Research Article |
Corresponding author: Xiaoqi Mi ( mixiaoqi1018@163.com ) Academic editor: Christine Rollard
© 2021 Cheng Wang, Jiahui Gan, Xiaoqi Mi.
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:
Wang C, Gan J, Mi X (2021) On four species of the genus Argiope Audouin, 1826 (Araneae, Araneidae) from China. ZooKeys 1019: 15-34. https://doi.org/10.3897/zookeys.1019.59521
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Based on morphological and molecular evidence, Argiope macrochoera Thorell, 1891 from China is found to be the unknown female of A. cameloides Zhu & Song, 1994, the known male of A. perforata Schenkel, 1963 is mismatched and provisionally suggested to be the male of A. boesenbergi Levi, 1983, and the true male of A. perforata Schenkel, 1963 is described for the first time. Argiope abramovi Logunov & Jäger, 2015 is suggested to be a synonym of A. perforata Schenkel, 1963. Argiope chloreides Chrysanthus, 1961 and A. vietnamensis Ono, 2010 are newly recorded from China. The unknown male of A. vietnamensis Ono, 2010 is described for the first time.
DNA barcoding, orb-weaving spider, sexual dimorphism, taxonomy
Argiope Audouin, 1826 is nested in the subfamily Argiopinae Simon, 1890 together with the genera Gea C.L. Koch, 1843 and Neogea Levi, 1983, and it comprises sexually dimorphic species well known for their showy, colorful females and their unique web stabilimenta (
While examining Argiope specimens from southwest China, A. chloreides Chrysanthus, 1961, A. vietnamensis Ono, 2010, and the previously unknown male of the latter were recognized. An extended study of morphological and molecular evidence has revealed that A. macrochoera Thorell, 1891 from China is misidentified and conspecific with A. cameloides Zhu & Song, 1994, and that the male of A. perforata Schenkel, 1963 is mismatched and may be A. boesenbergi Levi, 1983. Moreover, the true male of A. perforata is revealed for the first time, and A. abramovi Logunov & Jäger, 2015 is synonymized with A. perforata Schenkel, 1963. The goals of the present paper are to revise and describe the misidentifications, mismatches, and the unknown sexes of A. cameloides, A. perforata, and A. vietnamensis, as well as to provide a distributional map of those species.
All specimens were collected by beating shrubs or hand collecting and were preserved in 75% ethanol, except for the specimens for DNA extraction which were preserved in absolute ethyl alcohol. All specimens were deposited in the Museum of Tongren University, China (TRU).
The specimens were examined with an Olympus SZ51 stereomicroscope. After dissection, the epigyna were cleared in a trypsin enzyme solution before examination and imaging. Left male palps were used for the descriptions and illustrations. Photographs of the copulatory organs and habitus were taken with a Kuy Nice CCD or an Olympus C7070 camera mounted on an Olympus BX51 compound microscope. Compound focus images were generated using Helicon Focus v. 6.7.1.
All measurements are given in millimeters. Leg measurements are given as total length (femur, patella + tibia, metatarsus, tarsus). Abbreviations used in the text and figures are as follows:
C conductor;
CD copulatory duct;
CO copulatory opening;
E embolus;
F flange;
FD fertilization duct;
MA median apophysis;
PC paracymbium;
Pd pendant;
PP posterior plate;
S spermatheca;
Sc scape;
Sep septum;
Sp spur of the median apophysis.
The total genomic DNA from spider legs was extracted using the Animal Genomic DNA Isolation Kit (Kangwei Biotech, Beijing, China) following the manufacturer’s protocols. The primer pair LCO1490/HCO2198 (
Argiope cameloides
Zhu & Song in
A. macrochoera
Yin et al., 1989: 63, fig. 3A–C (♀, the specimens from Guangdong Province of China, locality unspecified and deposited in Hunan Normal University, not examined);
China – Guangxi Zhuang Autonomous Region • 4♂1♀ (TRU-Araneidae-31–35), Beihai City, Yinhai District, Yajishan Forestry Station (21°35.37'N, 109°18.41'E, ca 30 m), 12.viii.2017, Xiaoqi Mi et al. leg. • 1♀ (TRU-Araneidae-36), same locality, night of 03.x.2018, Xiaoqi Mi et al. leg. • 2♂ (TRU-Araneidae-37–38), Beihai City, Tieshangang District, Xinggang Township, Xiaomatou Village, Caobiaotang (21°33.11'N, 109°29.22'E, ca 10 m), 04.x.2018, Xiaoqi Mi et al. leg. • 2♀ (TRU-Araneidae-39–40), Fangchenggang City, Shangsi County, Wenlingshan Park, night of 06.x.2018, Xiaoqi Mi et al. leg.
The male of this species resembles A. dang Jäger & Praxaysombath, 2009 in having a similarly shaped median apophysis and broad, flat conductor, but it differs in: 1) the embolus is almost directed towards 6 o’clock apically in prolateral view (Fig.
Male (TRU-Araneidae-37). Total length 2.62. Carapace 1.46 long, 1.34 wide; abdomen 1.38 long, 1.02 wide. Eye sizes and interdistances: AME 0.10, ALE 0.06, PME 0.11, PLE 0.09, AME–AME 0.13, AME–ALE 0.05, PME–PME 0.14, PME–PLE 0.16. Legs: I 5.92 (1.68, 1.85, 1.63, 0.76), II 5.78 (1.63, 1.85, 1.54, 0.76), III 3.22 (1.02, 0.93, 0.76, 0.51), IV 4.65 (1.46, 1.39, 1.24, 0.56). Carapace (Fig.
Palp (Fig.
Female (TRU-Araneidae-40). Total length 6.78. Carapace 2.72 long, 2.28 wide; abdomen 4.38 long, 3.22 wide. Eye sizes and interdistances: AME 0.18, ALE 0.09, PME 0.19, PLE 0.16, AME–AME 0.14, AME–ALE 0.15, PME–PME 0.24, PME–PLE 0.28. Legs: I 13.39 (3.88, 4.25, 3.88, 1.38), II 13.14 (3.75, 4.13, 3.88, 1.38), III 7.77 (2.63, 2.38, 1.75, 1.01), IV 12.39 (4.13, 3.63, 3.38, 1.25). Carapace (Fig.
China (Hainan, Guangdong, and Guangxi).
Argiope cameloides is known only from the descriptions of the holotype from Hainan, China (see
Argiope chloreides
Chrysanthus 1961: 197, figs 5–8 (♀, female types from Indonesia, locality the environs of Mindiptana and deposited in Rijksmuseum van Natuurlijke Historie, Leiden, not examined);
Argiope chloreis
China – Guangxi Zhuang Autonomous Region • 1♂ (TRU-Araneidae-41), Chongzuo City, Longzhou County, Zhubu Township, Nonggang Village (22°27.84'N, 106°56.52'E, ca 170 m), 06.vii.2019, Cheng Wang et al. leg.
Well described and illustrated by
China (Guangxi); Laos, Malaysia, Indonesia.
Argiope perforata
Schenkel 1963: 135, fig. 79a, b (♀, female holotype from China locality Lunan fu City, Szetchuan Province and deposited in the Museum National d'Histoire Naturelle, Paris, not examined);
Argiope abramovi
China – Guizhou Province • 1♂ (TRU-Araneidae-42), Tongren City, Bijiang District, Wenbi Park (27°43.26'N, 109°10.03'E, ca 460 m), 19.v.2013, Xiaoqi Mi et al. leg. • 1♂ (TRU-Araneidae-43), same locality, 10.vi.2017, Cheng Wang leg. • 3♂ (TRU-Araneidae-44–46), Tongren City, Jiangkou County, Dewang Township, Baxi Village, Nanmuping (27°51.68'N, 108°36.88'E, ca 900 m), 10.vi.2013, Xiaoqi Mi et al. leg. • 5♀4♂ (TRU-Araneidae-47–55), same locality, 14.vi.2013, Xiaoqi Mi et al. leg. • 6♀ (TRU-Araneidae-56–61), same locality, 15–16.vii.2015, Cheng Wang and Mingyong Liao leg. • 2♂ (TRU-Araneidae-62–63), Tongren City, Jiangkou County, Dewang Township, Baxi Village, Datuzu, 10.vi.2013, Xiaoqi Mi et al. leg. • 1♀ (TRU-Araneidae-64), Tongren City, Yinjiang County, Muhuang Township, Jinchang Village (28°01.37'N, 108°45.00'E, ca 1300 m), 14.vii.2013, Xiaoqi Mi et al. leg. • 1♀ (TRU-Araneidae-65), Tongren City, Songtao County, Wuluo Township, Lengjiaba Village (27°54.93'N, 108°36.70'E, ca 1150 m), 15.vii.2013, Xiaoqi Mi et al. leg. • 3♀ (TRU-Araneidae-66–68), same locality, 10.vii.2015, Cheng Wang and Mingyong Liao leg. • 3♂ (TRU-Araneidae-69–71), Qiannan Buyi and Miao Autonomous Prefecture, Libo County, Dongtang Township, Yaosuo Village, Bizuo, Maolan National Nature Reserve (25°16.37'N, 108°02.97'E, ca 550 m), 07–10.viii.2013, Xiaoqi Mi et al. leg. • 1♀ (TRU-Araneidae-72), Tongren City, Jiangkou County, near the Kaima primary school (27°50.77'N, 108°46.43'E, ca 530 m), 07.vii.2015, Cheng Wang and Mingyong Liao leg. • 2♀ (TRU-Araneidae-73–74), Tongren City, Jiangkou County, Taiping Township, Kuaichang Village, Macaohe (27°49.08'N, 108°51.52'E, ca 680 m), 08.vii.2015, Cheng Wang and Mingyong Liao leg. • 1♀ (TRU-Araneidae-75), Tongren City, Yinjiang County, Ziwei Township, Zhangjiaba Village (27°56.62'N, 108°36.60'E, ca 780 m), 12.vii.2015, Cheng Wang and Mingyong Liao leg. • 1♀ (TRU-Araneidae-76), Tongren City, Yinjiang County, Ziwei Township, Tuanlong Village (27°54.93'N, 108°42.70'E, ca 1150 m), 14.vii.2015, Cheng Wang et al. leg. • 3♀1♂ (TRU-Araneidae-77–80), Tongren City, Jiangkou County, Dengwang Township, Jinghe Village, Xujiagou (27°48.29'N, 108°37.45'E, ca 820 m), 20.vii.2015, Cheng Wang et al. leg. • 2♀ (TRU-Araneidae-81–82), Qiandongnan Miao and Dong Autonomous Prefecture, Shibing County, Yuntaishan Scenic Area (27°07.74'N, 108°06.57'E, ca 990 m), 30.vii.2015, Cheng Wang et al. leg. • 4♀1♂ (TRU-Araneidae-83–87), Qiannan Buyi and Miao Autonomous Prefecture, Libo County, Maolan National Nature Reserve (25°16.07'N, 108°59.07'E, ca 760 m), 30.iv.2016, Xiaoqi Mi et al. leg. • 1♀ (TRU-Araneidae-88), Zunyi City, Xishui County, Sanchahe Township, Sanchahe Village (28°29.04'N, 106°25.28'E, ca 800 m), 29.vii.2016, Cheng Wang et al. leg. • 4♂ (TRU-Araneidae-89–92), Zunyi City, Xishui County, Sanchahe Township, Sanchahe Village, Tiantangba (28°26.37'N, 106°24.96'E, ca 1000 m), 30.vii.2016, Cheng Wang et al. leg. • 2♀ (TRU-Araneidae-93–94), Tongren City, Bijiang District, Tianshengqiao Scenic Area (27°49.80'N, 109°12.77'E, ca 580 m), 12.v.2018, Xiaoqi Mi et al. leg. • 1♀1♂ (TRU-Araneidae-95–96), Tongren City, Shiqian County, Pingshan Township, Daping Village (27°17.97'N, 108°10.05'E, ca 720 m), 08.vi.2019, Cheng Wang et al. leg. • 1♂ (TRU-Araneidae-97), Tongren City, Songtao County, Wuluo Township, Taohuayuan Village (27°58.04'N, 108°47.73'E, ca 790 m), 29.v.2017, Xiaoqi Mi et al. leg. • 3♀ (TRU-Araneidae-98–100), Qiandongnan Miao and Dong Autonomous Prefecture, Leishan County, Fangxiang Township, Getou Village (26°24.09'N, 108°15.40'E, ca 1100 m), night of 22.vii.2017, Cheng Wang et al. leg. • 1♂ (TRU-Araneidae-101), Guiyang City, Guanshanhu District, Donglinsi Park (26°39.60'N, 106°38.00'E, ca 1300 m), 23.v.2018, Cheng Wang leg. • 1♂ (TRU-Araneidae-102), Qiandongnan Miao and Dong Autonomous Prefecture, Zhenyuan County, Dadi Township, Dadi Village (27°21.56'N, 108°12.29'E, ca 700 m), night of 23.v.2018, Xiaoqi Mi leg. • 3♀ (TRU-Araneidae-103–105), Tongren City, Shiqian County, Pingshan Township, Fodingshan Village, Yaoshang (27°20.54'N, 108°09.50'E, ca 640 m), night of 24.v.2018, Xiaoqi Mi et al. leg. • 3♀ (TRU-Araneidae-106–108), Tongren City, Yinjiang County, Yangxi Township (27°38.53'N, 108°26.43'E, ca 700 m), 17.vi.2018, Xiaoqi Mi et al. leg. • 1♂ (TRU-Araneidae-109), Tongren City, Shiqian County, Ganxi Township, near the 524 road (27°25.10'N, 108°07.97'E, ca 550 m), night of 06.vi.2019, Cheng Wang et al. leg. • 2♀ (TRU-Araneidae-110–111), Shibing County, Baiduo Township, Heichong (27°9.37'N, 108°07.40'E, ca 990 m), 20.vi.2019, Xiaoqi Mi et al. leg. • 1♀(TRU-Araneidae-112), Qiandongnan Miao and Dong Autonomous Prefecture, Shibing County, Chengguan Township (27°03.02'N, 108°7.78'E, ca 700 m), night of 21.vi.2019, Xiaoqi Mi et al. leg.; China – Guangxi Zhuang Autonomous Region • 1♀2♂ (TRU-Araneidae-113–115), Fangchenggang City, Shangsi County, Shiwandashan National Forest Park (21°53.87'N, 107°54.26'E, ca 370 m), 14.viii.2017, Xiaoqi Mi et al. leg.; China – Hainan Province – Ledong County – Jianfeng Township – Jianfengling National Nature Reserve • 1♀ (TRU-Araneidae-116), Mingfeng Valley (18°44.61'N, 108°51.24'E, ca 810 m), night of 12.iv.2019, Cheng Wang and Yuanfa Yang leg. • 1♀ (TRU-Araneidae-117), Sanfenqu (18°45.24'N, 108°51.57'E, ca 900 m), night of 13.iv.2019, Cheng Wang and Yuanfa Yang leg. • 1♂ (TRU-Araneidae-118), Tianchi (18°45.22'N, 108°51.53'E, ca 850 m), 14.iv.2019, Cheng Wang & Yuanfa Yang leg. • 2♀1♂ (TRU-Araneidae-119–121), Sanfenqu (18°45.24'N, 108°51.57'E, ca 900 m), night of 14.iv.2019, Cheng Wang and Yuanfa Yang leg. • 2♀ (TRU-Araneidae-122–123), Yulin Valley, Zijin Waterfull (18°44.79'N, 108°55.76'E, ca 630 m), 15.iv.2019, Cheng Wang and Yuanfa Yang leg. • 1♂ (TRU-Araneidae-124), Tianchi (18°45.22'N, 108°51.53'E, ca 850 m), night of 15.iv.2019, Cheng Wang and Yuanfa Yang leg.
The male of this species resembles A. aetheroides Yin, Wang, Zhang, Peng & Chen, 1989 in the general shape of the palp, but it differs in the distal end of embolus is slender (Fig.
Male (TRU-Araneidae-118). Total length 3.65. Carapace 1.98 long, 1.90 wide; abdomen 1.90 long, 1.37 wide. Eye sizes and interdistances: AME 0.14, ALE 0.07, PME 0.15, PLE 0.12, AME–AME 0.13, AME–ALE 0.07, PME–PME 0.18, PME–PLE 0.16. Legs: I 9.59 (2.67, 3.01, 2.81, 1.10), II 9.39 (2.67, 2.95, 2.67, 1.10), III 5.05 (1.67, 1.48, 1.14, 0.76), IV 7.59 (2.48, 2.24, 2.01, 0.86). Carapace (Fig.
Palp (Fig.
Female (TRU-Araneidae-122). Total length 9.86. Carapace 4.67 long, 4.14 wide; abdomen 5.76 long, 4.48 wide. Eye sizes and interdistances: AME 0.26, ALE 0.15, PME 0.27, PLE 0.26, AME–AME 0.25, AME–ALE 0.34, PME–PME 0.49, PME–PLE 0.51. Legs: I 16.77 (5.13, 5.38, 4.63, 1.63), II 16.51 (5.13, 5.25, 4.50, 1.63), III 9.32 (3.25, 2.69, 2.25, 1.13), IV 14.76 (5.25, 4.38, 3.88, 1.25). Carapace (Fig.
China (Hainan, Guangxi).
Argiope perforata was originally described based on the female holotype from Sichuan, China; the illustrations are poor. It was redescribed and better illustrated by
Argiope vietnamensis
China – Guangxi Zhuang Autonomous Region • 1♂, 1♀ (subadult) (TRU-Araneidae-125–126), Fangchengang City, Shangsi County, Shiwandashan National Forest Park (21°53.87'N, 107°54.26'E, ca 370 m), night of 14.viii.2017, Xiaoqi Mi et al. leg.; 1♀ (TRU-Araneidae-127), Chongzuo City, Jiangzhou District, Zuozhou Township, Guanghe Village (22°34.72'N, 107°24.94'E, ca 160 m), night of 03.vii.2019, Cheng Wang et al. leg.; 1♀ (TRU-Araneidae-128), Chongzuo City, Longzhou County, Kouke Village (22°18.18'N, 106°42.23'E, ca 960 m), 08.vii.2019, Cheng Wang et al. leg.; China – Guizhou Province – Qiannan Buyi and Miao Autonomous Prefecture – Libo County • 1♀ (TRU-Araneidae-129), Xiaoqikong Scenic Area (25°15.28'N, 107°42.96'E, ca 580 m), 08.vii.2017, Xiaoqi Mi et al. leg. • 5♀ (TRU-Araneidae-130–134), Dongtang Township, Yaosuo Village, Maolan National Nature Reserve (25°16.37'N, 108°02.97'E, ca 550 m), 07–10.viii.2013, Xiaoqi Mi et al. leg. • 1♂ (TRU-Araneidae-135), Weng’ang Township, Jilong Village, Maolan National Nature Reserve (25°13.53'N, 107°56.18'E, ca 840 m), 10–11.viii.2013, Xiaoqi Mi et al. leg.
The female was clearly diagnosed by
Female described and illustrated by
Male (TRU-Araneidae-125). Total length 5.21. Carapace 3.10 long, 2.88 wide; abdomen 2.37 long, 1.93 wide. Eye sizes and interdistances: AME 0.23, ALE 0.12, PME 0.24, PLE 0.21, AME–AME 0.23, AME–ALE 0.14, PME–PME 0.24, PME–PLE 0.22. Legs were used for DNA extraction. Carapace (Fig.
Palp (Fig.
China (Guizhou, Guangxi); Vietnam.
A male and a subadult female were collected on the same web from Shiwandashan National Forest Park during the night. The embolus of the male is structurally consistent with the broken embolus found in an A. vietnamensis female epigyne. Moreover, other male and females were also collected from Maolan National Nature Reserve. Based on these points of evidence, we propose the males as A. vietnamensis.
The manuscript benefited greatly from comments by Christine Rollard (Paris, France), Marc Milne (Indianapolis, USA), Ivan L. F. Magalhaes (Buenos Aires, Argentina). We are grateful to Bo Yu, Mingyong Liao, Yuanfa Yang, Hong Liu, Guijie Tian, Chaoguan Qin, and Siyi Yan for collecting the specimens. This research was supported by the National Natural Sciences Foundation of China (NSFC-31660609, 31301861) and the Science and Technology Project Foundation ([2020]1Z014) and Key Laboratory Project ([2020]2003) of Guizhou Province.