Khorata Huber, 2005

Li, Jinglin, Zhang, Baisensen, Li, Shuqiang & Yao, Zhiyuan, 2025, Taxonomic notes on four closely related spider species of the genus Khorata (Araneae, Pholcidae), ZooKeys 1244, pp. 281-291 : 281-291

Genus Khorata Huber, 2005

Khorata Huber, 2005: 79. Yao and Li 2010: 5. Xu et al. 2020: 159.

Type species.

Khorata khammouan Huber, 2005 from Laos.

Diagnosis.

The genus can be distinguished from Savarna Huber, 2005 by the male chelicera with a sclerotized distal apophysis (da) on the front-lateral surface and a hooked frontal apophysis (fa 2), by the male palpal trochanter apophysis extremely short (as long as wide) and not attached to the femur, by the male palpal femur with a retrolateral apophysis (ra), and by the epigyne with a posterior lip (pl) in nearly half of the species.

Remarks.

Species of Khorata are medium-sized with long legs, and construct domed webs at limestone cave entrances or between rocks ( Xu et al. 2020). Three notable exceptions, K. dupla Yao & Li, 2013 , K. bayeri Yao, Li & Jäger, 2014 and K. kep Lan, Jäger & Li, 2021 , possess shorter legs and inhabit leaf litter ( Yao and Li 2013; Yao et al. 2014; Lan et al. 2021). At present, the records from Kep ( Cambodia; K. kep ), Chiang Rai ( Thailand; K. schwendingeri Huber, 2005 ), Tongren (Guizhou, China; K. yuhaoi Xu, Zheng & Yao, 2020 ) and Wuyi Mt. (Fujian, China; K. zhui Zhang & Zhang, 2008 ) represent the southernmost, westernmost, northernmost and easternmost distribution limit for the genus, respectively (Fig. 1 ). Forty-two species (79 %) are known from a single locality and three species are sympatric (found in the same locality; K. dangi Yao, Pham & Li, 2015 , K. digitata Yao & Li, 2010 , K. huberi Yao, Pham & Li, 2015 ). Morphological data indicate a close relationship among the genera Khorata , Savarna , Hantu Huber, 2016 , and Aetana Huber, 2005 ( Huber et al. 2015); however, the first three genera were merely used as outgroups for the genus Aetana , and the evidence supporting the inter-genetic relationships was limited. The most recent molecular data suggest that the phylogenetic positions of Khorata , Savarna , and Hantu within the ‘ Pholcinae group 1 ’ are unstable and problematic, necessitating further analysis ( Eberle et al. 2018; Huber et al. 2018).

Thirty-seven species of this genus are here divided into nine formal species groups based on superficial similarity, while the remaining species are tentatively not assigned to any species groups.

The digitata group. This group includes four species: K. digitata , K. qian Yao & Li, 2019 , K. qianlei sp. nov. and K. yuhaoi . They share five putative synapomorphies on the procursus: two prolatero-distal apophyses (pda 1, pda 2), distal sclerite (ds), dorso-subdistal apophysis (dsa), and retrolatero-distal apophysis (rda) (Figs 2 C , 4 A, C, E ). In addition, endogynes of K. digitata , K. qian and K. yuhaoi have sclerites (as) anterior to pore plates (pp) (Fig. 4 B, D, F ).

The epunctata group. This group includes four species: K. circularis Yao & Li, 2013 , K. epunctata Yao & Li, 2010 , K. fusui Zhang & Zhu, 2009 and K. shao Yao & Li, 2010 . They share two putative synapomorphies on the procursus: lamellar ventro-distal apophysis (vda) and slightly sclerotized distal apophysis (da) (e. g., fig. 28 C in Yao and Li 2013).

The flabelliformis group. This group includes six species: K. dongkou Yao & Li, 2010 , K. flabelliformis Yao & Li, 2010 , K. guiensis Yao & Li, 2010 , K. macilenta Yao & Li, 2010 , K. miaoshanensis Yao & Li, 2010 and K. ningyuan Wei & Xu, 2014 . They share two putative synapomorphies: spine-shaped prolatero-ventral apophysis (pva) on the procursus (e. g., fig. 13 D in Yao and Li 2010) and posterior lip (pl) on the epigyne (e. g., fig. 14 A in Yao and Li 2010).

The khammouan group. This group includes two species: K. khammouan Huber, 2005 and K. protumida Yao, Pham & Li, 2015 . They share two putative synapomorphies: sclerotized prolatero-dorsal apophysis (pda) on the procursus (e. g., fig. 151 in Huber 2005) and a pair of protrusions on the epigyne (e. g., fig. 155 in Huber 2005).

The matang group. This group includes two species: K. matang Yao & Li, 2019 and K. wenshan Yao & Li, 2019 . They share three putative synapomorphies: bifurcated prolatero-distal apophysis (pda) and bifurcated retrolatero-distal apophysis (rda) on the procursus (e. g., fig. 7 C, D in Yao et al. 2019) and posterior lip (pl) on the epigyne (e. g., fig. 8 A in Yao et al. 2019).

The nanningensis group. This group includes eight species: K. bachma Yao & Li, 2018 , K. danxia Sheng & Xu, 2021 , K. kep , K. nanningensis Yao & Li, 2010 , K. ninhbinh Zhang, Li & Yao, 2024 , K. quangbinh Yao & Li, 2018 , K. robertmurphyi Yao & Li, 2010 and K. suwei Yao & Li, 2019 . They share two putative synapomorphies on the procursus: lamellar ventro-distal apophysis (vda) and dorso-distal apophysis (dda) bearing teeth (e. g., fig. 37 D in Yao and Li 2010).

The palace group. This group includes two species: K. luoping Yao & Li, 2019 and K. palace Yao & Li, 2018 . They share three putative synapomorphies on the procursus: bifurcated prolatero-distal apophysis (pda), ventro-distal apophysis (vda), and spine-shaped retrolatero-distal apophysis (rda) (e. g., fig. 7 C in Nie et al. 2018).

The schwendingeri group. This group includes three species: K. bangkok Huber, 2005 , K. musee Lan & Li, 2021 and K. schwendingeri . They share one putative synapomorphy on the procursus: spine-shaped prolatero-dorsal apophysis (pda) (e. g., fig. 161 in Huber 2005).

The triangula group. This group includes six species: K. libo Yao & Li, 2019 , K. liuzhouensis Yao & Li, 2010 , K. luojinensis Yao & Li, 2010 , K. paquini Yao & Li, 2010 , K. sancai Wei & Xu, 2014 and K. triangula Yao & Li, 2010 . They share one putative synapomorphy on the procursus: lamellar distal apophysis (da) (e. g., fig. 55 D in Yao and Li 2010).

Figures

Figure 1. Distribution records of Khorata species. 1 K. bachma Yao & Li, 2018 2 K. bangkok Huber, 2005 3 K. bayeri Yao, Li & Jäger, 2014 4 K. circularis Yao & Li, 2013 5 K. cucphuong Yao & Li, 2018 6 K. dangi Yao, Pham & Li, 2015 7 K. danxia Sheng & Xu, 2021 8 K. dawei Yao & Li, 2019 9 K. diaoluoshanensis Tong & Li, 2008 10 K. digitata Yao & Li, 2010 11 K. dongkou Yao & Li, 2010 12 K. dupla Yao & Li, 2013 13 K. epunctata Yao & Li, 2010 14 K. flabelliformis Yao & Li, 2010 15 K. fusui Zhang & Zhu, 2009 16 K. guiensis Yao & Li, 2010 17 K. huberi Yao, Pham & Li, 2015 18 K. jaegeri Huber, 2005 19 K. kep Lan, Jäger & Li, 2021 20 K. khammouan Huber, 2005 21 K. libo Yao & Li, 2019 22 K. liuzhouensis Yao & Li, 2010 23 K. luojinensis Yao & Li, 2010 24 K. luoping Yao & Li, 2019 25 K. macilenta Yao & Li, 2010 26 K. matang Yao & Li, 2019 27 K. miaoshanensis Yao & Li, 2010 28 K. musee Lan & Li, 2021 29 K. nani Xu, Zheng & Yao, 2020 30 K. nanningensis Yao & Li, 2010 31 K. ningming Zhang & Zhu, 2009 32 K. ningyuan Wei & Xu, 2014 33 K. ninhbinh Zhang, Li & Yao, 2024 34 K. palace Yao & Li, 2018 35 K. paquini Yao & Li, 2010 36 K. protumida Yao, Pham & Li, 2015 37 K. qian Yao & Li, 2019 38 K. qianlei sp. nov. 39 K. qiyunshanensis Zhou, 2024 40 K. quangbinh Yao & Li, 2018 41 K. robertmurphyi Yao & Li, 2010 42 K. rongshuiensis Yao & Li, 2010 43 K. sancai Wei & Xu, 2014 44 K. schwendingeri Huber, 2005 45 K. shao Yao & Li, 2010 46 K. suwei Yao & Li, 2019 47 K. triangula Yao & Li, 2010 48 K. vinhphuc Yao & Li, 2018 49 K. wangae Yao & Li, 2010 50 K. wenshan Yao & Li, 2019 51 K. xingyi Chen, Zhang & Zhu, 2009 52 K. yangchun Yao & Li, 2019 53 K. yuhaoi Xu, Zheng & Yao, 2020 54 K. zhui Zhang & Zhang, 2008. Arrow indicates the habitat.

Figure 2. Khorata qianlei sp. nov., holotype male. A, B. Palp (A. Prolateral view; B. Retrolateral view); C, D. Distal part of procursus (C. Prolateral view; D. Retrolateral view). Abbreviations: b = bulb, ds = distal sclerite, dsa = dorso-subdistal apophysis, e = embolus, fra = femoral retrolateral apophysis, pda = prolatero-distal apophysis, pr = procursus, rda = retrolatero-distal apophysis, tra = trochanteral retrolateral apophysis, va = ventral apophysis. Scale bars: 0.20 mm (A, B); 0.05 mm (C, D).

Figure 4. Khorata digitata (A, B), K. qian (C, D), K. yuhaoi (E, F), male (A, C, E) and female (B, D, F). A, C, E. Distal parts of procursus, prolateral views; B, D, F. Endogynes (F. Arrow points at slightly curved part). Abbreviations: aa = anterior arch, as = anterior sclerite, ds = distal sclerite, dsa = dorso-subdistal apophysis, pda = prolatero-distal apophysis, pl = posterior lip, pp = pore plate, psa = prolatero-subdistal apophysis, rda = retrolatero-distal apophysis. Scale bars: 0.05 mm (A, C, E); 0.10 mm (B, D, F).