﻿Midget cave spiders (Araneae, Leptonetidae) from Jiangxi and Fujian Province, China

﻿Abstract Eleven leptonetid species belonging to four genera collected in Jiangxi and Fujian Provinces, China are presented. Ten new species of midget cave spiders from southern China are diagnosed, described, and illustrated: Leptoneteladawu Yao & Liu, sp. nov., L.yuanhaoi Yao & Liu, sp. nov. and L.zuojiashanensis Yao & Liu, sp. nov. from Jiangxi; Longileptonetaguadunensis Yao & Liu, sp. nov., L.huboliao Yao & Liu, sp. nov., L.jiaxiani Yao & Liu, sp. nov., L.letuensis Yao & Liu, sp. nov., L.renzhouensis Yao & Liu, sp. nov., L.tianmenensis Yao & Liu, sp. nov., and Pararanamingxuani Yao & Liu, sp. nov. from Fujian. Furthermore, Falcileptonetamonodactyla (Yin, Wang & Wang, 1984) is recorded from Jiangxi province for the first time. Distributions records are given for all investigated species.


Introduction
The midget cave spider family Leptonetidae Simon, 1890 is one of the smallest taxa in haplogyne spider families, with 374 species belonging to 22 genera (WSC 2023).At present, eight genera and 135 species are known from China.Among these genera, species of Leptonetela Kratochvíl, 1978 have been reported as being the most diverse in China.
Most records and descriptions of this family from China were contributed by the Chinese arachnologist Shu-Qiang Li and his team, such as of the genera Jingneta Wang & Li, 2020, Leptonetela, Longileptoneta Seo, 2015, Pararana Lin & Li, 2022and Rhyssoleptoneta Tong & Li, 2007(Wang and Li 2011;Wang et al. 2017Wang et al. , 2020;;Lan et al. 2021;Zhu and Li 2021;Lin et al. 2022).In addition, many more genera have been recorded or described from China by other authors with eight species from northern provinces (Tong and Li 2008;Wang et al. 2020;Zhu and Li 2021;Liu and Zhang 2022), and the remaining species from the southern provinces of China.Despite advances in the taxonomic knowledge of the family, there are still many more genera and species to discover from southern China that have unusual morphological characteristics.
While working on the leptonetid fauna of the Jiangxi and Fujian provinces, southern China, we discovered and examined in detail eleven species including one known and ten new leptonetids.The goal of this paper is to formally describe the new species and to report the first species of Falcileptoneta Komatsu, 1970 from Jiangxi Province.

Materials and methods
Specimens were examined using a Zeiss Stereo Discovery V12 stereomicroscope with a Zoom Microscope System.Both male palps and female genitalia were detached and examined in 80% ethanol, using a Zeiss Axio Scope A1 compound microscope with a KUY NICE CCD.The female genitalia were cleared in trypsin enzyme solution to dissolve soft tissues.For SEM photographs, specimens were dried under natural conditions, coated with gold using a small ion-sputtering apparatus ETD-2000, or without coating, and examined with a ZEISS EVO LS15 scanning electron microscope.Specimens including detached male palps and female genitalia were stored in 75% ethanol after examination.All the specimens are deposited in Animal Specimen Museum, Life Science of College, Jinggangshan University (ASM-JGSU).
To maintain uniformity of genitalia terminology within these genera, including Falcileptoneta, Leptonetela, Longileptoneta, and Pararana, the terms that are used are primarily from the Spider Anatomy Ontology on BioPortal (Ramírez and Michalik 2019).In the past, different terms have been used to refer to the same structure, and terms have been used incorrectly.Although some of these terms have synonyms in both males and females, the ones used here will hopefully become a standard for future studies of these genera, if applicable.Measurements were taken with the Axio Vision software (SE64 Rel.4.8.3) and are given in millimeters.Leg measurements are given as total length (femur, patella, tibia, metatarsus, tarsus).Distribution.Known from Jiangxi (new record) and Hunan (Liu et al. 2020), China (Fig. 29).Diagnosis.The male of this species is similar to Leptonetela sexdentata Wang &Li, 2011 (see Wang andLi 2011: 15, figs 53A-D) in having a tongueshaped prolateral lobe, but can be distinguished from it by dorsal habitus with obvious black-brown stripes (vs pale in L. sexdentata) and the foot-shaped median apophysis (vs square-shaped) (Figs 3, 8A-D).Females resemble that of Leptonetela rudong Wang & Li, 2017(see Wang et al. 2017: 362, fig. 31C) in having a sub-rectangular atrium, but can be separated from it by the spermathecal stalk with seven spirals (vs six) and the slightly curved spermathecae (vs straight) (Fig. 4C).
Coloration (Fig. 3A).Carapace yellow to dark brown, with radial, dark brown, mottled markings on lateral margin and mottled stripes medially.Chelicerae, endites, labium, and sternum yellow-brown.Legs yellow, with distinct annulations.Abdomen pale to dark brown, with five dark chevron-shaped stripes.Palp .Tibia with four long setae retrolaterally, the basal one thick, two short spines distally; cymbium with one thick, conspicuous spine prolaterally, one short, thick spine retrolaterally, and one long spine distally.Tip of bulb: prolateral lobe finger-like; median apophysis relatively long, foot-shaped, distal margin with ten teeth, the retrolateral one very large with blunt tip; conductor long, membranous, apically curved; embolus spine-like, very short, under the conductor.
Distribution.Known only from the type locality in Jiangxi Province, China (Fig. 29).
Etymology.The name is taken from the type locality; noun in apposition.Diagnosis.The male of this species is similar to that of Leptonetela sexdentata Wang &Li, 2011 (Wang andLi 2011: 15, fig.53B-D) in having the tibia with a row of spines retrolaterally including one thick strong spine proximally and three thin spines, but can be separated from it by the tongue-shaped prolateral lobe (vs finger-like) and the median apophysis with narrow base (vs broad) and five teeth distally (under microscope) (vs six) (Figs 5B-D, 8E-L).The males also resemble that of L. dawu sp.nov. in having the conductor with curved apex and the spine-like embolus, but can be separated from it by the leaf-shaped median apophysis (vs foot-shaped) (Figs 5B-D, 8E-L).The female can be easily distinguished from L. sexdentata (Wang and Li 2011: 15, fig.54C) by the transversely extended spermathecal stalk (vs directed anteromedially) with four regular spirals (vs irregular) (Fig. 6C).
Coloration (Fig. 5A).Carapace yellow to dark brown, with dark radial stripes and mottled markings on lateral margin, and an oval dark brown band medially.Chelicerae yellow.Endites yellow to dark brown, mottled.Labium yellow to dark brown.Legs with dark annulations on each segment except tarsi.Abdomen with four dark chevron-shaped stripes.
Palp (Figs 5B-D, 8E-L).Tibia with five long setae retrolaterally, the proximal one very thick, long, strong, spine-like; cymbium with one long conspicuous seta prolaterally, one short, strong, thick spine retrolaterally and one long spine distally.Tip of bulb: prolateral lobe tongue-like, relatively short; median apophysis leaf-shaped, distal margin with four to ten teeth, prolateral one very small, retrolateral one very large with triangular tip; conductor membranous, relatively broad, near the base of median apophysis, longer than median apophysis; embolus short, transparent, broad, slightly bending retrolaterally.
Distribution.Known only from the type locality in Jiangxi Province, China (Fig. 29).
Etymology.The species is named after Mr Yuanhao Ying, who collected the type specimens.
Comments.We compared the palps of the new species with that of the very similar species L. dawu sp.nov.several times.This similarity is probably because the males of these two species have very similar characters: tibial spines, cymbial spine, and embolus.The distance between Dawu Mountain and Ziyao Mountain is approximately 25 km (linear distances), which is very close.Despite the close distance, we consider them as different species based on the morphological differences listed.This hypothesis will be confirmed or rejected in the future when molecular data and analysis can be provided.Diagnosis.The male of this species is similar to that of Leptonetela gubin Wang & Li, 2017(in Wang et al. 2017: 386, fig. 48B-D) in having the curved cymbium forming an angle of ca 100° with tibial axis and the horn-like  prolateral sclerite, but can be separated from it by the tibia having two spines including one very thick and strong spine proximally (vs a row of spines and lacking a thick and strong spine) (Figs 7B-D, 8M-P).It also resembles that of L. mengzongensis Wang &Li, 2011 (Wang andLi 2011: 10, fig. 24B-D) in having the horn-like prolateral sclerite, but can be easily distinguished from it by the tibia with a thick and strong proximal spine (vs slender) (Figs 7B-D, 8M, N).
Coloration (Fig. 7A).Carapace yellow to dark brown, with dark radial stripes and mottled markings on lateral margin.Chelicerae yellow.Endites yellow, with mottled dark spots.Labium, anterior part dark brown, posterior part yellow.Sternum dark brown, medially with a yellow stripe.Legs yellow to dark brown.Abdomen with three pairs of dark brown spots and three dark chevron-shaped stripes.
Female.Unknown.Distribution.Known only from the type locality in Jiangxi Province, China (Fig. 29).
Etymology.The name is taken from the type locality.Diagnosis.This species is similar to that of Longileptoneta shenxian Wang & Li, 2020(in Wang et al. 2020: 698, fig.12A-D) and L. yamasakii Ballarin &Eguchi, 2022 (Ballarin andEguchi 2022: 373, figs 1C, 3A-C) in having the banded median apophysis with transparent tip, but can be distinguished from it by the carapace with six eyes (vs absent in L. shenxian), the tibia with one canine tooth-like apophysis armed with a short straight spine (vs one columnar apophysis, armed with one long, curved spine in L. shenxian and L. yamasakii) and the wedge-shaped prolateral lobe (vs mastoid in L. shenxian and sub-triangular in L. yamasakii) (Figs 9C-E, 10D, E).
Palp (Figs 9C-E, 10).Femur with three rows of long strong spines ventrally, dorsally, and prolaterally; patella without spine; tibia with two spines: one canine tooth-like apophysis, armed with a short straight spine subapically, and a thick and long spine near the base of anterior one.Cymbium with a constriction subapically.Tip of bulb: one long banded median apophysis, distally fur- cate; prolateral lobe wedge-shaped, large; conductor membranous, anteriorly curved.Embolus indistinct, with broad base.
Female.Unknown.Distribution.Known only from the type locality in Fujian Province, China (Fig. 29).
Etymology.The name is taken from the type locality.Diagnosis.The male of this species is similar to that of Longileptoneta shenxian Wang & Li, 2020(in Wang et al. 2020: 698, fig.12A-D) in having the bulb with an extruded coniform anterior part and three rows of spines, but can be distinguished from it by the carapace with six eyes (vs absent), the tibia with one long columnar apophysis armed with a long spine (vs one short columnar apophysis, armed with one long, curved spine) and the long needle-like prolateral sclerite (vs thick) (Figs 11,12).The female can be easily separated from L. shenxian (Wang et al. 2020: 698, fig.13C) by the bell-shaped atrium, but can be separated by the carapace with six eyes (vs absent) and the distal spermathecal stalk lacking a coil (vs present) (Fig. 13).
Note.The right spermathecal stalk and spermathecae were extruded deformation after covering cover slip when we took a photo under microscope.
Distribution.Known only from the type locality in Fujian Province, China (Fig. 29).

Description. Male (holotype). Habitus as in
Coloration (Fig. 14A, B).Carapace yellowish to black, with radial dark stripes submedially and mottled markings on lateral margin.Chelicerae yellow to dark brown.Endites yellow.Labium yellow.Sternum yellow to black, mottled.Legs yellow.Abdomen, dorsally yellow, mottled in anterior and posterior parts; venter mottled.
Distribution.Known only from the type locality in Fujian Province, China (Fig. 29).
Etymology.The species is named after Mr Jiaxian Gong, who collected the type specimens.Diagnosis.The male of this species can be easily distinguished from other members of this genus by the very large curved tibial apophysis armed with a short spine-like tip (Fig. 17E).
Palp 18).Femur with two rows of short strong spines ventrally and prolaterally; patella without spine; tibia with a very long curved apophysis, distally armed with a short straight spine, longer than tibia.Cymbium with a distinct constriction medially.Tip of bulb: one long flagelliform prolateral sclerite; prolateral lobe oval; conductor membranous, medially with a groove.Embolus indistinct.
Female.Unknown.Distribution.Known only from the type locality in Fujian Province, China (Fig. 29).
Etymology.The name is taken from the type locality.Diagnosis.The male of this species can be easily distinguished from other members of this genus by the very long curved spine-like prolateral sclerite with a feathery tip 20).The female resembles L. zhuxian Wang & Li, 2020(Wang et al. 2020: 700, fig.16C) in having spheroid spermathecae and the subtrapezoid atrium, but can be separated by the slightly curved spermathecal stalk (vs waved) (Fig. 21C).
Palp 20).Femur with three rows of short strong spines ventrally, dorsally, and prolaterally; patella lacking spine; tibia with one columnar apophysis, armed with one straight spine, and one long and very thick spine, slightly shorter than the apophysis.Cymbium with a distinct constriction medially.Tip of bulb: one broad median apophysis, with a furcate tip, including one long membranous and one hook-shaped; prolateral lobe finger-like; conductor membranous, touching with prolateral sclerite; prolateral sclerite waved, needle-like, with a feathery tip.Embolus short hook-shaped, with a broad base.
Distribution.Known only from the type locality in Fujian Province, China (Fig. 29).
Etymology.The name is taken from the type locality.Diagnosis.The male of this species is similar to that of L. huboliao sp.nov. in having the femur with three rows of strong spines, one columnar tibial apophysis, armed with a short spine-like tip on palp and a needle-like prolateral sclerite, but can be easily distinguished from it by the rod-like median apophysis (vs lacking) the membranous conductor lacking a serrulate tip (vs present) 23).The female resembles L. huboliao sp.nov. in having a bell-like atrium and the spermathecal stalk lacking a spiral twist, but can be separated by the oval spermathecae (vs tube-shaped) (Fig. 24C).
Coloration (Fig. 22A, B).Carapace yellowish to black-brown, with dark radial stripes on lateral margin.Chelicerae yellow to brown, mottled.Labium dark brown.Sternum yellow to black, mottled.Legs yellow, mottled.Abdomen, dorsally yellowish, with two pairs of spots; venter dark.
Palp 23).Femur with three rows of short strong spines ventrally, dorsally, and prolaterally; patella lacking strong spine; tibia with a columnar apophysis, armed with one short straight spine.Cymbium with a distinct constriction medially.Tip of bulb: one rod-like median apophysis, thick; prolateral lobe oval, lamellar; conductor membranous, shorter than median apophysis.Embolus hook-shaped, beneath conductor.
Etymology.The name is taken from the type locality.Diagnosis.The male of this species is similar to that of Pararana gaofani Lin & Li, 2022 (Lin et al. 2022: 217, figs 17A-C, 18A, B) in having the cymbium with a notch and the swollen patella, but can be easily separated by the patella with seven short tooth-like spines (vs four long relatively thick spines), the tibia with a thick spine (vs absent), the long lamellar median apophysis (vs the relatively short horn-like median apophysis) and the slightly curved rod-like embolus (vs horn-like) 26).The female can be easily distinguished by the oval atrium and the short S-shaped spermathecal stalk (Fig. 27C).
Coloration (Fig. 25A).Carapace yellow, with dark radial stripes.Chelicerae yellow to brown.Endites yellow to dark brown.Labium yellow brown.Legs with dark annulations on each segment except tarsi.Abdomen yellow.
Palp 26).Femur with four rows of short strong spines ventrally, dorsally, and prolaterally; patella expanded, with seven stout spines; tibia with a very thick spine retrolaterally.Cymbium with a notch subapically.Bulb: prolateral lobe banded, long; embolus rod-like, slightly curved, with a broad base and a net-shaped surface; median apophysis lamellar, shorter than conductor; conductor membranous, with serrulate margin; retrolateral lobe blunt, tongueshaped, touching base of conductor.
Distribution.Known only from the type locality in Fujian Province, China (Fig. 29).
Etymology.The species is named after Mr Mingxuan Wu, who collected the type specimens.

Discussion
At present, China is the most diverse region for Leptonetidae (144 species), followed by the USA (60 species), Korea (58 species), Japan (53 species), France   2017) and the Japanese species by Ballarin and Eguchi (2022).These two studies have revealed remarkable sexually dimorphic traits and relevant  It is interesting to note that the species Falcileptoneta monodactyla has no also been found from Jiangxi Province.Considering the locality of the holotype, Yanling county in Hunan province, it is likely that this species is more widely distributed in the Hunan and Jiangxi provinces.
The Longileptoneta species are very difficult to differentiate as their embolus is hidden in the tegular apophyses.Although the genera Falcileptoneta and Longileptoneta are clearly distinguished from all other leptonetid genera, ambiguity can occur in cave species lacking eyes, such as Falcileptoneta taizhensis (Chen & Zhang, 1993), Longileptoneta gutan Wang &Li, 2020, andL. shenxian (Wang et al. 2020).A very obvious feature reveals that they are living in caves and become vestigial.Each of these two genera seems to be monophyletic, which need to be confirmed by future phylogenetic studies.
The genus Pararana Lin & Li, 2022 was monotypic before this work and described based on a single male specimen (Lin et al. 2022).The diagnostic characters of the genus are inadequate as no females are known for the type species Pararana gaofani Lin & Li, 2022(Lin et al. 2022).Based on the female of P. mingxuani sp.nov., this genus can be characterized by a long atrium and very short spermathecal stalks.Since more Pararana species from China can be expected to be discovered, this genus will be more easily understood in future research.

Figure 8 .
Figure 8. SEM micrographs of male palps, Leptonetela spp.A Leptonetela dawu sp.nov., left palp, ventral view B same, ventral view C same, detail of tegular apophysis, ventral view D same, detail of conductor and embolus, ventral view E L. yuanhaoi sp.nov., left palp, ventral view F same, ventral view G same, detail of tegular apophysis, ventral view H same, detail of conductor and embolus, ventral view I L. yuanhaoi sp.nov., right palp, ventral view J same, ventral view K same, detail of tegular apophysis, ventral view L same, detail of conductor, ventral view M L. zuojiashanensis sp.nov., right palp, tibial spine, prolateral view N same, detail of the tip, prolateral view O same, bulb, ventral view P same, detail of tegular apophysis, ventral view.Abbreviations: Con -conductor, Em -embolus, MA -medial apophysis, PS -prolateral sclerite.

Figure 10 .
Figure 10.SEM micrographs of Longileptoneta guadunensis sp.nov., right palp, male holotype A ventral view B tibial apophysis, ventral view C detail of the tip of tibial apophysis, ventral view D detail of tegular apophysis, ventral view E same, ventral view.Abbreviations: Con -conductor, Em -embolus, MA -medial apophysis.

Figure 18 .
Figure 18.SEM micrographs of Longileptoneta letuensis sp.nov., male right palp A femur, retrolateral view B tibial apophysis, retrolateral view C detail of the tip of tibial apophysis, retrolateral view D bulb, ventral view E same, detail of tegular apophysis, ventral view F same, detail of conductor, ventral view.Abbreviations: Con -conductor, PS -prolateral sclerite.

(
26), and Greece (16), with few species found in the remaining regions (29 species; WSC 2023).Within 15 years (2008 to 2022) the total number of leptonetid species recorded from China increased six times (WSC 2023), mostly after the profusely illustrated revisions of the Chinese representatives by Wang et al. (