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<mods:classification id="25AEA45ACF770E5FA7AC8F789C9E963E">journal article</mods:classification>
<mods:identifier id="72B6F2EF4C6BA51E2A3EFB888A357BC6" type="CLB-Dataset">307986</mods:identifier>
<mods:identifier id="FDE5A929B4F170217EA9682B8CBB2447" type="DOI">10.1093/zoolinnean/zlae143</mods:identifier>
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<mods:identifier id="830E0999BAD26612007B83720903E075" type="ISSN">0024-4082</mods:identifier>
<mods:identifier id="B0ED2C3E9DE46E59C31F3574809AF3B3" type="Zenodo-Dep">14811333</mods:identifier>
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<subSubSection id="E0D5652BFFD0FFB0FC78A512FAD60576" box="[946,1323,1815,1842]" pageId="7" pageNumber="8" type="nomenclature">
<paragraph id="A87036A0FFD0FFB0FC78A512FAD60576" blockId="7.[946,1323,1815,1842]" box="[946,1323,1815,1842]" pageId="7" pageNumber="8">
<heading id="F33881CCFFD0FFB0FC78A512FAD60576" bold="true" box="[946,1323,1815,1842]" centered="true" fontSize="11" level="2" pageId="7" pageNumber="8" reason="2">
<taxonomicName id="6FCF4D23FFD0FFB0FC78A512FAD60576" ID-CoL="37ZVP" authority="de Beauchamp, 1952" authorityName="de Beauchamp" authorityYear="1952" box="[946,1323,1815,1842]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="7" pageNumber="8" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFD0FFB0FC78A512FAD60576" bold="true" box="[946,1323,1815,1842]" pageId="7" pageNumber="8">
<emphasis id="9ABBEAB2FFD0FFB0FC78A512FBB40575" bold="true" box="[946,1097,1815,1842]" italics="true" pageId="7" pageNumber="8">Dugesia milloti</emphasis>
<bibRefCitation id="CC5E4B51FFD0FFB0FB84A512FAD60576" author="de Beauchamp P" box="[1102,1323,1815,1841]" pageId="7" pageNumber="8" pagination="362 - 70" refId="ref16540" refString="de Beauchamp P. Sur les Dugesia (Turbellaries Triclades) d'Afrique tropicale et de Madagascar. Bulletin de la Societe Zoologique de France 1952; 77: 362 - 70." type="journal article" year="1952">de Beauchamp, 1952</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFD0FFBFFBDFA544FE1807A6" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" type="materials_examined">
<paragraph id="A87036A0FFD0FFB0FBDFA544FB35051C" blockId="7.[1045,1224,1857,1883]" box="[1045,1224,1857,1883]" pageId="7" pageNumber="8">
<emphasis id="9ABBEAB2FFD0FFB0FBDFA544FB35051C" box="[1045,1224,1857,1883]" italics="true" pageId="7" pageNumber="8">Material examined</emphasis>
</paragraph>
<paragraph id="A87036A0FFD0FFB0FCE0A56DFB9F05F9" blockId="7.[810,1459,1896,1982]" pageId="7" pageNumber="8">
<materialsCitation id="18A73CFDFFD0FFB0FCE0A56DFBA305F9" collectingDate="1957-09-17" collectionCode="ZMA, V" country="Madagascar" latitude="-12.616667" location="Roussette" longLatPrecision="1" longitude="49.15" municipality="Montagne d'Ambre" pageId="7" pageNumber="8" specimenCount="1">
<collectionCode id="CEDEAE65FFD0FFB0FCE0A56DFC9E05C7" box="[810,867,1896,1920]" country="Netherlands" httpUri="http://grbio.org/cool/a65z-wv6r" name="Universiteit van Amsterdam, Zoologisch Museum" pageId="7" pageNumber="8">ZMA</collectionCode>
<collectionCode id="CEDEAE65FFD0FFB0FCBAA56DFC7E05C7" box="[880,899,1896,1920]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="7" pageNumber="8" type="Museum">V</collectionCode>
.Pl. 787.1, stream of
<location id="AD10607BFFD0FFB0FBA0A56DFB2D0538" LSID="urn:lsid:plazi:treatment:206687B6FFD0FFBDFC78A512FE76071D:AD10607BFFD0FFB0FBA0A56DFB2D0538" box="[1130,1232,1896,1919]" country="Madagascar" latitude="-12.616667" longLatPrecision="1" longitude="49.15" municipality="Montagne d'Ambre" name="Roussette" pageId="7" pageNumber="8">Roussette</location>
,
<collectingMunicipality id="4814ACDAFFD0FFB0FB2BA56DFA5205C7" box="[1249,1455,1896,1920]" pageId="7" pageNumber="8">Montagne dAmbre</collectingMunicipality>
,
<collectingCountry id="D0D87630FFD0FFB0FCE0A582FC5E05D8" box="[810,931,1927,1951]" name="Madagascar" pageId="7" pageNumber="8">Madagascar</collectingCountry>
(
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12°37
<emphasis id="9ABBEAB2FFD0FFB0FC3BA583FC0A05D8" box="[1009,1015,1926,1951]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis>
0.00
<emphasis id="9ABBEAB2FFD0FFB0FBEEA583FBD205D8" box="[1060,1071,1926,1951]" italics="true" pageId="7" pageNumber="8">ʹʹ</emphasis>
S
</geoCoordinate>
<geoCoordinate id="CDFB5067FFD0FFB0FB89A582FB3205D8" box="[1091,1231,1926,1951]" degrees="49" direction="east" minutes="8" orientation="longitude" pageId="7" pageNumber="8" precision="1" seconds="60.00" value="49.15">
49°8
<emphasis id="9ABBEAB2FFD0FFB0FBB8A583FB8505D8" box="[1138,1144,1926,1951]" italics="true" pageId="7" pageNumber="8">ʹ</emphasis>
60.00
<emphasis id="9ABBEAB2FFD0FFB0FB7BA583FB4105D8" box="[1201,1212,1926,1951]" italics="true" pageId="7" pageNumber="8">ʹʹ</emphasis>
E
</geoCoordinate>
),
<date id="DC711060FFD0FFB0FB29A582FA5205D8" box="[1251,1455,1927,1951]" pageId="7" pageNumber="8" value="1957-09-17">
<collectingDate id="CC35E988FFD0FFB0FB29A582FA5205D8" box="[1251,1455,1927,1951]" pageId="7" pageNumber="8" value="1957-09-17">17 September 1957</collectingDate>
</date>
, sagittal sections on nine slides
</materialsCitation>
.
</paragraph>
<caption id="FCB06628FFDFFFBFFF4AA696FE24075C" ID-DOI="http://doi.org/10.5281/zenodo.14811339" ID-Zenodo-Dep="14811339" httpUri="https://zenodo.org/record/14811339/files/figure.png" pageId="8" pageNumber="9" startId="8.[128,193,1171,1195]" targetBox="[131,1470,145,1140]" targetPageId="8" targetType="figure">
<paragraph id="A87036A0FFDFFFBFFF4AA696FE24075C" blockId="8.[128,1457,1171,1307]" pageId="8" pageNumber="9">
<emphasis id="9ABBEAB2FFDFFFBFFF4AA696FF2506EC" bold="true" box="[128,216,1171,1195]" pageId="8" pageNumber="9">Figure 2.</emphasis>
Maximum likelihood molecular phylogenetic tree obtained from the concatenated dataset (
<emphasis id="9ABBEAB2FFDFFFBFFBF0A691FB9E06EC" box="[1082,1123,1172,1195]" italics="true" pageId="8" pageNumber="9">COI</emphasis>
, ITS1, 18S, and 28S) using the program IQ-tree 2.2.2.6. Supports on each node correspond with the supports of bootstrap (BS) of the maximum likelihood-based RaxML and IQ-tree programs, and supports of posterior probability (PP) of the Bayesian inference-based program MrBayes, respectively. Dashes () indicate those nodes not recovered in the corresponding approaches. Black squares on some nodes show full support for all three approaches. Scale bar: substitutions per position.
</paragraph>
</caption>
<paragraph id="A87036A0FFDFFFBFFF56A749FE1807A6" blockId="8.[127,777,1354,1505]" pageId="8" pageNumber="9">
<materialsCitation id="18A73CFDFFDFFFBFFF56A749FE1C07A6" collectingDate="1892-03-25" collectorName="Mf N F &amp; F. Sikora" country="Madagascar" latitude="-19.033333" location="Andrangolaoka" longLatPrecision="1" longitude="47.916668" pageId="8" pageNumber="9" specimenCount="1">
<collectorName id="053A5376FFDFFFBFFF56A749FF120723" box="[156,239,1356,1380]" pageId="8" pageNumber="9">Mf N F</collectorName>
217-c,
<location id="AD10607BFFDFFFBFFEF4A749FE200723" LSID="urn:lsid:plazi:treatment:206687B6FFD0FFBDFC78A512FE76071D:AD10607BFFDFFFBFFEF4A749FE200723" box="[318,477,1356,1380]" country="Madagascar" latitude="-19.033333" longLatPrecision="1" longitude="47.916668" name="Andrangolaoka" pageId="8" pageNumber="9">Andrangolaoka</location>
,
<collectingCountry id="D0D87630FFDFFFBFFE25A749FD950723" box="[495,616,1356,1380]" name="Madagascar" pageId="8" pageNumber="9">Madagascar</collectingCountry>
(
<geoCoordinate id="CDFB5067FFDFFFBFFD4AA749FCF40723" box="[640,777,1354,1380]" degrees="19" direction="south" minutes="1" orientation="latitude" pageId="8" pageNumber="9" precision="1" seconds="60.00" value="-19.033333">
19°1
<emphasis id="9ABBEAB2FFDFFFBFFD7BA74FFD4A0724" box="[689,695,1354,1379]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis>
60.00
<emphasis id="9ABBEAB2FFDFFFBFFD3BA74FFD010724" box="[753,764,1354,1379]" italics="true" pageId="8" pageNumber="9">ʹʹ</emphasis>
S
</geoCoordinate>
<geoCoordinate id="CDFB5067FFDFFFBFFF4BA76EFEF107C4" box="[129,268,1386,1411]" degrees="47" direction="east" minutes="55" orientation="longitude" pageId="8" pageNumber="9" precision="1" seconds="0.00" value="47.916668">
47°55
<emphasis id="9ABBEAB2FFDFFFBFFF77A76FFF3E07C4" box="[189,195,1386,1411]" italics="true" pageId="8" pageNumber="9">ʹ</emphasis>
0.00
<emphasis id="9ABBEAB2FFDFFFBFFF25A76FFF0707C4" box="[239,250,1386,1411]" italics="true" pageId="8" pageNumber="9">ʹʹ</emphasis>
E
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), coll.
<collectorName id="053A5376FFDFFFBFFE84A769FE5B07C4" box="[334,422,1387,1411]" pageId="8" pageNumber="9">F. Sikora</collectorName>
(specimens registered at the Berlin Museum on
<date id="DC711060FFDFFFBFFECDA78FFE5207E4" box="[263,431,1418,1443]" pageId="8" pageNumber="9" value="1892-03-25">
<collectingDate id="CC35E988FFDFFFBFFECDA78FFE5207E4" box="[263,431,1418,1443]" pageId="8" pageNumber="9" value="1892-03-25">25 March 1892</collectingDate>
</date>
), sagittal sections on nine slides;
<collectorName id="053A5376FFDFFFBFFF4BA7AFFF340785" box="[129,201,1450,1474]" pageId="8" pageNumber="9">Mf N F</collectorName>
217-a, ibid., sagittal sections on eight slides;
<collectorName id="053A5376FFDFFFBFFD4BA7AFFD340785" box="[641,713,1450,1474]" pageId="8" pageNumber="9">Mf N F</collectorName>
217-b, ibid., sagittal sections on five slides
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFDFFFBFFE84A404FD95049B" pageId="8" pageNumber="9" type="distribution">
<paragraph id="A87036A0FFDFFFBFFE84A404FDC1045C" blockId="8.[129,780,1537,1756]" box="[334,572,1537,1563]" pageId="8" pageNumber="9">
<emphasis id="9ABBEAB2FFDFFFBFFE84A404FDC1045C" box="[334,572,1537,1563]" italics="true" pageId="8" pageNumber="9">Geographical distribution</emphasis>
</paragraph>
<paragraph id="A87036A0FFDFFFBFFF4BA42DFD95049B" blockId="8.[129,780,1537,1756]" pageId="8" pageNumber="9">
Known from the following localities: (i) the
<typeStatus id="77748802FFDFFFBFFD48A42DFD4D0407" box="[642,688,1576,1600]" pageId="8" pageNumber="9">type</typeStatus>
locality Morafenobé, forêt
<collectorName id="053A5376FFDFFFBFFE81A443FE290419" box="[331,468,1606,1630]" pageId="8" pageNumber="9">de Mahajeby</collectorName>
, stream of Roussette, northwest
<collectingCountry id="D0D87630FFDFFFBFFF7CA463FECE0439" box="[182,307,1638,1662]" name="Madagascar" pageId="8" pageNumber="9">Madagascar</collectingCountry>
; (ii)
<collectingCounty id="41114E2CFFDFFFBFFEADA463FCFB0439" box="[359,774,1638,1662]" pageId="8" pageNumber="9">Parc National de La Montagne dAmbre</collectingCounty>
,
<collectingMunicipality id="4814ACDAFFDFFFBFFF49A480FEE004DA" box="[131,285,1669,1693]" pageId="8" pageNumber="9">Diana Region</collectingMunicipality>
, northern
<collectingCountry id="D0D87630FFDFFFBFFE54A480FDE604DA" box="[414,539,1669,1693]" name="Madagascar" pageId="8" pageNumber="9">Madagascar</collectingCountry>
; (iii) Andrangolaoka,
<location id="AD10607BFFDFFFBFFF4BA4A1FEA304FB" LSID="urn:lsid:plazi:treatment:206687B6FFD0FFBDFC78A512FE76071D:AD10607BFFDFFFBFFF4BA4A1FEA304FB" box="[129,350,1700,1724]" country="Madagascar" county="Parc National de La Montagne d'Ambre" municipality="Diana Region" name="Central High Plateau" pageId="8" pageNumber="9" stateProvince="Mutsamudu, Anjouan">Central High Plateau</location>
, east to
<location id="AD10607BFFDFFFBFFE7CA4A0FDA504FB" LSID="urn:lsid:plazi:treatment:206687B6FFD0FFBDFC78A512FE76071D:AD10607BFFDFFFBFFE7CA4A0FDA504FB" box="[438,600,1700,1724]" country="Madagascar" county="Parc National de La Montagne d'Ambre" municipality="Diana Region" name="Mantasoa Lake" pageId="8" pageNumber="9" stateProvince="Mutsamudu, Anjouan">Mantasoa Lake</location>
; (iv) Cascade de
<collectingRegion id="6A0BF842FFDFFFBFFF4BA4C1FE9F049B" box="[129,354,1732,1756]" isEnumeration="true" pageId="8" pageNumber="9">Mutsamudu, Anjouan</collectingRegion>
, Comoro Islands (
<figureCitation id="30F42A25FFDFFFBFFDEAA4C1FDA5049B" box="[544,600,1732,1756]" captionStart="Figure 1" captionStartId="5.[113,178,1836,1860]" captionTargetBox="[177,1395,150,1801]" captionTargetId="figure-72@5.[175,1398,148,1804]" captionTargetPageId="5" captionText="Figure 1. Geographic distribution of the long-eared freshwater planarians of the genus Dugesia in Madagascar and adjacent islands." figureDoi="http://doi.org/10.5281/zenodo.14811337" httpUri="https://zenodo.org/record/14811337/files/figure.png" pageId="8" pageNumber="9">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFDFFFBDFE9DA4F9FE76071D" lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9" type="discussion">
<paragraph id="A87036A0FFDFFFBFFE9DA4F9FDCE0552" blockId="8.[129,779,1787,1975]" box="[343,563,1787,1814]" pageId="8" pageNumber="9">
<emphasis id="9ABBEAB2FFDFFFBFFE9DA4F9FDCE0552" box="[343,563,1787,1814]" italics="true" pageId="8" pageNumber="9">Comparative discussion</emphasis>
</paragraph>
<paragraph id="A87036A0FFDFFFBFFF4BA527FBA50478" blockId="8.[129,779,1787,1975]" lastBlockId="8.[824,1477,1355,1975]" pageId="8" pageNumber="9">
As
<bibRefCitation id="CC5E4B51FFDFFFBFFF6FA527FEAD057D" author="de Vries EJ" box="[165,336,1826,1850]" pageId="8" pageNumber="9" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries (1988)</bibRefCitation>
based her redescription of
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<emphasis id="9ABBEAB2FFDFFFBFFDB1A526FD26057D" box="[635,731,1826,1850]" italics="true" pageId="8" pageNumber="9">D. milloti</emphasis>
</taxonomicName>
primarily on specimens from Montagne dAmbre (in our material represented by specimen
<collectionCode id="CEDEAE65FFDFFFBFFE41A564FE39053E" box="[395,452,1889,1913]" country="Netherlands" httpUri="http://grbio.org/cool/a65z-wv6r" name="Universiteit van Amsterdam, Zoologisch Museum" pageId="8" pageNumber="9">ZMA</collectionCode>
<collectionCode id="CEDEAE65FFDFFFBFFE01A565FE23053F" box="[459,478,1888,1912]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="8" pageNumber="9" type="Museum">V</collectionCode>
.Pl. 787.1) her account still is basically correct, although she did not provide specific information on the testes and the position of the mouth opening. Examination of our material learned that the small testes are situated dorsally and that the follicles extend from a short distance behind the ovaries to the posterior end of the body. The ovaries are located at a short distance (~400 μm) behind the brain, i.e. at about one-ninth of the distance between the brain and the root of the pharynx. The mouth opening is located at about one-sixth of the distance between the hind wall of the pharyngeal pocket and the root of the pharynx.
</paragraph>
<paragraph id="A87036A0FFDFFFBFFC9FA443FA91053F" blockId="8.[824,1477,1355,1975]" pageId="8" pageNumber="9">
Characteristic for
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<emphasis id="9ABBEAB2FFDFFFBFFBD9A442FB8F0419" box="[1043,1138,1606,1630]" italics="true" pageId="8" pageNumber="9">D. milloti</emphasis>
</taxonomicName>
are the two annular atrial folds or valves, the posterior valve receiving a bundle of longitudinal muscles that extends posteriorly from the penis bulb (cf.
<bibRefCitation id="CC5E4B51FFDFFFBFFCF3A4A1FC3404FB" author="de Vries EJ" box="[825,969,1700,1724]" pageId="8" pageNumber="9" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: figs 9, 10;
<figureCitation id="30F42A25FFDFFFBFFB8CA4A1FB8204FB" box="[1094,1151,1700,1724]" captionStart="Figure 3" captionStartId="9.[113,178,1794,1818]" captionTargetBox="[114,1458,144,1766]" captionTargetId="figure-6@9.[114,1458,144,1766]" captionTargetPageId="9" captionText="Figure 3. Dugesia milloti. Specimen ZMA V.Pl. 787.1, photomicrographs of sagittal sections. A, B, male copulatory apparatus, showing the slim penis papilla (pp), diaphragm (d), seminal vesicle (sv), and atrial valves (av)." figureDoi="http://doi.org/10.5281/zenodo.14811345" httpUri="https://zenodo.org/record/14811345/files/figure.png" pageId="8" pageNumber="9">Fig. 3</figureCitation>
). These annular atrial folds are much less clear in the old specimens from Andrangolaoka, which suffer from preservation and contraction artefacts (
<figureCitation id="30F42A25FFDFFFBFFA9CA4E6FA6B04BC" box="[1366,1430,1763,1787]" captionStart="Figure 4" captionStartId="10.[129,194,1180,1204]" captionTargetBox="[129,1473,144,1152]" captionTargetId="figure-333@10.[129,1473,144,1152]" captionTargetPageId="10" captionText="Figure 4. Dugesia milloti. Specimen MfN F217-c, sagittal section of the male copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811350" httpUri="https://zenodo.org/record/14811350/files/figure.png" pageId="8" pageNumber="9">Figs 4</figureCitation>
,
<figureCitation id="30F42A25FFDFFFBFFA6EA4E7FA4C04BD" box="[1444,1457,1762,1786]" captionStart="Figure 5" captionStartId="11.[113,178,1151,1175]" captionTargetBox="[211,1362,146,1121]" captionTargetId="figure-498@11.[209,1364,144,1123]" captionTargetPageId="11" captionText="Figure 5. Dugesia milloti. Specimen MfN F217-c, sagittal reconstruction of the male copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811354" httpUri="https://zenodo.org/record/14811354/files/figure.png" pageId="8" pageNumber="9">5</figureCitation>
). However, in specimen
<collectionCode id="CEDEAE65FFDFFFBFFBE9A507FBA5055D" box="[1059,1112,1794,1818]" country="Germany" httpUri="http://grbio.org/cool/jp8y-i5nv" name="Museum für Naturkunde" pageId="8" pageNumber="9" type="Museum">MfN</collectionCode>
F217-a, folds in the atrium are still visible, presumably representing the most posterior valve—also receiving the longitudinal muscles from the penis bulb—as the anterior fold sits at the root of the penis papilla (
<figureCitation id="30F42A25FFDFFFBFFAE9A565FAA6053F" box="[1315,1371,1888,1912]" captionStart="Figure 6" captionStartId="12.[129,194,1046,1070]" captionTargetBox="[225,1377,144,1018]" captionTargetId="figure-583@12.[225,1377,144,1018]" captionTargetPageId="12" captionText="Figure 6. Dugesia milloti. Specimen MfN F217-a, photomicrograph of a sagittal section of the male copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811356" httpUri="https://zenodo.org/record/14811356/files/figure.png" pageId="8" pageNumber="9">Fig. 6</figureCitation>
).
</paragraph>
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As compared with the other long-eared species of
<taxonomicName id="6FCF4D23FFDFFFBFFA8FA585FA6E05D0" authorityName="Girard" authorityYear="1850" box="[1349,1427,1920,1943]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFDFFFBFFA8FA585FA6E05D0" box="[1349,1427,1920,1943]" italics="true" pageId="8" pageNumber="9">Dugesia</emphasis>
</taxonomicName>
(see below), the copulatory apparatus of
<taxonomicName id="6FCF4D23FFDFFFBFFB6DA59AFAFF05F0" authorityName="de Beauchamp" authorityYear="1952" box="[1191,1282,1951,1975]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="8" pageNumber="9" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDFFFBFFB6DA59AFAFF05F0" box="[1191,1282,1951,1975]" italics="true" pageId="8" pageNumber="9">D. milloti</emphasis>
</taxonomicName>
is characterized by a slim, conical penis papilla and a large, valve-like diaphragm, the latter receiving the erythrophil secretion of penial glands that contributes to the formation of the spermatophore (
<bibRefCitation id="CC5E4B51FFDDFFBDFD78A726FF48071D" author="de Vries EJ" pageId="10" pageNumber="11" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: figs 9, 10;
<figureCitation id="30F42A25FFDDFFBDFEEDA747FE9E071D" box="[295,355,1346,1370]" captionStart="Figure 5" captionStartId="11.[113,178,1151,1175]" captionTargetBox="[211,1362,146,1121]" captionTargetId="figure-498@11.[209,1364,144,1123]" captionTargetPageId="11" captionText="Figure 5. Dugesia milloti. Specimen MfN F217-c, sagittal reconstruction of the male copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811354" httpUri="https://zenodo.org/record/14811354/files/figure.png" pageId="10" pageNumber="11">Figs 5</figureCitation>
,
<figureCitation id="30F42A25FFDDFFBDFEA4A747FE86071D" box="[366,379,1346,1370]" captionStart="Figure 6" captionStartId="12.[129,194,1046,1070]" captionTargetBox="[225,1377,144,1018]" captionTargetId="figure-583@12.[225,1377,144,1018]" captionTargetPageId="12" captionText="Figure 6. Dugesia milloti. Specimen MfN F217-a, photomicrograph of a sagittal section of the male copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811356" httpUri="https://zenodo.org/record/14811356/files/figure.png" pageId="10" pageNumber="11">6</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -0,0 +1,787 @@
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<emphasis id="9ABBEAB2FFDAFFBAFC5BA78FFBCC07E3" bold="true" box="[913,1073,1417,1444]" italics="true" pageId="13" pageNumber="14">Dugesia insolita</emphasis>
Stocchino &amp; Sluys
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,
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<emphasis id="9ABBEAB2FFDAFFBAFBDFA40DFB350465" box="[1045,1224,1544,1570]" italics="true" pageId="13" pageNumber="14">Material examined</emphasis>
</paragraph>
<paragraph id="A87036A0FFDAFFBAFCE0A42BFAB70545" blockId="13.[810,1460,1544,1795]" pageId="13" pageNumber="14">
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<emphasis id="9ABBEAB2FFDAFFBAFCE0A42BFC770401" box="[810,906,1582,1606]" italics="true" pageId="13" pageNumber="14">
<typeStatus id="77748802FFDAFFBAFCE0A42BFC780401" box="[810,901,1582,1606]" pageId="13" pageNumber="14" type="holotype">Holotype</typeStatus>
:
</emphasis>
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(field code: GS32-Dmad5-6) one set of sagittal sections on 18 slides,
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,
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, small unnamed water course, tributary of
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,
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,
<quantity id="6F379B45FFDAFFBAFB22A488FAD904E3" box="[1256,1316,1676,1701]" metricMagnitude="3" metricUnit="m" metricValue="4.0" pageId="13" pageNumber="14" unit="km" value="4.0">
<locationDeviation id="B9C22F82FFDAFFBAFB22A488FAD904E3" box="[1256,1316,1676,1701]" metricMagnitude="3" metricUnit="m" metricValue="4.0" pageId="13" pageNumber="14" unit="km" value="4.0">4 km</locationDeviation>
</quantity>
upstream of
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(
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21°08
<emphasis id="9ABBEAB2FFDAFFBAFBEFA4AFFBD60484" box="[1061,1067,1706,1731]" italics="true" pageId="13" pageNumber="14">ʹ</emphasis>
S
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47°13
<emphasis id="9ABBEAB2FFDAFFBAFBB0A4AFFB7D0484" box="[1146,1152,1706,1731]" italics="true" pageId="13" pageNumber="14">ʹ</emphasis>
E
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), between Route National 7 to
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and Route national 25 to
<location id="AD10607BFFDAFFBAFB19A4CEFCA10545" LSID="urn:lsid:plazi:treatment:206687B6FFDAFFA3FC5BA78FFB4C0613:AD10607BFFDAFFBAFB19A4CEFCA10545" country="Madagascar" latitude="-21.133333" longLatPrecision="1265" longitude="47.216667" name="Ranomafana National Park" pageId="13" pageNumber="14" stateProvince="Upper Mangoky basin">Ranomafana National Park</location>
,
<collectingCountry id="D0D87630FFDAFFBAFCAEA4EFFC230545" box="[868,990,1770,1794]" name="Madagascar" pageId="13" pageNumber="14">Madagascar</collectingCountry>
,
<date id="DC711060FFDAFFBAFC2DA4EFFB6C0544" box="[999,1169,1770,1795]" pageId="13" pageNumber="14" value="2011-09">
<collectingDate id="CC35E988FFDAFFBAFC2DA4EFFB6C0544" box="[999,1169,1770,1795]" pageId="13" pageNumber="14" value="2011-09">September 2011</collectingDate>
</date>
, coll.
<collectorName id="053A5376FFDAFFBAFB00A4EEFABB0545" box="[1226,1350,1770,1794]" pageId="13" pageNumber="14">R. Manconi</collectorName>
.
</materialsCitation>
</paragraph>
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<paragraph id="A87036A0FFDAFFBAFCE0A521FC2D05FE" blockId="13.[810,1459,1828,1977]" pageId="13" pageNumber="14">
<emphasis id="9ABBEAB2FFDAFFBAFCE0A521FC29057B" box="[810,980,1828,1852]" italics="true" pageId="13" pageNumber="14">Other specimens:</emphasis>
Nine individuals were preserved in absolute ethanol, two of which (
<emphasis id="9ABBEAB2FFDAFFBAFBEEA546FBA2051C" box="[1060,1119,1859,1883]" italics="true" pageId="13" pageNumber="14">Dins1</emphasis>
and
<emphasis id="9ABBEAB2FFDAFFBAFB5CA546FB2C051C" box="[1174,1233,1859,1883]" italics="true" pageId="13" pageNumber="14">Dins2</emphasis>
) were used for DNA extraction and sequencing in the study of
<bibRefCitation id="CC5E4B51FFDAFFBAFB3BA567FA53053C" author="Sola E &amp; Leria L &amp; Stocchino GA" box="[1265,1454,1890,1915]" pageId="13" pageNumber="14" pagination="1219 - 33" refId="ref18745" refString="Sola E, Leria L, Stocchino GA et al. Three dispersal routes out of Africa: a puzzling biogeographical history in freshwater planarians. Journal of Biogeography 2022; 49: 1219 - 33. https: // doi. org / 10.1111 / jbi. 14371" type="journal article" year="2022">
Solà
<emphasis id="9ABBEAB2FFDAFFBAFAEDA566FAAE053D" box="[1319,1363,1890,1914]" italics="true" pageId="13" pageNumber="14">et al</emphasis>
. (2022)
</bibRefCitation>
. GenBank accession numbers of the obtained sequences are given in
<tableCitation id="E54D031BFFDAFFBAFC4AA5A4FC3705FE" box="[896,970,1953,1977]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Specimens/species used in the phylogenetic analysis, with their abbreviated names or codes, and GenBank accession numbers for the various genes/sequences (COI, ITS1, 18S, 28S, DUNUC3).Dashes (): unavailable sequences. Because DUNUC3 was not used in the present study, only newly obtained DUNUC3 are provided, for the others consult Solà et al. (2022)." httpUri="http://table.plazi.org/id/FCB06628FFD1FFB1FF4BA28AFBD00298" pageId="13" pageNumber="14" tableUuid="FCB06628FFD1FFB1FF4BA28AFBD00298">Table 2</tableCitation>
.
</paragraph>
</subSubSection>
<caption id="FCB06628FFD9FFB9FF4BA4E9FD810558" ID-DOI="http://doi.org/10.5281/zenodo.14811363" ID-Zenodo-Dep="14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="14" pageNumber="15" startId="14.[129,194,1772,1796]" targetBox="[194,1409,147,1741]" targetPageId="14" targetType="figure">
<paragraph id="A87036A0FFD9FFB9FF4BA4E9FD810558" blockId="14.[129,1448,1772,1824]" pageId="14" pageNumber="15">
<emphasis id="9ABBEAB2FFD9FFB9FF4BA4E9FF250543" bold="true" box="[129,216,1772,1796]" pageId="14" pageNumber="15">Figure 8.</emphasis>
<emphasis id="9ABBEAB2FFD9FFB9FF17A4E8FE570543" box="[221,426,1772,1796]" italics="true" pageId="14" pageNumber="15">
<taxonomicName id="6FCF4D23FFD9FFB9FF17A4E8FE5A0543" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[221,423,1772,1796]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="14" pageNumber="15" phylum="Platyhelminthes" rank="species" species="crassimentula">Dugesia crassimentula</taxonomicName>
.
</emphasis>
Holotype ZMA V.Pl. 786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right. A, male copulatory apparatus; B, female copulatory apparatus.
</paragraph>
</caption>
<subSubSection id="E0D5652BFFD9FFB9FE58A545FEF5059B" pageId="14" pageNumber="15" type="etymology">
<paragraph id="A87036A0FFD9FFB9FE58A545FE05051D" blockId="14.[129,778,1856,2012]" box="[402,504,1856,1882]" pageId="14" pageNumber="15">
<emphasis id="9ABBEAB2FFD9FFB9FE58A545FE05051D" box="[402,504,1856,1882]" italics="true" pageId="14" pageNumber="15">Etymology</emphasis>
</paragraph>
<paragraph id="A87036A0FFD9FFB9FF4BA563FEF5059B" blockId="14.[129,778,1856,2012]" pageId="14" pageNumber="15">
The specific epithet, derived from the Latin adjective
<emphasis id="9ABBEAB2FFD9FFB9FD7EA563FCFE0539" box="[692,771,1894,1918]" italics="true" pageId="14" pageNumber="15">insolitus</emphasis>
, unusual, uncommon, alludes to the remarkable situation that in this species the diaphragm has taken over the function of the penis papilla.
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFD9FFB8FB85A545FF0C0046" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15" type="diagnosis">
<paragraph id="A87036A0FFD9FFB9FB85A545FB51051E" blockId="14.[1103,1196,1856,1881]" box="[1103,1196,1856,1881]" pageId="14" pageNumber="15">
<emphasis id="9ABBEAB2FFD9FFB9FB85A545FB51051E" box="[1103,1196,1856,1881]" italics="true" pageId="14" pageNumber="15">Diagnosis</emphasis>
</paragraph>
<paragraph id="A87036A0FFD9FFB8FCF3A563FF0C0046" blockId="14.[825,1476,1894,2012]" lastBlockId="15.[112,764,144,513]" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15">
A
<emphasis id="9ABBEAB2FFD9FFB9FC9BA562FC6E0539" box="[849,915,1895,1918]" italics="true" pageId="14" pageNumber="15">Dugesi</emphasis>
a species characterized by the following features: high triangular head with a distinctly pointed auricle on either side; mouth opening at the posterior end of the pharyngeal pocket; vasa deferentia separately opening into anterior portion of the bulbar lumen; functional penis papilla a stubby, cone-shaped structure exhibiting an outbulging that provides it a shape in the form of a head of a spear; erythrophil penis glands discharging through the lining epithelium of the functional penis papilla; male atrium with three annular folds, the posterior-most one receiving longitudinal muscles that extend from the penis bulb; hyperplasic ovaries extending posterior to the level of the root of the pharynx; oviducts separately opening into the most proximal section of the bursal canal; poorly developed ectal reinforcement layer extending from about the knee-shaped bend in the bursal canal to about halfway between the bend and the copulatory bursa.
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFD8FFB8FEF5A02EFE28002D" pageId="15" pageNumber="16" type="distribution">
<paragraph id="A87036A0FFD8FFB8FEF5A02EFDD00002" blockId="15.[319,557,555,581]" box="[319,557,555,581]" pageId="15" pageNumber="16">
<emphasis id="9ABBEAB2FFD8FFB8FEF5A02EFDD00002" box="[319,557,555,581]" italics="true" pageId="15" pageNumber="16">Geographical distribution</emphasis>
</paragraph>
<paragraph id="A87036A0FFD8FFB8FFBBA057FE28002D" blockId="15.[113,469,594,618]" box="[113,469,594,618]" pageId="15" pageNumber="16">
Known only from the
<typeStatus id="77748802FFD8FFB8FE98A057FE7D002D" box="[338,384,594,618]" pageId="15" pageNumber="16">type</typeStatus>
locality.
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFD8FFB8FE5AA08CFDE30434" pageId="15" pageNumber="16" type="biology_ecology">
<paragraph id="A87036A0FFD8FFB8FE5AA08CFE2600E4" blockId="15.[400,475,649,675]" box="[400,475,649,675]" pageId="15" pageNumber="16">
<emphasis id="9ABBEAB2FFD8FFB8FE5AA08CFE2600E4" box="[400,475,649,675]" italics="true" pageId="15" pageNumber="16">Habitat</emphasis>
</paragraph>
<paragraph id="A87036A0FFD8FFB8FFBBA0B5FD3100A0" blockId="15.[113,762,688,743]" pageId="15" pageNumber="16">Planarians were found under pebbles and boulders in shallow water of a rivulet in the relict Rainforests of the Atsinanana.</paragraph>
<paragraph id="A87036A0FFD8FFB8FEFBA102FDC60166" blockId="15.[113,765,775,1652]" box="[305,571,775,801]" pageId="15" pageNumber="16">
<emphasis id="9ABBEAB2FFD8FFB8FEFBA102FDC60166" box="[305,571,775,801]" italics="true" pageId="15" pageNumber="16">Reproduction and behaviour</emphasis>
</paragraph>
<paragraph id="A87036A0FFD8FFB8FFBBA12BFF1D0646" blockId="15.[113,765,775,1652]" pageId="15" pageNumber="16">
At collection, all
<specimenCount id="BEC9FD29FFD8FFB8FEFEA128FE350102" box="[308,456,813,837]" count="20" pageId="15" pageNumber="16" type="generic">20 specimens</specimenCount>
of
<taxonomicName id="6FCF4D23FFD8FFB8FE39A12BFDA20102" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[499,607,813,837]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="15" pageNumber="16" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFD8FFB8FE39A12BFDA20102" box="[499,607,813,837]" italics="true" pageId="15" pageNumber="16">D. insolita</emphasis>
</taxonomicName>
were asexual. After four months of rearing under standard laboratory conditions, a single asexual specimen underwent a sexualization process, thus producing an ex-fissiparous individual characterized by hyperplasic ovaries and presence of a copulatory apparatus. Asexual individuals exhibited post-pharyngeal fissioning.
</paragraph>
<paragraph id="A87036A0FFD8FFB8FF47A60CFDD906BC" blockId="15.[113,765,775,1652]" pageId="15" pageNumber="16">
Just after collection and during rearing, both in the field and in the laboratory, a particular kind of movement was observed. Besides the usual gliding
<typeStatus id="77748802FFD8FFB8FEB7A64DFE560627" box="[381,427,1096,1120]" pageId="15" pageNumber="16">type</typeStatus>
of locomotion, these planarians showed a swimming-like movement with up-and-down undulations of the body when they were not adhering to the substrate and moved, for example, from the water surface to the bottom of the container. Moreover, during their usual gliding the body margins showed an undulating movement.
</paragraph>
<paragraph id="A87036A0FFD8FFB8FF47A706FDE30434" blockId="15.[113,765,775,1652]" pageId="15" pageNumber="16">Furthermore, on some occasions we observed a peculiar autotomy-like process when the worms were picked up softly with a paintbrush. Immediately after the planarians were touched, they quickly detached their tail, thus dividing at a post-pharyngeal level. The wound margins of the two resulting fragments were characterized by very irregular outlines, resembling those resulting from a forceful laceration. On one occasion, it happened that after this first division, touching of the anterior portion triggered another quick autotomy-like process, thus resulting in the detachment of another post-pharyngeal fragment. After these autotomy-like processes, each of the resulting fragments regenerated the missing body parts.</paragraph>
</subSubSection>
<caption id="FCB06628FFD8FFB8FCE1A43CFA780415" ID-DOI="http://doi.org/10.5281/zenodo.14811365" ID-Zenodo-Dep="14811365" box="[811,1413,1593,1618]" httpUri="https://zenodo.org/record/14811365/files/figure.png" pageId="15" pageNumber="16" startId="15.[811,876,1593,1617]" targetBox="[816,1452,147,1561]" targetPageId="15" targetType="figure">
<paragraph id="A87036A0FFD8FFB8FCE1A43CFA780415" blockId="15.[811,1413,1593,1618]" box="[811,1413,1593,1618]" pageId="15" pageNumber="16">
<emphasis id="9ABBEAB2FFD8FFB8FCE1A43CFC7F0416" bold="true" box="[811,898,1593,1617]" pageId="15" pageNumber="16">Figure 9.</emphasis>
<emphasis id="9ABBEAB2FFD8FFB8FC4CA43FFBE60415" box="[902,1051,1594,1618]" italics="true" pageId="15" pageNumber="16">
<taxonomicName id="6FCF4D23FFD8FFB8FC4CA43FFBEB0415" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[902,1046,1594,1618]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="15" pageNumber="16" phylum="Platyhelminthes" rank="species" species="insolita">Dugesia insolita</taxonomicName>
.
</emphasis>
Habitus of a live fissiparous specimen.
</paragraph>
</caption>
<subSubSection id="E0D5652BFFD8FFA5FE4AA491FAE000DA" lastPageId="18" lastPageNumber="19" pageId="15" pageNumber="16" type="description">
<paragraph id="A87036A0FFD8FFB8FE4AA491FE1104EA" blockId="15.[384,492,1684,1709]" box="[384,492,1684,1709]" pageId="15" pageNumber="16">
<emphasis id="9ABBEAB2FFD8FFB8FE4AA491FE1104EA" box="[384,492,1684,1709]" italics="true" pageId="15" pageNumber="16">Description</emphasis>
</paragraph>
<paragraph id="A87036A0FFD8FFB8FFBBA4BFFC85049F" blockId="15.[112,763,1722,1965]" lastBlockId="15.[809,1459,1665,1971]" pageId="15" pageNumber="16">
Live animals with a high triangular head, provided with distinctly pointed auricles and two prominent eyes set in pigment-free patches (
<figureCitation id="30F42A25FFD8FFB8FF05A4FCFEF50556" box="[207,264,1785,1809]" captionStart="Figure 9" captionStartId="15.[811,876,1593,1617]" captionTargetBox="[816,1452,147,1561]" captionTargetId="figure-565@15.[815,1455,144,1566]" captionTargetPageId="15" captionText="Figure 9. Dugesia insolita.Habitus of a live fissiparous specimen." figureDoi="http://doi.org/10.5281/zenodo.14811365" httpUri="https://zenodo.org/record/14811365/files/figure.png" pageId="15" pageNumber="16">Fig. 9</figureCitation>
). Body size of live, fissiparous specimens ranged from
<quantity id="6F379B45FFD8FFB8FF66A51DFEDB0568" box="[172,294,1816,1840]" metricMagnitude="-3" metricUnit="m" metricValue="9.5" metricValueMax="11.0" metricValueMin="8.0" pageId="15" pageNumber="16" unit="mm" value="9.5" valueMax="11.0" valueMin="8.0">8 to 11 mm</quantity>
in length and
<quantity id="6F379B45FFD8FFB8FE08A51DFDD60568" box="[450,555,1816,1840]" metricMagnitude="-3" metricUnit="m" metricValue="1.75" metricValueMax="2.0" metricValueMin="1.5" pageId="15" pageNumber="16" unit="mm" value="1.75" valueMax="2.0" valueMin="1.5">1.52 mm</quantity>
in width, while the single sexualized specimen measured 17 ×
<quantity id="6F379B45FFD8FFB8FDE8A532FD9D0508" box="[546,608,1847,1871]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="15" pageNumber="16" unit="mm" value="5.0">5 mm</quantity>
. Dorsal surface with numerous dark-brown specks and a dark mid-dorsal stripe that runs from behind the pharynx to the tip of the tail (
<figureCitation id="30F42A25FFD8FFB8FD7BA573FD1405C9" box="[689,745,1910,1934]" captionStart="Figure 9" captionStartId="15.[811,876,1593,1617]" captionTargetBox="[816,1452,147,1561]" captionTargetId="figure-565@15.[815,1455,144,1566]" captionTargetPageId="15" captionText="Figure 9. Dugesia insolita.Habitus of a live fissiparous specimen." figureDoi="http://doi.org/10.5281/zenodo.14811365" httpUri="https://zenodo.org/record/14811365/files/figure.png" pageId="15" pageNumber="16">Fig. 9</figureCitation>
). In some specimens this dorsal stripe runs from the anterior tip of the head to the very tip of the tail. Ventral surface pale and provided with dark brown specks, less numerous than on the dorsal surface.
</paragraph>
<paragraph id="A87036A0FFD8FFB8FC8FA4DAFC8B0532" blockId="15.[809,1459,1665,1971]" pageId="15" pageNumber="16">Pharynx located at about halfway the length of the body, measuring about one-sixth of the body length. No extra layer of longitudinal muscles in the outer pharynx musculature. The mouth opening is located at the posterior end of the pharyngeal pocket.</paragraph>
<paragraph id="A87036A0FFD8FFA7FC8FA579FE700302" blockId="15.[809,1459,1665,1971]" lastBlockId="16.[129,778,144,669]" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16">
The outer pharynx epithelium is underlain by a thin, subepidermal layer of circular muscle, followed by an equally thin layer of longitudinal muscle; this outer pharyngeal musculature is devoid of an extra, inner layer of longitudinal muscles. The lining epithelium of the pharyngeal lumen is underlain by a thick, subepithelial layer of circular muscles (~50 μm thick), followed by a much thinner layer of longitudinal muscles (
<figureCitation id="30F42A25FFC7FFA7FEE8A328FE860302" box="[290,379,301,325]" captionStart="Figure 10" captionStartId="16.[129,194,1306,1330]" captionTargetBox="[129,1473,773,1277]" captionTargetId="figure-339@16.[129,1473,773,1277]" captionTargetPageId="16" captionText="Figure 10. Dugesia insolita. Holotype RMNH.VER.21536.1, photomicrographs, sagittal sections. A, pharynx; B, hyperplasic ovary." figureDoi="http://doi.org/10.5281/zenodo.14811367" httpUri="https://zenodo.org/record/14811367/files/figure.png" pageId="16" pageNumber="17">Fig. 10A</figureCitation>
).
</paragraph>
<paragraph id="A87036A0FFC7FFA7FF56A349FE7003E4" blockId="16.[129,778,144,669]" pageId="16" pageNumber="17">The well-developed testes are located dorsally and extend from about the anterior level of the hyperplasic ovaries far into the tail region of the body.</paragraph>
<paragraph id="A87036A0FFC7FFA7FF56A3AFFB7C0386" blockId="16.[129,778,144,669]" lastBlockId="16.[825,1476,144,669]" pageId="16" pageNumber="17">
The vasa deferentia are expanded to form spermiducal vesicles—filled with sperm—which, in addition, may show considerable outbulgings. Upon penetrating the muscular penis bulb, the vasa deferentia gradually decrease in diameter and after having become very narrow, the ducts separately open into the anterior portion of the bulbar lumen or seminal vesicle. The wall of the latter is highly pleated, with the folds projecting into its lumen. One of the larger folds forms an elongated papilla that projects posteriad into the lumen of a structure that functions as penis papilla (see below); through this fold runs a minute duct by means of which the seminal vesicle communicates with the lumen of the functional penis papilla. The latter is a stubby, cone-shaped structure that has a more or less horizontal orientation and projects into the atrial lumen (
<figureCitation id="30F42A25FFC7FFA7FC46A349FC2C0323" box="[908,977,332,356]" captionStart="Figure 11" captionStartId="16.[129,194,1954,1978]" captionTargetBox="[160,1443,1377,1926]" captionTargetId="figure-355@16.[160,1443,1377,1926]" captionTargetPageId="16" captionText="Figure 11. Dugesia insolita. Holotype RMNH.VER.21536.1, sagittal reconstruction of the copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811369" httpUri="https://zenodo.org/record/14811369/files/figure.png" pageId="16" pageNumber="17">Fig. 11</figureCitation>
). At about one-third of the distance between its root and the tip, the papilla exhibits a fold or outbulging, thus providing the papilla—in sagittal view—with a hastate shape or the form of a head of a spear.
</paragraph>
<paragraph id="A87036A0FFC7FFA5FC9FA3CCFDF00341" blockId="16.[825,1476,144,669]" lastBlockId="18.[127,780,144,1797]" lastPageId="18" lastPageNumber="19" pageId="16" pageNumber="17">
Erythrophil penis glands discharge through the lining epithelium of the functional penis papilla, which is surrounded by the funnel-shaped portion of a sclerotic spermatophore (
<figureCitation id="30F42A25FFC7FFA7FAA5A00DFA410067" box="[1391,1468,520,544]" captionStart="Figure 11" captionStartId="16.[129,194,1954,1978]" captionTargetBox="[160,1443,1377,1926]" captionTargetId="figure-355@16.[160,1443,1377,1926]" captionTargetPageId="16" captionText="Figure 11. Dugesia insolita. Holotype RMNH.VER.21536.1, sagittal reconstruction of the copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811369" httpUri="https://zenodo.org/record/14811369/files/figure.png" pageId="16" pageNumber="17">Figs 11</figureCitation>
,
<figureCitation id="30F42A25FFC7FFA7FCF3A022FCAE0078" box="[825,851,551,575]" captionStart="Figure 12" captionStartId="17.[114,179,1834,1858]" captionTargetBox="[205,1368,142,1806]" captionTargetId="figure-6@17.[205,1368,142,1806]" captionTargetPageId="17" captionText="Figure 12. Dugesia insolita. Holotype RMNH.VER.21536.1, sagittal sections.A, male and female—partly—copulatory apparatus; B, detail of male copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811371" httpUri="https://zenodo.org/record/14811371/files/figure.png" pageId="16" pageNumber="17">12</figureCitation>
). It is here surmised that the secretion of these penial glands contributed to the formation of the spermatophore, in the same way that normally in
<taxonomicName id="6FCF4D23FFC7FFA7FBDCA062FB4F0039" box="[1046,1202,614,638]" pageId="16" pageNumber="17">
<emphasis id="9ABBEAB2FFC7FFA7FBDCA062FB990039" box="[1046,1124,615,638]" italics="true" pageId="16" pageNumber="17">Dugesia</emphasis>
species
</taxonomicName>
the spermatophore emanates from the penial diaphragm (see below). Therefore, it is here also surmised that the presumed penis papilla of
<taxonomicName id="6FCF4D23FFC5FFA5FDA5A294FD2C02EF" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[623,721,144,168]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC5FFA5FDA5A294FD2C02EF" box="[623,721,144,168]" italics="true" pageId="18" pageNumber="19">D. insolita</emphasis>
</taxonomicName>
is actually homologous to the diaphragm (see below). Therefore, in the above and below we refer to the papilla as the
<emphasis id="9ABBEAB2FFC5FFA5FDA6A2CAFD3002A0" box="[620,717,207,231]" italics="true" pageId="18" pageNumber="19">functional</emphasis>
penis papilla instead of the true penis papilla.
</paragraph>
<caption id="FCB06628FFC7FFA7FF4BA71FFAB60775" ID-DOI="http://doi.org/10.5281/zenodo.14811367" ID-Zenodo-Dep="14811367" box="[129,1355,1306,1330]" httpUri="https://zenodo.org/record/14811367/files/figure.png" pageId="16" pageNumber="17" startId="16.[129,194,1306,1330]" targetBox="[129,1473,773,1277]" targetPageId="16" targetType="figure">
<paragraph id="A87036A0FFC7FFA7FF4BA71FFAB60775" blockId="16.[129,1355,1306,1330]" box="[129,1355,1306,1330]" pageId="16" pageNumber="17">
<emphasis id="9ABBEAB2FFC7FFA7FF4BA71FFF180775" bold="true" box="[129,229,1306,1330]" pageId="16" pageNumber="17">Figure 10.</emphasis>
<taxonomicName id="6FCF4D23FFC7FFA7FF23A71FFE8A0775" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[233,375,1306,1330]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="16" pageNumber="17" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC7FFA7FF23A71FFE8A0775" box="[233,375,1306,1330]" italics="true" pageId="16" pageNumber="17">Dugesia insolita</emphasis>
</taxonomicName>
. Holotype RMNH.VER.21536.1, photomicrographs, sagittal sections. A, pharynx; B, hyperplasic ovary.
</paragraph>
</caption>
<caption id="FCB06628FFC7FFA7FF4BA5A7FAAF05FC" ID-DOI="http://doi.org/10.5281/zenodo.14811369" ID-Zenodo-Dep="14811369" box="[129,1362,1954,1979]" httpUri="https://zenodo.org/record/14811369/files/figure.png" pageId="16" pageNumber="17" startId="16.[129,194,1954,1978]" targetBox="[160,1443,1377,1926]" targetPageId="16" targetType="figure">
<paragraph id="A87036A0FFC7FFA7FF4BA5A7FAAF05FC" blockId="16.[129,1362,1954,1979]" box="[129,1362,1954,1979]" pageId="16" pageNumber="17">
<emphasis id="9ABBEAB2FFC7FFA7FF4BA5A7FF1805FC" bold="true" box="[129,229,1954,1979]" pageId="16" pageNumber="17">Figure 11.</emphasis>
<taxonomicName id="6FCF4D23FFC7FFA7FF23A5A6FE8A05FC" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[233,375,1955,1979]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="16" pageNumber="17" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC7FFA7FF23A5A6FE8A05FC" box="[233,375,1955,1979]" italics="true" pageId="16" pageNumber="17">Dugesia insolita</emphasis>
</taxonomicName>
. Holotype RMNH.VER.21536.1, sagittal reconstruction of the copulatory apparatus; anterior to the left.
</paragraph>
</caption>
<caption id="FCB06628FFC6FFA6FFB8A52FFE8C0519" ID-DOI="http://doi.org/10.5281/zenodo.14811371" ID-Zenodo-Dep="14811371" httpUri="https://zenodo.org/record/14811371/files/figure.png" pageId="17" pageNumber="18" startId="17.[114,179,1834,1858]" targetBox="[205,1368,142,1806]" targetPageId="17" targetType="figure">
<paragraph id="A87036A0FFC6FFA6FFB8A52FFE8C0519" blockId="17.[114,1444,1834,1886]" pageId="17" pageNumber="18">
<emphasis id="9ABBEAB2FFC6FFA6FFB8A52FFF2B0505" bold="true" box="[114,214,1834,1858]" pageId="17" pageNumber="18">Figure 12.</emphasis>
<taxonomicName id="6FCF4D23FFC6FFA6FF11A52EFE940505" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[219,361,1834,1858]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="17" pageNumber="18" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC6FFA6FF11A52EFE940505" box="[219,361,1834,1858]" italics="true" pageId="17" pageNumber="18">Dugesia insolita</emphasis>
</taxonomicName>
. Holotype RMNH.VER. 21536.1, sagittal sections. A, male and female—partly—copulatory apparatus; B, detail of male copulatory apparatus.
</paragraph>
</caption>
<paragraph id="A87036A0FFC5FFA5FF56A308FDD003C4" blockId="18.[127,780,144,1797]" pageId="18" pageNumber="19">
The large penis bulb is strongly muscular and consists of intermingled circular and longitudinal muscle fibres. From the outer surface of the penis bulb longitudinal fibres run posteriad and attach to the posterior atrial fold (
<figureCitation id="30F42A25FFC5FFA5FE1DA36EFDE103C4" box="[471,540,363,387]" captionStart="Figure 11" captionStartId="16.[129,194,1954,1978]" captionTargetBox="[160,1443,1377,1926]" captionTargetId="figure-355@16.[160,1443,1377,1926]" captionTargetPageId="16" captionText="Figure 11. Dugesia insolita. Holotype RMNH.VER.21536.1, sagittal reconstruction of the copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811369" httpUri="https://zenodo.org/record/14811369/files/figure.png" pageId="18" pageNumber="19">Fig. 11</figureCitation>
).
</paragraph>
<paragraph id="A87036A0FFC5FFA5FF56A38EFD66013E" blockId="18.[127,780,144,1797]" pageId="18" pageNumber="19">
The major part of the male atrium is lined with a tall epithelium, which, however, becomes very thin on its most anterior section near the base of the functional penis papilla. The atrium is surrounded by a subepithelial layer of circular muscles that is well-developed on its anterior section but rather thin on more posterior portions. The male atrium is provided with three annular folds or constrictions. One of the folds envelopes the root of the functional penis papilla, a second fold is located just anterior to the tip of the papilla, while the third, posterior fold is positioned at the posterior end of the male atrium. This third fold may indeed be considered as the separation between the male and the female atrium, since the openings of shell glands penetrate the dorsal epithelium of that part of the atrium that lies immediately posterior to this third fold. Basically, the openings of erythrophilic shell glands surround the opening of the bursal canal through the roof of the female atrium (
<figureCitation id="30F42A25FFC5FFA5FD8FA164FD77013E" box="[581,650,865,889]" captionStart="Figure 11" captionStartId="16.[129,194,1954,1978]" captionTargetBox="[160,1443,1377,1926]" captionTargetId="figure-355@16.[160,1443,1377,1926]" captionTargetPageId="16" captionText="Figure 11. Dugesia insolita. Holotype RMNH.VER.21536.1, sagittal reconstruction of the copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811369" httpUri="https://zenodo.org/record/14811369/files/figure.png" pageId="18" pageNumber="19">Fig. 11</figureCitation>
).
</paragraph>
<paragraph id="A87036A0FFC5FFA5FF56A185FE9D0672" blockId="18.[127,780,144,1797]" pageId="18" pageNumber="19">
The highly hyperplasic ovaries extend from ~800 μm behind the eyes and the brain to a little posterior (~700 μm) to the level of the root of the pharynx, occupying almost the entire dorsoventral space of the body (
<figureCitation id="30F42A25FFC5FFA5FE45A1DBFE1F01B1" box="[399,482,990,1014]" captionStart="Figure 10" captionStartId="16.[129,194,1306,1330]" captionTargetBox="[129,1473,773,1277]" captionTargetId="figure-339@16.[129,1473,773,1277]" captionTargetPageId="16" captionText="Figure 10. Dugesia insolita. Holotype RMNH.VER.21536.1, photomicrographs, sagittal sections. A, pharynx; B, hyperplasic ovary." figureDoi="http://doi.org/10.5281/zenodo.14811367" httpUri="https://zenodo.org/record/14811367/files/figure.png" pageId="18" pageNumber="19">Fig. 10B</figureCitation>
). The oocytes clearly exhibit a degenerative condition, as evidenced by vacuolization of their nuclei and cytoplasm.
</paragraph>
<paragraph id="A87036A0FFC5FFA5FF56A639FCF907CA" blockId="18.[127,780,144,1797]" pageId="18" pageNumber="19">The infranucleated oviducts run immediately dorsal to the ventral nerve cords; at the level of the gonopore the ducts turn medially to open—separately—into the most proximal section of the bursal canal, close to the opening of the latter through the roof of the female atrium. From the point of this opening, the bursal canal at first runs vertically towards the dorsal body surface and then exhibits a knee-shaped, anteriorly-directed bend, after which it follows its course more or less parallel to the dorsal and ventral body surfaces. The bursal canal communicates with a more or less ellipsoidal copulatory bursa that is located at a short distance behind the pharyngeal pocket and the mouth opening.</paragraph>
<paragraph id="A87036A0FFC5FFA5FF56A790FDDA0480" blockId="18.[127,780,144,1797]" pageId="18" pageNumber="19">The bursal canal is lined with a tall, infranucleated epithelium. The long horizontal portion of the bursal canal is surrounded by a single, subepithelial layer of longitudinal muscles, followed by a slightly thicker layer of circular muscles. The short, vertical portion of the bursal canal is surrounded by a thin, subepithelial layer of circular muscles, followed by a layer of obliquely oriented fibres. An extra outer layer of longitudinal muscles forms a poorly developed ectal reinforcement layer that runs from about the knee-shaped bend in the bursal canal to about halfway between the bend and the copulatory bursa.</paragraph>
<paragraph id="A87036A0FFC5FFA5FF56A4CBFD490542" blockId="18.[127,780,144,1797]" pageId="18" pageNumber="19">Very abundant and conspicuous cement glands discharge their orange-staining secretion into the short gonoduct.</paragraph>
<paragraph id="A87036A0FFC5FFA5FE59A520FE0B0578" blockId="18.[129,778,1829,1986]" box="[403,502,1829,1855]" pageId="18" pageNumber="19">
<emphasis id="9ABBEAB2FFC5FFA5FE59A520FE0B0578" box="[403,502,1829,1855]" italics="true" pageId="18" pageNumber="19">Karyology</emphasis>
</paragraph>
<paragraph id="A87036A0FFC5FFA5FF4BA54EFC460046" blockId="18.[129,778,1829,1986]" lastBlockId="18.[825,1476,144,669]" pageId="18" pageNumber="19">
Metaphasic plates revealed that specimens constantly showed a set of 18 chromosomes. Chromosomes from six metaphasic plates could be arranged, according to their length, into nine groups of two chromosomes with a diploid chromosome set of 2n = 18; n = 9. The analysis within each pair of chromosomes revealed uniformity in both length and centromeric position, except for chromosome 1, in which the first element is always longer than its homologue. The first two chromosomes were clearly identifiable. Instead, from the third to the ninth pair, the differences in length were so small that contiguous chromosomes could be overlapping (such as chromosomes 3 and 5, 6 and 7, 8 and 9). Chromosomal length decreased gradually, with low standard deviation values. In contrast, the centromeric indices showed great variation, resulting in high standard deviations, as, for example, for chromosome numbers 4 and 6 (
<figureCitation id="30F42A25FFC5FFA5FA56A3CCFCAE0046" captionStart="Figure 13" captionStartId="19.[115,180,1452,1476]" captionTargetBox="[146,1411,149,1421]" captionTargetId="figure-115@19.[143,1429,146,1424]" captionTargetPageId="19" captionText="Figure 13.Dugesia insolita. A, metaphasic plate; B, karyogram; C, idiogram." figureDoi="http://doi.org/10.5281/zenodo.14811373" httpUri="https://zenodo.org/record/14811373/files/figure.png" pageId="18" pageNumber="19">Fig. 13</figureCitation>
;
<tableCitation id="E54D031BFFC5FFA5FCAAA3ECFC560046" box="[864,939,489,513]" captionStart="Table 3" captionStartId="19.[113,168,1523,1547]" captionTargetPageId="19" captionText="Table 3. Dugesia insolita. Mean values and standard deviations of the relative lengths (r.l.) and centromeric indices (c.i.) of the haploid set of chromosomes." httpUri="http://table.plazi.org/id/FCB06628FFC4FFA4FFBBA7F6FF030460" pageId="18" pageNumber="19" tableUuid="FCB06628FFC4FFA4FFBBA7F6FF030460">Table 3</tableCitation>
).
</paragraph>
<paragraph id="A87036A0FFC5FFA5FC9FA00DFAE000DA" blockId="18.[825,1476,144,669]" pageId="18" pageNumber="19">
The karyometric data indicate that the diploid chromosome complement is characterized by metacentric heterobrachial chromosomes, with the exceptions of chromosome 3, which is submetacentric, and chromosomes 1 and 9 that are metacentric, bordering on metacentric isobrachial (
<tableCitation id="E54D031BFFC5FFA5FB0BA080FAF100DA" box="[1217,1292,645,669]" captionStart="Table 3" captionStartId="19.[113,168,1523,1547]" captionTargetPageId="19" captionText="Table 3. Dugesia insolita. Mean values and standard deviations of the relative lengths (r.l.) and centromeric indices (c.i.) of the haploid set of chromosomes." httpUri="http://table.plazi.org/id/FCB06628FFC4FFA4FFBBA7F6FF030460" pageId="18" pageNumber="19" tableUuid="FCB06628FFC4FFA4FFBBA7F6FF030460">Table 3</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFC5FFA3FBDAA0B8FB4C0613" lastPageId="20" lastPageNumber="21" pageId="18" pageNumber="19" type="discussion">
<paragraph id="A87036A0FFC5FFA5FBDAA0B8FB160090" blockId="18.[824,1476,701,1985]" box="[1040,1259,701,727]" pageId="18" pageNumber="19">
<emphasis id="9ABBEAB2FFC5FFA5FBDAA0B8FB160090" box="[1040,1259,701,727]" italics="true" pageId="18" pageNumber="19">Comparative discussion</emphasis>
</paragraph>
<paragraph id="A87036A0FFC5FFA5FCF3A0E1FB4601DF" blockId="18.[824,1476,701,1985]" pageId="18" pageNumber="19">
<taxonomicName id="6FCF4D23FFC5FFA5FCF3A0E1FC2F00BC" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[825,978,739,763]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC5FFA5FCF3A0E1FC2F00BC" box="[825,978,739,763]" italics="true" pageId="18" pageNumber="19">Dugesia insolita</emphasis>
</taxonomicName>
is included in a phylogenetic clade encompassing all long-eared
<taxonomicName id="6FCF4D23FFC5FFA5FC04A106FBE1015D" authorityName="Girard" authorityYear="1850" box="[974,1052,771,794]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC5FFA5FC04A106FBE1015D" box="[974,1052,771,794]" italics="true" pageId="18" pageNumber="19">Dugesia</emphasis>
</taxonomicName>
from
<collectingCountry id="D0D87630FFC5FFA5FB91A106FB29015C" box="[1115,1236,771,795]" name="Madagascar" pageId="18" pageNumber="19">Madagascar</collectingCountry>
with full support (
<figureCitation id="30F42A25FFC5FFA5FA56A106FCBB017D" captionStart="Figure 2" captionStartId="8.[128,193,1171,1195]" captionTargetBox="[131,1470,145,1140]" captionTargetId="figure-382@8.[129,1473,148,1143]" captionTargetPageId="8" captionText="Figure 2. Maximum likelihood molecular phylogenetic tree obtained from the concatenated dataset (COI, ITS1, 18S, and 28S) using the program IQ-tree 2.2.2.6. Supports on each node correspond with the supports of bootstrap (BS) of the maximum likelihood-based RaxML and IQ-tree programs, and supports of posterior probability (PP) of the Bayesian inference-based program MrBayes, respectively. Dashes () indicate those nodes not recovered in the corresponding approaches. Black squares on some nodes show full support for all three approaches. Scale bar: substitutions per position." figureDoi="http://doi.org/10.5281/zenodo.14811339" httpUri="https://zenodo.org/record/14811339/files/figure.png" pageId="18" pageNumber="19">Fig. 2</figureCitation>
). However, none of the phylogenetic approaches lent good support for the relationship of
<taxonomicName id="6FCF4D23FFC5FFA5FB45A147FB05011E" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[1167,1272,833,857]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC5FFA5FB45A147FB05011E" box="[1167,1272,833,857]" italics="true" pageId="18" pageNumber="19">D. insolita</emphasis>
</taxonomicName>
to the rest of the long-eared populations in this lineage. Thus, its precise position within this group remains unresolved.
</paragraph>
<paragraph id="A87036A0FFC5FFA5FC9FA19AFC480652" blockId="18.[824,1476,701,1985]" pageId="18" pageNumber="19">
The
<specimenCount id="BEC9FD29FFC5FFA5FC4AA1A5FBEA01F0" box="[896,1047,927,951]" count="2" pageId="18" pageNumber="19" type="generic">two specimens</specimenCount>
from the same population of
<taxonomicName id="6FCF4D23FFC5FFA5FAF6A1A5FA6301F0" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[1340,1438,927,951]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC5FFA5FAF6A1A5FA6301F0" box="[1340,1438,927,951]" italics="true" pageId="18" pageNumber="19">D. insolita</emphasis>
</taxonomicName>
are clustered together with full support in both ML and BI phylogenetic trees, indicating that they most probably also belong to this species.
</paragraph>
<paragraph id="A87036A0FFC5FFA5FC9FA618FBFA0696" blockId="18.[824,1476,701,1985]" pageId="18" pageNumber="19">
Sperm transfer by means of a sclerotic spermatophore is a common phenomenon among freshwater planarians (
<bibRefCitation id="CC5E4B51FFC5FFA5FA46A639FC900634" author="Sluys R" pageId="18" pageNumber="19" pagination="3 - 25" refId="ref18508" refString="Sluys R. Phylogenetic relationships of the triclads (Platyhelminthes, Seriata, Tricladida). Bijdragen tot de Dierkunde 1989; 59: 3 - 25. https: // doi. org / 10.1163 / 26660644 - 05901001" type="journal article" year="1989">Sluys 1989</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC5FFA5FCB6A65EFB8F0634" author="Sluys R &amp; Riutort M" box="[892,1138,1115,1139]" pageId="18" pageNumber="19" pagination="1 - 56" refId="ref18575" refString="Sluys R, Riutort M. Planarian diversity and phylogeny. In: Rink JC (ed.), Planarian Regeneration: Methods and Protocols. Methods in Molecular Biology, Vol. 1774. New York: Springer, 2018, 1 - 56. https: // doi. org / 10.1007 / 978 - 1 - 4939 - 7802 - 1 _ 1" type="book chapter" year="2018">Sluys and Riutort 2018</bibRefCitation>
). Such spermatophores may be simply sac-shaped or may possess a long, narrow, and hollow stalk, the latter situation being the case in species of
<taxonomicName id="6FCF4D23FFC5FFA5FAA5A69EFA4006F5" authorityName="Girard" authorityYear="1850" box="[1391,1469,1179,1202]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC5FFA5FAA5A69EFA4006F5" box="[1391,1469,1179,1202]" italics="true" pageId="18" pageNumber="19">Dugesia</emphasis>
</taxonomicName>
, including
<taxonomicName id="6FCF4D23FFC5FFA5FC55A6BFFBFF0696" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[927,1026,1209,1233]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC5FFA5FC55A6BFFBFF0696" box="[927,1026,1209,1233]" italics="true" pageId="18" pageNumber="19">D. insolita</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A87036A0FFC5FFA5FC9FA6DCFB93042E" blockId="18.[824,1476,701,1985]" pageId="18" pageNumber="19">
Spermatophores are formed in the ejaculatory duct running through the penis papilla, the secretions of various penial glands contributing to the formation of the spermatophore. In species of
<emphasis id="9ABBEAB2FFC5FFA5FC9FA732FC550709" box="[853,936,1335,1358]" italics="true" pageId="18" pageNumber="19">
<taxonomicName id="6FCF4D23FFC5FFA5FC9FA732FC590709" authorityName="Girard" authorityYear="1850" box="[853,932,1335,1358]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="genus">Dugesia</taxonomicName>
,
</emphasis>
the diaphragm in the proximal section of the ejaculatory duct makes a different contribution to the formation of the spermatophore as compared with the middle and distal parts of the duct. The secretions discharged through the tip of the lining epithelium of the conus of the diaphragm form an antiagglutination layer at the inner surface of the stalk of the spermatophore, while the actual wall of the spermatophore is formed by secretions discharged through a more basal, anterior portion of the diaphragm, as well as the most proximal part of the ejaculatory duct (
<bibRefCitation id="CC5E4B51FFC5FFA5FC72A454FBA3042E" author="Vreys C &amp; Steffanie N &amp; Gevaerts H" box="[952,1118,1617,1641]" pageId="18" pageNumber="19" pagination="24 - 34" refId="ref19553" refString="Vreys C, Steffanie N, Gevaerts H. Spermatophore formation and transfer in the freshwater flatworm Dugesia gonocephala (Platyhelminthes, Tricladida, Paludicola). Invertebrate Biology 1999; 118: 24 - 34. https: // doi. org / 10.2307 / 3226909" type="journal article" year="1999">
Vreys
<emphasis id="9ABBEAB2FFC5FFA5FC3CA454FBE3042E" box="[1014,1054,1617,1641]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
. 1999
</bibRefCitation>
).
</paragraph>
<paragraph id="A87036A0FFC5FFA3FC9FA475FE810341" blockId="18.[824,1476,701,1985]" lastBlockId="20.[127,780,144,1985]" lastPageId="20" lastPageNumber="21" pageId="18" pageNumber="19">
This construction method results in the situation that during the formation process the funnel-shaped basal portion of the stalk of the spermatophore envelopes the diaphragm (cf.
<bibRefCitation id="CC5E4B51FFC5FFA5FA41A4AAFC5804A1" author="Vreys C &amp; Schockaert ER &amp; Michiels NK" pageId="18" pageNumber="19" pagination="1479 - 86" refId="ref19498" refString="Vreys C, Schockaert ER, Michiels NK. Formation, transfer, and assimilation of the spermatophore of the hermaphroditic flatworm Dugesia gonocephala (Tricladida, Paludicola). Canadian Journal of Zoology 1997; 75: 1479 - 86. https: // doi. org / 10.1139 / z 97 - 771" type="journal article" year="1997">
Vreys
<emphasis id="9ABBEAB2FFC5FFA5FCF3A4CAFC9904A1" box="[825,868,1742,1766]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
. 1997
</bibRefCitation>
: fig. 2; 1999: fig. 4), a situation that, over the years, we have regularly observed in histological preparations of species of the genus
<taxonomicName id="6FCF4D23FFC5FFA5FC3CA508FBB90563" authorityName="Girard" authorityYear="1850" box="[1014,1092,1805,1828]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC5FFA5FC3CA508FBB90563" box="[1014,1092,1805,1828]" italics="true" pageId="18" pageNumber="19">Dugesia</emphasis>
</taxonomicName>
. A similar situation is present in
<taxonomicName id="6FCF4D23FFC5FFA5FA61A508FC7D0503" authorityName="Stocchino &amp; Sluys" authorityYear="2024" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="18" pageNumber="19" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC5FFA5FA61A508FC7D0503" italics="true" pageId="18" pageNumber="19">D. insolita</emphasis>
</taxonomicName>
(
<figureCitation id="30F42A25FFC5FFA5FC5EA529FC1C0503" box="[916,993,1836,1860]" captionStart="Figure 11" captionStartId="16.[129,194,1954,1978]" captionTargetBox="[160,1443,1377,1926]" captionTargetId="figure-355@16.[160,1443,1377,1926]" captionTargetPageId="16" captionText="Figure 11. Dugesia insolita. Holotype RMNH.VER.21536.1, sagittal reconstruction of the copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811369" httpUri="https://zenodo.org/record/14811369/files/figure.png" pageId="18" pageNumber="19">Figs 11</figureCitation>
,
<figureCitation id="30F42A25FFC5FFA5FC3BA529FBF60503" box="[1009,1035,1836,1860]" captionStart="Figure 12" captionStartId="17.[114,179,1834,1858]" captionTargetBox="[205,1368,142,1806]" captionTargetId="figure-6@17.[205,1368,142,1806]" captionTargetPageId="17" captionText="Figure 12. Dugesia insolita. Holotype RMNH.VER.21536.1, sagittal sections.A, male and female—partly—copulatory apparatus; B, detail of male copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811371" httpUri="https://zenodo.org/record/14811371/files/figure.png" pageId="18" pageNumber="19">12</figureCitation>
) in which the funnel-shaped base of the spermatophore covers a cone-shaped structure that resembles a short penis papilla, the covering epithelium of which is penetrated by the openings of erythrophil penial glands that contribute to the formation of the stalk of the spermatophore. We have never encountered the situation that in species of
<taxonomicName id="6FCF4D23FFC4FFA4FD64A512FD010569" authorityName="Girard" authorityYear="1850" box="[686,764,1815,1838]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="19" pageNumber="20" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC4FFA4FD64A512FD010569" box="[686,764,1815,1838]" italics="true" pageId="19" pageNumber="20">Dugesia</emphasis>
</taxonomicName>
the penis papilla contributed in this way to the formation of the spermatophore, as the latter is always formed within the ejaculatory duct. Therefore, it is here surmised (see above) that this cone-shaped, penis-like structure is actually homologous to the diaphragm in other species of
<taxonomicName id="6FCF4D23FFC4FFA4FBA2A512FB4B0569" authorityName="Girard" authorityYear="1850" box="[1128,1206,1815,1838]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="19" pageNumber="20" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC4FFA4FBA2A512FB4B0569" box="[1128,1206,1815,1838]" italics="true" pageId="19" pageNumber="20">Dugesia</emphasis>
</taxonomicName>
—as it is without doubt that
<taxonomicName id="6FCF4D23FFC4FFA4FC9DA533FC47050A" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[855,954,1845,1869]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="19" pageNumber="20" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC4FFA4FC9DA533FC47050A" box="[855,954,1845,1869]" italics="true" pageId="19" pageNumber="20">D. insolita</emphasis>
</taxonomicName>
belongs to this genus (see molecular results)—albeit it may function as a penis papilla. It should be noted that the conspicuous fold projecting into the lumen of the functional penis papilla and through which runs a minute duct, cannot be seen as phylogenetically homologous to the diaphragm, nor as being functionally equivalent to it, since it does not receive the secretion of penial glands and does not have any connection with the spermatophore.
</paragraph>
<caption id="FCB06628FFC4FFA4FFB9A7A9FCCA0783" ID-DOI="http://doi.org/10.5281/zenodo.14811373" ID-Zenodo-Dep="14811373" box="[115,823,1452,1477]" httpUri="https://zenodo.org/record/14811373/files/figure.png" pageId="19" pageNumber="20" startId="19.[115,180,1452,1476]" targetBox="[146,1411,149,1421]" targetPageId="19" targetType="figure">
<paragraph id="A87036A0FFC4FFA4FFB9A7A9FCCA0783" blockId="19.[115,823,1452,1477]" box="[115,823,1452,1477]" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FFB9A7A9FF2A0783" bold="true" box="[115,215,1452,1476]" pageId="19" pageNumber="20">Figure 13.</emphasis>
<taxonomicName id="6FCF4D23FFC4FFA4FF10A7A8FE950783" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[218,360,1452,1476]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="19" pageNumber="20" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC4FFA4FF10A7A8FE950783" box="[218,360,1452,1476]" italics="true" pageId="19" pageNumber="20">Dugesia insolita</emphasis>
</taxonomicName>
. A, metaphasic plate; B, karyogram; C, idiogram.
</paragraph>
</caption>
<caption id="FCB06628FFC4FFA4FFBBA7F6FF030460" ID-Table-UUID="FCB06628FFC4FFA4FFBBA7F6FF030460" httpUri="http://table.plazi.org/id/FCB06628FFC4FFA4FFBBA7F6FF030460" pageId="19" pageNumber="20" startId="19.[113,168,1523,1547]" targetBox="[113,1459,1596,1746]" targetIsTable="true" targetPageId="19" targetType="table">
<paragraph id="A87036A0FFC4FFA4FFBBA7F6FF030460" blockId="19.[113,1423,1523,1575]" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FFBBA7F6FF42044C" bold="true" box="[113,191,1523,1547]" pageId="19" pageNumber="20">Table 3.</emphasis>
<taxonomicName id="6FCF4D23FFC4FFA4FF0EA7F1FEAF044C" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[196,338,1523,1547]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="19" pageNumber="20" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC4FFA4FF0EA7F1FEAF044C" box="[196,338,1523,1547]" italics="true" pageId="19" pageNumber="20">Dugesia insolita</emphasis>
</taxonomicName>
. Mean values and standard deviations of the relative lengths (r.l.) and centromeric indices (c.i.) of the haploid set of chromosomes.
</paragraph>
</caption>
<paragraph id="A87036A0FFC4FFA4FD2DA439FA4E0495" pageId="19" pageNumber="20">
<table id="DACFC400FFC40048FFBBA439FA4E0495" box="[113,1459,1596,1746]" gridcols="10" gridrows="4" pageId="19" pageNumber="20">
<tr id="16FF34E2FFC40048FFBBA439FA4E0413" box="[113,1459,1596,1620]" gridrow="0" pageId="19" pageNumber="20">
<th id="552E5D9EFFC40048FFBBA439FA4E0413" box="[113,1459,1596,1620]" colspan="10" colspanRight="9" gridcol="0" gridrow="0" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FD2DA439FC8D0413" bold="true" box="[743,880,1596,1620]" pageId="19" pageNumber="20">Chromosome</emphasis>
</th>
</tr>
<tr id="16FF34E2FFC40048FFBBA46DFA4E04C6" box="[113,1459,1640,1665]" gridrow="1" pageId="19" pageNumber="20" rowspan-0="1">
<td id="552E5D9EFFC40048FF6EA46DFEE104C6" box="[164,284,1640,1665]" gridcol="1" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FF10A46DFF1A04C7" bold="true" box="[218,231,1640,1664]" pageId="19" pageNumber="20">1</emphasis>
</td>
<td id="552E5D9EFFC40048FEFEA46DFE5004C6" box="[308,429,1640,1665]" gridcol="2" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FEA1A46DFE8504C7" bold="true" box="[363,376,1640,1664]" pageId="19" pageNumber="20">2</emphasis>
</td>
<td id="552E5D9EFFC40048FE0FA46DFDC304C6" box="[453,574,1640,1665]" gridcol="3" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FE31A46DFDF504C7" bold="true" box="[507,520,1640,1664]" pageId="19" pageNumber="20">3</emphasis>
</td>
<td id="552E5D9EFFC40048FD9CA46DFD3204C6" box="[598,719,1640,1665]" gridcol="4" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FD46A46DFD6404C7" bold="true" box="[652,665,1640,1664]" pageId="19" pageNumber="20">4</emphasis>
</td>
<td id="552E5D9EFFC40048FD2DA46DFCA204C6" box="[743,863,1640,1665]" gridcol="5" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FCEFA46CFCCF04C6" bold="true" box="[805,818,1641,1665]" pageId="19" pageNumber="20">5</emphasis>
</td>
<td id="552E5D9EFFC40048FC4DA46DFBFD04C6" box="[903,1024,1640,1665]" gridcol="6" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FC74A46DFC3604C7" bold="true" box="[958,971,1640,1664]" pageId="19" pageNumber="20">6</emphasis>
</td>
<td id="552E5D9EFFC40048FBD2A46DFB6C04C6" box="[1048,1169,1640,1665]" gridcol="7" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FB84A46CFBA604C6" bold="true" box="[1102,1115,1641,1665]" pageId="19" pageNumber="20">7</emphasis>
</td>
<td id="552E5D9EFFC40048FB63A46DFADF04C6" box="[1193,1314,1640,1665]" gridcol="8" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FB15A46DFB1104C7" bold="true" box="[1247,1260,1640,1664]" pageId="19" pageNumber="20">8</emphasis>
</td>
<td id="552E5D9EFFC40048FAF0A46DFA4E04C6" box="[1338,1459,1640,1665]" gridcol="9" gridrow="1" pageId="19" pageNumber="20">
<emphasis id="9ABBEAB2FFC4FFA4FABAA46DFA8004C7" bold="true" box="[1392,1405,1640,1664]" pageId="19" pageNumber="20">9</emphasis>
</td>
</tr>
<tr id="16FF34E2FFC40048FFBBA49DFA4E04F7" box="[113,1459,1688,1712]" gridrow="2" pageId="19" pageNumber="20">
<th id="552E5D9EFFC40048FFBBA49DFF7004F7" box="[113,141,1688,1712]" gridcol="0" gridrow="2" pageId="19" pageNumber="20">r.l.</th>
<td id="552E5D9EFFC40048FF6EA49DFEE104F7" box="[164,284,1688,1712]" gridcol="1" gridrow="2" pageId="19" pageNumber="20">15.15 ± 0.74</td>
<td id="552E5D9EFFC40048FEFEA49DFE5004F7" box="[308,429,1688,1712]" gridcol="2" gridrow="2" pageId="19" pageNumber="20">13.07 ± 0.40</td>
<td id="552E5D9EFFC40048FE0FA49DFDC304F7" box="[453,574,1688,1712]" gridcol="3" gridrow="2" pageId="19" pageNumber="20">11.58 ± 0.60</td>
<td id="552E5D9EFFC40048FD9CA49DFD3204F7" box="[598,719,1688,1712]" gridcol="4" gridrow="2" pageId="19" pageNumber="20">11.29 ± 0.32</td>
<td id="552E5D9EFFC40048FD2DA49DFCA204F7" box="[743,863,1688,1712]" gridcol="5" gridrow="2" pageId="19" pageNumber="20">10.90 ± 0.52</td>
<td id="552E5D9EFFC40048FC4DA49DFBFD04F7" box="[903,1024,1688,1712]" gridcol="6" gridrow="2" pageId="19" pageNumber="20">10.25 ± 0.29</td>
<td id="552E5D9EFFC40048FBD2A49DFB6C04F7" box="[1048,1169,1688,1712]" gridcol="7" gridrow="2" pageId="19" pageNumber="20">9.88 ± 0.27</td>
<td id="552E5D9EFFC40048FB63A49DFADF04F7" box="[1193,1314,1688,1712]" gridcol="8" gridrow="2" pageId="19" pageNumber="20">9.28 ± 0.34</td>
<td id="552E5D9EFFC40048FAF0A49DFA4E04F7" box="[1338,1459,1688,1712]" gridcol="9" gridrow="2" pageId="19" pageNumber="20">8.84 ± 0.40</td>
</tr>
<tr id="16FF34E2FFC40048FFBBA4BFFA4E0495" box="[113,1459,1722,1746]" gridrow="3" pageId="19" pageNumber="20">
<th id="552E5D9EFFC40048FFBBA4BFFF700495" box="[113,141,1722,1746]" gridcol="0" gridrow="3" pageId="19" pageNumber="20">c.i.</th>
<td id="552E5D9EFFC40048FF6EA4BFFEE10495" box="[164,284,1722,1746]" gridcol="1" gridrow="3" pageId="19" pageNumber="20">48.93 ± 1.04</td>
<td id="552E5D9EFFC40048FEFEA4BFFE500495" box="[308,429,1722,1746]" gridcol="2" gridrow="3" pageId="19" pageNumber="20">42.55 ± 1.94</td>
<td id="552E5D9EFFC40048FE0FA4BFFDC30495" box="[453,574,1722,1746]" gridcol="3" gridrow="3" pageId="19" pageNumber="20">35.90 ± 1.33</td>
<td id="552E5D9EFFC40048FD9CA4BFFD320495" box="[598,719,1722,1746]" gridcol="4" gridrow="3" pageId="19" pageNumber="20">47.43 ± 3.76</td>
<td id="552E5D9EFFC40048FD2DA4BFFCA20495" box="[743,863,1722,1746]" gridcol="5" gridrow="3" pageId="19" pageNumber="20">45.94 ± 1.15</td>
<td id="552E5D9EFFC40048FC4DA4BFFBFD0495" box="[903,1024,1722,1746]" gridcol="6" gridrow="3" pageId="19" pageNumber="20">46.50 ± 3.97</td>
<td id="552E5D9EFFC40048FBD2A4BFFB6C0495" box="[1048,1169,1722,1746]" gridcol="7" gridrow="3" pageId="19" pageNumber="20">48.32 ± 2.31</td>
<td id="552E5D9EFFC40048FB63A4BFFADF0495" box="[1193,1314,1722,1746]" gridcol="8" gridrow="3" pageId="19" pageNumber="20">47.56 ± 2.39</td>
<td id="552E5D9EFFC40048FAF0A4BFFA4E0495" box="[1338,1459,1722,1746]" gridcol="9" gridrow="3" pageId="19" pageNumber="20">48.94 ± 1.44</td>
</tr>
</table>
</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A30BFEF60385" blockId="20.[127,780,144,1985]" pageId="20" pageNumber="21">In this context it is noteworthy that the epithelium of the first annular fold in the atrium, located at the base of the functional penis papilla, is much lower than that of the rest of the epithelium. Perhaps this lower epithelium signals that this proximal, basal fold actually is the remnant of what once was a normal penis papilla?</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A3CCFEA80039" blockId="20.[127,780,144,1985]" pageId="20" pageNumber="21">
Other notable features of
<taxonomicName id="6FCF4D23FFC3FFA3FE5DA3CFFE0503A6" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[407,504,457,481]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FE5DA3CFFE0503A6" box="[407,504,457,481]" italics="true" pageId="20" pageNumber="21">D. insolita</emphasis>
</taxonomicName>
are (i) shell glands opening through the roof of the female atrium, (ii) the knee-shaped bend in the bursal canal, (iii) muscle fibres extending from the penis bulb to the posterior wall of the male atrium, (iv) the presence of hyperplasic ovaries, and (v) a highly triangular head with long and pointed auricles.
</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A083FF3001B1" blockId="20.[127,780,144,1985]" pageId="20" pageNumber="21">
In species of
<taxonomicName id="6FCF4D23FFC3FFA3FEFEA083FE7F00DA" authorityName="Girard" authorityYear="1850" box="[308,386,646,669]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC3FFA3FEFEA083FE7F00DA" box="[308,386,646,669]" italics="true" pageId="20" pageNumber="21">Dugesia</emphasis>
</taxonomicName>
, shell glands usually open into the bursal canal, just ventral to the oviducal openings. However, frequently the openings of the shell glands are located very close to the point where the bursal canal communicates with the atrium, while occasionally part of the shell glands actually discharges into the female atrium, e.g. in
<taxonomicName id="6FCF4D23FFC3FFA3FDC0A126FCF9017D" authority="Pala et al., 1981" authorityName="Pala" authorityYear="1981" box="[522,772,802,826]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="hepta">
<emphasis id="9ABBEAB2FFC3FFA3FDC0A126FDA3017D" box="[522,606,802,826]" italics="true" pageId="20" pageNumber="21">D. hepta</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FDA9A127FCF9017D" author="Pala M &amp; Casu S &amp; Vacca RA" box="[611,772,802,826]" pageId="20" pageNumber="21" pagination="97 - 107" refId="ref18194" refString="Pala M, Casu S, Vacca RA. Dugesia hepta, nuova specie di planaria d'acqua dolce di Sardegna appartenente alla superspecie Dugesia gonocephala (Duges) (Turbellaria, Tricladida). Bollettino della Societa Sarda di Scienze Naturali 1981; 20: 97 - 107." type="journal article" year="1981">
Pala
<emphasis id="9ABBEAB2FFC3FFA3FD5EA126FD43017D" box="[660,702,802,826]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
., 1981
</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="6FCF4D23FFC3FFA3FF4BA147FE03011E" authority="Stocchino &amp; Sluys, 2013" authorityName="Stocchino &amp; Sluys" authorityYear="2013" box="[129,510,833,857]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="superioris">
<emphasis id="9ABBEAB2FFC3FFA3FF4BA147FF07011E" box="[129,250,834,857]" italics="true" pageId="20" pageNumber="21">D. superioris</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FECAA144FE03011E" author="Stocchino GA &amp; Sluys R &amp; Deri P" box="[256,510,833,857]" pageId="20" pageNumber="21" pagination="25 - 43" refId="ref19132" refString="Stocchino GA, Sluys R, Deri P et al. Integrative taxonomy of a new species of planarian from the Lake Ohrid basin, including an analysis of biogeographical patterns in freshwater triclads from the Ohrid region (Platyhelminthes, Tricladida, Dugesiidae). ZooKeys 2013; 313: 25 - 43. https: // doi. org / 10.3897 / zookeys. 313.5363" type="journal article" year="2013">Stocchino &amp; Sluys, 2013</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="6FCF4D23FFC3FFA3FDC2A147FD24011E" authority="Sluys, 2013" authorityName="Sluys" authorityYear="2013" box="[520,729,833,857]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="effusa">
<emphasis id="9ABBEAB2FFC3FFA3FDC2A147FDA0011E" box="[520,605,833,857]" italics="true" pageId="20" pageNumber="21">D. effusa</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FDA8A144FD24011E" author="Sluys R &amp; Sola E &amp; Gritzalis K" box="[610,729,833,857]" pageId="20" pageNumber="21" pagination="523 - 47" refId="ref18642" refString="Sluys R, Sola E, Gritzalis K et al. Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 2013; 169: 523 - 47. https: // doi. org / 10.1111 / zoj. 12077" type="journal article" year="2013">Sluys, 2013</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="6FCF4D23FFC3FFA3FF4BA164FF1B013E" authorityName="Sluys" authorityYear="2013" box="[129,230,865,889]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="naiadis">
<emphasis id="9ABBEAB2FFC3FFA3FF4BA164FF1B013E" box="[129,230,865,889]" italics="true" pageId="20" pageNumber="21">D. naiadis</emphasis>
</taxonomicName>
(see:
<bibRefCitation id="CC5E4B51FFC3FFA3FEECA164FDFC013F" author="Stocchino GA &amp; Corso G &amp; Manconi R" box="[294,513,864,889]" pageId="20" pageNumber="21" pagination="1947 - 60" refId="ref18980" refString="Stocchino GA, Corso G, Manconi R et al. Endemic freshwater planarians of Sardinia: Redescription of Dugesia hepta (Platyhelminthes, Tricladida) with a comparison of the Mediterranean species of the genus. Journal of Natural History 2005; 39: 1947 - 60." type="journal article" year="2005">
Stocchino
<emphasis id="9ABBEAB2FFC3FFA3FE5FA164FE42013E" box="[405,447,865,889]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. 2005
</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FDC4A164FDBF013E" author="Stocchino GA &amp; Sluys R &amp; Deri P" box="[526,578,865,889]" pageId="20" pageNumber="21" pagination="25 - 43" refId="ref19132" refString="Stocchino GA, Sluys R, Deri P et al. Integrative taxonomy of a new species of planarian from the Lake Ohrid basin, including an analysis of biogeographical patterns in freshwater triclads from the Ohrid region (Platyhelminthes, Tricladida, Dugesiidae). ZooKeys 2013; 313: 25 - 43. https: // doi. org / 10.3897 / zookeys. 313.5363" type="journal article" year="2013">2013</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FD85A164FD04013E" author="Sluys R &amp; Sola E &amp; Gritzalis K" box="[591,761,865,889]" pageId="20" pageNumber="21" pagination="523 - 47" refId="ref18642" refString="Sluys R, Sola E, Gritzalis K et al. Integrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae). Zoological Journal of the Linnean Society 2013; 169: 523 - 47. https: // doi. org / 10.1111 / zoj. 12077" type="journal article" year="2013">
Sluys
<emphasis id="9ABBEAB2FFC3FFA3FD46A164FD4A013E" box="[652,695,865,889]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. 2013
</bibRefCitation>
).
<taxonomicName id="6FCF4D23FFC3FFA3FF4BA184FEE701DF" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[129,282,896,920]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FF4BA184FEE701DF" box="[129,282,896,920]" italics="true" pageId="20" pageNumber="21">Dugesia insolita</emphasis>
</taxonomicName>
represents the most extreme condition, in which almost none of the shell glands open into the bursal canal and the majority of the glands open through the roof of the female atrium.
</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A1FBFEA60696" blockId="20.[127,780,144,1985]" pageId="20" pageNumber="21">
From its opening into the atrium, the bursal canal in species of
<taxonomicName id="6FCF4D23FFC3FFA3FF50A618FF150673" authorityName="Girard" authorityYear="1850" box="[154,232,1053,1076]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC3FFA3FF50A618FF150673" box="[154,232,1053,1076]" italics="true" pageId="20" pageNumber="21">Dugesia</emphasis>
</taxonomicName>
generally curves smoothly towards the anterior end of the body to open into the copulatory bursa. However, in some species the bursal canal bends much more abruptly in the anterior direction, as is the case in
<taxonomicName id="6FCF4D23FFC3FFA3FE0CA67EFD5706D4" authority="Sluys, 2007" authorityName="Sluys" authorityYear="2007" box="[454,682,1147,1171]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="capensis">
<emphasis id="9ABBEAB2FFC3FFA3FE0CA67EFDCD06D5" box="[454,560,1147,1170]" italics="true" pageId="20" pageNumber="21">D. capensis</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FDFCA67EFD5706D4" author="Sluys R" box="[566,682,1147,1171]" pageId="20" pageNumber="21" pagination="229 - 57" refId="ref18548" refString="Sluys R. Annotations on freshwater planarians (Platyhelminthes Tricladida Dugesiidae) from the Afrotropical Region. Tropical Zoology 2007; 20: 229 - 57." type="journal article" year="2007">Sluys, 2007</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="6FCF4D23FFC3FFA3FD7FA67EFE8A06F5" authority="Stocchino &amp; Sluys, 2014" authorityName="Stocchino &amp; Sluys" authorityYear="2014" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="bifida">
<emphasis id="9ABBEAB2FFC3FFA3FD7FA67EFCF406D4" box="[693,777,1147,1171]" italics="true" pageId="20" pageNumber="21">D. bifida</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FF4BA69FFE8A06F5" author="Stocchino GA &amp; Sluys R &amp; Manconi R" box="[129,375,1178,1202]" pageId="20" pageNumber="21" pagination="71 - 88" refId="ref19199" refString="Stocchino GA, Sluys R, Manconi R. A new and aberrant species of Dugesia (Platyhelminthes, Tricladida, Dugesiidae) from Madagascar. ZooKeys 2014; 425: 71 - 88. https: // doi. org / 10.3897 / zookeys. 425.7811" type="journal article" year="2014">Stocchino &amp; Sluys, 2014</bibRefCitation>
</taxonomicName>
(see:
<bibRefCitation id="CC5E4B51FFC3FFA3FE65A69FFDE006F5" author="Sluys R" box="[431,541,1178,1202]" pageId="20" pageNumber="21" pagination="229 - 57" refId="ref18548" refString="Sluys R. Annotations on freshwater planarians (Platyhelminthes Tricladida Dugesiidae) from the Afrotropical Region. Tropical Zoology 2007; 20: 229 - 57." type="journal article" year="2007">Sluys 2007</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FDE2A69FFD0406F5" author="Stocchino GA &amp; Sluys R &amp; Manconi R" box="[552,761,1178,1202]" pageId="20" pageNumber="21" pagination="71 - 88" refId="ref19199" refString="Stocchino GA, Sluys R, Manconi R. A new and aberrant species of Dugesia (Platyhelminthes, Tricladida, Dugesiidae) from Madagascar. ZooKeys 2014; 425: 71 - 88. https: // doi. org / 10.3897 / zookeys. 425.7811" type="journal article" year="2014">
Stocchino
<emphasis id="9ABBEAB2FFC3FFA3FD59A69EFD4606F5" box="[659,699,1178,1202]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. 2014
</bibRefCitation>
), and also in
<taxonomicName id="6FCF4D23FFC3FFA3FF39A6BFFEAB0696" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[243,342,1209,1233]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FF39A6BFFEAB0696" box="[243,342,1209,1233]" italics="true" pageId="20" pageNumber="21">D. insolita</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A6DCFEAE07EA" blockId="20.[127,780,144,1985]" pageId="20" pageNumber="21">
The situation in which longitudinal muscle fibres run posteriorly from the penis bulb and eventually attach to the wall of the atrium is an uncommon condition among species of
<taxonomicName id="6FCF4D23FFC3FFA3FD7CA71DFCF90768" authorityName="Girard" authorityYear="1850" box="[694,772,1304,1327]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC3FFA3FD7CA71DFCF90768" box="[694,772,1304,1327]" italics="true" pageId="20" pageNumber="21">Dugesia</emphasis>
</taxonomicName>
. This feature is more or less present in
<taxonomicName id="6FCF4D23FFC3FFA3FDCEA732FF480729" authority="de Vries, 1988" authorityName="de Vries" authorityYear="1988" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="debeauchampi">
<emphasis id="9ABBEAB2FFC3FFA3FDCEA732FD560708" box="[516,683,1335,1359]" italics="true" pageId="20" pageNumber="21">D. debeauchampi</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FD7BA732FF480729" author="de Vries EJ" pageId="20" pageNumber="21" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries, 1988</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFC3FFA3FF2FA752FE0A0729" authority="de Vries, 1988" authorityName="de Vries" authorityYear="1988" box="[229,503,1366,1390]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="liguriensis">
<emphasis id="9ABBEAB2FFC3FFA3FF2FA752FE9D0729" box="[229,352,1366,1390]" italics="true" pageId="20" pageNumber="21">D. liguriensis</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FEACA753FE0A0729" author="de Vries EJ" box="[358,503,1366,1390]" pageId="20" pageNumber="21" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries, 1988</bibRefCitation>
</taxonomicName>
but a situation very similar to that in
<taxonomicName id="6FCF4D23FFC3FFA3FF14A773FEC207CA" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[222,319,1397,1421]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FF14A773FEC207CA" box="[222,319,1397,1421]" italics="true" pageId="20" pageNumber="21">D. insolita</emphasis>
</taxonomicName>
is expressed in
<taxonomicName id="6FCF4D23FFC3FFA3FE12A773FCF407CA" authority="de Beauchamp, 1952" authorityName="de Beauchamp" authorityYear="1952" box="[472,777,1397,1421]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFC3FFA3FE12A773FDCF07CA" box="[472,562,1397,1421]" italics="true" pageId="20" pageNumber="21">D. milloti</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FDFDA770FCF407CA" author="de Beauchamp P" box="[567,777,1397,1421]" pageId="20" pageNumber="21" pagination="362 - 70" refId="ref16540" refString="de Beauchamp P. Sur les Dugesia (Turbellaries Triclades) d'Afrique tropicale et de Madagascar. Bulletin de la Societe Zoologique de France 1952; 77: 362 - 70." type="journal article" year="1952">de Beauchamp, 1952</bibRefCitation>
</taxonomicName>
(see:
<bibRefCitation id="CC5E4B51FFC3FFA3FF7DA790FEBE07EA" author="de Vries EJ" box="[183,323,1429,1453]" pageId="20" pageNumber="21" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
).
</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A7B1FD890562" blockId="20.[127,780,144,1985]" pageId="20" pageNumber="21">Presence of hyperplasic ovaries is characteristic of normally asexually reproducing specimens that have sexualized, i.e. have developed reproductive organs, either in the field or under laboratory conditions. In a similar way, after 4 months of rearing, a single specimen in our laboratory culture of animals of the population from the small tributary of the Matsiatra River developed reproductive organs (specimen RMNH.VER.21536.1, described above), including aberrant hyperplasic ovaries. In our experience, hyperplasic ovaries and, sometimes, poorly developed testes are the only abnormalities in such sexualized individuals, so that their copulatory apparatus can be interpreted as representing the normal condition of the species.</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A529FD6205C5" blockId="20.[127,780,144,1985]" pageId="20" pageNumber="21">
A high triangular head with long, pointed auricles has been described as a uniquely characteristic feature of
<taxonomicName id="6FCF4D23FFC3FFA3FDB5A549FF0705C4" authority="(de Vries 1988)" baseAuthorityName="de Vries" baseAuthorityYear="1988" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFC3FFA3FDB5A549FD230524" box="[639,734,1867,1891]" italics="true" pageId="20" pageNumber="21">D. milloti</emphasis>
(
<bibRefCitation id="CC5E4B51FFC3FFA3FD24A54EFF0E05C4" author="de Vries EJ" pageId="20" pageNumber="21" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
)
</taxonomicName>
, but now it is known also from
<taxonomicName id="6FCF4D23FFC3FFA3FDFDA56EFD6405C4" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[567,665,1899,1923]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FDFDA56EFD6405C4" box="[567,665,1899,1923]" italics="true" pageId="20" pageNumber="21">D. insolita</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A87036A0FFC3FFA3FF56A58FFAFC03E4" blockId="20.[127,780,144,1985]" lastBlockId="20.[824,1475,144,419]" pageId="20" pageNumber="21">
Despite the fact that
<taxonomicName id="6FCF4D23FFC3FFA3FEA3A58EFE3605E5" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[361,459,1930,1954]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FEA3A58EFE3605E5" box="[361,459,1930,1954]" italics="true" pageId="20" pageNumber="21">D. insolita</emphasis>
</taxonomicName>
shares with
<taxonomicName id="6FCF4D23FFC3FFA3FD8EA58EFD6305E5" baseAuthorityName="de Vries" baseAuthorityYear="1988" box="[580,670,1930,1954]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFC3FFA3FD8EA58EFD6305E5" box="[580,670,1930,1954]" italics="true" pageId="20" pageNumber="21">D. milloti</emphasis>
</taxonomicName>
its characteristic head shape, the atrial folds, and the layer of longitudinal muscles that runs between the wall of the atrium and the penis bulb, the two species are distinctly different.
<taxonomicName id="6FCF4D23FFC3FFA3FAC4A2B5FA560280" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[1294,1451,175,199]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FAC4A2B5FA560280" box="[1294,1451,175,199]" italics="true" pageId="20" pageNumber="21">Dugesia insolita</emphasis>
</taxonomicName>
is the only species of
<taxonomicName id="6FCF4D23FFC3FFA3FC34A2CAFBB102A1" authorityName="Girard" authorityYear="1850" box="[1022,1100,207,230]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFC3FFA3FC34A2CAFBB102A1" box="[1022,1100,207,230]" italics="true" pageId="20" pageNumber="21">Dugesia</emphasis>
</taxonomicName>
in which the diaphragm has evolved into a functional penis papilla. Nevertheless, this diaphragm has retained its primary function, formation of the spermatophore (see above). In
<taxonomicName id="6FCF4D23FFC3FFA3FC1BA328FBD70302" baseAuthorityName="de Vries" baseAuthorityYear="1988" box="[977,1066,301,325]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFC3FFA3FC1BA328FBD70302" box="[977,1066,301,325]" italics="true" pageId="20" pageNumber="21">D. milloti</emphasis>
</taxonomicName>
, the ancestral condition is still present in which the spermatophore is formed within the ejaculatory duct, with the funnel-shaped base of the spermatophore enveloping the diaphragm (see:
<bibRefCitation id="CC5E4B51FFC3FFA3FBC3A38EFB6803E4" author="de Vries EJ" box="[1033,1173,395,419]" pageId="20" pageNumber="21" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: fig. 14A).
</paragraph>
<paragraph id="A87036A0FFC3FFA3FB86A3C7FB52039B" blockId="20.[1100,1199,450,476]" box="[1100,1199,450,476]" pageId="20" pageNumber="21">
<emphasis id="9ABBEAB2FFC3FFA3FB86A3C7FB52039B" box="[1100,1199,450,476]" italics="true" pageId="20" pageNumber="21">Karyology</emphasis>
</paragraph>
<paragraph id="A87036A0FFC3FFA3FCF3A3ECFC1900BC" blockId="20.[824,1476,489,1108]" pageId="20" pageNumber="21">
As for karyology,
<taxonomicName id="6FCF4D23FFC3FFA3FC3DA3ECFBA00046" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[1015,1117,489,513]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FC3DA3ECFBA00046" box="[1015,1117,489,513]" italics="true" pageId="20" pageNumber="21">D. insolita</emphasis>
</taxonomicName>
shows an eudiploid chromosome complement of 18 chromosomes with basic number n = 9. Among
<taxonomicName id="6FCF4D23FFC3FFA3FC47A02DFBD00078" box="[909,1069,551,575]" pageId="20" pageNumber="21">
<emphasis id="9ABBEAB2FFC3FFA3FC47A02DFC260078" box="[909,987,552,575]" italics="true" pageId="20" pageNumber="21">Dugesia</emphasis>
species
</taxonomicName>
, this haploid number is shared by only six other species:
<taxonomicName id="6FCF4D23FFC3FFA3FC39A042FBB50018" authorityName="Lepori" authorityYear="1948" box="[1011,1096,583,607]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="sicula">
<emphasis id="9ABBEAB2FFC3FFA3FC39A042FBB50018" box="[1011,1096,583,607]" italics="true" pageId="20" pageNumber="21">D. sicula</emphasis>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFC3FFA3FBB6A042FC900039" authority="Stocchino et al., 2009" authorityName="Stocchino" authorityYear="2009" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="maghrebiana">
<emphasis id="9ABBEAB2FFC3FFA3FBB6A042FAE40018" box="[1148,1305,583,607]" italics="true" pageId="20" pageNumber="21">D. maghrebiana</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FAD4A042FC900039" author="Stocchino GA &amp; Manconi R &amp; Corso G" pageId="20" pageNumber="21" pagination="83 - 91" refId="ref19029" refString="Stocchino GA, Manconi R, Corso G et al. African planarians: morphology and karyology of Dugesia maghrebiana sp. n. (Platyhelminthes, Tricladida) from Tunisia. Italian Journal of Zoology 2009; 76: 83 - 91. https: // doi. org / 10.1080 / 11250000802141683" type="journal article" year="2009">
Stocchino
<emphasis id="9ABBEAB2FFC3FFA3FA46A042FA4A0018" box="[1420,1463,583,607]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
., 2009
</bibRefCitation>
</taxonomicName>
, from the Mediterranean Region;
<taxonomicName id="6FCF4D23FFC3FFA3FB1BA062FAC50039" authorityName="Harrath &amp; Sluys" authorityYear="2013" box="[1233,1336,614,638]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="arabica">
<emphasis id="9ABBEAB2FFC3FFA3FB1BA062FAC50039" box="[1233,1336,614,638]" italics="true" pageId="20" pageNumber="21">D. arabica</emphasis>
</taxonomicName>
from
<collectingCountry id="D0D87630FFC3FFA3FABFA063FA43003A" box="[1397,1470,614,637]" name="Yemen" pageId="20" pageNumber="21">Yemen</collectingCountry>
;
<taxonomicName id="6FCF4D23FFC3FFA3FCF1A083FC4600DA" authorityName="Stocchino, Corso, Manconi &amp; Pala" authorityYear="2002" box="[827,955,645,669]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="aethiopica">
<emphasis id="9ABBEAB2FFC3FFA3FCF1A083FC4600DA" box="[827,955,645,669]" italics="true" pageId="20" pageNumber="21">D. aethiopica</emphasis>
</taxonomicName>
,
<taxonomicName id="6FCF4D23FFC3FFA3FC03A083FA8F00DA" authority="Stocchino &amp; Sluys, 2012" authorityName="Stocchino &amp; Sluys" authorityYear="2012" box="[969,1394,645,669]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="afromontana">
<emphasis id="9ABBEAB2FFC3FFA3FC03A083FB9B00DA" box="[969,1126,645,669]" italics="true" pageId="20" pageNumber="21">D. afromontana</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FBA4A080FA8F00DA" author="Stocchino GA &amp; Sluys R &amp; Manconi R" box="[1134,1394,645,669]" pageId="20" pageNumber="21" pagination="43 - 58" refId="ref19083" refString="Stocchino GA, Sluys R, Manconi R. A new species of Dugesia (Platyhelminthes, Tricladida, Dugesiidae) from the Afromontane forest in South Africa, with an overview of freshwater planarians from the African continent. Zootaxa 2012; 3551: 43 - 58." type="journal article" year="2012">Stocchino &amp; Sluys, 2012</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="6FCF4D23FFC3FFA3FA61A083FB7F00FA" authority="Stocchino &amp; Sluys, 2014" authorityName="Stocchino &amp; Sluys" authorityYear="2014" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="bifida">
<emphasis id="9ABBEAB2FFC3FFA3FA61A083FC8C00FA" italics="true" pageId="20" pageNumber="21">D. bifida</emphasis>
<bibRefCitation id="CC5E4B51FFC3FFA3FCB1A0A0FB7F00FA" author="Stocchino GA &amp; Sluys R &amp; Manconi R" box="[891,1154,677,701]" pageId="20" pageNumber="21" pagination="71 - 88" refId="ref19199" refString="Stocchino GA, Sluys R, Manconi R. A new and aberrant species of Dugesia (Platyhelminthes, Tricladida, Dugesiidae) from Madagascar. ZooKeys 2014; 425: 71 - 88. https: // doi. org / 10.3897 / zookeys. 425.7811" type="journal article" year="2014">Stocchino &amp; Sluys, 2014</bibRefCitation>
</taxonomicName>
from the Afrotropical Region (cf.
<bibRefCitation id="CC5E4B51FFC3FFA3FCAAA0C1FBC8009B" author="Stocchino GA &amp; Corso G &amp; Manconi R" box="[864,1077,708,732]" pageId="20" pageNumber="21" pagination="45 - 51" refId="ref18877" refString="Stocchino GA, Corso G, Manconi R et al. African planarians: Dugesia aethiopica sp. n. (Platyhelminthes, Tricladida) from Lake Tana (NW Ethiopia). Italian Journal of Zoology 2002; 69: 45 - 51. https: // doi. org / 10.1080 / 11250000209356437" type="journal article" year="2002">
Stocchino
<emphasis id="9ABBEAB2FFC3FFA3FC07A0C0FC0B009B" box="[973,1014,708,732]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. 2002
</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FB8AA0C1FB89009B" author="Stocchino GA &amp; Manconi R &amp; Corso G" box="[1088,1140,708,732]" pageId="20" pageNumber="21" pagination="89 - 93" refId="ref18932" refString="Stocchino GA, Manconi R, Corso G et al. Karyology and karyometric analysis of an Afrotropical freshwater planarian (Platyhelminthes, Tricladida). Italian Journal of Zoology 2004; 71: 89 - 93. https: // doi. org / 10.1080 / 11250000409356557" type="journal article" year="2004">2004</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FBB5A0C1FB4E009B" author="Stocchino GA &amp; Manconi R &amp; Corso G" box="[1151,1203,708,732]" pageId="20" pageNumber="21" pagination="83 - 91" refId="ref19029" refString="Stocchino GA, Manconi R, Corso G et al. African planarians: morphology and karyology of Dugesia maghrebiana sp. n. (Platyhelminthes, Tricladida) from Tunisia. Italian Journal of Zoology 2009; 76: 83 - 91. https: // doi. org / 10.1080 / 11250000802141683" type="journal article" year="2009">2009</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FB75A0C1FB0E009B" author="Stocchino GA &amp; Sluys R &amp; Manconi R" box="[1215,1267,708,732]" pageId="20" pageNumber="21" pagination="43 - 58" refId="ref19083" refString="Stocchino GA, Sluys R, Manconi R. A new species of Dugesia (Platyhelminthes, Tricladida, Dugesiidae) from the Afromontane forest in South Africa, with an overview of freshwater planarians from the African continent. Zootaxa 2012; 3551: 43 - 58." type="journal article" year="2012">2012</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FB34A0C1FACF009B" author="Stocchino GA &amp; Sluys R &amp; Manconi R" box="[1278,1330,708,732]" pageId="20" pageNumber="21" pagination="71 - 88" refId="ref19199" refString="Stocchino GA, Sluys R, Manconi R. A new and aberrant species of Dugesia (Platyhelminthes, Tricladida, Dugesiidae) from Madagascar. ZooKeys 2014; 425: 71 - 88. https: // doi. org / 10.3897 / zookeys. 425.7811" type="journal article" year="2014">2014</bibRefCitation>
,
<bibRefCitation id="CC5E4B51FFC3FFA3FAF7A0C1FC9000BC" author="Harrath AH &amp; Sluys R &amp; Aldahmash W" pageId="20" pageNumber="21" pagination="502 - 8" refId="ref17363" refString="Harrath AH, Sluys R, Aldahmash W et al. Reproductive strategies, karyology, parasites, and taxonomic status of Dugesia populations from Yemen (Platyhelminthes, Tricladida, Dugesiidae). Zoological Science 2013; 30: 502 - 8. https: // doi. org / 10.2108 / zsj. 30.502" type="journal article" year="2013">
Harrath
<emphasis id="9ABBEAB2FFC3FFA3FA59A0C0FA41009B" box="[1427,1468,708,732]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. 2013
</bibRefCitation>
) (
<tableCitation id="E54D031BFFC3FFA3FC42A0E6FC2E00BC" box="[904,979,739,763]" captionStart="Table 4" captionStartId="22.[129,184,143,167]" captionTargetPageId="22" captionText="Table 4. Species of Dugesia for which karyological information is available, with their geographical distribution, haploid number of the chromosome complement, karyotype, and relevant references." httpUri="http://table.plazi.org/id/FCB06628FFC1FFA1FF4BA28AFD330284" pageId="20" pageNumber="21" tableUuid="FCB06628FFC1FFA1FF4BA28AFD330284">Table 4</tableCitation>
).
</paragraph>
<paragraph id="A87036A0FFC3FFA3FC9FA106FB4C0613" blockId="20.[824,1476,489,1108]" pageId="20" pageNumber="21">
<taxonomicName id="6FCF4D23FFC3FFA3FC9FA106FC0C015C" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[853,1009,771,795]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FC9FA106FC0C015C" box="[853,1009,771,795]" italics="true" pageId="20" pageNumber="21">Dugesia insolita</emphasis>
</taxonomicName>
shares with
<taxonomicName id="6FCF4D23FFC3FFA3FBBFA106FB31015C" authorityName="Stocchino &amp; Sluys" authorityYear="2014" box="[1141,1228,771,795]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="bifida">
<emphasis id="9ABBEAB2FFC3FFA3FBBFA106FB31015C" box="[1141,1228,771,795]" italics="true" pageId="20" pageNumber="21">D. bifida</emphasis>
</taxonomicName>
the condition in which fissiparous individuals exhibit a diploid chromosome complement, while in all of the other above-mentioned five species the fissiparous populations are always triploid and/or mixoploid, while species with sexual populations are diploid (viz.,
<taxonomicName id="6FCF4D23FFC3FFA3FABAA184FA3E01DF" authorityName="Lepori" authorityYear="1948" box="[1392,1475,896,920]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="sicula">
<emphasis id="9ABBEAB2FFC3FFA3FABAA184FA3E01DF" box="[1392,1475,896,920]" italics="true" pageId="20" pageNumber="21">D. sicula</emphasis>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFC3FFA3FCAEA1A5FC3401F0" authorityName="Harrath &amp; Sluys" authorityYear="2013" box="[868,969,927,951]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="arabica">
<emphasis id="9ABBEAB2FFC3FFA3FCAEA1A5FC3401F0" box="[868,969,927,951]" italics="true" pageId="20" pageNumber="21">D. arabica</emphasis>
</taxonomicName>
). However,
<taxonomicName id="6FCF4D23FFC3FFA3FB8FA1A5FB5A01F0" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[1093,1191,927,951]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FB8FA1A5FB5A01F0" box="[1093,1191,927,951]" italics="true" pageId="20" pageNumber="21">D. insolita</emphasis>
</taxonomicName>
differs from
<taxonomicName id="6FCF4D23FFC3FFA3FAE2A1A5FA8101F0" authorityName="Stocchino &amp; Sluys" authorityYear="2014" box="[1320,1404,927,951]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="bifida">
<emphasis id="9ABBEAB2FFC3FFA3FAE2A1A5FA8101F0" box="[1320,1404,927,951]" italics="true" pageId="20" pageNumber="21">D. bifida</emphasis>
</taxonomicName>
in that its diploid karyotype is characterized by metacentric chromosomes, excepting chromosome 3, which is submetacentric, while all chromosomes of
<taxonomicName id="6FCF4D23FFC3FFA3FBD9A1FBFB960652" authorityName="Stocchino &amp; Sluys" authorityYear="2014" box="[1043,1131,1021,1045]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="bifida">
<emphasis id="9ABBEAB2FFC3FFA3FBD9A1FBFB960652" box="[1043,1131,1021,1045]" italics="true" pageId="20" pageNumber="21">D. bifida</emphasis>
</taxonomicName>
are metacentric.
<taxonomicName id="6FCF4D23FFC3FFA3FAEEA1FBFA3E0652" authorityName="Stocchino &amp; Sluys" authorityYear="2024" box="[1316,1475,1021,1045]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="insolita">
<emphasis id="9ABBEAB2FFC3FFA3FAEEA1FBFA3E0652" box="[1316,1475,1021,1045]" italics="true" pageId="20" pageNumber="21">Dugesia insolita</emphasis>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFC3FFA3FCAFA618FC440672" authorityName="Stocchino &amp; Sluys" authorityYear="2014" box="[869,953,1053,1077]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="species" species="bifida">
<emphasis id="9ABBEAB2FFC3FFA3FCAFA618FC440672" box="[869,953,1053,1077]" italics="true" pageId="20" pageNumber="21">D. bifida</emphasis>
</taxonomicName>
represent the easternmost records of species with a basic chromosomal number of n = 9.
</paragraph>
</subSubSection>
</treatment>
</document>

View file

@ -0,0 +1,447 @@
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<mods:title id="A07C02D9644C013E15E7D17009760730">The long-eared freshwater planarians of Madagascar form a separate phylogenetic clade within the genus Dugesia (Platyhelminthes: Tricladida), with the description of two new species</mods:title>
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<paragraph id="A87036A0FFDDFFBDFF02A77CFD3F07D3" blockId="10.[129,778,1401,1495]" box="[200,706,1401,1428]" pageId="10" pageNumber="11">
<emphasis id="9ABBEAB2FFDDFFBDFF02A77CFD3F07D3" bold="true" box="[200,706,1401,1428]" pageId="10" pageNumber="11">
<taxonomicName id="6FCF4D23FFDDFFBDFF02A77CFD9107D4" authority="Sluys &amp; Stocchino" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[200,620,1401,1428]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="10" pageNumber="11" phylum="Platyhelminthes" rank="species" species="crassimentula" status="sp. nov.">
<emphasis id="9ABBEAB2FFDDFFBDFF02A77CFE5B07D3" bold="true" box="[200,422,1401,1428]" italics="true" pageId="10" pageNumber="11">Dugesia crassimentula</emphasis>
Sluys &amp; Stocchino
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,
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<paragraph id="A87036A0FFDDFFBDFEA6A7F2FDE30456" blockId="10.[129,777,1527,1652]" box="[364,542,1527,1553]" pageId="10" pageNumber="11">
<emphasis id="9ABBEAB2FFDDFFBDFEA6A7F2FDE30456" box="[364,542,1527,1553]" italics="true" pageId="10" pageNumber="11">Material examined</emphasis>
</paragraph>
<paragraph id="A87036A0FFDDFFBDFF4BA418FDBF0433" blockId="10.[129,777,1527,1652]" pageId="10" pageNumber="11">
<materialsCitation id="18A73CFDFFDDFFBDFF4BA418FDBF0433" collectionCode="ZMA" collectorName="J. Millot" country="Madagascar" latitude="-22.48105" location="Route Nationale d'Ivohibe" longLatPrecision="1" longitude="46.885864" pageId="10" pageNumber="11" specimenCode="ZMA V.Pl. 786.1" specimenCount="1" typeStatus="holotype">
<emphasis id="9ABBEAB2FFDDFFBDFF4BA418FF1C0472" box="[129,225,1565,1589]" italics="true" pageId="10" pageNumber="11">
<typeStatus id="77748802FFDDFFBDFF4BA418FF210472" box="[129,220,1565,1589]" pageId="10" pageNumber="11" type="holotype">Holotype</typeStatus>
:
</emphasis>
<specimenCode id="F8699EDBFFDDFFBDFF23A41BFE610472" box="[233,412,1565,1590]" pageId="10" pageNumber="11">ZMA V.Pl. 786.1</specimenCode>
,
<location id="AD10607BFFDDFFBDFE63A41BFD430472" LSID="urn:lsid:plazi:treatment:206687B6FFDDFFBAFF02A77CFA4E0723:AD10607BFFDDFFBDFE63A41BFD430472" box="[425,702,1565,1589]" country="Madagascar" latitude="-22.48105" longLatPrecision="1" longitude="46.885864" name="Route Nationale d'Ivohibe" pageId="10" pageNumber="11">Route Nationale dIvohibe</location>
(700), south-east
<collectingCountry id="D0D87630FFDDFFBDFF3DA438FE8D0412" box="[247,368,1597,1621]" name="Madagascar" pageId="10" pageNumber="11">Madagascar</collectingCountry>
(
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22°28
<emphasis id="9ABBEAB2FFDDFFBDFE0FA43EFE360413" box="[453,459,1595,1620]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis>
51.78
<emphasis id="9ABBEAB2FFDDFFBDFDCEA43EFDF20413" box="[516,527,1595,1620]" italics="true" pageId="10" pageNumber="11">ʹʹ</emphasis>
S
</geoCoordinate>
<geoCoordinate id="CDFB5067FFDDFFBDFDE0A438FD480412" box="[554,693,1595,1621]" degrees="46" direction="east" minutes="53" orientation="longitude" pageId="10" pageNumber="11" precision="1" seconds="9.10" value="46.885864">
46°53
<emphasis id="9ABBEAB2FFDDFFBDFDACA43EFD910413" box="[614,620,1595,1620]" italics="true" pageId="10" pageNumber="11">ʹ</emphasis>
9.10
<emphasis id="9ABBEAB2FFDDFFBDFD52A43EFD5E0413" box="[664,675,1595,1620]" italics="true" pageId="10" pageNumber="11">ʹʹ</emphasis>
E
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), 1950, coll.
<collectorName id="053A5376FFDDFFBDFF7BA458FEF80433" box="[177,261,1628,1652]" pageId="10" pageNumber="11">J. Millot</collectorName>
, sagittal sections on four slides.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFDDFFBDFE58A496FEA40556" pageId="10" pageNumber="11" type="etymology">
<paragraph id="A87036A0FFDDFFBDFE58A496FE0504EA" blockId="10.[402,504,1683,1709]" box="[402,504,1683,1709]" pageId="10" pageNumber="11">
<emphasis id="9ABBEAB2FFDDFFBDFE58A496FE0504EA" box="[402,504,1683,1709]" italics="true" pageId="10" pageNumber="11">Etymology</emphasis>
</paragraph>
<paragraph id="A87036A0FFDDFFBDFF4BA4BFFEA40556" blockId="10.[128,778,1722,1809]" pageId="10" pageNumber="11">
The specific epithet is based on the Latin adjective
<emphasis id="9ABBEAB2FFDDFFBDFD48A4BEFD380495" box="[642,709,1723,1746]" italics="true" pageId="10" pageNumber="11">crassus</emphasis>
, thick, stout, and the noun
<emphasis id="9ABBEAB2FFDDFFBDFE86A4DCFE6104B6" box="[332,412,1753,1777]" italics="true" pageId="10" pageNumber="11">mentula</emphasis>
, penis; it alludes to the plump penis papilla of the species.
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFDDFFBDFE5DA534FAB307FF" pageId="10" pageNumber="11" type="diagnosis">
<paragraph id="A87036A0FFDDFFBDFE5DA534FE09050D" blockId="10.[407,500,1841,1866]" box="[407,500,1841,1866]" pageId="10" pageNumber="11">
<emphasis id="9ABBEAB2FFDDFFBDFE5DA534FE09050D" box="[407,500,1841,1866]" italics="true" pageId="10" pageNumber="11">Diagnosis</emphasis>
</paragraph>
<paragraph id="A87036A0FFDDFFBDFF4BA552FAB307FF" blockId="10.[129,779,1879,1965]" lastBlockId="10.[824,1474,1252,1464]" pageId="10" pageNumber="11">
A
<taxonomicName id="6FCF4D23FFDDFFBDFF53A552FECA0528" box="[153,311,1879,1903]" pageId="10" pageNumber="11">
<emphasis id="9ABBEAB2FFDDFFBDFF53A552FF1A0529" box="[153,231,1879,1902]" italics="true" pageId="10" pageNumber="11">Dugesia</emphasis>
species
</taxonomicName>
characterized by the following features: high triangular head with a distinctly pointed auricle on either side; mouth opening at the hind end of the pharyngeal pocket; distinct layer of transverse muscles directly dorsally to the ventral nerve cords; plump, barrel-shaped penis papilla, provided with a large dorsal penial fold and a small ventral fold; bundles of longitudinal muscles traversing the penis papilla and attaching to the penial folds; large, conical diaphragm; distinct seminal vesicle absent; ectal reinforcement on bursal canal absent; highly asymmetrical openings of oviducts into bursal canal.
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFDDFFBDFBCCA7D2FB600451" pageId="10" pageNumber="11" type="distribution">
<paragraph id="A87036A0FFDDFFBDFBCCA7D2FB0807B6" blockId="10.[1030,1269,1495,1521]" box="[1030,1269,1495,1521]" pageId="10" pageNumber="11">
<emphasis id="9ABBEAB2FFDDFFBDFBCCA7D2FB0807B6" box="[1030,1269,1495,1521]" italics="true" pageId="10" pageNumber="11">Geographical distribution</emphasis>
</paragraph>
<paragraph id="A87036A0FFDDFFBDFCF3A7FAFB600451" blockId="10.[825,1181,1534,1559]" box="[825,1181,1534,1559]" pageId="10" pageNumber="11">
Known only from the
<typeStatus id="77748802FFDDFFBDFBD0A7FAFBB50450" box="[1050,1096,1535,1559]" pageId="10" pageNumber="11">type</typeStatus>
locality.
</paragraph>
</subSubSection>
<subSubSection id="E0D5652BFFDDFFBBFB82A433FDDF07E2" lastPageId="12" lastPageNumber="13" pageId="10" pageNumber="11" type="description">
<paragraph id="A87036A0FFDDFFBDFB82A433FB490408" blockId="10.[1096,1204,1590,1615]" box="[1096,1204,1590,1615]" pageId="10" pageNumber="11">
<emphasis id="9ABBEAB2FFDDFFBDFB82A433FB490408" box="[1096,1204,1590,1615]" italics="true" pageId="10" pageNumber="11">Description</emphasis>
</paragraph>
<paragraph id="A87036A0FFDDFFBDFCF3A459FA9C0495" blockId="10.[825,1476,1628,1965]" pageId="10" pageNumber="11">
As this specimen was originally assigned to the species
<taxonomicName id="6FCF4D23FFDDFFBDFABCA458FC8504D4" authorityName="de Beauchamp" authorityYear="1952" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="10" pageNumber="11" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDDFFBDFABCA458FC8504D4" italics="true" pageId="10" pageNumber="11">Dugesia milloti</emphasis>
</taxonomicName>
(cf.
<bibRefCitation id="CC5E4B51FFDDFFBDFC6EA47EFB9A04D4" author="de Vries EJ" box="[932,1127,1659,1683]" pageId="10" pageNumber="11" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988: 363</bibRefCitation>
), it is assumed that its external appearance is similar to that of the latter. Length of preserved specimens ~
<quantity id="6F379B45FFDDFFBDFC5CA4BFFC2C0495" box="[918,977,1722,1746]" metricMagnitude="-3" metricUnit="m" metricValue="9.0" pageId="10" pageNumber="11" unit="mm" value="9.0">9 mm</quantity>
(as measured on histological sections).
</paragraph>
<paragraph id="A87036A0FFDDFFBDFC9FA4DFFC2E0517" blockId="10.[825,1476,1628,1965]" pageId="10" pageNumber="11">
Apart from the usual subepidermal body musculature there is a distinct layer of transverse muscle located directly dorsally to the ventral nerve cords and extending throughout the body length (
<figureCitation id="30F42A25FFDDFFBDFC40A532FC3F0508" box="[906,962,1847,1871]" captionStart="Figure 7" captionStartId="13.[113,178,1178,1202]" captionTargetBox="[114,1458,142,1150]" captionTargetId="figure-382@13.[114,1458,142,1150]" captionTargetPageId="13" captionText="Figure 7. Dugesia crassimentula. Holotype ZMA V.Pl.786.1. Photomicrograph of the posterior end of the body in sagittal section, showing the penis papilla (pp) and the transverse muscles (tm)." figureDoi="http://doi.org/10.5281/zenodo.14811360" httpUri="https://zenodo.org/record/14811360/files/figure.png" pageId="10" pageNumber="11">Fig. 7</figureCitation>
).
</paragraph>
<paragraph id="A87036A0FFDDFFBCFC9FA552FEF40741" blockId="10.[825,1476,1628,1965]" lastBlockId="11.[113,763,1231,1975]" lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11">Pharynx located in the middle of the body, measuring about a quarter of the body length (as measured on histological sections). No extra layer of longitudinal muscle in the outer pharynx musculature. Mouth opening at the hind end of the pharyngeal pouch.</paragraph>
<caption id="FCB06628FFDCFFBCFFBBA67AFAF006D0" ID-DOI="http://doi.org/10.5281/zenodo.14811354" ID-Zenodo-Dep="14811354" box="[113,1293,1151,1175]" httpUri="https://zenodo.org/record/14811354/files/figure.png" pageId="11" pageNumber="12" startId="11.[113,178,1151,1175]" targetBox="[211,1362,146,1121]" targetPageId="11" targetType="figure">
<paragraph id="A87036A0FFDCFFBCFFBBA67AFAF006D0" blockId="11.[113,1293,1151,1175]" box="[113,1293,1151,1175]" pageId="11" pageNumber="12">
<emphasis id="9ABBEAB2FFDCFFBCFFBBA67AFF3406D0" bold="true" box="[113,201,1151,1175]" pageId="11" pageNumber="12">Figure 5.</emphasis>
<taxonomicName id="6FCF4D23FFDCFFBCFF04A67AFEA806D0" authorityName="de Beauchamp" authorityYear="1952" box="[206,341,1151,1175]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="11" pageNumber="12" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDCFFBCFF04A67AFEA806D0" box="[206,341,1151,1175]" italics="true" pageId="11" pageNumber="12">Dugesia milloti</emphasis>
</taxonomicName>
. Specimen Mf N F217-c, sagittal reconstruction of the male copulatory apparatus; anterior to the left.
</paragraph>
</caption>
<paragraph id="A87036A0FFDCFFBCFF47A70BFE850723" blockId="11.[113,763,1231,1975]" pageId="11" pageNumber="12">Many small, dorsal testes are distributed throughout the body length, extending from directly behind the ovaries far into the posterior end of the body.</paragraph>
<paragraph id="A87036A0FFDCFFBCFF47A769FE2B04DA" blockId="11.[113,763,1231,1975]" pageId="11" pageNumber="12">
At about the level of the root of the pharynx, the vasa deferentia expand to form spermiducal vesicles, which are filled with sperm. Posteriorly to the pharyngeal pocket the sperm ducts turn dorso-medially, after which they first decrease in diameter and then separately penetrate the antero-lateral wall of the penis bulb. Hereafter, the sperm ducts traverse the penis bulb to separately open into the funnel-shaped, proximal, posterior portion of the lumen of the diaphragm (
<figureCitation id="30F42A25FFDCFFBCFDDAA442FDA70418" box="[528,602,1607,1631]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="11" pageNumber="12">Fig. 8A</figureCitation>
). This lumen is lined with a cuboidal, nucleated epithelium that is penetrated by a fine-grained, cyanophil secretion.
</paragraph>
<paragraph id="A87036A0FFDCFFBCFF47A4A0FA770702" blockId="11.[113,763,1231,1975]" lastBlockId="11.[809,1461,1231,1975]" pageId="11" pageNumber="12">
The large, conical diaphragm is covered by a cuboidal, nucleated epithelium, which continues on the proximal, anterior half of the ejaculatory duct and is penetrated by the orange secretion of penis glands. The distal part of the wide ejaculatory duct shows a few folds before it opens at the blunt tip of the penis papilla. The latter is a plump, barrel-shaped structure that is provided with penial annexes (cf.
<bibRefCitation id="CC5E4B51FFDCFFBCFE6EA564FD85053E" author="Stocchino GA &amp; Sluys R &amp; Riutort M" box="[420,632,1889,1913]" pageId="11" pageNumber="12" pagination="727 - 56" refId="ref19246" refString="Stocchino GA, Sluys R, Riutort M et al. Freshwater planarian diversity (Platyhelminthes, Tricladida, Dugesiidae) in Madagascar: new species, cryptic species, with a redefinition of character states. Zoological Journal of the Linnean Society 2017; 181: 727 - 56. https: // doi. org / 10.1093 / zoolinnean / zlx 017" type="journal article" year="2017">
Stocchino
<emphasis id="9ABBEAB2FFDCFFBCFDDAA564FDC4053E" box="[528,569,1889,1913]" italics="true" pageId="11" pageNumber="12">et al</emphasis>
. 2017
</bibRefCitation>
) in the form of two penial folds. One, small fold is located at the ventral surface of the papilla, rather close to its root, while the other fold occurs dorsally and is much larger (
<figureCitation id="30F42A25FFDCFFBCFB51A6CAFB2506A1" box="[1179,1240,1231,1255]" captionStart="Figure 7" captionStartId="13.[113,178,1178,1202]" captionTargetBox="[114,1458,142,1150]" captionTargetId="figure-382@13.[114,1458,142,1150]" captionTargetPageId="13" captionText="Figure 7. Dugesia crassimentula. Holotype ZMA V.Pl.786.1. Photomicrograph of the posterior end of the body in sagittal section, showing the penis papilla (pp) and the transverse muscles (tm)." figureDoi="http://doi.org/10.5281/zenodo.14811360" httpUri="https://zenodo.org/record/14811360/files/figure.png" pageId="11" pageNumber="12">Figs 7</figureCitation>
and
<figureCitation id="30F42A25FFDCFFBCFAC6A6CAFAD406A0" box="[1292,1321,1231,1255]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="11" pageNumber="12">8A</figureCitation>
). In addition to these penial folds, the ventral lip of the penis papilla exhibits at its tip an invagination that may represent a non-permanent structure, presumably being based on a contraction artefact.
</paragraph>
<paragraph id="A87036A0FFDCFFBCFC8FA749FB1D0467" blockId="11.[809,1461,1231,1975]" pageId="11" pageNumber="12">
The musculature of the penis bulb extends posteriorly beyond the root of the penis papilla and attaches to two small, dorsally and ventrally located atrial folds, situated near the root of the papilla (
<figureCitation id="30F42A25FFDCFFBCFCB5A7AFFC410786" box="[895,956,1450,1474]" captionStart="Figure 7" captionStartId="13.[113,178,1178,1202]" captionTargetBox="[114,1458,142,1150]" captionTargetId="figure-382@13.[114,1458,142,1150]" captionTargetPageId="13" captionText="Figure 7. Dugesia crassimentula. Holotype ZMA V.Pl.786.1. Photomicrograph of the posterior end of the body in sagittal section, showing the penis papilla (pp) and the transverse muscles (tm)." figureDoi="http://doi.org/10.5281/zenodo.14811360" httpUri="https://zenodo.org/record/14811360/files/figure.png" pageId="11" pageNumber="12">Figs 7</figureCitation>
and
<figureCitation id="30F42A25FFDCFFBCFC24A7AFFBF10785" box="[1006,1036,1450,1474]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="11" pageNumber="12">8A</figureCitation>
). Bundles of well-developed longitudinal muscle fibres run from the penis bulb through the papilla and attach to the penial folds. The epithelium of the penis papilla is underlain by a thin layer of circular muscle.
</paragraph>
<paragraph id="A87036A0FFDCFFBCFC8FA422FC37053E" blockId="11.[809,1461,1231,1975]" pageId="11" pageNumber="12">The ovaries are situated at some distance (~500 μm) behind the brain, i.e. at about a quarter of the distance between the brain and the root of the pharynx. The infranucleated oviducts run posteriorly to about the level of the gonopore, from where they turn medio-dorsad. The left oviduct runs far towards the dorsal body surface and then bends antero-ventrally to open through the dorsal wall of the bursal canal. The right oviduct opens into the broad, basal portion of the bursal canal, not far from where it communicates with the atrium. Shell glands discharge an orange secretion into the bursal canal immediately ventrally to the oviducal openings.</paragraph>
<paragraph id="A87036A0FFDCFFBBFC8FA585FDDF07E2" blockId="11.[809,1461,1231,1975]" lastBlockId="12.[126,779,1139,1445]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
The bursal canal arises from the postero-lateral wall of the atrium and at first has an oblique orientation into postero-dorsal direction before it curves sharply anteriad.The obliquely oriented portion of the bursal canal is already rather wide and after the bend in anterior direction it gradually enlarges even more until it is as wide as the copulatory bursa, thus passing smoothly into the latter (
<figureCitation id="30F42A25FFDBFFBBFF0EA6F5FEF1074F" box="[196,268,1264,1288]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="12" pageNumber="13">Fig. 8B</figureCitation>
). The bursal canal is lined with an infranucleated epithelium that is underlain by a single, subepithelial layer of longitudinal muscle, followed by an only slightly thicker layer of circular muscle; ectal reinforcement is absent. Bursal canal and copulatory bursa contain a large spermatophore of which the sac-shaped capsule is packed with sperm.
</paragraph>
</subSubSection>
<caption id="FCB06628FFDBFFBBFF4BA613FB0F066A" ID-DOI="http://doi.org/10.5281/zenodo.14811356" ID-Zenodo-Dep="14811356" box="[129,1266,1046,1070]" httpUri="https://zenodo.org/record/14811356/files/figure.png" pageId="12" pageNumber="13" startId="12.[129,194,1046,1070]" targetBox="[225,1377,144,1018]" targetPageId="12" targetType="figure">
<paragraph id="A87036A0FFDBFFBBFF4BA613FB0F066A" blockId="12.[129,1266,1046,1070]" box="[129,1266,1046,1070]" pageId="12" pageNumber="13">
<emphasis id="9ABBEAB2FFDBFFBBFF4BA613FF250669" bold="true" box="[129,216,1046,1070]" pageId="12" pageNumber="13">Figure 6.</emphasis>
<taxonomicName id="6FCF4D23FFDBFFBBFF17A612FE990669" authorityName="de Beauchamp" authorityYear="1952" box="[221,356,1046,1070]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFF17A612FE990669" box="[221,356,1046,1070]" italics="true" pageId="12" pageNumber="13">Dugesia milloti</emphasis>
</taxonomicName>
. Specimen Mf N F217-a, photomicrograph of a sagittal section of the male copulatory apparatus.
</paragraph>
</caption>
<subSubSection id="E0D5652BFFDBFFBAFE9DA7C0FA4E0723" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13" type="discussion">
<paragraph id="A87036A0FFDBFFBBFE9DA7C0FDCE0799" blockId="12.[128,779,1476,1978]" box="[343,563,1476,1503]" pageId="12" pageNumber="13">
<emphasis id="9ABBEAB2FFDBFFBBFE9DA7C0FDCE0799" box="[343,563,1476,1503]" italics="true" pageId="12" pageNumber="13">Comparative discussion</emphasis>
</paragraph>
<paragraph id="A87036A0FFDBFFBBFF4BA7EEFF3F05FD" blockId="12.[128,779,1476,1978]" pageId="12" pageNumber="13">
This specimen initially was identified by
<bibRefCitation id="CC5E4B51FFDBFFBBFDE3A7EEFD320444" author="de Vries EJ" box="[553,719,1515,1539]" pageId="12" pageNumber="13" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries (1988)</bibRefCitation>
as
<taxonomicName id="6FCF4D23FFDBFFBBFD39A7E9FF3D0465" authorityName="de Beauchamp" authorityYear="1952" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFD39A7E9FF3D0465" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
, as she was of the opinion that contraction of the longitudinal muscles running from the penis bulb and attaching to the posterior atrial fold would considerably shorten the atrium, so that the anterior fold would sit at the base of the penis papilla. Further, she argued that contraction of the penis bulb musculature would narrow the seminal vesicle and alter the shape of the diaphragm. However, we believe that the morphology of this specimen cannot be explained as merely due to contraction artefacts but that it exhibits a suite of essentially different character states concerning the following structures: penis papilla, penial and atrial folds, diaphragm, seminal vesicle, ectal reinforcement, oviducts, mouth opening, and transverse musculature. These differences between
<taxonomicName id="6FCF4D23FFDBFFBBFEF9A586FE7005DD" authorityName="de Beauchamp" authorityYear="1952" box="[307,397,1922,1946]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFEF9A586FE7005DD" box="[307,397,1922,1946]" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFDBFFBBFE76A586FDA605DD" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[444,603,1922,1946]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDBFFBBFE76A586FDA605DD" box="[444,603,1922,1946]" italics="true" pageId="12" pageNumber="13">D. crassimentula</emphasis>
</taxonomicName>
will be discussed below.
</paragraph>
<paragraph id="A87036A0FFDBFFBBFC9FA676FB6B0721" blockId="12.[825,1477,1139,1978]" pageId="12" pageNumber="13">
An immediately obvious difference between
<taxonomicName id="6FCF4D23FFDBFFBBFAD4A671FA8706CC" authorityName="de Beauchamp" authorityYear="1952" box="[1310,1402,1139,1163]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFAD4A671FA8706CC" box="[1310,1402,1139,1163]" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFDBFFBBFA66A671FC4006ED" authorityName="Sluys &amp; Stocchino" authorityYear="2024" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDBFFBBFA66A671FC4006ED" italics="true" pageId="12" pageNumber="13">D. crassimentula</emphasis>
</taxonomicName>
concerns the size and shape of the penis papilla, which in the former is always a slim cone (cf.
<bibRefCitation id="CC5E4B51FFDBFFBBFACDA6B7FA6E068D" author="de Vries EJ" box="[1287,1427,1202,1226]" pageId="12" pageNumber="13" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: figs 9, 10;
<figureCitation id="30F42A25FFDBFFBBFCB0A6D4FC3E06AE" box="[890,963,1233,1257]" captionStart="Figure 3" captionStartId="9.[113,178,1794,1818]" captionTargetBox="[114,1458,144,1766]" captionTargetId="figure-6@9.[114,1458,144,1766]" captionTargetPageId="9" captionText="Figure 3. Dugesia milloti. Specimen ZMA V.Pl. 787.1, photomicrographs of sagittal sections. A, B, male copulatory apparatus, showing the slim penis papilla (pp), diaphragm (d), seminal vesicle (sv), and atrial valves (av)." figureDoi="http://doi.org/10.5281/zenodo.14811345" httpUri="https://zenodo.org/record/14811345/files/figure.png" pageId="12" pageNumber="13">Fig. 3A</figureCitation>
), whereas in the latter it is a plump, barrel-shaped structure that completely fills the male atrium (cf.
<bibRefCitation id="CC5E4B51FFDBFFBBFAFAA6F5FA47074F" author="de Vries EJ" box="[1328,1466,1264,1288]" pageId="12" pageNumber="13" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: fig. 14C;
<figureCitation id="30F42A25FFDBFFBBFC53A715FC2C0760" box="[921,977,1296,1320]" captionStart="Figure 7" captionStartId="13.[113,178,1178,1202]" captionTargetBox="[114,1458,142,1150]" captionTargetId="figure-382@13.[114,1458,142,1150]" captionTargetPageId="13" captionText="Figure 7. Dugesia crassimentula. Holotype ZMA V.Pl.786.1. Photomicrograph of the posterior end of the body in sagittal section, showing the penis papilla (pp) and the transverse muscles (tm)." figureDoi="http://doi.org/10.5281/zenodo.14811360" httpUri="https://zenodo.org/record/14811360/files/figure.png" pageId="12" pageNumber="13">Fig. 7</figureCitation>
). Further, in
<taxonomicName id="6FCF4D23FFDBFFBBFB97A715FB03076F" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[1117,1278,1296,1320]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDBFFBBFB97A715FB03076F" box="[1117,1278,1296,1320]" italics="true" pageId="12" pageNumber="13">D. crassimentula</emphasis>
</taxonomicName>
the penis papilla is provided with two folds, a large dorsal one and a small ventral fold, which are absent in
<taxonomicName id="6FCF4D23FFDBFFBBFBFCA74AFB6D0721" authorityName="de Beauchamp" authorityYear="1952" box="[1078,1168,1358,1382]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFBFCA74AFB6D0721" box="[1078,1168,1358,1382]" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A87036A0FFDBFFBBFC9FA76BFB010465" blockId="12.[825,1477,1139,1978]" pageId="12" pageNumber="13">
The atrium of
<taxonomicName id="6FCF4D23FFDBFFBBFC2DA76BFBBF07C1" authorityName="de Beauchamp" authorityYear="1952" box="[999,1090,1390,1414]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFC2DA76BFBBF07C1" box="[999,1090,1390,1414]" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
is provided with two annular folds or valves, with the posteriorly extending musculature of the penis bulb attaching to the posterior valve (cf.
<bibRefCitation id="CC5E4B51FFDBFFBBFB06A7A9FAAB0783" author="de Vries EJ" box="[1228,1366,1452,1476]" pageId="12" pageNumber="13" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: figs 9, 10;
<figureCitation id="30F42A25FFDBFFBBFCF3A7C9FC6607A4" box="[825,923,1484,1508]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="12" pageNumber="13">Fig. 8A, B</figureCitation>
). In contrast,
<taxonomicName id="6FCF4D23FFDBFFBBFBE3A7C9FB3407A3" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[1065,1225,1484,1508]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDBFFBBFBE3A7C9FB3407A3" box="[1065,1225,1484,1508]" italics="true" pageId="12" pageNumber="13">D. crassimentula</emphasis>
</taxonomicName>
only has two small atrial folds, not valves, one dorsal and one ventral, receiving the longitudinal muscles coming from the penis bulb.
</paragraph>
<paragraph id="A87036A0FFDBFFBBFC9FA42FFAA104B9" blockId="12.[825,1477,1139,1978]" pageId="12" pageNumber="13">
The diaphragm of
<taxonomicName id="6FCF4D23FFDBFFBBFBD7A42FFB800405" authorityName="de Beauchamp" authorityYear="1952" box="[1053,1149,1578,1602]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFBD7A42FFB800405" box="[1053,1149,1578,1602]" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
is of the stubby, valve-shaped
<typeStatus id="77748802FFDBFFBBFCF3A44FFC9A0425" box="[825,871,1610,1634]" pageId="12" pageNumber="13">type</typeStatus>
(cf.
<bibRefCitation id="CC5E4B51FFDBFFBBFC53A44CFBDA0426" author="de Vries EJ" box="[921,1063,1609,1633]" pageId="12" pageNumber="13" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: figs 9, 10;
<figureCitation id="30F42A25FFDBFFBBFB6EA44CFAF30426" box="[1188,1294,1609,1633]" captionStart="Figure 3" captionStartId="9.[113,178,1794,1818]" captionTargetBox="[114,1458,144,1766]" captionTargetId="figure-6@9.[114,1458,144,1766]" captionTargetPageId="9" captionText="Figure 3. Dugesia milloti. Specimen ZMA V.Pl. 787.1, photomicrographs of sagittal sections. A, B, male copulatory apparatus, showing the slim penis papilla (pp), diaphragm (d), seminal vesicle (sv), and atrial valves (av)." figureDoi="http://doi.org/10.5281/zenodo.14811345" httpUri="https://zenodo.org/record/14811345/files/figure.png" pageId="12" pageNumber="13">Figs 3A, B</figureCitation>
,
<figureCitation id="30F42A25FFDBFFBBFAD1A44CFAD50426" box="[1307,1320,1609,1633]" captionStart="Figure 5" captionStartId="11.[113,178,1151,1175]" captionTargetBox="[211,1362,146,1121]" captionTargetId="figure-498@11.[209,1364,144,1123]" captionTargetPageId="11" captionText="Figure 5. Dugesia milloti. Specimen MfN F217-c, sagittal reconstruction of the male copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811354" httpUri="https://zenodo.org/record/14811354/files/figure.png" pageId="12" pageNumber="13">5</figureCitation>
), whereas that of
<taxonomicName id="6FCF4D23FFDBFFBBFC92A46CFC0004C7" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[856,1021,1640,1664]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDBFFBBFC92A46CFC0004C7" box="[856,1021,1640,1664]" italics="true" pageId="12" pageNumber="13">D. crassimentula</emphasis>
</taxonomicName>
is of the pointed, conical
<typeStatus id="77748802FFDBFFBBFAEAA46CFAB304C6" box="[1312,1358,1641,1665]" pageId="12" pageNumber="13">type</typeStatus>
(
<figureCitation id="30F42A25FFDBFFBBFAA9A46DFA4C04C7" box="[1379,1457,1640,1664]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="12" pageNumber="13">Fig. 8A</figureCitation>
). Another aspect of the penis that differs between
<taxonomicName id="6FCF4D23FFDBFFBBFAEBA48DFA8604E7" authorityName="de Beauchamp" authorityYear="1952" box="[1313,1403,1672,1696]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFAEBA48DFA8604E7" box="[1313,1403,1672,1696]" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFDBFFBBFA61A48DFC4004F8" authorityName="Sluys &amp; Stocchino" authorityYear="2024" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDBFFBBFA61A48DFC4004F8" italics="true" pageId="12" pageNumber="13">D. crassimentula</emphasis>
</taxonomicName>
is the presence of a well-developed seminal vesicle in the former (cf.
<bibRefCitation id="CC5E4B51FFDBFFBBFC3EA4C3FB7C0499" author="de Vries EJ" box="[1012,1153,1734,1758]" pageId="12" pageNumber="13" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: figs 9, 10;
<figureCitation id="30F42A25FFDBFFBBFB37A4C3FA9C0499" box="[1277,1377,1734,1758]" captionStart="Figure 3" captionStartId="9.[113,178,1794,1818]" captionTargetBox="[114,1458,144,1766]" captionTargetId="figure-6@9.[114,1458,144,1766]" captionTargetPageId="9" captionText="Figure 3. Dugesia milloti. Specimen ZMA V.Pl. 787.1, photomicrographs of sagittal sections. A, B, male copulatory apparatus, showing the slim penis papilla (pp), diaphragm (d), seminal vesicle (sv), and atrial valves (av)." figureDoi="http://doi.org/10.5281/zenodo.14811345" httpUri="https://zenodo.org/record/14811345/files/figure.png" pageId="12" pageNumber="13">Fig. 3A, B</figureCitation>
) and the virtual absence of such a vesicle in the latter (
<figureCitation id="30F42A25FFDBFFBBFAC8A4E3FAB604B9" box="[1282,1355,1766,1790]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="12" pageNumber="13">Fig. 8A</figureCitation>
).
</paragraph>
<paragraph id="A87036A0FFDBFFBAFC9FA503FD240439" blockId="12.[825,1477,1139,1978]" lastBlockId="13.[113,764,1294,1976]" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13">
In
<taxonomicName id="6FCF4D23FFDBFFBBFCBEA503FC2F055A" authorityName="de Beauchamp" authorityYear="1952" box="[884,978,1797,1821]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDBFFBBFCBEA503FC2F055A" box="[884,978,1797,1821]" italics="true" pageId="12" pageNumber="13">D. milloti</emphasis>
</taxonomicName>
the connection between copulatory bursa and bursal canal is established in the usual manner, while the diameter of the latter also shows the usual dimension (cf.
<bibRefCitation id="CC5E4B51FFDBFFBBFAA6A541FC90053C" author="de Vries EJ" pageId="12" pageNumber="13" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">de Vries 1988</bibRefCitation>
: figs 9, 10;
<figureCitation id="30F42A25FFDBFFBBFC14A566FBE8053C" box="[990,1045,1891,1915]" captionStart="Figure 5" captionStartId="11.[113,178,1151,1175]" captionTargetBox="[211,1362,146,1121]" captionTargetId="figure-498@11.[209,1364,144,1123]" captionTargetPageId="11" captionText="Figure 5. Dugesia milloti. Specimen MfN F217-c, sagittal reconstruction of the male copulatory apparatus; anterior to the left." figureDoi="http://doi.org/10.5281/zenodo.14811354" httpUri="https://zenodo.org/record/14811354/files/figure.png" pageId="12" pageNumber="13">Fig. 5</figureCitation>
). In contrast, in
<taxonomicName id="6FCF4D23FFDBFFBBFB76A561FAA6053C" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[1212,1371,1891,1915]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="12" pageNumber="13" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDBFFBBFB76A561FAA6053C" box="[1212,1371,1891,1915]" italics="true" pageId="12" pageNumber="13">D. crassimentula</emphasis>
</taxonomicName>
the diameter of the bursal canal expands until it has reached the same size as the copulatory bursa, after which it almost imperceptibly grades into the bursa (
<figureCitation id="30F42A25FFDAFFBAFE9EA70BFE670761" box="[340,410,1294,1318]" captionStart="Figure 8" captionStartId="14.[129,194,1772,1796]" captionTargetBox="[194,1409,147,1741]" captionTargetId="figure-80@14.[191,1412,144,1744]" captionTargetPageId="14" captionText="Figure 8. Dugesia crassimentula. Holotype ZMA V.Pl.786.1, sagittal reconstruction of the copulatory apparatus; anterior to the right.A, male copulatory apparatus; B, female copulatory apparatus." figureDoi="http://doi.org/10.5281/zenodo.14811363" httpUri="https://zenodo.org/record/14811363/files/figure.png" pageId="13" pageNumber="14">Fig. 8B</figureCitation>
). With respect to the female copulatory apparatus, another difference between
<taxonomicName id="6FCF4D23FFDAFFBAFD87A72BFD510702" authorityName="de Beauchamp" authorityYear="1952" box="[589,684,1325,1349]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDAFFBAFD87A72BFD510702" box="[589,684,1325,1349]" italics="true" pageId="13" pageNumber="14">D. milloti</emphasis>
</taxonomicName>
and
<taxonomicName id="6FCF4D23FFDAFFBAFD2EA72BFF080723" authorityName="Sluys &amp; Stocchino" authorityYear="2024" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDAFFBAFD2EA72BFF080723" italics="true" pageId="13" pageNumber="14">D. crassimentula</emphasis>
</taxonomicName>
resides in the development of the ectal reinforcement on the bursal canal. In
<taxonomicName id="6FCF4D23FFDAFFBAFE6BA768FE0307C3" authorityName="de Beauchamp" authorityYear="1952" box="[417,510,1388,1412]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDAFFBAFE6BA768FE0307C3" box="[417,510,1388,1412]" italics="true" pageId="13" pageNumber="14">D. milloti</emphasis>
</taxonomicName>
this extra layer of longitudinal muscles extends from the vaginal area to the copulatory bursa or at least to about halfway along the bursal canal, whereas in
<taxonomicName id="6FCF4D23FFDAFFBAFF5BA7CEFECA07A5" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[145,311,1482,1506]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDAFFBAFF5BA7CEFECA07A5" box="[145,311,1482,1506]" italics="true" pageId="13" pageNumber="14">D. crassimentula</emphasis>
</taxonomicName>
ectal reinforcement is completely absent. Another aspect of the female reproductive system concerns the oviduct, which consists of nucleated cells in
<taxonomicName id="6FCF4D23FFDAFFBAFDA4A40CFD310467" authorityName="de Beauchamp" authorityYear="1952" box="[622,716,1544,1568]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDAFFBAFDA4A40CFD310467" box="[622,716,1544,1568]" italics="true" pageId="13" pageNumber="14">D. milloti</emphasis>
</taxonomicName>
and infranucleated cells in
<taxonomicName id="6FCF4D23FFDAFFBAFE96A42CFE010407" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[348,508,1576,1600]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDAFFBAFE96A42CFE010407" box="[348,508,1576,1600]" italics="true" pageId="13" pageNumber="14">D. crassimentula</emphasis>
</taxonomicName>
, while in the former the ducts open symmetrically into the bursal canal, whereas in
<taxonomicName id="6FCF4D23FFDAFFBAFD2EA44DFF080439" authorityName="Sluys &amp; Stocchino" authorityYear="2024" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDAFFBAFD2EA44DFF080439" italics="true" pageId="13" pageNumber="14">D. crassimentula</emphasis>
</taxonomicName>
the oviducal openings are highly asymmetrical.
</paragraph>
<caption id="FCB06628FFDAFFBAFFBBA69FFDAC0689" ID-DOI="http://doi.org/10.5281/zenodo.14811360" ID-Zenodo-Dep="14811360" httpUri="https://zenodo.org/record/14811360/files/figure.png" pageId="13" pageNumber="14" startId="13.[113,178,1178,1202]" targetBox="[114,1458,142,1150]" targetPageId="13" targetType="figure">
<paragraph id="A87036A0FFDAFFBAFFBBA69FFDAC0689" blockId="13.[113,1444,1178,1230]" pageId="13" pageNumber="14">
<emphasis id="9ABBEAB2FFDAFFBAFFBBA69FFF3406F5" bold="true" box="[113,201,1178,1202]" pageId="13" pageNumber="14">Figure 7.</emphasis>
<emphasis id="9ABBEAB2FFDAFFBAFF04A69EFE6606F5" box="[206,411,1178,1202]" italics="true" pageId="13" pageNumber="14">
<taxonomicName id="6FCF4D23FFDAFFBAFF04A69EFE6506F5" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[206,408,1178,1202]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="crassimentula">Dugesia crassimentula</taxonomicName>
.
</emphasis>
Holotype ZMA V.Pl. 786.1. Photomicrograph of the posterior end of the body in sagittal section, showing the penis papilla (pp) and the transverse muscles (tm).
</paragraph>
</caption>
<paragraph id="A87036A0FFDAFFBAFF47A483FDF8055C" blockId="13.[113,764,1294,1976]" pageId="13" pageNumber="14">
In
<taxonomicName id="6FCF4D23FFDAFFBAFF60A482FEB104D9" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[170,332,1670,1694]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDAFFBAFF60A482FEB104D9" box="[170,332,1670,1694]" italics="true" pageId="13" pageNumber="14">D. crassimentula</emphasis>
</taxonomicName>
the mouth opening is located at the posterior end of the pharyngeal pouch, whereas in
<taxonomicName id="6FCF4D23FFDAFFBAFDA1A4A3FD3704FA" authorityName="de Beauchamp" authorityYear="1952" box="[619,714,1701,1725]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDAFFBAFDA1A4A3FD3704FA" box="[619,714,1701,1725]" italics="true" pageId="13" pageNumber="14">D. milloti</emphasis>
</taxonomicName>
it is shifted anteriad, being located at about one-sixth of the distance between the hind wall of the pharyngeal pouch and the root of the pharynx (see
<taxonomicName id="6FCF4D23FFDAFFBAFEEBA501FE86055C" authorityName="de Beauchamp" authorityYear="1952" box="[289,379,1795,1819]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDAFFBAFEEBA501FE86055C" box="[289,379,1795,1819]" italics="true" pageId="13" pageNumber="14">D. milloti</emphasis>
</taxonomicName>
description).
</paragraph>
<paragraph id="A87036A0FFDAFFBAFF47A526FE3605FF" blockId="13.[113,764,1294,1976]" pageId="13" pageNumber="14">
<bibRefCitation id="CC5E4B51FFDAFFBAFF47A526FEC4057C" author="de Vries EJ" box="[141,313,1826,1851]" pageId="13" pageNumber="14" pagination="345 - 82" refId="ref19690" refString="de Vries EJ. A synopsis of the nominal species of the subgenus Dugesia (Platyhelminthes, Tricladida, Paludicola) from Africa and Madagascar. Zoological Journal of the Linnean Society 1988; 92: 345 - 82." type="journal article" year="1988">De Vries (1988)</bibRefCitation>
already noted that the development of the transverse musculature, located directly dorsally to the ventral nerve cords, varies between species of
<taxonomicName id="6FCF4D23FFDAFFBAFDC1A567FDA4053E" authorityName="Girard" authorityYear="1850" box="[523,601,1890,1913]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="genus">
<emphasis id="9ABBEAB2FFDAFFBAFDC1A567FDA4053E" box="[523,601,1890,1913]" italics="true" pageId="13" pageNumber="14">Dugesia</emphasis>
</taxonomicName>
. In
<taxonomicName id="6FCF4D23FFDAFFBAFD4FA567FD1E053E" authorityName="de Beauchamp" authorityYear="1952" box="[645,739,1889,1913]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDAFFBAFD4FA567FD1E053E" box="[645,739,1889,1913]" italics="true" pageId="13" pageNumber="14">D. milloti</emphasis>
</taxonomicName>
it is poorly developed, whereas in
<taxonomicName id="6FCF4D23FFDAFFBAFE09A584FD9805DF" authorityName="Sluys &amp; Stocchino" authorityYear="2024" box="[451,613,1920,1944]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="crassimentula">
<emphasis id="9ABBEAB2FFDAFFBAFE09A584FD9805DF" box="[451,613,1920,1944]" italics="true" pageId="13" pageNumber="14">D. crassimentula</emphasis>
</taxonomicName>
the transverse muscles are quite distinct (
<figureCitation id="30F42A25FFDAFFBAFE49A5A5FE4605F0" box="[387,443,1952,1976]" captionStart="Figure 7" captionStartId="13.[113,178,1178,1202]" captionTargetBox="[114,1458,142,1150]" captionTargetId="figure-382@13.[114,1458,142,1150]" captionTargetPageId="13" captionText="Figure 7. Dugesia crassimentula. Holotype ZMA V.Pl.786.1. Photomicrograph of the posterior end of the body in sagittal section, showing the penis papilla (pp) and the transverse muscles (tm)." figureDoi="http://doi.org/10.5281/zenodo.14811360" httpUri="https://zenodo.org/record/14811360/files/figure.png" pageId="13" pageNumber="14">Fig. 7</figureCitation>
).
</paragraph>
<paragraph id="A87036A0FFDAFFBAFC8FA70BFA4E0723" blockId="13.[810,1459,1294,1381]" pageId="13" pageNumber="14">
Thus, there are ample anatomical differences between
<taxonomicName id="6FCF4D23FFDAFFBAFA90A70AFA4F0761" authorityName="de Beauchamp" authorityYear="1952" box="[1370,1458,1294,1318]" family="Dugesiidae" genus="Dugesia" kingdom="Animalia" order="Tricladida" pageId="13" pageNumber="14" phylum="Platyhelminthes" rank="species" species="milloti">
<emphasis id="9ABBEAB2FFDAFFBAFA90A70AFA4F0761" box="[1370,1458,1294,1318]" italics="true" pageId="13" pageNumber="14">D. milloti</emphasis>
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and
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<emphasis id="9ABBEAB2FFDAFFBAFC9FA72BFC0C0702" box="[853,1009,1325,1349]" italics="true" pageId="13" pageNumber="14">D. crassimentula</emphasis>
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, supporting the separate specific status of the latter, as it neither conforms to any other known species of
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.
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View file

@ -1,55 +1,55 @@
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@ -187,7 +187,7 @@ locality [Aescht].
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R e m a r k s:
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differs from

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Genus
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HAITLINGER &amp; ŠUNDIĆ
@ -64,7 +64,7 @@ HAITLINGER &amp; ŠUNDIĆ
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T y p e - s p e c i e s:
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balearica
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HAITLINGER &amp; ŠUNDIĆ 2016

View file

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<emphasis id="37D4E31FFF83B965FF600B8EFF38FCED" bold="true" box="[144,242,759,778]" pageId="2" pageNumber="209">Figure 1.</emphasis>
Photograph of the holotype of
<taxonomicName id="C2A0448EFF83B965FDD50B8EFD3DFCEE" authority="Wang &amp; Zhou &amp; Zhou, 2016" authorityName="Wang &amp; Zhou &amp; Zhou" authorityYear="2016" box="[549,759,759,778]" class="Insecta" family="Erebidae" genus="Bellulia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="209" phylum="Arthropoda" rank="species" species="rectusunguis" status="gen. et sp. nov.">
<taxonomicName id="C2A0448EFF83B965FDD50B8EFD3DFCEE" authority="Wang &amp; Zhou &amp; Zhou, 2016" authorityName="Wang &amp; Zhou &amp; Zhou" authorityYear="2016" box="[549,759,759,778]" class="Insecta" genus="Bellulia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="209" phylum="Chordata" rank="species" species="rectusunguis" status="gen. et sp. nov.">
<emphasis id="37D4E31FFF83B965FDD50B8EFD3DFCEE" box="[549,759,759,778]" italics="true" pageId="2" pageNumber="209">Bellulia rectusunguis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="2CE75E64FF83B965FF600A6DFEFEFCC0" box="[144,308,788,807]" pageId="2" pageNumber="209" rank="species">gen. et sp. nov.</taxonomicNameLabel>