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<mods:title id="9910B150A9EABE9B8FA96455977A0EDE">Revision of the dwarf cannibal snails (Nata s. l.) of southern Africa — Nata s. s. and Natella (Mollusca: Gastropoda: Rhytididae), with description of three new species</mods:title>
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<treatment id="03E687FCE03DD776FF1A142B086A9A1C" ID-DOI="http://doi.org/10.5281/zenodo.5615862" ID-GBIF-Taxon="119319669" ID-Zenodo-Dep="5615862" LSID="urn:lsid:plazi:treatment:03E687FCE03DD776FF1A142B086A9A1C" httpUri="http://treatment.plazi.org/id/03E687FCE03DD776FF1A142B086A9A1C" lastPageId="49" lastPageNumber="50" pageId="48" pageNumber="49">
<subSubSection id="C3556561E03DD777FF1A142B0AFF9D48" box="[151,501,1350,1376]" pageId="48" pageNumber="49" type="nomenclature">
<paragraph id="8BF036EAE03DD777FF1A142B0AFF9D48" blockId="48.[151,501,1350,1376]" box="[151,501,1350,1376]" pageId="48" pageNumber="49">
<heading id="D0B88186E03DD777FF1A142B0AFF9D48" bold="true" box="[151,501,1350,1376]" fontSize="11" level="1" pageId="48" pageNumber="49" reason="1">
<emphasis id="B93BEAF8E03DD777FF1A142B0AFF9D48" bold="true" box="[151,501,1350,1376]" pageId="48" pageNumber="49">
Genus
<taxonomicName id="4C4F4D69E03DD777FF63142B0AFF9D48" ID-CoL="7P9JX" authority="Watson, 1934" authorityName="Watson" authorityYear="1934" box="[238,501,1350,1376]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FF63142B0A4C9D48" bold="true" box="[238,326,1350,1376]" italics="true" pageId="48" pageNumber="49">Natella</emphasis>
<bibRefCitation id="EFDE4B1BE03DD777FEC0142B0AFF9D48" author="Watson" box="[333,501,1350,1376]" pageId="48" pageNumber="59" refString="Watson, H. (1934) Natalina and other South African snails. Proceedings of the Malacological Society of London, 21, 150 - 193." type="journal article" year="1934">Watson, 1934</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C3556561E03DD777FF1A14E70E6F9D88" box="[151,1381,1418,1440]" pageId="48" pageNumber="49" type="reference_group">
<paragraph id="8BF036EAE03DD777FF1A14E70E6F9D88" blockId="48.[151,1381,1418,1440]" box="[151,1381,1418,1440]" pageId="48" pageNumber="49">
<treatmentCitationGroup id="AB5F11C4E03DD777FF1A14E70E6F9D88" box="[151,1381,1418,1440]" pageId="48" pageNumber="49">
<taxonomicName id="4C4F4D69E03DD777FF1A14E70AA39D88" ID-CoL="7P9JX" authority="Watson, 1934: 160" authorityName="Watson" authorityPageNumber="160" authorityYear="1934" box="[151,425,1418,1440]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FF1A14E70BE89D88" box="[151,226,1418,1440]" italics="true" pageId="48" pageNumber="49">Natella</emphasis>
<treatmentCitation id="0AEE10FBE03DD777FF6514E60AA39D88" author="Watson" box="[232,425,1418,1440]" page="160" pageId="48" pageNumber="49" year="1934">
<bibRefCitation id="EFDE4B1BE03DD777FF6514E60A7C9D88" author="Watson" box="[232,374,1418,1440]" pageId="48" pageNumber="59" refString="Watson, H. (1934) Natalina and other South African snails. Proceedings of the Malacological Society of London, 21, 150 - 193." type="journal article" year="1934">Watson, 1934</bibRefCitation>
: 160
</treatmentCitation>
</taxonomicName>
.
<typeStatus id="54F48848E03DD777FE3F14E60AEE9D88" box="[434,484,1419,1440]" pageId="48" pageNumber="49">Type</typeStatus>
species (by original designation):
<taxonomicName id="4C4F4D69E03DD777FCCF14E70E6B9D88" authority="Melvill &amp; Ponsonby, 1891" authorityName="Melvill &amp; Ponsonby" authorityYear="1891" box="[834,1377,1418,1440]" class="Gastropoda" family="Helicidae" genus="Helix" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="species" species="viridescens" subGenus="Patula">
<emphasis id="B93BEAF8E03DD777FCCF14E708739D88" box="[834,889,1418,1440]" italics="true" pageId="48" pageNumber="49">Helix</emphasis>
(
<emphasis id="B93BEAF8E03DD777FC0A14E708C19D88" box="[903,971,1418,1440]" italics="true" pageId="48" pageNumber="49">Patula</emphasis>
)
<emphasis id="B93BEAF8E03DD777FC5714E70F469D88" box="[986,1100,1418,1440]" italics="true" pageId="48" pageNumber="49">viridescens</emphasis>
<bibRefCitation id="EFDE4B1BE03DD777FBDF14E70E6B9D88" author="Melvill" box="[1106,1377,1418,1440]" pageId="48" pageNumber="57" refString="Melvill, J. C. &amp; Ponsonby, J. H. (1891) Descriptions of nine new terrestrial and fluviatile mollusks from South Africa. Annals and Magazine of Natural History, Series 6, 8, 237 - 240. http: // dx. doi. org / 10.1080 / 00222939109460425" type="journal article" year="1891">Melvill &amp; Ponsonby, 1891</bibRefCitation>
</taxonomicName>
.
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</paragraph>
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<subSubSection id="C3556561E03DD777FF1A14A70FB09EBB" pageId="48" pageNumber="49" type="diagnosis">
<paragraph id="8BF036EAE03DD777FF1A14A70FB09EBB" blockId="48.[151,1437,1482,2035]" pageId="48" pageNumber="49">
<emphasis id="B93BEAF8E03DD777FF1A14A70A1B9DCB" bold="true" box="[151,273,1482,1507]" pageId="48" pageNumber="49">Diagnosis.</emphasis>
Shell small (adult diameter
<emphasis id="B93BEAF8E03DD777FDC614A1096C9DCB" box="[587,614,1484,1507]" italics="true" pageId="48" pageNumber="49">ca</emphasis>
5.0 mm), thin, discoidal, broadly umbilicate, glossy, pale yellowish; axial sculpture weak, spiral sculpture lacking; protoconch smooth. Radula beloglossan; labial palps not evident; penis stout, epiphallus absent, vas deferens short and not reflexed, inserting apically into penis; penis lumen not papillate, apical verge present; retractor muscle of right optic tentacle passes to left of penis; vagina broad, no atrial diverticulum or oviduct caecum present; kidney ovate to bean-shaped, primary ureter reflexed around anterior of kidney; suprapedal mucus gland not convoluted, the posterior portion lacking glandular tissue.
</paragraph>
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<paragraph id="8BF036EAE03DD777FF4A17F00BEA9F6B" blockId="48.[151,1437,1482,2035]" pageId="48" pageNumber="49">
<emphasis id="B93BEAF8E03DD777FF4A17F00A319E9E" bold="true" box="[199,315,1693,1718]" pageId="48" pageNumber="49">Remarks.</emphasis>
Watsons separation of '
<taxonomicName id="4C4F4D69E03DD777FDDB17F209869E9E" box="[598,652,1695,1718]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FDDB17F209869E9E" box="[598,652,1695,1718]" italics="true" pageId="48" pageNumber="49">Nata</emphasis>
</taxonomicName>
'
<taxonomicName id="4C4F4D69E03DD777FD1117F008109E9E" box="[668,794,1693,1718]" class="Gastropoda" family="Helicidae" genus="Helix" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="species" species="viridescens">
<emphasis id="B93BEAF8E03DD777FD1117F008109E9E" box="[668,794,1693,1718]" italics="true" pageId="48" pageNumber="49">viridescens</emphasis>
</taxonomicName>
in to a distinct, monotypic section [=subgenus]
<taxonomicName id="4C4F4D69E03DD777FAC417F00A4E9EF1" authority="Watson 1934" authorityName="Watson" authorityYear="1934" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FAC417F00E969E9E" box="[1353,1436,1693,1718]" italics="true" pageId="48" pageNumber="49">Natella</emphasis>
(
<bibRefCitation id="EFDE4B1BE03DD777FF1317AC0A369EF1" author="Watson" box="[158,316,1729,1753]" pageId="48" pageNumber="59" refString="Watson, H. (1934) Natalina and other South African snails. Proceedings of the Malacological Society of London, 21, 150 - 193." type="journal article" year="1934">Watson 1934</bibRefCitation>
)
</taxonomicName>
, is strongly supported by both the morphological data presented here and by molecular data (
<bibRefCitation id="EFDE4B1BE03DD777FF1317890ADB9ED5" author="Moussalli" box="[158,465,1764,1789]" pageId="48" pageNumber="58" refString="Moussalli, A. &amp; Herbert, D. G. (2016) Deep molecular divergence and exceptional morphological stasis in dwarf cannibal snails Nata sensu lato Watson, 1934 (Rhytididae) of southern Africa. Molecular Phylogenetics and Evolution, 95, 100 - 115. http: // dx. doi. org / 10.1016 / j. ympev. 2015.11.003" type="journal article" year="2016">Moussalli &amp; Herbert 2016</bibRefCitation>
). Furthermore, since the latter data show
<emphasis id="B93BEAF8E03DD777FC2017880F069ED5" box="[941,1036,1764,1789]" italics="true" pageId="48" pageNumber="49">
<taxonomicName id="4C4F4D69E03DD777FC20178808E99ED4" box="[941,995,1765,1788]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus" sensu="lato">Nata</taxonomicName>
<taxonomicNameLabel id="A2085783E03DD777FC6117890F069ED5" box="[1004,1036,1764,1789]" pageId="48" pageNumber="49" sensu="lato">s.l.</taxonomicNameLabel>
</emphasis>
to be a deeply divergent and quite possibly non-monophyletic taxon (
<taxonomicName id="4C4F4D69E03DD777FD93166A097B9F08" box="[542,625,1799,1824]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FD93166A097B9F08" box="[542,625,1799,1824]" italics="true" pageId="48" pageNumber="49">Natella</emphasis>
</taxonomicName>
is not sister to
<emphasis id="B93BEAF8E03DD777FC91166508779F37" box="[796,893,1800,1823]" italics="true" pageId="48" pageNumber="49">
<taxonomicName id="4C4F4D69E03DD777FC91166508599F37" box="[796,851,1800,1823]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus" sensu="stricto">Nata</taxonomicName>
<taxonomicNameLabel id="A2085783E03DD777FCD7166508779F37" box="[858,893,1800,1823]" pageId="48" pageNumber="49" sensu="stricto">s.s.</taxonomicNameLabel>
</emphasis>
), we consider
<taxonomicName id="4C4F4D69E03DD777FBAE166A0F7C9F08" box="[1059,1142,1799,1824]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FBAE166A0F7C9F08" box="[1059,1142,1799,1824]" italics="true" pageId="48" pageNumber="49">Natella</emphasis>
</taxonomicName>
should be treated as a full genus.
</paragraph>
<paragraph id="8BF036EAE03DD776FF4A162208079931" blockId="48.[151,1437,1482,2035]" lastBlockId="49.[151,1437,151,564]" lastPageId="49" lastPageNumber="50" pageId="48" pageNumber="49">
In addition to the characters given in the above diagnosis, Watson also listed the presence of a rachidian tooth in the radula as a feature distinctive of
<taxonomicName id="4C4F4D69E03DD777FDDE161C09AF9FA2" box="[595,677,1905,1930]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FDDE161C09AF9FA2" box="[595,677,1905,1930]" italics="true" pageId="48" pageNumber="49">Natella</emphasis>
</taxonomicName>
. While we can confirm that a rachidian tooth is present in
<taxonomicName id="4C4F4D69E03DD777FACF161C0E9E9FA2" box="[1346,1428,1905,1930]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FACF161C0E9E9FA2" box="[1346,1428,1905,1930]" italics="true" pageId="48" pageNumber="49">Natella</emphasis>
</taxonomicName>
, this can no longer be considered a distinctive character of the genus since such a tooth is also present in
<taxonomicName id="4C4F4D69E03DD777FAE816F80A299FF8" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="48" pageNumber="49" phylum="Mollusca" rank="species" species="aequiplicata" status="sp. nov.">
<emphasis id="B93BEAF8E03DD777FAE816F80A299FF8" italics="true" pageId="48" pageNumber="49">Nata aequiplicata</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2085783E03DD777FEA616DA0A8F9FF8" box="[299,389,1975,2000]" pageId="48" pageNumber="49" rank="species">
<emphasis id="B93BEAF8E03DD777FEA616DA0A8F9FF8" bold="true" box="[299,389,1975,2000]" pageId="48" pageNumber="49">sp. nov.</emphasis>
</taxonomicNameLabel>
, which genital morphology and molecular data show to belong to
<taxonomicName id="4C4F4D69E03DD777FBFB16D50FA69FE7" box="[1142,1196,1976,1999]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="48" pageNumber="49" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03DD777FBFB16D50FA69FE7" box="[1142,1196,1976,1999]" italics="true" pageId="48" pageNumber="49">Nata</emphasis>
</taxonomicName>
. (It is also present in some unusual specimens tentatively identified with
<taxonomicName id="4C4F4D69E03DD777FD5516B1089B9FDB" box="[728,913,2010,2035]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="48" pageNumber="49" phylum="Mollusca" rank="species" species="tarachodes">
<emphasis id="B93BEAF8E03DD777FD5516B1089B9FDB" box="[728,913,2010,2035]" italics="true" pageId="48" pageNumber="49">Nata tarachodes</emphasis>
</taxonomicName>
.)
<collectingCountry id="F358767AE03DD777FC2416B608C29FDB" box="[937,968,2011,2035]" name="American Samoa" pageId="48" pageNumber="49">As</collectingCountry>
stated in the Introduction, the presence of a rachidian tooth may represent an ancestral character state retained in juveniles and small-shelled species. The presence/absence of a rachidian tooth may thus not be phylogenetically informative. Watson also noted that the bursa copulatrix duct (as spermatheca duct) of
<taxonomicName id="4C4F4D69E03CD776FD3B11B3080398DF" box="[694,777,222,247]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="49" pageNumber="50" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03CD776FD3B11B3080398DF" box="[694,777,222,247]" italics="true" pageId="49" pageNumber="50">Natella</emphasis>
</taxonomicName>
was longer than in species of
<emphasis id="B93BEAF8E03CD776FBD711B20FBE98DE" box="[1114,1204,223,246]" italics="true" pageId="49" pageNumber="50">
<taxonomicName id="4C4F4D69E03CD776FBD711B20F9A98DE" box="[1114,1168,223,246]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="49" pageNumber="50" phylum="Mollusca" rank="genus" sensu="stricto">Nata</taxonomicName>
<taxonomicNameLabel id="A2085783E03CD776FB1511B20FBE98DE" box="[1176,1204,223,246]" pageId="49" pageNumber="50" sensu="stricto">s.s</taxonomicNameLabel>
</emphasis>
., but he provided no illustration of this and we do not find this to be the case.
</paragraph>
<paragraph id="8BF036EAE03CD776FF4A1048094199C5" blockId="49.[151,1437,151,564]" pageId="49" pageNumber="50">
A further unusual and possibly distinctive feature of the buccal apparatus of
<taxonomicName id="4C4F4D69E03CD776FBC110490F959915" box="[1100,1183,292,317]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="49" pageNumber="50" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03CD776FBC110490F959915" box="[1100,1183,292,317]" italics="true" pageId="49" pageNumber="50">Natella</emphasis>
</taxonomicName>
is its possession of a proboscis. First observed in a drowning individual as an extended oral tube with the radula near its tip (
<figureCitation id="13742A6FE03CD776FAB9102A0E859948" box="[1332,1423,327,352]" captionStart="FIGURE 2. A" captionStartId="6.[151,250,1952,1974]" captionTargetBox="[154,1433,1444,1921]" captionTargetId="figure@6.[151,1436,1439,1923]" captionTargetPageId="6" captionText="FIGURE 2. A, Natella viridescens (Melvill &amp; Ponsonby, 1891), drowning individual with proboscis everted, Ithala Game Res., KwaZulu-Natal, shell diameter 3.7 mm (NMSA W 7263). B, Nata vernicosa (Krauss, 1848), mating pair, Qora River mouth, E. Cape, respective shell diameter 20.5 mm and 14 mm (NMSA W 4015)." httpUri="https://zenodo.org/record/266546/files/figure.png" pageId="49" pageNumber="50">Fig. 2A</figureCitation>
), this structure is visible when retracted as a tube linking the mouth to the anterior end of the buccal mass, in the vicinity of the cerebral ganglia and anterior nerve ring. A similar structure has not been seen in species of
<taxonomicName id="4C4F4D69E03CD776FABB10E20E67998E" box="[1334,1389,399,422]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="49" pageNumber="50" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03CD776FABB10E20E67998E" box="[1334,1389,399,422]" italics="true" pageId="49" pageNumber="50">Nata</emphasis>
</taxonomicName>
and observations of the feeding behaviour of
<taxonomicName id="4C4F4D69E03CD776FDEB10DF081B99E1" box="[614,785,434,457]" class="Gastropoda" family="Rhytididae" genus="Nata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pulmonata" pageId="49" pageNumber="50" phylum="Mollusca" rank="species" species="vernicosa">
<emphasis id="B93BEAF8E03CD776FDEB10DF081B99E1" box="[614,785,434,457]" italics="true" pageId="49" pageNumber="50">Nata vernicosa</emphasis>
</taxonomicName>
suggest that there is very limited protrusion of the buccal tube beyond the mouth during feeding.
</paragraph>
<paragraph id="8BF036EAE03CD776FF4A109A086A9A1C" blockId="49.[151,1437,151,564]" pageId="49" pageNumber="50">
<taxonomicName id="4C4F4D69E03CD776FF4A109A0A109A38" box="[199,282,503,528]" class="Gastropoda" family="Rhytididae" genus="Natella" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="49" pageNumber="50" phylum="Mollusca" rank="genus">
<emphasis id="B93BEAF8E03CD776FF4A109A0A109A38" box="[199,282,503,528]" italics="true" pageId="49" pageNumber="50">Natella</emphasis>
</taxonomicName>
is monotypic and is endemic to southern Africa. To date it has been recorded only from north-eastern
<collectingCountry id="F358767AE03CD776FF1A13760A209A1C" box="[151,298,539,564]" name="South Africa" pageId="49" pageNumber="50">South Africa</collectingCountry>
, though its range may well extend into
<collectingCountry id="F358767AE03CD776FD6E137608569A1C" box="[739,860,539,564]" name="Swaziland" pageId="49" pageNumber="50">Swaziland</collectingCountry>
.
</paragraph>
</subSubSection>
</treatment>
</document>