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 : 5

Diagnosis of the ‘ Grosphus ’ group

The three buthid genera Grosphus Simon, 1880 , Neogrosphus Lourenço, 1995 , and Teruelius gen. n. comprise a distinct assemblage of Madagascar buthids (= ‘ Grosphus ’ group) sharing the following set of characters:

Carapace subrectangular, weakly trapezoidal or nearly parallel-sided, surface densely granular, carinae indistinct except for superciliary carinae; frontal region of carapace flat, not sloped towards anterior margin; median eyes large, median ocular tubercle prominent, located forward of the carapace centroid ( Figs. 165–180 ); 5 pairs of lateral eyes (3 large, 2 small) ( Figs. 227–230 ); chelicerae with typical buthid dentition on fixed and movable fingers ( Vachon, 1963), two enlarged denticles on ventral surface of fixed finger ( Figs. 231– 238 ); sternum type 1, subtriangular; tergites granular, tergites I–VI with single, weak median carina, tergite VII with weak median carina and 2 pairs of strong lateral carinae; metasoma moderately elongate, segments I–III with 8–10 carinae, IV with 8 carinae, V with 3–5 carinae; telson vesicle bulbous, ovoid or elongate, with or without subaculear tubercle ( Figs. 181–195 ); pectines with fulcra, 13–41 teeth, female with basal pectinal tooth dilated or elongated, lacking peg sensillae ( Figs. 40–51 , 196–210 ); hemispermatophore flagellum thicker at base, narrowed proximally, thickened distally ( Figs. 52, 58, 60, 67 , 71, 75, 78, 84 ); pedipalp chela elongate, smooth, carinae obsolete, surface typically with numerous short macrosetae ( Figs. 21–24 ); finger dentition composed of 8–15 discrete linear rows of granules or denticles, each slightly oblique with proximal ends directed externally; rows either non-overlapping or slightly imbricated, proximal 3 granules in each row enlarged, 2 of these slightly displaced outwards as ‘external accessory’ granules; series of large, dentate internal accessory granules present, offset from main rows; both chela fingers with enlarged apical teeth, 3–4 external subdistal granules; pedipalps sexually dimorphic, dentate margins of fingers weakly or strongly scalloped proximally in males, straight in females, manus of males broader than that of females; trichobothrial pattern orthobothriotaxic, type A ( Vachon, 1974), with femur d 1 - d 3 - d 4 in α-configuration ( Vachon, 1975), patella d 3 external to dorsomedian carina ( Fet et al., 2005); patella em much closer to est and et, than to esb 1 and esb 2, with em -est -et usually forming a compact triad ( Figs. 345 , 481a ); chela manus with Eb 1 - Eb 2 angled distally, Eb 1 - Eb 2 - Eb 3 acute angle opening in proximal direction (γ-configuration) ( Figs. 342 , 478a ); chela with db in proximal half to middle of fixed finger; legs III–IV with tibial spurs ( Figs. 211–226 , 261–262 , 318–319 , 364–365 , 414–417 , 487–488 , 514–515 , 540–541 , 578–579 , 618–619 ), tarsi without bristle-combs.

REMARKS. In describing the first ‘ Grosphus ’ group species, Scorpio (Androctonus) madagascariensis, Gervais (1844 : pl. XI, fig. 3) illustrated the carapace showing forward placement of the median eyes, and also accurately depicted five pairs of lateral eyes, now recognized to be the prevalent buthid configuration ( Loria & Prendini, 2014; Yang et al., 2013). In spite of this, Fage (1929) incorrectly declared that Grosphus (sensu lato) only bore 3 pairs of lateral eyes, and Lourenço (1996b) cited only 3–4 pairs. Moreover, only 3 pairs were described for: Grosphus ambre , G. darainensis , G. garciai , G. goudoti , G. halleuxi , G. hirtus , G. madagascariensis , G. makay , G. mandena , G. mayottensis , G. polskyi , G. rossii , G. simoni , G. rakotoariveloi , G. tavaratra , G. voahangyae , Teruelius ankarana , T. ankarafantsika , T. bemaraha , T. bicolor , T. bistriatus , T. eliseanneae , T. feti , T. ganzhorni , T. intertidalis , T. limbatus , T. magalieae , T. mahafaliensis , T. olgae , T. sabineae , T. waeberi ( Lourenço, 1996b, 1999, 2001b, 2003c, 2005, 2012c, 2013b, 2014; Lourenço & Goodman, 2006, 2009; Lourenço & Wilmé, 2015a, 2015b, 2016; Lourenço et al., 2004, 2007a, 2009b, 2016c, 2017, 2018b). We confirm here that 5 pairs are indeed present in all species that we have examined: G. garciai (= G. hirtus ), G. goudoti , G. ‘ halleuxi ’, G. hirtus , G. sp. nr hirtus , G. madagascariensis , G. ‘ mandena ’, G. voahangyae , Neogrosphus griveaudi , Teruelius ankarafantsika , T. ankarana , T. annulatus , T. bistriatus , T. feti , T. flavopiceus , T. grandidieri , T. intertidalis , T. limbatus , T. mahafaliensis and T. olgae (e.g., Figs. 227–230 ). We found only a few individual deviations from the standard pattern, such as 2 large and 2 small ocelli, that we regarded as developmental anomalies. We predict that other ‘ Grosphus ’ group species will also comply with the 5-eye pattern. Although undercounting of lateral eyes is perhaps attributable to overlooking of the smaller posterior and upper ocelli, 10 of the published 3-eye counts post-date introduction of the 5-eye model by Yang et al. (2013, coauthor Lourenço) and Loria & Prendini (2014). Paradoxically, Lourenço et al. (2007a) claimed 3 lateral eyes in boilerplate descriptions of G. hirtus and G. polskyi , yet their figures clearly depict all 5 lateral eyes as being present in both species. Vachon (1969) correctly reported 5 “nettement visibles” lateral eyes, 3 large and 2 small, in both sexes of Neogrosphus griveaudi . Although 3 pairs were described for N. blanci and N. andrafiabe ( Lourenço, 1996b; Lourenço et al., 2015), we are skeptical that these counts are accurate.

Figures

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 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 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 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 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 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 71–85. Hemispermatophores and capsule regions of Teruelius gen n. Multi-panel figures show: whole hemispermatophore; whole hemispermatophore and/or capsule with flagellum; capsule region in convex (or convex compressed), anterior and posterior views (panels in left to right sequence). Right hemispermatophores. Figure 71. T. ankarana, whole hemispermatophore. Scale bar: 4 mm. Figure 72. T. ankarana, capsule, left mirrored, Forêt d’Ankavanana, FMNH. Scale bar: 500 μm. Figure 73. T. grandidieri, whole hemispermatophore (flagellum truncated). Scale bar: 4 mm. Figure 74. T. grandidieri, capsule, Antsakabe River, FMNH. Scale bar: 500 μm. Figure 75. T. flavopiceus, whole hemispermatophore. Scale bar: 2 mm. Figure 76. T. flavopiceus, capsule, Madagascar, GLPC, FKCP. Scale bar: 500 μm. Figure 77. T. annulatus, capsule, Tsimanampetsotsa National Park, GLPC, FKCP. Scale bar: 500 μm. Figure 78. T. ankarafantsika, capsule and flagellum. Scale bar: 400 μm. Figure 79. T. ankarafantsika, capsule, Forêt d’Ankavanana, FMNH. Scale bar: 200 μm. Figure 80. T. ankarafantsika, capsule, Réserve Forestière de l’Ankarafantsika, FMNH. Scale bar: 200 μm. Figure 81. T. olgae, capsule, Itampolo village, FMNH. Scale bar: 500 μm. Figure 82. T. limbatus, whole hemispermatophore (flagellum truncated). Scale bar: 2 mm. Figure 83. T. limbatus, capsule, Forêt d’Ianasana, FMNH. Scale bar: 500 μm. Figure 84. T. mahafaliensis, capsule views, Zombitse-Vohibasia National Park, GLPC, FKCP. Scale bar: 500 μm. Figure 85. T. intertidalis, capsule, Madagascar, GLPC, FKCP. Scale bar: 500 μm.

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 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 477–490. Teruelius flavopiceus, male. Figures 477–486. Pedipalp chela, dorsal (477), external (478) and ventrointernal (479) views; pedipalp patella, dorsal (480), external (481) and ventral (482) views; pedipalp femur and trochanter, internal (483) and dorsal (484) views; pedipalp chela, movable (485) and fixed (486) finger dentate margin. The trichobothrial pattern is indicated by white circles in Figures 477a–484a. Figures 487–490. Right legs tibia, basitarsus and telotarsus, retrolateral views. Leg IV (487), leg III (488), leg. II (489), leg I (490).

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

Figures 316–319. Grosphus madagascariensis, male, right legs I–IV tibia, basitarsus and telotarsus, retrolateral views. Scale bar: 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 409–417. Teruelius ankarafantsika. Male paratype (409, 416–417) and female holotype (410–415). Figures 409–411. Carapace and tergites I–III (409–410) and sternopectinal region (411). Figures 412–417. Left legs tibia, basitarsus and telotarsus, retrolateral views. Leg I (412), leg II (413), leg III (414, 416), leg IV (415, 417).

Figures 512–515. Teruelius intertidalis, female holotype. Left legs tibia, basitarsus and telotarsus, retrolateral views. Leg I (512), leg II (513), leg III (514), leg IV (515).

Figures 538–541. Teruelius mahafaliensis, male holotype of G. rossii. Left legs tibia, basitarsus and telotarsus, retrolateral views. Leg I (538), leg II (539), leg. III (540), leg IV (541).

Figures 576–579. Teruelius olgae, male paratype. Right legs tibia, basitarsus and telotarsus, retrolateral and ventral views. Leg I (576), leg II (577), leg III (578), leg IV (579).

Figures 610–619. Neogrosphus griveaudi, male (610–613, 617–619) and female (614–616). Figures 610, 614. Sternopectinal region. Figures 611, 615. Carapace and tergites I–III. Figure 612. Pedipalp chela, movable finger dentate margin. Figures 613, 616. Pedipalp chela in dorsal views. Figure 617. Metasoma V and telson lateral view. Figures 618–619. Right leg III (618) and leg IV (619), tibia, basitarsus and telotarsus, retrolateral views.