taxonID	type	description	language	source
3E597234FFB3CB0961C3E5CEFCD183E6.taxon	materials_examined	TYPE SPECIES. Scorpio (Androctonus) madagascariensis Gervais, 1843.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB3CB0961C3E5CEFCD183E6.taxon	diagnosis	DIAGNOSIS. Small to medium-sized buthids, adult length 25 – 75 mm; anterosubmedian carinae of carapace absent; median ocular tubercle located in posterior 2 / 3 of carapace; fixed finger of chelicera with 2 denticles on ventral surface; pedipalp femur with trichobothria d 1 - d 3 - d 4 non-reflex angle opening externally (retrolaterally) (α-configuration; Vachon, 1975); femur petite ‘ trichobothrium’ d 2 position internal; pedipalp patella trichobothrium d 3 position external (retrolateral) to dorsomedian carina (DM c) (Fet et al., 2005); pedipalp patella trichobothrium esb 2 distal to esb 1 (mean distance> 0.18 esb 1 - em distance); pedipalp chela manus with trichobothrium Eb 2 distal to Eb 1, trichobothrium V 2 medial, located behind V 1 along proximo-distal axis of manus; chela manus with petite ‘ trichobothrium’ Eb 3 usually well separated from Eb 2, by more than half the distance between Eb 1 and Eb 2; pedipalp fixed finger with trichobothrium db in middle 30 % – 60 % of finger, trichobothrium it distal; pedipalp chela movable finger with 11 – 16 imbricated subrows of median denticles, each flanked proximally by 2 enlarged external accessory denticles; chela movable finger typically with 4 external subdistal granules; pedipalp manus with weak or obsolete carination; pectines with fulcra; internal and accessory internal fulcra present, rounded, sclerotized, fluorescent; female basal middle lamella (bml) not dilated, female basal pectinal tooth (bpt) modified but not distinctly longer than other teeth, dilated, oval, subrectangular or subtriangular; pectinal tooth count (excluding ♀ bpt): ♂ 15 – 24, ♀ 12 – 22; hemispermatophore capsule long or short, posterior lobe with long, lanceolate extension; legs III – IV with tibial spurs present; leg IV, mean ratio of tibial spur L / tibia distal D: <0.69; legs I – IV, telotarsi with ventral setation sparse, discrete with <25 setae in rows; tergites III – VI monocarinate; sternites with spiracles broad, hemi-elliptical or ovoid, sternite IV spiracle L / W <5; tergite VII, sternite VII and metasomal segments I – III without microsetal fringes on posterior margins; metasoma I ventrosubmedian carinae granulate or costate-granulate; telson with oval or bulbous vesicle, with or without subaculear tubercle in adults; cuticle with weak UV fluorescence.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB3CB0961C3E5CEFCD183E6.taxon	discussion	REMARKS. The above standard diagnosis is partly hypothetical because some characters have not been confirmed for all 14 species assigned to the genus. For a confirmed differential diagnosis, see below under AFFINITIES. SUBORDINATE TAXA.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB3CB0961C3E5CEFCD183E6.taxon	description	Grosphus ambre Lourenço, Wilmé & Waeber, 2018 Grosphus angulatus sp. n.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB3CB0961C3E5CEFCD183E6.taxon	description	Grosphus polskyi Lourenço, Qi & Goodman, 2007 Grosphus rakotoariveloi Lourenço, Wilmé, Soarimalala & Waeber, 2017 Grosphus simoni Lourenço, Goodman & Ramilijaona, 2004 Grosphus tavaratra Lourenço, Soarimalala & Goodman, 2009 Grosphus voahangyae Lourenço & Wilmé, 2015 AFFINITIES. The genus Grosphus belongs to the ‘ Charmus / Uroplectes ’ group of buthids (Fet et al., 2005; Štundlová et al. 2022). Grosphus is similar to Teruelius, and differentiated from other buthids, in the following combination of characters: pedipalp femur with trichobothria d 1 - d 3 - d 4 non-reflex angle opening externally (retrolaterally) (α-configuration; Vachon, 1975); pedipalp patella trichobothrium d 3 position external (retrolateral) to dorsomedian carina (DM c) (Fet et al., 2005); pedipalp patella trichobothrium esb 2 distal to esb 1 (mean distance> 0.18 esb 1 - em distance); pedipalp manus with trichobothrium Eb 2 distal to Eb 1, trichobothrium V 2 medial, located behind V 1 along proximo-distal axis of manus; pedipalp manus with weak or obsolete carination; pedipalp fixed finger with trichobothrium db in middle 30 % – 60 % of finger, trichobothrium it distal; pedipalp chela movable finger with 11 – 16 imbricated subrows of median denticles, each flanked proximally by 2 enlarged external accessory denticles; pectines with fulcra; internal and accessory internal fulcra present, rounded, sclerotized, fluorescent; female bml not dilated, female bpt modified, dilated or elongated; legs III – IV with tibial spurs present; tergites III – VI monocarinate; tergite VII, sternite VII and metasomal segments I – III without microsetal fringes on posterior margins. DIFFERENTIAL DIAGNOSIS. Grosphus is differentiated from Teruelius by any combination of two or more of the following characters: leg IV, mean ratio of tibial spur L / tibia distal D <0.69; legs I – IV, telotarsi with ventral setation sparse, discrete with <25 setae in rows; sternite IV spiracles broad, hemielliptical or ovoid, L / W <5; metasoma I ventrosubmedian carinae granulate or costate-granulate; PTC: ♂ <24, ♀ <22; and pedipalp femur petite ‘ trichobothrium’ d 2 position internal.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB6CB146328E0D1FD138229.taxon	description	(Figs. 13 – 15, 20, 23 – 28, 30 – 35, 43 – 44, 49 – 50, 54, 71 – 74, 93 – 94, 103 – 105, 160, 165 – 167, 175 – 178, 180, 182 – 183, 189, 196 – 209, 285 – 324, 326 – 327, 393, Tabs. 13 – 14) http: // zoobank. org / urn: lsid: zoobank. org: act: 8 CA 04 BBF- 5698 - 459 A- 8 E 69 - AA 16838667 E 1 Grosphus sp. nr hirtus Lowe & Kovařík, 2019: 13, 19, 21, 30 – 31, 41, 43 – 44, 54, 60; figs. 26, 36, 40, 107, 111, 158.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB6CB146328E0D1FD138229.taxon	materials_examined	TYPE LOCALITY AND TYPE REPOSITORY. Madagascar: Moramanga env., Anjiro, 1995; NZAC, GLPC. TYPE MATERIAL. Madagascar: Moramanga env., Anjiro, 10. II. 1995, 1 ♀ (holotype), 4 ♀ (paratypes) NZAC; 1 ♀ (paratype) GLPC.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB6CB146328E0D1FD138229.taxon	etymology	ETYMOLOGY. The species name refers to the angulate distal vertex of the modified basal pectinal tooth (bpt) in females.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB6CB146328E0D1FD138229.taxon	diagnosis	DIAGNOSIS (ADULT FEMALES). Medium-sized member of the genus, total length of adults around 50 mm; base color dark reddish-brown, carapace with weak variegated fuscous pattern; pedipalp patella with obsolete dorsointernal carina; femoral trichobothrium e 1 level with or slightly proximal to d 5; pedipalp chela movable finger with 11 – 12 median denticle subrows, fixed finger with 13; female bpt with angulate distal vertex; PTC 14 – 16; spiracles wide, ovoid in profile; metasoma II – IV with 3 – 4 robust, dentate granules on posterior dorsosubmedian carinae; telson vesicle hemielliptic in lateral profile, with small subaculear tubercle; morphometrics, L / W ratios (n = 6): metasoma I 0.86 – 0.92, metasoma II 1.06 – 1.10, metasoma III 1.18 – 1.20, metasoma IV 1.42 – 1.45, metasoma V 1.84 – 1.90, pedipalp chela 3.72 – 3.86, pedipalp femur 2.45 – 2.74, pedipalp patella 2.06 – 2.24.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFB6CB146328E0D1FD138229.taxon	description	DESCRIPTION (FEMALE). Coloration (Figs. 285 – 286, 310, 312, 314 – 317). Base color dark reddish-brown; carapace with weak variegated fuscosity; coxosternal area and sternites III – VI dark orange-brown; pectines yellow; legs dark-reddish brown to orange-brown, with more pale telotarsi; chelicerae dark reddish-brown, dorsal manus with fuscous anterior margin and reticulation. Carapace (Figs. 285, 287). Subrectangular, W / L 1.10 – 1.19; medial surface level along its entire length; anterior margin slightly concave with small epistomal process; preocular L / carapace L 0.36 - 0.38; surface mostly bearing fine granules of moderate density, except in some bilateral smooth strips and areas around central median, posterior median, posterior transverse and posterior marginal furrows; granulation more coarse and dense on preocular triangle; superciliary carinae granulate; macrosetae absent; lateral eye groups composed of either 3 large + 2 small ocelli (8 / 12 groups), or 2 large + 2 small ocelli (4 / 12 groups), i. e., type 5 or type 4 B, respectively (Loria & Prendini, 2014); 4 carapaces with type 5 / 5 pattern, 2 carapaces with type 4 B / 5 pattern, 1 carapace with type 4 B / 4 B pattern (left group / right group); median eyes of moderate size, eye diameter / carapace L 0.086. Chelicerae (Figs. 310 – 313). Dorsal surface of manus granulate on anterior 1 / 5, smooth on posterior 4 / 5; anterior granulate area with 8 – 9 macrosetae, 4 – 5 pale, fluorescent microsetae; dorsointernal carina strong, granulate; fingers with typical buthid dentition (Vachon, 1963), movable finger dorsal margin with two large subdistal denticles and two small basal denticles, ventral margin with subdistal and basal denticles (notched basal denticle in Figs. 312 – 313 is atypical or worn, and not present in the holotype and other paratypes), fixed finger with large subdistal denticle and proximal bicusp, two denticles on ventral surface; dorsal surface of movable finger smooth, with dorsal row of 5 – 7 pale, fluorescent microsetae. Coxosternal area (Figs. 286, 288). All coxae smooth with sparse macrosetae and fluorescent microsetae; sternum smooth, subtriangular, with narrow slit-like posteromedial depression, bearing 2 macrosetae; genital opercula smooth, divided. Pectines (Figs. 28, 32, 34 – 35, 43 – 44, 286, 288, 320 – 335). Basal piece smooth, with concave anterior margin, surface flat without groove or pit, pectines with 3 marginal lamellae, 4 – 6 middle lamellae, extending to distal end of coxa IV; marginal and middle lamellae with sparse cover of macrosetae and pale, fluorescent microsetae; fulcra with 2 fluorescent microsetae; bpt with angulate distal vertex. Hemispermatophore. Unknown. Mesosoma (Figs. 54, 71 – 74, 93 – 94, 285 – 288, 393). Tergites: pretergites smooth, with microsulcate posterior margins; tergites densely, finely granulated, with narrow, smooth transverse lateral strips on tergites II – VI; tergite I without discernible carinae, tergites II – VI with single weak, granulate median carina, tergite VII with medial hump and 2 pairs of granulate carinae; all tergites lacking macrosetae. Sternites: sternites III – VI smooth, acarinate; sternite VI smooth with two pairs of weak, granulate carinae; posterior margins of all sternites smooth; spiracles broad, hemi-elliptic; sternite III – VI macrosetae: one submedian pair, one lateral pair, two posterior marginal pairs; sternite VII macrosetae: one submedian carinal pair, one lateral pair; sternites III – VI glossy, sternite VII matte. Metasoma (Figs. 105, 160, 285 – 286, 301 – 303). Segments of uniform width, robust. Carination: segments I – III with 10 complete carinae, IV with 8 complete carinae (lateral median carinae indistinct posteriorly), V with 5 complete carinae; all carinae granulate; dorsosubmedian carinae on II – VI with enlarged dentate posterior granules; ventrolateral carinae on V strongly, uniformly granulate; dorsolateral carinae on V irregularly, coarsely granulate; lateral anal margin with 2 small granules, ventral anal margin with up to 18 granules. Intercarinal surfaces: moderately dense, fine granulation on lateral, ventrolateral and ventral surfaces of all segments; I – VI with dorsomedian surfaces finely granulated or shagreened, with decreasing density on more posterior segments, dorsolateral surfaces with sparse fine granules, mostly smooth; V with dorsomedian surface smooth, dorsolateral surfaces with sparse fine granules, mostly smooth. Setation: dorsal surfaces without setae; other surfaces with numerous short macrosetae and fluorescent microsetae, mostly associated with carinae. Telson (Figs. 285 – 286, 301 – 303). Vesicle dorsal surface smooth, lateral and ventral surfaces covered with numerous coarse and fine granules, except along lateral and paramedian longitudinal strips which are smooth; vesicle hemi-elliptic in lateral profile, with distinct subaculear tubercle; numerous short macrosetae and fluorescent microsetae on lateral and ventral surfaces; dorsal surface with posterior patch of short macrosetae; aculeus shorter than vesicle. Pedipalps (Figs. 289 – 300, 304 – 309, 318 – 319). Segments robust (see DIAGNOSIS for morphometrics). Femur: dorsointernal, dorsoexternal and ventrointernal carinae distinct, granulate; other carinae indistinct; intercarinal surfaces smooth; sparse short macrosetae and fluorescent microsetae present, mostly associated with granules and carinae. Patella: all carinae obsolete, smooth except for isolated granules on internal surface; numerous short macrosetae and fluorescent microsetae present. Chela: all carinae obsolete, surfaces smooth, with dense cover of short macrosetae and fluorescent microsetae; 11 – 12 median denticle subrows on fixed finger, 13 on movable finger including short subdistal row (excluding malformations), all subrows except proximal flanked by one mid-row internal and two proximal external accessory denticles. Trichobothriotaxy: orthobothriotaxic, type Aα (Vachon, 1974), femur d 2 internal, e 1 level with or slightly proximal to d 5; chela fixed finger db proximal to est. Legs (Figs. 285 – 286, 314 – 317). Femora and patellae with granulate carinae, prolateral surfaces sparsely granulate and matte, retrolateral surfaces smooth, glossy; tibial spurs present on legs III – IV; retrolateral tarsal spurs simple, prolateral tarsal spurs with very small basal bifurcation; basitarsi with 2 axial rows of macrosetae on ventral surface, irregular macrosetae on lateral and dorsal surfaces; telotarsi with 2 axial rows of up to 8 short macrosetae on ventral surface, lateral apices with 4 – 6 macrosetae; tarsal ungues stout. Measurements. See Tables 13 – 14. AFFINITIES. G. angulatus sp. n. is similar to three other species of the genus: G. hirtus, G. polskyi and G. voahangyae. The four species share the following characters: metasoma I stout, mean L / W ratio ♀ <0.97, ♂ <1.02 (Fig. 168); dorsointernal carina of pedipalp patella with granulation sparse or absent; pedipalp chela fixed finger with trichobothrium db level with or proximal to 0.92 est (Fig. 167); leg IV tibial spur L / tibia distal D, mean ratio 0.60 – 0.65 (Fig. 50). These four species were often recovered as a monophyletic ‘ hirtus ’ group in cladistic analyses (e. g., Figs. 196 – 200, 203 – 209). G. hirtus differs from G. angulatus sp. n. as follows: lighter base color of yellowish to reddish-yellow, with more distinct variegated fuscous patterns on body, pedipalps and legs; pedipalp patella with weakly granulate dorsointernal carina; femoral trichobothrium e 1 distal to d 5; female bpt rounded distally (Fig. 325 vs. Figs. 320 – 324), z-scores of Fourier harmonics clustered separately (Figs. 34 – 35), nonoverlapping in PC 1 - PC 2 plane (Fig. 326), disjunct in PC 3 - PC 4 plane (Fig. 327); metasoma II – IV with smaller dentate granules on posterior dorsosubmedian carinae; more elongate pedipalp patella, L / W ♀ 2.21 – 2.42. G. voahangyae differs from G. angulatus sp. n. as follows: smaller size, adult female total length around 40 mm; more distinct variegated fuscous patterns on body, pedipalps and legs; femoral trichobothrium e 1 distal to d 5; female bpt ovoid, rounded distally, z-scores of Fourier harmonics clustered separately (Figs. 34 – 35); more elongate pedipalp patella, L / W ♀ 2.22 – 2.30. Only the male is known for G. polskyi, but the following differences from G. angulatus sp. n. do not exhibit strong sexual dimorphism in species of Grosphus for which both sexes are known: smaller size, adult male total length around 33 mm; lighter base color of reddish-yellow, with more distinct variegated fuscous patterns on body; femoral trichobothrium e 1 distal to d 5; metasoma II – IV without enlarged dentate granules on posterior dorsosubmedian carinae; telson with larger subaculear tubercle. The species G. tavaratra is also similar, but only the male is known. However, G. tavaratra differs in having a granulate dorsointernal carina on the pedipalp patella, femoral trichobothrium e 1 distal to d 5, and narrower spiracles, traits that do not exhibit strong sexual dimorphism in species of Grosphus for which both sexes are known.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	materials_examined	TYPE SPECIES. Buthus limbatus Pocock, 1889.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	diagnosis	DIAGNOSIS. Medium- to large-sized buthids, adult length 35 – 120 mm; anterosubmedian carinae of carapace absent; median ocular tubercle located in posterior 2 / 3 of carapace; fixed finger of chelicera with 2 denticles on ventral surface; pedipalp femur with trichobothria d 1 - d 3 - d 4 non-reflex angle opening externally (retrolaterally) (α-configuration; Vachon, 1975); femur petite ‘ trichobothrium’ d 2 position dorsal; pedipalp patella trichobothrium d 3 position external (retrolateral) to dorsomedian carina (DM c) (Fet et al., 2005); pedipalp patella trichobothrium esb 2 distal to esb 1 (mean distance> 0.18 esb 1 - em distance); pedipalp chela manus with trichobothrium Eb 2 distal to Eb 1, trichobothrium V 2 medial, located behind V 1 along proximodistal axis of manus; chela manus with petite ‘ trichobothrium’ Eb 3 usually near Eb 2, closer than than half the distance between Eb 1 and Eb 2; pedipalp fixed finger with trichobothrium db in middle 30 % – 60 % of finger, trichobothrium it distal; pedipalp chela movable finger with 11 – 16 imbricated subrows of median denticles, each flanked proximally by 2 enlarged external accessory denticles; chela movable finger typically with 4 external subdistal granules; pedipalp manus with weak or obsolete carination; pectines with fulcra; internal and accessory internal fulcra present, rounded, sclerotized, fluorescent; female basal middle lamella (bml) not dilated, female basal pectinal tooth (bpt) modified with elongate, tapering distal extension, distinctly longer than other teeth; pectinal tooth count (excluding ♀ bpt): ♂ 25 – 42, ♀ 23 – 35; hemispermatophore capsule short, posterior lobe rounded without long, lanceolate extension; legs III – IV with tibial spurs present; leg IV, mean ratio of tibial spur L / tibia distal D:> 0.69; legs I – IV, telotarsi with ventral setation dense, irregular with broad, brush-like strips of> 25 long filiform macrosetae; tergites III – VI monocarinate; sternites with spiracles narrow, slit-like, sternite IV spiracle L / W> 5; tergite VII, sternite VII and metasomal segments I – III without microsetal fringes on posterior margins; metasoma I ventrosubmedian carinae costate-granulate, smooth or absent; telson with oval or bulbous vesicle, without subaculear tubercle in adults; cuticle with strong UV fluorescence.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	discussion	REMARKS. The above standard diagnosis is partly hypothetical because some characters have not been confirmed for all 22 species assigned to the genus. For a confirmed differential diagnosis, see below under AFFINITIES. SUBORDINATE TAXA.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	description	Teruelius ankarafantsika (Lourenço, 2003) Teruelius ankarana (Lourenço & Goodman, 2003) Teruelius annulatus (Fage, 1929)	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	description	Teruelius feti (Lourenço, 1996)	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	description	Teruelius intertidalis (Lourenço, 1999) Teruelius limbatus (Pocock, 1889)	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	description	Teruelius makay (Lourenço & Wilmé, 2015) Teruelius mavo (Lourenço & Rossi, 2020)	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFABCB1662C6E102FDFE8569.taxon	description	Teruelius waeberi (Lourenço & Wilmé, 2016) AFFINITIES. The genus Teruelius belongs to the ‘ Charmus / Uroplectes ’ group of buthids (Fet et al., 2005; Štundlová et al. 2022). Teruelius is similar to Grosphus, and differentiated from other buthids, in the following combination of characters: pedipalp femur with trichobothria d 1 - d 3 - d 4 non-reflex angle opening externally (retrolaterally) (α-configuration; Vachon, 1975); pedipalp patella trichobothrium d 3 position external (retrolateral) to dorsomedian carina (DM c) (Fet et al., 2005); pedipalp patella trichobothrium esb 2 distal to esb 1 (mean distance> 0.18 esb 1 - em distance); pedipalp manus with trichobothrium Eb 2 distal to Eb 1, trichobothrium V 2 medial, located behind V 1 along proximo-distal axis of manus; pedipalp manus with weak or obsolete carination; pedipalp fixed finger with trichobothrium db in middle 30 % – 60 % of finger, trichobothrium it distal; pedipalp chela movable finger with 11 – 16 imbricated subrows of median denticles, each flanked proximally by 2 enlarged external accessory denticles; pectines with fulcra; internal and accessory internal fulcra present, rounded, sclerotized, fluorescent; female bml not dilated, female bpt modified, dilated or elongated; legs III – IV with tibial spurs present; tergites III – VI monocarinate; tergite VII, sternite VII and metasomal segments I – III without microsetal fringes on posterior margins. DIFFERENTIAL DIAGNOSIS. Teruelius is differentiated from Grosphus by any combination of two or more of the following characters: leg IV, mean ratio of tibial spur L / tibia distal D> 0.69; legs I-IV, telotarsi with ventral setation dense, brush-like with> 25 irregular setae; sternite IV spiracles narrow, slitlike, L / W> 5; metasoma I ventrosubmedian costate-granulate, smooth or absent; PTC: ♂> 24, ♀> 22; and pedipalp femur petite ‘ trichobothrium’ d 2 position dorsal.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFA9CB186314E648FD5C82A9.taxon	description	(Figs. 13 – 16, 19 – 21, 23 – 28, 30 – 35, 49 – 50, 52, 137, 165, 167 – 168, 175 – 178, 180 – 183, 196 – 209, 328 – 392, Tab. 15) http: // zoobank. org / urn: lsid: zoobank. org: act: 7 CBD 4 E 06 - 8 D 6 C- 47 CB-BBD 4 - F 068 D 262 DF 50	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFA9CB186314E648FD5C82A9.taxon	materials_examined	TYPE LOCALITY AND TYPE REPOSITORY. Madagascar: Toliara Province, Tsimanampetsotsa National Park, Andranovao camp, FKCP. TYPE MATERIAL. Madagascar: Toliara Province, Tsimanampetsotsa National Park, Andranovao camp, 15 m a. s. l., 24 ° 01.505 ' S 43 ° 44.306 ' E, 1 ♂ (holotype), 3 ♂ 1 ♀ 1 juv ♂ (paratypes), 2014, FKCP, GLPC (1 hemispermatophore).	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFA9CB186314E648FD5C82A9.taxon	etymology	ETYMOLOGY. The species is named in honor of Czech entomologist, physician Martin Häckel.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFA9CB186314E648FD5C82A9.taxon	diagnosis	DIAGNOSIS. Small to medium-sized member of the genus, total length of adults 30 – 53 mm; body and appendages uniformly yellow, metasoma IV – V (or only V) and telson dark, blackbrown; carapace of males with granulate superciliary carinae; pedipalp patella with strong, costate-granulate dorsointernal carina in both sexes; pedipalp chela of males with internal surface of manus granulate, fingers without undulations on proximal dentate margins; leg III tibial spur L / tibia distal D> 0.73; female bpt falcate, without long narrow extension, shorter than basal comb width; PTC ♂ 37 – 40, ♀ 27 – 28, regular pectine tooth L / W ♂ 4.71, ♀ 3.89; hemispermatophore posterior lobe short, apically rounded, with two lateral carinae; metasoma III ventral intercarinal surface smooth, dorsosubmedian carinae of males bearing large dentate posterior terminal granule; metasoma V with dorsosubmedian carinae smooth, obsolete; telson with aculeus length equal to vesicle length, vesicle weakly granulate on ventral surface; morphometrics, L / W ratios (n = 4 ♂, 1 ♀): metasoma I ♂ 1.20 – 1.27, ♀ 1.195, metasoma II ♂ 1.49 – 1.54, ♀ 1.47, metasoma III ♂ 1.57 – 1.62, ♀ 1.56, metasoma IV ♂ 1.81 – 2.01, ♀ 1.89, metasoma V ♂ 2.03 – 2.19, ♀ 2.41, pedipalp chela ♂ 3.64 – 3.77, ♀ 5.16, pedipalp femur ♂ 2.78 – 3.44, ♀ 2.34, pedipalp patella ♂ 2.67 – 3.65, ♀ 2.64.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFA9CB186314E648FD5C82A9.taxon	description	DESCRIPTION. Coloration (Figs. 328 – 333, 338 – 357, 372 – 387). Base color yellow to orange-yellow; metasoma IV either dark brown (♂) or yellow and partially black ventrally (♀); metasoma V and telson either dark brown (♂) or black (♀). Leg femora and patellae with dark ventral margins (♀). Carapace (Figs. 328, 330, 334, 336). Subrectangular, W / L 1.06 – 1.08; medial surface level along its entire length; anterior margin slightly concave, male with epistomal process; preocular L / carapace L 0.40 – 0.43; surface densely, finely granulate in most areas; granulation more coarse on preocular triangle; granulation much weaker in female; superciliary carinae granulate in male, weakly granulate to smooth in female; anterior margin with several macrosetae; lateral eye groups composed of 3 large and 2 small ocelli (type 5; Loria & Prendini, 2014); median eyes large, eye diameter / carapace L 0.11 (♂), 0.15 (♀). Chelicerae (Figs. 332, 334 – 337). Dorsal surface of manus of male weakly granulate near anterior margin, smooth elsewhere, of female smooth throughout; several macrosetae and pale, fluorescent microsetae near anterior margin; dorsointernal carina strong, weakly granulate in male, smooth in female; fingers with typical buthid dentition (Vachon, 1963), movable finger dorsal margin with two large subdistal denticles and two small basal denticles, ventral margin with subdistal and basal denticles, fixed finger with large subdistal denticle and proximal bicusp, two denticles on ventral surface; dorsal surface of movable finger smooth, with dorsal row of 4 – 5 pale, fluorescent microsetae. Coxosternal area (Figs. 335, 337). All coxae smooth with sparse macrosetae and fluorescent microsetae; sternum smooth, subtriangular, with long medial depression, bearing 2 macrosetae; genital opercula smooth, divided in female. Pectines (Figs. 28, 32, 34 – 35, 323, 325, 327). Basal piece smooth, with deep anteromedian invagination, combs with 3 marginal lamellae, 10 – 12 middle lamellae; combs long, extending to distal limit (♂) or distal 2 / 3 (♀) of trochanter IV; marginal and middle lamellae with numerous small macrosetae, fewer fluorescent microsetae; fulcra with 4 – 6 short setae; female bpt falcate. Hemispermatophore. (Figs. 388 – 392). Flagelliform; trunk narrow, elongate; capsule short, with large, robust, hook-like basal lobe; posterior lobe rounded, with two carinate folds on convex surface; flagellum with short pars recta and pars reflecta (the latter probably incomplete in the examined specimen). Mesosoma (Figs. 332 – 337). Tergites: pretergites smooth, with microsulcate posterior margins; all tergites densely, finely granulated or shagreened, more weakly so in female; tergite I without distinct carinae, tergites II – VI with single weak, granulate median carina, VII with medial hump and 2 pairs of granulate carinae; all tergites lacking macrosetae. Sternites: all sternites smooth, glossy, acarinate; posterior margins of all sternites smooth; spiracles long, narrow, slit-like; sternite III – VI macrosetae: one submedian pair, one lateral pair, two posterior marginal pairs; sternite VII macrosetae: two submedian pairs, three lateral pairs; posteromedian margin of sternite V convex in female, forming a sensory patch with dense narrow, transverse band of microsetae along margin, and a wider, more sparse transverse band of microsetae slightly anterior to margin; posteromedian marginal setation on sternite V denser than on sternites IV and VI. Metasoma (Figs. 137, 372 – 374, 376 – 378). Elongate, segments I – IV uniform in width, segment V narrower posteriorly. Carination: segment I with 10 complete carinae, II with 8 complete carinae (lateral median carinae anteriorly indistinct), segments III – IV with 8 carinae, V with 5 carinae; dorsosubmedian and dorsolateral carinae on segments IIV and lateral median carinae on segments I – III granulate, crenulate or dentate-granulate in both sexes; ventrosubmedian and ventrolateral carinae on segments II – IV granulate or crenulate (♀) or smooth (♂); ventrosubmedian and ventrolateral carinae on segment I weakly crenulate to smooth in both sexes; segment V with dorsolateral carina granulate, ventrolateral and ventromedian carinae granulate with larger dentate granules in posterior half; lateral anal margin with 4 large granules, ventral anal margin with up to 20 granules. Intercarinal surfaces: dorsolateral, lateral, ventrolateral and ventral surfaces of segments I – VI smooth or almost smooth with sparse fine granules; dorsomedian surfaces of all segments smooth; segment V smooth laterally, with sparse coarse and fine granules ventrally. Setation: carinae bear regular series of long macrosetae, 3 – 6 on segments I – IV, up to 10 on segment V; posterior ventral margins of segments I – IV with several long macrosetae. Telson (Figs. 372 – 379). Vesicle dorsal surface smooth; lateral surfaces and ventral surface weakly granulate, with sparse, long macrosetae; vesicle hemi-elliptic or bulbous in lateral profile, without subaculear tubercle; aculeus shorter than vesicle. Pedipalps (Figs. 338 – 371). Femur: dorsointernal, dorsoexternal and ventrointernal carinae strong, coarsely granulate; other carinae indistinct; internal surface with 7 – 10 large granules; intercarinal surfaces smooth; sparse short macrosetae and fluorescent microsetae present. Patella: dorsointernal carina strong, costate-granulate in both sexes, weaker in female; dorsomedian carinae obsolete; dorsoexternal carina of male weakly granulate in distal half, obsolete in proximal half of segment, of female obsolete; ventroexternal and ventromedian carinae of male nearly obsolete, indicated by small granules, of female obsolete; ventrointernal carina granulate in both sexes, weaker in female; internal carina indicated by series of 4 – 5 enlarged dentate granules; sparse short and long macrosetae, and fluorescent microsetae present. Chela: all carinae obsolete, surfaces smooth except for finely granulate internal surface of manus in males; short macrosetae and fluorescent microsetae sparse on manus, dense on fingers; 11 – 12 median denticle ventral surfaces of telotarsi with dense brush of macrosetae subrows on fixed finger, 13 on movable finger including short irregularly arranged, lateral apices with conspicuous fringes subdistal row; subrows flanked by one mid-row internal and of long macrosetae; tarsal ungues stout. two proximal external accessory denticles (except for unfused Measurements. See Table 15. proximal subrow). Trichobothriotaxy: orthobothriotaxic, type Aα (Vachon, 1974), femur d 2 dorsal, e 1 distal to d 5; chela fixed AFFINITIES. Teruelius olgae (Lourenço, 2004) is similar to T. finger db proximal to est. haeckeli sp. n. in adult size, color pattern, and telson shape, Legs (Figs. 328 – 331, 380 – 387). Femora with crenulate but differs in having more slender metasomal segments, a lower ventral carinae; surfaces of all segments smooth; patellae range of male PTC (29 – 33), and a clavate female bpt with long with series of long macrosetae; tibia and tarsal segments curved extension. T. mahafaliensis is similar to T. haeckeli sp. bearing numerous short macrosetae; tibial spurs present on n. in having a higher range of male PTC (34 – 40), but differs legs III – IV; retrolateral tarsal spurs simple, prolateral tarsal in its larger adult size (55 – 60 mm), reddish coloration, lack of spurs basally bifurcate; ventral surfaces of basitarsi with black color on metasoma IV – V and telson vesicle, and a clavate numerous macrosetae arranged roughly in two axial series; female bpt with moderately long, curved extension.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
3E597234FFA9CB186314E648FD5C82A9.taxon	discussion	REMARKS. The convex profile of the posteromedian margin extends slightly anterior to the margin in females. In both of sternite V in females (Fig. 337) differs from the almost sexes, the posteromedian marginal setation is denser on straight posteromedian margin of sternite V in males (Fig. sternite V, than on sternites IV and VI. These anatomical 335). There is also sexual dimorphism of the setation, differences suggest functional specialization. Sternite V is which is strictly confined to the margin in males, but also modified in various other buthids, forming a smooth, pale or fluorescent posteromedian patch with possible glandular or sensory functions. We found differences in the relative density of posteromedian marginal setation of sternites IV – V between Grosphus and Teruelius in several examined species. In females, marginal setation on sternite V in Grosphus was similar in density to that of sternite IV (5 spp., ♀, Figs. 393 – 397), and marginal setation on sternite V in Teruelius was denser than that on sternite IV (5 spp., ♀, Figs. 398 – 402). In males of these species, denser marginal setation on sternite V was absent in G. hirtus, G. madagascariensis, G. simoni, G. voahangyae and T. mahafaliensis, and was present in T. ankarafantsika, T. ankarana, and T. limbatus. Other Teruelius with denser marginal setation on sternite V vs. IV include T. bistriatus (♀) (Lowe & Kovařík, 2019: 93, fig. 437). T. flavopiceus (♂, ♀; modestly so), T. intertidalis (♀), T. ganzhorni (♀) (Ref. MNHN-RS-RS 9080), T. grandidieri (♂, not ♀). These data suggest that dense marginal setation on sternite V is a potential diagnostic character or synapomorphy separating at least some Teruelius from Grosphus. However, we did not include this in our analyses, due to limited taxon sampling and character variability. There was variability in posterior marginal profiles (i. e., convex, linear or concave), and in the density and arrangement of setae. The denser setation could be associated with an increase in numbers of either macrosetae, or fluorescent microsetae. Further investigation of putative glandular or sensory specializations of sternite V, and their sexual dimorphism, is needed to establish character homologies for phylogenetic analysis.	en	Lowe, Graeme, KOvAřík, FRANtIšEk (2022): Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species. Euscorpius 356: 1-105, DOI: 10.5281/zenodo.7475051
