treatments-xml/data/CC/65/87/CC6587F70811FFF19BC0DE629F5DFE90.xml

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<mods:title id="CA928FDB093DFBD2A4278B65CA5DAA68">A molecular phylogeny of the African plated lizards, genus Gerrhosaurus Wiegmann, 1828 (Squamata: Gerrhosauridae), with the description of two new genera</mods:title>
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<mods:namePart id="0D88F715B7B657EDFD848B388A765C39">Tolley, Krystal A.</mods:namePart>
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<paragraph id="447336E10811FFE29BC0DE629C59F846" blockId="11.[151,537,1915,1942]" box="[151,537,1915,1942]" pageId="11" pageNumber="476">
<heading id="1F3B818D0811FFE29BC0DE629C59F846" bold="true" box="[151,537,1915,1942]" fontSize="11" level="1" pageId="11" pageNumber="476" reason="1">
<taxonomicName id="83CC4D620811FFE29BC0DE629C59F846" authority="Wiegmann, 1828" authorityName="Wiegmann" authorityYear="1828" box="[151,537,1915,1942]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="476" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30811FFE29BC0DE629C59F846" bold="true" box="[151,537,1915,1942]" pageId="11" pageNumber="476">
<emphasis id="76B8EAF30811FFE29BC0DE629F00F845" bold="true" box="[151,320,1915,1941]" italics="true" pageId="11" pageNumber="476">Gerrhosaurus</emphasis>
Wiegmann, 1828
</emphasis>
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<paragraph id="447336E10811FFE59BC0DEDB9F9DFF7E" blockId="11.[151,415,1986,2008]" lastBlockId="12.[151,477,152,174]" lastPageId="12" lastPageNumber="477" pageId="11" pageNumber="476">
<taxonomicName id="83CC4D620811FFE29BC0DEDB9FDFF808" authority="Smith, 1837" authorityName="Smith" authorityYear="1837" box="[151,415,1986,2008]" class="Squamata" family="Gerrhosauridae" genus="Pleurotuchus" higherTaxonomySource="CoL" kingdom="Animalia" pageId="11" pageNumber="476" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30811FFE29BC0DEDB9F5DF808" box="[151,285,1986,2008]" italics="true" pageId="11" pageNumber="476">Pleurotuchus</emphasis>
Smith, 1837
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<taxonomicName id="83CC4D620816FFE59BC0D9819F9DFF7E" authority="FitzSimons, 1953" authorityName="FitzSimons" authorityYear="1953" box="[151,477,152,174]" class="Reptilia" family="Gerrhosauridae" genus="Angolosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30816FFE59BC0D9819F64FF7E" box="[151,292,152,174]" italics="true" pageId="12" pageNumber="477">Angolosaurus</emphasis>
FitzSimons, 1953
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<subSubSection id="0CD6656A0816FFE59BC0D9C39F39FE53" pageId="12" pageNumber="477" type="materials_examined">
<paragraph id="447336E10816FFE59BC0D9C39D66FF23" blockId="12.[151,1436,218,675]" box="[151,806,218,243]" pageId="12" pageNumber="477">
<emphasis id="76B8EAF30816FFE59BC0D9C39F74FF23" bold="true" box="[151,308,218,243]" pageId="12" pageNumber="477">
<typeStatus id="9B7788430816FFE59BC0D9C39E92FF23" box="[151,210,218,243]" pageId="12" pageNumber="477">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="83CC4D620816FFE59A6CD9C39D66FF23" authority="Wiegmann, 1828" authorityName="Wiegmann" authorityYear="1828" box="[315,806,218,243]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30816FFE59A6CD9C39C20FF23" box="[315,608,218,243]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus flavigularis</emphasis>
Wiegmann, 1828
</taxonomicName>
</paragraph>
<paragraph id="447336E10816FFE59B90D9E79F39FE53" blockId="12.[151,1436,218,675]" pageId="12" pageNumber="477">
<emphasis id="76B8EAF30816FFE59B90D9E79F71FEC7" bold="true" box="[199,305,254,279]" pageId="12" pageNumber="477">Content:</emphasis>
<taxonomicName id="83CC4D620816FFE59A6BD9E79D78FEC7" authority="Wiegmann, 1828" authorityName="Wiegmann" authorityYear="1828" box="[316,824,254,279]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30816FFE59A6BD9E79C25FEC7" box="[316,613,254,279]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus flavigularis</emphasis>
Wiegmann, 1828
</taxonomicName>
; 
<taxonomicName id="83CC4D620816FFE5981ED9E79AAFFEC7" authority="Smith, 1837" authorityName="Smith" authorityYear="1837" box="[841,1263,254,279]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="typicus">
<emphasis id="76B8EAF30816FFE5981ED9E79A00FEC6" box="[841,1088,254,279]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus typicus</emphasis>
(Smith, 1837)
</taxonomicName>
; 
<taxonomicName id="83CC4D620816FFE59FA8D9E79FACFEEB" authority="Hallowell, 1857" authorityName="Hallowell" authorityYear="1857" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30816FFE59FA8D9E79F6CFEEB" italics="true" pageId="12" pageNumber="477">Gerrhosaurus nigrolineatus</emphasis>
Hallowell, 1857
</taxonomicName>
; 
<taxonomicName id="83CC4D620816FFE59AAED83B9D97FEEB" authority="Bocage, 1866" authorityName="Bocage" authorityYear="1866" box="[505,983,290,315]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF30816FFE59AAED83B9D70FEEB" box="[505,816,290,315]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus multilineatus</emphasis>
Bocage, 1866
</taxonomicName>
a; 
<taxonomicName id="83CC4D620816FFE598A6D83B9BD7FEEB" authority="Boettger, 1887" authorityName="Boettger" authorityYear="1887" box="[1009,1431,290,315]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF30816FFE598A6D83B9AA5FEEB" box="[1009,1253,290,315]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus auritus</emphasis>
Boettger, 1887
</taxonomicName>
; 
<taxonomicName id="83CC4D620816FFE59BC0D85F9CC7FE8F" authority="Lonnberg, 1907" authorityName="Lonnberg" authorityYear="1907" box="[151,647,326,351]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="intermedius" status="comb. nov.">
<emphasis id="76B8EAF30816FFE59BC0D85F9F83FE8F" box="[151,451,326,351]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus intermedius</emphasis>
Lönnberg, 1907
</taxonomicName>
<emphasis id="76B8EAF30816FFE599C4D85F9D58FE8F" bold="true" box="[659,792,326,351]" pageId="12" pageNumber="477">
<taxonomicNameLabel id="6D8B57880816FFE599C4D85F9D58FE8F" box="[659,792,326,351]" pageId="12" pageNumber="477">comb. nov.</taxonomicNameLabel>
</emphasis>
; 
<taxonomicName id="83CC4D620816FFE5987BD85F9AB0FE8F" authority="Andersson, 1916" authorityName="Andersson" authorityYear="1916" box="[812,1264,326,351]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="skoogi">
<emphasis id="76B8EAF30816FFE5987BD85F9A5DFE8F" box="[812,1053,326,351]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus skoogi</emphasis>
Andersson, 1916
</taxonomicName>
; 
<taxonomicName id="83CC4D620816FFE59FA8D85F9F35FE53" authority="Laurent, 1954" authorityName="Laurent" authorityYear="1954" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF30816FFE59FA8D85F9E8DFE53" italics="true" pageId="12" pageNumber="477">Gerrhosaurus bulsi</emphasis>
Laurent, 1954
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="0CD6656A0816FFE59B90D8979A98FD73" pageId="12" pageNumber="477" type="diagnosis">
<paragraph id="447336E10816FFE59B90D8979A98FD73" blockId="12.[151,1436,218,675]" pageId="12" pageNumber="477">
<emphasis id="76B8EAF30816FFE59B90D8979F03FE77" bold="true" box="[199,323,398,423]" pageId="12" pageNumber="477">Diagnosis:</emphasis>
The monophyly of 
<taxonomicName id="83CC4D620816FFE59976D8979CFEFE77" box="[545,702,398,423]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30816FFE59976D8979CFEFE77" box="[545,702,398,423]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus</emphasis>
</taxonomicName>
is established on the basis of a suite of nuclear and mitochondrial genetic characters (see above). These moderate-sized lizards are fairly well armoured and the head and body may be cylindrical, cyclotetragonal or slightly depressed; differentiated from the genera 
<taxonomicName id="83CC4D620816FFE59FE6D8CF9B21FE3F" box="[1201,1377,470,495]" class="Reptilia" family="Gerrhosauridae" genus="Broadleysaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30816FFE59FE6D8CF9B21FE3F" box="[1201,1377,470,495]" italics="true" pageId="12" pageNumber="477">Broadleysaurus</emphasis>
</taxonomicName>
and 
<taxonomicName id="83CC4D620816FFE59BC0D8E39F79FDC3" box="[151,313,506,531]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30816FFE59BC0D8E39F79FDC3" box="[151,313,506,531]" italics="true" pageId="12" pageNumber="477">Matobosaurus</emphasis>
</taxonomicName>
by its smaller size (maximum SVL 
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compared to 
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and 
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respectively for the latter two genera) and less robust appearance; most species of 
<taxonomicName id="83CC4D620816FFE5983CDB079A48FDE7" box="[875,1032,542,567]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30816FFE5983CDB079A48FDE7" box="[875,1032,542,567]" italics="true" pageId="12" pageNumber="477">Gerrhosaurus</emphasis>
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have only eight ventral scale rows longitudinally (but 
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<taxonomicName id="83CC4D620816FFE59AFBDB5A9C59FD8B" box="[428,537,579,603]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="typicus">
<emphasis id="76B8EAF30816FFE59AFBDB5A9C59FD8B" box="[428,537,579,603]" italics="true" pageId="12" pageNumber="477">G. typicus</emphasis>
</taxonomicName>
), whereas 
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<emphasis id="76B8EAF30816FFE599C5DB5B9D02FD8B" box="[658,834,578,603]" italics="true" pageId="12" pageNumber="477">Broadleysaurus</emphasis>
</taxonomicName>
has 9–10 and 
<taxonomicName id="83CC4D620816FFE598B5DB5B9AC3FD8B" box="[994,1155,578,603]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30816FFE598B5DB5B9AC3FD8B" box="[994,1155,578,603]" italics="true" pageId="12" pageNumber="477">Matobosaurus</emphasis>
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has 12–20; it also differs from 
<taxonomicName id="83CC4D620816FFE59B80DB7F9FC7FDAF" box="[215,391,614,639]" class="Reptilia" family="Gerrhosauridae" genus="Broadleysaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF30816FFE59B80DB7F9FC7FDAF" box="[215,391,614,639]" italics="true" pageId="12" pageNumber="477">Broadleysaurus</emphasis>
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by having 49–67 versus 31–38 transverse dorsal scale rows (Loveridge 1942; FitzSimons 1943, 1953; 
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Witte 1953; Laurent 1954, 1964; Broadley 1966; 
<collectingCountry id="3CDB76710816FFE59821DB929DD7FD72" box="[886,919,651,674]" name="South Georgia and the South Sandwich Islands" pageId="12" pageNumber="477">De</collectingCountry>
Waal 1978; Jacobsen 1989).
</paragraph>
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<paragraph id="447336E10816FFE59BC0DF0D9DBEF9FB" blockId="12.[151,1022,1556,1579]" box="[151,1022,1556,1579]" pageId="12" pageNumber="477">
<emphasis id="76B8EAF30816FFE59BC0DF0D9F59F9FB" bold="true" box="[151,281,1556,1579]" pageId="12" pageNumber="477">FIGURE 4.</emphasis>
<taxonomicName id="83CC4D620816FFE59A77DF0D9C4EF9FB" box="[288,526,1556,1579]" class="Reptilia" family="Gerrhosauridae" genus="Broadleysaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="major">
<emphasis id="76B8EAF30816FFE59A77DF0D9C4EF9FB" box="[288,526,1556,1579]" italics="true" pageId="12" pageNumber="477">Broadleysaurus major</emphasis>
</taxonomicName>
from Arusha, Tanzania. (Photo: W.R. Branch)
</paragraph>
</caption>
<subSubSection id="0CD6656A0816FFE49B90DF4C9DD9FF04" lastPageId="13" lastPageNumber="478" pageId="12" pageNumber="477" type="description">
<paragraph id="447336E10816FFE49B90DF4C9DD9FF04" blockId="12.[151,1436,1621,2006]" lastBlockId="13.[151,1436,151,392]" lastPageId="13" lastPageNumber="478" pageId="12" pageNumber="477">
<emphasis id="76B8EAF30816FFE59B90DF4C9F1BF9BE" bold="true" box="[199,347,1621,1646]" pageId="12" pageNumber="477">Description:</emphasis>
Head large, moderate or small, its length included in SVL 3.3–4.8 times (young lizards) to 4.0– 8.4 times (adults); head shields smooth or weakly striated; rostral in contact with, or separated from, the frontonasal; prefrontals well separated, slightly separated, in narrow contact, or in broad contact; supraoculars 4; supraciliaries 4–5 (rarely 3 or 6); tympanic shield narrow and band-like to broad and crescentic; body cyclotetragonal, slightly depressed in some 
<emphasis id="76B8EAF30816FFE599DADFFF9CB9F92D" box="[653,761,1766,1789]" italics="true" pageId="12" pageNumber="477">G. t y pi cu s</emphasis>
, or almost cylindrical (
<taxonomicName id="83CC4D620816FFE59F5ADFFF9A37F92E" box="[1037,1143,1765,1790]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="skoogi">
<emphasis id="76B8EAF30816FFE59F5ADFFF9A37F92E" box="[1037,1143,1765,1790]" italics="true" pageId="12" pageNumber="477">G. skoogi</emphasis>
</taxonomicName>
); dorsal scales weakly to strongly keeled, smooth or striated, in 20–28 (
<quantity id="83349B040816FFE599CBDE109D41F8F2" box="[668,769,1801,1826]" metricMagnitude="-1" metricUnit="m" metricValue="8.509" metricValueMax="8.89" metricValueMin="8.128" pageId="12" pageNumber="477" unit="in" value="33.5" valueMax="35.0" valueMin="32.0">32–35 in</quantity>
<taxonomicName id="83CC4D620816FFE59850DE139D2FF8F2" box="[775,879,1801,1826]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="skoogi">
<emphasis id="76B8EAF30816FFE59850DE139D2FF8F2" box="[775,879,1801,1826]" italics="true" pageId="12" pageNumber="477">G. skoogi</emphasis>
</taxonomicName>
) longitudinal and 49–67 transverse rows (usually counted from row posterior to nuchals to row above vent); lateral scales keeled, striated or smooth; ventral plates in 8 or 10 (
<taxonomicName id="83CC4D620816FFE59BA2DE4B9F22F8B9" box="[245,354,1874,1897]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="477" phylum="Chordata" rank="species" species="typicus">
<emphasis id="76B8EAF30816FFE59BA2DE4B9F22F8B9" box="[245,354,1874,1897]" italics="true" pageId="12" pageNumber="477">G. typicus</emphasis>
</taxonomicName>
only) longitudinal and 30–42 transverse rows (counted “from pectoral to anal shields” according to Loveridge 1942; i.e. from axilla to row before enlarged ventral plate); femoral pores 9–27 per thigh; fourth toe with 14–22 subdigital lamellae; largest known specimens: unknown sex 
<quantity id="83349B040816FFE5988BDE809A7BF861" box="[988,1083,1945,1970]" metricMagnitude="-1" metricUnit="m" metricValue="6.13" pageId="12" pageNumber="477" unit="mm" value="613.0">613 mm</quantity>
(
<quantity id="83349B040816FFE59F1ADE809AEEF861" box="[1101,1198,1945,1969]" metricMagnitude="-1" metricUnit="m" metricValue="2.13" pageId="12" pageNumber="477" unit="mm" value="213.0">213 mm</quantity>
SVL + 
<quantity id="83349B040816FFE59E58DE809B2FF861" box="[1295,1391,1945,1969]" metricMagnitude="-1" metricUnit="m" metricValue="4.0" pageId="12" pageNumber="477" unit="mm" value="400.0">400 mm</quantity>
tail length), male 
<quantity id="83349B040816FFE59A64DEA49FD1F805" box="[307,401,1981,2006]" metricMagnitude="-1" metricUnit="m" metricValue="4.85" pageId="12" pageNumber="477" unit="mm" value="485.0">485 mm</quantity>
(163 + 322), but another male had a SVL of 
<quantity id="83349B040816FFE598C3DEA49DB2F805" box="[916,1010,1981,2006]" metricMagnitude="-1" metricUnit="m" metricValue="1.75" pageId="12" pageNumber="477" unit="mm" value="175.0">175 mm</quantity>
female: 475 (142 + 333), but another female had a SVL of 
<quantity id="83349B040817FFE49ADFD98E9FA9FF7F" box="[392,489,151,176]" metricMagnitude="-1" metricUnit="m" metricValue="1.5699999999999998" pageId="13" pageNumber="478" unit="mm" value="157.0">157 mm</quantity>
; tail 1.0 to 2.5 times SVL (Loveridge 1942; FitzSimons 1943, 1953; 
<collectingCountry id="3CDB76710817FFE49FA7D9819B51FF7F" box="[1264,1297,152,175]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Witte 1953; Laurent 1954, 1964; Broadley 1966; 
<collectingCountry id="3CDB76710817FFE49961D9A49C17FF04" box="[566,599,189,212]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Waal 1978; Jacobsen 1989).
</paragraph>
</subSubSection>
<subSubSection id="0CD6656A0817FFE49B90D9C69D88FE58" pageId="13" pageNumber="478" type="distribution">
<paragraph id="447336E10817FFE49B90D9C69A7FFEB4" blockId="13.[151,1436,151,392]" pageId="13" pageNumber="478">
<emphasis id="76B8EAF30817FFE49B90D9C69F22FF28" bold="true" box="[199,354,223,248]" pageId="13" pageNumber="478">Distribution:</emphasis>
Widespread in Africa south of the equator, extending northwestwards into 
<collectingCountry id="3CDB76710817FFE49FE2D9C69B40FF28" box="[1205,1280,223,248]" name="Gabon" pageId="13" pageNumber="478">Gabon</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE49E6FD9C69BD8FF28" box="[1336,1432,223,248]" name="Angola" pageId="13" pageNumber="478">Cabinda</collectingCountry>
, and north-eastwards through 
<collectingCountry id="3CDB76710817FFE49ABED81D9C00FECD" box="[489,576,260,285]" name="Uganda" pageId="13" pageNumber="478">Uganda</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE49929D81C9C87FECD" box="[638,711,261,285]" name="Kenya" pageId="13" pageNumber="478">Kenya</collectingCountry>
to southern 
<collectingCountry id="3CDB76710817FFE4980AD81D9DE3FECD" box="[861,931,260,285]" name="Sudan" pageId="13" pageNumber="478">Sudan</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE498B6D81D9A00FECD" box="[993,1088,260,285]" name="Ethiopia" pageId="13" pageNumber="478">Ethiopia</collectingCountry>
(Loveridge 1942; FitzSimons 1943; 
<collectingCountry id="3CDB76710817FFE49B88D8319F40FEEF" box="[223,256,296,319]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Witte 1953; Mertens 1955; Broadley 1966, 1971; 
<collectingCountry id="3CDB76710817FFE49868D8319D20FEEF" box="[831,864,296,319]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Waal 1978; Auerbach 1987; Jacobsen 1989; Lang 1991; Branch 1998; Spawls 
<emphasis id="76B8EAF30817FFE49A83D8549C4CFEB5" box="[468,524,332,357]" italics="true" pageId="13" pageNumber="478">et al.</emphasis>
2002; Adolphs 2006, 2013; Bates 
<emphasis id="76B8EAF30817FFE498C5D8549A6FFEB4" box="[914,1071,332,357]" italics="true" pageId="13" pageNumber="478">et al. in press.</emphasis>
).
</paragraph>
<paragraph id="447336E10817FFE49B90D8769D88FE58" blockId="13.[151,1436,151,392]" box="[199,968,367,392]" pageId="13" pageNumber="478">
<emphasis id="76B8EAF30817FFE49B90D8769F48FE58" bold="true" box="[199,264,367,392]" pageId="13" pageNumber="478">Note:</emphasis>
Lizards in this genus are commonly known as ‘plated lizards’.
</paragraph>
</subSubSection>
<subSubSection id="0CD6656A0817FFF19BC0D8AE9F5DFE90" lastPageId="24" lastPageNumber="489" pageId="13" pageNumber="478" type="description">
<paragraph id="447336E10817FFE49BC0D8AE9C45FE00" blockId="13.[151,1436,439,861]" box="[151,517,439,464]" pageId="13" pageNumber="478">
<emphasis id="76B8EAF30817FFE49BC0D8AE9C45FE00" bold="true" box="[151,517,439,464]" pageId="13" pageNumber="478">
Status of ‘ 
<taxonomicName id="83CC4D620817FFE49A58D8AE9FBCFE00" box="[271,508,439,464]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="major">
<emphasis id="76B8EAF30817FFE49A58D8AE9FBCFE00" bold="true" box="[271,508,439,464]" italics="true" pageId="13" pageNumber="478">Gerrhosaurus major</emphasis>
</taxonomicName>
’
</emphasis>
</paragraph>
<paragraph id="447336E10817FFE49B90D8C59D2FFC8D" blockId="13.[151,1436,439,861]" pageId="13" pageNumber="478">
The 
<typeStatus id="9B7788430817FFE49BACD8C49F6BFE25" box="[251,299,477,501]" pageId="13" pageNumber="478">type</typeStatus>
locality of 
<taxonomicName id="83CC4D620817FFE49AF8D8C49C52FE24" box="[431,530,477,500]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="major">
<emphasis id="76B8EAF30817FFE49AF8D8C49C52FE24" box="[431,530,477,500]" italics="true" pageId="13" pageNumber="478">G. major</emphasis>
</taxonomicName>
(
<figureCitation id="DCF72A640817FFE49975D8C59C27FE24" box="[546,615,476,501]" captionStart="FIGURE 4" captionStartId="12.[151,255,1556,1579]" captionTargetBox="[151,1436,730,1530]" captionTargetId="figure@12.[151,1436,730,1531]" captionTargetPageId="12" captionText="FIGURE 4. Broadleysaurus major from Arusha, Tanzania. (Photo: W. R. Branch)" httpUri="https://zenodo.org/record/247070/files/figure.png" pageId="13" pageNumber="478">Fig. 4</figureCitation>
) is 
<collectingCountry id="3CDB76710817FFE499C5D8C59CB7FE25" box="[658,759,476,501]" name="Tanzania" pageId="13" pageNumber="478">Zanzibar</collectingCountry>
, an island off the coast of 
<collectingCountry id="3CDB76710817FFE49F72D8C59ACDFE25" box="[1061,1165,476,501]" name="Tanzania" pageId="13" pageNumber="478">Tanzania</collectingCountry>
, but 
<taxonomicName id="83CC4D620817FFE49F92D8C49B0CFE24" box="[1221,1356,477,500]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="major" subSpecies="major">
<emphasis id="76B8EAF30817FFE49F92D8C49B0CFE24" box="[1221,1356,477,500]" italics="true" pageId="13" pageNumber="478">G. m. major</emphasis>
</taxonomicName>
has an extensive range in the eastern half of Africa, from northern KwaZulu-Natal in 
<collectingCountry id="3CDB76710817FFE49F78D8E69A83FDC8" box="[1071,1219,511,536]" name="South Africa" pageId="13" pageNumber="478">South Africa</collectingCountry>
to 
<collectingCountry id="3CDB76710817FFE49FBAD8E69B10FDC8" box="[1261,1360,511,536]" name="Ethiopia" pageId="13" pageNumber="478">Ethiopia</collectingCountry>
; 
<taxonomicName id="83CC4D620817FFE49E09DB199EB3FDED" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="major" subSpecies="bottegoi">
<emphasis id="76B8EAF30817FFE49E09DB199EB3FDED" italics="true" pageId="13" pageNumber="478">G. m. bottegoi</emphasis>
</taxonomicName>
was described from Valley of Ghinda in 
<collectingCountry id="3CDB76710817FFE49991DB3D9D52FDED" box="[710,786,548,573]" name="Eritrea" pageId="13" pageNumber="478">Eritrea</collectingCountry>
and has a fragmented distribution, extending from northeast Africa (where it occurs together with the nominate subspecies in 
<collectingCountry id="3CDB76710817FFE498E5DB519DBDFDB0" box="[946,1021,584,608]" name="Kenya" pageId="13" pageNumber="478">Kenya</collectingCountry>
) across the continent to West Africa (Duméril 1851; Del Prato 1895; Loveridge 1942; FitzSimons 1943; Broadley 1966; Jacobsen 1989; Branch 1998; Spawls 
<emphasis id="76B8EAF30817FFE49BA6DB889F6CFD78" box="[241,300,655,680]" italics="true" pageId="13" pageNumber="478">et al.</emphasis>
2002; Adolphs 2006, 2013; Bates 
<emphasis id="76B8EAF30817FFE499E8DB889D24FD78" box="[703,868,655,680]" italics="true" pageId="13" pageNumber="478">et al. in press.</emphasis>
). The two subspecies are distinguishable only by their colour patterns (Broadley 1987). Our analysis included samples from southern and eastern Africa identifiable as 
<taxonomicName id="83CC4D620817FFE49BE1DBC19F7CFD20" box="[182,316,728,752]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="major" subSpecies="major">
<emphasis id="76B8EAF30817FFE49BE1DBC19F7CFD20" box="[182,316,728,752]" italics="true" pageId="13" pageNumber="478">G. m. major</emphasis>
</taxonomicName>
and one sample from Atakpame in 
<collectingCountry id="3CDB76710817FFE49983DBC19D4DFD20" box="[724,781,728,752]" name="Togo" pageId="13" pageNumber="478">Togo</collectingCountry>
referable to 
<taxonomicName id="83CC4D620817FFE498F7DBC19A7FFD20" box="[928,1087,727,752]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="major" subSpecies="bottegoi">
<emphasis id="76B8EAF30817FFE498F7DBC19A7FFD20" box="[928,1087,727,752]" italics="true" pageId="13" pageNumber="478">G. m. bottegoi</emphasis>
</taxonomicName>
(
<tableCitation id="094E035A0817FFE49F19DBCE9AE3FD20" box="[1102,1187,727,752]" captionStart="TABLE 1" captionStartId="4.[151,239,152,175]" captionText="TABLE 1. Sample numbers, collecting localities, museum voucher accession numbers (CAS = California Academy of" pageId="13" pageNumber="478">Table 1</tableCitation>
). The 
<collectingCountry id="3CDB76710817FFE49FA7DBC19B69FD20" box="[1264,1321,728,752]" name="Togo" pageId="13" pageNumber="478">Togo</collectingCountry>
sample is embedded within samples of 
<taxonomicName id="83CC4D620817FFE49AB2DBE49C2AFCC4" box="[485,618,765,788]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="major" subSpecies="major">
<emphasis id="76B8EAF30817FFE49AB2DBE49C2AFCC4" box="[485,618,765,788]" italics="true" pageId="13" pageNumber="478">G. m. major</emphasis>
</taxonomicName>
. Based on our molecular data, plus the weak morphological differences (i.e. colour variation) used for recognition of the two subspecies, we relegate 
<taxonomicName id="83CC4D620817FFE49F41DA399B11FCE8" authority="Del Prato, 1895" authorityName="Del Prato" authorityYear="1895" box="[1046,1361,799,824]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="bottegoi">
<emphasis id="76B8EAF30817FFE49F41DA399AD3FCE8" box="[1046,1171,799,824]" italics="true" pageId="13" pageNumber="478">G. bottegoi</emphasis>
Del Prato, 1895
</taxonomicName>
to the synonomy of 
<taxonomicName id="83CC4D620817FFE49A66DA5D9CA8FC8D" authority="Dumeril, 1851" authorityName="Dumeril" authorityYear="1851" box="[305,744,836,861]" class="Reptilia" family="Gerrhosauridae" genus="Broadleysaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="major" status="comb. nov.">
<emphasis id="76B8EAF30817FFE49A66DA5D9C6BFC8C" box="[305,555,836,861]" italics="true" pageId="13" pageNumber="478">Broadleysaurus major</emphasis>
(Duméril, 1851)
</taxonomicName>
<emphasis id="76B8EAF30817FFE499B8DA5D9D2FFC8D" bold="true" box="[751,879,836,861]" pageId="13" pageNumber="478">
<taxonomicNameLabel id="6D8B57880817FFE499B8DA5D9D2FFC8D" box="[751,879,836,861]" pageId="13" pageNumber="478">comb. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
<paragraph id="447336E10817FFE49BC0DA959C53FC75" blockId="13.[151,1436,907,1583]" box="[151,531,907,933]" pageId="13" pageNumber="478">
<emphasis id="76B8EAF30817FFE49BC0DA959C53FC75" bold="true" box="[151,531,907,933]" pageId="13" pageNumber="478">
Status of ‘ 
<taxonomicName id="83CC4D620817FFE49A58DA929C4AFC74" box="[271,522,907,932]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="validus">
<emphasis id="76B8EAF30817FFE49A58DA929C4AFC74" bold="true" box="[271,522,907,932]" italics="true" pageId="13" pageNumber="478">Gerrhosaurus validus</emphasis>
</taxonomicName>
’
</emphasis>
</paragraph>
<paragraph id="447336E10817FFE49B90DAB69FE8F9FF" blockId="13.[151,1436,907,1583]" pageId="13" pageNumber="478">
The two currently recognized subspecies of 
<emphasis id="76B8EAF30817FFE49994DAA99D71FC18" box="[707,817,943,968]" italics="true" pageId="13" pageNumber="478">G. va l i d us</emphasis>
each form separate monophyletic clades. In addition, sequence divergences between these taxa are much larger than would be expected for subspecies and instead are at the level of species (i.e. 8.5% ND2, 4.1% 16S). The two taxa are morphologically well differentiated (e.g. subocular excluded from lip by a labial in 
<taxonomicName id="83CC4D620817FFE4993ADD059CFEFBE5" box="[621,702,1052,1077]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="validus">
<emphasis id="76B8EAF30817FFE4993ADD059CFEFBE5" box="[621,702,1052,1077]" italics="true" pageId="13" pageNumber="478">validus</emphasis>
</taxonomicName>
, in contact with lip in 
<taxonomicName id="83CC4D620817FFE498E0DD059A65FBE5" box="[951,1061,1052,1077]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="maltzahni">
<emphasis id="76B8EAF30817FFE498E0DD059A65FBE5" box="[951,1061,1052,1077]" italics="true" pageId="13" pageNumber="478">maltzahni</emphasis>
</taxonomicName>
; longitudinal rows of dorsals 
<quantity id="83349B040817FFE49E25DD059E91FB88" metricMagnitude="-1" metricUnit="m" metricValue="7.874" metricValueMax="8.636" metricValueMin="7.112" pageId="13" pageNumber="478" unit="in" value="31.0" valueMax="34.0" valueMin="28.0">28– 34 in</quantity>
<taxonomicName id="83CC4D620817FFE49B8EDD269F6AFB88" box="[217,298,1087,1112]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="validus">
<emphasis id="76B8EAF30817FFE49B8EDD269F6AFB88" box="[217,298,1087,1112]" italics="true" pageId="13" pageNumber="478">validus</emphasis>
</taxonomicName>
, 
<quantity id="83349B040817FFE49A6EDD269FDDFB88" box="[313,413,1087,1112]" metricMagnitude="-1" metricUnit="m" metricValue="6.984999999999999" metricValueMax="7.62" metricValueMin="6.35" pageId="13" pageNumber="478" unit="in" value="27.5" valueMax="30.0" valueMin="25.0">25–30 in</quantity>
<taxonomicName id="83CC4D620817FFE49AF2DD269C53FB88" box="[421,531,1087,1112]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="maltzahni">
<emphasis id="76B8EAF30817FFE49AF2DD269C53FB88" box="[421,531,1087,1112]" italics="true" pageId="13" pageNumber="478">maltzahni</emphasis>
</taxonomicName>
; longitudinal rows of ventrals 
<quantity id="83349B040817FFE49825DD599D96FB88" box="[882,982,1087,1112]" metricMagnitude="-1" metricUnit="m" metricValue="4.318" metricValueMax="5.08" metricValueMin="3.556" pageId="13" pageNumber="478" unit="in" value="17.0" valueMax="20.0" valueMin="14.0">14–20 in</quantity>
<taxonomicName id="83CC4D620817FFE49889DD269A6FFB88" box="[990,1071,1087,1112]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="validus">
<emphasis id="76B8EAF30817FFE49889DD269A6FFB88" box="[990,1071,1087,1112]" italics="true" pageId="13" pageNumber="478">validus</emphasis>
</taxonomicName>
, 
<quantity id="83349B040817FFE49F69DD599AE3FB88" box="[1086,1187,1087,1112]" metricMagnitude="-1" metricUnit="m" metricValue="3.302" metricValueMax="3.556" metricValueMin="3.048" pageId="13" pageNumber="478" unit="in" value="13.0" valueMax="14.0" valueMin="12.0">12–14 in</quantity>
<taxonomicName id="83CC4D620817FFE49FFDDD269B58FB88" box="[1194,1304,1087,1112]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="maltzahni">
<emphasis id="76B8EAF30817FFE49FFDDD269B58FB88" box="[1194,1304,1087,1112]" italics="true" pageId="13" pageNumber="478">maltzahni</emphasis>
</taxonomicName>
; Loveridge 1942, FitzSimons 1943) and occur allopatrically. 
<taxonomicName id="83CC4D620817FFE49995DD7D9D8FFBAD" box="[706,975,1124,1149]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="validus" subSpecies="validus">
<emphasis id="76B8EAF30817FFE49995DD7D9D8FFBAD" box="[706,975,1124,1149]" italics="true" pageId="13" pageNumber="478">Gerrhosaurus v. validus</emphasis>
</taxonomicName>
occurs from Limpopo Province in 
<collectingCountry id="3CDB76710817FFE49E0CDD7D9E9FFB70" name="South Africa" pageId="13" pageNumber="478">South Africa</collectingCountry>
northwards to 
<collectingCountry id="3CDB76710817FFE49AC6DD9E9C68FB70" box="[401,552,1159,1184]" name="Mozambique" pageId="13" pageNumber="478">Mozambique</collectingCountry>
, 
<collectingCountry id="3CDB76710817FFE49961DD9E9CF1FB70" box="[566,689,1159,1184]" name="Zimbabwe" pageId="13" pageNumber="478">Zimbabwe</collectingCountry>
, 
<collectingCountry id="3CDB76710817FFE499E8DD9E9D55FB70" box="[703,789,1159,1184]" name="Zambia" pageId="13" pageNumber="478">Zambia</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE49806DD9E9DE9FB70" box="[849,937,1159,1184]" name="Malawi" pageId="13" pageNumber="478">Malawi</collectingCountry>
, while 
<taxonomicName id="83CC4D620817FFE49F56DD919AEDFB70" box="[1025,1197,1159,1184]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="variety" variety="maltzhani">
<emphasis id="76B8EAF30817FFE49F56DD919AEDFB70" box="[1025,1197,1159,1184]" italics="true" pageId="13" pageNumber="478">G. v. maltzhani</emphasis>
</taxonomicName>
(
<typeStatus id="9B7788430817FFE49FE9DD919AB0FB70" box="[1214,1264,1160,1184]" pageId="13" pageNumber="478">type</typeStatus>
locality: Farm Roidina, north of Omaruru, 
<collectingCountry id="3CDB76710817FFE49AB0DDB59C0BFB15" box="[487,587,1196,1221]" name="Namibia" pageId="13" pageNumber="478">Namibia</collectingCountry>
; 
<collectingCountry id="3CDB76710817FFE4990CDDB49C3CFB14" box="[603,636,1197,1220]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Grys 1938) is restricted to northern 
<collectingCountry id="3CDB76710817FFE49F6CDDB59ADBFB15" box="[1083,1179,1196,1221]" name="Namibia" pageId="13" pageNumber="478">Namibia</collectingCountry>
and southern 
<collectingCountry id="3CDB76710817FFE49E1DDDB59BDCFB15" box="[1354,1436,1196,1221]" name="Angola" pageId="13" pageNumber="478">Angola</collectingCountry>
(Loveridge 1942; FitzSimons 1943; Broadley 1966; Visser 1984a; Jacobsen 1989; Branch 1998; Spawls 
<emphasis id="76B8EAF30817FFE49E37DDC89BDCFB38" box="[1376,1436,1231,1256]" italics="true" pageId="13" pageNumber="478">et al.</emphasis>
2002; Adolphs 2006, 2013; Bates 
<emphasis id="76B8EAF30817FFE49972DDEC9C8CFADC" box="[549,716,1268,1293]" italics="true" pageId="13" pageNumber="478">et al. in press.</emphasis>
). The two taxa appear to be separated by the Kalahari Desert (Visser 1984a). Our samples of 
<taxonomicName id="83CC4D620817FFE49958DC019CDFFAE0" box="[527,671,1303,1328]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="validus" subSpecies="validus">
<emphasis id="76B8EAF30817FFE49958DC019CDFFAE0" box="[527,671,1303,1328]" italics="true" pageId="13" pageNumber="478">G. v. validus</emphasis>
</taxonomicName>
were from Limpopo Province in 
<collectingCountry id="3CDB76710817FFE49F78DC0E9A86FAE0" box="[1071,1222,1303,1328]" name="South Africa" pageId="13" pageNumber="478">South Africa</collectingCountry>
, 
<collectingCountry id="3CDB76710817FFE49F83DC0E9B28FAE0" box="[1236,1384,1303,1328]" name="Mozambique" pageId="13" pageNumber="478">Mozambique</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE49BC0DC259F53FA85" box="[151,275,1340,1365]" name="Zimbabwe" pageId="13" pageNumber="478">Zimbabwe</collectingCountry>
; 
<taxonomicName id="83CC4D620817FFE49A49DC249F83FA85" box="[286,451,1340,1365]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="subSpecies" species="validus" subSpecies="maltzahni">
<emphasis id="76B8EAF30817FFE49A49DC249F83FA85" box="[286,451,1340,1365]" italics="true" pageId="13" pageNumber="478">G. v. maltzahni</emphasis>
</taxonomicName>
was sampled in both 
<collectingCountry id="3CDB76710817FFE499EDDC259D5AFA85" box="[698,794,1340,1365]" name="Namibia" pageId="13" pageNumber="478">Namibia</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE49806DC259DE4FA85" box="[849,932,1340,1365]" name="Angola" pageId="13" pageNumber="478">Angola</collectingCountry>
(
<tableCitation id="094E035A0817FFE498E5DC259A47FA84" box="[946,1031,1340,1365]" captionStart="TABLE 1" captionStartId="4.[151,239,152,175]" captionText="TABLE 1. Sample numbers, collecting localities, museum voucher accession numbers (CAS = California Academy of" pageId="13" pageNumber="478">Table 1</tableCitation>
). The 
<typeStatus id="9B7788430817FFE49F05DC249AC2FA85" box="[1106,1154,1341,1365]" pageId="13" pageNumber="478">type</typeStatus>
locality for 
<taxonomicName id="83CC4D620817FFE49E5ADC249B3DFA85" box="[1293,1405,1340,1365]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="validus">
<emphasis id="76B8EAF30817FFE49E5ADC249B3DFA85" box="[1293,1405,1340,1365]" italics="true" pageId="13" pageNumber="478">G. validus</emphasis>
</taxonomicName>
of “towards the sources of the Garrep [Gariep], or Orange River” (Smith 1849, Appendix, p. 9), i.e. in 
<collectingCountry id="3CDB76710817FFE49FAADC469B1AFAA8" box="[1277,1370,1375,1400]" name="Lesotho" pageId="13" pageNumber="478">Lesotho</collectingCountry>
, must be in error―as noted by FitzSimons (1943)―as the species is not known to occur anywhere south of 28o latitude (Branch 1998; Bates 
<emphasis id="76B8EAF30817FFE49AD9DCB59C75FA13" box="[398,565,1450,1475]" italics="true" pageId="13" pageNumber="478">et al. in press.</emphasis>
). The combination of molecular, morphological and geographical evidence suggests that the two taxa represent separate evolutionary lineages, and we therefore revive 
<taxonomicName id="83CC4D620817FFE49FF5DCD69E8FF9DB" authority="De Grys, 1938" authorityName="De Grys" authorityYear="1938" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="maltzahni">
<emphasis id="76B8EAF30817FFE49FF5DCD69B6FFA37" box="[1186,1327,1486,1511]" italics="true" pageId="13" pageNumber="478">G. maltzahni</emphasis>
<collectingCountry id="3CDB76710817FFE49E61DCC99B17FA37" box="[1334,1367,1488,1511]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Grys, 1938
</taxonomicName>
as a full species, as 
<taxonomicName id="83CC4D620817FFE49AECDCEB9DD5F9DB" authority="De Grys, 1938" authorityName="De Grys" authorityYear="1938" box="[443,917,1522,1547]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="maltzahni" status="comb. nov.">
<emphasis id="76B8EAF30817FFE49AECDCEB9C93F9DB" box="[443,723,1522,1547]" italics="true" pageId="13" pageNumber="478">Matobosaurus maltzahni</emphasis>
(
<collectingCountry id="3CDB76710817FFE499B3DCEA9D46F9DB" box="[740,774,1523,1547]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Grys, 1938)
</taxonomicName>
<emphasis id="76B8EAF30817FFE498CADCEB9A5FF9DB" bold="true" box="[925,1055,1522,1547]" pageId="13" pageNumber="478">
<taxonomicNameLabel id="6D8B57880817FFE498CADCEB9A5FF9DB" box="[925,1055,1522,1547]" pageId="13" pageNumber="478">comb. nov.</taxonomicNameLabel>
</emphasis>
The two species in the genus are illustrated in 
<figureCitation id="DCF72A640817FFE49A7CDF0F9F30F9FF" box="[299,368,1558,1583]" captionStart="FIGURE 5" captionStartId="14.[151,255,1545,1568]" captionTargetBox="[151,1436,663,1520]" captionTargetId="figure@14.[151,1436,663,1520]" captionTargetPageId="14" captionText="FIGURE 5. Matobosaurus validus (NMB R 10893) from Greater Kuduland Safaris, east of Tshipise, Limpopo Province, South Africa. (Photo: M. Burger)" httpUri="https://zenodo.org/record/247071/files/figure.png" pageId="13" pageNumber="478">Figs 5</figureCitation>
&amp; 
<figureCitation id="DCF72A640817FFE49AC3DF0F9FE5F9FF" box="[404,421,1558,1583]" captionStart="FIGURE 6" captionStartId="15.[151,255,842,865]" captionTargetBox="[151,1436,218,818]" captionTargetId="figure@15.[151,1436,195,818]" captionTargetPageId="15" captionText="FIGURE 6. Matobosaurus maltzahni (PEM R 17984) from 0.5 km south of Tambor, Namibé district, Angola. (Photo: W. R. Branch)" httpUri="https://zenodo.org/record/247072/files/figure.png" pageId="13" pageNumber="478">6</figureCitation>
.
</paragraph>
<paragraph id="447336E10817FFE49BC0DF479DD2F9A7" blockId="13.[151,1436,1630,2015]" box="[151,914,1630,1655]" pageId="13" pageNumber="478">
<emphasis id="76B8EAF30817FFE49BC0DF479DD2F9A7" bold="true" box="[151,914,1630,1655]" pageId="13" pageNumber="478">
Status of taxa in the 
<taxonomicName id="83CC4D620817FFE49ADDDF479C8DF9A7" box="[394,717,1630,1655]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30817FFE49ADDDF479C8DF9A7" bold="true" box="[394,717,1630,1655]" italics="true" pageId="13" pageNumber="478">Gerrhosaurus nigrolineatus</emphasis>
</taxonomicName>
species complex
</emphasis>
</paragraph>
<paragraph id="447336E10817FFE49B90DF9B9AEFF847" blockId="13.[151,1436,1630,2015]" pageId="13" pageNumber="478">
The 
<typeStatus id="9B7788430817FFE49BA9DF9A9F6EF94B" box="[254,302,1667,1691]" pageId="13" pageNumber="478">type</typeStatus>
locality of 
<taxonomicName id="83CC4D620817FFE49AEADF9A9C34F94B" box="[445,628,1666,1691]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30817FFE49AEADF9A9C34F94B" box="[445,628,1666,1691]" italics="true" pageId="13" pageNumber="478">G. nigrolineatus</emphasis>
</taxonomicName>
is “ 
<collectingCountry id="3CDB76710817FFE499FEDF9B9D44F94B" box="[681,772,1666,1691]" name="Gabon" pageId="13" pageNumber="478">Gaboon</collectingCountry>
country, West Africa” (= 
<collectingCountry id="3CDB76710817FFE49F69DF9B9ACCF94B" box="[1086,1164,1666,1691]" name="Gabon" pageId="13" pageNumber="478">Gabon</collectingCountry>
; Hallowell 1857). This species has now been collected at several localities in 
<collectingCountry id="3CDB76710817FFE499B9DFBF9D79F96F" box="[750,825,1702,1727]" name="Gabon" pageId="13" pageNumber="478">Gabon</collectingCountry>
(Pauwels 
<emphasis id="76B8EAF30817FFE498FBDFB19DA4F96F" box="[940,996,1702,1727]" italics="true" pageId="13" pageNumber="478">et al.</emphasis>
2006), confirming its occurrence there. 
<collectingCountry id="3CDB76710817FFE49BC0DFD29EF6F933" box="[151,182,1739,1763]" name="American Samoa" pageId="13" pageNumber="478">As</collectingCountry>
currently understood it has a large distribution range, from 
<collectingCountry id="3CDB76710817FFE49820DFD39D82F933" box="[887,962,1738,1763]" name="Gabon" pageId="13" pageNumber="478">Gabon</collectingCountry>
and the lower 
<collectingCountry id="3CDB76710817FFE49F21DFD29A81F933" box="[1142,1217,1739,1763]" name="Republic of the Congo" pageId="13" pageNumber="478">Congo</collectingCountry>
eastwards through southern 
<collectingCountry id="3CDB76710817FFE49BA8DFF79CCCF8D7" box="[255,652,1774,1799]" name="Democratic Republic of the Congo" pageId="13" pageNumber="478">Democratic Republic of the Congo</collectingCountry>
(D.R.C.) to 
<collectingCountry id="3CDB76710817FFE4984CDFF79D32F8D7" box="[795,882,1774,1799]" name="Uganda" pageId="13" pageNumber="478">Uganda</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE498FDDFE99DB3F8D8" box="[938,1011,1776,1800]" name="Kenya" pageId="13" pageNumber="478">Kenya</collectingCountry>
in the east, then southwards as far as northern 
<collectingCountry id="3CDB76710817FFE49BABDE0B9F1FF8FB" box="[252,351,1810,1835]" name="Namibia" pageId="13" pageNumber="478">Namibia</collectingCountry>
, northern 
<collectingCountry id="3CDB76710817FFE49A98DE0A9C7DF8FA" box="[463,573,1811,1834]" name="Botswana" pageId="13" pageNumber="478">Botswana</collectingCountry>
and north-eastern 
<collectingCountry id="3CDB76710817FFE4985ADE0B9DDCF8FB" box="[781,924,1810,1835]" name="South Africa" pageId="13" pageNumber="478">South Africa</collectingCountry>
(Loveridge 1942; FitzSimons 1943; 
<collectingCountry id="3CDB76710817FFE49E6FDE0A9B19F8FA" box="[1336,1369,1811,1834]" name="South Georgia and the South Sandwich Islands" pageId="13" pageNumber="478">De</collectingCountry>
Witte 1953; Broadley 1966, 1971; Auerbach 1987; Jacobsen 1989; Branch 1998; Spawls 
<emphasis id="76B8EAF30817FFE49F32DE219AE2F89F" box="[1125,1186,1846,1871]" italics="true" pageId="13" pageNumber="478">et al.</emphasis>
2002; Bates 
<emphasis id="76B8EAF30817FFE49E68DE219E98F8A3" italics="true" pageId="13" pageNumber="478">et al. in press.</emphasis>
; Uetz 2013). Our samples were from Kouilou region, 
<collectingCountry id="3CDB76710817FFE49869DE439A7FF8A3" box="[830,1087,1882,1907]" name="Republic of the Congo" pageId="13" pageNumber="478">Republic of the Congo</collectingCountry>
(west-Central Africa) adjacent to 
<collectingCountry id="3CDB76710817FFE49BE3DE679F41F847" box="[180,257,1918,1943]" name="Gabon" pageId="13" pageNumber="478">Gabon</collectingCountry>
, and 
<collectingCountry id="3CDB76710817FFE49A6BDE679FE4F847" box="[316,420,1918,1943]" name="Tanzania" pageId="13" pageNumber="478">Tanzania</collectingCountry>
, 
<collectingCountry id="3CDB76710817FFE49AF9DE679C02F847" box="[430,578,1918,1943]" name="Mozambique" pageId="13" pageNumber="478">Mozambique</collectingCountry>
and 
<collectingCountry id="3CDB76710817FFE4992FDE679D49F847" box="[632,777,1918,1943]" name="South Africa" pageId="13" pageNumber="478">South Africa</collectingCountry>
(East and Southern Africa) (
<tableCitation id="094E035A0817FFE49F1DDE679ADEF847" box="[1098,1182,1918,1943]" captionStart="TABLE 1" captionStartId="4.[151,239,152,175]" captionText="TABLE 1. Sample numbers, collecting localities, museum voucher accession numbers (CAS = California Academy of" pageId="13" pageNumber="478">Table 1</tableCitation>
).
</paragraph>
<paragraph id="447336E10817FFE79B90DEBA9F44FDB0" blockId="13.[151,1436,1630,2015]" lastBlockId="14.[151,1436,151,608]" lastPageId="14" lastPageNumber="479" pageId="13" pageNumber="478">
Our analysis showed that 
<taxonomicName id="83CC4D620817FFE49ABDDEBA9CDDF86B" box="[490,669,1954,1979]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30817FFE49ABDDEBA9CDDF86B" box="[490,669,1954,1979]" italics="true" pageId="13" pageNumber="478">G. nigrolineatus</emphasis>
</taxonomicName>
as currently conceived is not monophyletic, although topology tests could not reject a monophyletic 
<taxonomicName id="83CC4D620817FFE49950DEDE9CFBF80F" box="[519,699,1990,2015]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="478" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30817FFE49950DEDE9CFBF80F" box="[519,699,1990,2015]" italics="true" pageId="13" pageNumber="478">G. nigrolineatus</emphasis>
</taxonomicName>
as presently defined. However, given the observed topology, the well-supported west-Central African clade of 
<taxonomicName id="83CC4D620814FFE799C1D9819D08FF60" box="[662,840,151,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE799C1D9819D08FF60" box="[662,840,151,176]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
is more closely related to 
<taxonomicName id="83CC4D620814FFE79F39D9819A9BFF60" box="[1134,1243,152,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF30814FFE79F39D9819A9BFF60" box="[1134,1243,152,176]" italics="true" pageId="14" pageNumber="479">G. auritus</emphasis>
</taxonomicName>
, rather than to 
<taxonomicName id="83CC4D620814FFE79ED3D9819F6CFF05" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79ED3D9819F6CFF05" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
from East and Southern Africa, and the nodes defining these groups are well-supported. Given the node support, as well as other lines of evidence (see below), we suggest that there is reasonably strong support that 
<taxonomicName id="83CC4D620814FFE79BC0D81C9F0DFECD" box="[151,333,260,285]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79BC0D81C9F0DFECD" box="[151,333,260,285]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
as currently defined is not monophyletic. Although the phylogeny of Lamb 
<emphasis id="76B8EAF30814FFE79F84D81C9B4FFECD" box="[1235,1295,260,285]" italics="true" pageId="14" pageNumber="479">et al.</emphasis>
(2003) also recovered a sister relationship between 
<taxonomicName id="83CC4D620814FFE79903D8319D48FE90" box="[596,776,295,320]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79903D8319D48FE90" box="[596,776,295,320]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
and 
<taxonomicName id="83CC4D620814FFE79868D8319DEDFE90" box="[831,941,296,320]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF30814FFE79868D8319DEDFE90" box="[831,941,296,320]" italics="true" pageId="14" pageNumber="479">G. auritus</emphasis>
</taxonomicName>
, only a single 
<taxonomicName id="83CC4D620814FFE79F03D8319B47FE90" box="[1108,1287,295,320]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79F03D8319B47FE90" box="[1108,1287,295,320]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
sample from 
<collectingCountry id="3CDB76710814FFE79BC0D8559F6BFEB5" box="[151,299,332,357]" name="Mozambique" pageId="14" pageNumber="479">Mozambique</collectingCountry>
was included. Because our analysis includes greater geographic coverage than previous studies, we were able to evaluate the status of 
<taxonomicName id="83CC4D620814FFE79976D8699C95FE58" box="[545,725,367,392]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79976D8699C95FE58" box="[545,725,367,392]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
. In addition to the lack of monophyly for 
<taxonomicName id="83CC4D620814FFE79FE1D8699B2AFE58" box="[1206,1386,367,392]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79FE1D8699B2AFE58" box="[1206,1386,367,392]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
, the west-Central African clade differs from the East and Southern African clade by large 
<emphasis id="76B8EAF30814FFE79F2CD88C9AC9FE7C" box="[1147,1161,405,428]" italics="true" pageId="14" pageNumber="479">p</emphasis>
-distances (13.0% ND2, 6.9% 16S). One individual (HB057, Arusha, 
<collectingCountry id="3CDB76710814FFE799C4D8AE9CBBFE00" box="[659,763,439,464]" name="Tanzania" pageId="14" pageNumber="479">Tanzania</collectingCountry>
; 
<figureCitation id="DCF72A640814FFE79850D8AE9D07FE00" box="[775,839,439,464]" captionStart="FIGURE 1" captionStartId="7.[151,255,1859,1882]" captionText="FIGURE 1. Maximum likelihood topology for Gerrhosauridae. Supported nodes are indicated by circles (black: ≥ 70 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities; grey: ≥ 65 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities). Genera and major clades are indicated, and corresponding letters A-F match the text." pageId="14" pageNumber="479">Fig. 1</figureCitation>
) was found less than 
<quantity id="83349B040814FFE79F69D8A19AD4FE00" box="[1086,1172,439,464]" metricMagnitude="5" metricUnit="m" metricValue="1.4" pageId="14" pageNumber="479" unit="km" value="140.0">140 km</quantity>
to the south-east of the approximate 
<typeStatus id="9B7788430814FFE79A7BD8C49F1CFE25" box="[300,348,477,501]" pageId="14" pageNumber="479">type</typeStatus>
locality of 
<taxonomicName id="83CC4D620814FFE79AB3D8C59A0DFE24" authority="Lonnberg, 1907" authorityName="Lonnberg" authorityYear="1907" box="[484,1101,476,501]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="intermedia">
<emphasis id="76B8EAF30814FFE79AB3D8C59DCEFE25" box="[484,910,476,501]" italics="true" pageId="14" pageNumber="479">Gerrhosaurus flavigularis intermedia</emphasis>
Lönnberg, 1907
</taxonomicName>
(i.e. “steppe near the Natron lakes, Kibonoto”, northern 
<collectingCountry id="3CDB76710814FFE79A8ED8E69C01FDC8" box="[473,577,511,536]" name="Tanzania" pageId="14" pageNumber="479">Tanzania</collectingCountry>
; p. 7). Taxonomic implications are that the East/Southern African clade represents a separate species, for which the name 
<taxonomicName id="83CC4D620814FFE799A2DB3D9AA9FDEC" authority="Lonnberg, 1907" authorityName="Lonnberg" authorityYear="1907" box="[757,1257,548,573]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="intermedius" status="comb. nov.">
<emphasis id="76B8EAF30814FFE799A2DB3D9A63FDED" box="[757,1059,548,573]" italics="true" pageId="14" pageNumber="479">Gerrhosaurus intermedius</emphasis>
Lönnberg, 1907
</taxonomicName>
<emphasis id="76B8EAF30814FFE79FA2DB3D9B3CFDED" bold="true" box="[1269,1404,548,573]" pageId="14" pageNumber="479">
<taxonomicNameLabel id="6D8B57880814FFE79FA2DB3D9B3CFDED" box="[1269,1404,548,573]" pageId="14" pageNumber="479">comb. nov.</taxonomicNameLabel>
</emphasis>
is available.
</paragraph>
<caption id="10B366690814FFE79BC0DF109FB9F990" httpUri="https://zenodo.org/record/247071/files/figure.png" pageId="14" pageNumber="479" targetBox="[151,1436,663,1520]" targetPageId="14">
<paragraph id="447336E10814FFE79BC0DF109FB9F990" blockId="14.[151,1436,1545,1600]" pageId="14" pageNumber="479">
<emphasis id="76B8EAF30814FFE79BC0DF109F59F9F0" bold="true" box="[151,281,1545,1568]" pageId="14" pageNumber="479">FIGURE 5.</emphasis>
<taxonomicName id="83CC4D620814FFE79A77DF109C4DF9F0" box="[288,525,1545,1568]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="validus">
<emphasis id="76B8EAF30814FFE79A77DF109C4DF9F0" box="[288,525,1545,1568]" italics="true" pageId="14" pageNumber="479">Matobosaurus validus</emphasis>
</taxonomicName>
(NMB R10893) from Greater Kuduland Safaris, east of Tshipise, Limpopo Province, South Africa. (Photo: M. Burger)
</paragraph>
</caption>
<paragraph id="447336E10814FFE79B90DF719D4DF839" blockId="14.[151,1437,1640,2026]" pageId="14" pageNumber="479">
Loveridge (1942) relegated 
<taxonomicName id="83CC4D620814FFE79947DF709C88F951" box="[528,712,1640,1665]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="intermedia">
<emphasis id="76B8EAF30814FFE79947DF709C88F951" box="[528,712,1640,1665]" italics="true" pageId="14" pageNumber="479">G. f. intermedia</emphasis>
</taxonomicName>
to the synonymy of 
<taxonomicName id="83CC4D620814FFE7989DDF709AE4F951" box="[970,1188,1640,1665]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="subSpecies" species="nigrolineatus" subSpecies="nigrolineatus">
<emphasis id="76B8EAF30814FFE7989DDF709AE4F951" box="[970,1188,1640,1665]" italics="true" pageId="14" pageNumber="479">G. n. nigrolineatus</emphasis>
</taxonomicName>
without explanation. Because of the similarity of taxa associated with the names 
<taxonomicName id="83CC4D620814FFE79865DF979D92F975" box="[818,978,1676,1701]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30814FFE79865DF979D92F975" box="[818,978,1676,1701]" italics="true" pageId="14" pageNumber="479">G. flavigularis</emphasis>
</taxonomicName>
and 
<taxonomicName id="83CC4D620814FFE79F5FDF979AFAF975" box="[1032,1210,1676,1701]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79F5FDF979AFAF975" box="[1032,1210,1676,1701]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
, the applicability of the name 
<taxonomicName id="83CC4D620814FFE79A58DFA89FF8F919" box="[271,440,1712,1737]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF30814FFE79A58DFA89FF8F919" box="[271,440,1712,1737]" italics="true" pageId="14" pageNumber="479">G. intermedius</emphasis>
</taxonomicName>
for eastern populations previously referred to 
<taxonomicName id="83CC4D620814FFE798B1DFA89AE0F919" box="[998,1184,1712,1737]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE798B1DFA89AE0F919" box="[998,1184,1712,1737]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
requires explanation. Although not mentioned in the text of Lönnberg’s (1907) description of 
<taxonomicName id="83CC4D620814FFE79891DFCF9A35F93E" box="[966,1141,1749,1774]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="intermedia">
<emphasis id="76B8EAF30814FFE79891DFCF9A35F93E" box="[966,1141,1749,1774]" italics="true" pageId="14" pageNumber="479">G. f. intermedia</emphasis>
</taxonomicName>
, it is evident from his fig. 1b (left side of head) that there are four supraciliaries as in 
<taxonomicName id="83CC4D620814FFE7983CDFE09A64F8C1" box="[875,1060,1784,1809]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE7983CDFE09A64F8C1" box="[875,1060,1784,1809]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
(usually five in 
<taxonomicName id="83CC4D620814FFE79FB8DFE09BD4F8C1" box="[1263,1428,1784,1809]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30814FFE79FB8DFE09BD4F8C1" box="[1263,1428,1784,1809]" italics="true" pageId="14" pageNumber="479">G. flavigularis</emphasis>
</taxonomicName>
; Loveridge 1942, FitzSimons 1943). The proportions and scutellation of the head (fig. 1a) are also very similar to FitzSimons’ (1943) fig. 157 of 
<taxonomicName id="83CC4D620814FFE79AAFDE589CECF889" box="[504,684,1856,1881]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79AAFDE589CECF889" box="[504,684,1856,1881]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
. In addition, Lönnberg’s description mentions that the flank scales of 
<taxonomicName id="83CC4D620814FFE79BEFDE7F9F2DF8AE" box="[184,365,1893,1918]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="intermedia">
<emphasis id="76B8EAF30814FFE79BEFDE7F9F2DF8AE" box="[184,365,1893,1918]" italics="true" pageId="14" pageNumber="479">G. f. intermedia</emphasis>
</taxonomicName>
are strongly keeled, and minium red in colour with dark bars extending from the back. The prefrontals are shown to be in good contact, with a long median suture (indicated in Lönnberg’s fig. 1a). All of these features are rare or absent in 
<taxonomicName id="83CC4D620814FFE79963DEB79C96F816" box="[564,726,1965,1990]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30814FFE79963DEB79C96F816" box="[564,726,1965,1990]" italics="true" pageId="14" pageNumber="479">G. flavigularis</emphasis>
</taxonomicName>
and often associated with 
<taxonomicName id="83CC4D620814FFE79F46DEB79A87F816" box="[1041,1223,1965,1990]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30814FFE79F46DEB79A87F816" box="[1041,1223,1965,1990]" italics="true" pageId="14" pageNumber="479">G. nigrolineatus</emphasis>
</taxonomicName>
, including eastern populations that we now refer to 
<taxonomicName id="83CC4D620814FFE7995EDECB9CECF83A" box="[521,684,2001,2026]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="14" pageNumber="479" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF30814FFE7995EDECB9CECF83A" box="[521,684,2001,2026]" italics="true" pageId="14" pageNumber="479">G. intermedius</emphasis>
</taxonomicName>
(
<figureCitation id="DCF72A640814FFE799ECDEC89CBFF839" box="[699,767,2001,2026]" captionStart="FIGURE 7" captionStartId="15.[151,255,1831,1854]" captionTargetBox="[151,1436,950,1806]" captionTargetId="figure@15.[151,1436,950,1807]" captionTargetPageId="15" captionText="FIGURE 7. Gerrhosaurus intermedius from 9 km south of Skukuza, Kruger National Park, Mpumalanga Province, South Africa. (Photo: W. R. Schmidt)" httpUri="https://zenodo.org/record/247073/files/figure.png" pageId="14" pageNumber="479">Fig. 7</figureCitation>
).
</paragraph>
<caption id="10B366690815FFE69BC0DA539F69FC51" httpUri="https://zenodo.org/record/247072/files/figure.png" pageId="15" pageNumber="480" targetBox="[151,1436,218,818]" targetPageId="15">
<paragraph id="447336E10815FFE69BC0DA539F69FC51" blockId="15.[151,1436,842,897]" pageId="15" pageNumber="480">
<emphasis id="76B8EAF30815FFE69BC0DA539F5CFCB1" bold="true" box="[151,284,842,865]" pageId="15" pageNumber="480">FIGURE 6.</emphasis>
<taxonomicName id="83CC4D620815FFE69A73DA539C6FFCB1" box="[292,559,842,865]" class="Reptilia" family="Gerrhosauridae" genus="Matobosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="480" phylum="Chordata" rank="species" species="maltzahni">
<emphasis id="76B8EAF30815FFE69A73DA539C6FFCB1" box="[292,559,842,865]" italics="true" pageId="15" pageNumber="480">Matobosaurus maltzahni</emphasis>
</taxonomicName>
(PEM R17984) from 0.5 km south of Tambor, Namibé district, Angola. (Photo: W.R. Branch)
</paragraph>
</caption>
<caption id="10B366690815FFE69BC0DE3E9C60F88E" httpUri="https://zenodo.org/record/247073/files/figure.png" pageId="15" pageNumber="480" targetBox="[151,1436,950,1806]" targetPageId="15">
<paragraph id="447336E10815FFE69BC0DE3E9C60F88E" blockId="15.[151,1436,1831,1886]" pageId="15" pageNumber="480">
<emphasis id="76B8EAF30815FFE69BC0DE3E9F5CF8EE" bold="true" box="[151,284,1831,1854]" pageId="15" pageNumber="480">FIGURE 7.</emphasis>
<taxonomicName id="83CC4D620815FFE69A70DE3E9C04F8EE" box="[295,580,1831,1854]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="480" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF30815FFE69A70DE3E9C04F8EE" box="[295,580,1831,1854]" italics="true" pageId="15" pageNumber="480">Gerrhosaurus intermedius</emphasis>
</taxonomicName>
from 9 km south of Skukuza, Kruger National Park, Mpumalanga Province, South Africa. (Photo: W.R. Schmidt)
</paragraph>
</caption>
<paragraph id="447336E10815FFF99B90DE919F76FEEF" blockId="15.[151,1436,1927,2024]" lastBlockId="16.[151,1436,151,789]" lastPageId="16" lastPageNumber="481" pageId="15" pageNumber="480">
In the 
<collectingCountry id="3CDB76710815FFE69A4CDE919F26F870" box="[283,358,1928,1952]" name="Republic of the Congo" pageId="15" pageNumber="480">Congo</collectingCountry>
and 
<collectingCountry id="3CDB76710815FFE69AFEDE9E9FB4F870" box="[425,500,1927,1952]" name="Gabon" pageId="15" pageNumber="480">Gabon</collectingCountry>
voucher specimens (
<taxonomicName id="83CC4D620815FFE699A3DE919DEDF870" box="[756,941,1927,1952]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="480" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30815FFE699A3DE919DEDF870" box="[756,941,1927,1952]" italics="true" pageId="15" pageNumber="480">G. nigrolineatus</emphasis>
</taxonomicName>
) examined (Appendix I) there were four supraciliaries on either side of the head (e.g. PEM R20067, 
<figureCitation id="DCF72A640815FFE6986FDEB29D39F814" box="[824,889,1963,1988]" captionStart="FIGURE 8" captionStartId="16.[151,255,1691,1714]" captionTargetBox="[151,1436,844,1666]" captionTargetId="figure@16.[151,1436,844,1667]" captionTargetPageId="16" captionText="FIGURE 8. Lateral view of the head of a Gerrhosaurus nigrolineatus (PEM R 20067) from Koutou base camp for Sintoukola Potash Project, Kouilou Province, Republic of the Congo. (Photo: M. Burger)" httpUri="https://zenodo.org/record/247074/files/figure.png" pageId="15" pageNumber="480">Fig. 8</figureCitation>
) in all but one specimen (PEM R20066, 
<collectingCountry id="3CDB76710815FFE69E1FDEB59BD4F814" box="[1352,1428,1964,1988]" name="Republic of the Congo" pageId="15" pageNumber="480">Congo</collectingCountry>
) which had five; flanks had weakly or moderately keeled scales in the two 
<collectingCountry id="3CDB76710815FFE69F4CDEC99A26F838" box="[1051,1126,2000,2024]" name="Republic of the Congo" pageId="15" pageNumber="480">Congo</collectingCountry>
specimens, weakly (5) or moderately (4) keeled in 
<collectingCountry id="3CDB7671080AFFF99A93D98E9C4FFF60" box="[452,527,151,176]" name="Gabon" pageId="16" pageNumber="481">Gabon</collectingCountry>
specimens; prefrontals in broad (PEM R20067) or moderate (PEM R20066) contact in 
<collectingCountry id="3CDB7671080AFFF99A58D9A59F1AFF04" box="[271,346,188,212]" name="Republic of the Congo" pageId="16" pageNumber="481">Congo</collectingCountry>
specimens, in broad (5) to moderate (4) contact in 
<collectingCountry id="3CDB7671080AFFF998F0D9A59DB2FF05" box="[935,1010,188,213]" name="Gabon" pageId="16" pageNumber="481">Gabon</collectingCountry>
specimens. We refer all of the above specimens to 
<emphasis id="76B8EAF3080AFFF99A65D9F99FADFF28" box="[306,493,223,248]" italics="true" pageId="16" pageNumber="481">
<taxonomicName id="83CC4D62080AFFF99A65D9F99FAAFF28" box="[306,490,223,248]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">G. nigrolineatus</taxonomicName>
.
</emphasis>
The vouchered 
<collectingCountry id="3CDB7671080AFFF999F1D9C69D7AFF28" box="[678,826,223,248]" name="Mozambique" pageId="16" pageNumber="481">Mozambique</collectingCountry>
sample of 
<taxonomicName id="83CC4D62080AFFF998EED9F99A1DFF28" box="[953,1117,223,248]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080AFFF998EED9F99A1DFF28" box="[953,1117,223,248]" italics="true" pageId="16" pageNumber="481">G. intermedius</emphasis>
</taxonomicName>
(TM 80959) from Moebase Village had four supraciliaries on either side of the head; flanks with strongly keeled scales; and prefrontals in broad contact.
</paragraph>
<paragraph id="447336E1080AFFF99B90D8559D18FCC5" blockId="16.[151,1436,151,789]" pageId="16" pageNumber="481">
Although Loveridge (1942: 511) was tempted to “separate an eastern race” of 
<taxonomicName id="83CC4D62080AFFF99F27D8549B69FEB5" box="[1136,1321,332,357]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080AFFF99F27D8549B69FEB5" box="[1136,1321,332,357]" italics="true" pageId="16" pageNumber="481">G. nigrolineatus</emphasis>
</taxonomicName>
, the only character he found useful was the number of longitudinal rows of dorsal scales, which numbered 
<quantity id="83349B04080AFFF99FBBD8699B10FE58" box="[1260,1360,367,392]" metricMagnitude="-1" metricUnit="m" metricValue="6.604" metricValueMax="7.112" metricValueMin="6.096" pageId="16" pageNumber="481" unit="in" value="26.0" valueMax="28.0" valueMin="24.0">24–28 in</quantity>
“West Africa” and 20–26 (but usually 22–24) in “East Africa”. Laurent (1954) later gave a count of 26 for a specimen from Dundo in north-eastern 
<collectingCountry id="3CDB7671080AFFF99AB0D8AE9C7AFE00" box="[487,570,439,464]" name="Angola" pageId="16" pageNumber="481">Angola</collectingCountry>
that he assigned to 
<taxonomicName id="83CC4D62080AFFF99875D8A19D96FE00" box="[802,982,439,464]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080AFFF99875D8A19D96FE00" box="[802,982,439,464]" italics="true" pageId="16" pageNumber="481">G. nigrolineatus</emphasis>
</taxonomicName>
. For southern Africa these counts were given as 22–24 (usually 22) by FitzSimons (1943) and 20–24 (mostly 22–23) by Jacobsen (1989). The 
<typeStatus id="9B778843080AFFF99E3BD8C49BDCFE25" box="[1388,1436,477,501]" pageId="16" pageNumber="481">type</typeStatus>
description of 
<taxonomicName id="83CC4D62080AFFF99A6EDB199CF4FDC8" authority="Hallowell 1857" authorityName="Hallowell" authorityYear="1857" box="[313,692,511,536]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080AFFF99A6EDB199FADFDC8" box="[313,493,511,536]" italics="true" pageId="16" pageNumber="481">G. nigrolineatus</emphasis>
(Hallowell 1857)
</taxonomicName>
refers to 25 longitudinal rows of dorsals, while the 
<typeStatus id="9B778843080AFFF99FABD8E69B1EFDC8" box="[1276,1374,511,536]" pageId="16" pageNumber="481" type="holotype">holotype</typeStatus>
of 
<taxonomicName id="83CC4D62080AFFF99ED3DB199FDFFDED" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="intermedia">
<emphasis id="76B8EAF3080AFFF99ED3DB199FDFFDED" italics="true" pageId="16" pageNumber="481">G. flavigularis intermedia</emphasis>
</taxonomicName>
has 22 such rows (Lönnberg 1907). Laurent (1964) later referred a specimen from Mayombe (lower 
<collectingCountry id="3CDB7671080AFFF99A33DB519FF0FDB0" box="[356,432,584,608]" name="Republic of the Congo" pageId="16" pageNumber="481">Congo</collectingCountry>
) with 25 such rows to 
<taxonomicName id="83CC4D62080AFFF999EADB519DD1FDB0" box="[701,913,583,608]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="subSpecies" species="nigrolineatus" subSpecies="nigrolineatus">
<emphasis id="76B8EAF3080AFFF999EADB519DD1FDB0" box="[701,913,583,608]" italics="true" pageId="16" pageNumber="481">G. n. nigrolineatus</emphasis>
</taxonomicName>
, and four specimens from Pweto in Katanga, D.R.C., with 24–26 such rows to 
<taxonomicName id="83CC4D62080AFFF9994EDB749C9BFD55" box="[537,731,620,645]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="subSpecies" species="nigrolineatus" subSpecies="intermedius">
<emphasis id="76B8EAF3080AFFF9994EDB749C9BFD55" box="[537,731,620,645]" italics="true" pageId="16" pageNumber="481">G. n. intermedius</emphasis>
</taxonomicName>
. The number of dorsal rows varied from 
<quantity id="83349B04080AFFF99FEFDB759B76FD55" box="[1208,1334,620,645]" metricMagnitude="-1" metricUnit="m" metricValue="6.096" metricValueMax="6.35" metricValueMin="5.842" pageId="16" pageNumber="481" unit="in" value="24.0" valueMax="25.0" valueMin="23.0">23 to 25 in</quantity>
both the 
<collectingCountry id="3CDB7671080AFFF99BC0DB899EA2FD78" box="[151,226,656,680]" name="Republic of the Congo" pageId="16" pageNumber="481">Congo</collectingCountry>
(
<emphasis id="76B8EAF3080AFFF99BA5DB889F45FD78" box="[242,261,657,680]" italics="true" pageId="16" pageNumber="481">N</emphasis>
= 2) and 
<collectingCountry id="3CDB7671080AFFF99A26DB969FFCFD78" box="[369,444,655,680]" name="Gabon" pageId="16" pageNumber="481">Gabon</collectingCountry>
(
<emphasis id="76B8EAF3080AFFF99A9BDB889F9FFD78" box="[460,479,657,680]" italics="true" pageId="16" pageNumber="481">N</emphasis>
= 9) specimens examined. The vouchered southern African sample of 
<taxonomicName id="83CC4D62080AFFF99FAEDB899BDCFD78" box="[1273,1436,655,680]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080AFFF99FAEDB899BDCFD78" box="[1273,1436,655,680]" italics="true" pageId="16" pageNumber="481">G. intermedius</emphasis>
</taxonomicName>
(TM 80959) had 24 longitudinal rows of ventrals. While there may be average differences in these counts between western and eastern populations, there is also some overlap, and the usefulness of this feature for separating 
<taxonomicName id="83CC4D62080AFFF99ED3DBC19F6CFCC5" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080AFFF99ED3DBC19F6CFCC5" italics="true" pageId="16" pageNumber="481">G. nigrolineatus</emphasis>
</taxonomicName>
and 
<taxonomicName id="83CC4D62080AFFF99A35DBE49C45FCC5" box="[354,517,764,789]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080AFFF99A35DBE49C45FCC5" box="[354,517,764,789]" italics="true" pageId="16" pageNumber="481">G. intermedius</emphasis>
</taxonomicName>
requires further investigation.
</paragraph>
<caption id="10B36669080AFFF99BC0DF829A0EF902" httpUri="https://zenodo.org/record/247074/files/figure.png" pageId="16" pageNumber="481" targetBox="[151,1436,844,1666]" targetPageId="16">
<paragraph id="447336E1080AFFF99BC0DF829A0EF902" blockId="16.[151,1436,1691,1746]" pageId="16" pageNumber="481">
<emphasis id="76B8EAF3080AFFF99BC0DF829F5EF962" bold="true" box="[151,286,1691,1714]" pageId="16" pageNumber="481">FIGURE 8.</emphasis>
Lateral view of the head of a 
<taxonomicName id="83CC4D62080AFFF9992BDF829DE9F962" box="[636,937,1691,1714]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080AFFF9992BDF829DE9F962" box="[636,937,1691,1714]" italics="true" pageId="16" pageNumber="481">Gerrhosaurus nigrolineatus</emphasis>
</taxonomicName>
(PEM R20067) from Koutou base camp for Sintoukola Potash Project, Kouilou Province, Republic of the Congo. (Photo: M. Burger)
</paragraph>
</caption>
<paragraph id="447336E1080AFFF89B90DFE29CFEFE24" blockId="16.[151,1436,1787,2028]" lastBlockId="17.[151,1436,151,501]" lastPageId="17" lastPageNumber="482" pageId="16" pageNumber="481">
According to Broadley (2007), 
<taxonomicName id="83CC4D62080AFFF9991FDFE59D43F8C4" box="[584,771,1787,1812]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080AFFF9991FDFE59D43F8C4" box="[584,771,1787,1812]" italics="true" pageId="16" pageNumber="481">G. nigrolineatus</emphasis>
</taxonomicName>
from 
<collectingCountry id="3CDB7671080AFFF9980FDFE29DE3F8C4" box="[856,931,1787,1812]" name="Gabon" pageId="16" pageNumber="481">Gabon</collectingCountry>
and the lower 
<collectingCountry id="3CDB7671080AFFF99F3DDFE29AF5F8C3" box="[1130,1205,1787,1811]" name="Republic of the Congo" pageId="16" pageNumber="481">Congo</collectingCountry>
region has ragged dorsolateral stripes and smooth plantar scales, features which he felt may distinguish it from populations of this species elsewhere in Africa. The plantar scales of eastern populations of 
<taxonomicName id="83CC4D62080AFFF998B7DE5D9AD6F88C" box="[992,1174,1859,1884]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080AFFF998B7DE5D9AD6F88C" box="[992,1174,1859,1884]" italics="true" pageId="16" pageNumber="481">G. nigrolineatus</emphasis>
</taxonomicName>
(= 
<emphasis id="76B8EAF3080AFFF99F95DE5D9B30F88C" box="[1218,1392,1859,1884]" italics="true" pageId="16" pageNumber="481">
<taxonomicName id="83CC4D62080AFFF99F95DE5D9B28F88C" box="[1218,1384,1859,1884]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="intermedius">G. intermedius</taxonomicName>
)
</emphasis>
are reportedly keeled (smooth and tubercular in 
<taxonomicName id="83CC4D62080AFFF999F4DE7E9D07F8AF" box="[675,839,1894,1919]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="16" pageNumber="481" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080AFFF999F4DE7E9D07F8AF" box="[675,839,1894,1919]" italics="true" pageId="16" pageNumber="481">G. flavigularis</emphasis>
</taxonomicName>
) (FitzSimons 1943; Broadley 1966). In the 
<collectingCountry id="3CDB7671080AFFF99E06DE7E9BDCF8AF" box="[1361,1436,1895,1919]" name="Republic of the Congo" pageId="16" pageNumber="481">Congo</collectingCountry>
specimens examined, the back and flanks were olive to light brown with distinct cream, black-bordered, dorsolateral stripes, with a similarly coloured vertebral stripe that was continuous in one specimen (PEM R20066) and broken in the other (PEM R20067). 
<collectingCountry id="3CDB7671080AFFF99939DECA9CF9F83C" box="[622,697,2003,2028]" name="Gabon" pageId="16" pageNumber="481">Gabon</collectingCountry>
specimens examined were light brown with scattered black and white lateral scales, and similar stripes, but the vertebral stripe was continuous in one specimen, broken in three and absent in five. 
<collectingCountry id="3CDB7671080BFFF89A20D9A59FD6FF04" box="[375,406,188,212]" name="American Samoa" pageId="17" pageNumber="482">As</collectingCountry>
shown in 
<figureCitation id="DCF72A64080BFFF89943D9A59C1AFF04" box="[532,602,188,213]" captionStart="FIGURE 9" captionStartId="17.[151,255,1049,1072]" captionTargetBox="[151,1436,539,1037]" captionTargetId="figure@17.[151,1436,535,1037]" captionTargetPageId="17" captionText="FIGURE 9. Gerrhosaurus nigrolineatus (MBUR 02993) from Tchiboula region, Kouilou Depratment, Republic of the Congo. (Photo: M. Burger)" httpUri="https://zenodo.org/record/247075/files/figure.png" pageId="17" pageNumber="482">Fig. 9</figureCitation>
, MBUR 02993―a specimen sampled for the current analysis―also has typical dorsolateral stripes as described above, with a broken vertebral stripe. The original description of 
<taxonomicName id="83CC4D62080BFFF89ED3D9F99F6CFECD" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF89ED3D9F99F6CFECD" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
refers to a yellow stripe on either side of the back, bordered internally (towards the centre of the back) by a black band; and also mentions that the centre of the back contains black spots in the form of longitudinal lines (Hallowell 1857). Colour photographs of the two 
<typeStatus id="9B778843080BFFF8985FD8549D2AFEB5" box="[776,874,333,357]" pageId="17" pageNumber="482" type="syntype">syntypes</typeStatus>
of 
<taxonomicName id="83CC4D62080BFFF898C6D8549A05FEB5" box="[913,1093,332,357]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF898C6D8549A05FEB5" box="[913,1093,332,357]" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
indicated that both specimens have faded somewhat, but their colour patterns were not dissimilar to the 
<collectingCountry id="3CDB7671080BFFF898A7D8699A7BFE58" box="[1008,1083,368,392]" name="Republic of the Congo" pageId="17" pageNumber="482">Congo</collectingCountry>
and 
<collectingCountry id="3CDB7671080BFFF89F20D8769A82FE58" box="[1143,1218,367,392]" name="Gabon" pageId="17" pageNumber="482">Gabon</collectingCountry>
material described above. ANSP 3729 had a pair of pale (cream) dorsolateral stripes with poorly defined black borders as well as a similar vertebral stripe anteriorly (not visible beyond the nape; 
<figureCitation id="DCF72A64080BFFF89831D8AE9DF9FE00" box="[870,953,439,464]" captionStart="FIGURE 10" captionStartId="18.[151,255,1786,1809]" captionTargetBox="[243,1344,700,1759]" captionTargetId="figure@18.[243,1344,700,1760]" captionTargetPageId="18" captionText="FIGURE 10. ANSP 3729, adult syntype of Gerrhosaurus nigrolineatus from Gabon. (Photo: N. Gilmore)" httpUri="https://zenodo.org/record/247076/files/figure.png" pageId="17" pageNumber="482">Fig. 10</figureCitation>
), while ANSP 8825 (juvenile) was similar but lacked a discernible vertebral stripe (
<figureCitation id="DCF72A64080BFFF89908D8C59CF1FE24" box="[607,689,476,501]" captionStart="FIGURE 11" captionStartId="19.[151,255,1999,2022]" captionTargetBox="[224,1361,179,1973]" captionTargetId="figure@19.[224,1363,159,1973]" captionTargetPageId="19" captionText="FIGURE 11. ANSP 8825, juvenile syntype of Gerrhosaurus nigrolineatus from Gabon. (Photo: N. Gilmore)" httpUri="https://zenodo.org/record/247077/files/figure.png" pageId="17" pageNumber="482">Fig. 11</figureCitation>
).
</paragraph>
<caption id="10B36669080BFFF89BC0DD009FF8FB80" httpUri="https://zenodo.org/record/247075/files/figure.png" pageId="17" pageNumber="482" targetBox="[151,1436,539,1037]" targetPageId="17">
<paragraph id="447336E1080BFFF89BC0DD009FF8FB80" blockId="17.[151,1436,1049,1104]" pageId="17" pageNumber="482">
<emphasis id="76B8EAF3080BFFF89BC0DD009F5BFBE0" bold="true" box="[151,283,1049,1072]" pageId="17" pageNumber="482">FIGURE 9.</emphasis>
<taxonomicName id="83CC4D62080BFFF89A74DD009C0DFBE0" box="[291,589,1049,1072]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF89A74DD009C0DFBE0" box="[291,589,1049,1072]" italics="true" pageId="17" pageNumber="482">Gerrhosaurus nigrolineatus</emphasis>
</taxonomicName>
(MBUR 02993) from Tchiboula region, Kouilou Depratment, Republic of the Congo. (Photo: M. Burger)
</paragraph>
</caption>
<paragraph id="447336E1080BFFF89B90DD6E9F6FFA03" blockId="17.[151,1436,1143,2032]" pageId="17" pageNumber="482">
Donald G. Broadley (in litt. 
<date id="30721021080BFFF8995BDD6E9CFEFB5F" box="[524,702,1143,1168]" pageId="17" pageNumber="482" value="2013-03-21">21 March 2013</date>
) noted that a specimen of 
<taxonomicName id="83CC4D62080BFFF898A3DD619AE9FB40" box="[1012,1193,1143,1168]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF898A3DD619AE9FB40" box="[1012,1193,1143,1168]" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
from Ponte Denis in 
<collectingCountry id="3CDB7671080BFFF89BC0DD829EA2FB64" box="[151,226,1179,1204]" name="Gabon" pageId="17" pageNumber="482">Gabon</collectingCountry>
in the collection of the Natural History Museum, 
<collectingCountry id="3CDB7671080BFFF8981CDD829D83FB64" box="[843,963,1179,1204]" name="Zimbabwe" pageId="17" pageNumber="482">Zimbabwe</collectingCountry>
(Bulawayo) had smooth plantar scales, differing somewhat from the weakly keeled plantar scales of PEM R20067 (a detailed photographic image was used for comparison) from 
<collectingCountry id="3CDB7671080BFFF89A9CDDFA9C8BFB2B" box="[459,715,1251,1276]" name="Republic of the Congo" pageId="17" pageNumber="482">Republic of the Congo</collectingCountry>
(Appendix I). In the 
<collectingCountry id="3CDB7671080BFFF898ECDDFA9A46FB2B" box="[955,1030,1251,1275]" name="Republic of the Congo" pageId="17" pageNumber="482">Congo</collectingCountry>
specimens examined, plantar scales were almost smooth or weakly keeled, while in the 
<collectingCountry id="3CDB7671080BFFF8998FDC1E9D63FAF0" box="[728,803,1287,1312]" name="Gabon" pageId="17" pageNumber="482">Gabon</collectingCountry>
sample they were weakly (7) or very weakly (2) keeled. Based on photographs of one foot of each of the 
<typeStatus id="9B778843080BFFF89997DC359D62FA94" box="[704,802,1324,1348]" pageId="17" pageNumber="482" type="syntype">syntypes</typeStatus>
of 
<taxonomicName id="83CC4D62080BFFF8981EDC359DBCFA94" box="[841,1020,1323,1348]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF8981EDC359DBCFA94" box="[841,1020,1323,1348]" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
, the scales on the soles were weakly keeled. The plantar scales of the sampled specimen (TM 80959) of 
<taxonomicName id="83CC4D62080BFFF898FADC499A13FAB8" box="[941,1107,1359,1384]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080BFFF898FADC499A13FAB8" box="[941,1107,1359,1384]" italics="true" pageId="17" pageNumber="482">G. intermedius</emphasis>
</taxonomicName>
were strongly keeled, while those of 10 additional specimens from 
<collectingCountry id="3CDB7671080BFFF89909DC6A9CB2FA5C" box="[606,754,1395,1420]" name="Mozambique" pageId="17" pageNumber="482">Mozambique</collectingCountry>
were moderately keeled; two out of three specimens from Limpopo Province in 
<collectingCountry id="3CDB7671080BFFF89AC3DC8E9C66FA60" box="[404,550,1431,1456]" name="South Africa" pageId="17" pageNumber="482">South Africa</collectingCountry>
had moderately keeled palmar scales, while one had distinctly keeled scales (Appendix I).
</paragraph>
<paragraph id="447336E1080BFFF89B90DCC69C22F97C" blockId="17.[151,1436,1143,2032]" pageId="17" pageNumber="482">
Although there was some variation in the extent and appearance of dorsal stripes and the keeling of plantar scales, the 
<collectingCountry id="3CDB7671080BFFF89A43DF1A9F1FF9CB" box="[276,351,1539,1563]" name="Republic of the Congo" pageId="17" pageNumber="482">Congo</collectingCountry>
and 
<collectingCountry id="3CDB7671080BFFF89ACFDF1A9FA3F9CC" box="[408,483,1539,1564]" name="Gabon" pageId="17" pageNumber="482">Gabon</collectingCountry>
samples (including material referred to by Broadley) are all considered conspecific and referable to 
<taxonomicName id="83CC4D62080BFFF89A18DF319C42F990" box="[335,514,1575,1600]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF89A18DF319C42F990" box="[335,514,1575,1600]" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
. Nevertheless, the smooth to feebly keeled plantar scales in 
<taxonomicName id="83CC4D62080BFFF89FFCDF319B1FF990" box="[1195,1375,1575,1600]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF89FFCDF319B1FF990" box="[1195,1375,1575,1600]" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
from 
<collectingCountry id="3CDB7671080BFFF89BC0DF529EA2F9B4" box="[151,226,1611,1636]" name="Gabon" pageId="17" pageNumber="482">Gabon</collectingCountry>
and 
<collectingCountry id="3CDB7671080BFFF89A72DF529F30F9B3" box="[293,368,1611,1635]" name="Republic of the Congo" pageId="17" pageNumber="482">Congo</collectingCountry>
is in contrast to the moderately to strongly keeled scales in populations referable to 
<taxonomicName id="83CC4D62080BFFF89ED3DF559F5CF958" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080BFFF89ED3DF559F5CF958" italics="true" pageId="17" pageNumber="482">G. intermedius</emphasis>
</taxonomicName>
(e.g. FitzSimons 1943), including those from 
<collectingCountry id="3CDB7671080BFFF89878DF769D83F958" box="[815,963,1647,1672]" name="Mozambique" pageId="17" pageNumber="482">Mozambique</collectingCountry>
(e.g. TM 80959 and the other specimens listed in Appendix I) as discussed above.
</paragraph>
<paragraph id="447336E1080BFFF89B90DFAE9D72F83F" blockId="17.[151,1436,1143,2032]" pageId="17" pageNumber="482">
The minium red to vermillion flanks (with pale spots or bars) of adult eastern 
<taxonomicName id="83CC4D62080BFFF89FC3DFA19B0FF900" box="[1172,1359,1719,1744]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF89FC3DFA19B0FF900" box="[1172,1359,1719,1744]" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
(= 
<taxonomicName id="83CC4D62080BFFF89ED3DFA19F5BF924" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080BFFF89ED3DFA19F5BF924" italics="true" pageId="17" pageNumber="482">G. intermedius</emphasis>
</taxonomicName>
) differ from the light and dark barred or mostly brown flanks of 
<taxonomicName id="83CC4D62080BFFF89F53DFC59AE5F924" box="[1028,1189,1755,1780]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080BFFF89F53DFC59AE5F924" box="[1028,1189,1755,1780]" italics="true" pageId="17" pageNumber="482">G. flavigularis</emphasis>
</taxonomicName>
(see descriptions and images in Jacobsen 1989; Branch 1998; Spawls 
<emphasis id="76B8EAF3080BFFF8998DDE199D57F8C8" box="[730,791,1791,1816]" italics="true" pageId="17" pageNumber="482">et al.</emphasis>
2002; Alexander &amp; Marais 2007). It should be noted however, that according to Broadley (1966), 
<taxonomicName id="83CC4D62080BFFF899C8DE3D9D01F8EC" box="[671,833,1827,1852]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080BFFF899C8DE3D9D01F8EC" box="[671,833,1827,1852]" italics="true" pageId="17" pageNumber="482">G. flavigularis</emphasis>
</taxonomicName>
from 
<collectingCountry id="3CDB7671080BFFF898DBDE3A9A60F8EC" box="[908,1056,1827,1852]" name="Mozambique" pageId="17" pageNumber="482">Mozambique</collectingCountry>
and adjacent parts of 
<collectingCountry id="3CDB7671080BFFF89E73DE3A9BDCF8EC" box="[1316,1436,1827,1852]" name="Zimbabwe" pageId="17" pageNumber="482">Zimbabwe</collectingCountry>
have vermillion flanks like 
<taxonomicName id="83CC4D62080BFFF89A9FDE519C3BF8B0" box="[456,635,1863,1888]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080BFFF89A9FDE519C3BF8B0" box="[456,635,1863,1888]" italics="true" pageId="17" pageNumber="482">G. nigrolineatus</emphasis>
</taxonomicName>
(= 
<taxonomicName id="83CC4D62080BFFF899F7DE519D02F8B0" box="[672,834,1863,1888]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080BFFF899F7DE519CF1F88F" box="[672,689,1864,1887]" italics="true" pageId="17" pageNumber="482">G</emphasis>
. 
<emphasis id="76B8EAF3080BFFF899E9DE5E9D02F8B0" box="[702,834,1863,1888]" italics="true" pageId="17" pageNumber="482">intermedius</emphasis>
</taxonomicName>
), although only in areas of allopatry. The same colour pattern has been recorded in 
<taxonomicName id="83CC4D62080BFFF89AB5DE759CC3F854" box="[482,643,1899,1924]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080BFFF89AB5DE759CC3F854" box="[482,643,1899,1924]" italics="true" pageId="17" pageNumber="482">G. flavigularis</emphasis>
</taxonomicName>
from eastern Limpopo Department and eastern North West Province, 
<collectingCountry id="3CDB7671080BFFF89BC0DE969F6CF878" box="[151,300,1935,1960]" name="South Africa" pageId="17" pageNumber="482">South Africa</collectingCountry>
, where the underside of the head is blue-grey in males (Jacobsen 1989). The possibility that such populations represent unique evolutionary lineages was not investigated in the present study, although some genetic structuring is evident within 
<taxonomicName id="83CC4D62080BFFF89966DEC19C90F820" box="[561,720,2007,2032]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="482" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080BFFF89966DEC19C90F820" box="[561,720,2007,2032]" italics="true" pageId="17" pageNumber="482">G. flavigularis</emphasis>
</taxonomicName>
(
<figureCitation id="DCF72A64080BFFF899B7DECE9D61F83F" box="[736,801,2007,2032]" captionStart="FIGURE 1" captionStartId="7.[151,255,1859,1882]" captionText="FIGURE 1. Maximum likelihood topology for Gerrhosauridae. Supported nodes are indicated by circles (black: ≥ 70 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities; grey: ≥ 65 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities). Genera and major clades are indicated, and corresponding letters A-F match the text." pageId="17" pageNumber="482">Fig. 1</figureCitation>
).
</paragraph>
<paragraph id="447336E10808FFFB9B90D98E9B1AFD54" blockId="18.[151,1436,151,645]" pageId="18" pageNumber="483">
According to Loveridge (1942), the scales on the flanks of 
<taxonomicName id="83CC4D620808FFFB982DD9819A70FF60" box="[890,1072,151,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30808FFFB982DD9819A70FF60" box="[890,1072,151,176]" italics="true" pageId="18" pageNumber="483">G. nigrolineatus</emphasis>
</taxonomicName>
(= G. 
<taxonomicName id="83CC4D620808FFFB9F28D98E9B43FF60" box="[1151,1283,151,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF30808FFFB9F28D98E9B43FF60" box="[1151,1283,151,176]" italics="true" pageId="18" pageNumber="483">intermedius</emphasis>
</taxonomicName>
) are striated, keeled, or more-or-less smooth, whereas those of adult 
<emphasis id="76B8EAF30808FFFB9855D9A49DF8FF05" box="[770,952,188,213]" italics="true" pageId="18" pageNumber="483">
G. f. 
<taxonomicName id="83CC4D620808FFFB9861D9A59DF8FF05" box="[822,952,188,213]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="flavigularis">flavigularis</taxonomicName>
</emphasis>
are smooth. For southern African material, FitzSimons (1943) noted that the laterals of 
<taxonomicName id="83CC4D620808FFFB99C6D9F99D05FF28" box="[657,837,223,248]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30808FFFB99C6D9F99D05FF28" box="[657,837,223,248]" italics="true" pageId="18" pageNumber="483">G. nigrolineatus</emphasis>
</taxonomicName>
(= 
<taxonomicName id="83CC4D620808FFFB9826D9F99A55FF28" box="[881,1045,223,248]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF30808FFFB9826D9F99A55FF28" box="[881,1045,223,248]" italics="true" pageId="18" pageNumber="483">G. intermedius</emphasis>
</taxonomicName>
) are keeled and sometimes feebly striated, while those of 
<taxonomicName id="83CC4D620808FFFB9AF4D81C9C05FECD" box="[419,581,260,285]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30808FFFB9AF4D81C9C05FECD" box="[419,581,260,285]" italics="true" pageId="18" pageNumber="483">G. flavigularis</emphasis>
</taxonomicName>
are smooth or feebly keeled and striated. However, Loveridge (1942: 515) also noted that in his “ill-defined race” 
<taxonomicName id="83CC4D620808FFFB990ED8319D3AFE90" box="[601,890,295,320]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="fitzsimonsi">
<emphasis id="76B8EAF30808FFFB990ED8319D3AFE90" box="[601,890,295,320]" italics="true" pageId="18" pageNumber="483">G. flavigularis fitzsimonsi</emphasis>
</taxonomicName>
(a synonym of 
<taxonomicName id="83CC4D620808FFFB9F78D8319A90FE90" box="[1071,1232,295,320]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30808FFFB9F78D8319A90FE90" box="[1071,1232,295,320]" italics="true" pageId="18" pageNumber="483">G. flavigularis</emphasis>
</taxonomicName>
) the laterals were striated and keeled, although occasionally almost smooth, whereas the prefrontals were in broad contact. The latter two features are consistent with 
<taxonomicName id="83CC4D620808FFFB995FD8699CFCFE58" box="[520,700,367,392]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30808FFFB995FD8699CFCFE58" box="[520,700,367,392]" italics="true" pageId="18" pageNumber="483">G. nigrolineatus</emphasis>
</taxonomicName>
. However, Loveridge (1942: 515) noted that his new subspecies had a short head (head length into SVL 4.75 times in young to 6 times in adults) as in 
<emphasis id="76B8EAF30808FFFB9F03D88C9B48FE7D" box="[1108,1288,404,429]" italics="true" pageId="18" pageNumber="483">
G. f. 
<taxonomicName id="83CC4D620808FFFB9FD0D88D9B48FE7D" box="[1159,1288,404,429]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="flavigularis">flavigularis</taxonomicName>
</emphasis>
, and “should not be confused with 
<taxonomicName id="83CC4D620808FFFB9ADBD8A19C7BFE00" box="[396,571,439,464]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="intermedia">
<emphasis id="76B8EAF30808FFFB9ADBD8A19C7BFE00" box="[396,571,439,464]" italics="true" pageId="18" pageNumber="483">G. f. intermedia</emphasis>
</taxonomicName>
…which, from his [Lönnberg 1907] figure, is a synonym of the long-headed 
<taxonomicName id="83CC4D620808FFFB9BC0D8C49F25FE25" box="[151,357,476,501]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="subSpecies" species="nigrolineatus" subSpecies="nigrolineatus">
<emphasis id="76B8EAF30808FFFB9BC0D8C49F25FE25" box="[151,357,476,501]" italics="true" pageId="18" pageNumber="483">G. n. nigrolineatus</emphasis>
</taxonomicName>
”. Head length into SVL was 4.7–5.0 times for the two 
<collectingCountry id="3CDB76710808FFFB9887D8C59A5BFE24" box="[976,1051,476,500]" name="Republic of the Congo" pageId="18" pageNumber="483">Congo</collectingCountry>
specimens examined, and 4.0–5.0 times (4.8–5.0 for three adults with SVL&gt; 
<quantity id="83349B040808FFFB99D6DB199CA2FDC7" box="[641,738,512,536]" metricMagnitude="-1" metricUnit="m" metricValue="1.0" pageId="18" pageNumber="483" unit="mm" value="100.0">100 mm</quantity>
, 4.0–4.6 for seven juveniles with SVL &lt;
<quantity id="83349B040808FFFB9FE3DB199B46FDC8" box="[1204,1286,512,536]" metricMagnitude="-2" metricUnit="m" metricValue="8.0" pageId="18" pageNumber="483" unit="mm" value="80.0">80 mm</quantity>
) for the nine 
<collectingCountry id="3CDB76710808FFFB9BC0DB3D9EA2FDED" box="[151,226,548,573]" name="Gabon" pageId="18" pageNumber="483">Gabon</collectingCountry>
samples. The vouchered 
<collectingCountry id="3CDB76710808FFFB9953DB3D9CD8FDED" box="[516,664,548,573]" name="Mozambique" pageId="18" pageNumber="483">Mozambique</collectingCountry>
sample of 
<taxonomicName id="83CC4D620808FFFB9841DB3C9DFAFDED" box="[790,954,548,573]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF30808FFFB9841DB3C9DFAFDED" box="[790,954,548,573]" italics="true" pageId="18" pageNumber="483">G. intermedius</emphasis>
</taxonomicName>
(TM 80959) was similar with head length into SVL 4.4 times. Therefore, we conclude that 
<taxonomicName id="83CC4D620808FFFB99B7DB519A1AFDB0" authority="Lonnberg, 1907" authorityName="Lonnberg" authorityYear="1907" box="[736,1114,583,608]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="subSpecies" species="flavigularis" subSpecies="intermedia">
<emphasis id="76B8EAF30808FFFB99B7DB519DD7FDB0" box="[736,919,583,608]" italics="true" pageId="18" pageNumber="483">G. f. intermedia</emphasis>
Lönnberg, 1907
</taxonomicName>
is conspecific with eastern populations currently referred to 
<taxonomicName id="83CC4D620808FFFB995EDB749D37FD55" authority="Hallowell, 1857" authorityName="Hallowell" authorityYear="1857" box="[521,887,620,645]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30808FFFB995EDB749CFCFD55" box="[521,700,620,645]" italics="true" pageId="18" pageNumber="483">G. nigrolineatus</emphasis>
Hallowell, 1857
</taxonomicName>
and which we now refer to 
<taxonomicName id="83CC4D620808FFFB9FE7DB749B13FD55" box="[1200,1363,620,645]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF30808FFFB9FE7DB749B13FD55" box="[1200,1363,620,645]" italics="true" pageId="18" pageNumber="483">G. intermedius</emphasis>
</taxonomicName>
.
</paragraph>
<caption id="10B366690808FFFB9BC0DFE39B43F8C1" box="[151,1283,1786,1809]" httpUri="https://zenodo.org/record/247076/files/figure.png" pageId="18" pageNumber="483" targetBox="[243,1344,700,1759]" targetPageId="18">
<paragraph id="447336E10808FFFB9BC0DFE39B43F8C1" blockId="18.[151,1283,1786,1809]" box="[151,1283,1786,1809]" pageId="18" pageNumber="483">
<emphasis id="76B8EAF30808FFFB9BC0DFE39F66F8C1" bold="true" box="[151,294,1786,1809]" pageId="18" pageNumber="483">FIGURE 10.</emphasis>
ANSP 3729, adult syntype of 
<taxonomicName id="83CC4D620808FFFB993ADFE39DD6F8C1" box="[621,918,1786,1809]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="18" pageNumber="483" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30808FFFB993ADFE39DD6F8C1" box="[621,918,1786,1809]" italics="true" pageId="18" pageNumber="483">Gerrhosaurus nigrolineatus</emphasis>
</taxonomicName>
from Gabon. (Photo: N. Gilmore)
</paragraph>
</caption>
<caption id="10B366690809FFFA9BC0DED69B62F836" box="[151,1314,1999,2022]" httpUri="https://zenodo.org/record/247077/files/figure.png" pageId="19" pageNumber="484" targetBox="[224,1361,179,1973]" targetPageId="19">
<paragraph id="447336E10809FFFA9BC0DED69B62F836" blockId="19.[151,1314,1999,2022]" box="[151,1314,1999,2022]" pageId="19" pageNumber="484">
<emphasis id="76B8EAF30809FFFA9BC0DED69F65F836" bold="true" box="[151,293,1999,2022]" pageId="19" pageNumber="484">FIGURE 11.</emphasis>
ANSP 8825, juvenile syntype of 
<taxonomicName id="83CC4D620809FFFA99DBDED69DF4F836" box="[652,948,1999,2022]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="19" pageNumber="484" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF30809FFFA99DBDED69DF4F836" box="[652,948,1999,2022]" italics="true" pageId="19" pageNumber="484">Gerrhosaurus nigrolineatus</emphasis>
</taxonomicName>
from Gabon. (Photo: N. Gilmore)
</paragraph>
</caption>
<paragraph id="447336E1080EFFFD9B90D9819A42FE00" blockId="20.[151,1436,151,464]" pageId="20" pageNumber="485">
In light of the phylogenetic and morphological differences mentioned above, we suggest that populations in 
<collectingCountry id="3CDB7671080EFFFD9BC0D9A59EA2FF05" box="[151,226,188,213]" name="Gabon" pageId="20" pageNumber="485">Gabon</collectingCountry>
and lower 
<collectingCountry id="3CDB7671080EFFFD9A3DD9A59FF5FF04" box="[362,437,188,212]" name="Republic of the Congo" pageId="20" pageNumber="485">Congo</collectingCountry>
(including Kouilou region) are all referable to 
<taxonomicName id="83CC4D62080EFFFD98B6D9A49AD8FF05" box="[993,1176,188,213]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="485" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080EFFFD98B6D9A49AD8FF05" box="[993,1176,188,213]" italics="true" pageId="20" pageNumber="485">G. nigrolineatus</emphasis>
</taxonomicName>
, and that all East and Southern African populations (
<collectingCountry id="3CDB7671080EFFFD9AAED9F99C06FF28" box="[505,582,224,248]" name="Kenya" pageId="20" pageNumber="485">Kenya</collectingCountry>
, 
<collectingCountry id="3CDB7671080EFFFD9903D9C69CEEFF28" box="[596,686,223,248]" name="Uganda" pageId="20" pageNumber="485">Uganda</collectingCountry>
, 
<collectingCountry id="3CDB7671080EFFFD99ECD9C69D5AFF28" box="[699,794,223,248]" name="Rwanda" pageId="20" pageNumber="485">Rwanda</collectingCountry>
, 
<collectingCountry id="3CDB7671080EFFFD987FD9C69DCFFF28" box="[808,911,223,248]" name="Tanzania" pageId="20" pageNumber="485">Tanzania</collectingCountry>
, 
<collectingCountry id="3CDB7671080EFFFD98CAD9C69DB5FF28" box="[925,1013,223,248]" name="Malawi" pageId="20" pageNumber="485">Malawi</collectingCountry>
, 
<collectingCountry id="3CDB7671080EFFFD9F54D9C69ADAFF28" box="[1027,1178,223,248]" name="Mozambique" pageId="20" pageNumber="485">Mozambique</collectingCountry>
, 
<collectingCountry id="3CDB7671080EFFFD9FF0D9C69B5FFF28" box="[1191,1311,223,248]" name="Zimbabwe" pageId="20" pageNumber="485">Zimbabwe</collectingCountry>
and 
<collectingCountry id="3CDB7671080EFFFD9E0DD9C69EA1FECD" name="South Africa" pageId="20" pageNumber="485">South Africa</collectingCountry>
) formerly identified as 
<taxonomicName id="83CC4D62080EFFFD9946D81C9C8EFECD" box="[529,718,260,285]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="485" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080EFFFD9946D81C9C8EFECD" box="[529,718,260,285]" italics="true" pageId="20" pageNumber="485">G. nigrolineatus</emphasis>
</taxonomicName>
be referred to 
<taxonomicName id="83CC4D62080EFFFD98CBD81C9A09FECD" box="[924,1097,260,285]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="485" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080EFFFD98CBD81C9A09FECD" box="[924,1097,260,285]" italics="true" pageId="20" pageNumber="485">G. intermedius</emphasis>
</taxonomicName>
. Accurate determination of geographical boundaries for these two species, especially in Central Africa (
<collectingCountry id="3CDB7671080EFFFD9F47D83E9A27FE90" box="[1040,1127,295,320]" name="Angola" pageId="20" pageNumber="485">Angola</collectingCountry>
, D.R.C., 
<collectingCountry id="3CDB7671080EFFFD9F81D83E9B6EFE90" box="[1238,1326,295,320]" name="Zambia" pageId="20" pageNumber="485">Zambia</collectingCountry>
, northern 
<collectingCountry id="3CDB7671080EFFFD9BC0D8549F48FEB4" box="[151,264,333,356]" name="Botswana" pageId="20" pageNumber="485">Botswana</collectingCountry>
, northern 
<collectingCountry id="3CDB7671080EFFFD9A2DD8559F9FFEB5" box="[378,479,332,357]" name="Namibia" pageId="20" pageNumber="485">Namibia</collectingCountry>
), will require additional sampling on a finer scale than presently available, as well as additional morphological examination of specimens from throughout their extensive ranges. The assignment of Angolan specimens referred to 
<taxonomicName id="83CC4D62080EFFFD9AACD88C9CEFFE7D" box="[507,687,404,429]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="485" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080EFFFD9AACD88C9CEFFE7D" box="[507,687,404,429]" italics="true" pageId="20" pageNumber="485">G. nigrolineatus</emphasis>
</taxonomicName>
(e.g. Hellmich 1957; Manaças 1963; Parker 1936; Schmidt 1933; Laurent 1964), and their relationship to 
<taxonomicName id="83CC4D62080EFFFD9901D8A19D46FE00" box="[598,774,439,464]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="485" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080EFFFD9901D8A19D46FE00" box="[598,774,439,464]" italics="true" pageId="20" pageNumber="485">G. multilineatus</emphasis>
</taxonomicName>
, remains problematic.
</paragraph>
<caption id="10B36669080EFFFD9BC0DD839EADFB01" httpUri="https://zenodo.org/record/247078/files/figure.png" pageId="20" pageNumber="485" targetBox="[151,1436,518,1156]" targetPageId="20">
<paragraph id="447336E1080EFFFD9BC0DD839EADFB01" blockId="20.[151,1436,1178,1233]" pageId="20" pageNumber="485">
<emphasis id="76B8EAF3080EFFFD9BC0DD839F68FB61" bold="true" box="[151,296,1178,1201]" pageId="20" pageNumber="485">FIGURE 12.</emphasis>
<taxonomicName id="83CC4D62080EFFFD9A66DD839C42FB61" box="[305,514,1178,1201]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="485" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080EFFFD9A66DD839C42FB61" box="[305,514,1178,1201]" italics="true" pageId="20" pageNumber="485">Gerrhosaurus bulsi</emphasis>
</taxonomicName>
(PEM R18941) from Kalumbila Village, Northwest Province, Zambia. (Photo: W.R. Branch)
</paragraph>
</caption>
<caption id="10B36669080EFFFD9BC0DE989A29F848" box="[151,1129,1921,1944]" httpUri="https://zenodo.org/record/247079/files/figure.png" pageId="20" pageNumber="485" targetBox="[230,1356,1285,1889]" targetPageId="20">
<paragraph id="447336E1080EFFFD9BC0DE989A29F848" blockId="20.[151,1129,1921,1944]" box="[151,1129,1921,1944]" pageId="20" pageNumber="485">
<emphasis id="76B8EAF3080EFFFD9BC0DE989F66F848" bold="true" box="[151,294,1921,1944]" pageId="20" pageNumber="485">FIGURE 13.</emphasis>
<taxonomicName id="83CC4D62080EFFFD9A7ADE989C54F848" box="[301,532,1921,1944]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="20" pageNumber="485" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF3080EFFFD9A7ADE989C54F848" box="[301,532,1921,1944]" italics="true" pageId="20" pageNumber="485">Gerrhosaurus auritus</emphasis>
</taxonomicName>
from near Kome Pan, Botswana. (Photo: W.D. Haacke)
</paragraph>
</caption>
<paragraph id="447336E1080FFFFC9B90D98E9A48FEB4" blockId="21.[151,1436,151,752]" pageId="21" pageNumber="486">
<taxonomicName id="83CC4D62080FFFFC9B90D98E9FE0FF60" box="[199,416,151,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080FFFFC9B90D98E9FE0FF60" box="[199,416,151,176]" italics="true" pageId="21" pageNumber="486">Gerrhosaurus bulsi</emphasis>
</taxonomicName>
, sister taxon to all other taxa in the 
<taxonomicName id="83CC4D62080FFFFC987AD9819D9FFF60" box="[813,991,151,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080FFFFC987AD9819D9FFF60" box="[813,991,151,176]" italics="true" pageId="21" pageNumber="486">G. nigrolineatus</emphasis>
</taxonomicName>
complex, is well supported as a distinct lineage (
<figureCitation id="DCF72A64080FFFFC9BA0D9A59F78FF04" box="[247,312,188,213]" captionStart="FIGURE 1" captionStartId="7.[151,255,1859,1882]" captionText="FIGURE 1. Maximum likelihood topology for Gerrhosauridae. Supported nodes are indicated by circles (black: ≥ 70 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities; grey: ≥ 65 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities). Genera and major clades are indicated, and corresponding letters A-F match the text." pageId="21" pageNumber="486">Fig. 1</figureCitation>
), and is easily identifiable from others in the complex by its distinct, largely uniform, brown or grey dorsal colour pattern in adults (
<figureCitation id="DCF72A64080FFFFC9AA1D9C69C0AFF28" box="[502,586,223,248]" captionStart="FIGURE 12" captionStartId="20.[151,255,1178,1201]" captionTargetBox="[151,1436,518,1156]" captionTargetId="figure@20.[151,1436,518,1156]" captionTargetPageId="20" captionText="FIGURE 12. Gerrhosaurus bulsi (PEM R 18941) from Kalumbila Village, Northwest Province, Zambia. (Photo: W. R. Branch)" httpUri="https://zenodo.org/record/247078/files/figure.png" pageId="21" pageNumber="486">Fig. 12</figureCitation>
). The 
<typeStatus id="9B778843080FFFFC99C6D9F99C81FF28" box="[657,705,224,248]" pageId="21" pageNumber="486">type</typeStatus>
locality of 
<emphasis id="76B8EAF3080FFFFC9814D9F99DD5FF28" box="[835,917,223,248]" italics="true" pageId="21" pageNumber="486">G. b ul s i</emphasis>
is Dundo, north-east 
<collectingCountry id="3CDB7671080FFFFC9FDBD9C69AA2FF28" box="[1164,1250,223,248]" name="Angola" pageId="21" pageNumber="486">Angola</collectingCountry>
; the species also occurs in 
<collectingCountry id="3CDB7671080FFFFC9A51D81D9F1BFECD" box="[262,347,260,285]" name="Zambia" pageId="21" pageNumber="486">Zambia</collectingCountry>
and the D.R.C. (Laurent 1954, 1964; Broadley 1966; Haagner 
<emphasis id="76B8EAF3080FFFFC9F7ED81C9A23FECD" box="[1065,1123,260,285]" italics="true" pageId="21" pageNumber="486">et al.</emphasis>
2000; Broadley &amp; Cotterill 2004; Adolphs 2006, 2013). Our samples were from Kalumbila Village in North West Province, 
<collectingCountry id="3CDB7671080FFFFC9F80D83E9B70FE90" box="[1239,1328,295,320]" name="Zambia" pageId="21" pageNumber="486">Zambia</collectingCountry>
; and near Lake Carumbo, 
<collectingCountry id="3CDB7671080FFFFC9A1BD8559FE1FEB5" box="[332,417,332,357]" name="Angola" pageId="21" pageNumber="486">Angola</collectingCountry>
, i.e. about 
<quantity id="83349B04080FFFFC994CD8559C30FEB5" box="[539,624,332,357]" metricMagnitude="5" metricUnit="m" metricValue="1.0" pageId="21" pageNumber="486" unit="km" value="100.0">100 km</quantity>
WSW of the 
<typeStatus id="9B778843080FFFFC985DD8549D7AFEB5" box="[778,826,333,357]" pageId="21" pageNumber="486">type</typeStatus>
locality (
<tableCitation id="094E035A080FFFFC98F4D8559DB7FEB4" box="[931,1015,332,357]" captionStart="TABLE 1" captionStartId="4.[151,239,152,175]" captionText="TABLE 1. Sample numbers, collecting localities, museum voucher accession numbers (CAS = California Academy of" pageId="21" pageNumber="486">Table 1</tableCitation>
).
</paragraph>
<paragraph id="447336E1080FFFFC9B90D8769DA9FD20" blockId="21.[151,1436,151,752]" pageId="21" pageNumber="486">
<taxonomicName id="83CC4D62080FFFFC9B90D8769F86FE58" box="[199,454,367,392]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF3080FFFFC9B90D8769F86FE58" box="[199,454,367,392]" italics="true" pageId="21" pageNumber="486">Gerrhosaurus auritus</emphasis>
</taxonomicName>
appears to be closely related to 
<taxonomicName id="83CC4D62080FFFFC98D0D8699A07FE58" box="[903,1095,367,392]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080FFFFC98D0D8699A07FE58" box="[903,1095,367,392]" italics="true" pageId="21" pageNumber="486">G. nigrolineatus</emphasis>
</taxonomicName>
, but morphologically it is distinguishable by its broad and crescentic (versus narrow) tympanic shield, smooth (versus keeled) lateral scales, and lack (versus presence) of distinct dorsolateral stripes in adults (Loveridge 1942; FitzSimons 1943; Broadley 2007). Its back is usually pale brown, often with 3–4 narrow, pale, black-bordered dorsolateral stripes (Broadley 1966; Branch 1998; 
<figureCitation id="DCF72A64080FFFFC9A28D8E69F93FDC8" box="[383,467,511,536]" captionStart="FIGURE 13" captionStartId="20.[151,255,1921,1944]" captionTargetBox="[230,1356,1285,1889]" captionTargetId="figure@20.[230,1356,1285,1889]" captionTargetPageId="20" captionText="FIGURE 13. Gerrhosaurus auritus from near Kome Pan, Botswana. (Photo: W. D. Haacke)" httpUri="https://zenodo.org/record/247079/files/figure.png" pageId="21" pageNumber="486">Fig. 13</figureCitation>
). The 
<typeStatus id="9B778843080FFFFC994BDB199C0CFDC8" box="[540,588,512,536]" pageId="21" pageNumber="486">type</typeStatus>
locality of 
<taxonomicName id="83CC4D62080FFFFC9987DB199D7EFDC8" box="[720,830,512,536]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF3080FFFFC9987DB199D7EFDC8" box="[720,830,512,536]" italics="true" pageId="21" pageNumber="486">G. auritus</emphasis>
</taxonomicName>
is Ondonga, Ovamboland, northern 
<collectingCountry id="3CDB7671080FFFFC9F89D8E69B01FDC8" box="[1246,1345,511,536]" name="Namibia" pageId="21" pageNumber="486">Namibia</collectingCountry>
, but the species also occurs in southern 
<collectingCountry id="3CDB7671080FFFFC9942DB3D9C2AFDED" box="[533,618,548,573]" name="Angola" pageId="21" pageNumber="486">Angola</collectingCountry>
, south-western 
<collectingCountry id="3CDB7671080FFFFC9872DB3D9D3BFDED" box="[805,891,548,573]" name="Zambia" pageId="21" pageNumber="486">Zambia</collectingCountry>
(where our single sample is from― 
<tableCitation id="094E035A080FFFFC9E64DB3D9BCAFDEC" box="[1331,1418,548,573]" captionStart="TABLE 1" captionStartId="4.[151,239,152,175]" captionText="TABLE 1. Sample numbers, collecting localities, museum voucher accession numbers (CAS = California Academy of" pageId="21" pageNumber="486">Table 1</tableCitation>
), 
<collectingCountry id="3CDB7671080FFFFC9BC0DB519F48FD8F" box="[151,264,584,607]" name="Botswana" pageId="21" pageNumber="486">Botswana</collectingCountry>
, western 
<collectingCountry id="3CDB7671080FFFFC9A27DB5E9FA8FDB0" box="[368,488,583,608]" name="Zimbabwe" pageId="21" pageNumber="486">Zimbabwe</collectingCountry>
and northern Limpopo Province in 
<collectingCountry id="3CDB7671080FFFFC9820DB5E9A46FDB0" box="[887,1030,583,608]" name="South Africa" pageId="21" pageNumber="486">South Africa</collectingCountry>
(Loveridge 1942; FitzSimons 1943; Broadley 1966; Visser 1984a; Broadley &amp; Rasmussen 1995; Branch 1998; Broadley &amp; Cotterill 2004; Adolphs 2006, 2013; Bates 
<emphasis id="76B8EAF3080FFFFC9A3ADB889C4FFD78" box="[365,527,655,680]" italics="true" pageId="21" pageNumber="486">et al. in press.</emphasis>
). The four species 
<taxonomicName id="83CC4D62080FFFFC99BBDB899DE0FD78" box="[748,928,655,680]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080FFFFC99BBDB899DE0FD78" box="[748,928,655,680]" italics="true" pageId="21" pageNumber="486">G. nigrolineatus</emphasis>
</taxonomicName>
, 
<taxonomicName id="83CC4D62080FFFFC98E7DB899A15FD78" box="[944,1109,655,680]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080FFFFC98E7DB899A15FD78" box="[944,1109,655,680]" italics="true" pageId="21" pageNumber="486">G. intermedius</emphasis>
</taxonomicName>
, 
<taxonomicName id="83CC4D62080FFFFC9F33DB899A93FD78" box="[1124,1235,656,680]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF3080FFFFC9F33DB899A93FD78" box="[1124,1235,656,680]" italics="true" pageId="21" pageNumber="486">G. auritus</emphasis>
</taxonomicName>
and 
<taxonomicName id="83CC4D62080FFFFC9E5ADB899B23FD78" box="[1293,1379,655,680]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080FFFFC9E5ADB899B23FD78" box="[1293,1379,655,680]" italics="true" pageId="21" pageNumber="486">G. bulsi</emphasis>
</taxonomicName>
, and possibly the morphologically and geographically allied form 
<taxonomicName id="83CC4D62080FFFFC9805DBAC9A45FD1D" box="[850,1029,692,717]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080FFFFC9805DBAC9A45FD1D" box="[850,1029,692,717]" italics="true" pageId="21" pageNumber="486">G. multilineatus</emphasis>
</taxonomicName>
(if valid, see below), constitute the ‘ 
<taxonomicName id="83CC4D62080FFFFC9BF7DBC19F13FD20" box="[160,339,727,752]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080FFFFC9BF7DBC19F13FD20" box="[160,339,727,752]" italics="true" pageId="21" pageNumber="486">G. nigrolineatus</emphasis>
</taxonomicName>
species complex’ with a widespread distribution in Africa.
</paragraph>
<caption id="10B36669080FFFFC9BC0DC349D39FAB4" httpUri="https://zenodo.org/record/247080/files/figure.png" pageId="21" pageNumber="486" targetBox="[230,1342,807,1300]" targetPageId="21">
<paragraph id="447336E1080FFFFC9BC0DC349D39FAB4" blockId="21.[151,1436,1325,1380]" pageId="21" pageNumber="486">
<emphasis id="76B8EAF3080FFFFC9BC0DC349F69FA94" bold="true" box="[151,297,1325,1348]" pageId="21" pageNumber="486">FIGURE 14.</emphasis>
Lateral view of the head and anterior part of the body of topotypical 
<taxonomicName id="83CC4D62080FFFFC9F61DC349B20FA94" box="[1078,1376,1325,1348]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080FFFFC9F61DC349B20FA94" box="[1078,1376,1325,1348]" italics="true" pageId="21" pageNumber="486">Gerrhosaurus multilineatus</emphasis>
</taxonomicName>
(BM 1904.5.2.32) from Duque de Bragança, Angola. (Photo: P. Campbell)
</paragraph>
</caption>
<caption id="10B36669080FFFFC9BC0DE7B9D39F849" httpUri="https://zenodo.org/record/247081/files/figure.png" pageId="21" pageNumber="486" targetBox="[243,1323,1431,1868]" targetPageId="21">
<paragraph id="447336E1080FFFFC9BC0DE7B9D39F849" blockId="21.[151,1436,1890,1945]" pageId="21" pageNumber="486">
<emphasis id="76B8EAF3080FFFFC9BC0DE7B9F69F8A9" bold="true" box="[151,297,1890,1913]" pageId="21" pageNumber="486">FIGURE 15.</emphasis>
Dorsal view of the head and anterior part of the body of topotypical 
<taxonomicName id="83CC4D62080FFFFC9F62DE7B9B20F8A9" box="[1077,1376,1890,1913]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="21" pageNumber="486" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080FFFFC9F62DE7B9B20F8A9" box="[1077,1376,1890,1913]" italics="true" pageId="21" pageNumber="486">Gerrhosaurus multilineatus</emphasis>
</taxonomicName>
(BM 1904.5.2.32) from Duque de Bragança, Angola. (Photo: P. Campbell)
</paragraph>
</caption>
<paragraph id="447336E1080CFFFF9BC0D98E9C07FF60" blockId="22.[151,1436,151,752]" box="[151,583,151,176]" pageId="22" pageNumber="487">
<emphasis id="76B8EAF3080CFFFF9BC0D98E9C07FF60" bold="true" box="[151,583,151,176]" pageId="22" pageNumber="487">
Status of 
<taxonomicName id="83CC4D62080CFFFF9A52D98E9C07FF60" box="[261,583,151,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080CFFFF9A52D98E9C07FF60" bold="true" box="[261,583,151,176]" italics="true" pageId="22" pageNumber="487">Gerrhosaurus multilineatus</emphasis>
</taxonomicName>
</emphasis>
</paragraph>
<paragraph id="447336E1080CFFFF9B90D9A59CBBFEB5" blockId="22.[151,1436,151,752]" pageId="22" pageNumber="487">
The taxonomic status of 
<taxonomicName id="83CC4D62080CFFFF9A8AD9A49CCEFF05" box="[477,654,188,213]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080CFFFF9A8AD9A49CCEFF05" box="[477,654,188,213]" italics="true" pageId="22" pageNumber="487">G. multilineatus</emphasis>
</taxonomicName>
has been confused in the literature and remains uncertain. According to Haagner 
<emphasis id="76B8EAF3080CFFFF9A75D9F89F1EFF28" box="[290,350,223,248]" italics="true" pageId="22" pageNumber="487">et al.</emphasis>
(2000), “Broadley (1999) notes that the taxon 
<taxonomicName id="83CC4D62080CFFFF98DBD9F99AE0FF28" authority="Bocage" authorityName="Bocage" box="[908,1184,223,248]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080CFFFF98DBD9F99A01FF28" box="[908,1089,223,248]" italics="true" pageId="22" pageNumber="487">G. multilineatus</emphasis>
Bocage
</taxonomicName>
is based on a hybrid specimen. The name is therefore unavailable.” However, this was in fact a reference to an unpublished manuscript (D.G. Broadley in litt. 
<date id="30721021080CFFFF9AC5D8319C10FE90" box="[402,592,295,320]" pageId="22" pageNumber="487" value="2012-02-08">8 February 2012</date>
). According to Article 17.2 of the Code (ICZN 1999), even if the specimen was a hybrid, the name would in fact still be available.
</paragraph>
<paragraph id="447336E1080CFFFF9B90D8699D4AFD20" blockId="22.[151,1436,151,752]" pageId="22" pageNumber="487">
In his description of 
<taxonomicName id="83CC4D62080CFFFF9AF9D8699C1EFE58" box="[430,606,367,392]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080CFFFF9AF9D8699C1EFE58" box="[430,606,367,392]" italics="true" pageId="22" pageNumber="487">G. multilineatus</emphasis>
</taxonomicName>
, based mainly on colour pattern, Bocage (1866a) noted that this form was similar to 
<taxonomicName id="83CC4D62080CFFFF9A41D88C9F90FE7D" box="[278,464,404,429]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080CFFFF9A41D88C9F90FE7D" box="[278,464,404,429]" italics="true" pageId="22" pageNumber="487">G. nigrolineatus</emphasis>
</taxonomicName>
, of which it may be merely a well characterised variety. Loveridge (1942) and FitzSimons (1943) subsequently relegated 
<taxonomicName id="83CC4D62080CFFFF9920D8A19D67FE00" box="[631,807,439,464]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080CFFFF9920D8A19D67FE00" box="[631,807,439,464]" italics="true" pageId="22" pageNumber="487">G. multilineatus</emphasis>
</taxonomicName>
to the synonymy of 
<taxonomicName id="83CC4D62080CFFFF9F47D8A19A82FE00" box="[1040,1218,439,464]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080CFFFF9F47D8A19A82FE00" box="[1040,1218,439,464]" italics="true" pageId="22" pageNumber="487">G. nigrolineatus</emphasis>
</taxonomicName>
. Although the 
<typeStatus id="9B778843080CFFFF9E3BD8A19BDCFE00" box="[1388,1436,440,464]" pageId="22" pageNumber="487">type</typeStatus>
series of 
<taxonomicName id="83CC4D62080CFFFF9BACD8C49FEBFE25" box="[251,427,476,501]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080CFFFF9BACD8C49FEBFE25" box="[251,427,476,501]" italics="true" pageId="22" pageNumber="487">G. multilineatus</emphasis>
</taxonomicName>
(Duque de Bragança district [region], interior of 
<collectingCountry id="3CDB7671080CFFFF9898D8C59A63FE25" box="[975,1059,476,501]" name="Angola" pageId="22" pageNumber="487">Angola</collectingCountry>
) was destroyed in the 1978 fire at Museu Bocage in Lisbon (Almaca &amp; Neves 1987; Madruga 2012), we examined colour photographs of two ‘virtual’ topotypes (‘Duque de Bragança’) in the collection of the Natural History Museum (London). In terms of morphology and colour pattern (e.g. 
<figureCitation id="DCF72A64080CFFFF9978DB5E9CC1FDB0" box="[559,641,583,608]" captionStart="FIGURE 14" captionStartId="21.[151,255,1325,1348]" captionTargetBox="[230,1342,807,1300]" captionTargetId="figure@21.[230,1356,807,1300]" captionTargetPageId="21" captionText="FIGURE 14. Lateral view of the head and anterior part of the body of topotypical Gerrhosaurus multilineatus (BM 1904.5.2.32) from Duque de Bragança, Angola. (Photo: P. Campbell)" httpUri="https://zenodo.org/record/247080/files/figure.png" pageId="22" pageNumber="487">Figs 14</figureCitation>
&amp; 
<figureCitation id="DCF72A64080CFFFF99F3DB5E9C81FDB0" box="[676,705,583,608]" captionStart="FIGURE 15" captionStartId="21.[151,255,1890,1913]" captionTargetBox="[243,1323,1431,1868]" captionTargetId="figure@21.[243,1343,1431,1868]" captionTargetPageId="21" captionText="FIGURE 15. Dorsal view of the head and anterior part of the body of topotypical Gerrhosaurus multilineatus (BM 1904.5.2.32) from Duque de Bragança, Angola. (Photo: P. Campbell)" httpUri="https://zenodo.org/record/247081/files/figure.png" pageId="22" pageNumber="487">15</figureCitation>
) these specimens agree well with Bocage’s description. Although somewhat faded, cream coloured longitudinal stripes, with black borders, are present on the back, at least anteriorly. In BM 1904.5.2.32 there are dorsolateral stripes as well as a vertebral stripe (
<figureCitation id="DCF72A64080CFFFF9FD5DB969A96FD78" box="[1154,1238,655,680]" captionStart="FIGURE 15" captionStartId="21.[151,255,1890,1913]" captionTargetBox="[243,1323,1431,1868]" captionTargetId="figure@21.[243,1343,1431,1868]" captionTargetPageId="21" captionText="FIGURE 15. Dorsal view of the head and anterior part of the body of topotypical Gerrhosaurus multilineatus (BM 1904.5.2.32) from Duque de Bragança, Angola. (Photo: P. Campbell)" httpUri="https://zenodo.org/record/247081/files/figure.png" pageId="22" pageNumber="487">Fig. 15</figureCitation>
), as described by Bocage (1866a), whereas BM 1904.5.2.33 appears to have only dorsolateral stripes. The two specimens (about 
<quantity id="83349B04080CFFFF9E25DBAD9E83FD3F" metricMagnitude="-1" metricUnit="m" metricValue="1.7" pageId="22" pageNumber="487" unit="mm" value="170.0">170 mm</quantity>
and 
<quantity id="83349B04080CFFFF9BAEDBCE9F16FD3F" box="[249,342,727,752]" metricMagnitude="-1" metricUnit="m" metricValue="1.5" pageId="22" pageNumber="487" unit="mm" value="150.0">150 mm</quantity>
SVL respectively) appear to be adults.
</paragraph>
<caption id="10B36669080CFFFF9BC0DF8A9FA9F91A" httpUri="https://zenodo.org/record/247082/files/figure.png" pageId="22" pageNumber="487" targetBox="[151,1436,807,1661]" targetPageId="22">
<paragraph id="447336E1080CFFFF9BC0DF8A9FA9F91A" blockId="22.[151,1436,1683,1738]" pageId="22" pageNumber="487">
<emphasis id="76B8EAF3080CFFFF9BC0DF8A9F6DF97A" bold="true" box="[151,301,1683,1706]" pageId="22" pageNumber="487">FIGURE 16.</emphasis>
Juvenile 
<taxonomicName id="83CC4D62080CFFFF9AC8DF8A9C35F97A" box="[415,629,1683,1706]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080CFFFF9AC8DF8A9C35F97A" box="[415,629,1683,1706]" italics="true" pageId="22" pageNumber="487">Gerrhosaurus bulsi</emphasis>
</taxonomicName>
from Lake Carumbo base camp, Lunda Norte Province, Angola (PEM R19475). (Photo: W.R. Branch)
</paragraph>
</caption>
<paragraph id="447336E1080CFFFE9B90DFE19A79FD1D" blockId="22.[151,1436,1783,2024]" lastBlockId="23.[151,1437,151,789]" lastPageId="23" lastPageNumber="488" pageId="22" pageNumber="487">
Laurent (1964) presented data for a large series of 
<taxonomicName id="83CC4D62080CFFFF985ADFEE9DEAF8C0" box="[781,938,1783,1808]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="genus">
<emphasis id="76B8EAF3080CFFFF985ADFEE9DEAF8C0" box="[781,938,1783,1808]" italics="true" pageId="22" pageNumber="487">Gerrhosaurus</emphasis>
</taxonomicName>
from 
<collectingCountry id="3CDB7671080CFFFF98A5DFEE9A04F8C0" box="[1010,1092,1783,1808]" name="Angola" pageId="22" pageNumber="487">Angola</collectingCountry>
which he referred to 
<taxonomicName id="83CC4D62080CFFFF9E69DFE19BD4F8C0" box="[1342,1428,1783,1808]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080CFFFF9E69DFE19BD4F8C0" box="[1342,1428,1783,1808]" italics="true" pageId="22" pageNumber="487">G. bulsi</emphasis>
</taxonomicName>
, contrasting these with a specimen from ‘Mayombe’ (may refer to the region from western 
<collectingCountry id="3CDB7671080CFFFF9FFDDE029AB5F8E4" box="[1194,1269,1819,1844]" name="Gabon" pageId="22" pageNumber="487">Gabon</collectingCountry>
southwards to western D.R.C., or to the Mayombe massif in 
<collectingCountry id="3CDB7671080CFFFF998CDE269D81F887" box="[731,961,1855,1880]" name="Republic of the Congo" pageId="22" pageNumber="487">Republic of Congo</collectingCountry>
) which he referred to 
<taxonomicName id="83CC4D62080CFFFF9FB6DE599F6CF8AC" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="subSpecies" species="nigrolineatus" subSpecies="nigrolineatus">
<emphasis id="76B8EAF3080CFFFF9FB6DE599F6CF8AC" italics="true" pageId="22" pageNumber="487">G. nigrolineatus nigrolineatus</emphasis>
</taxonomicName>
(because of its “blackish colouration”, p. 54), and four specimens from Pweto at the northern end of Lake Mweru in Katanga Province, D.R.C. which he referred to 
<taxonomicName id="83CC4D62080CFFFF9824DE919AF4F870" box="[883,1204,1927,1952]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="subSpecies" species="nigrolineatus" subSpecies="intermedius">
<emphasis id="76B8EAF3080CFFFF9824DE919AF4F870" box="[883,1204,1927,1952]" italics="true" pageId="22" pageNumber="487">G. nigrolineatus intermedius</emphasis>
</taxonomicName>
. Laurent (1964: 54) noted that if the “ 
<typeStatus id="9B778843080CFFFF9A00DEB29FCBF813" box="[343,395,1963,1987]" pageId="22" pageNumber="487">type</typeStatus>
” of 
<taxonomicName id="83CC4D62080CFFFF9AEEDEB59C29F814" box="[441,617,1963,1988]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080CFFFF9AEEDEB59C29F814" box="[441,617,1963,1988]" italics="true" pageId="22" pageNumber="487">G. multilineatus</emphasis>
</taxonomicName>
was a young 
<taxonomicName id="83CC4D62080CFFFF9855DEB59D16F814" box="[770,854,1963,1988]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080CFFFF9855DEB59D16F814" box="[770,854,1963,1988]" italics="true" pageId="22" pageNumber="487">G. bulsi</emphasis>
</taxonomicName>
, the former name would have priority. According to Laurent (1964), young 
<taxonomicName id="83CC4D62080CFFFF9AF4DEC99FBBF838" box="[419,507,1999,2024]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080CFFFF9AF4DEC99FBBF838" box="[419,507,1999,2024]" italics="true" pageId="22" pageNumber="487">G. bulsi</emphasis>
</taxonomicName>
have a (striped) dorsal colour pattern similar to that of 
<taxonomicName id="83CC4D62080CFFFF9FD0DEC99B7CF838" box="[1159,1340,1999,2024]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="22" pageNumber="487" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080CFFFF9FD0DEC99B7CF838" box="[1159,1340,1999,2024]" italics="true" pageId="22" pageNumber="487">G. nigrolineatus</emphasis>
</taxonomicName>
(striped throughout life), but this pattern gradually fades and adult 
<taxonomicName id="83CC4D62080DFFFE986BD9819DD3FF60" box="[828,915,151,176]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080DFFFE986BD9819DD3FF60" box="[828,915,151,176]" italics="true" pageId="23" pageNumber="488">G. bulsi</emphasis>
</taxonomicName>
(
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SVL and larger) display a uniformly coloured and unpatterned dorsum (
<figureCitation id="DCF72A64080DFFFE9975D9A59C36FF04" box="[546,630,188,213]" captionStart="FIGURE 12" captionStartId="20.[151,255,1178,1201]" captionTargetBox="[151,1436,518,1156]" captionTargetId="figure@20.[151,1436,518,1156]" captionTargetPageId="20" captionText="FIGURE 12. Gerrhosaurus bulsi (PEM R 18941) from Kalumbila Village, Northwest Province, Zambia. (Photo: W. R. Branch)" httpUri="https://zenodo.org/record/247078/files/figure.png" pageId="23" pageNumber="488">Fig. 12</figureCitation>
). However, juvenile 
<taxonomicName id="83CC4D62080DFFFE9835D9A49DF7FF05" box="[866,951,188,213]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080DFFFE9835D9A49DF7FF05" box="[866,951,188,213]" italics="true" pageId="23" pageNumber="488">G. bulsi</emphasis>
</taxonomicName>
are not always striped: PEM R19475 from Lake Carumbo base camp in 
<collectingCountry id="3CDB7671080DFFFE9AB5D9C69C73FF28" box="[482,563,223,248]" name="Angola" pageId="23" pageNumber="488">Angola</collectingCountry>
(
<figureCitation id="DCF72A64080DFFFE9913D9C69CD9FF28" box="[580,665,223,248]" captionStart="FIGURE 16" captionStartId="22.[151,255,1683,1706]" captionTargetBox="[151,1436,807,1661]" captionTargetId="figure@22.[151,1436,807,1661]" captionTargetPageId="22" captionText="FIGURE 16. Juvenile Gerrhosaurus bulsi from Lake Carumbo base camp, Lunda Norte Province, Angola (PEM R 19475). (Photo: W. R. Branch)" httpUri="https://zenodo.org/record/247082/files/figure.png" pageId="23" pageNumber="488">Fig. 16</figureCitation>
), used for the molecular analysis, is brown with dark bands on the flanks and scatterered dark scales on the back, but it lacks any distinct dorsolateral (and vertebral) stripes, whether pale, black, or pale with black borders. Bocage’s (1866a) description of 
<taxonomicName id="83CC4D62080DFFFE98BAD8319AE2FE90" box="[1005,1186,295,320]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080DFFFE98BAD8319AE2FE90" box="[1005,1186,295,320]" italics="true" pageId="23" pageNumber="488">G. multilineatus</emphasis>
</taxonomicName>
was based on “three specimens of identical colour” (see Bocage 1866b: 44, a paper preceding the description). However, although Bocage (1866a) described all three 
<typeStatus id="9B778843080DFFFE9974D8699CC5FE58" box="[547,645,368,392]" pageId="23" pageNumber="488" type="syntype">syntypes</typeStatus>
as having olive backs with narrow yellow, black-bordered dorsolateral and vertebral stripes, he noted that the largest (apparently adult) specimen (
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SVL + 
<quantity id="83349B04080DFFFE9FEFD88D9B57FE7C" box="[1208,1303,404,429]" metricMagnitude="-1" metricUnit="m" metricValue="2.5" pageId="23" pageNumber="488" unit="mm" value="250.0">250 mm</quantity>
tail length) also had three similar but narrower stripes in each of the interspaces between dorsolateral and vertebral stripes, while in the other (smaller) specimens these intermediate stripes were replaced by black markings. From the available information it therefore seems that, at the very least, the largest specimen examined by Bocage (1866a) is not conspecific with 
<taxonomicName id="83CC4D62080DFFFE9AC4DB3C9FADFDED" box="[403,493,548,573]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="bulsi">
<emphasis id="76B8EAF3080DFFFE9AC4DB3C9FADFDED" box="[403,493,548,573]" italics="true" pageId="23" pageNumber="488">G. bulsi</emphasis>
</taxonomicName>
(adults are unstriped). Whether or not Bocage’s specimens are referable to 
<taxonomicName id="83CC4D62080DFFFE9ED3DB3C9F6BFDB0" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080DFFFE9ED3DB3C9F6BFDB0" italics="true" pageId="23" pageNumber="488">G. nigrolineatus</emphasis>
</taxonomicName>
, or a separate species, is unclear. However, the two topotypes of 
<taxonomicName id="83CC4D62080DFFFE9F56DB519AF2FDB0" box="[1025,1202,583,608]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080DFFFE9F56DB519AF2FDB0" box="[1025,1202,583,608]" italics="true" pageId="23" pageNumber="488">G. multilineatus</emphasis>
</taxonomicName>
do not have the very large heads typical of adult 
<taxonomicName id="83CC4D62080DFFFE9A98DB749CC2FD55" box="[463,642,620,645]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="nigrolineatus">
<emphasis id="76B8EAF3080DFFFE9A98DB749CC2FD55" box="[463,642,620,645]" italics="true" pageId="23" pageNumber="488">G. nigrolineatus</emphasis>
</taxonomicName>
, 
<taxonomicName id="83CC4D62080DFFFE99C6DB749D74FD55" box="[657,820,620,645]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="76B8EAF3080DFFFE99C6DB749D74FD55" box="[657,820,620,645]" italics="true" pageId="23" pageNumber="488">G. intermedius</emphasis>
</taxonomicName>
and 
<taxonomicName id="83CC4D62080DFFFE983CDB749D9AFD54" box="[875,986,621,644]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="auritus">
<emphasis id="76B8EAF3080DFFFE983CDB749D9AFD54" box="[875,986,621,644]" italics="true" pageId="23" pageNumber="488">G. auritus</emphasis>
</taxonomicName>
(i.e. head length into SVL five times or less; see Loveridge 1942, FitzSimons 1943). Based on scaled photographs of BM 1904.5.2.33 and BM 1904.5.2.32, head length is contained in SVL about 5.6 and 5.7 times, respectively.
</paragraph>
<paragraph id="447336E1080DFFFE9B90DBCE9A31FCC5" blockId="23.[151,1437,151,789]" pageId="23" pageNumber="488">
Resolution of the taxonomic status of 
<taxonomicName id="83CC4D62080DFFFE99D0DBC19D7BFD20" box="[647,827,727,752]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="multilineatus">
<emphasis id="76B8EAF3080DFFFE99D0DBC19D7BFD20" box="[647,827,727,752]" italics="true" pageId="23" pageNumber="488">G. multilineatus</emphasis>
</taxonomicName>
must await the collection of material from the 
<typeStatus id="9B778843080DFFFE9E3BDBC19BDCFD20" box="[1388,1436,728,752]" pageId="23" pageNumber="488">type</typeStatus>
locality for molecular analysis, and a detailed morphological evaluation of the complex.
</paragraph>
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<paragraph id="447336E1080DFFFE9BC0DFA59F75F923" blockId="23.[151,1437,1724,1779]" pageId="23" pageNumber="488">
<emphasis id="76B8EAF3080DFFFE9BC0DFA59F68F903" bold="true" box="[151,296,1724,1747]" pageId="23" pageNumber="488">FIGURE 17.</emphasis>
<taxonomicName id="83CC4D62080DFFFE9A67DFA59C09F903" box="[304,585,1724,1747]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080DFFFE9A67DFA59C09F903" box="[304,585,1724,1747]" italics="true" pageId="23" pageNumber="488">Gerrhosaurus flavigularis</emphasis>
</taxonomicName>
from Suikerbosrand Nature Reserve, Gauteng Province, South Africa. (Photo: W.R. Schmidt)
</paragraph>
</caption>
<paragraph id="447336E1080DFFFE9BC0DE059C73F8E5" blockId="23.[151,1436,1820,2026]" box="[151,563,1820,1845]" pageId="23" pageNumber="488">
<emphasis id="76B8EAF3080DFFFE9BC0DE059C73F8E5" bold="true" box="[151,563,1820,1845]" pageId="23" pageNumber="488">
Status of 
<taxonomicName id="83CC4D62080DFFFE9A52DE059C73F8E5" box="[261,563,1820,1845]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080DFFFE9A52DE059C73F8E5" bold="true" box="[261,563,1820,1845]" italics="true" pageId="23" pageNumber="488">Gerrhosaurus flavigularis</emphasis>
</taxonomicName>
</emphasis>
</paragraph>
<paragraph id="447336E1080DFFF19B90DE599F5DFE90" blockId="23.[151,1436,1820,2026]" lastBlockId="24.[151,1436,151,320]" lastPageId="24" lastPageNumber="489" pageId="23" pageNumber="488">
While there is some sub-structuring within 
<taxonomicName id="83CC4D62080DFFFE9991DE589D2AF889" box="[710,874,1856,1881]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080DFFFE9991DE589D2AF889" box="[710,874,1856,1881]" italics="true" pageId="23" pageNumber="488">G. flavigularis</emphasis>
</taxonomicName>
(
<figureCitation id="DCF72A64080DFFFE9829DE599D94F889" box="[894,980,1856,1881]" captionStart="FIGURE 17" captionStartId="23.[151,255,1724,1747]" captionTargetBox="[151,1436,844,1700]" captionTargetId="figure@23.[151,1436,844,1701]" captionTargetPageId="23" captionText="FIGURE 17. Gerrhosaurus flavigularis from Suikerbosrand Nature Reserve, Gauteng Province, South Africa. (Photo: W. R. Schmidt)" httpUri="https://zenodo.org/record/247083/files/figure.png" pageId="23" pageNumber="488">Fig. 17</figureCitation>
), with populations from Eastern 
<collectingCountry id="3CDB7671080DFFFE9E0DDE599EA2F8AE" name="South Africa" pageId="23" pageNumber="488">South Africa</collectingCountry>
, Northern 
<collectingCountry id="3CDB7671080DFFFE9A0EDE7C9FAAF8AE" box="[345,490,1893,1918]" name="South Africa" pageId="23" pageNumber="488">South Africa</collectingCountry>
and East Africa (
<tableCitation id="094E035A080DFFFE99F8DE7C9D43F8AD" box="[687,771,1893,1918]" captionStart="TABLE 1" captionStartId="4.[151,239,152,175]" captionText="TABLE 1. Sample numbers, collecting localities, museum voucher accession numbers (CAS = California Academy of" pageId="23" pageNumber="488">Table 1</tableCitation>
) all identifiable as subclades in the phylogeny (
<figureCitation id="DCF72A64080DFFFE9E76DE7C9B22F8AD" box="[1313,1378,1893,1918]" captionStart="FIGURE 1" captionStartId="7.[151,255,1859,1882]" captionText="FIGURE 1. Maximum likelihood topology for Gerrhosauridae. Supported nodes are indicated by circles (black: ≥ 70 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities; grey: ≥ 65 % likelihood bootstrap and ≥ 95 % Bayesian posterior probabilities). Genera and major clades are indicated, and corresponding letters A-F match the text." pageId="23" pageNumber="488">Fig. 1</figureCitation>
), we consider this assemblage a single species pending a more detailed phylogeographical and morphological analysis. The 
<typeStatus id="9B778843080DFFFE9B9EDEB79EB9F816" box="[201,249,1966,1990]" pageId="23" pageNumber="488">type</typeStatus>
locality for 
<taxonomicName id="83CC4D62080DFFFE9AD5DEB79C62F816" box="[386,546,1965,1990]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="23" pageNumber="488" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF3080DFFFE9AD5DEB79C62F816" box="[386,546,1965,1990]" italics="true" pageId="23" pageNumber="488">G. flavigularis</emphasis>
</taxonomicName>
of “South [or southern] Africa” (see Bauer 
<emphasis id="76B8EAF3080DFFFE9F5EDEB79A01F816" box="[1033,1089,1965,1990]" italics="true" pageId="23" pageNumber="488">et al.</emphasis>
1994) was restricted to an area in the central Eastern Cape Province, 
<collectingCountry id="3CDB7671080DFFFE9901DEC89CAAF83A" box="[598,746,2001,2026]" name="South Africa" pageId="23" pageNumber="488">South Africa</collectingCountry>
(Bauer 2000), but the species occurs extensively from the Western Cape (
<collectingCountry id="3CDB76710802FFF19A1AD98E9FA6FF60" box="[333,486,151,176]" name="South Africa" pageId="24" pageNumber="489">South Africa</collectingCountry>
) northwards through southern and eastern Africa to 
<collectingCountry id="3CDB76710802FFF19F08D98E9AFEFF60" box="[1119,1214,151,176]" name="Ethiopia" pageId="24" pageNumber="489">Ethiopia</collectingCountry>
(Loveridge 1942). Although not sampled for this study, should the apparently disjunct population in the Western and Eastern Cape Provinces be found to represent a unique lineage, the name 
<taxonomicName id="83CC4D620802FFF19812D9F99DA7FF28" box="[837,999,223,248]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="489" phylum="Chordata" rank="species" species="flavigularis">
<emphasis id="76B8EAF30802FFF19812D9F99DA7FF28" box="[837,999,223,248]" italics="true" pageId="24" pageNumber="489">G. flavigularis</emphasis>
</taxonomicName>
would be applicable to it. If the other population in southern and East Africa proves to be a separate species, the name 
<taxonomicName id="83CC4D620802FFF19F4ED81D9BDCFECC" authority="Smith, 1844" authorityName="Smith" authorityYear="1844" box="[1049,1436,260,285]" class="Reptilia" family="Gerrhosauridae" genus="Gerrhosaurus" kingdom="Animalia" order="Squamata" pageId="24" pageNumber="489" phylum="Chordata" rank="species" species="bibroni">
<emphasis id="76B8EAF30802FFF19F4ED81D9B4DFECD" box="[1049,1293,260,285]" italics="true" pageId="24" pageNumber="489">Gerrhosaurus bibroni</emphasis>
Smith, 1844
</taxonomicName>
is available.
</paragraph>
</subSubSection>
</treatment>
</document>