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Research Article
Taxonomic revision of Telemidae (Arachnida, Araneae) from East and Southeast Asia
expand article infoHuifeng Zhao§, Shuqiang Li|, Aibing Zhang
‡ Capital Normal University, Beijing, China
§ Langfang Normal University, Langfang, China
| Institute of Zoology, Chinese Academy of Sciences, Beijing, China
Open Access

Abstract

Species of the spider family Telemidae Fage, 1913 from East and Southeast Asia are revised. Four new genera are erected: Mekonglema Zhao & Li, gen. nov. with the type species Mekonglema bailang Zhao & Li, sp. nov. (♂♀, Yunnan, China), Siamlema Zhao & Li, gen. nov. with the type species Siamlema changhai Zhao & Li, sp. nov. (♂♀, southern Thailand), Sundalema Zhao & Li, gen. nov. with the type species Sundalema bonjol Zhao & Li, sp. nov. (♂♀, Sumatra), and Zhuanlema Zhao & Li, gen. nov. with the type species Zhuanlema peteri Zhao & Li, sp. nov. (♂♀, northern Laos). Eight additional new species are described: Mekonglema kaorao Zhao & Li, sp. nov. (♂♀, northern Laos), M. walayaku Zhao & Li, sp. nov. (♂♀, Yunnan, China), M. yan Zhao & Li, sp. nov. (♂♀, Yunnan, China), Pinelema daguaiwan Zhao & Li, sp. nov. (♂♀, Guangxi, China), P. shiba Zhao & Li, sp. nov. (♂♀, Guangxi, China), P. tham Zhao & Li, sp. nov. (♂♀, northern Laos), Siamlema suea Zhao & Li, sp. nov. (♂♀, southern Thailand), and Sundalema khaorakkiat Zhao & Li, sp. nov. (♂♀, southern Thailand). Thirty species are transferred from the genus Telema Simon, 1882 to the genera Pinelema Wang & Li, 2012, Sundalema gen. nov., and Telemofila Wunderlich, 1995. Seychellia xinpingi Lin & Li, 2008 is transferred to Mekonglema gen. nov. as M. xinpingi comb. nov. Furthermore, the genus Pinelema is divided into seven species groups based on male morphological characters.

Keywords

Haplogynae, molecular phylogeny, new genus, new species, new combination

Introduction

The spider family Telemidae Fage, 1913 currently includes 85 species in ten genera (Li 2020). Telemids are tiny spiders, whose body length ranges from 0.9 to 2.2 mm, with much longer legs relative to their body. The dispersal ability of telemids is poor, resulting in high endemism (Lin and Li 2010; Wang and Li 2010b; Zhao et al. 2018a, b).

Telemids occur in tropical rainforests and karst caves in the southern Holarctic, Ethiopian, Oriental, Neotropical, and Australasian Realms (except Australia and New Zealand) (WSC 2020). Telema tenella Simon, 1882 occurs in Spain and France and is the type species of the genus Telema Simon, 1882 and the only known telemid species from Europe. Thirty-four Asian species were classified in the poorly defined genus Telema prior to the current study, and a revision of Asian Telema is necessary. Furthermore, the placement of Seychellia xinpingi Lin & Li, 2008 from southwestern China requires examination because the shape of the bulbal apophysis of the male differs from the generotype Seychellia wiljoi Saaristo, 1978, from Seychelles.

The goal of this paper is to revise all Asian telemid species using combined morphological and molecular approaches, adding new materials collected from Southeast Asia and southwestern China.

Materials and methods

Specimens used in this paper were collected by sifting leaf litter in rainforests or collected by hand from caves. All samples were examined and measured using a LEICA M205C stereomicroscope. The habitus, left male palp, and endogyne were photographed using an Olympus C7070 wide zoom digital camera. Images were montaged using Helicon Focus Lite 7.5.6 software. Female genitalia were removed and treated in lactic acid before being photographed. All measurements are given in millimetres. Leg measurements are shown as: total length (femur, patella, tibia, metatarsus, tarsus). For SEM images, the tibial glands on leg III were photographed using a Hitachi SU8010 Environmental Scanning Electron Microscope.

For molecular phylogenetic analyses, we used all available materials of Asian telemids as well as the type species Telema tenella and Seychellia wiljoi. Our analyses contain 57 of 62 known Asian species from the type localities and 12 potentially new species. Five species for which we did not obtain molecular data are Sundalema acicularis (Wang & Li, 2010) comb. nov., Pinelema claviformis (Tong & Li, 2008) comb. nov., Telemofila malaysiaensis (Wang & Li, 2010) comb. nov., Telema nipponica (Yaginuma, 1972), and Pinelema spina (Tong & Li, 2008) comb. nov. We used two segestriid species as outgroups, as Segestriidae is considered the sister lineage of Telemidae (Shao and Li 2018). In total, 73 taxa were included in our molecular dataset.

Genomic DNA was extracted from the legs or prosoma using TIANamp Genomic DNA Kit DP304 (TIANGEN Co., Beijing, China). Two nuclear loci, Histone 3 and Wingless (H3 and Wnt), were amplified for subsequent molecular phylogenetic analyses. Primer information and PCR protocols are shown in Suppl. material 1: Table S1. All amplicons were sequenced using an ABI 3730 automated sequencer, and raw sequences were corrected manually in BioEdit (Hall 1999). Sequence alignments were produced using Clustal W in MEGA 5 (Tamura et al. 2011), and the sequences were checked for stop codons after translation to amino acid sequences.

Phylogenetic relationships were inferred using maximum likelihood (ML) and Bayesian inference (BI). ML was performed in RAxML 7.0.4 (Stamatakis 2014) using the default rapid hill-climbing algorithm and the GTRGAMMA model to search for the best tree. Clade support was assessed using 1000 rapid bootstrap replicates. BI was performed in MrBayes 3.2.6 (Ronquist et al. 2012) using the best model selected by jModeltest 2.1.7 (Darriba et al. 2012) based on the Akaike information criterion (AIC). BI analysis was run for 10 million generations, sampling every 1000 generations, and was checked to ensure the average standard deviation of split frequencies was less than 0.01.

References to figures from the cited papers are listed in lowercase (fig. or figs), and figures in this paper are noted with an initial capital (Fig. or Figs). The following abbreviations are used:

Ca Cymbial apophysis

Em Embolus

BA Bulbal apophysis

Re Receptacle

SR Spiral ridge on embolus

Abbreviations of institutes:

AMNH American Museum of Natural History, New York, USA

IZCAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China

MHBU Museum of Hebei University, Baoding, Hebei, China

MLR Maolan National Natural Reserve, Libo, Guizhou, China

RMNH National Museum of Natural History, Leiden, the Netherlands

SMF Senckenberg Research Institute in Frankfurt, Germany

Types are deposited in IZCAS, except Mekonglema kaorao sp. nov. and Zhuanlema peteri sp. nov., which are lodged in SMF.

Taxonomy

Family Telemidae Fage, 1913

Key to telemid genera occurring in East and Southeast Asia (Males)

(Apneumonella Fage, 1921 was excluded as the male of A. jacobsoni Brignoli, 1977 is unknown).

1 Tibial glands plate-shaped (Fig. 1A; Emerit 1984: fig. A), cymbial apophysis absent (Wang et al. 2012: fig. 2C) Telema Simon, 1882
Tibial glands belt-shaped (Fig. 1B–G; Emerit 1984: fig. C, D), cymbial apophysis present (Figs 12C, 20C, 25C, 28C; Wang and Li 2012: fig. 2; Wunderlich 1995: fig. 16) 2
2 Palpal tibia with a dorso-distal spine, cymbium shorter than femur (Fig. 20C, D) Siamlema gen. nov.
Palpal tibia without dorso-distal spine, cymbium longer than femur 3
3 Cymbial apophysis located baso-prolaterally (Fig. 28C), embolus twisted (Fig. 24C, D) Zhuanlema gen. nov.
Cymbial apophysis located sub-baso-, meso-, or sub-disto-prolaterally, embolus not twisted 4
4 Bulb spherical, embolus sickle-shaped (Wunderlich 1995: fig. 17), leg formula: 1-4-2-3 Telemofila Wunderlich, 1995
Bulb ellipsoidal or nearly ellipsoidal, embolus not sickle-shaped, leg formula: 1-2-4-3 5
5 Bulb with one apophysis, and tip of embolus directed ventrally (Fig. 12B–D); or bulb without apophysis, and tip of embolus directed dorsally (Fig. 14C, D) Mekonglema gen. nov.
Bulb without apophysis, and tip of embolus directed ventrally 6
6 Embolus with spiral ridge, triangular, trapezoidal, or tube-like Pinelema Wang & Li, 2012
Embolus without spiral ridge, nearly L-shaped Sundalema gen. nov.

Key to telemid genera occurring in East and Southeast Asia (Females)

1 Tibial glands plate-shaped (Fig. 1A; Emerit 1984: fig. A), and endogyne walking-stick shaped, with membranous tubes (Wang et al. 2012: fig. 3C, D) Telema Simon, 1882
Tibial glands belt-shaped (Fig. 1B–G; Emerit 1984: fig. C, D), and endogyne not walking-stick shaped, without tube inside, or with sclerotized or membranous tubes 2
2 Leg formula: 1-4-2-3 Telemofila Wunderlich, 1995
Leg formula: 1-2-4-3 3
3 Receptacle sclerotized 4
Receptacle membranous 5
4 Receptacle long and coiled 1.25 to several loops, not swollen distally (Fig. 24A) Sundalema gen. nov.
Receptacle globular, swollen distally 6
5 Receptacle without tubes inside 7
Receptacle with several membranous tubes inside Pinelema Wang & Li, 2012 and Apneumonella Fage, 19211
6 Neck of receptacle sclerotized, umbrella-shaped (Fig. 23A) Siamlema gen. nov. part I (S. suea sp. nov.)
Neck of receptacle membranous, tube-shaped (Fig. 28A) Zhuanlema gen. nov.
7 Receptacle swollen distally Mekonglema gen. nov. part I2
Receptacle not swollen distally Mekonglema gen. nov. part II and Siamlema gen. nov. part II3

Pinelema Wang & Li, 2012

Pinelema Wang and Li 2012: 76; Wang and Li 2016: 547; Zhao et al. 2018a: 14; Zhao et al. 2018b: 10.

Type species

Pinelema bailongensis Wang & Li, 2012 from Guangxi, China.

Diagnosis

Pinelema can be distinguished from Telema by the following: belt-shaped tibial glands (arrows on Fig. 1B) (vs. plate-shaped), the presence of a cymbial apophysis (vs. absent), and the triangular, trapezoidal, or tube-like embolus (vs. duckbill shaped). The endogyne is extended distally (vs. no extension).

Figure 1. 

Tibial glands on leg III of telemids in East and Southeast Asia (Zhuanlema gen. nov. is not included). A Telema guihua B Pinelema bailongensis C Mekonglema xinpingi comb. nov. D Siamlema changhai sp. nov. E Sundalema anguina comb. nov. F Apneumonella jacobsoni G Telemofila samosirensis.

Description

Total length: 1.07–1.77 (male), 1.20–2.02 (female). Carapace 0.48–0.82 long. Sternum with several long setae. Six eyes ringed with black, vestigial, or completely absent in some species. Legs thin, long relative to body. Leg formula: 1-2-4-3. Tibia I 0.94–2.20 long, glands of legs belt-shaped (Fig. 1B). Cymbial apophysis distinct, bulb large relative to body, oval, kidney-shaped, droplet-shaped, etc.; embolus triangular, tube-shaped, needle-shaped, etc., embolus short, medium, or long relative to cymbium. Receptacle with several membranous tubes inside and typically swollen distally.

Distribution

China (Guangxi, Guizhou, Yunnan, Hainan), Laos (Vien Tiane), and Vietnam (northern and central Vietnam) (Figs 3032).

Comments

All 26 new combinations are supported by morphological characters, such as the presence of a cymbial apophysis and the extended tip of the receptacle. Including the new species described in this paper increases the total number of Pinelema species to 54, making it the most speciose genus in Telemidae.

Composition

According to the morphological characters of the male palp, 54 Pinelema species have been divided into seven species groups as well as six species not attached to a species group. The composition of species groups is discussed below.

adunca -group

Figures 2A, 30

Diagnosis

This group resembles the pacchanensis-group by having a long bulb but can be distinguished by the bulb is protruding ventro-basally and concave dorso-sub-basally (Fig. 2A) (vs. concave ventro-sub-basally and protruding dorso-mesially).

Description

Body length 1.40–1.98. Carapace 0.60–0.82 long. Six eyes ringed with black or absent. Tibia I 1.40–2.20. Ratio of bulbal length/width 1.96–2.35, bulb protruding ventro-basally and concave dorso-subbasally, embolus membranous (except in P. tortutheca (Lin & Li) comb. nov.), length ratio of embolus/bulb 0.38–0.71. Receptacle J-shaped.

Distribution

China (Guangxi, sites 1–5 in Fig. 30).

Composition

Pinelema adunca (Wang & Li, 2010) comb. nov., P. qingfengensis Zhao & Li, 2017, P. renalis (Wang & Li, 2010) comb. nov., P. tortutheca comb. nov., and P. yashanensis (Wang & Li, 2010) comb. nov.

Figure 2. 

Pinelema spp., palp, retrolateral view. A The adunca-group B the bailongensis-group C the cunfengensis-group.

Pinelema adunca (Wang & Li, 2010), comb. nov.

Figures 2A, 30

Telema adunca Wang and Li 2010b: 2, figs 1–6 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Pingguo County, Bafeng Mountain, Guandi Cave, 23.5670N, 107.6794E, elevation ca. 285 m, 3.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema adunca comb. nov. resembles P. yashanensis comb. nov. but can be distinguished by the following: the smaller length ratio of the embolus/bulb (0.64, Fig. 2A, and cf. Wang and Li 2010b: fig. 1A, B) (vs. 0.85), the cymbial apophysis which is as long as the cymbial base (cf. Wang and Li 2010b: fig. 1B) (vs. ¼ times shorter); the receptacle tip is 3.20 times wider than the neck (cf. Wang and Li 2010b: fig. 3C) (vs. 1.20 times).

Description

See Wang and Li (2010b).

Distribution

China (Guangxi, site 1 in Fig. 30), known only from the type locality.

Pinelema qingfengensis Zhao & Li, 2017

Figures 2A, 30

Pinelema qingfengensis Song et al. 2017: 91, figs 5, 6, 7D, 8D, 9D, 10D, 11D, 12D (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Chongzuo Prefecture, Tiandeng County, Qingfeng Cave, 23.1720N, 107.1565E, elevation ca. 444 m, 26.XII.2012, Z. Chen and Z. Zhao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema qingfengensis resembles P. renalis comb. nov. but can be distinguished by the following: the absence of eyes (cf. Song et al. 2017: fig. 5A) (vs. present); the ventrally bent embolus, and the obtuse-angled bulb (Fig. 2A, and cf. Song et al. 2017: fig. 5C, D) (vs. right-angled), the bulb is not protruding ventro-distally (Fig. 2A, and cf. Song et al. 2017: fig. 5C, D) (vs. protruding), and the width ratio of the basal embolus/distal bulb is 1.0 (cf. Song et al. 2017: fig. 5C, D) (vs. 0.5).

Description

See Song et al. (2017).

Distribution

China (Guangxi, site 2 in Fig. 30), known only from the type locality.

Pinelema renalis (Wang & Li, 2010), comb. nov.

Figures 2A, 30

Telema renalis Wang and Li 2010b: 24, figs 24–27 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hezhou Prefecture, Zhongshan County, Guanyin Cave, 24.5233N, 111.3242E, elevation ca. 196 m, 25.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema renalis comb. nov. resembles P. qingfengensis but can be distinguished by the following: six eyes ringed with black (cf. Wang and Li 2010b: fig. 24A) (vs. no eyes); the embolus bent ventrally at a right angle (Fig. 2A, and cf. Wang and Li 2010b: fig. 24C, D) (vs. obtuse angle), the bulb protrudes ventro-distally (Fig. 2A, and cf. Wang and Li 2010b: fig. 24C, D) (vs. not protruding), and the width ratio of the basal embolus/distal bulb is 0.5 (cf. Wang and Li 2010b: fig. 24C, D) (vs. 1.0).

Description

See Wang and Li (2010b).

Distribution

China (Guangxi, site 3 in Fig. 30), known only from the type locality.

Pinelema tortutheca (Lin & Li, 2010), comb. nov.

Figures 2A, 30

Telema tortutheca Lin and Li 2010: 26, figs 16, 17 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Nanning Prefecture, Mashan County, Guling Town, Yangyu Village, Jinlun Cave, 23.5663N, 107.2622E, elevation ca. 1490 m, 6.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema tortutheca comb. nov. can be distinguished from all four other species in this group by the sclerotized embolus (Fig. 2A, and cf. Lin and Li 2010: fig. 16C) (vs. membranous).

Description

See Lin and Li (2010).

Distribution

China (Guangxi, site 4 in Fig. 30), known only from the type locality.

Pinelema yashanensis (Wang & Li, 2010), comb. nov.

Figures 2A, 30

Telema yashanensis Wang and Li 2010b: 33, figs 28–32 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Laibin Prefecture, Yashan County, Yashan Cave, 23.6025N, 108.9124E, elevation ca. 115 m, 16.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema yashanensis comb. nov. resembles P. adunca comb. nov. but can be distinguished by the following: the larger length ratio of the embolus/bulb (0.85, Fig. 2A, and cf. Wang and Li 2010b: fig. 28A, B) (vs. 0.64); the cymbial apophysis ¼ as long as the cymbial base (cf. Wang and Li 2010b: fig. 28A) (vs. equal length); and the receptacle tip is 1.20 times wider than the neck (cf. Wang and Li 2010b: fig. 30C) (vs. 3.20 times).

Description

See Wang and Li (2010b).

Distribution

China (Guangxi, site 5 in Fig. 30), known only from the type locality.

bailongensis -group

Figures 2B, 30

Diagnosis

This group can be distinguished from other species groups in Pinelema by the following: the length ratio of the embolus/bulb is 1.19–1.80 (Fig. 2B) (vs. 0.37–0.84), the junction of the bulb and cymbium is located ventro-mesially on the bulb (Fig. 2B) (vs. ventro-basally, except the cunfengensis-group).

Description

Body length 1.11–1.75. Carapace 0.50–0.82 long. Tibia I 0.90–2.18 long. Six eyes ringed with black or absent. Embolus longer than bulb, length ratio of embolus/bulb is 1.19–1.80, junction of bulb and cymbium located ventro-mesially on bulb (Fig. 2B), bulb with papillae proximo-retrolaterally (except P. curcici Wang & Li, 2016, P. huoyan Zhao & Li, 2018, and P. xiushuiensis Wang & Li, 2016). Endogyne U-shaped, J-shaped, or spiralled.

Distribution

China (Guangxi, Guizhou, Yunnan, sites 6–18 in Fig. 30).

Composition

Pinelema bailongensis, P. cheni Zhao & Li, 2018, P. cordata (Wang & Li, 2010), P. curcici, P. huoyan, P. liangxi (Zhu & Chen, 2002), P. lizhuang Zhao & Li, 2018, P. strentarsi (Lin & Li, 2010), P. wangshang Zhao & Li, 2018, P. wenyang Zhao & Li, 2018, P. xiushuiensis, P. yunchuni Zhao & Li, 2018, and P. zhewang (Chen & Zhu, 2009).

Remarks

For the diagnoses and descriptions of this group (except P. bailongensis and P. curcici), see Zhao et al. (2018b).

Pinelema bailongensis Wang & Li, 2012

Figures 1B, 2B, 30

Pinelema bailongensis Wang and Li 2012: 82, figs 1–17 (♂♀); Song et al. 2017: 85, figs 7A, 8A, 9A, 10A, 11A, 12A (♂); Zhao et al. 2018b: fig. 1 (♂).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Pingguo County, Bailong Cave, 23.3182N, 107.5731E, elevation ca. 111 m, 1.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

For differences between P. bailongensis and P. curcici, see Wang and Li (2016); for differences between P. bailongensis and the other eleven species in this group, see Zhao et al. (2018b).

Description-amendments

Tibial glands belt-shaped (Fig. 1B), the arrangement of secretory orifices linear within a smooth, striped tegument (arrows on Fig. 1B). For a more detailed description, see Wang and Li (2012), Song et al. (2017), and Zhao et al. (2018b).

Distribution

China (Guangxi, site 6 in Fig. 30), known only from the type locality.

Pinelema curcici Wang & Li, 2016

Figures 2B, 30

Pinelema curcici Wang and Li 2016: 547, figs 1–4 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Yunnan Province, Wenshan Zhuang and Miao Autonomous Prefecture, Qiubei County, Shuanglongying Town, Fengwei Cave, 24.3361N, 104.2862E, elevation ca. 1372 m, 20.VIII.2010, Z. Yao, C. Wang and X. Wang leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema curcici resembles P. liangxi but can be distinguished by the following: the presence of eyes (vs. absent); the embolic tip is much narrower than the embolic base (Fig. 2B) (vs. equal), the smaller ratio of the bulbal length/width (1.13, Fig. 2B, and cf. Wang and Li 2016: fig. 1C, D) (vs. 1.27), the greater length ratio of the embolus/bulb (1.31, Fig. 2B, and cf. Wang and Li 2016: fig. 1C, D) (vs. 1.19); the J-shaped receptacle (vs. C-shaped), with the distal part of the receptacle 1.75 times wider than the neck of the receptacle (cf. Wang and Li 2016: fig. 2C) (vs. three times).

Description

See Wang and Li (2016).

Distribution

China (Yunnan, site 9 in Fig. 30), known only from the type locality.

cunfengensis -group

Figures 2C, 30

Diagnosis

This group resembles the feilong-group by having a triangular and short embolus relative to the bulb length but can be distinguished by the following: the bulb is bent at a right angle dorso-distally (Fig. 2C) (vs. not bent), and the junction of the bulb and cymbium is located dorso-mesially on the bulb (Fig. 2C) (vs. dorso-basally); the receptacle is U-shaped or coiled (vs. bag-like or globular).

Description

Body length 1.33–1.48. Carapace 0.48–0.64 long. Tibia I 1.15–1.98 long. Six eyes ringed with black (P. cunfengensis Zhao & Li, 2017) or absent (P. spirae (Lin & Li, 2010) comb. nov.). Bulb bent dorso-distally at a right angle (Fig. 2C), ratio of bulbal length/width 1.22–1.39, length ratio of embolus/bulb 0.38–0.58, embolus triangular. Receptacle U-shaped (P. cunfengensis) or coiled (P. spirae comb. nov.).

Distribution

China (Guangxi, sites 19–20 in Fig. 30).

Composition

Pinelema cunfengensis and P. spirae comb. nov.

Pinelema cunfengensis Zhao & Li, 2017

Figures 2C, 30

Pinelema cunfengensis Song et al. 2017: 85, figs 1, 2, 7B, 8B, 9B, 10B, 11B, 12B (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Nanning Prefecture, Long’an County, Nanxu Town, Nawan Village, Feng Cave, 23.2098N, 107.5906E, elevation ca. 115 m, 13.V.2015, Z. Chen and Y. Li. leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema cunfengensis resembles P. spirae comb. nov. but can be distinguished by the following: the presence of eyes (cf. Song et al. 2017: fig. 1A) (vs. absence); the different shape of the embolus (Fig. 2C, and cf. Song et al. 2017: fig. 1C, D), the ratio of bulbal length/width is 1.22 (vs. 1.39), and the length ratio of the embolus/bulb is 0.58 (Fig. 2C, and cf. Song et al. 2017: fig. 1C, D) (vs. 0.38); the U-shaped receptacle (cf. Song et al. 2017: fig. 2C) (vs. coiled).

Description

See Song et al. (2017).

Distribution

China (Guangxi, site 19 in Fig. 30), known only from the type locality.

Pinelema spirae (Lin & Li, 2010), comb. nov.

Figures 2C, 30

Telema spirae Lin and Li 2010: 21, figs 12, 13 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Bama County, Poyue Town, Poyue Village, Dawan Cave, 24.3016N, 107.1155E, elevation ca. 438 m, 16.VIII.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema spirae comb. nov. resembles P. cunfengensis but can be distinguished by the following: the absence of eyes (vs. presence); the different shape of the embolus (Fig. 2C), the ratio of bulbal length/width is 1.39 (vs. 1.22), and the length ratio of the embolus/bulb is 0.38 (Fig. 2C) (vs. 0.58); the coiled receptacle (cf. Lin and Li 2010: fig. 13B, C) (vs. U-shaped).

Description

See Lin and Li (2010).

Distribution

China (Guangxi, site 20 in Fig. 30), known only from the type locality.

feilong -group

Figures 3A, 31

Diagnosis

This group resembles the xiezi-group by the short embolus relative to the bulb but can be distinguished by the triangular shape of the embolus (Fig. 3A) (vs. trapezoidal).

Description

Body length 0.98–1.85. Carapace length 0.47–0.76. Tibia I length 0.85–2.20. Six eyes ringed with black (P. bella (Tong & Li, 2008) comb. nov., P. damtaoensis Zhao & Li, 2018, and P. spina comb. nov.) or absent (other species in this group), bulb oval, the junction of the bulb and the cymbium is located ventro-basally, the embolus is triangular, and the length ratio of the embolus/bulb is 0.33–0.49. The receptacle is stick-shaped (P. bella comb. nov., P. claviformis comb. nov., and P. feilong (Chen & Zhu, 2009) comb. nov.), boot-shaped (P. circularis (Tong & Li, 2008) comb. nov.), or globular.

Distribution

China (Guangxi, Yunnan, Hainan) and Vietnam (Vinh Phuc) (Fig. 31).

Composition

Pinelema bella comb. nov., P. circularis comb. nov., P. claviformis comb. nov., P. damtaoensis, P. feilong comb. nov., P. huobaensis Wang & Li, 2016, P. spina comb. nov., P. vesiculata (Lin & Li, 2010) comb. nov., and P. yaosaensis Wang & Li, 2016.

Figure 3. 

Pinelema spp., palp, retrolateral view. A The feilong-group B the pacchanensis-group C the podiensis-group. P. claviformis comb. nov. and P. spina comb. nov. are modified from Tong and Li 2008b and 2008a, respectively.

Pinelema bella (Tong & Li, 2008), comb. nov.

Figures 3A, 31

Telema bella Tong and Li 2008a: 68, figs 1A, 2A–G, 6 (♂♀); Tong 2013: 71, figs 31M, 86A–G (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Hainan Province, Dongfang County, Datian National Natural Reserve, Mihou Cave, 18.9114N, 109.0639E, elevation ca. 293 m, 28.V.2005, Y. Song, X. Han, G. Deng and Y. Tong leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema bella comb. nov. resembles P. spina comb. nov. but can be distinguished by the following: the larger ratio of the bulbal length/width (1.75, Fig. 3A, and cf. Tong and Li 2008a: fig. 2C) (vs. 1.45); the obtuse-angled ventral bend between the embolus and bulb (Fig. 3A, and cf. Tong and Li 2008a: fig. 2B, C) (vs. right-angled), and the blunt tip of the embolus (Fig. 3A, and cf. Tong and Li 2008a: fig. 2B, C) (vs. sharp).

Description

See Tong and Li (2008a).

Distribution

China (Hainan, site 1 in Fig. 31), known only from the type locality.

Pinelema circularis (Tong & Li, 2008), comb. nov.

Figures 3A, 31

Telema circularis Tong and Li 2008b: 363, figs 1A, 2 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guizhou Province, Anshun Prefecture, Guanling County, Shangguan Town, Ganzhishu Cave, 25.8120N, 105.6716E, elevation ca. 987 m, 8.V.2005, Y Tong and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema circularis comb. nov. resembles P. claviformis comb. nov. but can be distinguished by the following: the sharp tip of the embolus (Fig. 3A, and cf. Tong and Li 2008b: fig. 2B, C) (vs. blunt); the boot-shaped receptacle (cf. Tong and Li 2008b: fig. 2D, E) (vs. stick-shaped).

Description

See Tong and Li (2008b).

Distribution

China (Guizhou, site 2 in Fig. 31), known only from the type locality.

Pinelema claviformis (Tong & Li, 2008), comb. nov.

Figures 3A, 31

Telema claviformis Tong and Li 2008b: 364, figs 1B, 3 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guizhou Province, Qianxinan Buyei and Miao Autonomous Prefecture, Xingyi County, Maling Town, Qiuxiang Cave, 25.2000N, 104.8833E, 12.V.2005, Y. Tong and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS). Examined.

Diagnosis

Pinelema claviformis comb. nov. resembles P. circularis comb. nov. but can be distinguished by the following: the blunt tip of the embolus (Fig. 3A) (vs. sharp); the stick-shaped receptacle (cf. Tong and Li 2008b: fig. 3D, E) (vs. boot-shaped).

Description

See Tong and Li (2008b).

Distribution

China (Guizhou, site 3 in Fig. 31), known only from the type locality.

Pinelema damtaoensis Zhao & Li, 2018

Figures 3A, 31

Pinelema damtaoensis Zhao et al. 2018a: 15, figs 1–3 (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Vinh Phuc Province, Dam Tao National Park, leaf litter, 21.4600N, 105.6480E, elevation ca. 999 m, 1.XI.2012, H. Zhao and Z. Chen leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema damtaoensis resembles P. bella comb. nov. but can be distinguished by the following: a black spot and radial stripes on the carapace (cf. Zhao et al. 2018a: fig. 1A) (vs. carapace without pattern); the spiral ridge of the embolus is sclerotized (Fig. 3A, and cf. Zhao et al. 2018a: fig. 1B) (vs. membranous); the receptacle is globular (cf. Zhao et al. 2018a: fig. 3C) (vs. stick-shaped).

Description

See Zhao et al. (2018a).

Distribution

Vietnam (Vinh Phuc, site 4 in Fig. 31), known only from the type locality.

Pinelema feilong (Chen & Zhu, 2009), comb. nov.

Figures 3A, 31

Telema feilong Chen and Zhu 2009: 1707, fig. 2A–K (♂♀).

Type material

Holotype: ♂ (MLR), China, Guizhou Province, Qianxinan Buyei and Miao Autonomous Prefecture, Xingyi County, Feilong Cave, 24.9166N, 104.8833E, elevation ca. 1335 m, 25.V.2004, H. Chen and Y. Zhang leg. Paratypes: 3♂ and 1♀ (MHBU), same data as holotype. Not examined.

Other material examined

1♂ and 2♀ (including molecular voucher, IZCAS), same data as the type locality, 8.III.2011, C. Wang and L. Lin leg.

Diagnosis

Pinelema feilong comb. nov. resembles P. vesiculata comb. nov. but can be distinguished by the following: the smaller ratio of bulbal length/width (1.31, Fig. 3A, and cf. Chen and Zhu 2009: fig. 2C,) (vs. 1.44), the smaller length ratio of the embolus/bulb (0.36, Fig. 3A, and cf. Chen and Zhu 2009: fig. 2C, D) (vs. 0.48); the unmodified stick-shaped receptacle (cf. Chen and Zhu 2009: fig. 2J) (vs. receptacle with a vesicle ventro-distally).

Description

See Chen and Zhu (2009).

Distribution

China (Guizhou, site 5 in Fig. 31), known only from the type locality.

Pinelema huobaensis Wang & Li, 2016

Figures 3A, 31

Pinelema huobaensis Wang and Li 2016: 556, figs 5–8 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Yunnan Province, Wenshan Zhuang and Miao Autonomous Prefecture, Qiubei County, Shuanglongying Town, Puzhehei Village, Huoba Cave, 24.1385N, 104.1126E, elevation ca. 1457 m, 18.VIII.2009, Z. Yao, C. Wang and X. Wang leg. Paratypes: 1♂ and 1♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema huobaensis resembles P. yaosaensis but can be distinguished by the following: the bulb protrudes ventro-subdistally (Fig. 3A, and cf. Wang and Li 2016: fig. 5B, C) (vs. protrudes dorso-distally); the distal part of the receptacle is five times wider than the neck of the receptacle (cf. Wang and Li 2016: fig. 6C) (vs. eight times).

Description

See Wang and Li (2016).

Distribution

China (Yunnan, site 6 in Fig. 31), known only from the type locality.

Pinelema spina (Tong & Li, 2008), comb. nov.

Figures 3A, 31

Telema spina Tong and Li 2008a: 73, figs 1D, 5, 6 (♂); Tong 2013: 73, figs 31P, 89A–C (♂).

Type material

Holotype: ♂ (IZCAS), China, Hainan Province, Wuzhishan County, Wuzhi Mountain, leaf litter, 18.8167N, 109.6500E, Y. Song leg. Examined.

Diagnosis

Pinelema spina comb. nov. resembles P. bella comb. nov. but can be distinguished by the following: the smaller ratio of the bulbal length/width (1.45, Fig. 3A) (vs. 1.75); the obtuse-angled ventral bend between the embolus and bulb (Fig. 3A) (vs. right-angled), the sharp tip of the embolus (Fig. 3A) (vs. blunt).

Description

See Tong and Li (2008a).

Distribution

China (Hainan, site 7 in Fig. 31), known only from the type locality.

Pinelema vesiculata (Lin & Li, 2010), comb. nov.

Figures 3A, 31

Telema vesiculata Lin and Li 2010: 29, figs 18–20 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Yunnan Province, Qujing Prefecture, Luoping County, Luoxiong Town, Pingtian Village, Laobie Cave, 24.8419N, 104.2678E, elevation ca. 1490 m, 20.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema vesiculata comb. nov resembles P. feilong comb. nov. but can be distinguished by the following: the larger ratio of the bulbal length/width (1.44, Fig. 3A, and cf. Lin and Li 2010: fig. 18D) (vs. 1.31), the larger length ratio of the embolus/bulb (Fig. 3A, and 0.48, cf. Lin and Li 2010: fig. 18E) (vs. 0.36); the receptacle with a vesicle ventro-distally (cf. Lin and Li 2010: fig. 20B–D) (vs. unmodified).

Description

See Lin and Li (2010).

Distribution

China (Yunnan, site 8 in Fig. 31), known only from the type locality.

Pinelema yaosaensis Wang & Li, 2016

Figures 3A, 31

Pinelema yaosaensis Wang and Li 2016: 561, figs 13–17 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Yunnan Province, Wenshan Zhuang and Miao Autonomous Prefecture, Xichou County, Niuchangba Township, Mosa Village, Yaosa Cave, 23.5072N, 104.9023E, elevation ca. 1297 m, 6.VIII.2009, Z. Yao, C. Wu and X. Wang leg. Paratypes: 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema yaosaensis resembles P. huobaensis but can be distinguished by the following: the bulb does not protrude ventro-subdistally but protrudes dorso-distally (Fig. 3A, and cf. Wang and Li 2016: fig. 13B, C) (vs. protruding ventro-subdistally but not dorso-distally); the distal part of the receptacle is eight times wider than the neck of the receptacle (cf. Wang and Li 2016: fig. 14C) (vs. five times).

Description

See Wang and Li (2016).

Distribution

China (Yunnan, site 9 in Fig. 31), known only from the type locality.

pacchanensis -group

Figures 3B, 31

Diagnosis

This group resembles the adunca-group by having a long bulb but can be distinguished by the bulb being slightly concave ventrally and protruding dorso-mesially (Fig. 3B) (vs. protrude ventro-basally and concave dorso-subbasally).

Description

Body length 1.33–1.60. Carapace 0.56–0.71 long. Eyes absent. Tibia I 1.33–2.00 long. Ratio of bulbal length/width 1.80–2.11, bulb slightly concave ventrally and protruding dorso-mesially (Fig. 3B), embolus triangular or tube-like. Receptacle swollen distally and globular.

Distribution

China (Guizhou) and Vietnam (Bac Kan) (sites 10–11 in Fig. 31)

Composition

Pinelema daguaiwan sp. nov. and P. pacchanensis Zhao & Li, 2018.

Pinelema daguaiwan Zhao & Li, sp. nov.

Figures 3B, 4, 5, 31

Type material

Holotype: ♂ (IZCAS), China, Guizhou Province, Qianxinan Buyi and Miao Autonomous Prefecture, Wangmo County, Sanglang Town, Xinghe Village, Daguaiwan Cave. 25.2706N, 106.4328E, elevation ca. 886 m, 29.XII.2010, Z. Zha and Z. Chen leg. Paratypes: 1♂ and 5♀ (IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

Pinelema daguaiwan sp. nov. resembles P. pacchanensis by the bulb being concave ventrally but can be distinguished by the following: the bulb is more deeply concave (arrow in Fig. 4C, D) (vs. weaker), the smaller length ratio of the embolus/bulb (0.48, Figs 3B, 4C, D) (vs. 0.71), the triangular embolus (Figs 3B, 4C, D) (vs. tube-shaped); the diameter ratio of the receptacle tip/neck is 7.0 (Fig. 5A) (vs. 2.0).

Figure 4. 

Pinelema daguaiwan sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Description

Male (holotype). Total length 1.60. Carapace 0.71 long, 0.67 wide. Abdomen 0.85 long, 0.76 wide. Carapace light brown (Fig. 4A). Eyes absent (Fig. 4A). Chelicerae, legs, labium, and endites brown. Sternum light brown with sparse setae. Leg measurements: I 6.25 (1.80, 0.26, 2.00, 1.41, 0.78); II 5.46 (1.56, 0.26, 1.74, 1.19, 0.71); III (1.20, 0.25, 1.14, –, –); IV 4.96 (1.52, 0.24, 1.44, 1.13, 0.63). Abdomen pale brown (Fig. 4A).

Palp. Tibia two times longer than patella, cymbium 2.47 times longer than tibia, 1.88 times longer than femur, cymbial apophysis finger shaped (Fig. 4C); bulb concave ventrally, and shaped as in Fig. 4C, D, length ratio of bulb/cymbium about 0.9; embolus triangular and membranous, 1/5 as long as bulb (Fig. 4C, D), spiral ridge brown (Fig. 4B).

Female. Total length 1.72. Carapace 0.66 long, 0.59 wide. Abdomen 1.04 long, 0.87 wide. Coloration as in male (Fig. 5B, C). Leg measurements: I 5.33 (1.56, 0.24, 1.68, 1.15, 0.70); II 4.59 (1.33, 0.24, 1.41, 0.99, 0.62); III 3.37 (1.05, 0.22, 0.94, 0.67, 0.49); IV 4.15 (1.25, 0.22, 1.21, 0.91, 0.56). Abdomen pale brown (Fig. 5B, C). Neck of receptacle membranous, as long as diameter of receptacle (Fig. 5A); receptacle tip globular with several membranous tubes, much wider than neck (Fig. 5A).

Distribution

China (Guizhou, site 10 in Fig. 31), known only from the type locality.

Figure 5. 

Pinelema daguaiwan, sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Pinelema pacchanensis Zhao & Li, 2018

Figures 3B, 31

Pinelema pacchanensis Zhao et al. 2018a: 26, figs 10–12 (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Bac Kan Province, Cho Don District, Pac Chan Cave, 22.3790N, 105.6130E, elevation ca. 225 m, 18.X.2012, H. Zhao and Z. Chen leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema pacchanensis resembles P. daguaiwan sp. nov. but can be distinguished by the following: the ventrally concave part of the bulb is weaker (Fig. 3B, and cf. Zhao et al. 2018a: fig. 11A, B) (vs. stronger), the larger length ratio of the embolus/bulb (0.71, Fig. 3B, and cf. Zhao et al. 2018a: fig. 11A, B) (vs. 0.48), and the triangular embolus (cf. Zhao et al. 2018a: fig. 11 A–D) (vs. tube-shaped); the diameter ratio of the receptacle tip/neck is 2.0 (cf. Zhao et al. 2018a: fig. 12C) (vs. 7.0).

Description

See Zhao et al. (2018a).

Distribution

Vietnam (Bac Kan, site 11 in Fig. 31), known only from the type locality.

podiensis -group

Figures 3C, 31

Diagnosis

This group resembles the feilong-group by the embolus which is short relative to the bulb but can be distinguished by the following: the length ratio of the embolus/bulb ranges from 0.57 to 0.79 (Fig. 3C) (vs. 0.33 to 0.49), and the shape of the embolus is a long isosceles triangle (Fig. 3C) (vs. equilateral triangle).

Description

Body length 1.22–1.75. Carapace 0.48–0.75 long. Tibia I 1.15–1.87 long. Six eyes ringed with black (P. biyunensis (Wang & Li, 2010) comb. nov., P. shiba sp. nov., and P. zonaria (Wang & Li, 2010) comb. nov.), vestigial (P. podiensis Zhao & Li, 2017), or absent (P. bifida (Lin & Li, 2010) comb. nov.). Ratio of bulbal length/width 1.31–1.67, embolus shorter than bulb, length ratio of embolus/bulb 0.57–0.79, embolic shape long isosceles triangle (Fig. 3C). Receptacle J-shaped, slightly swollen distally.

Distribution

China (Guangxi, sites 10–14 in Fig. 31).

Composition

Pinelema bifida comb. nov., P. biyunensis comb. nov., P. podiensis, P. shiba sp. nov., and P. zonaria comb. nov.

Pinelema bifida (Lin & Li, 2010), comb. nov.

Figures 3C, 31

Telema bifida Lin and Li 2010: 5, figs 2, 3 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Dahua County, Qibainong Town, Qiaoxu Village, Qiaoxu Cave, 24.0761N, 107.6706E, elevation ca. 550 m, 9.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

. Pinelema bifida comb. nov. resembles P. zonaria comb. nov. but can be distinguished by the following: the absence of eyes (vs. present); the bulb does not protrude ventro-distally (Fig. 3C, and cf. Lin and Li 2010: fig. 2C, D) (vs. protruding), the ratio of bulbal length/width is larger (1.43, cf. Lin and Li 2010: fig. 2C, D) (vs. 1.31).

Description

See Lin and Li (2010).

Distribution

China (Guangxi, site 12 in Fig. 31), known only from the type locality.

Pinelema biyunensis (Wang & Li, 2010), comb. nov.

Figures 3C, 31

Telema biyunensis Wang and Li 2010b: 9, figs 7–10 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hezhou Prefecture, Zhongshan County, Biyun Cave, 24.357N, 111.1923E, elevation ca. 131 m, 25.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema biyunensis comb. nov. resembles P. zonaria comb. nov. but can be distinguished by the following: the larger ratio of bulbal length/width (1.59, Fig. 3C, and cf. Wang and Li 2010b: fig. 7C, D) (vs. 1.31), the smaller length ratio of the embolus/bulb (0.57, Fig. 3C, and cf. Wang and Li 2010b: fig. 7C, D) (vs. 0.76); the tip of the receptacle is 2.10 times wider than the neck (cf. Wang and Li 2010b: fig. 8C) (vs. 1.10 times).

Description

See Wang and Li (2010b).

Distribution

China (Guangxi, site 13 in Fig. 31), known only from the type locality.

Pinelema podiensis Zhao & Li, 2017

Figures 3C, 31

Pinelema podiensis Song et al. 2017: 88, figs 3, 4, 7C, 8C, 9C, 10C, 11C, 12C (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Debao County, Podi Cave, 23.3919N, 106.6400E, elevation ca. 578 m, 4.VIII.2011, C. Wang leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema podiensis resembles P. shiba sp. nov. but can be distinguished by the following: the weakly concave dorsum of the bulb (Fig. 3C, and cf. Song et al. 2017: fig. 3C, D) (vs. strongly concave), the larger bulbal length/width ratio (1.75, Fig. 3C, and cf. Song et al. 2017: fig. 3C, D) (vs. 1.33); the tip of the receptacle is 2.20 times wider than the neck (cf. Song et al. 2017: fig. 4C) (vs. 4.10 times).

Description

See Song et al. (2017).

Distribution

China (Guangxi, site 14 in Fig. 31), known only from the type locality.

Pinelema shiba Zhao & Li, sp. nov.

Figures 3C, 6, 7, 31

Type material

Holotype: ♂ (IZCAS); China, Guangxi Zhuang Autonomous Region, Chongzuo Prefecture, Daxin County, Hucheng Town, Baoxian Village, Shiba Cave. 22.8133N, 107.1632E, elevation ca. 157 m. 17.XI.2011, C. Wang leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

Pinelema shiba sp. nov. resembles P. podiensis but can be distinguished by the following: the strongly concave dorso-mesial part of the bulb (Fig. 6C, D) (vs. weakly concave), the ratio of the bulbal length/width is 1.33 (Fig. 6C, D) (vs. 1.75); the tip of the receptacle is 4.10 times wider than the neck (Fig. 7C) (vs. 2.20 times).

Description

Male (holotype). Total length 1.22. Carapace 0.51 long, 0.48 wide. Abdomen 0.71 long, 0.63 wide. Carapace brown (Fig. 6A). Six vestigial eyes (Fig. 6A). Chelicerae, legs, labium, and endites light brown (Fig. 6A). Sternum light brown with sparse setae. Leg measurements: I 5.00 (1.47, 0.21, 1.60, 1.06, 0.66); II 4.26 (1.28, 0.21, 1.33, 0.88, 0.56); III 3.05 (0.97, 0.18, 0.89, 0.56, 0.45); IV 3.55 (1.23, 0.17, 1.00, 0.65, 0.50). Abdomen pale yellow with a few long setae.

Palp. Tibia 2.30 times longer than patella, cymbium 1.91 times longer than tibia, 1.90 times longer than femur, cymbial apophysis short, as long as 1/4 width of cymbium base (Fig. 6C); bulb strongly concave dorsally (arrow in Fig. 6C, D); embolus triangular and half as long as cymbium (Fig. 6C, D), spiral ridge brown (Fig. 6B).

Figure 6. 

Pinelema shiba sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Female. Total length 1.39. Carapace 0.54 long, 0.50 wide. Abdomen 0.82 long, 0.73 wide. Six eyes ringed with black (Fig. 7B). Coloration similar to male (Fig. 7B, C). Leg measurements: I 4.94 (1.52, 0.21, 1.56, 1.00, 0.65); II 4.18 (1.31, 0.20, 1.27, 0.85, 0.55); III 3.08 (1.00, 0.19, 0.91, 0.53, 0.45); IV 3.92 (1.25, 0.19, 1.19, 0.78, 0.51). Abdomen brown (Fig. 7B, C). Receptacle U-shaped with several membranous tubes, distally swollen (Fig. 7A).

Figure 7. 

Pinelema shiba sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Distribution

China (Guangxi, site 15 in Fig. 31), known only from the type locality.

Pinelema zonaria (Wang & Li, 2010), comb. nov.

Figures 3C, 31

Telema zonaria Wang and Li 2010b: 33, figs 33–38 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Yizhou County, Xiannv Cave, 24.4887N, 108.5701E, elevation ca. 205 m, 28.VII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema zonaria comb. nov. resembles P. bifida comb. nov. but can be distinguished by the following: eyes are present (vs. absent); the bulb protrudes ventro-distally (Fig. 3C, and cf. Wang and Li 2010b: fig. 33C, D) (vs. not protrude), the ratio of the bulbal length/width is smaller (1.31, Fig. 3C, and cf. Wang and Li 2010b: fig. 33C, D) (vs. 1.43).

Description

See Wang and Li (2010b).

Distribution

China (Guangxi, site 16 in Fig. 31), known only from the type locality.

xiezi -group

Figures 8, 32

Diagnosis

This group resembles the feilong-group by the short embolus relative to the bulb but can be distinguished by the trapezoidal shape of the embolus (Fig. 8) (vs. triangular).

Description

Body length 0.98–2.05. Carapace 0.51–0.82 long. Tibia I 0.81–2.13 long. Six eyes ringed with black, vestigial (P. exiloculata (Lin, Pham & Li, 2009) comb. nov.), or absent, bulb oval, junction of bulb and cymbium located ventro-basally on bulb, embolus trapezoidal, length ratio of embolus/bulb 0.30–0.59. Receptacle J-shaped.

Distribution

China (Guangxi, Guizhou, Hainan) and Vietnam (Hai Phong, Ninh Binh, Phu Tho, Quang Binh) (Sites 1–12 in Fig. 32).

Composition

Pinelema breviseta (Tong & Li, 2008) comb. nov., P. conglobare (Lin & Li, 2010) comb. nov., P. cucphongensis (Lin, Pham & Li, 2009) comb. nov., P. cucurbitina (Wang & Li, 2010) comb. nov., P. dongbei (Wang & Ran, 1998) comb. nov., P. exiloculata comb. nov., P. grandidens (Tong & Li, 2008) comb. nov., P. laensis Zhao & Li, 2018, P. oculata (Tong & Li, 2008) comb. nov., P. pedati (Lin & Li, 2010) comb. nov., P. spinafemora (Lin & Li, 2010) comb. nov., and P. xiezi Zhao & Li, 2018.

Figure 8. 

Pinelema spp., palp, retrolateral view. The xiezi-group. P. spinafemora comb. nov. is modified from Lin and Li 2010.

Pinelema breviseta (Tong & Li, 2008), comb. nov.

Figures 8, 32

Telema breviseta Tong and Li 2008a: 69, figs 1B, 3A–I, 6 (♂♀); Tong 2013: 72, figs 31O, 87A–I (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Hainan Province, Dongfang County, Donghe Town, Yalong Village, Yalong Cave, 18.9770N, 108.8935E, elevation ca. 273 m, Y. Song, X. Han, G. Deng and Y. Tong leg. 1.IV.2005. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema breviseta comb. nov. resembles P. laensis but can be distinguished by the following: the ventro-distal bulb does not protrude (Fig. 8, and cf. Tong and Li 2008a: fig. 3B, C) (vs. protrude), the embolus lacks modification retrolaterally (Fig. 8, and cf. Tong and Li 2008a: fig. 3B, C) (vs. embolus with a vertical groove retrolaterally); the tip of the receptacle is three times wider than the neck (cf. Tong and Li 2008a: fig. 3H) (vs. four times).

Description

See Tong and Li (2008a).

Distribution

China (Hainan, site 1 in Fig. 32), known only from the type locality.

Pinelema conglobare (Lin & Li, 2010), comb. nov.

Figures 8, 32

Telema conglobare Lin and Li 2010: 8, figs 4, 5 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Fengshan County, Fengcheng Town, Songren Village, Xi’an Cave, 24.5657N, 107.0411E, elevation ca. 574 m, 12.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema conglobare comb. nov. resembles P. pedati comb. nov. but can be distinguished by the following: the larger ratio of bulbal length/width (1.77, Fig. 8, and cf. Lin and Li 2010: fig. 4D, E) (vs. 1.60), the ventral bend of the embolus and bulb is right-angled (Fig. 8, and cf. Lin and Li 2010: fig. 4D, E) (vs. acute-angled); the distal part of the receptacle is globular (cf. Lin and Li 2010: fig. 5B–E) (vs. boot-shaped).

Description

See Lin and Li (2010).

Distribution

China (Guangxi, site 2 in Fig. 32), known only from the type locality.

Pinelema cucphongensis (Lin, Pham & Li, 2009), comb. nov.

Figures 8, 32

Telema cucphongensis Lin et al. 2009: 327, figs 5A–E, 6A–I (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Cuc Phuong National Park, Prehistoric Man Cave, 20.2930N, 105.6670E, elevation ca. 256 m, 19.VII.2008, S. Li leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema cucphongensis comb. nov. resembles P. xiezi but can be distinguished by the following: a pair of lateral scutae on abdomen (cf. Lin et al. 2009: fig. 5H, I) (vs. absent); the different shape of the embolus, the larger ratio of the bulbal length/width (1.49, Fig. 8, and cf. Lin et al. 2009: fig. 5A–C) (vs. 1.28), and the smaller length ratio of the embolus/bulb (0.36, Fig. 8, and cf. Lin et al. 2009: fig. 5A–C) (vs. 0.63); the distal part of the receptacle is seven times wider than the neck of the receptacle (cf. Lin et al. 2009: fig. 6F) (vs. five times).

Description

See Lin et al. (2009).

Distribution

. Vietnam (Cuc Phuong National Park, site 3 in Fig. 32), known only from the type locality.

Pinelema cucurbitina (Wang & Li, 2010), comb. nov.

Figures 8, 32

Telema cucurbitina Wang and Li 2010b: 19, figs 16–19 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Guilin Prefecture, Lingui County, Shuixian Cave, 25.2137N, 110.2008E, elevation ca. 161 m, 18.VII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema cucurbitina comb. nov. resembles P. spinafemora comb. nov. but can be distinguished by the following: the bulb does not protrude ventro-distally (Fig. 8, and cf. Wang and Li 2010b: fig. 16C, D) (vs. protrude), the ventral bend between the embolus and the bulb is acute-angled (Fig. 8, and cf. Wang and Li 2010b: fig. 16C, D) (vs. right-angled).

Description

See Wang and Li (2010b).

Distribution

China (Guangxi, site 4 in Fig. 32).

Pinelema dongbei (Wang & Ran, 1998), comb. nov.

Figures 8, 32

Telema dongbei Wang and Ran 1998: 94, figs 1–5 (♂♀); Song et al. 1999: 51, fig. 21R–U (♂♀).

Type material

Holotype: ♂ (AMNH), China, Guizhou Province, Qiannan Buyei and Miao Autonomous Prefecture, Libo County, Yuping Town, Dongbei Cave, 25.4804N, 107.8959E, elevation ca. 812 m, 13.IX.1996, J. Ran leg. Paratype: 1♀ (AMNH), same data as holotype. Not examined.

Other material examined

1♂ and 2♀ (including molecular voucher, IZCAS), from the type locality, 18.III.2011, C. Wang and L. Lin leg.

Diagnosis

Pinelema dongbei comb. nov. resembles P. exiloculata comb. nov. and P. oculata comb. nov. but can be distinguished by the following: eyes are absent (vs. vestigial); the embolus protrudes dorsally (Fig. 8, and cf. Wang and Ran 1998: figs 1, 2) (vs. not protruding); the receptacle is slightly swollen distally (cf. Wang and Ran 1998: fig. 5) (vs. distinctively swollen).

Description

See Wang and Ran (1998).

Distribution

China (Guizhou, site 5 in Fig. 32), known only from the type locality.

Pinelema exiloculata (Lin, Pham & Li, 2009), comb. nov.

Figures 8, 32

Telema exiloculata Lin et al. 2009: 332, figs 7A–F, 8A–G (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Hai Phong Province, Cat Ba National Park, Trung Trang Cave, 20.8000N, 106.9833E, elevation ca. 256 m, 16.VII.2008, S. Li leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema exiloculata comb. nov. resembles P. dongbei comb. nov. but can be distinguished by the following: vestigial eyes (vs. absent); the embolus does not protrude dorsally (Fig. 8, and cf. Lin et al. 2009: fig. 7B) (vs. protruding); the receptacle is distinctively swollen distally (cf. Lin et al. 2009: fig. 8D–G) (vs. swollen slightly).

Description

See Lin et al. (2009).

Distribution

Vietnam (Cat Ba National Park, site 6 in Fig. 32), known only from the type locality.

Pinelema grandidens (Tong & Li, 2008), comb. nov.

Figures 8, 32

Telema grandidens Tong and Li 2008b: 366, figs 1C, 4 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guizhou Province, Qiannan Buyei and Miao Autonomous Prefecture, Dushan County, Xiasi Town, Xinhe Village, Bayoudadong Cave, 25.4457N, 107.4316E, elevation ca. 929 m, 21.V.2005, Y. Tong and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema grandidens comb. nov. resembles P. oculata comb. nov. but can be distinguished by the following: the eyes are absent (vs. present); the dorsal bend between the embolus and bulb is ca. 100° (Fig. 8, and cf. Tong and Li 2008b: fig. 4C) (vs. ca. 180°); the distal part of the receptacle is two times wider than the receptacle neck (cf. Tong and Li 2008b: fig. 4E) (vs. three times).

Description

See Tong and Li (2008b).

Distribution

China (Guizhou, site 7 in Fig. 32), known only from the type locality.

Pinelema laensis Zhao & Li, 2018

Figures 8, 32

Pinelema laensis Zhao et al. 2018a: 19, figs 7–9 (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Phu Tho Province, Tan Son District, Xuan Dai, Xuan Son National Park, La Cave, 21.1380N, 104.9390E, elevation ca. 424 m, 27.X.2012, H. Zhao and Z. Chen leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema laensis resembles P. breviseta comb. nov. but can be distinguished by the following: the bulb does not protrude ventro-distally (Fig. 8, and cf. Zhao et al. 2018a: fig. 7C, D) (vs. protrude), the embolus with a retrolateral vertical groove (Fig. 8, and cf. Zhao et al. 2018a: fig. 8B) (vs. embolus without any structure retrolaterally); the tip of receptacle is four times wider than the neck (cf. Zhao et al. 2018a: fig. 9C) (vs. three times).

Description

See Zhao et al. (2018a).

Distribution

Vietnam (Phu Tho, site 8 in Fig. 32), known only from the type locality.

Pinelema oculata (Tong & Li, 2008), comb. nov.

Figures 8, 32

Telema oculata Tong and Li 2008b: 369, figs 1D, 5 (♀).

Type material

Paratypes: 11 ♀ (IZCAS), China, Guizhou Province, Qiannan Buyei and Miao Autonomous Prefecture, Dushan County, Xiasi Town, Guojiafen Cave, 25.4833N, 107.4500E, 24.V.2005, Y. Tong and Y. Lin leg. Examined.

Other material examined

1♂ and 2♀ (including one molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema oculata comb. nov. resembles P. grandidens comb. nov. but can be distinguished by the following: the eyes are present (vs. absent); the dorsal bend between the embolus and bulb is ca. 180° (Fig. 8) (vs. ca. 100°); the distal part of the receptacle is three times wider than the receptacle neck (cf. Tong and Li 2008b: fig. 5C) (vs. two times).

Description

See Tong and Li (2008b).

Distribution

China (Guizhou, site 9 in Fig. 32), known only from the type locality.

Pinelema pedati (Lin & Li, 2010), comb. nov.

Figures 8, 32

Telema pedati Lin and Li 2010: 15, figs 8, 9 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Hechi Prefecture, Nandan County, Chengguan Town, En Village, Encun Cave, 25.0693N, 107.6033E, elevation ca. 605 m, 3.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema pedati comb. nov. resembles P. conglobare comb. nov. but can be distinguished by the following: the smaller ratio of the bulbal length/width (1.60, Fig. 8, and cf. Lin and Li 2010: fig. 8A, B) (vs. 1.77), the ventral bend of the embolus and bulb is acute-angled (Fig. 8, and cf. Lin and Li 2010: fig. 8A, B) (vs. right-angled); the distal part of the receptacle is boot-shaped (cf. Lin and Li 2010: fig. 9B–D) (vs. globular).

Description

See Lin and Li (2010).

Distribution

China (Guangxi, site 10 in Fig. 32), known only from the type locality.

Pinelema spinafemora (Lin & Li, 2010), comb. nov.

Figures 8, 32

Telema spinafemora Lin and Li 2010: 18, figs 10, 11 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Baise Prefecture, Lingyun County, Sicheng Town, Shuiyuan Cave, 24.3464N, 106.5579E, elevation ca. 573 m, 14.III.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema spinafemora comb. nov. resembles P. cucurbitina comb. nov. but can be distinguished by the following: the bulb protrudes ventro-distally (Fig. 8) (vs. not protrude), the ventral bend between the embolus and bulb is right-angled (Fig. 8) (vs. acute-angled).

Description

See Lin and Li (2010).

Distribution

China (Guangxi, site 11 in Fig. 32), known only from the type locality.

Pinelema xiezi Zhao & Li, 2018

Figures 8, 32

Pinelema xiezi Zhao et al. 2018a: 34, figs 16–18 (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Quang Binh Province, Phong Nha-Ke Bang National Park, Tien Son Cave, 17.5800N, 106.2820E, elevation ca. 102 m, 17.V.2016, Z. Chen and Q. Zhao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema xiezi resembles P. cucphongensis comb. nov. but can be distinguished by the following: the lateral scuta on the abdomen is absent (cf. Zhao et al. 2018a: figs 16A, 17A) (vs. present); the different shape of the embolus, the smaller ratio of the bulbal length/width (1.28, Fig. 8, and cf. Zhao et al. 2018a: fig. 16C, D) (vs. 1.49), the larger length ratio of the embolus/bulb (0.63, Fig. 8, and cf. Zhao et al. 2018a: fig. 16C, D) (vs. 0.36); the distal part of the receptacle is five times wider than the neck of the receptacle (cf. Zhao et al. 2018a: fig. 18C) (vs. seven times).

Description

See Zhao et al. (2018a).

Distribution

Vietnam (Quang Binh, site 12 in Fig. 32), known only from the type locality.

Species group uncertain

Figures 9, 32

Remarks.

Six species: Pinelema dengi (Tong & Li, 2008) comb. nov., P. mikrosphaira (Wang & Li, 2010) comb. nov., P. nuocnutensis Zhao & Li, 2018, P. spirulata Zhao & Li, 2018, P. tham sp. nov., and P. zhenzhuang Zhao & Li, 2018 are not grouped and seem to represent groups of their own.

Figure 9. 

Pinelema spp., palp, retrolateral view. Six species not attached to a species group.

Pinelema dengi (Tong & Li, 2008), comb. nov.

Figures 9, 32

Telema dengi Tong and Li 2008a: 69, figs 1C, 4A–H, 6 (♂♀); Tong 2013: 73, figs 31N, 88A–H (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Hainan Province, Sanya Prefecture, Lizhigou Town, Luobi Cave, 18.3318N, 109.5491E, elevation ca. 46 m, 10.IV.2005, X. Han, Y. Song, G. Deng and Y. Tong leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema dengi comb. nov. resembles P. spirulata but can be distinguished by the following: the cylindrical embolus (Fig. 9, and cf. Tong and Li 2008a: fig. 4B, C) (vs. twisted); the distal part of the receptacle is seven times wider than the receptacle neck (cf. Tong and Li 2008a: fig. 4D, E) (vs. five times).

Description

See Tong and Li (2008a).

Distribution

China (Hainan, site 13 in Fig. 32), known only from the type locality.

Pinelema mikrosphaira (Wang & Li, 2010), comb. nov.

Figures 9, 32

Telema mikrosphaira Wang and Li 2010b: 24, figs 20–23 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guangxi Zhuang Autonomous Region, Chongzuo Prefecture, Pinxiang County, Yinglong Cave, 22.1426N, 106.7128E, elevation ca. 200 m, 9.VIII.2009, C. Wang and Z. Yao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema mikrosphaira comb. nov. resembles P. nuocnutensis but can be easily distinguished by the following: the eyes are encircled by black rings (vs. vestigial), the width ratio of the bulb/palpal tibia is 4.0 (Fig. 9) (vs. 2.0); the embolus is shaped like an equilateral-triangle (Fig. 9) (vs. beak-shaped); the distal part of the receptacle is swollen and globular (cf. Wang and Li 2010b: fig. 21C, D) (vs. not swollen).

Description

See Wang and Li (2010b).

Distribution

China (Guangxi, site 14 in Fig. 32), known only from the type locality.

Pinelema nuocnutensis Zhao & Li, 2018

Figures 9, 32

Pinelema nuocnutensis Zhao et al. 2018a: 19, figs 4–6 (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Quang Binh Province, Phong Nha-Ke Bang National Park, Nuoc Nut Cave, 17.4940N, 106.2940E, elevation ca. 143 m, 25.V.2016, Z. Chen and Q. Zhao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema nuocnutensis resembles P. mikrosphaira comb. nov. but can be distinguished by the following: the eyes are vestigial (vs. present), the width ratio of the bulb/palpal tibia is 2.0 (Fig. 9) (vs. 4.0); the beak-shaped embolus (Fig. 9) (vs. equilateral triangle); the distal part of the receptacle is not swollen (cf. Zhao et al. 2018a: fig. 6C) (vs. swollen).

Description

See Zhao et al. (2018a).

Distribution

Vietnam (Quang Binh, site 15 in Fig. 32), known only from the type locality.

Pinelema spirulata Zhao & Li, 2018

Figures 9, 32

Pinelema spirulata Zhao et al. 2018a: 30, figs 13–15 (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Phu Tho Province, Xuan Son National Park, Lap Cave, 21.1400N, 104.9430E, elevation ca. 403 m, 2.X.2012, H. Zhao and Z. Chen leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema spirulata resembles P. dengi comb. nov. but can be distinguished by the following: the embolus is twisted (Fig. 9) (vs. cylindrical); the distal part of the receptacle is five times wider than the receptacle neck (cf. Zhao et al. 2018a: fig. 15C) (vs. seven times).

Description

See Zhao et al. (2018a).

Distribution

Vietnam (Phu Tho, site 16 in Fig. 32), known only from the type locality.

Pinelema tham Zhao & Li, sp. nov.

Figures 9, 10, 11, 32

Type material

Holotype: ♂ (IZCAS), Laos, Vien Tiane Province, Vang Vieng District, 1.54 km south of Vieng keo Village, Tham Cave, 18.9092N, 102.4421E, elevation ca. 270 m, XI.2012, S. Li and Z. Yao leg. Paratypes: 4♂ and 4♀ (IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

Pinelema tham sp. nov. resembles P. zhenzhuang but can be easily distinguished by the following: the embolus is hawk-beak-shaped (Figs 9, 10B) (vs. needle-shaped), the tip of the embolus is directed ventro-prolaterally (Figs 9, 10B) (vs. ventrally). The terminal part of the receptacle is 3.50 times wider than the neck (Fig. 11A) (vs. six times).

Figure 10. 

Pinelema tham sp. nov., male holotype. A Habitus, dorsal view B palp, ventral view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.1 mm (B–D).

Description

Male (holotype). Total length 1.23. Carapace 0.48 long, 0.47 wide. Abdomen 0.71 long, 0.59 wide. Carapace pale brown (Fig. 10A). Six vestigial eyes (Fig. 10A). Chelicerae, labium, and endites light brown, legs milky white (Fig. 10A). Sternum brown with sparse setae. Leg measurements: I 3.77 (1.08, 0.18, 1.19, 0.76, 0.56); II 3.20 (0.95, 0.19, 0.95, 0.60, 0.51); III 2.41 (0.75, 0.17, 0.63, 0.45, 0.41); IV 2.84 (0.89, 0.16, 0.79, 0.55, 0.45). Abdomen grey (Fig. 10A).

Palp. Tibia 1.96 times longer than patella, cymbium 1.67 times longer than tibia, 1.45 times longer than femur, cymbial apophysis short, as long as half width of cymbium base (Fig. 10C); bulb bean shaped (Fig. 10C, D); spiral ridge brown; embolus short relative to bulb, tip well sclerotized, hawk-beak-shaped, directed ventro-prolaterally (Fig. 10B–D).

Female. Total length 1.24. Carapace 0.44 long, 0.43 wide. Abdomen 0.74 long, 0.60 wide. Coloration as in male (Fig. 11B, C). Leg measurements: I 3.31 (0.93, 0.18, 1.03, 0.63, 0.55); II 2.79 (0.82, 0.17, 0.82, 0.51, 0.47); III 2.18 (0.67, 0.16, 0.57, 0.40, 0.38); IV 2.55 (0.79, 0.16, 0.71, 0.49, 0.40). Abdomen pale brown. Receptacle with two membranous tubes, swollen distally (Fig. 11A), tip of receptacle four times wider than the neck (Fig. 11A).

Figure 11. 

Pinelema tham sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Distribution

Laos (Vien Tiane, site 17 in Fig. 32), known only from the type locality.

Pinelema zhenzhuang Zhao & Li, 2018

Figures 9, 32

Pinelema zhenzhuang Zhao et al. 2018a: 35, figs 19–22 (♂♀).

Type material

Holotype: ♂ (IZCAS), Vietnam, Quang Binh Province, Phong Nha-Ke Bang National Park, Tien Duong Cave, 17.5190N, 106.2230E, elevation ca. 133 m, 18.V.2016, Z. Chen and Q. Zhao leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

Pinelema zhenzhuang resembles P. tham sp. nov. but can be distinguished by the following: the embolus is needle shaped (cf. Zhao et al. 2018a: fig. 19C, D) (vs. hawk beak shaped), the tip of the embolus is directed ventrally (Fig. 9) (vs. ventro-prolaterally). The terminal part of the receptacle is six times wider than the neck (cf. Zhao et al. 2018a: fig. 21C) (vs. 3.50 times).

Description

See Zhao et al. (2018a).

Distribution

Vietnam (Quang Binh, site 18 in Fig. 32), known only from the type locality.

Apneumonella Fage, 1921

Apneumonella Fage, 1921: 620; Brignoli 1978: 113; Song et al. 2017: 14.

Type species

Apneumonella oculata Fage, 1921 from Tanzania, Africa; Simon and Fage 1922: 528, fig. II.

Comments

This genus currently includes three species: two from Africa and one from Sumatra. The Sumatran species, A. jacobsoni Brignoli, 1977, is known by the female only and is most likely misplaced in this genus (see comments below).

Apneumonella jacobsoni Brignoli, 1977

Figure 1F

Apneumonella jacobsoni Brignoli, 1977: 221, figs 1–6 (♀); Lehtinen 1986: 155, fig. 6 (♀).

Type material

Holotype: ♀ (RMNH), Indonesia, Sumatra, West Sumatra Province, Fort de Kock, 0.2484S, 100.4832E, elevation ca. 920 m, 1926, E. Jacobson leg. Not examined.

Other material examined

3♀ (including molecular voucher, IZCAS) from the type locality: I. 2014, H. Zhao leg.

Distribution

Indonesia (Sumatra, West Sumatra, site 1 in Fig. 33).

Comments

The placement of this species in Apneumonella is doubtful because the males of both A. oculata Fage, 1921 (the type species of Apneumonella) and A. jacobsoni are unknown, and females of the above two species provide little information regarding their generic belonging. To test the relationship of A. jacobsoni to A. oculata, molecular data of A. oculata is necessary.

Mekonglema Zhao & Li, gen. nov.

Type species

Mekonglema bailang sp. nov. from Yunnan, China.

Etymology

The generic name is a combination of “Mekong” referring to the Mekong-Lancang River which encompasses the distributional range of the genus, and “-lema”, a convention used because it is part of the genus Telema, which was the first genus described in Telemidae. The gender is feminine.

Diagnosis

Mekonglema gen. nov. resembles Pinelema but can be distinguished by the following: Males of Mekonglema gen. nov. have a bulbal apophysis (M. bailang sp. nov., M. xinpingi comb. nov., and M. yan sp. nov.) (Fig. 12D) (vs. absent), or the tip of the embolus is directed dorsally (M. kaorao sp. nov. and M. walayaku sp. nov.) (Figs 14C, D, 16C, D) (vs. tip of embolus directed ventrally). Females of Mekonglema gen. nov. can be distinguished from those of Pinelema by the receptacle lacking tubes inside (Fig. 13A) (vs. receptacle with several membranous tubes inside).

Figure 12. 

Mekonglema bailang sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Description

Total length: 1.06–1.50 (male), 1.15–1.70 (female). Carapace without pattern in troglobitic species or with radial striae in rainforest species, i.e. M. xinpingi comb. nov. Sternum with sparse setae, milky white or light brown in troglobitic species, or nearly black in M. xinpingi comb. nov. Eyes ringed with black, vestigial, or absent; Leg formula: 1-2-4-3, tibial glands belt-shaped (Fig. 1C). Abdomen of males with three different types of scutae (except in M. yan sp. nov.): the first scuta connects to pedicel, dorsal (arrow 1 on Fig. 12A); the second scuta is posterior to the first one, dorsal (arrow 2 on Fig. 12A); and the third scutae are paired, lateral (arrows 3 in Fig. 12A). Abdomen of females without scuta. Male palp: cymbial apophysis cone shaped and located medially (Fig. 12C); bulb ellipsoid or nearly ellipsoid, bulbal apophysis present (except M. kaorao sp. nov. and M. walayaku sp. nov.); embolus short relative to cymbium, sclerotized, directed outward from the cymbium in M. bailang sp. nov., M. xinpingi comb. nov., and M. yan sp. nov.; embolus long relative to cymbium, unsclerotized, directed toward cymbium in M. kaorao sp. nov. and M. walayaku sp. nov. Endogyne: composed of single tube-like or globular receptacle with short neck, membranous tubes absent.

Figure 13. 

Mekonglema bailang sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Figure 14. 

Mekonglema kaorao sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Composition

Mekonglema bailang sp. nov., M. kaorao sp. nov., M. walayaku sp. nov., M. xinpingi comb. nov., and M. yan sp. nov. (Fig. 34).

Distribution

China (Yunnan) and Laos (Luang Prabang) (sites 2–6 in Fig. 33).

Mekonglema bailang Zhao & Li, sp. nov.

Figures 12, 13, 33

Type material

Holotype: ♂ (IZCAS), China, Yunnan Province, Baoshan Prefecture, Shidian County, Bailang Town, Xianren Cave, 24.6536N, 99.2645E, elevation ca. 1987 m, 29.VII.2010, C. Wang and Q. Zhao leg. Paratypes: 5♂ and 1♀ (IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

This species resembles M. yan sp. nov. but can be distinguished by the following characters: abdominal scutae present in the male (vs. absent); the embolus is sclerotized (Fig. 12B–D) (vs. unsclerotized), the bulb length/width ratio is ca. 1.2 (Fig. 12C, D) (vs. ca. 1.7); the receptacle is short and almost straight (Fig. 13A) (vs. long and U-shaped). This species also resembles M. xinpingi comb. nov. but can be differentiated by the following: the absence of eyes (vs. presence), the nearly ellipsoid shape of the bulb (Fig. 12C, D) (vs. droplet shaped), the fin-like embolus (Fig. 12C) (vs. cone shaped), and the receptacle is short and not swollen distally (vs. receptacle long and swollen distally).

Description

Male (holotype). Total length 1.50. Carapace 0.65 long, 0.61 wide. Abdomen 0.85 long, 0.79 wide. Carapace brown (Fig. 12A). Four vestigial eyes (Fig. 12A). Chelicerae, legs, labium, and endites light brown. Sternum bright brown with sparse setae (Fig. 12A). Leg measurements: I 5.05 (1.45, 0.23, 1.60, 1.13, 0.64); II 4.54 (1.36, 0.23, 1.41, 0.95, 0.59); III 3.20 (0.98, 0.21, 0.92, 0.62, 0.47); IV 4.17 (1.30, 0.20, 1.24, 0.88, 0.55). Abdomen pale brown (Fig. 12A).

Palp. Tibia 2.12 times longer than patella, cymbium 1.74 times longer than tibia, cymbial apophysis length 2/3 as wide as cymbial base (Fig. 12C); bulb shaped as shown in Fig. 12C, D; bulbal apophysis sclerotized and semi-circular with claw-like tip, embolus sclerotized (Fig. 12B–D).

Female. Total length 1.48. Carapace 0.65 long, 0.63 wide. Abdomen 0.81 long, 0.80 wide. Coloration as in male (Fig. 13B, C). Leg measurements: I 4.78 (1.47, 0.24, 1.45, 1.00, 0.62); II 4.35 (1.34, 0.22, 1.33, 0.88, 0.58); III 3.04 (0.96, 0.22, 0.87, 0.54, 0.45); IV 4.05 (1.28, 0.19, 1.18, 0.85, 0.55). Abdomen pale grey. Insemination entrance membranous, two times thinner than receptacle (Fig. 13A); receptacle tube-like and straight (Fig. 13A).

Distribution

China (Yunnan, Baoshan, site 2 in Fig. 33), known only from the type locality.

Mekonglema kaorao Zhao & Li, sp. nov.

Figures 14, 15, 33

Type material

Holotype: ♂ (SMF), Laos, Luang Prabang Province, Vieng Phoukha town, Ban Nam Eng, Kao Rao Cave, 20.7251N, 101.1541E, elevation ca. 729 m, P. Jäger leg. Paratype: 1♀ (SMF), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

This species resembles M. walayaku sp. nov. but can be distinguished by the following characters: the bulb is nearly globular (Fig. 14C, D) (vs. ellipsoidal), the tip of the embolus is not sclerotized (Fig. 14B) (vs. well sclerotized). The tip of receptacle is 1.50 times wider than the neck (Fig. 15A) (vs. three times).

Figure 15. 

Mekonglema kaorao sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Description

Male (holotype). Total length 1.13. Carapace 0.49 long, 0.44 wide. Abdomen 0.58 long, 0.45 wide. Carapace brown (Fig. 14A). Six vestigial eyes (Fig. 14A). Chelicerae, legs, labium, and endites light brown. Sternum bright brown with sparse setae. Leg measurements: I 3.06 (0.80, 0.18, 0.98, 0.62, 0.48); II 2.84 (0.85, 0.17, 0.84, 0.53, 0.45); III 2.06 (0.63, 0.15, 0.56, 0.37, 0.35); IV 2.73 (0.81, 0.15, 0.81, 0.56, 0.40). Abdomen light brown (Fig. 14A).

Palp. Tibia 2.71 times longer than patella, cymbium 1.65 times longer than tibia, length of cymbial apophysis as wide as cymbial base (Fig. 14C); bulb nearly globular (Fig. 14C, D); embolus 2/3 as long as cymbium, directed dorsally (Fig. 14C, D).

Female. Total length 1.18. Carapace 0.46 long, 0.45 wide. Abdomen 0.65 long, 0.52 wide. Eyes vestigial (Fig. 15B). Coloration as in male (Fig. 15B, C). Leg measurements: I 2.95 (0.88, 0.18, 0.88, 0.56, 0.45); II 2.62 (0.79, 0.17, 0.75, 0.50, 0.41); III 1.92 (0.59, 0.14, 0.53, 0.34, 0.32); IV 2.62 (0.79, 0.15, 0.75, 0.53, 0.40). Abdomen grey. Receptacle membranous, without tubes inside, neck 1.50 times thinner than tip (Fig. 15A).

Distribution

Laos (Luang Prabang, site 3 in Fig. 33), known only from the type locality.

Mekonglema walayaku Zhao & Li, sp. nov.

Figures 16, 17, 33

Type material

Holotype: ♂ (IZCAS), China, Yunnan Province, Nujiang Lisu Autonomous Prefecture, Lushui County, Daxingdi Township, Walayaku Cave. 26.1215N, 98.8581E, elevation ca. 895 m, 24.VI.2016, Y. Li and J. Liu leg. Paratypes: 2♂ and 4♀ (IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

This species resembles M. kaorao sp. nov. but can be distinguished by the following characters: the bulb is ellipsoidal (Fig. 16C, D) (vs. nearly globular), the tip of the embolus is well sclerotized (arrow on Fig. 16B) (vs. unsclerotized); the neck of the receptacle is ca. three times thinner (vs. 1.50 times thinner) than the distal part of the receptacle (Fig. 17A).

Figure 16. 

Mekonglema walayaku sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Description

Male (holotype). Total length 1.06. Carapace 0.49 long, 0.43 wide. Abdomen 0.56 long, 0.55 wide. Carapace light brown (Fig. 16A). Eyes vestigial (Fig. 16A). Chelicerae, legs, labium, and endites light yellow. Sternum bright brown with sparse setae. Leg measurements: I 3.65 (1.03, 0.18, 1.13, 0.78, 0.53); II 3.20 (0.96, 0.17, 0.95, 0.65, 0.47); III 2.26 (0.67, 0.16, 0.61, 0.44, 0.38); IV 2.86 (0.85, 0.16, 0.83, 0.59, 0.43). Abdomen dark blue (Fig. 16A).

Palp: Tibia 1.88 times longer than patella, cymbium 1.34 times longer than tibia, cymbial apophysis length as wide as cymbial base (Fig. 16C); bulb ellipsoidal as in Fig. 16C, D; embolus 5/7 as long as cymbium, directed dorsally (Fig. 16C, D).

Female. Total length 1.15. Carapace 0.50 long, 0.47 wide. Abdomen 0.63 long, 0.58 wide. Coloration as in male (Fig. 17B, C). Leg measurements: I 3.36 (0.97, 0.19, 1.03, 0.69, 0.48); II 3.00 (0.89, 0.19, 0.88, 0.59, 0.45); III 2.15 (0.65, 0.17, 0.59, 0.40, 0.34); IV 2.80 (0.85, 0.18, 0.79, 0.56, 0.42). Abdomen blue. Receptacle membranous, neck much longer than tip, and tip three times wider than neck (Fig. 17A).

Distribution

China (Yunnan, Nujiang, site 4 in Fig. 33), known only from the type locality.

Figure 17. 

Mekonglema walayaku sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Mekonglema xinpingi (Lin & Li, 2008), comb. nov.

Figure 1C

Seychellia xinpingi Lin and Li 2008: 650, figs 1–9 (♂♀).

Type material

Paratypes: 1♂ and 1♀ (IZCAS), China, Yunnan Province, Xishuangbanna Autonomous Prefecture, Mengla County, Menglun Town, rainforest, leaf litter, 21.9001N, 101.1833E, V–VII.2005, G. Zheng leg. Examined.

Other material examined

3♀ (including one molecular voucher, IZCAS), same data as paratypes.

Diagnosis

Mekonglema xinpingi comb. nov. resembles M. bailang sp. nov. but can be distinguished by the following characters: eyes are present (vs. absent); carapace with distinct radial striae (vs. no pattern); bulb is droplet-shaped (cf. Lin and Li 2008: fig. 2) (vs. nearly ellipsoidal).

Description

See Lin and Li (2008).

Distribution

China (Yunnan, Xishuangbanna, site 5 in Fig. 33), known only from the type locality.

Comments

This species is transferred to Mekonglema gen. nov. because it shares a similar shape of the copulatory organs with M. bailang sp. nov., the type species of the genus. This placement is also supported by molecular phylogenetic analyses (Fig. 34).

Mekonglema yan Zhao & Li, sp. nov.

Figures 18, 19, 33

Type material

Holotype: ♂ (IZCAS), China, Yunnan Province, Baoshan Prefecture, Tengchong County, Diantan Town, Lianzu Village, Yan Cave, 25.5501N, 98.4452E, elevation ca. 1867 m, 27.XI.2013, Y. Li and J. Liu leg. Paratypes: 2♂ and 4♀ (IZCAS), same data as holotype.

Etymology

The species refers to the type locality; noun in apposition.

Diagnosis

Mekonglema yan sp. nov. resembles M. bailang sp. nov. but can be distinguished by the following characters: an abdominal scuta is absent in the male (vs. present); the tip of the embolus is unsclerotized (Fig. 18B–D) (vs. sclerotized), the bulbal length/width ratio is approximately 1.7 (Fig. 18C, D) (vs. 1.2); the receptacle is long and U-shaped (Fig. 19A) (vs. short and nearly straight).

Figure 18. 

Mekonglema yan sp. nov., male holotype A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Description

Male (holotype). Total length 1.48. Carapace 0.69 long, 0.57 wide. Abdomen 0.77 long, 0.63 wide. Carapace brown (Fig. 18A). Four vestigial eyes (Fig. 18A). Chelicerae, labium, and endites brown, legs yellow. Sternum bright brown with sparse setae. Leg measurements: I 4.86 (1.44, 0.23, 1.52, 1.00, 0.67); II 4.33 (1.34, 0.23, 1.25, 0.91, 0.60); III 3.20 (1.00, 0.21, 0.88, 0.61, 0.50); IV 4.04 (1.25, 0.20, 1.16, 0.87, 0.56). Abdomen light brown (Fig. 18A).

Palp: Tibia 2.05 times longer than patella, cymbium 1.73 times longer than tibia, length of cymbial apophysis as wide as cymbial base (Fig. 18C); bulb shaped as in Fig. 18C, D; bulbal apophysis sclerotized and looped about 270° (Fig. 18B–D); embolus membranous and finger-like (Fig. 18C, D).

Female: Total length 1.70. Carapace 0.71 long, 0.61 wide. Abdomen 0.97 long, 0.90 wide. Coloration lighter than in male (Fig. 19B, C). Leg measurements: I 4.72 (1.44, 0.23, 1.45, 0.95, 0.65); II 4.24 (1.30, 0.23, 1.25, 0.85, 0.61); III 3.49 (1.33, 0.20, 0.89, 0.59, 0.48); IV 4.05 (1.25, 0.20, 1.19, 0.85, 0.56). Abdomen grey. Receptacle membranous, U-shaped, neck as wide as distal part (Fig. 19A).

Distribution

China (Yunnan, Baoshan, site 6 in Fig. 33), known only from the type locality.

Figure 19. 

Mekonglema yan sp. nov., female paratype A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Siamlema Zhao & Li, gen. nov.

Type species

Siamlema changhai sp. nov. from Trang Province, Thailand.

Etymology

The generic name is derived from “Siam”, referring to the old name of Thailand, and “-lema” is a convention from Telema, the type genus of the family. Feminine in gender.

Diagnosis

Siamlema gen. nov. can be distinguished from Telema by the following: belt-shaped tibial glands (Fig. 1E) (vs. plate-shaped); males can be distinguished from those of Telema by having a cymbial apophysis (Fig. 20C) (vs. absent), a distinct dorsal spine present on the palpal tibia (Fig. 20C, D) (vs. absent), and the palpal femur is longer than the cymbium (Fig. 20C, D) (vs. shorter). Receptacle without tubes inside (Fig. 21A) (vs. receptacle with membranous tubes).

Figure 20. 

Siamlema changhai sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Description

Total length: 0.92–1.11 (male), 1.04–1.20 (female). Carapace 0.41–0.48 long. Sternum with sparse setae. Eyes normally developed, vestigial, or absent. Leg formula: 1-2-4-3, tibia I 0.97–1.05 long, leg glands belt shaped (arrows on Fig. 1D). Male palp: cymbium, tibia, patella, and femur robust relative to bulb, femur longer than tibia and cymbium, dorsal spine present on distal part of palpal tibia, distinct cymbial apophysis located prolaterally on mesial part; bulb droplet-shaped or nearly globular; embolus sclerotized. Endogyne: receptacle tube-like or globular, without tubes inside (Fig. 21A).

Distribution

Thailand (Trang, Yala, sites 7, 8 in Fig. 33).

Composition

Siamlema changhai sp. nov. and S. suea sp. nov. (Fig. 34).

Siamlema changhai Zhao & Li, sp. nov.

Figures 1D, 20, 21, 33

Type material

Holotype: ♂ (IZCAS), Thailand, Trang Province, Noyong District, Chang Hai Cave, 7.5893N, 99.6688E, elevation ca. 32 m, XI.2015, Z. Chen, G. Zhou, and Q. Zhao leg. Paratypes: 2♂ and 4♀ (IZCAS), same data as holotype.

Other material examined

1♀ (molecular voucher, IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

This species resembles S. suea sp. nov. but can be distinguished by the following: the absence of eyes (Figs 20A, 21B) (vs. presence), the presence of a bulbal apophysis (Fig. 20B–D) (vs. absence), the presence of a dorsal spine on the palpal femur distally (arrows on Fig. 20C, D) (vs. spine absent), and the absence of a spine on the palpal tibia retrolaterally (Fig. 20D) (vs. spine present). Receptacle is tube-shaped (Fig. 21A) (vs. globular).

Figure 21. 

Siamlema changhai sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Description

Male (holotype). Total length 0.92. Carapace 0.41 long, 0.40 wide. Abdomen 0.50 long, 0.44 wide. Carapace brown (Fig. 20A). Eyes absent (Fig. 20A). Chelicerae, legs, labium, and endites brown. Sternum bright brown. Leg measurements: I 3.28 (0.95, 0.17, 1.00, 0.69, 0.47); II 2.93 (0.86, 0.16, 0.85, 0.63, 0.43); III 2.17 (0.66, 0.14, 0.58, 0.43, 0.36); IV 2.73 (0.86, 0.14, 0.75, 0.56, 0.42). Abdomen brown with a few long setae (Fig. 20A).

Palp. Tibia 1.57 times longer than patella, cymbium 1.22 times longer than tibia, femur 1.06 times longer than cymbium (Fig. 20C, D); cymbial apophysis present at sub-distal part (Fig. 20C), one femoral distal spine (Fig. 20C, D), a semi-round extension on patella proximally (Fig. 20D); bulb shaped as in Fig. 20C, D; bulbal apophysis nearly transparent, with 4 sclerotized teeth (Fig. 20B–D); embolus sclerotized, finger shaped (Fig. 20B–D).

Female. Total length 1.04. Carapace 0.48 long, 0.41 wide. Abdomen 0.56 long, 0.50 wide. Carapace, sternum, legs milky white (Fig. 21B, C). Leg measurements: I 3.31 (1.00, 0.17, 0.99, 0.67, 0.48); II 2.97 (0.90, 0.16, 0.86, 0.60, 0.45); III 2.18 (0.67, 0.14, 0.59, 0.42, 0.36); IV 2.73 (0.86, 0.15, 0.75, 0.56, 0.41). Abdomen dark brown with some long setae (Fig. 21B, C). Receptacle tube-shaped, neck two times thinner than distal part (Fig. 21A).

Distribution

Thailand (Trang, site 9 in Fig. 33), known only from the type locality.

Siamlema suea Zhao & Li, sp. nov.

Figures 22, 23, 33

Type material

Holotype: ♂ (IZCAS), Thailand, Yala Province, Mueang District, Suea Cave, 6.5226N, 101.2311E, elevation ca. 43 m, XI.2015, Z. Chen, G. Zhou, and Q. Zhao leg. Paratypes: 1♂ and 4♀ (IZCAS), same data as holotype.

Other material examined

1♀ (molecular voucher, IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

This species resembles S. changhai sp. nov. but can be distinguished by the following: six eyes ringed with black or eyes vestigial (Figs 22A, 23B) (vs. absent); the lack of a bulbal apophysis (Fig. 22B–D) (vs. present), having a spine on the palpal tibia retrolaterally (Fig. 22D) (vs. absent), the absence of a femoral dorsal spine (Fig. 22C, D) (vs. present); and a globular receptacle (Fig. 23A) (vs. tube-shaped).

Figure 22. 

Siamlema suea sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Description

Male (holotype). Total length 1.11. Carapace 0.46 long, 0.41 wide. Abdomen 0.60 long, 0.50 wide. Carapace light brown (Fig. 22A). Six vestigial eyes (Fig. 22A). Chelicerae, legs, labium, and endites brown. Sternum bright brown with sparse setae. Leg measurements: I 3.46 (1.03, 0.16, 1.05, 0.74, 0.48); II 2.92 (0.90, 0.16, 0.83, 0.60, 0.43); III 2.07 (0.63, 0.13, 0.59, 0.40, 0.32); IV 2.77 (0.85, 0.16, 0.77, 0.59, 0.40). Abdomen brown with a few long setae (Fig. 22A).

Palp. Tibia 1.38 times longer than patella, cymbium 0.94 times shorter than tibia, femur 1.17 times longer than cymbium (Fig. 22C, D); cymbial apophysis stout and cone shaped (Fig. 22C); tibia with 2 spines, one dorsal spine distally, the other bifurcate, located retrolaterally medially (Fig. 22D); patella with retrolateral semi-round extension mesially (Fig. 22D); bulb droplet shaped as in Fig. 22C, D; embolus strongly sclerotized, and tiny in comparison to bulb.

Female. Total length 1.20. Carapace 0.45 long, 0.41 wide. Abdomen 0.70 long, 0.66 wide. Coloration as in male (Fig. 23B, C). Six eyes ringed with black (Fig. 23B). Leg measurements: I 3.15 (0.94, 0.16, 0.97, 0.61, 0.47); II 2.79 (0.84, 0.16, 0.81, 0.55, 0.43); III 2.02 (0.62, 0.13, 0.56, 0.38, 0.33); IV 2.73 (0.85, 0.14, 0.76, 0.56, 0.42). Abdomen brown (Fig. 23B). Neck of receptacle umbrella-shaped, and distal part of receptacle globular (Fig. 23A).

Distribution

Thailand (Yala, site 8 in Fig. 33), known only from the type locality.

Figure 23. 

Siamlema suea sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Sundalema Zhao & Li, gen. nov.

Type species

Sundalema bonjol sp. nov. from West Sumatra Province, Indonesia.

Etymology

The generic name is derived from “Sunda”, referring to Sundaland (distributional range of this genus), and “-lema” is a convention from the type genus of the family. Feminine in gender.

Diagnosis

Females belonging to Sundalema gen. nov. can be distinguished from those of other genera by the following: a sclerotized and spiral receptacle (Fig. 24A) (vs. unsclerotized or not spiral). Males belonging to Sundalema gen. nov. resemble species in the bailongensis-group by having a long embolus but can be distinguished by the embolus lacking a spiral ridge and being nearly L-shaped (Fig. 25B–D) (vs. a distinct spiral ridge present on the nearly straight embolus).

Figure 24. 

Sundalema bonjol sp. nov., female holotype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Description

Total length: 0.98–1.10 (male), 1.05–1.25 (female). Carapace 0.44– 0.58 long. Sternum light brown or milky white, with several sparse setae. Six vestigial eyes or ringed with black. Leg formula: 1-2-4-3, tibia I 0.90–1.27 long, leg glands belt-shaped (Fig. 1E). For females, neck of receptacle membranous, receptacle spiral, its dorsal part sclerotized, coiled into 1.25 to 2.5 loops. For males, cymbial apophysis tiny, present meso-prolaterally; embolus long relative to bulb, nearly L-shaped, lacking spiral ridge. Abdomen blue or pale brown with several long setae.

Composition

Sundalema acicularis comb. nov., S. anguina (Wang & Li, 2010) comb. nov., S. bonjol sp. nov., and S. khaorakkiat sp. nov.

Distribution

Southeast Asia (Thailand and Indonesia, sites 9–12 in Fig. 33).

Comments

The females of this genus are easily distinguished from all other genera, as their receptacles are long, spiral, and sclerotized; thus, a female has been chosen to represent the holotype.

Sundalema bonjol Zhao & Li, sp. nov.

Figures 24, 25, 33

Type material

Holotype: ♀ (IZCAS), Indonesia, Sumatra, West Sumatra Province, Payakumbuh, Koto Tinggi Village, Imam Bonjol Cave, 0.0637S, 100.3451E, elevation ca. 962 m, III.2014, Z. Yao leg. Paratypes: 2♂ and 3♀ (IZCAS), same data as holotype.

Other material examined

1♀ (molecular voucher, IZCAS), same data as holotype.

Etymology

The species refers to the type locality; noun in apposition.

Diagnosis

The species resembles S. anguina comb. nov. but can be distinguished by the following characters: the eyes are vestigial (Figs 24B, 25A) (vs. ringed with black); the receptacle is coiled into 1.5 loops (Fig. 24A) (vs. 2 loops, cf. Wang and Li 2010a: fig. 4C); the embolus is thin and bent at an obtuse angle (Fig. 25B–D) (vs. wide and bent at a right angle, cf. Wang and Li 2010a: fig. 4A, B).

Figure 25. 

Sundalema bonjol sp. nov., male paratype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Description

Female (holotype). Total length 1.09. Carapace 0.53 long, 0.45 wide. Abdomen 0.64 long, 0.56 wide. Carapace light brown (Fig. 24B). Four vestigial eyes (Fig. 24B). Chelicerae, legs, labium, endites, and sternum bright brown. Sternum with sparse setae (Fig. 24B, C). Leg measurements: I 3.45 (1.04, 0.18, 1.06, 0.67, 0.50); II 3.10 (0.96, 0.17, 0.92, 0.60, 0.45); III 2.17 (0.68, 0.17, 0.58, 0.40, 0.34); IV 2.89 (0.90, 0.16, 0.79, 0.61, 0.43). Abdomen brown (Fig. 24B, C). Receptacle coiled into 1.5 loops, and neck as wide as distal part (Fig. 24A).

Male. Total length 1.03. Carapace 0.47 long, 0.40 wide. Abdomen 0.53 long, 0.44 wide. Coloration as in female (Fig. 25A). Leg measurements: I 3.34 (0.94, 0.16, 1.04, 0.71, 0.49); II 3.05 (0.90, 0.16, 0.91, 0.63, 0.45); III 2.16 (0.67, 0.14, 0.58, 0.42, 0.35); IV 2.82 (0.87, 0.15, 0.79, 0.59, 0.42). Palp: tibia 1.80 times longer than patella, cymbium 1.71 times longer than tibia, 1.69 times longer than femur; length of cymbial apophysis as wide as 1/2 width of cymbial base (Fig. 25C); bulb ellipsoidal (Fig. 25C, D); embolus tube-like, as long as major-axis of bulb (Fig. 25B–D).

Distribution

Indonesia (Sumatra, site 11 in Fig. 33), known only from the type locality.

Sundalema khaorakkiat Zhao & Li, sp. nov.

Figures 26, 27, 33

Type material

Holotype: ♀ (IZCAS), Thailand, Songkhla Province, Rattaphum District, Khao Rak Kiat Cave, 7.0724N, 100.2502E, elevation ca. 52 m, XI.2015, Z. Chen, G. Zhou and Q. Zhao leg. Paratypes: 2♂ and 2♀ (IZCAS), same data as holotype.

Other material examined

1♀ (molecular voucher, IZCAS), same data as holotype.

Etymology

The species name refers to the type locality; noun in apposition.

Diagnosis

This species resembles S. acicularis comb. nov. but can be distinguished by the following characters: the eyes are vestigial (Figs 26B, 27A) (vs. ringed with black); the receptacle is coiled into 1.25 loops (Fig. 26A) (vs. 2.5 loops); the embolus is longer than the cymbium (Fig. 27C, D) (vs. shorter), and the palpal tibia/cymbium ratio is 0.4 (Fig. 27C, D) (vs. 0.45).

Figure 26. 

Sundalema khaorakkiat sp. nov., female holotype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Description

Female (holotype). Total length 1.25. Carapace 0.58 long, 0.50 wide. Abdomen 0.63 long, 0.55 wide. Carapace pale white (Fig. 26B). Six vestigial eyes (Fig. 26B). Chelicerae, legs, labium milky white, endites brown (Fig. 26B). Sternum milky white with sparse setae (Fig. 26B). Leg measurements: I 3.93 (1.21, 0.19, 1.21, 0.80, 0.52); II 3.52 (1.10, 0.19, 1.06, 0.72, 0.45); III 2.45 (0.73, 0.18, 0.69, 0.49, 0.36); IV 3.28 (1.03, 0.18, 0.92, 0.70, 0.45). Abdomen pale brown with a few long setae (Fig. 26B, C). Receptacle coiled into 1.25 loops, and neck much thinner than mesial part (Fig. 26A).

Male. Total length 1.17. Carapace 0.54 long, 0.48 wide. Abdomen 0.63 long, 0.50 wide. Eyes and coloration as in female (Fig. 27A). Leg measurements: I 4.08 (1.24, 0.19, 1.27, 0.86, 0.53); II 3.70 (1.13, 0.19, 1.15, 0.75, 0.48); III 2.67 (0.81, 0.16, 0.76, 0.56, 0.39); IV 3.48 (1.06, 0.17, 1.00, 0.76, 0.49). Palp: Tibia 1.65 times longer than patella, cymbium 2.06 times longer than tibia, 1.50 times longer than femur; cymbial apophysis dark brown and cone shaped (Fig. 27C); bulb ellipsoidal; embolus longer than cymbium (Fig. 27C, D).

Figure 27. 

Sundalema khaorakkiat sp. nov., male paratype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Distribution

Thailand (Songkhla, site 12 in Fig. 33), known only from the type locality.

Sundalema acicularis (Wang & Li, 2010), comb. nov.

Telema acicularis Wang and Li 2010a: 2, figs 1–3 (♂♀).

Type material

Holotype: ♂ (RMNH), Thailand, Prachuap Khiri Khan Province, Hua Hin, Tham Kaew Cave, 12.5947N, 99.9571E, 3.I.1989, C.L. Deeleman-Reinhold leg. Paratypes: 1♂ and 1♀ (RMNH), same data as holotype. Examined.

Diagnosis

This species resembles S. khaorakkiat sp. nov. but can be distinguished by the following characters: six eyes ringed with black (vs. eyes vestigial); receptacle coiled into 2.5 loops (cf. Wang and Li 2010a: fig. 1B) (vs. 1.25 loops); embolus shorter than cymbium (cf. Wang and Li 2010a: fig. 1A) (vs. longer), and the palpal tibia/cymbium length ratio is 0.45 (cf. Wang and Li 2010a: fig. 1A) (vs. 0.4).

Description-amendments

Palpal tibia/cymbium ratio 0.45 in the male. Receptacle coiled into 2.5 loops (cf. Wang and Li 2010a: fig. 1B). For a more detailed description, see Wang and Li (2010a).

Distribution

Thailand (Prachuap Khiri Khan, site 9 in Fig. 33), known only from the type locality.

Sundalema anguina (Wang & Li, 2010), comb. nov.

Figure 1E

Telema anguina Wang and Li 2010a: 2, figs 4–7 (♂♀).

Type material

Holotype: ♂ (RMNH), Thailand, Krabi Province, Diamond Cave, 8.0670N, 98.9164E. 3.I.1989, C.L. Deeleman-Reinhold leg. Paratypes 1♂ and 1♀ (RMNH), same data as holotype. Examined.

Other material examined

3♀ (including one molecular voucher, IZCAS) from the type locality, X.2014, Z. Chen and Y. Li leg.

Diagnosis

This species resembles S. bonjol sp. nov. but can be distinguished by the following characters: the eyes are ringed with black (cf. Wang and Li 2010a: fig. 6B) (vs. vestigial); the receptacle is coiled into two loops (cf. Wang and Li 2010a: figs 4C, 7B) (vs. 1.5 loops); the embolus is thin and bent at a right angle (cf. Wang and Li 2010a: fig. 4A, B) (vs. much wider and obtuse angled).

Description

See Wang and Li (2010a).

Distribution

Thailand (Krabi, site 10 in Fig. 33), known only from the type locality.

Telema Simon, 1882

Telema Simon 1882: 205; Lehtinen 1967: 301; Gertsch 1973: 154; Yaginuma 1973: 17; Wang et al. 2012: 18; Dupérré and Tapia 2015: 191; Zhao et al. 2018a: 14.

Type species

Telema tenella Simon, 1882 from southern France.

Diagnosis

Telema resembles Telemofila but can be distinguished by the following: plate-shaped tibial glands (Fig. 1A; cf. Emerit 1984: fig. A) (vs. belt-shaped); lacking a cymbial apophysis (cf. Wang et al. 2012: fig. 2C) (vs. cymbial apophysis present); the endogyne is walking-stick shaped (cf. Wang et al. 2012: fig. 3C, D) (vs. bean-shaped or globular).

Description

Total length: 1.24–1.48 (male), 1.06–1.90 (female). Carapace 0.45–0.78 long. In T. auricoma Lin & Li, 2010 and T. nipponica, carapace with radial shaded pattern, six eyes ringed with black, sternum dark brown, abdomen blue; in T. tenella, T. guihua Lin & Li, 2010, and T. wunderlichi Song & Zhu, 1992, carapace without pattern, eyes absent, sternum and abdomen bright brown. Leg formula: 1-2-4-3, tibia I 1.32–2.45, tibia with many plate-shaped glands (Fig. 1A; cf. Emerit 1984: fig. A). In male, length of cymbium > femur > tibia > patella; bulb ellipsoid, large relative to body; embolus short relative to bulb, duckbill shaped. Receptacle walking-stick shaped, with a few membranous tubes inside.

Composition

Telema auricoma, T. guihua, T. nipponica, T. tenella and T. wunderlichi. The composition is supported not only by morphological characters but also by molecular phylogenetic analyses (Fig. 34; T. nipponica is not included).

Distribution

Eurasian disjunctive range, known from Southern Europe (France and Spain, one species) and East Asia (Japan and southwestern China, four species, sites 13–16 in Fig. 33).

Telema auricoma Lin & Li, 2010

Telema auricoma Lin and Li 2010: 3, fig. 1 (♀).

Type material

Holotype: ♀ (IZCAS), China, Guizhou Province, Dafang County, Xiaotun Town, Shiqiang Village, Daxiao Cave, 27.0925N,105.5551E, 1168m, 4.V.2007, J. Liu and Y. Lin leg. Paratypes 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♀ (molecular voucher, IZCAS), same data as holotype; 2♀ (IZCAS), Guizhou Province, Bijie County, Xiaoyan Cave, 27.1181N, 105.2367E; 1♀ (IZCAS) Guizhou Province, Dafang County, Shilongshang Cave, 27.0925N, 105.5551E; 3♀ (IZCAS) Guizhou Province, Hezhang County, Tanjiayan Cave, 27.2003N, 104.5910E; 1♀ (IZCAS) Guizhou Province, Weining County, Banbianshan Cave, 33.9061N, 104.5396E; 2♀ (IZCAS), Yunnan Province, Xuanwei County, Jianjiao Cave, 33.3085N, 104.3900E; 3♀ (IZCAS), Yunnan Province, Xuanwei County, Fengchao Cave, 33.3909N, 104.2093E.

Diagnosis and description

See Lin and Li (2010).

Distribution

China (Yunnan-Guizhou Plateau).

Comments

All known specimens of this species are female, and molecular barcoding data from all eight populations examined shows no differences (unpublished data). This indicates that the species may be parthenogenetic, a character that may allow it to easily disperse more broadly than gametogenetic species.

Telema guihua Lin & Li, 2010

Figures 1A, 33

Telema guihua Lin and Li 2010: 12, figs 6, 7 (♂♀).

Type material

Holotype: ♂ (IZCAS), China, Guizhou Province, Suiyang County, Wenquan Town, Guihua Village, Mahuang Cave, 28.2437N,107.2891E, elevation ca. 730 m, 4.V.2007, J. Liu and Y. Lin leg. Paratypes: 1♂ and 2♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♀ (molecular voucher, IZCAS), same data as holotype.

Diagnosis

This species resembles T. wunderlichi but can be distinguished by the following characters: the small body size (vs. larger size); the embolus is membranous (vs. sclerotized), and the tip of the embolus is blunt (cf. Lin and Li 2010: fig. 6E) (vs. sharply pointed).

Description

See Lin and Li (2010).

Distribution

China (Guizhou, site 14 in Fig. 33), known only from the type locality.

Telema nipponica (Yaginuma, 1972)

Merizocera nipponica Yaginuma 1972: 286, figs 10–14 (♀).

Telema nipponica: Yaginuma 1973: 22, figs 1–6 (♂♀); Yaginuma 1974: 14, figs 4–6 (♂); Shinkai 1977: 322, figs 1–3 (♂); Yaginuma 1986: 20, fig. 13.3 (♂); Chikuni 1989: 26, fig. 1 (♂); Ono 2009: 121, figs 1–6 (♂♀).

Type material

Holotype: ♀, Japan, Yamanashi Prefecture, Narusawa-mura, Karumizu, Shoiko-daini-fuketsu Cave, 22.IX.1969, S. Ueno and K. Kato leg. Not examined.

Diagnosis

This species resembles T. tenella but can be distinguished by the thumb-like shape of the embolus (cf. Yaginuma 1974: fig. 4), whereas in T. tenella, the embolus is short and triangular (cf. Wang et al. 2012: fig. 2C, D).

Description

See Yaginuma (1972, 1973).

Distribution

Japan (Site 15 in Fig. 33).

Telema wunderlichi Song & Zhu, 1994

Telema wunderlichi Song and Zhu 1994: 36, fig. 1A–E (♂♀); Song et al. 1999: 51, fig. 11G, 22A–C (♂♀); Yin et al. 2012: 161, fig. 29a, d (♂♀).

Type material

Holotype: ♀ (IZCAS), China, Hunan Province, Zhangjiajie Prefecture, Sangzhi County, Wulingyuan Scientific and Historic Interest Area, 29.1171N, 110.4792E, elevation ca. 280 m, 27.XI.1992. D. Wang leg. Paratypes: 1♂ and 1♀ (IZCAS), same data as holotype. Examined.

Other material examined

1♂ and 2♀ (including molecular voucher, IZCAS) from the type locality: XII.2015, Z. Chen leg.

Diagnosis

This species resembles T. guihua but can be distinguished by the following characters: the larger body size (vs. smaller), the slightly sclerotized embolus (cf. Song and Zhu 1994: fig. 1E) (vs. membranous), the sharply pointed embolus tip (cf. Song and Zhu 1994: fig. 1E) (vs. blunt).

Description

See Song and Zhu (1994), and Lin and Li (2010).

Distribution

China (Hunan, site 16 in Fig. 33).

Telemofila Wunderlich, 1995

Telemofila Wunderlich 1995: 562.

Type species

Telemofila samosirensis Wunderlich, 1995 from Sumatra, Indonesia.

Diagnosis

Telemofila can be distinguished from Telema by the following characters: leg formula is 1-4-2-3 (vs. 1-2-4-3), tibial glands are belt-shaped (Fig. 1G) (vs. plate-shaped); a cymbial apophysis is present (vs. absent), the embolus is sickle shaped or claw-like (vs. duckbill-shaped), and the length of the embolus is three times shorter than the diameter of the bulb (vs. two times shorter); the distal part of the receptacle is swollen (vs. not swollen).

Description

Total length: 0.90–1.27 (male), 0.98–1.11 (female). Carapace 0.37–0.50 long. Sternum 0.20–0.25 long, with several sparse setae. Tibia I 0.51–0.82 long, leg formula: 1-4-2-3, belt-shaped glands present (Fig. 1G). Six eyes ringed with black, body blue or brown. For males, cymbial apophysis present mesially, length as wide as cymbial base (cf. Wunderlich 1995: fig. 16), embolus sickle-shaped, length of embolus 1/3 as long as diameter of bulb (cf. Wunderlich 1995: fig. 17). For females, receptacle bean shaped or globular, neck narrower than distal part (cf. Wang and Li 2010a: figs 11A, B, 15A, B).

Composition

Telemofila fabata (Wang & Li, 2010) comb. nov., T. malaysiaensis comb. nov., T. pecki (Brignoli, 1980), and T. samosirensis Wunderlich, 1995.

Distribution

Rainforests in Southeast Asia (Indonesia, Singapore, Malaysian Borneo, sites 17–19 in Fig. 33) and a cave in New Caledonia.

Comments

The placement of T. pecki in this genus is dubious because its leg formula is 1-2-4-3 (Brignoli 1980) and the embolus is triangular (cf. Brignoli 1980: figs 1, 2). These characters are inconsistent with the genus characters of Telemofila. However, we have been unable to examine the types of T. pecki, and molecular data from this species is lacking.

Telemofila fabata (Wang & Li, 2010), comb. nov.

Telema fabata Wang and Li 2010a: 10, figs 8–11 (♂♀).

Type material

Holotype: ♂ (RMNH), Singapore, Bukit Timah Nature Reserve, Seraya Loop, 1.3521N, 103.8198E, 2.II.1983, P.R. Deeleman leg. Paratypes: 1♂ and 1♀ (RMNH), same data as holotype. Examined.

Other material examined

1♂ and 3♀ (including molecular voucher, IZCAS) from the type locality, VIII.2015, S. Li and Y. Tong leg.

Diagnosis

This species resembles T. malaysiaensis comb. nov. but can be distinguished by the following: bigger body size, bean-shaped receptacle (cf. Wang and Li 2010a: fig. 8C, D) (vs. globular); claw-like embolus (cf. Wang and Li 2010a: fig. 8A, B) (vs. sickle-shaped). This species can be distinguished from T. samosirensis by the embolus being three times shorter than the bulb (cf. Wang and Li 2010a: fig. 9A, B) (vs. two times shorter).

Description

See Wang and Li (2010a).

Distribution

Singapore (Site 17 in Fig. 33).

Telemofila malaysiaensis (Wang & Li, 2010), comb. nov.

Telema malaysiaensis Wang and Li 2010a: 10, figs 12–15 (♂♀).

Type material

Holotype: ♂ (RMNH), Malaysian Borneo, Sarawak Province, swampy lowland rainforest of Bako National Park, 1.7167N, 110.4667E, 29.III.1985, P.R. & C.L. Deeleman-Reinhold leg. Paratypes: 1♂ and 1♀ (RMNH), same data as holotype. Examined.

Diagnosis

This species resembles T. fabata comb. nov. but can be distinguished by the following: the smaller body size, the shape of the receptacle is globular (cf. Wang and Li 2010a: fig. 15B) (vs. receptacle bean-shaped); the embolus is sickle-shaped (cf. Wang and Li 2010a: fig. 12A–C) (vs. claw-like embolus).

Description

See Wang and Li (2010a).

Distribution

Malyasia (Borneo, Sarawak, site 18 in Fig. 33).

Telemofila samosirensis Wunderlich, 1995

Figures 1G, 33

Telemofila samosirensis Wunderlich 1995: 562, figs 10–17 (♂).

Type material

Holotype: ♂ (SMF), Indonesia, Sumatra, North Sumatra Province, Lake Toba, Samosir Village, 2.7424N, 98.7699E, elevation ca. 916 m, VIII. 1994, J. Wunderlich leg. Not examined.

Other material examined

2♀ (including molecular voucher, IZCAS) from the type locality: I.2014, H. Zhao leg.

Diagnosis

This species can be distinguished from T. fabata comb. nov. and T. malaysiaensis comb. nov. by the length of the embolus which is equal to the radius of the bulb (cf. Wunderlich 1995: fig. 17) (vs. length of embolus 1/3 as long as the diameter of the bulb in two similar species).

Descriptions

See Wunderlich (1995).

Distribution

Indonesia (Sumatra, North Sumatra, site 19 in Fig. 33).

Zhuanlema Zhao & Li, gen. nov.

Type species

Zhuanlema peteri sp. nov. from Luang Prabang Province, Laos.

Etymology

The generic name is derived from “Zhuan”, referring to the Chinese pinyin “zhuan”, indicating that the apex of the embolus is twisted, and “-lema” which is a convention from the type genus of the family. Feminine in gender.

Diagnosis

The new genus resembles species in the bailongensis-group but can be distinguished by the following characters: the apex of the embolus is twisted (Fig. 28C) (vs. tube-like embolus), the cymbial apophysis is located basally and four times shorter than the width of the cymbial base (Fig. 28C, D) (vs. cymbial apophysis located mesially or sub-basally, and longer or 1–3 times shorter than the width of the cymbial base). Females can be distinguished by the sclerotized receptacle (Fig. 29A) (vs. membranous).

Description

See species description.

Composition

Zhuanlema peteri sp. nov.

Distribution

Laos (Luang Prabang, site 20 in Fig. 33), known only from the type locality.

Zhuanlema peteri Zhao & Li, sp. nov.

Figures 28, 29, 33

Type material

Holotype: ♂ (SMF): Laos, Luang Prabang Province, NE Luang Prabang, Nam Ou, Nong Khiao, Tham Pathok, 20.5514N, 102.6321E, elevation ca. 373 m, 13.III.2007, P. Jäger leg. Paratypes: 1♂ and 2♀ (SMF), same data as holotype.

Other material examined

1♀ (molecular voucher, IZCAS), same data as holotype.

Etymology

The species is named in honor of Peter Jäger (Frankfurt am Main, Germany), a prolific spider taxonomist.

Diagnosis

See genus diagnosis.

Description

Male (holotype): Total length unknown. Carapace 0.49 long, 0.41 wide. Abdomen lost. Carapace brown (Fig. 28A). Six eyes ringed with black (Fig. 28A). Chelicerae, legs, labium, and endites brown. Sternum light brown with sparse setae. Glands belt-shaped, leg measurements: I 2.44 (0.71, 0.14, 0.73, 0.46, 0.40); II 2.23 (0.67, 0.14, 0.61, 0.43, 0.38); III 1.65 (0.49, 0.14, 0.43, 0.31, 0.28); IV 2.20 (0.65, 0.15, 0.61, 0.44, 0.35).

Palp: tibia 2.15 times longer than patella, cymbium 2.80 times longer than tibia, two times longer than femur. Cymbium bent, cymbial apophysis tiny, about 1/4 of cymbial base width (Fig. 28C); bulb shaped as in Fig. 28C, D with very long and twisted embolus, bent at right-angle dorsally on bulb (arrow 1 in Fig. 28D). Embolus with a right-angled bend (arrow 2 in Fig. 28D), its tip twisted and sclerotized slightly (Fig. 28C, D), spiral ridge originates from base of embolus (Fig. 28B).

Female: Total length 1.18. Carapace 0.49 long, 0.43 wide. Abdomen 0.63 long, 0.56 wide. Coloration as in male (Fig. 29B, C). Leg measurements: I 2.57 (0.77, 0.15, 0.77, 0.48, 0.40); II 2.34 (0.71, 0.14, 0.67, 0.44, 0.38); III 1.72 (0.54, 0.12, 0.45, 0.33, 0.28); IV 2.32 (0.71, 0.13, 0.66, 0.47, 0.35). Abdomen dark brown. Endogyne as in Fig. 29A, receptacle with sclerotized tube inside, comma-shaped, swollen distally, distal part of receptacle two times wider than neck.

Distribution

Laos (Luang Prabang, site 20 in Fig. 33), known only from the type locality.

Figure 28. 

Zhuanlema peteri sp. nov., male holotype. A Habitus, dorsal view B embolus, apical view C palp, prolateral view D palp, retrolateral view. Scale bars: 0.2 mm (A), 0.05 mm (B), 0.1 mm (C, D).

Figure 29. 

Zhuanlema peteri sp. nov., female paratype. A Endogyne, lateral view B habitus, dorsal view C habitus, ventral view. Scale bars: 0.05 mm (A), 0.2 mm (B, C).

Figure 30. 

Distribution records of Pinelema spp., the adunca-group (green), the bailongensis-group (orange), and the cunfengensis-group (blue): 1 P. adunca comb. nov. 2 P. qingfengensis 3 P. renalis comb. nov. 4 P. tortutheca comb. nov. 5 P. yashanensis comb. nov. 6 P. bailongensis 7 P. cheni 8 P. cordata 9 P. curcici 10 P. huoyan 11 P. liangxi 12 P. lizhuang 13 P. strentarsi 14 P. wangshang 15 P. wenyang 16 P. xiushuiensis 17 P. yunchuni 18 P. zhewang 19 P. cunfengensis 20 P. spirae comb. nov.

Figure 31. 

Distribution records of Pinelema spp., the feilong-group (green), the pacchanensis-group (orange), and the podiensis-group (blue): 1 P. bella comb. nov. 2 P. circularis comb. nov. 3 P. claviformis comb. nov. 4 P. damtaoensis 5 P. feilong comb. nov. 6 P. huobaensis 7 P. spina comb. nov. 8 P. vesiculata comb. nov. 9 P. yaosaensis 10 P. daguaiwan sp. nov. 11 P. pacchanensis 12 P. bifida comb. nov. 13 P. biyunensis comb. nov. 14 P. podiensis 15 P. shiba sp. nov.16 P. zonaria comb. nov.

Figure 32. 

Distribution records of Pinelema spp., the xiezi-group (green) and species not attached to a species group (red): 1 P. breviseta comb. nov. 2 P. conglobare comb. nov. 3 P. cucphongensis comb. nov. 4 P. cucurbitina comb. nov. 5 P. dongbei comb. nov. 6 P. exiloculata comb. nov. 7 P. grandidens comb. nov. 8 P. laensis 9 P. oculata comb. nov. 10 P. pedati comb. nov. 11 P. spinafemora comb. nov. 12 P. xiezi 13 P. dengi comb. nov. 14 P. mikrosphaira comb. nov. 15 P. nuocnutensis 16 P. spirulata 17 P. tham sp. nov. 18 P. zhenzhuang.

Figure 33. 

Distribution records of telemid species in East and Southeast Asia (except Pinelema). 1 Apneumonella jacobsoni 2 Mekonglema bailang sp. nov. 3 M. kaorao sp. nov. 4 M. walayaku sp. nov. 5 M. xinpingi comb. nov. 6 M. yan sp. nov. 7 Siamlema changhai sp. nov. 8 S. suea sp. nov. 9 Sundalema acicularis comb. nov. 10 S. anguina comb. nov. 11 S. bonjol sp. nov. 12 S. khaorakkiat sp. nov. 13 Telema auricoma 14 T. guihua 15 T. nipponica 16 T. wunderlichi 17 Telemofila fabata comb. nov. 18 T. malaysiaensis comb. nov. 19 T. samosirensis 20 Zhuanlema peteri sp. nov.

Molecular phylogenetic analyses

Sequence data and model selection

For 71 telemid taxa, a total of 71 and 67 sequences were successfully generated for H3 (331 base pairs, bp) and Wnt (330 bp), respectively. All sequences were submitted to GenBank (Suppl. material 1: Table S2). The best-fit Akaike information criterion (AIC) model of the concatenated BI dataset was SYM+I+G.

Molecular phylogenetic results

The topology of both the ML and BI trees are consistent at the genus level, branches to all tips are long, indicating distinct genetic divergence of each lineage (Fig. 34). Telemidae consists of two major clades: one of Telema and one of the other eight genera.

Telema includes four species (molecular data of T. nipponica was not acquired), and the remaining Telema species are clustered in Pinelema, Sundalema gen. nov., and Telemofila (new combinations in Fig. 34), supporting the hypothesis of previously incorrect generic placement of those species.

Pinelema is monophyletic with high support values (BS = 98 and PP = 100), although most interior nodes are not well supported (Fig. 34). Telemofila is robustly monophyletic (BS = 100 and PP = 100), and molecular data support the transfer of Telemofila fabata comb. nov. to this genus (Fig. 34). Mekonglema xinpingi comb. nov. did not cluster with the type species of Seychellia, so the hypothesis of the incorrect generic placement of M. xinpingi comb. nov. is supported (Fig. 34). The generic placement of Apneumonella jacobsoni is ambiguous because material of the type species A. oculata is lacking. The four new genera (Mekonglema gen. nov., Siamlema gen. nov., Sundalema gen. nov., and Zhuanlema gen. nov.) are monophyletic and (excluding Siamlema gen. nov.) strongly supported by both ML and BI (Fig. 34). This result is consistent with morphological delimitation (Fig. 34).

Figure 34. 

Maximum likelihood tree of Asian Telemidae. Each genus is indicated by black bar or solid line. BS < 70 and PP < 95 are not shown, taxa followed by an asterisk indicate the type species of the corresponding genus. BS: bootstrap support; PP: posterior possibility.

Discussion

In this paper we constructed a molecular phylogeny of Telemidae from East and Southeast Asia for the first time, and our results have changed the taxonomic framework of Asian telemids. We revised the taxonomic status of these Asian telemids, introducing four new genera and 12 new species, and 31 species were transferred to other genera.

When researchers are delimiting genera of Telemidae, qualitative morphological characters should be considered. These characters include: for both sexes, the leg formula and the shape of tibial glands; for males, the presence or absence of a cymbial apophysis, the length ratio of the palpal femur/cymbium, and the accessory structures on the palpal tibia or bulb; for females, the shape of the receptacle and the presence/absence of membranous/sclerotized tubes within the receptacle. Quantitative morphological characters of reproductive organs should be considered when researchers are delimiting different species in the same genus. These characters include the ratio of the embolus/bulb length, the angle between the embolus and bulb, and the width ratio of the receptacle tip/neck, etc.

Despite the large genetic difference between Telema species in East Asia and T. tenella (e.g. the long branch between T. tenella and Asian Telema spp. in Fig. 34) this genus occupies a small morphospace. This is also supported by paleontological evidence, as the morphological characters of T. moritzi Wunderlich, 2004 (extinct species in Baltic amber, ca. 45 million years ago) are difficult to distinguish from extant Telema spp. (Wunderlich 2004). The occurrence of Telema spp. in the entire Palearctic is intriguing given their extremely weak dispersal ability and requires further investigation. It is possible that they dispersed from Europe to Asia (or vice versa) in ancient warm stage, then many species became extinct when the climate cooled. Only a few species survived in refuges like southern Europe and the tropical and subtropical areas of Asia, and this has resulted in the current fragmented distribution.

Acknowledgements

The manuscript benefited greatly from comments by Yuri Marusik, Kadir Boğaç Kunt, and an anonymous reviewer. Sarah Crews kindly checked the language. Yang Song helped to prepare the figures for this study. Peter Jäger collected Mekonglema kaorao sp. nov. and Zhuanlema peteri sp. nov. This study was supported by the National Natural Science Foundation of China to Aibing Zhang (NSFC-31772501, 31425023) and Shuqiang Li (NSFC-31530067, 31970396, 31572304).

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1 Females of Pinelema and Apneumonella are indistinguishable.
2 Mekonglema gen. nov. part I includes M. kaorao sp. nov., M. walayaku sp. nov., and M. xinpingi comb. nov.
3 Females of Mekonglema gen. nov. part II (M. bailang sp. nov. and M. yan sp. nov.) and Siamlema gen. nov. part II (S. changhai sp. nov.) are indistinguishable.

Supplementary material

Supplementary material 1 

Taxonomic revision on Telemidae (Arachnida, Araneae) from East and Southeast Asia

Huifeng Zhao, Shuqiang Li, Aibing Zhang

Data type: Primer/specimen/DNA sequence

Explanation note: Table S1. The locus, primer pairs, and PCR protocols used in this study (Nested PCR, the product of round 1 was used as the template for round 2). Table S2. List of voucher information and GenBank accession numbers (na, indicates the molecular data is not acquired).

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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