Teruelius haeckeli, Lowe & KOvAřík, 2022

Lowe, Graeme & KOvAřík, FRANtIšEk, 2022, Reanalysis of Teruelius and Grosphus (Scorpiones Buthidae with descriptions of two new species, Euscorpius 356, pp. 1-105 : 82-86

Teruelius haeckeli sp. n.

( Figs. 13–16 , 19–21 , 23–28 , 30–35 , 49–50, 52 , 137 , 165, 167–168 , 175–178 , 180–183 , 196–209 , 328–392 , Tab. 15) http://zoobank.org/urn:lsid:zoobank.org:act:7CBD4E06-

8D6C-47CB-BBD4-F068D262DF50

Teruelius annulatus Lowe & Kovařík, 2019: 25 , 61, 64–65, 91–92; figs. 77, 203, 218, 422–432 (misidentification).

TYPE LOCALITY AND TYPE REPOSITORY. Madagascar: Toliara Province, Tsimanampetsotsa National Park, Andranovao camp, FKCP .

TYPE MATERIAL. Madagascar: Toliara Province, Tsimanampetsotsa National Park, Andranovao camp, 15 m a. s. l., 24°01.505'S 43°44.306'E, 1♂ (holotype), 3♂ 1♀ 1juv ♂ (paratypes), 2014, FKCP, GLPC (1 hemispermatophore) .

ETYMOLOGY. The species is named in honor of Czech entomologist, physician Martin Häckel.

DIAGNOSIS. Small to medium-sized member of the genus, total length of adults 30–53 mm; body and appendages uniformly yellow, metasoma IV–V (or only V) and telson dark, blackbrown; carapace of males with granulate superciliary carinae; pedipalp patella with strong, costate-granulate dorsointernal carina in both sexes; pedipalp chela of males with internal surface of manus granulate, fingers without undulations on proximal dentate margins; leg III tibial spur L/ tibia distal D> 0.73; female bpt falcate, without long narrow extension, shorter than basal comb width; PTC ♂ 37–40, ♀ 27–28, regular pectine tooth L/W ♂ 4.71, ♀ 3.89; hemispermatophore posterior lobe short, apically rounded, with two lateral carinae; metasoma III ventral intercarinal surface smooth, dorsosubmedian carinae of males bearing large dentate posterior terminal granule; metasoma V with dorsosubmedian carinae smooth, obsolete; telson with aculeus length equal to vesicle length, vesicle weakly granulate on ventral surface; morphometrics, L/W ratios (n = 4 ♂, 1 ♀): metasoma I ♂ 1.20– 1.27, ♀ 1.195, metasoma II ♂ 1.49–1.54, ♀ 1.47, metasoma III ♂ 1.57–1.62, ♀ 1.56, metasoma IV ♂ 1.81–2.01, ♀ 1.89, metasoma V ♂ 2.03–2.19, ♀ 2.41, pedipalp chela ♂ 3.64– 3.77, ♀ 5.16, pedipalp femur ♂ 2.78–3.44, ♀ 2.34, pedipalp patella ♂ 2.67–3.65, ♀ 2.64.

DESCRIPTION. Coloration ( Figs. 328–333 , 338–357 , 372– 387 ). Base color yellow to orange-yellow; metasoma IV either dark brown (♂) or yellow and partially black ventrally (♀); metasoma V and telson either dark brown (♂) or black (♀). Leg femora and patellae with dark ventral margins (♀).

Carapace ( Figs. 328, 330 , 334 , 336 ). Subrectangular, W/L 1.06–1.08; medial surface level along its entire length; anterior margin slightly concave, male with epistomal process; preocular L/ carapace L 0.40–0.43; surface densely, finely granulate in most areas; granulation more coarse on preocular triangle; granulation much weaker in female; superciliary carinae granulate in male, weakly granulate to smooth in female; anterior margin with several macrosetae; lateral eye groups composed of 3 large and 2 small ocelli (type 5; Loria & Prendini, 2014); median eyes large, eye diameter/ carapace L 0.11 (♂), 0.15 (♀).

Chelicerae ( Figs. 332 , 334–337 ). Dorsal surface of manus of male weakly granulate near anterior margin, smooth elsewhere, of female smooth throughout; several macrosetae and pale, fluorescent microsetae near anterior margin; dorsointernal carina strong, weakly granulate in male, smooth in female; fingers with typical buthid dentition (Vachon, 1963), movable finger dorsal margin with two large subdistal denticles and two small basal denticles, ventral margin with subdistal and basal denticles, fixed finger with large subdistal denticle and proximal bicusp, two denticles on ventral surface; dorsal surface of movable finger smooth, with dorsal row of 4–5 pale, fluorescent microsetae.

Coxosternal area ( Figs. 335 , 337 ). All coxae smooth with sparse macrosetae and fluorescent microsetae; sternum smooth, subtriangular, with long medial depression, bearing 2 macrosetae; genital opercula smooth, divided in female.

Pectines ( Figs. 28 , 32 , 34–35 , 323, 325, 327 ). Basal piece smooth, with deep anteromedian invagination, combs with 3 marginal lamellae, 10–12 middle lamellae; combs long, extending to distal limit (♂) or distal 2/3 (♀) of trochanter IV; marginal and middle lamellae with numerous small macrosetae, fewer fluorescent microsetae; fulcra with 4–6 short setae; female bpt falcate.

Hemispermatophore. ( Figs. 388–392 ). Flagelliform; trunk narrow, elongate; capsule short, with large, robust, hook-like basal lobe; posterior lobe rounded, with two carinate folds on convex surface; flagellum with short pars recta and pars reflecta (the latter probably incomplete in the examined specimen).

Mesosoma ( Figs. 332–337 ). Tergites: pretergites smooth, with microsulcate posterior margins; all tergites densely, finely granulated or shagreened, more weakly so in female; tergite I without distinct carinae, tergites II–VI with single weak, granulate median carina, VII with medial hump and 2 pairs of granulate carinae; all tergites lacking macrosetae. Sternites: all sternites smooth, glossy, acarinate; posterior margins of all sternites smooth; spiracles long, narrow, slit-like; sternite III–VI macrosetae: one submedian pair, one lateral pair, two posterior marginal pairs; sternite VII macrosetae: two submedian pairs, three lateral pairs; posteromedian margin of sternite V convex in female, forming a sensory patch with dense narrow, transverse band of microsetae along margin, and a wider, more sparse transverse band of microsetae slightly anterior to margin; posteromedian marginal setation on sternite V denser than on sternites IV and VI.

Metasoma ( Figs. 137 , 372–374, 376–378 ). Elongate, segments I–IV uniform in width, segment V narrower posteriorly. Carination: segment I with 10 complete carinae, II with 8 complete carinae (lateral median carinae anteriorly indistinct), segments III–IV with 8 carinae, V with 5 carinae; dorsosubmedian and dorsolateral carinae on segments IIV and lateral median carinae on segments I–III granulate, crenulate or dentate-granulate in both sexes; ventrosubmedian and ventrolateral carinae on segments II–IV granulate or crenulate (♀) or smooth (♂); ventrosubmedian and ventrolateral carinae on segment I weakly crenulate to smooth in both sexes; segment V with dorsolateral carina granulate, ventrolateral and ventromedian carinae granulate with larger dentate granules in posterior half; lateral anal margin with 4 large granules, ventral anal margin with up to 20 granules. Intercarinal surfaces: dorsolateral, lateral, ventrolateral and ventral surfaces of segments I–VI smooth or almost smooth with sparse fine granules; dorsomedian surfaces of all segments smooth; segment V smooth laterally, with sparse coarse and fine granules ventrally. Setation: carinae bear regular series of long macrosetae, 3–6 on segments I–IV, up to 10 on segment V; posterior ventral margins of segments I–IV with several long macrosetae.

Telson ( Figs. 372–379 ). Vesicle dorsal surface smooth; lateral surfaces and ventral surface weakly granulate, with sparse, long macrosetae; vesicle hemi-elliptic or bulbous in lateral profile, without subaculear tubercle; aculeus shorter than vesicle.

Pedipalps ( Figs. 338–371 ). Femur: dorsointernal, dorsoexternal and ventrointernal carinae strong, coarsely granulate; other carinae indistinct; internal surface with 7–10 large granules; intercarinal surfaces smooth; sparse short macrosetae and fluorescent microsetae present. Patella: dorsointernal carina strong, costate-granulate in both sexes, weaker in female; dorsomedian carinae obsolete; dorsoexternal carina of male weakly granulate in distal half, obsolete in proximal half of segment, of female obsolete; ventroexternal and ventromedian carinae of male nearly obsolete, indicated by small granules, of female obsolete; ventrointernal carina granulate in both sexes, weaker in female; internal carina indicated by series of 4–5 enlarged dentate granules; sparse short and long macrosetae, and fluorescent microsetae present. Chela: all carinae obsolete, surfaces smooth except for finely granulate internal surface of manus in males; short macrosetae and fluorescent microsetae sparse on manus, dense on fingers; 11–12 median denticle ventral surfaces of telotarsi with dense brush of macrosetae subrows on fixed finger, 13 on movable finger including short irregularly arranged, lateral apices with conspicuous fringes subdistal row; subrows flanked by one mid-row internal and of long macrosetae; tarsal ungues stout.

two proximal external accessory denticles (except for unfused Measurements. See Table 15.

proximal subrow). Trichobothriotaxy: orthobothriotaxic, type

Aα (Vachon, 1974), femur d 2 dorsal, e 1 distal to d 5; chela fixed AFFINITIES. Teruelius olgae ( Lourenço, 2004) is similar to T. finger db proximal to est. haeckeli sp. n. in adult size, color pattern, and telson shape, Legs ( Figs. 328–331 , 380–387 ). Femora with crenulate but differs in having more slender metasomal segments, a lower ventral carinae; surfaces of all segments smooth; patellae range of male PTC (29–33), and a clavate female bpt with long with series of long macrosetae; tibia and tarsal segments curved extension. T. mahafaliensis is similar to T. haeckeli sp. bearing numerous short macrosetae; tibial spurs present on n. in having a higher range of male PTC (34–40), but differs legs III–IV; retrolateral tarsal spurs simple, prolateral tarsal in its larger adult size (55–60 mm), reddish coloration, lack of spurs basally bifurcate; ventral surfaces of basitarsi with black color on metasoma IV–V and telson vesicle, and a clavate numerous macrosetae arranged roughly in two axial series; female bpt with moderately long, curved extension.

REMARKS. The convex profile of the posteromedian margin extends slightly anterior to the margin in females. In both of sternite V in females ( Fig. 337 ) differs from the almost sexes, the posteromedian marginal setation is denser on straight posteromedian margin of sternite V in males (Fig. sternite V, than on sternites IV and VI. These anatomical 335). There is also sexual dimorphism of the setation, differences suggest functional specialization. Sternite V is which is strictly confined to the margin in males, but also modified in various other buthids, forming a smooth, pale or fluorescent posteromedian patch with possible glandular or sensory functions. We found differences in the relative density of posteromedian marginal setation of sternites IV–V between Grosphus and Teruelius in several examined species. In females, marginal setation on sternite V in Grosphus was similar in density to that of sternite IV (5 spp., ♀, Figs. 393–397), and marginal setation on sternite V in Teruelius was denser than that on sternite IV (5 spp., ♀, Figs. 398–402). In males of these species, denser marginal setation on sternite V was absent in G. hirtus , G. madagascariensis , G. simoni , G. voahangyae and T. mahafaliensis , and was present in T. ankarafantsika , T. ankarana , and T. limbatus . Other Teruelius with denser marginal setation on sternite V vs. IV include T. bistriatus (♀) (Lowe & Kovařík, 2019: 93, fig. 437). T. flavopiceus (♂, ♀; modestly so), T. intertidalis (♀), T. ganzhorni (♀) (Ref. MNHN-RS-RS9080), T. grandidieri (♂, not ♀). These data suggest that dense marginal setation on sternite V is a potential diagnostic character or synapomorphy separating at least some Teruelius from Grosphus . However, we did not include this in our analyses, due to limited taxon sampling and character variability. There was variability in posterior marginal profiles (i.e., convex, linear or concave), and in the density and arrangement of setae. The denser setation could be associated with an increase in numbers of either macrosetae, or fluorescent microsetae. Further investigation of putative glandular or sensory specializations of sternite V, and their sexual dimorphism, is needed to establish character homologies for phylogenetic analysis.

Figures

Figure 34: Phenetic analysis of shapes of female basal pectinal teeth (bpt). Ultrametric tree obtained from hierarchical cluster analysis by UPGMA of the Euclidean distance matrix of z-scores of 32 Fourier coefficients. Font colors: Grosphus, blue; Teruelius flavopiceus, orange; T. ankarana, magenta; T. grandidieri, red; other Teruelius spp., dark yellow. Pectine images: Grosphus angulatus sp. n. (upper), Teruelius grandidieri (lower).

Figure 35: Phylogram of shapes of female basal pectinal teeth (bpt). Tree obtained from neighbor-joining cluster analysis of the Euclidean distance matrix of z-scores of 32 Fourier coefficients. Outgroup taxon: Pseudolychas ochraceus. Font colors: as in Fig. 34, with Pseudolychas black. Pectine images: Teruelius grandidieri (upper), Grosphus angulatus sp. n. (lower).

Figures 13–16. Morphometric analyses of carapace and hemispermatophores. Figure 13. Horizontal bar plot of mean carapace length (mm) (character 0) of Grosphus (n = 46, 14 spp.), Teruelius (n = 70, 21 spp.), Pseudolychas (n = 8, 3 spp.), and other outgroup taxa (n = 36, 9 spp.). Data from both sexes pooled. Error bars are standard errors. Discretization thresholds at step changes in ranked length. Figure 14. Horizontal bar plot of mean concavity angle (°) (character 4) of Grosphus (n = 49, 14 spp.), Teruelius (n = 72, 19 spp.), Pseudolychas (n = 8, 3 spp.) and other outgroup taxa (n = 22, 9 spp.). Data from both sexes were pooled. Error bars indicate standard errors. Discretization threshold at step transition in rank slope. Inset: angle defined by tangent line at midpoint between anterior-most lateral eye and carapace center. Figure 15. Horizontal bar plot of mean ratio of carapace preocular L/ carapace L (character 5) of Grosphus (n = 39, 14 spp.), Teruelius (n = 71, 21 spp.), Pseudolychas (n = 8, 3 spp.) and other outgroup taxa (n = 23, 9 spp.). Data from both sexes pooled.Error bars are standard errors.Discretization threshold at a minor step transition in rank slope. Figure 16. Bivariate logarithmic scatter plot of hemispermatophore posterior lobe width/ length ratio vs. hemispermatophore capsule length/ posterior lobe length for Grosphus (n = 13, 6 spp.), Teruelius (n = 10, 9 spp.) and outgroup Pseudolychas (n = 1, 1 sp.). Color codes of symbols or bars as indicated in Fig. 16 legend: Grosphus ‘hirtus’ group (G. angulatus sp. n., G. hirtus, G. polskyi, G. voahangyae), blue; other Grosphus spp., cyan; Teruelius flavopiceus, orange; T. ankarana, magenta; T. grandidieri, red; other Teruelius spp., yellow; Pseudolychas spp., black; other outgroups, gray.

Figures 19–22. Morphometric analyses of regular pectine teeth. Figures 19–20. Horizontal logarithmic bar plots comparing length/ width (L/W) ratios of regular pectine teeth in males (19) and females (20) of Grosphus, Teruelius and outgroup taxa. Bars are rank ordered means, error bars are standard errors; ♂ n = 52, 30 spp.; ♀ n = 66, 31 spp. Figure 21. Bivariate logarithmic scatter plot comparing pectinal tooth L/W ratios of males (ordinate) vs. females (abscissa) (21 spp.). Gray line is diagonal. Plotted values and error bars as in Figs. 19–20. Fig. 22. Regular pectine teeth (♀) of Grosphus voahangyae (left) and Teruelius ankarafantsika (right), showing measurements of length (L) and width (W). W is equal to inter-fulcral spacing. UV fluorescence. Measurements were taken at> 3 teeth away from most proximal or most distal teeth. Color codes of symbols and bars as in Fig. 16–17 legends.

Figures 23–28: Ratiometric analysis of shapes of female basal pectinal teeth (bpt). Figures 23–26. Bivariate logarithmic scatter plots of six ratiometric shape variables: roundness vs. solidity (23), circularity vs. roundness (24), maximum/ minimum caliper diameter vs. solidity (25), basal tooth width/ regular tooth width vs. perimeter attachment ratio (26). Figures 27–28. Bivariate scatter plots of first two principal components (PC2 vs. PC1) obtained from PCA of standardized logarithms of all six ratiometric variables, accounting for 85.31% and 8.43% of variance, respectively. Individual cases plotted in Fig. 27, means and standard errors for each species in Fig. 28. Profile silhouette examples are shown for analyzed species in Fig. 28. Data from 106 bpt from Grosphus (n = 31, 8 spp.), Teruelius (n = 70, 18 spp.) and Pseudolychas (n = 5, 2 spp.). Symbol colors indicated in legend of Fig. 23.

Figures 29–33: Elliptic Fourier analysis (EFA) of shapes of female basal pectinal teeth (bpt). Figure 29. Logarithmic plots of harmonic power (sums of squares of Fourier coefficients) vs. harmonic order for fits to bpt outlines of Grosphus (blue symbols) and Teruelius (yellow symbols). Error bars indicate ranges (minimum to maximum). Upper inset: examples of EFA fits to bpt from Grosphus (G. voahangyae) (left) and Teruelius (T. olgae) (right) by Fourier series with cumulative terms up to and including second (blue), fourth (green) and eighth (red) order harmonics. Contours of bpt oriented with perimeter attachment horizontal, start point at proximal vertex, area normalized.Figures 30–31. Bivariate scatter plots of bpt scores for first three principal components,PC2 vs. PC1 (Fig.30) and PC3 vs. PC2 (Fig.31), obtained from PCA of 32standardized Fourier coefficients from up to eighth order harmonic terms, accounting for 31.02%, 19.14% and 8.20% of variance, respectively (total variance 58.36%). Lower inset in Fig. 31: scree plot of eigenvalue vs. PC number.Figure 32. Bivariate scatter plot of means and standard errors of bpt scores of first two principal components, PC2 vs. PC1, for each species in Fig. 30.Figure 33. Trivariate scatter plot of bpt scores of first three principal components (PC1, PC2, PC3) rendered as 3D cross stereoscopic pair. Symbol colors as in legend of Fig. 30.Analyzed data set as in Figs. 23–28.

Figures 45–52. Tibial spurs and tarsal setation. Figures 45–46. Tibial and tarsal segments of leg III (45) and leg IV (46) of Grosphus madagascariensis in retrolateral view. Figures 47–48. Tibial and tarsal segments of leg III (47) and leg IV (48) of Teruelius limbatus in retrolateral view.Abbreviations:Tb, tibia; Bt, basitarsus (tarsomere I); Tt, telotarsus (tarsomere II); ts, tibial spur; vs, ventral setae of telotarsus. Figures 49–50. Horizontal bar plots of the mean ratio of tibial spur L/ tibia distal D for leg III (49) and leg IV (50) (characters 10 and 11, respectively) of Grosphus (n = 34, 9 spp.), Teruelius (n = 66, 21 spp.), Pseudolychas (n = 2, 2 spp.) and other outgroup taxa (n = 15, 8 spp.). Data from both sexes pooled. Error bars are standard errors. Discretization thresholds shown at step changes in ranked length. Color codes of bars as indicated in Fig. 16 legend. Figures 51–52. Ventral telotarsal setation. Figure 51. Logarithmic distribution of number of macrosetae on ventral telotarsus III (character 12) in Grosphus (cyan bars and symbols, n = 36, 8 spp.) and Teruelius (yellow bars and symbols, n = 12, 10 spp.). Macrosetal counts on abscissa plotted on logarithmic scale. Insets: UV fluorescence photomicrographs and maps of macroseta sockets (red dots) of ventral telotarsus III in Grosphus simoni (left) and Teruelius flavopiceus (right). Figure 52. Horizontal bar plot of ratios of mean blue channel intensities of dorsal vs. ventral regions-of-interest (ROI) (white boxes) in telotarsal images of Grosphus (cyan bars) and Teruelius (yellow bars). Insets: control images (*) of T. haeckeli sp. n. (a, leg III) and G. madagascariensis (c, leg III), original (upper) and resampled (lower); test images of T. sabineae (b, leg IV) and G. ambre (d, leg IV) (resampled from: Lourenço & Wilmé, 2016; Lourenço et al., 2018). Error bars: standard deviations of ROI pixels.

Figures 130–149. Ventral views of metasomal segment I showing variation in granulation and development of paired ventrosubmedian carinae in Teruelius. Labels and arrows as in Figs. 105–129. Reflected white light illumination: 130, 134, 137–138, 141, 146–149; UV fluorescence: 131–133, 135–136, 139–140, 142–145; resampled from published images: 130, 134, 141, 146, 148–149 (Lourenço & Wilmé, 2016; Lourenço et al., 2016; Lourenço, 2014; Lourenço & Wilmé, 2015b).

Figures 165–168. Metasoma and pedipalp characters. Figure 165. Horizontal bar plot of mean ratio R = d(Eb2, Eb3)/ d(Eb1, Eb2), of inter123 trichobothrial distances on pedipalp chela manus (character 36) for Grosphus (n = 50, 7 spp.), Teruelius (n = 61, 13 spp.), Pseudolychas (n = 6, 3 spp.) and other outgroup taxa (n = 25, 8 spp.). Error bars are standard errors. Discretization threshold at step change in ranked ratio. Figure 166. Bivariate logarithmic scatter plot of positions of pedipalp fixed finger trichobothria db vs. est for Grosphus (n = 40, 13 spp.) and Teruelius (n = 57, 21 spp.). Distances of db and est to distal terminus of fixed finger normalized to corresponding distance of manus Et (inset diagrams). Figure 167. Horizontal logarithmic bar plot of mean ratio of distances of trichobothria db and est from tip of fixed finger (character 35). Data as in Fig. 166. Error bars are standard errors. Discretization threshold at step change in ranked ratio. Color codes of symbols in Fig. 166 and bars in Figs. 165, 167–168 as in Fig. 16 legend. Figure 168. Horizontal bar plot of mean ratio of metasoma I L/W in males (character 18) of Grosphus (n = 20, 12 spp.), Teruelius (n = 28, 18 spp.), Pseudolychas (n = 5, 2 spp.) and other outgroup taxa (n = 12, 9 spp.). Error bars are standard errors. Discretization thresholds at step changes in ranked ratio.

Figures 173–178: Analysis of position of petite ‘trichobothrium’ d on pedipalp femur of Grosphus, Teruelius and Pseudolychas. Figures 2 173–174. Cartesian x-y coordinates for digitizing positions of d and dorsointernal carina in Grosphus madagascariensis (173) and Teruelius 2 olgae (174). Coordinates of d 2 are (d 2x, d 2y) indicated by cyan (173) or yellow (174) symbol. Positions of granules of dorsointernal carina indicated by black symbols. Granules fitted by B-spline (magenta curves) or parametric function (green curves). Ymax: asymptotic value of parametric function. Figure 175. Bivariate scatter plot of minimum distances, s, of d from parametric curve vs. d from B-spline curve (both 2 2 normalized against Y) for femora of Grosphus (n = 34, 14 spp.), Teruelius (n = 49, 20 spp.) and Pseudolychas (n = 3, 3 spp.). Gray diagonal max line: least squares regression for Grosphus and Teruelius data, R = 0.8131, P <0.0001. Symbol colors indicated in legend. Black arrows mark isolated outlier points. Figure 176. Bivariate scatter plot of means of minimum distances for species of Grosphus and Teruelius, summarizing data of Fig. 175. Error bars are standard errors. Gray diagonal line: least squares regression, R = 0.9700, P <0.0001. Figure 177. Bivariate scatter plot of normalized x-y coordinates of d 2 for data in Fig. 175. Abscissas normalized against femur length, L femur, ordinates against Y max. Figure 178. Bivariate scatter plot of means of normalized x-y coordinates for species of Grosphus and Teruelius, summarizing data of Fig. 177. Error bars are standard errors. Data extracted from images of specimens and published figures of femur in dorsal aspect showing granules and trichobothria, in which d 2 could be identified.

Figures 179–182. Morphometrics of telson lateral profiles. Figure 179. Elliptic Fourier analysis (EFA) of lateral profiles. Upper panel: EFA curve fits to profiles of Grosphus hirtus, G. mandena, Teruelius ankarafantsika and T. annulatus by Fourier series with cumulative terms up to and including second (blue), fourth (green), eighth (red) and sixteenth (orange) order harmonics. Telson profiles oriented with dorsal surface horizontal, start point at anterior limit of vesicle, peduncle truncated, area normalized. Lower panel: Histogram plots of harmonic loadings of first two principal components, PC1 and PC2, obtained from PCA of 64 standardized Fourier coefficients from up to sixteenth order harmonic terms, accounting for 30.33% and 19.54% of variance, respectively (total variance 49.87%). Bars are heat map coded by loading values, harmonics with highest positive loadings labelled (red bars). Figure 180. Bivariate scatter plot of subspace of first two principal components rotated (PC1*, PC2*) to minimize variance of Teruelius spp. (yellow symbols) along vertical axis. Data from 117 telson profiles from Grosphus (n = 35, 14 spp.), Teruelius (n = 65, 21 spp.), Pseudolychas (n = 6, 3 spp.) and other outgroup taxa (n = 11, 4 spp.), both males and females. Upper inset: scree plot of eigenvalue vs. PC number. Lower inset: legend for symbol colors: Grosphus ‘hirtus’ group (= G. angulatus sp. n., G. hirtus, G. polskyi, G. tavaratra, G. voahangyae), blue; other Grosphus spp., cyan; Teruelius flavopiceus, orange; T. ankarana, magenta; T. grandidieri, red; other Teruelius spp., yellow; Pseudolychas spp., black; other outgroup taxa, gray. Figure 181. Bivariate scatter plot of mean values of PC2* vs. PC1* for male telson profiles of Grosphus (12 spp.), Teruelius (16 spp.), Pseudolychas (3 spp.) and other outgroup taxa (4 spp.). Figure 182. Bivariate scatter plot of mean values of PC2* vs. PC1* for female telson profiles of Grosphus (7 spp.), Teruelius (15 spp.), Pseudolychas (3 spp.) and other outgroup taxa (4 spp.). Profile silhouette examples shown for analyzed species in Figs. 181–182. Error bars in Fig. 182 are standard errors. Symbol colors in Figs. 181–182 as in legend of Fig. 180.

Figure 183. Morphometrics of telson lateral profiles. Trivariate scatter plot of first three unrotated principal components (PC1, PC2, PC3) of harmonics extracted by Elliptic Fourier analysis (Fig. 179), rendered as a 3D cross stereoscopic pair. Symbol colors as in legend of Fig. 180.

Figures 196–199. Examples of MPTs retrieved by phylogenetic analyses of Grosphus and Teruelius, rooted by 4 different outgroup taxa: Charmus laneus (196), Karasbergia methueni (197), Lychas mucronatus (198) and Microcharmus variegatus (199). Weighting schemes and data matrices indicated above each tree (see Table 7). Numbers above nodes are jackknife by symmetric resampling supports, those below relative Bremer supports. Color coding of groups according to legend in Fig. 16.

Figures 200–203. Examples of MPTs retrieved by phylogenetic analyses of Grosphus and Teruelius, rooted by 4 different outgroup taxa: Pseudolychas ochraceus (200), P. pegleri (201), Tityobuthus monodi (202) and Somalicharmus whitmanae (203). Weighting schemes and data matrices indicated above each tree (see Table 7). Numbers above nodes are jackknife by symmetric resampling supports, those below relative Bremer supports. Color coding of groups according to legend in Fig. 16.

Figures 204–205. Mapping of unambiguous synapomorphies in two example MPTs retrieved by phylogenetic analyses of Grosphus and Teruelius, rooted by 2 different outgroup taxa: Charmus laneus (204) and Pseudolychas ochraceus (205). Numbers above nodes are discrete characters (Table 1), those below are discrete character states (Table 2).

Figures 206–209. Examples of MPTs retrieved by phylogenetic analyses of Grosphus and Teruelius, with multiple outgroup taxa in topologies fixed by backbone constraints. Figures 206–207. MPTs retrieved with 11 outgroup taxa in constrained topologies determined by the buthid MPT of Fig. 276, from discrete characters (206) or discrete and continuous characters (207), analyzed under implied weights. Figures 208–209. MPTs retrieved with 7 outgroup taxa in constrained topologies determined by molecular phylogenetic analyses of buthids (Fig. 190), from discrete characters (208) or discrete and continuous characters (209), analyzed under prior (209) or implied (208) weights. Numbers above nodes are jackknife by symmetric resampling supports, those below relative Bremer supports. Color coding of groups according to legend in Fig. 16.

Figures 328–331. Teruelius haeckeli sp. n., habitus. Figures 328– 329. Holotype male in dorsal (328) and ventral (329) views. Figures 330–331. Paratype female in dorsal (330) and ventral (331) views. Scale bars: 10 mm.

Figures 332–333. Teruelius haeckeli sp. n., holotype male. Carapace and tergites (332) and pectinal area and sternites (333).

Figures 334–335. Teruelius haeckeli sp. n., holotype male. Carapace and tergites (334) and coxosternal area and sternites (335). UV fluorescence.

Figures 336–337. Teruelius haeckeli sp. n., paratype female. Carapace and tergites (336) and coxosternal area and sternites (337). UV fluorescence.

Figures 338–357. Teruelius haeckeli sp. n., pedipalp.Figures 338–348. Holotype male. Right chela (338, 339, 340) and patella (341, 342, 343) in dorsal, external and ventral views, respectively. Right femur (344, 345, 346) in dorsal, ventrointernal and internal views, respectively. Dentition of right chela, movable (347) and fixed (348) fingers. Figures 349–357. Paratype female. Right chela (349, 350, 351) and patella (352, 353, 354) in dorsal, external and ventral views, respectively. Right femur (355, 356, 357) in dorsal, ventrointernal and internal views, respectively.

Figures 358–371. Teruelius haeckeli sp. n., pedipalp segments of male holotype (358–364) and female paratype (365–371) with trichobothrial pattern indicated. Chela in dorsal (358, 365), external (359, 366) and ventral (360, 367) views. Patella in dorsal (361, 368) and external (362, 369) views. Femur in dorsal (363, 370) and internal (364, 370) views.

Figures 372–379. Teruelius haeckeli sp. n., metasoma and telson. Figures 372–375. Paratype female. Metasoma and telson in right lateral (372), dorsal (373) and ventral (374) views. Telson in left lateral view (375). Figures 376–379. Holotype male. Metasoma and telson in right lateral (376), dorsal (377) and ventral (378) views. Telson in left lateral view (379). Scale bars: 10 mm (372–374), 10 mm (376–378).

Figures 380–387. Teruelius haeckeli sp. n., tarsi. Left basitarsi and telotarsi, legs I (380), II (382), III (384) and IV (386) in ventral (380, 382) and retrolateral (384, 386) views. Left telotarsi, legs I (381), II (383), III (385) and IV (387) in proventral (381, 383) and retroventral (385, 387) views.

Figures 388–392. Teruelius haeckeli sp. n., right hemispermatophore, paratype male. Figure 388. Capsule, part of flagellum and part of trunk, convex view. Pedicel truncated and lost during extraction. Scale bar: 1 mm. Figures 389–392. Capsule and part of flagellum, in convex compressed (389), convex (390), anterior (391) and posterior (392) views. Scale bar: 500 μm.

Figures 320–327. Grosphus angulatus sp. n., variation in femalebpt. Figures 320– 324. Basal pectines showing bpt shapesof holotype(322)and four paratypes (320–321, 323– 324). White arrow (320) indicates partially modified intermediate tooth bearing a sensorial area (cf. Figs. 43–44). Figure 325. Basal pectines of female G. hirtus showing typical bpt shape.UV fluorescence (320–325).Figures 326– 327. Bivariate scatter plots of bpt scores of G. angulatus sp. n. (black circles) and G. hirtus (gray circles) for principal components PC2 vs. PC1 (326) and PC4 vs. PC3 (327) obtained from PCA of 32 Fourier coefficients from up to eighth order harmonic terms in EFA of bpt shapes of Grosphus and Teruelius (cf. Figs. 29– 33).