Research Article |
Corresponding author: Yu Peng ( pengyu@hubu.edu.cn ) Corresponding author: Jie Liu ( sparassidae@aliyun.com ) Academic editor: Dimitar Dimitrov
© 2021 Zichang Li, Ingi Agnarsson, Yu Peng, Jie Liu.
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:
Li Z, Agnarsson I, Peng Y, Liu J (2021) Eight cobweb spider species from China building detritus-based, bell-shaped retreats (Araneae, Theridiidae). ZooKeys 1055: 95-121. https://doi.org/10.3897/zookeys.1055.67620
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Eight cobweb spider species building a detritus-based, bell-shaped retreat from China are reported in the current paper, including five new Campanicola species and three known species: Campanicola anguilliformis Li & Liu, sp. nov., Campanicola falciformis Li & Liu, sp. nov., Campanicola heteroidea Li & Liu, sp. nov., Campanicola tauricornis Li & Liu, sp. nov., Campanicola volubilis Li & Liu, sp. nov., Campanicola campanulata (Chen, 1993), Campanicola ferrumequina (Bösenberg & Strand, 1906), and Parasteatoda ducta (Zhu, 1998). Among them, the male of Parasteatoda ducta (Zhu, 1998) is described for the first time. We provide photographs of all species and descriptions for new species in the current paper. The type of bell-shaped retreat is rare in theridiid, and found only in four related genera. A natural next step upon completing this taxonomic study would be to analyse and understand the evolution of the retreat and related traits.
Bell-shaped retreat, Campanicola, new species, taxonomy, theridiid spiders, trash-decorating behaviour
Orb-weaving spiders often employ a variety of ‘trash’ materials such as soil particles, plant detritus, prey remains and egg sacs into webs, which can effectively deceive predators, thereby increasing their chances of survival (
In the past three years, a series of surveys on Chinese theridiid spiders were conducted by colleagues of Hubei University in China and yielded numerous new species. This is our first paper on Chinese cobweb spiders with the aim to describe the species building bell-shaped retreats including seven Campanicola species and one Parasteatoda species.
All specimens were kept in 100% ethanol and examined with an Olympus SZX16 stereomicroscope. Further details were studied under an Olympus BX51 compound microscope. Male and female genitalia were examined and illustrated after dissection. Epigynes were cleaned by Proteinase K solution. Male left palps and female epigynes were imaged with an Olympus BX51 compound microscope in ethanol and in Arabic gum. For SEM, specimens were treated according to
ALE anterior lateral eyes
AME anterior median eyes
C conductor
CD copulatory duct
Chd cymbial hood
CP copulatory pore
Cy cymbium
E embolus
FD fertilization duct
PLE posterior lateral eyes
PME posterior median eyes
S spermathecae
ST subtegulum
T tegulum
Ti tibia
I, II, III, IV legs I to IV
Campanicola is similar to some Achaearanea, Cryptachaea, Parasteatoda in building a detritus-based and bell-shaped retreat, but can be distinguished morphologically from Achaearanea by the cymbium with cymbial hood and without a distal projection in Campanicola, but with a cymbial hook and large distal projections in Achaearanea; from Cryptachaea by the cymbium not extending beyond the alveolus, the tegulum depressed and the copulatory ducts long in Campanicola, but the cymbium extending beyond alveolus, the tegulum spherical and the copulatory ducts short in Cryptachaea; from Parasteatoda by the conductor tip curved dorsally, the embolus and the copulatory ducts thin and the atrium small in Campanicola, but the conductor tip curved ventrally, the embolus and the copulatory ducts thick, the atrium large in Parasteatoda (
Achaearanea campanulata Chen, 1993: 36, f. 1–5 (description of male and female).
Achaearanea campanulata:
Parasteatoda campanulata:
Campanicola campanulata:
China, Guizhou Province: 1 ♂, 13 ♀, Suiyang County, Wenquan Town, Shuanghe Village, Shanwang Cave (27°57'56"N, 107°9'26"E, 756 m), 8–10 May 2018, F.X. Liu, W. Ding and Z.C. Li leg.; 1 ♂, 3 ♀, Suiyang County, Wenquan Town, Dishui Village, Manwang Cave and village road (28°14'30"N, 107°0'56"E, 650 m), 11 May 2018, F.X. Liu, W. Ding and Z.C. Li leg.; 1 ♂, 6 ♀, Fuquan City, Kouhuang Cave (26°33'31"N, 107°12'55"E, 1030 m), 13 May 2018, F.X. Liu, W. Ding and Z.C. Li leg.; Hunan Province: 15 ♀, Sangzhi County, Tianpingshan Forest Farm (29°47'N, 110°5'12"E, 1359 m), 1–3 June 2018, F.X. Liu, J.S. Lu, J.C. Zhang, R. Zhong and Z.C. Li leg.; 11 ♀, Sangzhi County, Meijiashan Park (29°24'24"N, 110°9'28"E, 336 m), 4 June 2018, F.X. Liu, J.S. Lu, J.C. Zhang, R. Zhong and Z.C. Li leg.; 2 ♀, Zhangjiajie City, Zhangjiajie National Forest Park, footpath of The Yellow Rock Village and The Golden Whip Brook (29°19'22"N, 110°25'38"E, 452–780 m), 6 June 2018, F.X. Liu, J.S. Lu, J.C. Zhang, R. Zhong and Z.C. Li leg.; 1 ♀, Changsha City, Yuelu Mountain (28°11'33"N, 112°56'6"E, 210 m), 12 August 2018, Z.W. Deng and Z.C. Li leg.; 1 ♀, Hengyang City, Hengshan scenic spot, Fanyin Valley (27°16'22"N, 112°42'41"E, 410 m), 17 August 2018, Z.W. Deng and Z.C. Li leg.; Sichuan Province: 8 ♀, Emeishan City, Emei Mountain Scenic Spot, footpath from Wuxian Gang to Wannian Temple (29°35'3"N, 103°22'55"E, 940 m), 21 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; 17 ♀, Leshan City, Leshan Giant Buddha Scenic Spot (29°32'28"N, 103°46'19"E, 380 m), 24 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; 1 ♀, Ya’an City, Baoxing County, Panda Square (30°22'10"N, 102°48'50"E, 1060 m), 27 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; 5 ♀, Ya’an City, Baoxing County, Longmen Town (30°15'14"N, 103°1'20"E, 810 m), 28 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; Hubei Province: 7 ♀, Lichuan City, Tenglongdong Scenic Spot (30°16'10"N, 108°56'15"E, 1070 m), 5 October 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; 2 ♀, Jianshi County, Chaoyangguan (30°35'45"N, 109°42'52"E, 650 m), F.X. Liu, Z.W. Deng and Z.C. Li leg.
The females of C. campanulata differ from all other Campanicola species by the copulatory pores that are circular and not clinging to the atrium margin, and the copulatory ducts that curve only twice (Fig.
Campanicola campanulata (Chen, 1993) A–C female habitus (A dorsal view B lateral view C ventral view) D–F female epigynum (D ventral view, uncleared E ventral view, cleared and embedded in Arabic gum F dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–F).
See
Theridion ferrum-equinum
Bösenberg & Strand, 1906: 139, pl. 12, f. 261 (description of male and female);
Theridion meum Bösenberg & Strand, 1906: 145, pl. 12, f. 294 (description of female).
Theridion ferrumequinum: Saito 1959: 70, f. 65A–C (female);
Achaearanea ferrumequina:
Parasteatoda ferrumequina:
Campanicola ferrumequina:
China, Hunan Province: 3 ♂, 32 ♀, Zhangjiajie City, Zhangjiajie National Forest Park, footpath of The Yellow Rock Village and The Golden Whip Brook (29°19'22"N, 110°25'38"E, 452–780 m), 6 June 2018, F.X. Liu, J.S. Lu, J.C. Zhang, R. Zhong and Z.C. Li leg.; Fujian Province: 1 ♂, 15 ♀, Wuyishan City, Wuyi Mountain Natural Reserves (26°39'42"N, 117°56'24"E, 399 m), 28–31 August 2019, Y. Zhong and F.J. Liu leg.
The females of C. ferrumequina differ from all other Campanicola species by the almost overlapping copulatory ducts except the short segment starting from the copulatory pores (Fig.
Campanicola ferrumequina (Bösenberg & Strand, 1906) A–C female habitus (A dorsal view B lateral view C ventral view) D–H female epigynum (D ventral view, uncleared E ventral view, cleared F dorsal view, cleared G ventral view, cleared and embedded in Arabic gum H dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–H).
See
Holotype: ♀, China, Fujian Province: Wuyishan City, Wuyi Mountain Natural Reserves (26°39'42"N, 117°56'24"E, 399 m), 28 August 2019, Y. Zhong and F.J. Liu leg. Paratypes: 1 ♀, same data as holotype.
This new species is similar to C. ferrumequina, C. tauricornis sp. nov. and C. volubilis sp. nov. in having spherical spermathecae, and long and looped copulatory ducts, but can be distinguished from them by the direction of the copulatory duct: it starts from the middle and anterior part of the atrium, extends posteriorly, then curves several times and enters the spermathecae laterally and ventrally in C. anguilliformis sp. nov. (Fig.
Campanicola anguilliformis sp. nov. A–C female habitus (A dorsal view B ventral view C lateral view) D–H female epigynum (D ventral view, not dissected E ventral view, cleared F dorsal view, cleared G ventral view, cleared and embedded in Arabic gum H dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–H).
The specific name is derived from the Latin adjective anguilliformis, meaning eel-like, referring to the eel-like copulatory ducts; adjective.
Male unknown. Female (holotype): Total length 2.06. Prosoma 0.91 long, 0.80 wide, brownish yellow, with black margin. Sternum 0.55 long, 0.51 wide, yellow, peltate, with sparse setae. Opisthosoma 1.45 long, 1.16 wide, dorsum black, with two pairs of transverse patterns in the shape of mustache and a longitudinal stripe, irregular white spots distributed in the patterns and the stripe; venter yellowish, with few bright white spots. Anal tubercle yellow. Spinnerets surrounded with black ring, without colulus (Fig.
China (Sichuan) (Fig.
Holotype ♀, China, Sichuan Province: Emeishan City, Emei Mountain Scenic Spot, footpath from Wuxian Gang to Wannian Temple (29°35'3"N, 103°22'55"E, 940 m), 21 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg. Paratypes: 12 ♀, same data as holotype; Sichuan Province: 2 ♀, Leshan City, Leshan Giant Buddha Scenic Spot (29°32'28"N, 103°46'19"E, 380 m), 24 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; 1 ♀, Ya’an City, Baoxing County, Fengtong Stronghold (30°34'21"N, 102°52'59"E, 1540 m), 26 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; 3 ♀, Ya’an City, Baoxing County, Panda Square (30°22'10"N, 102°48'50"E, 1060 m), 27 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.; 1♀, Ya’an City, Baoxing County, Longmen Town (30°15'14"N, 103°1'20"E, 810 m), 28 September 2018, F.X. Liu, Z.W. Deng and Z.C. Li leg.
This new species can be distinguished from other Campanicola species, in the female, by the position of the copulatory pores: located on the lateral and posterior part of atrium in this new species while located anteriorly in other Campanicola species; and the relative position of the copulatory ducts to the spermathecae: partial copulatory ducts overlap the posterior-ventral half of the spermathacea in this new species (Fig.
Campanicola falciformis sp. nov. A–C female habitus (A dorsal view B ventral view C lateral view) D–H female epigynum (D ventral view, uncleared E ventral view, cleared F dorsal view, cleared G ventral view, cleared and embedded in Arabic gum H dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–H).
The specific name is derived from the Latin adjective falciformis, meaning doubly falcate, referring to the falcate copulatory ducts in ventral view; adjective.
Male unknown. Female (holotype): Total length 2.29. Prosoma 0.92 long, 0.79 wide, brown, with dark brown margin. Sternum 0.49 long, 0.47 wide, yellow, peltate, with sparse setae. Opisthosoma 1.45 long, 1.38 wide, yellowish-brown, dorsum center with an angle bracket-shaped pale-yellow stripe either side of the midline, dorsum posterior with lateral pale-yellow stripe, and many bright white spots are distributed in the stripes (Fig.
China (Sichuan) (Fig.
Holotype: ♀, China, Guizhou Province: Liupanshui City, Zhongshan District (26°35'50"N, 104°49'10"E, 1760 m), 26 August 2020, B. Liang and J.H. Wang leg.
This new species can be distinguished from other Campanicola species by the following characteristics: epigynum dissymmetric; copulatory pores located on the anterior margin of the atrium; copulatory ducts short, winding simply (Fig.
Campanicola heteroidea sp. nov. A–C female habitus (A dorsal view B lateral view C ventral view) D–G female epigynum (D ventral view, cleared E dorsal view, cleared F ventral view, cleared and embedded in Arabic gum G dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–G).
The specific name is derived from the Latin adjective heteroideus, meaning asymmetrical, referring in essence to the dissymmetric epigynum structure; adjective.
Male unknown. Female (holotype): Total length 4.12. Prosoma 1.66 long, 1.54 wide, brown, with few black patches in the centre. Sternum 0.90 long, 0.93 wide, yellow, with black patches and sparse setae. Opisthosoma 2.45 long, 2.22 wide, oval, armed with black setae, dorsum yellowish with a middle longitudinal black belt which is surrounded with bright white spots, and more laterally mottled. Venter yellowish-brown, with a broad black transverse stripe next to epigastric furrow. Anal tubercle yellowish. Spinnerets yellow, surrounded with black ring, without colulus (Fig.
China (Guizhou) (Fig.
Holotype: ♀, China, Hainan Province: Limushan National Forest Park (19°10'N, 109°39'E), 20 July 2020, J. Liu leg.
This new species is similar to C. volubilis sp. nov. in having the spherical spermathecae, the long and looped copulatory ducts which start from the lateral and anterior part of the atrium, extend anteriorly, then curve several times and enter the spermathecae laterally and ventrally, but can be distinguished from the latter by the following characteristics: 1. The copulatory pores are slightly separated from each other in this new species (Fig.
Campanicola tauricornis sp. nov. A–C female habitus (A dorsal view B ventral view C lateral view) D–G female epigynum (D ventral view, cleared E dorsal view, cleared F ventral view, cleared and embedded in Arabic gum G dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–G).
The specific name is derived from the Latin adjective tauricornis, meaning ox horn-shaped, referring to the shape of the copulatory ducts in dorsal view; adjective.
Male unknown. Female (holotype): Total length 4.02. Prosoma 1.59 long, 1.52 wide, brownish black. Sternum 0.86 long, 0.95 wide, yellow, with sparse setae. Opisthosoma 2.18 long, 2.16 wide, oval, mottled, armed with sparse black setae, both dorsum and venter with irregular black plaque and white spots. Anal tubercle yellowish. Spinnerets brownish, without colulus (Fig.
China (Hainan) (Fig.
Holotype: ♀, China, Yunnan Province: Xishuangbanna Dai Autonomous Prefecture, Menghai county, Bulang Nationality Township (21°34' 37"N, 100°20'24"E, 1130 m), 29 July 2020, Z.W. Deng, W. Zhang, Y.T. Zhang, R. Zhong and Z.C. Li leg. Paratypes: 2 ♀, same data as holotype.
See the diagnosis under C. tauricornis sp. nov.
The specific name is derived from the Latin adjective volubilis, meaning winding, referring to the shape of the part of the copulatory ducts near the copulatory pores in ventral view; adjective.
Male unknown. Female (holotype): Total length 3.74. Prosoma 1.74 long, 1.44 wide, brownish black. Sternum 0.99 long, 0.88 wide, yellow, some reddish-brown patches on the margin, with sparse setae. Opisthosoma 2.02 long, 1.96 wide, oval, brownish black, with black setae, both dorsum and venter with white spots. Anal tubercle and spinnerets brownish black, without colulus (Fig.
Campanicola volubilis sp. nov. A–C female habitus (A dorsal view B lateral view C ventral view) D–G female epigynum (D ventral view, cleared E dorsal view, cleared F ventral view, cleared and embedded in Arabic gum G dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–G).
China (Yunnan) (Fig.
Achaearanea ducta Zhu, 1998: 107, f. 64A–C (description of female); Song, Zhu and Chen 1999: 90, f. 39E, F (female).
Parasteatoda ducta:
China, Hainan Province: 2 ♂, 10 ♀, Lingshui County, Diaoluoshan National Forest Park (18°40'02"N, 109°55'26"E, 80 m), 24 April 2019, F.X. Liu, J. Liu and F.J. Liu leg.
Males of P. ducta are similar to Parasteatoda cingulata (Zhu, 1998) and Parasteatoda transipora (Zhu & Zhang, 1992) in having a long and looped embolus (Figs
Parasteatoda ducta (Zhu, 1998) A–C female habitus (A dorsal view B ventral view C lateral view) D–G female epigynum (D ventral view, cleared E dorsal view, cleared F ventral view, cleared and embedded in Arabic gum G dorsal view, cleared and embedded in Arabic gum). Scale bars: 0.5 mm (A–C); 0.1 mm (D–G).
Male: Total length 1.25. Prosoma 0.71 long, 0.56 wide, brownish yellow, with dark brown margin. Sternum 0.41 long, 0.36 wide, yellow, peltate. Opisthosoma 0.59 long, 0.48 wide, oval, dorsum yellow to brown, with long setae and black patches, venter black. Anal tubercle yellow. Spinnerets brown, without colulus (Fig.
Female: Total length 2.04. Prosoma 0.92 long, 0.78 wide, brownish yellow, with dark brown margin. Sternum 0.49 long, 0.47 wide, yellow and peltate, with sparse setae. Opisthosoma 1.47 long, 1.18 wide, oval, dorsum brownish black, with a discontinuous wavy yellow transverse near the midpoint which contains sporadic white spots, venter brown. Anal tubercle and spinnerets yellow, surrounded with black ring, without colulus (Fig.
China (Hainan) (Fig.
Eight cobweb spider species building detritus-based, bell-shaped retreats from China are reported in the current paper, including five new Campanicola species, two known Campanicola species and one known Parasteatoda species. However, all new species are reported only based on the female specimens because the male individuals are relatively difficult to collected in the field. According to our experience both in field collection and feeding in the lab, the males after maturity often stop weaving webs and leave their retreats to search for females. We investigated more than 500 retreats of C. campanulata in the field for research focused on its web-weaving behaviour, and only 21 males were collected, often found together with females (manuscript in preparation). Therefore, more thorough collecting or rearing juveniles in the lab may be needed to find further male individuals for these new species in the future.
The type of bell-shaped retreat is rare in the family Theridiidae, and found only in four related genera: Achaearanea, Cryptachaea, Parasteatoda and Campanicola (
We thank Mr Fengxiang Liu, and He Zhang, Yang Zhong, Wei Ding, Jingsi Lu, Rui Zhong, Jiacheng Zhang, Zhongwei Deng, Fengjie Liu, Wang Zhang, Yuting Zhang, Bing Liang and Jinhui Wang for the spider collections. This study was financially supported by CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences (19CAS-TFE-3) and the National Natural Sciences Foundation of China (NSFC-31573236/31273268/31772420).