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<document id="7A69FA3E639480BBE906C664CDF47700" ID-CLB-Dataset="239620" ID-DOI="10.1080/00222933.2023.2168573" ID-GBIF-Dataset="83f7b49d-c03b-4658-883f-abbd0c538834" ID-ISSN="1464-5262" ID-Zenodo-Dep="7778206" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1679557092330" checkinUser="plazi" docAuthor="Falniowski, Andrzej, Jaszczyńska, Aleksandra, Osikowski, Artur &amp; Hofman, Sebastian" docDate="2023" docId="C92D878F3C66D354255DFB3DFE30FD64" docLanguage="en" docName="JNATHIST.57.5-8.299-329.pdf" docOrigin="Journal of Natural History 57 (5 - 8)" docSource="http://dx.doi.org/10.1080/00222933.2023.2168573" docStyle="DocumentStyle:3C762F9783B768A90048B82ACD135A62.3:JNatHist.2017-.journal_article.0cover" docStyleId="3C762F9783B768A90048B82ACD135A62" docStyleName="JNatHist.2017-.journal_article.0cover" docStyleVersion="3" docTitle="Litthabitellidae Falniowski and Hofman 2023, fam. nov." docType="treatment" docVersion="7" lastPageNumber="307" masterDocId="3514FFF73C60D35C244DFFDDFFC1FFB6" masterDocTitle="Litthabitellidae: a new family of the Truncatelloidea (Mollusca: Caenogastropoda)" masterLastPageNumber="329" masterPageNumber="299" pageNumber="305" updateTime="1732721511443" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="D3385BCCE287152C59FD25249CA53026">Litthabitellidae: a new family of the Truncatelloidea (Mollusca: Caenogastropoda)</mods:title>
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<mods:namePart id="3A7672772A8D12C2E1F55FFABB404264">Osikowski, Artur</mods:namePart>
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<mods:date id="01BD70B4D1E0D7D81920AEF38253B956">2023</mods:date>
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<treatment id="C92D878F3C66D354255DFB3DFE30FD64" ID-DOI="http://doi.org/10.5281/zenodo.7778208" ID-GBIF-Taxon="209469834" ID-Zenodo-Dep="7778208" LSID="urn:lsid:zoobank.org:act:A8E1F6A7-EEAE-4B8D-A82D-19A7144283C6" httpUri="http://treatment.plazi.org/id/C92D878F3C66D354255DFB3DFE30FD64" lastPageId="8" lastPageNumber="307" pageId="6" pageNumber="305" scope_class="Gastropoda" scope_order="Littorinimorpha" scope_phylum="Mollusca" scope_subClass="Caenogastropoda" scope_superFamily="Truncatelloidea">
<subSubSection id="099E65123C66D35A255DFB3DFBCFFA99" pageId="6" pageNumber="305" type="nomenclature">
<paragraph id="413B36993C66D35A255DFB3DFBD5FB4D" blockId="6.[272,1044,1248,1275]" box="[272,1044,1248,1275]" pageId="6" pageNumber="305">
<heading id="1A7381F53C66D35A255DFB3DFBD5FB4D" box="[272,1044,1248,1275]" centered="true" fontSize="11" level="1" pageId="6" pageNumber="305" reason="6">
Family
<taxonomicName id="86844D1A3C66D35A2525FB3DFC4CFB4D" ID-CoL="a52cdf20-a21b-4957-8208-f32fa60ee921" authority="Falniowski and Hofman" authorityName="Falniowski and Hofman" authorityYear="2023" box="[360,909,1248,1275]" class="Gastropoda" family="Litthabitellidae" higherTaxonomySource="CoL" kingdom="Animalia" order="Littorinimorpha" pageId="6" pageNumber="305" phylum="Mollusca" rank="family" status="fam. nov.">
<emphasis id="73F0EA8B3C66D35A2525FB3DFD9FFB4D" bold="true" box="[360,606,1248,1275]" pageId="6" pageNumber="305">LITTHABITELLIDAE</emphasis>
Falniowski and Hofman
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,
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<taxonomicNameLabel id="68C357F03C66D35A27D5FB3DFBD5FB4D" box="[920,1044,1248,1275]" pageId="6" pageNumber="305" rank="family">fam. nov.</taxonomicNameLabel>
</emphasis>
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urn:lsid:zoobank.org:act:
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<typeStatus id="9E3F883B3C66D35A24EDFAB9FF19FAC8" box="[160,216,1380,1406]" pageId="6" pageNumber="305">Type</typeStatus>
genus
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<taxonomicName id="86844D1A3C66D35A24EDFA55FE00FA17" ID-CoL="7P7R8" authority="Boeters, 1970" authorityName="Boeters" authorityYear="1970" box="[160,449,1416,1441]" class="Gastropoda" family="Hydrobiidae" genus="Litthabitella" kingdom="Animalia" order="Littorinimorpha" pageId="6" pageNumber="305" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C66D35A24EDFA55FEE0FA17" box="[160,289,1416,1441]" italics="true" pageId="6" pageNumber="305">Litthabitella</emphasis>
<bibRefCitation id="25154B683C66D35A2564FA55FE00FA17" author="Boeters HD" box="[297,449,1416,1441]" pageId="6" pageNumber="305" pagination="113 - 145" refId="ref10202" refString="Boeters HD. 1970. Die Gattung Microna Clessin, 1890 (Prosobranchia, Hydrobiidae). Archiv fur Molluskenkunde. 100: 113 - 145." type="journal article" year="1970">Boeters, 1970</bibRefCitation>
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<subSubSection id="099E65123C66D35A24EDFA0BFD31F976" pageId="6" pageNumber="305" type="diagnosis">
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<emphasis id="73F0EA8B3C66D35A24EDFA0BFED5FA46" bold="true" box="[160,276,1494,1520]" italics="true" pageId="6" pageNumber="305">Diagnosis</emphasis>
</paragraph>
<paragraph id="413B36993C66D35A24EDFA27FD31F976" blockId="6.[160,1156,1494,1728]" pageId="6" pageNumber="305">Minute truncatelloid snails with the following synapomorphic character states: osphradium broadly ovate; pallial oviduct gland complex short; gonoporus deep inside the mantle cavity; the ventral channel not on the ventral side of the capsule gland, but somewhat lateral; thickened oviduct circular; the distal termination of the penis wide and blunt, with sharp and short filament harbouring the vas deferens situated laterally, and with two flat, cockscomb-like, four-folded lobes.</paragraph>
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<subSubSection id="099E65123C67D35424EDFF53FE30FD64" lastPageId="8" lastPageNumber="307" pageId="7" pageNumber="306" type="description">
<paragraph id="413B36993C67D35B24EDFF53FEE7FF1E" blockId="7.[160,1157,142,1243]" box="[160,294,142,168]" pageId="7" pageNumber="306">
<emphasis id="73F0EA8B3C67D35B24EDFF53FEE7FF1E" bold="true" box="[160,294,142,168]" italics="true" pageId="7" pageNumber="306">Description</emphasis>
</paragraph>
<paragraph id="413B36993C67D35B24EDFF6FFEDFFE56" blockId="7.[160,1157,142,1243]" pageId="7" pageNumber="306">
Shell (
<figureCitation id="D9BF2A1C3C67D35B24A9FF6FFE8AFF7D" box="[228,331,178,203]" captionStart="Figure 3" captionStartId="13.[160,225,707,730]" captionTargetBox="[163,1153,142,675]" captionTargetId="figure-186@13.[163,1153,142,675]" captionTargetPageId="13" captionText="Figure 3. Holotypes of Litthabitella: (a) L. aquaadcrucem; (b) L. cetinensis; (c) L. levkadensis. Scale bar: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7778216" httpUri="https://zenodo.org/record/7778216/files/figure.png" pageId="7" pageNumber="306">Figures 3</figureCitation>
and
<figureCitation id="D9BF2A1C3C67D35B25CEFF6FFE50FF7D" box="[387,401,178,203]" captionStart="Figure 4" captionStartId="14.[160,225,1444,1467]" captionTargetBox="[163,1153,142,1413]" captionTargetId="figure-3@14.[163,1153,142,1413]" captionTargetPageId="14" captionText="Figure 4. Shells of Litthabitella. (al) L. chilodia: (ad) locality 3, (ej) locality 5, (k,l) locality 2; (ms) L. aquaadcrucem: (mq) locality 6, (r) locality 8, (s) locality 9; (tv) L. cetinensis, locality 1. Sequence numbers: a, 2G39; b, 2G40; c, 2G43; d, 2G45; e, 2G4; f, 2G1; g, 2G3; h, 2G2; i, 2F99; j, 2F100; k, 2D3; l, 2D5; m, 1T10; n, 1T11; o, 1T12; p, 2G15; q, 2G16; r, 8F3; s, 1M25; t, 8G1; u, 8G2; v, 6D8. Scale bar: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7778220" httpUri="https://zenodo.org/record/7778220/files/figure.png" pageId="7" pageNumber="306">4</figureCitation>
) ovate conical, convex in outline, thick-walled; the border between the proto- and teleoconch is easily discernible, the initial part of the protoconch narrow, on the protoconch surface only delicate irregularities (
<figureCitation id="D9BF2A1C3C67D35B2740FF2AFCABFEA6" box="[781,874,247,272]" captionStart="Figure 5" captionStartId="15.[160,225,1165,1188]" captionTargetBox="[170,1147,149,1128]" captionTargetId="figure-73@15.[163,1153,142,1134]" captionTargetPageId="15" captionText="Figure 5. Protoconchs and radulae of Litthabitella. (ac) Protoconchs: (a) L. chilodia, locality 5; (b) L. aquaadcrucem, locality 6; (c) L. cetinensis, locality 1; (di) radulae: (d,g) L. chilodia, locality 5; (e,h) L. aquaadcrucem, locality 6; (f,i) L. cetinensis, locality 1. Scale bars: ac = 400 μm; df = 30 μm; gi = 10 μm." figureDoi="http://doi.org/10.5281/zenodo.7778224" httpUri="https://zenodo.org/record/7778224/files/figure.png" pageId="7" pageNumber="306">Figure 5</figureCitation>
(ac)), aperture pyriform, teleoconch smooth with delicate growth lines. Operculum (
<figureCitation id="D9BF2A1C3C67D35B2716FEC7FC7AFE85" box="[859,955,282,307]" captionStart="Figure 6" captionStartId="16.[160,225,820,843]" captionTargetBox="[162,1154,142,789]" captionTargetId="figure-3@16.[162,1154,142,789]" captionTargetPageId="16" captionText="Figure 6. Operculum of Litthabitella chilodia. Scale bar: 300 μm." figureDoi="http://doi.org/10.5281/zenodo.7778226" httpUri="https://zenodo.org/record/7778226/files/figure.png" pageId="7" pageNumber="306">Figure 6</figureCitation>
) thin and horny, colourless and translucent, elongate-ellipsoidal, spiral, paucispiral, with submarginal nucleus. Head and tentacles white, with no pigment, no hyperciliation (long and dense cilia at one or two cephalic tentacles, characteristic of e.g. some
<taxonomicName id="86844D1A3C67D35B27D9FE5FFBE0FE2D" authorityName="Stimpson" authorityYear="1865" box="[916,1057,386,411]" class="Gastropoda" family="Hydrobiidae" kingdom="Animalia" order="Littorinimorpha" pageId="7" pageNumber="306" phylum="Mollusca" rank="family">Hydrobiidae</taxonomicName>
); mantle partly black pigmented. Osphradium (
<figureCitation id="D9BF2A1C3C67D35B2635FE79FD1EFE0B" box="[632,735,420,445]" captionStart="Figure 7" captionStartId="16.[160,225,1577,1600]" captionTargetBox="[166,1143,883,1543]" captionTargetId="figure-18@16.[163,1153,881,1546]" captionTargetPageId="16" captionText="Figure 7. Female reproductive organs of Litthabitella aquaadcrucem (Sotonići,locality 8): (a) female organs with rectum, anus, ctenidium and osphradium; (b) pallial section of the organs; (c) complex of bursa copulatrix, two receptacles and oviduct; (d) perpendicular sections through the capsule gland. Abbreviations: a, anus; bc, bursa copulatrix; cbc, bursal duct; ct, ctenidium; ga, albumen gland; gn, capsule gland; gp, gonoporus;osp,osphradium;ov, oviduct; rec, rectum;rs1, seminal receptacle close to junction of bursal duct and albumen gland; rs2, seminal receptacle adjoining renal oviduct; vc, ventral channel; vc̍, ventral channel separated with broad folds, imitating spermathecal duct) (after Szarowska 2006)." figureDoi="http://doi.org/10.5281/zenodo.7778230" httpUri="https://zenodo.org/record/7778230/files/figure.png" pageId="7" pageNumber="306">Figure 7</figureCitation>
(a)) broadly ovate; ctenidium well developed.
</paragraph>
<paragraph id="413B36993C67D35B248DFE37FEF2FD43" blockId="7.[160,1157,142,1243]" pageId="7" pageNumber="306">
In the radula (
<figureCitation id="D9BF2A1C3C67D35B253DFE37FE21FDB5" box="[368,480,490,515]" captionStart="Figure 5" captionStartId="15.[160,225,1165,1188]" captionTargetBox="[170,1147,149,1128]" captionTargetId="figure-73@15.[163,1153,142,1134]" captionTargetPageId="15" captionText="Figure 5. Protoconchs and radulae of Litthabitella. (ac) Protoconchs: (a) L. chilodia, locality 5; (b) L. aquaadcrucem, locality 6; (c) L. cetinensis, locality 1; (di) radulae: (d,g) L. chilodia, locality 5; (e,h) L. aquaadcrucem, locality 6; (f,i) L. cetinensis, locality 1. Scale bars: ac = 400 μm; df = 30 μm; gi = 10 μm." figureDoi="http://doi.org/10.5281/zenodo.7778224" httpUri="https://zenodo.org/record/7778224/files/figure.png" pageId="7" pageNumber="306">Figures 5</figureCitation>
(di) and 8(b)) the central tooth with deep sinus at the proximal part of the cutting plate and one pair of large and prominent basal cusps. The median cusp narrow and slender, more than twice as long as the adjacent cusps, similar in shape. On each side of the median cusp are four to five cusps; the cutting edge is straight, with no cusps. In the lateral tooth, the biggest cusp is triangular, twice as long and broad as the adjacent ones. The formula of the tooth is (5)415; on the inner marginal tooth 2628 long and slender cusps. The caecum is absent (
<figureCitation id="D9BF2A1C3C67D35B24E5FD01FEC8FD43" box="[168,265,732,757]" captionStart="Figure 8" captionStartId="17.[160,225,926,949]" captionTargetBox="[166,1147,144,885]" captionTargetId="figure-3@17.[163,1153,142,894]" captionTargetPageId="17" captionText="Figure 8. Histology of L. aquaadcrucem: (a) head; (b) radula and ctenidium; (c) stomach; (d) eye. Abbreviations: bc, buccal cavity; c, cuticle; cce, ciliated columnar epithelium; ct, ctenidium; e, eye; es, oesophagus; l, lens; m, mantle; o, odontophores; r, retina; ra, radulae; sg, salivary glands; ss, style sac; vb, vitreous body. Scale bars: a = 150 μm; bd = 100 μm." figureDoi="http://doi.org/10.5281/zenodo.7778233" httpUri="https://zenodo.org/record/7778233/files/figure.png" pageId="7" pageNumber="306">Figure 8</figureCitation>
(c)).
</paragraph>
<paragraph id="413B36993C67D35B248DFD22FC63FB6D" blockId="7.[160,1157,142,1243]" pageId="7" pageNumber="306">
The pallial oviduct gland complex (
<figureCitation id="D9BF2A1C3C67D35B262CFD22FD03FCAE" box="[609,706,767,792]" captionStart="Figure 7" captionStartId="16.[160,225,1577,1600]" captionTargetBox="[166,1143,883,1543]" captionTargetId="figure-18@16.[163,1153,881,1546]" captionTargetPageId="16" captionText="Figure 7. Female reproductive organs of Litthabitella aquaadcrucem (Sotonići,locality 8): (a) female organs with rectum, anus, ctenidium and osphradium; (b) pallial section of the organs; (c) complex of bursa copulatrix, two receptacles and oviduct; (d) perpendicular sections through the capsule gland. Abbreviations: a, anus; bc, bursa copulatrix; cbc, bursal duct; ct, ctenidium; ga, albumen gland; gn, capsule gland; gp, gonoporus;osp,osphradium;ov, oviduct; rec, rectum;rs1, seminal receptacle close to junction of bursal duct and albumen gland; rs2, seminal receptacle adjoining renal oviduct; vc, ventral channel; vc̍, ventral channel separated with broad folds, imitating spermathecal duct) (after Szarowska 2006)." figureDoi="http://doi.org/10.5281/zenodo.7778230" httpUri="https://zenodo.org/record/7778230/files/figure.png" pageId="7" pageNumber="306">Figure 7</figureCitation>
(a)) is relatively short, lying far posteriorly of the mantle margin, thus the gonopore is at the terminal part of the capsule gland and situated deep inside the mantle cavity, far behind the anus. The ventral channel is not on the ventral side of the capsule gland, but lies somewhat laterally (
<figureCitation id="D9BF2A1C3C67D35B24E5FC57FED2FC15" box="[168,275,906,931]" captionStart="Figure 7" captionStartId="16.[160,225,1577,1600]" captionTargetBox="[166,1143,883,1543]" captionTargetId="figure-18@16.[163,1153,881,1546]" captionTargetPageId="16" captionText="Figure 7. Female reproductive organs of Litthabitella aquaadcrucem (Sotonići,locality 8): (a) female organs with rectum, anus, ctenidium and osphradium; (b) pallial section of the organs; (c) complex of bursa copulatrix, two receptacles and oviduct; (d) perpendicular sections through the capsule gland. Abbreviations: a, anus; bc, bursa copulatrix; cbc, bursal duct; ct, ctenidium; ga, albumen gland; gn, capsule gland; gp, gonoporus;osp,osphradium;ov, oviduct; rec, rectum;rs1, seminal receptacle close to junction of bursal duct and albumen gland; rs2, seminal receptacle adjoining renal oviduct; vc, ventral channel; vc̍, ventral channel separated with broad folds, imitating spermathecal duct) (after Szarowska 2006)." figureDoi="http://doi.org/10.5281/zenodo.7778230" httpUri="https://zenodo.org/record/7778230/files/figure.png" pageId="7" pageNumber="306">Figure 7</figureCitation>
(b,d)). There is a bursa copulatrix and two receptacula typical of the
<taxonomicName id="86844D1A3C67D35B24ECFC71FE94FC73" authorityName="Gray" authorityYear="1840" box="[161,341,940,965]" class="Gastropoda" kingdom="Animalia" order="Littorinimorpha" pageId="7" pageNumber="305" phylum="Mollusca" rank="superFamily" superFamily="Truncatelloidea">Truncatelloidea</taxonomicName>
: a distal one (rs
<subScript id="DD0034DC3C67D35B2658FC6AFDDEFC71" attach="left" box="[533,543,951,967]" fontSize="7" pageId="7" pageNumber="306">1</subScript>
after
<bibRefCitation id="25154B683C67D35B2626FC71FCDBFC73" author="Radoman P" box="[619,794,940,965]" pageId="7" pageNumber="306" pagination="1 - 256" refId="ref12728" refString="Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Serbian Academy of Sciences and Arts, Monograph 547, Department of Sciences. 57: 1 - 256." type="journal article" year="1983">Radoman 1983</bibRefCitation>
) situated near the junction of the oviduct and the duct of the bursa, and a proximal one (rs
<subScript id="DD0034DC3C67D35B2736FC07FC44FC5C" attach="left" box="[891,901,986,1002]" fontSize="7" pageId="7" pageNumber="306">2</subScript>
after
<bibRefCitation id="25154B683C67D35B2783FC12FBBDFC5E" author="Radoman P" box="[974,1148,975,1000]" pageId="7" pageNumber="306" pagination="1 - 256" refId="ref12728" refString="Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Serbian Academy of Sciences and Arts, Monograph 547, Department of Sciences. 57: 1 - 256." type="journal article" year="1983">Radoman 1983</bibRefCitation>
) situated at the proximal end of thickened oviduct; in
<taxonomicName id="86844D1A3C67D35B271FFC2FFC18FBBD" authorityName="Boeters" authorityYear="1970" box="[850,985,1010,1035]" class="Gastropoda" family="Hydrobiidae" genus="Litthabitella" kingdom="Animalia" order="Littorinimorpha" pageId="7" pageNumber="306" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C67D35B271FFC2FFC18FBBD" box="[850,985,1010,1035]" italics="true" pageId="7" pageNumber="306">Litthabitella</emphasis>
</taxonomicName>
the proximal receptaculum is vestigial. The thickened oviduct is circular, forming neither a spiral nor a horseshoe-like loop (
<figureCitation id="D9BF2A1C3C67D35B25ADFBEAFD80FBE6" box="[480,577,1079,1104]" captionStart="Figure 7" captionStartId="16.[160,225,1577,1600]" captionTargetBox="[166,1143,883,1543]" captionTargetId="figure-18@16.[163,1153,881,1546]" captionTargetPageId="16" captionText="Figure 7. Female reproductive organs of Litthabitella aquaadcrucem (Sotonići,locality 8): (a) female organs with rectum, anus, ctenidium and osphradium; (b) pallial section of the organs; (c) complex of bursa copulatrix, two receptacles and oviduct; (d) perpendicular sections through the capsule gland. Abbreviations: a, anus; bc, bursa copulatrix; cbc, bursal duct; ct, ctenidium; ga, albumen gland; gn, capsule gland; gp, gonoporus;osp,osphradium;ov, oviduct; rec, rectum;rs1, seminal receptacle close to junction of bursal duct and albumen gland; rs2, seminal receptacle adjoining renal oviduct; vc, ventral channel; vc̍, ventral channel separated with broad folds, imitating spermathecal duct) (after Szarowska 2006)." figureDoi="http://doi.org/10.5281/zenodo.7778230" httpUri="https://zenodo.org/record/7778230/files/figure.png" pageId="7" pageNumber="306">Figure 7</figureCitation>
(c)). The prostate is small, nearly vestigial, observable as slight outgrowth of the vas deferens. The penis is big and massive (
<figureCitation id="D9BF2A1C3C67D35B2057FB87FB45FBC5" box="[1050,1156,1114,1139]" captionStart="Figure 9" captionStartId="17.[160,227,1700,1723]" captionTargetBox="[162,1152,1102,1669]" captionTargetId="figure-71@17.[162,1152,1102,1669]" captionTargetPageId="17" captionText="Figure 9. Penes of Litthabitella: (a) L. chilodia, Trsteno, locality 5, scale bar: 500 μm; (be) L. aquaadcrucem: (bd) Sotonići, locality 8; (e) Budva, locality 6; (f) L. cetinensis, Kostanje, locality 1; (b, d,e) bent, in relaxed habitus. Arrows: glandular swellings." figureDoi="http://doi.org/10.5281/zenodo.7778235" httpUri="https://zenodo.org/record/7778235/files/figure.png" pageId="7" pageNumber="306">Figures 9</figureCitation>
and
<figureCitation id="D9BF2A1C3C67D35B2498FBA1FF30FB23" box="[213,241,1148,1173]" captionStart="Figure 10" captionStartId="18.[160,225,925,948]" captionTargetBox="[167,1151,145,887]" captionTargetId="figure-237@18.[163,1153,142,893]" captionTargetPageId="18" captionText="Figure 10. (ac) Penes of L. aquaadcrucem histology: asterix, lobes; f, filament. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.7778237" httpUri="https://zenodo.org/record/7778237/files/figure.png" pageId="7" pageNumber="306">10</figureCitation>
), filled with glandular tissue (
<figureCitation id="D9BF2A1C3C67D35B2600FBA1FD7AFB23" box="[589,699,1148,1173]" captionStart="Figure 10" captionStartId="18.[160,225,925,948]" captionTargetBox="[167,1151,145,887]" captionTargetId="figure-237@18.[163,1153,142,893]" captionTargetPageId="18" captionText="Figure 10. (ac) Penes of L. aquaadcrucem histology: asterix, lobes; f, filament. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.7778237" httpUri="https://zenodo.org/record/7778237/files/figure.png" pageId="7" pageNumber="306">Figure 10</figureCitation>
), with a distal termination that is wide and blunt, with a sharp and short filament harbouring the vas deferens, which is situated laterally, and two flat, cockscomb-like, four-folded lobes.
</paragraph>
<paragraph id="413B36993C67D35B24EDFACBFE59FA86" blockId="7.[160,1156,1302,1675]" box="[160,408,1302,1328]" pageId="7" pageNumber="306">
<emphasis id="73F0EA8B3C67D35B24EDFACBFE59FA86" bold="true" box="[160,408,1302,1328]" italics="true" pageId="7" pageNumber="306">Molecular phylogeny</emphasis>
</paragraph>
<paragraph id="413B36993C67D35B24EDFAE7FEF2F93D" blockId="7.[160,1156,1302,1675]" pageId="7" pageNumber="306">
We obtained 26 new sequences of COI (457 bp, GenBank accession numbers ON661272ON661297), 16 of H3 (310 bp, GenBank accession numbers ON661565 ON661580) and 24 of 18S (401 bp, GenBank accession numbers ON667998 ON668023). The tests by
<bibRefCitation id="25154B683C67D35B259EFA7FFD52FA0D" author="Xia X &amp; Xie Z &amp; Salemi M &amp; Chen L &amp; Wang Y" box="[467,659,1442,1467]" pageId="7" pageNumber="306" pagination="1 - 7" refId="ref15048" refString="Xia X, Xie Z, Salemi M, Chen L, Wang Y. 2003. An index of substitution saturation and its application. Mol Phylogenet Evol. 26: 1 - 7. doi: 10.1016 / S 1055 - 7903 (02)." type="journal article" year="2003">Xia et al. (2003)</bibRefCitation>
revealed no saturation. Results from the substitution saturation analysis showed an Index of substitution saturation (ISS) (0.71 for COI; 0.48 for H3) significantly smaller than the critical ISS value (ISSC: 0.97 for COI; 0.59 for H3), indicating that all sequences are useful in phylogenetic reconstruction. In all analyses, the topologies of the resulting phylograms were identical in the ML and BI phylogram analyses; thus, we present the phylogram computed with RAxML.
</paragraph>
<paragraph id="413B36993C68D354248DFF53FEE2FEC1" blockId="8.[160,1156,142,722]" pageId="8" pageNumber="307">
The COI phylogram (
<figureCitation id="D9BF2A1C3C68D35425E2FF53FDDAFF11" box="[431,539,142,167]" captionStart="Figure 11" captionStartId="19.[160,225,1321,1344]" captionTargetBox="[162,1153,143,1288]" captionTargetId="figure-105@19.[162,1154,142,1289]" captionTargetPageId="19" captionText="Figure 11. Maximum likelihood phylogram computed for COI sequences; bootstrap support and Bayesian posterior probabilities are shown when bootstrap supports&gt;65%. Results of delimitation analysis are also shown." figureDoi="http://doi.org/10.5281/zenodo.7778239" httpUri="https://zenodo.org/record/7778239/files/figure.png" pageId="8" pageNumber="307">Figure 11</figureCitation>
) clearly shows all the populations of
<taxonomicName id="86844D1A3C68D3542787FF53FB8EFF11" authorityName="Boeters" authorityYear="1970" box="[970,1103,142,167]" class="Gastropoda" family="Hydrobiidae" genus="Litthabitella" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C68D3542787FF53FB8EFF11" box="[970,1103,142,167]" italics="true" pageId="8" pageNumber="307">Litthabitella</emphasis>
</taxonomicName>
as a distinct, highly supported group (bootstrap 99%). As usual with COI, deeper nodes are not well supported; thus, the relationships among the
<taxonomicName id="86844D1A3C68D3542749FF09FC48FF5B" authorityName="Boeters" authorityYear="1970" box="[772,905,212,237]" class="Gastropoda" family="Hydrobiidae" genus="Litthabitella" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C68D3542749FF09FC48FF5B" box="[772,905,212,237]" italics="true" pageId="8" pageNumber="307">Litthabitella</emphasis>
</taxonomicName>
remain unresolved. In the 18S phylogram of
<bibRefCitation id="25154B683C68D35425F0FF2BFD4FFEB9" author="Szarowska M" box="[445,654,246,271]" pageId="8" pageNumber="307" pagination="99 - 168" refId="ref13640" refString="Szarowska M. 2006. Molecular phylogeny, systematics and morphological character evolution in the Balkan Rissooidea (Caenogastropoda). Folia Malacol. 14 (3): 99 - 168. doi: 10.12657 / folmal. 014.014." type="journal article" year="2006">Szarowska (2006)</bibRefCitation>
,
<taxonomicName id="86844D1A3C68D35426ECFF2BFCD9FEB9" authorityName="Brusina" authorityYear="1870" box="[673,792,246,271]" class="Gastropoda" family="Emmericiidae" genus="Emmericia" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C68D35426ECFF2BFCD9FEB9" box="[673,792,246,271]" italics="true" pageId="8" pageNumber="307">Emmericia</emphasis>
</taxonomicName>
was a sister clade to all the
<taxonomicName id="86844D1A3C68D35424ECFEC4FEEEFE84" authorityName="Stimpson" authorityYear="1865" box="[161,303,281,306]" class="Gastropoda" family="Hydrobiidae" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="family">Hydrobiidae</taxonomicName>
(including Hydrobiinae and Sadlerianinae
<emphasis id="73F0EA8B3C68D3542775FEC4FCB7FE84" box="[824,886,281,306]" italics="true" pageId="8" pageNumber="307">sensu</emphasis>
<bibRefCitation id="25154B683C68D35427CFFEC4FBFEFE84" author="Szarowska M" box="[898,1087,281,306]" pageId="8" pageNumber="307" pagination="99 - 168" refId="ref13640" refString="Szarowska M. 2006. Molecular phylogeny, systematics and morphological character evolution in the Balkan Rissooidea (Caenogastropoda). Folia Malacol. 14 (3): 99 - 168. doi: 10.12657 / folmal. 014.014." type="journal article" year="2006">Szarowska 2006</bibRefCitation>
). The time of divergence of the
<taxonomicName id="86844D1A3C68D3542582FEE1FDBAFEE3" authorityName="Falniowski and Hofman" authorityYear="2023" box="[463,635,316,341]" class="Gastropoda" family="Litthabitellidae" higherTaxonomySource="CoL" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="323" phylum="Mollusca" rank="family">Litthabitellidae</taxonomicName>
was estimated as 8.69 ± 1.87 Mya in the COI phylogram.
</paragraph>
<paragraph id="413B36993C68D354248DFE5CFE30FD64" blockId="8.[160,1156,142,722]" pageId="8" pageNumber="307">
Our 18S phylogram (
<figureCitation id="D9BF2A1C3C68D354258DFE5CFDF3FE2C" box="[448,562,385,410]" captionStart="Figure 12" captionStartId="20.[160,225,1723,1746]" captionTargetBox="[316,999,143,1690]" captionTargetId="figure-1@20.[315,1001,142,1692]" captionTargetPageId="20" captionText="Figure 12. Maximum likelihood phylogram computed for 18S sequences; bootstrap support and Bayesian posterior probabilities are shown when bootstrap supports&gt;65%." figureDoi="http://doi.org/10.5281/zenodo.7778241" httpUri="https://zenodo.org/record/7778241/files/figure.png" pageId="8" pageNumber="307">Figure 12</figureCitation>
) shows a similar picture: a well-supported clade (bootstrap 83%) includes all the
<taxonomicName id="86844D1A3C68D3542668FE79FD70FE0B" authorityName="Stimpson" authorityYear="1865" box="[549,689,420,445]" class="Gastropoda" family="Hydrobiidae" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="family">Hydrobiidae</taxonomicName>
and
<taxonomicName id="86844D1A3C68D35426BFFE79FCB6FE0B" authorityName="Boeters" authorityYear="1970" box="[754,887,420,445]" class="Gastropoda" family="Hydrobiidae" genus="Litthabitella" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C68D35426BFFE79FCB6FE0B" box="[754,887,420,445]" italics="true" pageId="8" pageNumber="307">Litthabitella</emphasis>
</taxonomicName>
, with the latter as the sister clade of all the
<taxonomicName id="86844D1A3C68D35425ECFE1BFDF1FE69" authorityName="Stimpson" authorityYear="1865" box="[417,560,454,479]" class="Gastropoda" family="Hydrobiidae" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="family">Hydrobiidae</taxonomicName>
. The set of available sequences of histone H3 was very restricted, but the relationships (
<figureCitation id="D9BF2A1C3C68D354263BFE34FD28FDB4" box="[630,745,489,514]" captionStart="Figure 13" captionStartId="21.[160,226,1553,1576]" captionTargetBox="[164,1150,143,1518]" captionTargetId="figure-1@21.[163,1153,142,1521]" captionTargetPageId="21" captionText="Figure 13. Maximum likelihood phylogram computed for H3 sequences; bootstrap support and Bayesian posterior probabilities are shown when bootstrap supports&gt;65%." figureDoi="http://doi.org/10.5281/zenodo.7778243" httpUri="https://zenodo.org/record/7778243/files/figure.png" pageId="8" pageNumber="307">Figure 13</figureCitation>
) are similar to those in the 18S phylogram; the well-supported clade (bootstrap 97%) includes
<taxonomicName id="86844D1A3C68D35427CEFDD1FBC9FD93" authorityName="Boeters" authorityYear="1970" box="[899,1032,524,549]" class="Gastropoda" family="Hydrobiidae" genus="Litthabitella" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C68D35427CEFDD1FBC9FD93" box="[899,1032,524,549]" italics="true" pageId="8" pageNumber="307">Litthabitella</emphasis>
</taxonomicName>
as a sister clade of all the
<taxonomicName id="86844D1A3C68D3542523FDF3FE3CFDF1" authorityName="Stimpson" authorityYear="1865" box="[366,509,558,583]" class="Gastropoda" family="Hydrobiidae" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="family">Hydrobiidae</taxonomicName>
, but also of
<taxonomicName id="86844D1A3C68D35426E8FDF3FC7CFDF1" authorityName="Stimpson" authorityYear="1865" box="[677,957,558,583]" class="Gastropoda" family="Cochliopidae" genus="Heleobia" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="genus">
<emphasis id="73F0EA8B3C68D35426E8FDF3FCC7FDF1" box="[677,774,558,583]" italics="true" pageId="8" pageNumber="307">Heleobia</emphasis>
(Cochliopidae)
</taxonomicName>
. The phylogram inferred for the three concatenated loci (
<figureCitation id="D9BF2A1C3C68D35426DEFD8CFCC5FDDC" box="[659,772,593,618]" captionStart="Figure 14" captionStartId="22.[160,225,1353,1376]" captionTargetBox="[163,1152,144,1319]" captionTargetId="figure-80@22.[162,1154,142,1322]" captionTargetPageId="22" captionText="Figure 14. Maximum likelihood phylogram computed for all concatenated sequences; bootstrap support and Bayesian posterior probabilities are shown when bootstrap supports&gt;65%." figureDoi="http://doi.org/10.5281/zenodo.7778245" httpUri="https://zenodo.org/record/7778245/files/figure.png" pageId="8" pageNumber="307">Figure 14</figureCitation>
) also shows the
<taxonomicName id="86844D1A3C68D3542795FD8CFB45FDDC" authorityName="Falniowski and Hofman" authorityYear="2023" box="[984,1156,593,618]" class="Gastropoda" family="Litthabitellidae" higherTaxonomySource="CoL" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="323" phylum="Mollusca" rank="family">Litthabitellidae</taxonomicName>
as a sister taxon to all the
<taxonomicName id="86844D1A3C68D3542597FDA9FDA8FD3B" authorityName="Stimpson" authorityYear="1865" box="[474,617,628,653]" class="Gastropoda" family="Hydrobiidae" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="family">Hydrobiidae</taxonomicName>
, but the clade joining the
<taxonomicName id="86844D1A3C68D35427EEFDA9FB8EFD3B" authorityName="Falniowski and Hofman" authorityYear="2023" box="[931,1103,628,653]" class="Gastropoda" family="Litthabitellidae" higherTaxonomySource="CoL" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="323" phylum="Mollusca" rank="family">Litthabitellidae</taxonomicName>
and
<taxonomicName id="86844D1A3C68D35424EDFD4BFEEEFD19" authorityName="Stimpson" authorityYear="1865" box="[160,303,662,687]" class="Gastropoda" family="Hydrobiidae" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="307" phylum="Mollusca" rank="family">Hydrobiidae</taxonomicName>
is not well supported (only 62%); the clade
<taxonomicName id="86844D1A3C68D3542738FD4BFBE4FD19" authorityName="Falniowski and Hofman" authorityYear="2023" box="[885,1061,662,687]" class="Gastropoda" family="Litthabitellidae" higherTaxonomySource="CoL" kingdom="Animalia" order="Littorinimorpha" pageId="8" pageNumber="323" phylum="Mollusca" rank="family">Litthabitellidae</taxonomicName>
is well supported (bootstrap 100%).
</paragraph>
</subSubSection>
</treatment>
</document>