Buthus Leach, 1815

Kovařík, František, Šťáhlavský, František & Elmi, Hassan Sh Abdirahman, 2020, Scorpions of the Horn of Africa (Arachnida Scorpiones) Part XXIII Buthus (Buthidae), with description of two new species, Euscorpius 307, pp. 1-32 : 1-3

publication ID	https://doi.org/10.5281/zenodo.4648379
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DOI	https://doi.org/10.5281/zenodo.4770176
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scientific name	Buthus Leach, 1815
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Buthus Leach, 1815 View in CoL

( Figures 1–211 View Figures 1–4 View Figures 5–6 View Figures 7–15 View Figures 16–24 View Figures 25–28 View Figures 29–36 View Figures 37–53 View Figures 54–61 View Figures 62–65 View Figures 66–73 View Figures 74–89 View Figures 90–99 View Figures 100–101 View Figures 102–105 View Figures 106–113 View Figures 114–131 View Figures 132–139 View Figures 140–143 View Figures 144–151 View Figures 152–167 View Figures 168–175 View Figures 176–190 View Figures 191–194 View Figures 195–209 View Figures 210–211 , Tables 1–3 View Table 1 View Table 2 View Table 3 )

Buthus Leach, 1815: 391 View in CoL ; Fet & Lowe, 2000: 91–97 (complete references list until 1998); Sousa et al., 2017: 15–84 View Cited Treatment , figs. 1–16.

TYPE SPECIES. Scorpio occitanus Amoreux, 1789

DIAGNOSIS. Total length 25–90 mm. Carapace with distinct carinae joined to form a lyre-shaped configuration, in lateral view with entire dorsal surface horizontal or nearly so. Five pairs of lateral eyes and eyespot present. Pectines with fulcra. Pectine teeth number ca 20–40. Mesosoma tergites I–VI with three carina which do not project beyond posterior margin as distinct spiniform processes. Telson without subaculear tubercle. Chelicera with typical buthid dentition, fixed finger with two ventral denticles. Orthobothriotaxic type B, dorsal trichobothria of pedipalp femur arranged in beta -configuration. Patellar trichobothrium d 2 located externally to dorsomedian carina. Trichobothrium eb located on fixed finger of chela. Dentate margin of pedipalp chela movable finger with distinct granules divided into 9–14 rows, 3 terminal granules and one basal terminal granule. Tibial spurs present on third and fourth pairs of legs.

REMARKS ON KARYOTYPES ( Figs. 195–209 View Figures 195–209 ). We analyzed male karyotypes of four Buthus species from the Horn of Africa ( Table 3 View Table 3 ). The cytogenetic characteristics of all species correspond to the typical features of the family Buthidae such as holocentric organization, achiasmatic meiosis in males, and lower number of chromosomes (e. g. Mattos et al., 2013). All examined specimens of Buthus awashensis , B. pococki sp. n. and B. zeylensis possess 2n=22 ( Figs. 195, 201, 204, 205, 208 View Figures 195–209 ). We found only 21 chromosomes in karyotype of analysed Buthus berberensis ( Fig. 198 View Figures 195–209 ). The reduction of the diploid number is probably consequence of heterozygous fusion of chromosomes. This type of chromosomal rearrangement forms conspicuous trivalent during postpachytene ( Fig. 199 View Figures 195–209 ). Moreover, fusion of two chromosomes form one extra large chromosome in the karyotype of this species ( Fig. 200 View Figures 195–209 ). Except Buthus awashensis , the karyotypes of the all analysed species are typical with one pair of chromosomes with distinctive length whereas the subsequent chromosomes are shorter and gradually decrease in length ( Figs. 200, 203, 206, 209 View Figures 195–209 ). Similar one longer pair of chromosomes is known also in karyotypes of all karyotyped Androctonus species ( SadÍlek et al., 2015). This similarity of karyotypes may reflect phylogenetic relationships that was already proposed in previous phylogenetic analysis ( Fet et al., 2003). We observed multivalent association (ten chromosomes of different length) in Buthus awashensis during postpachytene ( Figs. 196–197 View Figures 195–209 ). This chain of chromosomes is consequence of multiple reciprocal translocations. Although this type of chromosome rearrangement does not affect chromosome numbers, it may however considerably change the size of the chromosomes (e. g. KovařÍk et al., 2015). It is probably the reason why this species has not one extra large pair of chromosomes, the typical feature of the remaining species with only bivalents ( Figs. 202, 205, 208 View Figures 195–209 ). Despite of the mentioned small differences among analysed species, the karyotypes of species from the Horn of Africa fully correspond to those of Buthus occitanus from France ( Guénin, 1961), and the karyotypes seem to be very conservative within the genus Buthus .

References

FET, V., B. GANTENBEIN, A. V. GROMOV, G. LOWE & W. R. LOURENCO. 2003. The first molecular phylogeny of Buthidae (Scorpiones). Euscorpius, 4: 1 - 10.

FET, V. & G. LOWE. 2000. Family Buthidae C. L. Koch, 1837. Pp. 54 - 286 in Fet, V., W. D. Sissom, G. Lowe & M. E. Braunwalder. Catalog of the Scorpions of the World (1758 - 1998). New York: The New York Entomological Society, 689 pp.

GUENIN, H. A. 1961. Contribution a la connaissance cytologique des scorpions: les chromosomes de Buthus occitanus Amor. (I). Vie et Milieu, 12: 89 - 96.

KOVARIK, F., G. LOWE, M. SEITER, J. PLISKOVA & F. STAHLAVSKY. 2015. Scorpions of Ethiopia (Arachnida: Scorpiones). Part II. Genus Babycurus Karsch, 1886 (Buthidae), with description of two new species. Euscorpius, 196: 1 - 31.

LEACH, W. E. 1815. A tabular view of the external characters of four classes of animals, which Linne arranged under Insecta; with the distribution of the genera composing three of these classes into orders, etc. and descriptions of several new genera and species. Transactions of the Linnean Society of London, 11: 306 - 400.

MATTOS, V. F., D. M. CELLA, L. S. CARVALHO, D. M. CANDIDO & M. C. SCHNEIDER. 2013. High chromosome variability and the presence of multivalent associations in buthid scorpions. Chromosome Research, 21: 121 - 136.

SADILEK, D., P. NGUYEN, H. KOC, F. KOVARIK, E. A. YAGMUR & F. STAHLAVSKY. 2015. Molecular cytogenetics of Androctonus scorpions: an oasis of calm in the turbulent karyotype evolution of the diverse family Buthidae. Biological Journal of the Linnean Society, 115: 69 - 76.

SOUSA P., M. A. ARNEDO & D. J. HARRIS. 2017. Updated catalogue and taxonomic notes on the Old-World scorpion genus Buthus Leach, 1815 (Scorpiones, Buthidae). ZooKeys, 686: 15 - 84.

Figures

Figures 1–4: Buthus awashensis. Figures 2–3. Male paratopotype in dorsal (1) and ventral (2) views. Figures 3–4. Female paratopotype in dorsal (3) and ventral (4) views. Scale bars: 10 mm.

Figures 5–6. Buthus berberensis, male holotype in dorsal (5) and ventral (6) views.

Figures 7–15. Buthus berberensis, male holotype, carapace and tergites I–IV (7), coxosternal area and sternites (8), left legs II–IV, retrolateral aspect (9–11 respectively), metasoma and telson lateral (12), and metasoma and telson lateral (13), dorsal (14), and ventral (15) views. Scale bar: 10 mm (13–15).

Figures 16–24: Buthus berberensis. Figures 16–23. Male holotype, pedipalp segments. Chela dorsal (16), external (17) and ventral (18) views. Patella dorsal (19), external (20) and ventral (21) views. Trochanter and femur dorsal (22) and ventral (23) views. Trichobothrial pattern is indicated by white circles (16–20, 22). Figure 24. Male from locality 19SO, in vivo habitus.

Figures 25–28: Buthus berberensis from locality 19SO. Figures 25–26. Male in dorsal (25) and ventral (26) views. Figures 27–28. Female in dorsal (27) and ventral (28) views. Scale bar: 10 mm.

Figures 29–36: Buthus berberensis from locality 19SO. Figures 29, 31–33. Male, telson lateral (29), and metasoma and telson lateral (31), dorsal (32), and ventral (33) views. Figures 30, 34–36. Female, telson lateral (30), and metasoma and telson lateral (34), dorsal (35), and ventral (36) views. Scale bars: 10 mm (31–33, 34–36).

Figures 37–53: Buthus berberensis from locality 19SO, pedipalp segments. Figures 37–46. Male. Chela dorsal (37), external (38) and ventral (39) views. Patella dorsal (40), external (41) and ventral (42) views.Trochanter and femur dorsal (43) and ventral (44) views. Movable (45) and fixed (46) finger dentition.Trichobothrial pattern is indicated by white circles (37–41, 43). Figures 47–53. Female. Chela dorsal (47), external (48) and ventral (49) views. Patella dorsal (50), external (51) and ventral (52) views. Trochanter and femur dorsal (53).

Figures 54–61: Buthus berberensis from locality 19SO. Figures 54, 56, 58–61. Male, carapace and tergites I–III (54), coxosternal area and sternites (56), left legs I–IV, retrolateral aspect (58–61 respectively). Figures 55, 57. Female, carapace and tergites I–IV (55), coxosternal area and sternites III–V (57).

Figures 62–65: Buthus pococki sp. n. Figures 62–63. Male holotype in dorsal (62) and ventral (63) views. Figures 64–65. Female paratopotype in dorsal (64) and ventral (65) views. Scale bar: 10 mm.

Figures 66–73: Buthus pococki sp. n. Figures 66, 68–70. Male holotype, telson lateral (66), and metasoma and telson lateral (68), dorsal (69), and ventral (70) views. Figures 67, 71–73. Female paratopotype, telson lateral (67), and metasoma and telson lateral (71), dorsal (72), and ventral (73) views. Scale bars: 10 mm (68–70, 71–73).

Figures 74–89: Buthus pococki sp. n., pedipalp segments. Figures 74–83. Male holotype. Chela dorsal (74), external (75) and ventral (76) views. Patella dorsal (77), external (78) and ventral (79) views. Trochanter and femur dorsal (80) and ventral (81) views. Movable (82) and fixed (83) finger dentition. Trichobothrial pattern is indicated by white circles (74–78, 80). Figures 84–89. Female paratopotype. Chela dorsal (84), external (85) and ventral (86) views. Patella dorsal (87), external (88) and ventral (89) views.

Figures 90–99: Buthus pococki sp. n. Figures 90, 92, 94–97. Male holotype, carapace and tergites I–III (90), coxosternal area and sternites (92), left legs I–IV, retrolateral aspect (94–97 respectively). Figures 91, 93, 98–99. Female paratopotype, carapace and tergites I–III (91), coxosternal area and sternite III (93), and right chelicera in dorsal (98) and ventral (99) views.

Figures 100–101: Localities. Figure 100. Buthus pococki sp. n., type locality (19SC). Figure 101. B. zeylensis, locality 19SH.

Figures 102–105: Buthus somalilandus sp. n. Figures 102–103. Male in dorsal (102) and ventral (103) views. Figures 104–105. Female in dorsal (104) and ventral (105) views. Scale bar: 10 mm.

Figures 106–113: Buthus somalilandus sp. n. Figures 106, 108–110. Male, telson lateral (106), and metasoma and telson lateral (108), dorsal (109), and ventral (111) views. Figures 107, 111–113. Female, telson lateral (107), and metasoma and telson lateral (111), dorsal (112), and ventral (113) views. Scale bar: 10 mm (108–113).

Figures 114–131: Buthus somalilandus sp. n. Figures 114–123. Male. Chela dorsal (114), external (115) and ventral (116) views. Patella dorsal (117), external (118) and ventral (119) views. Trochanter and femur dorsal (120) and femur ventral (121) views. Movable (122) and fixed (123) finger dentition. Trichobothrial pattern is indicated by white circles (114–118, 120). Figures 124–129. Female. Chela dorsal (124), external (125) and ventral (126) views. Patella dorsal (127), external (128) and ventral (129) views. Trochanter and femur dorsal (130) and ventral (131) views.

Figures 132–139: Buthus somalilandus sp. n. Figures 132, 136–139. Male holotype, carapace and tergites I–IV (132), coxosternal area and sternites (134), left legs I–IV, retrolateral aspect (136–139 respectively). Figures 133–134. Female paratopotype, carapace and tergites I–III (133), coxosternal area and sternites III–VI (134).

Figures 140–143: Buthus zeylensis from locality 19SH. Figures 140–141. Male in dorsal (140) and ventral (141) views. Figures 142–143. Female in dorsal (142) and ventral (143) views. Scale bar: 10 mm.

Figures 144–151: Buthus zeylensis from locality 19SH. Figures 144, 146–148. Male, telson lateral (144), and metasoma and telson lateral (146), dorsal (147), and ventral (148) views. Figures 145, 149–151. Female, telson lateral (145), and metasoma and telson lateral (149), dorsal (150), and ventral (151) views. Scale bar: 10 mm (146–151).

Figures 152–167: Buthus zeylensis from locality 19SH, pedipalp segments. Figures 152–161. Male. Chela dorsal (152), external (153) and ventral (154) views. Patella dorsal (155), external (156) and ventral (157) views. Trochanter and femur dorsal (158) and ventral (159) views. Movable (160) and fixed (161) finger dentition. Trichobothrial pattern is indicated by white circles (152–156, 158). Figures 162–167. Female. Chela dorsal (162), external (163) and ventral (164) views. Patella dorsal (165), external (166) and ventral (167) views.

Figures 168–175: Buthus zeylensis from locality 19SH. Figures 168, 170, 172–175. Male, carapace and tergites I–IV (168), coxosternal area and sternites (170), left legs I–IV, retrolateral aspect (172–175 respectively). Figures 169, 171. Female, carapace and tergites I–III (169), coxosternal area and sternites III–VI (171).

Figures 176–190: Comparison of pdipalp chela (176, 178, 180, 182, 184), pedipalp patella (177, 179, 181, 183, 185), and telson (186–190) of males of Buthus. Figures 176–177, 186. B. awashensis, holotype. Figures 178–179, 187. B. berberensis from locality 19SO. Figures 180–181, 188. B. pococki sp. n., holotype. Figures 182–183, 189. B. somalilandus sp. n., holotype. Figures 184–185, 190. B. zeylensis from locality 19SH.

Figures 191–194: Buthus, in vivo habitus. Figure 191. B. awashensis, female paratype. Figure 192. B. pococki sp. n., male paratype. Figure 193. B. somalilandus sp. n., female, paratype. Figure 194. B. zeylensis from locality 19SH.

Figures 195–209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195–197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198–200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201–203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204–206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207–209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195–196, 198–199, 201–202, 204–205, 207–208).