Grosphus madagascariensis (Gervais, 1843)

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 : 60

Grosphus madagascariensis (Gervais, 1843)

3♂ ( Fig. 56 ) 6♀ 2juvs., ‘ Madagascar’ , leg. Saussure, det. M. Vachon ( MHNG); 11♂ 41♀ ( Figs. 1–4 , 9 , 22 , 41 , 54 , 94 , 123 , 135 , 145 , 159–160 , 165 , 183 , 199 , 214 , 227 , 231–234 ), Moramanga env., Toamasina, Anjiro, 18°52’S 47°59’E, 1995 ( FKCP, GLPC) ; 1♂ ( Figs. 52–53 , 86 ), Andasibe, Marie Guest House , 18.94727°S 048.41782°E, No. 1197 ( FKCP, GLPC) ; 1♂ ( G. mandena holotype, Figs. 215 , 348–349 ), 2♂ 1♀ 1juv. ( G. mandena paratypes, Figs. 55 , 200 , 350–351 ), Toliara Province, Mandena - Fort Dauphin, littoral forest 10 km north of Fort Dauphin, 6-12/I/1999, leg. J.-B. Ramanamanjato ( MHNG); 1♂ 2♀ ( Figs. 341–347 ), Moramanga env., 1997, ( FKCP) ; 1♂ 1♀, Andasibe-Mantadia, Anamalazaotra forest ( FKCP) ; 1♀, Fianarantsoa district, Ranomafana env., 21°13’S 47°25’E, 1995 ( FKCP) ; 5♂ 1♀ ( Figs. 10 , 95, 124 , 136 , 148 , 168 , 184 , 306–340 ), Toamasina Province, Ambalafary Forest , 14.5 km SW Andasibe, 19°02’38”S 48°20’55”E, 995 m a. s. l., 11. III.2012, dense, humid lowland and montane forest, leg. V. Soarimalala VS-2142 ( FMNH 3482757 ), det. as G. simoni ; 2♂ 3♀ 1juv. ( Fig. 57 ), Toamasina Province, Andasibe- Mantadia, near Andasibe, Camp Feon’ny Ala , 938 m a. s. l., 18°56.836’S 48°25.063’E, ( FKCP, GLPC) .

Figures

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 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 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 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 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.

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 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 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 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 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.