On eight species of the spider genus Synagelides Strand, 1906 from China (Araneae: Salticidae)

Six new species of Synagelides Strand, 1906 are diagnosed and described: S. bohdanowiczi sp. nov. (♂♀), S. leigongensis sp. nov. (♂♀), S. logunovi sp. nov. (♂♀), S. subgambosus sp. nov. (♂♀), S. wuliangensis sp. nov. (♂♀) and S. xingdouensis sp. nov. (♂♀). The female of S. forkiforma Yang, Zhu & Song, 2007 and the male of S. longus Song & Chai, 1992 are described for the fi rst time. Photos of the habitus and copulatory organs, as well as a distributional map, are provided.


Introduction
The genus Synagelides Strand, 1906 comprises a group of ant-like spiders distributed from the Russian Far East to Southeast Asia (Maddison 2015;WSC 2020). A total of 51 nominal species with diagnostic illustrations have been assigned to the genus, of which 27 were described only from a single sex (13 only from males and 14 only from females) (WSC 2020). Almost all species were originally described as a member of this genus except Synagelides cavaleriei (Schenkel, 1963), which was transferred from Tagoria Schenkel, 1963, a synonym of Synagelides. Most species of Synagelides have very limited distributional ranges (46 species are endemics and known only from a single country and three species are known from two countries). Bohdanowicz (1978Bohdanowicz ( , 1979Bohdanowicz ( , 1987 has done extensive work on this genus, describing all species from Bhutan, Japan and Nepal, and also re-describing the type species. Seven new species and six synonyms were added by Logunov & Hereward (2006). To date, this genus exhibits the highest species diversity in China, with 24 species, including 19 endemic ones (Song 1990;Xie & Yin 1990;Song & Chai 1992;Peng et al. 1998Peng et al. , 2002Peng et al. , 2003Peng et al. , 2008Song & Zhu 1998;Zhu et al. 2005;Yang et al. 2007;Liu et al. 2017;Lin & Li 2020;Peng 2020), followed by Nepal, with 10 species, including nine endemic ones (Bohdanowicz 1979(Bohdanowicz , 1987Logunov & Hereward 2006).
While examining Synagelides specimens from South China (Yunnan, Guizhou, Hubei and Guangxi), more than twenty species, including six new to science with both sexes, the previously unknown female of S. forkiforma Yang, Zhu & Song, 2007 and the previously unknown male of S. longus Song & Chai, 1992, have been identifi ed and are described here. A re-examination of the types of S. gambosus Xie & Yin, 1990 reveals that the retrolateral tibial apophysis of the male palp has been ignored and never illustrated or described until now. The goals of this paper are to describe and illustrate the new species as well as the previously unknown sexes of S. longus and S. forkiforma, and to present the retrolateral tibial apophysis of S. gambosus.

Material and methods
Specimens were collected mainly by beating shrubs and screening leaf-litter. All specimens were preserved in 75% ethanol and are deposited in the museum of Tongren University (TRU), Tongren, China except the types of Synagelides xingdouensis sp. nov. and the new material of S. forkiforma Yang, Zhu & Song, 2007, which are deposited at the College of Life Sciences, Hunan Normal University (HNU), Changsha, Hunan, China.
The specimens were examined with an Olympus SZ51 stereo microscope. After dissecting from the body, epigynes were cleared in trypsin enzyme solution before examination and imaging. Left male palps, legs I and chelicerae were used for illustration. Photos were taken with a Kuy Nice CCD mounted on an Olympus BX51 compound microscope except the photos of Fig. 17, which were taken with a Canon G12 camera mounted on an Olympus BX53 compound microscope. Compound focus images were generated using Helicon Focus 6.7.1 and 3.1 software.
All measurements are given in millimeters. Leg measurements are giving as: total length (femur, patella + tibia, metatarsus, tarsus). References to fi gures in the literature are listed in lowercase type (fi g. or fi gs); fi gures in this paper are noted with an initial capital (Fig. or Figs). Terminology follows Lin & Li (2020). Abbreviations used in the text and fi gures are as follows: AERW = anterior eye row width AR = arcuated rim AME = anterior median eye At = atrium CD = copulatory duct CO = copulatory opening DCA = dorsal cymbial apophysis DTA = dorsal tibial apophysis EFL = eye fi eld length Em = embolus FD = fertilization duct GD = gland duct Ho = hood ITA = intermediate tibial apophysis MA = median apophysis MS = median septum PERW = posterior eye row width PLE = posterior lateral eye PME = posterior median eye PCA = prolateral cymbial apophysis RPA = retrolateral patellar apophysis RTA = retrolateral tibial apophysis S = spermatheca SD = sperm duct sRTA = second retrolateral tibial apophysis.

Differential diagnosis
Synagelides bohdanowiczi sp. nov. resembles S. gambosus (Xie & Yin 1990: fi gs 1-7; Fig. 11D) and S. subgambosus sp. nov. in the S-shaped retrolateral tibial apophysis and the long and narrow median septum in the epigyne, but differs from these two species by: 1) retrolateral tibial apophysis originates from the terminal of the tibia, with the tip directed upwards in retrolateral view in S. bohdanowiczi sp. nov. (Fig. 1B), whereas originates from the middle of the tibia, with the tip directed horizontally towards the genital bulb in S. gambosus (Fig. 11D) and S. subgambosus sp. nov. (Fig. 11B); 2) median apophysis is horn-shaped, pointed and far away from the embolus apically in S. bohdanowiczi sp. nov. (Fig. 1B), whereas more broadened and very close to or partially covering the embolus in S. gambosus (Fig. 11D) and S. subgambosus sp. nov. (Fig. 11B); 3) atrium is elongated oval, longer than wide, lacking wrinkles in S. bohdanowiczi sp. nov. (Fig. 2A), whereas wider than long, with distinct wrinkles in S. subgambosus sp. nov. (Fig. 12A); 4) spermathecae extend obliquely in dorsal view in S. bohdanowiczi sp. nov. (Fig. 2C), whereas extend transversely in S. gambosus (Xie & Yin 1990: fi g. 7) and S. subgambosus sp. nov. (Fig. 12C).  2D) stippled, red-brown, anterior surface and eye base dark, covered by brown hairs anteriorly and sparse white hairs posteriorly. Fovea oval, hollowed. Chelicerae (Fig. 2G) yellow, with two promarginal teeth and one retromarginal tooth fi ssident. Endites (Fig. 2E) yellowbrown, slightly longer than wide, inner margins pale with dense hairs. Labium (Fig. 2E) gradually narrowed, apically bearing dense hairs. Sternum (Fig. 2E) yellow, scutiform. Legs yellow to red-yellow except the basal half of femur I dark. Spination of leg I (Fig. 2H): femur v2-2-2-2; metatarsus v0-1-1. Abdomen ( Fig. 2D-E) elongated; dorsum dark-brown, the posterior one-third portion darker, with a pair of round spots antero-medially, and an inconsecutive, longitudinal pale-brown stripe extending from the anterior margin to the posterior one-third portion; venter gray, with two longitudinal dark-brown stripes bilaterally and two longitudinal lines medially. Palp (Figs 1A -D, 17C): patella about 1.5 times longer than wide in retrolateral view; retrolateral tibial apophysis S-shaped, broadened at base; cymbium hairy, with sclerotized prolateral apophysis tapering to a pointed tip in dorsal view; bulb bulged, with the sperm duct extending along the margin; embolus with fl atted, semicircular base, distal portion fl agelliform, and tip slightly pointed, reaching the cymbium apex; median apophysis ox horn-shaped in retrolateral view, the distal half sclerotized.  2F) similar to that of male except paler and with a pair of white oval patches on the dorsum of the abdomen. Epigyne ( Fig. 2A-C) wider than long; atrium large, separated by a ) (-shaped median septum; copulatory openings located bilaterally; copulatory ducts long, descending obliquely before extending transversely, and then descending posteriorly along the longitudinal axis, terminally with short gland ducts; spermathecae long, oval, touching each other anteriorly; fertilization ducts anterior to spermathecae.

Comments
At fi rst glance, the male and female look different in the markings of the abdomen. However, the male has blurry markings almost identical to the female. Furthermore, both of them were collected from the same locality and the sexual dimorphism in the morphology of the copulatory organs is consistent with sexual dimorphism in related species (S. gambosus and S. subgambosus sp. nov.). Based on this evidence, they are proposed to be the same species. Yang, Zhu & Song, 2007 Figs 3-4, 17B

Differential diagnosis
The male of Synagelides forkiforma can easily be distinguished from all other congeners by the presence of a short spine-shaped prong on the ventral surface of the retrolateral tibial apophysis. The female of S. forkiforma resembles that of S. lushanensis Xie & Yin, 1990(Xie & Yin 1990: fi gs 8-15) in having an anterior epigynal hood and a pair of lateral arcuated rims, but differs in: 1) epigynal hood is about one-third as long as epigynum in S. forkiforma (Fig. 4A), whereas about one-fi fth as long as epigynum in S. lushanensis (Xie & Yin 1990: fi g. 14); 2) median septum is almost triangular in S. forkiforma (Fig. 4A), whereas trapeziform in S. lushanensis (Xie & Yin 1990: fi g. 14). : patella slightly longer than wide in retrolateral view; retrolateral tibial apophysis slender and more than half as long as cymbium, with a ventral, short spine-shaped prong; the second retrolateral tibial apophysis paliform; cymbium hairy, with dorsal and prolateral apophyses; bulb infl ated, with sperm duct sinuous in retrolateral view; embolus spiraling and tapering to a slightly pointed tip that reaches cymbium apex; median apophysis complicated, with processes.

Material examined
Female (HNU-20040428-   1.22, 0.41). Habitus (Fig. 4F) similar to that of male except paler and lacking the pale transverse band on the dorsum of abdomen. Epigyne ( Fig. 4A-C) longer than wide, with an anterior hood about 2.5 times longer than wide and a pair of lateral arcuated rims; atrium large, separated by a triangular median septum; copulatory openings located postero-laterally; copulatory ducts almost S-shaped, terminating with gland ducts, which almost extend transversely; spermathecae reniform, touching each other anteriorly; fertilization ducts slightly longer than gland ducts, extending antero-obliquely.

Comments
The known drawings of the holotype of S. forkiforma are diffi cult to interpret due to poor print quality. Although the type specimens were not examined in the present contribution, the clear, original drawings of the holotype provided by Prof. Zi-Zhong Yang, who described the species, were compared with the specimens examined here from Fugong County. The male from Fugong is almost indistinguishable from the holotype, and the female from Fugong shares similar markings with the Fugong male. Fugong County is also near the type locality (the distance between the collecting site in Fugong and type locality is less than 150 kilometers) and both localities belong to Gaoligong Mountain and have similar climatic and environmental conditions. Based on the above evidence, the specimens collected from Fugong County are identifi ed here as new material of S. forkiforma.

Differential diagnosis
Synagelides leigongensis sp. nov. can easily be distinguished from all other congeners by: 1) retrolateral tibial apophysis is bifurcated basally (not bifurcated, or bifurcated medially or apically in that of other congeners); 2) epigyne lacks median septum and has a large anterior arcuated rim covering almost half the epigynal length.

Comments
The pairing of this species and the following ones are mainly based on the fact that both sexes were collected from the same sites and share similar markings. Moreover, the sexual dimorphism between the male and female of S. leigongensis sp. nov. is consistent with sexual dimorphism found in related species .

Differential diagnosis
The male of Synagelides logunovi sp. nov. can be easily distinguished from congeners by the presence of a retrolateral patellar apophysis, which is absent in all other congeners. The female of S. logunovi sp. nov. resembles that of S. yinae Liu, Chen, Xu & Peng, 2017(Liu et al. 2017: fi gs 7a-d, 8a-b) in the general shape of the copulatory organs, but differs in: 1) epigyne is wider than long in S. logunovi sp. nov. (Fig. 8A), whereas longer than wide in S. yinae (Liu et al. 2017: fi gs 7c, 8a); 2) the distance between the hood and the apex of median septum is almost as long as hood in S. logunovi sp. nov. (Fig. 8A), whereas more than two times as long as hood in S. yinae (Liu et al. 2017: fi gs 7c, 8a); 3) spermathecae are pyriform in S. logunovi sp. nov. (Fig. 8B-C), whereas elongated oval in S. yinae (Liu et al. 2017: fi gs 7d, 8b).

Etymology
The specifi c name is the patronym in honor of Dr Dmitri V. Logunov (Manchester, UK), who contributed signifi cantly to the taxonomy of the genus Synagelides.    (Fig. 8D) stippled, yellow-brown to dark-brown, eye base black, without distinct patches. Fovea oval, hollowed. Chelicerae (Fig. 8G) pale yellow, with two promarginal teeth and one retromarginal tooth. Endites (Fig. 8E) yellow, with paler prolateral side. Labium (Fig. 8E) brown, paler apically. Sternum (Fig. 8E) yellow. Legs pale yellow to yellow except femur I and the venter of patella I brown. Spination of leg I (Fig. 8H): femur v2-2-2-2-2; metatarsus v0-2-2. Abdomen ( Fig. 8D-E) elongated, contracted medially and slightly broadened posteriorly; dorsum dark-brown, with a narrow transverse pale stripe and two pairs of muscle depressions, posterior herringbone stripes indistinct; venter pale brown. Palp (Figs 7A-D, 17A): patella slightly longer than wide, distally with sclerotized and fl attened retrolateral apophysis; retrolateral tibial apophysis slender, about half as long as cymbium and pointed apically; cymbium hairy, with a distal apophysis; embolus coiled spirally, the basal half almost completely covered by median apophysis in retrolateral view; median apophysis sclerotized and with processes.  (Fig. 8F) similar to that of male except paler, lacking abdominal contraction and the transverse pale stripe on the dorsum of the abdomen. Epigyne (Fig. 8A-C) wider than long, with an anterior hood slightly longer than wide and a pair of lateral arcuated rims; atrium separated by a big, irregular median septum; copulatory openings located bilaterally; copulatory ducts S-shaped, terminally with short gland ducts extending obliquely; spermathecae pyriform, touching each other antero-medially; fertilization ducts extending transversely.

Differential diagnosis
Synagelides subgambosus sp. nov. closely resembles S. gambosus (Xie & Yin 1990: fi gs 1-7; Fig. 11D) in the shape of the copulatory organs and habitus but differs in: 1) the terminal edge of median apophysis is smooth in retrolateral view in S. subgambosus sp. nov. (Fig. 11B), whereas it is distinctly serrated in S. gambosus (Fig. 11D); 2) the middle portion of median apophysis is about four-fi fths the width of the terminal portion in retrolateral view in S. subgambosus sp. nov. (Fig. 11B), whereas it is about half the width of terminal portion in S. gambosus (Fig. 11D); 3) gland ducts are more than two-thirds as long as spermatheca in S. subgambosus sp. nov. (Fig. 12C), whereas less than one-fourth as long as spermatheca in S. gambosus (Xie & Yin 1990: fi g. 7). Synangelides subgambosus sp. nov. also somewhat resembles S. bohdanowiczi sp. nov. and the diagnostic features of this species are discussed above.

Etymology
The specifi c name refers to its similarity with S. gambosus, noun.  (Fig. 12D) stippled, red-brown, darker anteriorly, covered with thin hairs antero-medially. Fovea oval, hollowed. Chelicerae (Fig. 12G) yellow, with two promarginal teeth and one retromarginal tooth fi ssident. Endites, labium and sternum (Fig. 12E) yellow. Legs pale yellow to red-yellow except femora I and IV red-brown. Spination of leg I (Fig. 12H): femur v2-2-2-2; metatarsus v0-2-2. Abdomen ( Fig.  12D-E) elongated, dorsum with 2 pairs of muscle depressions and 2 transverse pale patches medially; venter gray-white, with a pair of pale brown lines extending from epigastric furrow to the terminus. Palp (Fig. 11A-C): patella swollen; tibia almost as long as wide; retrolateral tibial apophysis S-shaped, originating from the middle part of tibia; cymbium with a well-developed retrolateral apophysis; embolus enlarged at base, strongly curved medially and blunt at tip; median apophysis fl attened, slightly broadened terminally, and with a triangular process at the anterior edge in retrolateral view.  (Fig. 12F) similar to that of male except paler. Epigyne (Fig. 12A-C) slightly wider than long, with a pair of lateral arcuated rims; atrium large, almost covering two-thirds the epigyne, separated by a narrow median septum; copulatory openings located postero-laterally; copulatory ducts extending transversely at antero-median part and then descending posteriorly along the longitudinal axis, terminally with well-developed gland ducts which are longer than the fertilization ducts; spermathecae elongated, extending transversely and touching each other prolaterally; fertilization ducts short.

Differential diagnosis
Synagelides wuliangensis sp. nov. resembles S. hamatus Zhu, Zhang, Zhang & Chen, 2005(Zhu et al. 2005: fi g. 12a-e) in the shape of the copulatory organs and habitus, but differs in: 1) the second retrolateral tibial apophysis is about two times longer than wide in S. wuliangensis sp. nov. (Fig. 13B), whereas slightly wider than long in S. hamatus (Zhu et al. 2005: fi g. 12e); 2) retrolateral tibial apophysis tapers to a tail-shaped terminus in S. wuliangensis sp. nov. (Fig. 13B), whereas it becomes short, spine-shaped and terminates abruptly in S. hamatus (Zhu et al. 2005: fi g. 12e); 3) epigynal hood is more than three times longer than wide in S. wuliangensis sp. nov. (Fig. 14A), whereas almost as long as wide in S. hamatus (Zhu et al. 2005: fi g. 12b); 4) spermathecae are oval in S. wuliangensis sp. nov. (Fig. 14C), whereas reniform in S. hamatus (Zhu et al. 2005: fi g. 12c). Synagelides wuliangensis sp. nov. also somewhat resembles S. logunovi sp. nov. in the general shape of copulatory organs, but can be distinguished by the absence of the retrolateral patellar apophysis and the epigynal hood being more than three times longer than wide in S. wuliangensis sp. nov. (Fig. 174A), whereas the patella retrolateral apophysis is present and the epigynal hood is slightly longer than wide in S. logunovi (Fig. 8A).

Differential diagnosis
Synagelides xingdouensis sp. nov. can be easily distinguished from all other congeners by: male palp has three tibial apophyses in S. xingdouensis sp. nov. whereas none, one or two tibial apophyses in other congeners; spermathecae are mainly anterior to copulatory ducts and about three-fourth as long as epigynum in S. xingdouensis sp. nov., whereas mainly posterior to copulatory ducts and shorter than half the epigynal length in all other congeners.

Discussion
Within the genus Synagelides, fi ve groups were established by Bohdanowicz (1987): S. agriformis-group, S. annae-group, S. nishikawai-group, S. cavaleriei-group, and S. palpalis-group. Since then, surprisingly, no other species have been assigned to these groups. Based on morphology, many species, including some of the six new species described here, can be placed into one of these fi ve groups. For example, S. longus and S. palpaloides can be placed into the S. palpalis-group given that the habitus and copulatory organs of these two species are similar to S. palpalis. Other species can be placed together in new groups. S. subgambosus sp. nov. and S. gambosus can be placed together into a new group given that for both species, the male palp has a S-shaped retroalteral tibial apophysis and the female epigyne has a long and narrow median septum and a pair of lateral arcuated rims. Nevertheless, several species (e.g., S. xingdouensis sp. nov., S. leigongensis sp. nov. and S. yunnan Song & Zhu, 1998) are so unique that they cannot be placed into a group with any other species. Moreover, it is challenging to group species exclusively based on morphological characters from a limited number of specimens . Based on that, no species are formally assigned to existing groups or placed in new groups in the present contribution.
Like most salticid genera, one major problem with the taxonomy of Synagelides is that the known diversity may be overestimated given that many species are known only from a single sex and further investigations may show that some species known only from males may prove to be synonymous with other species known only from females and vice versa. However, most species of Synagelides have very localized distributions and mainly occur in mountains above 1000 meters, indicating more species could be discovered from remote areas and isolated mountains. Therefore, there is no doubt that the true diversity of the genus remains poorly studied.