treatments-xml/data/03/BB/87/03BB8789FFB7FFAF2720D4F6FD65FF44.xml

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<mods:title id="123860E42606740932385977EFD7099A">Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae)</mods:title>
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<emphasis id="B966EA8DFFB7FFA12720D4F6FDAAF9EF" bold="true" box="[113,536,1423,1450]" pageId="6" pageNumber="5">
Diagnosis of the ‘ 
<taxonomicName id="4C124D1CFFB7FFA12618D4F6FE0CF9EC" ID-CoL="4R7C" authorityName="Simon" authorityYear="1880" box="[329,446,1423,1450]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B966EA8DFFB7FFA12618D4F6FE0CF9EC" bold="true" box="[329,446,1423,1450]" italics="true" pageId="6" pageNumber="5">Grosphus</emphasis>
</taxonomicName>
’ group
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C3086514FFB7FFA127F0D4C8FA86F8C5" pageId="6" pageNumber="5" type="diagnosis">
<paragraph id="8BAD369FFFB7FFA127F0D4C8FDECFA6F" blockId="6.[111,773,1423,2025]" pageId="6" pageNumber="5">
The three buthid genera 
<taxonomicName id="4C124D1CFFB7FFA126B7D4C8FD4DF98F" authority="Simon, 1880" authorityName="Simon" authorityYear="1880" box="[486,767,1457,1481]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B966EA8DFFB7FFA126B7D4C8FDFFF98F" box="[486,589,1457,1481]" italics="true" pageId="6" pageNumber="5">Grosphus</emphasis>
<bibRefCitation id="EF834B6EFFB7FFA12535D4C8FD4DF98F" author="Simon" box="[612,767,1457,1481]" firstAuthor="Simon" pageId="6" pageNumber="5" pagination="377 - 398" refId="ref42670" refString="SIMON, E. 1880. Etudes arachnologiques 12 e Memorie (1). XVIII. Descriptions de genres et especes de l'ordre des Scorpiones. Annales de la Societe Entomologique de France, (5), 10: 377 - 398." type="journal article" year="1880">Simon, 1880</bibRefCitation>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12720D4A8FE1DF9AF" authority="Lourenco, 1995" authorityName="Lourenco" authorityYear="1995" box="[113,431,1489,1513]" class="Arachnida" family="Buthidae" genus="Neogrosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B966EA8DFFB7FFA12720D4A8FF4FF9AF" box="[113,253,1489,1513]" italics="true" pageId="6" pageNumber="5">Neogrosphus</emphasis>
<bibRefCitation id="EF834B6EFFB7FFA12655D4ABFE1DF9AF" author="LOURENCO, W. R." box="[260,431,1489,1513]" journalOrPublisher="Bulletin du Museum National d'Histoire naturelle, Paris (Zoologie, Biologie et Ecologie Animale)" pageId="6" pageNumber="5" pagination="95 - 106" part="4" refId="ref37469" refString="LOURENCO, W. R. 1995 a. Description de trois noveaux genres et quatre nouvelles especes de scorpions Buthidae de Madagascar. Bulletin du Museum National d'Histoire naturelle, Paris (Zoologie, Biologie et Ecologie Animale), (4), 17 A (1 - 2): 95 - 106." title="Description de trois noveaux genres et quatre nouvelles especes de scorpions Buthidae de Madagascar" type="journal article" year="1995">Lourenço, 1995</bibRefCitation>
</taxonomicName>
, and 
<taxonomicName id="4C124D1CFFB7FFA126B6D4A8FDFBF9AF" authority="Lowe &amp; Kovařík, 2019" authorityName="Lowe &amp; Kovařík" authorityYear="2019" box="[487,585,1489,1513]" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B966EA8DFFB7FFA126B6D4A8FDFBF9AF" box="[487,585,1489,1513]" italics="true" pageId="6" pageNumber="5">Teruelius</emphasis>
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<emphasis id="B966EA8DFFB7FFA12501D4A8FDCAF9AF" bold="true" box="[592,632,1489,1513]" pageId="6" pageNumber="5">gen</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA125D4D4A8FD26F9AF" bold="true" box="[645,660,1489,1513]" pageId="6" pageNumber="5">n</emphasis>
.
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comprise a distinct assemblage of 
<collectingCountry id="F305760FFFB7FFA126D7D488FDB5FA4F" box="[390,519,1521,1545]" name="Madagascar" pageId="6" pageNumber="5">Madagascar</collectingCountry>
buthids (= ‘ 
<taxonomicName id="4C124D1CFFB7FFA125C4D488FD4EFA4F" authorityName="Simon" authorityYear="1880" box="[661,764,1521,1545]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B966EA8DFFB7FFA125C4D488FD4EFA4F" box="[661,764,1521,1545]" italics="true" pageId="6" pageNumber="5">Grosphus</emphasis>
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’ group) sharing the following set of characters:
</paragraph>
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Carapace subrectangular, weakly trapezoidal or nearly parallel-sided, surface densely granular, carinae indistinct except for superciliary carinae; frontal region of carapace flat, not sloped towards anterior margin; median eyes large, median ocular tubercle prominent, located forward of the carapace centroid (
<figureCitation id="13292A1AFFB7FFA12611D7A8FE53FAAF" box="[320,481,1745,1769]" captionStart="Figures 165–180" captionStartId="63.[113,190,1958,1980]" captionTargetBox="[113,1473,312,1921]" captionTargetId="figure-73@63.[113,1473,312,1921]" captionTargetPageId="63" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143694" httpUri="https://zenodo.org/record/7143694/files/figure.png" pageId="6" pageNumber="5">Figs. 165–180</figureCitation>
); 5 pairs of lateral eyes (3 large, 2 small) (
<figureCitation id="13292A1AFFB7FFA12649D788FE04FB4F" box="[280,438,1777,1801]" captionStart="Figures 227–230" captionStartId="67.[113,190,1951,1973]" captionTargetBox="[113,1474,234,1902]" captionTargetId="figure-2@67.[113,1474,234,1902]" captionTargetPageId="67" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143712" httpUri="https://zenodo.org/record/7143712/files/figure.png" pageId="6" pageNumber="5">Figs. 227–230</figureCitation>
); chelicerae with typical buthid dentition on fixed and movable fingers (
<bibRefCitation id="EF834B6EFFB7FFA1257AD668FD74FB6F" author="Vachon" box="[555,710,1809,1833]" firstAuthor="Vachon" pageId="6" pageNumber="5" pagination="161 - 166" refId="ref43960" refString="VACHON, M. 1963. De l'utilite, en systematique, d'une nomenclature des dents de cheliceres chez les scorpions. Bulletin du Museum national d'Histoire Naturelle, Paris, (2), 35 (2): 161 - 166." type="journal article" year="1963">Vachon, 1963</bibRefCitation>
), two enlarged denticles on ventral surface of fixed finger (
<figureCitation id="13292A1AFFB7FFA125C5D648FF2FFB2F" captionStart="Figures 231–238" captionStartId="68.[112,189,1958,1980]" captionTargetId="figure-61@68.[113,1474,234,1421]" captionTargetPageId="68" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143714" httpUri="https://zenodo.org/record/7143714/files/figure.png" pageId="6" pageNumber="5">Figs. 231– 238</figureCitation>
); sternum 
<typeStatus id="54A9883DFFB7FFA1265FD62BFE8EFB2C" box="[270,316,1874,1898]" pageId="6" pageNumber="5">type</typeStatus>
1, subtriangular; tergites granular, tergites I–VI with single, weak median carina, tergite VII with weak median carina and 2 pairs of strong lateral carinae; metasoma moderately elongate, segments I–III with 8–10 carinae, IV with 8 carinae, V with 3–5 carinae; telson vesicle bulbous, ovoid or elongate, with or without subaculear tubercle (
<figureCitation id="13292A1AFFB7FFA122DDD192FC23FD64" captionStart="Figures 181–195" captionStartId="64.[113,190,1953,1975]" captionTargetId="figure-71@64.[113,1474,242,1670]" captionTargetPageId="64" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143700" httpUri="https://zenodo.org/record/7143700/files/figure.png" pageId="6" pageNumber="5">Figs. 181–195</figureCitation>
); pectines with fulcra, 13–41 teeth, female with basal pectinal tooth dilated or elongated, lacking peg sensillae (
<figureCitation id="13292A1AFFB7FFA122DDD052FCC4FD24" captionStart="Figures 40–51" captionStartId="22.[113,188,1930,1952]" captionTargetBox="[113,1474,234,1920]" captionTargetId="figure-124@22.[113,1474,234,1922]" captionTargetPageId="22" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143634" httpUri="https://zenodo.org/record/7143634/files/figure.png" pageId="6" pageNumber="5">Figs. 40–51</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA124D6D032FC58FD25" box="[903,1002,331,355]" captionStart="Figures 196–210" captionStartId="65.[113,190,1950,1972]" captionTargetId="figure-77@65.[113,1474,234,1589]" captionTargetPageId="65" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143706" httpUri="https://zenodo.org/record/7143706/files/figure.png" pageId="6" pageNumber="5">196–210</figureCitation>
); hemispermatophore flagellum thicker at base, narrowed proximally, thickened distally (
<figureCitation id="13292A1AFFB7FFA12267D012FCCEFDE5" captionStart="Figures 52–70" captionStartId="24.[113,190,1591,1613]" captionTargetBox="[116,1470,248,1526]" captionTargetId="figure-361@24.[113,1474,248,1527]" captionTargetPageId="24" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143636" httpUri="https://zenodo.org/record/7143636/files/figure.png" pageId="6" pageNumber="5">Figs. 52, 58, 60, 67</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA124DCD0F2FB85FDE5" box="[909,1079,394,419]" captionStart="Figures 71–85" captionStartId="26.[113,190,1701,1723]" captionTargetBox="[149,1434,231,1678]" captionTargetId="figure-287@26.[146,1446,225,1690]" captionTargetPageId="26" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143641" httpUri="https://zenodo.org/record/7143641/files/figure.png" pageId="6" pageNumber="5">71, 75, 78, 84</figureCitation>
); pedipalp chela elongate, smooth, carinae obsolete, surface typically with numerous short macrosetae (
<figureCitation id="13292A1AFFB7FFA124E9D0B2FB8EFDA5" box="[952,1084,459,483]" captionStart="Figures 21–27" captionStartId="14.[113,188,1859,1881]" captionTargetBox="[114,1474,227,1840]" captionTargetId="figure-206@14.[113,1474,222,1851]" captionTargetPageId="14" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143620" httpUri="https://zenodo.org/record/7143620/files/figure.png" pageId="6" pageNumber="5">Figs. 21–24</figureCitation>
); finger dentition composed of 8–15 discrete linear rows of granules or denticles, each slightly oblique with proximal ends directed externally; rows either non-overlapping or slightly imbricated, proximal 3 granules in each row enlarged, 2 of these slightly displaced outwards as ‘external accessory’ granules; series of large, dentate internal accessory granules present, offset from main rows; both chela fingers with enlarged apical teeth, 3–4 external subdistal granules; pedipalps sexually dimorphic, dentate margins of fingers weakly or strongly scalloped proximally in males, straight in females, manus of males broader than that of females; trichobothrial pattern orthobothriotaxic, 
<typeStatus id="54A9883DFFB7FFA12250D252FA9DFF05" box="[1281,1327,811,835]" pageId="6" pageNumber="5">type</typeStatus>
A (
<bibRefCitation id="EF834B6EFFB7FFA1223BD252FCDBFF25" author="Vachon" firstAuthor="Vachon" pageId="6" pageNumber="5" pagination="857 - 958" refId="ref44039" refString="VACHON, M. 1974. Etude des caracteres utilises pour classer les familles et les genres de Scorpions (Arachnides). 1. La trichobothriotaxie enArachnologie, Sigles trichobothriaux et types de trichobothriotaxie chez les Scorpions. Bulletin du Museum National d'Histoire naturelle, Paris, (3), 140 (Zool. 104), mai-juin 1973: 857 - 958." type="journal article" year="1974">Vachon, 1974</bibRefCitation>
), with femur 
<emphasis id="B966EA8DFFB7FFA124A8D232FBB4FF25" box="[1017,1030,843,867]" italics="true" pageId="6" pageNumber="5">d</emphasis>
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- 
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- 
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in
</quantity>
α-configuration (
<bibRefCitation id="EF834B6EFFB7FFA1224FD232FA04FF24" author="Vachon" box="[1310,1462,842,867]" firstAuthor="Vachon" pageId="6" pageNumber="5" pagination="1597 - 1599" refId="ref44105" refString="VACHON, M. 1975. Sur l'utilisation de la trichobothriotaxie du bras des pedipalpes des Scorpions (Arachnides) dans le classement des genres de famille des Buthidae Simon. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, (D), 281 (21): 1597 - 1599." type="journal article" year="1975">Vachon, 1975</bibRefCitation>
), patella 
<emphasis id="B966EA8DFFB7FFA124D3D212FC3DFFC5" box="[898,911,875,899]" italics="true" pageId="6" pageNumber="5">d</emphasis>
<subScript id="179634DAFFB7FFA124DED202FC25FFCF" attach="left" box="[911,919,891,905]" fontSize="6" pageId="6" pageNumber="5">3</subScript>
external to dorsomedian carina (
<bibRefCitation id="EF834B6EFFB7FFA12257D212FA04FFC4" author="FET, V. &amp; M. E. SOLEGLAD &amp; G. LOWE" box="[1286,1462,874,899]" journalOrPublisher="Euscorpius" pageId="6" pageNumber="5" pagination="1 - 40" part="23" refId="ref33769" refString="FET, V., M. E. SOLEGLAD &amp; G. LOWE. 2005. A new trichobothrial character for the high-level systematics of Buthoidea (Scorpiones: Buthida). Euscorpius, 23: 1 - 40." title="A new trichobothrial character for the high-level systematics of Buthoidea (Scorpiones: Buthida)" type="journal article" year="2005">Fet et al., 2005</bibRefCitation>
); patella 
<emphasis id="B966EA8DFFB7FFA1242CD2F2FC2EFFE4" box="[893,924,907,930]" italics="true" pageId="6" pageNumber="5">em</emphasis>
much closer to 
<emphasis id="B966EA8DFFB7FFA12313D2F2FBD2FFE4" box="[1090,1120,907,930]" italics="true" pageId="6" pageNumber="5">est</emphasis>
and 
<emphasis id="B966EA8DFFB7FFA123C0D2F2FB16FFE4" box="[1169,1188,907,930]" italics="true" pageId="6" pageNumber="5">et</emphasis>
, than to 
<emphasis id="B966EA8DFFB7FFA123AFD2F2FA90FFE5" box="[1278,1314,907,931]" italics="true" pageId="6" pageNumber="5">esb</emphasis>
<subScript id="179634DAFFB7FFA12273D2E2FA98FFEF" attach="left" box="[1314,1322,923,937]" fontSize="6" pageId="6" pageNumber="5">1</subScript>
and 
<emphasis id="B966EA8DFFB7FFA1220AD2F2FACDFFE5" box="[1371,1407,907,931]" italics="true" pageId="6" pageNumber="5">esb</emphasis>
<subScript id="179634DAFFB7FFA1222ED2E2FA35FFEF" attach="left" box="[1407,1415,923,937]" fontSize="6" pageId="6" pageNumber="5">2</subScript>
, with 
<emphasis id="B966EA8DFFB7FFA12461D2D2FC20FF84" box="[816,914,939,962]" italics="true" pageId="6" pageNumber="5">em -est -et</emphasis>
usually forming a compact triad (
<figureCitation id="13292A1AFFB7FFA12259D2D2FAC2FF84" box="[1288,1392,938,963]" captionStart="Figures 341–347" captionStartId="81.[113,190,1954,1976]" captionTargetBox="[136,1456,253,1871]" captionTargetId="figure-48@81.[123,1497,234,1879]" captionTargetPageId="81" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143752" httpUri="https://zenodo.org/record/7143752/files/figure.png" pageId="6" pageNumber="5">Figs. 345</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA1222FD2D2FA04FF85" box="[1406,1462,939,963]" captionStart="Figures 477–490" captionStartId="100.[113,190,1928,1950]" captionTargetBox="[110,1468,256,1846]" captionTargetId="figure-92@100.[110,1471,244,1880]" captionTargetPageId="100" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143803" httpUri="https://zenodo.org/record/7143803/files/figure.png" pageId="6" pageNumber="5">481a</figureCitation>
); chela manus with 
<emphasis id="B966EA8DFFB7FFA124A6D2B2FBA7FFA5" box="[1015,1045,971,995]" italics="true" pageId="6" pageNumber="5">Eb</emphasis>
<subScript id="179634DAFFB7FFA12344D2A2FBAFFFAF" attach="left" box="[1045,1053,987,1001]" fontSize="6" pageId="6" pageNumber="5">1</subScript>
- 
<emphasis id="B966EA8DFFB7FFA12377D2B2FBF6FFA5" box="[1062,1092,971,995]" italics="true" pageId="6" pageNumber="5">Eb</emphasis>
<subScript id="179634DAFFB7FFA12312D2A2FBF9FFAF" attach="left" box="[1091,1099,987,1001]" fontSize="6" pageId="6" pageNumber="5">2</subScript>
angled distally, 
<emphasis id="B966EA8DFFB7FFA123AED2B2FAAFFFA5" box="[1279,1309,971,995]" italics="true" pageId="6" pageNumber="5">Eb</emphasis>
<subScript id="179634DAFFB7FFA1224CD2A2FA97FFAF" attach="left" box="[1309,1317,987,1001]" fontSize="6" pageId="6" pageNumber="5">1</subScript>
- 
<emphasis id="B966EA8DFFB7FFA1227FD2B2FAFEFFA5" box="[1326,1356,971,995]" italics="true" pageId="6" pageNumber="5">Eb</emphasis>
<subScript id="179634DAFFB7FFA1221AD2A2FAE1FFAF" attach="left" box="[1355,1363,987,1001]" fontSize="6" pageId="6" pageNumber="5">2</subScript>
- 
<emphasis id="B966EA8DFFB7FFA1220DD2B2FAC8FFA5" box="[1372,1402,971,995]" italics="true" pageId="6" pageNumber="5">Eb</emphasis>
<subScript id="179634DAFFB7FFA12228D2A2FA33FFAF" attach="left" box="[1401,1409,987,1001]" fontSize="6" pageId="6" pageNumber="5">3</subScript>
acute angle opening in proximal direction (γ-configuration) (
<figureCitation id="13292A1AFFB7FFA122DDD292FCE9F865" captionStart="Figures 341–347" captionStartId="81.[113,190,1954,1976]" captionTargetBox="[136,1456,253,1871]" captionTargetId="figure-48@81.[123,1497,234,1879]" captionTargetPageId="81" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143752" httpUri="https://zenodo.org/record/7143752/files/figure.png" pageId="6" pageNumber="5">Figs. 342</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA12436D572FC2CF865" box="[871,926,1035,1059]" captionStart="Figures 477–490" captionStartId="100.[113,190,1928,1950]" captionTargetBox="[110,1468,256,1846]" captionTargetId="figure-92@100.[110,1471,244,1880]" captionTargetPageId="100" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143803" httpUri="https://zenodo.org/record/7143803/files/figure.png" pageId="6" pageNumber="5">478a</figureCitation>
); chela with 
<emphasis id="B966EA8DFFB7FFA12378D572FBF6F865" box="[1065,1092,1035,1059]" italics="true" pageId="6" pageNumber="5">db</emphasis>
in proximal half to middle of fixed finger; legs III–IV with tibial spurs (
<figureCitation id="13292A1AFFB7FFA123E6D552FAE0F805" box="[1207,1362,1067,1091]" captionStart="Figures 211–226" captionStartId="66.[113,190,1920,1942]" captionTargetBox="[117,1466,234,1743]" captionTargetId="figure-88@66.[113,1474,234,1743]" captionTargetPageId="66" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143708" httpUri="https://zenodo.org/record/7143708/files/figure.png" pageId="6" pageNumber="5">Figs. 211–226</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA1220FD552FA0CF805" box="[1374,1470,1067,1091]" captionStart="Figures 259–262" captionStartId="71.[116,193,2009,2031]" captionTargetBox="[130,1463,270,1921]" captionTargetId="figure-20@71.[113,1474,266,1956]" captionTargetPageId="71" captionText="Figures 259–262. Grosphus goudoti, male holotype, distal segments of right legs I–IV, retrolateral views." figureDoi="http://doi.org/10.5281/zenodo.7143726" httpUri="https://zenodo.org/record/7143726/files/figure.png" pageId="6" pageNumber="5">261–262</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA12461D532FC22F825" box="[816,912,1099,1123]" captionStart="Figures 316–319" captionStartId="79.[113,190,1998,2020]" captionTargetBox="[117,1450,260,1895]" captionTargetId="figure-23@79.[113,1474,234,1908]" captionTargetPageId="79" captionText="Figures 316–319. Grosphus madagascariensis, male, right legs I–IV tibia, basitarsus and telotarsus, retrolateral views. Scale bar: 2 mm." figureDoi="http://doi.org/10.5281/zenodo.7143748" httpUri="https://zenodo.org/record/7143748/files/figure.png" pageId="6" pageNumber="5">318–319</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA124F1D533FBB2F824" box="[928,1024,1098,1122]" captionStart="Figures 362–365" captionStartId="85.[113,190,1983,2005]" captionTargetBox="[114,1468,253,1900]" captionTargetId="figure-36@85.[113,1474,234,1926]" captionTargetPageId="85" captionText="Figures 362–365. Grosphus voahangyae, male, left legs I–IV tibia, basitarsus and telotarsus, retrolateral views. Leg I (362), leg II (363), leg III (364), leg IV (365). Scale bar: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7143766" httpUri="https://zenodo.org/record/7143766/files/figure.png" pageId="6" pageNumber="5">364–365</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA1235ED532FBC2F825" box="[1039,1136,1099,1123]" captionStart="Figures 409–417" captionStartId="89.[113,190,1958,1980]" captionTargetBox="[113,1474,234,1814]" captionTargetId="figure-58@89.[113,1474,234,1838]" captionTargetPageId="89" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143778" httpUri="https://zenodo.org/record/7143778/files/figure.png" pageId="6" pageNumber="5">414–417</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA1232ED532FB6DF825" box="[1151,1247,1099,1123]" captionStart="Figures 477–490" captionStartId="100.[113,190,1928,1950]" captionTargetBox="[110,1468,256,1846]" captionTargetId="figure-92@100.[110,1471,244,1880]" captionTargetPageId="100" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143803" httpUri="https://zenodo.org/record/7143803/files/figure.png" pageId="6" pageNumber="5">487–488</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA123BFD533FAFCF824" box="[1262,1358,1098,1122]" captionStart="Figures 512–515" captionStartId="104.[117,194,1981,2003]" captionTargetBox="[146,1452,271,1937]" captionTargetId="figure-34@104.[135,1459,234,1949]" captionTargetPageId="104" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143813" httpUri="https://zenodo.org/record/7143813/files/figure.png" pageId="6" pageNumber="5">514–515</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA1220FD533FA0CF824" box="[1374,1470,1098,1122]" captionStart="Figures 538–541" captionStartId="108.[113,190,1981,2003]" captionTargetBox="[129,1451,241,1964]" captionTargetId="figure-37@108.[123,1459,234,1977]" captionTargetPageId="108" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143821" httpUri="https://zenodo.org/record/7143821/files/figure.png" pageId="6" pageNumber="5">540–541</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA12461D513FC22F8C4" box="[816,912,1130,1154]" captionStart="Figures 576–579" captionStartId="113.[113,190,1981,2003]" captionTargetBox="[122,1462,264,1906]" captionTargetId="figure-33@113.[110,1474,245,1928]" captionTargetPageId="113" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7143831" httpUri="https://zenodo.org/record/7143831/files/figure.png" pageId="6" pageNumber="5">578–579</figureCitation>
, 
<figureCitation id="13292A1AFFB7FFA124CAD512FC4FF8C5" box="[923,1021,1131,1155]" captionStart="Figures 610–619" captionStartId="126.[113,190,1939,1961]" captionTargetBox="[119,1437,234,1911]" captionTargetId="figure-79@126.[119,1452,234,1925]" captionTargetPageId="126" captionText="Figures 610–619. Neogrosphus griveaudi, male (610–613, 617–619) and female (614–616). Figures 610, 614. Sternopectinal region. Figures 611, 615. Carapace and tergites I–III. Figure 612. Pedipalp chela, movable finger dentate margin. Figures 613, 616. Pedipalp chela in dorsal views. Figure 617. Metasoma V and telson lateral view. Figures 618–619. Right leg III (618) and leg IV (619), tibia, basitarsus and telotarsus, retrolateral views." figureDoi="http://doi.org/10.5281/zenodo.7143863" httpUri="https://zenodo.org/record/7143863/files/figure.png" pageId="6" pageNumber="5">618–619</figureCitation>
), tarsi without bristle-combs.
</paragraph>
</subSubSection>
<subSubSection id="C3086514FFB7FFAF2461D5D5FD65FF44" lastPageId="8" lastPageNumber="7" pageId="6" pageNumber="5" type="discussion">
<paragraph id="8BAD369FFFB7FFAF2461D5D5FD65FF44" blockId="6.[816,1474,1195,2022]" lastBlockId="8.[113,772,235,771]" lastPageId="8" lastPageNumber="7" pageId="6" pageNumber="5">
REMARKS. In describing the first ‘ 
<taxonomicName id="4C124D1CFFB7FFA123FED5D7FAA4F880" authorityName="Simon" authorityYear="1880" box="[1199,1302,1198,1222]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B966EA8DFFB7FFA123FED5D7FAA4F880" box="[1199,1302,1198,1222]" italics="true" pageId="6" pageNumber="5">Grosphus</emphasis>
</taxonomicName>
’ group species, 
<taxonomicName id="4C124D1CFFB7FFA12461D5B7FA29F8A0" authority=", Gervais" authorityName=", Gervais" authorityYear="1843" box="[816,1435,1230,1254]" class="Arachnida" family="Scorpionidae" genus="Scorpio" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="madagascariensis" subGenus="Androctonus">
<emphasis id="B966EA8DFFB7FFA12461D5B7FC31F8A3" box="[816,899,1230,1253]" italics="true" pageId="6" pageNumber="5">Scorpio</emphasis>
(
<emphasis id="B966EA8DFFB7FFA124C4D5B7FBAEF8A0" box="[917,1052,1230,1254]" italics="true" pageId="6" pageNumber="5">Androctonus</emphasis>
) 
<emphasis id="B966EA8DFFB7FFA1237CD5B7FB5CF8A0" box="[1069,1262,1230,1254]" italics="true" pageId="6" pageNumber="5">madagascariensis</emphasis>
, 
<bibRefCitation id="EF834B6EFFB7FFA123ACD5B7FA29F8A0" author="GERVAIS, P. M." bookContentInfo="418 pp." box="[1277,1435,1230,1254]" editor="Walckenaer, C. A." journalOrPublisher="Librarie Encyclopedique de Roret, Paris" pageId="6" pageNumber="5" pagination="14 - 74" part="3" refId="ref34383" refString="GERVAIS, P. M. 1844. Scorpions. Pp. 14 - 74 in: Walckenaer, C. A. (Ed.). Histoire Naturelle des Insectes. Apteres. Librarie Encyclopedique de Roret, Paris, 3 (8), 418 pp." title="Scorpions" type="book chapter" volumeTitle="Histoire Naturelle des Insectes. Apteres" year="1844">Gervais (1844</bibRefCitation>
</taxonomicName>
: pl. XI, fig. 3) illustrated the carapace showing forward placement of the median eyes, and also accurately depicted five pairs of lateral eyes, now recognized to be the prevalent buthid configuration (
<bibRefCitation id="EF834B6EFFB7FFA1249FD437FB63F920" author="LORIA, S. F. &amp; L. PRENDINI" box="[974,1233,1358,1382]" journalOrPublisher="PLoS ONE" pageId="6" pageNumber="5" pagination="112913" part="9" refId="ref37170" refString="LORIA, S. F. &amp; L. PRENDINI. 2014. Homology of the lateral eyes of Scorpiones: a six-ocellus model. PLoS ONE 9 (12): e 112913. doi: 10.1371 / journal. pone. 0112913." title="Homology of the lateral eyes of Scorpiones: a six-ocellus model" type="journal article" year="2014">Loria &amp; Prendini, 2014</bibRefCitation>
; 
<bibRefCitation id="EF834B6EFFB7FFA1238AD437FA2BF920" author="Yang" box="[1243,1433,1358,1382]" etAl="et al." firstAuthor="Yang" pageId="6" pageNumber="5" pagination="55125" refId="ref44730" refString="YANG, X., Y. NORMA-RASHID, W. R. LOURENCO &amp; M. ZHU. 2013. True lateral eye numbers for extant buthids: a new discovery on an old character. PLoS ONE, 8 (1): e 55125. doi: 10.1371 / journal. pone. 0055125" type="journal article" year="2013">Yang et al., 2013</bibRefCitation>
). In spite of this, 
<bibRefCitation id="EF834B6EFFB7FFA12491D417FBF6F9C0" author="FAGE, L." box="[960,1092,1390,1414]" journalOrPublisher="Societe d'Editions Geographiques Maritimes et Coloniales, Paris" pageId="6" pageNumber="5" pagination="637 - 693" refId="ref33536" refString="FAGE, L. 1929. Les scorpions de Madagascar, leur affinites, leur distribution geographique. Pp. 637 - 693 in: Faune des Colonies Francaises. Societe d'Editions Geographiques Maritimes et Coloniales, Paris." title="Les scorpions de Madagascar, leur affinites, leur distribution geographique" type="book chapter" volumeTitle="Faune des Colonies Francaises" year="1929">Fage (1929)</bibRefCitation>
incorrectly declared that 
<taxonomicName id="4C124D1CFFB7FFA1220AD417FA70F9C0" authorityName="Simon" authorityYear="1880" box="[1371,1474,1390,1414]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B966EA8DFFB7FFA1220AD417FA70F9C0" box="[1371,1474,1390,1414]" italics="true" pageId="6" pageNumber="5">Grosphus</emphasis>
</taxonomicName>
(sensu lato) only bore 3 pairs of lateral eyes, and 
<bibRefCitation id="EF834B6EFFB7FFA1220DD4F7FC37F980" author="LOURENCO, W. R." bookContentInfo="102 pp." journalOrPublisher="Museum National d'Histoire Naturelle, Paris" pageId="6" pageNumber="5" refId="ref37632" refString="LOURENCO, W. R. 1996 b. Faune de Madagascar. 87. Scorpions (Chelicerata, Scorpiones). Museum National d'Histoire Naturelle, Paris. 102 pp." title="Faune de Madagascar. 87. Scorpions (Chelicerata, Scorpiones)" type="book" year="1996">Lourenço (1996b)</bibRefCitation>
cited only 3–4 pairs. Moreover, only 3 pairs were described for: 
<taxonomicName id="4C124D1CFFB7FFA12483D4B7FB35F9A0" authorityName="Lourenco, Wilme &amp; Waeber" authorityYear="2018" box="[978,1159,1486,1510]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="ambre">
<emphasis id="B966EA8DFFB7FFA12483D4B7FB35F9A0" box="[978,1159,1486,1510]" italics="true" pageId="6" pageNumber="5">Grosphus ambre</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA123C9D4B7FA8BF9A0" authorityName="Lourenco, Goodman &amp; Ramilijaona" authorityYear="2004" box="[1176,1337,1486,1510]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="darainensis">
<emphasis id="B966EA8DFFB7FFA123C9D4B7FB19F9A3" box="[1176,1195,1486,1509]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA123ECD4B7FA8BF9A0" box="[1213,1337,1486,1510]" italics="true" pageId="6" pageNumber="5">darainensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA1221AD4B7FA09F9A0" authorityName="Lourenco" authorityYear="2001" box="[1355,1467,1486,1510]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="garciai">
<emphasis id="B966EA8DFFB7FFA1221AD4B7FAECF9A3" box="[1355,1374,1486,1509]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1223ED4B6FA09F9A0" box="[1391,1467,1487,1510]" italics="true" pageId="6" pageNumber="5">garciai</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12461D497FC17FA40" authorityName="Lourenco &amp; Goodman" authorityYear="2006" box="[816,933,1518,1542]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="goudoti">
<emphasis id="B966EA8DFFB7FFA12461D497FCF1FA43" box="[816,835,1518,1541]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12405D497FC17FA40" box="[852,933,1518,1542]" italics="true" pageId="6" pageNumber="5">goudoti</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA124E7D497FB82FA40" authorityName="Lourenco, Wilme, Soarimalala &amp; Waeber, Previous" authorityYear="2017" box="[950,1072,1518,1542]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="halleuxi">
<emphasis id="B966EA8DFFB7FFA124E7D497FC7BFA43" box="[950,969,1518,1541]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1248BD497FB82FA40" box="[986,1072,1518,1542]" italics="true" pageId="6" pageNumber="5">halleuxi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12310D497FB10FA40" baseAuthorityName="Lourenco" baseAuthorityYear="2000" box="[1089,1186,1518,1542]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="hirtus">
<emphasis id="B966EA8DFFB7FFA12310D497FBE6FA43" box="[1089,1108,1518,1541]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12335D497FB10FA40" box="[1124,1186,1518,1542]" italics="true" pageId="6" pageNumber="5">hirtus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA123E2D497FA2AFA40" authorityName=", Gervais" authorityYear="1843" box="[1203,1432,1518,1542]" class="Arachnida" family="Scorpionidae" genus="Scorpio" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="madagascariensis">
<emphasis id="B966EA8DFFB7FFA123E2D497FB74FA43" box="[1203,1222,1518,1541]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12386D497FA2AFA40" box="[1239,1432,1518,1542]" italics="true" pageId="6" pageNumber="5">madagascariensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA122F9D497FCC4FA60" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="makay">
<emphasis id="B966EA8DFFB7FFA122F9D497FA09FA43" box="[1448,1467,1518,1541]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12461D777FCC4FA60" box="[816,886,1550,1574]" italics="true" pageId="6" pageNumber="5">makay</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA124D9D777FBA2FA60" authorityName="Lourenco" authorityYear="2005" box="[904,1040,1550,1574]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="mandena">
<emphasis id="B966EA8DFFB7FFA124D9D777FC29FA63" box="[904,923,1550,1573]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA124FFD777FBA2FA60" box="[942,1040,1550,1574]" italics="true" pageId="6" pageNumber="5">mandena</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12373D777FB74FA60" authorityName="Lourenco &amp; Goodman" authorityYear="2009" box="[1058,1222,1550,1574]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="mayottensis">
<emphasis id="B966EA8DFFB7FFA12373D777FB87FA63" box="[1058,1077,1550,1573]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12319D776FB74FA60" box="[1096,1222,1551,1574]" italics="true" pageId="6" pageNumber="5">mayottensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12389D777FAF8FA60" authorityName="Lourenco, Qi &amp; Goodman" authorityYear="2007" box="[1240,1354,1550,1574]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="polskyi">
<emphasis id="B966EA8DFFB7FFA12389D777FB59FA63" box="[1240,1259,1550,1573]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA123AFD777FAF8FA60" box="[1278,1354,1550,1574]" italics="true" pageId="6" pageNumber="5">polskyi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA1220AD777FA09FA60" authorityName="Lourenco" authorityYear="2013" box="[1371,1467,1550,1574]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="rossii">
<emphasis id="B966EA8DFFB7FFA1220AD777FADCFA63" box="[1371,1390,1550,1573]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA122D0D776FA09FA60" box="[1409,1467,1551,1574]" italics="true" pageId="6" pageNumber="5">rossii</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12461D757FC2EFA00" authorityName="Lourenco, Goodman &amp; Ramilijaona" authorityYear="2004" box="[816,924,1582,1606]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="simoni">
<emphasis id="B966EA8DFFB7FFA12461D757FCF1FA03" box="[816,835,1582,1605]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12404D756FC2EFA00" box="[853,924,1583,1606]" italics="true" pageId="6" pageNumber="5">simoni</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA124FCD757FBD9FA00" authorityName="Lourenco, Wilme, Soarimalala &amp; Waeber" authorityYear="2017" box="[941,1131,1582,1606]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="rakotoariveloi">
<emphasis id="B966EA8DFFB7FFA124FCD757FC72FA03" box="[941,960,1582,1605]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12483D757FBD9FA00" box="[978,1131,1582,1606]" italics="true" pageId="6" pageNumber="5">rakotoariveloi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA1232DD757FAB7FA00" authorityName="Lourenco, Soarimalala &amp; Goodman" authorityYear="2009" box="[1148,1285,1582,1606]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="tavaratra">
<emphasis id="B966EA8DFFB7FFA1232DD757FB3DFA03" box="[1148,1167,1582,1605]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA123F1D756FAB7FA00" box="[1184,1285,1583,1606]" italics="true" pageId="6" pageNumber="5">tavaratra</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12247D757FA0EFA00" authorityName="Lourenco &amp; Wilme" authorityYear="2015" box="[1302,1468,1582,1606]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="voahangyae">
<emphasis id="B966EA8DFFB7FFA12247D757FA9BFA03" box="[1302,1321,1582,1605]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1226AD757FA0EFA00" box="[1339,1468,1582,1606]" italics="true" pageId="6" pageNumber="5">voahangyae</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12461D737FBB8FA20" authorityName="Lowe &amp; Kovařík" authorityYear="2019" baseAuthorityName="Lourenco &amp; Goodman" baseAuthorityYear="2003" box="[816,1034,1614,1638]" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="ankarana">
<emphasis id="B966EA8DFFB7FFA12461D737FBB8FA20" box="[816,1034,1614,1638]" italics="true" pageId="6" pageNumber="5">Teruelius ankarana</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12373D736FB5BFA20" authorityName="Lourenco" authorityYear="2003" box="[1058,1257,1614,1638]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="ankarafantsika">
<emphasis id="B966EA8DFFB7FFA12373D736FB83FA20" box="[1058,1073,1615,1638]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12318D737FB5BFA20" box="[1097,1257,1614,1638]" italics="true" pageId="6" pageNumber="5">ankarafantsika</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12250D736FA26FA20" authorityName="Lourenco, Wilme &amp; Waeber" authorityYear="2018" box="[1281,1428,1614,1638]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="bemaraha">
<emphasis id="B966EA8DFFB7FFA12250D736FAA2FA20" box="[1281,1296,1615,1638]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12279D737FA26FA20" box="[1320,1428,1614,1638]" italics="true" pageId="6" pageNumber="5">bemaraha</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA122FCD736FCCFFAC0" authorityName="Lourenco" authorityYear="2012" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="bicolor">
<emphasis id="B966EA8DFFB7FFA122FCD736FA0EFA20" box="[1453,1468,1615,1638]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12461D717FCCFFAC0" box="[816,893,1646,1670]" italics="true" pageId="6" pageNumber="5">bicolor</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA124DDD716FBBCFAC0" baseAuthorityName="Kraepelin" baseAuthorityYear="1900" box="[908,1038,1646,1670]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="bistriatus">
<emphasis id="B966EA8DFFB7FFA124DDD716FC29FAC0" box="[908,923,1647,1670]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA124F8D717FBBCFAC0" box="[937,1038,1646,1670]" italics="true" pageId="6" pageNumber="5">bistriatus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA1234CD716FB0AFAC0" authorityName="Lourenco &amp; Wilme" authorityYear="2016" box="[1053,1208,1646,1670]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="eliseanneae">
<emphasis id="B966EA8DFFB7FFA1234CD716FB9EFAC0" box="[1053,1068,1647,1670]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1236BD717FB0AFAC0" box="[1082,1208,1646,1670]" italics="true" pageId="6" pageNumber="5">eliseanneae</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12397D716FAB4FAC0" authorityName="Lowe &amp; Kovařík" authorityYear="2019" baseAuthorityName="Lourenco" baseAuthorityYear="1996" box="[1222,1286,1646,1670]" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="feti">
<emphasis id="B966EA8DFFB7FFA12397D716FB67FAC0" box="[1222,1237,1647,1670]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA123B5D717FAB4FAC0" box="[1252,1286,1646,1670]" italics="true" pageId="6" pageNumber="5">feti</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12245D716FA2CFAC0" authorityName="Lourenco, Wilme &amp; Waeber" authorityYear="2016" box="[1300,1438,1646,1670]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="ganzhorni">
<emphasis id="B966EA8DFFB7FFA12245D716FA91FAC0" box="[1300,1315,1647,1670]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12263D717FA2CFAC0" box="[1330,1438,1646,1670]" italics="true" pageId="6" pageNumber="5">ganzhorni</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA122FCD716FC17FAE0" authorityName="Lourenco" authorityYear="1999" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="intertidalis">
<emphasis id="B966EA8DFFB7FFA122FCD716FA0EFAC0" box="[1453,1468,1647,1670]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12461D7F7FC17FAE0" box="[816,933,1678,1702]" italics="true" pageId="6" pageNumber="5">intertidalis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA124E6D7F6FB86FAE0" authorityName=", Pocock" authorityYear="1889" box="[951,1076,1678,1702]" class="Arachnida" family="Buthidae" genus="Buthus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="limbatus">
<emphasis id="B966EA8DFFB7FFA124E6D7F6FC74FAE0" box="[951,966,1679,1702]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12489D7F7FB86FAE0" box="[984,1076,1678,1702]" italics="true" pageId="6" pageNumber="5">limbatus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12314D7F6FB67FAE0" authorityName="Lourenco" authorityYear="2014" box="[1093,1237,1678,1702]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="magalieae">
<emphasis id="B966EA8DFFB7FFA12314D7F6FBE6FAE0" box="[1093,1108,1679,1702]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12337D7F7FB67FAE0" box="[1126,1237,1678,1702]" italics="true" pageId="6" pageNumber="5">magalieae</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA123B7D7F6FA29FAE0" box="[1254,1435,1678,1702]" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="mahafaliensis">
<emphasis id="B966EA8DFFB7FFA123B7D7F6FB47FAE0" box="[1254,1269,1679,1702]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12256D7F7FA29FAE0" box="[1287,1435,1678,1702]" italics="true" pageId="6" pageNumber="5">mahafaliensis</emphasis>
</taxonomicName>
, 
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<emphasis id="B966EA8DFFB7FFA122FCD7F6FA0EFAE0" box="[1453,1468,1679,1702]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12461D7D7FCD9FA80" box="[816,875,1710,1734]" italics="true" pageId="6" pageNumber="5">olgae</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12426D7D6FC42FA80" authorityName="Lourenco &amp; Wilme" authorityYear="2016" box="[887,1008,1710,1734]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="sabineae">
<emphasis id="B966EA8DFFB7FFA12426D7D6FC34FA80" box="[887,902,1711,1734]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA124C0D7D7FC42FA80" box="[913,1008,1710,1734]" italics="true" pageId="6" pageNumber="5">sabineae</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA124AAD7D6FBD9FA80" authorityName="Lourenco &amp; Wilme" authorityYear="2016" box="[1019,1131,1710,1734]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="waeberi">
<emphasis id="B966EA8DFFB7FFA124AAD7D6FBB8FA80" box="[1019,1034,1711,1734]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12344D7D7FBD9FA80" box="[1045,1131,1710,1734]" italics="true" pageId="6" pageNumber="5">waeberi</emphasis>
</taxonomicName>
(
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). We confirm here that 5 pairs are indeed present in all species that we have examined: 
<taxonomicName id="4C124D1CFFB7FFA1235FD637FBCFFB20" authorityName="Lourenco" authorityYear="2001" box="[1038,1149,1870,1894]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="garciai">
<emphasis id="B966EA8DFFB7FFA1235FD637FB93FB23" box="[1038,1057,1870,1893]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12360D636FBCFFB20" box="[1073,1149,1871,1894]" italics="true" pageId="6" pageNumber="5">garciai</emphasis>
</taxonomicName>
(= 
<taxonomicName id="4C124D1CFFB7FFA123F9D637FAB8FB20" baseAuthorityName="Lourenco" baseAuthorityYear="2000" box="[1192,1290,1870,1894]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="hirtus">
<emphasis id="B966EA8DFFB7FFA123F9D637FB09FB23" box="[1192,1211,1870,1893]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1239DD637FAB8FB20" box="[1228,1290,1870,1894]" italics="true" pageId="6" pageNumber="5">hirtus</emphasis>
</taxonomicName>
), 
<taxonomicName id="4C124D1CFFB7FFA12272D637FA25FB20" authorityName="Lourenco &amp; Goodman" authorityYear="2006" box="[1315,1431,1870,1894]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="goudoti">
<emphasis id="B966EA8DFFB7FFA12272D637FA84FB23" box="[1315,1334,1870,1893]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12217D637FA25FB20" box="[1350,1431,1870,1894]" italics="true" pageId="6" pageNumber="5">goudoti</emphasis>
</taxonomicName>
, 
<emphasis id="B966EA8DFFB7FFA122F9D637FA09FB23" box="[1448,1467,1870,1893]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. ‘ 
<taxonomicName id="4C124D1CFFB7FFA12468D617FC3DFBC0" authorityName="Lourenco, Wilme, Soarimalala &amp; Waeber, Previous" authorityYear="2017" box="[825,911,1902,1926]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="halleuxi">
<emphasis id="B966EA8DFFB7FFA12468D617FC3DFBC0" box="[825,911,1902,1926]" italics="true" pageId="6" pageNumber="5">halleuxi</emphasis>
</taxonomicName>
’, 
<taxonomicName id="4C124D1CFFB7FFA124F4D617FBB1FBC0" baseAuthorityName="Lourenco" baseAuthorityYear="2000" box="[933,1027,1902,1926]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="hirtus">
<emphasis id="B966EA8DFFB7FFA124F4D617FC0AFBC3" box="[933,952,1902,1925]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12494D617FBB1FBC0" box="[965,1027,1902,1926]" italics="true" pageId="6" pageNumber="5">hirtus</emphasis>
</taxonomicName>
, 
<emphasis id="B966EA8DFFB7FFA12341D617FB91FBC3" box="[1040,1059,1902,1925]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. sp. nr 
<taxonomicName id="4C124D1CFFB7FFA12321D617FB1CFBC0" baseAuthorityName="Lourenco" baseAuthorityYear="2000" box="[1136,1198,1902,1926]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="hirtus">
<emphasis id="B966EA8DFFB7FFA12321D617FB1CFBC0" box="[1136,1198,1902,1926]" italics="true" pageId="6" pageNumber="5">hirtus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA123EAD617FA2EFBC0" authorityName=", Gervais" authorityYear="1843" box="[1211,1436,1902,1926]" class="Arachnida" family="Scorpionidae" genus="Scorpio" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="madagascariensis">
<emphasis id="B966EA8DFFB7FFA123EAD617FB7CFBC3" box="[1211,1230,1902,1925]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1238AD617FA2EFBC0" box="[1243,1436,1902,1926]" italics="true" pageId="6" pageNumber="5">madagascariensis</emphasis>
</taxonomicName>
, 
<emphasis id="B966EA8DFFB7FFA122F9D617FA09FBC3" box="[1448,1467,1902,1925]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. ‘ 
<taxonomicName id="4C124D1CFFB7FFA12468D6F7FC29FBE0" authorityName="Lourenco" authorityYear="2005" box="[825,923,1934,1958]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="mandena">
<emphasis id="B966EA8DFFB7FFA12468D6F7FC29FBE0" box="[825,923,1934,1958]" italics="true" pageId="6" pageNumber="5">mandena</emphasis>
</taxonomicName>
’, 
<taxonomicName id="4C124D1CFFB7FFA124E1D6F7FBE2FBE0" authorityName="Lourenco &amp; Wilme" authorityYear="2015" box="[944,1104,1934,1958]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="voahangyae">
<emphasis id="B966EA8DFFB7FFA124E1D6F7FC71FBE3" box="[944,963,1934,1957]" italics="true" pageId="6" pageNumber="5">G</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1249ED6F7FBE2FBE0" box="[975,1104,1934,1958]" italics="true" pageId="6" pageNumber="5">voahangyae</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA1230DD6F7FAE1FBE0" baseAuthorityName="Vachon" baseAuthorityYear="1969" box="[1116,1363,1934,1958]" class="Arachnida" family="Buthidae" genus="Neogrosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="griveaudi">
<emphasis id="B966EA8DFFB7FFA1230DD6F7FAE1FBE0" box="[1116,1363,1934,1958]" italics="true" pageId="6" pageNumber="5">Neogrosphus griveaudi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12231D6F7FC62FB80" authorityName="Lowe &amp; Kovařík" authorityYear="2019" baseAuthorityName="Lourenco" baseAuthorityYear="2003" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="ankarafantsika">
<emphasis id="B966EA8DFFB7FFA12231D6F7FC62FB80" italics="true" pageId="6" pageNumber="5">Teruelius ankarafantsika</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA124B2D6D6FBDFFB80" authorityName="Lowe &amp; Kovařík" authorityYear="2019" baseAuthorityName="Lourenco &amp; Goodman" baseAuthorityYear="2003" box="[995,1133,1966,1990]" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="ankarana">
<emphasis id="B966EA8DFFB7FFA124B2D6D6FC40FB80" box="[995,1010,1967,1990]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12355D6D7FBDFFB80" box="[1028,1133,1966,1990]" italics="true" pageId="6" pageNumber="5">ankarana</emphasis>
</taxonomicName>
<emphasis id="B966EA8DFFB7FFA1233CD6D7FAB6FB80" box="[1133,1284,1966,1990]" italics="true" pageId="6" pageNumber="5">
, 
<taxonomicName id="4C124D1CFFB7FFA1232DD6D6FAB6FB80" authorityName="Fage" authorityYear="1929" box="[1148,1284,1966,1990]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="annulatus">T. annulatus</taxonomicName>
</emphasis>
, 
<taxonomicName id="4C124D1CFFB7FFA12247D6D6FA29FB80" baseAuthorityName="Kraepelin" baseAuthorityYear="1900" box="[1302,1435,1966,1990]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="bistriatus">
<emphasis id="B966EA8DFFB7FFA12247D6D6FA29FB80" box="[1302,1435,1966,1990]" italics="true" pageId="6" pageNumber="5">T. bistriatus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA122FCD6D6FCE0FBA0" authorityName="Lowe &amp; Kovařík" authorityYear="2019" baseAuthorityName="Lourenco" baseAuthorityYear="1996" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="feti">
<emphasis id="B966EA8DFFB7FFA122FCD6D6FA0EFB80" box="[1453,1468,1967,1990]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12461D6B7FCE0FBA0" box="[816,850,1998,2022]" italics="true" pageId="6" pageNumber="5">feti</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12431D6B6FC41FBA0" baseAuthorityName="Kraepelin" baseAuthorityYear="1900" box="[864,1011,1998,2022]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="flavopiceus">
<emphasis id="B966EA8DFFB7FFA12431D6B6FC41FBA0" box="[864,1011,1998,2022]" italics="true" pageId="6" pageNumber="5">T. flavopiceus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12350D6B6FB27FBA0" authorityName="Kraepelin" authorityYear="1900" box="[1025,1173,1998,2022]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="grandidieri">
<emphasis id="B966EA8DFFB7FFA12350D6B6FB27FBA0" box="[1025,1173,1998,2022]" italics="true" pageId="6" pageNumber="5">T. grandidieri</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA123F2D6B6FA87FBA0" authorityName="Lourenco" authorityYear="1999" box="[1187,1333,1998,2022]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="intertidalis">
<emphasis id="B966EA8DFFB7FFA123F2D6B6FB00FBA0" box="[1187,1202,1999,2022]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA12391D6B7FA87FBA0" box="[1216,1333,1998,2022]" italics="true" pageId="6" pageNumber="5">intertidalis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB7FFA12212D6B6FA09FBA0" authorityName=", Pocock" authorityYear="1889" box="[1347,1467,1998,2022]" class="Arachnida" family="Buthidae" genus="Buthus" kingdom="Animalia" order="Scorpiones" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="limbatus">
<emphasis id="B966EA8DFFB7FFA12212D6B6FAE0FBA0" box="[1347,1362,1999,2022]" italics="true" pageId="6" pageNumber="5">T</emphasis>
. 
<emphasis id="B966EA8DFFB7FFA1220ED6B7FA09FBA0" box="[1375,1467,1998,2022]" italics="true" pageId="6" pageNumber="5">limbatus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C124D1CFFB9FFAF2720D192FE93FD45" box="[113,289,235,259]" class="Arachnida" family="Buthidae" genus="Teruelius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="species" species="mahafaliensis">
<emphasis id="B966EA8DFFB9FFAF2720D192FF32FD44" box="[113,128,235,258]" italics="true" pageId="8" pageNumber="7">T</emphasis>
. 
<emphasis id="B966EA8DFFB9FFAF27DCD192FE93FD45" box="[141,289,235,259]" italics="true" pageId="8" pageNumber="7">mahafaliensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C124D1CFFB9FFAF2605D192FE19FD45" authorityName="Lourenco" authorityYear="2004" box="[340,427,235,259]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="species" species="olgae">
<emphasis id="B966EA8DFFB9FFAF2605D192FED1FD44" box="[340,355,235,258]" italics="true" pageId="8" pageNumber="7">T</emphasis>
. 
<emphasis id="B966EA8DFFB9FFAF2621D192FE19FD45" box="[368,427,235,259]" italics="true" pageId="8" pageNumber="7">olgae</emphasis>
</taxonomicName>
(e.g., 
<figureCitation id="13292A1AFFB9FFAF26BFD192FD39FD45" box="[494,651,235,259]" captionStart="Figures 227–230" captionStartId="67.[113,190,1951,1973]" captionTargetBox="[113,1474,234,1902]" captionTargetId="figure-2@67.[113,1474,234,1902]" captionTargetPageId="67" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7143712" httpUri="https://zenodo.org/record/7143712/files/figure.png" pageId="8" pageNumber="7">Figs. 227–230</figureCitation>
). We found only a few individual deviations from the standard pattern, such as 2 large and 2 small ocelli, that we regarded as developmental anomalies. We predict that other ‘ 
<taxonomicName id="4C124D1CFFB9FFAF2683D032FD8BFD25" authorityName="Simon" authorityYear="1880" box="[466,569,331,355]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="B966EA8DFFB9FFAF2683D032FD8BFD25" box="[466,569,331,355]" italics="true" pageId="8" pageNumber="7">Grosphus</emphasis>
</taxonomicName>
’ group species will also comply with the 5-eye pattern. Although undercounting of lateral eyes is perhaps attributable to overlooking of the smaller posterior and upper ocelli, 10 of the published 3-eye counts post-date introduction of the 5-eye model by 
<bibRefCitation id="EF834B6EFFB9FFAF25E1D0B2FF67FE45" author="Yang" etAl="et al." firstAuthor="Yang" pageId="8" pageNumber="7" pagination="55125" refId="ref44730" refString="YANG, X., Y. NORMA-RASHID, W. R. LOURENCO &amp; M. ZHU. 2013. True lateral eye numbers for extant buthids: a new discovery on an old character. PLoS ONE, 8 (1): e 55125. doi: 10.1371 / journal. pone. 0055125" type="journal article" year="2013">Yang et al. (2013</bibRefCitation>
, coauthor Lourenço) and 
<bibRefCitation id="EF834B6EFFB9FFAF26A0D092FCB2FE45" author="LORIA, S. F. &amp; L. PRENDINI" box="[497,768,491,515]" journalOrPublisher="PLoS ONE" pageId="8" pageNumber="7" pagination="112913" part="9" refId="ref37170" refString="LORIA, S. F. &amp; L. PRENDINI. 2014. Homology of the lateral eyes of Scorpiones: a six-ocellus model. PLoS ONE 9 (12): e 112913. doi: 10.1371 / journal. pone. 0112913." title="Homology of the lateral eyes of Scorpiones: a six-ocellus model" type="journal article" year="2014">Loria &amp; Prendini (2014)</bibRefCitation>
. Paradoxically, 
<bibRefCitation id="EF834B6EFFB9FFAF2640D372FDBDFE65" author="LOURENCO, W. R. &amp; J. - X. QI &amp; S. M. GOODMAN" box="[273,527,523,547]" journalOrPublisher="Boletin de la Sociedad Entomologica Aragonesa" pageId="8" pageNumber="7" pagination="171 - 177" part="40" refId="ref39785" refString="LOURENCO, W. R., J. - X. QI &amp; S. M. GOODMAN. 2007 a. Scorpions of south-western Madagascar. A new species of Grosphus Simon, 1880 (Scorpiones, Buthidae). Boletin de la Sociedad Entomologica Aragonesa, 40: 171 - 177." title="Scorpions of south-western Madagascar. A new species of Grosphus Simon, 1880 (Scorpiones, Buthidae)" type="journal article" year="2007">Lourenço et al. (2007a)</bibRefCitation>
claimed 3 lateral eyes in boilerplate descriptions of 
<taxonomicName id="4C124D1CFFB9FFAF26EFD352FD93FE05" baseAuthorityName="Lourenco" baseAuthorityYear="2000" box="[446,545,555,579]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="species" species="hirtus">
<emphasis id="B966EA8DFFB9FFAF26EFD352FE63FE04" box="[446,465,555,578]" italics="true" pageId="8" pageNumber="7">G</emphasis>
. 
<emphasis id="B966EA8DFFB9FFAF26B2D352FD93FE05" box="[483,545,555,579]" italics="true" pageId="8" pageNumber="7">hirtus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C124D1CFFB9FFAF250ED352FD63FE05" authorityName="Lourenco, Qi &amp; Goodman" authorityYear="2007" box="[607,721,555,579]" class="Arachnida" family="Buthidae" genus="Grosphus" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="species" species="polskyi">
<emphasis id="B966EA8DFFB9FFAF250ED352FDC0FE04" box="[607,626,555,578]" italics="true" pageId="8" pageNumber="7">G</emphasis>
. 
<emphasis id="B966EA8DFFB9FFAF25D4D352FD63FE05" box="[645,721,555,579]" italics="true" pageId="8" pageNumber="7">polskyi</emphasis>
</taxonomicName>
, yet their figures clearly depict all 5 lateral eyes as being present in both species. 
<bibRefCitation id="EF834B6EFFB9FFAF2657D312FE14FEC5" author="Vachon" box="[262,422,619,643]" firstAuthor="Vachon" pageId="8" pageNumber="7" pagination="476 - 483" refId="ref44004" refString="VACHON, M. 1969. Grosphus griveaudi, nouvelle espece de Scorpion Buthidae malgache. Bulletin du Museum national d'Histoire naturelle, Paris, (2), 41: 476 - 483." type="journal article" year="1969">Vachon (1969)</bibRefCitation>
correctly reported 5 “nettement visibles” lateral eyes, 3 large and 2 small, in both sexes of 
<taxonomicName id="4C124D1CFFB9FFAF2720D3D2FEDEFE85" baseAuthorityName="Vachon" baseAuthorityYear="1969" box="[113,364,683,707]" class="Arachnida" family="Buthidae" genus="Neogrosphus" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="species" species="griveaudi">
<emphasis id="B966EA8DFFB9FFAF2720D3D2FEDEFE85" box="[113,364,683,707]" italics="true" pageId="8" pageNumber="7">Neogrosphus griveaudi</emphasis>
</taxonomicName>
. Although 3 pairs were described for 
<taxonomicName id="4C124D1CFFB9FFAF2720D3B2FF64FEA5" authorityName="Lourenco" authorityYear="1996" box="[113,214,715,739]" class="Arachnida" family="Buthidae" genus="Neogrosphus" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="species" species="blanci">
<emphasis id="B966EA8DFFB9FFAF2720D3B2FF31FEA4" box="[113,131,715,738]" italics="true" pageId="8" pageNumber="7">N</emphasis>
. 
<emphasis id="B966EA8DFFB9FFAF27C2D3B2FF64FEA5" box="[147,214,715,739]" italics="true" pageId="8" pageNumber="7">blanci</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C124D1CFFB9FFAF265ED3B2FE17FEA5" authorityName="Lourenco, Wilme &amp; Waeber" authorityYear="2015" box="[271,421,715,739]" class="Arachnida" family="Buthidae" genus="Neogrosphus" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="7" phylum="Arthropoda" rank="species" species="andrafiabe">
<emphasis id="B966EA8DFFB9FFAF265ED3B2FE93FEA4" box="[271,289,715,738]" italics="true" pageId="8" pageNumber="7">N</emphasis>
. 
<emphasis id="B966EA8DFFB9FFAF2660D3B2FE17FEA5" box="[305,421,715,739]" italics="true" pageId="8" pageNumber="7">andrafiabe</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF834B6EFFB9FFAF26E4D3B2FDC1FEA5" author="LOURENCO, W. R." bookContentInfo="102 pp." box="[437,627,715,739]" journalOrPublisher="Museum National d'Histoire Naturelle, Paris" pageId="8" pageNumber="7" refId="ref37632" refString="LOURENCO, W. R. 1996 b. Faune de Madagascar. 87. Scorpions (Chelicerata, Scorpiones). Museum National d'Histoire Naturelle, Paris. 102 pp." title="Faune de Madagascar. 87. Scorpions (Chelicerata, Scorpiones)" type="book" year="1996">Lourenço, 1996b</bibRefCitation>
; 
<bibRefCitation id="EF834B6EFFB9FFAF25D1D3B2FF60FF45" author="LOURENCO, W. R. &amp; A. BEIGEL" journalOrPublisher="Beitrage zur Araneologie" pageId="8" pageNumber="7" pagination="476 - 480" part="9" refId="ref39065" refString="LOURENCO, W. R. &amp; A. BEIGEL. 2015. A new genus and species of Palaeoburmesebuthinae Lourenco, 2015 (Scorpiones: Archaeobuthidae) from Cretaceous amber of Myanmar. Beitrage zur Araneologie, 9: 476 - 480." title="A new genus and species of Palaeoburmesebuthinae Lourenco, 2015 (Scorpiones: Archaeobuthidae) from Cretaceous amber of Myanmar" type="journal article" year="2015">Lourenço et al., 2015</bibRefCitation>
), we are skeptical that these counts are accurate.
</paragraph>
</subSubSection>
</treatment>
</document>