 1.00. Carapace 0.40 long, 0.36 wide, 0.40 high. Clypeus 0.16 high. Sternum 0.24 long, 0.24 wide. Abdomen 0.68 long, 0.68 wide, 0.76 high. Length of legs: I 1.40 (0.46, 0.16, 0.32, 0.20, 0.26); II 1.18 (0.36, 0.14, 0.26, 0.18, 0.24); III 0.82 (0.20, 0.12, 0.16, 0.14, 0.20); IV 1.04 (0.30, 0.14, 0.24, 0.16, 0.20).

Somatic characters (Fig. 16D–F). Coloration: prosoma and legs as in male. Abdomen dark, with tiny light yellow dots. Prosoma: carapace nearly pear shaped, ocular arrangement and modification as in male. Cephalic area lower than in male. PER straight. Mouthparts and sternum as in male. Abdomen: globular dorsally. Spinnerets weakly sclerotized.

Epigyne (Fig. 17D–F): epigynal area distinctly sclerotized, with a few setae. Scape short, strongly sclerotized. Copulatory openings separated, resembling a pig snout. Vulva visible via translucent tegument. Paired spermathecae separated by less than ½ their diameter. Fertilization ducts originate inside posterior edge of spermathecae, curving toward center of spermathecae. Copulatory ducts connected to dorsal center surface of spermathecae, twisting 3 ×, then forming a nearly closed rhombic area in center of vulva.

Etymology

The specific epithet is a Latin adjective (= nasal) and refers to the shape of the copulatory openings of the epigyne.

Distribution

China (Sichuan) (Fig. 38).

Crassignatha pianma Miller, Griswold & Yin, 2009

Figs 18, 19, 38

Crassignatha pianma Miller et al., 2009: 70, figs 67I, 74A–E, 75A, B, 76A, B, 77A–F, 78A–D, 79A, B (♂♀).

Type material

Holotype ♂ (HNU-CASENT 9022600) and paratypes 2♂ 6♀ (HNU-CASENT 9022360), China: Yunnan Province, Lushui County, Pianma Township, Changyan River, 9.3 km ESE Pianma, mixed broadleaf deciduous and evergreen forest, dusting small webs near ground in forest understory (25.99363°N, 98.66651°E; 2470 m), 13–15.V.2005, C. Griswold leg. Examined.

Figure 18. 

Crassignatha pianma Miller, Griswold & Yin, 2009 A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Other material examined

21♂ 93♀ (NHMSU-HA113), China: Yunnan Province, Lushui County, Pianma Town, Changyan River, mixed broadleaf deciduous and evergreen forest, dusting small webs near ground in forest (25.99363°N, 98.66651°E; 2470 m), 10.VIII.2018, Y. Lin et al. leg.; 1♂ (NHMSU-HA113) and 1♀ (NHMSU-HA113) used for sequencing, GenBank: MT992009 and MT992008, same data as for preceding.

Diagnosis

The male of C. pianma is similar to C. quanqu in the form of the palp but can be distinguished from the latter by the subquadrate median apophysis with a hooked process, rather than a subtriangular median apophysis with a truncated process (Figs 19A, B, 23A, B). The female of C. pianma seems closest to that of C. bangbie sp. nov. but differs by the more slender, more twisted copulatory ducts than in the latter, and the smaller copulatory openings (Figs 19E–G, 3E–G).

Figure 19. 

Crassignatha pianma Miller, Griswold & Yin, 2009 A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Description

See Miller et al. (2009).

Distribution

China (Yunnan) (Fig. 38).

Crassignatha quadriventris (Lin & Li, 2009), comb. nov.

Figs 20, 21, 38

Patu quadriventris Lin & Li, 2009: 55, figs 7A, B, 8A, B, 9A–E, 10A, B (♂♀).

Type material

Holotype ♂ and paratypes 2♂ 2♀ (IZCAS), China: Hainan Province, Wuzhishan City, Mt. Wuzhishan Nature Reserve (18.90000°N, 109.65000°E), 9.VIII.2007, S. Li and C. Wang leg.; paratype 1♀ (IZCAS), China: Hainan Province, Qiongzhong County, Limushan Nature Reserve (19.18333°N, 109.73333°E; 655 m), 12.VIII.2007, S. Li and C. Wang leg. Examined.

Figure 20. 

Crassignatha quadriventris (Lin & Li, 2009) comb. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Other material examined

1♂ 4♀ (IZCAS-Ar 41030–41034), China: Hainan Province, Dongfang City, Donghe Town, Nanlang Village, at the foot of Exianling Mountain (19.00633°N, 109.08383°E; 214 m), 16.XII.2014, Q. Zhao and L. Shao leg.; 1♀ (NHMSU-HA025) used for sequencing, GenBank: MT991990, same data as for preceding.

Diagnosis

This species differs from other congeneric species except C. baihua sp. nov. by the short, stiff embolus (Fig. 21A, B). It is most similar to C. baihua sp. nov. in the form of the palp and the vulva configuration but can be easily distinguished by the sloped embolic apex, rather than flat as in the latter, and by having six twists of the copulatory ducts, rather than four in the latter (Figs 21A, B, F, G, 2A, B, F, G).

Description

Male (IZCAS-Ar 41030). Total length 0.80. Carapace 0.36 long, 0.36 wide, 0.40 high. Clypeus 0.16 high. Sternum 0.28 long, 0.24 wide. Abdomen 0.52 long, 0.48 wide, 0.52 high. Length of legs: I 1.34 (0.42, 0.14, 0.34, 0.20, 0.24); II 1.10 (0.30, 0.14, 0.26, 0.16, 0.24); III 0.90 (0.24, 0.12, 0.18, 0.14, 0.22); IV 1.02 (0.30, 0.12, 0.22, 0.18, 0.20).

Somatic characters (Fig. 20A–C). Coloration: prosoma brown. Legs pale brown. Abdomen dorsally dark, with pale stripes and speckles, ventrally pale brown. Prosoma: carapace sub-rounded, granular, sculptured, with two cephalic setae. Cephalic area elevated. Clypeus concave. ALE protruded. PER strongly recurved. Chelicerae covered with few setae anteriorly. Labium tongue shaped, wider than long. Sternum heart shaped, surface rough, slightly plump, truncated posteriorly. Legs: tibia II with two clasping spurs. Abdomen: anteriorly round, nearly square posteriorly, with sparse, long setae, lateral scutum distinct. Spinnerets slightly sclerotized, surrounded by a circular plate.

Palp (Fig. 21A–C): tibia with four distal, dorsal, short setae. Cymbium bears a few distal setae. Apical cymbial tooth hook shaped. Tegulum broad, smooth, globular. Plate-like median apophysis with a sclerotized, longitudinal central ridge line and a prolaterally odontoid process. Embolic membrane arises near anterior part of median apophysis. Embolus short, stiff, basally constricted, mesally wide, distally coracoid.

Female (IZCAS-Ar 41031). Total length 1.04. Carapace 0.44 long, 0.40 wide, 0.40 high. Clypeus 0.14 high. Sternum 0.28 long, 0.28 wide. Abdomen 0.72 long, 0.68 wide, 0.80 high. Length of legs: I 1.72 (0.60, 0.18, 0.40, 0.26, 0.28); II 1.42 (0.44, 0.16, 0.34, 0.20, 0.28); III 0.98 (0.26, 0.10, 0.22, 0.16, 0.24); IV 1.18 (0.34, 0.12, 0.28, 0.18, 0.26).

Somatic characters (Fig. 20D–F). Coloration: prosoma and legs as in male. Abdomen dorsally darker than in male, ventrally lighter. Prosoma: modification and arrangement of eyes as in male, cephalic area lower than in male. Clypeus slightly concave. Mouthparts and sternum as in male. Abdomen: anteriorly round and posteriorly square, surface bears sparse, long setae, lateral scutum and circular plate absent. Spinnerets slightly sclerotized.

Epigyne (Fig. 21D–F): epigynal area weakly sclerotized, bears a few setae. Scape developed, protruded, longer than wide. Copulatory openings located at terminus. Internal structures faintly visible via translucent tegument. Paired spermathecae globose, widely separated by at least 1.5× their diameter. Fertilization ducts originating from the lower inside margin of spermathecae, bent downwards and laterally. Copulatory ducts long, connected to dorsal surface of spermathecae, from below spermathecae to the center of vulva, making six bends, then reaching copulatory openings.

Taxonomic justification

The shape of the male palps, the configuration of the epigyne, the modified carapace, and the male abdominal scutum and clasping spurs on tibia II leave no doubt that this species is a member of Crassignatha and not Patu. Therefore, we propose a new combination, C. quadriventris (Lin & Li, 2009) comb. nov., transferring it from Patu.

Distribution

China (Hainan) (Fig. 38).

Figure 21. 

Crassignatha quadriventris (Lin & Li, 2009) comb. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Crassignatha quanqu Miller, Griswold & Yin, 2009

Figs 22, 23, 38

Crassignatha quanqu Miller et al., 2009: 72, figs 76E–G, 78F, 79E, F, 83A–E, 84A, B, 85A–F (♂♀).

Type material

Holotype ♂ (HNU-CASENT 9029323) and paratypes 1♂ 1♀ (HNU-CASENT 9022388), China: Yunnan Province, Longling County, Zhen’an Township, Bangbie Village, at stream at km 6.8 on Route S317, shady embankments along stream, dusting webs in understory (24.81333°N, 98.83280°E, 1552.5 ± 7.5 m), 24.V.2005, C. Griswold leg. Examined.

Figure 22. 

Crassignatha quanqu A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female prosoma, dorsal E female prosoma, ventral F female abdomen, dorsal G female abdomen, ventral. Scale bars: 0.50 mm (A–G).

Diagnosis

The male of Crassignatha quanqu is similar to C. gucheng sp. nov. but can be distinguished by the median apophysis with two tapered distal processes and the details of the palp (Fig. 23A–C vs. Fig. 10A–C). The female of C. quanqu is similar to C. yinzhi and C. thamphra sp. nov. in the vulva configuration, but differs by the nearly vertical connection of copulatory ducts and copulatory openings, rather than connected diagonally as in the latter two, and differs by the fertilization ducts starting at the posterior margin of the spermathecae, rather than the inside medial margin as in the latter two (Fig. 23D, E vs. Figs 32F, G, 37F, G).

Description

See Miller et al. (2009).

Distribution

China (Yunnan) (Fig. 38).

Figure 23. 

Crassignatha quanqu A male palp, prolateral B male palp, ventral C male palp, retrolateral D vulva, ventral E vulva, dorsal. Scale bars: 0.10 mm (A–E).

Crassignatha rostriformis Y. Lin & S. Li, sp. nov.

Figs 24, 25, 38

Type material

Holotype ♂ (NHMSU Ar 036) and paratypes 2♂ 34♀ (NHMSU Ar 037–072), China: Yunnan Province, Wenshan Prefecture, Xichou County, Dongma Township, Xianrendong Village, Xianren Cave (23.50193°N, 104.86810°E; 1326 m), 6.VIII.2010, Z. Yao, X. Wang and C. Wu leg.; 2♂ 16♀ (NHMSU-HA078), 3♂ 2♀ (NHMSU-HA103), same data as holotype; 1♂ (NHMSU-HA079) and 1♀ (NHMSU-HA079) used for sequencing, GenBank: MT991998 and MT991997, same data as for preceding.

Figure 24. 

Crassignatha rostriformis sp. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Diagnosis

The male of C. rostriformis sp. nov. is similar to C. nasalis sp. nov. in the form of the palp but differs from the latter by the large, hooked cymbial tooth and the wider embolic base (Fig. 25B vs. Fig. 17A, B). The female is most similar to that of C. mengla sp. nov. in the vulva configuration but can be easily distinguished by spermathecae separated by less than their diameter, and the low second turn of the copulatory duct, rather than wide intervals of spermathecae and the high second turn of the copulatory duct (Fig. 25D–F vs. Fig. 13F, G).

Description

Male (holotype). Total length 0.64. Carapace 0.28 long, 0.28 wide, 0.32 high. Clypeus 0.10 high. Sternum 0.20 long, 0.20 wide. Abdomen 0.40 long, 0.36 wide, 0.48 high. Length of legs: I 1.04 (0.32, 0.12, 0.24, 0.16, 0.20); II 0.84 (0.22, 0.12, 0.20, 0.12, 0.18); III 0.66 (0.20, 0.10, 0.12, 0.10, 0.14); IV 0.80 (0.26, 0.10, 0.16, 0.12, 0.16).

Somatic characters (Fig. 24A–C). Coloration: prosoma dark brown, ventrally darker than dorsally. Legs brown, with black pigmentation. Abdomen dark, laterally and ventrally darker than dorsally, with light brown speckles. Prosoma: carapace sub-rounded, surface sculptured. Cephalic area elevated. Clypeus concave. ALE slightly protruded. PER recurved. Mouthparts distinctly sclerotized. Labium nearly semilunar. Sternum scutiform, surface subtly textured, slightly plump, truncated posteriorly. Legs: tibia II with two clasping spurs. Abdomen: rounded dorsally, abdominal lateral scutum weakly sclerotized, circular plate absent. Spinnerets tiny.

Palp (Fig. 25A–C): tibia as long as patella. Cymbium wider than femur, bears some distal setae, with a dorsal hooked tooth near distal margin. Tegulum plump and globular. Disciform median apophysis with an odontoid prolateral process. Embolic membrane arises from behind median apophysis. Embolus short, rigid, basally wide, mesally and distally narrow.

Female (one of paratypes). Total length 0.92. Carapace 0.40 long, 0.36 wide, 0.36 high. Clypeus 0.12 high. Sternum 0.24 long, 0.24 wide. Abdomen 0.60 long, 0.72 wide, 0.68 high. Length of legs: I 1.36 (0.46, 0.12, 0.32, 0.22, 0.24); II 1.12 (0.38, 0.14, 0.26, 0.16, 0.18); III 0.82 (0.22, 0.12, 0.14, 0.16, 0.18); IV 1.02 (0.30, 0.12, 0.24, 0.16, 0.20).

Somatic characters (Fig. 24D–F). Coloration: prosoma as in male. Abdomen darker than in male. Prosoma: carapace nearly pear shaped, weakly granular. Cephalic area lower than in male. PER slightly recurved. Mouthparts and sternum as in male. Abdomen: nearly globose, dorsally speckled. Spinnerets weakly sclerotized.

Epigyne (Fig. 25D–F): slightly sclerotized, with a few setae at lateral margins. Scape short, slightly protruded. Copulatory openings large, flat, beak shaped, located at the terminus of scape. Internal structures faintly visible via translucent tegument. Spermathecae globose, separated by ~0.8× their diameter. Fertilization ducts starting at inside posterior margin of spermathecae and bending below the venter of spermathecae. Copulatory ducts relatively long, connected to lower dorsal surface of spermathecae, bypassing around spermathecae, forming three bends, fusing before copulatory openings.

Figure 25. 

Crassignatha rostriformis sp. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E vulva, ventral F vulva, dorsal. Scale bars: 0.10 mm (A–F).

Etymology

The specific epithet is derived from the Latin adjective rostriformis (rostriform), in reference to the shape of the copulatory openings.

Distribution

China (Yunnan) (Fig. 38).

Crassignatha shiluensis (Lin & Li, 2009), comb. nov.

Figs 26, 27, 38

Patu shiluensis Lin & Li, 2009: 59, figs 11A, B, 12A, B, 13A–D (♂♀).

Type material

Holotype ♂ and paratypes 4♂ 9♀ (IZCAS), China: Hainan Province, Changjiang Lizu Autonomous County, Shilu Town, in leaf litter in rainforest (19.20000°N, 109.06667°E), 22.III.2005, Y. Tong, Y. Song and X. Han leg. Examined.

Figure 26. 

Crassignatha shiluensis (Lin & Li, 2009) comb. nov. A male habitus, dorsal, B male habitus, ventral C female habitus, dorsal D female habitus, ventral. Scale bars: 0.50 mm (A–D).

Other material examined

3♂ 8♀ (NHMSU-HA081), China: Yunnan Province, Mengla County, Menglun Town, Xishuangbanna Tropical Botanic Garden, tropical rainforest (21.917°N, 101.275°E; 558 m), 5.X.2017, Y. Lin and Y. Li leg.; 1♂ juvenile (NHMSU-HA081) and 1♀ (NHMSU-HA081) used for sequencing, GenBank: MT992002 and MT992001, same data as for preceding; 1♂ 3♀ (NHMSU-HA059), China: Yunnan Province, Menglun Nature Reserve, secondary tropical seasonal moist forest (21.91197°N, 101.28233°E; 645 ± 17 m), 5–12.I.2007, G. Zheng leg.

Diagnosis

This species differs from all other species of Crassignatha by the long embolus coiling into two loops (Fig. 27A) and by the long copulatory ducts connected to the anterolateral margin of the spermathecae, coiled into three loops below spermathecae (Fig. 27D, E). Its dorsoventral dichroism is also a prominent feature (Fig. 26A–D).

Figure 27. 

Crassignatha shiluensis (Lin & Li, 2009) comb. nov. A male palp, prolateral B male palp, retrolateral C epigyne, ventral D vulva, ventral E vulva, dorsal. Scale bars: 0.10 mm (A–E).

Description

See Lin and Li (2009).

Taxonomic justification

A series of combinations: the form of the male palp and the configuration of the epigyne, the chelicerae fused at the base, and the male clasping setae distoventrally on tibia II suggest that this species is more similar to Crassignatha than Patu. It shares homologous characters of Crassignatha, such as a large median apophysis on a slightly oblate male palpal bulb and globular spermathecae rather than a nearly oviform male palpal bulb and claviform spermathecae as in Patu. Thus, we propose a new combination, Crassignatha shiluensis (Lin & Li, 2009) comb. nov., transferring it from Patu.

Distribution

China (Hainan, Yunnan) (Fig. 38).

Crassignatha shunani Y. Lin & S. Li, sp. nov.

Figs 28, 29, 38

Type material

Holotype ♂ (NHMSU Ar 073) and paratypes 4♂ 49♀ (NHMSU Ar 074–126), China: Sichuan Province, Luzhou City, Gulin County, Jianzhu Township, Wenyi Village, Dahei Cave (28.06134°N, 105.58015°E; 852 m), 23.IV.2014, Y. Lin, H. Zhao and J. Wu leg.; 1♂ juvenile (NHMSU-HA046) and 1♀ (NHMSU-HA046) used for sequencing, GenBank: MT991994 and MT991993, same data as for preceding.

Diagnosis

This species is similar to C. nantou sp. nov. in the form of the male palp and the vulva configuration but can be distinguished by having a sharp cymbial tooth and lacking a hooked process on the median apophysis (Fig. 29A vs. Fig. 15A), and by the more widely separated spermathecae and lower inflection point of the copulatory ducts in the center of the vulva (Figs 29F, 15G).

Figure 28. 

Crassignatha shunani sp. nov. A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Description

Male (holotype). Total length 0.68. Carapace 0.32 long, 0.28 wide, 0.32 high. Clypeus 0.12 high. Sternum 0.20 long, 0.20 wide. Abdomen 0.40 long, 0.40 wide, 0.48 high. Length of legs: I 1.02 (0.32, 0.12, 0.24, 0.16, 0.18); II 0.88 (0.24, 0.12, 0.20, 0.16, 0.16); III 0.66 (0.18, 0.10, 0.12, 0.10, 0.16); IV 0.76 (0.22, 0.10, 0.16, 0.12, 0.16).

Somatic characters (Fig. 28A–C). Coloration: prosoma brown. Legs pale brown. Abdomen grayish yellow dorsally, dark brownish laterally and ventrally, with weakly sclerotized dots dorsally. Prosoma: carapace sub-rounded, cephalic and thoracic area granular, thoracic center and clypeus smooth. Cephalic area strongly elevated. Clypeus concave. ALE protruded. PER distinctly recurved. Chelicerae bears short setae anteriorly. Labium subtriangular. Sternum heart shaped, surface textured, slightly swollen, truncated posteriorly. Legs: tibia II with two clasping spurs. Abdomen: nearly globular, posterolateral scutum weakly sclerotized. Spinnerets without circular plate.

Palp (Fig. 29A–C): tibia laminar, subequal to patella in length. Cymbium with some setae at distal margin, horn-shaped cymbial tooth near cymbial apex. Tegulum globose, swollen, surface rugose. Nearly rounded median apophysis prolaterally on bulb, with short, straight distal process. Embolic membrane arises beside median apophysis. Embolus long, spiral, basally and mesally wide, distally narrow and bent.

Female (one of paratypes). Total length 0.80. Carapace 0.36 long, 0.32 wide, 0.32 high. Clypeus 0.12 high. Sternum 0.24 long, 0.20 wide. Abdomen 0.56 long, 0.60 wide, 0.64 high. Length of legs: I 1.12 (0.38, 0.12, 0.26, 0.18, 0.18); II 0.94 (0.26, 0.12, 0.22, 0.14, 0.20); III 0.74 (0.18, 0.10, 0.16, 0.12, 0.18); IV 0.88 (0.26, 0.12, 0.18, 0.12, 0.20).

Somatic characters (Fig. 28D–F). Coloration: prosoma and legs as in male. Abdomen black, with sclerotized dots dorsally. Prosoma: carapace pear shaped, surface modification and arrangement of eyes as in male. Cephalic area lower than in male. Mouthparts and sternum as in male. Abdomen: globular. Spinnerets tiny, lacking circular plate.

Epigyne (Fig. 29D–G): epigynal area slightly sclerotized. Scape protruded, as wide as long. Copulatory openings located at terminus of scape. Internal structures faintly visible via translucent tegument. Spermathecae strongly sclerotized, separated by their diameter. Fertilization ducts originating posteromedially from spermathecae. Copulatory ducts long, connected to posterior margin of spermathecae, curved upward and inward to center of vulva, then turned sharply downward to copulatory openings. The base of copulatory ducts unfused.

Figure 29. 

Crassignatha shunani sp. nov. A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Etymology

The specific name is from the Chinese pinyin shŭ nán, referring to the collection locality of this new spider species from southern Sichuan; noun in apposition.

Distribution

China (Sichuan) (Fig. 38).

Crassignatha si Y. Lin & S. Li, sp. nov.

Figs 30, 31, 38

Type material

Holotype ♂ (NHMSU Ar 127) and paratypes 2♀ (NHMSU Ar 128–129), China: Yunnan Province, Yiliang County, Jiuxiang Town, near entrance of Baiyan Cave, in low bush (25.15100°N, 103.40100°E; 1875 m), 24.VIII.2018, Y. Lin et al. leg.; 1♂ juvenile (NHMSU-HA141) and 1♀ juvenile (NHMSU-HA141) used for sequencing, GenBank: MT992017 and MT992016, same data as for preceding.

Figure 30. 

Crassignatha si sp. nov. A male habitus, dorsal, B male habitus, ventral C female habitus, dorsal D female habitus, ventral. Scale bars: 0.50 mm (A–D).

Diagnosis

The male of C. si sp. nov. differs from all other congeners by the presence of a helical embolus coiled slightly more than twice (Fig. 31B, C). The female of C. si sp. nov. is similar to C. gudu and C. yamu in the vulva configuration and the course of the copulatory ducts but can be distinguished from the latter two by the longer columnar atrium formed by the union of the copulatory ducts (Fig. 31F vs. Figs 11F, 35F).

Description

Male (holotype). Total length 0.56. Carapace 0.20 long, 0.24 wide, 0.28 high. Clypeus 0.04 high. Sternum 0.16 long, 0.16 wide. Abdomen 0.36 long, 0.36 wide, 0.48 high. Length of legs: I 0.88 (0.24, 0.10, 0.20, 0.14, 0.20); II 0.76 (0.24, 0.08, 0.16, 0.10, 0.18); III 0.6 (0.18, 0.08, 0.12, 0.10, 0.12); IV 0.68 (0.18, 0.08, 0.14, 0.10, 0.18).

Somatic characters (Fig. 30A, B). Coloration: prosoma and legs fuscous. Abdomen grayish green, ventrally darker than dorsally, with pale yellow speckles. Prosoma: carapace nearly round, surface weakly sculptured. ALE protruded. PER slightly recurved. Cephalic area elevated. Clypeus concave. Chelicerae fused near base, covered with long setae anteriorly. Labium triangular. Sternum heart shaped, swollen, surface pitted. Legs: femora and tibiae rough, granular. Tibia II with a clasping spur. Abdomen: globose and rugose, lacking lateral scutum and circular plate. Spinnerets pale brown.

Palp (Fig. 31A–C): femur swollen, wider than patella. Tibia flat. Cymbium weakly sclerotized, with a few setae distally. Cymbial tooth small, inconspicuous. Tegulum narrow, slightly swollen. Laminar median apophysis sub-rounded, translucent, with a small, hooked process on prolateral margin. Embolic membrane arises from behind median apophysis. Embolus long, helical, circling clockwise into almost two full coils.

Female (one of paratypes). Total length 0.92. Carapace 0.36 long, 0.32 wide, 0.36 high. Clypeus 0.06 high. Sternum 0.24 long, 0.24 wide. Abdomen 0.56 long, 0.56 wide, 0.52 high. Length of legs: I 1.22 (0.40, 0.16, 0.28, 0.18, 0.20); II 1.02 (0.28, 0.14, 0.22, 0.18, 0.20); III 0.66 (0.16, 0.08, 0.18, 0.10, 0.14); IV 0.86 (0.24, 0.08, 0.22, 0.14, 0.18).

Somatic characters (Fig. 30C, D). Coloration: prosoma and legs dark brown. Abdomen color and modification as in male. Prosoma: carapace nearly pear shaped, surface granular and sculptured, two strong setae medially. ALE slightly protruded. PER slightly recurved. Mouthparts as in male. Sternum scutiform, surface textured. Abdomen: globose, with small sclerotized patches dorsally and ventrally. Spinnerets weakly sclerotized.

Epigyne (Fig. 31D–F): epigynal area lightly sclerotized. Scape slightly protruded, as wide as long. Internal structures faintly visible via translucent tegument. Paired spermathecae separated by their diameter. Fertilization ducts slender, starting at the inside medial margin of spermathecae. Copulatory ducts long and extremely tortuous, connected to posterior margin of spermathecae, extending from under spermathecae to venter, forming two open loops, curving upward to center of vulva, then fusing into a columnar atrium that reaches copulatory opening at scape terminus.

Figure 31. 

Crassignatha si sp. nov. A right male palp, prolateral B right male palp, ventral C right male palp, retrolateral D epigyne, ventral E vulva, ventral F vulva, dorsal. Scale bars: 0.10 mm (A–F).

Etymology

The specific name is derived from the Chinese pinyin word for “spiral” (si), referring to the shape of the embolus, and is a noun in apposition.

Distribution

China (Yunnan) (Fig. 38).

Crassignatha thamphra Y. Lin & S. Li, sp. nov.

Figs 32, 38

Type material

Holotype ♀ (IZCAS-Ar 41035), Thailand: Khon Kaen Province, Phu Pha Man District, Phu Pha Man Subdistrict, Tham Phra Cave (16.66603°N, 101.89623°E; 262 m), 10.XI.2016, H. Zhao, Y. Li and Z. Chen leg.; 1♀ (NHMSU-HA089) used for sequencing, GenBank: MT992003, same data as for preceding.

Diagnosis

This species differs other Crassignatha species, except C. yinzhi, by the copulatory ducts diagonally connected to the copulatory opening and not fused before reaching copulatory opening. It can be easily distinguished from C. yinzhi by the larger spermathecae separated by less than their diameter and the tighter turns of the copulatory ducts at center of the vulva (Fig. 32G vs. Fig. 37G).

Figure 32. 

Crassignatha thamphra sp. nov. A female habitus, dorsal B female habitus, ventral C female habitus, lateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.50 mm (A–C); 0.10 mm (D–G).

Description

Female (holotype). Total length 0.64. Carapace 0.32 long, 0.32 wide, 0.28 high. Clypeus 0.10 high. Sternum 0.20 long, 0.20 wide. Abdomen 0.44 long, 0.48 wide, 0.48 high. Length of legs: I 0.92 (0.28, 0.12, 0.20, 0.16, 0.16); II 0.74 (0.22, 0.10, 0.14, 0.12, 0.16); III 0.64 (0.16, 0.10, 0.12, 0.12, 0.14); IV 0.84 (0.24, 0.12, 0.18, 0.12, 0.18).

Somatic characters (Fig. 32A–C). Coloration: prosoma yellowish brown. Legs dark brownish. Abdomen black with faint, sclerotized dots. Prosoma: carapace nearly pear shaped, surface indistinctly textured. Cephalic area elevated. ALE slightly protruded. PER straight. Chelicerae bears sparse, short setae anteriorly. Labium tongue shaped. Sternum heart shaped, slightly plump, surface smooth, truncated posteriorly. Legs: covered with setae and bristles. Abdomen: abdominal shape irregular and surface rugose (caused by alcohol immersion). Spinnerets weakly sclerotized.

Epigyne (Fig. 32D–G): epigynal area slightly sclerotized, bears a few long setae. Scape protruded, longer slightly than wide. Internal structures faintly visible via translucent tegument. Spermathecae separated by ~ ½ their diameter. Fertilization ducts thin, slender, originating from posteromedial margin of spermathecae, forming a U-shape. Copulatory ducts long, connected to the posterior surface of spermathecae, curving twice below spermathecae, linked diagonally, and fused to copulatory opening.

Male. Unknown.

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

Thailand (Fig. 38).

Crassignatha xichou Y. Lin & S. Li, sp. nov.

Figs 33, 38

Type material

Holotype ♀ (NHMSU Ar 130), China: Yunnan Province, Wenshan Prefecture, Xichou County, near to radio and television transmitting tower, in leaf litter (23.43302°N, 104.67320°E; 1556 m), 17.V.2015, Z. Chen and Y. Li leg.; paratype 1♀ (NHMSU Ar 131), China: Yunnan Province, Nanjian County, Xiaowan Township, Huilongshan Village, near to entrance of Banpoyan Cave, in bushes (24.93353°N, 100.31443°E; 1990 m), 23.VIII.2018, Y. Lin et al. leg.

Diagnosis

This new species can be easily distinguished from all species of Crassignatha by the lack of a scape, the fertilization ducts starting at the posterolateral margin of the spermathecae, the copulatory ducts connecting to the anterolateral margin of spermathecae, fusing into an H-shaped atrium above copulatory opening (Fig. 33E, F).

Figure 33. 

Female of Crassignatha xichou sp. nov. A habitus, dorsal B habitus, ventral C habitus, lateral D epigyne, ventral E vulva, ventral F vulva, dorsal. Scale bars: 0.50 mm (A–C); 0.10 mm (D–F).

Description

Female (holotype). Total length 0.60. Carapace 0.24 long, 0.28 wide, 0.24 high. Clypeus 0.06 high. Sternum 0.16 long, 0.16 wide. Abdomen 0.44 long, 0.40 wide, 0.48 high. Length of legs: I 0.88 (0.24, 0.10, 0.20, 0.14, 0.20); II 0.78 (0.20, 0.10, 0.18, 0.12, 0.18); III 0.64 (0.14, 0.10, 0.14, 0.10, 0.16); IV 0.74 (0.20, 0.10, 0.16, 0.10, 0.18).

Somatic characters (Fig. 33A–C). Coloration: carapace dark brownish, darker in thoracic center and margins. Mouthparts and sternum dark. Abdomen dark grayish. Prosoma: carapace nearly pear shaped, surface indistinctly textured. Cephalic part raised. ALE slightly protruded. PER straight. Chelicerae lighter than endites and labium, bears short setae anteriorly. Labium nearly semicircular. Sternum scutiform, slightly swollen, surface pitted, truncated posteriorly. Legs: metatarsi and tarsi yellow-brown, tibiae and femora dark. Abdomen: oval dorsally, with light brown sclerotized dots. Spinnerets tiny.

Epigyne (Fig. 33D–F): epigynal area dark, slightly sclerotized, with sparse, short setae. Scape absent. Internal structures faintly visible via translucent tegument. Paired spermathecae small, globose, separated by their diameter. Fertilization ducts originating posteromedially from spermathecae, coiled below spermathecae. Copulatory ducts long, connected to outer lateral margin of spermathecae, making four turns before merging into an H-shaped atrium.

Male. Unknown.

Etymology

The specific name is derived from the type locality; noun in apposition.

Distribution

China (Yunnan) (Fig. 38).

Crassignatha yamu Miller, Griswold & Yin, 2009

Figs 34, 35, 38

Crassignatha yamu Miller et al., 2009: 73, figs 76H, I, 86A–C, 87A, B (♂♀).

Type material

Holotype ♂ (HNU-CASENT 9029321) and paratype 1♀ (HNU-CASENT 9020752), China: Yunnan Province, Fugong County, S fork Yamu River, 1.51 km 150° SW of confluence [with N Fork], Gaoligongshan, moist shaded embankments (27.11905°N, 98.83108°E; 1723 m), 26.IV.2004, C. Griswold leg.; paratypes 2♀ (HNU-CASENT 9020736), China: Yunnan Province, Fugong County, 10th km W NuJiang on Shibali Rd., N fork Yamu River, Gaoligongshan, moist earthen embankments (27.13795°N, 98.82240°E; 1850 m), 25.IV.2004, C. Griswold leg. Examined.

Figure 34. 

Crassignatha yamu Miller, Griswold & Yin, 2009 A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Other material examined

6♂ 8♀ (NHMSU-HA115), China: Yunnan Province, Fugong County, Yamu River, ca. 1.51 km on 150° SW from confluence [with N fork], Gaoligongshan, moist shaded embankments (27.11905°N, 98.83108°E; 1723 m), 18.VIII.2018, Y. Lin et al.; 1♂ (NHMSU-HA115) and 1♀ (NHMSU-HA115) used for sequencing, GenBank: MT992011 and MT992010, same data as for preceding; 3♂ 7♀ (NHMSU-HA116), China: Yunnan Province, Fugong County, Shilajia Village, near estuary of Yamu River (27.13440°N, 98.82625°E; 1792 m), 18.VIII.2018, Y. Lin et al. leg.

Diagnosis

The male of C. yamu is most similar to that of C. haeneli, C. danaugirangensis, and C. shiluensis in the form of the palp and the long, linear embolus but differs from C. haeneli and C. danaugirangensis by the spiral embolus on the ventral portion of the palpal bulb (Fig. 25B vs. Wunderlich 1995: figs 18, 19; Miller et al. 2004: fig. 4); from C. shiluensis by having fewer loops of the embolus (Fig. 35B vs. Fig. 27A). The female is similar to that of C. gudu and C. si in vulva configuration and the course of the copulatory ducts but can be easily distinguished by having the columnar atrium longer than in C. gudu and shorter than in C. si (Fig. 35G vs. Figs 11F, 31F).

Figure 35. 

Crassignatha yamu Miller, Griswold & Yin, 2009 A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Description

See Miller et al. (2009).

Distribution

China (Yunnan) (Fig. 38).

Crassignatha yinzhi Miller, Griswold & Yin, 2009

Figs 36, 37, 38

Crassignatha yinzhi Miller et al., 2009: 71, figs 76C, D, 78E, 79C, D, 80A–E, 81A, B, 82A–F (♂♀).

Type material

Holotype ♂ (HNU-CASENT 9029322) and paratypes 5♀ (HNU-CASENT 9022376), China: Yunnan Province, Longling County, Longjiang Township, Xiaoheishan Nature Reserve, 1.2 km SSE of Route S317 at 23.5 km, good primary broadleaf forest, dusting webs in understory (24.82888°N, 98.76001°E; 2020 m), 27–28.V.2005, C. Griswold leg.; 1♂ 1♀ (HNU-CASENT 9022354), China: Yunnan Province, Longling County, Longjiang Township, Xiaoheishan Nature Reserve, 1.2 km SSE of Route S317 at 23.5 km, good primary broadleaf forest, night collecting (24.82888°N, 98.76001°E; 2020 m), 28.V.2005, C. Griswold and D. Kavanaugh leg.; 1♀ (HNU-CASENT 9022396), China: Yunnan Province, Longling County, Longjiang Township, Xiaoheishan Nature Reserve, 1.2 km SSE of Route S317 at 23.5 km, good primary broadleaf forest, night collecting (24.82888°N, 98.76001°E; 2020 m), 26.V.2005, C. Griswold and D. Kavanaugh leg. Examined.

Figure 36. 

Crassignatha yinzhi Miller, Griswold & Yin, 2009 A male habitus, dorsal B male habitus, ventral C male habitus, lateral D female habitus, dorsal E female habitus, ventral F female habitus, lateral. Scale bars: 0.50 mm (A–F).

Figure 37. 

Crassignatha yinzhi Miller, Griswold & Yin, 2009 A male palp, prolateral B male palp, ventral C male palp, retrolateral D epigyne, ventral E epigyne, lateral F vulva, ventral G vulva, dorsal. Scale bars: 0.10 mm (A–G).

Other material examined

6♂ 6♀ (NHMSU-HA117), China: Yunnan Province, Longling County, Longjiang Township, Xiaoheishan Nature Reserve (24.82888°N, 98.76001°E; 2020 m), 22.VIII.2018, Y. Lin et al. leg.; 1♂ (NHMSU-HA117) and 1♀ (NHMSU-HA117) used for sequencing, GenBank: MT992013 and MT992012, same data as for preceding; 1♂ 1♀ (NHMSU-HA127), China: Yunnan Province, Longling County, Longjiang Township, Xiaoheishan Nature Reserve, Gucheng Hill, good forest (24.82886°N, 98.75917°E; 2010 m), 22.VIII.2018, Y. Lin et al. leg.

Diagnosis

The male of C. yinzhi is similar to C. ertou but can be distinguished by the nearly straight embolus and the caniniform cymbial tooth, rather than spiraled embolus and hooked cymbial tooth in the latter (Fig. 37A, B vs. Fig. 8A, B). The female of C. yinzhi is most similar to that of C. thamphra sp. nov. in vulva configuration but differs from the latter by the more widely separated spermathecae and the widely separated inflection points of the copulatory ducts (Fig. 37G vs. Fig. 32G).

Description

See Miller et al. (2009).

Distribution

China (Yunnan) (Fig. 38).

Figure 38. 

Distribution records of Crassignatha spp. in Asia. 1 C. baihua sp. nov. 2 C. bangbie sp. nov. 3 C. bicorniventris 4 C. changyan sp. nov. 5 C. danaugirangensis 6 C. dongnai sp. nov. 7 C. ertou 8 C. gucheng sp. nov. 9 C. gudu 10 C. haeneli 11 C. mengla sp. nov. 12 C. nantou sp. nov. 13 C. nasalis sp. nov. 14 C. pianma 15 C. quadriventris 16 C. quanqu 17 C. rostriformis sp. nov. 18 C. shiluensis 19 C. shunani sp. nov. 20 C. si sp. nov. 21 C. thamphra sp. nov. 22 C. xichou sp. nov. 23 C. yamu 24 C. yinzhi.

Acknowledgements

The manuscript benefited greatly from comments by Yuri Marusik (Magadan, Russia) and Akio Tanikawa (Tokyo, Japan). Theo Blick (Hummeltal, Germany) checked etymology. Sarah Crews (San Francisco, USA) corrected the English of the final manuscript. Permission for field surveys was granted by Biodiversity Institute of Gongshan Administration Bureau (Director: Yingchun Li). Xianjin Peng and Xiang Xu (Changsha, China) facilitated the loan of type material of Chinese symphytognathids from the Gaoligong Mountains. This study was supported by the National Natural Science Foundation of China (NSFC-31772410, 31750002, 31972870).

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Appendix 1

Table A1.

Uncorrected genetic pairwise distance (below diagonal) and standard error (above diagonal) of a partial fragment of COI from the seventeen species discussed in this text.

Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
♂J ♂J ♂J ♀J ♂J
1 C. baihua sp. nov. 0.003 0.012 0.008 0.013 0.005 0.008 0.008 0.012 0.012 0.012 0.012 0.011 0.011 0.007 0.010 0.011 0.016 0.016 0.013 0.013 0.012 0.012 0.012 0.012 0.012 0.012 0.012
0.005 0.012 0.008 0.013 0.005 0.008 0.008 0.012 0.012 0.012 0.012 0.011 0.010 0.007 0.010 0.010 0.016 0.016 0.013 0.013 0.011 0.011 0.012 0.012 0.011 0.012 0.012
2 C. bangbie sp. nov. 0.090 0.088 0.012 0.013 0.012 0.011 0.011 0.012 0.012 0.011 0.011 0.010 0.010 0.012 0.010 0.011 0.015 0.015 0.012 0.011 0.011 0.011 0.013 0.012 0.012 0.012 0.012
3 C. dongnai sp. nov. 0.045 0.044 0.090 0.013 0.009 0.007 0.007 0.012 0.012 0.012 0.011 0.011 0.011 0.008 0.011 0.011 0.016 0.016 0.013 0.013 0.012 0.012 0.012 0.012 0.012 0.012 0.012
4 C. ertou 0.104 0.099 0.101 0.106 0.013 0.013 0.013 0.012 0.012 0.013 0.013 0.013 0.013 0.013 0.012 0.012 0.017 0.017 0.013 0.013 0.013 0.013 0.013 0.013 0.013 0.011 0.011
5 C. gucheng sp. nov. 0.022 0.021 0.087 0.044 0.103 0.008 0.008 0.012 0.012 0.012 0.012 0.011 0.011 0.006 0.011 0.011 0.016 0.016 0.012 0.012 0.011 0.011 0.012 0.012 0.011 0.012 0.012
6 C. mengla sp. nov. 0.040 0.039 0.080 0.026 0.103 0.037 0.000 0.011 0.011 0.011 0.011 0.011 0.011 0.007 0.010 0.010 0.016 0.016 0.012 0.012 0.011 0.011 0.012 0.012 0.011 0.012 0.012
♂J 0.040 0.039 0.080 0.026 0.103 0.037 0.000 0.011 0.011 0.011 0.011 0.011 0.011 0.007 0.010 0.010 0.016 0.016 0.012 0.012 0.011 0.011 0.012 0.012 0.011 0.012 0.012
7 C. nantou sp. nov. 0.097 0.096 0.090 0.094 0.101 0.097 0.090 0.090 0.000 0.010 0.010 0.012 0.012 0.012 0.011 0.012 0.016 0.016 0.008 0.008 0.012 0.012 0.013 0.013 0.012 0.012 0.012
0.097 0.096 0.090 0.094 0.101 0.097 0.090 0.090 0.000 0.010 0.010 0.012 0.012 0.012 0.011 0.012 0.016 0.016 0.008 0.008 0.012 0.012 0.013 0.013 0.012 0.012 0.012
8 C. nasalis sp. nov. 0.088 0.087 0.081 0.085 0.097 0.080 0.085 0.085 0.071 0.071 0.004 0.011 0.011 0.012 0.011 0.011 0.015 0.015 0.010 0.010 0.011 0.011 0.012 0.012 0.011 0.011 0.011
0.083 0.081 0.078 0.080 0.096 0.078 0.080 0.080 0.069 0.069 0.010 0.011 0.011 0.011 0.011 0.011 0.015 0.015 0.010 0.011 0.011 0.011 0.012 0.012 0.011 0.011 0.011
9 C. pianma 0.083 0.081 0.062 0.085 0.105 0.076 0.083 0.083 0.105 0.105 0.085 0.085 0.003 0.011 0.011 0.011 0.016 0.016 0.011 0.012 0.011 0.011 0.013 0.013 0.012 0.011 0.011
0.080 0.078 0.057 0.083 0.099 0.076 0.080 0.080 0.101 0.101 0.081 0.081 0.005 0.011 0.010 0.011 0.016 0.016 0.011 0.011 0.011 0.011 0.013 0.012 0.012 0.011 0.011
10 C. quadriventris 0.032 0.031 0.088 0.044 0.104 0.027 0.035 0.035 0.101 0.101 0.088 0.083 0.081 0.078 0.010 0.011 0.016 0.016 0.012 0.012 0.010 0.010 0.012 0.012 0.011 0.012 0.012
11 C. rostriformis sp. nov. 0.068 0.066 0.071 0.074 0.094 0.071 0.066 0.066 0.088 0.088 0.074 0.071 0.073 0.067 0.068 0.001 0.016 0.016 0.012 0.012 0.010 0.010 0.011 0.011 0.010 0.012 0.011
0.069 0.068 0.073 0.076 0.096 0.073 0.067 0.067 0.090 0.090 0.076 0.073 0.074 0.069 0.069 0.002 0.016 0.016 0.012 0.012 0.010 0.010 0.011 0.011 0.010 0.012 0.011
12 C. shiluensis 0.144 0.150 0.155 0.147 0.175 0.147 0.151 0.151 0.153 0.153 0.140 0.136 0.157 0.155 0.149 0.155 0.155 0.000 0.015 0.015 0.015 0.015 0.015 0.017 0.016 0.016 0.016
♂J 0.144 0.150 0.155 0.147 0.175 0.147 0.151 0.151 0.153 0.153 0.140 0.136 0.157 0.155 0.149 0.155 0.155 0.000 0.015 0.015 0.015 0.015 0.015 0.017 0.016 0.016 0.016
13 C. shunani sp. nov. 0.099 0.097 0.088 0.101 0.108 0.092 0.094 0.094 0.041 0.041 0.068 0.071 0.096 0.092 0.092 0.092 0.094 0.134 0.134 0.004 0.011 0.011 0.013 0.013 0.012 0.011 0.012
♂J 0.101 0.099 0.087 0.103 0.110 0.094 0.096 0.096 0.042 0.042 0.069 0.073 0.097 0.094 0.094 0.094 0.096 0.142 0.142 0.010 0.011 0.011 0.013 0.013 0.012 0.012 0.012
14 C. si sp. nov. ♀J 0.088 0.087 0.078 0.083 0.094 0.081 0.081 0.081 0.097 0.097 0.080 0.078 0.083 0.078 0.074 0.064 0.066 0.149 0.149 0.088 0.087 0.000 0.013 0.007 0.006 0.011 0.011
♂J 0.088 0.087 0.078 0.083 0.094 0.081 0.081 0.081 0.097 0.097 0.080 0.078 0.083 0.078 0.074 0.064 0.066 0.149 0.149 0.088 0.087 0.000 0.013 0.007 0.006 0.011 0.011
15 C. thamphra sp. nov. 0.090 0.088 0.094 0.094 0.104 0.083 0.094 0.094 0.108 0.108 0.088 0.087 0.095 0.092 0.088 0.080 0.081 0.142 0.142 0.104 0.110 0.095 0.095 0.013 0.012 0.011 0.011
16 C. yamu 0.096 0.096 0.096 0.101 0.101 0.085 0.090 0.090 0.109 0.109 0.087 0.083 0.103 0.097 0.090 0.076 0.078 0.163 0.163 0.101 0.103 0.036 0.036 0.097 0.003 0.012 0.012
0.094 0.092 0.087 0.097 0.096 0.081 0.087 0.087 0.099 0.099 0.082 0.078 0.094 0.088 0.087 0.071 0.073 0.155 0.155 0.092 0.094 0.027 0.027 0.094 0.008 0.011 0.011
17 C. yinzhi 0.092 0.090 0.090 0.088 0.076 0.081 0.087 0.087 0.097 0.097 0.074 0.069 0.083 0.081 0.087 0.081 0.083 0.151 0.151 0.083 0.088 0.083 0.083 0.080 0.094 0.088 0.004
0.092 0.090 0.090 0.088 0.073 0.081 0.087 0.087 0.097 0.097 0.071 0.069 0.080 0.078 0.087 0.074 0.076 0.153 0.153 0.088 0.090 0.080 0.080 0.078 0.094 0.088 0.010
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