Scorpiopidae Kraepelin , 1905

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 : 31

Family Scorpiopidae Kraepelin, 1905

( Figures 1–81 , 90–815, Tables 1 –9)

Scorpiopsinae Kraepelin, 1905: 341 (incorrect original spelling); Birula, 1917: 58; Werner, 1934: 282; Kästner, 1941: 237; Millot & Vachon, 1949: 428; Stahnke, 1974: 112, 116; Tikader & Bastawade, 1983: 380; Sissom, 1990: 114; Nenilin & Fet, 1992: 6.

Scorpiopsidae Stockwell, 1989 ; Stockwell, 1992: 411; Kovařík, 1998: 141; Lourenço, 1998: 245.

Scorpiopidae Fet, 2000: 487 (corrected spelling); Kovařík, 2000: 154; Prendini & Wheeler, 2005: 482, tab. 10; Kamenz & Prendini, 2008: 6, 10, 25, 44; Volschenk et al., 2008: 655, 665, 675, tab. 1–2; Prendini, 2011: 117; Loria & Prendini, 2014: 5, 20, 22, tab. 4, S1-10, S2-8; Beron, 2015: 11, 16; Lourenço, 2015: 9; Santibáñez-López et al., 2016: 2, tab. 1; Monod et al., 2017: 2–3, 10, 13, 15, 18, 38; Monod et. al., 2019: 110, 183.

Scorpiopini Soleglad & Sissom, 2001: 89, 93, 96; Soleglad & Fet, 2003b: 88, 106, 120, tab. 9; Fet & Soleglad, 2005: 12.

Scorpiopinae Soleglad & Fet, 2003a: 25 (in part); Coddington et al., 2004: 309 (in part).

DIAGNOSIS. Total length 24–75 mm. Carapace anterior margin with deep median notch. Inner accessory denticles present on pedipalp chelal fingers. Median eyes and median ocular tubercle present. Lateral eyes with 2–3 larger ocelli, one small ocellus may also be present. Cheliceral movable finger with dorsal distal denticle slightly shorter than ventral distal denticle, dorsal margin with two large subdistal denticles, ventral margin with 4–7 smaller denticles or crenulations. Ventral aspect of cheliceral fixed finger smooth without denticles. Sternum pentagonal. Pectines short, with 4–14 teeth, fulcra present or absent. Hemispermatophore lamelliform, distal lamina long, slender, terminally curved and tapered, capsule conforming to 2-folds bauplan, truncal flexure absent, terminal membrane of sperm duct spiculate, trunk with mid-axial rib. Ovariuterus 6-celled, oocyte development apoikogenic. Pedipalp chela more or less compressed, flattened. Pedipalp fingers with median denticles aligned in linear row (or rows), outer denticles usually present and displaced externally, inner accessory denticles present. Trichobothrial pattern type C, with additive neobothriotaxy, trichobothrial counts: patella 16–75 external, 6–28 ventral, chela manus 4–16 ventral. External surface of chela with Eb 3 level with or distal to Db. Legs with two pedal spurs, tibial spurs absent. Tarsomere II with row of 3–11 stout spinules on ventral surface. Metasoma with paired ventrosubmedian carinae. Telson without subaculear tubercle, often with annular construction at base of aculeus .

Type genus. Scorpiops Peters, 1861 .

SUBORDINATE TAXA. Scorpiops Peters, 1861 and Parascorpiops Banks, 1928 .Validity of the monophyletic genus Parascorpiops was not tested, either here or in Štáhlavský et al. (in press). Banks (1928) defined the genus Parascorpiops with type species P. montanus , by the possession of two pairs of lateral eyes (vs. three pairs in Scorpiops ). Francke (1976) questioned whether lateral eye counts were a reliable generic character, and several species of the genus Scorpiops are now known to have a third pair of lateral eyes that are greatly reduced, giving the impression of only 2 pairs present. We found only one unique character for the genus Parascorpiops : dentition of the pedipalp movable finger composed of irregular denticles in three or four rows ( Fig. 104 ). All species of Scorpiops have fingers with median denticles aligned in a regular linear row, or two rows ( Figs. 105–121 ).

REMARKS ON KARYOTYPES. The karyotypes have already been described in 19 scorpiopid species ( Kovařík et al., 2013a, 2015a, 2015c; Šťáhlavský et al., in press). In our present study we supplemented previous information with the karyograms in 16 of them ( Figs. 800–815 ). All chromosomes of scorpiopid species are monocentric, i. e. chromosomes have a defined centromere. This cytogenetic feature is typical for the whole parvorder Iurida (e.g. Shanahan, 1989; Schneider et al., 2009; Štundlová et al., 2019). Because all karyotyped scorpiopid species are now classified into the genus Scorpiops , the 2n of this genus ranges from 48 to 147 (Šťáhlavský et al., in press). Such a high variability of diploid numbers are known also in other genera from different families belonging to the Iurida such as Hadogenes (Hormuridae) : 2n=48–174, Urodacus (Urodacidae) : 2n=29–175, Alpiscorpius (Euscorpiidae) : 2n=46–90, and Heterometrus (Scorpionidae) : 2n=54–112 (see Schneider et al., 2020). High interspecific karyotype variability in these entire groups, including the genus Scorpiops , underscores the importance of cytogenetic markers as an important tool in species delimitation of scorpions with monocentric chromosomes ( Plíšková et al., 2016). Despite the high differences of karyotypes among scorpiopid species, their karyotypes usually possess a predominance of one-armed (acrocentric) chromosomes (see Šťáhlavský et al., in press). The high number of bi-armed chromosomes (almost one third) has been documented only in the species Scorpiops neradi (2n=48) with the lowest known number of chromosomes within scorpiopid species. This high number of bi-armed chromosomes in this species and also the high incidence (in 75% of species) of one trivalent during meiosis I comprising one extra-large bi-armed chromosome and two shorter acrocentrics, indicates Robertsonian translocations in the karyotype evolution of this group. Moreover, a high variability of nucleolus organizer regions (NORs) location within scorpiopid species was documented. Despite their stable number (always one pair), the NORs changed position from terminal to pericentromeric location and were identified on chromosomes of different lengths among analyzed species (from the longest chromosomes in Scorpiops citadelle and S. leptochirus to almost the shortest chromosomes in S. montanus ) (see Figs. 800–815 ). These differences in NOR locations and predominance of one-armed chromosomes also indicate an important role of inversions during the karyotype evolution. It is evident that the differentiation of karyotypes within scorpiopid scorpions is a dynamic process that includes complex chromosomal rearrangements that leads to the specific features of all recognized species. For this reason, we fully recommend implementing the characterization of karyotypes as one of the important characters in the description of new scorpiopid species in the future.

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.

Figures 104–121: Pedipalp movable finger dentition under UV light. Figure 104. Parascorpiops montanus, female, Malaysia, Borneo, Sarawak, Matang, ca. 20 km W of Kuching, FKCP. Figures 105–121: Scorpiops spp. Figure 105. S. satarensis, female, topotype, India, Maharashtra State, Mahabaleshwar, FKCP. Figure 106. S. grandjeani, female (1823), Thailand, Kapang, 8.004608°N 99.744415°E, FKCP. Figure 107. S. anthracinus, male (885), Thailand, Phetchaburi Province, Kaeng Krachan, 450 m a. s. l., FKCP. Figure 108. S. asthenurus, male, India, Meghalaya State, Nong Poh env., FKCP. Figure 109. S. sejnai, male, holotype. Figure 110. S. afghanus, female, holotype. Figure 111. S. braunwalderi, male, holotype. Figure 112. S. dastychi, male, paratype. Figure 113. S. hardwickii, female, India, Uttaranchal State, ca. 30 km N of Bageshwar, SE of Dhakuri vill., 2600-2800 m a. s. l., FKCP. Figure 114. S. irenae, female, holotype. Figure 115. S. jendeki, male, holotype. Figure 116. S. leptochirus, male (306), India, Meghalaya State, Nong Poh env., FKCP. Figure 117. S. lindbergi, female, Pakistan, northern Balochistan, Ziarat, FKCP. Figure 118. S. margerisonae, male, holotype. Figure 119. S. oligotrichus, male (508), Laos, Xieng Khouang Province, Tham Xang Caves, Phong Savan, near Muang Khoun, FKCP. Figure 120. S. petersii, male, India, Himachal Pradesh State, Molta, FKCP. Figure 121. S. zubairahmedi, male, holotype.

Figures 800–807: Male karyotypes of scorpiopid species based on postpachytene. Figure 800. Scorpiops anthracinus (2n=105, 51II+III). Figure 801. Scorpiops birulai sp. n. (2n=99, 96II+III). Figure 802. Scorpiops citadelle (2n=104, 52II). Figure 803. Scorpiops dii sp. n. (2n=109, 53II+III). Figure 804. Scorpiops kautti sp. n. (2n=87, 42II+III). Figure 805. Scorpiops krabiensis sp. n. (2n=81, 39II+III). Figure 806. Scorpiops leptochirus (2n=96, 48II). Figure 807. Scorpiops montanus (2n=70, 35II). Insets show the location of 18S rDNA (red signal) on the same chromosome as in the karyogram (ppach) and additionally on chromosomes during pachytene (pach) or metaphase II (met II). Abbreviation of postpachytene configuration: II – bivalent, III – trivalent. Scale bar: 10 μm.

Figures 808–815: Male karyotypes of scorpiopid species based on postpachytene. Figure 808. Scorpiops oligotrichus (2n=105, 51II+III). Figure 809. Scorpiops pakseensis sp. n. (2n=87, 42II+III). Figure 810. Scorpiops phatoensis sp. n. (2n=105, 51II+III). Figure 811. Scorpiops problematicus (2n=97, 47II+III). Figure 812. Scorpiops schumacheri sp. n. (2n=63, 60II+III). Figure 813. Scorpiops sherwoodae sp. n. (2n=104, 52II). Figure 814. Scorpiops thailandus sp. n. (2n=147, 72II+III). Figure 815. Scorpiops thaomischi (2n=94, 47II). Insets show the location of 18S rDNA (red signal) on the same chromosome as in the karyogram (ppach) and additionally on chromosomes during pachytene (pach) or metaphase II (met II). Abbreviation of postpachytene configuration: II – bivalent, III – trivalent. Scale bar: 10 μm.