Sarotes promptus O. Pickard-Cambridge, 1885, from Murree (Pakistan) and Himachal Pradesh (India).

See Zhang et al. (2023a) and Wu et al. (2024).

Its distribution range extends west to Murree in Pakistan (approximately 73°E), east to the Ryukyu Islands in Japan (approximately 128°E), south to Kaeng Krachan National Park in Phetchaburi Province, Thailand (approximately 13°N), and north to Taibaishan National Forest Park in Shaanxi Province, China (approximately 34°N). Also can refer to MAP1 in Zhang et al. (2023a).

Up to now, a total of nine Pseudopoda species groups have been proposed, and 67 species were clearly assigned. Among them, the daliensis-group (five species), signata-group (seven species), and interposita-group (two species) were established based on both molecular and morphological characteristics by Zhang et al. (2017, 2019) and Li et al. (2019), respectively. The remaining six were all established by Jäger (2001) based on genitalic morphological characters to accommodate 42 species: the diversipunctata-group (three species), latembola-group (seven species), martensi-group (13 species), parvipunctata-group (eight species), prompta-group (nine species), and schwendingeri-group (two species). After the establishment of these species groups, at least 11 species were clearly assigned to these groups in the original papers: four were allocated to the diversipunctata group by Jäger et al. (2006) and Yang et al. (2022), four were allocated to the martensi group by Jäger (2008), Zhang et al. (2013b), and Caleb et al. (2018), two were allocated to the prompta group by Jäger (2008) and Zhang et al. (2023a), and one was allocated to the schwendingeri group by Jäger et al. (2015).

It is evident that the number of species clearly assigned to these species groups accounts for about a quarter of the total species in the genus. In contrast, the remaining nearly 200 species, including a large number of recently described species, cannot be allocated (Jiang et al. 2018; Deng et al. 2023; Gong et al. 2023; Zhang et al. 2023a; Chang et al. 2024; Wen et al. 2024; Wu et al. 2024). Sorting Pseudopoda species into species groups is highly challenging. The possible reasons for this have been discussed in detail in Zhang et al. (2023a) and will not be repeated here.

We have also attempted to group the six species treated in this paper based on both morphological and molecular data but were unable to do so. While P. ying sp. nov. can be clearly assigned to the daliensis group (as it exhibits typical features of the daliensis group and resembles P. sicyoidea Zhang, Jäger & Liu, 2017, the core species of this group, and the grouping is also supported by molecular data), the remaining five species cannot be assigned to any of the existing nine species groups. Furthermore, reviewing the species groups of the genus is not within the scope of this work. Therefore, we refrain from assigning species (except P. ying sp. nov.) to species groups in the current paper.

♀ (CBEE, LJ202002838), China: • Yunnan Prov.: Honghe Hani and Yi Autonomous Prefecture, Luechun Co., Huanglianshan Mt., 22.99°N, 102.46°E, c. 1940 m, by hand, 13 VII 2020, R. Zhong et al. leg. Examined.

• 2 ♂ 2 ♀ (YNZY013, YNZY014, YNZY024, YNZY025). Same locality as holotype, by hand, 16 IV 2024, Y. Zhong & S. Yang leg.

Males of this species can be easily distinguished from those of all other congeners by the embolus (E) shaped like the lowercase letter ‘y’ in ventral view, the embolic tip (ET) strongly torqued along its length, with distinct subterminal torsion, and by the rhombic tip of conductor (C) (Figs 4A, 5A, B, 6A–C). In contrast, the embolus (E) and conductor (C) of all other species do not exhibit these characteristics. In most Pseudopoda species, such as P. mamillaris and P. yangae sp. nov., the embolic tip (ET) is not torqued (as shown in Figs 8A, 9A, B, 10A–C, 20A, 21A, B, 22A–C), or the embolic tip (ET) is moderately torqued with no distinct subterminal torsion, as in a few species like P. oliviformis and P. xiaozhua sp. nov. (as shown in Figs 12A, 13A, B, 14A–C, 17A, B, 18C). Female of P. huanglianensis resembles that of P. anfracta Zhang, Jäger & Liu, 2023, in having the similarly shaped median field (MF) and the relatively sclerotized, nearly funnel-shaped first windings (FD), but can be recognised by the median field (MF) relatively narrower, ca. 2/5 epigyne width (vs. wider, more than 2/3 epigyne width) (cf. Fig. 7A, B and Zhang et al. 2023a: fig. 12A). Female also resembles that of P. cangschana Jäger & Vedel, 2007 (Zhang et al. 2023a: 50, figs 39A, B, 40C, D); see Zhang et al. (2023a) for the diagnosis.

Figure 4. 

Male palp of the topotype of Pseudopoda huanglianensis. A. Ventral; B. Dorsal. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 5. 

Male palp of the topotype of Pseudopoda huanglianensis. A. Prolateral; B. Retrolateral. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 6. 

Pseudopoda huanglianensis, male topotype, palpal bulb (A–C) and habitus (D, E). A. Prolateral; B. Ventral; C. Retrolateral; D. Dorsal; E. Ventral. Abbreviations: C = conductor; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum. Scale bars: 0.2 mm (equal for A–C); 1 mm (equal for D, E).

Figure 7. 

Pseudopoda huanglianensis, female topotype, epigyne (A–C) and habitus (D, E). A. Intact, ventral; B. Cleared and macerated, ventral; C. Cleared and macerated, dorsal; D. Dorsal; E. Ventral. Abbreviations: AB = anterior band; aEF = anterior margin of epigynal field; amLL = anterior margin of lateral lobes; CO = copulatory opening; FD = fertilization duct; FW = first winding; LL = lateral lobe; MF = median field of epigyne; mmLL = median margin of lateral lobes; MS = membranous sac; PI = posterior incision; pmLL = posterior margin of lateral lobes; S = spermatheca; SA = spermathecal appendage. Scale bars: 0.2 mm (equal for A–C); 2 mm (equal for D, E).

Male (YNZY013). Total length 7.1. Carapace 3.6 long, 3.5 wide, anterior width 1.7. Opisthosoma 3.5 long, 2.2 wide. Eye sizes and interdistances: AME 0.18, ALE 0.30, PME 0.24, PLE 0.30, AME–AME 0.10, AME–ALE 0.06, PME–PME 0.22, PME–PLE 0.24, AME–PME 0.25, ALE–PLE 0.25, CH AME 0.32, CH ALE 0.25. Spination: palp: 131, 101, 2111; Fe: I–II 323, III 322, IV 321; Pa: I–IV 101; Ti: I–II 2228, III 2226, IV 2126; Mt: I–II 2024, III–IV 3036. Measurements of palp and legs: palp 4.9 (1.6, 0.7, 0.8, 1.8), I 15.3 (4.3, 1.9, 4.1, 3.6, 1.4), II 16.5 (4.5, 1.9, 4.3, 4.2, 1.6), III 12.8 (3.9, 1.3, 3.3, 3.2, 1.1), IV 14.9 (4.4, 1.4, 3.6, 4.1, 1.4). Cheliceral furrow with ~34 denticles.

(Fig. 6D, E). DS light brown, lateral bands and margin slightly darker, clothed with fine setae; median band bright yellowish-brown, not distinctly delimited to lateral bands, with indistinct Ψ-shaped markings starting from behind PER, almost reaching indistinct cervical groove; fovea and striae distinctly marked. Cheliceral base coloured similarly to median band, with red fangs. Sternum uniformly light brown. Endites light brown. Labium coloured similarly to endites. Legs yellowish-brown, with numerous brown spots, and covered with short spines. OS elongate-oval; dorsum with median band starting from behind pedicel, reaching 4/5 of abdomen length, with two pairs of circular dots on each side; median band with diamond-shaped anterior part, cross-shaped stem, and ‘˽’-shaped, black posterior part, all three parts fused; dorsum with transverse yellow line located posterior to median band; ventral OS marked with numerous brown spots.

Palp (Figs 4, 5, 6A–C, 28A). Femur and patella unmodified. Tibia (Ti) relatively long, ca. 1/2 cymbium length, with retrolateral apophysis (RTA) arising mesially to distally; RTA bifurcated, with ventral part (vRTA) and dorsal branch (dRTA): dRTA finger-like, slightly curved and tapering, ca. 3/4 tibia length, extending to cymbial base; vRTA laminar, humble, and broad in retrolateral view, with blunt, round apex. Cymbium (Cy) approximately 2.4 times as long as wide, retrolaterally with indistinct bulge (CB). Tegulum (T) nearly egg-shaped, ca. 1.3× longer than wide, relatively flat, marginally with distinct, ɔ-shaped spermophor (Sp) in ventral view. Embolus (E) shaped like the lowercase letter ‘y’ in ventral view, ca. as long as tegulum (T), arising from tegulum at nearly the 8–9 o’clock position, terminating at ca. 11:30 o’clock position; the proximal first half of embolus (E) columnar, while the second half thin and flat, ribbon-shaped; embolic tip (ET) strongly torqued along its length, with subterminal torsion and distal, rostrate bend. Conductor (C) membranous, ca. 1/2 of embolus length, inserted anterodorsally on tegulum, extending obliquely, the latter half of conductor (C) widens and diamond-shaped, directing prolaterally.

Female (YNZY025). Total length 9.1. Carapace 4.1 long, 3.8 wide, anterior width 2.3. Opisthosoma 5.0 long, 3.6 wide. Eye sizes and interdistances: AME 0.22, ALE 0.34, PME 0.25, PLE 0.34, AME–AME 0.14, AME–ALE 0.08, PME–PME 0.32, PME–PLE 0.34, AME–PME 0.32, ALE–PLE 0.32, CH AME 0.35, CH ALE 0.26. Spination: palp: 131, 101, 2121, 1014; Fe: I–II 323, III 322, IV 321; Pa: I–IV 101; Ti: I–II 2228, III–IV 2126; Mt: I–II 2024, III–IV 3036. Measurements of palp and legs: palp 5.4 (1.7, 0.9, 1.1, 1.7), I 14.4 (4.3, 1.8, 3.3, 3.7, 1.3), II 15.1 (4.4, 2.0, 3.8, 3.6, 1.3), III 12.0 (3.7, 1.5, 2.9, 2.8, 1.1), IV 14.0 (3.9, 1.8, 3.6, 3.4, 1.3). Cheliceral furrow with ~50 denticles. Colouration in ethanol as in males, but body slightly darker (Fig. 7D, E; see Zhang et al. (2023a) for others described).

Epigyne (Fig. 7A–C). Epigynal field ca. 1.25× wider than long; anterior margin (aEF) distinctly delimited, mesially distinctly recurved; anterior bands (AB) indistinct. Median field (MF) shaped like a ginkgo leaf (more or less fan-shaped), ca. 2/5 epigyne length and 2/5 epigyne width. Lateral lobes (LL) nearly as wide as long, slightly converged along the axis; anterior margins (amLL) slightly recurved; median margins (mmLL) touching each other along middle line in anterior half; posterior margins (pmLL) with distinct incision (PI). Copulatory openings (CO) indistinct, located at basolateral borders of median field (MF). First windings (FW) weakly sclerotized, nearly funnel-shaped; starting from near copulatory openings (CO), descending obliquely, posteriorly with U-turns; two first windings (FW) separated by ca. 0.75× diameters. Spermathecae (S) represented by thick, ‘∩’-shaped tubes, forming letter ‘m’; laterally covered by first windings (FW) and with globular appendage (SA), medially exposed and touching each other along middle line in posterior half. Membranous sac (MS) nearly trapeziform, located at posterior portion of vulva; anterior margin recurved, separated from epigastric fold by ca. 1/4 epigyne length, reaching contact point of spermathecae; posterior margin nearly straight, close to epigastric fold. Fertilization ducts (FD) acicular, membranous, ca. 1/2 length of first windings.

Known only from the type locality (Fig. 1).

♀ (CBEE, LJ202002615), China: • Yunnan Pro.: Honghe Hani and Yi Autonomous Prefecture, Luechun Co., Martyr Cemetery, 22.99°N, 102.45°E, c. 1940 m, by hand, 11 VII 2020, R. Zhong et al. leg. Examined.

• 2 ♂, 2 ♀ (YNZY009, YNZY010, YNZY022, YNZY023), same locality as holotype, by hand, 15 IV 2024, Y. Zhong & S. Yang leg.

Male of P. mamillaris is similar to that of P. platembola Jäger, 2001, in having a similar sickle-shaped embolus (E) and petal-shaped conductor (C). However, it can be distinguished from the latter by the following characteristics: (1) in ventral view, embolus (E) is apically rugged and as wide as its middle section (vs. apically not rugged, pointed) (cf. Figs 8A, 10B, and Jäger 2001: figs 35b, g, i); (2) dorsal branch of RTA (dRTA) tip not curved (vs. slightly or distinctly curved) (cf. Figs 8A, B, 9B, 28B, and Jäger 2001: figs 35a–c, f–h); (3) RTA inserted closer to cymbium (Cy), both dRTA and vRTA separated from cymbium (Cy) by approximately half of the tibial (Ti) diameter (vs. inserted farther, both dRTA and vRTA separated from cymbium (Cy) by more than one tibial (Ti) diameter) (cf. Figs 8A, B and Jäger 2001: figs 35b, g). For the female diagnosis, see Zhang et al. (2023a).

Figure 8. 

Male palp of the topotype of Pseudopoda mamillaris. A. Ventral; B. Dorsal. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral branch of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 9. 

Male palp of the topotype of Pseudopoda mamillaris. A. Prolateral; B. Retrolateral. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral branch of RTA. Scale bar: 0.5 mm (equal for A, B).

Male (YNZY009). Total length 7.3. Carapace 3.6 long, 3.5 wide, anterior width 1.8. Opisthosoma 3.8 long, 2.6 wide. Eye sizes and interdistances: AME 0.19, ALE 0.30, PME 0.23, PLE 0.31, AME–AME 0.14, AME–ALE 0.06, PME–PME 0.23, PME–PLE 0.26, AME–PME 0.28, ALE–PLE 0.23, CH AME 0.32, CH ALE 0.27. Spination: palp: 131, 101, 2111; Fe: I–III 323, IV 321; Pa: I–III 101, IV 100; Ti: I–II 2226, III–IV 2126; Mt: I–II 2024, III–IV 3036. Measurements of palp and legs: palp 5.2 (1.6, 0.9, 1.0, 1.7), I 15.3 (4.4, 1.5, 3.8, 4.2, 1.4), II 15.9 (4.5, 1.9, 4.2, 3.8, 1.5), III 13.1 (3.8, 1.4, 3.4, 3.3, 1.2), IV 14.7 (4.0, 1.7, 4.0, 3.6, 1.4). Cheliceral furrow with ~26 denticles.

Colouration in ethanol (Fig. 10D, E). DS yellowish brown, with numerous dark spots; lateral bands and margin slightly darker, clothed with thin hairs; median band bright yellowish-brown, without distinct pattern, not distinctly delimited to lateral bands; fovea and radial grooves distinctly marked. Cheliceral base yellowish white, with red fangs. Sternum uniformly yellowish. Endites and labium reddish orange, darker distally. Legs coloured as DS, with numerous brown spots, and covered by short spines. OS elongate-oval; dorsum with indistinct median band starting from behind pedicel, extending to 4/5 of abdomen length, almost reaching the indistinct transverse yellow line; median band anteriorly represented by a longitudinal brown line, with two light brown and reniform patches on each side, the latter half distinctly widened and not clearly delimited, with three pairs of circular spots located at lateral part; ventral OS basically yellowish, centrally marked with three pairs of purplish dots.

Figure 10. 

Pseudopoda mamillaris, male topotype, palpal bulb (A–C) and habitus (D, E). A. Prolateral; B. Ventral; C. Retrolateral; D. Dorsal; E. Ventral. Abbreviations: C = conductor; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum. Scale bars: 0.2 mm (equal for A–C); 1 mm (equal for D, E).

Palp (Figs 8, 9, 10A–C, 28B). Femur and patella unmodified. Tibia (Ti) moderately long, ca. 3/5 cymbium length, with retrolateral apophysis (RTA) arising mesially; RTA bifurcated, ɔ-shaped in retrolateral view, both ventral and dorsal branches distinctly protruding: dorsal branch (dRTA) finger-like, slightly curved and tapering, ca. 3/5 of tibia length, extending to cymbial base; ventral branch (vRTA) thumb-like, relatively short, ca. 1/2 length of dRTA, apex round. Cymbium (Cy) ca. 1.9× longer than wide, cymbial bulge (CB) indistinct. Tegulum (T) oval, ca. 1.2× longer than wide, posteriorly slightly bulged, slightly excavated on prolatero-apical side to accommodate embolus (E) and conductor (C); spermophor (Sp) sinuate, forming a loop along tegular margin. Embolus (E) wide and flattened, nearly Ƨ-shaped in ventral view, ca. 3/4 of tegulum length and 1/6–1/5 of tegulum width; the embolic base (EB) situated prolateral on the tegulum (ca. 9–10 o’clock on tegulum); the free part of the embolus (E) sickle-shaped; the embolic tip (ET) apically rugged, terminated at ca. 12 o’clock position. Conductor (C) membranous, ca. 1/3 of embolus length, thick, more or less petal-shaped, inserted apically (approximately 12 o’ lock relative to tegulum), covering embolic tip (ET) in prolateral and ventral views.

Female (YNZY010). Total length 9.2. Carapace 3.8 long, 3.4 wide, anterior width 2.0. Opisthosoma 5.4 long, 4.0 wide. Eye sizes and interdistances: AME 0.20, ALE 0.31, PME 0.25, PLE 0.33, AME–AME 0.16, AME–ALE 0.06, PME–PME 0.21, PME–PLE 0.29, AME–PME 0.31, ALE–PLE 0.31, CH AME 0.32, CH ALE 0.23. Spination: palp: 131, 101, 2121, 1014; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–III 2026, IV 2126; Mt: I–II 2024, III 2026, IV 3036. Measurements of palp and legs: palp: 4.3 (1.3, 0.7, 0.9, 1.4), I 11.0 (3.4, 1.2, 2.6, 2.8, 1.0), II 12.1 (3.7, 1.6, 3.1, 2.6, 1.1), III 9.6 (2.9, 1.1, 2.4, 2.3, 0.9), IV 10.6 (3.1, 1.3, 2.7, 2.5, 1.0). Cheliceral furrow with ~26 denticles. Colouration in ethanol as in males, but body slightly darker (Fig. 11D, E; see Zhang et al. (2023a) for others described).

Figure 11. 

Pseudopoda mamillaris, female topotype, epigyne (A–C) and habitus (D, E). A. Intact, ventral; B. Cleared and macerated, ventral; C. Cleared and macerated, dorsal; D. Dorsal; E. Ventral. Abbreviations: AB = anterior band; aEF = anterior margin of epigynal field; amLL = anterior margin of lateral lobes; CO = copulatory opening; FD = fertilization duct; FW = first winding; LL = lateral lobe; MF = median field of epigyne; mmLL = median margin of lateral lobes; MS = membranous sac; pmLL = posterior margin of lateral lobes; S = spermatheca; SA = spermathecal appendage. Scale bars: 0.2 mm (equal for A–C); 2 mm (equal for D, E).

Epigyne (Fig. 11A–C). Epigynal field ca. 1.27× wider than long; anterior margin (aEF) distinctly delimited, trilobate, anterolaterally with two large v-shaped incisions that are well separated by at ca. 2× widths; anterior bands (AB) indistinct, situated at the two incisions. Median field (MF) anterior margin invisible, large, ca. 1/2 epigyne length and as wide as epigyne. Lateral lobes (LL) nearly as wide as long; anterior margins (amLL) distinctly recurved; median margins (mmLL) entirely touching each other along the middle line; posterior margins (pmLL) distinctly procurved; posterior incision (PI) absent. Copulatory openings (CO) indistinct, located at basolateral borders of median field (MF). First windings (FW) hyaline, shaped like inverted triangles, starting from copulatory openings (CO), descending longitudinally, tapering posteriorly; two first windings (FW) separated by ca. one diameter. Spermatheca (S) nearly ‘∞’-shaped, consisting of inner part and lateral part; inner part spherical, touching median margin of lateral lobe (mmLL), entirely covered by membranous sac (MS) in dorsal view; lateral part shaped like water droplet, anterolaterally with globular appendage (SA), entirely covered by first winding (FW) in dorsal view. Membranous sac (MS) nearly trapeziform, located at posterior portion of vulva; anterior margin almost straight, ca. 1/3 epigyne width, separated from epigastric fold by ca. 2/5 epigyne length; posterior margin also nearly straight, ca. 2/3 epigyne width, reaching posterior margins of lateral lobes (pmLL). Fertilization ducts (FD) acicular, membranous, ca. 1/2 length of first windings.

Known only from the type locality (Fig. 1).

♀ (CBEE, LJ2140), China: • Yunnan Pro.: Honghe Hani and Yi Autonomous Prefecture, Luechun Co., Martyr Cemetery, 22.99°N, 102.45°E, c. 1934 m, by hand, 30 X 2015, Y. Zhong & Y. Zhu leg. Examined.

• 2 ♂, 2 ♀ (YNZY011, YNZY012, YNZY015, YNZY016), same locality as holotype, by hand, 15 IV 2024, Y. Zhong & S. Yang leg.

Both sexes of P. oliviformis resemble those of P. zixiensis Zhao & Li, 2018 in the general shape of the male palp, the epigynal plate, and vulva. The palps of the two species share the similarly shaped embolus (E), which has a torsional tip, and the finger-like dorsal branch of RTA (dRTA), but differ in the following: (1) ventral branch of RTA (vRTA) subtriangular, apex sharp in retrolateral view (vs. humble and broad, with a blunt apex) (cf. Fig. 13B and Jiang et al. 2018: fig. 34C); (2) retrolateral rim of embolus (E) distinctly curved in ventral view (vs. almost straight) (cf. Figs 12A, 13A, B, 14A–C and Jiang et al. 2018: figs 35A, B); (3) embolic projection absent (vs. present) (cf. Figs 12A, 13A, 14B and Jiang et al. 2018: figs 34B, 35A). Female resembles P. zixiensis in having the similarly shaped lateral lobes (LL) and the spherical spermathecae (S), but can be recognised by: (1) anterior band (AB) and anterior margin of epigynal field (aEF) indistinct (vs. both distinct) (cf. Fig. 15A, B and Jiang et al. 2018: fig. 36A); (2) posterior margins of lateral lobes (pmLL) with distinct posterior incision (PI) on each side, respectively (vs. PI absent) (cf. Fig. 15A, B and Jiang et al. 2018: fig. 36A); (3) spermathecae (S) surface smooth, without coiling ducts embedded (vs. surface wrinkled, with coiling ducts embedded) (cf. Fig. 15C and Jiang et al. 2018: fig. 36B).

Figure 12. 

Male palp of the topotype of Pseudopoda oliviformis A. Ventral; B. Dorsal. Abbreviations: C = conductor; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 13. 

Male palp of the topotype of Pseudopoda oliviformis A. Prolateral; B. Prolateral. Abbreviations: C = conductor; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 14. 

Pseudopoda oliviformis, male topotype, palpal bulb (A–C) and habitus (D, E). A. Prolateral; B. Ventral; C. Retrolateral; D. Dorsal; E. Ventral. Abbreviations: C = conductor; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum. Scale bars: 0.2 mm (equal for A–C); 1 mm (equal for D, E).

Figure 15. 

Pseudopoda oliviformis, female topotype, epigyne (A–C) and habitus (D, E). A. Intact, ventral; B. Cleared and macerated, ventral; C. Cleared and macerated, dorsal; D. Dorsal; E. Ventral. Abbreviations: AB = anterior band; aEF = anterior margin of epigynal field; amLL = anterior margin of lateral lobes; CO = copulatory opening; FD = fertilization duct; FW = first winding; LL = lateral lobe; MF = median field of epigyne; mmLL = median margin of lateral lobes; MS = membranous sac; PI = posterior incision; pmLL = posterior margin of lateral lobes; S = spermatheca. Scale bars: 0.2 mm (equal for A–C); 2 mm (equal for D, E).

Male (YNZY011). Total length 8.4. Carapace 4.3 long, 4.4 wide; anterior width 2.3. Opisthosoma 4.1 long, 2.6 wide. Eye sizes and interdistances: AME 0.21, ALE 0.34, PME 0.27, PLE 0.32, AME–AME 0.13, AME–ALE 0.05, PME–PME 0.26, PME–PLE 0.32, AME–PME 0.34, ALE–PLE 0.30, CH AME 0.42, CH ALE 0.33. Spination: palp: 131, 101, 2101; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–II 2026, III–IV 2126; Mt: I–II 1014, III 2026, IV 3036. Measurements of palp and legs: palp 5.9 (2.0, 1.0, 1.1, 1.8), I 19.1 (5.0, 2.0, 5.2, 5.1, 1.8), II 20.2 (5.5, 2.0, 5.5, 5.2, 2.0), III 16.1 (4.6, 1.7, 4.1, 4.2, 1.5), IV 18.4 (5.3, 1.6, 4.5, 5.2, 1.8). Cheliceral furrow with ~42 denticles.

Colouration in ethanol (Fig. 14D, E). DS yellowish white, with numerous indistinct, irregularly shaped patches; lateral bands and margin slightly darker, relatively smooth, sparsely covered with hairs; median band bright yellowish, with distinct Ψ-shaped markings starting from behind PME, almost reaching reddish fovea; fovea and radial furrows distinctly marked. Cheliceral base coloured as DS, cheliceral fang red. Sternum mainly yellowish, with two pairs of indistinct dots located at lateral parts. Endites and labium yellowish, slightly darker on inner margin. Legs coloured as DS, with numerous black spots, and bearing short spines. OS oval; dorsum anteriorly with nearly ‘⨅‘-shaped bright region, centrally with indistinct ‘⚲’-shaped median band, posteriorly with large ‘)(‘-shaped black pattern located on both sides of median band, distally marked with more or less semicircular transverse yellow band; venter of OS centrally with inverted trapezoidal black patch.

Palp (Figs 12, 13, 14A–C, 28C). Femur and patella unmodified. Tibia (Ti) moderately long, ca. 2/3 cymbium length, with retrolateral apophysis (RTA) arising mesially; RTA bifurcated, with ventral part (vRTA) and dorsal branch (dRTA): dRTA ɔ-shaped in retrolateral view, curved and tapering, ca. 2/5 of tibia length, extending to cymbial base; vRTA represented by a distinctly short triangular lamina, ca. 1/3 length of dRTA, apex sharp. Cymbium (Cy) distinctly slender, ca. 2.7× longer than wide, cymbial bulge (CB) indistinct. Tegulum (T) oval, ca. 1.35× longer than wide, relatively flattened, proximally slightly bulged and prolapsed, slightly excavated on prolatero-apical side to accommodate embolus (E) and conductor (C); spermophor (Sp) sinuate, not distinct, oriented clockwise along the margin of the tegulum (T). Embolus (E) nearly Ƨ-shaped in ventral view, ca. 2/3 of tegulum length; embolic base (EB) situated meso-prolaterally on the tegulum (T) (approximately the 9 o’clock position); mesially broadened and flattened, nearly 1/2 of tegulum width; embolic tip (ET) distinctly narrowed, with subterminal torsion and distal beak-shaped bend, apex sharp, terminated at ~ 11 o’clock position. Conductor (C) membranous, irregularly shaped, inserted apically (at approximately 11–12 o’clock position relative to the tegulum).

Female (YNZY012). Total length 10.5. Carapace 4.3 long, 4.0 wide, anterior width 2.4. Opisthosoma 6.2 long, 4.8 wide. Eye sizes and interdistances: AME 0.21, ALE 0.34, PME 0.27, PLE 0.33, AME–AME 0.18, AME–ALE 0.07, PME–PME 0.28, PME–PLE 0.35, AME–PME 0.32, ALE–PLE 0.31, CH AME 0.40, CH ALE 0.35. Spination: palp: 131, 101, 2121, 1014; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–IV 2126; Mt: I–II 2024, III–IV 3036. Measurements of palp and legs: palp 5.6 (1.7, 0.9, 1.2, 1.8), I 15.2 (4.3, 1.9, 3.9, 3.7, 1.4), II 16.7 (4.8, 2.1, 4.3, 4.0, 1.5), III 13.6 (4.3, 1.8, 3.1, 3.2, 1.2), IV 14.7 (4.6, 1.4, 3.5, 4.0, 1.2). Cheliceral furrow with ~46 denticles. Colouration in ethanol as in males, but body slightly darker (Fig. 15D, E; see Zhang et al. (2023a) for others described).

Epigyne (Fig. 15A–C). Epigynal field ca. 1.17× wider than long; anterior margin (aEF) indistinct, trilobate, anterolaterally with two large U-shaped incisions that are well separated by ca. one width; anterior bands (AB) indistinct. Median field (MF) nearly fan-shaped, large, ca. 1/2 epigyne length and 2/3 epigyne width; anterior margin invisible, posteriorly with a circular patch anterior to lateral lobes (LL) with special surface structure of cuticle. Lateral lobes (LL) distinctly longer than wide; anterior margins (amLL) slightly recurved, almost straight, forming the letter ‘V’; median margins (mmLL) entirely touching each other along the middle line; posterior margins (pmLL) ‘∧’-shaped, with distinct posterior incision (PI) on each side, respectively. Copulatory openings (CO) indistinct, located at basolateral borders of median field (MF). First windings (FW) membranous, tubular, slightly curved; starting from copulatory openings (CO), descending obliquely, then connecting with spermathecae at mid length of epigyne. Spermathecae (S) spherical, not subdivided, widely separated by ca. 1.5 diameters. Membranous sac (MS) large, nearly disc-shaped, located at posterior portion of vulva; anterior margin distinctly recurved, ca. 3/4 epigyne width, separated from epigastric fold by ca. 1/2 epigyne length, reaching cross point of anterior margins of lateral lobes (amLL); posterior margin distinctly procurved, as long as anterior margin, separated from epigastric fold by ca. 1/6 epigyne length. Fertilization ducts (FD) hyaline, slightly curved, relatively long, nearly 1/2 epigyne width; arising at central axis of vulva, extending laterally, terminating at lateral margin of epigynal field.

Known only from the type locality (Fig. 1).

♂ (YNZY003), China: • Yunnan Pro.: Honghe Hani and Yi Autonomous Prefecture, Luechun Co., Huanglianshan Mt., 22.99°N, 102.46°E, c. 1940 m, by hand, 16 IV 2024, Y. Zhong & S. Yang leg. Paratypes: • 1 ♂ 2 ♀ (YNZY004, YNZY018, YNZY019), same data as holotype.

The specific name is derived from the Chinese pinyin xiǎo zhuǎ, which means ‘small claw’, referring to the embolic tip, which is shaped like an unguiculus; noun in apposition.

The males of new species can be easily distinguished from those of all other congeners by a combination of the following characters: (1) embolus (E) almost as long as tegulum (T), nearly column-shaped, distally with a small claw-shaped tip (ET), as in Figs 16A, 17A, B, 18A–C (vs. embolus (E) not as above); (2) tegulum (T) with a heavily sclerotised, strongly expanded tegular apophysis (TA), as in Figs 16A, 17A, B, 18A–C (vs. tegular apophysis (TA) absent). Female of P. xiaozhua sp. nov. is very similar to that of P. mingshengi Yang & Zhang, 2022 in the general appearance of the median field (MF) and vulva, but can be recognised by: (1) anterior margin of epigyne (aEF) with two small v-shaped incisions that are well separated by at least 3× widths (vs. two wide incisions closely spaced, aEF nearly W-shaped) (cf. Fig. 19A, B and Yang et al. 2022: figs 2D, 3D); (2) membranous sac (MS) disc-shaped, located at posterior portion of vulva, anterior margin not beyond the contact point of lateral lobes (LL) (vs. membranous sac (MS) triangular, located at central portion, posterior margin beyond the contact point) (cf. Fig. 19C and Yang et al. 2022: figs 2E, 3E).

Figure 16. 

Male palp of the holotype of Pseudopoda xiaozhua sp. nov. A. Ventral; B. Dorsal. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; TA = tegular apophysis; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 1 mm (equal for A, B).

Figure 17. 

Male palp of the holotype of Pseudopoda xiaozhua sp. nov. A. Prolateral; B. Retrolateral. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; TA = tegular apophysis; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 1 mm (equal for A, B).

Figure 18. 

Pseudopoda xiaozhua sp. nov., male holotype, palpal bulb (A–C) and habitus (D, E). A. Prolateral; B. Ventral; C. Retrolateral; D. Dorsal; E. Ventral. Abbreviations: C = conductor; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; TA = tegular apophysis. Scale bars: 0.5 mm (equal for A–C); 2 mm (equal for D, E).

Figure 19. 

Pseudopoda xiaozhua sp. nov., female paratype, epigyne (A–C) and habitus (D, E). A. Intact; ventral; B. Cleared and macerated; ventral; C. Cleared and macerated; dorsal; D. Dorsal; E. Ventral. Abbreviations: AB = anterior band; aEF = anterior margin of epigynal field; amLL = anterior margin of lateral lobes; CO = copulatory opening; FD = fertilization duct; FW = first winding; LL = lateral lobe; MF = median field of epigyne; mmLL = median margin of lateral lobes; MS = membranous sac; PI = posterior incision; pmLL = posterior margin of lateral lobes; S = spermatheca. Scale bars: 0.2 mm (equal for A–C); 2 mm (equal for D, E).

Male (YNZY003). Total length 10.9. Carapace 5.5 long, 5.2 wide; anterior width 2.7. Opisthosoma 5.4 long, 3.6 wide. Eye sizes and interdistances: AME 0.31, ALE 0.44, PME 0.36, PLE 0.42, AME–AME 0.19, AME–ALE 0.09, PME–PME 0.27, PME–PLE 0.41, AME–PME 0.37, ALE–PLE 0.37, CH AME 0.58, CH ALE 0.45. Spination: palp: 131, 101, 2101; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–II 2026, III–IV 2126; Mt: I–II 1014, III 2024, IV 3036. Measurements of palp and legs: palp 8.4 (2.7, 1.3, 1.4, 3.0), I 27.7 (7.4, 3.5, 6.3, 7.8, 2.7), II 30.3 (8.1, 3.0, 8.2, 8.3, 2.7), III 23.0 (6.5, 2.4, 6.0, 6.2, 1.9), IV 26.6 (7.6, 2.4, 6.5, 7.9, 2.2). Cheliceral furrow with ~50 denticles.

Colouration in ethanol (Fig. 18D, E). DS yellowish brown, marked with numerous small spots along radial grooves, clothed with fine hairs; lateral bands and median band inconspicuous, not distinctly delimited; cervical groove not distinct, fovea and radial grooves distinct. Cheliceral base light brown, with red fang. Sternum yellowish-white, margin slightly darker. Endites and labium coloured as cheliceral base. Legs coloured as DS, with numerous spots, and bearing short spines. OS oval, dorsum laterally with bright patterns, centrally with Y-shaped median band, with a pair of circular dots on each side of the median band, posteriorly with large ‘)(‘-shaped black pattern, transverse line indistinct; venter of OS medially with a pair of diagonal broken lines.

Palp (Figs 16, 17, 18A–C, 28D). Femur and patella unmodified. Tibia (Ti) moderately long, ca. 2/5 cymbium length, with retrolateral apophysis (RTA) arising proximally to medially; RTA bifurcated, with ventral part (vRTA) and dorsal branch (dRTA): dRTA finger-like, distinctly long, nearly as long as tibia, almost reaching cymbial bulge (CB); vRTA humble and broad base and papilliform apex, ca.1/2 of dRTA length. Cymbium (Cy) ca. 2.1× longer than wide, basoretrolaterally with distinct, nearly triangular bulge (CB). Tegulum (T) egg-shaped, ca. 1.15× longer than wide, relatively flattened, prolatero-apically slightly excavated, with heavily sclerotised tegular apophysis (TA); spermophor (Sp) distinct, V-shaped in ventral view. Tegular apophysis (TA) strongly expanded, inserted at apico-prolateral portion of tegulum; proximally exposed and petal-shaped; mesially and distally digitiform, hidden behind embolus. Embolus (E) almost as long as tegulum (T); basally and mesially thick and robust, nearly column-shaped, originated at ~8–9 o’clock; embolic tip (ET) distinctly narrowed and curved, claw-shaped in ventral view, apex sharp, terminated at ~11 o’clock position. Conductor (C) membranous, ca. 1/2 of the embolus length, extending obliquely, arising at ca. 1 o’clock position from tegulum, terminating at c. 11 o’clock position; conductor (C) proximally narrowed, its tip widened, shaped like the membranous wing of hymenoptera, directed prolaterally and apically beyond embolic tip (ET).

Female (YNZY004). Total length 13.4. Carapace 6.3 long, 5.5 wide; anterior width 3.3. Opisthosoma 7.1 long, 5.1 wide. Eye sizes and interdistances: AME 0.32, ALE 0.47, PME 0.37, PLE 0.41, AME–AME 0.18, AME–ALE 0.07, PME–PME 0.32, PME–PLE 0.51, AME–PME 0.42, ALE–PLE 0.42, CH AME 0.63, CH ALE 0.51. Spination: palp: 131, 101, 2121, 1014; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–IV 2026; Mt: I–II 1014, III 2024, IV 3036. Measurements of palp and legs: palp 8.0 (2.5, 1.3, 1.7, 2.5), I 21.4 (6.4, 2.8, 5.1, 5.3, 1.8), II 22.4 (6.8, 2.6, 5.6, 5.7, 1.7), III 18.9 (6.0, 2.5, 4.8, 4.5, 1.1), IV 21.0 (6.5, 2.4, 5.3, 5.5, 1.3). Cheliceral furrow with ~56 denticles. Colouration in ethanol as in males, but generally distinctly darker (Fig. 19D, E).

Epigyne (Fig. 19A–C). Epigynal field ca. 1.5× wider than long; anterior margin (aEF) trilobate, with two small v-shaped incisions that are well separated by at least 3× widths; anterior bands (AB) indistinct, situated at the two incisions. Median field (MF) nearly heart-shaped, relatively small, no more than 1/3 epigyne length and 1/4 epigyne width. Lateral lobes (LL) distinctly longer than wide, slightly converged on the central axis; anterior margins (amLL) distinctly procurved and delimited; median margins (mmLL) touching each other along the middle line in anterior half; posterior margins (pmLL) with incision (PI) on each side. Copulatory openings (CO) indistinct, located at anterolateral margins of median field (MF). First windings (FW) represented by translucent, membranous short tube, starting from near copulatory openings (CO), descending obliquely, moving laterally to basolateral surfaces of spermathecae (S), ca. 1/2 of epigyne length. Spermathecae (S) clavate, at least 2.7 longer than diameters; surface wrinkled, provided with several depressions and anterior hump; spermathecae (S) widely separated by ca. 2.4× diameters. Membranous sac (MS) disc-shaped, located at posterior portion of vulva; anterior margin separated from epigastric fold by ca. 1/3 epigyne length, reaching the contact point of median margins of lateral lobes (mmLL); posterior margin close to the epigastric fold. Fertilization ducts (FD) acicular, membranous, nearly 1/2 spermathecae length, proximally covered by membranous sac (MS).

Presently known only from the type locality (Fig. 1).

♂ (YNZY001), China: • Yunnan Pro.: Honghe Hani and Yi Autonomous Prefecture: Pingbian Co., Daweishan National Park, 22.94°N, 103.70°E, c. 2365 m, by hand, 14 IV 2024, Y. Zhong & S. Yang leg. Paratypes: • 3 ♂ 4 ♀ (YNZY002, YNZY005, YNZY006, YNZY008, YNZY017, YNZY020, YNZY021), same data as holotype.

The specific name is dedicated to Ms. Siyu Yang (Xianning, China), collector of several specimens examined in this study.

The males of the new species resemble those of P. amelia Jäger & Vedel, 2007; P. putaoensis Zhao & Li, 2018; and P. zhangi Fu & Zhu, 2008 in having a ɔ-shaped embolus (E) (Figs 20A, 22B; Jäger and Vedel 2007: figs 32, 33; Jiang et al. 2018: figs 18A, B, 19A, B; Fu and Zhu 2008: fig. 4), but differs by: (1) dorsal branch of RTA (dRTA) blade-shaped, reaching basal part of cymbium (Cy) (vs. distinctly shorter, not reaching cymbial base in P. amelia; finger-like in P. putaoensis and P. zhangi) (cf. Figs 20A, B, 21B, 28E, F and Jäger and Vedel 2007: figs 33, 34 and Jiang et al. 2018: figs 18B, C and Fu and Zhu 2008: figs 4, 5); (2) in retrolateral view, ventral branch of RTA (vRTA) claviform and distinctly protruding (vs. papilliform in P. amelia, laminar in P. putaoensis and P. zhangi, humble and barely protruding in P. amelia, P. putaoensis and P. zhangi) (cf. Figs 21B, 28E, F and Jäger and Vedel 2007: fig. 34 and Jiang et al. 2018: fig. 18C and Fu and Zhu 2008: fig. 5). Female of P. yangae sp. nov. is similar to that of P. emei F. Zhang, B. S. Zhang & Z. S. Zhang, 2013 by the similarly shaped median field (MF), but can be recognised by: (1) anterior bands (AB) indistinct (vs. distinct) (cf. Fig. 23A, B, D, E and Zhang et al. 2013b: fig. 25 and Jäger et al. 2015: figs 24, 29); (2) in dorsal view, lateral lobes (LL) without ridges, length of lateral margin of lateral lobes distinctly longer than that of median margin (mmLL) (vs. with distinct ridges, length of lateral margin almost equal to median margin) (cf. Fig. 23C, F and Zhang et al. 2013b: figs 26, 31 and Jäger et al. 2015: fig. 25); (3) first winding (FW) distinctly thinner, ca. 1/15–1/20 of epigyne width, widely separated by ca. 15–17× diameters (vs. relatively thicker, ca. 1/10–1/12 of epigyne width, separated by ca. 5× diameters) (cf. Fig. 23C, F and Zhang et al. 2013b: figs 26, 31 and Jäger et al. 2015: fig. 25); (4) first winding (FW) entirely covered by lateral lobes (LL) (vs. anterior half of first winding (FW) exposed, not covered by lateral lobes (LL)) (cf. Fig. 23C, F and Zhang et al. 2013b: figs 26, 31 and Jäger et al. 2015: fig. 25).

Figure 20. 

Male palp of the holotype of Pseudopoda yangae sp. nov. A. Ventral; B. Dorsal. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral branch of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 21. 

Male palp of the holotype of Pseudopoda yangae sp. nov. A. Prolateral; B. Retrolateral. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal branch of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral branch of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 22. 

Pseudopoda yangae sp. nov., male holotype, palpal bulb (A–C) and habitus (D, E). A. Prolateral; B. Ventral; C. Retrolateral; D. Dorsal; E. Ventral. Abbreviations: C = conductor; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum. Scale bars: 0.2 mm (equal for A–C); 2 mm (equal for D, E).

Figure 23. 

Pseudopoda yangae sp. nov., female paratypes, YNZY002 (A–C) and YNZY006 (D–E), epigyne, showing the intraspecific variations exhibited by different females, related to different degrees of sclerotization of amMF. A, D. Intact, ventral; B, E. Cleared and macerated, ventral; C, F. Cleared and macerated, dorsal. Abbreviations: aEF = anterior margin of epigynal field; amLL = anterior margin of lateral lobes; amMF = anterior margin of median field; CO = copulatory opening; FD = fertilization duct; FW = first winding; LL = lateral lobe; MF = median field of epigyne; mmLL = median margin of lateral lobes; MS = membranous sac; PI = posterior incision; pmLL = posterior margin of lateral lobes; S = spermatheca. Scale bars: 0.5 mm (equal for A–C, equal for D–E).

Figure 24. 

Pseudopoda yangae sp. nov., female paratypes, YNZY002 (A, B) and YNZY006 (C, D), habitus, showing the intraspecific variations exhibited by different females, related to different abdominal pattern. A, C. Dorsal; B, D. Ventral. Scale bars: 2 mm (equal for A, B; equal for C, D).

Male (YNZY001). Total length 7.5. Carapace 3.9 long, 3.6 wide, anterior width 1.9. Opisthosoma 3.6 long, 2.4 wide. Eye sizes and interdistances: AME 0.19, ALE 0.31, PME 0.24, PLE 0.31, AME–AME 0.13, AME–ALE 0.04, PME–PME 0.20, PME–PLE 0.27, AME–PME 0.29, ALE–PLE 0.26, CH AME 0.29, CH ALE 0.20. Spination: palp: 131, 101, 2101; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–II 2126, III–IV 2226; Mt: I–II 1014, III 2024, IV 3036. Measurements of palp and legs: palp 6.0 (2.1, 1.0, 1.1, 1.8), I 19.2 (5.4, 1.8, 5.2, 5.1, 1.7), II 20.9 (5.7, 1.9, 5.8, 5.5, 2.0), III 15.3 (4.0, 1.4, 4.3, 4.1, 1.5), IV 17.6 (5.2, 1.6, 4.7, 4.5, 1.6). Cheliceral furrow with ~32 denticles.

Colouration in ethanol (Fig. 22D, E). DS yellowish, marked with numerous small spots along radial grooves, lateral bands slightly darker, clothed with fine hairs; median band bright yellowish-brown, not distinctly delimited to lateral bands, with indistinct Ψ-shaped markings starting from behind PER, almost reaching fovea; fovea and striae distinctly marked. Cheliceral base yellowish white, fang reddish. Sternum yellowish white, margin slightly darker. Endites and labium light orange. Legs yellowish white, marked with numerous spots. OS oval, dorsum laterally with a pair of arc-shaped stripes, centrally with short, longitudinal median band, posteriorly with thick transverse line; venter of OS medially with a pair of indistinct, diagonal broken lines, posteriorly with triangular marking.

Palp (Figs 20, 21, 22A–C, 28E). Femur and patella unmodified. Tibia (Ti) moderately long, ca. 2/3 cymbium length, with retrolateral apophysis (RTA) arising proximally to medially; RTA bifurcated, both ventral and dorsal branches distinctly protruding: dorsal branch (dRTA) blade-shaped, basally slightly curved, no more than 1/2 of the tibia length; ventral branch (vRTA) almost as long as dRTA, claviform, basally and distally slightly widened, medially slightly narrowed. Cymbium (Cy) ca. 2.1× longer than wide, retrolaterally with distinct blunt bulge (CB). Tegulum (T) elongate-oval, proximally strongly bulged and prolapsed, distinctly excavated on prolatero-apical side to accommodate embolus (E); spermophor (Sp) distinct, U-shaped in ventral view, oriented clockwise along tegular margin. Embolus (E) robust, slightly longer than tegulum, more or less ɔ-shaped in ventral view and Ƨ-shaped in prolateral view; embolic base (EB) broadened, inserted prolaterally (approximately 9 o’clock relative to tegulum); embolic tip (ET) distinctly curved, terminated at ~12 o’clock position, apex sharply pointed, directed prolaterally. Conductor (C) situated apically, irregularly shaped, covering embolic tip (ET) in prolateral and ventral views.

Female (YNZY002). Total length 9.7. Carapace 4.5 long, 4.3 wide, anterior width 2.5. Opisthosoma 5.2 long, 3.8 wide. Eye sizes and interdistances: AME 0.22, ALE 0.36, PME 0.28, PLE 0.34, AME–AME 0.20, AME–ALE 0.11, PME–PME 0.26, PME–PLE 0.39, AME–PME 0.38, ALE–PLE 0.37, CH AME 0.45, CH ALE 0.36. Spination: palp: 131, 101, 2121, 1014; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–II 2026, III–IV 2126; Mt: I–II 1014, III 2026, IV 3036. Measurements of palp and legs: palp 6.0 (1.9, 1.1, 1.2, 1.9), I 16.7 (4.8, 2.1, 4.3, 4.1, 1.4), II 17.6 (5.2, 2.2, 4.6, 4.2, 1.5), III 14.1 (4.4, 1.8, 3.5, 3.2, 1.3), IV 15.8 (4.9, 1.7, 3.8, 4.1, 1.5). Cheliceral furrow with ~44 denticles. Colouration in ethanol as in males, but generally distinctly darker (Fig. 24A, B).

Epigyne (Fig. 23 A–C). Epigynal field nearly as wide as long; anterior margin (aEF) recurved, without incision; anterior bands (AB) indistinct. Median field (MF) more or less cordiform, large, more than 1/2 epigyne length and 2/3 epigyne width, anterior margin (amMF) indistinct. Lateral lobes (LL) distinctly longer than wide; anterior margins (amLL) distinctly delimited, V-shaped; median margins (mmLL) touching each other along the middle line in anterior half; posterior margins (pmLL) curved, with distinct median indentation and distinct posterior incisions (PI) on each side. Copulatory openings (CO) indistinct, located at anterolateral borders of median field (MF). First windings (FW) entirely covered by lateral lobes (LL), represented by translucent, slightly curved long tubes, starting from near copulatory openings (CO), descending longitudinally, almost extending posteriorly to level of posterior parts of spermathecae (S), nearly as long as epigyne. Spermathecae (S) fist-shaped, ca. 1.5× wider than long; relatively sclerotised, surface wrinkled, provided with several depressions and lateral hump; inner parts of the spermathecae covered by membranous sac (MS), separated by 0.3× widths. Membranous sac (MS) hyaline, nearly trapeziform, located at posterior portion of vulva; anterior margin separated from epigastric fold by ca. 1/2 epigyne length, beyond the contact point of lateral lobes (LL); posterior margin close to the epigastric fold. Fertilization ducts (FD) acicular, membranous, nearly 1/2 spermathecae length.

Presently known only from the type locality (Fig. 1).

Both males and females exhibit some morphological variation among different individuals: for males, mostly related to different shapes of vRTA and dRTA (vRTA medially slightly narrowed, and dRTA apically more sharp in some males, such as in holotype, YNZY001 vs. vRTA medially more narrowed, dRTA apically more blunt in some males, such as in YNZY005; cf. Fig. 28E and Fig. 28F); for females, mostly related to different degrees of sclerotization of amMF and curve of amLL (amMF indistinct, amLL relatively straight in some females, such as in YNZY002 vs. amMF distinct, amLL distinctly curved in some females, such as in YNZY006; cf. Fig. 23A, B and Fig. 23D, E), and the different abdominal pattern (dorsum of abdomen posteriorly with a narrow, W-shaped transverse line in some females, such as in YNZY002 vs. with a broad, W-shaped transverse band in some females, such as in YNZY006; cf. Fig. 24A and Fig. 24C). However, all molecular species delimitation analyses results show that the different individuals exhibiting considerable morphological variations should be classified as the same species (Fig. 2; for details see the results and discussion section). After careful examination, the variations listed above were determined to be intraspecific differences.

♂ (YNZY007), China: • Yunnan Pro.: Honghe Hani and Yi Autonomous Prefecture: Pingbian Co., Daweishan National Park, 22.94°N, 103.70°E, c. 2365 m, by hand, 14 IV 2024, Y. Zhong & S. Yang leg. Paratype: • 1 ♂ (YNZY026), same data as holotype.

The specific name is derived from the Chinese pinyin ‘yìng’, which means ‘hard’, referring to the conductor heavily sclerotized and well developed; adjective.

Median-sized Sparassidae with body length of males 7.8 mm, belonging to the daliensis species group. Male of the new species can be easily distinguished from all other congroupers, with the exception of P. sicyoidea, by having similar palp with Ƨ-shaped, wide embolus (E), and broad ventral part of RTA (vRTA) with two distinct margins that resemble mountains, but can be distinguished by: (1) conductor (C) exhibits high degree of sclerotization (vs. membranous) (cf. Figs 25A, 26A, B, 27A–C and Zhang et al. 2017: 274, figs 13A, 14A); (2) embolic tip (ET) blunt, slightly curved, directing prolatero-distally, terminating at c. 12 o’clock position (vs. sharp and peak-shaped, distinctly curved, directing proximally, terminating at c. 10–11 o’clock position) (cf. Figs 25A, 27B and Zhang et al. 2017: 274, figs 12A, 14A).

Figure 25. 

Male palp of the holotype of Pseudopoda ying sp. nov. A. Ventral; B. Dorsal. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal part of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 26. 

Male palp of the holotype of Pseudopoda ying sp. nov. A. Prolateral; B. Retrolateral. Abbreviations: C = conductor; CB = cymbial bulge; Cy = cymbium; dRTA = dorsal part of RTA; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum; Ti = palpal tibia; vRTA = ventral part of RTA. Scale bar: 0.5 mm (equal for A, B).

Figure 27. 

Pseudopoda ying sp. nov., male holotype, palpal bulb (A–C) and habitus (D, E). A. Prolateral; B. Ventral; C. Retrolateral; D. Dorsal; E. Ventral. Abbreviations: C = conductor; E = embolus; EB = embolic base; ET = embolic tip; Sp = spermophor; St = subtegulum; T = tegulum. Scale bars: 0.5 mm (equal for A–C); 1 mm (equal for D, E).

Male (YNZY007). Total length 7.8. Carapace 3.9 long, 3.6 wide, anterior width 2.0. Opisthosoma 3.9 long, 2.4 wide. Eye sizes and interdistances: AME 0.28, ALE 0.35, PME 0.29, PLE 0.30, AME–AME 0.18, AME–ALE 0.07, PME–PME 0.24, PME–PLE 0.36, AME–PME 0.34, ALE–PLE 0.28, CH AME 0.34, CH ALE 0.28. Spination: palp: 131, 101, 2101; Fe: I–III 323, IV 321; Pa: I–IV 101; Ti: I–IV 2226; Mt: I–II 1014, III 2026, IV 3036. Measurements of palp and legs: palp 6.0 (2.1, 0.7, 1.1, 2.1) I 24.3 (7.7, 2.0, 6.7, 6.2, 1.7); II 28.0 (7.9, 2.7, 8.2, 7.1, 2.1); III 19.6 (5.7, 1.7, 5.5, 5.2, 1.5); IV 23.4 (7.3, 1.4, 6.2, 6.7, 1.8). Cheliceral furrow with ~26 denticles.

Colouration in ethanol (Fig. 27D, E): DS yellowish brown, darker anteriorly and marginally; median band bright yellowish-brown, distinctly delimited to black lateral bands, with distinct Ψ-shaped markings starting from behind PER, almost reaching distinct fovea; fovea and radial grooves distinctly marked. Cheliceral base yellowish white, with red fangs. Sternum yellowish white, marked with numerous purplish spots. Endites and labium coloured as sternum. Legs dark yellowish-brown, with numerous brown spots, and covered by short spines. OS elongate-oval, dorsum anteriorly with median band, reaching 4/5 of abdomen length; median band consisting of cross-shaped anterior stripe and nearly square-shaped posterior stripe, the two stripes fused; ventral OS medially with V-shaped markings.

Palp (Figs 25, 26, 27A–C, 28G). Femur and patella unmodified. Tibia (Ti) relatively short, ca. 2/5 cymbium length, with retrolateral apophysis (RTA) arising proximally to mesially; RTA subdivided, with ventral part (vRTA) and dorsal part (dRTA): dRTA basally wide, mesially and distally narrowed, apex nearly triangular, ca. 1/2 length of tibia; vRTA broad and slightly shorter than dRTA, with two distinct margins that look like mountains, among them the dorsal margin with quadrilateral-shaped apophysis. Cymbium (Cy) relatively short and wide, ca. 2.1× longer than wide, retrolaterally with a large bulge (CB). Tegulum (T) elongate-oval, ca. 1.3× longer than wide, strongly bulged and prolapsed. Spermophor (Sp) sinuate, forming a loop along tegular margin. Embolus (E) broad, ca. 4/5 of tegulum length, and wider than 1/2 of tegulum width, Ƨ-shaped in ventral view, arising at approximately the 8–9 o’clock position, terminating at c. 12 o’clock position; embolic tip (ET) wide and blunt, apex rugged. Conductor (C) heavily sclerotized, c. 1/2 of the embolus length, originating at 12–1 o’clock position portion of tegulum; the base of conductor (C) relatively narrow, inserted dorsally to embolus; the tip of conductor (C) relatively wide and extending above apex of embolus.

Figure 28. 

RTA of Pseudopoda spp. treated in this paper, retrolateral-dorsal A. P. huanglianensis; B. P. mamillaris; C. P. oliviformis; D. P. xiaozhua sp. nov.; E, F. P. yangae sp. nov., YNZY001 (E) and YNZY005 (F), showing the intraspecific variations exhibited by different males, related to different shapes of vRTA and dRTA; G. P. ying sp. nov. Abbreviations: dRTA = dorsal branch/part of RTA; vRTA = ventral branch/part of RTA. Scale bars: 0.2 mm (A–C, G, equal for E, F); 5 mm (D).

Female. Unknown.

A total of four Pseudopoda species are known only from females in Honghe Prefecture: P. daweiensis, P. rhopalocera, P. zhaoae, and P. zuoi (Table 1). However, none of them could be matched with P. ying sp. nov. due to their different habitus: abdomen with a distinct median band, which consists of a cross-shaped anterior stripe and a nearly square-shaped posterior stripe in P. ying sp. nov. vs. abdominal median band indistinct and not as above in P. daweiensis, P. rhopalocera, P. zhaoae, and P. zuoi (cf. Fig. 21D and Zhang et al. 2023a: figs 85A, 206A, 278A, 282A and Yang et al. 2009: fig. F). Moreover, males of P. ying sp. nov. exhibit typical daliensis-group features (diagnosis of P. daliensis-group, see Zhang et al. 2017: 273) and resemble the core species of the daliensis-group, such as P. sicyoidea, for their characteristic embolus and RTA (for a detailed diagnosis, see above). In contrast, females of P. daweiensis, P. rhopalocera, P. zhaoae and P. zuoi exhibit the following distinctive suite of characters, here contrasted with the corresponding condition in daliensis-group: amLL distinctly oblique in P. daweiensis and P. zuoi (vs. amLL almost transversal to body length axis in daliensis-group species) (cf. Zhang et al. 2023a: figs 84A, 281A and Zhang et al. 2017: figs 2B, 4C, 6A, 9A, 12B, 14C, 16A, 17A); spermathecae covered by first windings in P. rhopalocera, located internally to first windings in P. zhaoae, not extending laterally beyond first winding (vs. extending laterally beyond first winding in daliensis-group species) (cf. Zhang et al. 2023a: figs 205B, 277C and Zhang et al. 2017: figs 2D, 4D, 6D, 9D, 12D, 14D, 16B, 17B). Therefore, we can rule out the possibility that the above four species are conspecific with P. ying sp. nov.

Presently known only from the type locality (Fig. 1).