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<mods:title id="568D7B88BA5BC227CB5ABD539DEA1E91">Where the snails have no name: a molecular phylogeny of Raphitomidae (Neogastropoda: Conoidea) uncovers vast unexplored diversity in the deep seas of temperate southern and eastern Australia</mods:title>
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<mods:namePart id="9269DA2BBD76BCADB2F3163944E578B2">Criscione, Francesco</mods:namePart>
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<mods:namePart id="E7FBB36BA0D49F1DA7B4C2D8DBFB928C">Hallan, Anders</mods:namePart>
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<mods:namePart id="E68FB35B14B1A8DFB48F7FA447804A77">Puillandre, Nicolas</mods:namePart>
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<mods:namePart id="4883ADD88D25E8176C8FB6BC767D0DD6">Fedosov, Alexander</mods:namePart>
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<subSubSection id="4BFB05CAFFBCF843FF0EFC70FFEC3FA2" box="[233,709,993,1017]" pageId="25" pageNumber="986" type="nomenclature">
<paragraph id="035E5641FFBCF843FF0EFC70FFEC3FA2" blockId="25.[233,709,993,1017]" box="[233,709,993,1017]" pageId="25" pageNumber="986">
<heading id="5816E12DFFBCF843FF0EFC70FFEC3FA2" box="[233,709,993,1017]" centered="true" fontSize="9" level="2" pageId="25" pageNumber="986" reason="2">
<taxonomicName id="C4E12DC2FFBCF843FF0EFC70FF733FAC" ID-CoL="7Q9DL" authority="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov, 2021" authorityName="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov" authorityYear="2021" box="[233,602,993,1017]" class="Gastropoda" family="Raphitomidae" genus="Austrotheta" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="993" phylum="Mollusca" rank="species" species="crassidentata" status="sp. nov.">AUSTROTHETA CRASSIDENTATA</taxonomicName>
<emphasis id="31958A53FFBCF843FD85FC75FFEC3FA2" bold="true" box="[610,709,993,1017]" pageId="25" pageNumber="986">
<taxonomicNameLabel id="2AA63728FFBCF843FD85FC75FFEC3FA2" box="[610,709,993,1017]" pageId="25" pageNumber="986" rank="species">SP. NOV.</taxonomicNameLabel>
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<subSubSection id="4BFB05CAFFBCF843FE61FB9EF9B93B15" pageId="25" pageNumber="986" type="description">
<paragraph id="035E5641FFBCF843FE61FB9EFF0E3873" blockId="25.[390,551,1039,1064]" box="[390,551,1039,1064]" pageId="25" pageNumber="986">
(
<figureCitation id="9BDA4AC4FFBCF843FE68FB9EFCC73873" box="[399,494,1039,1064]" captionStart="Figure 3" captionStartId="15.[163,241,1570,1592]" captionTargetBox="[163,1443,195,1530]" captionTargetId="figure-87@15.[163,1443,195,1530]" captionTargetPageId="15" captionText="Figure 3. Shells of species described herein. A, Notothauma magnifica, holotype AMS C.532707; B, Fusobela parvioculata, holotype TMAG E59231; C, Aplotoma brevitentaculata, holotype AMS C.571635; D, Biconitoma cretosa, holotype AMS C.482313. E, Pagodibela maia, holotype AMS C.571678; F, Austrobela rufa, holotype AMS C.571709; G, Glaciotomella investigator, holotypeAMSC.571621;H, Austrotheta crassidentata, holotype AMSC.519302.Scale bar20 mm (A), 10 mm (B–H)." pageId="25" pageNumber="986">FIGS 3H</figureCitation>
, 
<figureCitation id="9BDA4AC4FFBCF843FE1BFB9EFF363873" box="[508,543,1039,1064]" captionStart="Figure 6" captionStartId="18.[145,227,1378,1400]" captionTargetBox="[145,1425,195,1338]" captionTargetId="figure-24@18.[145,1425,195,1338]" captionTargetPageId="18" captionText="Figure 6. Hypodermic teeth of species described herein. A, Aplotoma brevitentaculata, holotype AMS C.571635; B, Gladiobela angulata, paratype AMS C.571737; C, Pueridaphne cirrisulcata, paratype AMS C.571609; D: Austrobela rufa, holotype AMS C.571709 (left), paratype AMS C.571679 (right); E, Pagodibela maia, holotype AMS C.571678; F, Fusobela parvioculata, holotype TMAG E59231; G, Biconitoma cretosa, holotype AMS C.482313; H, Austrotheta crassidentata, holotype AMS C.519302. Scale bars = 50 μm." pageId="25" pageNumber="986">6H</figureCitation>
)
</paragraph>
<paragraph id="035E5641FFBCF843FF44FBC8FE2F38D7" blockId="25.[163,780,1112,1164]" pageId="25" pageNumber="986">
<emphasis id="31958A53FFBCF843FF44FBC8FCF63835" box="[163,479,1112,1134]" italics="true" pageId="25" pageNumber="986">Z o o B a n k r e g i s t r a t i o n</emphasis>
: u r n: l s i d: z o o b a n k. org:act: 
<uuid id="77476C94FFBCF843FF15FBE7FE2A38D7" box="[242,771,1142,1164]" pageId="25" pageNumber="986">4EF95055-B0D2-4151-89B1-762BD62DF8DE</uuid>
.
</paragraph>
<paragraph id="035E5641FFBCF843FF44FB27FFB83953" blockId="25.[163,779,1205,1289]" pageId="25" pageNumber="986">
<emphasis id="31958A53FFBCF843FF44FB27FC003890" box="[163,297,1205,1227]" italics="true" pageId="25" pageNumber="986">H o l o t y p e:</emphasis>
A u s t r a l i a, T a s m a n i a, F r e y c i n e t Commonwealth Marine Reserve, 
<quantity id="C419FBA4FFBCF843FDDDFB45FFBC38B2" box="[570,661,1236,1257]" metricMagnitude="3" metricUnit="m" metricValue="2.82" pageId="25" pageNumber="986" unit="m" value="2820.0">2820 m</quantity>
, IN2017_ V03_004, (–41.731, 149.12), AMS C.519302.
</paragraph>
<paragraph id="035E5641FFBCF843FF44FAA0FFE1391C" blockId="25.[163,712,1329,1351]" box="[163,712,1329,1351]" pageId="25" pageNumber="986">
<emphasis id="31958A53FFBCF843FF44FAA0FC1D391D" box="[163,308,1329,1350]" italics="true" pageId="25" pageNumber="986">Distribution:</emphasis>
Known only from the 
<typeStatus id="DC5AE8E3FFBCF843FDDFFAA3FF40391C" box="[568,617,1330,1351]" pageId="25" pageNumber="986">type</typeStatus>
locality.
</paragraph>
<paragraph id="035E5641FFBCF843FF44FAE1FF4E3998" blockId="25.[163,777,1392,1475]" pageId="25" pageNumber="986">
<emphasis id="31958A53FFBCF843FF44FAE1FC0339DE" box="[163,298,1392,1413]" italics="true" pageId="25" pageNumber="986">Etymology:</emphasis>
The epithet is composed of the Latin adjectives 
<emphasis id="31958A53FFBCF843FECCFA1EFCAE39FF" box="[299,391,1423,1444]" italics="true" pageId="25" pageNumber="986">crassus</emphasis>
, thick, and 
<emphasis id="31958A53FFBCF843FDC0FA1FFFBA39F8" box="[551,659,1422,1443]" italics="true" pageId="25" pageNumber="986">dentatus</emphasis>
, toothed, referring to the thick hypodermic tooth.
</paragraph>
<paragraph id="035E5641FFBCF843FF44FA6EFC063A4D" blockId="25.[163,303,1535,1558]" box="[163,303,1535,1558]" pageId="25" pageNumber="986">
<emphasis id="31958A53FFBCF843FF44FA6EFC063A4D" box="[163,303,1535,1558]" italics="true" pageId="25" pageNumber="986">Description</emphasis>
</paragraph>
<paragraph id="035E5641FFBCF843FF44F9B7F9753E10" blockId="25.[163,780,1574,1872]" lastBlockId="25.[827,1444,197,894]" pageId="25" pageNumber="986">
Shell (
<figureCitation id="9BDA4AC4FFBCF843FF16F9B7FC603A67" box="[241,329,1574,1596]" captionStart="Figure 3" captionStartId="15.[163,241,1570,1592]" captionTargetBox="[163,1443,195,1530]" captionTargetId="figure-87@15.[163,1443,195,1530]" captionTargetPageId="15" captionText="Figure 3. Shells of species described herein. A, Notothauma magnifica, holotype AMS C.532707; B, Fusobela parvioculata, holotype TMAG E59231; C, Aplotoma brevitentaculata, holotype AMS C.571635; D, Biconitoma cretosa, holotype AMS C.482313. E, Pagodibela maia, holotype AMS C.571678; F, Austrobela rufa, holotype AMS C.571709; G, Glaciotomella investigator, holotypeAMSC.571621;H, Austrotheta crassidentata, holotype AMSC.519302.Scale bar20 mm (A), 10 mm (B–H)." pageId="25" pageNumber="986">Fig. 3H</figureCitation>
) (SL = 20.9, SW = 9.6) fusiform, thinwalled, semitranslucent to opaque. Protoconch orange, multispiral (at least 2.5 whorls), with arcuate cordlets on adapical half to two-thirds of whorl, with diagonally cancellate sculpture below. Teleoconch of 4.4 whorls; subsutural ramp distinctly concave in early whorls, in more mature whorls less distinct and with less marked concavity.Whorl profile with prominent shoulder on early teleoconch whorls, situated at adapical third of whorl; in penultimate whorl more rounded, in last adult whorl indistinct; whorl periphery nearly cylindrical in early teleconch,more convex in mature whorls.Early teleoconch whorls with about 12 sharp, weakly opisthocline axials, vanishing well above suture, indistinct in later whorls. Spiral sculpture of dense, slightly undulating striae pronounced throughout last whorl. Microsculpture of collabral growth lines, forming distinct, raised cordlets on subsutural ramp in early whorls, weaker in more mature whorls. Last adult whorl evenly convex below subsutural ramp, with long, slender siphonal canal. Aperture wide, pyriform, about half of shell length. Inner lip with whitish callus, straight. Outer lip thin, unsculptured. Anal sinus shallow, weakly U-shaped.
</paragraph>
<paragraph id="035E5641FFBCF843FCB4FDC5FE823EFC" blockId="25.[827,1444,197,894]" pageId="25" pageNumber="986">Head wide, eyes small. Muscular bulb extremely large; proboscis short, wide; radular sac thin, bearing few teeth.</paragraph>
<paragraph id="035E5641FFBCF843FCB4FD21F9E23F25" blockId="25.[827,1444,197,894]" pageId="25" pageNumber="986">
Radula (
<figureCitation id="9BDA4AC4FFBCF843FC25FD21F9083E9D" box="[962,1057,688,710]" captionStart="Figure 6" captionStartId="18.[145,227,1378,1400]" captionTargetBox="[145,1425,195,1338]" captionTargetId="figure-24@18.[145,1425,195,1338]" captionTargetPageId="18" captionText="Figure 6. Hypodermic teeth of species described herein. A, Aplotoma brevitentaculata, holotype AMS C.571635; B, Gladiobela angulata, paratype AMS C.571737; C, Pueridaphne cirrisulcata, paratype AMS C.571609; D: Austrobela rufa, holotype AMS C.571709 (left), paratype AMS C.571679 (right); E, Pagodibela maia, holotype AMS C.571678; F, Fusobela parvioculata, holotype TMAG E59231; G, Biconitoma cretosa, holotype AMS C.482313; H, Austrotheta crassidentata, holotype AMS C.519302. Scale bars = 50 μm." pageId="25" pageNumber="986">Fig. 6H</figureCitation>
) of thick, straight cylindrical hypodermic teeth exceeding 175 µm in length; two weak distal, lateral barbs; adapical opening short, lateral (i.e. orientation of barbs and adapical opening offset to point of overlap between margins near base); base broad, inflated, with extremely coarse external sculpture (
<figureCitation id="9BDA4AC4FFBCF843FC52FCF9F9203F25" box="[949,1033,872,894]" captionStart="Figure 6" captionStartId="18.[145,227,1378,1400]" captionTargetBox="[145,1425,195,1338]" captionTargetId="figure-24@18.[145,1425,195,1338]" captionTargetPageId="18" captionText="Figure 6. Hypodermic teeth of species described herein. A, Aplotoma brevitentaculata, holotype AMS C.571635; B, Gladiobela angulata, paratype AMS C.571737; C, Pueridaphne cirrisulcata, paratype AMS C.571609; D: Austrobela rufa, holotype AMS C.571709 (left), paratype AMS C.571679 (right); E, Pagodibela maia, holotype AMS C.571678; F, Fusobela parvioculata, holotype TMAG E59231; G, Biconitoma cretosa, holotype AMS C.482313; H, Austrotheta crassidentata, holotype AMS C.519302. Scale bars = 50 μm." pageId="25" pageNumber="986">Fig. 6H</figureCitation>
); ligament large.
</paragraph>
<paragraph id="035E5641FFBCF843FCDCFC27FE813F95" blockId="25.[827,936,950,974]" box="[827,936,950,974]" pageId="25" pageNumber="986">
<emphasis id="31958A53FFBCF843FCDCFC27FE813F95" box="[827,936,950,974]" italics="true" pageId="25" pageNumber="986">Remarks</emphasis>
</paragraph>
<paragraph id="035E5641FFBCF843FCDCFC4EF93A38B1" blockId="25.[827,1444,990,1871]" pageId="25" pageNumber="986">
<taxonomicName id="C4E12DC2FFBCF843FCDCFC4EF9B23FAF" authorityName="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov" authorityYear="2021" box="[827,1179,990,1012]" class="Gastropoda" family="Raphitomidae" genus="Austrotheta" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="crassidentata">
<emphasis id="31958A53FFBCF843FCDCFC4EF9B23FAF" box="[827,1179,990,1012]" italics="true" pageId="25" pageNumber="986">Austrotheta crassidentata</emphasis>
</taxonomicName>
differs from other raphitomids in the following combined characters: a fusiform shell with sharp, weakly opisthocline axials on early teleoconch whorls and weakly sculptured late teleoconch whorls; slender siphonal canal; thick hypodermic teeth with two weak laterally orientated barbs and a short laterally orientated adapical opening, as well as a broad base with extremely coarse external sculpture.
</paragraph>
<paragraph id="035E5641FFBCF843FCB4FB63F9B93B15" blockId="25.[827,1444,990,1871]" pageId="25" pageNumber="986">
Based on its shell morphology, 
<taxonomicName id="C4E12DC2FFBCF843FB3CFB62F88A395C" authorityName="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov" authorityYear="2021" box="[1243,1443,1266,1288]" class="Gastropoda" family="Raphitomidae" genus="Austrotheta" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="crassidentata">
<emphasis id="31958A53FFBCF843FB3CFB62F88A395C" box="[1243,1443,1266,1288]" italics="true" pageId="25" pageNumber="986">A. crassidentata</emphasis>
</taxonomicName>
resembles 
<taxonomicName id="C4E12DC2FFBCF843FC56FA80F8B6397C" authority="(Kuroda &amp; Habe, 1961)" baseAuthorityName="Kuroda &amp; Habe" baseAuthorityYear="1961" box="[945,1439,1297,1319]" class="Gastropoda" family="Raphitomidae" genus="Gymnobela" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="yoshidai">
<emphasis id="31958A53FFBCF843FC56FA80F9BA397D" box="[945,1171,1297,1318]" italics="true" pageId="25" pageNumber="986">Gymnobela yoshidai</emphasis>
(Kuroda &amp; 
<bibRefCitation id="67702BB0FFBCF843FAFFFA80F8BC397C" author="Habe T" box="[1304,1429,1297,1319]" pageId="25" pageNumber="986" refId="ref28572" refString="Habe T. 1961. Coloured illustrations of the shells of Japan (II). Hoikusha, Osaka. xii + 183 + 42 pp., 66 pls." type="book" year="1961">Habe, 1961</bibRefCitation>
)
</taxonomicName>
, in which strongly shouldered subcylindrical early teleoconch whorls with sharp ribs are succeeded by - in 
<bibRefCitation id="67702BB0FFBCF843FCBEFAFCFEF039D8" author="Habe T" box="[857,985,1389,1411]" pageId="25" pageNumber="986" refId="ref28572" refString="Habe T. 1961. Coloured illustrations of the shells of Japan (II). Hoikusha, Osaka. xii + 183 + 42 pp., 66 pls." type="book" year="1961">Habe, 1961</bibRefCitation>
evenly convex and finely striate whorls. However, the latter species has a narrower shell with a pointed spire, and a less convex, more elongated shell base, as well as a yellowish staining of the shell. A similar change in the whorl outline and sculptural pattern can be observed in 
<taxonomicName id="C4E12DC2FFBCF843FC3FF996F9BE3A40" baseAuthorityName="Habe" baseAuthorityYear="1962" box="[984,1175,1542,1564]" class="Gastropoda" family="Raphitomidae" genus="Spergo" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="fusiformis">
<emphasis id="31958A53FFBCF843FC3FF996F9BE3A40" box="[984,1175,1542,1564]" italics="true" pageId="25" pageNumber="986">Spergo fusiformis</emphasis>
</taxonomicName>
[based on the 
<typeStatus id="DC5AE8E3FFBCF843FAC9F996F8773A47" box="[1326,1374,1543,1564]" pageId="25" pageNumber="986">type</typeStatus>
series illustrated by 
<bibRefCitation id="67702BB0FFBCF843FC3AF9B4F9D23A60" author="Bouchet P &amp; Sysoev A" box="[989,1275,1573,1595]" pageId="25" pageNumber="986" pagination="1693 - 1715" refId="ref27640" refString="Bouchet P, Sysoev A. 2001. Typhlosyrinx-like tropical deep-water turriform gastropods (Mollusca, Gastropoda, Conoidea). Journal of Natural History 35: 1693 - 1715." type="journal article" year="2001">Sysoev &amp; Bouchet (2001)</bibRefCitation>
], although the latter can be readily differentiated from 
<taxonomicName id="C4E12DC2FFBCF843FB15F9D5F88A3A02" authorityName="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov" authorityYear="2021" box="[1266,1443,1604,1625]" class="Gastropoda" family="Raphitomidae" genus="Austrotheta" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="crassidentata">
<emphasis id="31958A53FFBCF843FB15F9D5F88A3A02" box="[1266,1443,1604,1625]" italics="true" pageId="25" pageNumber="986">A. crassidentata</emphasis>
</taxonomicName>
by its larger size and narrower, lanceolate aperture. 
<taxonomicName id="C4E12DC2FFBCF843FA6BF9F2FEFB3ACD" authorityName="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov" authorityYear="2021" class="Gastropoda" family="Raphitomidae" genus="Austrotheta" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="crassidentata">
<emphasis id="31958A53FFBCF843FA6BF9F2FEFB3ACD" italics="true" pageId="25" pageNumber="986">A. crassidentata</emphasis>
</taxonomicName>
also has some resemblance to 
<taxonomicName id="C4E12DC2FFBCF843FACDF910FE423AEE" authorityName="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov" authorityYear="2021" class="Gastropoda" family="Raphitomidae" genus="Austrobela" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="rufa">
<emphasis id="31958A53FFBCF843FACDF910FE423AEE" italics="true" pageId="25" pageNumber="986">Austrobela rufa</emphasis>
</taxonomicName>
, but can be readily differentiated by its diagonally cancellate protoconch, sharp axial ribs, a more convex shell base, and its whitish shell. When compared to 
<taxonomicName id="C4E12DC2FFBCF843FCDCF96DFEE13B4A" baseAuthorityName="A. H. Clarke" baseAuthorityYear="1959" box="[827,968,1788,1809]" class="Gastropoda" family="Raphitomidae" genus="Theta" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="lyronuclea">
<emphasis id="31958A53FFBCF843FCDCF96DFEE13B4A" box="[827,968,1788,1809]" italics="true" pageId="25" pageNumber="986">T. lyronuclea</emphasis>
</taxonomicName>
, it differs in its more elongate shell and in its diagonally cancellate protoconch (which is arcuate in both 
<taxonomicName id="C4E12DC2FFBCF843FC95F8A8FE973B15" authorityName="Criscione &amp; Hallan &amp; Puillandre &amp; Fedosov" authorityYear="2021" box="[882,958,1849,1870]" class="Gastropoda" family="Raphitomidae" genus="Austrobela" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="rufa">
<emphasis id="31958A53FFBCF843FC95F8A8FE973B15" box="[882,958,1849,1870]" italics="true" pageId="25" pageNumber="986">A. rufa</emphasis>
</taxonomicName>
and 
<taxonomicName id="C4E12DC2FFBCF843FC12F8ABF9A83B15" baseAuthorityName="A. H. Clarke" baseAuthorityYear="1959" box="[1013,1153,1849,1871]" class="Gastropoda" family="Raphitomidae" genus="Theta" kingdom="Animalia" order="Neogastropoda" pageId="25" pageNumber="986" phylum="Mollusca" rank="species" species="lyronuclea">
<emphasis id="31958A53FFBCF843FC12F8ABF9A83B15" box="[1013,1153,1849,1871]" italics="true" pageId="25" pageNumber="986">T. lyronuclea</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
</treatment>
</document>