treatments-xml/data/F3/48/42/F34842265DF98E6CAE07D28F9BC1D50E.xml
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<mods:title id="48CD4DBC4478174F33BE27CA4541BBF6">New records of the endemic Sicilian land snail species Marmorana (Murella) muralis (O. F. Mueller, 1774) from the north of Tunisia (Pulmonata, Gastropoda)</mods:title>
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<mods:namePart id="025D2135911454DE6125D4B9C28BB255">Ezzine, Issaad Kawther</mods:namePart>
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<paragraph id="76CFA3FDCE45AD09E6F3D7C7B802E9B0" pageId="0" pageNumber="131">
<taxonomicName id="FD84F0F3A9351815692F4877B68ADE9D" ID-CoL="3Y826" authority="O. F. Mueller, 1774" authorityName="O. F. Mueller" authorityYear="1774" class="Gastropoda" family="Helicidae" genus="Marmorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Marmorana ((Murella)) muralis" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="muralis" subGenus="(Murella)">
Marmorana (Murella) muralis (O. F.
<normalizedToken id="E08B278E79105BEC8DEF59AE9839A9ED" originalValue="Müller">Mueller</normalizedToken>
, 1774)
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</paragraph>
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<subSubSection id="07AF0F832BE03C1B27FC673215489DAA" pageId="0" pageNumber="131" type="description">
<paragraph id="C4BBEF3768379DA288D0B4EBD84BBF6E" pageId="0" pageNumber="131">Description.</paragraph>
<paragraph id="5C698E65628F9670703E89E7FADCCB3B" pageId="0" pageNumber="131">
Shell medium-sized, depressed globular, thick, solid basic colour beige; large protoconch, clear, smooth, consisting of
<normalizedToken id="BFF2B4C5BA585CC52472C3F4F2FBB537" originalValue="1½">11/2</normalizedToken>
whorls; teleoconch consisting of
<normalizedToken id="82228DDDEE3C046961860B96C3DB56F6" originalValue="3½">31/2</normalizedToken>
slightly flattened whorls, distinctly ribbed; last whorl slightly keeled, larger than the rest whorls, descending towards aperture; aperture sub-spherical; peristome thick, white; suture moderately deep; underside with single interrupted spiral band; moderately ribbed, umbilicus completely covered by the reflected columellar margin (Fig. 1).
</paragraph>
<caption id="75C631963AA0D4BDDBE01BD19EB1D0C6" pageId="0" pageNumber="131">
<paragraph id="95A838B4341423A227900DA1EDC90AAD" pageId="0" pageNumber="131">
Figure 1.
<taxonomicName id="74E297D640121A250E4CDF6F07C8B29B" class="Gastropoda" family="Helicidae" genus="Marmorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Marmorana (Murella) muralis" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="muralis" subGenus="Murella">Marmorana (Murella) muralis</taxonomicName>
(O. F.
<normalizedToken id="F69ABA5A62F609EEE1A61751D3077C31" originalValue="Müller">Mueller</normalizedToken>
, 1774) and
<taxonomicName id="E0FA0FD8E894964A80765CFEFB4120EB" class="Gastropoda" family="Helicidae" genus="Murella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Murella nicollei" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="nicollei">Murella nicollei</taxonomicName>
Pallary, 1926. A Menzel Jemil, Bizerte, 17.ii.2015, NMBE 534231, leg. Ezzine, D = 16.79 mm B Kelibiya, 10.i.2016, NMBE 551457, leg. Ezzine, D = 16.98 mm. Photographs Bochud &amp; Ezzine C
<taxonomicName id="C1C6D6D0EF1FA11E50A2E85448C13AB3" class="Gastropoda" family="Helicidae" genus="Murella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Murella nicollei" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="nicollei">Murella nicollei</taxonomicName>
, Tabarka. Scale bar: D 15.5 mm (copy of the original publication).
</paragraph>
</caption>
<paragraph id="64B30269E86DD25EC005C83FD5253D42" pageId="0" pageNumber="131">Male genital anatomy. Penis club-shaped, thick; epiphallus as long as penis; retractor muscle inserting into the distal part of the epiphallus; flagellum twice the length of epiphallus; penial papilla elongated, with a slit-like pore on one side.</paragraph>
<paragraph id="B5A3E235D11D3D96EF7D06602799E546" pageId="0" pageNumber="131">Female genital anatomy. Dart sac simple, well developed, two glandulae mucosae, non-ramified, inserting into the middle part of the vagina near the base of the dart sac; bursa copulatrix and diverticulum inserting into the proximal part of the vagina (Fig. 2).</paragraph>
<caption id="BE6BDDB1974E50B302880686AB64834A" pageId="0" pageNumber="131">
<paragraph id="7875B9E151A3D031C6907B1776E0A1DF" pageId="0" pageNumber="131">
Figure 2. Anatomy of genital organs of
<taxonomicName id="435EA20BB51F1FEB949CC04868F02674" class="Gastropoda" family="Helicidae" genus="Marmorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Marmorana (Murella) muralis" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="muralis" subGenus="Murella">Marmorana (Murella) muralis</taxonomicName>
(
<normalizedToken id="EBFCACFDB9012AC3FBA3FA24C9BFEB29" originalValue="Müller">Mueller</normalizedToken>
, 1774). A Situs B Details of dart sac C Details of epiphallus D Penial papilla. Abbreviations: A. atrium, Bc. Bursa copulatrix, D. Diverticulum, Ds. Dart sac, E. Epiphallus, F. Flagellum, Gm. Glandulae mucosae, mrp. Penial retractor muscle, P. Penis, Vd. Vas deferens.
</paragraph>
</caption>
<subSection id="79935E6404DA547AF4517852497FCB9C" pageId="0" pageNumber="131" type="haplotype network and genetic diversity">
<paragraph id="3FF9165C94A5F0778AE73A2676712BCB" pageId="0" pageNumber="131">Haplotype network and genetic diversity</paragraph>
<paragraph id="224201DCA5B6E6C25E136BA13F41B57A" pageId="0" pageNumber="131">
Among nine Tunisian and 58 Italian partial COI sequences of
<taxonomicName id="506396C962717AC136692EA3D82CDBE0" lsidName="M. (M.) muralis" pageId="0" pageNumber="131" rank="species" species="muralis" subGenus="M.">M. (M.) muralis</taxonomicName>
(Fig. 3), 46 distinct haplotypes were found, suggesting an extremely high haplotype diversity (Hd = 0.9815) (Fig. 4A). With 45 haplotypes detected, Italian sequences are highly diverse (Hd = 0.9903), while only 3 haplotypes were found in Tunisia (Hd = 0.5596). The haplotype network therefore suggests a relatively low genetic variability of COI sequences from Tunisian specimens compared to sequences from Italian specimens. Tunisian and Italian specimens share two haplotypes: the first is represented by the COI sequences of the samples collected from Manzel Jemil, Manzel Abderrahmen and the sequence from Selinunte. The second haplotype is represented by the sequence of the sample collected at Haouaria and the sequences H1, H4, and H50 from Erice (
<bibRefCitation id="1B0C6352B7FB65ADF673E85D73CBC910" author="Fiorentino, V" journalOrPublisher="Molecular Ecology" pageId="0" pageNumber="131" pagination="170 - 186" title="A question of, time: the land snail Murellamuralis (Gastropoda: Pulmonata) reveals constraints on past ecological speciation." url="https://doi.org/10.1111/mec.12107" volume="22" year="2013">Fiorentino et al. 2013</bibRefCitation>
). The sequences of the samples collected at Kelibiya represent a unique haplotype that is neither shared with the other specimens from Tunisia nor with any of the specimens from Italy. Within the Tunisian sequences, the highest K2P value (0.078) was recorded between the haplotype of the sequence from Haouaria and the sequences from Kelibiya however; the lowest value (0.01) was reported between the sequences from Kelibiya and those from Manzel Jemil and Manzel Abderrahmen. Between Tunisian and Italian populations, the highest K2P value (0.081) was registered between the sequences from Kelibiya and the sequences from Erice (H5, H10) and Monte Monaco (H22, H28, H30, H38, H52). The lowest was recorded between the sequence from Haouaria and the sequences H1, H4 and H50 (
<bibRefCitation id="CC7836E867A4BC94C2AE64A94B05B0D9" author="Fiorentino, V" journalOrPublisher="Molecular Ecology" pageId="0" pageNumber="131" pagination="170 - 186" title="A question of, time: the land snail Murellamuralis (Gastropoda: Pulmonata) reveals constraints on past ecological speciation." url="https://doi.org/10.1111/mec.12107" volume="22" year="2013">Fiorentino et al. 2013</bibRefCitation>
) on the one hand and the sequences from Manzel Jemil- Manzel Abderrahmen and the sequences from Selinunte on the other hand (
<bibRefCitation id="CB118A9FE2A8B245AB7DC00E5FB08EBF" author="Fiorentino, V" journalOrPublisher="Molecular Ecology" pageId="0" pageNumber="131" url="https://doi.org/10.1016/j.ympev.2009.11.024" year="2010">Fiorentino et al. 2010</bibRefCitation>
). The nucleotide divergence within Tunisian population reached a value of 0.0178 but the divergence was slightly higher (0.0316) within Italian population. Moreover, the divergence between Tunisian and Italian populations reached a value of 0.0353.
</paragraph>
<caption id="DB15E77900BFD0A03409C4F5198AE92C" pageId="0" pageNumber="131">
<paragraph id="0690AD61D7D7015C3014BE5856FB4BDF" pageId="0" pageNumber="131">Figure 3. Localisation of Tunisian and Italian specimens used in the study.</paragraph>
</caption>
<caption id="77C2F6EDD54F5F86981DB9862920158E" pageId="0" pageNumber="131">
<paragraph id="0B373764E51DA380E5A606B0F6452C87" pageId="0" pageNumber="131">
Figure 4. Haplotype network showing the relationships among Italian and Tunisian specimens of
<taxonomicName id="CA77AC11E7D3D2884894313863B96356" lsidName="M. (M.) muralis" pageId="0" pageNumber="131" rank="species" species="muralis" subGenus="M.">M. (M.) muralis</taxonomicName>
. A Haplotype network based on partial COI sequences B Haplotype network based on partial 16S sequences C Haplotype network based on concatenated partial COI and 16S sequences.
</paragraph>
</caption>
<paragraph id="C8E303ACD902ACBA7F4111A931177AD4" pageId="0" pageNumber="131">The analysis of ten Tunisian and six Italian 16S partial fragments shows five haplotypes suggesting a low haplotype diversity (0.450) (Fig. 4B). Sequences of Italian specimens represent four haplotypes (Hd = 0.80), while sequences from Tunisian specimens represent only two haplotypes (Hd = 0.20). Tunisian and Italian samples share one haplotype, which was represented by sequences of Tunisian specimens from Kelibiya, Manzel Jemil, Manzel Abderrahmen and sequences of Italian specimens from Lazio, Selinunte and Marsala. The sequences of specimens from Haouaria, Fiumedinisi, Caltabellotta and Joppolo represented four different haplotypes. The maximum value of K2P distance (0.02), within Tunisian sequences, is recorded between the sequence from Haouaria and the rest. While the maximum value recorded between Tunisian and Italian populations is 0.044 between the sequence from Haouaria and the sequences from Fiumedinisi and Caltabellotta. The nucleotide divergence of the 16S partial fragment is remarkably low within Tunisian population (0.00409), as well as, between Tunisian and Italian populations (0.00828).</paragraph>
<paragraph id="D9B6AB1F20719F477833EB3D34387DBC" pageId="0" pageNumber="131">The analysis of nine Tunisian and six Italian concatenated sequences (COI, 16S) recovered seven different haplotypes among them (Fig. 4C). One haplotype is shared by Tunisian and Italian populations. The rest is divided into two Tunisian and four Italian haplotypes.</paragraph>
</subSection>
<subSection id="151A68BAD0D866976703A10E29BEF3B5" pageId="0" pageNumber="131" type="phylogeny">
<paragraph id="3351B87B18AA9DD781A513B782AF4AB1" pageId="0" pageNumber="131">Phylogeny</paragraph>
<paragraph id="1E5F584EDB3FDFCE60C531043FF37AB3" pageId="0" pageNumber="131">
The topology, obtained by Bayesian inference based on the concatenated COI and 16S data set was rooted with
<taxonomicName id="777ECBCA99DFCB055473672A2BA92F92" class="Gastropoda" family="Helicidae" genus="Macularia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macularia sylvatica" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="sylvatica">Macularia sylvatica</taxonomicName>
and
<taxonomicName id="E5091D9D2ABB07C09C745F6998E0B9AC" class="Gastropoda" family="Helicidae" genus="Macularia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macularia niciensis" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="niciensis">Macularia niciensis</taxonomicName>
as outgroups (Fig. 5). The
<taxonomicName id="FBD779F49EE6AF3C2FE4F3BAE818008A" class="Gastropoda" family="Helicidae" genus="Marmorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Marmorana" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="genus">Marmorana</taxonomicName>
species form two opposite clades well supported (PP = 1): The first one is formed by the samples of
<taxonomicName id="999C4A41788367A07B7492E836336CE7" lsidName="M. (M.) serpentina" pageId="0" pageNumber="131" rank="species" species="serpentina" subGenus="M.">M. (M.) serpentina</taxonomicName>
and the second is formed by both Tunisian and Italian
<taxonomicName id="FDBEA0EC5CE7FA783CCCFE4EB75CF957" lsidName="M. (M.) muralis" pageId="0" pageNumber="131" rank="species" species="muralis" subGenus="M.">M. (M.) muralis</taxonomicName>
. Within the
<taxonomicName id="C674AD3139B1950146A6BECDB8727EBC" lsidName="M. (M.) muralis" pageId="0" pageNumber="131" rank="species" species="muralis" subGenus="M.">M. (M.) muralis</taxonomicName>
clade,
<taxonomicName id="10055AF83D30729B91001C1E20CC55A5" lsidName="M. globularis" pageId="0" pageNumber="131" rank="species" species="globularis">M. cf. globularis</taxonomicName>
and the
<taxonomicName id="EC1C3B9BD34172DC77C9A930E451B51D" lsidName="M. (M.) muralis" pageId="0" pageNumber="131" rank="species" species="muralis" subGenus="M.">M. (M.) muralis</taxonomicName>
of Fiumedinisi are situated at the base of the clade with a high value of posterior probability (1-0.93). The rest sequences form three well supported clades. The first is composed by the sequences of Marsala and Lazio, the second is formed by the sequences of Kelibiya, Joppolo, and Selinunte (0.92) and the third clade is formed by the sequences of Manzel Abderrahmen, Manzel Jemil, Haouaria and Erice (0.82).
</paragraph>
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<paragraph id="33052916AE4ADD8C4B6FB022761E070C" pageId="0" pageNumber="131">
Figure 5. Bayesian 50% majority-rule consensus tree based on the analysis of concatenated partial COI and 16S sequences showing the relationships among Tunisian and Italian
<taxonomicName id="035CC4AE71EC24C7587DCD311B51C8DA" class="Gastropoda" family="Helicidae" genus="Marmorana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Marmorana (Murella) muralis" order="Stylommatophora" pageId="0" pageNumber="131" phylum="Mollusca" rank="species" species="muralis" subGenus="Murella">Marmorana (Murella) muralis</taxonomicName>
samples.
</paragraph>
</caption>
</subSection>
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</treatment>
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