Teruelius limbatus (Pocock, 1889) Lowe & Kovařík, 2019

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 : 61

Teruelius limbatus (Pocock, 1889) comb. n.

8♂ 6♀ ( Figs. 5–8 , 18 , 23 , 49 , 82, 83 , 91 , 103, 126 , 139 , 157 , 159–160 , 176 , 192 , 229 , 235–238 ), Fianarantsoa Province, Forêt d’Ianasana, 7 km W Itremo, at source of Atsirakamhaity River, 20°36.1’S 46°34.3’E, 1630 m a. s. l., 6.II.1999, under rocks in Tapia forest , leg. S. M. Goodman ( FMNH 73449 ) ; 2♂ 7♀ ( Fig. 208 , 225 , 516–521 ), Central region, south of Antsirabe , 2006 ( FKCP) ; 1♂ 3♀, Central region, south ofAntsirabe, 2010 ( FKCP) .

Figures

Figures 5–8. Teruelius limbatus. Type species of Teruelius gen. n. Habitus. Male (5–6) and female (7–8), dorsal (5, 7) and ventral (6, 8) views. Scale bars: 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 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 90–93. Hemispermatophores and capsule regions of Teruelius gen. n. Cross stereoscopic convex views. Figure 90. T. ankarana, left mirrored, Forêt d’Ankavanana, FMNH. Figure 91. T. limbatus, Forêt d’Ianasana, FMNH. Figure 92. T. flavopiceus, Madagascar, GLPC, FKCP. Figure 93. T. ankarafantsika, Forêt d’Ankavanana, FMNH. Scale bars: 200 μm.

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 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 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 516–521. Teruelius limbatus. Figures 516–519. Habitus. Male (516–517) and female (518–519), dorsal (516, 518) and ventral (517, 519) views. Figure 520. Pedipalp chela external, male. Figure 521. Pedipalp chela, movable finger dentate margin, female. Scale bar: 10 mm (516–519).