Teruelius, 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 : 12-20

Genus Teruelius gen. n.

( Figs. 5–8 , 13–20 , 23–39 , 44–51 , 71–85 , 90–93 , 99–105, 106–121, 137–144 , 150–160 , 170–180 , 186–195 , 201–210 , 216–226 , 229–230 , 235–238 , 387–579 , 584–605 , Tabs. 1–4 )

http://zoobank.org/urn:lsid:zoobank.org:act:54CB8128- BCFD-4B1F-A947-153C7CDD5B83

Grosphus Vachon, 1940: 254 , 256; Lourenço et al., 2007b: 375; Prendini & Esposito, 2010: 675–676; Loria & Prendini, 2014: 25; Loria & Prendini, 2018: 184.

TYPE SPECIES. Buthus limbatus Pocock, 1889 .

ETYMOLOGY. The generic epithet Teruelius (masculine) is a patronym honoring Rolando Teruel from Cuba in recognition of his many important contributions to the knowledge of scorpions.

DIAGNOSIS. A member of the ‘ Grosphus ’ group differentiated as follows: medium-sized to large-sized scorpions, adults ca. 35–120 mm in length; pedipalp finger granule rows 10–15 ( Figs. 402 , 431 , 452 , 485–486 , 508 , 521 , 529 , 560 ), movable finger typically with 4 external subdistal granules; femur trichobothrium d 2 straddling dorsointernal carina, or located on dorsal surface ( Figs. 13–20 ); chela manus with petite trichobothrium Eb 3 near Eb 2, closer than half the distance between Eb 1 and Eb 2 ( Figs. 23–24 ); manus trichobothrium V 2 roughly collinear with V 1 along chela axis or slightly displaced internally; higher pectinal tooth counts: ♂ 25– 41, ♀ 24–35 ( Figs. 28–31 ); basal pectinal tooth of females wide, with elongate, tapering distal extension, distinctly longer than other teeth ( Figs. 44–51 , 201–210 , 411 , 510 , 526 ); hemispermatophore capsule short, carinate, posterior lobe rounded, without lanceolate extension ( Figs. 71–85 ); sternites with narrow, slit-like spiracles ( Figs. 99–105); metasoma I with ventromedian carinae moderately to weakly crenulate or smooth to obsolete ( Figs. 126–132 ); telson with oval or bulbous vesicle, without subaculear tubercle in adults ( Figs. 186–195 ); legs with ventral surface of telotarsus densely setose or scopulate, with broad, brush-like strips of> 20 long filiform macrosetae ( Figs. 138–144 , 216–226 , 409–417 , 487–490 , 512–515 , 538–541 , 576–579 ); telotarsus with dorsal terminal process of normal size; cuticle with strong UV fluorescence ( Figs. 150–157 ).

SUBORDINATE TAXA.

Teruelius ankarafantsika ( Lourenço, 2003) comb. n. Teruelius ankarana ( Lourenço & Goodman, 2003) comb. n. Teruelius annulatus ( Fage, 1929) comb. n.

Teruelius bemaraha ( Lourenço, Wilmé & Waeber, 2018) comb. n.

Teruelius bicolor ( Lourenço, 2012) comb. n.

Teruelius bistriatus ( Kraepelin, 1900) comb. n.

Teruelius eliseanneae ( Lourenço & Wilmé, 2016) comb. n. Teruelius feti ( Lourenço, 1996) comb. n.

Teruelius flavopiceus ( Kraepelin, 1900) comb. n.

Teruelius ganzhorni ( Lourenço, Wilmé & Waeber, 2016) comb. n.

Teruelius grandidieri ( Kraepelin, 1900) comb. n.

Teruelius intertidalis ( Lourenço, 1999) comb. n.

Teruelius limbatus ( Pocock, 1889) comb. n.

Teruelius magalieae ( Lourenço, 2014) comb. n.

(= T. mahafaliensis ?)

Teruelius mahafaliensis ( Lourenço, Goodman & Ramilijaona, 2004) comb. n.

Teruelius olgae ( Lourenço, 2004) comb. n.

Teruelius sabineae ( Lourenço & Wilmé, 2016) comb. n. Teruelius waeberi ( Lourenço & Wilmé, 2016) comb. n.

See Tables 1–3 for diagnostic characters used to place the above species under Teruelius gen. n.

REMARKS. Recognition of Teruelius gen. n. as a separate genus, distinct from Grosphus , necessitates revision of some previous concepts about taxonomy and biogeography of Grosphus . Lourenço et al. (2017) associated G. ‘ simoni ’(= G. madagascariensis ) with G. rakotoariveloi and G. ‘ halleuxi ’, and subsequently Lourenço et al. (2018b) elaborated on a ‘ Grosphus simoni ’ group, treating it as a monophyletic unit of closely related “sister” species including the aforementioned three, plus G. ambre , G. bemaraha and G. mahafaliensis . A group diagnosis was not provided but, for G. bemaraha , mention was made of “a number of features such as spiniform granules on the dorsal carinae of metasomal segments II-IV and on internal carinae of pedipalp femur and patella”. We place G. bemaraha under Teruelius gen n., on the basis of narrow spiracles, dense tarsal setation and high pectinal tooth count ( Table 3 ), and consider spiniform granules shared with G. simoni to be a homoplasy. G. bemaraha was claimed to be closer to G. rakotoariveloi , but these species belong to different genera. Lourenço et al. (2018b: 74, fig.1) included G. mahafaliensis in the ‘ simoni ’group, perhaps due to similarities to G. bemaraha , noting in particular a high number of pectine teeth (but it lacks spiniform granules on metasomal carinae). In contrast, we find that T. mahafaliensis comb. n. is very far removed from G. ‘ simoni ’ (= G. madagascariensis ), differing in all nine genus-level diagnostic characters, and in the fundamental architecture of the hemispermatophore capsule (cf. Fig. 84 vs. Figs. 52–57 ).

Lourenço et al. (2018b) discussed biogeographic hypotheses attempting to explain the distribution of the incongruous, polyphyletic ‘ Grosphus simoni ’ group. The very wide distribution of the group meant that it was “adapted to humid, dry and subarid environments”, and the two most disjunct species, G. bemaraha and G. mahafaliensis from western and southern localities, were speculated to “belong to relict populations, which may have survived in humid refugia encountered in the sedimentary basins during the dry episodes of the paleoclimate oscillations”. However, these two species are not closely related to the other four group members, but belong to Teruelius gen. n., whose ancestors may have already been adapted to dry environments. Although Pleistocene climatic fluctuations could be relevant for recent speciation events in Grosphus and Teruelius gen. n., the many correlated characters separating these two genera suggest a far earlier split, as in other taxa. For example, dated molecular phylogenies of Madagascar archaeid spiders (Wood et al., 2015), Brookesia and other chameleons ( Tolley et al., 2013; Townsend et al., 2009) and Zonosaurus plated lizards ( Blair et al., 2015) have revealed that the majority of divergences in these other endemic taxa are quite deep, occurring long before the advent of Pleistocene climate cycles.

Species of Teruelius gen. n. can be loosely subdivided by size and coloration: large species, T. flavopiceus , T. ankarana , T. grandidieri and T. bicolor ; species with patterns of dark stripes on tergites (‘ bistriatus ’ group of Lourenço & Wilmé, 2016): T. ankarafantsika , T. bistriatus , T. eliseanneae , T. feti , T. limbatus , T. sabineae and T. waeberi ; species with almost uniform yellow, orange or brown tergites, and maybe darker metasoma IV or V: T. annulatus , T. bemaraha , T. ganzhorni , T. intertidalis , T. magalieae , T. mahafaliensis and T. olgae . These groupings have been used to construct species keys, in conjunction with some other characters including shapes of female basal pectine teeth ( Fage, 1929; Lourenço, 2003c, 2004a, 2014; Lourenço et al., 2007b; Vachon, 1969). Monophyly of these groupings remains to be tested.

NEW SYNONYMIES.

Grosphus makay Lourenço & Wilmé, 2015 = Teruelius feti ( Lourenço, 1996) comb. n., syn. n.

Grosphus feti was described by Lourenço (1996b) from a juvenile male holotype, ostensibly collected from “Prov. Tulear, Tanjon’ I Vohimena [= Cap Sainte Marie] Réserve spéciale, X.1995 ” and deposited in FMNH. We loaned and studied the holotype and a second juvenile male labeled as “ ♂ paratype ” (which we concur is conspecific with the holotype). Until now, this species was only known from these two types. Associated with the type, we found locality labels ( Fig. 459 ) that differ from the published type locality: “ MADAGASCAR: Province de Toliara, Fôret de Vohimena, 35 km SE Sakaraha, 17-24.i.1996, MyrCE-7541, 22°41.0’S 44°49.8’E, 780 m, S. M. Goodman 0000 011 031 FMNH- INS Grosphus feti Lourenço HOLOTYPE: det. 1996 ”. This locality is ca. 325 km roughly north of the published type locality at Cap Sainte Marie and the collection date of January 1996 is several months later. Either a labeling error occurred after description, or the published type locality is incorrect. Cap Sainte Marie is a biological study area and frequent source of scorpion materials (e.g., Lourenço & Wilmé, 2016) so data labels of specimens could have been confused.

The FMNH label site is ca. 145 km south and slightly west of the type locality of Grosphus makay ( Lourenço & Wilmé, 2015b) : “Region Atsimo-Andrefana, ex Province of Toliara, Makay Mts., General Collection, dry-Forest on sandy soil, 12/III/2010 (B. L. Fisher et al.). BLF25549. Female holotype (CAS)”. These two localities have sandstone substrates, similar elevation and are located in the same general bioclimatic region. Comparison of the published habitus of the holotype adult female of T. makay and the holotype juvenile male of T. feti revealed very similar morphology and morphometrics even though sex and age differ. Most notably, their color patterns are identical in all details including: pattern of fuscosity on interocular triangle of carapace with pale cut-out behind lateral eyes; thin median line, precise fuscous banding patterns and transverse lateral striping on all tergites; darkly marked ventrolateral and ventromedian carinae on metasoma I–IV; fuscous patterns on metasoma V and telson; short fuscous strip on interno-proximal margin of pedipalp patella; leg femora with distal short, pale cut-outs on distal dark areas of prolateral surfaces and pale narrow lines on dorsal margins; and leg patellae with fuscous ventral margins on prolateral surfaces (compare Fig. 459 to fig. 13 of Lourenço & Wilmé, 2015b). These very particular details of pigmentation pattern are not found in T. limbatus which was regarded as a closely related species ( Figs. 5–8 , 516–521 ). We also examined two adult females, near topotypes of G. makay from the Makay Mountains , that exhibited the same coloration patterns and morphometrics as T. feti ( Fig. 459a ). We allowed for the fact that juveniles of pigmented scorpions usually display darker, more intense markings, and that in adults these color patterns are somewhat faded. Details of coloration pattern have been given high priority as characters for species-level taxonomy of Grosphus ( Lourenço et al., 2009b; Lourenço, 2014). We therefore consider Grosphus makay to be a junior synonym of T. feti , and the correct type locality of the latter to be that indicated on FMNH data labels. T. feti was never again collected from Cap Sainte Marie in over two decades of fieldwork since its description, although other scorpion species (e.g., T. sabineae ) were discovered there. The adult male remains unknown. Our opinion could be verified by collection and analysis of topotypic adult and juvenile specimens from the FMNH locality. The description of T. feti ( Lourenço, 1996b: 14) noted the juvenile status of the holotype which is ca. 30 mm, but Lourenço (2014: 636) diagnosed the species as “of small size with a total length of 30 to 40 mm ”. Small size cannot be a species diagnostic character if it is a property of the juvenile, and adult males are probably medium-sized, comparable to an adult female of G. makay , ca. 56 mm in body length. Our synonymy also implies that this is not a microendemic species of the upper Central Menabe, as suggested of Lourenço & Wilmé (2015b). The T. feti emended type locality lies a significant distance south of the Makay mountains , in the Mangoky watershed ( Wilmé et al., 2006).

Grosphus rossii Lourenço, 2013 = Teruelius mahafaliensis ( Lourenço, Goodman & Ramilijaona, 2004) comb. n., syn. n.

Grosphus rossii was described by Lourenço (2013b) from a single adult male holotype collected from “ Central region, NE Manandona, S of Antsirabe, in secondary growth forest, under log, 8 August 2004, W. R Lourenço” and deposited in ZMUH. We loaned and studied the holotype and found that it was virtually identical in coloration, external morphological characters and morphometrics to Teruelius mahafaliensis ( Lourenço, Goodman & Ramilijaona, 2004) comb. n. Our comparative materials of the latter included a male collected near the type locality of that species ( Figs. 522–525 ), and determined material loaned from FMNH including two adult males. In his description, Lourenço (2013b: 59) compared and contrasted G. rossii to T. limbatus , but not to T. mahafaliensis . A potential diagnostic difference is the pectinal tooth count (= 28) in G. rossii being lower than the range (35–40) reported for male T. mahafaliensis by Lourenço et al., (2007b: 373, tab. III). However, we examined a male T. mahafaliensis collected near the species type locality with PTC of 29–33 ( Figs. 524– 525 ). The type locality of G. rossii on the central plateau at ca. 1400 m a. s. l., is in a cooler, more humid zone, quite far from the other records of T. mahafaliensis concentrated on the Mahafaly Plateau, a region of subarid thorn scrub along the southwest coast (ca. 120 m a.s.l.). It was suggested that G. rossii was evidence of microendemism. We take a more conservative position and interpret the very close morphologies of G. rossii and T. mahafaliensis as indicative of a eurytopic species with wider distribution. Broad elevation ranges of> 1400 m are known for some widely distributed scorpions that inhabit varied bioclimatic zones (e.g., Anuroctonus pococki Soleglad & Fet, 2004 , 300– 1850 m a.s.l., Soleglad & Fet, 2004; Bothriurus burmeisteri Kraepelin, 1894 and Brachistosternus weijenberghi (Thorell, 1876), 1000 –3000 m a.s.l., Campón et al., 2014; Compsobuthus maindroni (Kraepelin, 1901) , Hottentotta jayakari (Pocock, 1895) , Nebo omanensis Francke, 1980 and Orthochirus glabrifrons (Kraepelin, 1903) , 0–1850 m a.s.l., Lowe, 2010c). The status of G. rossii should be reviewed when topotypic females are collected and their basal pectinal tooth compared to that of T. mahafaliensis , as this is a more reliable diagnostic character in Teruelius gen. n.

Another potential synonym of T. mahafaliensis is T. magalieae ( Lourenço, 2014) . According to its description, the morphometrics, coloration, and meristics of T. magalieae are very close to those of T. mahafaliensis . The type locality of Cap Saint Marie (holotype male as only known specimen) lies on the southwestern coast in the same bioclimatic region as the latter species. Lourenço (2014: 633) did not compare T. magalieae to T. mahafaliensis , but claimed that the most closely related species was G. rossii , which we here synonymize under T. mahafaliensis . The diagnostic differences between T. magalieae and G. rossii are not compelling: (i) pectines with 36 vs. 28 teeth (a range of variation allowed here for T. mahafaliensis ); (ii) pedipalp fingers with 12–13 vs. 12–12 granule rows (overlapping counts); and (iii) overall paler coloration (differences in color shade are not uncommon for different populations of a species inhabiting areas with different substrates). We provisionally list this species, until it can be critically evaluated by study of more material and analysis of variation. The female of T. magalieae is unknown, and the species might be better diagnosed if the female basal pectinal tooth were determined to be unique.

Figures

Figure 459. Teruelius feti. Holotype juvenile male. Habitus, dorsal view, with specimen labels (FMNH). Scale bar: 5 mm (holotype). Figure 459a. Adult female from Makay Mts., carapace and tergites I–II.

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 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 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 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 387–390. Teruelius ankarafantsika. Habitus. Male paratype (387–388) and female holotype (389–390), dorsal (387, 389) and ventral (388, 390) views. Scale bar: 10 mm.

Figures 391–408. Teruelius ankarafantsika. Male paratype (391–393, 404–405) and female holotype (394–403, 406–408). Figures 391–402. Pedipalp.Chela in dorsal (391, 394), external (392, 395) and ventral (396) views. Patella in dorsal (397), external (398) and ventral (399) views. Femur and trochanter in internal (393, 400) and dorsal (401) views. Pedipalp chela, movable finger dentate margin (402). Figures 403–408. Telson in lateral (403, 404) views. Metasoma and telson in lateral (405, 406), dorsal (407) and ventral (408) views.Scale bar: 10 mm (406–408).

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 418–419. Teruelius ankarana. Habitus. Male in dorsal (418) and ventral (419) views. Scale bar: 10 mm.

Figures 420–421. Teruelius ankarana. Habitus. Female in dorsal (420) and ventral (421) views. Scale bar: 10 mm.

Figures 422–425. Teruelius annulatus. Habitus. Male (422–423) and female (424–425), dorsal (422, 424) and ventral (423, 425) views. Scale bar: 10 mm.

Figures 426–435. Figures 426–432. Teruelius annulatus. Male (426, 428, 430–431) and female (427, 429). Sternopectinal region (426–427). Carapace and tergites I–III (428). Pedipalp chela dorsal (429–430). Pedipalp chela, movable finger dentate margin (431). Metasoma V and telson lateroventral view (432). Figures 433–435. Teruelius bistriatus. Female (433–434) and male (435) syntypes. Sternopectinal region (433, 435). Carapace and tergites I–III (434).

Figures 436–438. Teruelius bistriatus. Habitus. Female syntype in dorsal (436) and ventral (437) views, and original labels (438). Scale bar: 10 mm.

Figures 439–446. Teruelius bistriatus. Metasoma and telson. Male (439, 441–443) and female syntypes (440, 444–446), lateral (439–441, 444), ventral (442, 445) and dorsal (443, 446) views. Scale bars: 10 mm (441–443, 444–446).

Figures 447–458. Teruelius bistriatus. Pedipalp. Female (447–452) and male (453–458) syntypes. Pedipalp chela, dorsal (447, 453), external (448, 454) and ventrointernal (449, 455) views; pedipalp patella, dorsal (450, 456) and ventral (457) views; pedipalp femur and trochanter in dorsal (451, 458) views; pedipalp chela, movable finger dentate margin (452). Pedipalp of male is mirrored.

Figures 460–472. Teruelius feti. Holotype male. Metasoma and pedipalp. Figures 460–462. Metasoma in dorsal (460), lateral (461) and ventral (462) views. Figures 463–472. Pedipalp. Chela in dorsal (463), external (464) and ventral (465) views. Patella in dorsal (466), external (467) and ventral (468) views. Femur and trochanter in internal (469), dorsal (470), ventral (471) and external (472) views. Right femur shows developmental anomaly (Figs. 469, 472). Scale bars: 2 mm (460–462, 463–472).

Figures 473–476. Teruelius flavopiceus. Habitus. Male (473–474) and female (475–476), dorsal (473, 475) and ventral (474, 476) views. Scale bar: 10 mm.

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 491–494. Teruelius grandidieri. Habitus. Male (491–492) and female (493–494), dorsal (491, 493) and ventral (492, 494) views. Scale bar: 10 mm.

Figures 495–499. Teruelius intertidalis, female holotype. Figures 495–496. Habitus in dorsal (495) and ventral (496) views, and original labels. Figures 497–499. Metasoma in lateral (497), ventral (498) and dorsal (499) views. Scale bars: 10 mm.

Figures 500–511. Teruelius intertidalis, female holotype. Figures 500–508. Pedipalp chela, dorsal (500), external (501) and ventrointernal (502) views; pedipalp patella, dorsal (503), external (504) and ventral (505) views; pedipalp femur and trochanter, internal (506) and dorsal (507) views; pedipalp chela, movable finger dentate margin (508). Figure 509. Carapace and tergites I–III. Figure 510. Sternopectinal region. Figure 511. Telson lateral view.

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 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).

Figures 522–525. Teruelius mahafaliensis. Habitus. Male holotype of G. rossii (522–523) and male from near the type locality of T. mahafaliensis (524–525), dorsal (522, 524) and ventral (523, 525) views. Scale bar: 10 mm.

Figures 526–537. Teruelius mahafaliensis, female (526, 529–530, 535), male (527–528, 531–534) and male holotype of G. rossii (536–537). Sternopectinal region (526–527). Carapace and tergites I–II (528). Pedipalp chela, movable finger dentate margin (529). Pedipalp chela dorsal (530–531) and external (532) views. Pedipalp patella dorsal (533) view. Pedipalp femur and trochanter dorsal (534) view. Metasoma V and telson lateral (535) view. Metasoma and telson dorsal (536) and ventral (537) views. Scale bar: 10 mm (536–537).

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 542–546. Teruelius olgae, male paratype.Figures 542–543. Habitus, dorsal (542) and ventral (543) views, and original labels. Figures 544–546. Metasoma and telson in lateral (544), ventral (545) and dorsal (546) views. Scale bars: 10 mm (542–543, 544–546).

Figures 547–551. Teruelius olgae, female. Figures 547–548. Habitus, dorsal (547) and ventral (548) views. Scale bar: 10 mm. Figures 549–551. Metasoma and telson. lateral (549), ventral (550) and dorsal (551) views. Scale bar: 5 mm.

Figures 552–563. Teruelius olgae, male paratype. Figures 552–560. Pedipalp chela, dorsal (552), external (553) and ventrointernal (554) views; pedipalp patella, dorsal (555), external (556) and ventral (557) views; pedipalp femur and trochanter, internal (558) and dorsal (559) views; pedipalp chela, movable finger dentate margin (560). Figure 561. Carapace and tergites I–II. Figure 562. Sternopectinal region. Figure 563. Telson lateral view.

Figures 564–575. Teruelius olgae. Female pedipalp. Chela in dorsal (564), external (565), ventral (566) and internal (567) views. Patella in dorsal (568), external (569), ventral (570) and internal (571) views. Femur and trochanter in dorsal (572), internal (573), external (574) and ventral (575) views. Scale bar: 2 mm.

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 584–587. Teruelius ankarana. Juveniles before second (584 top), after second (584 below and 585), third (586) and fourth (587) ecdysis.

Figures 588–589. Teruelius ankarana. Male (588) and female (589), in vivo habitus.

Figures 590–592. Teruelius flavopiceus. Juveniles after second (590), third (591) and fourth (592) ecdysis.

Figures 593–594. Teruelius flavopiceus. Male (593) and female (594), in vivo habitus.

Figures 595–596. Teruelius flavopiceus. Female with newborns (595) and with juveniles after first ecdysis (596).

Figures 597–599. Teruelius flavopiceus. Juvenile with duplicated metasoma (597, 599) and its unsuccesful second ecdysis (598).

Figures 600–601. Teruelius grandidieri. Male (600) and female (601) in vivo habitus.

Figures 602–603. Teruelius grandidieri. Female with newborns (602) and with juveniles after first ecdysis (603).

Figures 604–605. Teruelius limbatus. Male (in the middle) and females (604) and female with newborns (605).

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 127–132. Ventral aspect of metasoma and telson of adult males (♂) or females (♀) of representative Teruelius gen. n under UV fluorescence to reveal carination and granulation. T. ankarafantsika (127), T. ankarana (128), T. flavopiceus (129), T. grandidieri (130), T. mahafaliensis (131), and T. olgae (132). Scale bars: 4 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.