Sixteen new species of the genus Pseudopoda Jäger, 2000 from China, Myanmar, and Thailand (Sparassidae, Heteropodinae)

Abstract Sixteen new species of Pseudopoda Jäger, 2000 (Sparassidae, Heteropodinae) are described. Among them, eight species were collected from China: P.chayuensis Zhao & Li, sp. n. (♂), P.conaensis Zhao & Li, sp. n. (♂), P.medogensis Zhao & Li, sp. n. (♂), P.nyingchiensis Zhao & Li, sp. n. (♂), P.shacunensis Zhao & Li, sp. n. (♂), P.shuo Zhao & Li, sp. n. (♂♀), P.yuanjiangensis Zhao & Li, sp. n. (♀) and P.zixiensis Zhao & Li, sp. n. (♂); seven from Myanmar: P.colubrina Zhao & Li, sp. n. (♂♀), P.daxing Zhao & Li, sp. n. (♂), P.gexiao Zhao & Li, sp. n. (♂), P.putaoensis Zhao & Li, sp. n. (♂), P.subbirmanica Zhao & Li, sp. n. (♂♀), P.titan Zhao & Li, sp. n. (♂♀), P.xia Zhao & Li, sp. n. (♂); and one from Thailand: P.maeklongensis Zhao & Li, sp. n. (♂). A distribution map of the new species is also provided.


Introduction
Pseudopoda Jäger, 2000 is currently the third largest genus in the family Sparassidae Bertkau, 1872, containing 124 known species. A molecular phylogeny of Sparassidae asserted that Pseudopoda belongs to the subfamily Heteropodinae, and is closely related to Heteropoda Latreille, 1804 and Sinopoda Jäger, 1999(Moradmand et al. 2014. Along with the description of 49 new species from Himalayas and adjacent mountains, Jäger (2001) proposed six species-groups mainly according to the features of male pedipalp and female epigyne: P. diversipunctata-group, P. latembola-group, P. martensi-group, P. parvipunctata-group, P. prompta-group and P. schwendingeri-group. Based on both molecular and morphological characteristics, Zhang et al. (2017) proposed the seventh species group: P. daliensis-group and described three new species from Yunnan, China.
Currently, all of the Pseudopoda species are found in Asian countries: Bhutan, China, India, Indonesia, Japan, Laos, Myanmar, Nepal, Pakistan, Thailand, and Vietnam. To date, 54 species have been reported from China, while 14 from Myanmar and six species from Thailand (World Spider Catalog 2018). A considerable number of them are recorded from high altitude mountain regions, such as the Himalayas and Yunnan-Guizhou Plateau in China. Most of the Pseudopoda species exhibit very small-ranged distributions, but a high local diversity. A previous research explored on the application of DNA barcoding in taxonomic assessment in this genus, and indicated there is a greater species diversity remaining to be discovered (Cao et al. 2016). Here, we described 16 newly discovered species collected from southern China (Yunnan Province, Jiangxi Province and Tibet Autonomous Region), northern Myanmar (Kachin State), and Thailand (Tak Province).

Material and methods
All specimens were examined and measured with a Leica M205C stereomicroscope. Images of male pedipalps and female epigynes were taken with an Olympus C7070 wide zoom digital camera (7.1 megapixels) mounted on an Olympus BX51 compound light microscope after removing them from the spiders' bodies. Images of bodies were taken with an Olympus C7070 camera mounted on an Olympus SZX12 dissecting microscope. Epigynes were cleaned and treated in trypsin and if necessary, in boiling solution of potassium hydroxide (KOH) before taking images of the vulvae. All images were assembled using Helicon Focus 6.7.1 software.
All measurements are in millimeters. Leg formula, spination, and measurements of palp and legs follow Jäger and Vedel (2007). Arising points of tegular appendices (i.e. embolus, conductor) are given as 'clock positions' on the left palp in ventral view. When the left palp is lost or incomplete, the images of right palp will be taken and flipped horizontally for the sake of comparison. In this case, the right palp will be treated as the left one.
Abbreviations used in the text and figures are given below: AB anterior bands of epigynal field ALE anterior lateral eye AME anterior median eye BP basal part of embolic projection C conductor Diagnosis. Exclusively distributed in Asia. Small to large Heteropodinae. Male palp with membranous conductor (but sometimes absent), embolus at least in its basal part broadened and flattened, RTA arising basally or mesially from tibia; female epigyne with lateral lobes rising distantly beyond epigastric furrow, and in most cases covering median septum (modified from Jäger 2000). Etymology. The specific name refers to the type locality; adjective. Diagnosis. Medium-sized Pseudopoda species. Male resembles P. gongschana Jäger &Vedel, 2007 (see Jäger andVedel 2007: 6, figs 10-15) and P. platembola Jäger, 2001(see Jäger 2001: 57, figure 35a-e) by: 1. embolus sickle-shaped, tapering very moderately ( Figure 2A); 2. dRTA well developed and finger-like, curving distally ( Figure  1A-C). It can be distinguished from the two congeners by the following combination of characters: 1. embolic projection near the tip of embolus, making the tip look  somewhat incised ( Figure 1A; embolic projection absent in P. platembola); 2. embolus curving more intensely than in P. gongschana (Figure 2A).
Palp as in diagnosis. Cymbium distally slender and elongated, with a small retrobasal projection in ventral view. RTA arising basally to mesially from tibia, vRTA thumb-like, shorter than dRTA ( Figure 1A-C). Sperm duct running submarginally retrolaterally in tegulum. Embolus sickle-shaped, arising from tegulum at 10 o'clock position. The embolus tapering and very moderately curved. Embolic projection emerging at the prolateral margin of embolus as a blunt hump. Conductor arising from tegulum at 12 o'clock position, slightly leaning prolaterally and covering the tip of embolus (Figure 2A, B).
Coloration in ethanol: carapace yellow. Radial furrows and fovea dark brown. Dorsal opisthosoma brown with black pattern. Legs yellowish brown, with reddish brown dots and patches ( Figure 2C, D Etymology. The specific name is derived from the Latin word colubrinus, -a, -um, meaning 'serpentine, winding', and referring to the shape of embolus in this species, which coils at the basal part and erects distally and looks like an alarmed snake; adjective. Diagnosis. Small to median-sized Pseudopoda species. Male resembles P. wu Jäger, Li & Krehenwinkel, 2015(see Jäger et al. 2015 and P. tji Jäger, 2015(see Jäger 2015 by: 1. embolus robust but twisted, forming loops ( Figure 4A, B; rarely seem in other Pseudopoda spp.); 2. conductor absent ( Figure  4A, B). It can be easily distinguished from the two congeners by the following combination of characters: 1. only basal part of embolus twisted, distal part elongated and mildly bent ( Figure 4A, B; distal part coiled in P. tji and P. wu); 2. tegulum occupying two third of alveolus ( Figure 3B; covering whole or most of alveolus in P. tji and P. wu).
Female resembles P. hyatti Jäger, 2001(see Jäger 2001 by: 1. posterior part of lateral lobes surpassing the epigastric furrow; 2. loops of internal duct system mainly winding near the central axis, running transversally ( Figure 5A, B, E). It can be distinguished from the latter by the following combination of characters: 1. copulatory opening located at the middle to posterior part of epigyne ( Figure 5A; located near the anterior margin of lateral lobe in P. hyatti); 2. anterior margin of epigynal field truncated, anterior bands absent ( Figure 5A; anterior margin of epigynal field trilobate with short anterior bands in P. hyatti).
Epigyne as in diagnosis. Epigynal field with nearly equal length in transverse and longitudinal axis. Lateral lobes longer in longitudinal axis. Median margin of lateral lobes touching each other medially. Internal duct system with loops looming through the lateral lobes in ventral view ( Figure 5A). A pair of small appendages present ( Figure  5E).
Coloration in ethanol: As in male, but generally darker with more dots and patches ( Figure 5C, D).
Palp as in diagnosis. Cymbium relatively widened, with distinct retrolateral bulge beside bulb. RTA arising basally from tibia, well developed. Subtegulum extended, covering the base of conductor in prolateral view ( Figure 6A-C). Sperm duct running submarginally retrolaterally in tegulum. Embolus long, sickle-shaped, arising from tegulum at 9 o'clock position. Conductor arising from tegulum at 12 o'clock position, leaning prolaterally and covering the tip of embolus ( Figure 7A, B).
Coloration in ethanol: carapace yellowish brown, with a pair of dark longitudinal lateral bands. Radial furrows and fovea dark brown. Dorsal opisthosoma reddish brown. Legs yellowish brown, with darker brown dots and patches ( Figure 7C, D).
Female. Unknown. Distribution. Known only from the type locality. Etymology. The specific name is derived from the Chinese Pinyin word for 'large size' (dà xíng), referring to the relatively large body size of the species; noun in apposition.
Coloration in ethanol: carapace yellowish brown. Radial furrows and fovea dark brown. Dorsal opisthosoma reddish brown. Ventral opisthosoma with a pair of longitudinal bright lines. Legs yellowish brown, with randomly distributed brown dots ( Figure 9C, D).
Female. Unknown. Distribution. Known only from the type locality. Etymology. The specific name is derived from the Chinese Pinyin word for 'smallsize' (gè xiǎo), referring to the relatively small body size of the species; noun in apposition.
Palp as in diagnosis. Retrolateral margin of cymbium swollen. Distal part of cymbium sub-triangular. RTA arising mesially to distally from tibia, dRTA needlelike, while vRTA broad ( Figure 10A-C). Sperm duct running submarginally retrolaterally in tegulum. Embolus sickle-shaped, arising from tegulum at 9 o'clock position. Basal part of embolus broad, then tapering as it runs and coils, resulting in a filiform tip. Conductor arising from tegulum at 11 o'clock position, leaning prolaterally and then bent in a right angle, with its end covering the tip of embolus ( Figure 11A, B).
Coloration in ethanol: carapace yellow, with a pair of dark longitudinal lateral bands. Dorsal opisthosoma reddish brown with a bright transverse band in the posterior half. Legs yellowish brown, with reddish brown dots and patches ( Figure 11C, D).
Female. Unknown. Distribution. Known only from the type locality.    Jäger and Ono 2002: 112, figs 11-14). It can be distinguished from the two congeners by the following combination of characters: 1. tegulum small, leaning towards the retrolateral margin of cymbium ( Figure 12B); 2. embolic projection long, arising from the basal part of embolus at 9 o'clock position, forming a semicircle with its basal part running along with embolus and covering a part of it like a sheath ( Figure 13A, B; absent in P. spirembolus and P. parvipunctata); 3. embolus extremely long, forming five loops ( Figure 13A, B; forming three loops in P. spirembolus; two in P. parvipunctata); 4. cymbium flattened and broadened without any bulges (Figure 12A-C; elongated and with one bulge on the retrolateral margin in P. parvipunctata; broadened and with one bulge on the retrolateral margin in P. spirembolus).
Palp as in diagnosis. Cymbium large. RTA arising basally from tibia. Both vRTA and dRTA flattened and blunt in ventral view ( Figure 12A-C). Sperm duct S-shaped, running retrolaterally in tegulum. Embolus arising from tegulum at 9 o'clock position, extremely elongated. Conductor large and elongated, arising from the tegulum at 10 to 12 o'clock position ( Figure 13A, B).
Coloration in ethanol: carapace yellow. Radial furrows and fovea brown. Dorsal opisthosoma yellowish to reddish brown. Legs yellow, with randomly distributed brown dots ( Figure 13C, D).

Pseudopoda medogensis
Palp as in diagnosis. Cymbium slender. RTA almost straight, arising mesially from tibia ( Figure 14A-C). Sperm duct running submarginally retrolaterally in tegulum. Embolus arising from tegulum at 10 to 11 o'clock position with its basal part broadened. Distal part of embolus curved intensely, with its tip pointing at the base of embolus. Conductor arising from tegulum at 11 o'clock position (Figure 15A, B).
Coloration in ethanol: carapace bright brown. Radial furrows and fovea darker. Dorsal opisthosoma dark brown with black pattern. Legs bright brown, with dark brown patches ( Figure 15C, D).
Female. Unknown. Distribution. Known only from the type locality.    Figure 18B, C); 2. embolus sickle-shaped ( Figure 19A, B). It can be distinguished from the three congeners by the following combination of characters: 1. embolic projection pronounced, emerging from the prolateral margin of embolus ( Figure 19A, B; absent in P. platembola); 2. cymbium slender and elongated ( Figure 18B; shorter and wider in P. nyingchiensis Zhao & Li, sp. n. and P. platembola); 3. flange absent near the tip of embolus (present in P. huberi).
Palp as in diagnosis. RTA arising mesially from tibia ( Figure 20A-C). Sperm duct running submarginally retrolaterally in tegulum. Embolus arising from tegulum at 9-10 o'clock position with its basal part broadened and its distal part filiform. Embolic projection arising mesially from embolus, steeply narrowed at its distal half. Distal part of embolic projection filiform, curved, and running backwards to the tegulum. Conductor arising from tegulum at 12 o'clock position, leaning prolaterally and covering the tip of embolus ( Figure 21A, B).
Coloration in ethanol: carapace yellow. Radial furrows and fovea dark brown. Dorsal opisthosoma bright brown with reddish brown pattern composed of dense reddish brown dots. Legs yellow, with reddish brown dots and patches ( Figure 21C, D).
Female. Unknown. Distribution. Known only from the type locality. Etymology. The specific name is derived from the Chinese Pinyin word for 'gigantism' (shuò), referring to the relatively larger bulb on male palp than other Pseudopoda species; noun in apposition.
Palp as in diagnosis. Cymbium relatively shortened compared to other Pseudopoda species. RTA arising basally from tibia ( Figure 22A-C). Sperm duct running submarginally and retrolaterally in tegulum. Embolus arising from tegulum at 10-11 o'clock position. Angle between the tip of embolus and the broad part of embolus is ca. 180°. Conductor arising from tegulum at 12 o'clock position ( Figure 23A, B).
Coloration in ethanol: carapace bright brown with dark brown lateral bands. Radial furrows and fovea darker. Dorsal opisthosoma reddish brown with black pattern and a bright transverse band in the posterior half. Legs bright brown, with reddish brown patches ( Figure 23C, D).
Epigyne as in diagnosis. Epigynal field longer in transverse axis, with poorly developed anterior bands and trilobate anterior margin. Lateral lobes longer in transverse axis, curved. Median margin of lateral lobe converged, with the posterior part V-shaped. Posterior incision of lateral lobe indistinct or absent ( Figure 24A, B).
Coloration in ethanol: as in male, but generally darker. Ventral opisthosoma with a pair of bright, longitudinal, dashed lines ( Figure 24C, D).
Distribution. Known only from the type locality.  Etymology. The specific name refers to the similarity of its female individual to P. birmanica Jäger, 2001;adjective. Diagnosis. Small to median-sized Pseudopoda species. Male resembles P. digitata Vedel, 2007 (see Jäger andVedel 2007: 29, figs 105-113) by: embolus with prolateral projection near the tip ( Figure 26A, B). It can be distinguished from the latter    Etymology. The specific name is derived from the name of giants in Greek myth, referring to the gigantic size of this species; noun in apposition. Diagnosis. Large-sized Pseudopoda species. Male resembles P. emei Zhang, Zhang & Zhang, 2013(see Zhang et al. 2013b, P. namkhan Jäger, Pathoumthong & Vedel, 2006(see Jäger et al. 2006 and P. mediana (see Quan et al. 2014: 562, figs 6A-C, 7A-C, 8A-D, 9A-C) by: tip of embolus sharply curved and pointing prolaterally ( Figure 29A, B). It can be distinguished from the three congeners by the following combination of characters: 1. dRTA well developed, curved, and finger-like (Figure 28A-C; straight and significantly shorter in P. emei and P. mediana; broadened in P. namkhan); 2. tip of embolus slightly broadened ( Figure 29B; filiform in P. emei); 3. significantly larger in body size.
Palp as in diagnosis. Cymbium slender, with distinct retrolateral bulge beside bulb. RTA arising basally from tibia ( Figure 28A-C). Sperm duct running submarginally retrolaterally in tegulum. Embolus arising from tegulum at 10 o'clock position, broad, almost straight in ventral view. Tip of embolus leaf-like, sharply curved, and pointing prolaterally. Embolic projection present as two additional triangular rims near the tip. Conductor arising from the tegulum at 12 to 1 o'clock position ( Figure 29A, B).
Coloration in ethanol: carapace yellowish brown. Radial furrows and fovea dark brown. Dorsal opisthosoma reddish brown, with white dots and yellow patches. Legs orange. Ventral opisthosoma with two pairs of longitudinal lines composed of orange dots ( Figure 29C, D).
Epigyne as in diagnosis. Epigynal field longer in transverse axis, with distinct anterior bands and trilobate anterior margin. Lateral lobes longer in transverse axis, subrectangular but narrower laterally. Posterior incision of lateral lobe distinct, near the posterior meeting point of lateral lobes ( Figure 30A). Lateral loops of internal duct system running transversally, covered by first winding in dorsal view ( Figure 30B, E).
Coloration in ethanol: as in male but generally darker ( Figure 30C, D). Distribution. Known only from the type locality.  Jäger et al. 2006: 220, figs 1-13, 29-32) by: embolus running near the prolateral margin of tegulum in ventral view. It can be distinguished from the four congeners by the following combination of characters: 1. RTA simple, with only one apex ( Figure 31B, C; RTA with two apices in P. confusa); 2. tegulum with a distinct sub-triangular protrusion near the retrolateral margin ( Figure  32A; absent in P. prompta and P. confusa; a blunt hump present on tegulum near the basal part of embolus in P. trisuliensis and P. brauni); 3. embolus with an extra rim running along the distal part of it ( Figure 32B; absent or indistinct in P. prompta and P. confusa).
Palp as in diagnosis. Cymbium slender, slightly elongated distally. RTA arising basally from tibia ( Figure 31A-C). Tegulum with an additional ridge emerging basally, and running distally, ending with a sub-triangular protrusion pointing at the basal part of embolus. Sperm duct running submarginally retrolaterally in tegulum, visible near the base of embolus as an S-shaped duct. Embolus arising from tegulum at 9 to 10 o'clock position. Wrinkles present below the distal part on embolus. Tip of embolus with indention. Conductor arising from tegulum at 1 to 2 o'clock position, slender, bent basally and then directed prolaterally ( Figure 32A, B).   Jäger and Vedel 2007: 15, figs 44-59) by: internal duct system with distinct lateral loops visible through cuticle in ventral view as rounded patches ( Figure 33A). It can be distinguished from the latter species by the following combination of characters: 1. anterior bands distinct ( Figure 33A; absent in P. bibulba); 2. lateral lobes much longer in transverse axis, with anterior margins bending posteriolaterally ( Figure 33A; anterior margins bending anteriolaterally and then directed medially in P. bibulba).
Epigyne as in diagnosis. Epigynal field longer in transverse axis, with anterior bands and trilobate anterior margin. Lateral lobes slightly converged on the central axis. Posterior incision of lateral lobe distinct, near the meeting point of lateral lobes. (Figure 33A, B).
Coloration in ethanol: carapace yellowish brown. Radial furrows and fovea dark brown. Dorsal opisthosoma reddish brown, with a bright transverse band in the posterior half. Legs yellowish brown, with randomly distributed reddish brown dots ( Figure 33C, D).
Male. Unknown. Distribution. Known only from the type locality.   Epigyne as in diagnosis. Epigynal field with nearly equal length in transverse and longitudinal axis. Anterior bands distinct, anterior margin slightly trilobate. Lateral lobes longer in longitudinal axis. Lateral lobes converged on the central axis, with both anterior and posterior part V-shaped. Spermathecae exposed in dorsal view. Spermathecae oval, with coiling ducts embedded ( Figure 36B, E).
Coloration in ethanol: as in male, but generally darker ( Figure 36C, D). Distribution. Known only from the type locality.