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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.953.53977" ID-GBIF-Dataset="77c65687-4a7f-4b63-911d-1d096aaaad76" ID-PMC="PMC7398950" ID-Pensoft-Pub="1313-2970-953-1" ID-Pensoft-UUID="C7001D577310538DBE2A4736802E1BF4" ID-PubMed="32821193" ID-ZooBank="BA81E87988A2495A92FB98D1F2909BA7" ModsDocID="1313-2970-953-1" checkinTime="1595913774766" checkinUser="pensoft" docAuthor="Wesener, Thomas" docDate="2020" docId="B286BFDB5D4F5709B5DEF0A154F890A2" docLanguage="en" docName="ZooKeys 953: 1-29" docOrigin="ZooKeys 953" docSource="http://dx.doi.org/10.3897/zookeys.953.53977" docTitle="Riotintobolus tsimelahy Wesener 2020, sp. nov." docType="treatment" docUuid="672376C7-CC91-4259-B896-B45229104743" docUuidSource="ZooBank" docVersion="4" id="C7001D577310538DBE2A4736802E1BF4" lastPageNumber="1" masterDocId="C7001D577310538DBE2A4736802E1BF4" masterDocTitle="Ecotone shifts in southern Madagascar: first barcoding data and six new species of the endemic millipede genus Riotintobolus (Spirobolida, Pachybolidae)" masterLastPageNumber="29" masterPageNumber="1" pageNumber="1" updateTime="1668169162946" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Ecotone shifts in southern Madagascar: first barcoding data and six new species of the endemic millipede genus Riotintobolus (Spirobolida, Pachybolidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wesener, Thomas</mods:namePart>
<mods:affiliation>Zoological Research Museum Alexander Koenig (ZFMK), Leibniz Institute for Animal Biodiversity, Adenauerallee 160, D- 53113, Bonn, Germany</mods:affiliation>
<mods:nameIdentifier type="email">t.wesener@leibniz-zfmk.de</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="volume">
<mods:number>953</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>29</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.953.53977</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.953.53977</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-953-1</mods:identifier>
<mods:identifier type="ZooBank">BA81E87988A2495A92FB98D1F2909BA7</mods:identifier>
<mods:identifier type="Pensoft-UUID">C7001D577310538DBE2A4736802E1BF4</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="166016731" LSID="urn:lsid:zoobank.org:act:672376C7-CC91-4259-B896-B45229104743" httpUri="http://treatment.plazi.org/id/B286BFDB5D4F5709B5DEF0A154F890A2" lastPageNumber="1" pageId="0" pageNumber="1">
<subSubSection pageId="0" pageNumber="1" type="nomenclature">
<paragraph pageId="0" pageNumber="1">
<taxonomicName LSID="B286BFDB-5D4F-5709-B5DE-F0A154F890A2" authority="Wesener, 2020" authorityName="Wesener" authorityYear="2020" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus tsimelahy" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="tsimelahy" status="sp. nov.">Riotintobolus tsimelahy</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="1">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">Figure 3</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="materials_examined">
<paragraph pageId="0" pageNumber="1">Material examined.</paragraph>
<paragraph pageId="0" pageNumber="1">
1 ♂
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">holotype</emphasis>
,
<emphasis bold="true" pageId="0" pageNumber="1">ZFMK MYR9940</emphasis>
, Madagascar, PN Andohahela, Tsimelahy,
<geoCoordinate degrees="24" direction="south" minutes="57.296" orientation="latitude" precision="1" value="-24.954933">24°57.296'S</geoCoordinate>
,
<geoCoordinate degrees="046" direction="east" minutes="37.214" orientation="longitude" precision="1" value="46.62023">046°37.214'E</geoCoordinate>
, 135 m, spiny forest, coll. Wesener and
<normalizedToken originalValue="Schütte">Schuette</normalizedToken>
, 24.v.2007.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Paratypes</emphasis>
: 9 ♂, 12 ♀,
<emphasis bold="true" pageId="0" pageNumber="1">ZFMK MYR941</emphasis>
, same data as holotype; 1 ♂,
<emphasis bold="true" pageId="0" pageNumber="1">ZFMK MYR9950</emphasis>
, same data as holotype; 1 ♂ with damaged gonopods;
<emphasis bold="true" pageId="0" pageNumber="1">ZFMK MYR9949</emphasis>
, same data as holotype.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="materials_examined">
<paragraph pageId="0" pageNumber="1">Other material examined.</paragraph>
<paragraph pageId="0" pageNumber="1">
3 ♂ and ♀,
<emphasis bold="true" pageId="0" pageNumber="1">BLF 5239 (CASENT9032808)</emphasis>
, Madagascar, Toliara,
<normalizedToken originalValue="Forêt">Foret</normalizedToken>
de Mahavelo, Isantoria River,
<geoCoordinate degrees="24" direction="south" minutes="45" orientation="latitude" precision="15" seconds="30" value="-24.758333">24°45'30&quot;S</geoCoordinate>
,
<geoCoordinate degrees="46" direction="east" minutes="9" orientation="longitude" precision="15" seconds="26" value="46.157223">46°9'26&quot;E</geoCoordinate>
, 110 m, 28.i-1.ii.2002, spiny forest/thicket, EH18 pitfall trap, coll. B. L. Fisher et al.; 1 ♂, 1 imm.,
<emphasis bold="true" pageId="0" pageNumber="1">MZUF Fi-10</emphasis>
, Madagascar; Andohahela, 6-12.xii.1991, leg B. Randriamampionona.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="etymology">
<paragraph pageId="0" pageNumber="1">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="1">
Tsimelahy, after the type locality (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Distribution of the newly described Riotintobolus species, as well as of the still undescribed species. Vegetation marks modified from Moat and Smith 2007." figureDoi="10.3897/zookeys.953.53977.figure2" httpUri="https://binary.pensoft.net/fig/436163" pageId="0" pageNumber="1">2</figureCitation>
), spiny forests next to the Tsimelahy River, Andohahela National Parc. Noun in apposition.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="diagnosis">
<paragraph pageId="0" pageNumber="1">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Wesener" authorityYear="2020" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus tsimelahy" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="tsimelahy">
<emphasis italics="true" pageId="0" pageNumber="1">Riotintobolus tsimelahy</emphasis>
</taxonomicName>
sp. nov. shares the absence of a projecting epiproct on the telson only with
<taxonomicName lsidName="R. anomalus" pageId="0" pageNumber="1" rank="species" species="anomalus">
<emphasis italics="true" pageId="0" pageNumber="1">R. anomalus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="R. antafoky" pageId="0" pageNumber="1" rank="species" species="antafoky">
<emphasis italics="true" pageId="0" pageNumber="1">R. antafoky</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="R. bovinus" pageId="0" pageNumber="1" rank="species" species="bovinus">
<emphasis italics="true" pageId="0" pageNumber="1">R. bovinus</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="R. mangatsiaka" pageId="0" pageNumber="1" rank="species" species="mangatsiaka">
<emphasis italics="true" pageId="0" pageNumber="1">R. mangatsiaka</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName lsidName="R. lavanono" pageId="0" pageNumber="1" rank="species" species="lavanono">
<emphasis italics="true" pageId="0" pageNumber="1">R. lavanono</emphasis>
</taxonomicName>
sp. nov. The posterior telopod featuring two slender, sharp projections is only shared with
<taxonomicName lsidName="R. bovinus" pageId="0" pageNumber="1" rank="species" species="bovinus">
<emphasis italics="true" pageId="0" pageNumber="1">R. bovinus</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="R. mangatsiaka" pageId="0" pageNumber="1" rank="species" species="mangatsiaka">
<emphasis italics="true" pageId="0" pageNumber="1">R. mangatsiaka</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName lsidName="R. lavanono" pageId="0" pageNumber="1" rank="species" species="lavanono">
<emphasis italics="true" pageId="0" pageNumber="1">R. lavanono</emphasis>
</taxonomicName>
sp. nov. A posterior gonopod separated into three parts is only shared with
<taxonomicName lsidName="R. mangatsiaka" pageId="0" pageNumber="1" rank="species" species="mangatsiaka">
<emphasis italics="true" pageId="0" pageNumber="1">R. mangatsiaka</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName lsidName="R. lavanono" pageId="0" pageNumber="1" rank="species" species="lavanono">
<emphasis italics="true" pageId="0" pageNumber="1">R. lavanono</emphasis>
</taxonomicName>
sp. nov., whose habitus and gonopods look very similar to those of
<taxonomicName lsidName="R. tsimelahy" pageId="0" pageNumber="1" rank="species" species="tsimelahy">
<emphasis italics="true" pageId="0" pageNumber="1">R. tsimelahy</emphasis>
</taxonomicName>
sp. nov. Both species differ in details of the tip of the posterior gonopod and in the colour of their antennae and legs, which are red in
<taxonomicName lsidName="R. tsimelahy" pageId="0" pageNumber="1" rank="species" species="tsimelahy">
<emphasis italics="true" pageId="0" pageNumber="1">R. tsimelahy</emphasis>
</taxonomicName>
sp. nov. and dark grey in
<taxonomicName lsidName="R. lavanono" pageId="0" pageNumber="1" rank="species" species="lavanono">
<emphasis italics="true" pageId="0" pageNumber="1">R. lavanono</emphasis>
</taxonomicName>
sp. nov.
<taxonomicName lsidName="R. tsimelahy" pageId="0" pageNumber="1" rank="species" species="tsimelahy">
<emphasis italics="true" pageId="0" pageNumber="1">R. tsimelahy</emphasis>
</taxonomicName>
sp. nov. differs from
<taxonomicName lsidName="R. mangatsiaka" pageId="0" pageNumber="1" rank="species" species="mangatsiaka">
<emphasis italics="true" pageId="0" pageNumber="1">R. mangatsiaka</emphasis>
</taxonomicName>
sp. nov. in the presence of two lateral processes on the posterior gonopod. All three species differ by&gt;11% uncorrected p-distance in the COI barcoding gene.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="description">
<paragraph pageId="0" pageNumber="1">Description.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Measurements</emphasis>
: Telson not included in counts of segments. Male holotype with 50+0 segments, ca. 44 mm long, 4.3 mm wide. Largest females of type series with 49 or 50+0 segments, up to 50 mm long, 5.3 mm wide.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Colour</emphasis>
(in living specimens): Body rings grey, appendages red. Head, paraprocts and posterior margins of body segments darker grey to black. Ozopore openings highlighted by black spot (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3A, C</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Head</emphasis>
: each eye with 30-35 ommatidia in six rows. Incisura lateralis open (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3A</figureCitation>
). Labrum with standard three irregular teeth and a single row of 10-12 stout marginal setae. Clypeus with two setiferous foveolae on each side (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3B</figureCitation>
). Antennae long, protruding back to segment 5. Length of antennomeres: 1&lt;2&gt;3=4=5=6. Second antennomere slenderer but twice as long as first. Terminal antennomere with four large sensory cones located together inside a membranous area.
</paragraph>
<caption doi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1" start="Figure 3" startId="F3">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 3.</emphasis>
<taxonomicName authorityName="Wesener" authorityYear="2020" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus tsimelahy" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="tsimelahy">
<emphasis italics="true" pageId="0" pageNumber="1">Riotintobolus tsimelahy</emphasis>
</taxonomicName>
sp. nov., holotype (ZFMK MYR99940) (
<emphasis bold="true" pageId="0" pageNumber="1">A-F</emphasis>
) and male paratype (ZFMK MYR9949) (
<emphasis bold="true" pageId="0" pageNumber="1">G, H</emphasis>
)
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
multi-layer photograph, male anterior body, lateral view
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
multi-layer photograph, male leg pairs 1-7, ventral view
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
multi-layer photograph, male posterior body end with telson
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
anterior gonopod, anterior view
<emphasis bold="true" pageId="0" pageNumber="1">E</emphasis>
left posterior gonopod, posterior view
<emphasis bold="true" pageId="0" pageNumber="1">F</emphasis>
right posterior gonopod, anterior view
<emphasis bold="true" pageId="0" pageNumber="1">G</emphasis>
damaged? left posterior gonopod, posterior view
<emphasis bold="true" pageId="0" pageNumber="1">H</emphasis>
damaged? right posterior gonopod, anterior view.
<emphasis bold="true" pageId="0" pageNumber="1">Abbreviations</emphasis>
: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous
<normalizedToken originalValue="flag">'flag'</normalizedToken>
; numbers refer to leg pair number. Scale bars: 1 mm.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Gnathochilarium</emphasis>
: lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Palpi of similar size. Endochilarium not dissected.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Mandible</emphasis>
not dissected.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Collum</emphasis>
: smooth, laterally not protruding as far as ring 2 (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3A</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Body rings</emphasis>
: ozopores starting at segment 6, marked by a black spot. Located on suture between meso- and metazonite. Rings with smooth, but irregular coriaceous surface, ventrally on metazona with transverse ridges.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Telson</emphasis>
: paraprocts elongated, with weak lips, abundant micropunctation especially towards edges (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3C</figureCitation>
). Epiproct well-rounded, covering, but not reaching above paraproct (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3C</figureCitation>
). Hypoproct inconspicuous (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3C</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Legs</emphasis>
: leg 1 with a large cylindrical coxa, twice as long as other podomeres. Tarsus with three pairs of ventral spines and an apical spine beyond claw. Leg 2 with an elongated coxa. Tarsus with three pairs of ventral spines and a short apical spine. Midbody legs with a rectangular coxa, as long as other podomeres. Each podomere ventrally with a single or a pair of apical setae, tarsus with a single apical and three pairs of ventral spines. Length of midbody legs ca. 1.2 times body diameter in males.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">
<emphasis italics="true" pageId="0" pageNumber="1">Female sexual characters</emphasis>
.
</emphasis>
No tarsal pads, antennae shorter than male, only protruding back to ring 2. Vulvae not dissected.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Male sexual characters</emphasis>
: tarsal pads present from leg 3 to midbody, small, inconspicuous (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3B</figureCitation>
). Coxae 3-7 without coxal processes, but coxae 3-5 swollen (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3B</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Anterior gonopod</emphasis>
sternite massive (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3D</figureCitation>
), elongated into a wide, well-rounded triangular lobe (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3D</figureCitation>
). Sternite in anterior view well-visible, without discernible apodemes, protruding almost higher than coxal processes. Coxite with a large, well-rounded mesal process (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3D</figureCitation>
). Telopodite with process arising mesally (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3D</figureCitation>
), process apically curved with a large triangular projection (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3D</figureCitation>
), tip well-rounded, slightly protruding above lateral margin of telopodite (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3D</figureCitation>
). Whole telopodite process resembling an even-sided triangle.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Posterior gonopods</emphasis>
consisting of three parts, separated by sutures or articulations: a basal coxite with a slender coxite projection and a slightly shorter telopodite, efferent duct discharging laterally (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3E, F</figureCitation>
). Process of coxite and telopodite standing in same axis (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3E, F</figureCitation>
). Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite. Basal part of coxite wide, mesally with a large triangular sclerite located on lower level than remaining part. Coxite elongated. Efferent duct running at mesal margin of coxite (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3E</figureCitation>
) before curving to the lateral margin at beginning of telopodite (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3E</figureCitation>
). Telopodite as wide as but much shorter than coxite, standing in same axis (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3F</figureCitation>
), apically membranous, with one triangular apical process and two slender lateral processes (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3E, F</figureCitation>
). Lateral processes straight, running almost parallel to one another, slender and sclerotised, efferent duct seems to be ending at base of lateral process (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3E</figureCitation>
). Base of lateral process with a short, membranous-white projection (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR 99940) (A-F) and male paratype (ZFMK MYR 9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1 - 7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ' flag'; numbers refer to leg pair number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure3" httpUri="https://binary.pensoft.net/fig/436164" pageId="0" pageNumber="1">3E</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="intraspecific variation">
<paragraph pageId="0" pageNumber="1">Intraspecific variation.</paragraph>
<paragraph pageId="0" pageNumber="1">
The number of segments varies, even within one population, between 47 and 51. The population from the
<normalizedToken originalValue="Forêt">Foret</normalizedToken>
de Mahavelo (Fig. 1) differs by 1.5% uncorrected p-distance of the COI gene to those from the type locality, Tsimelahy (Suppl. material 1). One small male from the type series (ZFMK MYR9949) has incompletely developed posterior gonopods, either a sign of healed damage (there are black spots on it), or maybe not fully developed (FiIg. 3G, H).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="live observations">
<paragraph pageId="0" pageNumber="1">Live observations.</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName lsidName="R. tsimelahy" pageId="0" pageNumber="1" rank="species" species="tsimelahy">
<emphasis italics="true" pageId="0" pageNumber="1">R. tsimelahy</emphasis>
</taxonomicName>
sp. nov. could be found in great numbers in the early morning (7-9 a.m.) on the forest floor of the spiny bush. The otherwise dry spiny bush was still quite wet because of dew. No juveniles were observed. Contrary to other
<taxonomicName authorityName="Wesener" authorityYear="2009" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Riotintobolus</emphasis>
</taxonomicName>
species, such as
<taxonomicName lsidName="R. mandenensis" pageId="0" pageNumber="1" rank="species" species="mandenensis">
<emphasis italics="true" pageId="0" pageNumber="1">R. mandenensis</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="R. minutus" pageId="0" pageNumber="1" rank="species" species="minutus">
<emphasis italics="true" pageId="0" pageNumber="1">R. minutus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="R. tsimelahy" pageId="0" pageNumber="1" rank="species" species="tsimelahy">
<emphasis italics="true" pageId="0" pageNumber="1">R. tsimelahy</emphasis>
</taxonomicName>
sp. nov. did not remain stiff like a stick when disturbed, but rolled-up into a spiral, a common defence behaviour for juliform millipedes.
</paragraph>
</subSubSection>
</treatment>
</document>