Phylogenetic analysis and revision of the linyphiid spider genus Solenysa (Araneae: Linyphiidae: Erigoninae) Author Tu, Lihong Author Hormiga, Gustavo text Zoological Journal of the Linnean Society 2011 2011-02-24 161 3 484 530 http://dx.doi.org/10.1111/j.1096-3642.2010.00640.x journal article 10.1111/j.1096-3642.2010.00640.x 0024-4082 5494095 SOLENYSA PROTRUDENS GROUP Composition: Two species, S. protrudens Gao, Zhu & Sha, 1993 and S. lanyuensis Tu sp. nov. Diagnosis: Males of the S. protrudens group can be distinguished by the proximal tibial process with three bristles ( Fig. 17A ), by the paracymbial proximal curve posteriorly orientated ( Fig. 17A ), and by the straight sword-shaped posterior branch of lamella characteristica ( Fig. 17C ). Females are distinguished by the dorsally connected solenoid transversely twisted, anterior to the epigynum ( Fig. 17F–G ). Description: Both sexes have typical Solenysa appearance. Somatic characters as in the genus description. Male palp ( Figs 8A–D , 17A, C, D ): tibia twice as long as patella with proximal process bearing three long bristles and a ventral extension distally. Cymbial probasal process outstanding. Paracymbium with additional proximal curve posteriorly orientated. Solenysa tegular triangle long. Radix with a blunt anterior process. Embolus translucent, with a membranous projection arising from its distal part, covering embolic opening (the distal embolar membrane). Lamella characteristica with three well-developed branches extending straight and almost in parallel: anterior branch and posterior branch sword-shaped, strongly sclerotized, second branch ribbon-like with dentil apex. Terminal apophysis three free ends combined with embolus, the median one a large sclerite · Figure 11. A–G, Solenysa yangmingshana sp. nov. , H, Solenysa melloteei , I, Solenysa partibilis . A–D, male palp: A, retrolateral; B, ventral; C, prolateral; D, embolus division, ventral. E–I, epigynum: E, ventral; F, lateral; G–I, dorsal (cleared). Abbreviations: ARP, anterior radical process; ATA, anterior terminal apophysis; CD, copulatory duct; CO, copulatory opening; DP, dorsal plate; DSA, distal suprategular apophysis; E, embolus; EC, epigynal collar; FD, fertilization duct; FO, fertilization opening; LC 1 , anterior branch of lamella characteristica; LC 2 , median branch of lamella characteristica; LC 3 , posterior branch of lamella characteristica; M, distal membrane of embolus; MTA, median terminal apophysis; P, paracymbium; PBP, cymbial probasal process; PTA, posterior terminal apophysis; R, radix; S, spermatheca; SC, solenoid cover; SL, solenoid; STT, Solenysa tegular triangle. Scale bars: A–G = 0.1 mm; H–I = 0.05 mm. covering embolus, leaving anterior and posterior terminal apophysis behind at each side. Epigynum ( Fig. 17F–G ): about twice as wide as long. Solenoid connecting epigynum from dorsal side, twisting anteriorly in nonfunctional stage. A pair of transverse seams located on dorsal surface. Copulatory ducts extending forward, and then turning back, entering spermathecae from inner side ( Fig. 17I ). Fertilization ducts mesially directed. Distribution: China ( Zhejiang , Jiangsu , Taiwan) ( Fig. 22 ). SOLENYSA PROTRUDENS GAO ET AL ., 1993 ( FIGS 9 , 16 , 17 ) Solenysa protrudens Gao et al ., 1993: 65 , figs 1–6; Song et al ., 1999: 204 , fig. 116L–M, P, S. Solenysa circularis Gao et al ., 1993: 66 , figs 7–10; Song et al ., 1999: 204 , fig. 116H–I, N–O; Tu & Li, 2006b: 90 ,figs 1–11 (misidentification). New synonymy. Types: See comments below. Female holotype of S. circularis , CHINA : Mt. Tianmushan ( 30.40°N , 119.50°E ), Zhejiang Province , 10.viii.1992 (JLU). One female paratype S. circularis , CHINA : Mt. Putuoshan ( 30.00°N , 122.40°E ), Zhejiang Province , 4.viii.1992 (JLU). Material examined: CHINA : Shangsong Village , Mt. Longaoshan ( 29.60°N , 121.40°E ), Fenghua City , Zhejiang Province , coll. Y. Song , 25.iv.2006 , three males and three females ( IZCAS ) ; CHINA : Zhongshan Park ( 30.20°N , 120.10°E ), Hangzhou City , Zhejiang Province , coll. Q. Wang & Y. Han , 1.vii.2006 , two males and two females ( IZCAS ) ; CHINA : Mt. Qixiashan ( 32.10°N , 118.90°E ), Nanjing City , Jiangsu Province , coll. Q. Wang & Y. Han , 21.vi.2006 , three males and three females ( IZCAS ) . Diagnosis: Males of S. protrudens differ from those of S. lanyuensis in the pointed cymbial probasal process ( Fig. 17A ), which is stouter in S. lanyuensis ( Tu & Li, 2006b : fig. 34); by the large median terminal apophysis extending out and covering the embolus ( Fig. 17C, D ), which is shorter in S. lanyuensis ( Fig. 7C ); by the stout horn-shaped posterior terminal apophysis ( Fig. 17D ), which is a smaller triangular apophysis in S. lanyuensis ( Fig. 7C ). The females differ in the morphology of the epigynum in ventral view: the lateral sides are more rounded in S. protrudens ( Fig. 17F ) and straight in S. lanyuensis ( Tu & Li, 2006b : fig. 39). Description: Male. Total length 1.50. Carapace 0.90 long, 0.55 wide. Abdomen 0.65 long, 0.50 wide. Chelicerae with four promarginal and two retromarginal teeth. Length of legs: I 3.47 (0.90 + 1.17 + 0.80 + 0.60); II 2.80 (0.77 + 0.88 + 0.65 + 0.50); III 2.13 (0.60 + 0.65 + 0.50 + 0.43); IV 2.75 (0.75 + 0.80 + 0.68 + 0.52). Tm I 0.18, Tm IV absent. Other somatic characters as reported in the genus description. Female. Total length 1.60. Carapace 0.90 long, 0.50 wide. Abdomen 0.85 long, 0.60 wide. Chelicerae with four promarginal and two retromarginal teeth. Length of legs: I 3.05 (0.85 + 0.95 + 0.70 + 0.55); II 2.60 (0.75 + 0.80 + 0.60 + 0.45); III 1.95 (0.60 + 0.50 + 0.45 + 0.40); IV 2.53 (0.70 + 0.83 + 0.55 + 0.45). Tm I 0.19, Tm IV absent. Other somatic characters as in male. Other genitalic characters as reported in the description of the S. protrudens group above. Variation: Male carapace length is 0.85–0.90, total length 1.50–1.55 ( N = 8). Female carapace length is 0.85–0.90, total length 1.50–1.60 ( N = 8). Remarks: We have not been able to locate the type material of S. protrudens . New specimens were recently collected in the type locality ( Jiangsu ), as well as in Zhejiang (southern China ). Careful examination shows that the specimens from the type locality belong to a different species from those collected in Taiwan which was formally identified as S. protrudens by Tu & Li (2006b) . Although the original description by Gao et al . (1993) is rather superficial, comparison to their illustrations (particularly the pointed cymbial probasal process of male palp) suggests that the new specimens from Jiangsu and Zhejiang belong to S. protrudens Gao et al ., 1993 , whereas the Taiwan specimens were misidentified and belong to a new species, S. lanyuensis sp. nov. (see description below). Figure 12. Solenysa partibilis . A–F, male palp: A, retrolateral; B, ventral; C, detail of A; D, anterior; E, detail of B; F, detail of D (arrow indicates sperm duct in embolus). G, anterior part of male abdomen showing epiandrous fusules absent and smooth book lung cover. H, detail of G. Abbreviations: ATA, anterior terminal apophysis; DSA, distal suprategular apophysis; E, embolus; LC 1 , anterior branch of lamella characteristica; LC 2 , median branch of lamella characteristica; LC 3 , posterior branch of lamella characteristica; MTA, median terminal apophysis; P, paracymbium; PBP, cymbial probasal process; PTA, posterior terminal apophysis; PTP, proximal tibial process; RLP, cymbial retrolateral process; STT, Solenysa tegular triangle; T, tegulum. Scale bars in mm. Figure 13. Solenysa partibilis . A–D, epigynum: A, ventral; B, dorsal; C, caudal; D, detail of C. E–H, spinnerets: E, male; F, female; G, female ALS; H, female PLS. Abbreviations: AC, aciniform gland spigot(s); AG, aggregate gland spigot(s); ALS, anterior lateral spinneret; CY, cylindrical gland spigot(s); DP, dorsal plate; FO, fertilization opening; FL, flagelliform gland spigot(s); mAP, minor ampullate gland spigot(s); MAP, major ampullate gland spigot(s); PI, piriform gland spigot(s); PLS, posterior lateral spinneret; PMS, posterior median spinneret; SL, solenoid; VP, ventral plate. Scale bars in mm. Figure 14. Solenysa longqiensis . A–C, female prosoma: A, dorsal; B, ventral; C, frontal. D, female chelicera, ectal. E, female palp. F, solenoid cover, anterior. G–H, epigynum: G, ventral; H, dorsal. Abbreviations: CO, copulatory opening; DP, dorsal plate; SC, solenoid cover; SL, solenoid. Scale bars in mm. Furthermore, comparison of the new female specimens of S. protrudens including those from Zhejiang , the type locality of S. circularis , with the female holotype of S. circularis suggests that they are conspecific. The two species were first described in the same paper ( Gao et al ., 1993 ), but the description of S. circularis was posterior to that of S. protrudens . The original descriptions and illustrations fail to provide any information to distinguish these two species from each other. Our result suggests that the female holotype of S. circularis is conspecific with S. protrudens and thus S. circularis is a junior synonym of S. protrudens . The male paratype of S. circularis belongs to a new species S . tianmushana sp. nov. (see below). Figure 15. Solenysa longqiensis . A–C, male palp with embolic division removed: A, prolateral; B, retrolateral (arrow indicates cymbial basal excavation); C, dorsal. D–F, male embolic division: D, ventral; E, dorsal; F, detail of D. G, female spinnerets. H, anterior part of male abdomen showing epiandrous fusules absent. Abbreviations: AC, aciniform gland spigot(s); AG, aggregate gland spigot(s); ALS, anterior lateral spinneret; ARP, anterior radical process; ATA, anterior terminal apophysis; CY, cylindrical gland spigot(s); DSA, distal suprategular apophysis; E, embolus; FL, flagelliform gland spigot(s); LC 1 , anterior branch of lamella characteristica; LC 2 , median branch of lamella characteristica; LC 3 , posterior branch of lamella characteristica; M, distal membrane of embolus; MAP, major ampullate gland spigot(s); MTA, median terminal apophysis; P, paracymbium; PBP, cymbial probasal process; PI, piriform gland spigot(s); PLS, posterior lateral spinneret; PMS, posterior median spinneret; PTA, posterior terminal apophysis; R, radix; RLP, cymbial retrolateral process; SPT, suprategulum; STT, Solenysa tegular triangle; T, tegulum. Scale bars in mm. Figure 16. Solenysa protrudens . A, female body, dorsal (anterior arrow, lateral lobe of carapace; posterior arrow, median pair tracheae extending into prosoma through petiole). B–C, female prosoma: B, lateral; C, ventral (arrow, sternum connected with carapace around legs coxae). D, female palp. E–H, spinnerets: E, female ALS; F, female PLS; G, female PMS; H, male PMS and PLS (arrow, triplet nubbin). Abbreviations: AC, aciniform gland spigot(s); AG, aggregate gland spigot(s); CY, cylindrical gland spigot(s); FL, flagelliform gland spigot(s); mAP, minor ampullate gland spigot(s); MAP, major ampullate gland spigot(s); PI, piriform gland spigot(s); PLS, posterior lateral spinneret; PMS, posterior median spinneret. Scale bars in mm. Figure 17. Solenysa protrudens . A, male palp, retrolateral (arrow, paracymbial proximal curve). B, female chelicerae, ectal. C, male palpal embolic division, ventral. D, detail of C. E, anterior part of male abdomen showing epiandrous fusules absent. F–H, epigynum: F, ventral; G, dorsal; H, caudal. Abbreviations: ARP, anterior radical process; ATA, anterior terminal apophysis; CO, copulatory opening; DSA, distal suprategular apophysis; E, embolus; LC 1 , anterior branch of lamella characteristica; LC 2 , median branch of lamella characteristica; LC 3 , posterior branch of lamella characteristica; M, distal membrane of embolus; MTA, median terminal apophysis; P, paracymbium; PBP, cymbial probasal process; PTA, posterior terminal apophysis; PTP, proximal tibial process; R, radix; RLP, cymbial retrolateral process; SL, solenoid; STT, Solenysa tegular triangle; T, tegulum. Scale bars in mm. Figure 18. Solenysa retractilis sp. nov. A, male chelicerae, ectal. B, pits on carapace. C, female tracheal system. D, epigynum, ventral. E–H, spinnerets: E, male; F, female; G, male PLS; H, female PMS. Abbreviations: AG, aggregate gland spigot(s); ALS, anterior lateral spinneret; CY, cylindrical gland spigot(s); mAP, minor ampullate gland spigot(s); PLS, posterior lateral spinneret; PMS, posterior median spinneret. Scale bars in mm. Figure 19. Solenysa retractilis sp. nov. A–D, male palp: A, retrolateral (arrow, paracymbial proximal curve); B, prolateral; C, ventral (arrow, basal sclerotic extension of anterior terminal apophysis); D, detail of A. E, male palpal embolic division, ventral. F, detail of C (arrow, embolus opening). G, apex of anterior terminal apophysis. H, detail of E. Abbreviations: ATA, anterior terminal apophysis; DSA, distal suprategular apophysis; E, embolus; LC 1 , anterior branch of lamella characteristica; LC 2 , median branch of lamella characteristica; LC 3 , posterior branch of lamella characteristica; MTA, median terminal apophysis; P, paracymbium; PBP, cymbial probasal process; PTA, posterior terminal apophysis; RLP, cymbial retrolateral process; STT, Solenysa tegular triangle. Scale bars in mm. Figure 20. Solenysa wulingensis . A, male body, dorsal (left arrow, median pair tracheae extending into prosoma through petiole; right arrow, points to location of special glands). B–D, male prosoma. B, lateral; C, frontal; D, ventral. E, inner view of basal part of PLS showing special glands. F–H, spinnerets, F, male PLS and PMS; G, female PMS; H, female ALS and PLS. Abbreviations: AC, aciniform gland spigot(s); AG, aggregate gland spigot(s); ALS, anterior lateral spinneret; CY, cylindrical gland spigot(s); FL, flagelliform gland spigot(s); mAP, minor ampullate gland spigot(s); MAP, major ampullate gland spigot(s); PI, piriform gland spigot(s); PLS, posterior lateral spinneret; PMS, posterior median spinneret; SG, special gland. Scale bars in mm. Natural history: This species can be found amongst grasses, near the soil surface. Distribution: China ( Jiangsu , Zhejiang ) ( Fig. 22 ).