treatments-xml/data/03/D1/87/03D187A3FFF5EC44572E3D31C403FA65.xml

<document ID-DOI="http://doi.org/10.5281/zenodo.5695569" ID-GBIF-Dataset="7cb9ff08-b69f-4a75-b5c9-64aeff699f40" ID-GBIF-Taxon="127623528" ID-Zenodo-Dep="5695569" LSID="urn:lsid:zoobank.org:act:E94496B3-AC04-4FE2-AFA6-8CC7DBE3A63A" checkinTime="1485191403157" checkinUser="plazi" docAuthor="Sara Ruane, Frank T. Burbrink, Bernard Randriamahatantsoa &amp; Christopher J. Raxworthy" docDate="2016" docId="03D187A3FFF5EC44572E3D31C403FA65" docLanguage="en" docName="copeia.104.3.712.pdf" docOrigin="Copeia 104 (3)" docStyle="DocumentStyle{}" docTitle="Madagascarophis lolo Ruane, Burbrink, Randriamahatantsoa &amp; Raxworthy, 2016, new species" docType="treatment" docUuid="E94496B3-AC04-4FE2-AFA6-8CC7DBE3A63A" docUuidSource="ZooBank" docVersion="7" lastPageId="8" lastPageNumber="719" masterDocId="FFE8FFDBFFF0EC4C541A3B68C545FFAC" masterDocTitle="The Cat-eyed Snakes of Madagascar: Phylogeny and Description of a New Species of Madagascarophis (Serpentes: Lamprophiidae) from the Tsingy of Ankarana" masterLastPageNumber="721" masterPageNumber="711" pageId="5" pageNumber="716" updateTime="1636818723269" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>The Cat-eyed Snakes of Madagascar: Phylogeny and Description of a New Species of Madagascarophis (Serpentes: Lamprophiidae) from the Tsingy of Ankarana</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Sara Ruane</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Frank T. Burbrink</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bernard Randriamahatantsoa</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Christopher J. Raxworthy</mods:namePart>
</mods:name>
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<mods:title>Copeia</mods:title>
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<mods:part>
<mods:date>2016</mods:date>
<mods:detail type="volume">
<mods:number>104</mods:number>
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<mods:detail type="issue">
<mods:number>3</mods:number>
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<mods:start>711</mods:start>
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<mods:identifier type="DOI">10.1643/Ch-15-346</mods:identifier>
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<mods:identifier type="Zenodo-Dep">269699</mods:identifier>
<mods:identifier type="ZooBank">7D899252-C0B1-40CE-9332-ACBC7A6BFA2B</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5695569" ID-GBIF-Taxon="127623528" ID-Zenodo-Dep="5695569" LSID="urn:lsid:zoobank.org:act:E94496B3-AC04-4FE2-AFA6-8CC7DBE3A63A" httpUri="http://treatment.plazi.org/id/03D187A3FFF5EC44572E3D31C403FA65" lastPageId="8" lastPageNumber="719" pageId="5" pageNumber="716">
<subSubSection pageId="5" pageNumber="716" type="nomenclature">
<paragraph blockId="5.[820,1449,1625,1737]" box="[820,1231,1625,1648]" pageId="5" pageNumber="716">
<heading bold="true" box="[820,1231,1625,1648]" fontSize="9" level="4" pageId="5" pageNumber="716" reason="6">
<emphasis bold="true" box="[820,1231,1625,1648]" pageId="5" pageNumber="716">
<taxonomicName box="[820,1071,1625,1648]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="716" phylum="Chordata" rank="species" species="lolo" status="new species">
<emphasis bold="true" box="[820,1071,1625,1648]" italics="true" pageId="5" pageNumber="716">Madagascarophis lolo</emphasis>
</taxonomicName>
, 
<taxonomicNameLabel box="[1087,1231,1625,1648]" pageId="5" pageNumber="716" rank="species">new species</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph blockId="5.[820,1449,1625,1737]" pageId="5" pageNumber="716">urn:lsid:zoobank.org:act:E94496B3-AC04-4FE2-AFA6- 8CC7DBE3A63A</paragraph>
</subSubSection>
<subSubSection box="[820,1125,1714,1737]" pageId="5" pageNumber="716" type="description">
<paragraph blockId="5.[820,1449,1625,1737]" box="[820,1125,1714,1737]" pageId="5" pageNumber="716">
Figures 1̡ 2̡ 3̡ 4̡ 5; 
<tableCitation box="[1046,1125,1714,1737]" captionStart="Table 1" captionStartId="4.[142,190,130,150]" captionTargetBox="[142,1529,222,410]" captionText="Table 1. Genetic similarity of Madagascarophis by maximum percent uncorrected pairwise identity between / within species by gene; locus order is COI / Cmos / Rag 2. Note that as M. fuchsi and M. lolo are represented by one individual, there is no maximum pairwise identity available for within species variation." httpUri="http://table.plazi.org/id/DF07663DFFF4EC4854943BEAC474FF67" pageId="5" pageNumber="716" tableUuid="DF07663DFFF4EC4854943BEAC474FF67">Table 1</tableCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="5" pageNumber="716" type="materials_examined">
<paragraph blockId="5.[820,1490,1768,1940]" pageId="5" pageNumber="716">
<materialsCitation ID-GBIF-Occurrence="1425979787" collectingDate="2014-02-09" collectionCode="AMNH, RAX" collectorName="B. Randriamahatantsoa &amp; C. Raxworthy &amp; S. Ruane" country="Madagascar" elevation="102" latitude="-12.9421" location="Ankarana National Park, ~ 4 km northwest of the village of Mahamasina, tsingy karst trail" longLatPrecision="1" longitude="49.11507" municipality="Diana Region" pageId="5" pageNumber="716" specimenCode="AMNH 176422, RAX 12626" specimenCount="2" specimenCount-adult="1" specimenCount-male="1" stateProvince="Antsiranana Province" typeStatus="holotype">
<emphasis bold="true" box="[820,945,1768,1791]" italics="true" pageId="5" pageNumber="716">
<typeStatus box="[820,917,1768,1791]" pageId="5" pageNumber="716">Holotype</typeStatus>
.—
</emphasis>
<specimenCode box="[945,1112,1768,1791]" pageId="5" pageNumber="716">AMNH 176422</specimenCode>
(
<specimenCode box="[1127,1258,1768,1791]" pageId="5" pageNumber="716">RAX 12626</specimenCode>
), 
<specimenCount box="[1279,1335,1768,1791]" pageId="5" pageNumber="716" type="adult">adult</specimenCount>
<specimenCount box="[1344,1397,1768,1791]" pageId="5" pageNumber="716" type="male">male</specimenCount>
(Fig. 1), 
<collectingCountry box="[820,950,1798,1821]" name="Madagascar" pageId="5" pageNumber="716">Madagascar</collectingCountry>
, 
<collectingRegion box="[963,1205,1798,1821]" country="Madagascar" name="Antsiranana" pageId="5" pageNumber="716">Antsiranana Province</collectingRegion>
, 
<collectingMunicipality box="[1218,1371,1798,1821]" pageId="5" pageNumber="716">Diana Region</collectingMunicipality>
, 
<location LSID="urn:lsid:plazi:treatment:03D187A3FFF5EC44572E3D31C403FA65:8EA7606EFFF5EC4951733C6EC10FF8F4" country="Madagascar" latitude="-12.9421" longLatPrecision="1" longitude="49.11507" municipality="Diana Region" name="Ankarana National Park, ~ 4 km northwest of the village of Mahamasina, tsingy karst trail" pageId="5" pageNumber="716" stateProvince="Antsiranana Province">
Ankarana National Park, 
<emphasis bold="true" box="[989,1007,1828,1850]" pageId="5" pageNumber="716">~</emphasis>
<quantity box="[1007,1065,1828,1851]" metricMagnitude="3" metricUnit="m" metricValue="4.0" pageId="5" pageNumber="716" unit="km" value="4.0">4 km</quantity>
northwest of the village of Mahamasina, tsingy karst trail
</location>
, 
<elevation box="[1123,1204,1858,1880]" pageId="5" pageNumber="716">
<quantity box="[1151,1204,1858,1880]" metricMagnitude="0" metricUnit="m" metricValue="2.0" pageId="5" pageNumber="716" unit="m" value="2.0">102 m</quantity>
</elevation>
elevation, 
<geoCoordinate box="[1353,1483,1857,1880]" direction="east" orientation="longitude" pageId="5" pageNumber="716" precision="1" value="49.11507">49.11507°E</geoCoordinate>
, 
<geoCoordinate box="[820,945,1887,1909]" direction="south" orientation="latitude" pageId="5" pageNumber="716" precision="1" value="-12.9421">12.94210°S</geoCoordinate>
, 
<collectingDate box="[960,1146,1887,1910]" pageId="5" pageNumber="716" value="2014-02-09">
<date box="[960,1146,1887,1910]" pageId="5" pageNumber="716" value="2014-02-09">9 February 2014</date>
</collectingDate>
, 1930 hours, 
<collectorName pageId="5" pageNumber="716">B. Randriamahatantsoa</collectorName>
, 
<collectorName box="[916,1069,1917,1940]" pageId="5" pageNumber="716">C. Raxworthy</collectorName>
, 
<collectorName box="[1081,1180,1918,1939]" pageId="5" pageNumber="716">S. Ruane</collectorName>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="717" pageId="5" pageNumber="716" type="diagnosis">
<paragraph blockId="5.[820,1490,1971,2024]" lastBlockId="6.[142,813,868,2024]" lastPageId="6" lastPageNumber="717" pageId="5" pageNumber="716">
<emphasis bold="true" box="[820,953,1971,1994]" italics="true" pageId="5" pageNumber="716">Diagnosis.—</emphasis>
A new species of 
<taxonomicName box="[1160,1338,1971,1993]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="716" phylum="Chordata" rank="genus">
<emphasis box="[1160,1338,1971,1993]" italics="true" pageId="5" pageNumber="716">Madagascarophis</emphasis>
</taxonomicName>
than can be diagnosed from its congeners by the following combination of characters: an overall gray body color with a black vertebral stripe and alternating light gray blotches down the dorsum, 25 scale rows at midbody, 189 ventral scales and 56 divided subcaudal scales, with extended contact of the posterior inframaxillary scales. 
<emphasis box="[512,749,987,1009]" italics="true" pageId="6" pageNumber="717">
<taxonomicName box="[512,743,987,1009]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo" status="new species">Madagascarophis lolo</taxonomicName>
̗
</emphasis>
<taxonomicNameLabel pageId="6" pageNumber="717" rank="species">new species</taxonomicNameLabel>
, differs from all other species of 
<taxonomicName box="[634,812,1017,1039]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="genus">
<emphasis box="[634,812,1017,1039]" italics="true" pageId="6" pageNumber="717">Madagascarophis</emphasis>
</taxonomicName>
having a gray body color combined with an alternating pattern of pale gray blotches along the vertebral column and the presence of black scales on the vertebral row scales, giving the appearance of a thin black dorsal stripe (Figs. 1, 3, 4). This overall combination of color and pattern is unique among species of 
<taxonomicName box="[336,514,1196,1218]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="genus">
<emphasis box="[336,514,1196,1218]" italics="true" pageId="6" pageNumber="717">Madagascarophis</emphasis>
</taxonomicName>
(see 
<bibRefCitation author="Domergue, C. A." box="[568,751,1196,1219]" journalOrPublisher="Bulletin du Museum national d'Histoire Naturelle, Paris" pageId="6" pageNumber="717" pagination="455 - 489" part="9" refString="Domergue, C. A. 1987. Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens, 1952. Bulletin du Museum national d'Histoire Naturelle, Paris 9: 455 - 489." title="Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens" type="journal article" year="1987">Domergue, 1987</bibRefCitation>
:fig. 4 for comparison). 
<emphasis box="[343,577,1226,1248]" italics="true" pageId="6" pageNumber="717">
<taxonomicName box="[343,571,1226,1248]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo" status="new species">Madagascarophis lolo</taxonomicName>
̗
</emphasis>
<taxonomicNameLabel box="[589,727,1226,1249]" pageId="6" pageNumber="717" rank="species">new species</taxonomicNameLabel>
, differs from all other species with the exception of 
<taxonomicName box="[671,772,1256,1278]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[671,772,1256,1278]" italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
by having extended contact of the posterior inframaxillary scales (
<figureCitation box="[219,281,1315,1338]" captionStart="Fig. 5" captionStartId="6.[142,170,777,797]" captionTargetBox="[145,1528,131,748]" captionTargetId="figure@6.[144,1528,130,748]" captionTargetPageId="6" captionText="Fig. 5. Ventral head view, showing the posterior infralabial scale contact of Madagascarophis lolo (A; RAX 12626), M. fuchsi (B; RAX 12424), M. colubrinus (C; RAX 10540); a scale bar is provided for each species. Photos by SR (A, C) and FTB (B)." httpUri="https://zenodo.org/record/269703/files/figure.png" pageId="6" pageNumber="717">Fig. 5</figureCitation>
). We note the specimen of 
<emphasis box="[588,667,1315,1337]" italics="true" pageId="6" pageNumber="717">
<taxonomicName box="[588,661,1315,1337]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo" status="new species">M. lolo</taxonomicName>
̗
</emphasis>
<taxonomicNameLabel box="[675,808,1315,1338]" pageId="6" pageNumber="717" rank="species">new species</taxonomicNameLabel>
, has a single gular scale that infringes on the posterior end of the posterior inframaxillaries. However, with the exception of 
<taxonomicName box="[174,272,1405,1427]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[174,272,1405,1427]" italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
, the posterior inframaxillary contact of 
<emphasis box="[730,812,1405,1427]" italics="true" pageId="6" pageNumber="717">
<taxonomicName box="[730,806,1405,1427]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo" status="new species">M. lolo</taxonomicName>
̗
</emphasis>
<taxonomicNameLabel box="[142,274,1434,1457]" pageId="6" pageNumber="717" rank="species">new species</taxonomicNameLabel>
, is still much greater than for the other species of 
<taxonomicName box="[142,320,1464,1486]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="genus">
<emphasis box="[142,320,1464,1486]" italics="true" pageId="6" pageNumber="717">Madagascarophis</emphasis>
</taxonomicName>
(
<figureCitation box="[335,398,1464,1487]" captionStart="Fig. 5" captionStartId="6.[142,170,777,797]" captionTargetBox="[145,1528,131,748]" captionTargetId="figure@6.[144,1528,130,748]" captionTargetPageId="6" captionText="Fig. 5. Ventral head view, showing the posterior infralabial scale contact of Madagascarophis lolo (A; RAX 12626), M. fuchsi (B; RAX 12424), M. colubrinus (C; RAX 10540); a scale bar is provided for each species. Photos by SR (A, C) and FTB (B)." httpUri="https://zenodo.org/record/269703/files/figure.png" pageId="6" pageNumber="717">Fig. 5</figureCitation>
; see 
<bibRefCitation author="Glaw, F. &amp; C. Kucharzewski &amp; J. Koehler &amp; M. Vences &amp; Z. T. Nagy" box="[452,653,1464,1487]" journalOrPublisher="Zootaxa" pageId="6" pageNumber="717" pagination="317 - 332" part="3630" refString="Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332." title="Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar" type="journal article" year="2013" yearSuffix="a">Glaw et al., 2013a</bibRefCitation>
for additional examples).
</paragraph>
<caption httpUri="https://zenodo.org/record/269703/files/figure.png" pageId="6" pageNumber="717" targetBox="[145,1528,131,748]" targetPageId="6">
<paragraph blockId="6.[142,1529,777,824]" pageId="6" pageNumber="717">
Fig. 5. Ventral head view, showing the posterior infralabial scale contact of 
<taxonomicName box="[880,1093,777,797]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo">
<emphasis bold="true" box="[880,1093,777,797]" italics="true" pageId="6" pageNumber="717">Madagascarophis lolo</emphasis>
</taxonomicName>
(A; RAX12626), 
<taxonomicName box="[1256,1342,777,797]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="fuchsi">
<emphasis bold="true" box="[1256,1342,777,797]" italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
(B; RAX12424), 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="colubrinus">
<emphasis bold="true" italics="true" pageId="6" pageNumber="717">M. colubrinus</emphasis>
</taxonomicName>
(C; RAX10540); a scale bar is provided for each species. Photos by SR (A, C) and FTB (B).
</paragraph>
</caption>
<paragraph blockId="6.[142,813,868,2024]" pageId="6" pageNumber="717">
<emphasis box="[166,395,1524,1546]" italics="true" pageId="6" pageNumber="717">
<taxonomicName box="[166,389,1524,1546]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo" status="new species">Madagascarophis lolo</taxonomicName>
̗
</emphasis>
<taxonomicNameLabel box="[402,535,1524,1547]" pageId="6" pageNumber="717" rank="species">new species</taxonomicNameLabel>
, differs from 
<taxonomicName box="[681,777,1524,1547]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[681,777,1524,1547]" italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
by having a lower number of infralabial scales (10 
<taxonomicName box="[659,730,1554,1576]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[659,730,1554,1576]" italics="true" pageId="6" pageNumber="717">M. lolo</emphasis>
</taxonomicName>
vs. 12– 13 
<taxonomicName box="[179,277,1584,1606]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[179,277,1584,1606]" italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
) and a higher ventral scale count (171–174 
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<emphasis italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
vs. 1 8 9 
<taxonomicName box="[325,408,1614,1636]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[325,408,1614,1636]" italics="true" pageId="6" pageNumber="717">M. lolo</emphasis>
</taxonomicName>
). However, this 1 5 ventral scale difference falls within the intraspecific range of other species (e.g., 35 ventral scales in 
<taxonomicName box="[426,589,1673,1696]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[426,589,1673,1696]" italics="true" pageId="6" pageNumber="717">M. meridionalis</emphasis>
</taxonomicName>
). It differs from the other species of 
<taxonomicName box="[343,521,1703,1725]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="genus">
<emphasis box="[343,521,1703,1725]" italics="true" pageId="6" pageNumber="717">Madagascarophis</emphasis>
</taxonomicName>
except 
<taxonomicName box="[621,770,1703,1725]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[621,770,1703,1725]" italics="true" pageId="6" pageNumber="717">M. colubrinus</emphasis>
</taxonomicName>
by having a lower ventral scale count (189 
<taxonomicName box="[591,663,1733,1755]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[591,663,1733,1755]" italics="true" pageId="6" pageNumber="717">M. lolo</emphasis>
</taxonomicName>
): 183–209 in 
<taxonomicName box="[142,287,1763,1785]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[142,287,1763,1785]" italics="true" pageId="6" pageNumber="717">M. colubrinus</emphasis>
</taxonomicName>
, 205–224 in 
<taxonomicName box="[445,573,1763,1785]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[445,573,1763,1785]" italics="true" pageId="6" pageNumber="717">M. ocellatus</emphasis>
</taxonomicName>
, and 197–232 in 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="meridionalis">
<emphasis italics="true" pageId="6" pageNumber="717">M. meridionalis</emphasis>
</taxonomicName>
. A general difference between 
<emphasis box="[660,748,1793,1815]" italics="true" pageId="6" pageNumber="717">
<taxonomicName box="[660,742,1793,1815]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo" status="new species">M. lolo</taxonomicName>
̗
</emphasis>
<taxonomicNameLabel pageId="6" pageNumber="717" rank="species">new species</taxonomicNameLabel>
, and most other 
<taxonomicName box="[422,600,1822,1844]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="genus">
<emphasis box="[422,600,1822,1844]" italics="true" pageId="6" pageNumber="717">Madagascarophis</emphasis>
</taxonomicName>
is the dorsal scale count at midbody. 
<emphasis box="[355,585,1852,1874]" italics="true" pageId="6" pageNumber="717">
<taxonomicName box="[355,579,1852,1874]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo" status="new species">Madagascarophis lolo</taxonomicName>
̗
</emphasis>
<taxonomicNameLabel box="[594,727,1852,1875]" pageId="6" pageNumber="717" rank="species">new species</taxonomicNameLabel>
, has 25 dorsal scale rows as does 
<taxonomicName box="[426,524,1882,1904]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[426,524,1882,1904]" italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
, in contrast to the 27–29 typically found in 
<taxonomicName box="[351,494,1912,1934]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[351,494,1912,1934]" italics="true" pageId="6" pageNumber="717">M. colubrinus</emphasis>
</taxonomicName>
(rarely 25, and not syntopically), 29–33 in 
<taxonomicName box="[332,499,1942,1964]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[332,499,1942,1964]" italics="true" pageId="6" pageNumber="717">M. meridionalis</emphasis>
</taxonomicName>
, and 29–31 in 
<taxonomicName box="[683,812,1942,1964]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[683,812,1942,1964]" italics="true" pageId="6" pageNumber="717">M. ocellatus</emphasis>
</taxonomicName>
(
<bibRefCitation author="Glaw, F. &amp; M. Vences" box="[149,419,1971,1994]" journalOrPublisher="Verlag, Cologne, Germany" pageId="6" pageNumber="717" refString="Glaw, F., and M. Vences. 2007. Amphibians and Reptiles of Madagascar. Third edition. Verlag, Cologne, Germany." title="Amphibians and Reptiles of Madagascar. Third edition" type="book" year="2007">Glaw and Vences, 2007</bibRefCitation>
; 
<bibRefCitation author="Glaw, F. &amp; C. Kucharzewski &amp; J. Koehler &amp; M. Vences &amp; Z. T. Nagy" box="[433,643,1971,1994]" journalOrPublisher="Zootaxa" pageId="6" pageNumber="717" pagination="317 - 332" part="3630" refString="Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332." title="Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar" type="journal article" year="2013" yearSuffix="a">Glaw et al., 2013a</bibRefCitation>
). It also differs genetically from all other species in the genus, e.g., 
<taxonomicName box="[705,776,2001,2023]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[705,776,2001,2023]" italics="true" pageId="6" pageNumber="717">M. lolo</emphasis>
</taxonomicName>
vs.
</paragraph>
<paragraph blockId="6.[859,1529,868,979]" pageId="6" pageNumber="717">
<taxonomicName box="[859,954,868,890]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="717" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[859,954,868,890]" italics="true" pageId="6" pageNumber="717">M. fuchsi</emphasis>
</taxonomicName>
, 
<emphasis box="[968,1010,869,890]" italics="true" pageId="6" pageNumber="717">COI</emphasis>
uncorrected pairwise distance =9.6% (
<tableCitation box="[1435,1512,868,891]" captionStart="Table 1" captionStartId="4.[142,190,130,150]" captionTargetBox="[142,1529,222,410]" captionText="Table 1. Genetic similarity of Madagascarophis by maximum percent uncorrected pairwise identity between / within species by gene; locus order is COI / Cmos / Rag 2. Note that as M. fuchsi and M. lolo are represented by one individual, there is no maximum pairwise identity available for within species variation." httpUri="http://table.plazi.org/id/DF07663DFFF4EC4854943BEAC474FF67" pageId="6" pageNumber="717" tableUuid="DF07663DFFF4EC4854943BEAC474FF67">Table 1</tableCitation>
). For specimens not examined here (
<figureCitation box="[1260,1390,897,920]" captionStart="Appendix 1" captionStartId="10.[142,226,130,150]" captionTargetBox="[0,1530,55,101]" captionText="Appendix 1. Information for genetic samples used in this study; both field numbers and associated museum numbers are included when available (tissue sample only / not yet vouchered individuals have only field numbers). Institutional abbreviations follow Sabaj Pérez (2014), with the addition of ELO (collection of E. Courtois), RAN (collection of R. A. Nussbaum, University of Michigan), RAX (collection of C. J. Raxworthy, American Museum of Natural History), and TJP (collection of T. Papenfuss). * Collected on the 2014 Madagascar Expedition." httpUri="https://zenodo.org/record/3098139/files/figure.png" pageId="6" pageNumber="717">Appendix 1</figureCitation>
), additional data were used from 
<bibRefCitation author="Domergue, C. A." box="[1122,1323,927,949]" journalOrPublisher="Bulletin du Museum national d'Histoire Naturelle, Paris" pageId="6" pageNumber="717" pagination="455 - 489" part="9" refString="Domergue, C. A. 1987. Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens, 1952. Bulletin du Museum national d'Histoire Naturelle, Paris 9: 455 - 489." title="Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens" type="journal article" year="1987">Domergue (1987)</bibRefCitation>
and 
<bibRefCitation author="Glaw, F. &amp; C. Kucharzewski &amp; J. Koehler &amp; M. Vences &amp; Z. T. Nagy" journalOrPublisher="Zootaxa" pageId="6" pageNumber="717" pagination="317 - 332" part="3630" refString="Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332." title="Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar" type="journal article" year="2013">Glaw et al. (2013a)</bibRefCitation>
for the diagnosis.
</paragraph>
</subSubSection>
<subSubSection lastPageId="7" lastPageNumber="718" pageId="6" pageNumber="717" type="description">
<paragraph blockId="6.[859,1529,1009,1795]" pageId="6" pageNumber="717">
<emphasis bold="true" box="[859,1008,1009,1032]" italics="true" pageId="6" pageNumber="717">Description.—</emphasis>
Adult male in excellent state of preservation, tail complete, short ventral slit midbody for DNA tissue sample, lower body slit for assessing gonad development (fully formed mature testes; 10 mm length, 2 mm width). Snout–vent length 426 mm, tail length 65 mm, tail short (13% of total body length). Head length 20 mm, width 12 mm. Head distinct from neck. Eyes large, 3 mm horizontal diameter, pupil vertically elliptical. Supralabials 8, not in contact with the eye. Infralabials 10, first pair in contact behind mental, infralabials 1–5 in contact with inframaxillaries. Rostral broader than high, 3 mm wide/1.5 mm high, visible from above. Nasal divided below nostril, in contact with 1st and 2nd supralabials. Single loreal, in contact with nasal, preoculars, prefrontal, and supralabials 2 and 3. Circumoculars 9, 1 supraocular, 2 preoculars, 3 suboculars, and 3 postoculars. Temporals 4 + 4/4 + 5. Dorsal surface of head includes pair of internasals (width 1.7 mm/length of suture 2 mm), pair of prefrontals (width 2 mm/length of suture 2.1 mm), pair of supraoculars (width 2.8 mm/length 4.8 mm), frontal longer than wide (length 5.7 mm/anterior width 2.9 mm), pair of parietals (length of suture 4.6 mm). Two pairs of inframaxillaries (anterior inframaxillary length 4.9 mm, posterior inframaxillary length 3.1 mm), posterior inframaxillaries substantially in contact with each other excepting small gular scale at posterior end (
<figureCitation box="[1370,1435,1713,1736]" captionStart="Fig. 5" captionStartId="6.[142,170,777,797]" captionTargetBox="[145,1528,131,748]" captionTargetId="figure@6.[144,1528,130,748]" captionTargetPageId="6" captionText="Fig. 5. Ventral head view, showing the posterior infralabial scale contact of Madagascarophis lolo (A; RAX 12626), M. fuchsi (B; RAX 12424), M. colubrinus (C; RAX 10540); a scale bar is provided for each species. Photos by SR (A, C) and FTB (B)." httpUri="https://zenodo.org/record/269703/files/figure.png" pageId="6" pageNumber="717">Fig. 5</figureCitation>
). Dorsal scale rows 23-25-19 at 10th ventral from anterior, midbody, and 10th ventral anterior to cloaca.
</paragraph>
<paragraph blockId="6.[859,1529,1825,2024]" lastBlockId="7.[103,773,131,508]" lastPageId="7" lastPageNumber="718" pageId="6" pageNumber="717">
<emphasis bold="true" box="[859,1147,1825,1848]" italics="true" pageId="6" pageNumber="717">Coloration and pattern.—</emphasis>
Dorsal ground color gray in life, alternating lighter gray blotches/squares alongside vertebral column, many vertebral row scales black in coloration, giving general appearance of black dorsal line interrupted occasionally by gray scales (Fig. 1). At roughly the posterior 1/3 of the body, dorsal scale rows 7 and 8 occasionally black, giving spotted appearance in lateral view. Overall coloration pale gray in preservation. Color of iris in life gray/silver with gold flecking (Fig. 1), opaque gray in preservation. Dorsum of head, including rostral, internasals, prefrontals, frontal, supraoculars, and parietals gray. Supralabials 4–8 with gray and white mottling, infralabials with gray and white mottling. Slightly darker diffuse brown line runs from the posterior of eye to posterior margin of mouth. Tail gray with black mottling, darker and more contrasting compared to body, with slight flush of pale orange towards tail tip. Ventral scales cream with no pattern anteriorly, small amounts of gray flecking on ventral scales beginning at the posterior 1/3 of the body, continuing and increasing in intensity onto the subcaudal scales.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="718" type="biology_ecology">
<paragraph blockId="7.[103,773,539,1065]" pageId="7" pageNumber="718">
<emphasis bold="true" box="[103,294,539,562]" italics="true" pageId="7" pageNumber="718">Natural history.—</emphasis>
Similar to other species of 
<taxonomicName box="[588,766,539,561]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[588,766,539,561]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
, 
<taxonomicName box="[103,180,569,591]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[103,180,569,591]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
appears to be crepuscular/nocturnal; the specimen was found active on the ground at 1930 hours on tsingy karst rocks, in an exposed area with low scrub habitat. This is very similar to what has been described for 
<taxonomicName authority="Glaw et al., 2013a" authorityName="Glaw et al." authorityYear="2013" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[534,631,657,680]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
(
<bibRefCitation author="Glaw, F. &amp; C. Kucharzewski &amp; J. Koehler &amp; M. Vences &amp; Z. T. Nagy" journalOrPublisher="Zootaxa" pageId="7" pageNumber="718" pagination="317 - 332" part="3630" refString="Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332." title="Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar" type="journal article" year="2013" yearSuffix="a">Glaw et al., 2013a</bibRefCitation>
)
</taxonomicName>
<emphasis box="[178,185,688,710]" italics="true" pageId="7" pageNumber="718">̗</emphasis>
and our own observation of the 
<taxonomicName box="[582,683,687,709]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[582,683,687,709]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
sample included here, which we found outside a small cave in the karst system of the Montagne des Français massif, approximately 7 0 km away. By contrast, the other species of 
<taxonomicName box="[103,285,805,827]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[103,285,805,827]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
found at Ankarana, 
<taxonomicName box="[554,708,805,828]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[554,708,805,828]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
, was common in canyon forests and surrounding relict forests, as well as in anthropogenically disturbed habitat. We suspect the reason that 
<taxonomicName box="[287,362,894,916]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[287,362,894,916]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
has gone undetected for so long at Ankarana is that the exposed tsingy plateau has been poorly surveyed at night in previous expeditions due to problems of gaining safe access to these areas. 
<taxonomicName box="[491,715,983,1005]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[491,715,983,1005]" italics="true" pageId="7" pageNumber="718">Madagascarophis lolo</emphasis>
</taxonomicName>
may be endemic to the karst habitats of Ankarana, and possibly Analamerana, which is the closest karst system to the east.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="718" type="etymology">
<paragraph blockId="7.[103,773,1095,1266]" pageId="7" pageNumber="718">
<emphasis bold="true" box="[103,248,1095,1118]" italics="true" pageId="7" pageNumber="718">Etymology.—</emphasis>
The species name, 
<taxonomicName box="[489,528,1095,1117]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[489,528,1095,1117]" italics="true" pageId="7" pageNumber="718">lolo</emphasis>
</taxonomicName>
, is taken from the Malagasy word for ghost; it is a noun in apposition to the genus name. This name refers to 1) the ghostly pale gray color of the holotype, and 2) that 
<taxonomicName box="[470,541,1184,1206]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[470,541,1184,1206]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
has eluded discovery for so long at Ankarana, arguably one of the better surveyed sites in Madagascar.
</paragraph>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="719" pageId="7" pageNumber="718" type="discussion">
<paragraph blockId="7.[103,246,1305,1328]" box="[103,246,1305,1328]" pageId="7" pageNumber="718">
<heading allCaps="true" bold="true" box="[103,246,1305,1328]" fontSize="9" level="3" pageId="7" pageNumber="718" reason="2">
<emphasis bold="true" box="[103,246,1305,1328]" pageId="7" pageNumber="718">DISCUSSION</emphasis>
</heading>
</paragraph>
<paragraph blockId="7.[103,774,1351,2024]" pageId="7" pageNumber="718">
High levels of microendemicity may be common for certain regions in Madagascar (
<bibRefCitation author="Brown, J. L. &amp; N. Sillero &amp; F. Glaw &amp; P. Bora &amp; D. R. Vieites &amp; M. Vences" box="[358,559,1380,1403]" journalOrPublisher="PLOS ONE" pageId="7" pageNumber="718" part="11" refString="Brown, J. L., N. Sillero, F. Glaw, P. Bora, D. R. Vieites, and M. Vences. 2016. Spatial biodiversity patterns of Madagascar's amphibians and reptiles. PLOS ONE 11: e 0144076." title="Spatial biodiversity patterns of Madagascar's amphibians and reptiles" type="journal volume" year="2016">Brown et al., 2016</bibRefCitation>
), such as Ankarana, where several reptile genera have endemic representatives (e.g., 
<taxonomicName box="[168,283,1439,1461]" class="Reptilia" family="Lamprophiidae" genus="Alluaudina" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[168,283,1439,1461]" italics="true" pageId="7" pageNumber="718">Alluaudina</emphasis>
</taxonomicName>
, 
<taxonomicName box="[301,398,1439,1461]" class="Reptilia" family="Chamaeleonidae" genus="Brookesia" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[301,398,1439,1461]" italics="true" pageId="7" pageNumber="718">Brookesia</emphasis>
</taxonomicName>
, 
<taxonomicName box="[417,549,1439,1461]" class="Reptilia" family="Gekkonidae" genus="Lygodactylus" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[417,549,1439,1461]" italics="true" pageId="7" pageNumber="718">Lygodactylus</emphasis>
</taxonomicName>
; 
<bibRefCitation author="Glaw, F. &amp; M. Vences" journalOrPublisher="Verlag, Cologne, Germany" pageId="7" pageNumber="718" refString="Glaw, F., and M. Vences. 2007. Amphibians and Reptiles of Madagascar. Third edition. Verlag, Cologne, Germany." title="Amphibians and Reptiles of Madagascar. Third edition" type="book" year="2007">Glaw and Vences, 2007</bibRefCitation>
). The new species of 
<taxonomicName box="[397,575,1469,1491]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[397,575,1469,1491]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
described here is, as far as we currently know, restricted to the tsingy habitat of Ankarana. It seems unlikely that this snake has never been recorded previously, given that this particular area of Ankarana is one of the most accessible areas in the national park and a popular ecotourist destination in Madagascar. However, the trail where we collected this snake was only created in the last ten years and traverses a previously inaccessible area of exposed tsingy plateau which is otherwise difficult to access, especially at night. Because this new snake is found in a national park, and its habitat is naturally well-protected from anthropogenic degradation, we do not consider this species to be vulnerable to extinction. However more survey work is needed to establish the population size and distribution limits of this snake.
</paragraph>
<paragraph blockId="7.[103,774,1351,2024]" lastBlockId="7.[820,1491,131,2024]" pageId="7" pageNumber="718">
Both the morphological and genetic results indicate a close/sister taxa relationship of 
<taxonomicName box="[451,522,1942,1964]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[451,522,1942,1964]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
with 
<taxonomicName box="[587,683,1942,1964]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[587,683,1942,1964]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
(
<figureCitation box="[697,760,1942,1965]" captionStart="Fig. 4" captionStartId="5.[103,131,1066,1086]" captionTargetBox="[103,1492,130,1036]" captionTargetId="figure@5.[103,1492,130,1036]" captionTargetPageId="5" captionText="Fig. 4. Species tree from * BEAST showing the relationships of species of Madagascarophis with posterior probability support for each node indicated; scale bar indicates substitutions per site. Photos from top to bottom from FTB, SR, CJR, SR, SR." httpUri="https://zenodo.org/record/269702/files/figure.png" pageId="7" pageNumber="718">Fig. 4</figureCitation>
). With just a single specimen, more individuals are needed to describe the intraspecific variation of 
<taxonomicName box="[532,607,2001,2024]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[532,607,2001,2024]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
, which is also true for the recently described 
<taxonomicName box="[1173,1272,131,153]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[1173,1272,131,153]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
, known from only four specimens. There is the possibility that these two sister taxa represent populations of the same species; however, the genetic differentiation between 
<taxonomicName box="[1185,1261,218,240]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1185,1261,218,240]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1326,1426,218,240]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[1326,1426,218,240]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
(e.g., 
<emphasis box="[820,862,248,269]" italics="true" pageId="7" pageNumber="718">COI</emphasis>
= 9.6%; 
<tableCitation box="[961,1038,248,271]" captionStart="Table 1" captionStartId="4.[142,190,130,150]" captionTargetBox="[142,1529,222,410]" captionText="Table 1. Genetic similarity of Madagascarophis by maximum percent uncorrected pairwise identity between / within species by gene; locus order is COI / Cmos / Rag 2. Note that as M. fuchsi and M. lolo are represented by one individual, there is no maximum pairwise identity available for within species variation." httpUri="http://table.plazi.org/id/DF07663DFFF4EC4854943BEAC474FF67" pageId="7" pageNumber="718" tableUuid="DF07663DFFF4EC4854943BEAC474FF67">Table 1</tableCitation>
) is higher than or similar to the mtDNA divergence found among many species of snakes, e.g., 
<taxonomicName authority="Zamudio and Greene, 1997" authorityName="Zamudio and Greene" authorityYear="1997" box="[820,1249,306,329]" class="Reptilia" family="Viperidae" genus="Lachesis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[820,906,306,328]" italics="true" pageId="7" pageNumber="718">Lachesis</emphasis>
(
<bibRefCitation author="Zamudio, K. R. &amp; H. W. Greene" box="[925,1240,306,329]" journalOrPublisher="Biological Journal of the Linnean Society" pageId="7" pageNumber="718" pagination="421 - 442" part="62" refString="Zamudio, K. R., and H. W. Greene. 1997. Phylogeography of the bushmaster (Lachesis muta: Viperidae): implications for neotropical biogeography, systematics, and conservation. Biological Journal of the Linnean Society 62: 421 - 442." title="Phylogeography of the bushmaster (Lachesis muta: Viperidae): implications for neotropical biogeography, systematics, and conservation" type="journal article" year="1997">Zamudio and Greene, 1997</bibRefCitation>
)
</taxonomicName>
, 
<taxonomicName authority="Slowinski and Wuster, 2000" authorityName="Slowinski and Wuster" authorityYear="2000" class="Reptilia" family="Elapidae" genus="Naja" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[1264,1314,307,328]" italics="true" pageId="7" pageNumber="718">Naja</emphasis>
(
<bibRefCitation author="Burbrink, F. T. &amp; R. Lawson &amp; J. B. Slowinski" journalOrPublisher="Evolution" pageId="7" pageNumber="718" pagination="2107 - 2118" part="54" refString="Burbrink, F. T., R. Lawson, and J. B. Slowinski. 2000. Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): a critique of the subspecies concept. Evolution 54: 2107 - 2118." title="Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): a critique of the subspecies concept" type="journal article" year="2000">Slowinski and Wüster, 2000</bibRefCitation>
)
</taxonomicName>
, and 
<taxonomicName authority="Burbrink et al., 2000" authorityName="Burbrink et al." authorityYear="2000" box="[1034,1415,335,358]" class="Reptilia" family="Colubridae" genus="Pantherophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[1034,1168,335,357]" italics="true" pageId="7" pageNumber="718">Pantherophis</emphasis>
(
<bibRefCitation author="Burbrink, F. T. &amp; R. Lawson &amp; J. B. Slowinski" box="[1181,1408,335,358]" journalOrPublisher="Evolution" pageId="7" pageNumber="718" pagination="2107 - 2118" part="54" refString="Burbrink, F. T., R. Lawson, and J. B. Slowinski. 2000. Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): a critique of the subspecies concept. Evolution 54: 2107 - 2118." title="Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): a critique of the subspecies concept" type="journal article" year="2000">Burbrink et al., 2000</bibRefCitation>
)
</taxonomicName>
, and is beyond what is found within other species of 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
, including the highly variable 
<taxonomicName box="[1236,1381,394,416]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[1236,1381,394,416]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
(
<tableCitation box="[1399,1483,394,417]" captionStart="Table 1" captionStartId="4.[142,190,130,150]" captionTargetBox="[142,1529,222,410]" captionText="Table 1. Genetic similarity of Madagascarophis by maximum percent uncorrected pairwise identity between / within species by gene; locus order is COI / Cmos / Rag 2. Note that as M. fuchsi and M. lolo are represented by one individual, there is no maximum pairwise identity available for within species variation." httpUri="http://table.plazi.org/id/DF07663DFFF4EC4854943BEAC474FF67" pageId="7" pageNumber="718" tableUuid="DF07663DFFF4EC4854943BEAC474FF67">Table 1</tableCitation>
; 
<bibRefCitation author="Nagy, Z. T. &amp; F. Glaw &amp; F. Andreone &amp; M. Wink &amp; M. Vences" box="[820,1019,423,446]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="7" pageNumber="718" pagination="241 - 251" part="7" refString="Nagy, Z. T., F. Glaw, F. Andreone, M. Wink, and M. Vences. 2007. Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers. Organisms Diversity &amp; Evolution 7: 241 - 251." title="Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers" type="journal article" year="2007">Nagy et al., 2007</bibRefCitation>
). 
<taxonomicName box="[1041,1266,423,445]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1041,1266,423,445]" italics="true" pageId="7" pageNumber="718">Madagascarophis lolo</emphasis>
</taxonomicName>
can also be readily identified from all other species in the genus, including 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
, based on morphology alone (see Key to the species of 
<taxonomicName box="[820,998,511,533]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[820,998,511,533]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
). Our coalescent species delimitation analyses also indicate 
<taxonomicName box="[1007,1080,540,562]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1007,1080,540,562]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
is a distinct species, although this is best supported under small ancestral population sizes and shallow divergences (Pp = 100%). This scenario may be the most realistic for the 
<taxonomicName box="[1060,1238,627,649]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[1060,1238,627,649]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
, where populations of the range-restricted 
<taxonomicName box="[1038,1109,657,679]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1038,1109,657,679]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
as well as 
<taxonomicName box="[1220,1315,657,679]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[1220,1315,657,679]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
are likely small.
</paragraph>
<paragraph blockId="7.[820,1491,131,2024]" pageId="7" pageNumber="718">
The new species described here is unique with respect to coloration. Although species of 
<taxonomicName box="[1192,1370,715,737]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[1192,1370,715,737]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
(with the exception of 
<taxonomicName box="[971,1098,744,767]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[971,1098,744,767]" italics="true" pageId="7" pageNumber="718">M. ocellatus</emphasis>
</taxonomicName>
) have extremely variable intraspecific color patterns, we have not observed any species or individuals with the same coloration seen in the specimen of 
<taxonomicName box="[820,892,832,854]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[820,892,832,854]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
; it possesses a pattern that appears to be well matched to the tsingy rock habitat with varying and alternating shades of gray (Figs. 1, 2). By contrast, no other species of 
<taxonomicName box="[820,998,920,942]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[820,998,920,942]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
has a predominantly gray dorsal ground coloration—they are either brown, blackish brown, orange, or yellowish brown.
</paragraph>
<paragraph blockId="7.[820,1491,131,2024]" pageId="7" pageNumber="718">
Although it is beyond the scope of this study to validate the status of previously described subspecies of 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis italics="true" pageId="7" pageNumber="718">Madagascarophis colubrinus</emphasis>
</taxonomicName>
, we discuss them here to avoid problems with synonymy with respect to 
<taxonomicName box="[1138,1216,1095,1117]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1138,1216,1095,1117]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
. In the most complete examination of the genus, 
<bibRefCitation author="Domergue, C. A." box="[1121,1315,1125,1147]" journalOrPublisher="Bulletin du Museum national d'Histoire Naturelle, Paris" pageId="7" pageNumber="718" pagination="455 - 489" part="9" refString="Domergue, C. A. 1987. Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens, 1952. Bulletin du Museum national d'Histoire Naturelle, Paris 9: 455 - 489." title="Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens" type="journal article" year="1987">Domergue (1987)</bibRefCitation>
recognized five subspecies of 
<taxonomicName box="[972,1114,1154,1176]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[972,1114,1154,1176]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
and an additional full species, 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="citrinus">
<emphasis italics="true" pageId="7" pageNumber="718">M. citrinus</emphasis>
</taxonomicName>
(as well as 
<taxonomicName box="[1043,1210,1183,1205]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[1043,1210,1183,1205]" italics="true" pageId="7" pageNumber="718">M. meridionalis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1276,1405,1183,1205]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[1276,1405,1183,1205]" italics="true" pageId="7" pageNumber="718">M. ocellatus</emphasis>
</taxonomicName>
). More recent work on these snakes (
<bibRefCitation author="Nagy, Z. T. &amp; F. Glaw &amp; F. Andreone &amp; M. Wink &amp; M. Vences" box="[1154,1349,1212,1235]" journalOrPublisher="Organisms Diversity &amp; Evolution" pageId="7" pageNumber="718" pagination="241 - 251" part="7" refString="Nagy, Z. T., F. Glaw, F. Andreone, M. Wink, and M. Vences. 2007. Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers. Organisms Diversity &amp; Evolution 7: 241 - 251." title="Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers" type="journal article" year="2007">Nagy et al., 2007</bibRefCitation>
; 
<bibRefCitation author="Glaw, F. &amp; C. Kucharzewski &amp; J. Koehler &amp; M. Vences &amp; Z. T. Nagy" journalOrPublisher="Zootaxa" pageId="7" pageNumber="718" pagination="317 - 332" part="3630" refString="Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332." title="Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar" type="journal article" year="2013" yearSuffix="a">Glaw et al., 2013a</bibRefCitation>
) proposed: 1) 
<taxonomicName box="[1053,1217,1241,1263]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[1053,1217,1241,1263]" italics="true" pageId="7" pageNumber="718">M. meridionalis</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1277,1403,1241,1263]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[1277,1403,1241,1263]" italics="true" pageId="7" pageNumber="718">M. ocellatus</emphasis>
</taxonomicName>
remain distinct species; 2) 
<emphasis box="[1026,1202,1271,1293]" italics="true" pageId="7" pageNumber="718">M. c. occidentalis</emphasis>
is a junior synonym of 
<emphasis italics="true" pageId="7" pageNumber="718">
M. c. 
<taxonomicName box="[849,955,1300,1322]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">colubrinus</taxonomicName>
</emphasis>
; 3) 
<emphasis box="[1009,1166,1300,1322]" italics="true" pageId="7" pageNumber="718">M. c. insularis</emphasis>
is a junior synonym of 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="citrinus">
<emphasis italics="true" pageId="7" pageNumber="718">M. citrinus</emphasis>
</taxonomicName>
, but 
<taxonomicName box="[964,1080,1330,1351]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="citrinus">
<emphasis box="[964,1080,1330,1351]" italics="true" pageId="7" pageNumber="718">M. citrinus</emphasis>
</taxonomicName>
is a subspecies of 
<taxonomicName box="[1304,1449,1329,1351]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[1304,1449,1329,1351]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
; 4) There is some genetic structure and corresponding morphological variation indicating that 
<emphasis box="[1176,1383,1387,1410]" italics="true" pageId="7" pageNumber="718">M. c. septentrionalis</emphasis>
and 
<emphasis italics="true" pageId="7" pageNumber="718">
M. c. 
<taxonomicName box="[820,896,1418,1439]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="citrinus">citrinus</taxonomicName>
</emphasis>
are distinct from the nominant 
<emphasis box="[1265,1439,1417,1439]" italics="true" pageId="7" pageNumber="718">
M. c. 
<taxonomicName box="[1326,1433,1417,1439]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">colubrinus</taxonomicName>
;
</emphasis>
and 5) 
<taxonomicName box="[848,943,1446,1468]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[848,943,1446,1468]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
is a distinct species that occurs sympatrically with 
<taxonomicName box="[820,962,1475,1497]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[820,962,1475,1497]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
at Montagne des Français.
</paragraph>
<paragraph blockId="7.[820,1491,131,2024]" pageId="7" pageNumber="718">
Importantly, in our genetic analyses and in the aforementioned studies, all of the subspecies of 
<taxonomicName box="[1262,1405,1534,1556]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[1262,1405,1534,1556]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
form a separate and distinct 
<taxonomicName box="[1061,1204,1563,1585]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[1061,1204,1563,1585]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
clade, which is the sister taxon to 
<taxonomicName box="[923,1086,1592,1614]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[923,1086,1592,1614]" italics="true" pageId="7" pageNumber="718">M. meridionalis</emphasis>
</taxonomicName>
. Our study, like 
<bibRefCitation author="Glaw, F. &amp; C. Kucharzewski &amp; J. Koehler &amp; M. Vences &amp; Z. T. Nagy" box="[1272,1486,1592,1615]" journalOrPublisher="Zootaxa" pageId="7" pageNumber="718" pagination="317 - 332" part="3630" refString="Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332." title="Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar" type="journal article" year="2013">Glaw et al. (2013a)</bibRefCitation>
, finds 
<taxonomicName box="[894,1003,1621,1644]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[894,1003,1621,1644]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
falling outside the 
<taxonomicName box="[1260,1415,1621,1644]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[1260,1415,1621,1644]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
+ 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="meridionalis">
<emphasis italics="true" pageId="7" pageNumber="718">M. meridionalis</emphasis>
</taxonomicName>
clade, with the addition that 
<taxonomicName box="[1284,1356,1650,1673]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1284,1356,1650,1673]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
is the sister taxon to 
<taxonomicName box="[919,1014,1680,1702]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[919,1014,1680,1702]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
. Therefore, all taxa within 
<taxonomicName box="[1311,1451,1680,1702]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[1311,1451,1680,1702]" italics="true" pageId="7" pageNumber="718">M. colubrinus</emphasis>
</taxonomicName>
are distinct from 
<taxonomicName box="[980,1057,1709,1731]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[980,1057,1709,1731]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1123,1224,1709,1731]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[1123,1224,1709,1731]" italics="true" pageId="7" pageNumber="718">M. fuchsi</emphasis>
</taxonomicName>
. One subspecies, 
<emphasis italics="true" pageId="7" pageNumber="718">M. c. pastoriensis̗</emphasis>
has not been included in any molecular phylogenetic studies (due to a lack of genetic material); however, this taxon is restricted to the Antananarivo region in central Madagascar and is characterized by having a nearly black body and yellow eyes (
<bibRefCitation author="Domergue, C. A." box="[1150,1339,1856,1878]" journalOrPublisher="Bulletin du Museum national d'Histoire Naturelle, Paris" pageId="7" pageNumber="718" pagination="455 - 489" part="9" refString="Domergue, C. A. 1987. Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens, 1952. Bulletin du Museum national d'Histoire Naturelle, Paris 9: 455 - 489." title="Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens" type="journal article" year="1987">Domergue, 1987</bibRefCitation>
), which does not correspond to 
<taxonomicName box="[1028,1101,1884,1907]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1028,1101,1884,1907]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="7.[820,1491,131,2024]" lastBlockId="8.[142,813,131,1481]" lastPageId="8" lastPageNumber="719" pageId="7" pageNumber="718">
The other 
<taxonomicName box="[963,1141,1914,1936]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="genus">
<emphasis box="[963,1141,1914,1936]" italics="true" pageId="7" pageNumber="718">Madagascarophis</emphasis>
</taxonomicName>
taxon also found at Ankarana, 
<emphasis box="[820,1033,1943,1965]" italics="true" pageId="7" pageNumber="718">M. c. septentrionalis̗</emphasis>
is distinct from 
<taxonomicName box="[1213,1284,1943,1965]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="718" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[1213,1284,1943,1965]" italics="true" pageId="7" pageNumber="718">M. lolo</emphasis>
</taxonomicName>
in that it does not typically have 25 dorsal scale rows (rather 27, or 29 from sites further north) or contact of the posterior inframaxillaries (see 
<bibRefCitation author="Glaw, F. &amp; C. Kucharzewski &amp; J. Koehler &amp; M. Vences &amp; Z. T. Nagy" box="[142,344,131,154]" journalOrPublisher="Zootaxa" pageId="8" pageNumber="719" pagination="317 - 332" part="3630" refString="Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332." title="Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar" type="journal article" year="2013" yearSuffix="a">Glaw et al., 2013a</bibRefCitation>
). We included 
<taxonomicName box="[509,651,131,153]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[509,651,131,153]" italics="true" pageId="8" pageNumber="719">M. colubrinus</emphasis>
</taxonomicName>
(which would correspond to 
<emphasis box="[299,503,160,182]" italics="true" pageId="8" pageNumber="719">M. c. septentrionalis</emphasis>
) from the collecting locality of 
<taxonomicName box="[171,243,189,212]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[171,243,189,212]" italics="true" pageId="8" pageNumber="719">M. lolo</emphasis>
</taxonomicName>
in our genetic analyses and demonstrate that these are not sister taxa.
</paragraph>
<paragraph blockId="8.[142,813,131,1481]" pageId="8" pageNumber="719">
This study is the first to include genetic data for 
<taxonomicName box="[689,812,249,271]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[689,812,249,271]" italics="true" pageId="8" pageNumber="719">M. ocellatus</emphasis>
</taxonomicName>
and all other described species of 
<taxonomicName box="[567,745,278,300]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="genus">
<emphasis box="[567,745,278,300]" italics="true" pageId="8" pageNumber="719">Madagascarophis</emphasis>
</taxonomicName>
in a coalescent-based species tree (
<figureCitation box="[494,565,308,331]" captionStart="Fig. 4" captionStartId="5.[103,131,1066,1086]" captionTargetBox="[103,1492,130,1036]" captionTargetId="figure@5.[103,1492,130,1036]" captionTargetPageId="5" captionText="Fig. 4. Species tree from * BEAST showing the relationships of species of Madagascarophis with posterior probability support for each node indicated; scale bar indicates substitutions per site. Photos from top to bottom from FTB, SR, CJR, SR, SR." httpUri="https://zenodo.org/record/269702/files/figure.png" pageId="8" pageNumber="719">Fig. 4</figureCitation>
). This tree was well supported at almost all nodes (Pp ± 0.99), and importantly, it provides information on the sister taxa relationship of the two most recently described species, 
<taxonomicName box="[566,642,396,418]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[566,642,396,418]" italics="true" pageId="8" pageNumber="719">M. lolo</emphasis>
</taxonomicName>
and 
<taxonomicName box="[705,805,396,418]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[705,805,396,418]" italics="true" pageId="8" pageNumber="719">M. fuchsi</emphasis>
</taxonomicName>
. Each of these species occurs sympatrically with the widespread 
<taxonomicName box="[223,367,455,477]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[223,367,455,477]" italics="true" pageId="8" pageNumber="719">M. colubrinus</emphasis>
</taxonomicName>
(which is the sister taxon to the more southerly distributed 
<taxonomicName box="[383,547,485,507]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[383,547,485,507]" italics="true" pageId="8" pageNumber="719">M. meridionalis</emphasis>
</taxonomicName>
), but our tree indicates that no sister species of 
<taxonomicName box="[407,585,514,536]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="genus">
<emphasis box="[407,585,514,536]" italics="true" pageId="8" pageNumber="719">Madagascarophis</emphasis>
</taxonomicName>
occur sympatrically. These findings are similar to other studies that suggest that recently diverged sister taxa are typically allopatric and that similarity in niche may limit sympatry (e.g., 
<bibRefCitation author="Peterson, A. T." box="[641,807,603,625]" journalOrPublisher="Science" pageId="8" pageNumber="719" pagination="1265 - 1267" part="285" refString="Peterson, A. T. 1999. Conservatism of ecological niches in evolutionary time. Science 285: 1265 - 1267." title="Conservatism of ecological niches in evolutionary time" type="journal article" year="1999">Peterson, 1999</bibRefCitation>
; McCormack et al., 2 0 10; Pigot and Tobias, 2 0 1 3). The possible karst specialization of 
<taxonomicName box="[505,582,662,684]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="lolo">
<emphasis box="[505,582,662,684]" italics="true" pageId="8" pageNumber="719">M. lolo</emphasis>
</taxonomicName>
and 
<taxonomicName box="[650,751,662,684]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="fuchsi">
<emphasis box="[650,751,662,684]" italics="true" pageId="8" pageNumber="719">M. fuchsi</emphasis>
</taxonomicName>
may allow it to occur in the same area with the more distantly related generalist 
<taxonomicName box="[338,480,721,743]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[338,480,721,743]" italics="true" pageId="8" pageNumber="719">M. colubrinus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="8.[142,813,131,1481]" pageId="8" pageNumber="719">
The exception to the generally high support values across the species tree is the placement of 
<taxonomicName box="[546,672,780,802]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[546,672,780,802]" italics="true" pageId="8" pageNumber="719">M. ocellatus</emphasis>
</taxonomicName>
as the sister taxon to 
<taxonomicName box="[248,391,809,832]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[248,391,809,832]" italics="true" pageId="8" pageNumber="719">M. colubrinus</emphasis>
</taxonomicName>
+ 
<taxonomicName box="[424,588,809,832]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[424,588,809,832]" italics="true" pageId="8" pageNumber="719">M. meridionalis</emphasis>
</taxonomicName>
(Pp = 0.62). Unlike the more widespread 
<taxonomicName box="[391,536,839,861]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[391,536,839,861]" italics="true" pageId="8" pageNumber="719">M. colubrinus</emphasis>
</taxonomicName>
and 
<taxonomicName box="[600,766,839,861]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[600,766,839,861]" italics="true" pageId="8" pageNumber="719">M. meridionalis</emphasis>
</taxonomicName>
, 
<taxonomicName class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="ocellatus">
<emphasis italics="true" pageId="8" pageNumber="719">M. ocellatus</emphasis>
</taxonomicName>
is poorly known and is found only in the dry regions of southwestern Madagascar. Our results suggest 
<taxonomicName box="[687,812,898,920]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[687,812,898,920]" italics="true" pageId="8" pageNumber="719">M. ocellatus</emphasis>
</taxonomicName>
is the sister taxon to 
<taxonomicName box="[381,524,927,950]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="colubrinus">
<emphasis box="[381,524,927,950]" italics="true" pageId="8" pageNumber="719">M. colubrinus</emphasis>
</taxonomicName>
+ 
<taxonomicName box="[555,719,927,950]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="meridionalis">
<emphasis box="[555,719,927,950]" italics="true" pageId="8" pageNumber="719">M. meridionalis</emphasis>
</taxonomicName>
(
<figureCitation box="[734,798,927,950]" captionStart="Fig. 4" captionStartId="5.[103,131,1066,1086]" captionTargetBox="[103,1492,130,1036]" captionTargetId="figure@5.[103,1492,130,1036]" captionTargetPageId="5" captionText="Fig. 4. Species tree from * BEAST showing the relationships of species of Madagascarophis with posterior probability support for each node indicated; scale bar indicates substitutions per site. Photos from top to bottom from FTB, SR, CJR, SR, SR." httpUri="https://zenodo.org/record/269702/files/figure.png" pageId="8" pageNumber="719">Fig. 4</figureCitation>
), though we expect future studies using larger genetic datasets may be able to provide more robust support for the phylogenetic placement of 
<taxonomicName box="[447,572,1016,1038]" class="Reptilia" family="Lamprophiidae" genus="Madagascarophis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="ocellatus">
<emphasis box="[447,572,1016,1038]" italics="true" pageId="8" pageNumber="719">M. ocellatus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="8.[142,813,131,1481]" pageId="8" pageNumber="719">
Finally, this study demonstrates that snake species new to science are likely waiting to be discovered in Madagascar, even among commonly encountered taxa, and we expect there are still high numbers of endemic, undescribed squamates in Madagascar. As many of the currently recognized squamate species in Madagascar have very large ranges (e.g., 
<taxonomicName box="[205,418,1222,1244]" class="Reptilia" family="Gekkonidae" genus="Geckolepis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="maculata">
<emphasis box="[205,418,1222,1244]" italics="true" pageId="8" pageNumber="719">Geckolepis maculata</emphasis>
</taxonomicName>
, 
<taxonomicName box="[434,743,1222,1245]" class="Reptilia" family="Gerrhosauridae" genus="Zonosaurus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis box="[434,743,1222,1245]" italics="true" pageId="8" pageNumber="719">Zonosaurus madagascariensis</emphasis>
</taxonomicName>
, 
<taxonomicName class="Reptilia" family="Gekkonidae" genus="Phelsuma" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="719" phylum="Chordata" rank="species" species="lineata">
<emphasis italics="true" pageId="8" pageNumber="719">Phelsuma lineata</emphasis>
</taxonomicName>
; Glaw and Vences, 200 7), phylogeographic studies using modern genomic techniques in an integrative context with morphology and ecology will likely discover additional taxa. Future work in Madagascar and other tropical regions worldwide should focus not only on the discovery of obvious, morphologically differentiated species, but also consider widespread taxa with potential cryptic diversity as well.
</paragraph>
</subSubSection>
</treatment>
</document>