Thirty-one new species of the spider genus Leclercera from Southeast Asia (Araneae, Psilodercidae)

Abstract Thirty-one new species of the genus Leclercera Deeleman-Reinhold, 1995 from China, Indonesia, Malaysia, Myanmar, Nepal, and Thailand are described: L. mianqiusp. nov. (♂♀), L. thamsangensissp. nov. (♂♀), L. yandousp. nov. (♂♀), L. thamkaewensissp. nov. (♂♀), L. xiangbabangsp. nov. (♂♀), L. jianzuiyusp. nov. (♂♀), L. yamaensissp. nov. (♂♀), L. banensissp. nov. (♂♀), L. dumuzhousp. nov. (♀), L. suwanensissp. nov. (♂♀), L. maochongsp. nov. (♀), L. shanzisp. nov. (♀), L. duandaisp. nov. (♂♀), L. hponensissp. nov. (♂♀), L. lizisp. nov. (♂), L. xiaodaisp. nov. (♀), L. yanjingsp. nov. (♀), L. ekteenensissp. nov. (♂), L. zhamensissp. nov. (♂), L. sanjiaosp. nov. (♀), L. selasihensissp. nov. (♂♀), L. paiensissp. nov. (♀), L. yuanzhuisp. nov. (♀), L. zanggaensissp. nov. (♀), L. aniensissp. nov. (♂♀), L. renqinensissp. nov. (♂♀), L. shergylaensissp. nov. (♂♀), L. pulongensissp. nov. (♂), L. tudaosp. nov. (♂♀), L. duibaensissp. nov. (♂), and L. jiazhongensissp. nov. (♂♀). Types are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS) in Beijing.


Introduction
The spider family Psilodercidae Machado, 1951 was previously a subfamily of Ochyroceratidae Fage, 1912until Wunderlich (2004 elevated it to the family level. To date, Psilodercidae includes a total of 165 species in 11 genera (WSC 2019), and the family has been shown to be monophyletic (Li and Li 2018). It is distributed in Southeast Asia, southern China, and parts of South Asia (WSC 2019, Li et al. 2020). The number of species has ballooned to almost three times its size in the 21 st century, from only 53 species known by the end of 20 th century (Platnick 2000). More than half of the psilodercid genera, including Flexicrurum Tong & Li, 2007, Luzonacera Li & Li, 2017, Priscaleclercera Wunderlich, 2017, Qiongocera Li & Li, 2017, Relictocera Li & Li, 2017, Sinoderces Li & Li, 2017, and Thaiderces Li & Li, 2017, have been described only recently.
While examining spider collections from tropical Asia, we found 31 new species of Leclercera from China, Indonesia, Malaysia, Myanmar, Nepal, and Thailand. The goal of this paper is to provide detailed descriptions of these new species with images of their copulatory organs and chelicerae.

Materials and methods
Types are deposited in the Institute of Zoology, Chinese Academy of Sciences (IZ-CAS) in Beijing. All specimens collected were studied and preserved in 95% ethanol. The specimens were measured and examined with a Leica M205 C stereomicroscope, and further morphological details were observed with an Olympus BX41 compound microscope. Male palps were detached from the left side of the animal for further examination (except for Leclercera xiangbabang sp. nov. whose right palp was detached). Carapace length was measured excluding the clypeus. Internal genitalia of the female and palpal bulbs were dissected and immersed in lactic acid. An Olympus C7070 wide zoom digital camera (7.1 megapixels) mounted on an Olympus SZX12 stereomicroscope was used to take photos at different focal planes. The photos were then transferred to the image stacking software Helicon Focus 6.7.1 to generate photos with a greater depth of field before further processing with Adobe Photoshop CC 2014. Leg measurements are shown as total length: femur, patella, tibia, metatarsus, and tarsus. Leg segments were measured from their retrolateral side. All measurements are given in millimetres (mm). All terminology follows that of Li et al. (2014).    Etymology. The species name is an adjective referring to the type locality. Diagnosis. Males of L. thamsangensis sp. nov. can be distinguished from congeners by the presence of two spines on a small retrolateral protrusion (retrolateral apophysis) of the cymbium (Fig. 4D) (vs. the absence of a retrolateral apophysis with two spines on the cymbium), the conductor and embolus are not widely separated, appearing to be similar in length and width (Fig. 4B) (vs. conductor and embolus different in congeners); females can be differentiated from congeners by rectangular, sheet-like spermathecae ( Fig. 3A) (vs. absence of sheet-like spermathecae in congeners).
Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "yāndǒu" (smoking pipe) and refers to the palpal bulb which resembles a smoking pipe (Fig. 6B).  Diagnosis. Males of L. yandou sp. nov. can be distinguished from congeners by the structure of the bulb, with a rounded base bearing a slightly curved and elongated embolus (Fig. 6B), the presence of a swollen triangular tibia with a retrolateral apophysis ( Fig. 6D) (vs. absence of swollen triangular tibia in congeners); females can be differentiated from congeners by a pair of saucer-shaped, sinuous spermathecae (Fig. 5A) Description. Male (Holotype). Total length 2.50; carapace 0.75 long, 0.94 wide; abdomen 1.75 long, 0.80 wide. Carapace round and brown, with three longitudinal dark brown bands, median band twice as wide as lateral band (Fig. 5C). Chelicerae brown (Fig. 55I). Clypeus brown. Endites dark brown, light brown basally. Labium dark brown. Sternum brown, with dark brown patched laterally. Abdomen elongated, dorsum with 3 pairs of dark brown spots medially, median dark brown bands concentrated posteriorly, antero-ventrally brown with elliptical patch, posterior part with indistinct dark and light brown pattern. Legs uniformly brown; measurements: I 8. 54 (2.81,0.25,2.97,1.88,0.63),II missing,III missing,IV 8.09 (2.40,0.25,2.19,2.25,1.00). Palp ( Fig. 6A-D): femur slender, five times longer than patella; patella not swollen; tibia swollen, 1.5 times shorter than femur, forming a triangular shape with a retrolateral apophysis bearing a spine; cymbium two times shorter than femur, dark brown anteriorly; bulb spatulate with circular base, elongated embolus arises distally; embolus slightly bent, two times longer than the length of rounded tegulum and half the width of tegulum (Fig. 6B).
Female ( Etymology. The species name is a noun in apposition derived from the Chinese pinyin "xiàngbábàng" (geoduck) and refers to the entire structure of the bulb which resembles the appearance of a Pacific geoduck bivalve.
Female ( Etymology. The species name is a noun in apposition derived from the Chinese pinyin "jiānzuĭyú" (bird wrasse -a type of fish) and refers to the entire structure of bulb which resembles the mouth of a bird wrasse.
Diagnosis. Diagnostic features of males and females are discussed in the diagnosis of L. xiangbabang sp. nov.
Female ( 11B): a pair of dark brown horizontal patches laterally. Endogyne (Fig. 11A): a pair of circular spermathecae with thin strips laterally, wavy patterns in spermathecae, lateral stripes two times longer than circular spermathecae.
Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is an adjective referring to the type locality. Diagnosis. Males of L. yamaensis sp. nov. can be distinguished from congeners by an indentation on the anterior end of the palpal tibia formed by the presence of a retrolateral apophysis bearing a spine that is longer than the tegulum (Fig. 14D) (vs. the absence of such an indentation from a retrolateral apophysis in congeners), the presence of two strong setae on the retrolateral apophysis ( Fig. 14D) (vs. the absence of setae on the retrolateral apophysis in congeners), the presence of a laminar apophysis adhering to the embolus (vs. absence of laminar apophysis, or if present, widely separated from the embolus in congeners); females can be differentiated from congeners by a pair of hook-like spermathecae (Fig. 13A).
Female ( Etymology. The species name is an adjective referring to the type locality. Diagnosis. Males of L. banensis sp. nov. resemble L. suwanensis sp. nov. but can be distinguished by an embolus that is longer and almost as wide as the conductor (Fig. 16B) (vs. an embolus that is equally as long as the conductor but almost four times wider basally (Fig. 19B)), bulb rather slender (vs. bulb rather expanded), patella swollen, cymbium not swollen (vs. patella not swollen, cymbium swollen), hexagonal swollen tibia (vs. pentagonal swollen tibia), tibia with three retrolateral apophyses, each bearing a spine ( Fig. 16D) (vs. tibia with a single retrolateral apophysis bearing a spine and a strong seta (Fig. 19D)); females can be differentiated by a pair of flattened spermathecae with an anterior extension laterally ( Fig. 15A) (vs. a pair of bulging, curled spermathecae with posterior extension laterally (Fig. 18A)).
Female (Paratype). General features and coloration similar to those of the male (Fig. 15D  Etymology. The species name is a noun in apposition derived from the Chinese pinyin "dúmùzhōu" (canoe) and refers to the structure of the spermathecae which resembles a canoe.
Diagnosis. Female of L. dumuzhou sp. nov. resembles the female of L. banensis sp. nov. but can be differentiated by the antero-ventral dark brown band surrounding the external genitalia which does not extend across the entire width of the abdomen ( Fig. 17B) (vs. the dark brown transverse band fully covering the entire width of the abdomen (Fig. 15B)), spermathecae slightly curved anteriorly with tip directed laterally ( Fig. 17A) (vs. spermathecae flattened with tubular extensions laterally (Fig. 15A)).
Female (Paratype). General features and coloration similar to those of male (   Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "máochóng" (caterpillar) and refers to the structure of spermathecae which resemble a caterpillar in lateral view.
Diagnosis. Males of L. maochong sp. nov. resemble L. shanzi sp. nov. but can be distinguished by a pair of divided conductors (Fig. 21B) (i.e. -consisting of two components) (vs. a slightly twisted undivided conductor (Fig. 23B)), an absence of a laminar apophysis adjacent to embolus (vs. presence of a laminar apophysis adjacent to embolus), embolus two times longer than tegulum (vs. embolus of similar length to tegulum), presence of two dorsal apophyses anteriorly on tibia (Fig. 21D) (vs. the presence of two retrolateral tibial apophyses anteriorly (Fig. 23C)); females can be recognized by having more coils of the spermathecae and by having a pair of lateral, spherical structures connected via the duct system to the posterior ends of the spermathecae ( Description. Male (Holotype). Total length 2.58; carapace 0.88 long, 1.20 wide; abdomen 1.70 long, 0.80 wide. Carapace round and brown, with three dark brown longitudinal bands, median band two times wider than lateral band (Fig. 20C). Chelicerae brown (Fig. 55G). Clypeus with dark brown band medially. Endites dark brown, light brown basally. Labium dark brown. Sternum dark brown with vertically thin median light brown band. Abdomen elongated, dorsum with pairs of dark brown spots, posterior with dark brown horizontal stripes, antero-ventrally with a pair of ovoid brown patches followed by rectangular dark brown patch, posterior with complicated dark brown pattern. Legs uniformly brown; measurements: I-IV missing. Palp (Fig. 21A-D): femur slender, four times longer than patella; patella not swollen; tibia swollen, 1.5 times shorter and 3 times wider than femur, with two dorsal apophyses anteriorly (apophyses almost as long as cymbium); cymbium three times shorter than femur, dark brown distally; bulb brown, pyriform, with conductor and embolus arising distally; conductor comprises two components, basally and distally merged with embolus; embolus elongated and sheet-like, slightly twisted, two times longer than tegulum (Fig. 21B).    (Fig. 20B): rectangular dark brown patch anteriorly, followed by brown crescent-shaped slit posteriorly. Endogyne (Fig. 20A): spermathecae resembles a crawling caterpillar in lateral view, with pointed ends, connected posteriorly to a pair of lateral spherical bodies by a duct system double-looped like a shallow "W".
Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "shànzĭ" (fan) and refers to the resemblance of the external genitalia to a hand fan.
Diagnosis. Diagnostic features of males and females are discussed in the diagnosis of L. maochong sp. nov.
Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "duàndài" (ribbon) and refers to the structure of the spermathecae resembling a ribbon knot (Fig. 24A).
Diagnosis. Females of L. duandai sp. nov. can be distinguished from other congeners by the unique orange coloration of the external genitalia and the spermathecae (Fig. 24A, B) which resemble a pair of orange-coloured ribbon knots (vs. absence of orange coloration on the external genitalia or spermathecae in congeners).
Male. Unknown. Distribution. Known only from the type locality (Fig. 58).  Etymology. The species name is an adjective referring to the type locality. Diagnosis. Males of L. hponensis sp. nov. can be distinguished from congeners by the presence of a three-branched, laminar apophysis on the distal end of the bulb (Fig. 26B) (vs. the absence of a three-branched laminar apophysis in congeners), a spheroid bulb (vs. the bulb of congeners have other shapes), tibia with a retrolateral apophysis bearing a spine that is three times shorter than the apophysis, the entire apophysis, including the spine, is two times longer than the tegulum (Fig. 26D) (vs. the absence of such a combination of a tibia apophysis and spine in congeners); the female can be differentiated from congeners by the pair of stalked spermathecae with a triangular distal part (Fig. 25A).
Female (Paratype). General features and coloration similar to those of male (Fig.  25D (Fig. 25B): dark brown patch delimiting a horizontal light brown slit. Endogyne (Fig. 25A): a pair of stalked spermathecae with anterior stalks and posteriorly a triangular distal part.
Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "lízĭ" (pear) and refers to the structure of the bulb resembling a pear (Fig. 28B). Diagnosis. Males of L. lizi sp. nov. can be distinguished from congeners by the abundance of apophyses and spines on the palp (Fig. 28C, D): cymbium with two retrolateral apophyses posteriorly, seven retrolateral apophyses on the swollen tibia with an anterior dorsal apophysis bearing two spines; embolus almost as long as the tegulum (Fig. 28B) (vs. the absence of congeners with such a profusion of apophyses and spines).
Female. Unknown. Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "xiǎodài" (small pouch) and refers to the distinct, pouch-like structure of the external genitalia (Fig. 29B).
Diagnosis. Females of L. duandai sp. nov. can be distinguished from other congeners by the unique, pouch-like external genitalia (Fig. 29B) and a pair of fusiform spermathecae with two pairs of apophyses extending both anteriorly and posteriorly (Fig. 29A) (vs. an absence of pouch-like external genitalia in congeners).
Description. Female. Total length 2.44; carapace 0.88 long, 0.90 wide; abdomen 1.56 long, 1.25 wide. Carapace round and brown, with three longitudinal dark brown bands, median band three times wider than lateral bands (Fig. 29C). Chelicerae brown (Fig. 55A). Clypeus light brown. Endites, labium, and sternum dark brown. Abdomen elongated, dorsum with indistinct dark brown spots (Fig. 29C), antero-ventrally with a pair of rounded dark brown patches laterally, followed by cone-shaped external genitalia resembling a small pouch, posterior dark brown (Fig. 29D)   Etymology. The species name is a noun in apposition derived from the Chinese pinyin "yǎnjìng" (spectacles) and refers to the structure of the spermathecae resembling a pair of spectacles (Fig. 30A).
Diagnosis. Females of L. yanjing sp. nov. can be distinguished from other congeners by a unique curved 'x' spot on the external genitalia (Fig. 30B) and the spermathecae which resemble a pair of aviator glasses (Fig. 30A) (vs. absence of such characteristics in congeners).
Male. Unknown. Distribution. Known only from the type locality (Fig. 58).  (Fig. 32B) (vs. the absence of a two-branched conductor in congeners), the embolus arising from medial tegulum (vs. embolus arising distally in congeners), the presence of a distinct darkened apophysis anteriorly on the tibia (Fig. 32D) (vs. the absence of a darkened tibial apophysis in congeners).
Description. Male (Holotype). Total length 3.09; carapace 0.90 long, 1.00 wide; abdomen 2.19 long, 0.94 wide. Carapace round and brown, with three dark brown longitudinal bands, median band five times wider than the lateral bands (Fig. 31A). Chelicerae brown (Fig. 56B). Clypeus light brown, with dark brown band medially. Endites dark brown, light brown basally. Labium dark brown. Sternum with dark brown stripes laterally, delimiting light brown anterior and median region. Abdomen elongated, antero-dorsally with three pairs of dark brown spots laterally, light brown medially, posterior with dark brown stripes medially, antero-ventrally with dark brown elliptical patch, posterior with indistinct dark and light brown spots, posterior edge : femur slender, four times longer than patella; patella not swollen; tibia swollen, 1.5 times shorter and wider than femur, antero-dorsally with a black apophysis, almost as long as tegulum; cymbium with dark brown spots, two times shorter than femur; bulb light brown, pyriform, with conductor arising distally, embolus arising medially; conductor dark and thin with two branches, one shorter than the other, short branch bends towards long branch; embolus arising from median tegulum, attached to short branch of conductor, almost as long as long branch of the conductor (Fig. 32B).
Female. Unknown. Distribution. Known only from the type locality (Fig. 58).   Etymology. The species name is an adjective referring to the type locality. Diagnosis. Males of L. zhamensis sp. nov. can be distinguished from congeners by the laminar-shaped embolus that is basally fused with the conductor (Fig. 34B) (vs. the absence of a basally fused conductor and embolus in congeners); the presence of a retrolateral apophysis on the tibia with a spine four times shorter than the apophysis (Fig. 34D) (vs. the absence of a spine with such characteristics in congeners).
Female. Unknown. Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "sānjiǎo" (triangle) and refers to the distinct triangular shape of the external genitalia.
Diagnosis. Females of L. sanjiao sp. nov. can be distinguished from other congeners by the distinct triangular form of the external genitalia (Fig. 35B) (vs. the absence of triangular external genitalia in congeners), and a pair of transverse linear spermathecae (Fig. 35A) (vs. the absence of linear spermathecae in congeners).
Male. Unknown. Distribution. Known only from the type locality (Fig. 58).  Etymology. The species name is an adjective referring to the type locality. Diagnosis. Diagnostic features of males and females are discussed in the diagnosis of L. mianqiu sp. nov.
Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "yuánzhuī" (conical) and refers to the shape of the external genitalia with a genitalic lobe that greatly resembles the shape of a conical flask (Fig. 38B, C).

Leclercera yuanzhui
Diagnosis. Females of L. yuanzhui sp. nov. resemble L. maochong sp. nov. and L. shanzi sp. nov. but can be distinguished by the presence of a protruded genitalic lobe ( Fig. 38C) (vs. the absence of a genitalic lobe), the presence of a pair of widely separated semi-circular bodies posteriorly (vs. the absence of semi-circular bodies or with circular bodies), and entire spermathecae rather thin and narrow (Fig. 38A) (vs. entire spermathecae rather thick and wide).
Male. Unknown. Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is an adjective referring to the type locality. Diagnosis. Females of L. paiensis sp. nov. can be distinguished from other congeners by the presence of a distinct horizontal, thick lip posterior to the external genitalia region (Fig. 39B) (vs. the absence of a thick lip on the external genitalia region in congeners), and twisted, stalked spermathecae with ovoid bases (Fig. 39A) (vs. an absence of twisted, stalked spermathecae in congeners).

Leclercera zanggaensis
Male. Unknown. Distribution. Known only from the type locality (Fig. 58).   Diagnosis. Diagnostic features of males and females are discussed in the diagnosis of L. jiazhongensis sp. nov.
Female. Unknown. Distribution. Known only from the type locality (Fig. 58). Etymology. The species name is a noun in apposition derived from the Chinese pinyin "túdāo" (cleaver) and refers to the structure of the laminar apophysis and embolus together resembling a pair of cleavers (Fig. 50A).
Diagnosis. Diagnostic features of the males are discussed in the diagnosis of L. renqinensis sp. nov.
Distribution. Known only from the type locality (Fig. 58).

Discussion
This study describes 31 new species, yielding a total of 42 species in the genus Leclercera. Leclercera species are reported for the first time from the Tibet Autonomous Region of China, Malaysia, Indonesia, and Myanmar. The large amount of new species discovered in Tibet (15 new species in this study) are noteworthy. The new Tibetan species occur in close proximity to one another, but this study surprisingly reveals large differences in genital morphology. This is congruent with previous studies of the abundant biodiversity and biological diversification due to the uplift of the Tibetan Plateau (Zhang et al. 2002, Zhao andLi 2017). This study provides a strong case for taxonomic studies in Southeast Asia, especially Tibet. Based on this work and additional observations, we predict that there are additional species in this genus awaiting discovery.