Research Article |
Corresponding author: Qiaoqiao He ( heqq@synu.edu.cn ) Corresponding author: Zhiyuan Yao ( yaozy@synu.edu.cn ) Academic editor: Danilo Harms
© 2024 Lan Yang, Qiaoqiao He, Zhiyuan Yao.
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:
Yang L, He Q, Yao Z (2024) Taxonomic study of four closely-related species of the Pholcus yichengicus species group (Araneae, Pholcidae) from China’s Qinling Mountains: An integrated morphological and molecular approach. Zoosystematics and Evolution 100(1): 279-289. https://doi.org/10.3897/zse.100.115633
|
Four morphologically similar species of the Pholcus yichengicus species group, occurring in geographic proximity of China’s Qinling Mountains, were recognised, based on morphology and four methods of molecular species delimitation. They comprise two new species, namely Pholcus ankang sp. nov. and P. baoji sp. nov. and two previously described species: P. ovatus Yao & Li, 2012 and P. taibaiensis Wang & Zhu, 1992. Their DNA barcodes were obtained to estimate p-distances and K2P distances. In addition, an identification key for the four closely-related species is presented.
Biodiversity, daddy-long-legs spider, identification key, molecular species delimitation, new species
The family Pholcidae C.L. Koch, 1850 is a highly diverse group of spiders, with 97 genera and 1,937 species (
Bestriding the Palaearctic and Oriental Regions, China harbours a high diversity of Pholcus spiders. Recently, a large number of new species of Pholcus have been reported from northern China, based on morphological and molecular data. For instance, the extensive 2020 expedition into the Changbai Mountains, at the border between north-eastern China and North Korea, brings the species count of Pholcus in the Changbai Mountains to 27 species, including 13 new species (
China’s Qinling Mountains is generally regarded as a geographical dividing line between northern China and southern China. It straddles the Provinces of Shanxi, Shaanxi and Henan. To date, 14 species of Pholcus have been found to be recorded in the Qinling Mountains (
Specimens were examined and measured with a Leica M205 C stereomicroscope. Left male pedipalps were photographed. Epigynes were photographed before dissection. Vulvae were illustrated after treating them in a 10% warm solution of potassium hydroxide (KOH) to dissolve soft tissues. Images were captured with a Canon EOS 750D wide zoom digital camera (24.2 megapixels) mounted on the stereomicroscope mentioned above and assembled using Helicon Focus v. 3.10.3 image stacking software (
Terminology and taxonomic descriptions follow
The mitochondrial gene fragment encoding COI and two nuclear gene fragments encoding H3 and wnt were obtained for 19 samples (Table
Species | Voucher code | GenBank accession number | Collection locality | ||
---|---|---|---|---|---|
COI | H3 | wnt | |||
P. ankang sp. nov. | W265 | PP082941 | PP349964 | PP349983 | China, Shaanxi, Ankang |
W266 | PP082942 | PP349965 | PP349984 | ||
W267 | PP082943 | PP349966 | PP349985 | ||
W268 | PP082944 | PP349967 | PP349986 | ||
W269 | PP082945 | PP349968 | PP349987 | ||
P. baoji sp. nov. | W270 | PP082946 | PP349969 | PP349988 | China, Shaanxi, Baoji |
W271 | PP082947 | PP349970 | PP349989 | ||
W272 | PP082948 | PP349971 | PP349990 | ||
W273 | PP082949 | PP349972 | PP349991 | ||
W274 | PP082950 | PP349973 | PP349992 | ||
P. ovatus | W220 | PP082951 | PP349955 | PP349974 | China, Shaanxi, Xi’an |
W221 | PP082952 | PP349956 | PP349975 | ||
W222 | PP082953 | PP349957 | PP349976 | ||
W223 | PP082954 | PP349958 | PP349977 | ||
P. taibaiensis | W224 | PP082955 | PP349959 | PP349978 | China, Shaanxi, Baoji |
W225 | PP082956 | PP349960 | PP349979 | ||
W226 | PP082957 | PP349961 | PP349980 | ||
W227 | PP082958 | PP349962 | PP349981 | ||
W228 | PP082959 | PP349963 | PP349982 | ||
P. paralinzhou | y046 | MW721825 | ON375203 | ON375294 | China, Henan, Jiaozuo |
P. taishan | y133 | MW721826 | ON375204 | ON375293 | China, Shandong, Taian |
Gene | Primer | F/R | Sequence 5’–3’ | Reference |
---|---|---|---|---|
COI | LCO1490 | F | GGTCAACAAATCATAAAGATATTGG |
|
C1-N-2776 | R | GGATAATCAGAATANCGNCGAGG |
|
|
H3 | H3af | F | ATGGCTCGTACCAAGCAGACVGC |
|
H3ar | R | ATATCCTTRGGCATRATRGTGAC | ||
wnt | Spwgf1 | F | GYAAATGCCAYGGWATGTCMGG |
|
Spwgr1 | R | ACTTGRCAACACCARTGAAAWG | ||
Wnt2f | F | CAGTGRAATGTRCARTTG | ||
Wnt2r | R | CNGTTCAAACTTGYTGGATG |
We applied four methods for molecular species delimitation. The Automatic Barcode Gap Discovery (ABGD) analyses were conducted using both Jukes–Cantor and Kimura 2-P distance matrices with options: Pmin = 0.001, Pmax = 0.1, Steps = 10, X = 1.0, Nb bins = 20 (
We obtained a concatenated alignment of 1767 bp (COI, 1184 bp; H3, 293 bp; wnt, 290 bp). Separate phylogenetic analyses of the individual gene COI and concatenated data found compatible topologies. Fig.
The average uncorrected p-distances (below diagonal) and K2P distances (above diagonal) amongst the species and the maximum p-distances (on diagonal) within each species.
P. ankang sp. nov. | P. baoji sp. nov. | P. ovatus | P. taibaiensis | |
---|---|---|---|---|
P. ankang sp. nov. | 0.001 | 0.099 | 0.102 | 0.119 |
P. baoji sp. nov. | 0.092 | 0 | 0.096 | 0.102 |
P. ovatus | 0.094 | 0.090 | 0.001 | 0.072 |
P. taibaiensis | 0.109 | 0.095 | 0.068 | 0.001 |
Family Pholcidae C.L. Koch, 1850
Subfamily Pholcinae C.L. Koch, 1850
Aranea phalangioides Fuesslin, 1775
Pholcus yichengicus species group
Diagnosis and description. See
Note that males and females must be present for this key to work.
1 | Sclerotised prolatero-subdistal apophysis of procursus prolatero-proximally strongly widened (figs 134C, 137A in |
P. ovatus |
– | Sclerotised prolatero-subdistal apophysis of procursus not widened prolatero-proximally (e.g. fig. 169C in |
2 |
2 | Raised prolatero-subdistal membranous edge of procursus laterally rectangular (Fig. |
P. baoji sp. nov. |
– | Raised prolatero-subdistal membranous edge of procursus laterally angular; epigynal plate posteriorly strongly curved (e.g. Fig. |
3 |
3 | Sclerotised prolatero-subdistal apophysis of procursus not protruding latero-distally (arrow 4 in Fig. |
P. ankang sp. nov. |
– | Sclerotised prolatero-subdistal apophysis of procursus latero-distally protruding (fig. 169D in |
P. taibaiensis |
Holotype ♂ (SYNU-Ar00398) and Paratypes 2♂ (SYNU-Ar00399–400) 2♀ (SYNU-Ar00401–02), China, Shaanxi, Ankang, Shiquan County, Chengguan Town, Guiguling Scenic Spot (33°12.32'N, 108°20.15'E, 1585 m elev.), 24 July 2022, Z. Yao, L. Yang & L. Zhang leg.
The specific name refers to the type locality and is a noun in apposition.
The species resembles P. baoji sp. nov. with similar male chelicerae and uncus (Fig.
Pholcus ankang sp. nov., holotype male A, B. Pedipalp (A. Prolateral view; B. Retrolateral view); C, D. Distal part of procursus (C. Prolateral view, arrow 1 indicates distal membranous process, arrow 2 indicates sclerotised prolatero-subdistal apophysis; D. Dorsal view, arrows 1, 2 indicate dorsal spines, arrow 3 indicates angular part of raised prolatero-subdistal membranous edge, arrow 4 indicates latero-distal part of sclerotised prolatero-subdistal apophysis). Abbreviations: a = appendix, b = bulb, e = embolus, pr = procursus, u = uncus. Scale bars: 0.20 mm (A, B); 0.10 mm (C, D).
Pholcus ankang sp. nov., holotype male (C–F) and paratype female (A, B, G, H) A. Epigyne, ventral view; B. Vulva, dorsal view; C. Bulbal apophyses, prolateral view, arrow 1 indicates latero-median protrusion, arrow 2 indicates angular median branch; D. Chelicerae, frontal view; E–H. Habitus (E, G. Dorsal view; F. Lateral view; H. Ventral view). Abbreviations: a = appendix, b = bulb, da = distal apophysis, e = embolus, fa = frontal apophysis, pa = proximo-lateral apophysis, pp = pore plate, u = uncus. Scale bars: 0.20 mm (A–D); 1.00 mm (E–H).
Male (holotype): Total length 5.17 (5.26 with clypeus), carapace 1.53 long, 1.62 wide, opisthosoma 3.64 long, 1.58 wide. Leg I: 39.46 (10.13, 0.66, 9.94, 16.54, 2.19), leg II: 26.96 (7.50, 0.63, 6.79, 10.57, 1.47), leg III: 18.30 (5.38, 0.60, 4.45, 6.79, 1.08), leg IV: 24.35 (7.24, 0.61, 6.03, 9.29, 1.18); tibia I L/d: 70. Eye interdistances and diameters: PME-PME 0.25, PME 0.19, PME-ALE 0.06, AME-AME 0.06, AME 0.15. Sternum width/length: 1.12/0.94. Habitus as in Fig.
Female (paratype, SYNU-Ar00401): Similar to male, habitus as in Fig.
Tibia I in two paratype males (SYNU-Ar00399–400): 9.04, 9.36. Tibia I in another paratype female (SYNU-Ar00402): 7.12.
The species was found on the underside of an overhang on rocky cliffs.
China (Shaanxi, type locality; Fig.
Holotype ♂ (SYNU-Ar00403) and Paratypes 1♂ (SYNU-Ar00404) 2♀ (SYNU-Ar00405–06), China, Shaanxi, Baoji, Long County, Xinjichuan Town, Longmendong Scenic Spot (35°2.33'N, 106°40.22'E, 1489 m elev.), 29 July 2022, Z. Yao, L. Yang & L. Zhang leg.
The specific name refers to the type locality and is a noun in apposition.
The species resembles P. ankang sp. nov. with similar male chelicerae and uncus (Fig.
Pholcus baoji sp. nov., holotype male A, B. Pedipalp (A. Prolateral view; B. Retrolateral view); C, D. Distal part of procursus (C. Prolateral view, arrow 1 indicates distal membranous process, arrow 2 indicates sclerotised prolatero-subdistal apophysis; D. Dorsal view, arrows 1, 2 indicate dorsal spines, arrow 3 indicates rectangular part of raised prolatero-subdistal membranous edge, arrow 4 indicates latero-distal part of sclerotised prolatero-subdistal apophysis). Abbreviations: a = appendix, b = bulb, e = embolus, pr = procursus, u = uncus. Scale bars: 0.20 mm (A, B); 0.10 mm (C, D).
Pholcus baoji sp. nov., holotype male (C–F) and paratype female (A, B, G, H) A. Epigyne, ventral view; B. Vulva, dorsal view; C. Bulbal apophyses, prolateral view, arrow 1 indicates latero-median protrusion, arrow 2 indicates slender median branch; D. Chelicerae, frontal view; E–H. Habitus (E, G. Dorsal view; F. Lateral view; H. Ventral view). Abbreviations: a = appendix, b = bulb, da = distal apophysis, e = embolus, fa = frontal apophysis, pa = proximo-lateral apophysis, pp = pore plate, u = uncus. Scale bars: 0.20 mm (A–D); 1.00 mm (E–H).
Male (holotype): Total length 4.96 (5.05 with clypeus), carapace 1.48 long, 1.78 wide, opisthosoma 3.48 long, 1.52 wide. Leg I: 42.86 (10.77, 0.75, 11.15, 17.56, 2.63), leg II: 28.13 (7.82, 0.70, 6.98, 10.96, 1.67), leg III missing, leg IV: 25.71 (7.69, 0.65, 6.28, 9.62, 1.47); tibia I L/d: 70. Eye interdistances and diameters: PME-PME 0.26, PME 0.16, PME-ALE 0.06, AME-AME 0.05, AME 0.12. Sternum width/length: 1.08/0.93. Habitus as in Fig.
Female (paratype, SYNU-Ar00405): Similar to male, habitus as in Fig.
Tibia I in paratype male (SYNU-Ar00404): 10.44. Tibia I in another paratype female (SYNU-Ar00406): 7.18.
The species was found on the underside of an overhang on rocky cliffs.
China (Shaanxi, type locality; Fig.
Pholcus ovatus
1♂ (SYNU-Ar00120F) 1♀ (SYNU-Ar00121F), China, Shaanxi, Xi’an, Zhouzhi County, Banfangzi Town (type locality), roadside of G108 (33°48.02'N, 107°59.08'E, 1165 m elev.), 31 July 2022, Z. Yao, L. Yang & L. Zhang leg.
The species resembles P. taibaiensis Wang & Zhu, 1992 (
The species was found on the underside of an overhang on rock cliffs.
China (Shaanxi, Fig.
Pholcus taibaiensis
3♂ (SYNU-Ar00130F–Ar00132F) 3♀ (SYNU-Ar00133F–Ar00135F), China, Shaanxi, Baoji, Mei County, Yingtou Town, near Haopingsi Temple (type locality) (34°5.32'N, 107°42.33'E, 1101 m elev.), 30 July 2022, Z. Yao, L. Yang & L. Zhang leg.
The species resembles P. ovatus Yao & Li, 2012 (
The species was found on the underside of an overhang on rocky cliffs.
China (Shaanxi, Fig.
The manuscript benefits greatly from comments by Danilo Harms, Bernhard Huber, Yanfeng Tong and an anonymous reviewer. Joseph KH Koh checked the English. This study was supported by the Science & Technology Fundamental Resources Investigation Program of China (2023FY100200) and the National Natural Science Foundation of China (NSFC-32170461, 31872193).