Trypanothacus barnesi, Lowe & Kovařík & Stockmann & Šťáhlavský, 2019

Lowe, Graeme, Kovařík, František, Stockmann, Mark & Šťáhlavský, František, 2019, Trypanothacus gen. n., a new genus of burrowing scorpion from the Arabian Peninsula (Scorpiones: Buthidae), Euscorpius 277, pp. 1-30 : 5-21

Trypanothacus barnesi sp. n.

( Figs. 1–42 , 47–77 , 84–87 , 107–108 , 111–118 , Tables 1–2 ) http://zoobank.org/ urn:lsid:zoobank.org:act:

B0598EB1-8878-4F09-A850-E36129A16ED0

TYPE LOCALITY AND TYPE DEPOSITORY. Oman: Dhofar Province, S of Shalim , 18.041070°N 55.612262°E, FKCP.

TYPE MATERIAL EXAMINED. Oman: 1♀ (paratype), Yalooni, Jiddat Al Harasis , 19°57'N 57°06'E, XII.1988, ONHM 1186 , Yalooni 185, NHMB; 1♀ (paratype), Dhofar, Thamarit, 17°39'N 54°02'E, 10.XI.1989, leg. I. A. J. Brown, ONHM 1416 , NHMB; 3♂ (paratypes), Thumrait, wadi area SE of Thumrait in slightly raised sedimentary plateau, under sheet wood, and in aircraft shelter, 17°42'N 53°59'E, VII.1997, leg. J. N. Barnes, NHMB, GLPC, ONHM; 1♂ subadult (paratype), Thumrait airfield, in aircraft shelter, 17°42'N, 53°49'E, 23. VII.1998, leg. J. N. Barnes, NHMB; 3♂ (holotype No. 1121 and 2 paratypes No. 1594 ), S of Shalim, 18.041070°N 55.612262°E, 6.IX.2016, 21:00–23:00 h, full moon, UV detection, FKCP, GLPC (hemispermatophore); 1♀ (paratype), same locality as holotype, 27–28.X.2017, 23:00–01:00 h, 62% moon, UV detection, leg. M. Stockmann, FKCP.

OTHERMATERIALEXAMINED. Oman: 1juv, nearThamarit, Dhofar, under rock, 450 m a.s.l., 17°40'N 54°02'E, 22.III.1980, leg. J. N. Barnes, Sc74, Barnes 36, MNHN RS8787 ; 1 ♀ fragment, Yalooni, Jiddat Al Harasis , found dead on level limestone rock and sand, 19°56'N 57°06'E, 10.XI.1980, leg. M. D. Gallagher, MDG6020 , NHMB.

DISTRIBUTION. OMAN. Ranging from the inland plateau of Jiddat al-Harasis in central Oman to the Nejd Desert of Dhofar, southern Oman.

ETYMOLOGY. The specific epithet is a patronym dedicated to naturalist J. Neil Barnes, Airworks Ltd, Salalah, Oman, who first collected this species in 1980 from around the Thumrait airbase, and contributed other important specimens of Oman scorpions.

DIAGNOSIS. A member of Trypanothacus gen. n. differentiated from its congener as follows: pedipalp femur L/W ♂ 2.6–3.1, ♀ 2.5–2.7, patella L/W ♂ 2.5–2.7, ♀ 2.4–2.7, chela L/W ♂ 3.5–4.0, ♀ 4.4–4.6; metasoma IV L/D ♂ 2.0–2.2, ♀ 1.9–2.0, V L/D ♂ 2.7–3.0, ♀ 2.5–2.9.

DESCRIPTION. Length of adults: ♂ 40–51 mm, ♀ 43–57 mm; habitus as shown in Figs. 1–4 .

Coloration ( Figs. 1–10 , 15–22 , 29–42 , 71–72 , 84–87 ). Base color yellow or yellowish brown to orange; chelicerae yellow with or without reticulation, dentition reddish to black; median eyes may be surrounded by moderate fuscosity, with weak fuscosity extending forward towards lateral eyes; articular condyles of pedipalp chela fingers and legs reddish brown; macrosetae of body and appendages dark reddish brown; denticles on pedipalp chela fingers and aculeus of telson dark reddish brown.

Carapace ( Figs. 5–6 , 11 , 13 ). Trapezoidal, anterior margin straight or almost straight, with weak fine denticulation, 6–12 marginal macrosetae, bordered with row of coarse granules; surface with sparse to moderately dense coarse and fine granulation; anterior median carinae indicated by coarse granules, other carinae of carapace obsolete; lateral areas of interocular triangle with coarse granules, area between anterior median carinae smooth; median ocular tubercle smooth, except for a few posterior granules; median eyes large, well separated; posterolateral areas of interocular triangle, median postocular area and posterior marginal furrow smooth; posterior margin bordered by row of coarse granules; 5 lateral eyes (3 larger, 2 smaller; in ‘ type 5’ configuration of Loria & Prendini, 2014).

Chelicera ( Figs. 71–73 ). Fingers with typical buthid dentition ( Vachon, 1963); fixed finger with large distal denticle, one subdistal denticle and two basal denticles fused into bicusp, two denticles on ventral surface, one at level of bicusp, other slightly proximal to subdistal denticle; dorsal margin of movable finger with 5 denticles: one large distal denticle, medium-sized subdistal denticle, large medial denticle, and two small, partially fused basal denticles; ventral margin with 3 denticles: one large distal denticle, and two smaller denticles in medial and basal positions.

Mesosoma ( Figs. 1–4 , 6, 8 , 11–14 ). Tergites I–VI tricarinate with median carina and anteriorly diverging pair of lateral carinae; carinae coarsely granular, the lateral pair on the tergites I–II possibly indistinct; all carinae short with only ca. 2–5 granules, confined to posterior half or third of tergite; lateral flanks of tergites with sparse to moderately dense, coarse granulation on posterior half, posterior margins bordered with row of granules; other tergite surfaces smooth to faintly shagreened; tergite VII pentacarinate, median carina a granulated hump, lateral carinae well developed, coarsely granular; posterior margin of tergite VII without rim of granules, intercarinal surfaces smooth or with sparse, coarse granulation; coxae mostly smooth, with few sparse isolated patches of granules, coxal margins smooth or with weak granulation; sternum subtriangular, smooth, with deep posteromedian pit; sternites with smooth surfaces and posterior margins, III–VI without carinae, VII with 2 pairs of weak to moderate, smooth to granulated carinae; sternal chaetotaxy: sternite III-VI posterior margins conspicuously setose, bearing approximately 20 macrosetae, III bearing about a dozen macrosetae on medial surface, IV-VII with 4 medial and 2 lateral macrosetae, VII with single anterior macroseta on each of the 4 carinae, plus a lateral pair; pectines with anterior margins extending to midpoint (♂) or to approximately proximal 1/6 (♀) of trochanter IV, with 3 marginal lamellae, 4–9 middle lamellae; lamellae and fulcra bear numerous short, fine, dark macrosetae; pectine basal piece and genital opercula smooth with fine macrosetae; pectinal tooth counts, ♂ 22–26, ♀ 17–21.

Hemispermatophore ( Figs. 74–77 ). Flagelliform, moderately elongated, trunk ca. 6 times length of capsule region as measured from proximal flagellum base; flagellum well separated from capsule lobes, pars recta short, ca. 2.7 times length of capsule and lobes, 50% length of trunk, with weak fin along anterior margin; pars reflecta long, narrow, gradually tapered, hyaline, 0.88 times length of trunk; capsule region with 4 lobes arranged in ‘3 +1’ configuration, posterior lobe the largest, a lanceolate lamina with pointed apex; median lobe smallest, acuminate, attached to base of posterior lobe along short, suture, but a sclerotized carina not clearly visible; anterior lobe of intermediate length, laminate, apically acuminate; basal lobe a small, rounded knob-like process situated well proximal from point of splitting of posterior and median lobes.

Metasoma and telson ( Figs. 15–28 ). Metasoma I with 10 granulated or crenulated carinae, median lateral carinae complete, ventromedian carinae may be weak or obsolete in males; II–III with 10 granulated or crenulated carinae, median lateral carinae incomplete, indicated by ca. 5–14 granules on posterior 1/3 to 2/3 of segment; ventromedian and ventrolateral carinae on II–III stronger in females, with conspicuously enlarged, dentate or lobate granules, increasing in size posteriorly; coarsely granulose anterior ventral margins present on III–IV; IV with 8 granulated or crenulated carinae; V with 5 carinae, dorsolateral carinae weak, granulated, ventrolateral carinae strong, irregularly crenulated with several enlarged, lobate granules that become larger posteriorly; ventromedian carina of V composed of series of large granules interspersed with finer granulation; intercarinal surfaces of I–IV smooth or almost smooth dorsally, almost smooth or with sparse granules dorsolaterally and laterally; ventrolateral and ventral surfaces of I smooth, of II–III almost smooth with sparse fine granulation, of IV with weak, fine granulation or shagreened; segment V smooth dorsally with scattered small granules, with weak fine granulation laterally, slightly denser fine granulation ventrally; lateral anal arch divided into 2–3 lobes; ventral anal arch armed with regular series of ca. 5–7 coarse granules; telson with distinctly bulbous vesicle bearing coarse granules on ventral surface; granulation more dense in males, more sparse in females, with smooth areas; lateral surface of vesicle with sparser granulation, dorsal surface smooth; aculeus equal to or shorter than vesicle in length, moderately curved; subaculear tubercle absent; chaetotaxy: metasomal segments and telson sparsely setose; long macrosetae located along carinae on metasoma I–IV, typically 2–3 per carina; V with several setae along dorsolateral carina, ca. 6 setae in longitudinal series on lower lateral surface, several setae on ventral surface associated with enlarged granules; telson vesicle with a dozen or more long macrosetae scattered over ventral and ventrolateral surfaces.

Pedipalps ( Figs. 29–40 , 47–70 ). Segments short, robust; femur with 3 strong, granulated carinae: dorsoexternal, dorsointernal, and ventrointernal, other carinae obsolete; dorsal, lateral and ventral surfaces smooth except for few small solitary granules, internal surface smooth except for several solitary coarse granules; distal external surface with group of ca. dozen macrosetae; patella with 7 carinae: dorsointernal, internal and ventrointernal carinae coarsely granulose, internal carina with sparse, non-contiguous granules; dorsomedian and dorsoexternal carinae weak, smooth or almost smooth with traces of coarse granules; external and ventroexternal carinae weak, smooth; ventromedian carina obsolete, indicated by series of few granules; all intercarinal surfaces smooth; setation very sparse, with few large solitary macrosetae on carinae; chela smooth, carinae obsolete with residual traces of carinae in females; few large macrosetae present, ventral surface of movable finger with numerous short macrosetae, increasing in density distally; dentate margin of movable and fixed fingers with 7–8 rows of granules, each with a single external and internal accessory granule, and 3–6 subterminal granules (typically 3–4 of them enlarged); trichobothrial pattern orthobothriotaxic type Aβ, chela fixed finger with db near base of fixed finger.

Legs ( Figs. 1–4 , 9–10 ). Legs with long femora and robust patellae, tibiae and tarsi; femora with few solitary macrosetae; tibiae I–III with dorsal (retrosuperior) linear series of long macrosetae (= tibial ‘bristle comb’, 8–12 setae in sample of N=13 leg III segments), and shorter macrosetae on other surfaces; basitarsi I–III compressed, with two linear series of shorter ventral (proinferior and retroinferior) macrosetae, single linear series of longer dorsal (retrosuperior) macrosetae (= basitarsal ‘bristle comb’, 17–23 setae in sample of N=12 leg III segments); macrosetae long and thin in both sexes; leg IV without basitarsal compression, longer than legs I–III, long macrosetae more sparse, not arranged in linear series; leg I–IV femora and patella with indications of 4–6 carinae, which are usually obsolete; paired ventral carinae on femora granulate or denticulate, prolateral surfaces of femora with weak granulation localized on inferior proximal areas of I–II; tibial spur on leg IV strong, longer than spur on leg III; prolateral pedal spurs basally bifurcate, setose (leg III spur with 6–8 short macrosetae in sample of N=10 segments); retrolateral pedal spurs simple.

Sexual dimorphism. Compared to females, males have longer pectines ( Figs. 2, 4 , 7–8 , 12 , 14 ), higher pectinal tooth counts, and broader pedipalp chelae; chela fingers are straight in both sexes, and males lack recess or scalloping of the proximal dentate margins ( Figs. 29–31, 39–40 , 47–50 , 59–62 , Table 1 ); the male telson vesicle is slightly less bulbous, granulation on the carapace and tergites is more dense; sternite VII carinae of the male are weaker and smoother, and metasomal carinae are not as robust as in females, especially the ventromedian carinae on metasoma II–III; intercarinal granulation is more dense on the male metasoma. For data on morphometric sexual dimorphism, see under species diagnosis and Table 2 .

Measurements. See Table 1 .

Morphometric variation. See Table 2 .

Karyotypes ( Figs. 111–113 ). We analyzed two males (holotype and paratype from type locality) using standard cytogenetic methods (e. g. Kovařík et al., 2009). The diploid number of both specimens is 26 chromosomes ( Figs. 111–112 ). We did not observe chiasmata during male meiosis and centromeres on chromosomes. These characteristics are typical for the scorpions from the family Buthidae (e.g. Mattos et al., 2013). The chromosomes of Trypanothacus barnesi sp. n. gradually decrease in length from 8.69 % to 5.92 % of the halpoid set ( Fig. 113 ).

AFFINITIES. In the diagnosis, we separated T. barnesi sp. n. from its similar allopatric congener, T. buettikeri comb. n. by measurable differences in the shapes of the pedipalp and metasoma, indicated by non-overlapping ranges of morphometric ratios. T. barnesi sp. n. is a less robust, more slender scorpion with narrower pedipalps and posterior metasomal segments. Only limited numbers of specimens of T. buettikeri comb. n. were available to us for study and confirmation of species identity, and additional material may extend morphometric variation in this species. However, samples from around the type locality and a well separated western site (Wadi Turabah) both have robust segments, so the morphometric diagnosis is likely to remain valid. We hypothesize that T. barnesi sp. n. and T. buettikeri comb. n. are sister species that diverged after southern and northern populations were isolated from each other by formation of the Rub‘ al Khali (Empty Quarter) dune system during the Quaternary. Soft unstabilized sands may be an effective physical barrier blocking dispersal of ancestral Trypanothacus adapted to firmer substrates. This model of vicariant speciation by dune barriers has also been suggested for Leiurus macroctenus , a sympatric scorpion that may have substrate preferences similar to those of T. barnesi sp. n. ( Lowe et al., 2014).

COMMENTS ON LOCALITIES AND LIFE STRATEGY.The type locality is a flat desert plateau along route 42, the road between Shalim and Ash Shuwaymiyyah near the coast. The substrate consists of reddish soil alternating with bare rock, clay and sand. Shrubs and trees are scattered and provide only slight patches of humidity in an otherwise arid area. There are few if any loose rocks, except in disturbed areas where stones may be piled near the road by construction activity. The weather on both collection dates was mild at night (ca. 22 °C) with high humidity, barely any wind and no clouds to block the bright moon. At night, Trypanothacus barnesi sp. n. was the most active scorpion in this area. Males were highly active on open ground, whilst females were resting in ambush position at the entrances of their burrows. The burrows were located in sandy areas at the bases of shrubs or rocks, and only rarely on open ground. Only small juveniles to medium-sized immatures were found under rocks without a burrow. Far more males than females were observed on both nights. Remarkably, many were observed actively foraging and feeding on termites, close to the insect nests and trails. Several termites were captured and consumed simultaneously by one scorpion. Other potential prey, including various insects ( Formicidae, Heteroptera, Coleoptera , Blattodea , lepismatids) and arachnids (ticks, spiders and pseudoscorpions) were abundant in the area. Mobile white mites were observed on many individuals of T. barnesi sp. n. (cf. Fig. 85 , an adult female with mites on tergite II and carapace), but not on other arthropods in this habitat. Some individual scorpions were densely covered with mites, whereas others bore only a few. Other species of scorpions found in this area were: Androctonus crassicauda (Olivier, 1807) , Leiurus macroctenus Lowe, Yağmur & Kovařík, 2014 , Femtobuthus shutuae Lowe, 2010 , Nebo sp. , and Compsobuthus acutecarinatus (Simon, 1882) .

Both the type locality and the paratype collection site of Yalooni (Arabian Oryx sanctuary) are located on the Jiddat Al Harasis, an extensive stony limestone plateau in central Oman. The region is ecologically classified as coastal fog desert. Moisture is deposited by condensation of humid air swept inland from the sea during the summer monsoon (khareef). This sustains local vegetation and animal populations, including the scorpion fauna. The other collection sites around Thumrait are located on the Nejd Desert plain in the rain shadow of the Jabal Qara mountains. Without monsoon mist, the terrain is more arid, consisting mostly of sand and gravel flats with sporadic rocky outcrops. Drainage in some shallow wadi beds supports scattered trees and shrubs, with xerophytic herbs and grass as low ground cover. In this area, T. barnesi sp. n. was reported to be present but uncommon (J.N. Barnes, personal communication, 05.VII.1992, Figs. 86– 87 ). Other scorpions recorded from there were: Androctonus crassicauda , Leiurus macroctenus , Orthochirus innesi Simon, 1910 and Xenobuthus anthracinus (Pocock, 1895) . The first two of these were the most common.

CAPTIVE REARING AND BREEDING.Two medium-sized immatures (1♂, 1♀) were reared to maturity in captivity. They were housed in 11 cm × 11cm × 6 cm plastic boxes with two perforated sides, filled with a thin layer of loose sand, and were provided with a small shelter. A burrowing substrate was not provided in order to more easily observe behavior. Average temperatures in summer were 32°C (day) and 24°C (night). In winter, a mild diapause was simulated with temperatures of about 25°C (day) and 20°C (night). One corner of the box was slightly moistened by spraying water once a week to provide drinking water. The water completely evaporated in the following day and conditions were kept dry for the rest of the week to prevent mycosis. Insects ( Thermobia domestica , Shelfordella lateralis and Acheta domesticus ) of adequate sizes were offered as prey.

Three weeks after attaining maturity, the female was mated (XI. 2017). Upon being introduced to the female, a male immediately initiated courtship without preliminary behavior like juddering. The promenade à deux began with the male grasping the female pedipalps, and continued with a series of back and forth pulling motions by both sexes ( Fig. 114 ). Male pectinal movements occurred almost continually, without lengthy interruptions. Remarkably, both sexes engaged in sand scraping with legs I–III ( Fig. 115 ), sometimes settling their bodies into depressions they excavated in the loose sand to rest for a few seconds ( Fig. 116 ) before continuing the promenade à deux. Several episodes of cheliceral massage lasting several seconds were observed ( Fig. 117 ), but there was no sexual sting. After 17 min 48 s, a spermatophore was deposited by the male on a flat rock ( Fig. 118 ) and taken up by the female 3 s later. Both individuals then immediately separated, and the female began to display aggressive behavior towards the male, but did not attempt mate cannibalism. The female was subsequently reared under the same conditions as described above, and gave birth to 23 juveniles in the following season (VII. 2018). These were separated from the parent one week after their first ecdysis, and successfully reared under the same conditions as the parental generation.

Figures

Figures 1–4: Trypanothacus barnesi gen. et sp. n. Figures 1–2. Male paratype from the type locality in dorsal (1) and ventral (2) views. Figures 3–4: Female paratype from the type locality in dorsal (3) and ventral (4) views. Scale bar: 10 mm.

Figures 5–10: Trypanothacus barnesi gen. et sp. n. Figures 5, 7, 9. Male holotype, carapace and tergites I–V (5), coxosternal area and sternites (7), right legs I–IV, retrolateral aspect (9.I–IV). Figures 6, 8, 10. Female paratype from the type locality, carapace and tergites I–III (6), coxosternal area and sternites III–IV (8), right legs III–IV, retrolateral aspect (10.III–IV).

Figures 11–12: Trypanothacus barnesi gen. et sp. n. Male paratype from Thumrait (VII.1997), carapace and tergites I–VII (11), coxosternal area and sternites (12). UV fluorescence. Scale bars: 2 mm.

Figures 13–14: Trypanothacus barnesi gen. et sp. n. Female paratype from Yalooni (XII.1988), carapace and tergites I–VII (13), coxosternal area and sternites (14). UV fluorescence. Scale bar: 2 mm.

Figures 15–22: Trypanothacus barnesi gen. et sp. n. Figures 15, 20–22. Male holotype, metasoma V and telson lateral view (15), metasoma and telson lateral (20), ventral (21), and dorsal (22) views. Figures. 16–19. Female paratype from type locality, metasoma V and telson lateral view (16), metasoma and telson lateral (17), ventral (18), and dorsal (19) views. Scale bars: 4 mm (17–19, 20–21).

Figures 23–28: Trypanothacus barnesi gen. et sp. n. Figures 23–25. Male paratype from Thumrait (VII.1997), metasoma and telson dorsal (23), lateral (24) and ventral (25) views. Figures 26–28. Female paratype from Yalooni (XII.1988), metasoma and telson dorsal (26), lateral (27) and ventral (28) views. UV fluorescence. Scale bar: 4 mm.

Figures 29–46: Figures 29–42: Trypanothacus barnesi gen. et sp. n. Figures 29–40. Pedipalp segments. Figures 29–38. Male holotype, chela dorsal (29), external (30), and ventral (31) views. Patella dorsal (32), external (33) and ventral (34) views. Femur and trochanter internal (35), dorsal (36), and ventral (37) views. Movable finger dentition (38). Trichobothrial pattern is indicated in Figures 30–33 and 35–36. Figures 39– 40. Female paratype from the type locality, chela dorsal (39) and external (40) views. Figures 41–42. Telson, male holotype (41) and female paratype from type locality (42), lateral views. Figures 43–44: T. buettikeri comb. n., pedipalp chela dorsal, male holotype (43) and female paratype (44). Figures 45–46: Buthacus stockmanni Kovařík et al., 2016, telson, male holotype (45) and female paratype (46), lateral views.

Figures 47–58: Trypanothacus barnesi gen. et sp. n. Pedipalp segments of male paratype from Thumrait (VII.1997), chela dorsal (47), external (48), ventral (49) and internal (50) views. Patella dorsal (51), external (52), ventral (53) and internal (54) views. Femur and trochanter dorsal (55), external (56), ventral (57) and internal (58) views. UV fluorescence. Scale bar: 2 mm.

Figures 59–70: Trypanothacus barnesi gen. et sp. n. Pedipalp segments of female paratypes, chela dorsal (59), external (60), ventral (61) and internal (62) views. Patella dorsal (63), external (64), ventral (65) and internal (66) views. Femur and trochanter dorsal (67), external (68), ventral (69) and internal (70) views. Dorsal views (59, 63, 67): paratype female from Thumrait (XI.1989) (with normal dorsointernal carina on femur). External (60, 64, 68), ventral (61, 65, 69) and internal (62, 66, 70) views: paratype female from Yalooni (XII.1988) (with malformed dorsointernal carina on femur). UV fluorescence. Scale bars: 2 mm (59, 63, 67), 2 mm (60–62, 64–66, 69–70).

Figures 71–77: Trypanothacus barnesi gen. et sp. n. Figures 71–73. Paratype male, right chelicera, dorsal (71), ventral (72) and ventral UV fluorescence (73) views. Scale bar: 1 mm. Figures 74–77. Holotype male, right hemispermatophore, capsule region with lobes in anterior (74), convex compressed (75) and posterior (76) views, and whole hemispermatophore in convex view (77). Scale bars: 1 mm (71–73), 500 µm (74–76), 1 mm (77).

Figures 84–85: Trypanothacus barnesi gen. et sp. n., in vivo habitus, male holotype (84) and female paratype from type locality (85).

Figures 86–89: Trypanothacus barnesi gen. et sp. n.. Figures 86–87. Male (86) and female (87) from Thumrait, photos by J. N. Barnes. Figures 88–89. Type locality, Oman, S of Schelim, 18.041070°N 55.612262°E.

Figures 107–110: Morphometric scaling relations of the telson of fossorial ‘Buthus group’ scorpions. Figure 107. Double logarithmic scatterplot of telson vesicle length/ depth (L/ D) ratio vs. aculeus length normalized to vesicle length (aculeus L/ vesicle L), for large pelophiles/ semipsammophiles (4 genera, 53 species) (Buthus, Odontobuthus, Pantobuthus: orange circles; Trypanothacus gen. n.: blue circles) and large psammophiles/ ultrapsammophiles (6 genera, 29 species) (Buthacus: cyan circles; Vachoniolus: dark green circles; Apistobuthus, Buthiscus, Liobuthus, and Plesiobuthus: yellow circles). Dark line: linear least squares regression fit to all data shown (Pearson correlation coefficient, R = 0.57). Upper right inset: diagram of measurements estimated from inverted telson lateral profiles, with dorsal surface of vesicle approximately leveled; SA = ventral profile length of aculeus from base point to tip, LV = length of vesicle from anterior limit or inflection point to aculeus base point; HV = vesicle depth or height (distance between dorsal and ventral tangent planes). Base point was placed at local minimum of inverted ventral profile curve. SA was calculated by cubic spline fit (>10 anchor points). For a minority of broken tips, SA was estimated by intersecting extrapolated dorsal and ventral profile curves (<5%). Figure 108. Scatterplot of fossorial buthids in Figure 107, superposed upon larger scale plot of Order Scorpiones (gray circles). Gray line: linear least squares regression fit to Order Scorpiones data (including data in Figure 107) (Pearson correlation coefficient, R = –0.62).Also plotted are data from small ‘Buthus group’ psammophiles/ ultrapsammophiles (5 genera, 10 species: Compsobuthus arabicus, Lanzatus somalilandus, Neobuthus kutcheri, Picobuthus dundoni, P. wahibaensis, Anomalobuthus krivochatskyi, A. pavlovskyi, A. rickmersi, A. lowei, A. talebii) (dark yellow circles). Figure 109. Double logarithmic scatterplot of carapace length (L) vs. aculeus length normalized to vesicle length (i.e., aculeus L/ vesicle L) for psammophile/ ultrapsammophile members of the ‘Buthus group’, including large species in Figure 107, and small species in Figure 108 (dark yellow circles). Blue line: linear least squares regression fit to all data shown (Pearson correlation coefficient, R = 0.74). Figure 110. Scatterplot of psammophile buthid data in Figure 109 (all in one color, blue circles) superposed upon larger scale plot of Order Scorpiones (gray circles). Gray line: linear least squares regression to Order Scorpiones data (including data in Figure 109) (Pearson correlation coefficient, R = –0.01). Also plotted are data from bothriurid genus Brachistosternus (43 species) (magenta triangles), and corresponding linear least squares regression fit (R = 0.39). All plots in Figures 107-110 include data from both males and females. Data were assembled by mining taxonomic literature or directly measuring specimens. Included are 1854 species or subspecies (1331 ♂, 1201 ♀) representing all known extant genera (1841 species or subspecies) plus 7 fossil genera (13 species).

Figures 111–113: Trypanothacus barnesi gen. et sp. n., mitotic metaphase (111), postpachytene (112) and ideogram (113) (y axis - % of the chromosome length of the haploid set) of male holotype. Scale bar = 5 μm.

Figures 114–116: Trypanothacus barnesi gen. et sp. n., promenade à deux (courtship dance) in a mating pair of individuals collected from the type locality. Photos taken in laboratory. Male is on the right.

Figures 117–118: Trypanothacus barnesi gen. et sp. n., cheliceral massage (117) and spermatophore deposition (118) in a mating pair of individuals collected from the type locality. Photos taken in laboratory. Male is on the right.