Anomalobuthus lowei Teruel, Kovařík et Fet, 2018

Teruel, Rolando, Kovařík, František & Fet, Victor, 2018, Revision of the Central Asian scorpion genus Anomalobuthus Kraepelin, 1900, with descriptions of three new species and a generic synonymy (Scorpiones: Buthidae), Euscorpius 270, pp. 1-45 : 20-28

Anomalobuthus lowei Teruel, Kovařík et Fet , sp. n.

Figures 65–78, 141; Tables 3–5

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5C-4E36-4D73-ABCC-DCCE1A6E9DB9

Anomalobuthus rickmersi: Gromov & Kopdykbaev, 1994: 20 (in part); Gromov, 2005: 39 (in part); Gromov & Kazenas, 2006: 39 (in part); Mityaev et al., 2005: 39 (in part); Gromov, 2006: 44–45 (in part); Kamenz & Prendini, 2008: 6, 8, 40, 56–57, fig. 15a, tabs. 1–2.

HOLOTYPE ♀ ( GLPC). Kazakhstan, Almaty Province, Kapchagay [43°53'N, 77°05'E], Ili River , 75 km north of Almaty, 17 May 1993, leg. A. Feodorov.

ETYMOLOGY. We are pleased to name this species after our friend and colleague, Graeme Lowe (Philadelphia, USA), who contributed greatly to the knowledge of the Palearctic desert scorpions. His assistance to the present paper was crucial, by selflessly spending his scarce free time in exhausting photographic sessions providing all the images of the holotype of A. lowei ( Figs. 65–78).

DIAGNOSIS (based on one adult female). Adult size 35 mm. Coloration yellow, essentially immaculate, only with some irregular blackish spots on carapace, tergites, pedipalps, legs, and metasoma; metasomal segment V deeply infuscate, telson blackish. Pedipalp fingers with 7–8 principal rows of denticles and 6–7 internal accessory denticles. Pectines with 21/20 teeth. Tibial spur on leg IV only, small. Metasoma robust, with most carinae moderately developed; ventral lateral carinae of segment V composed of narrowly lobate denticles, dorsal lateral and lateral supramedian carinae of segments I–IV with terminal denticle greatly enlarged; intercarinal spaces essentially smooth on dorsal areas of all segments, densely granulose on ventral and lateral areas of I and V, sparsely granulose on ventral and lateral areas of II–IV. Telson vesicle short oval and sparsely setose.

DESCRIPTION (adult female holotype). Coloration ( Fig. 65) base light yellow, somewhat translucent due to non-optimal preservation. Chelicerae immaculate, except for blackish finger teeth. Pedipalp femur with a conspicuous blackish spot on dorsodistal apex, which diffusely continues as a broad stripe over almost all internal surface; patella with internal surface blackish; chela with internal surface diffusely reticulate with dark brown, finger denticles blackish. Carapace with a large blackish spot that widely surrounds median ocular tubercle and extends to anterior margin and also diffusely to every group of lateral eyes. Tergites diffusely marbled with dark brown. Coxosternal region and genital operculum immaculate. Pectines pale yellowish, immaculate. Sternites immaculate. Legs spotted with dark brown, more densely on trochanter and femur of I– III; claws with distal third dark brown to blackish. Metasoma bicolor, segments I–IV yellow with a diffuse annulated appearance (basal and distal parts of each segment infuscate in the shape of thin blackish rings, which become somewhat larger and more diffuse dorsally and basally) and segment V very densely reticulated with blackish brown, becoming gradually paler and sparser towards basal half; carinae not infuscate nor underlined with dark pigment. Telson vesicle blackish, aculeus with basal half brownish and distal half blackish.

Chelicerae ( Fig. 67). With dentition typical for the genus, as described for A. krivochatskyi sp. n. (see above).

Pedipalps ( Figs. 70–74). Relatively short but slender, essentially bare. Trichobothrial pattern A-β neobothriotaxic, with the following trichobothria missing: femoral d 2 and chelal esb (both pedipalps), chelal Et and chelal V 1 (right pedipalp). Chelal it in subdistal position, midway between second and third enlarged principal denticle from tip (left chela) or just distal to third enlarged principal denticle from tip (right chela). Femur essentially straight, with carinae weak to moderate, granulose to subdenticulate; intercarinal tegument smooth and glossy. Patella essentially straight, with carinae obsolete to absent, smooth; intercarinal tegument smooth and glossy. Chela elongate and slender; manus conspicuously narrower than patella (ratio 0.70), cylindrical (1.94 times longer than wide, 1.13 times deeper than wide), with carinae obsolete to absent, smooth; intercarinal tegument smooth and glossy; fingers long (movable finger 2.31 times longer than underhand), only subtly curved and with 8–9 principal rows of denticles (the two basalmost rows are poorly defined), basal lobe/notch combination absent, external accessory denticles absent, internal accessory denticles very large and claw-like (increasing in size distally), numbering seven and six on fixed and movable fingers, respectively, movable finger with two claw-like accessory denticles basal to the very large terminal denticle.

Carapace ( Fig. 67). Very strongly trapezoidal (much narrower anteriorly) and wider than long; anterior margin shallowly convex, with 5–6 pairs of thin macrosetae and some very short microsetae. Carination essentially absent: the only carinae present are the superciliaries, which are vestigially granulose to smooth. Furrows: anterior median, median ocular, central median, posterior median and posterior marginal fused, wide and very shallow; lateral oculars, lateral centrals, central transverse, and posterior laterals long, narrow and shallow. Tegument very finely and densely granulose, with many small to medium-sized granules scattered all over except on ocular triangle, where the granules are coarser and glossy.

Sternum ( Fig. 68). Standard for the genus 1, very small, and narrowly pentagonal in shape, with two pairs of macrosetae.

Genital operculum ( Fig. 68). Very large, each half widely crescent-shaped, with 3–4 pairs of inconspicuous macrosetae, plus a few short microsetae. Genital papillae absent.

Pectines ( Fig. 68). Somewhat small for the genus (just reaching leg IV coxa-trochanter joint), subrectangular and densely setose. Tooth count 21/20 (terminal tooth of right pecten missing, but base remains allows unequivocal count). Basal plate heavily sclerotized and densely granulose, much wider than long, anterior margin with a very deep, narrow anteromedian furrow, posterior margin very shallowly convex.

Legs ( Figs. 65–66). Very slender, with all carinae weak to obsolete and subgranulose to smooth; intercarinal tegument smooth and glossy. Tibial spurs absent from both legs III and right leg IV, but moderate-sized on left leg IV.

Mesosoma ( Figs. 67–68). Tergites with the same sculpture as on carapace; I–VI essentially monocarinate: the median longitudinal carina is moderately strong, long, and formed by irregular medium-sized granules that do not project beyond posterior margin and the submedian carinae are undefined on I–VI, although suggested by a few slightly coarser, irregular granules; tergite VII with five well-defined carinae (median, submedians and laterals), which are long, strong and serrate to denticulate. Sternites essentially bare except on posterior margin, which is conspicuously setose; III–VI acarinate, glossy but with posterior and lateral margins finely and densely granulose, spiracles relatively short and slit-like, transversely arranged (not oblique), V with smooth patch absent; VII with two pairs of carinae: the submedians are long and finely denticulate and the laterals are short and coarsely denticulate, intercarinal tegument very densely granulose.

Metasoma ( Figs. 69, 75–78). Robust and slightly wider both basally and distally; with 10/10/10/8/5 complete to almost complete carinae, all except dorsal laterals and laterals supramedians, formed by contiguous, sharply serrate to denticulate granulation: dorsal laterals moderate on I, strong on II–III, moderate on IV (with terminal denticle greatly enlarged as a coarse, conical tubercle), absent on V; lateral supramedians moderately strong on I, weak on II–III, obsolete to very weak on IV (with 1–2 terminal denticles enlarged), vestigial as rounded ridges on V; lateral inframedians strong on I, moderate on II (undefined on basal 1/4), weak on III (undefined on basal 1/3), absent on IV–V; ventral laterals strong on I–II, moderate on III–IV, very strong on V, where become progressively stronger and somewhat flared distally, formed by round, subequal denticles; ventral submedians very strong on I–III, strong on III, moderate on IV, absent on V; ventral median absent on I–IV, moderate on V (poorly defined from intercarinal granulation). Intercarinal tegument smooth and glossy, with fine granules scattered all over except dorsally on I–IV, on V such granules are restricted to ventral surface but are much denser. Dorsal furrow wide and moderately deep on all segments. Setation moderately dense, mostly represented by 6–8 dark macrosetae over every carina. Setae on lateral surfaces of IV and V, and ventral surface of V arising from anterior edges of shallow pits or depressions. Lateral anal margins of V with 3 lobes, ventral anal arc heavily granulated.

Telson ( Figs. 69, 75–78). Moderately setose, with some setae scattered over dorsal and lateral surfaces. Vesicle short oval (1.47 times longer than wide, 1.13 times wider than deep), tegument smooth and glossy, with vestiges of coarse granules arranged into five vestigial longitudinal carinae (ventral median, ventral submedians, and ventral laterals), and some coarse punctations ventrally. Subaculear tubercle absent. Aculeus shorter than vesicle, thick and moderately curved.

MALE. Unknown.

VARIATION. Unknown, no other specimens available to us.

COMPARISON (females only). A. lowei sp. n. can be very easily distinguished from all other Anomalobuthus spp. by its very robust appearance, well evident in its many unique morphometric proportions ( Tab. 4 ). For example, it is the only species of this genus with metasomal segment I about as long as wide (ratio = 1.02 vs. 1.09–1.44 in all other congeners), it has the shortest metasoma + telson when compared to carapace length (ratio = 5.40 vs. 5.68–6.38), the shortest metasomal segment V (length/depth ratio = 1.85 vs. 2.00–2.50) and the deepest telson vesicle (length/depth ratio = 1.66 vs. 1.89–2.17).

DISTRIBUTION ( Fig. 141). Sands of the Ili River valley in southeastern Kazakhstan. It is the easternmost species of Anomalobuthus , occurring at the northern piedmont of the Tian Shan Range.

NOTES. Based on geographical occurrence, the male recorded by Kamenz & Prendini (2008) as A. rickmersi most likely corresponds to A. lowei sp. n.

Anomalobuthus pavlovskyi Teruel, Kovařík et Fet , sp. n.

Figures 79–89, 107–116, 141; Tables 3–5

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Anomalobuthus rickmersi: Birula, 1911b: 171 (in part; “Bajgakum” [=Baigakum]); Pavlovsky, 1916a: 35 (in part; Dzhulek); Pavlovsky, 1916b: 243, fig. 1 (in part; Dzhulek); Fet, 1989: 80, unnumbered figure in page 138 (in part; Dzhulek and Kunya-Urgench); Gromov & Kopdykbaev, 1994: 20 (in part; Dzhulek); Gromov, 2005: 39 (in part); Mityaev et al., 2005: 39 (in part); Gromov, 2006: 44–45 (in part); Graham et al., 2012: 95–106, figs. 1–2, 6 (in part; Baigakum).

HOLOTYPE ♂ ( RTOC: Sco-0476). Kazakhstan, Kizylkum Desert , Kyzyl-Orda Province, Chiili District, ca 2.5 km NW of Baigakum (=Djulek or Dzhulek), 44°20'37'' to 44°20'29''N, 66°27'09'' to 66°27'07''E, 127– 143 m a.s.l., 25 May 2002, leg. V. Fet & A. V. Gromov.

PARATYPES. Same data as holotype , 2♂♂, 2♀♀ ( FKCP) , 1♂ ( ZMUH) , 1♂ ( SOFM) , 1♂ ( NMPC) , 3♀♀ ( RTOC: Sco-0476). Same data as holotype except 6–21 June [19 June –4 July] 1907, leg. D. Glazunov , 2♂♂ ( ZISP 810 , one examined by Max Vachon and bearing his handwritten code VA-1667). Turkmenistan, Daşoguz (= Tashauz ), Province, Köneürgenç (= Kunya-Urgench), 42º23'18"N 59º16'14"E, 3 June 1972, leg. G. S. Medvedev , 1♂, 1♀ ( ZISP 1699 ) .

OTHER MATERIAL. Turkmenistan, Daşoguz (=Tashauz) Province, Shakhsenem [old fortress ruins], 8 October 1983, leg. O. Soyunov, 1♀ (ZMMSU Tb-374).

ETYMOLOGY. We dedicate this species to the memory of the famous Russian zoologist Evgenii Nikanorovich Pavlovsky (1884–1965), who produced many well-known contributions to scorpion anatomy and histology. He was also the Director of ZISP for 20 years (after Alexei Birula was arrested in 1931 during the purges), and was the first one to observe and collect this species at the type locality, more than 100 years ago ( Pavlovsky, 1916a, 1916b).

DIAGNOSIS. Adult size 22–30 mm in males, 23–36 mm in females. Coloration light yellow, essentially immaculate, only with some regular blackish spots on pedipalps and metasoma (darker and more extended in female); metasomal segment V bicolor, blackish with yellowish red basal third (male) to basal half (female), telson blackish. Pedipalp fingers with 8–9 principal rows of denticles and 7–8 internal accessory denticles. Pectines with 24–27 teeth in males and 22–23 in females. Tibial spur in males entirely absent to present only in leg IV, in females absent to present in legs III–IV. Metasoma moderately robust, with most carinae obsolete to vestigial; ventral lateral carinae of segment V composed of narrowly conical denticles, dorsal lateral and lateral supramedian carinae of segments I–IV with terminal denticle enlarged; intercarinal areas of segments I–IV with granulation sparse ventrally (male) to dense ventrally and laterally (female), of segment V moderately dense ventrally (male) to very dense ventrally and laterally (female). Telson vesicle elongate oval and sparsely setose.

DESCRIPTION (adult male holotype, except figures of adult male paratopotype). Coloration ( Figs. 79–80) base light yellowish, with a subtle orange shade; in general, the base color is paler on pedipalp chelae, legs, and pectines. Chelicerae subtly reticulate with dark brown, finger teeth blackish. Pedipalp femur with a conspicuous brown spot on dorsodistal apex, which diffusely continues as a broad stripe over almost all internal surface; patella with internal surface brown; chela immaculate, only with finger denticles blackish. Carapace largely immaculate, only with a blackish spot under every ocular group and interocular triangle moderately infuscate. Tergites immaculate. Coxosternal region and genital operculum immaculate. Pectines pale yellowish, immaculate. Sternites immaculate. Legs immaculate; claws with distal half dark brown. Metasoma bicolor, segment V blackish with basal third yellowish red, I–IV yellow with a diffuse annulated appearance: basal and distal parts of each segment infuscate in the shape of thin blackish rings, which become somewhat larger and more diffuse dorsally and basally; carinae not infuscate nor underlined with dark pigment, except for darkened terminal denticles on dorsal laterals and lateral supramedians. Telson blackish, with basal half of aculeus yellowish.

Chelicerae. With dentition typical for the genus, as described for A. krivochatskyi sp. n. (see above).

Pedipalps ( Fig. 83). Relatively short but very slender, essentially bare. Femur subtly curved inwards, with carinae weak, granulose to subdenticulate; intercarinal tegument smooth and glossy. Patella straight, with carinae vestigial to weak, smooth; intercarinal tegument smooth and glossy. Chela elongate and very slender; manus conspicuously narrower than patella (ratio 0.71), cylindrical (2.00 times longer than wide, 1.20 times deeper than wide), with carinae vestigial to weak, smooth; intercarinal tegument smooth and glossy; fingers very long (movable finger 2.40 times longer than underhand), only subtly curved and with 8/9 principal rows of denticles (the two basalmost rows are poorly defined on movable finger), basal lobe/notch combination absent, external accessory denticles absent, internal accessory denticles very large and claw-like (increasing in size distally), numbering eight and seven on fixed and movable fingers, respectively, movable finger with three claw-like accessory denticles basal to the very large terminal denticle.

Carapace. Very strongly trapezoidal (much narrower anteriorly) and wider than long; anterior margin straight, with 5–6 pairs of thin macrosetae and some very short microsetae. Carination essentially absent: the only carinae present are the superciliaries, which are weakly granulose. Furrows: anterior median, median ocular, central median, posterior median and posterior marginal fused, wide and moderately deep; lateral oculars, lateral centrals, central transverse, and posterior laterals long, narrow and moderately deep. Tegument very finely and densely granulose, with several small to medium-sized granules scattered all over

Sternum. Standard for the genus, relatively small and widely triangular in shape, with one pair of inconspicuous macrosetae.

Genital operculum. Relatively large, each half roundly subtriangular in shape, without macrosetae but with a few short microsetae. Genital papillae present.

Pectines. Standard-sized for the genus: long, extending beyond leg IV coxa-trochanter joint), subrectangular and densely setose. Tooth count 26/27. Basal plate heavily sclerotized, much wider than long, anterior margin with a very deep, narrow anteromedian furrow, posterior margin very shallowly convex.

Legs ( Figs. 79–80). Very slender, with all carinae weak to obsolete and subgranulose to smooth; intercarinal tegument smooth and glossy. Tibial spurs present only on right leg IV.

Mesosoma ( Figs. 79–80). Tergites with the same sculpture as on carapace; I–VI irregularly tricarinate: the median longitudinal carina is moderately strong, short, and formed by irregular medium-sized granules that do not project beyond posterior margin, but the submedian carinae are poorly defined on I–VI; tergite VII with five well-defined carinae (median, submedians and laterals), which are long, strong and finely serrate. Sternites essentially bare; III–VI glossy and with a pair of weak, smooth submedian carinae, spiracles relatively short and slit-like, transversely arranged (not oblique), V with smooth patch absent; VII with two pairs of carinae (submedians and laterals) which are long and finely crenulate, intercarinal tegument coriaceous to minutely granulose.

Metasoma ( Figs. 84–86). Slightly elongated and essentially parallel-sided; with 10/10/10/8/5 complete to almost complete carinae, most formed by conspicuously isolated, sharply serrate to denticulate granulation: dorsal laterals vestigial on I–III, obsolete on IV (with 1–2 terminal denticle slightly enlarged), absent on V; lateral supramedians vestigial to obsolete on I–IV (with 1–2 terminal denticles slightly enlarged), vestigial as rounded ridges on V; lateral inframedians weak on I, obsolete to vestigial on II–III, absent on IV–V; ventral laterals weak on I–IV, moderate to strong on V, where become progressively stronger and somewhat flared distally, formed by sharp, subequal denticles; ventral submedians weak on I–III, vestigial on IV, absent on V (indicated by somewhat raised tegument and irregular granulation on basal half, but largely obscured by intercarinal granulation); ventral median absent on I–IV, moderate on V but largely obscured by intercarinal granulation. Intercarinal tegument smooth and glossy, with sparse granulation of different sizes ventrally. Dorsal furrow wide and shallow on all segments. Setation moderately dense, represented by 6–9 mostly pale macrosetae over every carina.

Telson ( Figs. 84–86). Sparsely setose, with some setae scattered all over dorsal and lateral surfaces. Vesicle short oval (2.00 times longer than wide, 0.83 times wider than deep), tegument smooth and glossy, with vestiges of coarse granules arranged into three obsolete longitudinal carinae (ventral median and ventral submedians), some also on dorsal surface, and some coarse punctations all over. Subaculear tubercle absent, but subtly suggested by a vestigial granule. Aculeus conspicuously shorter than vesicle, thick and shallowly curved.

FEMALE (paratopotype). Very similar to described male, sexual dimorphism evident by: 1) size comparatively larger inside each size-class; 2) mesosoma and metasoma slightly less slender; 3) genital papillae absent; 4) pectines smaller, with consistently lower tooth counts; 5) metasomal segments with ventral and lateral intercarinal tegument much more densely granulose. See Figs. 81–82, 87–89, 114 and Tabs. 3–5

VARIATION. Adult size varied from 22.6–25.5 mm in males and 23.2–35.6 mm in females.

Pectinal tooth count varied as follows ( Tab. 5 ): in males 25 (5), 26 (5), 27 (2), and in females 22 (5) and 23 (3). No significant differences detected among specimens from the localities examined by us.

The tibial spur is highly variable amongst paratypes: it is entirely absent from all legs (one male), present on a single leg IV (three males including holotype, two females), present on each leg III–IV of the same side (left, one female), or present on both legs IV (two males, one female). In all cases where it is present, it is small to vestigial.

COMPARISON. A. pavlovskyi sp. n. can be very easily distinguished by having intercarinal spaces of metasoma much more densely granulose in females than in males ( Figs. 84–89); in all other Anomalobuthus spp. , the metasomal segments are mostly sparsely granulated to smooth and glossy in females ( A. krivochatskyi sp. n., A. lowei sp. n. and A. talebii , see Figs. 62–64, 69, 75– 78, 98–100), or more densely granulose in males ( A. rickmersi , see Figs. 23–25, 27–29, 31–33).

Moreover, this species and A. lowei sp. n. are the only that still retain as adult a typically juvenile character: the segment V with basal third yellowish red ( Figs. 84–89); in all other congeners it is entirely reddish or blackish, or only subtly faded to dark reddish brown at it extreme base ( Figs. 1–2, 4–5, 8–9, 23–45, 59–64, 90–91, 98–100). But A. lowei sp. n. is very different in many morphometric ratios (see Tab. 4 and Comparison section of that species).

DISTRIBUTION ( Fig. 141). Lowland sands of the Aral Basin (Kizylkum Desert), from south-central Kazakhstan through extreme northern Turkmenistan.

NOTES.

1. The first specimens of this species were recorded by Birula (1911b) as A. rickmersi ; all were collected from Baigakum Sands (Dzhulek) in 1907 by Dmitri Konstantinovich Glazunov (1869–1914), a Russian entomologist and brother of the composer Alexander K. Glazunov.

2. The pioneering observations of Pavlovsky (1916b) in Dzhulek (now Baigakum) remain the only data on this species' biology and behavior.

3. The Baigakum specimens of this new species were included (under A. rickmersi ) in a DNA phylogeographic study by Graham et al. (2012), which demonstrated a derived position of this taxon within the genus.