Research Article |
Corresponding author: Feng Zhang ( dudu06042001@163.com ) Corresponding author: Zhisheng Zhang ( zhangzs327@qq.com ) Academic editor: Ingi Agnarsson
© 2025 Yannan Mu, Luyu Wang, Feng Zhang, Zhisheng Zhang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Mu Y, Wang L, Zhang F, Zhang Z (2025) Mutatus gen. nov., a new genus of Phrurolithidae (Arachnida, Araneae) from China, with descriptions of four new species. ZooKeys 1226: 121-137. https://doi.org/10.3897/zookeys.1226.137752
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Mutatus gen. nov. is described from southwest China, including four new species: M. gelao sp. nov. (♂♀, Guizhou Province), M. tianyan sp. nov. (♂♀, Hubei Province), M. tujia sp. nov. (♂♀, Hubei Province), and M. yintiaoling sp. nov. (♂, Chongqing Municipality). This new genus can be differentiated from other phrurolithid genera by the male palp which has a prolateral tegular apophysis and tegular sheet, and by a small protuberance on the female epigynal plate.
Asia, guardstone spiders, morphology, taxonomy
The family Phrurolithidae Banks, 1892 comprises 405 species in 25 genera and is mainly distributed in the Holarctic and Southeast Asia, but with a few species known from Africa and Australia. Members of the family Phrurolithidae usually inhabit leaf litter on the forest floor, the undersides of stones and tree bark (
While examining phrurolithid specimens collected in Chongqing Municipality, and Hubei and Guizhou provinces, we identified species exhibiting a unique combination of traits. The presence of a tegular sheet suggested a close relationship with Lingulatus Mu & Zhang, 2022, while the prolateral tegular apophysis, indicates an affinity with Xilithus Liu & Li, 2023. Based on this unique combination of characteristics, we describe four new species as members of a new genus, Mutatus gen. nov.
All specimens are preserved in 95% alcohol. All measurements in the text are given in millimeters. The measurements of legs are shown as total length (femur, patella, tibia, metatarsus, tarsus). The palps were dissected from the femur. The epigyna were removed and cleared in a pancreatin solution (
The following abbreviations are used: AER—anterior eye row; ALE—anterior lateral eye; AME—anterior median eye; CH—clypeal height; CRW—cephalic region width; CT—connecting tube; CW—carapace width; EAW—eye area width; FA—femur apophysis; MOA—median ocular area; MS—median septum; PLE—posterior lateral eye; PME—posterior median eye; PER—posterior eye row; PTA—prolateral tegular apophysis; RTA—retrolateral tibial apophysis; TS—tegular sheet; RvTA—retroventral tibial apophysis. Spination: d—dorsal; pl—prolateral; pv—proventral; rv—retroventral.
Phrurolithidae Banks, 1892
The genus name is derived from a Latin word “mutatio”, meaning “variable”, referring to the unique combination of characters in this new genus. Unlike the similar genera Lingulatus, which possesses only a tegular sheet, and Xilithus, which features only prolateral tegular apophysis, the new genus exhibits both characteristics in the male palp. Gender masculine.
The male of new genus resembles those of the genera Lingulatus and Xilithus in having both tegular sheet and prolateral tegular apophysis, but Mutatus can be differentiated by the absence of a ventral tibial apophysis (present both in Lingulatus and Xilithus), tegular sheet present in both Mutatus gen. nov. and Lingulatus (absent in Xilithus), and sperm duct V-shaped in both Mutatus gen. nov. and Lingulatus (U-shaped in Xilithus). The female of new genus resembles Xilithus in having a pair of large concavities (atria), a pair of small bursae, and clavate spermathecae, but it can be differentiated by the presence of a protuberance at the middle part of the lateral margin of the atrium (absent in Xilithus).
Small, total length 3.18–4.78 mm. Carapace oval, smooth, brownish to black, widest at coxae II and III, highest near fovea, with several markings resembling flowing water droplets beside fovea. Fovea dark-red. PER slightly wider than AER, PER recurved in dorsal view, AER recurved in dorsal view. Chelicerae brown, with 1 long and 1 short spine anteriorly. Endites longer than wide, labium wider than long. Sternum brown, shield-shaped, smooth and without pattern, longer than wide. Legs dark brown, all femora with 1 dorsal spine; femur I with 3–5 prolateral spines distally, femur II with 0–2 prolateral spines distally; tibiae I–II usually with 6–7 pairs of ventral spines, tibiae III–IV without ventral spines; metatarsus I with 4–5 pairs of ventral spines, metatarsus II usually with 4 proventral spines and 3 retroventral spines, metatarsi III–IV without ventral spines but with distal preening brush. Leg formula 4123. Abdomen oval, gray to black in dorsal view, with dorsal scutum covering almost entire abdomen in males, scutum in females absent.
Male palp: femur with well-developed apophysis located at middle part; patella short, as long as tibia; tibia slightly bulged ventrally, with retroventral and retrolateral apophyses; cymbium longer than femur; bulb pyriform; prolateral tegular apophysis and tegular sheet (spade-shaped) originating near base of embolus; embolus acicular; conductor absent.
Epigyne: epigynal plate weakly sclerotized, with pair of atria; septum dumbbell-shaped; copulatory openings small, separated; copulatory ducts thin; connecting tubes thick, shorter than copulatory ducts; bursae small; spermathecae clavate, located posteriorly; glandular appendages mastoid-shaped.
Mutatus gelao sp. nov.
Mutatus gelao sp. nov., M. tujia sp. nov., M. tianyan sp. nov., and M. yintiaoling sp. nov.
China (Hubei Province, Guizhou Province, Chongqing Municipality).
Holotype : China • ♂; Guizhou Province, Tongren City, Shiqian County, Gelao Village; 27°2'3.12"N, 108°9'6.94"E, elev. 647 m, 08 May 2023; Z.Y. Li, W.H. Wang leg. Paratype: 1♀, with same data as holotype.
The specific name comes from the word “Gelao”. The name is one of the Chinese ethnic minorities that live adjacent to the area inhabited by the new species; a noun in apposition.
The new species can be differentiated from Mutatus tujia sp. nov. and M. tianyan sp. nov. by the ventrally bulging tibia of the male palp (vs not inflated; compare Fig.
Male (holotype, Fig.
Palp (Fig.
Copulatory organs of Mutatus gelao sp. nov. A male left palp, prolateral view B same, ventral view C same, retrolateral view D same, dorsal view E RTA, retrolateral view F femur, prolateral view G epigyne, ventral view H same, dorsal view. Abbreviations: E—embolus; FA—femoral apophysis; PTA—prolateral tegular apophysis; RTA—retrolateral tibial apophysis; RvTA—retroventral tibial apophysis; SD—sperm duct; TS—tegular sheet; B—bursa; CD—copulatory duct; CO—copulatory opening; CT—connecting tube; FD—fertilization duct; GA—glandular appendage; MS—median septum; S—spermathecae.
Female (Fig.
Epigyne (Fig.
Measurement of legs:
male/female | Fe | Pa | Ti | Me | Ta | Total |
---|---|---|---|---|---|---|
Leg I | 1.67/1.71 | 0.60/0.56 | 1.80/1.65 | 1.58/1.51 | 0.89/0.85 | 6.54/6.27 |
Leg II | 1.54/1.63 | 0.47/0.61 | 1.44/1.49 | 1.20/1.38 | 0.86/0.93 | 5.51/6.04 |
Leg III | 1.31/1.43 | 0.51/0.54 | 1.07/1.07 | 1.22/1.33 | 0.70/0.77 | 4.81/5.14 |
Leg IV | 1.96/2.10 | 0.58/0.65 | 1.79/1.73 | 2.01/2.06 | 1.02/0.97 | 7.36/7.51 |
Spination of legs:
Fe | Pa | Ti | Me | Ta | ||
---|---|---|---|---|---|---|
male | Leg I | d 1 pl 4 | – | pv 7 rv 7 | pv 4 rv 4 | – |
Leg II | d 1 pl 0 | – | pv 7 rv 5 | pv 4 rv 4 | – | |
Leg III | d 1 | – | – | – | – | |
Leg IV | d 1 | – | – | – | – | |
female | Leg I | d 1 pl 5 | – | pv 7 rv 7 | pv 5 rv 5 | – |
Leg II | d 1 pl 2 | – | pv 7 rv 7 | pv 4 rv 3 | – | |
Leg III | d 1 | – | – | – | – | |
Leg IV | d 1 | – | – | – | – |
Known only from the type locality.
Holotype : China • ♂; Hubei province, Shennongjia Forest District, Xiaguping, Tujiazu Township, Huangxi River; 31°21'59.87"N, 110°11'18.32"E, elev. 1059 m, 19 May 2023; Z.Y. Li, Z.Y. Yang leg. Paratypes: • 1♂ 7♀, with same data as holotype; • 2♂ 4♀ same locality, Dongkou Village (31°22'54.25"N, 110°9'48.53"E, 1053 m a.s.l.), 19 May 2023, Z.Y. Li, Z.Y. Yang leg.
The specific name comes from the word “Tujia”. The name is one of the Chinese ethnic minorities that live adjacent to the area inhabited by the new species; a noun in apposition.
The new species resembles M. tianyan sp. nov. in having similar truncated prolateral tegular apophysis and embolus, but it can be differentiated by the following: posterior of tegular sheet tail-shaped (vs tongue-shaped; compare Fig.
Male (holotype, Fig.
Palp (Fig.
Copulatory organs of Mutatus tujia sp. nov. A male left palp, prolateral view B same, retrolateral view C same, dorsal view D same, ventral view E epigyne, ventral view F same, dorsal view. Abbreviations: E—embolus; FA—femoral apophysis; PTA—prolateral tegular apophysis; RTA—retrolateral tibial apophysis; RvTA—retroventral tibial apophysis; SD—sperm duct; TS—tegular sheet; B—bursa; CD—copulatory duct; CO—copulatory opening; CT—connecting tube; FD—fertilization duct; GA—glandular appendage; MS—median septum; S—spermathecae.
Female (one of the paratypes; Fig.
Epigyne (Fig.
Measurement of legs:
male/female | Fe | Pa | Ti | Me | Ta | Total |
---|---|---|---|---|---|---|
Leg I | 1.38/1.44 | 0.43/0.46 | 1.42/1.58 | 1.14/1.34 | 0.61/0.69 | 4.98/5.51 |
Leg II | 1.20/1.19 | 0.45/0.44 | 1.10/1.21 | 0.96/1.07 | 0.58/0.58 | 4.29/4.40 |
Leg III | 1.00/1.01 | 0.39/0.39 | 0.80/0.81 | 0.91/1.01 | 0.57/0.58 | 3.67/3.80 |
Leg IV | 1.50/1.62 | 0.44/0.46 | 1.32/1.53 | 1.45/1.60 | 0.77/0.89 | 5.48/6.10 |
Spination of legs:
Fe | Pa | Ti | Me | Ta | ||
---|---|---|---|---|---|---|
male | Leg I | d 1 pl 3 | – | pv 7 rv 7 | pv 4 rv 4 | – |
Leg II | d 1 pl 0 | – | pv 6 rv 6 | pv 4 rv 3 | – | |
Leg III | d 1 | – | – | – | – | |
Leg IV | d 1 | – | – | – | – | |
female | Leg I | d 1 pl 3 | – | pv 6 rv 6 | pv 4 rv 4 | – |
Leg II | d 1 pl 0 | – | pv 6 rv 6 | pv 4 rv 4 | – | |
Leg III | d 1 | – | – | – | – | |
Leg IV | d 1 | – | – | – | – |
Known only from the type locality.
Holotype : China • ♂ Hubei province: Shennongjia Forest District, Tianyan Primitive Ecotourism Area; 31°43'16"N, 110°28'31"E, elev. 1871 m, 14 June 2023, Z.S. Zhang, Y.J. Cai leg. Paratype: • 7♀, with same data as holotype.
This specific name is derived from the type locality; a noun in apposition.
See the diagnosis of M. tujia sp. nov.
Male (holotype, Fig.
Palp (Fig.
Copulatory organs of Mutatus tianyan sp. nov. A male left palp, prolateral view B same, retrolateral view C same, ventral view D same, dorsal view E epigyne, ventral view F same, dorsal view. Abbreviations: E—embolus; FA—femoral apophysis; PTA—prolateral tegular apophysis; RTA—retrolateral tibial apophysis; RvTA—retroventral tibial apophysis; SD—sperm duct; TS—tegular sheet; B—bursa; CD—copulatory duct; CO—copulatory opening; CT—connecting tube; FD—fertilization duct; GA—glandular appendage; MS—median septum; S—spermathecae.
Female (one of paratype, Fig.
Epigyne (Fig.
Measurement of legs:
male/female | Fe | Pa | Ti | Me | Ta | Total |
---|---|---|---|---|---|---|
Leg I | 1.49/1.43 | 0.53/0.57 | 1.45/1.55 | 1.30/1.37 | 0.73/0.77 | 5.50/5.69 |
Leg II | 1.28/1.27 | 0.49/0.54 | 1.09/1.13 | 1.07/1.10 | 0.70/0.75 | 4.63/4.79 |
Leg III | 1.10/1.10 | 0.46/0.43 | 0.84/0.82 | 1.02/1.10 | 0.61/0.68 | 4.03/4.13 |
Leg IV | 1.59/1.68 | 0.53/0.57 | 1.34/1.45 | 1.64/1.69 | 0.84/0.93 | 5.94/6.32 |
Spination of legs:
Fe | Pa | Ti | Me | Ta | ||
---|---|---|---|---|---|---|
male | Leg I | d 1 pl 4 | – | pv 6 rv 6 | pv 4 rv 4 | – |
Leg II | d 1 pl 1 | – | pv 6 rv 4 | pv 4 rv 3 | – | |
Leg III | d 1 | – | – | – | – | |
Leg IV | d 1 | – | – | – | – | |
female | Leg I | d 1 pl 3 | – | pv 7 rv 7 | pv 4 rv 4 | – |
Leg II | d 1 pl 2 | – | pv 6 rv 6 | pv 4 rv 3 | – | |
Leg III | d 1 | – | – | – | – | |
Leg IV | d 1 | – | – | – | – |
Known only from the type locality.
Holotype ♂ China • Chongqing Municipality: Yintiaoling National Nature Reserve, Zhuanping (31°29'43.12"N, 109°55'17.96"E, elev. 1752 m), 23 May 2022, H.L. Zhu leg.
This specific name is derived from the type locality; a noun in apposition.
This new species can be differentiated from M. gelao sp. nov., M. tujia sp. nov., and M. tianyan sp. nov. by the shorter prolateral tegular apophysis (vs long; compare Fig.
Habitus and copulatory organs of Mutatus yintiaoling sp. nov. A male holotype, dorsal view B male left palp, ventral view C same, dorsal view D same, prolateral view E same, retrolateral view. Abbreviations: E—embolus; FA—femoral apophysis; PTA—prolateral tegular apophysis; RTA—retrolateral tibial apophysis; RvTA—retroventral tibial apophysis; SD—sperm duct; TS—tegular sheet.
Male (holotype, Fig.
Palp (Fig.
Female: unknown.
Measurement of legs:
male | Fe | Pa | Ti | Me | Ta | Total |
---|---|---|---|---|---|---|
Leg I | 1.62 | 0.62 | 1.61 | 1.60 | 0.74 | 6.19 |
Leg II | 1.41 | 0.61 | 1.21 | 1.11 | 0.73 | 5.07 |
Leg III | 1.26 | 0.52 | 0.94 | 1.19 | 0.76 | 4.67 |
Leg IV | 1.81 | 0.67 | 1.41 | 1.64 | 0.86 | 6.39 |
Spination of legs:
Fe | Pa | Ti | Me | Ta | ||
---|---|---|---|---|---|---|
male | Leg I | d 1 pl 3 | – | pv 6 rv 6 | pv 4 rv 3 | – |
Leg II | d 1 pl 2 | – | pv 6 rv 5 | pv 4 rv 3 | – | |
Leg III | d 1 | – | – | – | – | |
Leg IV | d 1 | – | – | – | – |
Known only from the type locality.
Mutatus gen. nov., resembles Lingulatus and Xilithus in having tegular sheet and prolateral tegular apophysis on the male palp, respectively. In Lingulatus, the position of the tegular sheet seems variable and aligns with the position of embolus, which suggests a functional relationship between the tegular sheet and the embolus. In Mutatus gen. nov., the position of the tegular sheet and embolus are fixed and resembling the embolus and distal tegular apophysis in Xilithus. The distal tegular apophysis appears morphologically matched with the atria in Xilithus, while significant differences exist between tegular sheet and atria in Mutatus gen. nov. Additionally, the presence of a prolateral tegular apophysis and similar epigynal features further indicate a close relationship between Mutatus gen. nov. and Xilithus.
If the tegular sheet in Lingulatus and Mutatus gen. nov. and distal tegular apophysis in Xilithus were considered homologous, it could suggest that Mutatus gen. nov. and Lingulatus are subgroups of Xilithus. However, substantial differences between the Lingulatus and Xilithus argue against this hypothesis. These differences include the presence of a prolateral tegular apophysis and atria in Xilithus (absent/occasional in Lingulatus), as well as ventral tibial apophysis, tegular sheet, and large bursae found in Lingulatus (small or absent in Xilithus). Furthermore, the length and spatial path of sperm duct differs significantly between Lingulatus and Xilithus. So, we believe that the tegular sheet and distal tegular apophysis are not homologous, and the establishment of Mutatus gen. nov. as a valid genus is well supported.
We thank Zhaoyi Li, Weihang Wang, Jainzhou Sun, Jiaqi Zhao, Long Hao, Xiaoru Qi, and Zhiyong Yang (Hebei University), and Xulong Chen, Qianle Lu, Yujun Cai (Southwest University) for collecting valuable specimens. Many thanks go to subject editor Ingi Agnarsson and the two reviewers, David Chamé Vázquez and an anonymous reviewer for their valuable comments, which greatly improved the quality of the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was supported by the Science & Technology Fundamental Resources Investigation Program (Grant No. 2022FY202100), the National Natural Science Foundation of China (No. 32170468), Project of Background Resources Survey in Shennongjia National Park (SNJNP2022009), and Open Project Fund of Hubei Provincial Key Laboratory on Conservation Biology of the Shennongjia Golden Snub-nosed Monkey (SNJGKL2022009).
All authors have contributed equally.
Yannan Mu https://orcid.org/0000-0002-2504-673X
Luyu Wang https://orcid.org/0000-0002-5250-3473
Feng Zhang https://orcid.org/0000-0002-3347-1031
Zhisheng Zhang https://orcid.org/0000-0002-9304-1789
All of the data that support the findings of this study are available in the main text.