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1 Zootaxa 4379 (2): Copyright 2018 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Two new species of Sabacon Simon, 1879 (Opiliones: Dyspnoi: Sabaconidae) from Yunnan Province, China LI KUN ZHAO 1, JOCHEN MARTENS 2 & CHAO ZHANG 1,3 1 The Key Laboratory of Invertebrate Systematics and Application, College of Life Sciences, Hebei University, Baoding, Hebei , China 2 Institut für Organismische und Molekulare Evolutionsbiologie, D Mainz, Germany. martens@uni-mainz.de 3 Corresponding author. opiliones@163.com Abstract Two new species of the harvestman genus Sabacon Simon, 1879 are recorded from Yunnan Province, southern China, Sabacon monacanthus sp. n. (male and female; relationships unclear) and S. suzukii sp. n. (male and female; closely related to S. kangding Martens, 2015 from Sichuan Province, southwestern China). Key words: Arachnida, harvestmen, taxonomy, genitalia, evolutionary centres, relationships Introduction Sabaconidae are average-sized harvestmen with a body length of mm (Gruber 2007). They are defined by the morphology of the pedipalps, the male genitalia and by molecular phylogenetic evidence (Schӧnhofer 2013; Fernandéz et al. 2017). The family is currently reverted to its original monogeneric state (Kury 2013). Sabacon Simon occurs in the Palearctic and Nearctic in an irregular, scattered distribution and is absent in large areas (rough map in Schönhofer et al. 2013). Individual species often have small, restricted distributions (Schӧnhofer 2013, Martens 2015), with a few exceptions in Japan (Suzuki 1974) and Southwest Europe (Martens 1983), a pattern indicative of low dispersal capacities. These characteristics make the genus attractive for zoogeographers and evolutionary biologists. A molecular genetic analysis discovered massive intercontinental dispersal movements that likely occurred during the Eocene-Oligocene (Schönhofer et al. 2013). Based on strong morphological differences between various species or species groups, subgeneric (or even generic) division of the genus (eventually supported by genetic markers) is overdue, but comparison of morphological characters, especially of the male genitalia, and the results of genetic analyses thus far being unsuccessful. Presumably, old clades (or single species) occur in Japan and China (see Discussion). Concerning present knowledge, most species of the genus are concentrated in the Himalayas (Nepal and India: 18 species), North America mainly in the west coast states (eight species), and Japan (seven species, with only one being added since Suzuki s (1974) fundamental paper [i.e., a new subspecies of S. makinoi, S. m. sugimotoi Suzuki and Tsurusaki, 1983]). The mountains of Southwest and South China, where most of the Chinese species are located until now, likely form another centre of Sabacon distribution and evolution; 11 species are now known from there. The soil faunas of these regions are little studied and further investigations may provide deeper insight, as seen from the present paper. The taxonomic catalogue of the suborder Dyspnoi Hansen and Sørensen, 1904, (Schӧnhofer 2013) included 35 Sabacon species. Subsequently, Martens (2015) described an additional 20 new species from the Palearctic region with focus on Nepal and China. Among the 55 species of Sabacon presently known, only nine occur in China and many additional ones are to be expected. The Chinese species are distributed as follows: Liaoning Province (S. okadai Suzuki, 1941), Qinghai Province (S. beishanensis Martens, 2015), Shaanxi Province (S. pauperoserratus Martens, 2015; S. multiserratus Martens, 2015), and Sichuan Province (S. gonggashan Tsurusaki and Song, 1993; S. martensi Tsurusaki and Song, 1993; S. kangding Martens, 2015; S. jaegeri Martens, 2015; S. minshanensis Martens, 2015). Accepted by J. Shultz: 12 Dec. 2017; published: 14 Feb

2 Although Yunnan is one of the best-investigated regions in South China, the harvestmen fauna remains rather poorly known. Recently, some Dyspnoi specimens from Yunnan Province became available courtesy of the Arachnological Laboratory of Hebei University, among them two Sabacon species new to science, which are described here. Materials and methods Specimens were collected mainly by pitfall traps and were preserved in 75% ethanol. Line drawings were produced under a Leica M205A stereomicroscope equipped with a drawing tube. Photographs were taken using a Leica M205A stereomicroscope equipped with a DFC450 CCD. The terminology follows Gruber (2007) and Martens (1978, 2015). The type specimens are deposited in the Museum of Hebei University, Baoding, China (MHBU). All measurements are given in mm. Taxonomy Sabacon Simon, 1879 Sabacon Simon, 1879: 266; Shear, 1975: 11 12; Schӧnhofer, 2013: 16; Martens, 2015: 170. Phlegmacera Packard, 1884: 203. Tomicomerus Pavesi, 1899: 533. Parasabacon Hansen & Sørensen, 1904: 83. Type species: Sabacon paradoxus Simon, 1879, by monotypy (southern France). Sabacon suzukii sp. n. (Figs 1 8, 10 15, 17 34, 53 55) Type material. Holotype male, CHINA: Yunnan Province, Lijiang, Mt. Yulongxue Shan, [N 27 11, E ], alt m, August 2012, A. Nakamura leg. (MHBU-Opi ). Paratypes: one male (MHBU-Opi ) and one female (MHBU-Opi ), CHINA: Yunnan Province, Qianjiazai, Mt. Ailao Shan [N 24 17, E ], alt m, August 2011, A. Nakamura leg.; one male (MHBU-Opi ), CHINA: Yunnan Province, Dali City, Mt. Cang Shan [N 25 58, E ], alt m, June 2, 2008, P. Feng leg. Diagnosis. Opisthosomal dorsum with scutum parvum (male) or scutum laminatum (female). The second segment of male chelicerae latero-basally with one or three stout thorns. Pedipalps elongated, thin and filiform. Male pedipalpal patellae distally with six to 11 ventral teeth. Penial glans dorsally with two spoon-like integrated appendages. Ovipositor of elongated type. S. suzukii sp. n. probably is closely related to S. kangding Martens from Sichuan. Male genital morphological characters are uniquely similar and unknown from any other Sabacon species of the Holarctic: the two spoon-like and punch-like appendages on the glans penis, and in addition the extremely thin and slender pedipalps of both sexes. S. kangding carries vestigial claws on the pedipalp tarsus, a truly plesiomorph character within Opiliones in general, which are lacking in S. suzukii sp. n. Etymology. The species is named in honor of Professor Dr. Seisho Suzuki ( ), esteemed and wellknown Japanese arachnologist. In many taxonomic papers he described and revised the whole Japanese opilionid fauna and, among others, paid much attention to the local Sabacon fauna. The first Sabacon species known from China was published by him (Suzuki 1941). In addition, in numerous, often voluminous papers he brought to our attention the rich fauna of Southeast Asian opilionids always accompanied by meticulous drawings of the habitus, external and genital morphology showing a wealth of details. In this, he stands among the pioneers of Asian arachnology. Description. It is based on male holotype and a female paratype from Mt. Ailaoshan. Male habitus as in Figs 1, 8, 53, 55. Coloration (Figs 53, 55): medial area of carapace with yellowish brown trapezoidal area and two conspicuous yellowish brown spots on dull white soft cuticle laterally. Ocularium dark brown, with blackish eye 200 Zootaxa 4379 (2) 2018 Magnolia Press ZHAO ET AL.

3 rings and a pale dorsal band. Meso- and metapeltidium greyish white and anteriorly each with a transverse row of yellowish brown stripes, also and mesopeltidium with longitudinal brown short stripes. Opisthosomal dorsum dark brown medially and greyish white laterally. Coxae of legs and genital operculum yellowish brown. Sternites greyish white, anteriorly each with a yellowish brown streak. Chelicerae, pedipalps and legs yellowish brown to dark brown. Dorsum (Figs 8, 53). Dorsal scutum oval in shape. Carapace smooth, only a few setae on each side of ocularium. Anterior margin of carapace with a median notch, and curved inward on each side. Ozopores small and visible above the first coxae. Ocularium canaliculate, smooth. Metapeltidium medially with a pair of sensory cones. Opisthosomal dorsum with scutum parvum and area I V each with transverse rows of setae. Free tergites each with a transverse row of setae. Venter. Surface of all coxae and genital operculum with many scattered setae. Free sternites smooth, each only with a transverse row of setae along its anterior margin. Chelicerae (Figs 2 4). Basal segment smooth, apophysis absent, dorsally with many scattered setae and ventrally with a few similar setae. Second segment latero-basally with three stout, massive cuticular projections (Fig. 3); many long setae on the frontal surface, and clusters of short ones at base of fixed finger. Chela fingers relatively short, with diaphanous narrow teeth and a subapical coarse dark tooth (Fig. 4): the coarse tooth on moveable finger pointed; a similar molar tooth on the fixed finger. Pedipalps (Figs 5 7; Table 1). Slender. Trochanter ventrally and dorsally with scattered setae. Femur ventrally with brush-like setae and remaining surfaces covered with setae. Patella distally with a longitudinal row of six medio-ventral teeth, one inconspicuous tooth in distal third (Fig. 5), these and the brush-like setae restricted to the base. Tibia and tarsus throughout with dense brush-like setae. Tarsus extremely slender, without claw, and a length: width ratio of 0.76: Legs (Table 1). Relatively long, cylindrical in cross section, all articles with dark, short, scattered setae. Femora, patellae, tibiae, metatarsi and tarsi with short setae. Penis (Figs 20 25). Shaft slender and flattened dorso-ventrally; nearly parallel-sided, widened basally, then distended in apical portion (dorsal and ventral view); basal portion (fork) bent dorsally against main portion of shaft by almost 90. Glans dorsally with a large and a small spoon-like membranous appendage at the distal margin: the bigger one bent and opened dorsally, proximally widened and distally narrowed; the smaller one bent ventrally. Stylus asymmetrically twisted, gradually tapering distally. Glans with 20 setae: four short dorsal ones, two small dorsal ones, 10 short ventral ones, two short lateral ones, and two long lateral ones. TABLE 1. Pedipalp and legs measurements of Sabacon suzukii sp. n., male holotype from Mt. Yulongxue Shan, length/ width Trochanter Femur Patella Tibia Metatarsus Tarsus Total Pedipalp 0.48/ / / / / Leg I 0.30/ / / / / / Leg II 0.33/ / / / / / Leg III 0.32/ / / / / / Leg IV 0.35/ / / / / / Female (Figs 15, 17 19, 54; Table 2). In general appearance similar to the male but opisthosomal dorsum with scutum laminatum. The second segment of chelicerae unarmed, latero-basally without stout thorns, only with setae (Fig. 18). Pedipalpal articles less slender than in males, patellae unarmed, distally without any medio-ventral teeth (Fig. 17). Dorsum light purplish brown instead of dull white as in the male (Fig. 54). Ovipositor as illustrated (Fig. 19), elongate type. Measurements. Male holotype (female paratype from Mt. Ailaoshan): body 3.16 (4.19) long, 1.65 (2.29) wide at the widest portion. Ocularium 0.27 (0.34) long, 0.37 (0.47) wide, 0.15 (0.19) high. Penis 1.57 long (including glans), 0.14 wide at base, fork 0.47 long. Ovipositor 1.70 long. Measurements of pedipalp and legs as in Tables 1-2. Habitat. All the specimens were collected by pitfall traps in coniferous forest. TWO NEW SPECIES OF SABACON Zootaxa 4379 (2) 2018 Magnolia Press 201

4 FIGURES 1 7. Sabacon suzukii sp. n., male holotye (from Mt. Yulongxue Shan). 1. Body, lateral view. 2. Left chelicera, medial view. 3. Same, ectal view. 4. Cheliceral fingers, frontal view. 5. Left pedipalp, medial view (arrow indicates the inconspicuous tooth). 6. Same, ectal view. 7. Right pedipalpal patella, medial view. Scale bars: 1 mm (1 6); 0.5 mm (7). 202 Zootaxa 4379 (2) 2018 Magnolia Press ZHAO ET AL.

5 FIGURES 8, Sabacon suzukii sp. n.; 9. Pattern of scutum parvum (from Martens, 1978). 8, male holotype (from Mt. Yulongxue Shan). ); 10, 13 14, male paratype from Mt. Ailao Shan (MHBU-Opi ); 11 12, male paratype from Mt. Cang Shan (MHBU-Opi ). 8. Body, dorsal view Left chelicera, ectal view. 12. Left pedipalpal patella, medial view. 13. Left pedipalp, medial view. 14. Same, ectal view. Scale bars: 1 mm (8, 13 14); 0.5 mm (10 12). TWO NEW SPECIES OF SABACON Zootaxa 4379 (2) 2018 Magnolia Press 203

6 FIGURES 15, Sabacon suzukii sp. n., female paratype (from Mt. Ailao Shan); 16. Pattern of scutum laminatum (from Martens, 1978). 15. Body, dorsal view. 17. Left pedipalp, medial view. 18. Left chelicera, ectal view. 19. Ovipositor. Scale bars: 2 mm (15); 1 mm (17, 19); 0.5 mm (18). 204 Zootaxa 4379 (2) 2018 Magnolia Press ZHAO ET AL.

7 FIGURES Sabacon suzukii sp. n., male penis , holotype from Mt. Yulongxue Shan (MHBU-Opi ); 26 29, paratype from Mt. Cang Shan (MHBU-Opi ); 30 34, paratype from Mt. Ailao Shan (MHBU-Opi ). 20. Ventral view. 21. Lateral view. 22, 26, 31. Distal part, ventral view. 23, 27, 32. Distal part, dorsal view , 28 29, Distal part, lateral view. 30. Lateral view. LLS, long lateral setae; LVS, long ventral setae; SDS, short dorsal setae; SLS, short lateral setae; SmS, small setae; SVS, short ventral setae. Scale bars: 0.2 mm. TWO NEW SPECIES OF SABACON Zootaxa 4379 (2) 2018 Magnolia Press 205

8 Variation. Two male specimens were examined (MHBU-Opi from Mt. Ailao Shan, from Mt. Cang Shan). Body 3.03 and 3.70 long, 1.68 and 1.97 wide at the widest portion. Second segment of the male chelicerae latero-basally with one thorn (Figs 10, 11). The male (MHBU-Opi ) pedipalpal patellae distally with a longitudinal row of 11 medio-ventral teeth (Figs 13 14). The distal part of penis shaft (MHBU-Opi ) bent to ventral side against main portion of shaft by almost 90, probably an artificial state (Fig. 30). The glans of penis (MHBU-Opi ) basally with two additional long ventral setae (Fig. 26) compared with the other male (Figs 22, 31). The male pedipalpal patellae from Mt. Yulongxue Shan (Figs 5 6) and Mt. Cang Shan (Fig. 12) with six teeth, that of the specimens from Mt. Ailao Shan with 11 teeth (Figs 13 14). The other variation is that teeth on specimens from Mt. Yulongxue Shan and Mt. Cang Shan are more robust than in the specimens from Mt. Ailao Shan. Left pedipalpal patella of male specimens from Mt. Yulongxue Shan with six teeth (Fig. 5), the right one with three teeth (Fig. 7). Although the difference in the shape of the teeth on the male pedipalpal patellae is conspicuous, this does not appear to indicate species level differences. The crucial male genitalia are quite homogeneous though slight differences in penial glans setation are obvious: four small setae on the distal truncus from Mt. Cang Shan (Fig. 27); two small setae and eight small setae in the same position from Mt. Yulongxue Shan (Fig. 23) and Mt. Ailao Shan (Fig. 32), respectively; glans lacks two long ventral setae in both specimens from Mt. Yulongxue Shan (Fig. 22) and Mt. Ailao Shan (Fig. 31). Because the three mountains are quite isolated and population exchange may be reduced due to areas inhospitable to Sabacon species, which prefer cool and moist mountainous habitats, these differences reflect geographic variation. The linear distances between the three localities are as follows: 143 km between Mt. Yulongxue Shan and Mt. Cang Shan; 230 km between Mt. Cang Shan and Mt. Ailao Shan; 336 km between Mt. Yulongxue Shan and Mt. Ailao Shan. Distribution. Known from three mountain stocks in Yunnan Province, China. TABLE 2. Pedipalp and legs measurements of Sabacon suzukii sp. n., female paratype from Mt. Ailao Shan, length/ width Trochanter Femur Patella Tibia Metatarsus Tarsus Total Pedipalp 0.54/ / / / / Leg I 0.43/ / / / / / Leg II 0.48/ / / / / / Leg III 0.43/ / / / / / Leg IV 0.48/ / / / / / Sabacon monacanthus sp. n. (Figs 35 42, 44 52, 56 58) Type material. Holotype male, CHINA: Yunnan Province, Lijiang City, Mt. Yulongxue Shan [N27 11, E ], alt m, August 2012, A. Nakamura leg. (MHBU-Opi ). Paratypes: one male (MHBU- Opi ) and one female (MHBU-Opi ), same collecting data as holotype. Diagnosis. Opisthosomal dorsum with scutum dissectum (male and female). The second segment of male chelicerae latero-basally with one stout thorn. Pedipalps short and robust. Male pedipalpal patellae distally with one ventral thorn. Scanty penial spination. The shaft of penis nearly parallel-sided up to glans. Stylus enlarged on proximal third of length. Sabacon monacanthus sp. n. can easily be distinguished from North American, European, Russian, and Japanese species by the lack of a dorsal apophysis on the basal segment of the male chelicerae and by male genital morphology (Shear 1975; Martens 1969, 1983, 1989, 2015; Suzuki 1974; Luque 1991). Similarly, the morphology of the male genitalia and the pedipalp of the new species are helpful to distinguish it from the set of Chinese species. Sabacon monacanthus sp. n. possesses short and robust pedipalps (Figs 39 40) instead of extremely slender ones (cf. S. okadai and S. kangding; Suzuki 1941: 16 19, f.1 3; Martens 2015: , f ); the penis lacks spines and thorns along the distal part of the penis truncus (cf. S. beishanensis, S. 206 Zootaxa 4379 (2) 2018 Magnolia Press ZHAO ET AL.

9 jaegeri, S. martensi, S. minshanensis, S. multiserratus, S. pauperoserratus and S. kangding; Martens 2015: 196; Tsurusaki and Song, 1993: , f.1 2); S. gonggashan shows only one pair of setae on the penis (Tsurusaki and Song 1993: , f. 3 5). Sabacon monacanthus sp. n. on the other hand is morphologically similar to Nepalese species, i.e., the Himalayan chomolongmae species group (12 species) (Martens 1972, 2015). One of these species, S. sineglandula Martens, 2015 appears to be most similar to S. monacanthus sp. n. in lacking the dorsal cheliceral apophysis of males and the gland normally encapsulated there. S. sineglandula differs, however, in having an apophysis-like structure on the second segment of the male chelicerae (Martens 2015: 186, 189, f. 116). Also, the broadest part of penis is situated below the glans in S. sineglandula (Martens 2015: 189, f. 123), while in S. monacanthus sp. n. the shaft of the penis (Figs 46, 48) is nearly parallel-sided (dorsal and ventral view). Further, the stylus is enlarged from base to apical one-third (Figs 49, 52) which is absent in S. sineglandula (dorsal and ventral view); glans spination differs as well (Martens 2015: 189, f. 121). Etymology. The specific name is derived from the Greek terms μόνος (monos) for one and ἄκανθα (akantha) for thorn. The Latinized monacanthus means single thorn and refers to one stout thorn each on the male cheliceral second segment and the male pedipalp. Noun in apposition. Description. Description below is based on male holotype and a female paratype. Male habitus as in Figs 35, 41, Coloration (Figs 56 57): medial area of carapace with yellow trapezoidal area and two purplish brown stripes laterally. Ocularium dark brown. Meso- and metapeltidium each with a transverse row of purplish brown streak. Opisthosomal dorsum with an irregular light median stripe from metapeltidium to area II, following posterior areas also light, many white spots laterally. Coxae of legs and genital operculum yellowish brown. Sternites grayish white, medially with a longitudinal light purplish brown stripe, and two dark ones laterally. Chelicerae, pedipalps and legs yellowish white to yellowish brown. Dorsum (Figs 41, 57). Scutum oval in shape. Carapace smooth, only a few setae along anterior margin; its anterior margin with a median notch. Ozopores small and visible from above, located close to first and second coxae. Ocularium unarmed, canaliculate. Metapeltidium free, with a median pair of sensory cones. Opisthosomal dorsum with scutum dissectum and only a few small scattered setae. Venter. Surface of all coxae, genital operculum and free sternites with many scattered setae. Chelicerae (Figs 36 38). Basal segment smooth, dorsal apophysis absent, dorsally with many scattered setae. Second segment latero-basally with a stout, pointed and hook-like thorn pointing to the basal segment, many long setae on the frontal surface and a few short ones at base of fixed finger. Chela finger short, with diaphanous narrow teeth and subapical coarse dark teeth (Fig. 38): one pointed tooth on moveable finger and two on fixed finger. Pedipalps (Figs 39 40; Table 3). Stout. Trochanter ventrally with scattered setae, dorsally with only a few short setae. Femur distally broadened and with scattered setae. Patella distally with a ventro-mesal strong thorn, setae similar to those of femur. Tibia and tarsus inflated, throughout with dense brush-like setae. Tarsus stout, without claw, and a length: width ratio of 0.35: Legs (Table 3). Short, cylindrical in cross section, all articles with dark, short, scattered setae. TABLE 3. Pedipalp and legs measurements of Sabacon monacanthus sp. n., male holotype from Mt. Yulongxue Shan, length/width Trochanter Femur Patella Tibia Metatarsus Tarsus Total Pedipalp 0.25/ / / / / Leg I 0.23/ / / / / / Leg II 0.26/ / / / / / Leg III 0.24/ / / / / / Leg IV 0.26/ / / / / / Penis (Figs 46 52). Shaft slender and flattened dorso-ventrally, nearly parallel-sided up to glans; basal portion (fork) bent dorsally against main portion of shaft by almost 150. Glans slightly asymmetrical, stylus enlarged on basal one third. Small-scale spination scanty, all spines minute, concentrated at basis of stylus, at both sides of joint area and below joint area (ventral view). Truncus distally with eight ventral setae (Figs 49 50), glans with eight lateral setae on each side (Fig. 52). TWO NEW SPECIES OF SABACON Zootaxa 4379 (2) 2018 Magnolia Press 207

10 FIGURES Sabacon monacanthus sp. n., male holotype (from Mt. Yulongxue Shan). 35. Body, lateral view. 36. Left chelicera, medial view. 37. Same, ectal view. 38. Cheliceral fingers, frontal view. 39. Left pedipalp, medial view. 40. Same, ectal view. Scale bars: 1 mm (35); 0.5 mm (36 40). 208 Zootaxa 4379 (2) 2018 Magnolia Press ZHAO ET AL.

11 FIGURES 41 42, Sabacon monacanthus sp. n.; 43. Pattern of scutum dissectum (adapted from Martens, 1978). 41. Male holotype. 42, Female paratype (from Mt. Yulongxue Shan) Body, dorsal view. 44. Left pedipalp, medial view. 45. Left chelicera, ectal view. Scale bars: 1 mm (41 42); 0.5 mm (44 45). TWO NEW SPECIES OF SABACON Zootaxa 4379 (2) 2018 Magnolia Press 209

12 FIGURES Sabacon monacanthus sp. n., penis of male holotype (from Mt. Yulongxue Shan). 46. Ventral view. 47. Lateral view. 48. Dorsal view. 49. Distal part, ventral view Same, lateral view (different angles from the same penis). 52. Same, dorsal view. Scale bars: 0.5 mm (46 48); 0.2 mm (49 52). 210 Zootaxa 4379 (2) 2018 Magnolia Press ZHAO ET AL.

13 FIGURES Photographs of body and parts of appendages of Sabacon suzukii sp. n. and Sabacon monacanthus sp. n. 53, 55. Male holotype of Sabacon suzukii sp. n., from Mt. Yulongxue Shan. 54. Female paratype of Sabacon suzukii sp. n., from Mt. Ailao Shan Sabacon monacanthus sp. n., from Mt. Yulongxue Shan. 53, 57. Male holotype, dorsal view. 54, 58. Female paratype, dorsal view Male holotype, lateral view. Scale bars: 1 mm (53 55); 0.5 mm (56 58). Female (Figs 42, 44 45, 58; Table 4). In general appearance similar to the male. The second segment of chelicerae and pedipalps unarmed. Ovipositor. No information; lost. Measurements. Male holotype (female paratype): body 2.04 (2.00) long, 1.39 (1.21) wide at the widest portion. Ocularium 0.22 (0.22) long, 0.34 (0.30) wide, 0.16 (0.16) high. Penis 1.66 long (including glans), 0.10 wide at base, fork 0.41 long. Measurements of pedipalp and legs as in Tables 3 4. TWO NEW SPECIES OF SABACON Zootaxa 4379 (2) 2018 Magnolia Press 211

14 Habitat. The specimens were collected by pitfall traps in conifer forest. Variation (male paratype). Body 1.97 long, 1.13 wide at the widest portion. Distribution. Known only from the type locality in Yunnan Province, China. Notes. The four most common patterns of the dorsal opisthosomal surface of harvestmen are scutum completum, scutum magnum, scutum laminatum, and scutum parvum (Martens 1978: 16 17, figs 2 5; Shultz and Pinto-da-Rocha 2007: 21 22, fig. 2.4). Scutum laminatum and scutum parvum are used here just to describe the morphological patterns of Sabacon suzukii sp. n. (Figs 9, 16). However, there is an additional term to describe variation of opithosomal sclerites of harvestmen. One of the patterns proposed here, scutum dissectum, found in Sabacon monacanthus sp. n., is similar to the scutum parvum. In scutum dissectum the distal five opisthosomal tergites are disintegrated and form rows or groups of small sclerites per opisthosomal area. (Fig. 43). TABLE 4. Pedipalp and legs measurements of Sabacon monacanthus sp. n., female paratype from the type locality, length/width Trochanter Femur Patella Tibia Metatarsus Tarsus Total Pedipalp 0.28/ / / / / Leg I 0.22/ / / / / / Leg II 0.25/ / / / / / Leg III 0.22/ / / / / / Leg IV 0.24/ / / / / / Discussion Obviously, knowledge of Sabacon in China is still in its infancy. Eleven species (including the new ones presented here) are known to date and they span a large geographic arc from the northeastern to the southwestern provinces, including mountain ranges in Shaanxi (Qin Ling), Qinghai (Bei Shan), and Sichuan (Min Shan), with Yunnan here adding further mountain ranges. In the collection of harvestmen made in China by J. Martens are more Sabacon specimens but they are only females and juveniles, which are not useful for delimiting species. The network of localities is even narrower than presently published and includes southern Gansu (Lianhua Shan; Schönhofer et al. 2013, fig. 3). Phylogenetic relationships of Chinese species, or Asian species in general, are not yet worked out. Molecular phylogenetic relationships based on a few of the Japanese, Chinese and Nepal species point to a number of Asian evolutionary centres of speciation and various pathways of intercontinental exchange between all presently known more or less isolated distributional areas. However, the few species investigated do not allow us to draw detailed conclusions, but a first step has been achieved (Schönhofer et al. 2013). However, we tentatively try to judge some character states of Sabacon morphology. Apparently, basal states are thin pedipalps as in Dyspnoan harvestmen in general (vs. inflated ones in most of the known Sabacon species). Also the presence of pedipalpal claws as in harvestmen in general is basal (vs. its lack in all Sabacon species except for the single one presently known and in all Dyspnoi). The general coincidence of thin (basal) pedipalps and complicated male genital morphology with various appendages on the glans area in several Chinese and Japanese species is another hint that the latter character is basal, plesiomorphic. But due to many peculiarities in the genitalic character set of the relevant species in China and Japan these may belong to different clades in the early evolution of the genus. Among the 18 Himalayan species, Martens (2015) was able to separate two species sets. One is characterized by complicated male genital morphology (but always without glans appendages) and dark body coloration. The species of this set are confined to comparatively low-elevation distributional areas between 2400 m and about 2900 m. The other set has simpler male genital morphology (glans and stylus drawn out lanceolate in prolongation of the truncus) and light body coloration. These species live at high altitudes from about 2400 m to 4600 m, even considerably beyond the timberline. The species of both species sets dispose of inflated pedipalps, but much less so in the first group (hardly approaching Chinese species S. kangding and S. suzukii sp. n.). 212 Zootaxa 4379 (2) 2018 Magnolia Press ZHAO ET AL.

15 Here we hypothesize that both species groups represent different evolutionary clades. The first one including low-altitude species may belong to an older clade with still (but unusual in present Sabacon) subtropical ecological affinities and more basal morphological characters (and dark body and appendages coloration as an additional external sign). The other group shows more derived characters and adapted to severe high-elevational biota of the Himalayas and/or may have been uplifted with the growing mountain chain. This group may represent more recent settlers in the Himalayas and it underwent multiple speciation events resulting in a number of highly similar species. Unfortunately, Schönhofer et al. (2013) included only specimens of the low-altitude Nepal group in their molecular genetic study thus making it presently impossible to confirm our hypothesis. From this morphological point of view, rather plesiomorphic species seem to occur in China and in Japan. The only Sabacon species known to date with vestigial palpal claws is S. kangding from Sichuan. In addition, its pedipalps are extremely slender and its genital morphology most complicated. In S. okadai from Northeast China, pedipalps are slender as in S. kangding, but its genital morphology is undescribed. It may join Japanese species, several of them carrying slender pedipalps as well. The genital morphology of S. kangding is rather complicated and basal (as in S. suzukii sp. n.). In Japan comparable characteristics prevail in slender pedipalps and complicated genital morphology as well, but genitalic traits differ in many details. At this time, close relationships between Japanese and Chinese species are unproven. Within China, a vast morphological diversity within Sabacon emerges even among the few species known to date. Apparently, S. kangding and S. suzukii sp. n. are basal and S. beishanensis and S. monacanthus sp. n. are more derived, with massively inflated pedipalps and simple male genital morphology with inconspicuous setal armament. In between these, there exists a range of species with variously and often heavily specialized sets of penial setae, indicating long periods of separate species evolution in various mountain stocks of different ages. This scenario emerges only dimly, and we are far from a detailed understanding of Asian Sabacon diversity, distribution and evolution. Acknowledgements For access to rare specimens we are deeply indebted to Wenzhi Wang, Zizhong Yang, Akihiro Nakamura, Zongxu Li and Ping Feng. Cordial thanks are due to Peter Jäger for providing literature. This work was supported by the National Natural Science Foundation of China (No ). Jochen Martens was supported by Feldbausch Foundation and Wagner Foundation, both at Fachbereich Biologie at Mainz University, to carry out field research in Asia. We thank William Shear, Jeffrey Shultz, and an anonymous reviewer for useful comments on a former draft of this paper, and special thanks are due to Jeffrey Shultz for his careful editing. References Fernandéz, R, Sharma, P.P., Tourinho, A.L. & Giribet, G. (2017) The Opiliones tree of life: Shedding light on harvestmen relationships through transcriptomics. Proceedings of the Royal Society B, 284, Gruber, J. (2007) Sabaconidae Dresco, In: Pinto-da-Rocha, R., Machado, G. & Giribet, G. (Eds.), Harvestmen: The biology of Opiliones. Harvard University Press, Cambridge & London, pp Hansen, H.J. & Sørensen, W. (1904) On two orders of Arachnida. Opiliones, especially the suborder Cyphophthalmi, and Ricinulei, namely the family Cryptostemmatoidae. Cambridge University Press, Cambridge, pp Kury, A.B. (2013) Order Opiliones Sundevall, In: Zhang, Z.-Q. (Ed.), Animal Biodiversity: An Outline of Higher-level Classification and Survey of Taxonomic Richness (Addenda 2013). Zootaxa, 3703 (1), Luque, C.G. (1991) Los Ischyropsalidoidea de la Cornisa Cantabrica. Cantabria. Arquenas. In: Fauna Iberica Subterranea. Cantubria. Vol.1. Editorial Impresion, Santander, pp Martens, J. (1969) Die Abgrenzung von Biospezies auf biologisch-ethologischer und morphologischer Grundlage am Beispiel der Gattung Ischyropsalis C. L. Koch 1839 (Opiliones, Ischyropsalididae). Zoologische Jahrbücher, Jena, Abteilung für Systematik, Ökologie und Geographie der Tiere, 96 (2), Martens, J. (1972) Opiliones aus dem Nepal-Himalaya. I. Das Genus Sabacon Simon (Arachnida: Ischyropsalididae). Senckenbergiana biologica, 53 (3/4), Martens, J. (1978) Spinnentiere, Arachnida: Weberknechte, Opiliones. In: Senglaub, K., Hannenmann, H.J. & Schumann, H. TWO NEW SPECIES OF SABACON Zootaxa 4379 (2) 2018 Magnolia Press 213

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