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<document ID-DOI="10.3897/zookeys.19.221" ID-GBIF-Dataset="e6ea8eea-7156-4785-8313-f4bd88dadad1" ID-ISSN="13132970" ID-Zenodo-Dep="576503" ID-ZooBank="C473F9F6-1AE7-4B3F-B17F-CA1C2709010C" checkinTime="1587956031210" checkinUser="plazi" docAuthor="Wesener, Thomas, Enghoff, Henrik &amp; Sierwald, Petra" docDate="2009" docId="1772122E7835FFF5FF013D72AEB8E8A5" docLanguage="en" docName="ZK_article_2037.pdf" docOrigin="ZooKeys 19 (19)" docStyle="DocumentStylede.uka.ipd.idaho.easyIO.settings.Settings@11b73717" docStyleName="zookeys.2008.journal_article" docTitle="Riotintobolus mandenensis Wesener &amp; Enghoff &amp; Sierwald 2009, sp. n." docType="treatment" docUuid="7C0503C7-C70F-4FA4-A9A8-4E9F049E5391" docUuidSource="ZooBank" docVersion="11" lastPageNumber="52" masterDocId="EB4B6A567807FFC6FF8D3F74AF59EA51" masterDocTitle="Review of the Spirobolida on Madagascar, with descriptions of twelve new genera, including three genera of ' fire millipedes' (Diplopoda)" masterLastPageNumber="128" masterPageNumber="1" pageNumber="51" updateTime="1636630017160" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-3.0">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Review of the Spirobolida on Madagascar, with descriptions of twelve new genera, including three genera of ' fire millipedes' (Diplopoda)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wesener, Thomas</mods:namePart>
<mods:nameIdentifier type="LSID">urn:lsid:zoobank.org:author:</mods:nameIdentifier>
<mods:affiliation>Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605, U. S. A. &amp; Zoologisches Forschungsmuseum Alexander Koenig, Museumsmeile Bonn, Adenauerallee 160, D- 53113 Bonn, Germany. &amp; B 54 D 3623309</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Enghoff, Henrik</mods:namePart>
<mods:nameIdentifier type="LSID">urn:lsid:zoobank.org:author:</mods:nameIdentifier>
<mods:affiliation>Natural History Museum of Denmark, Universitetsparken 15, DK- 2100 Copenhagen, Denmark</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Sierwald, Petra</mods:namePart>
<mods:nameIdentifier type="LSID">urn:lsid:zoobank.org:author:</mods:nameIdentifier>
<mods:affiliation>Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605, U. S. A.</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2009</mods:date>
<mods:detail type="pubDate">
<mods:number>2009-09-04</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>19</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>19</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>128</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.3897/zookeys.19.221</mods:identifier>
<mods:identifier type="GBIF-Dataset">e6ea8eea-7156-4785-8313-f4bd88dadad1</mods:identifier>
<mods:identifier type="ISSN">13132970</mods:identifier>
<mods:identifier type="Zenodo-Dep">576503</mods:identifier>
<mods:identifier type="ZooBank">C473F9F6-1AE7-4B3F-B17F-CA1C2709010C</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.3791419" ID-GBIF-Taxon="163747641" ID-Zenodo-Dep="3791419" LSID="urn:lsid:zoobank.org:act:7C0503C7-C70F-4FA4-A9A8-4E9F049E5391" httpUri="http://treatment.plazi.org/id/1772122E7835FFF5FF013D72AEB8E8A5" lastPageId="51" lastPageNumber="52" pageId="50" pageNumber="51">
<subSubSection box="[140,646,518,545]" pageId="50" pageNumber="51" type="nomenclature">
<paragraph blockId="50.[140,963,518,581]" box="[140,646,518,545]" pageId="50" pageNumber="51">
<heading box="[140,646,518,545]" fontSize="11" level="2" pageId="50" pageNumber="51" reason="3">
<emphasis bold="true" box="[140,646,518,545]" pageId="50" pageNumber="51">
<taxonomicName authority="Wesener" authorityName="Wesener &amp; Enghoff &amp; Sierwald" authorityYear="2009" box="[140,569,518,545]" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" kingdom="Animalia" order="Spirobolida" pageId="50" pageNumber="51" phylum="Arthropoda" rank="species" status="sp. n.">
<emphasis bold="true" box="[140,460,518,545]" italics="true" pageId="50" pageNumber="51">Riotintobolus mandenensis</emphasis>
Wesener
</taxonomicName>
,
<taxonomicNameLabel box="[581,646,519,545]" pageId="50" pageNumber="51" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[140,963,554,581]" pageId="50" pageNumber="51" type="description">
<paragraph blockId="50.[140,963,518,581]" box="[140,963,554,581]" pageId="50" pageNumber="51">
<uri box="[140,963,554,581]" pageId="50" pageNumber="51">
urn:lsid:zoobank.org:act:
<uuid box="[420,963,554,581]" pageId="50" pageNumber="51">7C0503C7-C70F-4FA4-A9A8-4E9F049E5391</uuid>
</uri>
</paragraph>
</subSubSection>
<subSubSection pageId="50" pageNumber="51" type="materials_examined">
<paragraph blockId="50.[140,1108,624,1672]" pageId="50" pageNumber="51">
<emphasis bold="true" box="[140,375,624,650]" pageId="50" pageNumber="51">
<emphasis box="[140,241,624,650]" inLineHeading="true" pageId="50" pageNumber="51" reason="1">Material</emphasis>
examined:
</emphasis>
<materialsCitation ID-GBIF-Occurrence="2608818276" collectingDate="2007-06-02" collectionCode="FMMC" collectorName="T. Wesener" elevation="34" latitude="-24.954334" location="Mandena" longLatPrecision="1" longitude="46.99165" municipality="Province" pageId="50" pageNumber="51" specimenCode="W040" specimenCount="36" specimenCount-female="16" stateProvince="Toliara" typeStatus="holotype">
1 ♂,
<specimenCount box="[450,512,624,651]" pageId="50" pageNumber="51" type="female">16 ♀</specimenCount>
,
<specimenCount box="[530,632,624,650]" pageId="50" pageNumber="51" type="generic">20 imm.</specimenCount>
<emphasis box="[643,745,624,650]" italics="true" pageId="50" pageNumber="51">
<typeStatus box="[643,740,624,650]" pageId="50" pageNumber="51">Holotype</typeStatus>
:
</emphasis>
1 ♂ (
<quantity box="[823,911,624,651]" metricMagnitude="-2" metricUnit="m" metricValue="3.6" pageId="50" pageNumber="51" unit="mm" value="36.0">36 mm</quantity>
long, dissected),
<collectionCode box="[140,230,660,686]" pageId="50" pageNumber="51">FMMC</collectionCode>
<specimenCode box="[235,311,660,686]" collectionCode="FMMC" pageId="50" pageNumber="51">W040</specimenCode>
C,
<collectingMunicipality box="[342,440,659,685]" pageId="50" pageNumber="51">Province</collectingMunicipality>
<collectingRegion box="[444,526,659,685]" country="Madagascar" name="Toliara" pageId="50" pageNumber="51">Toliara</collectingRegion>
,
<location LSID="urn:lsid:plazi:treatment:1772122E7835FFF5FF013D72AEB8E8A5:9A04F5E37835FFF4FD953DE7ADDEE8FC" box="[536,647,659,685]" latitude="-24.954334" longLatPrecision="1" longitude="46.99165" municipality="Province" name="Mandena" pageId="50" pageNumber="51" stateProvince="Toliara">Mandena</location>
, littoral forest on sand,
<quantity box="[909,971,660,686]" metricMagnitude="1" metricUnit="m" metricValue="3.4" pageId="50" pageNumber="51" unit="m" value="34.0">
<elevation box="[909,971,660,686]" metricMagnitude="1" metricUnit="m" metricValue="3.4" pageId="50" pageNumber="51" unit="m" value="34.0">34 m</elevation>
</quantity>
,
<geoCoordinate degrees="24" direction="south" minutes="57.260" orientation="latitude" pageId="50" pageNumber="51" precision="1" value="-24.954334">24°57.260 S</geoCoordinate>
,
<geoCoordinate box="[170,325,694,721]" degrees="46" direction="east" minutes="59.499" orientation="longitude" pageId="50" pageNumber="51" precision="1" value="46.99165">46°59.499 E</geoCoordinate>
, leg.
<collectorName box="[385,511,695,721]" pageId="50" pageNumber="51">T. Wesener</collectorName>
et al.,
<date box="[592,728,695,721]" pageId="50" pageNumber="51" value="2007-06-02">
<collectingDate box="[592,728,695,721]" pageId="50" pageNumber="51" value="2007-06-02">02.VI.2007</collectingDate>
</date>
</materialsCitation>
.
<materialsCitation ID-GBIF-Occurrence="2608818272" collectingDate="2007-06-02" collectionCode="FMMC" collectorName="T. Wesener" elevation="34" latitude="-24.954334" location="Mandena" longLatPrecision="1" longitude="46.99165" municipality="Province" pageId="50" pageNumber="51" specimenCode="W039" specimenCount="10" specimenCount-female="5" stateProvince="Toliara" typeStatus="paratype">
<emphasis box="[741,847,695,720]" italics="true" pageId="50" pageNumber="51">
<typeStatus box="[741,843,695,720]" pageId="50" pageNumber="51">Paratypes</typeStatus>
:
</emphasis>
<specimenCount box="[857,901,694,721]" pageId="50" pageNumber="51" type="female">5 ♀</specimenCount>
,
<specimenCount box="[917,1005,694,721]" pageId="50" pageNumber="51" type="generic">5 imm.</specimenCount>
,
<collectionCode box="[1018,1108,695,721]" pageId="50" pageNumber="51">FMMC</collectionCode>
<specimenCode box="[140,216,730,756]" collectionCode="FMMC" pageId="50" pageNumber="51">W039</specimenCode>
C, same data as holotype
</materialsCitation>
;
<materialsCitation ID-GBIF-Occurrence="2608818297" box="[508,1081,730,756]" collectingDate="2007-06-02" collectionCode="CAS" collectorName="T. Wesener" elevation="34" latitude="-24.954334" location="Mandena" longLatPrecision="1" longitude="46.99165" municipality="Province" pageId="50" pageNumber="51" specimenCode="W039" specimenCount="3" specimenCount-female="1" stateProvince="Toliara" typeStatus="paratype">
<specimenCount box="[508,552,730,756]" pageId="50" pageNumber="51" type="female">1 ♀</specimenCount>
,
<specimenCount box="[566,652,730,756]" pageId="50" pageNumber="51" type="generic">2 imm.</specimenCount>
,
<collectionCode box="[664,717,730,756]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15690" lsid="urn:lsid:biocol.org:col:34700" name="California Academy of Sciences" pageId="50" pageNumber="51" type="Museum">CAS</collectionCode>
<specimenCode box="[724,800,730,756]" collectionCode="CAS" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15690" lsid="urn:lsid:biocol.org:col:34700" name="California Academy of Sciences" pageId="50" pageNumber="51" type="Museum">W039</specimenCode>
C, same data as holotype
</materialsCitation>
;
<materialsCitation ID-GBIF-Occurrence="2608818232" collectingDate="2007-06-02" collectionCode="ZMUC" collectorName="T. Wesener" elevation="34" latitude="-24.954334" location="Mandena" longLatPrecision="1" longitude="46.99165" municipality="Province" pageId="50" pageNumber="51" specimenCode="W39" specimenCount="3" specimenCount-female="1" stateProvince="Toliara" typeStatus="paratype">
<specimenCount pageId="50" pageNumber="51" type="female">1 ♀</specimenCount>
,
<specimenCount box="[176,262,765,791]" pageId="50" pageNumber="51" type="generic">2 imm.</specimenCount>
,
<collectionCode box="[274,362,765,791]" country="Denmark" httpUri="http://grbio.org/cool/mci8-ehqk" name="Zoological Museum, University of Copenhagen" pageId="50" pageNumber="51">ZMUC</collectionCode>
<specimenCode box="[370,430,765,791]" collectionCode="ZMUC" country="Denmark" httpUri="http://grbio.org/cool/mci8-ehqk" name="Zoological Museum, University of Copenhagen" pageId="50" pageNumber="51">W39</specimenCode>
C, same data as holotype
</materialsCitation>
;
<materialsCitation ID-GBIF-Occurrence="2608818299" collectingDate="2007-06-02" collectionCode="ZMH" collectorName="T. Wesener" elevation="34" latitude="-24.954334" location="Mandena" longLatPrecision="1" longitude="46.99165" municipality="Province" pageId="50" pageNumber="51" specimenCode="W39" specimenCount="3" specimenCount-female="1" stateProvince="Toliara" typeStatus="paratype">
<specimenCount box="[725,768,765,792]" pageId="50" pageNumber="51" type="female">1 ♀</specimenCount>
,
<specimenCount box="[783,870,765,791]" pageId="50" pageNumber="51" type="generic">2 imm.</specimenCount>
,
<collectionCode box="[882,951,766,792]" country="Germany" httpUri="http://grbio.org/cool/kdnf-0379" name="Zoologisches Museum Hamburg" pageId="50" pageNumber="51" type="Museum">ZMH</collectionCode>
<specimenCode box="[958,1019,765,791]" collectionCode="ZMH" country="Germany" httpUri="http://grbio.org/cool/kdnf-0379" name="Zoologisches Museum Hamburg" pageId="50" pageNumber="51" type="Museum">W39</specimenCode>
C, same data as holotype
</materialsCitation>
;
<materialsCitation ID-GBIF-Occurrence="2608818259" box="[337,917,800,827]" collectingDate="2007-06-02" collectionCode="ZSM" collectorName="T. Wesener" elevation="34" latitude="-24.954334" location="Mandena" longLatPrecision="1" longitude="46.99165" municipality="Province" pageId="50" pageNumber="51" specimenCode="W039" specimenCount="3" specimenCount-female="1" stateProvince="Toliara" typeStatus="paratype">
<specimenCount box="[337,380,800,827]" pageId="50" pageNumber="51" type="female">1 ♀</specimenCount>
,
<specimenCount box="[395,481,800,826]" pageId="50" pageNumber="51" type="generic">2 imm.</specimenCount>
,
<collectionCode box="[493,553,800,826]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="50" pageNumber="51" type="Museum">ZSM</collectionCode>
<specimenCode box="[560,635,800,826]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="50" pageNumber="51" type="Museum">W039</specimenCode>
C, same data as holotype
</materialsCitation>
.
</paragraph>
<paragraph blockId="50.[140,1108,624,1672]" pageId="50" pageNumber="51">
<emphasis bold="true" box="[185,492,835,861]" pageId="50" pageNumber="51">
<emphasis box="[185,257,835,861]" inLineHeading="true" pageId="50" pageNumber="51" reason="1">Other</emphasis>
material examined:
</emphasis>
<specimenCount box="[499,541,836,862]" pageId="50" pageNumber="51" type="female">7 ♀</specimenCount>
,
<specimenCount box="[556,641,835,862]" pageId="50" pageNumber="51" type="generic">7 imm.</specimenCount>
,
<emphasis bold="true" box="[653,747,835,861]" pageId="50" pageNumber="51">W039C</emphasis>
, same data as
<typeStatus box="[907,1009,835,861]" pageId="50" pageNumber="51">holotype</typeStatus>
, University
<collectingRegion box="[186,343,871,897]" country="Madagascar" name="Antananarivo" pageId="50" pageNumber="51">Antananarivo</collectingRegion>
.
</paragraph>
</subSubSection>
<subSubSection pageId="50" pageNumber="51" type="diagnosis">
<paragraph blockId="50.[140,1108,624,1672]" pageId="50" pageNumber="51">
<emphasis bold="true" box="[185,449,905,931]" pageId="50" pageNumber="51">
<emphasis box="[185,323,905,931]" inLineHeading="true" pageId="50" pageNumber="51" reason="1">Differential</emphasis>
diagnosis:
</emphasis>
the size and special colour pattern are unique for Malagasy millipedes. The gonopods of this species are similar in some aspects to those of
<taxonomicName authority="Wesener &amp; Enghoff &amp; Sierwald, 2009" authorityName="Wesener &amp; Enghoff &amp; Sierwald" authorityYear="2009" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" kingdom="Animalia" order="Spirobolida" pageId="50" pageNumber="51" phylum="Arthropoda" rank="species" status="sp. n.">
<emphasis italics="true" pageId="50" pageNumber="51">R. minutus</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[235,294,977,1003]" pageId="50" pageNumber="51" rank="species">sp. n.</taxonomicNameLabel>
and
<taxonomicName authority="Wesener &amp; Enghoff &amp; Sierwald, 2009" authorityName="Wesener &amp; Enghoff &amp; Sierwald" authorityYear="2009" box="[348,481,976,1002]" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" kingdom="Animalia" order="Spirobolida" pageId="50" pageNumber="51" phylum="Arthropoda" rank="species" status="sp. n.">
<emphasis box="[348,481,976,1002]" italics="true" pageId="50" pageNumber="51">R. anomalus</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[487,549,977,1003]" pageId="50" pageNumber="51" rank="species">sp. n.</taxonomicNameLabel>
, but the posterior gonopods are of a unique shape.
</paragraph>
</subSubSection>
<subSubSection lastPageId="51" lastPageNumber="52" pageId="50" pageNumber="51" type="description">
<paragraph blockId="50.[140,1108,624,1672]" pageId="50" pageNumber="51">
<emphasis bold="true" box="[185,335,1011,1037]" pageId="50" pageNumber="51">
<emphasis box="[185,330,1011,1037]" inLineHeading="true" pageId="50" pageNumber="51" reason="1">Description</emphasis>
.
</emphasis>
<emphasis box="[346,498,1012,1037]" italics="true" pageId="50" pageNumber="51">Measurements</emphasis>
:
<typeStatus box="[516,615,1011,1037]" pageId="50" pageNumber="51">holotype</typeStatus>
with 41 podous and 2 apodous rings,
<quantity metricMagnitude="-2" metricUnit="m" metricValue="3.6" pageId="50" pageNumber="51" unit="mm" value="36.0">36 mm</quantity>
long,
<quantity box="[262,355,1047,1073]" metricMagnitude="-3" metricUnit="m" metricValue="3.1" pageId="50" pageNumber="51" unit="mm" value="3.1">3.1 mm</quantity>
wide. Females with 4143 podous and 02 apodous body rings,
<quantity box="[140,267,1082,1108]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" metricValueMax="4.3" metricValueMin="3.7" pageId="50" pageNumber="51" unit="mm" value="40.0" valueMax="43.0" valueMin="37.0">3743 mm</quantity>
long,
<quantity box="[339,480,1082,1108]" metricMagnitude="-3" metricUnit="m" metricValue="4.3" metricValueMax="4.4" metricValueMin="4.2" pageId="50" pageNumber="51" unit="mm" value="4.3" valueMax="4.4" valueMin="4.2">4.24.4 mm</quantity>
wide.
</paragraph>
<paragraph blockId="50.[140,1108,624,1672]" pageId="50" pageNumber="51">
<emphasis box="[185,305,1117,1143]" italics="true" pageId="50" pageNumber="51">Coloration</emphasis>
black, dorsally with an extraordinary wide, light brown stripe (
<figureCitation captionStart="Figure 24" captionStartId="44.[140,212,1590,1612]" captionTargetBox="[140,1108,447,1570]" captionTargetId="figure@44.[140,1108,446,1570]" captionTargetPageId="44" captionText="Figure 24. A Riotintobolus mandenensis sp. n., living specimens B pseudosteppe and burned shrubs dominating the area between the littoral forests of Mandena and Sainte Luce, SE Madagascar C Sainte Luce littoral forest, manioc plantation and destroyed trees inside conservation zone S8." figureDoi="http://doi.org/10.5281/zenodo.3767302" httpUri="https://zenodo.org/record/3767302/files/figure.png" pageId="50" pageNumber="51">Fig. 24A</figureCitation>
). Posterior half of collum dorsally black (
<figureCitation box="[686,795,1152,1179]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26A</figureCitation>
). Head, legs and antennae light brown.
<emphasis box="[291,336,1188,1213]" italics="true" pageId="50" pageNumber="51">Eyes</emphasis>
with 1620 almost fused ocelli arranged in 4 or 5 rows (
<figureCitation captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Figs 26A, C</figureCitation>
).
<emphasis box="[187,293,1223,1249]" italics="true" pageId="50" pageNumber="51">Antennae</emphasis>
protruding back to body ring 2 (
<figureCitation box="[690,799,1223,1250]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26A</figureCitation>
).
<emphasis box="[820,867,1223,1248]" italics="true" pageId="50" pageNumber="51">Legs</emphasis>
without tarsal pads. Male legs reach 0.7 times, female legs 0.6 times body diameter.
<emphasis box="[936,1107,1258,1284]" italics="true" pageId="50" pageNumber="51">Preanal process</emphasis>
sharp-edged, straight, protruding above anal valves (
<figureCitation box="[765,873,1293,1320]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26B</figureCitation>
). Anal valves small, between anterior part and lips with a deep groove, margin towards groove sharpedged (
<figureCitation box="[224,331,1363,1390]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26B</figureCitation>
).
</paragraph>
<paragraph blockId="50.[140,1108,624,1672]" pageId="50" pageNumber="51">
<emphasis box="[185,372,1399,1425]" italics="true" pageId="50" pageNumber="51">Anterior gonopod</emphasis>
sternite basally wide, apically elevated into a lobe with a triangular, well-rounded tip (
<figureCitation box="[389,484,1434,1461]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26I</figureCitation>
). Sternite tip almost as high as mesal process of coxite; latter relatively slender, elongated, longer than sternite (
<figureCitation box="[762,856,1469,1496]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26I</figureCitation>
). Telopodite on posterior side apically with a large triangular, retrorse process (
<figureCitation box="[780,885,1504,1531]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26K</figureCitation>
). Process projecting above coxite and telopodite (
<figureCitation box="[465,570,1539,1566]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26K</figureCitation>
).
</paragraph>
<paragraph blockId="50.[140,1108,624,1672]" lastBlockId="51.[140,1108,167,756]" lastPageId="51" lastPageNumber="52" pageId="50" pageNumber="51">
<emphasis box="[185,386,1575,1601]" italics="true" pageId="50" pageNumber="51">Posterior gonopods</emphasis>
telopodite laterally with a single, large, finger-shaped process (
<figureCitation box="[148,276,1610,1636]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26J, x</figureCitation>
). Second small projection, where sperm canal is discharging, is located close to apex (
<figureCitation box="[300,438,1645,1672]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="50" pageNumber="51">Fig. 26M, y</figureCitation>
). A large, bi-lobical membrane present apically. Membrane apically three times wider than basally (
<figureCitation box="[614,749,167,193]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="51" pageNumber="52">Fig. 26L, z</figureCitation>
). A small, white, swollen area (
<figureCitation box="[148,256,202,229]" captionStart="Figure 26" captionStartId="46.[140,212,1398,1420]" captionTargetBox="[184,1064,165,1374]" captionTargetId="figure@46.[183,1065,165,1100]" captionTargetPageId="46" captionText="Figure 26. Riotintobolus mandenensis sp. n., A, B, IM male holotype, CH female paratype. A head, lateral view B telson C head, detail of incisura lateralis D head, frontal view E second leg with vulva F gnathochilarium, overview G gnathochilarium, detail of mentum H left mandible, mesal view I anterior gonopod, anterior view Į left posterior gonopod, anterior view K anterior gonopod, posterior view L telopodite of left posterior gonopod, posterior view M left posterior gonopod, overview, posterior view. 3iT = 3-combed internal tooth; av = anal valves; Co = collum; Cx = coxite; eT = external tooth; Gn = gnathochilarium; Gr = groove; IL = incisura lateralis; L = ledge; LL = lamella linguales; LT = lateral tooth; Md = mandible; Me = mentum; mp = molar plate; pL = pectinate lamella; Pre = preanal ring; sf = setiferous foveolae; St = sternite; ST = stipites; sub = subanal scale; T = telopodite; Vu = vulva. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3767304" httpUri="https://zenodo.org/record/3767304/files/figure.png" pageId="51" pageNumber="52">Fig. 26M</figureCitation>
) is also located basally of membrane.
</paragraph>
<paragraph blockId="51.[140,1108,167,756]" pageId="51" pageNumber="52">
<emphasis bold="true" box="[185,456,237,263]" pageId="51" pageNumber="52">
<emphasis box="[185,333,237,263]" inLineHeading="true" pageId="51" pageNumber="52" reason="1">Intraspecific</emphasis>
variation:
</emphasis>
females exist in different size classes. The presence of apodous rings in front of the telson in mature specimens hint to the existence of postmaturity moults.
</paragraph>
</subSubSection>
<subSubSection pageId="51" pageNumber="52" type="distribution">
<paragraph blockId="51.[140,1108,167,756]" pageId="51" pageNumber="52">
<emphasis bold="true" box="[185,486,342,369]" pageId="51" pageNumber="52">
<emphasis box="[185,336,342,368]" inLineHeading="true" pageId="51" pageNumber="52" reason="1">Distribution</emphasis>
and ecology
</emphasis>
: this species was only collected in the littoral rainforest on sand of Mandena (
<figureCitation box="[386,469,378,405]" captionStart="Figure 25" captionStartId="45.[140,212,1654,1676]" captionTargetBox="[162,1133,1244,1638]" captionTargetId="figure@45.[162,1130,1244,1634]" captionTargetPageId="45" captionText="Figure 25. Distribution of the genus Riotintobolus gen. n." figureDoi="http://doi.org/10.5281/zenodo.3820816" httpUri="https://zenodo.org/record/3820816/files/figure.png" pageId="51" pageNumber="52">Fig. 25</figureCitation>
). Only 160 ha of this forest will remain after a large-scale mining project (
<bibRefCitation author="Vincelette" box="[326,581,413,440]" etAl="et al." firstAuthor="Vincelette" pageId="51" pageNumber="52" pagination="1535 - 1537" refId="ref57793" refString="Vincelette M, Randrihasipara L, Ramanamanjato J-B, Lowry (II) PP, Ganzhorn JU (2003) Mining and environmental conservation: The case of QIT Madagascar Minerals in the Southeast. In: Goodman SM, Benstead JP (Eds) The Natural History of Madagascar. University of Chicago Press, Chicago, pp. 1535 - 1537." type="book chapter" year="2003">Vincelette et al. 2003</bibRefCitation>
). All specimens were collected directly on the root layer under a thin layer of leaf litter. Disturbed specimens often did not curl into a spiral but remained motionless and stiff like a stick even when picked up. Because of their black/light-brown colour pattern (
<figureCitation box="[595,700,519,545]" captionStart="Figure 24" captionStartId="44.[140,212,1590,1612]" captionTargetBox="[140,1108,447,1570]" captionTargetId="figure@44.[140,1108,446,1570]" captionTargetPageId="44" captionText="Figure 24. A Riotintobolus mandenensis sp. n., living specimens B pseudosteppe and burned shrubs dominating the area between the littoral forests of Mandena and Sainte Luce, SE Madagascar C Sainte Luce littoral forest, manioc plantation and destroyed trees inside conservation zone S8." figureDoi="http://doi.org/10.5281/zenodo.3767302" httpUri="https://zenodo.org/record/3767302/files/figure.png" pageId="51" pageNumber="52">Fig. 24A</figureCitation>
), specimens of
<taxonomicName authorityName="Wesener &amp; Enghoff &amp; Sierwald" authorityYear="2009" box="[877,1046,519,545]" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" kingdom="Animalia" order="Spirobolida" pageId="51" pageNumber="52" phylum="Arthropoda" rank="species">
<emphasis box="[877,1046,519,545]" italics="true" pageId="51" pageNumber="52">R. mandenensis</emphasis>
</taxonomicName>
were quite difficult to detect against the background consisting of sand with some darker top soil inclusion. The giant pill-millipede species
<taxonomicName authority="Wesener &amp; Sierwald, 2005" authorityName="Wesener &amp; Sierwald" authorityYear="2005" class="Diplopoda" family="Sphaerotheriidae" genus="Sphaeromimus" kingdom="Animalia" order="Sphaerotheriida" pageId="51" pageNumber="52" phylum="Arthropoda" rank="species">
<emphasis box="[708,1005,589,615]" italics="true" pageId="51" pageNumber="52">Sphaeromimus inexpectatus</emphasis>
<bibRefCitation author="Wesener &amp; Sierwald" firstAuthor="Wesener" pageId="51" pageNumber="52" pagination="557 - 599" refId="ref57920" refString="Wesener T, Sierwald P (2005) The giant pill-millipedes of Madagascar: Revision of the genus Sphaeromimus, with a review of the morphological terminology (Diplopoda, Sphaerotheriida, Sphaerotheriidae). Proceedings of the California Academy of Sciences 56 (29): 557 - 599. [http: // research. calacademy. org / research / scipubs / pdfs / v 56 / proccas _ v 56 _ n 29. pdf]" type="journal article" year="2005">Wesener &amp; Sierwald, 2005</bibRefCitation>
</taxonomicName>
is also only known from a small area surrounding Mandena (Wesen- er and Sierwald 2005).
</paragraph>
</subSubSection>
<subSubSection pageId="51" pageNumber="52" type="etymology">
<paragraph blockId="51.[140,1108,167,756]" pageId="51" pageNumber="52">
<emphasis bold="true" box="[185,323,695,721]" pageId="51" pageNumber="52">
<emphasis box="[185,318,695,721]" inLineHeading="true" pageId="51" pageNumber="52" reason="1">Etymology</emphasis>
:
</emphasis>
<taxonomicName authorityName="Wesener &amp; Enghoff &amp; Sierwald" authorityYear="2009" box="[330,466,695,721]" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" kingdom="Animalia" order="Spirobolida" pageId="51" pageNumber="52" phylum="Arthropoda" rank="species">
<emphasis box="[330,466,695,721]" italics="true" pageId="51" pageNumber="52">mandenensis</emphasis>
</taxonomicName>
, adjective, refers to the only area where this species lives, the littoral forest of Mandena.
</paragraph>
</subSubSection>
</treatment>
</document>