Scorpiops Peters, 1861

Kovařík, František, Lowe, Graeme, Stockmann, Mark & Šťáhlavský, František, 2020, Revision of genus-group taxa in the family Scorpiopidae Kraepelin, 1905, with description of 15 new species (Arachnida Scorpiones), Euscorpius 325, pp. 1-140 : 32-33

Genus Scorpiops Peters, 1861

( Figures 1–75 , 77–81 , 90–103, 105–127, 129–254, 257–815, Tables 1 –9) http://zoobank.org/urn:lsid:zoobank.org:act:45E3D60F-

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Scorpiops Peters, 1861: 510 ; Kraepelin, 1899: 179–182; Vachon, 1980: 143–159, figs. 1–40; Tikader & Bastawade, 1983: 403–452, figs. 1129–1246; Francke, 1985: 13 (in part); Sissom, 1990: 114; Kovařík, 1998: 142; Lourenço, 1998: 246; Fet, 2000: 491–495 (in part); Kovařík, 2000: 162–198, figs. 1–7, 14–22, 25, 28, 29, 31–72 (in part); Soleglad & Sissom, 2001: 93–97, figs. 3, 13–14, 86, 96, 101, 114, 147, 159, 183, 202, 220 (in part); Qi et al., 2005: 2–14, figs. 1–46; Dupré, 2007: 10 (in part); Kovařík, 2009: 27, 32; Kovařík & Ahmed, 2009: 1–10, figs. 1–26; Di et al., 2011a: 1–2 (in part); Francke, 2019: 15; Kovařík, 2020: 1–43 , figs. 1–241, tables 1–3.

= Dasyscorpiops Vachon, 1974: 952–953 , figs. 142, 147, 160– 162 (type species Dasyscorpiops grandjeani Vachon,

1974); Francke, 1976: 82; Francke, 1985: 7; Kovařík, 1998: 141; Fet, 2000: 488; Kovařík, 2000: 156; Soleglad & Sissom, 2001: 93–97, figs. 5, 16, 95, 104; Dupré, 2007: 5; Kovařík, 2009: 32; Francke, 2019: 7. Syn. n.

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= Scorpiops (Neoscorpiops) Vachon, 1980: 150 , 151, figs. 10– 16 (type species Scorpiops montanus satarensis Pocock, 1900 ); Tikader & Bastawade, 1983: 382–483, figs. 1060– 1128; Francke, 1985: 13. Syn. n.

http://zoobank.org/ urn:lsid:zoobank.org:act:E04E0206-554E-47BC-9BCC-28D94DA02843

= Scorpiops (Alloscorpiops) Vachon, 1980: 151 , figs. 18–25 (type species Scorpiops anthracinus Simon, 1887 ); Francke, 1985: 13. Syn. n.

http://zoobank.org/ urn:lsid:zoobank.org:act:7EDB9233-E168-4E52-AB25-4B5066112656

= Scorpiops (Euscorpiops) Vachon, 1980: 155 , figs. 26–32 (type species Scorpiops asthenurus Pocock, 1900 ); Tikader & Bastawade, 1983: 452–476, figs. 1247–1305; Francke, 1985: 13 (syn. by Kovařík, 2000: 164, restored to generic status by Soleglad & Sissom, 2001: 93, and synonymized here again).

http://zoobank.org/ urn:lsid:zoobank.org:act:AE9E0E91-2234-424D-89FE-6716B2ACCF38

Alloscorpiops: Stockwell, 1989: 120 ; Fet, 2000: 487–488; Kovařík, 2000: 155–156, figs. 8, 24, 26; Soleglad & Sissom, 2001: 49, 93, figs. 4, 61, 99, 103, 117, 136, 143, 177, 188–189, 201; Dupré, 2007: 2; Kovařík, 2009: 32; Kovařík, 2013: 1–11, figs. 1–24.

Euscorpiops: Stockwell, 1989: 120 ; Fet, 2000: 488–489; Soleglad & Sissom, 2001: 93–97, figs. 6, 15, 60, 65, 97, 115, 145, 178, 185–186, 190, 203, 222; Dupré, 2007: 5; Kovařík, 2009: 32; Di et al., 2010 b: 50; 2011b: 5; Francke, 2019: 8.

Neoscorpiops: Stockwell, 1989: 120 ; Fet, 2000: 489–490; Kovařík, 2000: 158–162, figs. 9–12, 27, 30; Soleglad & Sissom, 2001: 93–97, figs. 61, 67, 98, 102, 113, 125, 131– 132, 176, 184, 200, 221, 223; Dupré, 2007: 7; Kovařík, 2009: 32; Francke, 2019: 11.

= Alloscorpiops (Laoscorpiops) Lourenço, 2013a: 52–55 , figs. 2–3 (type species Alloscorpiops (Laoscorpiops) calmonti Lourenço, 2013 ); Francke, 2019: 3 (syn. by Kovařík et al., 2013: 1).

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= Scorpiops (Vietscorpiops) Lourenço & Pham, 2015a: 212– 217 , figs. 1–6 (type species Scorpiops (Vietscorpiops) dentidactylus Lourenço & Pham, 2015 ). Syn. n.

http://zoobank.org/ urn:lsid:zoobank.org:act:91B89979-0C30-4BB0-A7AB-0B1FACF81886

= Plethoscorpiops Lourenço, 2017: 352–357 , figs. 6–10 (type species Plethoscorpiops profusus Lourenço, 2017 ); Francke, 2019: 14. Syn. n.

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Vietscorpiops: Pham et al., 2017: 136 .

Alloscorpiops (Alloscorpiops) : Francke, 2019: 3.

Type species. Scorpio hardwickii Gervais, 1843 .

DIAGNOSIS. See family diagnosis.

SUBORDINATE TAXA.

Scorpiops affinis Kraepelin, 1898

Scorpiops afghanus Lourenço & Qi, 2006

Scorpiops alexandreanneorum (Lourenço, 2013) , comb. n. Scorpiops anthracinus Simon, 1887 , comb. n.

= Scorpiops lindstroemii Thorell, 1889

= Scorpiops lugubris Thorell, 1889

Scorpiops artemisae (Kovařík et al., 2015) , comb. n. Scorpiops asthenurus Pocock, 1900

Scorpiops bastawadei sp. n.

Scorpiops beccaloniae (Kovařík, 2005) , comb. n.

Scorpiops bhutanensis Tikader & Bastawade, 1983 Scorpiops binghamii Pocock, 1893

Scorpiops birulai sp. n.

Scorpiops braunwalderi Kovařík, 2000

Scorpiops calmonti (Lourenço, 2013) , comb. n.

Scorpiops cavernicola (Lourenço & Pham, 2013) , comb. n. Scorpiops ciki sp. n.

Scorpiops chiangmai (Lourenço, 2019) , comb. n., nomen dubium

Scorpiops citadelle (Kovařík, 2013) , comb. n.

Scorpiops dakrong (Lourenço & Pham, 2014)

Scorpiops dastychi Kovařík, 2000

Scorpiops deccanensis Tikader & Bastawade, 1977 Scorpiops demisi Kovařík, 2005

Scorpiops dii sp. n.

Scorpiops dunlopi sp. n.

Scorpiops farkaci Kovařík, 1993

= Scorpiops (Vietscorpiops) dentidactylus Lourenço & Pham, 2015 , syn. n.

Scorpiops feti Kovařík, 2000

Scorpiops furai Kovařík, 2020

Scorpiops grandjeani ( Vachon, 1974) , comb. n.

Scorpiops grosseri Kovařík, 2020

Scorpiops hardwickii ( Gervais, 1843)

= Scorpiops crassimanus Pocock, 1899

=? Scorpiops insculptus Pocock, 1900

=? Chaerilus pirpanjalus Mani, 1959

Scorpiops harmsi Kovařík, 2020

Scorpiops hofereki Kovařík, 2020

Scorpiops ingens Yin et al., 2015

Scorpiops irenae Kovařík, 1994

Scorpiops jendeki Kovařík, 1994

Scorpiops kaftani Kovařík, 1993

Scorpiops kamengensis (Bastawade, 2006) , comb. n. Scorpiops kautti sp. n.

Scorpiops kejvali Kovařík, 2020

Scorpiops krabiensis sp. n.

Scorpiops kubani ( Kovařík, 2004) , comb. n.

Scorpiops langxian Zhu, Qi & Lourenço, 2005

Scorpiops leptochirus Pocock, 1893

Scorpiops lii (Di & Qiao, 2020) , comb. n.

Figures

Figures 1–2. Scorpiops hardwickii, India, Uttaranchal State, ca. 30 km N of Bageshwar, SE of Dhakuri, 2600-2800 m a. s. l., FKCP, ♀, dorsal (1) and ventral (2) views. Scale bar: 10 mm.

Figures 3–9: Variation in positions of trichobothria Eb and Dt on the manus of the pedipalp chela of scorpiopids. Figure 3. Bivariate logarithmic 3 scatter plot of normalized positions of Eb 3 vs. Dt along the proximal-distal axis of the manus (normalized by manus length, mL), for males (triangles, 61 species) and females (circles, 56 species). Biometrics defined in right page inset. Data compiled from published trichobothrial maps, and study of our own material. Each symbol represents one species with a total of 84 species (= 90% of the 93 known scorpiopids). For illustrative purposes, symbols are color-coded (color key in lower right page inset) by old generic assignments determined by trichobothriotaxic scheme of Soleglad & Sissom (2001), including new species described herein under Scorpiops. Eb is level with Dt along the black diagonal line, distal to it 3 above the line, and proximal to it below the line. Gray line is least squares regression fit to all 117 points, showing weak or insignificant correlation between Eb 3 and Dt position. “Alloscorpiops” calmonti (indicated arrow) clusters high with other “Alloscorpiops”, assuming homology of midmanus trichobothrium with Eb 3. Figure 4. Scatter plot of Fig. 3, with 5 specimens of S. montanus highlighted in black symbols. Intraspecific variation encompasses cases with Eb distal to, level with, or proximal to Dt, invalidating the use of the relative positions of these two trichobothria 3 as a taxonomic character. Figure 5. Histogram showing distribution of position of Eb relative to Dt. Vertical separation from the diagonal of each 3 point in Fig. 3 was calculated and binned at intervals of 0.05. Zero value on abscissa corresponds to Eb 3 level with Dt. Figures 6–7. Logarithmic scatter plots of normalized position of Eb 3 along proximal-distal axis vs. L/D ratio of manus for 61 male (6) and 56 female (7) scorpiopids. Female “A.” calmonti (7, upper arrow) clusters with other “Alloscorpiops” and with “Euscorpiops” having elongated manus, if we interpret the mid-manus trichobothrium as Eb. Female “A.” citadelle (7, lower arrow) is separated from other “Alloscorpiops” by having less distal Eb, although it has 3 3 an elongated manus. Gray lines are least squares regressions, showing significant positive correlation between distal placement of Eb and manus 3 elongation. Figures 8–9. Scatter plots of position of Eb 3 relative to Dt along proximal-distal axis vs. L/D ratio of manus for 60 male (8) and 56 female (9) scorpiopids. Abscissa scale is logarithmic. Gray lines are least squares regressions, showing significant positive correlation between distal placement of Eb 3 relative to Dt and manus elongation. R=Pearson´s correlation coefficient.

Figures 10–20: Scorpiops spp., pedipalp chela external (10–16, 18 and 20) and dorsal (17 and 19) views. Trichobothria Dt and Eb are 3 indicated by white open circles. Figures 10–14. S. montanus, juvenile, holotype, ZMHB (10), female (11) and male reared from litter of female 11 (12), India, Uttarakhand State, Dhika forest, 29.65°N 78.94°E, FKCP, female (13) and male (14) from India, Uttarakhand State, Rishikesh, 30.13°N 78.32°E, FKCP. In the juvenile holotype, Eb 3 is located distal to Dt (Fig. 10); in the female from Dhika forest, Eb 3 is located slightly distal to Dt (Fig. 11) but the male from her litter has Eb 3 proximal to Dt (Fig. 12); in both specimens from Rishikesh, Eb 3 is proximal to Dt but its relative position differs between individuals (Figs. 13–14). Figure 15. S. hardwickii, female, India, Uttaranchal State, ca. 30 km of Bageshwar, SE of Dhakuri vill., 2600–2800 m a. s. l., FKCP. Figure 16. S. asthenurus, male, India, Meghalaya State, Nong Poh env., FKCP. Figures 17–18. S. petersii, male, India, Himachal Pradesh State, Molta, FKCP. Figures 19–20. S. vonwicki, female, holotype, ZISP.

Figures 21–52: Scorpiops spp., telson lateral. Figure 21. S. affinis, male, lectotype. Figure 22. S. afghanus, female, holotype. Figure 23. S. braunwalderi, male, holotype. Figure 24. S. dastychi, female, paratype. Figure 25. S. demisi, female, holotype. Figure 26. S. farkaci, male, holotype.Figure 27. S.feti, male,paratype.Figure 28. S.furai, female, holotype.Figure 29. S.grosseri, female, holotype.Figure 30. S.hardwickii, male, holotype. Figure 31. S. harmsi, female, holotype. Figure 32. S. hofereki, female, holotype. Figure 33. S. irenae, female, holotype. Figure 34. S. jendeki, male, holotype. Figure 35. S. kejvali, male, holotype. Figure 36. S. langxian, male, China, Tibet, W. Kangtissu Shan Mts., Nakaeh Cushul, 4200 m a. s. l., FKCP. Figure 37. S. leptochirus, male (306), India, Meghalaya State, Nong Poh env., FKCP. Figure 38. S. lhasa, female, topotype, China, Tibet, Lhasa, 3700 m a. s. l., FKCP. Figure 39. S. lindbergi, male, holotype of S. kraepelini. Figure 40. S. margerisonae, male, holotype. Figure 41. S. oligotrichus, male (508), Laos, Xieng Khouang Province, Tham Xang Caves, Phone Savan, near Muang Khoun, FKCP. Figure 42. S.pakistanus, male, holotype.Figure 43. S. petersii, male, India, Himachal PradeshState, Molta,FKCP.Figure 44. S. pseudomontanus, male, holotype. Figure 45. S. solidus, male, holotype. Figure 46. S. tibetanus, male, topotype of S. pococki, China, Tibet, Gyaca district, 29°08'N 92°43'E, FKCP. Figure 47. S. tryznai, female, holotype. Figure 48. S. vonwicki, female, holotype. Figure 49. S. wrzecionkoi, female, holotype. Figure 50. S. yagmuri, male, holotype. Figure 51. S. zubairahmedi, male, holotype. Figure 52. S. zubairi, male, holotype.

Figures 53–76: Scorpiops spp., telson lateral. Figures 53–68: Species previously placed in genus Euscorpiops. Figure 53. S. asthenurus, female, India, Meghalaya State, Nong Poh env., FKCP. Figure 54. S. beccaloniae, male, holotype. Figure 55. S. kaftani, female, holotype. Figure 56. S. kamengensis, male, India, Arunachal Pradesh State, Hunli vicinity, 28°19'32"N 95°57'31"E, FKCP. Figure 57. S. kubani, male, paratype. Figure 58–59. S. longimanus, male (59), India, Assam State, 50 km E of Tezpur, 1800 m a. s. l., FKCP and female (59), India, Maghalaya State, West Garo-Hills, FKCP. Figures 60–61. S. montanus, male (60), and female (61), India, Uttarakhand State, Dhika forest 29.65°N 78.94°E, FKCP. Figure 62. S. neradi, male, holotype. Figure 63. S. novaki, male, holotype. Figure 64. S. orioni, male, holotype. Figure 65. S. problematicus, male (1265) topotype, Thailand, Chiang Mai Province, Doi Poi mt., FKCP. Figure 66. S. sejnai, male, holotype. Figure 67. S. shidian, male, topotype, China, Yunnan Province, Shidian district, Jiucheng town, 24.43°N 99.09°E, FKCP. Figure 68. S. thaomischi, male, holotype. Figures 69–72: Species previously placed in genus Alloscorpiops. Figures 69–70. S. anthracinus, female (69) and male (70), Thailand, Phetchaburi Province, Kaeng Krachan, 450 m a. s. l., FKCP. Figure 71. S. citadelle, male (1632) topotype. Figure 72. S. wongpromi, male, holotype. Figure 73. S. grandjeani (previously placed in genus Dasyscorpiops), female (1823), Thailand, Kapang, 8.004608°N 99.744415°E, FKCP. Figures 74–75: S. satarensis (previously placed in genus Neoscorpiops), female (74), India, Maharashtra State, Panrel env., FKCP and female (75), topotype, India, Maharashtra State, Mahabaleshwar, FKCP. Figure 76. Parascorpiops montanus, female, Malaysia, Borneo, Sarawak, Matang, ca. 20 km W of Kuching, FKCP.

Figures 77–81: Variation in positions of trichobothria db and eb on the fixed finger of the pedipalp chela of scorpiopids. Figure 77. Bivariate scatter plot of normalized distal positions of db vs. eb referred to trichobothrium Et5 on the distal chela manus (a proxy landmark for base of fixed finger) for males (triangles, 57 species) and females (circles, 57 species). Biometrics and symbol color indicating trichobothrial generic assignments according to old trichobothrial diagnoses, in upper right page inset. Trichobothrium db is level with eb along the black diagonal line, distal to it above the line, and proximal to it below the line. Figures 78–79. Relation of distal positioning of db vs. eb to manus elongation. Scatter plots of distal position of db relative to eb vs. L/D ratio of manus for 58 male (78) and 54 female (79) adult scorpiopids. Abscissa scale is logarithmic. Gray lines are least squares regressions, showing correlation is weak for males and strong for females. Figures 80–81. Relation of distal positioning of db vs. eb to positions of trichobothria Dt and Eb 3 on manus. Scatter plots of distal position of db relative to eb vs. normalized position of Dt (80) and Eb 3 (81) for males (triangles, 56 and 57 species) and females (circles, 54 and 52 species). Least squares regression lines are shown for each sex (males, dark gray; females, light gray). Arrows in Figs. 77, 79–81 indicate “Alloscorpiops” citadelle with more proximal db and Eb , separated from other “Alloscorpiops” with distal db. R=Pearson´s correlation coefficient.