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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="351673C0372A5B059A310174A6FBC82D" docLanguage="en" docName="ZooKeys 953: 1-29" docOrigin="ZooKeys 953" docSource="http://dx.doi.org/10.3897/zookeys.953.53977" docTitle="Riotintobolus makayi Wesener 2020, sp. nov." docType="treatment" docUuid="233B0416-1281-4AB0-8D12-72688CFEE696" 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="166016735" LSID="urn:lsid:zoobank.org:act:233B0416-1281-4AB0-8D12-72688CFEE696" httpUri="http://treatment.plazi.org/id/351673C0372A5B059A310174A6FBC82D" lastPageNumber="1" pageId="0" pageNumber="1">
<subSubSection pageId="0" pageNumber="1" type="nomenclature">
<paragraph pageId="0" pageNumber="1">
<taxonomicName LSID="351673C0-372A-5B05-9A31-0174A6FBC82D" authority="Wesener, 2020" authorityName="Wesener" authorityYear="2020" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus makayi" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="makayi" status="sp. nov.">Riotintobolus makayi</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="1">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Riotintobolus makayi sp. nov., male holotype (ZFMK MYR 939) A multi-layer photograph, anterior body, lateral view B multi-layer photograph, leg pairs 1 - 7, ventral view C multi-layer photograph, posterior body end with telson D anterior gonopod, anterior view E anterior gonopod, posterior view F right posterior gonopod, anterior view G right posterior gonopod, posterior view. Abbreviations: Apo = apodeme; av = paraprocts; Co = collum (ring 1); Cx = coxite; Gn = gnathochilarium; Md = basal joints of mandible; Pre = epiproct; St = sternite; T = telopodite; x = lateral swollen process; y = opening of efferenct duct; z = apical membranous ' flag'; numbers refer to leg pair or segment number. Scale bars: 1 mm." figureDoi="10.3897/zookeys.953.53977.figure10" httpUri="https://binary.pensoft.net/fig/436171" pageId="0" pageNumber="1">Figure 10</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 MYR7173</emphasis>
, Madagascar, Province de Toliara, Makay Mts.,
<normalizedToken originalValue="forêt">foret</normalizedToken>
de galerie,
<geoCoordinate degrees="21" direction="south" minutes="13" orientation="latitude" precision="1" seconds="25.1" value="-21.223639">21°13'25.1&quot;S</geoCoordinate>
,
<geoCoordinate degrees="045" direction="east" minutes="19" orientation="longitude" precision="1" seconds="36.0" value="45.326664">045°19'36.0&quot;E</geoCoordinate>
, 512 m, coll. Jean Noel, 29.xi.2010. Paratypes: 1 ♂, 1 ♀,
<emphasis bold="true" pageId="0" pageNumber="1">ZFMK MYR939</emphasis>
, Madagascar, Province de Toliara, Makay Mts.,
<normalizedToken originalValue="forêt">foret</normalizedToken>
de galerie,
<geoCoordinate degrees="21" direction="south" minutes="13" orientation="latitude" precision="1" seconds="27.5" value="-21.224306">21°13'27.5&quot;S</geoCoordinate>
,
<geoCoordinate degrees="045" direction="east" minutes="19" orientation="longitude" precision="1" seconds="35.4" value="45.3265">045°19'35.4&quot;E</geoCoordinate>
, 531 m, coll. Jean Noel, 01.xii.2010.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="etymology">
<paragraph pageId="0" pageNumber="1">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="1">
Makayi, after the type locality, the area of Makay (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>
). 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 lsidName="R. makayi" pageId="0" pageNumber="1" rank="species" species="makayi">
<emphasis italics="true" pageId="0" pageNumber="1">R. makayi</emphasis>
</taxonomicName>
sp. nov. shares the flag-like membranous tip of the posterior gonopod with
<taxonomicName lsidName="R. mandenensis" pageId="0" pageNumber="1" rank="species" species="mandenensis">
<emphasis italics="true" pageId="0" pageNumber="1">R. mandenensis</emphasis>
</taxonomicName>
,
<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. aridus" pageId="0" pageNumber="1" rank="species" species="aridus">
<emphasis italics="true" pageId="0" pageNumber="1">R. aridus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="R. anomalus" pageId="0" pageNumber="1" rank="species" species="anomalus">
<emphasis italics="true" pageId="0" pageNumber="1">R. anomalus</emphasis>
</taxonomicName>
and
<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 authorityName="Wesener" authorityYear="2020" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus makayi" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="makayi">
<emphasis italics="true" pageId="0" pageNumber="1">Riotintobolus makayi</emphasis>
</taxonomicName>
sp. nov. shares the wide dorsal stripe, presence of tarsal pads on at least male legs 3-7, and a projecting epiproct only with
<taxonomicName lsidName="R. mandenensis" pageId="0" pageNumber="1" rank="species" species="mandenensis">
<emphasis italics="true" pageId="0" pageNumber="1">R. mandenensis</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="R. minutus" pageId="0" pageNumber="1" rank="species" species="minutus">
<emphasis italics="true" pageId="0" pageNumber="1">R. minutus</emphasis>
</taxonomicName>
, and
<taxonomicName lsidName="R. aridus" pageId="0" pageNumber="1" rank="species" species="aridus">
<emphasis italics="true" pageId="0" pageNumber="1">R. aridus</emphasis>
</taxonomicName>
.
<taxonomicName lsidName="R. makayi" pageId="0" pageNumber="1" rank="species" species="makayi">
<emphasis italics="true" pageId="0" pageNumber="1">R. makayi</emphasis>
</taxonomicName>
sp. nov. differs from
<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>
in the configuration of the flag of the posterior telopod, which consists only of a single fold, while it has two folds in the latter two species. The curved lateral process and the freely ending efferent duct are unique characters for
<taxonomicName lsidName="R. makayi" pageId="0" pageNumber="1" rank="species" species="makayi">
<emphasis italics="true" pageId="0" pageNumber="1">R. makayi</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Description</emphasis>
(based on male holotype):
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Measurements</emphasis>
: Male holotype: 38+0 segments. Ca. 22 mm long (broken), 2.4 mm wide. Female paratype: 38+0 segments. Ca. 33 mm long (broken), 3.5 mm wide.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Colour</emphasis>
(after ten years in ethanol): Head except for light clypeus black, body rings black, appendages red (Fig. 10A, B). Anterior margin of collum light brown. Dorsally with two wide light brown-reddish stripes, divided by a black stripe, all stripes even crossing the epiproct. Anal valves black, margin light brown, hypoproct light brown (Fig. 10C).
</paragraph>
<caption doi="10.3897/zookeys.953.53977.figure10" httpUri="https://binary.pensoft.net/fig/436171" pageId="0" pageNumber="1" start="Figure 10" startId="F10">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 10.</emphasis>
<taxonomicName authorityName="Wesener" authorityYear="2020" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus makayi" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="makayi">
<emphasis italics="true" pageId="0" pageNumber="1">Riotintobolus makayi</emphasis>
</taxonomicName>
sp. nov., male holotype (ZFMK MYR939)
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
multi-layer photograph, anterior body, lateral view
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
multi-layer photograph, leg pairs 1-7, ventral view
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
multi-layer photograph, 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>
anterior 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>
right posterior gonopod, posterior view.
<emphasis bold="true" pageId="0" pageNumber="1">Abbreviations</emphasis>
: Apo = apodeme; av = paraprocts; Co = collum (ring 1); Cx = coxite; Gn = gnathochilarium; Md = basal joints of mandible; Pre = epiproct; St = sternite; T = telopodite; x = lateral swollen process; y = opening of efferenct duct; z = apical membranous
<normalizedToken originalValue="flag">'flag'</normalizedToken>
; numbers refer to leg pair or segment number. Scale bars: 1 mm.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Head</emphasis>
: each eye with 26-28 ommatidia in six rows. Incisura lateralis open (Fig. 10A). 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. 10B). Antennae short, protruding back to segment 3. Length of antennomeres: 1&lt;2=3=4=5=6. Terminal antennomere with four large sensory cones located together inside a membranous area (Fig. 10B). Antennomere 5 and 6 latero-apically with sensilla basiconica.
</paragraph>
<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. Endochilarium not dissected.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Mandible</emphasis>
: Stipes without projection, well rounded (Fig. 10A, B). Gnathal lobe 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. 10A).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Body rings</emphasis>
: ozopores starting at segment 6, located slightly before, but touching 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 with lips, abundant micropunctation especially towards edges (Fig. 10C). Epiproct well-rounded, reaching slightly above paraproct with a massive process (Fig. 10C). Hypoproct inconspicuous (Fig. 10C).
</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 two 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 of leg 3- midbody with a tarsal pad, a single apical and three or four pairs of ventral spines (Fig. 10B). Legs on posterior part of body without a tarsal pad and two or three pairs of ventral spines. Length of midbody legs ca. 1.2 times body diameter in male.
</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 even shorter than male, only protruding back to collum. Female vulva simple, bivalve-like.
</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 (Fig. 10B). Coxae 3-7 without coxal processes (Fig. 10B).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Anterior gonopod</emphasis>
sternite massive, elongated into a wide, well-rounded, triangular lobe (Fig. 10D). Sternite in anterior view well-visible, without discernible apodemes, protruding almost as high as coxal processes. Coxite with sharp triangular mesal process (Fig. 10D, E). Telopodite with slender process arising mesally (Fig. 10E), process apically curved with a strong triangular projection (Fig. 10D), tip well-rounded tip, protruding above lateral margin of telopodite (Fig. 10C).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Posterior gonopods</emphasis>
consisting of two parts, separated by an articulation: a long coxite and a slightly shorter telopodite, efferent duct discharging apically (Fig. 10G). Process of coxite and telopodite standing in same axis (Fig. 10F, G). Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite (Fig. 10G). Basal part of coxite wide, mesally with a triangular sclerite located on lower level than remaining part (Fig. 10G). Coxite elongated. Efferent duct running at mesal margin of coxite (Fig. 10F, G). Telopodite as wide as but slightly shorter than coxite, standing in same axis, apically with a membranous
<normalizedToken originalValue="flag">'flag'</normalizedToken>
(Fig. 10F, G). Laterally with sclerotised projection with a slender mesad-orientated branch (Fig. 10F, G). Membranous
<normalizedToken originalValue="flag">'flag'</normalizedToken>
longer than lateral process. Efferent duct ending freely, projecting above 'flag apically (Fig. 10F, G).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="comments">
<paragraph pageId="0" pageNumber="1">Comments.</paragraph>
<paragraph pageId="0" pageNumber="1">
this species lives in direct sympatry with
<taxonomicName authorityName="Wesener" authorityYear="2020" class="Diplopoda" family="Pachybolidae" genus="Riotintobolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Riotintobolus bovinus" order="Spirobolida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="bovinus">
<emphasis italics="true" pageId="0" pageNumber="1">Riotintobolus bovinus</emphasis>
</taxonomicName>
sp. nov. The female carried eggs in the body.
</paragraph>
</subSubSection>
</treatment>
</document>