Grosphus Simon, 1880

Lowe, Graeme & Kovařík, František, 2019, Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae), Euscorpius 281, pp. 1-128 : 7-12

Genus Grosphus Simon, 1880

( Figs. 1–4 , 9–12 , 21–22, 25–43 , 52–68 , 86 , 94–98, 106–125 , 133–136 , 145–149 , 158–160 , 165–169 , 181–185 , 196–200 , 211–215 , 227–228 , 231–234 , 239–386 , 580–583 , Tabs. 1–4 )

Grosphus Simon, 1880: 377–378 ; Karsch, 1886: 77; Pocock, 1889a: 348–349; Kraepelin, 1891: 70 (in part); Pocock, 1893: 312 (in part); Kraepelin, 1895: 84 (in part); Kraepelin, 1899: 32 (in part); Kraepelin, 1900: 11–12 (in part); Birula, 1917a: 164 (in part); Birula, 1917b: 55 (in part); Fage, 1929: 640–642 (in part); Werner, 1934: 270 (in part); Vachon, 1969: 483 (in part); Legendre, 1972: 428 (in part); Stahnke, 1972: 130 (in part); Vachon, 1974: 906 (in part); Vachon, 1975: 1598 (in part); Lamoral & Reynders, 1975: 507 (in part); Francke, 1985: 8, 15 (in part); Sissom, 1990: 101 (in part); Lourenço, 1995a: 101 (in part); Lourenço, 1996a: 44; Lourenço, 1996b: 5, 8 (in part); Kovařík, 1998: 109 (in part); Fet & Lowe, 2000: 130 (in part); Lourenço, 2001b: 640 (in part); Prendini, 2001: 16–17, 32, 33–35; Fet et al., 2003: 2, 5–6; Lourenço, 2003a: 577 (in part); Lourenço, 2003c: 153–154 (in part); Lourenço & Goodman, 2003a: 26–27; Soleglad & Fet, 2003a: 26; Soleglad & Fet, 2003b: 19, 66–68, 78–79, 88, 90, 154 (in part); Lourenço, 2004a: 31– 33 (in part); Lourenço et al., 2004: 232 (in part); Prendini, 2004a: 39, 41–42; Prendini, 2004b: 115; Fet et al., 2005: 3, 7–8, 10, 23, 26, 29; Prendini & Wheeler, 2005: 481 (in part); Dupré, 2007: 5, 13, 17 (in part); Lourenço et al., 2007a: 176 (in part); Lourenço et al., 2007b: 369 (in part); Kamenz & Prendini, 2008: 6, 8 (in part); Volschenk et al., 2008: 63 (in part); Kovařík, 2009: 22, 31 (in part); Lourenço et al., 2009b: 145 (in part); Lourenço & Wilmé, 2015: 209, 211; Lourenço et al., 2017: 62; Lourenço et al., 2018b: 74 (in part).

Buthus (Grosphus) : Pocock, 1890: 123 (in part).

TYPE SPECIES. Scorpio (Androctonus) madagascariensis Gervais, 1843 .

DIAGNOSIS. A member of the ‘ Grosphus ’ group differentiated as follows: medium-sized scorpions, adults ca. 25–75 mm in length; pedipalp finger granule rows 11–14 ( Figs. 252 , 286 , 302 , 330 , 376 ), movable finger typically with 4 external subdistal granules; femur trichobothrium d 2 located on internal surface, or straddling dorsointernal carina ( Figs. 9–12 ); 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 ( Figs. 21–22 ); manus trichobothrium V 2 roughly collinear with V 1 along chela axis or slightly displaced internally; lower pectinal tooth counts: ♂ 15–23, ♀ 12–19 ( Figs. 28–31 ); basal pectinal tooth of females wide, oval to subrectangular, not distinctly longer than other teeth ( Figs. 40–43 , 196–200 , 289 ); hemispermatophore capsule long or short, posterior lobe with long, lanceolate extension ( Figs. 52–68 ); sternites with broad ovoid, elliptical or hemi-elliptical spiracles ( Figs. 94–98 ); metasoma I with ventromedian carinae moderately to strongly crenulate or granulate ( Figs. 122–125 ); telson with oval or bulbous vesicle, with or without subaculear tubercle in adults ( Figs. 181–185 ); legs with ventral surface of telotarsus sparsely setose, with two rows of <20 short, setiform macrosetae ( Figs. 133–137 , 211–215 , 259–262 , 316–319 , 362–365 ); telotarsus with dorsal terminal process of normal size; cuticle with weak UV fluorescence ( Figs. 145–149 ).

SUBORDINATE TAXA.

Grosphus ambre Lourenço, Wilmé & Waeber, 2018

Grosphus darainensis Lourenço, Goodman & Ramilijaona, 2004

Grosphus goudoti Lourenço & Goodman, 2006

Grosphus hirtus Kraepelin, 1900

Grosphus madagascariensis ( Gervais, 1843)

Grosphus mayottensis Lourenço & Goodman, 2009 Grosphus polskyi Lourenço, Qi & Goodman, 2007

Grosphus rakotoariveloi Lourenço, Wilmé, Soarimalala & Waeber, 2017

Grosphus tavaratra Lourenço, Soarimalala & Goodman, 2009 Grosphus voahangyae Lourenço & Wilmé, 2015

See Tables 1–3 for diagnostic characters used to place the above taxa under Grosphus .

REMARKS. We consider Grosphus paraphyletic and define two species groups distinguished by major differences in hemispermatophore capsule form: (i)‘ madagascariensis ’ group: capsule elongate, monocarinate, with basal lobe located far proximal to base of flagellum ( G. madagascariensis ); (ii) ‘ hirtus ’ group: capsule short, carination variable, with basal lobe located distally near base of flagellum ( G. goudoti , G. hirtus and G. voahangyae ). Phylogenetic polarity of capsule form is unclear. Possible group affiliations of other species are suggested by some similarities in external characters, e.g.: (i) ‘ madagascariensis ’ group: elliptic spiracles, more elongate metasomal segments, maculation patterns weak or absent, subaculear tubercle small or absent; may include G. ambre , G. mayottensis and G. rakotoariveloi ; (ii) ‘ hirtus ’ group: ovoid spiracles, more stout metasomal segments, stronger maculation patterns, subaculear tubercle more developed; may include G. polskyi , G. tavaratra . However, external characters can be misleading and definitive group assignment requires study of hemispermatophore capsules. For example, G. goudoti resembles species of the ‘ madagascariensis ’ group in external characters (metasoma slender, weak maculation, elliptic spiracles, lack of subaculear tubercle) but possesses a ‘ hirtus ’ group type of capsule.

NEW SYNONYMIES.

Grosphus halleuxi Lourenço, Wilmé, Soarimalala & Waeber, 2017 = Grosphus madagascariensis ( Gervais, 1843) , syn. n.

Grosphus mandena Lourenço, 2005 = Grosphus madagascariensis ( Gervais, 1843) , syn. n.

Grosphus simoni Lourenço, Goodman & Ramilijaona, 2004 = Grosphus madagascariensis ( Gervais, 1843) , syn. n.

The single holotype male of G. madagascariensis used for description by Gervais (1843, 1844) is in poor condition after 176 years. It is disarticulated into several fragments: metasoma III–V + telson, metasoma I–II, hollowed carapace and tergites with most of coxosternal area and sternites III–VI missing, and 4 partial leg fragments (cf. https://science.mnhn. fr/taxon/species/grosphus/madagascariensis). The type locality is given only as ‘Madagascar’. Gervais (1844: pl. XI, figs. 1–3) published a color painting of the dorsal habitus, and drawings of two consecutive metasomal segments in lateral view (segments not specified, but possibly III–IV), showing enlarged spiniform granules on posterior dorsal carinae, and the carapace with median and lateral eyes. With only limited information available about the holotype, which has lost many body parts bearing key taxonomic characters, it is difficult to precisely pin down the identity of G. madagascariensis in relation to a group of several other currently-named similar taxa (i.e., G. darainensis , G. halleuxi , G. mandena , G. rakotoariveloi and G. simoni ). Previous diagnoses of Kraepelin (1900), Fage (1929) and Lourenço (1996b) listed some characters that differentiate G. madagascariensis from G. hirtus or Teruelius gen. n. Meristic characters were: PTC ♂ 18–20, ♀ 16–18, and pedipalp movable finger granule rows 12. Lourenço & Goodman (2006) suggested that Goudot, collector of the holotype, travelled in the north eastern region. They selected a male and female from Forêt de Plateau de Makira, in humid northeastern forest near Antongil Bay, as reference material for a redescription. The redescription is generic for the group, with few diagnostic characters: PTC ♂ 20, ♀ 15–16, pedipalp movable finger granule rows 13. Lateral eyes were incorrectly cited only as only 3 pairs, contradicting Gervais (1844).

Grosphus simoni was described by Lourenço, et al. (2004) from two specimens: the holotype male from Forêt de Plateau de Makira, Forêt de Sahantaha, which is humid tropical forest in the northeast; and a paratype male from Station Forestière d’Ampijoroa, Ankarafantsika National Park, which is dry deciduous forest in the northwest. The differential diagnosis was brief: paler coloration, metasoma with strong granules and carinae, including several larger posterior spiniform granules on dorsal carinae on segments II–IV. Recently, Lourenço, et al. (2017) moved the paratype to a different species, G. rakotoariveloi , invalidating the original diagnosis of G. simoni based on both specimens. They revised the diagnosis of G. simoni as: moderately darker coloration, several larger posterior spiniform granules on dorsal carinae of metasoma II–IV, PTC ♂ 15–17, ♀ 14–15, pedipalp finger granule rows ♂ 11–12, ♀ 12– 13, male chela with weak to moderate scalloping. Photographs were included for G. simoni specimens of both sexes from Forêt de Sahantaha (figs. 2–5), although the male was misidentified as a female, and the female misidentified as a male. The holotype male of G. simoni is well documented in high resolution images published on the FMNH website: https://collections-zoology. fieldmuseum.org/catalogue/963985. We studied FMNH materials (5♂, 1♀) from Andasibe determined as G. simoni .We further compared other materials, including a male determined by M. Vachon as G. madagascariensis (MHNG) . We found no convincing diagnostic characters to support a distinction between G. simoni and G. madagascariensis . Diagnostic characters for G. simoni involve relatively minor differences in darker vs. lighter shades of color, differences in size of spiniform granules on dorsal metasomal segments, meristic differences of one or two pectine teeth with contiguous or overlapping ranges of PTC, and/ or pedipalp finger granule row counts. These characters are subject to inter-population and geographic variation in many scorpion taxa. Allowing for typical genetic variation, the metasoma and telson of holotypes of G. simoni and G. madagascariensis do not differ significantly in carination, spination or morphometrics. In the absence of quantitative analysis showing discontinuous variation either in characters or morphometrics to support splitting into discrete species, we regard them as synonyms. Localities of G. simoni overlap or are sympatric with the distribution for G. madagascariensis in northeastern humid forests.

The species G. rakotoariveloi has meristics (PTC ♂ 18– 19, pedipalp granule rows ♂ 13–14) that also overlap or are contiguous with those of G. madagascariensis . However, the type (and only known) locality is in a different bioclimatic region with dry deciduous forest, disjunct from eastern humid forests. It has much lighter coloration and relatively wide pedipalp chelae. We provisionally list this species, until additional data are available.

Grosphus halleuxi was described by Lourenço, et al. (2017) from a series of males from Torotorofotsy Forest, ca. 20 km NW of Moramanga, in a central humid forest area that is locally less humid than other eastern forests. The diagnostic characters for differentiating it from G. simoni were: darker coloration, smaller size of 55 mm, PTC ♂ 16–19, pedipalp granule rows ♂ 11–12, and weaker scalloping of pedipalp fingers. The meristic counts do not yield a differential diagnosis as they overlap those of G. simoni (= G. madagascariensis ). We analyzed a series of near topotypic specimens (5♂, 6♀) from Moramanga and ca. 30 km E of Moramanga, whose males closely match photos of the G. halleuxi male holotype ( Figs. 1–2 , cf. Lourenço, et al., 2017: figs. 16–17). We obtained meristics: PTC ♂ 15–18, ♀ 13–15, pedipalp finger granule rows ♂ ♀ 12, which are not distinguishable from meristic ranges of G. madagascariensis . Other characters of darker or lighter shades, and degree of pedipalp finger scalloping can also be variable between populations. For example, in Figs. 1 & 3 , a female from Moramanga area is darker than a male from the same area, showing that intensity of coloration varies within the same population, weakening this diagnostic character. G. halleuxi was diagnosed as “much darker”, but ‘ G. simoni ’ appears as dark, if not darker (Lourenço, et al., 2017: figs. 2–5 vs. 16–17). It was argued that G. halleuxi is a narrow-ranged species adapted to a less humid local microclimate. The existence of local microendemic taxa should be supported by strong diagnostic characters. Until such characters are defined, we regard this species as a local population of G. madagascariensis .

Grosphus mandena was described by Lourenço (2005) from near Fort Dauphin, in the Mandena region of southeastern coastal rainforest. Differential diagnostic characters were: lighter coloration, weaker metasomal carination, one larger spiniform granule on metasoma II–IV, and a more granulated telson. Meristics were: PTC ♂ 19–20, ♀ 15–17, pedipalp granule rows ♂ 12–13. As discussed above, these characters fall within ranges of variation for G. madagascariensis , including local populations given other species names. We loaned and studied the male holotype and a female paratype from MHNG and found them to be indistinguishable from G. madagascariensis . We note that the female paratype is considerably darker than the male holotype. This shows that intensity of coloration varies even within the type population, and is not a reliable diagnostic character ( Figs. 348–351 , 580–583 ). Lourenço & Wilmé (2016: fig. 36) showed a nonoverlapping discontinuous transition, at latitude ca. 22°S between the northern range of G. madagascariensis and the southern range of G. mandena . The transition latitude does not correspond to any boundary between centers of endemism as defined by the watershed model of Wilmé et al. (2006). Lourenço et al., (2009b) suggested that the disjunction is recent, due to extirpation of south littoral rainforest by humans. More robust diagnostic characters and analysis of clinal vs. discontinuous variation is necessary to delimit the southern populations as a distinctive species. Until such characters are defined, we regard this species as a southern population of G. madagascariensis . The synonymies of G. halleuxi and G. mandena with G. madagascariensis are further supported by their identical hemispermatophores, all of which have a unique, elongated capsule architecture with a proximal basal lobe (cf. Figs. 52–53, 56–57 vs. Figs. 54–55 ).

Grosphus hirtus garciai Lourenço, 2001 = Grosphus hirtus Kraepelin, 1900 , syn. n.

Grosphus garciai was described by Lourenço (2001b) from Station Forestière d’Ampijoroa, Ankarafantsika National Park, based on an adult male holotype and a juvenile, collected by García Herrero. In the diagnosis, it was differentiated from G. madagascariensis , a quite different species, by: smaller size, maculated light and dark pigmentation, pedipalp granule rows ♂ 13, weaker spiniform granules on pedipalp and metasoma, and weaker scalloping of pedipalp fingers. Curiously, it was not compared to G. hirtus , which bears a much greater similarity. Subsequently, Lourenço & Goodman (2006) redescribed G. hirtus based on material also from Station Forestière d’Ampijoroa, Ankarafantsika National Park, also collected by García Herrero. G. hirtus was separated from G. garciai by yellowish rather than reddish brown color, and larger size (40–50 mm vs. 32 mm). Subsequently, Lourenço & Wilmé, (2015a) downgraded G. garciai to the status of subspecies, G. hirtus garciai , supposedly a microendemic taxon in a “local isolated population”. It is morphologically identical to, and differentiated from the nominotypical G. hirtus only by smaller size. The subspecies is known only from the type locality and the nominotypical G. hirtus also occurs in the same area ( Lourenço & Goodman, 2006). The species G. hirtus is distributed more widely over northwest Madagascar ( Lourenço & Wilmé, 2016: fig. 36). We loaned and studied the holotype of G. garciai , as well as male and female topotypes from FMNH. We confirmed that it is morphologically indistinguishable from G. hirtus ( Figs. 263–305 ). We question whether an animal population of somewhat smaller average body size compared to closely neighboring conspecifics, but otherwise identical to them, merits subspecies status. Local size variations of species may be caused by varying environmental conditions that limit growth rates and development. The size differential is exaggerated by the reported body lengths, a measurement that can vary with mesosomal expansion: G. hirtus 40–50 mm ( Lourenço & Goodman, 2006); G. h. garciai 28–32 mm ( Lourenço & Wilmé, 2015a). These numbers imply that G. hirtus is at least 25% longer. A more reliable size comparison for morphometrically similar scorpions would use carapace length. For example, we measured carapaces of G. hirtus : ♂ 5.5 mm, ♀ 5.0 mm, vs. G. h. garciai : ♂ 5.2 mm, ♀ 4.4 mm, i.e., only 6–12% longer. Lourenço et al. (2007a) redescribed G. hirtus with body lengths of ♂ 34.3 mm, ♀ 31.8 mm, and carapace lengths ♂ 4.1 mm, ♀ 4.3 mm. These are small enough to overlap measurements for G. h. garciai . Considering this overlap, we regard G. h. garciai as a synonym of G. hirtus .

Grosphus polskyi is another species that is quite similar to G. hirtus . It was diagnosed by having paler color, with weaker, more diffuse maculate patterns restricted to carapace and tergites, slightly more elongate metasoma segment I, weak spination on dorsal metasomal carinae, and a slightly larger subaculear tubercle. The only known record is the single male holotype from Mikea Forest near Toliara, on the southwestern coast. Although this is quite far south of the southern-most record of G. hirtus ( Lourenço & Wilmé, 2016) , records of the latter are sparse, so it is unclear if it represents a disjunction. We provisionally list this species, until it can be critically evaluated by more material and analysis of variation.

Figures

Figures 1–4. Grosphus madagascariensis. Type species of Grosphus Simon, 1880. Habitus. Male (1–2) and female (3–4), in dorsal (1, 3) and ventral (2, 4) views. Scale bar: 10 mm.

Figures 9–20. Position of femur trichobothrium d 2in Grosphus and Teruelius gen. n. Dorsal surfaces of proximal pedipalp femur of adult females (♀, left panels) and adult males (♂, right panels), underUV fluorescence to highlightgranulation,carinae and trichobothrial areolae.Whitearrows indicate positions of trichobothrium d in each image. G. madagascariensis (9, 10; 2 samples from Anjiro (9) and Andasibe (10) show consistency of d 2 2 position),G. hirtus(11), G. voahangyae (12), T. ankarafantsika (13, 15; 2 samples fromAmpijoroa show consistency ofd 2position),T. ankarana (14), T. flavopiceus (16), T. grandidieri (17), T. limbatus (18), T. mahafaliensis (19), and T. olgae (20). Scale bars: 1 mm (9–20♀) or 500 μm (20 ♂).

Figures 21–27. Positions of trichobothria Eb, Eb and petite Eb on manus of pedipalp chela in Grosphus, Neogrosphus and Teruelius gen. 1 2 3 n. Figures 21–24. External views of pedipalp chela, shown under UV fluorescence to highlight trichobothrial areolae. G. hirtus ♂ (21), G. madagascariensis ♀ (22), T. limbatus ♂ (23), T. ankarafantsika ♀ (24). Positions of Eb 1, Eb 2 and Eb 3 and lines joining them shown as white overlays. Scale bars: 2 mm. Figures 25–26. Horizontal histograms comparing ratios, R = d(Eb, Eb)/ d(Eb, Eb), of Eb -Eb distance, to Eb - 123 2 3 1 2 2 3 1 Eb distance, in males (25), and females (26) of Grosphus, Neogrosphus and Teruelius gen. n. Error bars indicate ranges (minimum, maximum), 2 histogram bars mid-range values. Figure 27. Scatter plot of male vs. female ratios R. Ratio is larger in males if points fall above the diagonal 123 (gray) line, larger in females if they fall below it.

Figures 28–29. Variation in pectinal tooth count (PTC) in Grosphus, Neogrosphus and Teruelius gen. n. Horizontal histograms comparing PTC in males (28), and females (29) of Grosphus, Neogrosphus and Teruelius gen. n. Error bars indicate ranges (minimum, maximum), bars mid-range values.

Figures 30–35. Variation in pectinal tooth count (PTC) in Grosphus, Neogrosphus and Teruelius gen. n. Figures 30–31. Logarithmic scatter plots showing scaling of PTC vs. carapace length in males (30), and females (31) of Grosphus, Neogrosphus and Teruelius gen. n. Color lines are least squares linear regression fits of scaling trends for the different genera. Figures 32–33. Logarithmic scatter plots showing scaling of PTC vs. carapace length in Figs. 30–31 superposed upon overall scaling trends for a larger sample of male (32) and female (33) buthids including both α and β trichobothrial configurations (gray symbols; 1055♂, 1046♀). Dark lines: least squares linear regression fits of scaling trends for larger sample of buthids. Figures 34–35. Logarithmic scatter plots showing scaling of PTC vs. carapace length in Figs. 30–31 superposed upon overall scaling trends for a larger sample of male (34) and female (35) α-buthids restricted to α trichobothrial configurations (gray symbols; 445♂, 371♀). Dark lines: least squares linear regression fits of scaling trends for α-buthids. Numbers: slopes of regression lines (= allometric exponents). Purple symbols and lines: outgroup Pseudolychas.

Figures 36–39. Comparative biometrics of female basal pectinal tooth and hemispermatophore trunk of Grosphus, Neogrosphus and Teruelius gen. n. Figures 36–37. Logarithmic scatter plots showing scaling of length of female basal pectinal tooth vs. carapace length (36), and hemispermatophore trunk length vs. carapace length (37). Color lines: least squares linear regression fits of scaling trends for the different genera. Figures 38–39. Logarithmic scatter plots of female basal pectinal tooth length vs. hemispermatophore trunk length (38), and the ratio of basal pectinal tooth length/ hemispermatophore trunk length (%) vs. hemispermatophore trunk length, showing fraction of trunk spanned by the female tooth (39). Female basal pectinal tooth length measured as chord length, from more proximal corner (along comb axis) at base of tooth, to more distal corner (along comb axis) at tip of tooth. Trunk length measured from proximal edge of basal lobe to pedicel (foot). Color lines: least squares linear regression fits of scaling trends for the different genera.

Figures 40–51. Female basal pectinal teeth in Grosphus and Teruelius gen. n. Ventral views of proximal left pectine of females shown under UV fluorescence to highlight cuticular surface texture, setation and absence of peg sensillae on basal tooth vs. their presence on other teeth. G. sp. nr hirtus (40), G. madagascariensis (41), G. hirtus (42), G. voahangyae (43), T. ankarafantsika (44–45; 2 samples fromAmpijoroa show variation in tooth shape), T. ankarana (46), T. flavopiceus (47), T. grandidieri (48), T. limbatus (49), T. mahafaliensis (50), and T. olgae (51). Scale bars: 1 mm.

Figures 52–70. Hemispermatophores and capsule regions of Grosphus and Neogrosphus. Multi-panel figures show: whole hemispermatophore; whole hemispermatophore and capsule with flagellum; capsule region in convex (and/ or convex compressed), anterior and posterior views (panels in left to right sequence). Right hemispermatophores unless indicated as mirrored left images. Figure 52. G. madagascariensis, whole hemispermatophore (scale bar: 2 mm), capsule and flagellum (scale bar: 1 mm). Figure 53. G. madagascariensis, capsule, Sc1197, Andasibe, GLPC, FKCP. Scale bar: 500 μm. Figure 54. G. madagascariensis, capsule, Anjiro, G. halleuxi nr topotype, GLPC. Scale bar: 500 μm. Figure 55. G. madagascariensis, capsule, Mandena- Fort Dauphin, G. mandena paratype, MHNG.Scale bar: 500 μm.Figure 56. G. madagascariensis, capsule, Madagascar, det. Vachon, MHNG. Scale bar: 500 μm. Figure 57. G. madagascariensis, capsule, Andasibe, GLPC, FKCP. Scale bar: 500 μm. Figure 58. G. goudoti, whole hemispermatophore. Scale bar: 2 mm. Figure 59. G. goudoti, capsule, Forêt de Bobankota, holotype, MHNG. Scale bar: 500 μm. Figure 60. G. hirtus, whole hemispermatophore (scale bar: 2 mm), capsule and flagellum (scale bar: 1 mm), Antsiranana, Ramena vill., mirrored left, GLPC, FKCP. Figure 61. G. hirtus, capsule, Mahajamba River, GLPC, FKCP. Scale bar: 500 μm. Figures 62–65. G. hirtus, capsules in convex view.Antsiranana, Ramena vill., mirrored left, GLPC, FKCP (62), Jardin Botanique, MHNG (63), Ranohira-Llakaka, ZMUH, mirrored left (64), Forêt de Vohitaly, MHNG (65). Scale bars: 500 μm. Figure 66. G. hirtus, capsule, Forest Station Ampijoroa, G. garciai holotype, MHNG. Scale bar: 500 μm. Figure 67. G. voahangyae, whole hemispermatophore. Scale bar: 2 mm. Figure 68. G. voahangyae, capsule, Analamy Forest, FMNH. Scale bar: 500 μm. Figure 69. N. griveaudi, whole hemispermatophore (flagellum truncated). Scale bar: 2 mm. Figure 70. N. griveaudi, capsule, mirrored left, Tsimanampetsotsa National Park, GLPC, FKCP.

Figures 86–89. Hemispermatophores and capsule regions of Grosphus and Neogrosphus. Cross stereoscopic convex views. Scale bars: 500 μm. Figure 86. G. madagascariensis, Sc1197, Andasibe, GLPC, FKCP. Figure 87. N. griveaudi, mirrored left, Tsimanampetsotsa National Park, GLPC, FKCP. Figure 88. G. goudoti, Forêt de Bobankota, holotype, MHNG. Figure 89. G. voahangyae, Analamy Forest, FMNH.

Figures 94–105. Spiracles on sternite IV in Grosphus and Teruelius gen. n. Spiracle on right side of sternite IV in adult females (♀, left panels) and adult males (♂, right panels), shown under UV fluorescence. G. madagascariensis (94 from Anjiro, 95 from Andasibe), G. hirtus (96), G. hirtus (= G. h. garciai) (97), G. voahangyae (98), T. ankarafantsika (99), T. ankarana (100), T. flavopiceus (101), T. grandidieri (102), T. limbatus (103), T. mahafaliensis (104), and T. olgae (105). Scale bars: 200 μm (96–98), 400 μm (94–95, 99, 103–105) or 1 mm (100–102).

Figures 133–144. Ventral setation of telotarsus III in Grosphus and Teruelius gen. n. Ventral surfaces of right telotarsus III of adult males (♂) or females (♀), shown under UV fluorescence to highlight setation. Macrosetae appear dark with strongly fluorescent cuticular sockets at their base. Putative chemosensory microsetae appear bright. G. hirtus (= G. h. garciai) (133), G. hirtus (134), G. madagascariensis (135, from Anjiro, 136 from Andasibe), G. voahangyae (137), T. ankarafantsika (138), T. limbatus (139), T. mahafaliensis (140), T. ankarana (141), T. grandidieri (142), T. flavopiceus (143), and T. olgae (144). Scale bars: 500 μm (133–140, 144) and 1 mm (141–143).

Figures 145–157. Comparative intensity of UV fluorescence in Grosphus and Teruelius gen. n. Photographic comparison of fluorescence emission intensities of representative species of each genus, including adult males (♂) or females (♀). G. madagascariensis (145 from Anjiro, 148 from Andasibe), G. hirtus (146), G. voahangyae (147), G. hirtus (= G. h. garciai) (149), T. ankarafantsika (150), T. grandidieri (151), T. olgae (152), T. flavopiceus (153), T. ankarana (154–155), T. mahafaliensis (156), and T. limbatus (157). Images acquired under identical intensities of UV excitation (395 nm LED source, 0.35 A current) and camera exposure (Canon EOS 7D Mark II, 100 mm f/13, 0.5 s, ISO 320), with 475 nm longpass filter to block excitation wavelengths. Scale bars: 10 mm (145–150, 152–153, 156–157), 20 mm (151, 154–155).

Figures 158-160. Comparative intensity of UV fluorescence in Grosphus and Teruelius gen. n. Figure 158. Horizontal histograms comparing relative fluorescence emission intensities measured by photodiode current, for Grosphus and Teruelius species.Histogram bars are means, error bars are SE, species averages over all samples, male and female (sexes not significantly different, P> 0.05). Sample sizes: G. hirtus, 1♂, 1♀; G. sp. nr hirtus, 6♀; G. madagascariensis, 9♂, 1♀; G. voahangyae, 4♂, 1♀; T. ankarafantsika, 2♂, 2♀; T. ankarana, 1♂, 1♀; T. flavopiceus, 1♂, 1♀; T. grandidieri, 1♂, 1♀; T. limbatus, 6♂, 5♀; T. mahafaliensis, 2♂, 2♀; T. olgae, 1♂, 1♀. Sample means may not reflect single specimen intensities recorded in Figs. 145–157. Figure 159. Time course of photodiode current in UV fluorescence emission measurement for T. limbatus (upper curve), and two specimens of G. madagascariensis (middle and lower curves) over 620 s. Arrow indicates time of shutter opening to initiate UV excitation.160. Photobleaching time course for UV fluorescence emission of T. limbatus (middle curve) and G. madagascariensis (upper and lower curves) over 600 s following shutter opening (see Fig. 159). Ordinate is percentage bleaching relative to initial fluorescence after shutter opening. Blue curves are mono-exponential fits with decay time constants: T. limbatus, 640.2 s; G. madagascariensis, #1 654.4 s and #2 347.5 s.

Figures 165–180. Carapace of representative Grosphus and Teruelius gen. n. G. madagascariensis (165, 168), G. hirtus (166–167), G. voahangyae (169), T. ankarafantsika (170–171), T. ankarana (172), T. flavopiceus (173–174), T. grandidieri (175), T. limbatus (176), T.mahafaliensis (177–178) andT. olgae (179–180).UV fluorescence,♂male, ♀female.Scale bars:2mm (165–171, 176–180),4 mm (172–175).

Figures 181–195. Telson of representativeGrosphus andTeruelius gen. n. G. hirtus (181–182), G. madagascariensis (183–184), G. voahangyae (185), T. ankarafantsika (186–187), T. ankarana (188), T. flavopiceus (189), T. grandidieri (190–191), T. limbatus (192), T. mahafaliensis (193) and T. olgae (194–195). UV fluorescence, ♂ male, ♀ female. Scale bars: 2 mm (181–187, 192–195), 4 mm (188–191).

Figures 196–210. Female basal pectinal teeth of representative Grosphus and Teruelius gen. n. G. hirtus (196–198), G. madagascariensis (199), G. madagascariensis, paratype of G. mandena(200), T. ankarafantsika (201),T. ankarana (202), T. annulatus (203),T. bistriatus (204), T. flavopiceus (205), T. grandidieri (206), T. intertidalis (207), T. limbatus (208), T. mahafaliensis (209), T. feti (210).

Figures 211–226. Ventral tarsal setation of legs III or IV in Grosphus and Teruelius gen. n. G. goudoti (211), G. hirtus (212), G. hirtus (G. garciai) (213), G. madagascariensis (214), G. madagascariensis (G. mandena) (215), Teruelius ankarana (216), T. flavopiceus (217), T. annulatus (218), T. ankarafantsika (219–220), T. bistriatus (221), T. intertidalis (222), T. grandidieri (223), T. mahafaliensis (224), T. limbatus (225), T. feti (226).

Figures 227–230. Right lateral eyes of Grosphus and Teruelius gen. n. G. madagascariensis (227), G. voahangyae (228), T. limbatus (229) and T. flavopiceus (230). All species comply with the 5-eye buthid pattern with series of 3 larger ocelli in lower position, and two smaller ocelli in posterior and upper positions. UV fluorescence, males. Scale bars: 500 μm.

Figures 231–238. Males. Right chelicera. Figures 231–234. Grosphus madagascariensis, dorsal (231, 233) and ventral (232, 234) views, under white light (231–232) and UV fluorescence (233–234). Figures 235–238. Teruelius limbatus, dorsal (235, 237) and ventral (236, 238) views, under white light (235–236) and UV fluorescence (237–238). Scale bars: 1 mm.

Figures 239–243. Grosphus goudoti, male holotype, dorsal (239) and ventral (240) views, sternopectinal region and sternites III–IV (241), carapace and tergites I–III (242), and original label (243). Scale bar: 10 mm (239–240).

Figures 244–258. Grosphus goudoti, male holotype, pedipalp chela, dorsal (244), external (245), and ventrointernal (246) views; pedipalp patella, dorsal (247), external (248), and ventral (249) views; pedipalp femur and trochanter, internodorsal (250) and dorsal (251) views; pedipalp chela, movable finger dentate margin (252); right chelicera, dorsal (253) and ventral (254) views; telson lateral view (255); metasoma and telson, lateral (256), dorsal (257) and ventral (258) views.

Figures 259–262. Grosphus goudoti, male holotype, distal segments of right legs I–IV, retrolateral views.

Figures 263–266. Grosphus hirtus, habitus. Male (263–264) and female (265–266), in dorsal (263, 265) and ventral (264, 2662) views. Scale bar: 10 mm.

Figures 267–274. Grosphus hirtus, telson lateral view in male (267) and female (268); metasoma and telson in male and female, lateral (269, 272), ventral (270, 273) and dorsal (271, 274) views. Scale bar: 10 mm (269–274).

Figures 275–290. Grosphus hirtus. Figures 275–277, 289–290. Female, pedipalp chela, dorsal (275) and external (276) views, pedipalp femur and trochanter, internodorsal (277) view; sternopectinal region and sternite III (289) and carapace and tergites I–III (290). Figures 278–288. Male, pedipalp chela, dorsal (278), external (279) and ventrointernal (280) views; pedipalp patella, dorsal (281), external (282) and ventral (283) views; pedipalp femur and trochanter, internodorsal (284) and dorsoexternal (285) views; pedipalp chela, movable finger dentate margin (286); sternopectinal region and sternite III (287) and carapace and tergites I–III (288).

Figures 291–292. Grosphus hirtus, male holotype of G. garciai, dorsal (291) and ventral (292) views. Scale bar: 10 mm.

Figures 293–305. Grosphus hirtus. Figure 293. Juvenile paratype of G. garciai, pedipalp chela dorsal. Figures 294–305. Grosphus hirtus, male holotype of G. garciai. Figures 294–302. Left pedipalp (mirrored), pedipalp chela, dorsal (294), external (295) and ventrointernal (296) views; pedipalp patella, dorsal (297), external (298) and ventral (299) views; pedipalp femur and trochanter, internodorsal (300) and dorsal (301) views; pedipalp chela, movable finger dentate margin (302). Figures 303–305. Carapace and tergites I–III (303), sternopectinal region and sternites III–VI (304) and telson lateral view (305).

Figures 306–309. Grosphus madagascariensis. Habitus. Male (306–307) and female (308–309), dorsal (306, 308) and ventral (307, 309) views. Scale bars: 10 mm..

Figures 310–315. Grosphus madagascariensis. Metasoma and telson. Male (310–312) and female (313–315), dorsal (310, 313), lateral (311, 314) and ventral (312, 315) views. Scale bars: 4 mm.

Figures 316–319. Grosphus madagascariensis, male, right legs I–IV tibia, basitarsus and telotarsus, retrolateral views. Scale bar: 2 mm.

Figures 320–340. Grosphus madagascariensis. Pedipalp. Male (320–330) and female (331–340). Chela in dorsal (320, 331), external (321, 332) and ventrointernal (322, 333) views. Patella in dorsal (323, 334), external (324, 335) and ventral (325, 336) views. Femur and trochanter in internal (326, 337), dorsal (327, 338), ventral (329, 339) and external (329, 340) views. Pedipalp chela, movable finger dentate margin (330). Scale bar: 4 mm.

Figures 341–347. Grosphus madagascariensis. Pedipalp. Male. Chela in dorsal (341), external (342) and ventrointernal (343) views. Patella in dorsal (344) and external (345) views. Femur and trochanter in internal (346) and dorsoexternal (347) views. The trichobothrial pattern is indicated by white circles.

Figures 348–351. Grosphus madagascariensis. Habitus. Male holotype (348–349) and female paratype (350–351) of Grosphus mandena in dorsal (348, 350) and ventral (349, 351) views. Scale bar: 10 mm.

Figures 352–355. Grosphus voahangyae. Habitus. Male (352–353) and female (354–355), dorsal (352, 354) and ventral (353, 355) views. Scale bars: 5 mm.

Figures 356–361. Grosphus voahangyae. Metasoma. Male (356–358) and female (359–361), dorsal (356, 359), lateral (357, 360) and ventral (358, 361) views. Scale bars: 2 mm.

Figures 362–365. Grosphus voahangyae, male, left legs I–IV tibia, basitarsus and telotarsus, retrolateral views. Leg I (362), leg II (363), leg III (364), leg IV (365). Scale bar: 1 mm.

Figures 366–386. Grosphus voahangyae. Pedipalp. Male (366–376) and female (377–386). Chela in dorsal (366, 377), external (367, 378) and ventral (368, 379) views. Patella in dorsal (369, 380), external (370, 381) and ventral (371, 382) views. Femur and trochanter in internal (372, 383), dorsal (373, 385), external (375, 386) and ventral (374, 384) views. Pedipalp chela, movable finger dentate margin (376). Scale bars: 2 mm (366–375, 377–386).

Figures 580–581. Grosphus madagascariensis. Male and female, in vivo habitus (580) and female with newborns (581).

Figures 582–583. Grosphus madagascariensis. Female with newborns (582) and with juveniles after first ecdysis (583).

Figures 122–126. Ventral aspect of metasoma and telson of adult males (♂) or females (♀) of representative Grosphus spp. and Teruelius limbatus under UV fluorescence to reveal carination and granulation. G. hirtus (122), G. madagascariensis (123 from Anjiro, 124 from Andasibe), G. voahangyae (125), and T. limbatus (126). Scale bars: 4 mm.