Review of the genus Babycurus Karsch, 1886 (Arachnida, Scorpiones, Buthidae), with descriptions of Barbaracurus gen. n. and two new species from Oman and Yemen
Abstract
Kovařík, František, Lowe, Graeme, Šťáhlavský, František (2018): Review of the genus Babycurus Karsch, 1886 (Arachnida, Scorpiones, Buthidae), with descriptions of Barbaracurus gen. n. and two new species from Oman and Yemen. Euscorpius 267: 1-41, DOI: 10.5281/zenodo.6544157
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Review of the genus Babycurus Karsch, 1886 (Arachnida, Scorpiones, Buthidae), with descriptions of Barbaracurus gen. n. and two new species from Oman and Yemen František Kovařík, Graeme Lowe & František Šťáhlavský August 2018 – No. 267
Euscorpius Occasional Publications in Scorpiology EDITOR: Victor Fet, Marshall University, ‘[email protected]’ ASSOCIATE EDITOR: Michael E. Soleglad, ‘[email protected]’ Euscorpius is the first research publication completely devoted to scorpions (Arachnida: Scorpiones). Euscorpius takes advantage of the rapidly evolving medium of quick online publication, at the same time maintaining high research standards for the burgeoning field of scorpion science (scorpiology). Euscorpius is an expedient and viable medium for the publication of serious papers in scorpiology, including (but not limited to): systematics, evolution, ecology, biogeography, and general biology of scorpions. Review papers, descriptions of new taxa, faunistic surveys, lists of museum collections, and book reviews are welcome. Derivatio Nominis The name Euscorpius Thorell, 1876 refers to the most common genus of scorpions in the Mediterranean region and southern Europe (family Euscorpiidae). Euscorpius is located at: http://www.science.marshall.edu/fet/Euscorpius (Marshall University, Huntington, West Virginia 25755-2510, USA) ICZN COMPLIANCE OF ELECTRONIC PUBLICATIONS: Electronic (“e-only”) publications are fully compliant with ICZN (International Code of Zoological Nomenclature) (i.e. for the purposes of new names and new nomenclatural acts) when properly archived and registered. All Euscorpius issues starting from No. 156 (2013) are archived in two electronic archives: •Biotaxa, http://biotaxa.org/Euscorpius (ICZN-approved and ZooBank-enabled) •Marshall Digital Scholar, http://mds.marshall.edu/euscorpius/. (This website also archives all Euscorpius issues previously published on CD-ROMs.) Between 2000 and 2013, ICZN did not accept online texts as "published work" (Article 9.8). At this time, Euscorpius was produced in two identical versions: online (ISSN 1536-9307) and CD-ROM (ISSN 1536-9293) (laser disk) in archive-quality, read-only format. Both versions had the identical date of publication, as well as identical page and figure numbers. Only copies distributed on a CD-ROM from Euscorpius in 2001-2012 represent published work in compliance with the ICZN, i.e. for the purposes of new names and new nomenclatural acts. In September 2012, ICZN Article 8. What constitutes published work, has been amended and allowed for electronic publications, disallowing publication on optical discs. From January 2013, Euscorpius discontinued CD-ROM production; only online electronic version (ISSN 1536-9307) is published. For further details on the new ICZN amendment, see http://www.pensoft.net/journals/zookeys/article/3944/. Publication date: 15 August 2018 http://zoobank.org/urn:lsid:zoobank.org:pub:6580A0C7-E18D-4B06-A158-522F089CEAF5
Euscorpius — Occasional Publications in Scorpiology. 2018, No. 267 Review of the genus Babycurus Karsch, 1886 (Arachnida, Scorpiones, Buthidae), with descriptions of Barbaracurus gen. n. and two new species from Oman and Yemen František Kovařík1, 3, Graeme Lowe2 & František Šťáhlavský3 1 P. O. Box 27, CZ-145 01 Praha 45, Czech Republic; www. scorpio.cz 2 Monell Chemical Senses Center, 3500 Market St., Philadelphia, PA 19104-3308, USA 3 Department of Zoology, Charles University, Viničná 7, CZ-128 44 Praha 2, Czech Republic http://zoobank.org/urn:lsid:zoobank.org:pub:6580A0C7-E18D-4B06-A158-522F089CEAF5 Summary The genus Babycurus Karsch, 1886 sensu lato is split into two genera, a strictly African genus Babycurus, and the new genus Barbaracurus gen. n., which mainly includes species from the Horn of Africa and Arabian Peninsula. Two new species Barbaracurus winklerorum sp. n. from Oman and B. yemenensis sp. n. from Yemen are described, compared with other species, and fully illustrated with color photos of morphology, habitus, live specimens and collection localities. Males of Barbaracurus somalicus (Hirst, 1907) comb. n. and Barbaracurus zambonellii (Borelli, 1902) comb. n. are recorded for the first time and fully illustrated. Babycurus ornatus Werner, 1936 from Mozambique is shown to be a junior synonym of Lychas burdoi (Simon, 1882), a species from the same area. Babycurus brignolii Lourenço et Rossi, 2017 is designated to be a nomen dubium. Hemispermatophores are described and illustrated to show their differences between the species and genera. Analyses of karyotypes reveal a similar degree of interspecific variability of diploid chromosomal numbers within the genera Babycurus (2n=16–30) and Barbaracurus gen. n. (2n=22–36). Introduction In the years 2011–2017, one of the authors (F.K.) had the opportunity to participate in expeditions to the Horn of Africa, study scorpions at 113 localities, and publish several articles (e. g. Kovařík, 2011; Kovařík et al., 2015, 2018). Scorpions of the genus Babycurus Karsch, 1886 sensu lato are relatively rare in the Horn of Africa. Only six of 113 sampled localities yielded a total of four species including both sexes of the very rare species, Barbaracurus somalicus (Hirst, 1907) comb. n. (Somaliland) and Barbaracurus zambonellii (Borelli, 1902) comb. n. (Eritrea), that were previously known only from female types. Here we present photographs of the males for the first time, characterize their sexual dimorphism, and correct the erroneous assumption in the literature that the female holotype of B. zambonellii is a male. We also studied specimens collected by Alexander Winkler and his wife Birgit Winkler in Oman, and by Petr Kabátek and David Král in Yemen. The chance to study such a rich series of Babycurus (sensu lato) materials enabled us to gain a much better understanding of the genus, and motivated us to split it into two genera: Babycurus which includes African species and Barbaracurus gen. n. which mainly includes species from the Horn of Africa and Arabian Peninsula. According to our diagnosis, we also transfer to this genus Barbaracurus prudenti (Lourenço, 2013) comb. n. from Cameroon and B. ugartei (Kovařík, 2000) comb. n. from Nigeria, species that we did not study in detail. Here we focus exclusively on species of Barbaracurus gen. n. from the Horn of Africa and Arabian Peninsula. Methods, Material & Abbreviations Nomenclature and measurements follow Vachon (1963), Stahnke (1971), Sissom (1990), Kovařík (2009), and Kovařík & Ojanguren Affilastro (2013), except for trichobothriotaxy (Vachon, 1974, 1975), and morphology of sternum (Soleglad & Fet, 2003), and hemispermatophore (Kovařík et al., 2018). The numbers of rows of granules on the pedipalp chela movable finger was specified differently by various authors, depending on whether they included the short apical row, and whether the most proximal row was counted as one row, or as two rows divided by
Euscorpius — 2018, No. 267 2 external and internal accessory granules. In this paper, we counted all rows except the short apical row, and we counted as a single row (i.e. formed by ‘fusion’) the most proximal row which in the genus Babycurus sensu lato lacks an internal accessory granule (Figs. 1–23). Hence, the number of rows reported here can differ from counts given in previous papers (e. g. Kovařík, 2000, 2015; Lourenço, 2013; Prendini, 2004). Karyotype analyses were based on chromosome preparations prepared by the spreading technique which is frequently used in scorpions (e. g. Kovařík et al., 2009; Plíšková et al., 2016). The chromosomes were stained by 5% Giemsa solution in Sörensen phosphate buffer for 20 min. Five spermatocyte nuclei were measured using the software Image J 1.45r (http:// rsbweb.nih.gov/ij) with the plugin Levan (Sakamoto & Zacaro, 2009). The relative length of the chromosomes was calculated for the diploid set. Specimens studied herein are preserved in 80% ethanol. Depositories: BMNH (The Natural History Museum, London, United Kingdom); FKCP (František Kovařík, private collection, Prague, Czech Republic); GLPC (Graeme Lowe, private collection, Philadelphia, USA); MCSN (Museo Civico de Storia Naturale “Giacomo Doria”, Genoa, Italy); MZUT (Museo Regionale di Scienze Naturali of Turin, Italy); NHMB (Naturhistorisches Museum Basel, Switzerland); ZSMC (Bavarian State Collection of Zoology, Munich, Germany). Systematics Family Buthidae C. L. Koch, 1837 Babycurus Karsch, 1886 (Figures 11–23, 29–31, 37–39, 255–260, 265) Babycurus Karsch, 1886: 77–79, figs. 1–2; Kraepelin, 1895: 88–89 (in part); Pocock, 1896: 427–431; Pocock, 1899: 835; Kraepelin, 1899: 61–64; Kraepelin, 1913: 179–183 (in part); Sissom, 1990: 101; Fet & Lowe, 2000: 76–80 (in part); Kovařík, 2000: 236–263, figs. 1–9, 11–12, 14–20, 23–25, 27–37, 39–40, tables 1–3 (in part); Prendini, 2004: 238– 250, figs. 1–10; Kovařík, 2009: 30 (in part); Loria & Prendini, 2014: 19, 25; Loria & Prendini, 2018: 184. Buthus (Rhoptrurus) (in part): Pocock, 1890: 122. Rhoptrurus: Kraepelin, 1891: 238–241 (in part); Kraepelin, 1898: 3 (in part) (syn. by Kraepelin, 1899: 61). TYPE SPECIES. Babycurus buettneri Karsch, 1886. DIAGNOSIS. Medium to large buthids, adults 30–100 mm. Carapace granular, lacking distinct carinae, flat, subrectangular with concave anterior margin. Median eyes on low ocular tubercle in anterior half of carapace; usually with 4, or sometimes 5 pairs of lateral eyes (3 major ocelli, 1–2 minor ocelli). Anterior, central and posterior median furrows distinct, connected by median groove running over ocular tubercle. Sternum type 1, triangular in shape. Tergites I–VI granular, with single median carina which may be obsolete on I–II, tergite VII with 5 carinae. Metasoma elongate, segment I with 10 carinae, II–IV with 8 carinae, lacking lateral median carina. Metasoma V convex, sometimes dilated, carinae present or obsolete. Telson ellipsoidal or pyriform in shape, with distinct subaculear tooth. Pectines with fulcra. Hemispermatophore capsule with 2-lobed sperm hemiduct, basal lobe atrophied to a short, weak carina. Chelicerae with typical buthid dentition: movable finger dorsal margin with large subdistal and medial denticles and two smaller basal denticles, ventral margin with two large denticles, dorsal distal tine slightly shorter than ventral counterpart; fixed finger with subdistal denticle, and median and basal denticles formed as a bicusp, ventral surface armed with two small denticles. Pedipalps orthobothriotaxic, type Aβ, femur trichobothrium d2 internal, patella d3 external to dorsomedian carina, chela db in distal half of fixed finger. Chela manus smooth, with carinae reduced or obsolete, dentate margins of chela movable finger armed with 6–9 imbricated rows of denticulate granules, successive rows overlapping by at least 2 granules, each row terminated proximally by an enlarged granule, in most cases flanked by two enlarged external accessory granules at oblique angle (single external accessory granule only in B. gigas), and single internal accessory granule displaced distally. Most proximal granule row with one or (rarely) two isolated external accessory granules midway along its length, no internal accessory granules. Pedipalp chelae sexually dimorphic, males with manus dilated and fingers proximally undulate on dentate margins, denticles of proximal granule rows bicuspid. Tibial spurs absent on leg III, present on leg IV, tibia and tarsus III– IV without bristle combs, ventral surfaces of tarsi equipped with two rows of setae, ungues stout. SUBORDINATE TAXA. Babycurus ansorgei Hirst, 1911 (Angola, Congo), B. buettneri Karsch, 1886 (West Africa), B. centrurimorphus Karsch, 1886 (Congo, Tanzania, ?Mozambique, Rwanda), B. dunlopi Kovařík et al., 2015 (Ethiopia), B. gigas Kraepelin, 1896 (Tanzania), B. jacksoni (Pocock, 1890) (Kenya, Tanzania), B. kirki (Pocock, 1890) (West Africa), B. melanicus Kovařík, 2000 (Congo), B. multisubaculeatus Kovařík, 2000 (Somalia), B. pictus Pocock, 1896 (Kenya, Tanzania), B. solegladi Lourenço, 2005 (Sudan), B. taramassoi Borelli, 1919 (Somalia), and B. wituensis Kraepelin, 1913 (Kenya). CYTOGENETICS. The karyotype analyses of three species Babycurus jacksoni (2n=16) (Figs. 255, 256), B. buett-
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 3 Figures 1–23: Pedipalp movable finger, comparison of Barbaracurus gen. n. (1–10) and Babycurus (11–23) species. Figures 1–4: Barbaracurus gen. n., species with 6 non-imbricated almost linear rows of granules. Figure 1. B. ugartei comb. n., female holotype. Figure 2. B. sofomarensis comb. n., male paratype. Figure 3. B. somalicus comb. n., female from Somaliland, locality 17SR. Figure 4. B. subpunctatus comb. n., female from Ethiopia, locality 14EI. Figures 5–10: Barbaracurus gen. n., species with 7 non-imbricated almost linear rows of granules. Figure 5. B. exquisitus comb. n., male holotype. Figures 6–7, 10. B. zambonellii comb. n., female (6 and 10) and male (7) from Eritrea, locality 15EH. Figure 8. B. winklerorum sp. n., female paratype. Figure 9. B. yemenensis sp. n., female holotype. Figures 12–14: Babycurus, species with 6 imbricated rows of movable fingers. Figure 12. B. ansorgei, female, Congo, Bukama env., FKCP. Figure 13. B. centrurimorphus, female lectotype. Figure 14. B. pictus, female, Congo, Sampwe env., FKCP. Figures 11, 15–21: Babycurus, species with 7–8 imbricated rows of movable fingers. Figure 15. B. buettneri, female, MCSN. Figure 11, 16. B. dunlopi, male paratype. Figure 17. B. jacksoni, male, Tanzania, FKCP, No. 1189. Figure 18. B. kirki, female, Cote d’Ivoire, Comoe, FKCP. Figure 19. Babycurus melanicus, female holotype. Figure 20. B. taramassoi, female, Somalia, Belet Amin, MCSN. Figure 21. Babycurus wituensis, female lectotype. Figures 22–23: Babycurus, species with 9 imbricated rows of movable fingers. Figure 22. B. gigas, female lectotype. Figure 23. B. multisubaculeatus, female holotype. Numbers in red indicate number of granule rows in series of fingers on the left of each number (red arrows).
Euscorpius — 2018, No. 267 4 neri (2n=28) (Figs. 257, 258) and B. gigas (2n=30) (Figs. 259, 260) show considerable interspecific variability of the chromosome diploid numbers within this genus. The chromosomes have holocentric organization which is typical feature of the family Buthidae (Schneider et al., 2009a). Males display achiasmatic meisois.which is characteristic to the whole order Scorpiones (Schneider et al., 2009b). Taxonomic position of Babycurus ornatus Werner, 1936 Babycurus ornatus was described by Werner in 1936 from Mozambique and has been accepted as valid by other authors (Vachon, 1940: 179; Weidner, 1959: 98; Lamoral & Reynders, 1975: 498; Kovařík, 1998: 104; Fet & Lowe, 2000: 79; Kovařík, 2000: 252). The first author (F.K.) recently studied the types (Figs. 40– 49) and found that legs III and IV both bear tibial spurs (Figs. 46–47). This character is diagnostic for the genus Lychas C. L. Koch, 1845, whereas in genera Babycurus and Barbaracurus gen. n., leg III lacks tibial spurs (only present on leg IV) (e.g. Figs. 99–106). Moreover, it was found that the types of Babycurus ornatus and Lychas burdoi are precisely matched in the following key characters: trichobothrial pattern, pedipalp finger dentation (Figs. 44–45), pectinal tooth count and pectine lamellar structure, body and appendage proportions, setation, carination and sculpture of pedipalps, carapace, tergites, sternites, form of metasoma and telson (Figs. 40–49), as well as armature of chelicerae and pedipalp fingers. The inevitable conclusion is that Babycurus ornatus Werner, 1936 is a junior synonym of Lychas burdoi (Simon, 1882) syn. n. Taxonomic position of Babycurus brignolii Lourenço et Rossi, 2017 Babycurus brignolii was based on a juvenile, which Lourenço & Rossi (2017) in the original description, incorrectly declared as a male. It is evident from the sternopectinal area and pectines that it is juvenile female, and it displays typical juvenile morphology and color, as shown in fig. 9 of Lourenço & Rossi (2017: 6). In addition to the above observations, we note that the juvenile holotype of B. brignolii and juveniles of B. gigas Kraepelin, 1896, that we have studied, match closely in the following key described characters: trichobothrial pattern, pectinal tooth count and lamellar structure, proportions, setation, carination and sculpture of pedipalps, carapace, tergites, sternites, and metasoma, shape of the telson, as well as armature of pedipalp fingers. However, the conclusion that B. brignolii is a junior synonym of B. gigas is complicated by the listed type locality of B. brignolii (Central African Republic, Bamingui-Bangoran Province) which lies outside the known range of distribution of B. gigas (Tanzania). We therefore regard Babycurus brignolii Lourenço et Rossi, 2017 as a nomen dubium until either the true distribution of both species is revised, or an adult topotype specimen of B. brignolii is described. Barbaracurus gen. n. (Figures 1–10, 24–28, 32–36, 50–254, 261–265, Tables 1–2) http://zoobank.org/urn:lsid:zoobank.org:act:75EF2A 5A-6CF8-4F3B-94AB-3374C2902E6C Babycurus: Kraepelin, 1913: 179–183 (in part); Fet & Lowe, 2000: 76–80 (in part); Kovařík, 2000: 244– 245, 255–256, 260–262, figs. 10, 13, 21–22, 26, 38– 40, tables 1–3 (in part); Kovařík, 2009: 30 (in part). TYPE SPECIES. Babycurus sofomarensis Kovařík, Lowe, Seiter, Plíšková et Šťáhlavský, 2015. ETYMOLOGY. The genus-group name is a patronym honoring Dr. Barbara York Main for her lifelong contributions to arachnology, especially the systematics and natural history of Australian mygalomorph spiders, and also to many other topics in ecology, biogeography, evolution and conservation. DIAGNOSIS. Small to medium buthids, adults 22–47 mm. Carapace granular, lacking distinct carinae, flat, subrectangular with concave anterior margin. Median eyes on low ocular tubercle in anterior half of carapace; usually with 4, or sometimes 5 pairs of lateral eyes (3 major ocelli, 1–2 minor ocelli). Anterior, central and posterior median furrows distinct, connected by median groove running over ocular tubercle. Sternum type 1, triangular in shape. Tergites I–VI granular, with single median carina which may be obsolete on I–II, tergite VII with 5 carinae. Metasoma elongate, segment I with 10 carinae, II–IV with 8 carinae, lacking lateral median carina. Metasoma V convex, sometimes dilated, carinae present or obsolete. Telson ellipsoidal, pyriform or slightly bulbous, with distinct subaculear tooth. Pectines with fulcra. Hemispermatophore capsule with 2-lobed sperm hemiduct and a long, carinate or scoop-like basal lobe. Chelicerae with typical buthid dentition: movable finger dorsal margin with large subdistal and medial denticles and two smaller basal denticles, ventral margin with two large denticles, dorsal distal tine slightly shorter than ventral counterpart; fixed finger with subdistal denticle, and median and basal denticles formed as a bicusp, ventral surface armed with two small denticles (Figs. 252–254). Pedipalps orthobothriotaxic, type Aβ, femur trichobothrium d2 internal, patella d3 external to dorsomedian carina, chela db in distal half of fixed finger. Chela manus smooth, with carinae reduced or obsolete, dentate margins of chela
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 5 Figures 24–39: Hemispermatophore capsules of Barbaracurus gen. n. and Babycurus species, convex and anterior views. In each figure. left image is convex view with capsule compressed to show shape of lobes, right image anterior view to show profile of basal lobe. Figures 24–31. Color images. Figures 24–28: Barbaracurus gen. n. Figure 24. B. winklerorum sp. n., holotype. Figure 25. B. exquisitus comb. n., Oman, Jabal Akhdar hotel, 23º04'N 57º38'E. Figure 26. B. sofomarensis comb. n., paratype from the type locality. Figure 27. B. somalicus comb. n., Somaliland, Borama, locality 17SR. Figure 28. B. zambonellii comb. n., Eritrea, Filfil, locality 15EH. Figures 29–31: Babycurus. Figure 29: B. jacksoni, Tanzania. Figure 30: B. gigas, Tanzania, Usambara Mts. Figure 31: B. buettneri, Cameroon, Mamba vill., 04°07.311'N 10°11.615'E. Scale bars: 200 µm. Small numbered labels indicate specimen numbers (see lists of Material Examined). Figures 32–39. Gray scale images with basal lobes outlined in red. Figures 32–36: Barbaracurus gen. n.: B. winklerorum sp. n. (32), B. exquisitus comb. n. (33), B. sofomarensis comb. n. (34), B. somalicus comb. n. (35) and B. zambonellii comb. n. (36). Figures 37–39: Babycurus: B. jacksoni (37), B. gigas (38) and B. buettneri (39). Solid red curves indicate either the distal margin of a scoop-like basal lobe (32, 34) or a well developed carina (35–37, 39). Dashed red curves indicate either the line of attachment of a scoop-like basal lobe (32, 34) or a weakly developed carina (35–36, 38).
Euscorpius — 2018, No. 267 6 Figures 40–49: Lychas burdoi (Simon, 1882) Figures 40, 42, 44, 48. Male paralectotype of Babycurus ornatus Werner, 1936, dorsal (40) and ventral (42) views, pedipalp chela dorsal (44), metasoma IV–V and telson lateral (48). Figures 41, 43, 45–47, 49. emale lectotype of Babycurus ornatus Werner, 1936, dorsal (41) and ventral (43) views, pedipalp chela dorsal (45), Distal near or contiguous rows of denticulate granules, each external accessory granule midway along its length. F segments of right legs III (46) and IV (47) retrolateral views, and metasoma IV and telson lateral (49). Scale bar: 10 mm (40–43). movable finger armed with 6–7 non-imbricated, almost cies from Cameroon and Nigeria) a single isolated li row terminated proximally by an enlarged granule flanked by adjacent single internal and external accessory granules. Most proximal granule row without internal accessory denticle, and either with (species from Horn of Africa and Arabian Peninsula) or without (spePedipalp chelae sexually dimorphic, males typically with manus dilated and fingers proximally undulate on dentate margins, denticles of proximal granule rows bicuspid. Tibial spurs absent on leg III, present on leg IV, tibia and tarsus III–IV without bristle combs, ventral
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 7 Figures 50–69: Pedipalp chela dorsal and external views, comparison of Barbaracurus gen. n. species. Figures 50–53: B. sofomarensis comb. n., male (50–51) and female (52–53) paratypes. Figures 54–57: B. somalicus comb. n., male (54–55) and emale (56–57) from Somaliland, locality 17SR. Figures 58–61: B. zambonellii comb. n., male (58–59) and female (60–61) from f Eritrea, locality 15 69: B. winklerorum sp. n., male holotype (66–67) and female (68–69) paratype. surfaces of tarsi equipped with two rows of setae, ngues stout. 264) show considerable interspecific variability of the diploid chrom EH. Figures 62–65: B. exquisitus comb. n., male holotype (62–63) and female (64–65) paratype. Figures 66– . prudenti (Lourenço, 2013) comb. n. ameroon), B. sofomarensis (Kovařík et al., 2015) is comb. n. (2n=22) (Kovařík t al., 2015), B. zambonelli comb. n. (2n=26) (Figs. 261, 262) and B. somalicus comb. n. (2n=36) (Figs. 263, osomal numbers within this genus. The chromosomes have holocentric organization which is the ainly in the Horn of Africa and the Arabian eninsula (Fig. 265). It is differentiated from Babycurus primarily by its pedipalp finger dentition. In Barbaracurus gen. n., the pedipalp chela movable finger is u SUBORDINATE TAXA. B. exquisitus (Lowe, 2000) comb. n. (Oman), B (C comb. n. (Ethiopia), B. somalicus (Hirst, 1907) comb. n. (Somaliland), B. subpunctatus (Borelli, 1925) comb. n. (Ethiopia, Somalia), B. ugartei (Kovařík, 2000) comb. n. (Nigeria), B. winklerorum sp. n. (Oman), B. yemenensis sp. n. (Yemen), B. zambonellii (Borelli, 1902) comb. n. (Eritrea). CYTOGENETICS. The karyotype analyses of three species Barbaracurus sofomarens e typical feature of the family Buthidae (Schneider et al., 2009a). Males display achiasmatic meisois that is characteristic to the whole order Scorpiones (Schneider et al., 2009b). AFFINITIES. Seven of the species in this genus were previously included in Babycurus sensu lato. Here we transfer them to a new genus and describe two additional species in that genus. Barbaracurus gen. n. is a rather morphologically uniform assemblage of species that is distributed m P
Euscorpius — 2018, No. 267 14 igures 107–112: Barbaracurus exquisitus comb. n. Figures 107–110. Metasoma V and telson. Dorsal (107). lateral (108) F amd ventral (109) views. UV fluorescence. Paratype male, Oman, Jabal Akhdar. Scale bar: 1 mm. Figures 110–111. In vivo habitus, holotype male (110) and topoparatype female (111). Figure 112. Habitat, Oman, Jabal Shams, Jabal Akhdar.
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 15 Figures 113–117: Barbaracurus somalicus comb. n. from Somaliland, locality 17SR. Figures 113–114. Male, dorsal (113) and ventral (114) views. Figures 115–116. Female, dorsal (115) and ventral (116) views. Figure 117. Right hemispermatophore, convex aspect. Scale bars: 10 mm (113–116), 1 mm (117). metasomal segments (Figs. 70–71); metasoma I with 10 arinae, II–IV with 8 carinae. Telson setose, bearing nummale; aculeus slender, curved, shorter than vesicle. Barbaracurus sofomarensis (Kovařík, Lowe, Seiter, Plíšková et Šťáhlavský, 2015) comb. n. –30, figs. 46–55, 58–61, 64–65, 67–77, 87–102, 123, table 2. c erous long macrosetae and short, pointed subaculear tubercle (Figs. 80–82, 107–109); vesicle smooth, elongate, pyriform, telson length/depth ratio 2.70 in male, 2.89 in (Figures 2, 26, 34, 50–53, 72–73, 265) Babycurus sofomarensis Kovařík et al., 2015: 5 fe
Euscorpius — 2018, No. 267 16 Figures 118–127: Barbaracurus somalicus comb. n. from S chelicerae, carapace and tergites I–III (118), sternopectinal regi dorsal (123), and ventral (124) views. Figures 119, 121, 1 sternopectinal region and sternites III–V (121), metasoma and bar: 10 mm (122–127). o on 25–127. F (119), telson, late . Scale maliland, locality 17SR. Figures 118, 120, 122–124. Male, and sternites III–V (120), metasoma and telson, lateral (122), emale, chelicerae, carapace and tergites I–III ral (125), dorsal (126), and ventral (127) views
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 17 Figures 128–142: Barbaracurus somalicus comb. n. from Somaliland, locality 17SR. Figures 128–133. Male, pedipalp chela, dorsal (128), external (129), and ventral (130) views, pedipalp patella, dorsal (131), external (132) and ventral (133) views. igures 134–142. FemaF ele, pedipalp chela, dorsal (134), external (135), and ventral (136) views, pedipalp patella, dorsal (137), xternal (138) and ventral (139) views, pedipalp femur and trochanter internal (140) and dorsal (141) views, pedipalp movable nger dentate margin (142). The trichobothrial pattern is indicated n Figures 135–138 and 140–141 (white circles). fi i
Euscorpius — 2018, No. 267 18 Figures 143–144: Barbaracurus somalicus comb. n., in vivo habitus. Male (143) and female (144) from Somaliland, locality 7SR. , 4♂ (para1 TYPE LOCALITY AND TYPE DEPOSITORY. Ethiopia, Oromia State, Arsi Province, Sof Omar, 06°54'19"N 40°51' 04"E, 1200 m a.s.l.; FKCP. MATERIAL EXAMINED (FKCP). Ethiopia, Oromia State, Arsi Province, Sof Omar, 06°54'19"N 40°51'04"E, 1200 m a.s.l. (locality No. 13EC), 24.-25.VI.2013
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 19 Figures 145–146: Figure 145. Locality of Barbaracurus somalicus comb. n., Somaliland, locality 17SR. Figure 146. Yemen, Jabal Bura NEE Al Hudaydah, 14°53'N 43°26'E, 557 m a.s.l., locality of paratype of Barbaracurus yemenensis sp. n.
Euscorpius — 2018, No. 267 20 B. zambonellii B. somalicus DIMENSIONS (MM) ♂ (876) ♀ ♂ ♀ Carapace L / W 4.050 / 3.575 5.375 / 5.015 4.075 / 3.725 4.750 / 4.500 Mesosoma L 8.400 13.600 9.550 11.250 Tergite VII L / W 2.200 / 3.500 3.850 / 5.360 2.525 / 3.550 3.500 / 4.650 Metasoma + telson L 20.875 27.225 21.875 24.650 Segment I L / W / D 2.550 / 2.375 / 2.150 3.350 / 2.762 / 2.350 2.700 / 2.275 / 2.175 3.050 / 2.500 / 2.325 Segment II L / W / D 3.150 / 2.250 / 2.175 4.100 / 2.575 / 2.375 3.200 / 2.175 / 2.125 3.500 / 2.350 / 2.125 Segment III L / W / D 3.400 / 2.250 / 2.215 4.450 / 2.550 / 2.400 3.475 / 2.225 / 2.150 3.850 / 2.300 / 2.250 Segment IV L / W / D 3.900 / 1.965 / 2.225 5.025 / 2.525 / 2.575 4.000 / 2.225 / 1.950 4.375 / 2.350 / 2.175 Segment V L / W / D 4.575 / 2.000 / 1.875 5.850 / 2.500 / 2.400 4.600 / 2.250 / 1.925 5.400 / 2.300 / 2.075 Telson L / W / D 3.300 / 1.400 / 1.450 4.450 / 1.762 / 1.875 3.900 / 1.275 / 1.400 4.475 / 1.625 / 1.550 Pedipalp L 14.950 19.337 15.450 17.550 Femur L / W 3.725 / 1.150 4.800 / 1.875 3.750 / 1.175 4.275 / 1.275 Patella L / W 4.250 / 1.650 5.537 / 1.850 4.450 / 1.600 5.025 / 1.850 Chela L 6.975 9.000 7.250 8.250 Manus W / D 2.038 / 2.000 2.100 / 1.9500 2.100 / 2.200 1.975 / 1.813 Movable finger L 4.100 5.800 4.450 5.450 Total L 33.33 46.20 35.50 40.65 able depth (D) 1: Measurements of Barbaracurus zambonellii comb. n. and B. somalicus comb. n. Abbreviations: length (L), width (W, in carapace it corresponds to posteri , . or width) T
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 21 Figures 147–152: Barbaracurus winklerorum sp. n. Figures 147–148. Male holotype, dorsal (147) and ventral (148) views. Figures 149–150. Female paratype, dorsal (149) and ventral (150) views. Figures 151 – 152. Right hemispermatophore, convex aspect (151), posterior aspect of capsule region (152). Scale bars: 10 mm (147–150), 1 mm (151), 200 µm (152). types) (UV detection), leg. F. Kovařík, J. Plíšková et P. Novák, 23.-24.XI.2014, 2♂ (paratypes) 1♀ (holotype) (UV detection), leg. F. Kovařík; Oromia State, West Harerge, 07°44'37"N 40°42'39.5"E, 1234 m a.s.l. (locality No. 14EO), 24.-25.XI.2014, 1♂ (paratype) (UV detection), Oromia State, West Harerge, 07°46'39.7"N 40°37'12.4"E, 800 m a.s.l. (locality No. 14EP), 25. XI.2014, 1juv. (paratype).
Euscorpius — 2018, No. 267 22 Figures 153–162: Barbaracurus winklerorum sp. n. Figures 153, 155, 157–159. type, chelicerae, carapace and tergites I–IV (153), sternopectinal region and sternites III–IV (155), metasoma and telson, lateral (157), ventral (158), and dorsal (159) views. Figures 154, 156, 160–162. Female, chelicerae, carapace and tergites I–IV nopectinal region and sternite III (156), metasoma and telson, lateral (160), ventral (161), and dorsal (162) views. Scale bar: 10 mm (157–162). Male holo (154), ster
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 23 Figures 163–181: Barbaracurus winklerorum sp. n. Figures 163–171, 181. Male holotype, pedipalp chela, dorsal (163), 1 a external (164), and ventral (165) views, pedipalp patella, dorsal ( 66), external (167) and ventral (168) views, pedipalp femur and trochanter ventral (169), internal (170) and dorsal (171) views, pedipalp movable finger dentate margin (181). Figures 172–180. Female, pedipalp chela, dorsal (172), external (173), and ventral (174) views, pedipalp patella, dorsal (175), external (176) and ventral (177) views, pedipalp femur and trochanter dorsal (178) and ventral (179) views, pedipalp movable finger dentate margin (180). The trichobothrial pattern is indicated in Figures 164–167 nd 170–171 (white circles).
Euscorpius — 2018, No. 267 30 Figures 219–225: Barba r c H. Figures 219, 221. Male, dorsal (219) and ventral (221) views. Figures 22 3. Right hemispermatophore, convex aspect. Figures 224–225 Barbaracurus yemenensis sp. n. (Figures 9, 78, 87, 103–106, 206–218, 245–247, 265, Table 2) http://zoobank.org/urn:lsid:zoobank.org:act:FBEDE C29-0D9D-40C4-A2EE-84AC7FAA257E Babycurus zambonellii n part, Yemen): Sissom, 1994: 5–6, figs. 1–7; řík, 1998: 104; Fet & Lowe, 2000: 80; Kov , 2000: 260–261, figs. 13, 26; Lowe, 2000: 18 91; Kovařík & Whitman, 2005: 106; Hendrixso : 109. racurus zambonellii 0 . Eritrea, comb . n. from Erit ea, lo ent ality 15 (22 E 2) v, 222. Female, dorsal (220) and 15EH. Scale bar v s: 10 mm (219–222), 1 ral iews. locality Figure 22 mm (223). (i Kova ařík 5–1 n, 2006
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 31 i d al (230) and external (231) views. The trichobothrial pattern is indicated in Figures 233–238 (white circles). Figures 226–238: Barbaracurus zambonellii comb. n. from chelicerae, carapace and tergites I–III (226), sternopectinal reg external (233), and ventrointernal (234) views, pedipalp patella trochanter internal (237) and dorsal (238) views. Figures 227, 2 sternopectinal region and sternites III–IV (229), pedipalp chela, Eritrea, locality 15EH. Figures 226, 228, 232–238. Male, on and sternite III (228), pedipalp chela, dorsoexternal (232), , dorsal (235) and external (236) views, pedipalp femur and 29–231. Female, chelicerae, carapace and tergites I–III (227), orsoextern
Euscorpius — 2018, No. 267 32 Figures 239–247: Figures 239–244: Barbaracurus zambonellii comb. n. from Eritrea, locality 15EH. Figures 239–241. Male, metasoma and telson, lateral (239), ventral (240), and dorsal (241) views. Figures 242–244. Female, metasoma and telson, lateral (242), ventral (243), and dorsal (244) views. Figures 245–247: Barbaracurus yemenensis sp. n., Female holotype, metasoma and lson, lateral (245), dorsal (246), and ventral (247) views. Scale bars: 10 mm (239–241, 242–244, 245–247). TYPE LOCALITY AND T SITORY. Yemen, Wadi FKCP. TYPE MATERIAL EXAMINED. Yemen, Wadi Dawan NW Al Mukalla, 15°09'N 48°26'E, 946 m a.s.l., 3.IV.2007, 1♀ (holotype), leg. P. Kabátek, FKCP; Jabal Bura NEE te YPE DEPO Dawan NW Al Mukalla, 15°09'N 48°26'E, 946 m a.s.l.;
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 33 Al Hudaydah, 14°53'N 43°26'E, 557 m a.s.l. (Fig. 146), 19.-21.III.2007, 1♀ (paratype), leg. P. Kabátek, FKCP; Hajjah gov., 2.–3. XI.2007, Halhal vill. env., NE Hajjah by road, 15°43'42"N 43°37'25"E, 998 m a.s.l., (locality No. 14), 1juv. (paratype), leg. D. Král, FKCP. ETYMOLOGY. Named after the country of occurence. DIAGNOSIS. Total length of adult females 40–42 mm, males unknown. Coloration pale yellow to light orange, chelicerae yellow without reticulation. Pedipalp chela length/width ratio 3.4–4.2 in females; pedipalp fingers of females straight (Fig. 211); dentate margin of movable finger armed with 7 rows of granules, and a short apical row of 4 denticles (Fig. 9); most proximal granule row with one external accessory granule. Pectines with 19– 21 teeth. Metasoma very narrow, metasoma V length/ width ratio is 2.30–2.46 in females (Figs. 79); metasoma I with 10 carinae, II–IV with 8 carinae. Telson setose, bearing numerous long macrosetae and short, pointed subaculear tubercle; vesicle smooth, ellipsoidal, slightly bulbous, telson length/depth ratio 2.78–2.88; aculeus slender, curved, shorter than vesicle. DESCRIPTION. Total length of adult females 40–42 mm, males unknown. Measurements of the carapace, telson, segments of the metasoma and segments of the pedipalps are given in Table 2. Base color base is pale yellow to light orange (Figs. 206–218). Chelicerae are yellow without reticulation. (Fig. 208). Sexual dimorphism unknown. PEDIPALP (Figs. 210–217). Pedipalp mostly very sparsely hirsute, more densely so on ventral surface of movable finger. Femur granulated, with five granulose carinae. Patella almost smooth with seven granulose carinae. Chela smooth with traces of carinae visible; fingers long, curved, with 7 granule rows and short apical row of 4 denticles on dentate margins; the most proximal row with one external and no internal granule. Pedipalp fingers straight in females. CARAPACE (Figs. 206, 208). Slightly trapezoidal (narrower anteriorly) and slightly longer than wide, or as long as wide; anterior margin convex, with some short microsetae. Carination absent. Median and posterior lateral furrows wide and deep, others vestigial to absent. Tegument densely and coarsely granulose. Median eyes large and raised; four or five pairs of lateral eyes: three major ocelli aligned along each anterolateral corner, plus two minor ocelli that are vestigial to absent. MESOSOMA (Figs. 206–209). Tergites I–VI bear a single conspicuous median carina; tergite VII bears five well nite III in females. Tooth count 19–21 (mean 20, 6 ♀combs). Pectines with 3 marginal lamellae and 7 middle lamellae. Sternites lack carinae, surfaces smooth and sparsely setose. Posterior margin of sternite V without smooth median patch in both sexes. Sternite VII with four well defined carinae. LEGS (Figs. 103–106). The tarsomeres bear two rows of macrosetae on their ventral surface and numerous macrosetae on other surfaces; bristle combs absent. Femur bears only solitary macrosetae. Femur coarsely granulose, femur and patella with carinae developed. Moderate tibial spurs present on leg IV. METASOMA AND TELSON (Figs. 78, 87, 245–247). All segments with granulate, completely developed carinae. The carinae are composed of minute, rounded, equalsized, and evenly spaced granules. The first metasomal segment has a total of 10 carinae, the second through fourth segments have eight carinae, and the fifth segment has five carinae. All metasomal segments are very sparsely granulated. Metasoma is very sparsely hirsute. Telson smooth with only a weak trace of a ventral carina and a dense cover of long setae mainly on the ventral surface. defined carinae (median, submedians and laterals), which are long and serrate to crenulate. All tergites densely and coarsely granulose mainly on posterior parts. Sternum type 1, triangular in shape; medial depression large. Pectines extending to around end of sterSubaculear tubercle short and pointed. Vesicle elongate, ellipsoidal, telson length/depth ratio 2.78–2.88 in females. Aculeus curved, shorter than vesicle. Barbaracurus zambonellii (Borelli, 1902) comb. n. (Figures 7, 28, 36, 58–61, 79–80, 88–89, 219–244, 248– 251, 261–262, 265, Table 1) Babycurus zambonellii Borelli, 1902: 1–4; Hirst, 1907: 209; Kraepelin, 1913: 181; Caporiacco, 1947: 232; Probst, 1973: 329; Lamoral & Reynders, 1975: 498; Kovařík, 1998: 104 (in part); Fet & Lowe, 2000: 80 (in part); Lowe, 2000: 185–191 (in part); Kovařík, 2000: 260–261 (in part); Kovařík, 2003: 137 (? in part); Chiarle et al., 2012: 21. TYPE LOCALITY AND HOLOTYPE DEPOSITORY. Eritrea, Chenafena; MZUT. MATERIAL EXAMINED. Eritrea, Chenafena (14.786N 39.018E), 1♀ (holotype), MZUT; Filfil, Salamuna river, 15°36'34.6"N 38°57'22.8"E, 817 m a.s.l., 3.-4.XI.2015, (Figs. 224–225, Locality No. 15EH), 3♂3♀1♀im.3juvs. (No. 876, Figs. 28, 36, 219–223, 226–244, 258–251, 258–259), leg. F. Kovařík, FKCP. DIAGNOSIS. Total length of adult males 32–35 mm, adult females 46–52 mm. Coloration yellowish brown to grey with darker markings, chelicerae yellow without or with traces of reticulation. Pedipalp chela manus much wider in male than female, chela length/width ratio 3.42 in males and 4.29 in female; proximal margins of pedipalp fingers of female straight (Figs. 61, 231), of male un-
Euscorpius — 2018, No. 267 34 v Figures 248–249: Barbaracurus zambonellii comb. n., in vi 15EH. o habitus. Male (248) and female (249) from Eritrea, locality
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 35 ea Figures 250–251: Barbaracurus zambonellii comb. n. female w with juveniles (second instar) after first ecdysis (251) from Eritr ith newborns (first instar) before first ecdysis (250) and female , locality 15EH.
Euscorpius — 2018, No. 267 36 Figures 252–254: Barbaracurus exquisitus comb. n. Right chelicera of male, dorsal (252), dorsointernal (253) and ventral (254) views. Paratype male, Jabal Shams, Oman, 14.X.1993. Scale bar: 500 µm. dulate so as to leave a gap with fingers closed (Figs. 59, 233); dentate margin of movable finger armed with 7 rows of granules, and a short apical row of 3–4 denticles (Fig. 7); most proximal granule row with one external accessory granule. Pectines with 17–19 teeth in both sexes. Hemispermatophore basal lobe a weak, oblique carina (Figs. 28, 36). Metasoma narrow, metasoma V length/width ratio is 2.40–2.46 in males and 2.56–2.58 in females (Figs. 76–77); metasoma I with 10 carinae, II–IV with 8 carinae. Telson setose, bearing numerous long macrosetae and short, pointed subaculear tubercle; vesicle smooth, elongate, ellipsoidal, slightly bulbous, telson length/depth ratio 2.27–2.37 in both sexes; aculeus slender, curved, shorter than vesicle. NOTE. In his original description, Borelli (1902: 3) assumed that the holotype was a male, and this was accepted by most subsequent authors. Only Lowe (2000: 190–191) questioned this assumption, observing that the holotype exhibited some characters more consistent with females of other species of Babycurus sensu lato, and noted that “Study of additional material is needed to larify variation and sexual dimorphism in B. zamaterial representing both sexes of B. zambonellii. The new specimens clearly demonstrated that the holotype is indeed female, not male. B. zambonellii is an Eritrean endemic and the female which Sissom (1994) cited from Yemen represents a new species which we describe here as B. yemenensis sp. n. COMMENTS ON LOCALITY AND LIFE STRATEGY. The first author (F.K.) visited the locality 15EH (Figs. 224–225), a montane, forested habitat along a riverbed of an occasional river, on 3–4 November 2015. At this locality, the author recorded a maximum daytime temperature of 30.3 ºC, and minimum nighttime temperatures of 19.6 ºC. The recorded humidity was between 46% and 92%. In addition to B. zambonellii the first author also recorded Hottentotta minax (L. Koch, 1875) at this locality. Acknowledgments Thanks are due to Marcel Bednář, Petra Frýdlová, Daniel Frynta, Martin Häckel, Pavel Just, Petr Kabátek, Hynek Kmoníček, David Král, Pavel Kučera, Pavel Novák, Vít Socha, Jana Štundlová, Vladimír Trailin, and Arab Emirates), Dereje Belay, Daneil Denbi, Aba c bonellii”. Recently, the first author (F.K.) was able to finally settle this question by collecting additional David Vašíček (Czech Republic), Alex and Birgit Winkler (Munich, Germany), Gary R. Feulner (United m
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 37 Figures 255–264: Male postpachytene (255, 257, 261, 263), mitotic metaphase (259), and ideograms (256, 258, 260, 262, 264) (y axis - % of the diploid chromosome length) of Babycurus (255–260) and Barbaracurus gen. n. (261–264) species. Babycurus jacksoni (1189), Tanzania (2n=16) (255, 256), B. buettneri (1190) from Cameroon, Mamba vill., 04°07.311'N 10°11.615'E (2n=28) (257, 258) and B. gigas (914) from Tanzania (2n=30) (259, 260); Barbaracurus zambonelli comb. n. (876) from Eritrea, locality 15EH (2n=26) (261, 262) and B. somalicus comb. n. (1332), Somaliland, locality 17SR (2n=36) (263, 264). Scale bar: 5 µm (255, 257, 259, 261, 263). ebede, and Zelalem Mandefro (EthiCommunication Directorate, University of Hargeisa), med A. Boqore (Vice President, AcaGragn, Zelalem K opia), Mohamud Yousuf Muse (President of University f Hargeisa), Mohamed A. Sulub (Director, Corporate Suleiman Ahmed Gulair (President of Amound University), Ah o
Euscorpius — 2018, No. 267 38 rba o Figure 265: Map showing the known distribution of genera Ba Horn of Africa, Kenya and partially in Tanzania. The upper phot n. demic Affairs of Amound University), and Yesuf Ahmed Ali (Director General of Higher Education, Hargeisa, Republic of Somaliland) for their help. Special thanks to Abdiqaadir Abdilahi, Abdisalaan Shabele, racurus gen. n. and Babycurus in the Arabian Peninsula, inset shows the type locality of Barbaracurus winklerorum sp. Omar Yussuf Hussein (Republic of Somaliland), Stefan Friedrich (ZSMC), Mark Stockmann (Germany) and Tomáš Mazuch (Czech Republic); Nadine Dupérré and Danilo Harms (Centrum für Naturkunde (CeNak),
Kovařík, Lowe & Šťáhlavský: New Genus Barbaracurus 39 Center of Natural History Universität Hamburg, Zoological Museum, Hamburg, Germany) for loans of Babycurus ornatus types; Victor Fet and Michael Soleglad for their help in processing the manuscript. We are grateful to Jana Štundlová (Czech Republic) for technical assistance during chromosome preparations. We are also indebted to Petr Kabátek for kindly giving permission to use his photo from Yemen herein as Figure 146, and to Birgit Winkler for contributing her photo in Fig. 265. Further, we thank two anonymous reviewers for their comments to the manuscript. References BORELLI, A. 1902. Di una nuova specie di scorpione della Colonia Eritrea. Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Universitá di Torino, 17 (422): 1–4. BORELLI, A. 1925. Di alcuni Scorpioni della Somalia Italiana. Annali del Museo Civico di Storia Naturale di Genova, 51: 316–326. CAPORIACCO, L. DI 1947. Scorpioni dell’Eritrea del Museo zoologici di Firenze. Acta Pontificae Academiae Scientiarum Novi Lyncei, 11(19): 227–233. CHIARLE, A., F. KOVAŘÍK, L. LEVI & E. GAVETTI. 2012. The scorpion collections (Arachnida, Scorpiones) held in the Museo Regionale di Scienze Naturali of Turin (Italy). Arachnologische Mitteilungen, 43: 17–23. ESPOSITO, L.A., H.Y. YAMAGUTI, C.A. SOUZA, R. PINTO-DA-ROCHA & L. PRENDINI. 2017. Systematic revision of the neotropical club-tailed scorpions, Physoctonus, Rhopalurus, and Troglorhopalurus, revalidation of Heteroctenus, and descriptions of two new genera and three new species (Buthidae: Rhopalurusinae). Bulletin of the American Museum Of Natural History, 145: 1–134. 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 r for the high-level systematics of Buthoidea (Scorpiones: Buthida). EusHENDRIXSON, B.E. 2006. Buthid scorpions of Saudi Arabia, with notes on other families (Scorpiones: Buthidae, Liochelidae, Scorpionidae). Fauna of Arabia, 21: 33–120. HIRST, S. 1907. Notes on scorpions, with descriptions of two new species. Annals and Magazine of Natural History, VII(19): 208–211. KARSCH, F. 1886. Skorpionologische Beiträge. Berliner Entomologische Zeitschrift, 30: 75–79. KOVAŘÍK, F. 1998. Štíři [Scorpiones]. Jihlava (Czech Republic): Publishing House ”Madagaskar”, 176 pp (in Czech). KOVAŘÍK, F. 2000. Revision of Babycurus with descriptions of three new species (Scorpiones: Buthidae). Acta Societatis Zoologicae Bohemicae, 64: 235–265. KOVAŘÍK, F. 2003. Scorpions of Djibouti, Eritrea, Ethiopia, and Somalia (Arachnida: Scorpiones) with a key and descriptions of three new species. Acta Societatis Zoologicae Bohemicae, 67: 133–159. KOVAŘÍK, F. 2009. Illustrated catalog of scorpions. Part I. Introductory remarks; keys to families and genera; subfamily Scorpioninae with keys to Hetero-metrus and Pandinus species. Prague: Clairon Production, 170 pp. KOVAŘÍK, F. 2011. Buthus awashensis sp. n. from Ethiopia (Scorpiones, Buthidae). Euscorpius, 128: 1–6. KOVAŘÍK F., G. LOWE, P. JUST, A. I. AWALE, H. SH A. ELMI & F. ŠŤÁHLAVSKÝ. 2018. Scorpions of the Horn of Africa (Arachnida: Scorpiones). Part XVI. Review of the genus Gint Kovařík et al., 2013, with description of three new species from Somaliland (Scorpiones, Buthidae). Euscorpius, 258: 1–41. KOVAŘÍK, F., G. LOWE, M. SEITER, J. PLÍŠKOVÁ & F. ŠŤÁHLAVSKÝ. 2015. Scorpions of Ethiopia (Arachnida: Scorpiones), Part II. Genus Babycurus Karsch, 1886 (Buthidae), with description of two Bothriuridae; Chaerilidae; Buthidae I. Genera Compsobuthus, Hottentotta, Isometrus, Lychas, and Scorpions of the World (1758–1998). New York: The New York Entomological Society, 689 pp. FET, V., M.E. SOLEGLAD & G. LOWE. 2005. A new trichobothrial characte new species. Euscorpius, 196: 1–31. KOVAŘÍK, F. & A. A. OJANGUREN AFFILASTRO. 2013. Illustrated catalog of scorpions. Part II. corpius, 23: 1–40. Sassanidotus. Prague: Clairon Production, 400 pp.