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Barbaracurus kabateki sp. n. (Figures 1–33, 101–103, 107, Table 1)

Key to species of Barbaracurus 1. Pedipalp movable finger without an external accessory granule midway along most proximal granule row......... 2 – Pedipalp movable finger with an external accessory granule midway along most proximal granule row...................... 3 2. Base color uniformly yellow or orange, without any darker markings; sternite VII with very weak carination.......................................................... B. prudenti (Lourenço, 2013) – Base color yellow with brown spots on carapace, dark stripes on tergites, dark pedipalp patella and metasoma V; sternite VII with 4 well developed carinae.......................................................................... B. ugartei ( KovařÍk, 2000) 3. Pedipalp movable finger with 6 rows of granules........... 4 – Pedipalp movable finger with 7 rows of granules........... 7 4. Pedipalp chela with narrower manus, chela length/ width ratio 4.3–5.6, finger margins weakly undulate at base, not leaving gap when closed (figs. 81, 84, 86 in KovařÍk et al., 2015)............................................................................... 5 – Pedipalp chela with broader manus, chela length/ width ratio 3.4–4.2, finger margins strongly undulate at base, leaving wide gap when closed.......................................... 6 5. Total length 22.5 mm in male, 32.1–32.25 mm in females; pectines with 16–17 teeth in both sexes; distributed in Ethiopia and Somalia..... B. subpunctatus ( Borelli, 1925) – Total length 31.25 mm in male, 38.80 mm in female; pectines with 25–27 teeth in both sexes; distributed in Somaliland.............................. B. feti KovařÍk et al., 2019 6. Telson vesicle pyriform in lateral profile, deeper anteriorly (figs. 83–84 in KovařÍk et al., 2018b); telson length/ depth ratio 2.75–2.89; pedipalp movable finger of female very weakly undulate at base; hemispermatophore basal lobe a weak carina (figs. 27, 35 in KovařÍk et al., 2018b)............................................................. B. somalicus (Hirst, 1907) – Telson vesicle symmetric in lateral profile (figs. 76–77 in KovařÍk et al., 2015); telson length/ depth ratio 2.60–2.73; pedipalp movable finger of female moderately undulate at base; hemispermatophore basal lobe a well developed scoop-like lamina (figs. 26, 34 in KovařÍk et al., 2018b)................................. B. sofomarensis ( KovařÍk et al., 2015) 7. Pedipalp chela with broader manus (Figs 53–55, and figs. 58, 60 in KovařÍk et al., 2018b), chela length/ width ratio 4.28–5.43 (♀), 3.2–3.42 (♂); proximal margins of pedipalp fingers of male strongly undulate, leaving gap with fingers closed (Fig. 54, and fig. 59 in KovařÍk et al., 2018b)...... 8 – Pedipalp chela with narrower manus (Figs. 7–9, figs. 62, 64, 66, 68, 210, 212 in KovařÍk et al., 2018b), chela length/ width ratio 4.07–6.12 (♀, ♂); proximal margins of pedipalp fingers of male weakly undulate, not leaving gap with fingers closed (Fig. 8, and figs. 62–67 in KovařÍk et al., 2018b)........................................................................ 9 8. Pectines with 17–19 teeth (♀, ♂); telson more bulbous, length/ depth ratio 2.27–2.37 (♀, ♂); distributed in Africa (Eritrea)............................. B. zambonellii (Borelli, 1902) – Pectines with 22–25 (♂) 19–23 (♀) teeth; telson less bulbous, length/ depth ratio 2.48–2.70 (♀, ♂); distributed in Arabian Peninsula............................................................................................... B. yemenensis KovařÍk et al., 2018 9. Metasoma V length/ width ratio 2.71 (♂); pedipalp chela length/ width ratio 4.07 (♂); hemispermatophore basal lobe obsolete, reduced to a weak ridge (Figs. 102–103); found in Saudi Arabia......................................... B. kabateki sp. n. – Metasoma V length/ width ratio 2.40–2.47 (♂); pedipalp chela length/ width ratio 4.24–4.70 (♂); hemispermatophore basal lobe a well developed scoop-like lamina (figs. 24– 25, 32–33 in KovařÍk et al., 2018b); found in Oman. – Telson less slender (figs. 85–87 in KovařÍk et al., 2018b), length/ depth ratio 2.70–2.72 (♂, ♀); not found in northern Oman....................... B. winklerorum KovařÍk et al., 2018, Published as part of Kovařík, František, Lowe, Graeme, Šťáhlavský, František & Just, Pavel, 2022, The genus Barbaracurus in Saudi Arabia (Scorpiones: Buthidae), with description of a new species., pp. 1-26 in Euscorpius 365 on page 23, DOI: 10.5281/zenodo.7616748, {"references":["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.","KOVARIK, F., G. LOWE, F. STAHLAVSKY & A. A. HURRE. 2019. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part XX. Barbaracurus feti sp. n. from Somaliland (Buthidae). Euscorpius, 280: 1 - 11.","KOVARIK, F., G. LOWE & F. STAHLAVSKY. 2018 b. Review of the genus Babycurus Karsch, 1886 (Arachnida, Scorpiones, Buthidae), with description of Barbaracurus gen. n. and two new species from Oman and Yemen. Euscorpius, 267: 1 - 41."]}

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GINT GAITAKO All specimens exhibit 2 n = 30 chromosomes, which decrease in length from 4.64% to 1.80% of DSL (Supporting Information, Fig. S2A; Table S2). The post-pachytene cells show 13 bivalents and one quadrivalent (chromosomes 1, 18, 25 and 30) (Fig. 2M; Supporting Information, Fig. S2A). The pair of 18S rDNA sites is located in the terminal region of the third largest bivalent (chromosomes 6 and 7) (Fig. 2N; Supporting Information, Fig. S2A)., Published as part of Just, Pavel, Šťáhlavský, František, Kovařík, František & Štundlová, Jana, 2022, Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae), pp. 885-901 in Zoological Journal of the Linnean Society 196 (2) on page 891, DOI: 10.1093/zoolinnean/zlac049, http://zenodo.org/record/7184492

GINT GUBANENSIS Karyotype of G. gubanensis comprises 2 n = 45 chromosomes, which decrease in length from 2.97% to 1.02% of DSL (Supporting Information, Fig. S2G; Table S2). Overall, 21 bivalents and a single trivalent, composed of chromosomes 7, 33 and 45, are detected in post-pachytene nuclei (Fig. 3I; Supporting Information, Fig. S2G). One pair of 18S rDNA loci is found on a terminal part of the heteromorphic chromosome pair (chromosomes 26 and 44). A single large 18S rDNA cluster is located on the overhanging part of chromosome 26 (the larger chromosome of the heteromorphic pair) (Fig. 3J; Supporting Information, Fig. S2G). MTDNA DIVERSITY AND PHYLOGENY The sequences for both 16S and COI were obtained from 22 Gint individuals. In G. calviceps, only the 16S gene fragment was successfully sequenced. The final alignment with a total length of 963 bp (16S: 358 bp; COI: 605 bp) consisted of 765 conserved sites, 194 variable sites and 172 parsimony-informative sites. Based on COI data, Gint exhibits genetic variation at both inter- and intraspecific levels, with the pairwise genetic distances ranging from 3.28% to 11.52% between species, and from 0.00% to 2.64% within species (Supporting Information, Tables S3, S4). The haplotype network reconstruction showed a presence of 15 COI sequence haplotypes in Gint (Supporting Information, (Fig. S3; Table S3). Within the species studied, G. banfasae shows the highest within-location haplotype diversity (four haplotypes per locality), but with minor pairwise genetic differences (p -distance: ≤0.66%). In this species, all individuals of the same cytotype have distinct haplotypes. The haplotypes of cytotype I individuals (S1531 and S1534) are separated from one another by four mutational steps. Cytotype II (S1532) shares its haplotype with the cytotype III individual (S1530), which is one mutational step distant from the haplotype of the cytotype III specimen (S1533) (Supporting Information, Fig. S3). Gint dabakalo, comprising three haplotypes, shows the highest intraspecific genetic variation with a p -distance of 0.17–2.64%. In this species, the genetic distance between individuals of cytotype I and cytotype II from the same locality is higher than between each of them and a specimen of cytotype III from a distinct locality (Supporting Information, Fig. S3; Table S1). Gint maidensis, a species with a stable karyotype 2 n = 34, is represented by three haplotypes with a p -distance of 0.33–0.50%. Gintgaitako presentstwohaplotypes, where one haplogroup, formed by five specimens, is distant from the other by one mutational step (p -distance: 0.17). In G. amoudensis, individuals of cytotype II, cytotype III and cytotype IV share the same haplotype, whereas the specimen of cytotype I differs from this haplogroup by eight point mutations (p -distance: 1.49%). Both BI and ML mtDNA analyses provide similar tree topologies and nodal supports for clades. Phylogenetic reconstruction depicts Gint as a monophyletic group (Fig. 5). Most of the Gint species form well-supported clades [bootstrap support (bs) ≥ 83, posterior probability (PP) ≥ 0.98]. The monophyly of the G. banfasae specimens remains unresolved due to low support values. Gint maidensis forms an early-branching clade, which is sister to the clade comprising all of the remaining Gint species. This clade is further subdivided into two groups: (1) a clade in which G. gubanensis is a sister-species to G. amoudensis, and (2) a clade consisting of G. gaitako, G. calviceps, G. dabakalo and G. banfasae. In the latter, G. gaitako forms a sister-lineage to the clade comprising G. calviceps, G. dabakalo and G. banfasae. The mutual relationships between the species in this group remain unresolved due to low support values. THE RECONSTRUCTION OF ANCESTRAL CHROMOSOME NUMBER AND POSITION OF 18S RDNA GENE CLUSTERS Maximum parsimony analysis inferred 2 n = 32–34 as the ancestral chromosome number for the genus Gint (Fig. 6A). Two independent increases of 2 n were detected within Gint: (1) the clade containing all cytotypes of G. amoudensis and G. gubanensis, and (2) cytotype III within the G. dabakalo clade. When compared with the ancestral 2 n, a tendency of decreasing of 2 n from 30 to 18 chromosomes can be observed in G. gaitako, G. dabakalo and G. banfasae. Concerning the position of 18S rRNA gene clusters, maximum parsimony inferred their terminal position as the ancestral state for the genus Gint (Fig. 6B). Two independent changes of the ancestral terminal position to a subterminal position were identified in G. amoudensis and in the clade containing G. dabakalo and G. banfasae., Published as part of Just, Pavel, Šťáhlavský, František, Kovařík, František & Štundlová, Jana, 2022, Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae), pp. 885-901 in Zoological Journal of the Linnean Society 196 (2) on page 893, DOI: 10.1093/zoolinnean/zlac049, http://zenodo.org/record/7184492

GINT DABAKALO In this species, the differences in the diploid number, multivalent associations and the appearance of 18S rDNA-bearing chromosomes are detected at inter- and intrapopulation levels, resulting in three distinguished cytotypes. Specimen S 1305 from the locality Burao is assigned to cytotype I, and specimens S1527 and S1199 from N of Burao, Togdheer represent cytotypes II and III, respectively. Cytotype I (1 Ƌ; S1305) has 2 n = 23 chromosomes, which decrease in length from 7.90% to 2.55% of DSL (Supporting Information, Fig. S1D; Table S2). The post-pachytene cells show nine bivalents and one pentavalent that is composed of chromosomes 1, 4, 5, 10 and 23 (Fig. 2G; Supporting Information, Fig. S1D). The pair of 18S rDNA sites is located in the subterminal region of the largest bivalent (Fig. 2H). Cytotype II (1 Ƌ; S1527) exhibits 2 n = 24 chromosomes, which decrease in length from 7.65% to 2.95% of DSL (Supporting Information, Fig. S1E; Table S2). The post-pachytene nuclei show a configuration of eight bivalents, one trivalent (chromosomes 2, 12 and 17) and one pentavalent (chromosomes 1, 3, 4, 7 and 20) (Fig. 2I; Supporting Information, Fig. S1E). The pair of 18S rDNA sites is situated in the subterminal region of chromosomes 2 and 12 of the trivalent (Fig. 2J; Supporting Information, Fig. S1E). Cytotype III (1 Ƌ; S1199) has 2 n = 27 chromosomes, which decrease in length from 8.30% to 1.80% of DSL (Supporting Information, Fig. S1F; Table S2). In the post-pachytene spermatocytes, we observe eight bivalents, two trivalents and one pentavalent (Fig. 2K). The larger trivalent is composed of chromosomes 2, 9 and 10, the smaller trivalent is composed of chromosomes 4, 22 and 25, and chromosomes 1, 3, 5, 8 and 17 form a pentavalent (Supporting Information, Fig. S1F). The 18S rDNA sites are located in the subterminal region of the chromosomes 2 and 9 of the larger trivalent (Fig. 2L; Supporting Information, Fig. S1F)., Published as part of Just, Pavel, Šťáhlavský, František, Kovařík, František & Štundlová, Jana, 2022, Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae), pp. 885-901 in Zoological Journal of the Linnean Society 196 (2) on pages 890-891, DOI: 10.1093/zoolinnean/zlac049, http://zenodo.org/record/7184492

GINT BANFASAE In this species, we detected intrapopulation variability in diploid number and/or multivalent associations showing the presence of three cytotypes. The position of a single pair of 18S rDNA sites, as well as the 18S rDNA-bearing chromosome pair involved in the multivalent chain, are consistent in all cytotypes. Cytotype I (2 ƋƋ; S1531, S1534) has 2 n = 18 chromosomes, which decrease in length from 9.85% to 1.81% of DSL (Supporting Information, Fig. S1A; Table S2). In post-pachytene nuclei, the chromosomes form seven bivalents and one quadrivalent that is composed of the four larger chromosomes (Fig. 2A). A single pair of 18S rDNA sites is located in the subterminal region of the chromosomes 3 and 4 involved in the quadrivalent (Fig. 2B; Supporting Information, Fig. S1A). Cytotype II (1 Ƌ; S1532) exhibits 2 n = 18 chromosomes, which decrease in length from 9.79% to 1.77% of DSL (Supporting Information, Fig. S1B; Table S2). The post-pachytene cells show six bivalents and one hexavalent (chromosomes 1, 2, 3, 4, 15 and 18) (Fig. 2C; Supporting Information, Fig. S1B). A pair of 18S rDNA sites is located in the subterminal region of the chromosomes 3 and 4 involved in the hexavalent (Fig. 2D). Cytotype III (2 ƋƋ; S1530, S1533) has 2 n = 19 chromosomes, which decrease in length from 9.66% to 1.78% of DSL (Fig. 2E; Supporting Information, Fig. S1C; Table S2). Post-pachytene spermatocytes exhibit five bivalents, one trivalent (chromosomes 7, 17 and 18) and one hexavalent (chromosomes 1, 2, 3, 4, 14 and 19). A pair of 18S rDNA sites is situated in the subterminal region of the chromosomes 3 and 4 involved in the hexavalent (Fig. 2F)., Published as part of Just, Pavel, Šťáhlavský, František, Kovařík, František & Štundlová, Jana, 2022, Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae), pp. 885-901 in Zoological Journal of the Linnean Society 196 (2) on page 889, DOI: 10.1093/zoolinnean/zlac049, http://zenodo.org/record/7184492

GINT MAIDENSIS The karyotype of G. maidensis comprises 2 n = 34 chromosomes, which decrease in length from 4.22% to 1.87% of DSL (Supporting Information, Fig. S2B; Table S2). The post-pachytene nuclei exhibit 17 bivalents (Fig. 2O). No multivalent associations are found in this species. One pair of 18S rDNA sites is located in the terminal region of the largest bivalent (Fig. 2P)., Published as part of Just, Pavel, Šťáhlavský, František, Kovařík, František & Štundlová, Jana, 2022, Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae), pp. 885-901 in Zoological Journal of the Linnean Society 196 (2) on page 892, DOI: 10.1093/zoolinnean/zlac049, http://zenodo.org/record/7184492

GINT AMOUDENSIS T h e m o s t s t r i k i n g i n t r a s p e c i f i c v a r i a b i l i t y i s discovered in G. amoudensis, which exhibits four cytotypes differing in the diploid number, multivalent associations and in the number and position of 18S rDNA sites. The cytotypes II, III and IV occur sympatrically, while the cytotype I (S1325) is found in a remote locality Borama c. 150 km far from the other cytotypes. Cytotype I (1Ƌ; S1325) exhibits 2 n = 36 chromosomes, which decrease in length from 4.49% to 1.63% of DSL (Supporting Information, Fig. S2C; Table S2). The post-pachytene nuclei show 16 bivalents and one quadrivalent (chromosomes 1, 2, 9 and 15) (Fig. 3A; Supporting Information, Fig. S2C). Three 18S rDNA sites are detected (Fig. 3B), two of them are located in the subterminal region of the chromosomes 1 and 2 involved in the quadrivalent and one in the terminal position of chromosome 2 in the quadrivalent (Supporting Information, Fig. S2C). Cytotype II (1 Ƌ; S1291) has 2 n = 35 chromosomes, which decrease in length from 5.01% to 1.45% of DSL (Supporting Information, Fig. S2D; Table S2). The post-pachytene spermatocytes exhibit 14 bivalents, one trivalent (chromosomes 4, 24 and 35) and one quadrivalent (chromosomes 1, 5, 8 and 23) (Fig. 3C; Supporting Information, Fig. S2D). Three 18S rDNA loci are present (Fig. 3D), one pair is located on the chromosomes 1 (subterminal site) and 5 (terminal site) involved in the quadrivalent, and one single heterozygous locus is situated in the interstitial region of the third largest bivalent (chromosome 9; Fig. 3D; Supporting Information, Fig. S2D). Cytotype III (1 Ƌ; S1293) has 2 n = 36 chromosomes, which decrease in length from 4.71% to 1.52% of DSL (Supporting Information, Fig. S2E; Table S2). The post-pachytene cells show 15 bivalents and one hexavalent (chromosomes 1, 2, 5, 6, 11 and 24) (Fig. 3E; Supporting Information, Fig. S2E). One pair of 18S rDNA sites is located in the subterminal region of the chromosomes 1 and 2 of the hexavalent (Fig. 3F; Supporting Information, Fig. S2E). Cytotype IV (1 Ƌ; S1292) exhibits 2 n = 35 chromosomes, which decrease in length from 4.66% to 1.54% of DSL (Supporting Information, Fig. S2F; Table S2). The post-pachytene nuclei show a configuration of 13 bivalents, one trivalent (chromosomes 7, 26 and 35) and one hexavalent (chromosomes 1, 4, 5, 6, 10 and 23) (Fig. 3G; Supporting Information, Fig. S2F). One pair of 18S rDNA sites is located in the subterminal region of the chromosomes 1 and 4 involved in the hexavalent (Fig. 3H; Supporting Information, Fig. S2F)., Published as part of Just, Pavel, Šťáhlavský, František, Kovařík, František & Štundlová, Jana, 2022, Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae), pp. 885-901 in Zoological Journal of the Linnean Society 196 (2) on pages 892-893, DOI: 10.1093/zoolinnean/zlac049, http://zenodo.org/record/7184492

Simple Summary Whip spiders (Amblypygi) are spectacular arachnids characterized by powerful raptorial pedipalps and exceptionally elongated forelegs. Although cytogenetic data from amblypygids might be, given their phylogenetic position, important for the reconstruction of arachnid ka-ryotype evolution, cytogenetics of this order is still largely understudied. Here, we applied conventional and molecular cytogenetics to describe the karyotype patterns in Charinidae—the family placed almost at the base of amblypygid phylogeny, thereby providing insights into the ancestral amblypygid karyotype traits. We surveyed four Charinus and five Sarax species and found a wide range of diploid chromosome numbers (2n) in both genera. Representatives with lower 2n possessed generally a higher proportion of biarmed (metacentric and submetacentric) chromosomes. Our results indicate the important roles of chromosome fusions and pericentric inversions in the karyotype differentiation of Charinidae, similarly to that suggested previously for neoamblypygids. Our data, gathered from C-banding, fluorescent banding, and chromosomal mapping of ribosomal DNA and telomeric repeats, bring evidence for the action of these rearrangements and suggest the general trajectory towards 2n reduction from ancestral high chromosome counts. However, we cannot rule out the contribution of chromosome fissions. Abstract Whip spiders (Amblypygi) represent an ancient order of tetrapulmonate arachnids with a low diversity. Their cytogenetic data are confined to only a few reports. Here, we analyzed the family Charinidae, a lineage almost at the base of the amblypygids, providing an insight into the ancestral traits and basic trajectories of amblypygid karyotype evolution. We performed Giemsa staining, selected banding techniques, and detected 18S ribosomal DNA and telomeric repeats by fluorescence in situ hybridization in four Charinus and five Sarax species. Both genera exhibit a wide range of diploid chromosome numbers (2n = 42–76 and 22–74 for Charinus and Sarax, respectively). The 2n reduction was accompanied by an increase of proportion of biarmed elements. We further revealed a single NOR site (probably an ancestral condition for charinids), the presence of a (TTAGG)n telomeric motif localized mostly at the chromosome ends, and an absence of heteromorphic sex chromosomes. Our data collectively suggest a high pace of karyotype repatterning in amblypygids, with probably a high ancestral 2n and its subsequent gradual reduction by fusions, and the action of pericentric inversions, similarly to what has been proposed for neoamblypygids. The possible contribution of fissions to charinid karyotype repatterning, however, cannot be fully ruled out.

Additional file 19: Table S5. Smeringopinae, summary of male cytogenetic data, including results of other authors. Doubtful data are not included. See database [24] for full list of published data on pholcid karyotypes including doubtful data. Abbreviations: a = acrocentric, bi = biarmed, CP = chromosome pair, m = metacentric, n = number of plates evaluated, p = short chromosome arm, q = long chromosome arm, SC = sex chromosome, SCS = sex chromosome system, sm = submetacentric, ® = revision of data of other authors, st = subtelocentric, t = terminal, ? = unknown, *X = data of other authors (X = reference number).

Additional file 32: Table S6. Species studied, their instar, sex, collecting data and depositories. Abbreviations: AM = Australian Museum, Sydney, Australia (specimen KS 128687), Co. = county, Hwy = highway, Isl. = island, m = male, Mts. = mountains, N = north, N.P. = national park, NW = northwest, S = south, SL = specimens lost or discarded, sm = subadult male, W = west, ZFMK = Zoological Research Museum Alexander Koenig.

Additional file 7: Table S3. Modisiminae and Ninetinae, summary of male cytogenetic data, including results of other authors. Doubtful data are not included. See database [24] for full list of published data on pholcid karyotypes, including doubtful data. Abbreviations: bi = biarmed, CP = chromosome pair, m = metacentric, n = number of plates evaluated, p = short chromosome arm, q = long chromosome arm, SC = sex chromosome, SCS = sex chromosome system, sm = submetacentric, st = subtelocentric, t = terminal, ? = unknown, *X = data of other authors (X = reference number).

Additional file 28: Appendix S1. Evolution of particular chromosomes of the pholcid X1X2Y system.

Euscorpius sadileki sp. n. (Figures 115–149, 184, 186, Tables 3–4) http: //zoobank. org / urn: lsid: zoobank. org: act: 382FE1BA- 8307-406D-A493-6EC930BAA309 TYPE LOCALITY AND TYPE REPOSITORY. Serbia, Sokobanja, 43.63°N 21.80°E; FKCP. TYPE MATERIAL. Serbia, Sokobanja, 43.63°N 21.80°E, 14. VI.2014, leg. D. SadÍlek, 1♂ 1♀ (holotype and paratype, 543), FKCP; Nišava Province, Gadžin Han env., road between Donji Dušnik and Supotnica, Suva Planina foothills, 43.155ºN 22.118ºE, 5. VI.2014, 1♀ (paratype), leg. P. ŠÍpek et al., FKCP. ETYMOLOGY. The species epithet is a patronym honoring David SadÍlek, the collector of holotype of the new species. DIAGNOSIS (♂ ♀). Total length 26–36 mm. Color reddish black, legs and telson yellow. Pedipalp patellar external trichobothria numbers: 4 eb, 4 eb a, 2 esb, 4 em, 4 est, 6 et, ventral aspect of patella with 8–9 trichobothria. Pectinal teeth number 9 in male, 6–8 in females. Chelicerae yellow, with reticulation. Male with pedipalp fingers type C, female with fingers undulate. Dorsal metasomal carinae on III–IV segments granulated. Dorsolateral metasomal carinae on II–IV segments absent. Ventrolateral metasomal carinae on II–IV segments present and smooth or granulate (mainly in male). Metasoma II–IV densely (roughly) granulated including lateral surface. Metasoma V ventrally granulated with median carina present. Chela length/width ratio, 2.2 in male and 2.4–2.5 in females. Metasoma IV length/ width ratio, 1.68 in male and 1.8 in females. Telson length/ depth ratio, 2.6 in male and 3–3.1 in females. DESCRIPTION (♀ ♂). Total length about 26–36 mm in both sexes. The habitus is shown in Figs. 115–118. For position and distribution of trichobothria of pedipalps, see Figs. 141–147. Sexual dimorphism, see below in the description. Coloration (Figs. 115–118, 149). Base color uniformly reddish black, sternites lighter, yellowish brown, legs and telson yellow. Chelicerae yellow, reticulate. Carapace and mesosoma (Figs. 115–122). Carapace finely granulated with several smooth areas; carinae are absent. The anterior margin of the carapace is straight. Carapace with two lateral eyes. Tergites finely granulated, without developed carina. Tergite VII lacking lateral and median carinal pairs. Sternites III–VII rather smooth in females and sparsely finely granulated in male, both lacking lateral and median carinae. Stigmata small, narrow ellipical. Pectinal teeth number 9 in male and 6–8 (1 x 6, 1 x 7, 2 x 8) in females, fulcra are present. Pectines have three marginal lamellae and 4–6 middle lamellae. Metasoma and telson (Figs. 126–133). Metasoma very sparsely hirsute. Metasoma I–V granulated finely in females and roughly in male. Dorsal metasomal carinae on I–V segments distinctively irregularly granulated; dorsolateral carinae absent; ventrolateral carinae present, smooth in females and granulated in male. Metasoma V completely granulated in both sexes with median carina developed, on metasoma I–IV ventral median carina is absent. Anal arch with small pigmented granules. Telson finely granulated, elongate in females and swollen in males, with annular ring present in both sexes. Aculeus short more curved in male. Pedipalps (Figs. 134–148). Pedipalps very sparsely hirsute. Patella with 24 (4 eb, 4 eb a, 2 esb, 4 em, 4 est, 6 et) external and 8–9 (3 x 8, 1 x 9) ventral trichobothria. Chela with 4 ventral trichobothria, of which V 4 is located on external surface near Eb 1. Female paratype has additional trichobothrium eb a on chela external surface (Fig. 142), which is absent in male holotype. Femur and patella are granulated, femur roughly and patella finely. Femur has granulated developed carinae; ventroexternal carina is incomplete to missing. Patella with 5 complete carinae including irregular wide externomedian. DPS weak to moderate. Manus dorsally with fine, rounded granules, which do not form median carina; only five chelal carinae are developed. Male with pedipalp fingers type C, weakly undulate, similar ar females. Legs (Figs. 123–125). Both pedal spurs present on all legs, lacking spinelets; tibial spurs absent. Tarsus with single row of spinules on ventral surface, terminating distally with two essentially adjacent spinules. Measurements. See Table 3. AFFINITIES. The combination of four characters (pedipalp patellar external trichobothria numbers: 4 eb, 4 eb a; metasoma IV length/width ratio 1.6–1.8; male with pedipalp fingers type C and female with fingers undulate; and metasoma II– IV densely (roughly) granulated including lateral surface) is unique in the entire genus Euscorpius. DISTRIBUTION. Serbia (Fig. 184)., Published as part of Kovařík, František & Šťáhlavský, František, 2020, Five new species of Euscorpius Thorell, 1876 (Scorpiones: Euscorpiidae) from Albania, Greece, North Macedonia, and Serbia, pp. 1-37 in Euscorpius 315 on pages 22-28, DOI: 10.5281/zenodo.4648646

Compsobuthus Vachon, 1949 (Figures 1–224, Tables 1–4) Compsobuthus Vachon, 1949: 93 (1952: 213); Fet & Lowe, 2000: 124 (complete reference list until 1998); KovařÍk, 2003: 88 (in part); KovařÍk, 2009: 31; KovařÍk & Ojanguren-Affilastro, 2013: 145–158, figs. 777–941; KovařÍk et al., 2016a: 1–21, figs. 1–77; KovařÍk, 2018a: 1–11, figs. 1–39, tab. 1; KovařÍk, 2018b: 4–6, figs. 1–9. TYPE SPECIES. Buthus acutecarinatus Simon, 1882. DIAGNOSIS. Small to medium-sized buthid scorpions, total length 20–55 mm. Carapace with distinct carinae, central lateral and posterior lateral carinae connected to form continuous linear series of granules, projecting beyond posterior margin as distinct spiniform processes. Carapace in lateral view with entire dorsal surface horizontal, 5 pairs of lateral eyes. Cheliceral fixed finger with two ventral denticles. Sternum type 1, sub-triangular. Pectines with fulcra, pectine teeth number 9–34. Hemispermatophore flagelliform, capsule in ‘3+1’-lobe configuration (‘ Buthus ’ group; KovařÍk et al., 2016a), with 3 sperm hemiduct lobes well separated from flagellum, basal lobe a well-developed acuminate hook. Tergites I–VI with three carinae projecting beyond posterior margins as distinct spiniform processes. Sternites III–VI with slit-like spiracles. Orthobothriotaxic type A. Pedipalp femur with dorsal trichobothria arranged in β -configuration. Pedipalp patella with trichobothrium d 3 internal to dorsomedian carina; chela with db basal to est, eb located on fixed finger; chela manus with Eb 1–3 triad angled proximally (δ -configuration) or almost collinear (λ -configuration). Dentate margin of pedipalp chela movable finger with distinct granules divided into 8–14 linear rows, with 4 terminal granules and one basal terminal granule. Tibial spurs present on third and fourth pairs of legs. REMARKS ON KARYOTYPES. We analyzed karyotypes of four Compsobuthus species (Table 4). All examined specimens of C. acutecarinatus, C. arabicus, C. maindroni and C. ullrichi sp. n. possess 2n=22 (Figs. 210, 213, 216, 219, 222) and their chromosomes gradually decrease in length (Figs. 212, 215, 218, 221, 224, Table 4). This number of chromosomes was also previously found in C. eritreaensis KovařÍk, Lowe, PlÍškovÁ & ŠťÁhlavský, 2016 (KovařÍk et al., 2016a) and C. matthiesseni (Birula, 1905) (ŠťÁhlavský et al., 2014) (Table 4). This genus seems to have a conserved diploid number, similar to the genus Androctonus, which typically has 2n= 24 in all analyzed species (SadÍlek et al. 2015). The chromosomes of all examined Compsobuthus species also exhibit typical buthid features such as holocentric organization and achiasmatic meiosis in males (e. g. Mattos et al., 2013). We observed only bivalents in one male of C. acutecarinatus from the locality W of Qairoon Hairitti (Fig. 211) and in all males of C. maindroni (Fig. 220) during pachytene and postpachytene. However, we found also one quadrivalent in another two males of C. acutecarinatus from the localities Dhalkut beach and Wadi Ash Shuwaymiyyah (Fig. 214), in two males of C. arabicus from localities Jabal Bani Jabir and E of Wahiba (Fig. 217) and in the male hototype of C. ullrichi sp. n. (Fig. 223) during meiosis. Interestingly, the lengths of the chromosomes that form these multivalent associations are different among species (Figs. 215, 221, 224). This fact attests to the independent origin of these quadrivalents via reciprocal translocations that are very frequent chromosomal rearrangements within buthid scorpions (e. g. ŠťÁhlavský et al., 2014; Mattos et al., 2018)., Published as part of Kovařík, František, Lowe, Graeme, Stockmann, Mark & Šťáhlavský, František, 2020, Notes on Compsobuthus: redescription of C. arabicus Levy et al., 1973 from Arabia, and description of two new species from North Africa (Scorpiones: Buthidae), pp. 1-40 in Euscorpius 298 on page 3, DOI: 10.5281/zenodo.5741445, {"references":["VACHON, M. 1952. Etudes sur les scorpions. Institut Pasteur d'Algerie, Alger, 1 - 482. (published 1948 - 1951 in Archives de l'Institut Pasteur d'Algerie, 1948, 26: 25 - 90, 162 - 208, 288 - 316, 441 - 481. 1949, 27: 66 - 100, 134 - 169, 281 - 288, 334 - 396. 1950, 28: 152 - 216, 383 - 413. 1951, 29: 46 - 104).","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.","KOVARIK, F. 2003. Eight new species of Compsobuthus Vachon, 1949 from Africa and Asia (Scorpiones: Buthidae). Serket, 8 (3): 87 - 112.","KOVARIK, F. 2009. Illustrated catalog of scorpions. Part I. Introductory remarks; keys to families and genera; subfamily Scorpioninae with keys to Heterometrus and Pandinus species. Prague: Clairon Production, 170 pp.","KOVARIK, F. & A. A. OJANGUREN-AFFILASTRO. 2013. Illustrated catalog of scorpions. Part II. Bothriuridae; Chaerilidae; Buthidae I. Genera Compsobuthus, Hottentotta, Isometrus, Lychas, and Sassanidotus. Prague: Clairon Production, 400 pp.","KOVARIK, F., G. LOWE, J. PLISKOVA & F. STAHLAVSKY. 2016 a. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part VI. Compsobuthus Vachon, 1949 (Buthidae) with description of C. eritreaensis sp. n. Euscorpius, 226: 1 - 21.","KOVARIK, F. 2018 a. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part XVI. Compsobuthus maidensis sp. n. (Buthidae) from Somalia. Euscorpius, 260: 1 - 11.","KOVARIK, F. 2018 b. Notes on the genera Buthacus, Compsobuthus, and Lanzatus with several synonymies and corrections of published characters (Scorpiones: Buthidae). Euscorpius, 269: 1 - 12.","STAHLAVSKY, F., H. KOC & E. A. YAGMUR. 2014. The first record of karyotypes in Leiurus abdullahbayrami and Compsobuthus matthiesseni (Scorpiones: Buthidae) from Turkey. Northwestern Journal of Zoology, 10 (2): 355 - 358.","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.","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.","MATTOS, V. F., L. S. CARVALHO, M. A. CARVALHO & M. C. SCHNEIDER. 2018. Insights into the origin of the high variability of multivalent meiotic associations in holocentric chromosomes of Tityus (Archaeotityus) scorpions. PloS ONE 13, e 0192070."]}

Euscorpius kabateki sp. n. (Figures 80–114, 184, Table 2) http: //zoobank. org / urn: lsid: zoobank. org: act: A813698C- 63F9-40B2-B2EA-45DE501FF7D4 Euscorpius sicanus (in part): Parmakelis et al. 2013: fig. 1 (Locality 60), fig. 2–3 (FESP5). Euscorpius sicanus complex (in part): Fet et al., 2018: 129, fig. 3. TYPE LOCALITY AND TYPE REPOSITORY. Greece, Mt. Parnassos, Ski Center Parnassos, 38°32.4‘N 22°34.8‘E, 1800 m a. s. l.; FKCP. TYPE MATERIAL. Greece, Mt. Parnassós, Ski Center Parnassou, 38°32.4'N 22°34.8'E, 1800 m a. s. l., 2♂ 4♀1♀ juv. (holotype and paratypes), 5– 6. VI.2007, leg. P. Kabátek, 6. VI.2007, 2♀1♀ juv. (paratypes), leg. V. Ryjáček, FKCP. ETYMOLOGY. The species epithet is a patronym honoring Petr Kabátek, the collector of holotype of the new species. DIAGNOSIS (♂ ♀). Total length 28–37 mm. Color reddish yellow. Pedipalp patellar external trichobothria numbers: 5–6 eb, 4 eb a, 2 esb, 4 em, 4 est, 7 et, ventral aspect of patella with 9–11 trichobothria. Pectinal teeth number 9–10 in males, 7–9 in females. Chelicerae yellow, without reticulation. Males with pedipalp fingers type C, females with fingers undulate. Dorsal metasomal carinae on III–IV segments granulated. Dorsolateral metasomal carinae on II–IV segments absent. Ventrolateral metasomal carinae on II–IV segments present and smooth. Metasoma V ventrally with median carina indicated. Chela length/width ratio, 2.3 in males and 2.4–2.5 in females. Metasoma IV length/width ratio, 2.1–2.2 in males and 1.7–1.8 in females. Telson length/depth ratio, 2.2–2.3 in males and 3.2–3.3 in females. DESCRIPTION (♀ ♂). Total length about 28–37 mm in both sexes. The habitus is shown in Figs. 80–83. For position and distribution of trichobothria of pedipalps, see Figs. 107–113. Sexual dimorphism, see below in the description. Coloration (Figs. 80–83). Base color uniformly reddish yellow to orange. Chelicerae yellow, without reticulation. Carapace and mesosoma (Figs. 80–88). Carapace finely granulated with several smooth areas; carinae are absent. The anterior margin of the carapace is almost straight. Carapace with two lateral eyes. Tergites finely granulated, without developed carina. Tergite VII lacking lateral and median carinal pairs. Sternites III–VII smooth and lustrous, lacking lateral and median carinae. Stigmata small, narrow ellipical. Pectinal teeth number 9–10 (1 x 9, 3 x 10) in males and 7–9 (2 x 7, 3 x 8, 7 x 9) in females, fulcra are present. Pectines have three marginal lamellae and 4–5 middle lamellae. Metasoma and telson (Figs. 91–98). Metasoma very sparsely hirsute. Metasoma, mainly in dorsal aspect and whole metasoma V very finely granulated. Dorsal metasomal carinae on II–V segments distinctively irregularly granulated; dorsolateral carinae absent; ventrolateral carinae present, smooth on segments II–IV and granulated on segment V. Metasoma V ventrally granulated with median carina developed, on metasoma I–IV ventral median carina is absent. Anal arch with small pigmented granules. Telson smooth, elongate in females and swollen in males, with annular ring indicated in females and developed in male. Aculeus short more curved in male. Pedipalps (Figs. 99–114). Pedipalps very sparsely hirsute. Patella with 26–27 (5–6 eb [8 x 5, 12 x 6], 4 eb a, 2 esb, 4 em, 4 est, 7 et) external and 9–10 (2 x 9, 15 x 10, 3 x 11) ventral trichobothria. Chela with 4 ventral trichobothria, of which V 4 is located on external surface near Eb 1. Femur granulated; patella rather smooth. Femur has granulated developed carinae; ventroexternal carina is incomplete. Patella with 5 complete carinae including irregular wide externomedian. DPS well developed. Manus dorsally with fine solitary granules, almost absent in females; median carina missing, other carinae are smooth. Male with pedipalp fingers type C, female with fingers undulate. Legs (Figs. 89–90). Both pedal spurs present on all legs, lacking spinelets; tibial spurs absent. Tarsus with single row of spinules on ventral surface, terminating distally with two essentially adjacent spinules. Measurements. See Table 2. AFFINITIES. This species belongs to the subgenus Euscorpius Thorell, 1876, s. str.; it is morphologically similar to E. sicanus (C. L. Koch, 1837) from Italy (Sicily), which differs by having darker coloration. The presence of more than 4 (usually 5) trichobothria in eb series is a signature trait of a monophyletic ‘ Euscorpius sicanus complex’ (see Fet et al., 2003; Parmakelis et al., 2013). COMMENTS. Fet et al. (2018: 129) mentioned that “‘ Euscorpius sicanus complex’ representatives are found across Central Greece, including Mt. Parnassos”.After E. sicanus was recently restricted to its type locality of Sicily (Tropea, 2017), the Greek populations of this diverse complex remained unnamed. DNA data are available for this species in Parmakelis et al. (2013) under FESP5 (Locality 60; a specimen was collected by J. O. Rein in Lilaia, Mt. Parnassos). DISTRIBUTION. Greece, Mt. Parnassos (Fig. 184)., Published as part of Kovařík, František & Šťáhlavský, František, 2020, Five new species of Euscorpius Thorell, 1876 (Scorpiones: Euscorpiidae) from Albania, Greece, North Macedonia, and Serbia, pp. 1-37 in Euscorpius 315 on pages 16-22, DOI: 10.5281/zenodo.4648646, {"references":["PARMAKELIS, A., I. STATHI, P. KOTSAKIOZI, S. POULIKARAKOU & V. FET. 2013. Hidden diversity of Euscorpius (Scorpiones: Euscorpiidae) in Greece revealed by multilocus species-delimitation approaches. Biological Journal of the Linnean Society, 110: 728 - 748.","THORELL, T. 1876. On the classification of scorpions. Annals and Magazine of Natural History, 4 (17): 1 - 15.","TROPEA, G. 2017. Reconsideration of some populations of Euscorpius sicanus complex in Italy (Scorpiones: Euscorpiidae). Arachnida - Rivista Aracnologica Italiana, 11: 2 - 60."]}

Chaerilus Simon, 1877 (Figures 1���86, Table 1) http://zoobank.org/urn:lsid:zoobank.org:act:52 CAD 12D- 165A-470D-BD01-9589DC5EFCB6 Chaerilus Simon, 1877: 238; Kovař��k & Ojanguren, 2013: 131���145, figs. 617���776 (complete reference list until 2013); Kovař��k et al, 2015: 1���21, figs. 1���91, tables 1���3; Kovař��k et al., 2018: 1���27, figs. 1���125, tables 1���2; Kovař��k, 2019: 1���9, figs. 1���58, table 1. = Chelomachus Thorell, 1889: 583 (sYn. bY Kraepelin, 1899: 157). = Uromachus Pocock, 1890: 250 (sYn. bY Kraepelin, 1899: 157). TYPE SPECIES. Chaerilus variegatus Simon, 1877 DIAGNOSIS. Total length 16���80 mm; trichobothrial pattern orthobothriotaxic tYpe B; pedipalp femur d 3 ���d 4 trichobothrial axis angled toward dorsoexternal carina; patella with 3 ventral trichobothria, femur with 9 trichobothria (4 dorsal); cheliceral fixed finger with median and basal denticles flush on surface, not fused into bicusp; ventral margin of cheliceral movable finger crenulated, dorsal margin with single subdistal denticle; ventral surface of cheliceral fixed finger with 4���8 denticles; sternum, type 1, exhibiting subtle, wide horizontal compression; lateral eYes usuallY composed of 2 large major ocelli, eYespot present; maxillarY lobes I spatulate; hemispermatophore fusiform; ovariuterus 6-celled, embrYonic development apoikogenic; lateral lYmphoid organs absent; stigmata oval or round; median denticle row (MD) of pedipalp chela finger arranged in oblique groups; pedipalp chela with 8-carina configuration; legs without tibial spurs, but with prolateral and retrolateral pedal spurs; tarsi of legs with 2���4 rows of ventral setae and a median row of spinules; fifth metasomal segment with a single ventral carina; telson without subaculear tubercle; sclerotized cuticle of integument with weak or no autofluorescence emission under ultraviolet (UV) excitation., Published as part of Kova����k, Franti��ek, Lowe, Graeme, Stockmann, Mark & ������hlavsk��, Franti��ek, 2020, Two new Chaerilus from Thailand and Laos (Scorpiones: Chaerilidae), pp. 1-20 in Euscorpius 324 on page 1, DOI: 10.5281/zenodo.4648882, {"references":["SIMON, E. 1877. Etudes Arachnologiques. Part X. Arachnides nouveaux et peu connus. Annales de la Societe Entomologique de France, 5 (7): 225 - 242.","KOVARIK, F., G. LOWE, D. HOFEREK, M. FORMAN & J. KRAL. 2015. Two new Chaerilus from Vietnam (Scorpiones, Chaerilidae), with observations on growth and maturation of Chaerilus granulatus sp. n. and C. hofereki Kovarik et al., 2014. Euscorpius, 213: 1 - 21.","KOVARIK, F., G. LOWE & F. STAHLAVSKY. 2018. Three new Chaerilus from MalaYsia (Tioman Island) and Thailand (Scorpiones, Chaerilidae), with a review of C. cimrmani, C. sejnai and C. tichyi. Euscorpius, 268: 1 - 27.","KOVARIK, F. 2019. Chaerilus alberti sp. n. from MalaYsia (Scorpiones: Chaerilidae). Euscorpius, 274: 1 - 9.","THORELL, T. 1889. Viaggio di Leonardo Fea in Birmanie e regioni vicine. XXI. - Aracnidi Artrogastri Birmani raccolti da L. Fea nel 1885 - 1887. Annali del Museo Civico di Storia Naturale di Genova, 27: 521 - 729.","KRAEPELIN, K. 1899. Scorpiones und Pedipalpi. In Dahl, F. (ed.), Das Tierreich. Herausgegeben von der Deutschen Zoologischen Gesellschaft. Berlin: R. Friedlander und Sohn Verlag, 8. Lieferung, 265 pp.","POCOCK, R. I. 1890. Description of a new genus and species of scorpion belonging to the group Jurini. Annals and Magazine of Natural History, 6 (5): 250 - 252."]}

Reddyanus justi sp. n. (Figures 1–51, Table 1) http://zoobank.org/urn:lsid:zoobank.org:act:3FB403D2- AF00-426F-9C4B-109 D98750 FD0 TYPE LOCALITY AND TYPE REPOSITORY. Laos, Savannakhet Province, Phine District, 16º53'N 106º01'E; FKCP. TYPE MATERIAL. Laos, Savannakhet Province, Phine District, 16º53'N 106º01'E, March 2020, 1♂ (holotype, No. 1849) 1♂ 3♀ (paratypes), leg. M. Černička, FKCP. ETYMOLOGY. The specific epithet is a patronym honoring Pavel Just, an arachnologist who has assisted us greatly with our studies on scorpions in the laboratory as well as in the field during the past several years. DIAGNOSIS (♂ ♀). Adult length up to 33 mm (female) and 47 mm (male). Males with more elongate metasomal segments and telson compared to females. Pedipalps and legs yellow to orange, with reduced brown spots. Manus of pedipalps yellow, fingers reddish black. Metasomal segment I with 10 carinae, segments II–IV with 8 carinae. Metasoma V with 5 carinae, which are reduced in males. Posterior terminal tubercle on dorsolateral carinae of metasoma II–III of both seXes very slightly enlarged. Subaculear tooth broad, in females with terminus rounded, in males beveled and irregular quadrilateral in shape, dorsally with 5–7 granules in 3 rows. Pectinal tooth count 13–14 in males, 12–14 in females. DESCRIPTION.The adults are 45–47 mm (males) and 30–33 mm (females) long. Habitus as shown in Figs. 3–6. For position and distribution of trichobothria of pedipalps see Figs. 24–28 and Figs. 30–31. SeXual dimorphism: adult male with longer metasomal segments and longer, narrower telson. Coloration (Figs. 1–6). Base color yellow to orange with dark spots. Chelicerae are yellow, strongly reticulated. Fingers of chelicerae are darkly marked. Carapace and mesosoma patterned with 3 dark longitudinal strips. Ventral side of mesosoma yellow to yellowish brown, with several fuscous markings on sternite VII. Pedipalp femur and patella yellow or orange with small reduced dark maculae which are almost missing on femur. Legs also yellow or orange with black spots. Metasoma is yellowish or reddish brown with black spots, metasoma V darker than metasoma I–IV, telson yellow, red to reddish black. Carapace and mesosoma (Figs. 16–19). Entire carapace covered with large granules; carinae absent. Anterior margin of carapace is medially emarginate. Tergites I–VI with 1 median carina, tergite VII pentacarinate. All tegites strongly granulated. Pectinal tooth count 13–14 in males, 12–14 in females. Pectine marginal tips eXtending to posterior end of sternite III in males, to 3/4 of sternite III in females. Pectines with 3 marginal lamellae, 6–7 middle lamellae. Lamellae and fulcra bear numerous pale microsetae. Sternites smooth with 2 parallel furrows, eXcept sternite VII which bears 4 carinae and is granulated. Granulation also present on lateral margins of sternite VI. Posterior medial area of sternite V with a glabrous zone well developed in males, reduced in females. Metasoma and telson (Figs. 7–15). Metasomal segment I with 10 carinae, segments II–IV with eight carinae. Metasoma V with 5 carinae which are reduced in males. Vesicle of telson in males with a ventral carina, in females with ventral and lateral carinae. Metasoma with intercarinal surfaces sparsely granulated, with the eXception of metasoma V of males which is smooth. Posterior terminal tubercle on dorsolateral carinae of metasoma II–III of both seXes very slightly enlarged. Telson longer and narrower in males than females. Subaculear tubercle broad in lateral profile, with terminus rounded in females, beveled in males which have a tubercle with irregular quadrilateral shape. Tubercle dorsally with 5–7 granules in three rows. Pedipalps (Figs. 24–43). Pedipalps very sparsely hirsute. Femur with 5 granulated carinae, patella with 7 granulated carinae. Chela with 4–5 carinae which are granulated in both seXes and more strongly developed in females. Chela sparsely granulated, patella and femur are rather densely granulated eXcept ventrally. Chela movable finger with 6 rows of granules, fiXed finger with 7 rows; fingers bearing 6 eXternal accessory granules, 5–6 internal accessory granules. Legs (Figs. 20–23). Femora with 3–4 carinae, patellae with 5 carinae, both granulated. Legs without tibial spurs. Tibia and tarsomeres of legs with macrosetae not arranged serially as bristle combs on dorsal surfaces. Tarsomere II of leg IV with two sparse rows of Hemispermatophore (Figs. 44–48). Flagelliform, with flagellum long, narrow and coiled. Capsule relatively large, ca. half as long as trunk. Sperm hemiduct simple, undivided, long, with one prominent carina; posteriorly fused to base of flagellum. Basal lobe large, triangular in profile with sharp verteX. Trunk short, broad. Measurements: trunk length 2.5 mm, capsule length 1.12 mm, uncoiled flagellum length 3.89 mm. Karyotype (Figs. 49–50). Diploid number of these specimens was 12 chromosomes. The first pair of chromosomes was slightly longer (each chromosome from this pair forms 10.10 % of the diploid set), whereas the remaining chromosomes gradually decreased in length from 8.68 % to 7.32 % of the diploid set. Only bivalents without chiasmata were observed during male meiosis, and no centromeres on the chromosomes. These characteristics are typical for scorpions in the family Buthidae (e. g. Mattos et al., 2013) and the lower number of chromosomes appears to be typical of the genus Reddyanus (2n=11–17) (KovařÍk & ŠťÁhlavský, 2019). Measurements. See Table 1. AFFINITIES. The described features distinguish Reddyanus justi sp. n. from all other species of the genus. The new species is characterized by the irregular quadrilateral shape in lateral profile of the subaculear tubercle in males, which is a unique character within the genus (Fig. 9 versus figs. 209–214 in KovařÍk & ŠťÁhlavský, 2019: 40). The morphologically closest species is R. petrzelkai (KovařÍk, 2003) (see also a key in KovařÍk & ŠťÁhlavský, 2019: 43) from Vietnam which has a large subaculear tubercle that is rounded (Fig. 9 versus fig. 210 in KovařÍk & ŠťÁhlavský, 2019: 40) and stronger dark maculation on its pedipalp segments (Figs. 24–30 versus fig. 199 in KovařÍk & ŠťÁhlavský, 2019: 39). REMARKS. The capsule structures and overall dimensions of the hemispermatophore of Reddyanus justi sp. n. are similar to those recorded for other species of the genus (i. e., R. basilicus (Karsch, 1879), R. ceylonensis KovařÍk et al., 2016 and R. loebli (Vachon, 1982); cf. KovařÍk et al, 2016). These species also eXhibit a simple, monocarinate sperm hemiduct, large triangular basal lobe and long, coiled flagellum. This provides additional support for our hypothesis that these features may be diagnostic for the genus (KovařÍk et al, 2016), although the hemispermatophores of other Reddyanus species have yet to be studied. DISTRIBUTION. Laos (Fig. 51).

Compsobuthus turieli sp. n. (Figures 110–163, Table 1) http://zoobank.org/urn:lsid:zoobank.org:act:1BCCBD64- 4FB0-45ED-B1E1-0C1A6DC01E50 TYPE LOCALITY AND TYPE DEPOSITORY. Western Sahara, 70 km S of Ad-Dakhla, 23°04.13'N 016°05.08'W; FKCP. TYPE MATERIAL EXAMINED. Western Sahara: 70 km S of Ad- Dakhla, 23°04.13'N 016°05.08'W, 7.II.2005, 1♂ (holotype) 1juv. (paratype), leg. R. et H. Fouquè & S. BečvÁř. Morocco: At Akka, 29.390083°N 8.268220°W, IX.2013, dry, barely vegetated farmland nr Qued Akka, collected by day under rock left from road construction, 1♀ (paratype), leg. C. Turiel and M. Stockman, FKCP; N of Zag, 28.24872°N 09.33291°W (Fig. 156), X.2016, in sandy flats with large black rocks, loose cover of shrubs and solitary trees, collected by day under rocks and UV detection by night sitting on rock, 2♀ 1juv. (paratypes), leg. M. Stockmann, FKCP. ETYMOLOGY. It is a pleasure to name this species after Carlos Turiel (Neuss, Germany). DIAGNOSIS. Total length 29–31 mm. Sexual dimorphism minor, pedipalp fingers straight in females, almost straight in male holotype, male chela slightly more robust, chela L/W ratio: ♂ 5.39, ♀ 5.59; metasomal segment proportions similar in both sexes. Base color uniformly yellow to yellowish brown. Carapace and tergites densely, finely granular. Anterior margin of carapace bearing 8 symmetrically distributed spinules. Pedipalp femur L/ Carapace L ratio: 0.86–0.93. Movable finger of pedipalp chela with 9–11 rows of granules, without external accessory denticles, with 10 internal accessory granules (‘ acutecarinatus ’ group; Levy & Amitai, 1980). Manus of pedipalp chela shorter than fixed finger. Pedipalp chela L/movable finger L ratio: 1.38–1.43. Metasoma I–II with 10 carinae, III with 8–10 carinae, IV with 8 carinae. All metasomal segments longer than wide; metasoma L/W ratios: III 1.85–1.92, IV 2.20–2.27, V 2.50–2.68. Metasoma V W/D ratio: 1.07–1.09. Ventral intercarinal surfaces of metasoma lacking macrosetae. Pectine teeth: ♂ 16, ♀ 13–14. Pectine L/ Metasoma V W ratio: ♂ 2.00, ♀ 1.46. Sternites and metasoma granulated, more strongly in females. Sternites VI–VII with 4 crenulate carinae. Telson rather elongate, aculeus shorter than vesicle. Subaculear tubercle absent. DESCRIPTION. Total length 29–31 mm in both sexes. The habitus is shown in Figs. 110–113. Trichobothriotaxy of pedipalps is shown in Figs. 135–142. Sexual dimorphism. Sex differences are minor, fingers of pedipalps straight in females and almost straight in the male holotype. No sex differences in proportions of metasomal segments. Female with considerably shorter pectines, smaller pectine teeth and larger genital opercula. Coloration (Figs. 110–113, 157–159). The base color is uniformly yellow to yellowish brown. Weak fuscosity may be present on anterior interocular triangle. Carapace and mesosoma (Figs. 123–126). The entire carapace is covered by granules of different sizes. The carinae are moderately to strongly developed and granular. The anterior margin of the carapace is weakly concave medially, and bears 8 symmetrically distributed spinules (macrosetae). The tergites are strongly granulated. Tergites I–VI are tricarinate, with strong, denticulate median and lateral carinae. Each carina terminates in a spiniform process that in the lateral carinae extends well past the posterior margin of the tergite. Tergite VII is pentacarinate, with lateral pairs of carinae strong, serratocrenulate, and median pairs moderate, crenulate; the median carina is weak and confined to the anterior half of the segment. Pectinal tooth counts: ♂ 16, ♀ 13–14 (3×13, 3×14). The pectine marginal tips extend barely to the posterior margin of sternite III in females, and to half the length of sternite IV in the male. The pectines have 3 marginal lamellae and 7–9 middle lamellae. The lamellae bear numerous dark setae, and each fulcrum bears 2–3 dark setae. All sternites are granulated. The posterior areas of sternites lack a broad glabrous patch. Sternites VI–VII bear 4 crenulate carinae, which are more strongly developed on VII. The other sternites bear one pair of carinae on the medial side of the spiracles. Metasoma and telson (Figs. 114–122). Metasomal segments I–II with 10 carinae, III with 8–10 carinae, IV with 8 carinae, and V with 5 carinae. Median lateral carinae of metasoma III are indicated by isolated granules that may partially form carinae. All segments sparsely setose and densely granulate, more so in females. Accessory rows of granules are present on dorsal surfaces of segments as well as on the ventral surface of the fifth segment. The telson is rather elongate, with the aculeus a little shorter than the vesicle. A subaculear tubercle is absent. Pedipalps (Figs. 135–155). The pedipalps are finely granulated and sparsely hirsute. The femur bears 5 carinae, the patella 7 granular carinae, the chela 5–7 carinae. The movable and fixed fingers bear 9–11 rows of granules, without external accessory granules, with 10 internal accessory granules on movable finger, 8–10 on fixed finger. Pedipalp chela L/W ratio: ♂ 5.39, ♀ 5.59. Manus of chela shorter than fixed finger. Pedipalp chela L/movable finger L ratio 1.38–1.43 in both sexes. Legs (Figs. 127–134). Legs III–IV bear small to moderate tibial spurs. Retrolateral and prolateral pedal spurs are present on all legs. The tarsomeres bear two rows of macrosetae on the ventral surface and several macrosetae on the other surfaces. Bristlecombs are absent. The femur bears 4 carinae, the patella 4–6 carinae. The femur and patella bear only solitary macrosetae and are granulated on prolateral surfaces, smooth on retrolateral surfaces. Tarsal ungues moderately elongated, curved. Hemispermatophore (Figs. 160–163). Flagelliform, elongate and slender, trunk ca. 7.7 times length of capsule region. Flagellum separated from external lobe, pars recta ca. 60% of trunk length, broad with anterior lamina, pars reflecta long, about the same length as trunk, thick, hyaline. Capsule region with 4 lobes at base of flagellum: posterior lobe longest, triangular, apically rounded, median lobe shortest, apically truncate, anterior lobe acuminate with long thin terminus. Basal lobe strong with broad base and sharp, falcate hook. Left and right hemispermatophores were similar. Measurements. See Table 1. AFFINITIES. The described features distinguish C. turieli sp. n. from all other species of the genus. In the region, the only other known member of the ‘ acutecarinatus group’ is C. berlandi Vachon, 1950, from Mauritania (type locality: Fort Gouraud, 400 km E of Villa-Cisneros) which is characterized by the presence of 7–8 rows of granules on the pedipalp movable finger, and total length 38 mm (Vachon, 1950). In contrast, C. turieli sp. n. has 9–11 rows of granules on the pedipalp movable finger, and total length ECOLOGY. Specimens were found under rocks, or on rocks at night, in arid sandy terrain with rather sparse vegetation. Water was available only in an oasis or date plantations, or transiently (a few hours) after a heavy storm. Loamy soil was present but C. turieli sp. n. was only found in sandy areas. Other scorpions observed together with C. turieli sp. n. were: in sandy areas, Androctonus amoreuxi (Audouin,1826), Buthacus stockmanni KovařÍk, Lowe & ŠťÁhlavský, 2016 and Lissothus occidentalis Vachon, 1950; in loamy areas, Buthus mariefranceae Lourenço, 2003.

Chaerilus kautti sp. n. (Figures 1–50, 84, Table 1) http://zoobank.org/urn:lsid:zoobank.org:act:8275997C- 69A8-4494-A6F1-ED502A69E1FF TYPE LOCALITY AND TYPE REPOSITORY. Thailand, Prachuap Khiri Khan Province, Khao Ma Rong Cave, 11.2021900°N 99.4946250°E, 56 m a. s. l., FKCP. TYPE MATERIAL (FKCP). Thailand, Prachuap Khiri Khan Province, Khao Ma Rong Cave, 11.2021900°N 99.4946250°E, 56 m a. s. l. (Fig. 48), 19.VIII.2018, 1♂ (holotYpe, 1770, Figs. 1, 3–4, 7–8, 15, 20–32, 44–47), leg. Peter Kautt, 1♀ (paratYpe, Figs. 2, 5–6, 9–14, 16–19, 33– 43, 49–50, scorpion born 27.IX.2020), leg. Britta & Mark Stockmann, 16 juvs. (paratYpes, Figs. 49–50, offspring of the female paratYpe 27.IX.2020, still alive), bred bY F. Kovařík and M. Stockmann. ETYMOLOGY. The specific epithet is a patronYm honoring Peter Kautt, the collector of the male holotYpe of the new species. DIAGNOSIS (♂ ♀). Total length 37–43 mm; two pairs of lateral eYes and one pair of median eYes; ventral surface of cheliceral fixed finger with 4 denticles; male differs from female in having pedipalp chela much narrower; chela length/width ratio ♂ 4.55, ♀ 2.88; movable finger of pedipalp with 11 imbricated rows of granules; fingers straight in both sexes; pedipalp chela with 7–8 carinae; pectinal tooth count ♂ 6, ♀ 4; carapace granulated sparselY in male, denselY in female; mesosomal tergites granulated, mainlY in the female; all sternites smooth without carinae and granules; metasoma I–III with 6 or 8 carinae, ventral carinae maY be absent. DESCRIPTION. Total length ♂ 37 mm, ♀ 43 mm; color reddish orange to brown, maculose; chelicerae (Figs. 42–43) sparselY granulated dorsallY, Yellow and stronglY reticulate, anteriorlY black; male differs from female in having pedipalp chela much narrower with short fingers; chela length/width ratio ♂ 4.55, ♀ 2.88; male with larger pectines (Figs. 8 and 10); no sexual dimorphism in shape of metasoma and telson; trichobothrial pattern as shown in Figs. 23–29, measurements in Table 1. Carapace and mesosoma (Figs. 7–10). Carapace covered bY large granules in female, almost smooth in male; anterior margin of carapace weaklY concave; two well developed pairs of lateral eYes and one pair of median eYes present; mesosomal tergites irregularlY granulated in female, almost smooth in male; all sternites smooth without carinae and granulation; sternite V with smooth patch present; pectinal tooth count ♂ 6, ♀ 4. Metasoma and telson (Figs. 15–22). Metasoma I–II with 6 or 8 incomplete carinae, ventral carinae reduced or absent; metasoma IV with 8 carinae, metasoma V with 5 carinae; all carinae composed of sparse, large granules; intercarinal surfaces sparselY, irregularlY granulated, more so on lateral surfaces of all segments, and ventral surfaces of segment V; metasomal segments I–III ventrallY smooth; all segments verY sparselY hirsute; telson elongate, ampullate, smooth, verY sparselY hirsute. Pedipalps (Figs. 23–41). Pedipalp chela elongated in male, rather stout in female; movable finger with 11 and fixed finger with 10 imbricated rows of granules; chela with 7 or 8 carinae, smooth in male, partlY finelY granulated in female; carina on dorsoexternal surface of the manus maY be incomplete; dorsal and internal surfaces of chela with reticulate granulation patterns; patella with 5–6 smooth carinae, surfaces of patella smooth except internal surface which is finelY granulated; femur denselY granulated with 4 carinae. Legs (Figs. 11–14). Hirsute, without bristlecombs and carinae; femora and patellae granulated dorsallY, other surfaces smooth; tarsomeres with 4 rows of spiniform setae; spiniform seta formula of inner rows 6–7/7–8: 7–8/6–7: 7–8/8–9: 6–8/8–9; of outer rows 3–4/3–4 on all legs. Hemispermatophore (Figs. 44–47). Fusiform; distal lamina short, broad, weaklY tapered, apex rounded; capsule with distal carina weaklY sclerotized; sperm hemiduct delineated bY two thin, sclerotized carinae, of which the distal is longer, the basal shorter; trunk long, as broad as capsule. Measurements of right hemispermatophore (mm): distal lamina length 0.64, capsule length 0.72, trunk length 1.96, pedicel length 0.42, capsule width 0.63. Left hemispermatophore with shorter trunk, length 1.70. Karyotype (Fig. 84). Male holotYpe was analYzed. During metaphase I 59 bivalents were observed (Fig. 84), implYing that 2n=118. Chiasmata were not observed during meiosis. AFFINITIES. Chaerilus kautti sp. n. is reliablY distinguished from all other Chaerilus species bY the following unique combination of two characters: movable pedipalp finger with 11 imbricated granule rows; and pedipalp chela length/width ratio 4.55 in the male. A movable pedipalp finger with 11 (or 10–11) imbricated granule rows is also found in six other species of the genus (C. cimrmani Kovařík, 2012 from Thailand, C. hofereki Kovařík, 2014 from Vietnam, C. julietteae Lourenço, 2011 from Vietnam, C. neradorum Kovařík et al., 2018 from Thailand, C. robinsoni Hirst, 1911 from MalaYsia and Indonesia, and C. stockmannorum Kovařík et al., 2018 from Thailand). However, males of all of these species have a pedipalp chela length/width ratio between 1.84 (C. hofereki) and 3.13 (C. julietteae). COMMENTS ON LOCALITIES AND LIFE STRATEGY. The tYpe localitY is located on an isolated karst mountain close to the eastern coast of Thailand, west of Bang Saphan. The entire mountain is clothed in primarY forest, but is surrounded bY cultivated farmland with mostlY oil palms and gumtrees. The tYpe localitY is close to two extensive caves, but C. kautti sp. n. does not appear to be associated with them. Climate conditions are warm and verY wet during the wet season from MaY to November, and slightlY cooler and drier during the drY season from December to April. Even during the drY season, the area is verY humid and water is alwaYs present. The male from 2018 was collected during heavY rain on a dark night, the female from 2019 on a drY night without wind during a new moon. The average temperatures on both nights were ca. 25–26°C with verY high humiditY. Both specimens were found at night on the ground, between leaf litter or sitting openlY on rocks. In captivitY, the scorpions concealed themselves under bark or in moss. The probablY do not excavate burrows, but instead shelter between rocks, or in wood and leaf litter in their habitat. Other scorpions observed in this habitat were Heterometrus cimrmani Kovařík, 2004, Lychas scutilus Koch 1845, and Liocheles australasiae Fabricius, 1775. An adult gravid female collected in 2019 gave birth in captivitY to 48 juveniles after 11 months.

Kova����k, Franti��ek, Lowe, Graeme, ������hlavsk��, Franti��ek (2020): Reddyanus justi sp n from Laos (Scorpiones Buthidae). Euscorpius 321: 1-11, DOI: http://doi.org/10.5281/zenodo.4648807, {"references":["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.","KOVARIK, F. 2003. A review of the genus Isometrus Ehrenberg, 1828 (Scorpiones: Buthidae) with descriptions of four new species from Asia and Australia. Euscorpius, 10: 1-19.","KOVARIK, F. 2009. Illustrated catalog of scorpions. Part I. Introductory remarks; keys to families and genera; subfamily Scorpioninae with keys to Heterometrus and Pandinus species. Prague: Clairon Production, 170 pp.","KOVARIK, F. G. LOWE, K. B. RANAWANA, D. HOFEREK, V. A. SANJEEWA JAYARATHNE, J. PLISKOVA & F. STAHLAVSKY 2016. Scorpions of Sri Lanka (Arachnida, Scorpiones: Buthidae, Chaerilidae, Scorpionidae) with description of four new species of the genera Charmus Karsch, 1879 and Reddyanus Vachon, 1972 stat. n. Euscorpius, 220: 1-133.","KOVARIK, F. & A. A. OJANGUREN AFFILASTRO. 2013. Illustrated catalog of scorpions. Part II. Bothriuridae; Chaerilidae; Buthidae I. Genera Compsobuthus, Hottentotta, Isometrus, Lychas, and Sassanidotus. Prague: Clairon Production, 400 pp.","KOVARIK, F. & STAHLAVSKY, F. 2019. Revision of the genus Reddyanus from Southeast Asia, with description of five new species from Cambodia, Malaysia, Thailand and Vietnam (Scorpiones: Buthidae). Euscorpius, 295: 1-45.","KOVARIK, F., F. STAHLAVSKY, T. KORINKOVA, J. KRAL & T. VAN DER ENDE. 2009. Tityus ythieri Lourenco, 2007 is a synonym of Tityus magnimanus Pocock, 1897 (Scorpiones: Buthidae): a combined approach using morphology, hybridization eXperiments, chromosomes, and mitochondrial DNA. Euscorpius, 7: 1-12.","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.","SAKAMOTO, Y. & A.A. ZACARO. 2009. LEVAN, an ImageJ plugin for morphological cytogenetic analysis of mitotic and meiotic chromosomes. Available at: http:// rsbweb.nih.gov/ij/plugins/levan/levan.html. Accessed 3rd June 2016.","SOLEGLAD, M. E. & V. FET. 2003. The scorpion sternum: structure and phylogeny (Scorpiones: Orthosterni). Euscorpius, 5: 1-34.","STAHNKE, H. L. 1971. Scorpion nomenclature and mensuration. Entomological News, 81: 297-316.","TIKADER, B. K. & D. B. BASTAWADE. 1983. Scorpions (Scorpionida: Arachnida). In The Fauna of India, Vol. 3. (Edited by the Director). Calcutta: Zoological Survey of India, 671 pp.","VACHON, M. 1972. Remarques sur les scorpions appartenant au genre Isometrus H. et E. (Buthidae) A propos de l`espece Isometrus maculatus (Geer) habitant l'ile de PAques. Cahiers Pacifique, 16: 169-180.","VACHON, M. 1974. Etudes des caracteres utilises pour classer les familles et les genres des scorpions (Arachnides). 1. La trichobothriotaXie en arachnologie. Sigles trichobothriauX et types de trichobothriotaXie chez les Scorpions. Bulletin du Museum national d'Histoire naturelle, 3e serie, 140 (Zoologie, 104): 857-958.","VACHON, M. 1982. Les scorpions de Sri Lanka (Recherches sur les scorpions appartenant ou deposes au Museum d`Histoire naturelle de Geneve III.). Revue suisse de Zoologie, 89: 77-114."]}

Compsobuthus ullrichi sp. n. (Figures 164–209, 222–224, Tables 3–4) http://zoobank.org/urn:lsid:zoobank.org:act:7EEF1 EA5294-4BAB-B92D-9F568946A2F4 TYPE LOCALITY AND TYPE DEPOSITORY. Egypt, DahÁb, 28°29'N 34°30'E; FKCP. TYPE MATERIAL EXAMINED. Egypt: DahÁb, 28°29'N 34°30'E, II.2019, 1♂ (holotype, 1597), leg. A. Ullrich, FKCP. ETYMOLOGY. It is a pleasure to name this species after Alex Ullrich (Germany), collector of the male holotype. DIAGNOSIS (♂). Total length 28 mm. Male pedipalp fingers with dentate margins proximally undulate, chela L/W ratio: 4.75. Base color uniform yellow to yellowish brown with dark spot on fifth metasomal segment. Carapace and tergites finely granular with some larger granules. Anterior margin of carapace bearing 8 symmetrically distributed spinules. Pedipalp femur L/ Carapace L ratio: ♂ 0.95. Movable finger of pedipalp chela with 10–11 rows of granules, all rows with external and internal accessory denticles (‘ werneri ’ group; Levy & Amitai, 1980). Manus of pedipalp chela shorter than fixed finger. Pedipalp chela L/movable finger L ratio: ♂ 1.47. Metasoma I–II with 10 carinae, III–IV with 8 carinae. All metasomal segments longer than wide; metasoma L/W ratios ♂: III 1.94, IV 2.25, V 2.80. Metasoma V W/D ratio ♂: 0.945. Ventral intercarinal surfaces of metasoma with macrosetae. Pectine teeth: ♂ 22–23. Pectine L/ Metasoma V W ratio: ♂ 2.31. Sternites smooth medially and finely granulated marginally; metasoma granulated. Sternites VI–VII with 4 crenulate carinae. Telson rather bulbous, aculeus shorter than vesicle. Subaculear tubercle weakly developed. DESCRIPTION (♂ HOLOTYPE). Total length 28.13 mm. Female unknown. The habitus is shown in Figs. 168–169. Trichobothriotaxy of pedipalps is shown in Figs. 190–197. Male with pedipalp fingers proximally undulate. Coloration (Figs. 168–169). The base color is uniform yellow to yellowish brown, with fuscosity on the anterior half of metasoma V. Carapace and mesosoma (Figs. 180–181). The carapace is covered by granules of different sizes. The carinae are moderately to strongly developed and granular. The anterior margin of the carapace is weakly concave medially, and bears 8 symmetrically distributed spinules (macrosetae). The tergites are partially, coarsely granulated. Tergites I–VI are tricarinate, with strong, denticulate median and lateral carinae. Each carina terminates in a spiniform process that in the lateral carinae extends well past the posterior margin of the tergite. Tergite VII is pentacarinate, with lateral pairs of carinae strong, serratocrenulate, median pairs moderate, crenulate; median carina is weak and confined to the anterior half of the segment. Pectinal tooth count: ♂ 22–23. The pectine marginal tips extend to the posterior margin of sternite IV. The pectines have 3 marginal lamellae and 7–9 middle lamellae. The lamellae bear numerous dark setae, and each fulcrum bears 2–3 dark setae. Sternites are smooth medially and finely granulated marginally. The posterior areas of sternites lack a broad glabrous patch. Sternites VI–VII bear 4 crenulate carinae which are more strongly granulated on VII. The other sternites bear one pair of carinae on the medial side of the spiracles. Metasoma and telson (Figs. 173, 177–179). Metasomal segments I–II with 10 carinae, III–IV with 8 carinae, and V with 5 carinae. Median lateral carinae of metasoma III reduced and indicated only by several granules posteriorly. All segments sparsely setose and granulate. Accessory rows of granules are present on dorsal surfaces of segments as well as on the ventral surface of segment V. The telson is rather bulbous, with the aculeus a little shorter than the vesicle. A subaculear tubercle is indicated. Pedipalps (Figs. 190–200). The pedipalps are smooth, partly finely granulated and sparsely hirsute. The femur bears 5 carinae, the patella 7 granular carinae, the chela 5 rather smooth carinae. The movable and fixed fingers bear 10–11 rows of granules, all with external and internal accessory granules. Pedipalp chela L/W ratio: ♂ 4.75. Manus of chela shorter than fixed finger. Pedipalp chela L/movable finger L ratio: ♂ 1.47. . Legs (Figs. 184–187). Legs III–IV bear moderate tibial spurs. Retrolateral and prolateral pedal spurs are present on all legs. The tarsomeres bear two rows of macrosetae on the ventral surface and several macrosetae on the other surfaces. Bristlecombs are absent. The femur bears 4 carinae, the patella 4–6 six carinae. The femur and patella bear only solitary macrosetae and are granulated except for external lateral surfaces which are smooth. Tarsal ungues moderately elongated, curved. Hemispermatophore (Figs. 164–167). Flagelliform, elongate and slender, trunk ca. 6 times length of capsule region. Flagellum separated from external lobe, pars recta ca. 50% of trunk length, broad with anterior lamina, pars reflecta ca. 75% of trunk length, narrow, hyaline. Capsule region with 4 lobes at base of flagellum: posterior lobe longest, triangular, median lobe shortest, apically truncate, anterior lobe acuminate with long thin terminus. Basal lobe strong with broad base and sharp, falcate hook. Left and right hemispermatophores were similar. Measurements. See Table 3. AFFINITIES. The described characters distinguish C. ullrichi sp. n. from all other known species of the genus. Among Compsobuthus species known from the region of Middle East and Egypt, C. ullrichi sp. n. is most similar to C. levyi KovařÍk, 2012, from Jordan (type locality: near Qasr Burqu). The two species can be separated by several characters: males of C. levyi have longer and narrower pedipalp and metasomal segments (see Table 1 and Figs. 174–179). Pedipalp patella L/W ratio is 3.06 in male holotype C. ullrichi, 3.25–3.26 in males of C. levyi; metasoma I L/W ratio is 1.377 in male C. ullrichi, 1.51–1.54 in males of C. levyi; and metasoma IV L/W ratio is 2.24 in male C. ullrichi, 2.46–2.48 in males of C. levyi. In this region are found five other species of Compsobuthus belonging to the ‘ werneri ’ group of Levy & Amitai (1980). From these, C. ullrichi sp. n. can be clearly differentiated by metasomal segment I being narrower than that of other species: C. carmelitis Levy, Amitai & Shulov, 1973 from Israel, and probably also Jordan and Syria, has metasoma I of males wider than long, or as wide as long; C. egyptiensis Lourenco et al., 2009 from Egypt (type locality: NW of Siwa) has metasoma I L/W ratio 1.1–1.17 in males; and C. kabateki KovařÍk, 2003 from Egypt (Luxor) has metasoma I L/W ratio 1.07 in males. In contrast, C. ullrichi sp. n. has metasoma I L/W ratio 1.37. Compsobuthus longipalpis Levy, Amitai & Shulov, 1973 from Egypt, Israel, Jordan, Saudi Arabia, and Syria, differs in having 14 rows of granules on the pedipalp movable finger and a total length of 40–50 mm (in C. ullrichi sp. n. there are 10–11 rows of granules on the pedipalp movable finger and total length is C. schmiedeknechti Vachon, 1949 from Israel, Jordan, Lebanon, Syria, and Turkey differs in having 15–18 pectinal teeth in males; C. kabateki KovařÍk, 2003 from Egypt (Luxor) differs in having 19 pectinal teeth in males (in C. ullrichi sp. n. there are 22–23 pectinal teeth in the male holotype)., Published as part of Kovařík, František, Lowe, Graeme, Stockmann, Mark & Šťáhlavský, František, 2020, Notes on Compsobuthus: redescription of C. arabicus Levy et al., 1973 from Arabia, and description of two new species from North Africa (Scorpiones: Buthidae), pp. 1-40 in Euscorpius 298 on pages 33-38, DOI: 10.5281/zenodo.5741445, {"references":["KOVARIK, F., G. LOWE, J. PLISKOVA & F. STAHLAVSKY. 2016 a. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part VI. Compsobuthus Vachon, 1949 (Buthidae) with description of C. eritreaensis sp. n. Euscorpius, 226: 1 - 21.","STAHLAVSKY, F., H. KOC & E. A. YAGMUR. 2014. The first record of karyotypes in Leiurus abdullahbayrami and Compsobuthus matthiesseni (Scorpiones: Buthidae) from Turkey. Northwestern Journal of Zoology, 10 (2): 355 - 358.","KOVARIK, F. 2012. Three new species of Compsobuthus Vachon, 1949 from Yemen, Jordan, Israel, and Somaliland (Scorpiones: Buthidae). Euscorpius, 150: 1 - 10.","KOVARIK, F. 2003. Eight new species of Compsobuthus Vachon, 1949 from Africa and Asia (Scorpiones: Buthidae). Serket, 8 (3): 87 - 112."]}

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