treatments-xml/data/03/86/35/03863547FFE7FF92FDF7F98A1052FB88.xml

<document ID-DOI="10.7717/peerj.2955" ID-GBIF-Dataset="fd10e47d-00c5-4129-97d3-306ee2afc719" ID-PMC="PMC5299998" ID-PubMed="28194313" ID-Zenodo-Dep="291225" checkinTime="1486703386375" checkinUser="plazi" docAuthor="Mark D. Scherz, Juan D. Daza, Jörn Köhler, Miguel Vences &amp; Frank Glaw" docDate="2017" docId="03863547FFE7FF92FDF7F98A1052FB88" docLanguage="en" docName="peerJ.2955.pdf" docOrigin="PeerJ 2955" docStyle="DocumentStyle{}" docTitle="Geckolepis megalepis Scherz, Daza, Köhler, Vences &amp; Glaw, 2017, sp. nov." docType="treatment" docVersion="8" lastPageNumber="31" masterDocId="FFBF4D3FFFE2FF8CFFF4FF9B1464FF93" masterDocTitle="Off the scale: a new species of fishscale gecko (Squamata: Gekkonidae: Geckolepis) with exceptionally large scales" masterLastPageNumber="35" masterPageNumber="1" pageNumber="6" updateTime="1668138776344" updateUser="ExternalLinkService">
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<mods:title>Off the scale: a new species of fishscale gecko (Squamata: Gekkonidae: Geckolepis) with exceptionally large scales</mods:title>
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<mods:namePart>Mark D. Scherz</mods:namePart>
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<mods:namePart>Juan D. Daza</mods:namePart>
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<mods:namePart>Jörn Köhler</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Miguel Vences</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Frank Glaw</mods:namePart>
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<taxonomicName box="[515,766,1553,1580]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[515,766,1553,1580]" italics="true" pageId="5" pageNumber="6">Geckolepis megalepis</emphasis>
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<taxonomicNameLabel box="[772,865,1553,1579]" pageId="5" pageNumber="6" rank="species">sp. nov.</taxonomicNameLabel>
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<taxonomicName box="[515,633,1589,1615]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[515,633,1589,1615]" italics="true" pageId="5" pageNumber="6">Geckolepis</emphasis>
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sp. OTU D—(
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[808,1021,1589,1616]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="5" pageNumber="6" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[808,1021,1589,1616]" italics="true" pageId="5" pageNumber="6">Lemme et al., 2013</emphasis>
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)
</paragraph>
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<paragraph blockId="5.[515,1416,1553,1688]" box="[515,1009,1625,1652]" pageId="5" pageNumber="6">
(
<figureCitation box="[526,629,1625,1652]" captionStart-0="Figure 3" captionStart-1="Figure 4" captionStart-2="Figure 5" captionStart-3="Figure 6" captionStart-4="Figure 7" captionStart-5="Figure 8" captionStart-6="Figure 9" captionStartId-0="6.[507,571,1215,1237]" captionStartId-1="7.[507,571,1752,1774]" captionStartId-2="8.[507,571,729,751]" captionStartId-3="13.[507,571,1713,1735]" captionStartId-4="15.[507,571,1713,1735]" captionStartId-5="16.[507,571,1352,1374]" captionStartId-6="17.[507,571,1453,1475]" captionTargetBox-0="[491,1541,229,1170]" captionTargetBox-1="[491,1512,230,1704]" captionTargetBox-2="[492,1541,229,684]" captionTargetBox-3="[491,1188,229,1668]" captionTargetBox-4="[491,1381,229,1668]" captionTargetBox-5="[491,1541,229,1307]" captionTargetBox-6="[492,1541,229,1408]" captionTargetId-0="figure@6.[491,1541,229,1170]" captionTargetId-1="figure@7.[491,1514,229,1708]" captionTargetId-2="figure@8.[491,1541,229,684]" captionTargetId-3="figure@13.[491,1188,229,1668]" captionTargetId-4="figure@15.[491,1381,229,1668]" captionTargetId-5="figure@16.[491,1541,229,1307]" captionTargetId-6="figure@17.[491,1541,229,1408]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionTargetPageId-2="8" captionTargetPageId-3="13" captionTargetPageId-4="15" captionTargetPageId-5="16" captionTargetPageId-6="17" captionText-0="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." captionText-1="Figure 4 Specimens of Geckolepis megalepis sp. nov. in preservative. (A – C) The holotype, ZSM 2126 / 2007; (D – F) paratype ZSM 232 / 2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale." captionText-2="Figure 5 Micro-CT images of mineraliZed scales of Geckolepis specimens. Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of Geckolepis megalepis (A – C, ZSM 2126 / 2007), Geckolepis OTU AB (D – E, ZSM 1520 / 2008), G. maculata (F – G, ZMB 9655), and G. humbloti (H – I, ZSM 80 / 2010; J – K, ZSM 81 / 2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3)." captionText-3="Figure 6 Micro-CT images of skull (cranium and jaw) of the holotype of Geckolepis megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal, (B) ventral, (C) lateral, (D) labial, and (E) lingual view. Abbreviations: adf, anterior inferior dental foramen; amf, anterior mylohyoid foramen; ar, articular facet; asnp, ascending nasal process of premaxilla; bo, basioccipital; bp, basipterygoid process of parabasisphenoid; ch, choana; cob, compound bone; cor, coronoid; d, dentary; dpp, (continued on next page ...) Figure 6 (... continued) descending parietal process; ect, ectopterygoid; ept, epipterygoid; f, frontal; hscc, horizontal semi-circular canal; if, incisive foramen; j, jugal; larst, lateral aperture of the recessus scalae tympani; mdf, mandibular fossa; mf, mental foramen; mko, opening of the Meckelian canal, mx, maxilla; mx. ap, anterior process of maxilla; mx. fp, facial process of maxilla; mx. pp, posterior process of maxilla; mx. ps, palatal shelf of maxilla; n, nasal; occ, occipital condyle; oto, otooccipital; pal, palatine; pal. mp, maxillary process of palatine; pal. vf, vomerine flange of palatine; par, parietal; pbsh, parabasisphenoid; pmx, premaxilla; pmx. pps, premaxilla palatal shelf; pof, postorbitofrontal; pop, paroccipital process; ppp, posterolateral process of parietal; prf, prefrontal; pro, prootic; pro. ca, crista alaris of prootic; pro. cp, crista prootica of prootic; psaf, posterior surangular foramen; pscc, posterior semicircular canal; pt, pterygoid; pt. qp, quadrate process of pterygoid; q, quadrate; q. ch, conch of quadrate; rap, retroarticular process; s, stapes; saf, surangular foramen; scr, sclerotic ring; smx, septomaxilla; spl, splenial; sof, suborbital fenestra; sq, squamosal, so, supraoccipital; spht, sphenooccipital tubercle; tbr, trabeculae; v, vomer; v. lp, lateral process of vomer; v. mp, maxillary process of vomer. A rotational video of the skull is provided in Video S 1." captionText-4="Figure 7 Micro-CT images of spinal column of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Lateral and (B) dorsal view. Abbreviations: lb, lumbar; sc, sacral. Regions are indicated by the posterodorsal-most point of the neural arches." captionText-5="Figure 8 Micro-CT images of pectoral girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Ventral and (B) lateral view. Abbreviations: 1 cf, primary coracoid fenestra; 2 cf, secondary coracoid fenestra; acr, acromion process; cf, clavicular fenestra; cl, clavicle; cor, coracoid; ic, interclavicle; gf, glenoid fossa; hu, humerus; hu. c, humeral condyle; hu. dtp, deltopectoral crest of humerus; hu. ec, ectepicondylar crest of humerus; hu. hc, humeral crest of humerus; mc, metacarpal; ms, mesosternum; p, phalanges; ps, presternum; pu, patella ulnaris; r, radius; rl, radiale; s, scapula; sc. 1 c, primary coracoid ray of scapulocoracoid; sc. 2 c, secondary coracoid ray of scapulocoracoid; scf, scapulocoracoid fenestra; sf, scapular fenestra; ss, suprascapula; sc. sr, scapular ray of scapulocoracoid; sr, sternal ribs; sucf, supracoracoid foramen; u, ulna; ul, ulnare; xs, xiphisternum." captionText-6="Figure 9 Micro-CT images of pelvic girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal and (B) lateral view. Abbreviations: ac, acetabulum; ag, astragalocalcaneum; ep, epipubis; f, femur; fb, fibula; f. it, internal trochanter; f. itf, intertrochanteric fossa; f. fc, femoral condyle; hyi, hypoischium; il, ilium; il. pap, preacetabular process of ilium; is, ischium; is. t, ischiadic tuberosity; ltp, lenticular tibial patella; mt, metatarsals; of, obturator foramen; p, phalanges; pic, proischiadic cartilage; pb, pubis; pb. pl, processus lateralis of pubis; pc, post-cloacal bone; pt, pubic tubercle; tb, tibia; tf, thyroid fenestra." httpUri-0="https://zenodo.org/record/291381/files/figure.png" httpUri-1="https://zenodo.org/record/291229/files/figure.png" httpUri-2="https://zenodo.org/record/291230/files/figure.png" httpUri-3="https://zenodo.org/record/291231/files/figure.png" httpUri-4="https://zenodo.org/record/291232/files/figure.png" httpUri-5="https://zenodo.org/record/291233/files/figure.png" httpUri-6="https://zenodo.org/record/291234/files/figure.png" pageId="5" pageNumber="6">Figs. 3–9</figureCitation>
, 
<tableCitation box="[642,728,1625,1652]" captionStart="Table 1" captionStartId="9.[106,162,246,267]" captionTargetBox="[113,1515,327,1147]" captionText="Table 1 Comparative morphological data on Geckolepis megalepis, and the holotype of G. maculata (ZMB 9655), with values for G. polylepis and G. typica given by Köhler et al. (2009). For raw measurements of individual G. megalepis specimens, see Appendix S 1." httpUri="http://table.plazi.org/id/DF50D4D9FFEBFF85FF9EFF6D109BFEBA" pageId="5" pageNumber="6" tableUuid="DF50D4D9FFEBFF85FF9EFF6D109BFEBA">Table 1</tableCitation>
, Appendices S1 and S2)
</paragraph>
</subSubSection>
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<subSubSection box="[515,1416,1661,1688]" pageId="5" pageNumber="6" type="nomenclature">LSID: urn:lsid:zoobank.org:act:08097CA5-172E-4F45-AE68-AC4E02BBC644</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6" type="materials_examined">
<materialsCitation ID-GBIF-Occurrence="1427603770" collectingDate="2007-03-03" collectionCode="ZSM, FGZC" collectorName="P Bora &amp; H Enting &amp; F Glaw &amp; A Knoll &amp; J Köhler" country="Madagascar" elevation="150" latitude="-12.9564" location="east side of Ankarana National Park" longLatPrecision="7" longitude="49.1172" pageId="5" pageNumber="6" specimenCode="ZSM 2126, FGZC 1144" specimenCount="1" specimenCount-adult="1" stateProvince="Antsiranana Province" typeStatus="holotype">
<typeStatus box="[523,625,1768,1794]" pageId="5" pageNumber="6">
<emphasis box="[523,625,1768,1794]" italics="true" pageId="5" pageNumber="6">Holotype</emphasis>
</typeStatus>
. 
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/2007 (
<specimenCode box="[842,987,1768,1794]" pageId="5" pageNumber="6">FGZC 1144</specimenCode>
), an 
<specimenCount box="[1044,1104,1768,1794]" pageId="5" pageNumber="6" type="adult">adult</specimenCount>
of unknown sex, from the 
<location LSID="urn:lsid:plazi:treatment:03863547FFE7FF92FDF7F98A1052FB88:8EF0D28AFFE7FF89FA67F9721742F8B9" country="Madagascar" latitude="-12.9564" longLatPrecision="7" longitude="49.1172" name="east side of Ankarana National Park" pageId="5" pageNumber="6" stateProvince="Antsiranana Province">east side of Ankarana National Park</location>
(
<geoCoordinate box="[820,942,1803,1834]" direction="south" orientation="latitude" pageId="5" pageNumber="6" precision="5" value="-12.9564">12.9564°S</geoCoordinate>
, 
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, ca. 
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<elevation box="[1126,1199,1807,1834]" metricMagnitude="2" metricUnit="m" metricValue="1.5" pageId="5" pageNumber="6" unit="m" value="150.0">150 m</elevation>
</quantity>
a.s.l.), 
<collectingRegion box="[1282,1537,1808,1834]" country="Madagascar" name="Antsiranana" pageId="5" pageNumber="6">Antsiranana Province</collectingRegion>
, north 
<collectingCountry box="[566,707,1847,1873]" name="Madagascar" pageId="5" pageNumber="6">Madagascar</collectingCountry>
, collected on 
<collectingDate box="[871,1035,1847,1873]" pageId="5" pageNumber="6" value="2007-03-03">
<date box="[871,1035,1847,1873]" pageId="5" pageNumber="6" value="2007-03-03">3 March 2007</date>
</collectingDate>
by 
<collectorName box="[1079,1162,1848,1874]" pageId="5" pageNumber="6">P Bora</collectorName>
, 
<collectorName box="[1175,1287,1847,1874]" pageId="5" pageNumber="6">H Enting</collectorName>
, 
<collectorName box="[1299,1386,1847,1874]" pageId="5" pageNumber="6">F Glaw</collectorName>
, 
<collectorName box="[1398,1491,1847,1873]" pageId="5" pageNumber="6">A Knoll</collectorName>
&amp; 
<collectorName box="[491,591,1887,1913]" pageId="5" pageNumber="6">J Köhler</collectorName>
.
</materialsCitation>
<caption httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="6" pageNumber="7" targetBox="[491,1541,229,1170]" targetPageId="6">
<paragraph blockId="6.[507,1517,1215,1354]" pageId="6" pageNumber="7">
<emphasis bold="true" box="[507,1096,1215,1237]" pageId="6" pageNumber="7">
Figure 3 Specimens of 
<taxonomicName box="[742,948,1215,1237]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[742,948,1215,1237]" italics="true" pageId="6" pageNumber="7">Geckolepis megalepis</emphasis>
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<taxonomicNameLabel box="[953,1029,1215,1236]" pageId="6" pageNumber="7" rank="species">sp. nov.</taxonomicNameLabel>
in life.
</emphasis>
(A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to 
<taxonomicName box="[1099,1216,1303,1324]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1099,1216,1303,1324]" italics="true" pageId="6" pageNumber="7">G. megalepis</emphasis>
</taxonomicName>
is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated.
</paragraph>
</caption>
<paragraph blockId="6.[491,1542,1408,1913]" pageId="6" pageNumber="7">
<emphasis box="[523,633,1409,1434]" italics="true" pageId="6" pageNumber="7">Paratypes</emphasis>
. ZSM 289/2004 (FGZC 554), probably a subadult, sex unknown, from between Mahamasina and the Petit Tsingy (exact coordinates not known, but ca. 12.9558°S, 49.1181°E, ca. 125 m a.s.l.), Ankarana National Park, Antsiranana Province, north Madagascar, collected 25 February 2004 by F Glaw, M Puente &amp; R Randrianiaina; ZSM 232/2016 (FGZC 5476), an adult of unknown sex, from the private forest of the Ankarana Lodge (12.9613°S, 49.1499°E, 134 m a.s.l.), Ankarana massif, Antsiranana Province, north Madagascar, collected 28 August 2016 by F Glaw, K Glaw, T Glaw, Jaques &amp; NA Raharinoro; FGZC 1606 (UADBA uncatalogued), sex and age unknown, from Petit Tsingy (ca. 12.9558°S, 49.1181°E, ca. 125 m a.s.l.), Ankarana National Park, Antsiranana Province, north Madagascar, collected 12 February 2008 by N D’Cruze, M Franzen, F Glaw &amp; J Köhler.
</paragraph>
</subSubSection>
</paragraph>
<subSubSection lastPageId="9" lastPageNumber="10" pageId="6" pageNumber="7" type="diagnosis">
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<emphasis box="[523,632,1808,1833]" italics="true" pageId="6" pageNumber="7">Diagnosis</emphasis>
. A species of the genus 
<taxonomicName box="[912,1029,1807,1833]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[912,1029,1807,1833]" italics="true" pageId="6" pageNumber="7">Geckolepis</emphasis>
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based on its overall morphology and large, fish-like scales (similar to cycloid scales in terms of the extent of overlap), as well as its phylogenetic position (
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[763,976,1886,1913]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="6" pageNumber="7" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[763,976,1886,1913]" italics="true" pageId="6" pageNumber="7">Lemme et al., 2013</emphasis>
</bibRefCitation>
; 
<figureCitation box="[990,1055,1886,1913]" captionStart="Figure 1" captionStartId="2.[507,571,950,972]" captionTargetBox="[1198,1540,229,885]" captionTargetPageId="2" captionText="Figure 1 Molecular phylogeny and distribution of Geckolepis OTUs based mainly on Lemme et al. (2013). Dotted lines in the phylogeny indicate uncertain placement. The phylogenetic position and distribution of Geckolepis humbloti is inferred from Hawlitschek et al. (2016)." httpUri="https://zenodo.org/record/291226/files/figure.svg" pageId="6" pageNumber="7">Fig. 1</figureCitation>
). 
<taxonomicName box="[1081,1314,1886,1912]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1081,1314,1886,1912]" italics="true" pageId="6" pageNumber="7">Geckolepis megalepis</emphasis>
</taxonomicName>
differs from all of its congeners by the possession of the following suite of characters: innermost pair of postmental scales in broad contact (condition A/B, 
<figureCitation box="[1099,1164,962,988]" captionStart="Figure 4" captionStartId="7.[507,571,1752,1774]" captionTargetBox="[491,1512,230,1704]" captionTargetId="figure@7.[491,1514,229,1708]" captionTargetPageId="7" captionText="Figure 4 Specimens of Geckolepis megalepis sp. nov. in preservative. (A – C) The holotype, ZSM 2126 / 2007; (D – F) paratype ZSM 232 / 2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale." httpUri="https://zenodo.org/record/291229/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
), SVL ≤ 69.5 mm, infralabials to anterior margin of eye 4.5–4.8, 17–18 scales rows around the midbody, 27–31 ventral scales between the postmentals and the vent, and the absence of a dark lateral stripe, and typical midbody dorsal scales measuring 7.3–8.3% of the SVL in length. Osteologically, 
<taxonomicName class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="megalepis">
<emphasis italics="true" pageId="8" pageNumber="9">G. megalepis</emphasis>
</taxonomicName>
is characterised by a narrow infraorbital fenestra, a bulging nasal cavity, nasals with straight sides, a well developed anterior extension of the subfrontal process, a notched premaxilla-vomer fenestra, scapular ray of scapulocoracoid not surpassing the clavicle, and posteriorly curved pubic tubercle of the pubis. Additionally, it is separated by an uncorrected pairwise genetic distance in the mitochondrial ND4 gene of ≥10.1% from all other lineages of 
<taxonomicName box="[691,808,1321,1347]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[691,808,1321,1347]" italics="true" pageId="8" pageNumber="9">Geckolepis</emphasis>
</taxonomicName>
and has a unique CMOS haplotype (
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[1243,1456,1321,1347]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="8" pageNumber="9" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[1243,1456,1321,1347]" italics="true" pageId="8" pageNumber="9">Lemme et al., 2013</emphasis>
</bibRefCitation>
).
</paragraph>
<caption httpUri="https://zenodo.org/record/291229/files/figure.png" pageId="7" pageNumber="8" targetBox="[491,1512,230,1704]" targetPageId="7">
<paragraph blockId="7.[507,1498,1752,1862]" pageId="7" pageNumber="8">
<emphasis bold="true" box="[507,1186,1752,1774]" pageId="7" pageNumber="8">
Figure 4 Specimens of 
<taxonomicName box="[742,948,1752,1774]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[742,948,1752,1774]" italics="true" pageId="7" pageNumber="8">Geckolepis megalepis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[953,1029,1753,1774]" pageId="7" pageNumber="8" rank="species">sp. nov.</taxonomicNameLabel>
in preservative.
</emphasis>
(A–C) The holotype, ZSM 2126/2007; (D–F) paratype ZSM 232/2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale.
</paragraph>
</caption>
<caption httpUri="https://zenodo.org/record/291230/files/figure.png" pageId="8" pageNumber="9" targetBox="[492,1541,229,684]" targetPageId="8">
<paragraph blockId="8.[507,1502,729,868]" pageId="8" pageNumber="9">
<emphasis bold="true" box="[507,1237,729,751]" pageId="8" pageNumber="9">
Figure 5 Micro-CT images of mineraliZed scales of 
<taxonomicName box="[1019,1123,729,751]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[1019,1123,729,751]" italics="true" pageId="8" pageNumber="9">Geckolepis</emphasis>
</taxonomicName>
specimens.
</emphasis>
Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of 
<taxonomicName box="[1268,1460,759,780]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1268,1460,759,780]" italics="true" pageId="8" pageNumber="9">Geckolepis megalepis</emphasis>
</taxonomicName>
(A– C, ZSM 2126/2007), 
<taxonomicName box="[706,802,788,809]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[706,802,788,809]" italics="true" pageId="8" pageNumber="9">Geckolepis</emphasis>
</taxonomicName>
OTU AB (D–E, ZSM 1520/2008), 
<taxonomicName box="[1133,1248,788,809]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1133,1248,788,809]" italics="true" pageId="8" pageNumber="9">G. maculata</emphasis>
</taxonomicName>
(F–G, ZMB 9655), and 
<taxonomicName class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="humbloti">
<emphasis italics="true" pageId="8" pageNumber="9">G. humbloti</emphasis>
</taxonomicName>
(H–I, ZSM 80/2010; J–K, ZSM 81/2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3).
</paragraph>
</caption>
<paragraph blockId="8.[491,1542,922,1866]" box="[523,1496,1361,1387]" pageId="8" pageNumber="9">
<taxonomicName box="[523,756,1361,1387]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[523,756,1361,1387]" italics="true" pageId="8" pageNumber="9">Geckolepis megalepis</emphasis>
</taxonomicName>
may be distinguished from 
<taxonomicName box="[1097,1236,1361,1387]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1097,1236,1361,1387]" italics="true" pageId="8" pageNumber="9">G. maculata</emphasis>
</taxonomicName>
(note: because of the
</paragraph>
<paragraph blockId="8.[491,1542,922,1866]" pageId="8" pageNumber="9">
substantial uncertainty surrounding the identity of 
<emphasis box="[1105,1251,1401,1427]" italics="true" pageId="8" pageNumber="9">
<taxonomicName box="[1105,1246,1401,1427]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="maculata">G. maculata</taxonomicName>
,
</emphasis>
we here compare 
<emphasis box="[1473,1500,1401,1427]" italics="true" pageId="8" pageNumber="9">G.</emphasis>
<taxonomicName box="[491,600,1440,1466]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[491,600,1440,1466]" italics="true" pageId="8" pageNumber="9">megalepis</emphasis>
</taxonomicName>
only to the holotype of that species, ZMB 9655, until such a time as its true affinities can be clarified; see 
<bibRefCitation author="Kohler G &amp; Diethert H-H &amp; Nussbaum RA &amp; Raxworthy CJ." box="[836,1055,1480,1507]" journalOrPublisher="Herpetologica" pageId="8" pageNumber="9" pagination="419 - 435" part="65" publicationUrl="10.1655/08-059.1." refString="Kohler G, Diethert H-H, Nussbaum RA, Raxworthy CJ. 2009. A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros. Herpetologica 65: 419 - 435 DOI 10.1655 / 08 - 059.1." title="A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros" type="journal article" year="2009">
<emphasis box="[836,1055,1480,1507]" italics="true" pageId="8" pageNumber="9">Köhler et al. (2009)</emphasis>
</bibRefCitation>
for a detailed morphological account of that specimen) by the combination of fewer scale rows around midbody (17–18 vs. 25), fewer ventral scales (27–31 vs. 32), larger relative scale size (typical midbody dorsal scale 7.3–8.3% of SVL vs. 5.4%), and the absence of a dark lateral head stripe (vs. presence); from 
<taxonomicName box="[556,657,1640,1666]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="typica">
<emphasis box="[556,657,1640,1666]" italics="true" pageId="8" pageNumber="9">G. typica</emphasis>
</taxonomicName>
by larger maximum size (SVL up to 69.5 mm vs. &lt;57 mm), fewer scale rows around midbody (17–18 vs. 28–32), fewer ventral scales (27–31 vs. 42–49), postmental scale condition (A/B vs. D), and the absence of dark longitudinal stripes on the dorsum (vs. presence); from 
<taxonomicName box="[685,815,1759,1786]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="polylepis">
<emphasis box="[685,815,1759,1786]" italics="true" pageId="8" pageNumber="9">G. polylepis</emphasis>
</taxonomicName>
by larger maximum size (SVL up to 69.5 mm vs. &lt;52 mm), fewer scale rows at midbody (17–18 vs. 30–37), fewer ventral scales (27–31 vs. 37–55), and the absence of dark longitudinal stripes on the dorsum (vs. presence); and from 
<emphasis box="[1495,1522,1839,1865]" italics="true" pageId="8" pageNumber="9">G.</emphasis>
</paragraph>
<caption ID-Table-UUID="DF50D4D9FFEBFF85FF9EFF6D109BFEBA" httpUri="http://table.plazi.org/id/DF50D4D9FFEBFF85FF9EFF6D109BFEBA" pageId="9" pageNumber="10" targetBox="[113,1515,327,1147]" targetIsTable="true" targetPageId="9">
<paragraph blockId="9.[106,1493,245,297]" pageId="9" pageNumber="10">
<emphasis bold="true" pageId="9" pageNumber="10">
Table 1 Comparative morphological data on 
<taxonomicName box="[561,767,245,267]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="megalepis">
<emphasis bold="true" box="[561,767,245,267]" italics="true" pageId="9" pageNumber="10">Geckolepis megalepis</emphasis>
</taxonomicName>
, and the holotype of 
<taxonomicName box="[974,1094,245,267]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="maculata">
<emphasis bold="true" box="[974,1094,245,267]" italics="true" pageId="9" pageNumber="10">G. maculata</emphasis>
</taxonomicName>
(ZMB 9655), with values for 
<taxonomicName box="[1379,1493,245,267]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="polylepis">
<emphasis bold="true" box="[1379,1493,245,267]" italics="true" pageId="9" pageNumber="10">G. polylepis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[149,236,275,296]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="typica">
<emphasis bold="true" box="[149,236,275,296]" italics="true" pageId="9" pageNumber="10">G. typica</emphasis>
</taxonomicName>
given by 
<emphasis bold="true" box="[328,516,275,297]" italics="true" pageId="9" pageNumber="10">Köhler et al. (2009)</emphasis>
.
</emphasis>
For raw measurements of individual 
<taxonomicName box="[884,1001,275,296]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[884,1001,275,296]" italics="true" pageId="9" pageNumber="10">G. megalepis</emphasis>
</taxonomicName>
specimens, see Appendix S1.
</paragraph>
</caption>
<paragraph pageId="9" pageNumber="10">
<table box="[113,1515,327,1147]" gridcols="5" gridrows="22" pageId="9" pageNumber="10">
<tr box="[113,1515,327,349]" gridrow="0" pageId="9" pageNumber="10">
<th box="[113,256,327,349]" gridcol="0" gridrow="0" pageId="9" pageNumber="10">
<heading bold="true" box="[113,190,327,349]" fontSize="9" level="4" pageId="9" pageNumber="10" reason="6">
<emphasis bold="true" box="[113,190,327,349]" pageId="9" pageNumber="10">Species:</emphasis>
</heading>
</th>
<th box="[363,589,327,349]" gridcol="1" gridrow="0" pageId="9" pageNumber="10">
<heading bold="true" box="[363,568,327,349]" fontSize="9" level="4" pageId="9" pageNumber="10" reason="6">
<emphasis bold="true" box="[363,568,327,349]" pageId="9" pageNumber="10">
<taxonomicName box="[363,488,327,349]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[363,488,327,349]" italics="true" pageId="9" pageNumber="10">G. megalepis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[492,568,328,349]" pageId="9" pageNumber="10" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</th>
<th box="[696,850,327,349]" gridcol="2" gridrow="0" pageId="9" pageNumber="10">
<heading bold="true" box="[696,816,327,349]" fontSize="9" level="4" pageId="9" pageNumber="10" reason="6">
<taxonomicName box="[696,816,327,349]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="maculata">
<emphasis bold="true" box="[696,816,327,349]" italics="true" pageId="9" pageNumber="10">G. maculata</emphasis>
</taxonomicName>
</heading>
</th>
<th box="[957,1183,327,349]" gridcol="3" gridrow="0" pageId="9" pageNumber="10">
<heading bold="true" box="[957,1071,327,349]" fontSize="9" level="4" pageId="9" pageNumber="10" reason="6">
<taxonomicName box="[957,1071,327,349]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="polylepis">
<emphasis bold="true" box="[957,1071,327,349]" italics="true" pageId="9" pageNumber="10">G. polylepis</emphasis>
</taxonomicName>
</heading>
</th>
<th box="[1290,1515,327,349]" gridcol="4" gridrow="0" pageId="9" pageNumber="10">
<heading bold="true" box="[1290,1377,328,349]" fontSize="9" level="4" pageId="9" pageNumber="10" reason="6">
<taxonomicName box="[1290,1377,328,349]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="typica">
<emphasis bold="true" box="[1290,1377,328,349]" italics="true" pageId="9" pageNumber="10">G. typica</emphasis>
</taxonomicName>
</heading>
</th>
</tr>
<tr box="[113,1515,366,388]" gridrow="1" pageId="9" pageNumber="10">
<th box="[113,256,366,388]" gridcol="0" gridrow="1" pageId="9" pageNumber="10">No. Specimens</th>
<td box="[363,589,366,388]" gridcol="1" gridrow="1" pageId="9" pageNumber="10">3</td>
<td box="[696,850,366,388]" gridcol="2" gridrow="1" pageId="9" pageNumber="10">Only ZMB 9655</td>
<td box="[957,1183,366,388]" gridcol="3" gridrow="1" pageId="9" pageNumber="10">11</td>
<td box="[1290,1515,366,388]" gridcol="4" gridrow="1" pageId="9" pageNumber="10">102</td>
</tr>
<tr box="[113,1515,402,425]" gridrow="2" pageId="9" pageNumber="10">
<th box="[113,256,402,425]" gridcol="0" gridrow="2" pageId="9" pageNumber="10">SVL (mm)</th>
<td box="[363,589,402,425]" gridcol="1" gridrow="2" pageId="9" pageNumber="10">57.8–69.5 (65.2 ± 6.46)</td>
<td box="[696,850,402,425]" gridcol="2" gridrow="2" pageId="9" pageNumber="10">58.5</td>
<td box="[957,1183,402,425]" gridcol="3" gridrow="2" pageId="9" pageNumber="10">41.1–50.2 (46.5 ± 2.8)</td>
<td box="[1290,1515,402,425]" gridcol="4" gridrow="2" pageId="9" pageNumber="10">31.1–56.3 (43.0 ± 5.02)</td>
</tr>
<tr box="[113,1515,438,461]" gridrow="3" pageId="9" pageNumber="10">
<th box="[113,256,438,461]" gridcol="0" gridrow="3" pageId="9" pageNumber="10">TL (mm)</th>
<td box="[363,589,438,461]" gridcol="1" gridrow="3" pageId="9" pageNumber="10">71.7–80.1 (75.3 ± 4.33)</td>
<td box="[696,850,438,461]" gridcol="2" gridrow="3" pageId="9" pageNumber="10">N/A</td>
<td box="[957,1183,438,461]" gridcol="3" gridrow="3" pageId="9" pageNumber="10">34.5–56.5 (48.0 ± 6.4)</td>
<td box="[1290,1515,438,461]" gridcol="4" gridrow="3" pageId="9" pageNumber="10">29.7–58.5 (43.6 ± 6.83)</td>
</tr>
<tr box="[113,1515,475,497]" gridrow="4" pageId="9" pageNumber="10">
<th box="[113,256,475,497]" gridcol="0" gridrow="4" pageId="9" pageNumber="10">Ax–Gr (mm)</th>
<td box="[363,589,475,497]" gridcol="1" gridrow="4" pageId="9" pageNumber="10">23.9–32.5 (27.9 ± 4.34)</td>
<td box="[696,850,475,497]" gridcol="2" gridrow="4" pageId="9" pageNumber="10">23.7</td>
<td box="[957,1183,475,497]" gridcol="3" gridrow="4" pageId="9" pageNumber="10">14.7–24.3 (19.8 ± 2.7)</td>
<td box="[1290,1515,475,497]" gridcol="4" gridrow="4" pageId="9" pageNumber="10">12.4–28.3 (19.0 ± 2.76)</td>
</tr>
<tr box="[113,1515,511,533]" gridrow="5" pageId="9" pageNumber="10">
<th box="[113,256,511,533]" gridcol="0" gridrow="5" pageId="9" pageNumber="10">ShL (mm)</th>
<td box="[363,589,511,533]" gridcol="1" gridrow="5" pageId="9" pageNumber="10">5.0–7.0 (5.9 ± 1.01)</td>
<td box="[696,850,511,533]" gridcol="2" gridrow="5" pageId="9" pageNumber="10">6.2</td>
<td box="[957,1183,511,533]" gridcol="3" gridrow="5" pageId="9" pageNumber="10">5.5–7.2 (6.5 ± 0.5)</td>
<td box="[1290,1515,511,533]" gridcol="4" gridrow="5" pageId="9" pageNumber="10">4.0–8.2 (5.7 ± 0.72)</td>
</tr>
<tr box="[113,1515,547,569]" gridrow="6" pageId="9" pageNumber="10">
<th box="[113,256,547,569]" gridcol="0" gridrow="6" pageId="9" pageNumber="10">HL (mm)</th>
<td box="[363,589,547,569]" gridcol="1" gridrow="6" pageId="9" pageNumber="10">14.9–17.1 (16.0 ± 1.10)</td>
<td box="[696,850,547,569]" gridcol="2" gridrow="6" pageId="9" pageNumber="10">15.2</td>
<td box="[957,1183,547,569]" gridcol="3" gridrow="6" pageId="9" pageNumber="10">9.7–12.0 (10.7 ± 0.8)</td>
<td box="[1290,1515,547,569]" gridcol="4" gridrow="6" pageId="9" pageNumber="10">6.9–12.8 (9.6 ± 1.08)</td>
</tr>
<tr box="[113,1515,583,605]" gridrow="7" pageId="9" pageNumber="10">
<th box="[113,256,583,605]" gridcol="0" gridrow="7" pageId="9" pageNumber="10">HW (mm)</th>
<td box="[363,589,583,605]" gridcol="1" gridrow="7" pageId="9" pageNumber="10">13.3–18.5 (16.0 ± 2.60)</td>
<td box="[696,850,583,605]" gridcol="2" gridrow="7" pageId="9" pageNumber="10">14.2</td>
<td box="[957,1183,583,605]" gridcol="3" gridrow="7" pageId="9" pageNumber="10">9.1–12.0 (10.6 ± 0.9)</td>
<td box="[1290,1515,583,605]" gridcol="4" gridrow="7" pageId="9" pageNumber="10">6.4–14.2 (9.8 ± 1.43)</td>
</tr>
<tr box="[113,1515,619,641]" gridrow="8" pageId="9" pageNumber="10">
<th box="[113,256,619,641]" gridcol="0" gridrow="8" pageId="9" pageNumber="10">HH (mm)</th>
<td box="[363,589,619,641]" gridcol="1" gridrow="8" pageId="9" pageNumber="10">7.4–10.1 (8.80 ± 1.35)</td>
<td box="[696,850,619,641]" gridcol="2" gridrow="8" pageId="9" pageNumber="10">7.6</td>
<td box="[957,1183,619,641]" gridcol="3" gridrow="8" pageId="9" pageNumber="10">4.8–6.4 (5.8 ± 0.5)</td>
<td box="[1290,1515,619,641]" gridcol="4" gridrow="8" pageId="9" pageNumber="10">3.4–8.8 (5.1 ± 0.77)</td>
</tr>
<tr box="[113,1515,655,677]" gridrow="9" pageId="9" pageNumber="10">
<th box="[113,256,655,677]" gridcol="0" gridrow="9" pageId="9" pageNumber="10">SnL (mm)</th>
<td box="[363,589,655,677]" gridcol="1" gridrow="9" pageId="9" pageNumber="10">7.0–7.9 (7.5 ± 0.47)</td>
<td box="[696,850,655,677]" gridcol="2" gridrow="9" pageId="9" pageNumber="10">7.0</td>
<td box="[957,1183,655,677]" gridcol="3" gridrow="9" pageId="9" pageNumber="10">4.8–5.8 (5.1 ± 0.3)</td>
<td box="[1290,1515,655,677]" gridcol="4" gridrow="9" pageId="9" pageNumber="10">3.2–6.0 (4.4 ± 0.54)</td>
</tr>
<tr box="[113,1515,691,714]" gridrow="10" pageId="9" pageNumber="10">
<th box="[113,256,691,714]" gridcol="0" gridrow="10" pageId="9" pageNumber="10">ED (mm)</th>
<td box="[363,589,691,714]" gridcol="1" gridrow="10" pageId="9" pageNumber="10">4.3–4.3 (4.3 ± 0.00)</td>
<td box="[696,850,691,714]" gridcol="2" gridrow="10" pageId="9" pageNumber="10">3.7</td>
<td box="[957,1183,691,714]" gridcol="3" gridrow="10" pageId="9" pageNumber="10">2.2–3.3 (2.6 ± 0.3)</td>
<td box="[1290,1515,691,714]" gridcol="4" gridrow="10" pageId="9" pageNumber="10">1.8–3.2 (2.4 ± 0.29)</td>
</tr>
<tr box="[113,1515,727,750]" gridrow="11" pageId="9" pageNumber="10">
<th box="[113,256,727,750]" gridcol="0" gridrow="11" pageId="9" pageNumber="10">IOD (mm)</th>
<td box="[363,589,727,750]" gridcol="1" gridrow="11" pageId="9" pageNumber="10">7.7–9.6 (8.7 ± 0.95)</td>
<td box="[696,850,727,750]" gridcol="2" gridrow="11" pageId="9" pageNumber="10">7.0</td>
<td box="[957,1183,727,750]" gridcol="3" gridrow="11" pageId="9" pageNumber="10">4.6–6.4 (5.3 ± 0.5)</td>
<td box="[1290,1515,727,750]" gridcol="4" gridrow="11" pageId="9" pageNumber="10">3.5–6.7 (4.9 ± 0.64)</td>
</tr>
<tr box="[113,1515,764,786]" gridrow="12" pageId="9" pageNumber="10">
<th box="[113,256,764,786]" gridcol="0" gridrow="12" pageId="9" pageNumber="10">EED (mm)</th>
<td box="[363,589,764,786]" gridcol="1" gridrow="12" pageId="9" pageNumber="10">4.9–6.3 (5.4 ± 0.78)</td>
<td box="[696,850,764,786]" gridcol="2" gridrow="12" pageId="9" pageNumber="10">4.8</td>
<td box="[957,1183,764,786]" gridcol="3" gridrow="12" pageId="9" pageNumber="10">2.8–4.0 (3.4 ± 0.4)</td>
<td box="[1290,1515,764,786]" gridcol="4" gridrow="12" pageId="9" pageNumber="10">2.2–4.1 (3.1 ± 0.41)</td>
</tr>
<tr box="[113,1515,800,822]" gridrow="13" pageId="9" pageNumber="10">
<th box="[113,256,800,822]" gridcol="0" gridrow="13" pageId="9" pageNumber="10">IOS</th>
<td box="[363,589,800,822]" gridcol="1" gridrow="13" pageId="9" pageNumber="10">9–10 (9.67 ± 0.58)</td>
<td box="[696,850,800,822]" gridcol="2" gridrow="13" pageId="9" pageNumber="10">9</td>
<td box="[957,1183,800,822]" gridcol="3" gridrow="13" pageId="9" pageNumber="10">9–12 (10.8 ± 1.09)</td>
<td box="[1290,1515,800,822]" gridcol="4" gridrow="13" pageId="9" pageNumber="10">9–13 (10.6 ± 0.93)</td>
</tr>
<tr box="[113,1515,836,894]" gridrow="14" pageId="9" pageNumber="10">
<th box="[113,256,836,894]" gridcol="0" gridrow="14" pageId="9" pageNumber="10">SPL IFL</th>
<td box="[363,589,836,894]" gridcol="1" gridrow="14" pageId="9" pageNumber="10">7–8 (7.33 ± 0.52) 4.5–4.8 (4.6 ± 0.15)a</td>
<td box="[696,850,836,894]" gridcol="2" gridrow="14" pageId="9" pageNumber="10">6 4.1</td>
<td box="[957,1183,836,894]" gridcol="3" gridrow="14" pageId="9" pageNumber="10">6.0–7.0 (6.91 ± 0.30) 4.0–5.5 (4.65 ± 0.45)</td>
<td box="[1290,1515,836,894]" gridcol="4" gridrow="14" pageId="9" pageNumber="10">5.0–7.0 (6.06 ± 0.44) 2.8–4.6 (3.72 ± 0.39)</td>
</tr>
<tr box="[113,1515,908,930]" gridrow="15" pageId="9" pageNumber="10">
<th box="[113,256,908,930]" gridcol="0" gridrow="15" pageId="9" pageNumber="10">CS</th>
<td box="[363,589,908,930]" gridcol="1" gridrow="15" pageId="9" pageNumber="10">6–7 (6.7 ± 0.58)</td>
<td box="[696,850,908,930]" gridcol="2" gridrow="15" pageId="9" pageNumber="10">6</td>
<td box="[957,1183,908,930]" gridcol="3" gridrow="15" pageId="9" pageNumber="10">6–7 (6.1 ± 0.30)</td>
<td box="[1290,1515,908,930]" gridcol="4" gridrow="15" pageId="9" pageNumber="10">4–7 (5.5 ± 0.58)</td>
</tr>
<tr box="[113,1515,944,967]" gridrow="16" pageId="9" pageNumber="10">
<th box="[113,256,944,967]" gridcol="0" gridrow="16" pageId="9" pageNumber="10">MBS</th>
<td box="[363,589,944,967]" gridcol="1" gridrow="16" pageId="9" pageNumber="10">17–18 (17.7 ± 0.58)</td>
<td box="[696,850,944,967]" gridcol="2" gridrow="16" pageId="9" pageNumber="10">25</td>
<td box="[957,1183,944,967]" gridcol="3" gridrow="16" pageId="9" pageNumber="10">30–37 (34.5 ± 2.46)</td>
<td box="[1290,1515,944,967]" gridcol="4" gridrow="16" pageId="9" pageNumber="10">26–36 (30.9 ± 1.98)</td>
</tr>
<tr box="[113,1515,980,1003]" gridrow="17" pageId="9" pageNumber="10">
<th box="[113,256,980,1003]" gridcol="0" gridrow="17" pageId="9" pageNumber="10">VS</th>
<td box="[363,589,980,1003]" gridcol="1" gridrow="17" pageId="9" pageNumber="10">27–31 (29.0 ± 2.00)</td>
<td box="[696,850,980,1003]" gridcol="2" gridrow="17" pageId="9" pageNumber="10">32</td>
<td box="[957,1183,980,1003]" gridcol="3" gridrow="17" pageId="9" pageNumber="10">40–50 (44.6 ± 3.01)</td>
<td box="[1290,1515,980,1003]" gridcol="4" gridrow="17" pageId="9" pageNumber="10">37–48 (41.6 ± 0.22)</td>
</tr>
<tr box="[113,1515,1017,1039]" gridrow="18" pageId="9" pageNumber="10">
<th box="[113,256,1017,1039]" gridcol="0" gridrow="18" pageId="9" pageNumber="10">VHL</th>
<td box="[363,589,1017,1039]" gridcol="1" gridrow="18" pageId="9" pageNumber="10">6.0–8.0 (7.0 ± 1.00)</td>
<td box="[696,850,1017,1039]" gridcol="2" gridrow="18" pageId="9" pageNumber="10">9</td>
<td box="[957,1183,1017,1039]" gridcol="3" gridrow="18" pageId="9" pageNumber="10">7.0–13.0 (10.41 ± 1.76)</td>
<td box="[1290,1515,1017,1039]" gridcol="4" gridrow="18" pageId="9" pageNumber="10">7.0–12.0 (8.78 ± 1.11)</td>
</tr>
<tr box="[113,1515,1053,1075]" gridrow="19" pageId="9" pageNumber="10">
<th box="[113,256,1053,1075]" gridcol="0" gridrow="19" pageId="9" pageNumber="10">L1TT</th>
<td box="[363,589,1053,1075]" gridcol="1" gridrow="19" pageId="9" pageNumber="10">14–15 (14.33 ± 0.58)</td>
<td box="[696,850,1053,1075]" gridcol="2" gridrow="19" pageId="9" pageNumber="10">12</td>
<td box="[957,1183,1053,1075]" gridcol="3" gridrow="19" pageId="9" pageNumber="10">8–15 (10.4 ± 1.86)</td>
<td box="[1290,1515,1053,1075]" gridcol="4" gridrow="19" pageId="9" pageNumber="10">8–13 (9.6 ± 1.03)</td>
</tr>
<tr box="[113,1515,1089,1111]" gridrow="20" pageId="9" pageNumber="10">
<th box="[113,256,1089,1111]" gridcol="0" gridrow="20" pageId="9" pageNumber="10">L4TT</th>
<td box="[363,589,1089,1111]" gridcol="1" gridrow="20" pageId="9" pageNumber="10">18–21 (19.67 ± 1.53)</td>
<td box="[696,850,1089,1111]" gridcol="2" gridrow="20" pageId="9" pageNumber="10">17</td>
<td box="[957,1183,1089,1111]" gridcol="3" gridrow="20" pageId="9" pageNumber="10">10–18 (15.3 ± 2.31)</td>
<td box="[1290,1515,1089,1111]" gridcol="4" gridrow="20" pageId="9" pageNumber="10">11–18 (14.9 ± 1.09)</td>
</tr>
<tr box="[113,1515,1125,1147]" gridrow="21" pageId="9" pageNumber="10">
<th box="[113,256,1125,1147]" gridcol="0" gridrow="21" pageId="9" pageNumber="10">SAE/SPE</th>
<td box="[363,589,1125,1147]" gridcol="1" gridrow="21" pageId="9" pageNumber="10">0.42–0.48 (0.45 ± 0.03)</td>
<td box="[696,850,1125,1147]" gridcol="2" gridrow="21" pageId="9" pageNumber="10">0.49</td>
<td box="[957,1183,1125,1147]" gridcol="3" gridrow="21" pageId="9" pageNumber="10">0.67–1.00 (0.79 ± 0.01)</td>
<td box="[1290,1515,1125,1147]" gridcol="4" gridrow="21" pageId="9" pageNumber="10">0.50–0.80 (0.63 ± 0.07)</td>
</tr>
</table>
</paragraph>
<tableNote pageId="9" pageNumber="10" targetBox="[113,1515,327,1147]" targetPageId="9">
<paragraph blockId="9.[91,1484,1170,1263]" box="[91,144,1170,1186]" pageId="9" pageNumber="10">
<heading bold="true" box="[91,144,1170,1186]" fontSize="7" level="5" pageId="9" pageNumber="10" reason="6">
<emphasis bold="true" box="[91,144,1170,1186]" pageId="9" pageNumber="10">Notes.</emphasis>
</heading>
</paragraph>
<paragraph blockId="9.[91,1484,1170,1263]" pageId="9" pageNumber="10">
Ranges for 
<taxonomicName box="[220,309,1196,1213]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="polylepis">
<emphasis box="[220,309,1196,1213]" italics="true" pageId="9" pageNumber="10">G. polylepis</emphasis>
</taxonomicName>
are probably reliable, but those for 
<taxonomicName box="[593,662,1196,1212]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="typica">
<emphasis box="[593,662,1196,1212]" italics="true" pageId="9" pageNumber="10">G. typica</emphasis>
</taxonomicName>
certainly come from several OTUs, and are therefore not actually representative of the variation of that species. Values for true 
<taxonomicName box="[318,387,1221,1237]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="typica">
<emphasis box="[318,387,1221,1237]" italics="true" pageId="9" pageNumber="10">G. typica</emphasis>
</taxonomicName>
, as for 
<taxonomicName box="[445,540,1221,1238]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[445,540,1221,1238]" italics="true" pageId="9" pageNumber="10">G. maculata</emphasis>
</taxonomicName>
, are currently certain only from its holotype. aThis value includes the specimen figured in 
<figureCitation box="[487,538,1246,1263]" captionStart="Figure 2" captionStartId="4.[507,571,1276,1298]" captionTargetBox="[491,1541,229,1231]" captionTargetId="figure@4.[491,1541,229,1231]" captionTargetPageId="4" captionText="Figure 2 Schematic drawings exemplifying morphological measurements and scale counts, following the scheme of Hawlitschek et al. (2016) based on Köhler et al. (2009). (A) Measurements and pholidosis of the head, drawn from the specimen in Fig. 3 A (small scales around the eye and nostril not depicted); (B) body measurements; (C) postmental scale conditions, adapted from Lemme et al. (2013); (D) postnasal scale configuration types, adapted from Köhler et al. (2009). Scale counts in A: purple, canthal scales; blue, supralabials; red, infralabials. In C, cyan indicates mental scales and purple indicates postmental scales. For abbreviations and unillustrated characters, see Materials and Methods." httpUri="https://zenodo.org/record/291227/files/figure.png" pageId="9" pageNumber="10">Figs. 2</figureCitation>
and 
<figureCitation box="[577,585,1246,1263]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="9" pageNumber="10">3</figureCitation>
A, and has a sample size of 4.
</paragraph>
</tableNote>
<paragraph blockId="9.[491,1541,1294,1913]" pageId="9" pageNumber="10">
<taxonomicName box="[491,594,1294,1320]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[491,594,1294,1320]" italics="true" pageId="9" pageNumber="10">humbloti</emphasis>
</taxonomicName>
by fewer scale rows around midbody (17–18 vs. 22–30), more infralabials to the anterior margin of the eye (4.5–4.8 vs. 3–4), and fewer ventral scales (27–31 vs. 33–41).
</paragraph>
<paragraph blockId="9.[491,1541,1294,1913]" pageId="9" pageNumber="10">
For comparison of the osteology of the new species with 
<taxonomicName box="[1228,1458,1373,1399]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1228,1458,1373,1399]" italics="true" pageId="9" pageNumber="10">Geckolepis maculata</emphasis>
</taxonomicName>
, 
<taxonomicName box="[491,628,1413,1439]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[491,628,1413,1439]" italics="true" pageId="9" pageNumber="10">G. humbloti</emphasis>
</taxonomicName>
, and a specimen of OTU AB, see the Osteology of 
<taxonomicName box="[1220,1337,1413,1439]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="10" phylum="Chordata" rank="genus">
<emphasis box="[1220,1337,1413,1439]" italics="true" pageId="9" pageNumber="10">Geckolepis</emphasis>
</taxonomicName>
section below.
</paragraph>
</subSubSection>
<subSubSection lastPageId="11" lastPageNumber="12" pageId="9" pageNumber="10" type="description">
<paragraph blockId="9.[491,1541,1294,1913]" pageId="9" pageNumber="10">
<emphasis box="[523,823,1452,1478]" italics="true" pageId="9" pageNumber="10">Description of the holotype</emphasis>
. (
<figureCitation box="[850,916,1452,1478]" captionStart="Figure 4" captionStartId="7.[507,571,1752,1774]" captionTargetBox="[491,1512,230,1704]" captionTargetId="figure@7.[491,1514,229,1708]" captionTargetPageId="7" captionText="Figure 4 Specimens of Geckolepis megalepis sp. nov. in preservative. (A – C) The holotype, ZSM 2126 / 2007; (D – F) paratype ZSM 232 / 2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale." httpUri="https://zenodo.org/record/291229/files/figure.png" pageId="9" pageNumber="10">Fig. 4</figureCitation>
) A large specimen in a moderately good state of preservation. Several scales missing from dorsum and the left knee, and a few older scars on the venter; tail detached but preserved, presumably autotomized during or after capture.
</paragraph>
<paragraph blockId="9.[491,1541,1294,1913]" lastBlockId="10.[491,1542,228,1913]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
SVL 68.4 mm; tail length 80.1 mm; axilla-groin distance 27.2 mm; shank length 5.8 mm; head length 15.9 mm; head width 16.1 mm; head height 8.9 mm; snout length 7.7 mm; eye diameter 4.3 mm; interorbital distance 8.7 mm; eye-ear distance 5.0 mm; rostral large, convex, distinctly visible from above; large postrostrals (4 supranasals) separated by a thin oblong scale anteriorly and two small scales posteriorly (type D); nostril bordered by rostral, first supralabial, three postnasals, and one supranasal (=postrostral); postnasals approximately equal in size to anterior loreals; postnasals and loreals separated by one row of small scales; scales on snout and on loreal region almost all with triangular posterior margins, slightly convex, imbricate; seven canthal scales in a line on the canthal ridge, including the postnasals, between nostril and anterior margin of eye; scales at supraorbital region similar to and continuous with those on top of head; scales increase by a factor of 1.68 from level of mideye to occipital region; ten scales along a straight line dorsally between orbits; three rows of small scales adjacent to anterior margin of eye, decreasing to two on the upper and two on the posterior margin of eye; pupil vertical; seven supralabials to below centre of eye (eight total) on left side, eight to below centre of eye (nine total) on right side, all of roughly equal size, the posterior-most two smaller than the rest; scales in temporal region larger by factor 2.25 than those in loreal region; ear opening much smaller than eye, horizontally oval; 4.5 infralabials to below level of anterior margin of eye, decreasing in size posteriorly; mental scale large, triangular, with convex anterior border; postmentals asymmetric: one large pair immediately posterior to mental scale with broad medial contact, followed on the right side by two postmentals of decreasing size, and on the left by one broader postmental with an irregular posteromedial border (condition A/B; 
<figureCitation box="[683,747,741,768]" captionStart="Figure 4" captionStartId="7.[507,571,1752,1774]" captionTargetBox="[491,1512,230,1704]" captionTargetId="figure@7.[491,1514,229,1708]" captionTargetPageId="7" captionText="Figure 4 Specimens of Geckolepis megalepis sp. nov. in preservative. (A – C) The holotype, ZSM 2126 / 2007; (D – F) paratype ZSM 232 / 2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale." httpUri="https://zenodo.org/record/291229/files/figure.png" pageId="10" pageNumber="11">Fig. 4</figureCitation>
); one row of small scales separating postmentals from anterior chin scales; chin and ventral body scales rhomboid with rounded triangular posterior margins, imbricate, gradually increasing to double size posteriorly, arranged in roughly regular rows; 29 scales along the midventral line (count somewhat inhibited by ventral scarring) between postmentals and vent, not including smaller scales adjacent to postmentals and anterior to vent; dorsal scales cycloid, larger than lateral or ventral scales; 18 scales around midbody; anterior half of tail flat and rather wide (55% of body width), decreasing in width gradually, down to a thin tip; tail covered with scales similar to body scales, but gradually decreasing in size posteriorly, a series of transversely expanded median subcaudals present, starting roughly 20 mm from the base of the tail; 14 lamellae under first toe, extending from the sole of the foot to the claw, and 21 under fourth toe, 12 of which are noticeably expanded; claws exposed, non-retractile. A single midbody scale measures 5.0 mm wide by 5.8 mm long, and is therefore 7.8% of the SVL in length.
</paragraph>
<paragraph blockId="10.[491,1542,228,1913]" pageId="10" pageNumber="11">Colouration in preservative: Head dorsally homogenously greyish brown, laterally greyishbrown flecked with darker and lighter areas posterior to eye and below mouth; no obvious dark lateral stripe; dorsal scales greyish brown flecked with dark and pale spots; legs as dorsum; ventrally dirty white; tail greyish brown with four dark transverse markings that do not continue onto the whitish ventral surface; exposed dorsal skin brown, ventral skin whitish. No information exists regarding the life colouration of the specimen.</paragraph>
<paragraph blockId="10.[491,1542,228,1913]" lastBlockId="11.[491,1542,228,1913]" lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11">
<emphasis box="[523,633,1493,1518]" italics="true" pageId="10" pageNumber="11">Variation</emphasis>
. (
<figureCitation box="[660,736,1492,1518]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="10" pageNumber="11">Figs. 3</figureCitation>
and 
<figureCitation box="[799,813,1492,1518]" captionStart="Figure 4" captionStartId="7.[507,571,1752,1774]" captionTargetBox="[491,1512,230,1704]" captionTargetId="figure@7.[491,1514,229,1708]" captionTargetPageId="7" captionText="Figure 4 Specimens of Geckolepis megalepis sp. nov. in preservative. (A – C) The holotype, ZSM 2126 / 2007; (D – F) paratype ZSM 232 / 2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale." httpUri="https://zenodo.org/record/291229/files/figure.png" pageId="10" pageNumber="11">4</figureCitation>
) Both ZSM paratypes strongly resemble the holotype, but one (ZSM 289/2004) is much smaller and presumed subadult. Their measurements and meristics are provided as part of 
<tableCitation box="[866,950,1571,1597]" captionStart="Table 1" captionStartId="9.[106,162,246,267]" captionTargetBox="[113,1515,327,1147]" captionText="Table 1 Comparative morphological data on Geckolepis megalepis, and the holotype of G. maculata (ZMB 9655), with values for G. polylepis and G. typica given by Köhler et al. (2009). For raw measurements of individual G. megalepis specimens, see Appendix S 1." httpUri="http://table.plazi.org/id/DF50D4D9FFEBFF85FF9EFF6D109BFEBA" pageId="10" pageNumber="11" tableUuid="DF50D4D9FFEBFF85FF9EFF6D109BFEBA">Table 1</tableCitation>
, and in full detail in Appendix S1. These specimens differ from the holotype in the following characters not provided in Appendix S1: scales increase by a factor of 2.12–2.13 from level of mideye to occipital region; eight or nine total supralabials on each side; postnasal configuration of ZSM 232/2016 type A, 289/2004 type D; postmentals large, three or four pairs present: one large pair immediately posterior to mental scale with broad medial contact, followed two or three pairs of decreasing size (condition A); mid-dorsal scales range from 7.3% to 8.3% of SVL; tail of ZSM 289/2004 not especially broadened (regenerated) and greyish brown with three darker transverse markings that do not continue onto the whitish ventral surface; tail of ZSM 232/2016 broad at its base and narrows rapidly (at least partially regenerated); overall body colouration of ZSM 232/2016 is greyer than the other specimens, though this may be because it was collected more recently. In ZSM 289/2004, one scale on the dorsal surface of the neck and one on left side of dorsal tail base are dark brown with a burned appearance. The animal in 
<figureCitation box="[522,583,386,412]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="11" pageNumber="12">Fig. 3</figureCitation>
A has 4.8 infralabials to the anterior edge of the eye (see 
<figureCitation box="[1234,1297,386,412]" captionStart="Figure 2" captionStartId="4.[507,571,1276,1298]" captionTargetBox="[491,1541,229,1231]" captionTargetId="figure@4.[491,1541,229,1231]" captionTargetPageId="4" captionText="Figure 2 Schematic drawings exemplifying morphological measurements and scale counts, following the scheme of Hawlitschek et al. (2016) based on Köhler et al. (2009). (A) Measurements and pholidosis of the head, drawn from the specimen in Fig. 3 A (small scales around the eye and nostril not depicted); (B) body measurements; (C) postmental scale conditions, adapted from Lemme et al. (2013); (D) postnasal scale configuration types, adapted from Köhler et al. (2009). Scale counts in A: purple, canthal scales; blue, supralabials; red, infralabials. In C, cyan indicates mental scales and purple indicates postmental scales. For abbreviations and unillustrated characters, see Materials and Methods." httpUri="https://zenodo.org/record/291227/files/figure.png" pageId="11" pageNumber="12">Fig. 2</figureCitation>
A), and we thus infer that this value can range from 4.5 to 4.8. From 
<figureCitation box="[1049,1122,425,452]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="11" pageNumber="12">Figs. 3</figureCitation>
A and 3B, the colouration in life is assessed to be mostly grey with dark spots on some scales, giving a mottled appearance.
</paragraph>
</subSubSection>
<subSubSection pageId="11" pageNumber="12" type="discussion">
<paragraph blockId="11.[491,1542,228,1913]" pageId="11" pageNumber="12">
<emphasis box="[523,812,504,530]" italics="true" pageId="11" pageNumber="12">Phylogenetic relationships</emphasis>
. 
<taxonomicName box="[828,1061,504,530]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[828,1061,504,530]" italics="true" pageId="11" pageNumber="12">
<taxonomicName box="[828,945,504,530]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Geckolepis</taxonomicName>
<taxonomicName box="[952,1061,504,530]" pageId="11" pageNumber="12" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
is closely related to a sister species pair formed by OTUs C and AB (
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[836,1049,544,570]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="11" pageNumber="12" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[836,1049,544,570]" italics="true" pageId="11" pageNumber="12">Lemme et al., 2013</emphasis>
</bibRefCitation>
; 
<figureCitation box="[1064,1129,544,570]" captionStart="Figure 1" captionStartId="2.[507,571,950,972]" captionTargetBox="[1198,1540,229,885]" captionTargetPageId="2" captionText="Figure 1 Molecular phylogeny and distribution of Geckolepis OTUs based mainly on Lemme et al. (2013). Dotted lines in the phylogeny indicate uncertain placement. The phylogenetic position and distribution of Geckolepis humbloti is inferred from Hawlitschek et al. (2016)." httpUri="https://zenodo.org/record/291226/files/figure.svg" pageId="11" pageNumber="12">Fig. 1</figureCitation>
), which are widely distributed in eastern and northern Madagascar. Their taxonomic status will need to be assessed in more detail in future work on this genus.
</paragraph>
<paragraph blockId="11.[491,1542,228,1913]" pageId="11" pageNumber="12">
<emphasis box="[523,1069,662,688]" italics="true" pageId="11" pageNumber="12">Habitat, natural history, and conservation status</emphasis>
. 
<taxonomicName box="[1080,1313,662,688]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1080,1313,662,688]" italics="true" pageId="11" pageNumber="12">
<taxonomicName box="[1080,1197,662,688]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Geckolepis</taxonomicName>
<taxonomicName box="[1204,1313,662,688]" pageId="11" pageNumber="12" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
was observed active at night both in the rainy and dry seasons, on trees (see 
<figureCitation box="[1143,1216,702,728]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="11" pageNumber="12">Figs. 3</figureCitation>
A–3B) and tsingy limestone rock. When captured, these geckos showed a strong tendency to autotomize large parts of their scales, leading to partly ‘naked’ geckos without any visible (bloody) lesions (
<figureCitation box="[1440,1503,781,807]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="11" pageNumber="12">Fig. 3</figureCitation>
C). In a subjective comparison this tendency appeared to be even more developed than in other 
<taxonomicName box="[561,678,860,886]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis box="[561,678,860,886]" italics="true" pageId="11" pageNumber="12">Geckolepis</emphasis>
</taxonomicName>
species.
</paragraph>
<paragraph blockId="11.[491,1542,228,1913]" pageId="11" pageNumber="12">
The new species occurs syntopically in Ankarana with at least one additional 
<taxonomicName box="[1424,1541,899,925]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis box="[1424,1541,899,925]" italics="true" pageId="11" pageNumber="12">Geckolepis</emphasis>
</taxonomicName>
species (OTU G). It is only known from the dry deciduous forest among the limestone tsingy karst of Ankarana Reserve and its immediate vicinity, an area of 182 km2. Due to its likely limited distribution (182 km2), knowledge from only two threat-defined localities in the Ankarana massif, and the potential for rapid decrease in quality of the forests of that reserve and the area around it due to illicit deforestation, anthropogenic fire, sapphire mining, and free-ranging grazing of livestock (e.g., 
<bibRefCitation author="Hawkins AFA &amp; Chapman P &amp; GanZhorn JU &amp; Bloxam QMC &amp; Barlow SC &amp; Tongue SJ." box="[1085,1315,1136,1162]" journalOrPublisher="Biological Conservation" pageId="11" pageNumber="12" pagination="83 - 110" part="43" publicationUrl="10.1016/0006-3207(90)90136-D." refString="Hawkins AFA, Chapman P, GanZhorn JU, Bloxam QMC, Barlow SC, Tongue SJ. 1990. Vertebrate conservation in Ankarana Special Reserve, northern Madagascar. Biological Conservation 43: 83 - 110 DOI 10.1016 / 0006 - 3207 (90) 90136 - D." title="Vertebrate conservation in Ankarana Special Reserve, northern Madagascar" type="journal article" year="1990">
<emphasis box="[1085,1315,1136,1162]" italics="true" pageId="11" pageNumber="12">Hawkins et al., 1990</emphasis>
</bibRefCitation>
) we propose that it be listed as Near Threatened under the IUCN criteria (
<bibRefCitation author="IUCN" box="[1132,1271,1176,1202]" journalOrPublisher="Gland, Cambridge: IUCN" pageId="11" pageNumber="12" refString="IUCN. 2012. IUCN red list categories and criteria: version 3.1. Gland, Cambridge: IUCN." title="IUCN red list categories and criteria: version 3.1" type="book" year="2012">
<emphasis box="[1132,1271,1176,1202]" italics="true" pageId="11" pageNumber="12">IUCN, 2012</emphasis>
</bibRefCitation>
).
</paragraph>
<paragraph blockId="11.[491,1542,228,1913]" pageId="11" pageNumber="12">
Two other geckos endemic to Ankarana Reserve are assessed as Near Threatened (
<taxonomicName authority="Pasteur &amp; Blanc, 1967" authorityName="Pasteur &amp; Blanc" authorityYear="1967" box="[502,1030,1255,1282]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="expectatus">
<emphasis box="[502,769,1255,1281]" italics="true" pageId="11" pageNumber="12">
<taxonomicName box="[502,647,1255,1281]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Lygodactylus</taxonomicName>
expectatus
</emphasis>
Pasteur &amp; Blanc, 1967
</taxonomicName>
) and Endangered (
<taxonomicName authority="Glaw et al., 2010" authorityName="Glaw et al." authorityYear="2010" class="Reptilia" family="Gekkonidae" genus="Phelsuma" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="roesleri">
<emphasis box="[1251,1448,1255,1281]" italics="true" pageId="11" pageNumber="12">
<taxonomicName box="[1251,1361,1255,1281]" class="Reptilia" family="Gekkonidae" genus="Phelsuma" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Phelsuma</taxonomicName>
roesleri
</emphasis>
<bibRefCitation author="Glaw F &amp; Gehring P-S &amp; Kohler J &amp; FranZen M &amp; Vences M." journalOrPublisher="Salamandra" pageId="11" pageNumber="12" pagination="83 - 92" part="46" refString="Glaw F, Gehring P-S, Kohler J, FranZen M, Vences M. 2010. A new dwarf species of day gecko, genus Phelsuma, from the Ankarana pinnacle karst in northern Madagascar. Salamandra 46: 83 - 92." title="A new dwarf species of day gecko, genus Phelsuma, from the Ankarana pinnacle karst in northern Madagascar" type="journal article" year="2010">Glaw et al., 2010</bibRefCitation>
</taxonomicName>
). We defend the choice of Near Threatened instead of Endangered for 
<taxonomicName box="[1399,1541,1294,1321]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1399,1541,1294,1321]" italics="true" pageId="11" pageNumber="12">G. megalepis</emphasis>
</taxonomicName>
on the following grounds: although it satisfies IUCN criterion B1 sub-criterion a, it fails to qualify for Endangered under sub-criteria b or c, as we can only identify potential threats; were these to be realised, then the species would immediately qualify for Endangered, but until that point, it remains Near Threatened. The same cannot be said of 
<taxonomicName box="[1356,1466,1452,1478]" class="Reptilia" family="Gekkonidae" genus="Phelsuma" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="roesleri">
<emphasis box="[1356,1466,1452,1478]" italics="true" pageId="11" pageNumber="12">P. roesleri</emphasis>
</taxonomicName>
, as it lives on 
<taxonomicName box="[586,699,1492,1518]" class="Liliopsida" family="Pandanaceae" genus="Pandanus" kingdom="Plantae" order="Pandanales" pageId="11" pageNumber="12" phylum="Tracheophyta" rank="genus">
<emphasis box="[586,699,1492,1518]" italics="true" pageId="11" pageNumber="12">Pandanus</emphasis>
</taxonomicName>
plants, which are less common in Ankarana and potentially harvested by humans, making the risk to it greater, while 
<taxonomicName box="[994,1136,1531,1558]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[994,1136,1531,1558]" italics="true" pageId="11" pageNumber="12">G. megalepis</emphasis>
</taxonomicName>
is more generalist in its habits. Our recent observation of several individuals in a short timespan suggests that the population of 
<taxonomicName authority="in Ankarana" authorityName="in Ankarana" box="[522,908,1610,1636]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[522,755,1610,1636]" italics="true" pageId="11" pageNumber="12">
<taxonomicName box="[522,639,1610,1636]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Geckolepis</taxonomicName>
megalepis
</emphasis>
in Ankarana
</taxonomicName>
is at least locally healthy.
</paragraph>
</subSubSection>
<subSubSection lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" type="etymology">
<paragraph blockId="11.[491,1542,228,1913]" pageId="11" pageNumber="12">
<emphasis box="[523,641,1650,1676]" italics="true" pageId="11" pageNumber="12">Etymology</emphasis>
. The specific epithet is derived from the two Greek stems 
<emphasis box="[1331,1417,1650,1676]" italics="true" pageId="11" pageNumber="12">µ´ εγ ας</emphasis>
(mégas) meaning ‘very large’ and 
<emphasis box="[788,862,1690,1716]" italics="true" pageId="11" pageNumber="12">λεπ´ ις</emphasis>
(lepís) meaning ‘scale’, and refers to the large size of the scales of this species in comparison to its congeners and other geckos, which aids also in its diagnosis.
</paragraph>
<paragraph blockId="11.[491,1542,228,1913]" lastBlockId="12.[491,1542,228,773]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
<emphasis box="[523,708,1808,1834]" italics="true" pageId="11" pageNumber="12">Available names</emphasis>
. Three junior synonyms currently exist within the genus 
<taxonomicName box="[1371,1488,1808,1834]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis box="[1371,1488,1808,1834]" italics="true" pageId="11" pageNumber="12">Geckolepis</emphasis>
</taxonomicName>
that must be considered as possible earlier names for 
<taxonomicName box="[1045,1187,1847,1874]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1045,1187,1847,1874]" italics="true" pageId="11" pageNumber="12">G. megalepis</emphasis>
</taxonomicName>
: 
<taxonomicName authority="Mocquard, 1909" authorityName="Mocquard" authorityYear="1909" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="subSpecies" species="typica" subSpecies="anomala">
<taxonomicName authority="Mocquard" authorityName="Mocquard" box="[1200,1537,1847,1874]" pageId="11" pageNumber="12" rank="subSpecies" species="typica" subSpecies="anomala">
<taxonomicName box="[1200,1406,1847,1874]" pageId="11" pageNumber="12" rank="subSpecies" species="typica" subSpecies="anomala">
<emphasis box="[1200,1406,1847,1874]" italics="true" pageId="11" pageNumber="12">G. typica anomala</emphasis>
</taxonomicName>
Mocquard
</taxonomicName>
, 1909
</taxonomicName>
, 
<taxonomicName authority="Methuen &amp; Hewitt, 1913" authorityName="Methuen &amp; Hewitt" authorityYear="1913" box="[562,1065,1887,1913]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="subSpecies" species="typica" subSpecies="modesta">
<taxonomicName authority="Methuen &amp; Hewitt" authorityName="Methuen &amp; Hewitt" box="[562,995,1887,1913]" pageId="11" pageNumber="12" rank="subSpecies" species="typica" subSpecies="modesta">
<taxonomicName authority="Methuen" authorityName="Methuen" box="[562,877,1887,1913]" pageId="11" pageNumber="12" rank="subSpecies" species="typica" subSpecies="modesta">
<emphasis box="[562,763,1887,1913]" italics="true" pageId="11" pageNumber="12">G. typica modesta</emphasis>
Methuen
</taxonomicName>
&amp; Hewitt
</taxonomicName>
, 1913
</taxonomicName>
, and 
<taxonomicName authority="Angel, 1942" authorityName="Angel" authorityYear="1942" box="[1129,1369,1887,1913]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="petiti">
<emphasis box="[1129,1220,1887,1913]" italics="true" pageId="11" pageNumber="12">G. petiti</emphasis>
<bibRefCitation author="Angel F." box="[1228,1369,1887,1913]" journalOrPublisher="Memoires de l'Academie Malgache" pageId="11" pageNumber="12" pagination="1 - 139" part="36" refString="Angel F. 1942. Les lezards de Madagascar. Memoires de l'Academie Malgache 36: 1 - 139." title="Les lezards de Madagascar" type="journal article" year="1942">Angel, 1942</bibRefCitation>
</taxonomicName>
. Synonymy of 
<emphasis box="[491,765,228,254]" italics="true" pageId="12" pageNumber="13">
<taxonomicName box="[491,627,228,254]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="anomala">G. anomala</taxonomicName>
, 
<taxonomicName box="[638,765,228,254]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="modesta">G. modesta</taxonomicName>
</emphasis>
, and 
<taxonomicName box="[830,921,228,254]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="petiti">
<emphasis box="[830,921,228,254]" italics="true" pageId="12" pageNumber="13">G. petiti</emphasis>
</taxonomicName>
with 
<taxonomicName box="[987,1088,228,254]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="typica">
<emphasis box="[987,1088,228,254]" italics="true" pageId="12" pageNumber="13">G. typica</emphasis>
</taxonomicName>
was discussed at length by 
<bibRefCitation author="Kohler G &amp; Diethert H-H &amp; Nussbaum RA &amp; Raxworthy CJ." journalOrPublisher="Herpetologica" pageId="12" pageNumber="13" pagination="419 - 435" part="65" publicationUrl="10.1655/08-059.1." refString="Kohler G, Diethert H-H, Nussbaum RA, Raxworthy CJ. 2009. A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros. Herpetologica 65: 419 - 435 DOI 10.1655 / 08 - 059.1." title="A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros" type="journal article" year="2009">
<emphasis italics="true" pageId="12" pageNumber="13">Köhler et al. (2009)</emphasis>
</bibRefCitation>
. While this placement needs to be re-analysed in light of the genetic information produced by 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[645,871,308,334]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="12" pageNumber="13" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[645,871,308,334]" italics="true" pageId="12" pageNumber="13">Lemme et al. (2013)</emphasis>
</bibRefCitation>
, 
<taxonomicName box="[885,1118,308,334]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[885,1118,308,334]" italics="true" pageId="12" pageNumber="13">
<taxonomicName box="[885,1002,308,334]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">Geckolepis</taxonomicName>
megalepis
</emphasis>
</taxonomicName>
can be distinguished easily from the type series of 
<taxonomicName box="[649,781,347,374]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="anomala">
<emphasis box="[649,781,347,374]" italics="true" pageId="12" pageNumber="13">G. anomala</emphasis>
</taxonomicName>
, 
<taxonomicName box="[796,922,347,374]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="modesta">
<emphasis box="[796,922,347,374]" italics="true" pageId="12" pageNumber="13">G. modesta</emphasis>
</taxonomicName>
, and 
<taxonomicName box="[987,1079,348,374]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="petiti">
<emphasis box="[987,1079,348,374]" italics="true" pageId="12" pageNumber="13">G. petiti</emphasis>
</taxonomicName>
by its postmental scales (condition A/B vs. D in 
<emphasis box="[586,859,387,414]" italics="true" pageId="12" pageNumber="13">
<taxonomicName box="[586,722,387,414]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="anomala">G. anomala</taxonomicName>
, 
<taxonomicName box="[733,859,387,414]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="modesta">G. modesta</taxonomicName>
</emphasis>
, and 
<taxonomicName box="[923,1015,388,414]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="petiti">
<emphasis box="[923,1015,388,414]" italics="true" pageId="12" pageNumber="13">G. petiti</emphasis>
</taxonomicName>
), and fewer scale rows at midbody (17–18 vs. 32 in 
<taxonomicName box="[558,690,427,454]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="anomala">
<emphasis box="[558,690,427,454]" italics="true" pageId="12" pageNumber="13">G. anomala</emphasis>
</taxonomicName>
, 22–25 in 
<taxonomicName box="[813,940,427,454]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="modesta">
<emphasis box="[813,940,427,454]" italics="true" pageId="12" pageNumber="13">G. modesta</emphasis>
</taxonomicName>
, and 28 in 
<taxonomicName box="[1072,1163,428,454]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="petiti">
<emphasis box="[1072,1163,428,454]" italics="true" pageId="12" pageNumber="13">G. petiti</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="30" lastPageNumber="31" pageId="12" pageNumber="13" type="discussion">
<paragraph blockId="12.[491,1542,228,773]" pageId="12" pageNumber="13">
<emphasis box="[523,621,467,493]" italics="true" pageId="12" pageNumber="13">Remarks</emphasis>
. The living specimen depicted in 
<figureCitation box="[1024,1090,467,493]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="12" pageNumber="13">Fig. 3</figureCitation>
of 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[1131,1357,467,493]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="12" pageNumber="13" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[1131,1357,467,493]" italics="true" pageId="12" pageNumber="13">Lemme et al. (2013)</emphasis>
</bibRefCitation>
as OTU D is misattributed and does not belong to 
<taxonomicName box="[945,1178,507,533]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[945,1178,507,533]" italics="true" pageId="12" pageNumber="13">
<taxonomicName box="[945,1062,507,533]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">Geckolepis</taxonomicName>
<taxonomicName box="[1069,1178,507,533]" pageId="12" pageNumber="13" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
. 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[1195,1421,507,533]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="12" pageNumber="13" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[1195,1421,507,533]" italics="true" pageId="12" pageNumber="13">Lemme et al. (2013)</emphasis>
</bibRefCitation>
report 17–20 scale rows at midbody for this species; the reason for this discrepancy could not be established here, but we are confident in our counts. However, we also emphasise that their higher number would remain diagnostic in all of the comparisons presented above. 
<bibRefCitation author="Kohler G &amp; Diethert H-H &amp; Nussbaum RA &amp; Raxworthy CJ." box="[491,711,666,692]" journalOrPublisher="Herpetologica" pageId="12" pageNumber="13" pagination="419 - 435" part="65" publicationUrl="10.1655/08-059.1." refString="Kohler G, Diethert H-H, Nussbaum RA, Raxworthy CJ. 2009. A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros. Herpetologica 65: 419 - 435 DOI 10.1655 / 08 - 059.1." title="A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros" type="journal article" year="2009">
<emphasis box="[491,711,666,692]" italics="true" pageId="12" pageNumber="13">Köhler et al. (2009)</emphasis>
</bibRefCitation>
probably did not include any specimens of 
<taxonomicName box="[1216,1449,666,692]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1216,1449,666,692]" italics="true" pageId="12" pageNumber="13">
Geckolepis 
<taxonomicName box="[1340,1449,666,692]" pageId="12" pageNumber="13" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
in their revision of the genus, as they did not consider any individuals with fewer than 22 scale rows at midbody.
</paragraph>
<paragraph blockId="12.[491,1542,806,1872]" box="[491,864,806,837]" pageId="12" pageNumber="13">
<heading bold="true" box="[491,864,806,837]" fontSize="12" level="2" pageId="12" pageNumber="13" reason="0">
<emphasis bold="true" box="[491,864,806,837]" pageId="12" pageNumber="13">
Osteology of 
<taxonomicName box="[695,864,806,837]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[695,864,806,837]" italics="true" pageId="12" pageNumber="13">Geckolepis</emphasis>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph blockId="12.[491,1542,806,1872]" pageId="12" pageNumber="13">
<emphasis box="[491,772,850,876]" italics="true" pageId="12" pageNumber="13">Osteological comparisons</emphasis>
. The scales of 
<taxonomicName box="[949,1066,850,876]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis box="[949,1066,850,876]" italics="true" pageId="12" pageNumber="13">Geckolepis</emphasis>
</taxonomicName>
geckos are mineralized and resemble osteoderms (
<figureCitation box="[646,713,889,916]" captionStart="Figure 5" captionStartId="8.[507,571,729,751]" captionTargetBox="[492,1541,229,684]" captionTargetId="figure@8.[491,1541,229,684]" captionTargetPageId="8" captionText="Figure 5 Micro-CT images of mineraliZed scales of Geckolepis specimens. Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of Geckolepis megalepis (A – C, ZSM 2126 / 2007), Geckolepis OTU AB (D – E, ZSM 1520 / 2008), G. maculata (F – G, ZMB 9655), and G. humbloti (H – I, ZSM 80 / 2010; J – K, ZSM 81 / 2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3)." httpUri="https://zenodo.org/record/291230/files/figure.png" pageId="12" pageNumber="13">Fig. 5</figureCitation>
; see also 
<bibRefCitation author="Schmidt WJ." box="[829,990,890,916]" editor="Voeltzkow A" journalOrPublisher="Stuttgart: Schweizerbart'sche Verlagsbuchhandlung" pageId="12" pageNumber="13" pagination="331 - 352" refString="Schmidt WJ. 1911. Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden. In: Voeltzkow A, ed. Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung, 331 - 352." title="Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden" type="book chapter" volumeTitle="Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow" year="1911">
<emphasis box="[829,990,890,916]" italics="true" pageId="12" pageNumber="13">Schmidt, 1911</emphasis>
</bibRefCitation>
). Among gekkotans, only 
<taxonomicName box="[1308,1445,890,916]" class="Reptilia" family="Gekkonidae" genus="Gekko" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="gecko">
<emphasis box="[1308,1445,890,916]" italics="true" pageId="12" pageNumber="13">Gekko gecko</emphasis>
</taxonomicName>
and 
<taxonomicName box="[491,602,929,955]" class="Reptilia" family="Phyllodactylidae" genus="Tarentola" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis box="[491,602,929,955]" italics="true" pageId="12" pageNumber="13">Tarentola</emphasis>
</taxonomicName>
species (
<bibRefCitation author="Bauer AM &amp; Russell AP." box="[710,958,929,956]" journalOrPublisher="Canadian Journal of Zoology" pageId="12" pageNumber="13" pagination="678 - 684" part="67" publicationUrl="10.1139/z89-098." refString="Bauer AM, Russell AP. 1989. Supraorbital ossifications in geckos (Reptilia: Gekkonidae). Canadian Journal of Zoology 67: 678 - 684 DOI 10.1139 / z 89 - 098." title="Supraorbital ossifications in geckos (Reptilia: Gekkonidae)" type="journal article" year="1989">
<emphasis box="[710,958,929,956]" italics="true" pageId="12" pageNumber="13">Bauer &amp; Russell, 1989</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="DaZa JD &amp; Bauer AM." box="[973,1203,930,956]" editor="Bininda-Emonds ORP" journalOrPublisher="Oldenburg: BIS-Verlag" pageId="12" pageNumber="13" pagination="399 - 405" refString="DaZa JD, Bauer AM. 2015. Cranial anatomy of the pygopodid lizard Aprasia repens, a gekkotan masquerading as a Scolecophidian. In: Bininda-Emonds ORP, Powell GL, Jamniczky HA, Bauer AM, Theodor J, eds. All animals are interesting: a Festschrift in honour of Anthony P Russell. Oldenburg: BIS-Verlag, 399 - 405." title="Cranial anatomy of the pygopodid lizard Aprasia repens, a gekkotan masquerading as a Scolecophidian" type="book chapter" volumeTitle="All animals are interesting: a Festschrift in honour of Anthony P Russell" year="2015">
<emphasis box="[973,1203,930,956]" italics="true" pageId="12" pageNumber="13">Daza &amp; Bauer, 2015</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Schmidt WJ." box="[1218,1380,929,956]" editor="Voeltzkow A" journalOrPublisher="Stuttgart: Schweizerbart'sche Verlagsbuchhandlung" pageId="12" pageNumber="13" pagination="331 - 352" refString="Schmidt WJ. 1911. Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden. In: Voeltzkow A, ed. Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung, 331 - 352." title="Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden" type="book chapter" volumeTitle="Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow" year="1911">
<emphasis box="[1218,1380,929,956]" italics="true" pageId="12" pageNumber="13">Schmidt, 1911</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Vickaryous MK &amp; Meldrum G &amp; Russell AP." journalOrPublisher="Journal of Morphology" pageId="12" pageNumber="13" pagination="1345 - 1357" part="276" publicationUrl="10.1002/jmor.20422." refString="Vickaryous MK, Meldrum G, Russell AP. 2015. Armored geckos: a histological investigation of osteoderm development in Tarentola (Phyllodactylidae) and Gekko (Gekkonidae) with comments on their regeneration and inferred function. Journal of Morphology 276: 1345 - 1357 DOI 10.1002 / jmor. 20422." title="Armored geckos: a histological investigation of osteoderm development in Tarentola (Phyllodactylidae) and Gekko (Gekkonidae) with comments on their regeneration and inferred function" type="journal article" year="2015">
<emphasis italics="true" pageId="12" pageNumber="13">Vickaryous, Meldrum &amp; Russell, 2015</emphasis>
</bibRefCitation>
) are known to develop similarly mineralised integumentary coverings. The dense scales of 
<taxonomicName box="[858,975,1009,1035]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis box="[858,975,1009,1035]" italics="true" pageId="12" pageNumber="13">Geckolepis</emphasis>
</taxonomicName>
differ from the osteoderms of 
<taxonomicName box="[1349,1443,1009,1036]" class="Reptilia" family="Gekkonidae" genus="Gekko" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="gecko">
<emphasis box="[1349,1443,1009,1036]" italics="true" pageId="12" pageNumber="13">G. gecko</emphasis>
</taxonomicName>
and 
<taxonomicName box="[491,663,1050,1075]" class="Reptilia" family="Phyllodactylidae" genus="Tarentola" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="mauritanica">
<emphasis box="[491,663,1050,1075]" italics="true" pageId="12" pageNumber="13">T. mauritanica</emphasis>
</taxonomicName>
in that they are imbricate, and not juxtaposed and adpressed against the skull bones. 
<taxonomicName box="[631,748,1089,1115]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis box="[631,748,1089,1115]" italics="true" pageId="12" pageNumber="13">Geckolepis</emphasis>
</taxonomicName>
also differ from the majority of extant gekkotans in having paired and unfused (both dorsally and ventrally) frontal bones; we only found fused frontal bones in a large specimen of the OTU AB sensu 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[919,1145,1169,1195]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="12" pageNumber="13" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[919,1145,1169,1195]" italics="true" pageId="12" pageNumber="13">Lemme et al. (2013)</emphasis>
</bibRefCitation>
from Montagne des Français (‘AB specimen’ henceforth), which is the most osteologically distinct specimen from our sample.
</paragraph>
<paragraph blockId="12.[491,1542,806,1872]" pageId="12" pageNumber="13">
<taxonomicName box="[523,756,1248,1274]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[523,756,1248,1274]" italics="true" pageId="12" pageNumber="13">Geckolepis megalepis</emphasis>
</taxonomicName>
and the AB specimen differ from other 
<taxonomicName box="[1205,1322,1248,1274]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis box="[1205,1322,1248,1274]" italics="true" pageId="12" pageNumber="13">Geckolepis</emphasis>
</taxonomicName>
in having a narrow infraorbital fenestra. In these two taxa, the nasal cavity also bulges slightly more than in smaller 
<taxonomicName box="[582,699,1328,1354]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis box="[582,699,1328,1354]" italics="true" pageId="12" pageNumber="13">Geckolepis</emphasis>
</taxonomicName>
specimens. There is some variation in the shape of the nasal bones, being rectangular (with straight sides) in 
<taxonomicName box="[901,1044,1368,1394]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[901,1044,1368,1394]" italics="true" pageId="12" pageNumber="13">G. megalepis</emphasis>
</taxonomicName>
and the AB specimen. 
<taxonomicName box="[1314,1541,1368,1394]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1314,1541,1368,1394]" italics="true" pageId="12" pageNumber="13">Geckolepis humbloti</emphasis>
</taxonomicName>
has nasal bones with a sigmoid lateral edge instead of straight. The holotype of 
<taxonomicName box="[1402,1542,1408,1434]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1402,1542,1408,1434]" italics="true" pageId="12" pageNumber="13">G. maculata</emphasis>
</taxonomicName>
has nasals with straight lateral edges (
<figureCitation box="[933,1012,1448,1474]" captionStart="Figure 10" captionStartId="18.[507,571,1747,1769]" captionTargetBox="[492,1509,230,1692]" captionTargetId="figure@18.[491,1512,229,1702]" captionTargetPageId="18" captionText="Figure 10 Comparative micro-CT images of the skull (cranium and jaw) of other Geckolepis species. Shown in dorsal (A, F, K, P), ventral (B, G, L, Q), lateral (C, H, M, R), labial (D, I, N, S), and lingual (E, J, O, T) view. Depicting Geckolepis OTU AB (A – E, ZSM 1520 / 2008), holotype of G. maculata (F – J, ZMB 9655), and G. humbloti (K – O, ZSM 80 / 2010; P – T, ZSM 81 / 2006). From volume-rendering of micro-CT scans. Rotational videos of these skulls are provided in Videos S 2 – S 5. For labels, see Fig. 6." httpUri="https://zenodo.org/record/291235/files/figure.png" pageId="12" pageNumber="13">Fig. 10</figureCitation>
). 
<taxonomicName box="[1038,1271,1448,1474]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1038,1271,1448,1474]" italics="true" pageId="12" pageNumber="13">Geckolepis megalepis</emphasis>
</taxonomicName>
and the AB specimen have a more anterior extension of the subfrontal process of the frontal in palatal view, fused in the AB specimen and not fused in 
<taxonomicName box="[923,1066,1527,1554]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[923,1066,1527,1554]" italics="true" pageId="12" pageNumber="13">G. megalepis</emphasis>
</taxonomicName>
; all others have a large notched area that does not extend anteriorly. Another distinct feature of 
<taxonomicName box="[1126,1268,1567,1594]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1126,1268,1567,1594]" italics="true" pageId="12" pageNumber="13">G. megalepis</emphasis>
</taxonomicName>
was found in the shape of the premaxilla-vomer fenestra, being notched instead of rounded as in other 
<taxonomicName box="[1417,1534,1607,1633]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis box="[1417,1534,1607,1633]" italics="true" pageId="12" pageNumber="13">Geckolepis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="12.[491,1542,806,1872]" lastBlockId="14.[491,1542,697,1838]" lastPageId="14" lastPageNumber="15" pageId="12" pageNumber="13">
There are also some differences between 
<taxonomicName box="[1004,1146,1647,1673]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1004,1146,1647,1673]" italics="true" pageId="12" pageNumber="13">G. megalepis</emphasis>
</taxonomicName>
and the other species examined, in the postcranium, the lateral processes of the first five caudal vertebrae (pygial series) are curved laterally (vs. straight in 
<taxonomicName box="[901,1040,1726,1753]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[901,1040,1726,1753]" italics="true" pageId="12" pageNumber="13">G. maculata</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1100,1237,1726,1753]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1100,1237,1726,1753]" italics="true" pageId="12" pageNumber="13">G. humbloti</emphasis>
</taxonomicName>
); the scapular ray of the scapulocoracoid does not surpass the clavicle (vs. surpassing the clavicle in 
<taxonomicName box="[1375,1515,1766,1793]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1375,1515,1766,1793]" italics="true" pageId="12" pageNumber="13">G. maculata</emphasis>
</taxonomicName>
); the secondary coracoid ray of the scapulocoracoid extends to the level of the posterior margin of the clavicular fenestra (vs. surpassing the posterior margin of the fenestra in 
<taxonomicName box="[491,631,697,723]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[491,631,697,723]" italics="true" pageId="14" pageNumber="15">G. maculata</emphasis>
</taxonomicName>
); and the pubic tubercle of the pubis is posteriorly curved (vs. more or less vertical in 
<taxonomicName box="[615,754,736,763]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[615,754,736,763]" italics="true" pageId="14" pageNumber="15">G. maculata</emphasis>
</taxonomicName>
and 
<taxonomicName box="[812,949,736,763]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[812,949,736,763]" italics="true" pageId="14" pageNumber="15">G. humbloti</emphasis>
</taxonomicName>
).
</paragraph>
<caption httpUri="https://zenodo.org/record/291231/files/figure.png" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14" targetBox="[491,1188,229,1668]" targetPageId="13">
<paragraph blockId="13.[507,1519,1713,1852]" pageId="13" pageNumber="14">
<emphasis bold="true" pageId="13" pageNumber="14">
Figure 6 Micro-CT images of skull (cranium and jaw) of the holotype of 
<taxonomicName box="[1228,1435,1713,1735]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[1228,1435,1713,1735]" italics="true" pageId="13" pageNumber="14">Geckolepis megalepis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1439,1515,1714,1735]" pageId="13" pageNumber="14" rank="species">sp. nov.</taxonomicNameLabel>
(ZSM 2126/2007).
</emphasis>
(A) Dorsal, (B) ventral, (C) lateral, (D) labial, and (E) lingual view. Abbreviations: 
<emphasis box="[1483,1513,1743,1764]" italics="true" pageId="13" pageNumber="14">adf</emphasis>
, anterior inferior dental foramen; 
<emphasis box="[826,862,1772,1793]" italics="true" pageId="13" pageNumber="14">amf</emphasis>
, anterior mylohyoid foramen; 
<emphasis box="[1158,1178,1773,1794]" italics="true" pageId="13" pageNumber="14">ar</emphasis>
, articular facet; 
<emphasis box="[1331,1374,1773,1794]" italics="true" pageId="13" pageNumber="14">asnp</emphasis>
, ascending nasal process of premaxilla; 
<emphasis box="[775,797,1801,1822]" italics="true" pageId="13" pageNumber="14">bo</emphasis>
, basioccipital; 
<emphasis box="[939,962,1801,1822]" italics="true" pageId="13" pageNumber="14">bp</emphasis>
, basipterygoid process of parabasisphenoid; 
<emphasis box="[1388,1409,1801,1822]" italics="true" pageId="13" pageNumber="14">ch</emphasis>
, choana; 
<emphasis box="[507,538,1830,1851]" italics="true" pageId="13" pageNumber="14">cob</emphasis>
, compound bone; 
<emphasis box="[719,747,1831,1852]" italics="true" pageId="13" pageNumber="14">cor</emphasis>
, coronoid; 
<emphasis box="[857,869,1830,1851]" italics="true" pageId="13" pageNumber="14">d</emphasis>
, dentary; 
<emphasis box="[965,1000,1830,1851]" italics="true" pageId="13" pageNumber="14">dpp</emphasis>
, (continued on next page...)
</paragraph>
<caption pageId="14" pageNumber="15">
<paragraph blockId="14.[507,1522,241,643]" box="[507,735,241,263]" pageId="14" pageNumber="15">Figure 6 (... continued)</paragraph>
<paragraph blockId="14.[507,1522,241,643]" pageId="14" pageNumber="15">
descending parietal process; 
<emphasis box="[779,804,271,292]" italics="true" pageId="14" pageNumber="15">ect</emphasis>
, ectopterygoid; 
<emphasis box="[958,986,271,292]" italics="true" pageId="14" pageNumber="15">ept</emphasis>
, epipterygoid; 
<emphasis box="[1129,1135,270,291]" italics="true" pageId="14" pageNumber="15">f</emphasis>
, frontal; 
<emphasis box="[1223,1260,270,291]" italics="true" pageId="14" pageNumber="15">hscc</emphasis>
, horizontal semi-circular canal; 
<emphasis box="[568,581,299,320]" italics="true" pageId="14" pageNumber="15">if</emphasis>
, incisive foramen; 
<emphasis box="[762,768,300,321]" italics="true" pageId="14" pageNumber="15">j</emphasis>
, jugal; 
<emphasis box="[837,877,299,320]" italics="true" pageId="14" pageNumber="15">larst</emphasis>
, lateral aperture of the recessus scalae tympani; 
<emphasis box="[1335,1372,299,320]" italics="true" pageId="14" pageNumber="15">mdf</emphasis>
, mandibular fossa; 
<emphasis box="[565,590,329,350]" italics="true" pageId="14" pageNumber="15">mf</emphasis>
, mental foramen; 
<emphasis box="[766,806,329,350]" italics="true" pageId="14" pageNumber="15">mko</emphasis>
, opening of the Meckelian canal, 
<emphasis box="[1128,1158,329,350]" italics="true" pageId="14" pageNumber="15">mx</emphasis>
, maxilla; 
<emphasis box="[1250,1308,329,350]" italics="true" pageId="14" pageNumber="15">mx.ap</emphasis>
, anterior process of maxilla; 
<emphasis box="[588,642,358,379]" italics="true" pageId="14" pageNumber="15">mx.fp</emphasis>
, facial process of maxilla; 
<emphasis box="[891,950,359,380]" italics="true" pageId="14" pageNumber="15">mx.pp</emphasis>
, posterior process of maxilla; 
<emphasis box="[1236,1291,359,380]" italics="true" pageId="14" pageNumber="15">mx.ps</emphasis>
, palatal shelf of maxilla; 
<emphasis box="[507,519,388,409]" italics="true" pageId="14" pageNumber="15">n</emphasis>
, nasal; 
<emphasis box="[590,618,388,409]" italics="true" pageId="14" pageNumber="15">occ</emphasis>
, occipital condyle; 
<emphasis box="[803,832,388,409]" italics="true" pageId="14" pageNumber="15">oto</emphasis>
, otooccipital; 
<emphasis box="[967,996,387,408]" italics="true" pageId="14" pageNumber="15">pal</emphasis>
, palatine; 
<emphasis box="[1095,1159,387,408]" italics="true" pageId="14" pageNumber="15">pal.mp</emphasis>
, maxillary process of palatine; 
<emphasis box="[1455,1506,387,408]" italics="true" pageId="14" pageNumber="15">pal.vf</emphasis>
, vomerine flange of palatine; 
<emphasis box="[779,810,417,438]" italics="true" pageId="14" pageNumber="15">par</emphasis>
, parietal; 
<emphasis box="[905,948,416,437]" italics="true" pageId="14" pageNumber="15">pbsh</emphasis>
, parabasisphenoid; 
<emphasis box="[1138,1179,417,438]" italics="true" pageId="14" pageNumber="15">pmx</emphasis>
, premaxilla; 
<emphasis box="[1304,1381,417,438]" italics="true" pageId="14" pageNumber="15">pmx.pps</emphasis>
, premaxilla palatal shelf; 
<emphasis box="[631,660,446,467]" italics="true" pageId="14" pageNumber="15">pof</emphasis>
, postorbitofrontal; 
<emphasis box="[848,882,446,467]" italics="true" pageId="14" pageNumber="15">pop</emphasis>
, paroccipital process; 
<emphasis box="[1094,1129,446,467]" italics="true" pageId="14" pageNumber="15">ppp</emphasis>
, posterolateral process of parietal; 
<emphasis box="[1461,1487,446,467]" italics="true" pageId="14" pageNumber="15">prf</emphasis>
, prefrontal; 
<emphasis box="[615,646,475,496]" italics="true" pageId="14" pageNumber="15">pro</emphasis>
, prootic; 
<emphasis box="[738,795,475,496]" italics="true" pageId="14" pageNumber="15">pro.ca</emphasis>
, crista alaris of prootic; 
<emphasis box="[1025,1082,475,496]" italics="true" pageId="14" pageNumber="15">pro.cp</emphasis>
, crista prootica of prootic; 
<emphasis box="[1341,1378,475,496]" italics="true" pageId="14" pageNumber="15">psaf</emphasis>
, posterior surangular foramen; 
<emphasis box="[708,744,505,526]" italics="true" pageId="14" pageNumber="15">pscc</emphasis>
, posterior semicircular canal; 
<emphasis box="[1033,1052,505,526]" italics="true" pageId="14" pageNumber="15">pt</emphasis>
, pterygoid; 
<emphasis box="[1166,1213,505,526]" italics="true" pageId="14" pageNumber="15">pt.qp</emphasis>
, quadrate process of pterygoid; 
<emphasis box="[507,518,534,555]" italics="true" pageId="14" pageNumber="15">q</emphasis>
, quadrate; 
<emphasis box="[626,664,533,554]" italics="true" pageId="14" pageNumber="15">q.ch</emphasis>
, conch of quadrate; 
<emphasis box="[859,890,534,555]" italics="true" pageId="14" pageNumber="15">rap</emphasis>
, retroarticular process; 
<emphasis box="[1117,1125,534,555]" italics="true" pageId="14" pageNumber="15">s</emphasis>
, stapes; 
<emphasis box="[1205,1230,533,554]" italics="true" pageId="14" pageNumber="15">saf</emphasis>
, surangular foramen; 
<emphasis box="[1442,1467,534,555]" italics="true" pageId="14" pageNumber="15">scr</emphasis>
, sclerotic ring; 
<emphasis box="[608,645,563,584]" italics="true" pageId="14" pageNumber="15">smx</emphasis>
, septomaxilla; 
<emphasis box="[788,813,562,583]" italics="true" pageId="14" pageNumber="15">spl</emphasis>
, splenial; 
<emphasis box="[909,934,562,583]" italics="true" pageId="14" pageNumber="15">sof</emphasis>
, suborbital fenestra; 
<emphasis box="[1135,1154,563,584]" italics="true" pageId="14" pageNumber="15">sq</emphasis>
, squamosal, 
<emphasis box="[1277,1295,563,584]" italics="true" pageId="14" pageNumber="15">so</emphasis>
, supraoccipital; 
<emphasis box="[1453,1491,562,583]" italics="true" pageId="14" pageNumber="15">spht</emphasis>
, sphenooccipital tubercle; 
<emphasis box="[752,779,592,613]" italics="true" pageId="14" pageNumber="15">tbr</emphasis>
, trabeculae; 
<emphasis box="[900,910,592,613]" italics="true" pageId="14" pageNumber="15">v</emphasis>
, vomer; 
<emphasis box="[994,1028,592,613]" italics="true" pageId="14" pageNumber="15">v.lp</emphasis>
, lateral process of vomer; 
<emphasis box="[1278,1324,592,613]" italics="true" pageId="14" pageNumber="15">v.mp</emphasis>
, maxillary process of vomer. A rotational video of the skull is provided in Video S1.
</paragraph>
</caption>
</caption>
<paragraph blockId="14.[491,1542,697,1838]" pageId="14" pageNumber="15">
For future reference we listed 24 variable skull traits (Appendix S2) among 
<taxonomicName box="[1411,1528,776,802]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis box="[1411,1528,776,802]" italics="true" pageId="14" pageNumber="15">Geckolepis</emphasis>
</taxonomicName>
that provide a baseline for future comparative studies.
</paragraph>
<paragraph blockId="14.[491,1542,697,1838]" pageId="14" pageNumber="15">
<emphasis box="[523,738,856,882]" italics="true" pageId="14" pageNumber="15">Skeletal description</emphasis>
. In the following section, we present a generalised skeletal description of the genus 
<taxonomicName box="[636,753,896,922]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis box="[636,753,896,922]" italics="true" pageId="14" pageNumber="15">Geckolepis</emphasis>
</taxonomicName>
. Data on 
<taxonomicName box="[864,1000,896,922]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[864,1000,896,922]" italics="true" pageId="14" pageNumber="15">G. humbloti</emphasis>
</taxonomicName>
is based on the scans produced for 
<bibRefCitation author="Hawlitschek O &amp; Glaw F." journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="14" pageNumber="15" pagination="289 - 298" part="16" publicationUrl="10.1007/s13127-015-0255-1." refString="Hawlitschek O, ScherZ MD, Straube N, Glaw F. 2016. Resurrection of the Comoran fish scale gecko Geckolepis humbloti Vaillant, 1887 reveals a disjointed distribution caused by natural overseas dispersal. Organisms Diversity &amp; Evolution 16: 289 - 298 DOI 10.1007 / s 13127 - 015 - 0255 - 1." title="Resurrection of the Comoran fish scale gecko Geckolepis humbloti Vaillant, 1887 reveals a disjointed distribution caused by natural overseas dispersal" type="journal article" year="2016">
<emphasis italics="true" pageId="14" pageNumber="15">Hawlitschek et al. (2016)</emphasis>
</bibRefCitation>
, re-analysed for this study. The postcranial skeleton of the AB specimen was not assessed; our postcranial osteological description pertains only to 
<taxonomicName box="[1246,1382,976,1002]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1246,1382,976,1002]" italics="true" pageId="14" pageNumber="15">G. humbloti</emphasis>
</taxonomicName>
, 
<taxonomicName box="[1395,1535,976,1002]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1395,1535,976,1002]" italics="true" pageId="14" pageNumber="15">G. maculata</emphasis>
</taxonomicName>
, and 
<taxonomicName box="[543,685,1015,1042]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[543,685,1015,1042]" italics="true" pageId="14" pageNumber="15">G. megalepis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="14.[491,1542,697,1838]" pageId="14" pageNumber="15">
As we have mentioned above, one key feature of 
<taxonomicName box="[1101,1218,1055,1081]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis box="[1101,1218,1055,1081]" italics="true" pageId="14" pageNumber="15">Geckolepis</emphasis>
</taxonomicName>
is the presence of a dense covering of mineralized scales, (
<figureCitation box="[858,920,1095,1122]" captionStart="Figure 5" captionStartId="8.[507,571,729,751]" captionTargetBox="[492,1541,229,684]" captionTargetId="figure@8.[491,1541,229,684]" captionTargetPageId="8" captionText="Figure 5 Micro-CT images of mineraliZed scales of Geckolepis specimens. Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of Geckolepis megalepis (A – C, ZSM 2126 / 2007), Geckolepis OTU AB (D – E, ZSM 1520 / 2008), G. maculata (F – G, ZMB 9655), and G. humbloti (H – I, ZSM 80 / 2010; J – K, ZSM 81 / 2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3)." httpUri="https://zenodo.org/record/291230/files/figure.png" pageId="14" pageNumber="15">Fig. 5</figureCitation>
). These mineralized scales contrary to the osteoderms of 
<taxonomicName box="[525,662,1135,1161]" class="Reptilia" family="Gekkonidae" genus="Gekko" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="gecko">
<emphasis box="[525,662,1135,1161]" italics="true" pageId="14" pageNumber="15">Gekko gecko</emphasis>
</taxonomicName>
and 
<taxonomicName authority="Vickaryous, Meldrum &amp; Russell, 2015" authorityName="Vickaryous, Meldrum &amp; Russell" authorityYear="2015" box="[727,1441,1135,1162]" class="Reptilia" family="Phyllodactylidae" genus="Tarentola" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="mauritanica">
<taxonomicName box="[727,984,1135,1161]" class="Reptilia" family="Phyllodactylidae" genus="Tarentola" higherTaxonomySource="CoL" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="mauritanica">
<emphasis box="[727,984,1135,1161]" italics="true" pageId="14" pageNumber="15">Tarentola mauritanica</emphasis>
</taxonomicName>
(
<bibRefCitation author="Vickaryous MK &amp; Meldrum G &amp; Russell AP." box="[1005,1431,1135,1161]" journalOrPublisher="Journal of Morphology" pageId="14" pageNumber="15" pagination="1345 - 1357" part="276" publicationUrl="10.1002/jmor.20422." refString="Vickaryous MK, Meldrum G, Russell AP. 2015. Armored geckos: a histological investigation of osteoderm development in Tarentola (Phyllodactylidae) and Gekko (Gekkonidae) with comments on their regeneration and inferred function. Journal of Morphology 276: 1345 - 1357 DOI 10.1002 / jmor. 20422." title="Armored geckos: a histological investigation of osteoderm development in Tarentola (Phyllodactylidae) and Gekko (Gekkonidae) with comments on their regeneration and inferred function" type="journal article" year="2015">
<emphasis box="[1005,1431,1135,1161]" italics="true" pageId="14" pageNumber="15">Vickaryous, Meldrum &amp; Russell, 2015</emphasis>
</bibRefCitation>
)
</taxonomicName>
are imbricate and not adpressed against the skull. 
<bibRefCitation author="Schmidt WJ." box="[1047,1223,1175,1201]" editor="Voeltzkow A" journalOrPublisher="Stuttgart: Schweizerbart'sche Verlagsbuchhandlung" pageId="14" pageNumber="15" pagination="331 - 352" refString="Schmidt WJ. 1911. Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden. In: Voeltzkow A, ed. Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung, 331 - 352." title="Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden" type="book chapter" volumeTitle="Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow" year="1911">
<emphasis box="[1047,1223,1175,1201]" italics="true" pageId="14" pageNumber="15">Schmidt (1911)</emphasis>
</bibRefCitation>
referred to these scales as osteoderm, but noted that they are unique in lacking bone cells (that is to say, they are not osteoderm in the strict sense). He found that these mineralizations, which he showed to be formed from calcified tissue fibres, roughly trace the outline of the scales, but are mostly confined to their middles and do not extend into the keratinous scale. Our micro-CT data indicates that mineralization of scales is more extensive, at least in the examined species; scales shown in 
<figureCitation box="[682,747,1414,1441]" captionStart="Figure 5" captionStartId="8.[507,571,729,751]" captionTargetBox="[492,1541,229,684]" captionTargetId="figure@8.[491,1541,229,684]" captionTargetPageId="8" captionText="Figure 5 Micro-CT images of mineraliZed scales of Geckolepis specimens. Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of Geckolepis megalepis (A – C, ZSM 2126 / 2007), Geckolepis OTU AB (D – E, ZSM 1520 / 2008), G. maculata (F – G, ZMB 9655), and G. humbloti (H – I, ZSM 80 / 2010; J – K, ZSM 81 / 2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3)." httpUri="https://zenodo.org/record/291230/files/figure.png" pageId="14" pageNumber="15">Fig. 5</figureCitation>
are very similar in dimensions to what they look like in life (e.g., compare 
<figureCitation box="[601,664,1454,1481]" captionStart="Figure 5" captionStartId="8.[507,571,729,751]" captionTargetBox="[492,1541,229,684]" captionTargetId="figure@8.[491,1541,229,684]" captionTargetPageId="8" captionText="Figure 5 Micro-CT images of mineraliZed scales of Geckolepis specimens. Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of Geckolepis megalepis (A – C, ZSM 2126 / 2007), Geckolepis OTU AB (D – E, ZSM 1520 / 2008), G. maculata (F – G, ZMB 9655), and G. humbloti (H – I, ZSM 80 / 2010; J – K, ZSM 81 / 2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3)." httpUri="https://zenodo.org/record/291230/files/figure.png" pageId="14" pageNumber="15">Fig. 5</figureCitation>
A with ZSM 2126/2007 in 
<figureCitation box="[977,1041,1454,1480]" captionStart="Figure 4" captionStartId="7.[507,571,1752,1774]" captionTargetBox="[491,1512,230,1704]" captionTargetId="figure@7.[491,1514,229,1708]" captionTargetPageId="7" captionText="Figure 4 Specimens of Geckolepis megalepis sp. nov. in preservative. (A – C) The holotype, ZSM 2126 / 2007; (D – F) paratype ZSM 232 / 2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale." httpUri="https://zenodo.org/record/291229/files/figure.png" pageId="14" pageNumber="15">Fig. 4</figureCitation>
). The reasons for this discrepancy are not clear, and will require further study. However, we can confirm that these mineralizations do not extend to the tip of the scales, as can be seen by the soft-looking distal edges of the scales in 
<figureCitation box="[597,662,1573,1600]" captionStart="Figure 5" captionStartId="8.[507,571,729,751]" captionTargetBox="[492,1541,229,684]" captionTargetId="figure@8.[491,1541,229,684]" captionTargetPageId="8" captionText="Figure 5 Micro-CT images of mineraliZed scales of Geckolepis specimens. Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of Geckolepis megalepis (A – C, ZSM 2126 / 2007), Geckolepis OTU AB (D – E, ZSM 1520 / 2008), G. maculata (F – G, ZMB 9655), and G. humbloti (H – I, ZSM 80 / 2010; J – K, ZSM 81 / 2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3)." httpUri="https://zenodo.org/record/291230/files/figure.png" pageId="14" pageNumber="15">Fig. 5</figureCitation>
. 
<bibRefCitation author="Schmidt WJ." box="[677,853,1573,1600]" editor="Voeltzkow A" journalOrPublisher="Stuttgart: Schweizerbart'sche Verlagsbuchhandlung" pageId="14" pageNumber="15" pagination="331 - 352" refString="Schmidt WJ. 1911. Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden. In: Voeltzkow A, ed. Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung, 331 - 352." title="Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden" type="book chapter" volumeTitle="Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow" year="1911">
<emphasis box="[677,853,1573,1600]" italics="true" pageId="14" pageNumber="15">Schmidt (1911)</emphasis>
</bibRefCitation>
also found that mineralization was lacking from specific scales, including the labials and postmentals and scales proximal to these, small scales of the head including those around the eye and ears, and fingers and toes; they also decrease in frequency in the tail scales beyond the first third. This pattern is recapitulated by our micro-CT scans. In a new study by Paluh and colleagues (in review), these mineralized scales are again suggested to be osteoderms, but we conservatively refer to them here as ‘mineralized scales’ until their study is published.
</paragraph>
<caption httpUri="https://zenodo.org/record/291232/files/figure.png" pageId="15" pageNumber="16" targetBox="[491,1381,229,1668]" targetPageId="15">
<paragraph blockId="15.[507,1522,1713,1794]" pageId="15" pageNumber="16">
<emphasis bold="true" pageId="15" pageNumber="16">
Figure 7 Micro-CT images of spinal column of 
<taxonomicName box="[980,1084,1713,1735]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[980,1084,1713,1735]" italics="true" pageId="15" pageNumber="16">Geckolepis</emphasis>
</taxonomicName>
, based on 
<taxonomicName box="[1187,1312,1713,1735]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[1187,1312,1713,1735]" italics="true" pageId="15" pageNumber="16">G. megalepis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1316,1392,1714,1735]" pageId="15" pageNumber="16" rank="species">sp. nov.</taxonomicNameLabel>
(ZSM 2126/2007).
</emphasis>
(A) Lateral and (B) dorsal view. Abbreviations: 
<emphasis box="[1085,1102,1743,1764]" italics="true" pageId="15" pageNumber="16">lb</emphasis>
, lumbar; 
<emphasis box="[1195,1211,1743,1764]" italics="true" pageId="15" pageNumber="16">sc</emphasis>
, sacral. Regions are indicated by the posterodorsal-most point of the neural arches.
</paragraph>
</caption>
<caption httpUri="https://zenodo.org/record/291233/files/figure.png" pageId="16" pageNumber="17" targetBox="[491,1541,229,1307]" targetPageId="16">
<paragraph blockId="16.[507,1497,1351,1608]" pageId="16" pageNumber="17">
<emphasis bold="true" pageId="16" pageNumber="17">
Figure 8 Micro-CT images of pectoral girdle of 
<taxonomicName box="[983,1087,1351,1373]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[983,1087,1351,1373]" italics="true" pageId="16" pageNumber="17">Geckolepis</emphasis>
</taxonomicName>
, based on 
<taxonomicName box="[1190,1314,1351,1373]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[1190,1314,1351,1373]" italics="true" pageId="16" pageNumber="17">G. megalepis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1319,1395,1352,1373]" pageId="16" pageNumber="17" rank="species">sp. nov.</taxonomicNameLabel>
(ZSM 2126/2007).
</emphasis>
(A) Ventral and (B) lateral view. Abbreviations: 
<emphasis box="[1092,1118,1381,1402]" italics="true" pageId="16" pageNumber="17">1cf</emphasis>
, primary coracoid fenestra; 
<emphasis box="[1387,1413,1381,1402]" italics="true" pageId="16" pageNumber="17">2cf</emphasis>
,
</paragraph>
<paragraph blockId="16.[507,1497,1351,1608]" pageId="16" pageNumber="17">
secondary coracoid fenestra; 
<emphasis box="[784,812,1411,1432]" italics="true" pageId="16" pageNumber="17">acr</emphasis>
, acromion process; 
<emphasis box="[1004,1019,1410,1431]" italics="true" pageId="16" pageNumber="17">cf</emphasis>
, clavicular fenestra; 
<emphasis box="[1214,1228,1410,1431]" italics="true" pageId="16" pageNumber="17">cl</emphasis>
, clavicle; 
<emphasis box="[1321,1349,1411,1432]" italics="true" pageId="16" pageNumber="17">cor</emphasis>
, coracoid; 
<emphasis box="[507,522,1440,1461]" italics="true" pageId="16" pageNumber="17">ic</emphasis>
, interclavicle; 
<emphasis box="[660,676,1440,1461]" italics="true" pageId="16" pageNumber="17">gf</emphasis>
, glenoid fossa; 
<emphasis box="[822,847,1440,1461]" italics="true" pageId="16" pageNumber="17">hu</emphasis>
, humerus; 
<emphasis box="[955,994,1440,1461]" italics="true" pageId="16" pageNumber="17">hu.c</emphasis>
, humeral condyle; 
<emphasis box="[1177,1238,1440,1461]" italics="true" pageId="16" pageNumber="17">hu.dtp</emphasis>
, deltopectoral crest of humerus; 
<emphasis box="[604,652,1469,1490]" italics="true" pageId="16" pageNumber="17">hu.ec</emphasis>
, ectepicondylar crest of humerus; 
<emphasis box="[981,1033,1469,1490]" italics="true" pageId="16" pageNumber="17">hu.hc</emphasis>
, humeral crest of humerus; 
<emphasis box="[1301,1328,1470,1491]" italics="true" pageId="16" pageNumber="17">mc</emphasis>
, metacarpal; 
<emphasis box="[507,533,1499,1520]" italics="true" pageId="16" pageNumber="17">ms</emphasis>
, mesosternum; 
<emphasis box="[687,699,1499,1520]" italics="true" pageId="16" pageNumber="17">p</emphasis>
, phalanges; 
<emphasis box="[815,834,1499,1520]" italics="true" pageId="16" pageNumber="17">ps</emphasis>
, presternum; 
<emphasis box="[969,993,1499,1520]" italics="true" pageId="16" pageNumber="17">pu</emphasis>
, patella ulnaris; 
<emphasis box="[1150,1158,1499,1520]" italics="true" pageId="16" pageNumber="17">r</emphasis>
, radius; 
<emphasis box="[1241,1255,1498,1519]" italics="true" pageId="16" pageNumber="17">rl</emphasis>
, radiale; 
<emphasis box="[1342,1350,1499,1520]" italics="true" pageId="16" pageNumber="17">s</emphasis>
, scapula; 
<emphasis box="[507,549,1527,1548]" italics="true" pageId="16" pageNumber="17">sc.1c</emphasis>
, primary coracoid ray of scapulocoracoid; 
<emphasis box="[958,1001,1527,1548]" italics="true" pageId="16" pageNumber="17">sc.2c</emphasis>
, secondary coracoid ray of scapulocoracoid; 
<emphasis box="[1429,1452,1527,1548]" italics="true" pageId="16" pageNumber="17">scf</emphasis>
, scapulocoracoid fenestra; 
<emphasis box="[755,769,1557,1578]" italics="true" pageId="16" pageNumber="17">sf</emphasis>
, scapular fenestra; 
<emphasis box="[952,967,1557,1578]" italics="true" pageId="16" pageNumber="17">ss</emphasis>
, suprascapula; 
<emphasis box="[1113,1151,1557,1578]" italics="true" pageId="16" pageNumber="17">sc.sr</emphasis>
, scapular ray of scapulocoracoid; 
<emphasis box="[1475,1491,1557,1578]" italics="true" pageId="16" pageNumber="17">sr</emphasis>
, sternal ribs; 
<emphasis box="[624,659,1586,1607]" italics="true" pageId="16" pageNumber="17">sucf</emphasis>
, supracoracoid foramen; 
<emphasis box="[905,918,1586,1607]" italics="true" pageId="16" pageNumber="17">u</emphasis>
, ulna; 
<emphasis box="[982,1000,1586,1607]" italics="true" pageId="16" pageNumber="17">ul</emphasis>
, ulnare; 
<emphasis box="[1084,1103,1586,1607]" italics="true" pageId="16" pageNumber="17">xs</emphasis>
, xiphisternum.
</paragraph>
</caption>
<paragraph blockId="16.[491,1530,1662,1888]" pageId="16" pageNumber="17">The mineralized scales were digitally removed from all the CT scans to facilitate rendering of the underlying bone surface and sculpturing. We also digitally removed the endolymphatic sacs.</paragraph>
<paragraph blockId="16.[491,1530,1662,1888]" lastBlockId="19.[491,1541,228,1888]" lastPageId="19" lastPageNumber="20" pageId="16" pageNumber="17">
Skull (
<figureCitation box="[601,719,1781,1808]" captionStart-0="Figure 5" captionStart-1="Figure 6" captionStart-2="Figure 7" captionStart-3="Figure 8" captionStart-4="Figure 9" captionStart-5="Figure 10" captionStartId-0="8.[507,571,729,751]" captionStartId-1="13.[507,571,1713,1735]" captionStartId-2="15.[507,571,1713,1735]" captionStartId-3="16.[507,571,1352,1374]" captionStartId-4="17.[507,571,1453,1475]" captionStartId-5="18.[507,571,1747,1769]" captionTargetBox-0="[492,1541,229,684]" captionTargetBox-1="[491,1188,229,1668]" captionTargetBox-2="[491,1381,229,1668]" captionTargetBox-3="[491,1541,229,1307]" captionTargetBox-4="[492,1541,229,1408]" captionTargetBox-5="[492,1509,230,1692]" captionTargetId-0="figure@8.[491,1541,229,684]" captionTargetId-1="figure@13.[491,1188,229,1668]" captionTargetId-2="figure@15.[491,1381,229,1668]" captionTargetId-3="figure@16.[491,1541,229,1307]" captionTargetId-4="figure@17.[491,1541,229,1408]" captionTargetId-5="figure@18.[491,1512,229,1702]" captionTargetPageId-0="8" captionTargetPageId-1="13" captionTargetPageId-2="15" captionTargetPageId-3="16" captionTargetPageId-4="17" captionTargetPageId-5="18" captionText-0="Figure 5 Micro-CT images of mineraliZed scales of Geckolepis specimens. Shown in dorsal (A, D, F, H, J) and ventral (B, E, G, I, K) view, and coronal cut at the parietal level (C) of Geckolepis megalepis (A – C, ZSM 2126 / 2007), Geckolepis OTU AB (D – E, ZSM 1520 / 2008), G. maculata (F – G, ZMB 9655), and G. humbloti (H – I, ZSM 80 / 2010; J – K, ZSM 81 / 2006). Note the high density of the scale covering in all the species, and the lack of mineralization in the postmental scales (compare with Figs. 2 and 3)." captionText-1="Figure 6 Micro-CT images of skull (cranium and jaw) of the holotype of Geckolepis megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal, (B) ventral, (C) lateral, (D) labial, and (E) lingual view. Abbreviations: adf, anterior inferior dental foramen; amf, anterior mylohyoid foramen; ar, articular facet; asnp, ascending nasal process of premaxilla; bo, basioccipital; bp, basipterygoid process of parabasisphenoid; ch, choana; cob, compound bone; cor, coronoid; d, dentary; dpp, (continued on next page ...) Figure 6 (... continued) descending parietal process; ect, ectopterygoid; ept, epipterygoid; f, frontal; hscc, horizontal semi-circular canal; if, incisive foramen; j, jugal; larst, lateral aperture of the recessus scalae tympani; mdf, mandibular fossa; mf, mental foramen; mko, opening of the Meckelian canal, mx, maxilla; mx. ap, anterior process of maxilla; mx. fp, facial process of maxilla; mx. pp, posterior process of maxilla; mx. ps, palatal shelf of maxilla; n, nasal; occ, occipital condyle; oto, otooccipital; pal, palatine; pal. mp, maxillary process of palatine; pal. vf, vomerine flange of palatine; par, parietal; pbsh, parabasisphenoid; pmx, premaxilla; pmx. pps, premaxilla palatal shelf; pof, postorbitofrontal; pop, paroccipital process; ppp, posterolateral process of parietal; prf, prefrontal; pro, prootic; pro. ca, crista alaris of prootic; pro. cp, crista prootica of prootic; psaf, posterior surangular foramen; pscc, posterior semicircular canal; pt, pterygoid; pt. qp, quadrate process of pterygoid; q, quadrate; q. ch, conch of quadrate; rap, retroarticular process; s, stapes; saf, surangular foramen; scr, sclerotic ring; smx, septomaxilla; spl, splenial; sof, suborbital fenestra; sq, squamosal, so, supraoccipital; spht, sphenooccipital tubercle; tbr, trabeculae; v, vomer; v. lp, lateral process of vomer; v. mp, maxillary process of vomer. A rotational video of the skull is provided in Video S 1." captionText-2="Figure 7 Micro-CT images of spinal column of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Lateral and (B) dorsal view. Abbreviations: lb, lumbar; sc, sacral. Regions are indicated by the posterodorsal-most point of the neural arches." captionText-3="Figure 8 Micro-CT images of pectoral girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Ventral and (B) lateral view. Abbreviations: 1 cf, primary coracoid fenestra; 2 cf, secondary coracoid fenestra; acr, acromion process; cf, clavicular fenestra; cl, clavicle; cor, coracoid; ic, interclavicle; gf, glenoid fossa; hu, humerus; hu. c, humeral condyle; hu. dtp, deltopectoral crest of humerus; hu. ec, ectepicondylar crest of humerus; hu. hc, humeral crest of humerus; mc, metacarpal; ms, mesosternum; p, phalanges; ps, presternum; pu, patella ulnaris; r, radius; rl, radiale; s, scapula; sc. 1 c, primary coracoid ray of scapulocoracoid; sc. 2 c, secondary coracoid ray of scapulocoracoid; scf, scapulocoracoid fenestra; sf, scapular fenestra; ss, suprascapula; sc. sr, scapular ray of scapulocoracoid; sr, sternal ribs; sucf, supracoracoid foramen; u, ulna; ul, ulnare; xs, xiphisternum." captionText-4="Figure 9 Micro-CT images of pelvic girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal and (B) lateral view. Abbreviations: ac, acetabulum; ag, astragalocalcaneum; ep, epipubis; f, femur; fb, fibula; f. it, internal trochanter; f. itf, intertrochanteric fossa; f. fc, femoral condyle; hyi, hypoischium; il, ilium; il. pap, preacetabular process of ilium; is, ischium; is. t, ischiadic tuberosity; ltp, lenticular tibial patella; mt, metatarsals; of, obturator foramen; p, phalanges; pic, proischiadic cartilage; pb, pubis; pb. pl, processus lateralis of pubis; pc, post-cloacal bone; pt, pubic tubercle; tb, tibia; tf, thyroid fenestra." captionText-5="Figure 10 Comparative micro-CT images of the skull (cranium and jaw) of other Geckolepis species. Shown in dorsal (A, F, K, P), ventral (B, G, L, Q), lateral (C, H, M, R), labial (D, I, N, S), and lingual (E, J, O, T) view. Depicting Geckolepis OTU AB (A – E, ZSM 1520 / 2008), holotype of G. maculata (F – J, ZMB 9655), and G. humbloti (K – O, ZSM 80 / 2010; P – T, ZSM 81 / 2006). From volume-rendering of micro-CT scans. Rotational videos of these skulls are provided in Videos S 2 – S 5. For labels, see Fig. 6." httpUri-0="https://zenodo.org/record/291230/files/figure.png" httpUri-1="https://zenodo.org/record/291231/files/figure.png" httpUri-2="https://zenodo.org/record/291232/files/figure.png" httpUri-3="https://zenodo.org/record/291233/files/figure.png" httpUri-4="https://zenodo.org/record/291234/files/figure.png" httpUri-5="https://zenodo.org/record/291235/files/figure.png" pageId="16" pageNumber="17">Figs. 5–10</figureCitation>
): The skull of 
<taxonomicName box="[895,1012,1781,1807]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis box="[895,1012,1781,1807]" italics="true" pageId="16" pageNumber="17">Geckolepis</emphasis>
</taxonomicName>
is the typical broad and depressed skull of geckos (
<bibRefCitation author="Kluge AG." box="[588,722,1821,1847]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="16" pageNumber="17" pagination="1 - 60" part="135" refString="Kluge AG. 1967. Higher taxonomic categories of gekkonid lizards and their evolution. Bulletin of the American Museum of Natural History 135: 1 - 60." title="Higher taxonomic categories of gekkonid lizards and their evolution" type="journal article" year="1967">
<emphasis box="[588,722,1821,1847]" italics="true" pageId="16" pageNumber="17">Kluge, 1967</emphasis>
</bibRefCitation>
), wedge shaped in lateral view. The left maxilla of the holotype of 
<taxonomicName box="[491,631,1861,1888]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[491,631,1861,1888]" italics="true" pageId="16" pageNumber="17">
G. 
<taxonomicName box="[525,631,1861,1887]" pageId="16" pageNumber="17" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
(ZMB 9655) is fractured, and the premaxillary and maxillary palatal shelves show some irregular holes. There is no trace of fracture bones associated with these holes, so the cracks may have been caused by an infection that healed during the animal’s life, or may have resulted from fixation in formalin and gradual decalcification over almost 150 years (although the skull and postcranial skeleton shows no other signs of extensive decalcification). The remaining specimens examined are intact (
<taxonomicName box="[1241,1383,1859,1885]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1241,1383,1859,1885]" italics="true" pageId="17" pageNumber="18">
G. 
<taxonomicName box="[1274,1383,1859,1885]" pageId="17" pageNumber="18" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
, 
<taxonomicName box="[1398,1534,1859,1885]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1398,1534,1859,1885]" italics="true" pageId="17" pageNumber="18">
G. 
<taxonomicName box="[1431,1534,1859,1885]" pageId="17" pageNumber="18" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
, and the AB specimen). 
<figureCitation box="[771,868,228,254]" captionStart="Figure 6" captionStartId="13.[507,571,1713,1735]" captionTargetBox="[491,1188,229,1668]" captionTargetId="figure@13.[491,1188,229,1668]" captionTargetPageId="13" captionText="Figure 6 Micro-CT images of skull (cranium and jaw) of the holotype of Geckolepis megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal, (B) ventral, (C) lateral, (D) labial, and (E) lingual view. Abbreviations: adf, anterior inferior dental foramen; amf, anterior mylohyoid foramen; ar, articular facet; asnp, ascending nasal process of premaxilla; bo, basioccipital; bp, basipterygoid process of parabasisphenoid; ch, choana; cob, compound bone; cor, coronoid; d, dentary; dpp, (continued on next page ...) Figure 6 (... continued) descending parietal process; ect, ectopterygoid; ept, epipterygoid; f, frontal; hscc, horizontal semi-circular canal; if, incisive foramen; j, jugal; larst, lateral aperture of the recessus scalae tympani; mdf, mandibular fossa; mf, mental foramen; mko, opening of the Meckelian canal, mx, maxilla; mx. ap, anterior process of maxilla; mx. fp, facial process of maxilla; mx. pp, posterior process of maxilla; mx. ps, palatal shelf of maxilla; n, nasal; occ, occipital condyle; oto, otooccipital; pal, palatine; pal. mp, maxillary process of palatine; pal. vf, vomerine flange of palatine; par, parietal; pbsh, parabasisphenoid; pmx, premaxilla; pmx. pps, premaxilla palatal shelf; pof, postorbitofrontal; pop, paroccipital process; ppp, posterolateral process of parietal; prf, prefrontal; pro, prootic; pro. ca, crista alaris of prootic; pro. cp, crista prootica of prootic; psaf, posterior surangular foramen; pscc, posterior semicircular canal; pt, pterygoid; pt. qp, quadrate process of pterygoid; q, quadrate; q. ch, conch of quadrate; rap, retroarticular process; s, stapes; saf, surangular foramen; scr, sclerotic ring; smx, septomaxilla; spl, splenial; sof, suborbital fenestra; sq, squamosal, so, supraoccipital; spht, sphenooccipital tubercle; tbr, trabeculae; v, vomer; v. lp, lateral process of vomer; v. mp, maxillary process of vomer. A rotational video of the skull is provided in Video S 1." httpUri="https://zenodo.org/record/291231/files/figure.png" pageId="19" pageNumber="20">Figure 6</figureCitation>
provides anatomical labels for most features based on the holotype of 
<taxonomicName box="[679,822,268,294]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[679,822,268,294]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[713,822,268,294]" pageId="19" pageNumber="20" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
, whereas 
<figureCitation box="[939,1019,268,294]" captionStart="Figure 10" captionStartId="18.[507,571,1747,1769]" captionTargetBox="[492,1509,230,1692]" captionTargetId="figure@18.[491,1512,229,1702]" captionTargetPageId="18" captionText="Figure 10 Comparative micro-CT images of the skull (cranium and jaw) of other Geckolepis species. Shown in dorsal (A, F, K, P), ventral (B, G, L, Q), lateral (C, H, M, R), labial (D, I, N, S), and lingual (E, J, O, T) view. Depicting Geckolepis OTU AB (A – E, ZSM 1520 / 2008), holotype of G. maculata (F – J, ZMB 9655), and G. humbloti (K – O, ZSM 80 / 2010; P – T, ZSM 81 / 2006). From volume-rendering of micro-CT scans. Rotational videos of these skulls are provided in Videos S 2 – S 5. For labels, see Fig. 6." httpUri="https://zenodo.org/record/291235/files/figure.png" pageId="19" pageNumber="20">Fig. 10</figureCitation>
provides comparative images of one adult specimen each of the three other species included for comparative purposes. Rotational videos of these scans are provided as Videos S1–S5. The skeleton of 
<taxonomicName box="[1286,1428,347,374]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1286,1428,347,374]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[1319,1428,347,373]" pageId="19" pageNumber="20" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
paratype ZSM 289/2004 is not figured, as the resolution of our micro-CT scan of it is too low, and only some of its character states could be accurately determined (see Appendix S2); these however largely agree with the holotype.
</paragraph>
<caption httpUri="https://zenodo.org/record/291234/files/figure.png" pageId="17" pageNumber="18" targetBox="[492,1541,229,1408]" targetPageId="17">
<paragraph blockId="17.[507,1511,1453,1651]" pageId="17" pageNumber="18">
<emphasis bold="true" pageId="17" pageNumber="18">
Figure 9 Micro-CT images of pelvic girdle of 
<taxonomicName box="[960,1064,1453,1475]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[960,1064,1453,1475]" italics="true" pageId="17" pageNumber="18">Geckolepis</emphasis>
</taxonomicName>
, based on 
<taxonomicName box="[1167,1291,1453,1475]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="megalepis" status="sp. nov.">
<emphasis bold="true" box="[1167,1291,1453,1475]" italics="true" pageId="17" pageNumber="18">G. megalepis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1296,1372,1454,1475]" pageId="17" pageNumber="18" rank="species">sp. nov.</taxonomicNameLabel>
(ZSM 2126/2007).
</emphasis>
(A) Dorsal and (B) lateral view. Abbreviations: 
<emphasis box="[1083,1103,1483,1504]" italics="true" pageId="17" pageNumber="18">ac</emphasis>
, acetabulum; 
<emphasis box="[1238,1259,1483,1504]" italics="true" pageId="17" pageNumber="18">ag</emphasis>
, astragalocalcaneum; 
<emphasis box="[1469,1490,1483,1504]" italics="true" pageId="17" pageNumber="18">ep</emphasis>
, epipubis; 
<emphasis box="[600,606,1512,1533]" italics="true" pageId="17" pageNumber="18">f</emphasis>
, femur; 
<emphasis box="[687,705,1512,1533]" italics="true" pageId="17" pageNumber="18">fb</emphasis>
, fibula; 
<emphasis box="[783,809,1512,1533]" italics="true" pageId="17" pageNumber="18">f.it</emphasis>
, internal trochanter; 
<emphasis box="[1012,1045,1512,1533]" italics="true" pageId="17" pageNumber="18">f.itf</emphasis>
, intertrochanteric fossa; 
<emphasis box="[1282,1310,1512,1533]" italics="true" pageId="17" pageNumber="18">f.fc</emphasis>
, femoral condyle; 
<emphasis box="[507,536,1541,1562]" italics="true" pageId="17" pageNumber="18">hyi</emphasis>
, hypoischium; 
<emphasis box="[683,695,1541,1562]" italics="true" pageId="17" pageNumber="18">il</emphasis>
, ilium; 
<emphasis box="[769,822,1541,1562]" italics="true" pageId="17" pageNumber="18">il.pap</emphasis>
, preacetabular process of ilium; 
<emphasis box="[1133,1147,1542,1563]" italics="true" pageId="17" pageNumber="18">is</emphasis>
, ischium; 
<emphasis box="[1246,1273,1542,1563]" italics="true" pageId="17" pageNumber="18">is.t</emphasis>
, ischiadic tuberosity; 
<emphasis box="[1481,1506,1541,1562]" italics="true" pageId="17" pageNumber="18">ltp</emphasis>
, lenticular tibial patella; 
<emphasis box="[732,758,1571,1592]" italics="true" pageId="17" pageNumber="18">mt</emphasis>
, metatarsals; 
<emphasis box="[888,905,1571,1592]" italics="true" pageId="17" pageNumber="18">of</emphasis>
, obturator foramen; 
<emphasis box="[1107,1119,1571,1592]" italics="true" pageId="17" pageNumber="18">p</emphasis>
, phalanges; 
<emphasis box="[1236,1263,1571,1592]" italics="true" pageId="17" pageNumber="18">pic</emphasis>
, proischiadic cartilage; 
<emphasis box="[507,530,1600,1621]" italics="true" pageId="17" pageNumber="18">pb</emphasis>
, pubis; 
<emphasis box="[605,651,1600,1621]" italics="true" pageId="17" pageNumber="18">pb.pl</emphasis>
, processus lateralis of pubis; 
<emphasis box="[928,948,1600,1621]" italics="true" pageId="17" pageNumber="18">pc</emphasis>
, post-cloacal bone; 
<emphasis box="[1139,1158,1600,1621]" italics="true" pageId="17" pageNumber="18">pt</emphasis>
, pubic tubercle; 
<emphasis box="[1317,1335,1600,1621]" italics="true" pageId="17" pageNumber="18">tb</emphasis>
, tibia; 
<emphasis box="[1401,1415,1600,1621]" italics="true" pageId="17" pageNumber="18">tf</emphasis>
, thyroid fenestra.
</paragraph>
</caption>
<caption httpUri="https://zenodo.org/record/291235/files/figure.png" pageId="18" pageNumber="19" targetBox="[492,1509,230,1692]" targetPageId="18">
<paragraph blockId="18.[507,1500,1747,1886]" pageId="18" pageNumber="19">
<emphasis bold="true" box="[507,1498,1747,1769]" pageId="18" pageNumber="19">
Figure 10 Comparative micro-CT images of the skull (cranium and jaw) of other 
<taxonomicName box="[1315,1419,1747,1769]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[1315,1419,1747,1769]" italics="true" pageId="18" pageNumber="19">Geckolepis</emphasis>
</taxonomicName>
species.
</emphasis>
Shown in dorsal (A, F, K, P), ventral (B, G, L, Q), lateral (C, H, M, R), labial (D, I, N, S), and lingual (E, J, O, T) view. Depicting 
<taxonomicName box="[738,834,1806,1827]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">
<emphasis box="[738,834,1806,1827]" italics="true" pageId="18" pageNumber="19">Geckolepis</emphasis>
</taxonomicName>
OTU AB (A–E, ZSM 1520/2008), holotype of 
<taxonomicName box="[1278,1393,1806,1827]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1278,1393,1806,1827]" italics="true" pageId="18" pageNumber="19">G. maculata</emphasis>
</taxonomicName>
(F–J, ZMB 9655), and 
<taxonomicName box="[615,727,1835,1856]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[615,727,1835,1856]" italics="true" pageId="18" pageNumber="19">G. humbloti</emphasis>
</taxonomicName>
(K–O, ZSM 80/2010; P–T, ZSM 81/2006). From volume-rendering of micro-CT scans. Rotational videos of these skulls are provided in Videos S2–S5. For labels, see Fig. 6.
</paragraph>
</caption>
<paragraph blockId="19.[491,1541,228,1888]" pageId="19" pageNumber="20">Cranium: Nares oriented anteriorly, bordered medially by premaxilla, ventrally by premaxilla and maxilla, laterally by facial process of the maxilla, and dorsally by nasals. The orbits are incomplete posteriorly, and they accommodate the majority of the circular eye (as defined by the sclerotic ring). The orbits are oriented anterolaterally, possibly enabling some field of vision overlap. They are formed by the maxilla and jugal ventrally, prefrontal anteriorly, frontal dorsally, and postfrontal posterodorsally. A sclerotic ring is present, composed of 14 bones.</paragraph>
<paragraph blockId="19.[491,1541,228,1888]" pageId="19" pageNumber="20">
The premaxilla is fused, with isodont, sharply pointed teeth with 13 tooth loci, this being a constant number among all 
<taxonomicName box="[916,1033,826,852]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis box="[916,1033,826,852]" italics="true" pageId="19" pageNumber="20">Geckolepis</emphasis>
</taxonomicName>
specimens examined. The ascending nasal process is short and forms a bony septum between the nares, tapering abruptly dorsally, where it briefly overlaps the nasals. The palatal shelf contacts the vomer, defines an incisive foramen, and contacts the maxillae laterally.
</paragraph>
<paragraph blockId="19.[491,1541,228,1888]" pageId="19" pageNumber="20">
The maxilla possesses a large facial process and a relatively narrow palatal shelf, as well as a long posterior process, an anterior process, and an anterior maxillary lappet on the lingual side of the anterior process. The alveolar border bears deeply pleurodont, sharply pointed isodont teeth. Tooth loci fluctuate between 35 and 40, 36 in 
<taxonomicName box="[1314,1457,1105,1131]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1314,1457,1105,1131]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[1348,1457,1105,1131]" pageId="19" pageNumber="20" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
and the AB specimen; 
<taxonomicName box="[707,846,1144,1171]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[707,846,1144,1171]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[740,846,1144,1170]" pageId="19" pageNumber="20" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
presents the lowest tooth count, with 35 tooth loci. The maxilla is pierced by four to six supralabial foramina. The posterior process is in contact with the jugal and ectopterygoid posteromedially. The palatal shelf contacts the anterior lateral process of the palatine posteriorly. The maxillary lappet contacts the vomer laterally and the premaxilla’s posterior palatine shelf ventrally, and does not extend to meet its contralateral. The anterior process contacts the premaxilla. The facial process is broad and dorsolaterally convex, its dorsal margin sloped downward from its posterior end to its anterior end, its posterior margin weakly (
<taxonomicName box="[991,1131,1423,1450]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[991,1131,1423,1450]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[1025,1131,1423,1449]" pageId="19" pageNumber="20" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
, one specimen of 
<taxonomicName box="[1346,1483,1423,1450]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1346,1483,1423,1450]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[1380,1483,1423,1449]" pageId="19" pageNumber="20" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
) or strongly curved (
<taxonomicName box="[693,836,1463,1490]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[693,836,1463,1490]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[727,836,1463,1489]" pageId="19" pageNumber="20" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
, the AB specimen, and one specimen of 
<taxonomicName box="[1320,1457,1463,1490]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1320,1457,1463,1490]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[1354,1457,1463,1489]" pageId="19" pageNumber="20" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
) and in contact with the prefrontal; posterodorsally in contact with the frontal, and dorsally in contact with the nasal.
</paragraph>
<paragraph blockId="19.[491,1541,228,1888]" pageId="19" pageNumber="20">
The nasal is nearly rectangular (except by the curved anterior edge that forms the posterodorsal margin of the nares), a small portion of the medial edge lies beneath the ascending nasal process of the premaxilla, and the anterolateral margin borders a small gap with the facial process of the maxilla; the lateral edge is straight in 
<taxonomicName box="[1314,1454,1702,1729]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1314,1454,1702,1729]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[1348,1454,1702,1728]" pageId="19" pageNumber="20" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
, bulges slightly outward in 
<taxonomicName box="[720,863,1742,1769]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[720,863,1742,1769]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[754,863,1742,1768]" pageId="19" pageNumber="20" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
and the AB specimen, and is curved with a lateral flange overlapping the maxillary facial process in 
<taxonomicName box="[999,1135,1782,1808]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[999,1135,1782,1808]" italics="true" pageId="19" pageNumber="20">
G. 
<taxonomicName box="[1032,1135,1782,1808]" pageId="19" pageNumber="20" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
(as seen in other geckos; 
<bibRefCitation author="Evans SE." editor="Gaunt AS" journalOrPublisher="Ithaca: Society for the Study of Amphibians and Reptiles" pageId="19" pageNumber="20" pagination="1 - 348" refString="Evans SE. 2008. The skull of lizards and tuatara. In: Gaunt AS, Adler K, eds. The skull of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 348." title="The skull of lizards and tuatara" type="book chapter" volumeTitle="The skull of Lepidosauria" year="2008">
<emphasis italics="true" pageId="19" pageNumber="20">Evans, 2008</emphasis>
</bibRefCitation>
); laterally in broad contact with maxillary facial process, and posteriorly in contact with the frontal. Nasals are partially fused in the AB specimen.
</paragraph>
<paragraph blockId="20.[491,1542,228,1913]" pageId="20" pageNumber="21">
The prefrontal is strongly convex and has an extensive overlap with the facial process of the maxilla, leaving the exposed surface roughly crescent-shaped in all species (slightly more crescentic in 
<taxonomicName box="[714,857,307,333]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[714,857,307,333]" italics="true" pageId="20" pageNumber="21">
G. 
<taxonomicName box="[748,857,307,333]" pageId="20" pageNumber="21" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
and some individuals of 
<taxonomicName box="[1153,1290,307,333]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1153,1290,307,333]" italics="true" pageId="20" pageNumber="21">
G. 
<taxonomicName box="[1187,1290,307,333]" pageId="20" pageNumber="21" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
). The posterior edge is weakly bowed and curves posteromedially forming the orbito-nasal flange. Dorsally it is distantly separated from the postorbitofrontal. The prefrontal and the maxilla bound the lacrimal foramen.
</paragraph>
<paragraph blockId="20.[491,1542,228,1913]" pageId="20" pageNumber="21">
The frontals remain paired and unfused—this state may however change with age, as the AB specimen has at least partially fused frontals, although a partial suture is still visible anteriorly and posteriorly (see 
<figureCitation box="[848,924,544,570]" captionStart="Figure 10" captionStartId="18.[507,571,1747,1769]" captionTargetBox="[492,1509,230,1692]" captionTargetId="figure@18.[491,1512,229,1702]" captionTargetPageId="18" captionText="Figure 10 Comparative micro-CT images of the skull (cranium and jaw) of other Geckolepis species. Shown in dorsal (A, F, K, P), ventral (B, G, L, Q), lateral (C, H, M, R), labial (D, I, N, S), and lingual (E, J, O, T) view. Depicting Geckolepis OTU AB (A – E, ZSM 1520 / 2008), holotype of G. maculata (F – J, ZMB 9655), and G. humbloti (K – O, ZSM 80 / 2010; P – T, ZSM 81 / 2006). From volume-rendering of micro-CT scans. Rotational videos of these skulls are provided in Videos S 2 – S 5. For labels, see Fig. 6." httpUri="https://zenodo.org/record/291235/files/figure.png" pageId="20" pageNumber="21">Fig. 10</figureCitation>
A). It is in anterior contact with the nasal (straight or slightly concave suture), lateral contact to the facial process of the maxilla (concave suture) and the prefrontal (convex suture), posterolateral contact with the postorbitofrontal (which clasps the frontoparietal suture), and an extensive frontoparietal suture that is weakly curved anteriorly. The anterior end is overlapped by the nasal bones, and the visible portion is roughly half the width of the posterior end, the narrowest point being at the interorbital point. The subolfactory processes of the frontals contact each other but they remain separated, so there is also no ventral fusion. This condition is extremely rare in gekkotans, known only in the fossil 
<taxonomicName authority="Daza, Bauer &amp; Snively, 2013" authorityName="Daza, Bauer &amp; Snively" authorityYear="2013" box="[912,1479,860,886]" class="Reptilia" genus="Gobekko" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="cretacicus">
<emphasis box="[912,1124,860,886]" italics="true" pageId="20" pageNumber="21">
Gobekko 
<taxonomicName authority="Daza, Bauer &amp; Snively, 2013" authorityName="Daza, Bauer &amp; Snively" authorityYear="2013" box="[1016,1479,860,886]" pageId="20" pageNumber="21" rank="species" species="cretacicus">
<taxonomicName box="[1016,1124,861,886]" pageId="20" pageNumber="21" rank="species" species="cretacicus">cretacicus</taxonomicName>
(
<emphasis box="[1142,1467,860,886]" italics="true" pageId="20" pageNumber="21">
Daza, 
<bibRefCitation author="DaZa JD &amp; Bauer AM &amp; Snively ED." box="[1215,1467,860,886]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="20" pageNumber="21" pagination="430 - 448" part="167" publicationUrl="10.1111/zoj.12012." refString="DaZa JD, Bauer AM, Snively ED. 2013. Gobekko cretacicus (Reptilia: Squamata) and its bearing on the interpretation of gekkotan affinities. Zoological Journal of the Linnean Society 167: 430 - 448 DOI 10.1111 / zoj. 12012." title="Gobekko cretacicus (Reptilia: Squamata) and its bearing on the interpretation of gekkotan affinities" type="journal article" year="2013">Bauer &amp; Snively, 2013</bibRefCitation>
</emphasis>
)
</taxonomicName>
</emphasis>
</taxonomicName>
. The 
<emphasis box="[491,624,900,925]" italics="true" pageId="20" pageNumber="21">crista cranii</emphasis>
of the frontals are sutured to the medial side of the posterodorsal process of the prefrontal, thereby forming the dorsal and anterior orbital ridge. The frontal lacks significantly extended anteromedial and anterolateral processes.
</paragraph>
<paragraph blockId="20.[491,1542,228,1913]" pageId="20" pageNumber="21">The jugal is elongated and slender with tapered ends. It extends from the posterior process of the maxilla anteriorly along its medial edge, in contact with the ectopterygoid ventromedially, almost extending far enough forward to meet the palatine and participate in the lacrimal foramen.</paragraph>
<paragraph blockId="20.[491,1542,228,1913]" pageId="20" pageNumber="21">
The parietal is in broad medial contact with its contralateral, the suture is straight in 
<taxonomicName box="[491,631,1215,1242]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[491,631,1215,1242]" italics="true" pageId="20" pageNumber="21">
G. 
<taxonomicName box="[525,631,1215,1241]" pageId="20" pageNumber="21" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
and zigzags in 
<emphasis box="[812,1105,1215,1242]" italics="true" pageId="20" pageNumber="21">
<taxonomicName box="[812,958,1215,1242]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="megalepis">
G. 
<taxonomicName box="[846,958,1215,1241]" pageId="20" pageNumber="21" rank="species" species="megalepis">megalepis</taxonomicName>
</taxonomicName>
, 
<taxonomicName box="[969,1105,1215,1242]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="humbloti">
G. 
<taxonomicName box="[1002,1105,1215,1241]" pageId="20" pageNumber="21" rank="species" species="humbloti">humbloti</taxonomicName>
</taxonomicName>
</emphasis>
, and the AB specimen, although in the lattermost there is also partial fusion, rendering the suture faint. The parietal also contacts the postorbitofrontal anterolaterally, 
<emphasis box="[1035,1164,1294,1320]" italics="true" pageId="20" pageNumber="21">crista alaris</emphasis>
of the prootic lateroventrally, and the squamosal posterolaterally. The bone is broad, curved downwards forming some lateral protection for the brain. It is subtrapezoidal in shape, its lateral and median margins subparallel, the anterior margin angled posteriorly along the frontoparietal suture, the posterior margin angled anteriorly. The posteroparietal process is long and thin in 
<taxonomicName box="[491,631,1492,1518]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[491,631,1492,1518]" italics="true" pageId="20" pageNumber="21">
G. 
<taxonomicName box="[525,631,1492,1518]" pageId="20" pageNumber="21" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
and one specimen of 
<taxonomicName box="[900,1037,1492,1518]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[900,1037,1492,1518]" italics="true" pageId="20" pageNumber="21">
G. 
<taxonomicName box="[934,1037,1492,1518]" pageId="20" pageNumber="21" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
, and broad and short in 
<emphasis box="[1344,1494,1492,1518]" italics="true" pageId="20" pageNumber="21">
<taxonomicName box="[1344,1490,1492,1518]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="megalepis">
G. 
<taxonomicName box="[1378,1490,1492,1518]" pageId="20" pageNumber="21" rank="species" species="megalepis">megalepis</taxonomicName>
</taxonomicName>
,
</emphasis>
one specimen of 
<taxonomicName box="[693,830,1531,1558]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[693,830,1531,1558]" italics="true" pageId="20" pageNumber="21">
G. 
<taxonomicName box="[727,830,1531,1557]" pageId="20" pageNumber="21" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
and the AB specimen, extending posterolaterally from the posterolateral corner of the parietal to contact with the squamosal.
</paragraph>
<paragraph blockId="20.[491,1542,228,1913]" pageId="20" pageNumber="21">
The postorbitofrontal is laminar: thin, short, and curved, extending just anterior and posterior to frontoparietal suture and bracing it (
<emphasis box="[1056,1246,1650,1676]" italics="true" pageId="20" pageNumber="21">Daza et al., 2008</emphasis>
; 
<bibRefCitation author="Rieppel O." box="[1258,1410,1650,1676]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="20" pageNumber="21" pagination="291 - 318" part="82" refString="Rieppel O. 1984. The structure of the skull and jaw adductor musculature of the Gekkota, with comments on the phylogenetic relationships of the Xantusiidae (Reptilia: Lacertilia). Zoological Journal of the Linnean Society 82: 291 - 318" title="The structure of the skull and jaw adductor musculature of the Gekkota, with comments on the phylogenetic relationships of the Xantusiidae (Reptilia: Lacertilia)" type="journal article" year="1984">
<emphasis box="[1258,1410,1650,1676]" italics="true" pageId="20" pageNumber="21">Rieppel, 1984</emphasis>
</bibRefCitation>
) in contact with frontal anteromedially and parietal posteromedially. It lacks a discrete free process for the attachment of the postorbital ligament (
<bibRefCitation author="Evans SE." box="[998,1133,1729,1755]" editor="Gaunt AS" journalOrPublisher="Ithaca: Society for the Study of Amphibians and Reptiles" pageId="20" pageNumber="21" pagination="1 - 348" refString="Evans SE. 2008. The skull of lizards and tuatara. In: Gaunt AS, Adler K, eds. The skull of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 348." title="The skull of lizards and tuatara" type="book chapter" volumeTitle="The skull of Lepidosauria" year="2008">
<emphasis box="[998,1133,1729,1755]" italics="true" pageId="20" pageNumber="21">Evans, 2008</emphasis>
</bibRefCitation>
), which might instead be anchored to the body of the bone.
</paragraph>
<paragraph blockId="20.[491,1542,228,1913]" pageId="20" pageNumber="21">
The squamosal is short, slender, and curved, contacting the posterolateral process of the parietal anteromedially, and the paroccipital process posteriorly. It is considerably reduced in 
<taxonomicName box="[522,662,1887,1913]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[522,662,1887,1913]" italics="true" pageId="20" pageNumber="21">
G. 
<taxonomicName box="[556,662,1887,1913]" pageId="20" pageNumber="21" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="21.[491,1542,228,1888]" pageId="21" pageNumber="22">The quadrate has a deep indentation in the conch. This bone meets the quadrate process of the pterygoid ventrally and has suspension formed by ligaments of the squamosal and the paroccipital processes; it is not in direct contact with any other bones. It has a thick central column and a thin, posterolaterally directed conch that lacks an obvious squamosal notch dorsally. Its cephalic condyle is dorsomedial and not strongly expanded. Its mandibular condyle is concave. It possesses a large foramen in the ventral half of the conch.</paragraph>
<paragraph blockId="21.[491,1542,228,1888]" pageId="21" pageNumber="22">The septomaxilla is very thin, U-shaped, in anterior contact with the premaxilla, otherwise suspended in the nasal capsule. Its medial arm contacts the contralateral,</paragraph>
<paragraph blockId="21.[491,1542,228,1888]" pageId="21" pageNumber="22">separated anteriorly by a small foramen lying dorsal to the incisive foramen of the vomer. The lateral arm ascends slightly, and is long and cuneate with a sculpted lateral surface.</paragraph>
<paragraph blockId="21.[491,1542,228,1888]" pageId="21" pageNumber="22">
The vomer is a thin, laminar bone. It is strongly fused to its contralateral, though a median ridge allows the individual bones to be distinguished despite strong ontogenetic fusion. Anteromedially an incisive foramen is present between the vomer and the posterior palatal shelf of the premaxilla, the shape of this foramen is variable among the species examined, being a v-shaped notch (
<taxonomicName box="[785,927,786,812]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[785,927,786,812]" italics="true" pageId="21" pageNumber="22">
G. 
<taxonomicName box="[818,927,786,812]" pageId="21" pageNumber="22" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
and 
<taxonomicName box="[988,1127,786,812]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[988,1127,786,812]" italics="true" pageId="21" pageNumber="22">
G. 
<taxonomicName box="[1021,1127,786,812]" pageId="21" pageNumber="22" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
) or oval shaped (
<taxonomicName box="[1336,1473,786,812]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1336,1473,786,812]" italics="true" pageId="21" pageNumber="22">
G. 
<taxonomicName box="[1370,1473,786,812]" pageId="21" pageNumber="22" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
and the AB specimen). The anterolateral extension of the vomer is in contact with the lingual maxillary lappet. Posteriorly it bears two slender lateral processes, and the paired elements form a broad median projection. The lateral process curves medially to join the vomerine process of the palatine. The median spur is bordered on either side by the distal tips of the palatine vomerine processes, and forms the anterior end of the interpterygoid vacuity. The vomer has also two foramina that might correspond to openings of the lacrimal duct.
</paragraph>
<paragraph blockId="21.[491,1542,228,1888]" pageId="21" pageNumber="22">The palatine is squarish, with rounded lateral and medial edges. The vomerine flange and maxillary process are slender and subequal in length, together forming the border of the choana. The vomerine flange lies parallel to the posterior processes of the vomer and rests on a notch on the body of the vomer; the maxillary process contacts the maxilla’s palatal shelf laterally. The palatine forms the anteromedial border of the suborbital fenestra. The bone is without an obvious pterygoid process but possesses a posteroventral shelf where the palatine process of the pterygoid overlaps it. Lateral to this overlap, the bone borders a slit extending medially from the suborbital fenestra between the palatine and pterygoid. The medial edge of the palatine forms the lateral border of the interpterygoid vacuity. The lateral face of the palatine is in contact with the anterolingual end of the ectopterygoid. The pterygopalatine joint is oblique.</paragraph>
<paragraph blockId="21.[491,1542,228,1888]" pageId="21" pageNumber="22">The pterygoid is roughly y-shaped, with a brief anteromedial articulation with the palatine and anterodorsal articulation with the ectopterygoid, articulating with the</paragraph>
<paragraph blockId="21.[491,1542,228,1888]" lastBlockId="22.[491,1542,228,1913]" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22">
epipterygoid at the 
<emphasis box="[725,904,1583,1609]" italics="true" pageId="21" pageNumber="22">fossa columellae</emphasis>
, and contacting the quadrate posterolaterally. The pterygoid has a palatine process anteromedially and a sculpted anterior border that is straight lateral to the palatine process, forming the posterior border of the slit extending medially from the suborbital fenestra—then concave, forming the posterior border of the suborbital fenestra—then extending anteriorly again to form the pterygoid flange in contact with ectopterygoid, practically excluding the ectopterygoid from the posterior margin of the suborbital fenestra; also forming the posterolateral border of the interpterygoid vacuity. The facet that contacts the basipterygoid process is porous. In lateral view, the quadrate process curves laterally beyond this point and the 
<emphasis box="[1079,1258,228,254]" italics="true" pageId="22" pageNumber="23">fossa columellae</emphasis>
to below the quadrate.
</paragraph>
<paragraph blockId="22.[491,1542,228,1913]" pageId="22" pageNumber="23">
The ectopterygoid is bent downward. It is anterolaterally in contact with the jugal, and posteriorly in ventrolateral contact with the anterolateral pterygoid flange. The bone’s downward bend is due to the more dorsal position of palatine and maxilla relative to the pterygoid. Its medial margin is sigmoid in 
<taxonomicName box="[1043,1182,386,412]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1043,1182,386,412]" italics="true" pageId="22" pageNumber="23">
G. 
<taxonomicName box="[1076,1182,386,412]" pageId="22" pageNumber="23" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1243,1379,386,412]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1243,1379,386,412]" italics="true" pageId="22" pageNumber="23">
G. 
<taxonomicName box="[1276,1379,386,412]" pageId="22" pageNumber="23" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
, variable in 
<taxonomicName box="[491,634,425,452]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[491,634,425,452]" italics="true" pageId="22" pageNumber="23">
G. 
<taxonomicName box="[525,634,425,451]" pageId="22" pageNumber="23" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
(wavy in ZSM 2126/2007 but sigmoid in ZSM 289/2004), and wavy in the AB specimen. The suborbital fenestra is roughly teardrop shaped, pointed anteriorly and rounded posteriorly, formed by the ectopterygoid laterally, pterygoid posteriorly, palatine anteromedially, and maxilla anterolaterally. The suborbital fenestra is broad in 
<taxonomicName box="[1399,1541,544,570]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1399,1541,544,570]" italics="true" pageId="22" pageNumber="23">G. megalepis</emphasis>
</taxonomicName>
and the AB specimen and narrow in all the remaining specimens examined.
</paragraph>
<paragraph blockId="22.[491,1542,228,1913]" pageId="22" pageNumber="23">
The epipterygoid is columnar, tilted posteriorly, and appears mildly medially bowed in 
<taxonomicName box="[491,634,662,689]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[491,634,662,689]" italics="true" pageId="22" pageNumber="23">
G. 
<taxonomicName box="[525,634,662,688]" pageId="22" pageNumber="23" rank="species" species="megalepis">megalepis</taxonomicName>
</emphasis>
</taxonomicName>
, vertical or even externally bowed in 
<taxonomicName box="[1070,1207,662,689]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1070,1207,662,689]" italics="true" pageId="22" pageNumber="23">G. humbloti</emphasis>
</taxonomicName>
, and vertical in 
<taxonomicName box="[1395,1535,662,689]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1395,1535,662,689]" italics="true" pageId="22" pageNumber="23">G. maculata</emphasis>
</taxonomicName>
. It extends from the 
<emphasis box="[721,900,702,728]" italics="true" pageId="22" pageNumber="23">fossa columellae</emphasis>
of the pterygoid toward, but not into contact with, the 
<emphasis box="[491,621,741,767]" italics="true" pageId="22" pageNumber="23">crista alaris</emphasis>
of the prootic. The dorsal end is somewhat broader than the rest of the bone. The interpterygoid vacuity is hourglass shaped, but broadens more rapidly posteriorly than anteriorly, and is anteriorly bifurcated as a result of the medial spur of the vomer. The stapes has an oval footplate that fits in the 
<emphasis box="[986,1144,860,886]" italics="true" pageId="22" pageNumber="23">fenestra ovalis</emphasis>
, and two posts extending laterally from footplate, one anterior, the other posterior, converge to form the stapedial stem, leaving an open stapedial foramen. The 
<emphasis box="[959,1117,939,965]" italics="true" pageId="22" pageNumber="23">fenestra ovalis</emphasis>
is posterior to the quadrate.
</paragraph>
<paragraph blockId="22.[491,1542,228,1913]" pageId="22" pageNumber="23">
The basioccipital underlies most of the braincase, is slightly wider than long, and lacks a distinct basal tubera. It is in contact with the parabasisphenoid anteriorly, otooccipitals laterally, and forms the ventral component of the 
<emphasis box="[1082,1285,1057,1083]" italics="true" pageId="22" pageNumber="23">foramen magnum</emphasis>
. It is excluded from participation in the lateral aperture of the 
<emphasis box="[997,1246,1097,1123]" italics="true" pageId="22" pageNumber="23">recessus scala tympani</emphasis>
by the otooccipital. It is posterolaterally bordered by anteroventral extensions of the otooccipitals forming the sphenooccipital tubercle, which is connected to a sharp 
<emphasis box="[1144,1308,1176,1202]" italics="true" pageId="22" pageNumber="23">crista tuberalis</emphasis>
.
</paragraph>
<paragraph blockId="22.[491,1542,228,1913]" pageId="22" pageNumber="23">
The parabasisphenoid is in contact with the prootic dorsally and basioccipital posteriorly. It possesses a short, pointed parasphenoid rostrum, which is an extension of the squared anterior ends of the 
<emphasis box="[729,924,1294,1320]" italics="true" pageId="22" pageNumber="23">cristae trabeculae</emphasis>
. The basipterygoid processes diverge anterolaterally, broadening distally, with flat, curved distal ends forming a synovial joint with the
</paragraph>
<paragraph blockId="22.[491,1542,228,1913]" pageId="22" pageNumber="23">
corresponding fossa of the posterior pterygoid (
<bibRefCitation author="Payne S &amp; Holliday CM &amp; Vickaryous MK." box="[1066,1475,1373,1400]" journalOrPublisher="The Anatomical Record" pageId="22" pageNumber="23" pagination="399 - 405" part="294" publicationUrl="10.1002/ar.21329." refString="Payne S, Holliday CM, Vickaryous MK. 2011. An osteological and histological investigation of cranial joints in geckos. The Anatomical Record 294: 399 - 405 DOI 10.1002 / ar. 21329." title="An osteological and histological investigation of cranial joints in geckos" type="journal article" year="2011">
<emphasis box="[1066,1475,1373,1400]" italics="true" pageId="22" pageNumber="23">Payne, Holliday &amp; Vickaryous, 2011</emphasis>
</bibRefCitation>
). The vidian bridge extends to the base of the basipterygoid process from the 
<emphasis box="[1379,1537,1414,1439]" italics="true" pageId="22" pageNumber="23">crista prootica</emphasis>
of the prootic. Posteriorly, the 
<emphasis box="[855,998,1452,1478]" italics="true" pageId="22" pageNumber="23">crista sellaris</emphasis>
forms the anterior wall of the 
<emphasis box="[1361,1493,1452,1478]" italics="true" pageId="22" pageNumber="23">sella turcica</emphasis>
. It has two pairs of anterior openings: carotid canals opening anteromedially, and the anterior openings of the Vidian canal anterolaterally, parallel to the basipterygoid processes.
</paragraph>
<paragraph blockId="22.[491,1542,228,1913]" pageId="22" pageNumber="23">
The supraoccipital contacts the prootic anteriorly and otooccipitals ventrally, forming the dorsal edge of the 
<emphasis box="[706,908,1610,1636]" italics="true" pageId="22" pageNumber="23">foramen magnum</emphasis>
. The posterior semi-circular canal extends posteriorly to the dorsal margin of the foramen magnum. The supraoccipital has a pair of dorsal tubercles that project dorsally without contacting the parietal; these tubercles are on either side of the midline of the holotype of 
<taxonomicName box="[930,1072,1729,1755]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[930,1072,1729,1755]" italics="true" pageId="22" pageNumber="23">G. megalepis</emphasis>
</taxonomicName>
, but are not strongly raised in any other specimen examined.
</paragraph>
<paragraph blockId="22.[491,1542,228,1913]" lastBlockId="23.[491,1542,228,1913]" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23">
The prootic is thin and has a prominent, triangular 
<emphasis box="[1121,1250,1808,1834]" italics="true" pageId="22" pageNumber="23">crista alaris</emphasis>
. It is in contract with the descending parietal process dorsally, the parabasisphenoid anteroventrally, supraoccipital posterodorsally, otooccipital posteroventrally, and almost in contact with the epipterygoid at the end of the 
<emphasis box="[698,827,228,254]" italics="true" pageId="23" pageNumber="24">crista alaris</emphasis>
. The posterolateral margin forms the anterior wall of the fenestra ovalis, and the posteromedial surface forms the anterolateral wall of the brain case. The anterior semi-circular canal runs through the base of the alary process and 
<emphasis box="[1405,1534,307,333]" italics="true" pageId="23" pageNumber="24">crista alaris</emphasis>
. The horizontal semicircular canal and the ampullar bulge are visible in the posterior edge of the prootic. A projection from the 
<emphasis box="[923,1052,386,412]" italics="true" pageId="23" pageNumber="24">crista alaris</emphasis>
extends anteromedially down to the 
<emphasis italics="true" pageId="23" pageNumber="24">crista sellaris</emphasis>
of the sphenoid and contains the trigeminal foramen (
<emphasis box="[1199,1524,425,452]" italics="true" pageId="23" pageNumber="24">
Daza, 
<bibRefCitation author="DaZa JD &amp; Bauer AM &amp; Snively ED." box="[1272,1524,425,452]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="23" pageNumber="24" pagination="430 - 448" part="167" publicationUrl="10.1111/zoj.12012." refString="DaZa JD, Bauer AM, Snively ED. 2013. Gobekko cretacicus (Reptilia: Squamata) and its bearing on the interpretation of gekkotan affinities. Zoological Journal of the Linnean Society 167: 430 - 448 DOI 10.1111 / zoj. 12012." title="Gobekko cretacicus (Reptilia: Squamata) and its bearing on the interpretation of gekkotan affinities" type="journal article" year="2013">Bauer &amp; Snively, 2013</bibRefCitation>
</emphasis>
), flaring also anterolaterally to the level of the epipterygoid from the base of the 
<emphasis box="[1405,1534,465,491]" italics="true" pageId="23" pageNumber="24">crista alaris</emphasis>
.
</paragraph>
<paragraph blockId="23.[491,1542,228,1913]" pageId="23" pageNumber="24">The otooccipital is in contact with the prootic anteriorly, basioccipital ventromedially, supraoccipital dorsally, and the squamosal on the anterior face of the distal end of the paroccipital process. The horizontal and posterior semi-circular canals are visible as a bulge in posterior view. The occipital recess is enclosed in its posteroventral face. Anterodistally it projects ventrally to participate in the sphenooccipital tubercle with the basioccipital. The paroccipital process is long and thin, but broad dorsoventrally.</paragraph>
<paragraph blockId="23.[491,1542,228,1913]" pageId="23" pageNumber="24">
The 
<emphasis box="[577,780,741,767]" italics="true" pageId="23" pageNumber="24">foramen magnum</emphasis>
is suboval, formed by the supraoccipital dorsally, otooccipitals lateroventrally, and basioccipital ventrally. The occipital condyles are double, formed by the otooccipitals laterally and the basioccipitals medially.
</paragraph>
<paragraph blockId="23.[491,1542,228,1913]" pageId="23" pageNumber="24">
Jaw (
<figureCitation box="[583,658,860,886]" captionStart="Figure 6" captionStartId="13.[507,571,1713,1735]" captionTargetBox="[491,1188,229,1668]" captionTargetId="figure@13.[491,1188,229,1668]" captionTargetPageId="13" captionText="Figure 6 Micro-CT images of skull (cranium and jaw) of the holotype of Geckolepis megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal, (B) ventral, (C) lateral, (D) labial, and (E) lingual view. Abbreviations: adf, anterior inferior dental foramen; amf, anterior mylohyoid foramen; ar, articular facet; asnp, ascending nasal process of premaxilla; bo, basioccipital; bp, basipterygoid process of parabasisphenoid; ch, choana; cob, compound bone; cor, coronoid; d, dentary; dpp, (continued on next page ...) Figure 6 (... continued) descending parietal process; ect, ectopterygoid; ept, epipterygoid; f, frontal; hscc, horizontal semi-circular canal; if, incisive foramen; j, jugal; larst, lateral aperture of the recessus scalae tympani; mdf, mandibular fossa; mf, mental foramen; mko, opening of the Meckelian canal, mx, maxilla; mx. ap, anterior process of maxilla; mx. fp, facial process of maxilla; mx. pp, posterior process of maxilla; mx. ps, palatal shelf of maxilla; n, nasal; occ, occipital condyle; oto, otooccipital; pal, palatine; pal. mp, maxillary process of palatine; pal. vf, vomerine flange of palatine; par, parietal; pbsh, parabasisphenoid; pmx, premaxilla; pmx. pps, premaxilla palatal shelf; pof, postorbitofrontal; pop, paroccipital process; ppp, posterolateral process of parietal; prf, prefrontal; pro, prootic; pro. ca, crista alaris of prootic; pro. cp, crista prootica of prootic; psaf, posterior surangular foramen; pscc, posterior semicircular canal; pt, pterygoid; pt. qp, quadrate process of pterygoid; q, quadrate; q. ch, conch of quadrate; rap, retroarticular process; s, stapes; saf, surangular foramen; scr, sclerotic ring; smx, septomaxilla; spl, splenial; sof, suborbital fenestra; sq, squamosal, so, supraoccipital; spht, sphenooccipital tubercle; tbr, trabeculae; v, vomer; v. lp, lateral process of vomer; v. mp, maxillary process of vomer. A rotational video of the skull is provided in Video S 1." httpUri="https://zenodo.org/record/291231/files/figure.png" pageId="23" pageNumber="24">Figs. 6</figureCitation>
D and 6E): The jaw curves medially anteriorly, and forms a weak symphysis with its counterpart. The dentary is the longest bone, being tubular and enclosing the Meckelian canal, which becomes broader posteriorly, as it approaches the mandibular fossa. It bears isodont, pleurodont, somewhat conical and some recurved teeth. Tooth loci varies considerably from 27 to 40, the smallest number on the left ramus of the AB specimen, but it seems that this specimens has a pathological condition since the number is higher on the other side, reaching the base of the coronoid eminence. The interdental space is larger in the AB specimen, which also explains the lowest number. Maximum tooth loci is roughly 34 in 
<taxonomicName box="[737,877,1176,1202]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[737,877,1176,1202]" italics="true" pageId="23" pageNumber="24">
G. 
<taxonomicName box="[771,877,1176,1202]" pageId="23" pageNumber="24" rank="species" species="maculata">maculata</taxonomicName>
</emphasis>
</taxonomicName>
, 37 in 
<taxonomicName box="[961,1104,1176,1202]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[961,1104,1176,1202]" italics="true" pageId="23" pageNumber="24">G. megalepis</emphasis>
</taxonomicName>
, and 40 in 
<taxonomicName box="[1241,1377,1176,1202]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1241,1377,1176,1202]" italics="true" pageId="23" pageNumber="24">
G. 
<taxonomicName box="[1274,1377,1176,1202]" pageId="23" pageNumber="24" rank="species" species="humbloti">humbloti</taxonomicName>
</emphasis>
</taxonomicName>
; with some teeth clearly missing in all specimens). Five mental foramina are present in the holotype, six in the paratype ZSM 289/2004 of 
<taxonomicName box="[932,1075,1255,1281]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[932,1075,1255,1281]" italics="true" pageId="23" pageNumber="24">G. megalepis</emphasis>
</taxonomicName>
. Posteriorly, the dentary contacts the surangular portion of the compound bone by superior and ventral processes, the latter extending considerably further than the former. The Meckelian canal is not outwardly pronounced, and opens anteriorly below the symphysis.
</paragraph>
<paragraph blockId="23.[491,1542,228,1913]" pageId="23" pageNumber="24">The splenial is a thin, triangular, and flat bone that forms the medial wall of the Meckelian canal, posterior to the tubular portion. It has two discrete foramina, the anterior inferior dental foramen and the anterior mylohyoid foramen.</paragraph>
<paragraph blockId="23.[491,1542,228,1913]" pageId="23" pageNumber="24">The coronoid has a strong and fin-like dorsal eminence with a broadened anterior edge, but its precise shape varies within species. It inserts into the dentary at the level of the last or penultimate tooth (except on one side of the jaw of the AB specimen, as mentioned above). The posteromedial process reaches the middle of the surangular, anterior to the distinct mandibular fossa. The triangular splenial is present on the lingual surface of the mandible, in contact with the posteromedial surface of the dentary, the lingual anteroventral face of the coronoid, and the lingual surface of the surangular.</paragraph>
<paragraph blockId="23.[491,1542,228,1913]" lastBlockId="24.[491,1542,228,1888]" lastPageId="24" lastPageNumber="25" pageId="23" pageNumber="24">
The dorsal edge of the surangular portion of the compound bone is concave. The Meckelian canal is closed, extending into the dentary from the adductor fossa. Surangular and posterior surangular foramina are located in the labial side of the compound bone. An external foramen for the 
<emphasis box="[784,962,228,254]" italics="true" pageId="24" pageNumber="25">chorda tympani</emphasis>
is present at the base of the retroarticular process of the compound bone. The length, width, and concavity of the retroarticular process are variable within species probably due to scaling of the jaw muscles. The retroarticular process is strongly laterally notched, with a medial ridge on its articular surface.
</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">
Axial Skeleton (
<figureCitation box="[704,767,387,414]" captionStart="Figure 7" captionStartId="15.[507,571,1713,1735]" captionTargetBox="[491,1381,229,1668]" captionTargetId="figure@15.[491,1381,229,1668]" captionTargetPageId="15" captionText="Figure 7 Micro-CT images of spinal column of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Lateral and (B) dorsal view. Abbreviations: lb, lumbar; sc, sacral. Regions are indicated by the posterodorsal-most point of the neural arches." httpUri="https://zenodo.org/record/291232/files/figure.png" pageId="24" pageNumber="25">Fig. 7</figureCitation>
): 26 presacral, two sacral, and a varying number of caudal vertebrae are present (the total number cannot be ascertained due to autotomized or regenerated tails in all scanned specimens). Of the presacrals, eight are cervical (defined as being anterior to the first vertebra possessing a rib reaching the sternum), sixteen or seventeen are thoracic (rib-bearing), and one or two lack ossified ribs and are thus considered lumbars.
</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">The atlas has an unfused neural arch, which is also not fused to the centrum, each side with a short dorsolateral posterior projection not overlying the axis. The odontoid process of the axis extends forward between the walls of atlas and into the braincase, fitting in between the occipital condyles. The anterior three cervical vertebrae (atlas, axis, and third cervical) lack ribs. The following five bear ribs of increasing length, all of which are to some degree dorsoventrally broadened.</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">
The vertebrae are notochordal amphicoelous type (
<bibRefCitation author="Romer AS." box="[1128,1272,825,851]" journalOrPublisher="Chicago: The University of Chicago Press" pageId="24" pageNumber="25" refString="Romer AS. 1956. The osteology of the reptiles. Chicago: The University of Chicago Press." title="The osteology of the reptiles" type="book" year="1956">
<emphasis box="[1128,1272,825,851]" italics="true" pageId="24" pageNumber="25">Romer, 1956</emphasis>
</bibRefCitation>
). The ribs of the first four thoracic vertebrae reach the sternum—the fourth via the xiphisternum—followed by seven vertebrae articulating with long, posteriorly arching ribs distally associated with postxiphisternal inscriptional ribs, followed by five or six vertebrae possessing shorter ribs gradually becoming more posteriorly curved (see 
<figureCitation box="[1077,1141,985,1012]" captionStart="Figure 7" captionStartId="15.[507,571,1713,1735]" captionTargetBox="[491,1381,229,1668]" captionTargetId="figure@15.[491,1381,229,1668]" captionTargetPageId="15" captionText="Figure 7 Micro-CT images of spinal column of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Lateral and (B) dorsal view. Abbreviations: lb, lumbar; sc, sacral. Regions are indicated by the posterodorsal-most point of the neural arches." httpUri="https://zenodo.org/record/291232/files/figure.png" pageId="24" pageNumber="25">Fig. 7</figureCitation>
B); one or two lumbar vertebrae follow that are similar in shape to the posterior thoracic vertebrae but lack ribs.
</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">
The sacral pleurapophysis of the first sacral vertebra juts slightly posteriorly, articulating distally with the pelvic girdle. The posterodistal edge is fused to the anterior edge of the pleurapophysis of the second sacral vertebra, forming the 
<emphasis box="[1182,1359,1144,1171]" italics="true" pageId="24" pageNumber="25">foramen sacrale</emphasis>
. The second sacral vertebra possesses a dorsoventrally thin posterior crest comprising almost half the distal breadth of the pleurapophysis (with asymmetrically emarginated distal edges in 
<taxonomicName box="[491,634,1264,1290]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[491,634,1264,1290]" italics="true" pageId="24" pageNumber="25">G. megalepis</emphasis>
</taxonomicName>
specimens: more emarginated on the left than the right in ZSM 2126/2007 and right than left in ZSM 289/2004);
</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">
The first five caudal vertebrae possess long thin lateral processes, initially extending beyond the sacrals, gradually decreasing in breadth, jutting posterolaterally, straight in 
<taxonomicName box="[491,631,1423,1450]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[491,631,1423,1450]" italics="true" pageId="24" pageNumber="25">G. maculata</emphasis>
</taxonomicName>
and 
<taxonomicName box="[694,831,1423,1450]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[694,831,1423,1450]" italics="true" pageId="24" pageNumber="25">G. humbloti</emphasis>
</taxonomicName>
, curved laterally in 
<taxonomicName box="[1072,1215,1423,1450]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1072,1215,1423,1450]" italics="true" pageId="24" pageNumber="25">G. megalepis</emphasis>
</taxonomicName>
, becoming increasingly posterior-jutting. The first three caudals lack hemal arches.
</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">
Pectoral Girdle (
<figureCitation box="[714,776,1503,1529]" captionStart="Figure 8" captionStartId="16.[507,571,1352,1374]" captionTargetBox="[491,1541,229,1307]" captionTargetId="figure@16.[491,1541,229,1307]" captionTargetPageId="16" captionText="Figure 8 Micro-CT images of pectoral girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Ventral and (B) lateral view. Abbreviations: 1 cf, primary coracoid fenestra; 2 cf, secondary coracoid fenestra; acr, acromion process; cf, clavicular fenestra; cl, clavicle; cor, coracoid; ic, interclavicle; gf, glenoid fossa; hu, humerus; hu. c, humeral condyle; hu. dtp, deltopectoral crest of humerus; hu. ec, ectepicondylar crest of humerus; hu. hc, humeral crest of humerus; mc, metacarpal; ms, mesosternum; p, phalanges; ps, presternum; pu, patella ulnaris; r, radius; rl, radiale; s, scapula; sc. 1 c, primary coracoid ray of scapulocoracoid; sc. 2 c, secondary coracoid ray of scapulocoracoid; scf, scapulocoracoid fenestra; sf, scapular fenestra; ss, suprascapula; sc. sr, scapular ray of scapulocoracoid; sr, sternal ribs; sucf, supracoracoid foramen; u, ulna; ul, ulnare; xs, xiphisternum." httpUri="https://zenodo.org/record/291233/files/figure.png" pageId="24" pageNumber="25">Fig. 8</figureCitation>
): The pectoral girdle is comprised of paired clavicles, epicoracoids, and scapulocoracoids, and a non-paired interclavicle and presternum.
</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">The presternum is kite-shaped, and varies in ossification levels from poorly to fully ossified. It has a synchondrotic articulation with the first three sternal ribs along its posterolateral border, but lacks distinct facets for these. Its anterolateral edges are thickened to form the coracosternal groove. No frontanelles are present. The mesosternal extension of the xiphisternum is variably long, but poorly ossified.</paragraph>
<paragraph blockId="24.[491,1542,228,1888]" pageId="24" pageNumber="25">The sagittal interclavicle is posteriorly arrowhead-shaped, and extends less than one third into the sternum. It is anteriorly elongated, tubular and tapering, extending between the clavicles but not beyond them.</paragraph>
<paragraph blockId="25.[491,1542,228,1913]" box="[523,1484,228,254]" pageId="25" pageNumber="26">The suprascapular and epicoracoid regions are at least partly ossified, but never</paragraph>
<paragraph blockId="25.[491,1542,228,1913]" pageId="25" pageNumber="26">
completely. The scapulocoracoid is typical in being composed of a horizontal plate (coracoid portion) and a vertical lateral ascending process (scapular portion). No clear suture of the scapula and coracoid is visible in the micro-CT scans. The coracoid portion is broad, plate-like, with a bulbous process at its posterolateral corner. The scapular portion is long, at least as long as the coracoid portion. Three rays are present, all of which are directed anteromedially: the scapular ray is slender, and passes dorsally anterior to the ascending lateral process of the clavicle in 
<taxonomicName box="[1072,1212,504,531]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1072,1212,504,531]" italics="true" pageId="25" pageNumber="26">G. maculata</emphasis>
</taxonomicName>
, but does not surpass the clavicle in 
<taxonomicName box="[616,758,544,570]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[616,758,544,570]" italics="true" pageId="25" pageNumber="26">G. megalepis</emphasis>
</taxonomicName>
or 
<taxonomicName box="[800,936,544,570]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[800,936,544,570]" italics="true" pageId="25" pageNumber="26">G. humbloti</emphasis>
</taxonomicName>
—it defines the dorsal edge of the scapulocoracoid fenestra, which is ventrally completed by the primary coracoid ray. The secondary coracoid ray extends to the level of the clavicular fenestra in 
<taxonomicName box="[1103,1242,623,649]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1103,1242,623,649]" italics="true" pageId="25" pageNumber="26">G. maculata</emphasis>
</taxonomicName>
, and to the level of the posterior margin of the clavicular fenestra in 
<taxonomicName box="[1014,1157,662,689]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1014,1157,662,689]" italics="true" pageId="25" pageNumber="26">G. megalepis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1214,1351,662,689]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[1214,1351,662,689]" italics="true" pageId="25" pageNumber="26">G. humbloti</emphasis>
</taxonomicName>
. The rays define four fenestrae: the secondary coracoid fenestra (medial scapulocoracoid + secondary coracoid ray); the primary coracoid fenestra (secondary coracoid ray + primary coracoid ray); the scapulocoracoid fenestra (primary coracoid ray + scapular ray); and the scapular fenestra (scapular ray+distal scapulocoracoid). Anteriorly, all four fenestrae are closed by the cartilaginous epicoracoids, which are not rendered in our micro-CT scans (see 
<figureCitation box="[1459,1523,860,886]" captionStart="Figure 8" captionStartId="16.[507,571,1352,1374]" captionTargetBox="[491,1541,229,1307]" captionTargetId="figure@16.[491,1541,229,1307]" captionTargetPageId="16" captionText="Figure 8 Micro-CT images of pectoral girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Ventral and (B) lateral view. Abbreviations: 1 cf, primary coracoid fenestra; 2 cf, secondary coracoid fenestra; acr, acromion process; cf, clavicular fenestra; cl, clavicle; cor, coracoid; ic, interclavicle; gf, glenoid fossa; hu, humerus; hu. c, humeral condyle; hu. dtp, deltopectoral crest of humerus; hu. ec, ectepicondylar crest of humerus; hu. hc, humeral crest of humerus; mc, metacarpal; ms, mesosternum; p, phalanges; ps, presternum; pu, patella ulnaris; r, radius; rl, radiale; s, scapula; sc. 1 c, primary coracoid ray of scapulocoracoid; sc. 2 c, secondary coracoid ray of scapulocoracoid; scf, scapulocoracoid fenestra; sf, scapular fenestra; ss, suprascapula; sc. sr, scapular ray of scapulocoracoid; sr, sternal ribs; sucf, supracoracoid foramen; u, ulna; ul, ulnare; xs, xiphisternum." httpUri="https://zenodo.org/record/291233/files/figure.png" pageId="25" pageNumber="26">Fig. 8</figureCitation>
). This formation is type 6 sensu 
<bibRefCitation author="Lecuru S." box="[856,1016,899,926]" journalOrPublisher="Annales des Sciences Naturelles-Zoologie et Biologie Animale" pageId="25" pageNumber="26" pagination="475 - 510" part="12" refString="Lecuru S. 1968. Remarques sur le scapulo-coracoide des lacertiliens. Annales des Sciences Naturelles-Zoologie et Biologie Animale 12: 475 - 510." title="Remarques sur le scapulo-coracoide des lacertiliens" type="journal article" year="1968">
<emphasis box="[856,1016,899,926]" italics="true" pageId="25" pageNumber="26">Lécuru (1968)</emphasis>
</bibRefCitation>
. The supracoracoid foramen is small, lying closer to first coracoid fenestra than to the glenoid fossa.
</paragraph>
<paragraph blockId="25.[491,1542,228,1913]" pageId="25" pageNumber="26">The clavicle curves posteriorly and dorsally from the midline. It articulates with the ossified acromion process of the poorly ossified suprascapula. It is angled posterolaterally, with a broadly expanded but dorsoventrally flat medial portion—containing a large, oblong clavicular fenestra—and slender curving lateral portion. It articulates at the midline with its contralateral and the interclavicle, and is dorsally exceeded or at least overlapped by the epicoracoid cartilage and parts of the suprascapular rays.</paragraph>
<paragraph blockId="25.[491,1542,228,1913]" pageId="25" pageNumber="26">
Forelimbs (
<figureCitation box="[661,727,1215,1241]" captionStart="Figure 8" captionStartId="16.[507,571,1352,1374]" captionTargetBox="[491,1541,229,1307]" captionTargetId="figure@16.[491,1541,229,1307]" captionTargetPageId="16" captionText="Figure 8 Micro-CT images of pectoral girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Ventral and (B) lateral view. Abbreviations: 1 cf, primary coracoid fenestra; 2 cf, secondary coracoid fenestra; acr, acromion process; cf, clavicular fenestra; cl, clavicle; cor, coracoid; ic, interclavicle; gf, glenoid fossa; hu, humerus; hu. c, humeral condyle; hu. dtp, deltopectoral crest of humerus; hu. ec, ectepicondylar crest of humerus; hu. hc, humeral crest of humerus; mc, metacarpal; ms, mesosternum; p, phalanges; ps, presternum; pu, patella ulnaris; r, radius; rl, radiale; s, scapula; sc. 1 c, primary coracoid ray of scapulocoracoid; sc. 2 c, secondary coracoid ray of scapulocoracoid; scf, scapulocoracoid fenestra; sf, scapular fenestra; ss, suprascapula; sc. sr, scapular ray of scapulocoracoid; sr, sternal ribs; sucf, supracoracoid foramen; u, ulna; ul, ulnare; xs, xiphisternum." httpUri="https://zenodo.org/record/291233/files/figure.png" pageId="25" pageNumber="26">Fig. 8</figureCitation>
): The humerus is marginally longer than the radius and ulna. It is somewhat sigmoidal in dorsal view, with expanded proximal and distal ends. The proximal end is slightly less broad than the distal end. It possesses prominent humeral and deltopectoral crests (the latter with a sharp break separating it from the rest of the proximal humerus dorsally), as well as a moderately developed ectepicondylar crest and ectepicondyle. The bicipital fossa is deeply concave. The ectepicondylar foramen is visible in posterior view. In summary, it is fairly typical of gekkonids (
<bibRefCitation author="Russell AP &amp; Bauer AM." box="[1229,1477,1452,1479]" editor="Gans C" journalOrPublisher="Ithaca: Society for the Study of Amphibians and Reptiles" pageId="25" pageNumber="26" pagination="1 - 465" refString="Russell AP, Bauer AM. 2008. The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates. In: Gans C, Gaunt AS, Adler K, eds. The skull and appendicular locomotor apparatus of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 465." title="The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates" type="book chapter" volumeTitle="The skull and appendicular locomotor apparatus of Lepidosauria" year="2008">
<emphasis box="[1229,1477,1452,1479]" italics="true" pageId="25" pageNumber="26">Russell &amp; Bauer, 2008</emphasis>
</bibRefCitation>
).
</paragraph>
<paragraph blockId="25.[491,1542,228,1913]" pageId="25" pageNumber="26">
The radius is long and thin, slightly dorsoventrally flattened and weakly curved, with its distal articulatory facet with a distinct 
<emphasis box="[989,1207,1531,1557]" italics="true" pageId="25" pageNumber="26">processus styloideus</emphasis>
; its distal end articulates with the radiale posteriorly. The ulna is slender, dorsoventrally flattened, and straight, narrowing distally, but flaring at its distal end, where it articulates with the ulnare laterally and pisciform ventrally. The olecranon process is clearly distinct, and proximal to it, on the articular surface of the humerus, lies the sesamoid patella ulnaris, which is rounded. The internal face of the olecranon process forms a smooth sigmoid notch.
</paragraph>
<paragraph blockId="25.[491,1542,228,1913]" pageId="25" pageNumber="26">The spatium interosseum is formed by the diverging radius and ulna proximally and the ulnare, centrale, and radiale distally, rendering it roughly teardrop shaped.</paragraph>
<paragraph blockId="25.[491,1542,228,1913]" lastBlockId="26.[491,1542,228,1913]" lastPageId="26" lastPageNumber="27" pageId="25" pageNumber="26">
Nine carpal elements are present: the ulnare and radiale are subequal in size. The centrale is thin and long, and lies between these two elements. The pisciform is small and rounded, lies below the ulnare, and is probably not a true carpal (
<bibRefCitation author="Russell AP &amp; Bauer AM." box="[1150,1398,228,254]" editor="Gans C" journalOrPublisher="Ithaca: Society for the Study of Amphibians and Reptiles" pageId="26" pageNumber="27" pagination="1 - 465" refString="Russell AP, Bauer AM. 2008. The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates. In: Gans C, Gaunt AS, Adler K, eds. The skull and appendicular locomotor apparatus of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 465." title="The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates" type="book chapter" volumeTitle="The skull and appendicular locomotor apparatus of Lepidosauria" year="2008">
<emphasis box="[1150,1398,228,254]" italics="true" pageId="26" pageNumber="27">Russell &amp; Bauer, 2008</emphasis>
</bibRefCitation>
). A further five distal carpals are identifiable, the first in contact with phalange I, second with phalange II, third with phalanges II and III, fourth with phalanges III and IV, and fifth with phalange V. The phalangeal formula is 2-2-3-4-2.
</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" pageId="26" pageNumber="27">
The first phalange of the first finger, first of the second finger, first and second of the third finger, first through third of the fourth finger, and first of the fifth finger, are dorsoventrally flattened and laterally broadened. The terminal phalanges of each digit are slender and arcuate, ending in a laterally compressed, square tip with a distal claw-like projection, underlying the claws proper. The digits are able to hyperextend significantly. These three characters are presumably related to the adhesive pads of the fingers (
<bibRefCitation author="Russell AP &amp; Bauer AM." editor="Gans C" journalOrPublisher="Ithaca: Society for the Study of Amphibians and Reptiles" pageId="26" pageNumber="27" pagination="1 - 465" refString="Russell AP, Bauer AM. 2008. The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates. In: Gans C, Gaunt AS, Adler K, eds. The skull and appendicular locomotor apparatus of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 465." title="The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates" type="book chapter" volumeTitle="The skull and appendicular locomotor apparatus of Lepidosauria" year="2008">
<emphasis italics="true" pageId="26" pageNumber="27">Russell &amp; Bauer, 2008</emphasis>
</bibRefCitation>
). No ossified paraphalangeal elements are present in the micro-CT scans.
</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" pageId="26" pageNumber="27">
Pelvis (
<figureCitation box="[611,677,662,689]" captionStart="Figure 9" captionStartId="17.[507,571,1453,1475]" captionTargetBox="[492,1541,229,1408]" captionTargetId="figure@17.[491,1541,229,1408]" captionTargetPageId="17" captionText="Figure 9 Micro-CT images of pelvic girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal and (B) lateral view. Abbreviations: ac, acetabulum; ag, astragalocalcaneum; ep, epipubis; f, femur; fb, fibula; f. it, internal trochanter; f. itf, intertrochanteric fossa; f. fc, femoral condyle; hyi, hypoischium; il, ilium; il. pap, preacetabular process of ilium; is, ischium; is. t, ischiadic tuberosity; ltp, lenticular tibial patella; mt, metatarsals; of, obturator foramen; p, phalanges; pic, proischiadic cartilage; pb, pubis; pb. pl, processus lateralis of pubis; pc, post-cloacal bone; pt, pubic tubercle; tb, tibia; tf, thyroid fenestra." httpUri="https://zenodo.org/record/291234/files/figure.png" pageId="26" pageNumber="27">Fig. 9</figureCitation>
): The pelvis is composed of fused paired ilia, ischia, and pubes. The ischiopubic fenestra formed by the ischia and pubes is cardioid in shape, anteriorly rounded at the medial symphysis of the pubes in 
<taxonomicName box="[954,1093,741,768]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[954,1093,741,768]" italics="true" pageId="26" pageNumber="27">G. maculata</emphasis>
</taxonomicName>
, but more pointed in 
<taxonomicName box="[1348,1491,741,768]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1348,1491,741,768]" italics="true" pageId="26" pageNumber="27">G. megalepis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[491,628,781,807]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[491,628,781,807]" italics="true" pageId="26" pageNumber="27">G. humbloti</emphasis>
</taxonomicName>
—this fenestra may be medially divided by a proischiadic cartilage, but only the posterior-most portion of this element is shown in our micro-CT scans.
</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" pageId="26" pageNumber="27">
The pubis and ischia are broad and thin, concave in ventral view. The ilium is long, dorsoventrally broadened but laterally thin, and curves posteriorly. In lateral view (
<figureCitation box="[1444,1506,899,925]" captionStart="Figure 9" captionStartId="17.[507,571,1453,1475]" captionTargetBox="[492,1541,229,1408]" captionTargetId="figure@17.[491,1541,229,1408]" captionTargetPageId="17" captionText="Figure 9 Micro-CT images of pelvic girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal and (B) lateral view. Abbreviations: ac, acetabulum; ag, astragalocalcaneum; ep, epipubis; f, femur; fb, fibula; f. it, internal trochanter; f. itf, intertrochanteric fossa; f. fc, femoral condyle; hyi, hypoischium; il, ilium; il. pap, preacetabular process of ilium; is, ischium; is. t, ischiadic tuberosity; ltp, lenticular tibial patella; mt, metatarsals; of, obturator foramen; p, phalanges; pic, proischiadic cartilage; pb, pubis; pb. pl, processus lateralis of pubis; pc, post-cloacal bone; pt, pubic tubercle; tb, tibia; tf, thyroid fenestra." httpUri="https://zenodo.org/record/291234/files/figure.png" pageId="26" pageNumber="27">Fig. 9</figureCitation>
B), the iliac blade is reminiscent of the shoes of the Greek god Hermes—it rises posterodorsally, and is in broad medial contact with the pleurapophysis of the first sacral, and brief medial contact with the anterior portion of the pleurapophysis of the second sacral. Anterolaterally the ilium participates in the dorsal portion of the acetabulum.
</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" box="[523,1507,1097,1123]" pageId="26" pageNumber="27">The epipubic cartilage anterior to the medial pubic symphysis is somewhat calcified.</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" pageId="26" pageNumber="27">
The pubis curves from the anterior acetabulum ventrally and medially to the anterior symphysis with its contralateral at the front of the pelvis. It has a strong, medioventrally jutting pubic tubercle on the posterior portion of its lateral edge (see p. 138–145 in 
<bibRefCitation author="Russell AP &amp; Bauer AM." editor="Gans C" journalOrPublisher="Ithaca: Society for the Study of Amphibians and Reptiles" pageId="26" pageNumber="27" pagination="1 - 465" refString="Russell AP, Bauer AM. 2008. The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates. In: Gans C, Gaunt AS, Adler K, eds. The skull and appendicular locomotor apparatus of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 465." title="The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates" type="book chapter" volumeTitle="The skull and appendicular locomotor apparatus of Lepidosauria" year="2008">
<emphasis italics="true" pageId="26" pageNumber="27">Russell &amp; Bauer (2008)</emphasis>
</bibRefCitation>
for discussion of terminology), which descends more or less vertically in 
<taxonomicName box="[491,631,1294,1321]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[491,631,1294,1321]" italics="true" pageId="26" pageNumber="27">G. maculata</emphasis>
</taxonomicName>
and 
<taxonomicName box="[687,824,1294,1321]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[687,824,1294,1321]" italics="true" pageId="26" pageNumber="27">G. humbloti</emphasis>
</taxonomicName>
, but is posteriorly curved in 
<taxonomicName box="[1157,1299,1294,1321]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1157,1299,1294,1321]" italics="true" pageId="26" pageNumber="27">G. megalepis</emphasis>
</taxonomicName>
. Medial to this is the concavo-convex pubic apron, the anterolateral edge of which runs anteromedially toward the medial symphysis with the contralateral pubis. The relatively large obturator foramen lies posterior to the pelvic tubercle, in line with the medial edge of the acetabulum.
</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" box="[523,1440,1452,1478]" pageId="26" pageNumber="27">Dorsolaterally, the ischium forms the posterior margin of the acetabulum.</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" pageId="26" pageNumber="27">Ventromedially, it is roughly equal in breadth to the pubis, broadening toward the posteromedial symphysis with its contralateral at the back of the girdle. It possesses an almost lateral–pointing ischiadic tuberosity, rendering the posterolateral margin of the ischium deeply concave. The anterior margin of the ischium is also concave, extending anteriorly to form a weak prong, associated with the proischiadic cartilage. The medial ischial symphysis is not strongly fused, and the cartilaginous hypoischium likely extends into it.</paragraph>
<paragraph blockId="26.[491,1542,228,1913]" pageId="26" pageNumber="27">
A pair of curved post-cloacal bones is present in one specimen (ZSM 2126/2007; 
<figureCitation box="[491,557,1808,1834]" captionStart="Figure 9" captionStartId="17.[507,571,1453,1475]" captionTargetBox="[492,1541,229,1408]" captionTargetId="figure@17.[491,1541,229,1408]" captionTargetPageId="17" captionText="Figure 9 Micro-CT images of pelvic girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal and (B) lateral view. Abbreviations: ac, acetabulum; ag, astragalocalcaneum; ep, epipubis; f, femur; fb, fibula; f. it, internal trochanter; f. itf, intertrochanteric fossa; f. fc, femoral condyle; hyi, hypoischium; il, ilium; il. pap, preacetabular process of ilium; is, ischium; is. t, ischiadic tuberosity; ltp, lenticular tibial patella; mt, metatarsals; of, obturator foramen; p, phalanges; pic, proischiadic cartilage; pb, pubis; pb. pl, processus lateralis of pubis; pc, post-cloacal bone; pt, pubic tubercle; tb, tibia; tf, thyroid fenestra." httpUri="https://zenodo.org/record/291234/files/figure.png" pageId="26" pageNumber="27">Fig. 9</figureCitation>
), but is absent from all other specimens. These may be sexually dimorphic and/or age dependent; see 
<bibRefCitation author="Russell AP &amp; Vickaryous MK &amp; Bauer AM." box="[720,1121,1847,1874]" journalOrPublisher="Journal of Morphology" pageId="26" pageNumber="27" pagination="264 - 277" part="277" publicationUrl="10.1002/jmor.20494." refString="Russell AP, Vickaryous MK, Bauer AM. 2016. The phylogenetic distribution, anatomy and histology of the post-cloacal bones and adnexa of geckos. Journal of Morphology 277: 264 - 277 DOI 10.1002 / jmor. 20494." title="The phylogenetic distribution, anatomy and histology of the post-cloacal bones and adnexa of geckos" type="journal article" year="2016">
<emphasis box="[720,1121,1847,1874]" italics="true" pageId="26" pageNumber="27">Russell, Vickaryous &amp; Bauer (2016)</emphasis>
</bibRefCitation>
for a review of their phylogenetic distribution and evolution.
</paragraph>
<paragraph blockId="27.[491,1542,228,1087]" pageId="27" pageNumber="28">
Hindlimbs (
<figureCitation box="[668,734,228,254]" captionStart="Figure 9" captionStartId="17.[507,571,1453,1475]" captionTargetBox="[492,1541,229,1408]" captionTargetId="figure@17.[491,1541,229,1408]" captionTargetPageId="17" captionText="Figure 9 Micro-CT images of pelvic girdle of Geckolepis, based on G. megalepis sp. nov. (ZSM 2126 / 2007). (A) Dorsal and (B) lateral view. Abbreviations: ac, acetabulum; ag, astragalocalcaneum; ep, epipubis; f, femur; fb, fibula; f. it, internal trochanter; f. itf, intertrochanteric fossa; f. fc, femoral condyle; hyi, hypoischium; il, ilium; il. pap, preacetabular process of ilium; is, ischium; is. t, ischiadic tuberosity; ltp, lenticular tibial patella; mt, metatarsals; of, obturator foramen; p, phalanges; pic, proischiadic cartilage; pb, pubis; pb. pl, processus lateralis of pubis; pc, post-cloacal bone; pt, pubic tubercle; tb, tibia; tf, thyroid fenestra." httpUri="https://zenodo.org/record/291234/files/figure.png" pageId="27" pageNumber="28">Fig. 9</figureCitation>
): The femur is slender and weakly sigmoidal, with broad terminal ends. The epiphyseal internal trochanter is strong, and lies distal to the femoral condyle, from which it is separated by a deep notch. Its shape differs slightly among species: in 
<taxonomicName box="[491,634,347,373]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[491,634,347,373]" italics="true" pageId="27" pageNumber="28">G. megalepis</emphasis>
</taxonomicName>
, it is distinct and bulbous, in 
<taxonomicName box="[990,1129,347,373]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[990,1129,347,373]" italics="true" pageId="27" pageNumber="28">G. maculata</emphasis>
</taxonomicName>
it is ridge-like without a bulbous end, and in 
<taxonomicName box="[635,771,386,413]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="humbloti">
<emphasis box="[635,771,386,413]" italics="true" pageId="27" pageNumber="28">G. humbloti</emphasis>
</taxonomicName>
it is variable, with one specimen (ZSM 81/2006) resembling 
<taxonomicName class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="megalepis">
<emphasis italics="true" pageId="27" pageNumber="28">G. megalepis</emphasis>
</taxonomicName>
and one (ZSM 80/2010) resembling 
<taxonomicName box="[1022,1162,426,453]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[1022,1162,426,453]" italics="true" pageId="27" pageNumber="28">G. maculata</emphasis>
</taxonomicName>
. The ventral face of the proximal end of the femur has a moderately deep intertrochanteric fossa. The lateral distal condyle is distinctly larger than the medial one. The distal intercondylar groove is pronounced, and the popliteal fossa is not strongly deepened.
</paragraph>
<paragraph blockId="27.[491,1542,228,1087]" pageId="27" pageNumber="28">The fibula articulates via a sesamoid cyamella (=parafibula) with the lateral surface of the posterior femoral condyle. Additional sesamoids include the lenticular tibial patella dorsal to the distal end of the femur, a spherical post-axial ligament sesamoid (fabella), and the tibial lunula between the tibia and femur.</paragraph>
<paragraph blockId="27.[491,1542,228,1087]" pageId="27" pageNumber="28">
The fibula is laterally flattened to a slender rod of bone, with a slightly broader distal than proximal end. The tibia is broad and dorsoventrally compressed, and bows slightly outward. The cnemial and ventral crests are not pronounced and may be absent. Together the tibia and fibula articulate with the subtriangular astragalocalcaneum, which in turn articulates with the first metatarsal and the fourth distal tarsal Two distal tarsals are present (third and fourth; see 
<bibRefCitation author="Russell AP &amp; Bauer AM." box="[757,1019,942,968]" editor="Gans C" journalOrPublisher="Ithaca: Society for the Study of Amphibians and Reptiles" pageId="27" pageNumber="28" pagination="1 - 465" refString="Russell AP, Bauer AM. 2008. The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates. In: Gans C, Gaunt AS, Adler K, eds. The skull and appendicular locomotor apparatus of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 465." title="The appendicular locomotor apparatus of Sphenodon and normal-limbed squamates" type="book chapter" volumeTitle="The skull and appendicular locomotor apparatus of Lepidosauria" year="2008">
<emphasis box="[757,1019,942,968]" italics="true" pageId="27" pageNumber="28">Russell &amp; Bauer (2008)</emphasis>
</bibRefCitation>
for discussion of terminology). The fifth metatarsal is L-shaped and bears distinctly raised anterior and posterior plantar tubercles. The second metatarsal is longer than the fourth, and the third is the longest. The pedal phalangeal formula is 2-2-3-4-3.
</paragraph>
<paragraph blockId="27.[491,722,1132,1168]" box="[491,722,1132,1168]" pageId="27" pageNumber="28">
<heading allCaps="true" bold="true" box="[491,722,1132,1168]" fontSize="14" level="1" pageId="27" pageNumber="28" reason="0">
<emphasis bold="true" box="[491,722,1132,1168]" pageId="27" pageNumber="28">DISCUSSION</emphasis>
</heading>
</paragraph>
<paragraph blockId="27.[491,1541,1190,1733]" pageId="27" pageNumber="28">
<taxonomicName box="[491,725,1190,1216]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[491,725,1190,1216]" italics="true" pageId="27" pageNumber="28">Geckolepis megalepis</emphasis>
</taxonomicName>
is the first 
<taxonomicName box="[859,976,1190,1216]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[859,976,1190,1216]" italics="true" pageId="27" pageNumber="28">Geckolepis</emphasis>
</taxonomicName>
species to be described in 75 years (and it has been 123 years since the last currently recognised species was described). Although far northern Madagascar is relatively well surveyed for reptiles, and numerous 
<taxonomicName box="[1389,1506,1270,1296]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[1389,1506,1270,1296]" italics="true" pageId="27" pageNumber="28">Geckolepis</emphasis>
</taxonomicName>
populations have been recorded from this area, the new species has not been found outside the Ankarana massif. Considering the increasing number of squamate species putatively endemic to this spectacular limestone formation (
<bibRefCitation author="Glaw F &amp; Gehring P-S &amp; Kohler J &amp; FranZen M &amp; Vences M." box="[1200,1390,1388,1415]" journalOrPublisher="Salamandra" pageId="27" pageNumber="28" pagination="83 - 92" part="46" refString="Glaw F, Gehring P-S, Kohler J, FranZen M, Vences M. 2010. A new dwarf species of day gecko, genus Phelsuma, from the Ankarana pinnacle karst in northern Madagascar. Salamandra 46: 83 - 92." title="A new dwarf species of day gecko, genus Phelsuma, from the Ankarana pinnacle karst in northern Madagascar" type="journal article" year="2010">
<emphasis box="[1200,1390,1388,1415]" italics="true" pageId="27" pageNumber="28">Glaw et al., 2010</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Glaw F &amp; Kohler J &amp; Townsend TM &amp; Vences M." journalOrPublisher="PLOS ONE" pageId="27" pageNumber="28" pagination="31314" part="7" refString="Glaw F, Kohler J, Townsend TM, Vences M. 2012. Rivaling the world's smallest reptiles: discovery of miniaturized and microendemic new species of leaf chameleons (Brookesia) from northern Madagascar. PLOS ONE 7: e 31314" title="Rivaling the world's smallest reptiles: discovery of miniaturized and microendemic new species of leaf chameleons (Brookesia) from northern Madagascar" type="journal article" year="2012">
<emphasis italics="true" pageId="27" pageNumber="28">Glaw et al., 2012</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Jono T &amp; Bauer AM &amp; Brennan I &amp; Mori A." box="[561,741,1428,1454]" journalOrPublisher="Zootaxa" pageId="27" pageNumber="28" pagination="406 - 416" part="3980" publicationUrl="10.11646/zootaxa.3980.3.4." refString="Jono T, Bauer AM, Brennan I, Mori A. 2015. New species of Blaesodactylus (Squamata: Gekkonidae) from Tsingy karstic outcrops in Ankarana National Park, northern Madagascar. Zootaxa 3980: 406 - 416 DOI 10.11646 / zootaxa. 3980.3.4." title="New species of Blaesodactylus (Squamata: Gekkonidae) from Tsingy karstic outcrops in Ankarana National Park, northern Madagascar" type="journal article" year="2015">
<emphasis box="[561,741,1428,1454]" italics="true" pageId="27" pageNumber="28">Jono et al., 2015</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Ruane S &amp; Burbrink FT &amp; Randriamahatantsoa B &amp; Raxworthy CJ." box="[755,958,1428,1454]" journalOrPublisher="Copeia" pageId="27" pageNumber="28" pagination="712 - 721" part="104" publicationUrl="10.1643/CH-15-346." refString="Ruane S, Burbrink FT, Randriamahatantsoa B, Raxworthy CJ. 2016. The cat-eyed snakes of Madagascar: phylogeny and description of a new species of Madagascarophis (Serpentes: Lamprophiidae) from the Tsingy of Ankarana. Copeia 104: 712 - 721 DOI 10.1643 / CH- 15 - 346." title="The cat-eyed snakes of Madagascar: phylogeny and description of a new species of Madagascarophis (Serpentes: Lamprophiidae) from the Tsingy of Ankarana" type="journal article" year="2016">
<emphasis box="[755,958,1428,1454]" italics="true" pageId="27" pageNumber="28">Ruane et al., 2016</emphasis>
</bibRefCitation>
), it is likely that 
<taxonomicName box="[1156,1298,1428,1455]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[1156,1298,1428,1455]" italics="true" pageId="27" pageNumber="28">G. megalepis</emphasis>
</taxonomicName>
will also turn out to be microendemic to the region; unusual among 
<taxonomicName box="[1057,1174,1468,1494]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[1057,1174,1468,1494]" italics="true" pageId="27" pageNumber="28">Geckolepis</emphasis>
</taxonomicName>
species. In Ankarana, it occurs in sympatry with another lineage of 
<taxonomicName box="[916,1033,1507,1533]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[916,1033,1507,1533]" italics="true" pageId="27" pageNumber="28">Geckolepis</emphasis>
</taxonomicName>
(OTU G in 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[1178,1390,1507,1534]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="27" pageNumber="28" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[1178,1390,1507,1534]" italics="true" pageId="27" pageNumber="28">Lemme et al., 2013</emphasis>
</bibRefCitation>
), which also may be microendemic to this area, but to which it is only distantly related (uncorrected pairwise distance 11.3% in 12S rRNA according to sequences published by 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="27" pageNumber="28" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis italics="true" pageId="27" pageNumber="28">Lemme et al. (2013)</emphasis>
</bibRefCitation>
. The new species can be distinguished from these geckos by the lack of a dark stripe from eye to ear opening (vs. presence), possession of 17–18 scale rows at midbody (vs. 25–28), and possession of 27–31 ventral scale rows (vs. 33–43) (
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[1238,1450,1706,1732]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="27" pageNumber="28" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[1238,1450,1706,1732]" italics="true" pageId="27" pageNumber="28">Lemme et al., 2013</emphasis>
</bibRefCitation>
).
</paragraph>
<paragraph blockId="27.[491,1508,1764,1913]" box="[491,1377,1764,1795]" pageId="27" pageNumber="28">
<heading bold="true" box="[491,1377,1764,1795]" fontSize="12" level="2" pageId="27" pageNumber="28" reason="0">
<emphasis bold="true" box="[491,1377,1764,1795]" pageId="27" pageNumber="28">
Extreme integumentary autotomy in 
<taxonomicName box="[1044,1377,1764,1795]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="megalepis">
<emphasis bold="true" box="[1044,1377,1764,1795]" italics="true" pageId="27" pageNumber="28">Geckolepis megalepis</emphasis>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph blockId="27.[491,1508,1764,1913]" lastBlockId="28.[491,1542,228,1913]" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28">
Many reptiles have evolved the ability to shed some part of their body in response to predator attack. The most widespread form is caudal autotomy, the shedding of all or part of the tail, which is widespread among Lepidosauria, from amphisbaenians to rhynchocephalians, even being found in some snakes (
<bibRefCitation author="Arnold EN." box="[1133,1280,228,254]" journalOrPublisher="Journal of Natural History" pageId="28" pageNumber="29" pagination="127 - 169" part="18" publicationUrl="10.1080/00222938400770131." refString="Arnold EN. 1984. Evolutionary aspects of tail shedding in lizards and their relatives. Journal of Natural History 18: 127 - 169 DOI 10.1080 / 00222938400770131." title="Evolutionary aspects of tail shedding in lizards and their relatives" type="journal article" year="1984">
<emphasis box="[1133,1280,228,254]" italics="true" pageId="28" pageNumber="29">Arnold, 1984</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Bateman PW &amp; Fleming PA." journalOrPublisher="Journal of Zoology" pageId="28" pageNumber="29" pagination="1 - 14" part="277" publicationUrl="10.1111/j.1469-7998.2008.00484.x." refString="Bateman PW, Fleming PA. 2009. To cut a long tail short: a review of lizard caudal autotomy studies carried out over the last 20 years. Journal of Zoology 277: 1 - 14 DOI 10.1111 / j. 1469 - 7998.2008.00484. x." title="To cut a long tail short: a review of lizard caudal autotomy studies carried out over the last 20 years" type="journal article" year="2009">
<emphasis italics="true" pageId="28" pageNumber="29">Bateman &amp; Fleming, 2009</emphasis>
</bibRefCitation>
). 
<taxonomicName box="[571,688,267,293]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis box="[571,688,267,293]" italics="true" pageId="28" pageNumber="29">Geckolepis</emphasis>
</taxonomicName>
species are also able to shed their tails, and indeed few specimens survive to adulthood with their original tails intact (see for instance 
<figureCitation box="[1195,1268,307,333]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="28" pageNumber="29">Figs. 3</figureCitation>
A and 3B). In addition, these geckos have evolved an even more extreme adaptation, i.e. the autotomy of virtually their entire integument when seized or even touched. Earlier studies have shown that the autotomized layers include epidermis, underlying connective tissue, and subcutaneous fat tissue, and that a layer between the integument and the underlying tissue represents a pre-formed splitting zone (
<bibRefCitation author="Schubert C &amp; Christophers E." box="[812,1156,504,531]" journalOrPublisher="Zoologischer Anzeiger" pageId="28" pageNumber="29" pagination="129 - 141" part="214" refString="Schubert C, Christophers E. 1985. '' Dermolytische Schreckhautung' ' - ein besonderes Autotomieverhalten von Geckolepis typica (Reptilia, Gekkonidae). Zoologischer Anzeiger 214: 129 - 141." title="'' Dermolytische Schreckhautung' ' - ein besonderes Autotomieverhalten von Geckolepis typica (Reptilia, Gekkonidae)" type="journal article" year="1985">
<emphasis box="[812,1156,504,531]" italics="true" pageId="28" pageNumber="29">Schubert &amp; Christophers, 1985</emphasis>
</bibRefCitation>
). The shedding process is most likely achieved by contraction of the network of myofibroblasts in the preformed splitting zone, with vasoconstriction in the most superficial vasculature of the dermis to avoid bleeding (
<bibRefCitation author="Schubert C &amp; Christophers E." box="[610,954,623,649]" journalOrPublisher="Zoologischer Anzeiger" pageId="28" pageNumber="29" pagination="129 - 141" part="214" refString="Schubert C, Christophers E. 1985. '' Dermolytische Schreckhautung' ' - ein besonderes Autotomieverhalten von Geckolepis typica (Reptilia, Gekkonidae). Zoologischer Anzeiger 214: 129 - 141." title="'' Dermolytische Schreckhautung' ' - ein besonderes Autotomieverhalten von Geckolepis typica (Reptilia, Gekkonidae)" type="journal article" year="1985">
<emphasis box="[610,954,623,649]" italics="true" pageId="28" pageNumber="29">Schubert &amp; Christophers, 1985</emphasis>
</bibRefCitation>
). This process is thus completely different from the normal skin shedding of squamate reptiles, which leads to a loss of keratinized epidermis only (
<bibRefCitation author="Schubert C &amp; Christophers E." box="[562,906,702,728]" journalOrPublisher="Zoologischer Anzeiger" pageId="28" pageNumber="29" pagination="129 - 141" part="214" refString="Schubert C, Christophers E. 1985. '' Dermolytische Schreckhautung' ' - ein besonderes Autotomieverhalten von Geckolepis typica (Reptilia, Gekkonidae). Zoologischer Anzeiger 214: 129 - 141." title="'' Dermolytische Schreckhautung' ' - ein besonderes Autotomieverhalten von Geckolepis typica (Reptilia, Gekkonidae)" type="journal article" year="1985">
<emphasis box="[562,906,702,728]" italics="true" pageId="28" pageNumber="29">Schubert &amp; Christophers, 1985</emphasis>
</bibRefCitation>
) The scarless regeneration of the whole integument occurs within a few weeks, apparently starting from stem cells of the deeper layers of the connecting tissue and is considered as unique among vertebrates (
<bibRefCitation author="Schubert C &amp; Steffen T &amp; Christophers E." journalOrPublisher="Zoologischer Anzeiger" pageId="28" pageNumber="29" pagination="175 - 192" part="224" refString="Schubert C, Steffen T, Christophers E. 1990. Weitere Beobachtungen zur '' dermolytischen Schreckhautung' ' bei Geckolepis typica (Reptilia, Gekkonidae). Zoologischer Anzeiger 224: 175 - 192." title="Weitere Beobachtungen zur '' dermolytischen Schreckhautung' ' bei Geckolepis typica (Reptilia, Gekkonidae)" type="journal article" year="1990">
<emphasis italics="true" pageId="28" pageNumber="29">Schubert, Steffen &amp; Christophers, 1990</emphasis>
</bibRefCitation>
). Superficially, no differences are apparent between regenerated and original scales, due to the irregularity of scalation patterns and some variability in scale size. The same is true for regenerated tails; indeed, it is often hard to be certain that a 
<taxonomicName box="[1424,1541,899,925]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis box="[1424,1541,899,925]" italics="true" pageId="28" pageNumber="29">Geckolepis</emphasis>
</taxonomicName>
tail has been regenerated without X-ray images showing that the vertebrae are absent.
</paragraph>
<paragraph blockId="28.[491,1542,228,1913]" pageId="28" pageNumber="29">
The new species is remarkable in the possession of proportionally larger scales than any of its congeners (especially in the dorsal cervical region, see 
<figureCitation box="[1185,1260,1018,1044]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="28" pageNumber="29">Figs. 3</figureCitation>
and 
<figureCitation box="[1317,1331,1018,1044]" captionStart="Figure 4" captionStartId="7.[507,571,1752,1774]" captionTargetBox="[491,1512,230,1704]" captionTargetId="figure@7.[491,1514,229,1708]" captionTargetPageId="7" captionText="Figure 4 Specimens of Geckolepis megalepis sp. nov. in preservative. (A – C) The holotype, ZSM 2126 / 2007; (D – F) paratype ZSM 232 / 2016. Shown in dorsal (A, D) and ventral (C, F) view, with a close-up view of the postmental scales (B, E) coloured for reference to Fig. 1. Scale bars indicate 10 mm. Chin insets are not to scale." httpUri="https://zenodo.org/record/291229/files/figure.png" pageId="28" pageNumber="29">4</figureCitation>
). Midbody dorsal scales of 
<taxonomicName box="[595,828,1057,1083]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[595,828,1057,1083]" italics="true" pageId="28" pageNumber="29">Geckolepis megalepis</emphasis>
</taxonomicName>
are 7.3–8.3% of the SVL (by comparison, a typical midbody dorsal scale of the holotype of 
<taxonomicName box="[856,995,1097,1123]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[856,995,1097,1123]" italics="true" pageId="28" pageNumber="29">G. maculata</emphasis>
</taxonomicName>
is just 5.4% of its SVL). Indeed as far as we have been able to determine, 
<taxonomicName box="[836,979,1136,1163]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[836,979,1136,1163]" italics="true" pageId="28" pageNumber="29">G. megalepis</emphasis>
</taxonomicName>
has the largest mid-body scales of any 
<taxonomicName box="[1437,1504,1136,1162]" class="Reptilia" family="Gekkonidae" genus="Gekko" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="gecko">gecko</taxonomicName>
in both relative and absolute terms, as its scales outstrip those of all known congeners, and only 
<taxonomicName box="[550,702,1216,1241]" class="Reptilia" family="Sphaerodactylidae" genus="Teratoscincus" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis box="[550,702,1216,1241]" italics="true" pageId="28" pageNumber="29">Teratoscincus</emphasis>
</taxonomicName>
may approach 
<taxonomicName box="[884,1001,1215,1241]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis box="[884,1001,1215,1241]" italics="true" pageId="28" pageNumber="29">Geckolepis</emphasis>
</taxonomicName>
in scale size. Remarkably, the latter genus has similarly fragile skin (
<bibRefCitation author="Bauer AM &amp; Russell AP &amp; Shadwick RE." box="[741,1111,1255,1281]" journalOrPublisher="Amphibia-Reptilia" pageId="28" pageNumber="29" pagination="321 - 331" part="14" refString="Bauer AM, Russell AP, Shadwick RE. 1993. Skin mechanics and morphology of the gecko Teratoscincus scincus. Amphibia-Reptilia 14: 321 - 331" title="Skin mechanics and morphology of the gecko Teratoscincus scincus" type="journal article" year="1993">
<emphasis box="[741,1111,1255,1281]" italics="true" pageId="28" pageNumber="29">Bauer, Russell &amp; Shadwick, 1993</emphasis>
</bibRefCitation>
), which may have evolved for similar antipredatory function.
</paragraph>
<paragraph blockId="28.[491,1542,228,1913]" box="[523,1496,1334,1360]" pageId="28" pageNumber="29">
The exceptionally large scales of 
<taxonomicName box="[916,1058,1334,1360]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[916,1058,1334,1360]" italics="true" pageId="28" pageNumber="29">G. megalepis</emphasis>
</taxonomicName>
lead to questions about the possible
</paragraph>
<paragraph blockId="28.[491,1542,228,1913]" lastBlockId="29.[491,1542,228,892]" lastPageId="29" lastPageNumber="30" pageId="28" pageNumber="29">
advantage of larger scales for species with autotomizable integument. As is visible in 
<figureCitation box="[1453,1514,1373,1399]" captionStart="Figure 3" captionStartId="6.[507,571,1215,1237]" captionTargetBox="[491,1541,229,1170]" captionTargetId="figure@6.[491,1541,229,1170]" captionTargetPageId="6" captionText="Figure 3 Specimens of Geckolepis megalepis sp. nov. in life. (A) A specimen observed by A. Anker (photograph used with permission); (B) a specimen observed by FG, and (C) a specimen photographed after scale loss, with inset indicating the transparent ‘ tear zone’ at the base of a scale. None of the photographed animals were collected, but their attribution to G. megalepis is clear on the basis of the large size of their scales. Note that the tails of all three specimens are regenerated." httpUri="https://zenodo.org/record/291381/files/figure.png" pageId="28" pageNumber="29">Fig. 3</figureCitation>
C, the large, imbricate body scales of 
<taxonomicName box="[888,1121,1413,1439]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[888,1121,1413,1439]" italics="true" pageId="28" pageNumber="29">Geckolepis megalepis</emphasis>
</taxonomicName>
are attached to the integument only superficially by a narrow transparent zone which covers less than 20% of the scale’s edge surface. 
<bibRefCitation author="Schmidt WJ." box="[588,764,1492,1518]" editor="Voeltzkow A" journalOrPublisher="Stuttgart: Schweizerbart'sche Verlagsbuchhandlung" pageId="28" pageNumber="29" pagination="331 - 352" refString="Schmidt WJ. 1911. Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden. In: Voeltzkow A, ed. Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung, 331 - 352." title="Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden" type="book chapter" volumeTitle="Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow" year="1911">
<emphasis box="[588,764,1492,1518]" italics="true" pageId="28" pageNumber="29">Schmidt (1911)</emphasis>
</bibRefCitation>
called this region the ‘Anwachsfläche’ (literally ‘attachment area’), and noted that it is much smaller relative to the size of the scales of 
<taxonomicName box="[1266,1383,1531,1557]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis box="[1266,1383,1531,1557]" italics="true" pageId="28" pageNumber="29">Geckolepis</emphasis>
</taxonomicName>
than in other lizards. With increasing scale diameter, the circumference of the scale and therefore its zone of connectivity increases linearly while the area of the scales increases exponentially (approximating the scales to a circular shape). This increases the surface area and therefore force of friction on scales exponentially while the ‘tear zone’ of the scales increases linearly, meaning that there is a smaller tear zone per unit area with greater scale size. Thus, less force, applied in a posterior or lateral direction, should be required to remove a larger scale than a smaller one. Additionally, as the scales are imbricate, leverage may play a role: an anteriorly directed pressure on the scales may cause them to lift and detach, much as one might detach a sticky note from a surface. The leverage of a longer scale is greater than a shorter one, and these may therefore lift more easily. This is probably further enhanced by the steeper angle of larger scales to the body (
<bibRefCitation author="Schmidt WJ." box="[1055,1216,268,294]" editor="Voeltzkow A" journalOrPublisher="Stuttgart: Schweizerbart'sche Verlagsbuchhandlung" pageId="29" pageNumber="30" pagination="331 - 352" refString="Schmidt WJ. 1911. Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden. In: Voeltzkow A, ed. Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung, 331 - 352." title="Beobachtungen an der Haut von Geckolepis und einigen anderen Geckoniden" type="book chapter" volumeTitle="Reise in Ostafrika in den Jahren 1903 - 1905 mit Mitteln der Hermann und Elise geb. Heckmann Wentzel-Stiftung ausgefuhrt: Wissenschaftliche Ergebnisse von Alfred Voeltzkow" year="1911">
<emphasis box="[1055,1216,268,294]" italics="true" pageId="29" pageNumber="30">Schmidt, 1911</emphasis>
</bibRefCitation>
). Together, these principles may explain why the scales of 
<taxonomicName box="[854,997,308,334]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="species" species="megalepis">
<emphasis box="[854,997,308,334]" italics="true" pageId="29" pageNumber="30">G. megalepis</emphasis>
</taxonomicName>
appear to come off more easily than those of other 
<taxonomicName box="[594,711,347,373]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[594,711,347,373]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
species, but further studies are clearly necessary to confirm or reject this hypothesis.
</paragraph>
<paragraph blockId="29.[491,1542,228,892]" pageId="29" pageNumber="30">
Although it is highly plausible to interpret their ability of scale autotomy as an antipredator defence mechanism, direct observations of predation events or attempts on 
<taxonomicName box="[491,608,507,533]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[491,608,507,533]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
individuals are scarce and include only a scorpion (
<taxonomicName box="[1233,1468,507,533]" class="Arachnida" family="Buthidae" genus="Grosphus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Scorpiones" pageId="29" pageNumber="30" phylum="Arthropoda" rank="species" species="flavopiceus">
<emphasis box="[1233,1468,507,533]" italics="true" pageId="29" pageNumber="30">Grosphus flavopiceus</emphasis>
</taxonomicName>
), a bird (
<taxonomicName box="[560,770,547,573]" class="Aves" family="Dicruridae" genus="Dicrurus" kingdom="Animalia" order="Passeriformes" pageId="29" pageNumber="30" phylum="Chordata" rank="species" species="forficatus">
<emphasis box="[560,770,547,573]" italics="true" pageId="29" pageNumber="30">Dicrurus forficatus</emphasis>
</taxonomicName>
) and a large nocturnal 
<taxonomicName box="[1052,1215,547,573]" class="Reptilia" family="Gekkonidae" genus="Blaesodactylus" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[1052,1215,547,573]" italics="true" pageId="29" pageNumber="30">Blaesodactylus</emphasis>
</taxonomicName>
<taxonomicName box="[1223,1290,547,573]" class="Reptilia" family="Gekkonidae" genus="Gekko" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="species" species="gecko">gecko</taxonomicName>
(
<bibRefCitation author="Gardner CJ &amp; Jasper L." journalOrPublisher="Malagasy Nature" pageId="29" pageNumber="30" pagination="105 - 106" part="8" refString="Gardner CJ, Jasper L. 2014. A record of vertebrate carnivory by the Crested Drongo (Dicrurus forficatus). Malagasy Nature 8: 105 - 106." title="A record of vertebrate carnivory by the Crested Drongo (Dicrurus forficatus)" type="journal article" year="2014">
<emphasis italics="true" pageId="29" pageNumber="30">Gardner &amp; Jasper, 2014</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Gardner CJ &amp; Jasper L." box="[561,828,586,613]" journalOrPublisher="Herpetology Notes" pageId="29" pageNumber="30" pagination="479 - 481" part="8" refString="Gardner CJ, Jasper L. 2015. A fish-scaled gecko (Geckolepis sp.) escapes predation by a velvet gecko (Blaesodactylus sp.) through skin shedding. Herpetology Notes 8: 479 - 481." title="A fish-scaled gecko (Geckolepis sp.) escapes predation by a velvet gecko (Blaesodactylus sp.) through skin shedding" type="journal article" year="2015">
<emphasis box="[561,828,586,613]" italics="true" pageId="29" pageNumber="30">Gardner &amp; Jasper, 2015</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Glaw F &amp; Vences M &amp; Lourenco WR." box="[843,1200,586,613]" journalOrPublisher="Herpetological Review" pageId="29" pageNumber="30" pagination="135" part="33" refString="Glaw F, Vences M, Lourenco WR. 2002. Geckolepis maculata (Spotted Fishscale Gecko) - Predation. Herpetological Review 33: 135." title="Geckolepis maculata (Spotted Fishscale Gecko) - Predation" type="journal article" year="2002">
<emphasis box="[843,1200,586,613]" italics="true" pageId="29" pageNumber="30">Glaw, Vences &amp; Lourenco, 2002</emphasis>
</bibRefCitation>
). In the lattermost case, the 
<taxonomicName box="[491,608,626,652]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[491,608,626,652]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
individual slipped from the mouth of the 
<taxonomicName box="[1117,1280,626,652]" class="Reptilia" family="Gekkonidae" genus="Blaesodactylus" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[1117,1280,626,652]" italics="true" pageId="29" pageNumber="30">Blaesodactylus</emphasis>
</taxonomicName>
ca. 30 s after being captured, and escaped denuded (
<bibRefCitation author="Gardner CJ &amp; Jasper L." box="[874,1141,666,693]" journalOrPublisher="Herpetology Notes" pageId="29" pageNumber="30" pagination="479 - 481" part="8" refString="Gardner CJ, Jasper L. 2015. A fish-scaled gecko (Geckolepis sp.) escapes predation by a velvet gecko (Blaesodactylus sp.) through skin shedding. Herpetology Notes 8: 479 - 481." title="A fish-scaled gecko (Geckolepis sp.) escapes predation by a velvet gecko (Blaesodactylus sp.) through skin shedding" type="journal article" year="2015">
<emphasis box="[874,1141,666,693]" italics="true" pageId="29" pageNumber="30">Gardner &amp; Jasper, 2015</emphasis>
</bibRefCitation>
), thereby providing the first direct evidence of successful escape by skin shedding. Further studies on the role of dermolytic scale autotomy by 
<taxonomicName box="[712,829,746,772]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[712,829,746,772]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
are clearly needed, in order to understand its functionality with a range of predators (its function against snakes, for instance, has not yet been observed), and to understand the pressures driving the evolution of greater scale size in this particular lineage of 
<taxonomicName box="[782,899,865,891]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[782,899,865,891]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="29.[491,1542,926,1910]" pageId="29" pageNumber="30">
<heading bold="true" fontSize="12" level="2" pageId="29" pageNumber="30" reason="0">
<emphasis bold="true" box="[491,1479,926,957]" pageId="29" pageNumber="30">
The osteology of 
<taxonomicName box="[755,924,926,957]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[755,924,926,957]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
and the next steps in resolving their
</emphasis>
<emphasis bold="true" box="[491,640,963,994]" pageId="29" pageNumber="30">taxonomy</emphasis>
</heading>
</paragraph>
<paragraph blockId="29.[491,1542,926,1910]" pageId="29" pageNumber="30">
Our osteological description of the genus 
<taxonomicName box="[969,1086,1007,1033]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[969,1086,1007,1033]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
, based on six specimens of four species, reveals strong morphological conservatism in this genus. Few characters show potentially diagnostic differences at the species level (see Appendix S2), and the degree of intra-specific variation is apparently quite high. However, through the use of micro-CT, we were able to include the holotype of 
<taxonomicName box="[771,911,1166,1192]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[771,911,1166,1192]" italics="true" pageId="29" pageNumber="30">G. maculata</emphasis>
</taxonomicName>
in our skeletal description. This will be an important step toward the resolution of its identity, despite our continued failure to trace its type locality (see Supplemental Information). 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[978,1204,1246,1272]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="29" pageNumber="30" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[978,1204,1246,1272]" italics="true" pageId="29" pageNumber="30">Lemme et al. (2013)</emphasis>
</bibRefCitation>
assigned their OTU AB to this species on the basis of its overall similar morphology, but our analysis of their skulls suggests that they are probably not conspecific. This means that the assignment of this name is still completely unknown; it belongs either to another of the known clades, or to one not yet characterised. A broader survey of the osteology of the genus will be required to resolve the identity of this species, and will in turn yield the total resolution of this genus.
</paragraph>
<paragraph blockId="29.[491,1542,926,1910]" lastBlockId="30.[491,1542,228,1051]" lastPageId="30" lastPageNumber="31" pageId="29" pageNumber="30">
Despite the detailed osteological description, we admit that 
<taxonomicName box="[1232,1349,1485,1511]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis box="[1232,1349,1485,1511]" italics="true" pageId="29" pageNumber="30">Geckolepis</emphasis>
</taxonomicName>
represents an extremely difficult taxonomic group that is hard to characterise. The high variability in scale number and enhanced ability to shed scales upon capture has misled taxonomists in the past into believing they were dealing with distinct new species, which have subsequently been synonymised (
<bibRefCitation author="Angel F." box="[727,860,1644,1670]" journalOrPublisher="Memoires de l'Academie Malgache" pageId="29" pageNumber="30" pagination="1 - 139" part="36" refString="Angel F. 1942. Les lezards de Madagascar. Memoires de l'Academie Malgache 36: 1 - 139." title="Les lezards de Madagascar" type="journal article" year="1942">
<emphasis box="[727,860,1644,1670]" italics="true" pageId="29" pageNumber="30">Angel, 1942</emphasis>
</bibRefCitation>
; 
<bibRefCitation author="Kohler G &amp; Diethert H-H &amp; Nussbaum RA &amp; Raxworthy CJ." box="[875,1080,1644,1670]" journalOrPublisher="Herpetologica" pageId="29" pageNumber="30" pagination="419 - 435" part="65" publicationUrl="10.1655/08-059.1." refString="Kohler G, Diethert H-H, Nussbaum RA, Raxworthy CJ. 2009. A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros. Herpetologica 65: 419 - 435 DOI 10.1655 / 08 - 059.1." title="A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros" type="journal article" year="2009">
<emphasis box="[875,1080,1644,1670]" italics="true" pageId="29" pageNumber="30">Köhler et al., 2009</emphasis>
</bibRefCitation>
). The trouble is further enhanced by multiple genetic lineages occurring in sympatry (
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[1070,1282,1684,1710]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="29" pageNumber="30" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[1070,1282,1684,1710]" italics="true" pageId="29" pageNumber="30">Lemme et al., 2013</emphasis>
</bibRefCitation>
; 
<figureCitation box="[1297,1362,1684,1710]" captionStart="Figure 1" captionStartId="2.[507,571,950,972]" captionTargetBox="[1198,1540,229,885]" captionTargetPageId="2" captionText="Figure 1 Molecular phylogeny and distribution of Geckolepis OTUs based mainly on Lemme et al. (2013). Dotted lines in the phylogeny indicate uncertain placement. The phylogenetic position and distribution of Geckolepis humbloti is inferred from Hawlitschek et al. (2016)." httpUri="https://zenodo.org/record/291226/files/figure.svg" pageId="29" pageNumber="30">Fig. 1</figureCitation>
), and further still by apparent osteological conservatism. However, we were able to show that this is not always the case, and members of the AB OTU of 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." box="[1132,1358,1764,1790]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="29" pageNumber="30" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis box="[1132,1358,1764,1790]" italics="true" pageId="29" pageNumber="30">Lemme et al. (2013)</emphasis>
</bibRefCitation>
for instance show strong osteological differences that will facilitate its description. Nevertheless, the osteology did not provide as many taxonomic characters as we had hoped. Admittedly our sample size is small, and therefore practically no data yet exist on the degree of inter- and intraspecific osteological variation in these geckos. Examination of many further specimens and other lineages will enhance our ability to use osteology as a source of characters in their taxonomic resolution.
</paragraph>
<paragraph blockId="30.[491,1542,228,1051]" pageId="30" pageNumber="31">
Thus, the next steps are now clear: (1) a survey of osteology in this genus in the context of molecular phylogenetic relationships of OTUs, and (2) a survey of intraspecific and sexual skeletal variability in at least one lineage, although this is generally minimal among gekkotans (
<emphasis box="[625,814,467,493]" italics="true" pageId="30" pageNumber="31">Daza et al., 2008</emphasis>
; 
<emphasis box="[826,1016,467,493]" italics="true" pageId="30" pageNumber="31">Daza et al., 2009</emphasis>
). Based on this data and corresponding other datasets, we must establish with a high degree of certainty which OTU from 
<bibRefCitation author="Lemme I &amp; Erbacher M &amp; Kaffenberger N &amp; Vences M &amp; Kohler J." journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="30" pageNumber="31" pagination="87 - 95" part="13" publicationUrl="10.1007/s13127-012-0098-y." refString="Lemme I, Erbacher M, Kaffenberger N, Vences M, Kohler J. 2013. Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis. Organisms Diversity &amp; Evolution 13: 87 - 95 DOI 10.1007 / s 13127 - 012 - 0098 - y." title="Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis" type="journal article" year="2013">
<emphasis italics="true" pageId="30" pageNumber="31">Lemme et al. (2013)</emphasis>
</bibRefCitation>
really corresponds to 
<taxonomicName box="[831,970,547,573]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="30" pageNumber="31" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[831,970,547,573]" italics="true" pageId="30" pageNumber="31">G. maculata</emphasis>
</taxonomicName>
(if any). Once this information is gathered, we may proceed with the resolution of the taxonomy of the genus.
</paragraph>
<paragraph blockId="30.[491,1542,228,1051]" pageId="30" pageNumber="31">
The framework of an integrative dataset composed of morphological, meristic, molecular phylogenetic, and osteological data has considerable potential for dealing with species complexes in squamates—even those as tortuous as 
<taxonomicName box="[1115,1232,706,732]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis box="[1115,1232,706,732]" italics="true" pageId="30" pageNumber="31">Geckolepis</emphasis>
</taxonomicName>
, and certainly in other species with fragile skin. However, it is clear that the robustness of conclusions strongly depends on the available sample size. In instances, such as this one, where sample size is limited to a low number of specimens, any osteological, morphological, or pholidotic feature identified as differing must first be highlighted as being potentially diagnostic, until more data becomes available to verify the value of each of these features. Nevertheless, the value of these data, especially when they are extracted from holotypes and old specimens in a non-destructive way, cannot be overstated. Micro-CT is therefore likely to have a pivotal role in resolving many difficult species complexes.
</paragraph>
</subSubSection>
</treatment>
</document>