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<mods:title id="A76A1A11DF2C5DC19BFFA5C9B6928D4C">Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae</mods:title>
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<mods:namePart id="AA3BD6A9B8A474AD0A75839513CE8B47">Simon, Eric</mods:namePart>
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<mods:namePart id="84F38B2FA952A7F831AC2A09C38B6604">Logan, Alan</mods:namePart>
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<mods:namePart id="5611A4050CC801AEF70CCC84258B59C5">Zuschin, Martin</mods:namePart>
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<mods:namePart id="1BF80EB0401154580EED696055982785">Mainguy, Jerome</mods:namePart>
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<mods:namePart id="C69EE2082AFB2B862FF08344BF754B91">Mottequin, Bernard</mods:namePart>
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<subSubSection id="C32165635D3AFF82FA8652B2FDAB76A6" box="[151,538,1231,1258]" pageId="6" pageNumber="7" type="nomenclature">
<paragraph id="8B8436E85D3AFF82FA8652B2FDAB76A6" blockId="6.[151,538,1231,1258]" box="[151,538,1231,1258]" pageId="6" pageNumber="7">
<heading id="D0CC81845D3AFF82FA8652B2FDAB76A6" bold="true" box="[151,538,1231,1258]" fontSize="11" level="1" pageId="6" pageNumber="7" reason="1">
<emphasis id="B94FEAFA5D3AFF82FA8652B2FDAB76A6" bold="true" box="[151,538,1231,1258]" pageId="6" pageNumber="7">
The genus
<taxonomicName id="4C3B4D6B5D3AFF82FB0F52ADFDAB76A6" ID-CoL="87H97" authority="King, 1850" authorityName="King" authorityYear="1850" box="[286,538,1231,1258]" class="Rhynchonellata" family="Kraussinidae" genus="Megerlia" kingdom="Animalia" order="Terebratulida" pageId="6" pageNumber="7" phylum="Brachiopoda" rank="genus">
<emphasis id="B94FEAFA5D3AFF82FB0F52ADFE3A76A5" bold="true" box="[286,395,1231,1257]" italics="true" pageId="6" pageNumber="7">Megerlia</emphasis>
<bibRefCitation id="EFAA4B195D3AFF82FB8352B2FDAB76A6" author="King" box="[402,538,1232,1258]" pageId="6" pageNumber="7" refString="King, W. (1850) A monograph of the Permian fossils of England. Palaeontographical Society Monograph, 3, xxxvii + 258 pp. http: // dx. doi. org / 10.5962 / bhl. title. 114608" type="book" year="1850">King, 1850</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C32165635D3AFF83FA865374FF5D7081" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7" type="description">
<paragraph id="8B8436E85D3AFF82FA865374FE8E7566" blockId="6.[151,1437,1302,2014]" pageId="6" pageNumber="7">
The main question of this paper is “to be or not to be
<taxonomicName id="4C3B4D6B5D3AFF82F9335374FC387763" box="[802,905,1302,1327]" class="Rhynchonellata" family="Kraussinidae" genus="Megerlia" kingdom="Animalia" order="Terebratulida" pageId="6" pageNumber="7" phylum="Brachiopoda" rank="genus">
<emphasis id="B94FEAFA5D3AFF82F9335374FC387763" box="[802,905,1302,1327]" italics="true" pageId="6" pageNumber="7">Megerlia</emphasis>
</taxonomicName>
”. The
<typeStatus id="5480884A5D3AFF82F9CF5375FBBF7763" box="[990,1038,1303,1327]" pageId="6" pageNumber="7">type</typeStatus>
species of this genus is
<taxonomicName id="4C3B4D6B5D3AFF82FF245374FE2C771F" authority="Linne, 1767" authorityName="Linne" authorityYear="1767" class="Rhynchonellata" family="Kraussinidae" genus="Megerlia" kingdom="Animalia" order="Terebratulida" pageId="6" pageNumber="7" phylum="Brachiopoda" rank="species" species="truncata">
<emphasis id="B94FEAFA5D3AFF82FF245374FF47771F" italics="true" pageId="6" pageNumber="7">Megerlia truncata</emphasis>
(
<bibRefCitation id="EFAA4B195D3AFF82FB175358FE25771F" author="Linne" box="[262,404,1338,1363]" pageId="6" pageNumber="7" refString="Linne, C. A. (1767) Systema Naturae, sive Regna tria Naturae systematicae proposita per Classes, Ordines, Genera, Species cum Characteribus &amp; Differentiis. 12 th edit., vol. 1, n ° 2, Holmiae, pp. 533 - 1327." type="book chapter" year="1767">Linné, 1767</bibRefCitation>
)
</taxonomicName>
. The original material described by Linné as
<taxonomicName id="4C3B4D6B5D3AFF82F9B25359FBD0771F" box="[931,1121,1339,1363]" class="Insecta" family="Cicadellidae" genus="Anomia" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="truncata">
<emphasis id="B94FEAFA5D3AFF82F9B25359FBD0771F" box="[931,1121,1339,1363]" italics="true" pageId="6" pageNumber="7">Anomia truncata</emphasis>
</taxonomicName>
apparently consists of two dried specimens (as indicated in the Catalogue of
<typeStatus id="5480884A5D3AFF82F8F7533DFCAF773B" box="[742,798,1375,1399]" pageId="6" pageNumber="7">Type</typeStatus>
specimens of the University of Uppsala: n°1907a and n°1907b) and should be Norwegian specimens (“
<emphasis id="B94FEAFA5D3AFF82F8C253E0FB6277D7" box="[723,1235,1410,1435]" italics="true" pageId="6" pageNumber="7">Habitat in Pelago Norvegico Supra corallia</emphasis>
<bibRefCitation id="EFAA4B195D3AFF82FEFB53E0FAC977D7" author="Linne" box="[1258,1400,1410,1435]" pageId="6" pageNumber="7" refString="Linne, C. A. (1767) Systema Naturae, sive Regna tria Naturae systematicae proposita per Classes, Ordines, Genera, Species cum Characteribus &amp; Differentiis. 12 th edit., vol. 1, n ° 2, Holmiae, pp. 533 - 1327." type="book chapter" year="1767">Linné, 1767</bibRefCitation>
; p. 1152, n° 229). However, the presence of this species off
<collectingCountry id="F32C76785D3AFF82F90853C5FCC277F3" box="[793,883,1447,1471]" name="Norway" pageId="6" pageNumber="7">Norway</collectingCountry>
was not mentioned by Logan (2007) or
<bibRefCitation id="EFAA4B195D3AFF82FF2E53C4FEBB77AF" pageId="6" pageNumber="7" refString="Hiller, N., MacKinnon, D. I. &amp; Nielsen, S. N. (2008) A review of the systematic, biogeography and evolutionary relationships of Recent and fossil brachiopods of the Superfamily Kraussinoidea Dall, with descriptions of two new fossil species from New-Zealand and Chile. Earth and Environmental Science transactions of the Royal Society of Edinburgh, 98, 379 - 390. http: // dx. doi. org / 10.1017 / S 1755691007078474" type="journal article">
Hiller
<emphasis id="B94FEAFA5D3AFF82FF9953C5FF1C77AF" italics="true" pageId="6" pageNumber="7">et al</emphasis>
. (2008)
</bibRefCitation>
. The species was not found in northerly latitudes such as the Irish coasts by Massy (1925),
<collectingCountry id="F32C76785D3AFF82FF0C53A9FAC677AF" box="[1309,1399,1483,1507]" name="Norway" pageId="6" pageNumber="7">Norway</collectingCountry>
by
<bibRefCitation id="EFAA4B195D3AFF82FA865393FE1D7446" author="Wesenberg-Lund" box="[151,428,1521,1546]" pageId="6" pageNumber="7" refString="Wesenberg-Lund, E. (1939) Recent Norwegian Brachiopods. Det Kongelige Norske Videnskabers-Selskab Forhandlinger, 11 (52), 201 - 204." type="journal article" year="1939">Wesenberg-Lund (1939)</bibRefCitation>
, the North Sea (
<bibRefCitation id="EFAA4B195D3AFF82F87B5393FD567446" author="Cori" box="[618,743,1521,1546]" pageId="6" pageNumber="7" refString="Cori, C. I. (1933) Brachiopoda. In: Grimpe, G &amp; Wagler, E. (Eds.), Die Tierwelt der Nord-und Ostsee, 24 (VIIc 3): 132 - 149." type="journal article" year="1933">Cori, 1933</bibRefCitation>
) and the
<collectingCountry id="F32C76785D3AFF82F9475390FC5F7446" box="[854,1006,1521,1546]" name="Faroe Islands" pageId="6" pageNumber="7">Faroe Islands</collectingCountry>
(
<bibRefCitation id="EFAA4B195D3AFF82F9EE5393FB047446" author="Thomsen" box="[1023,1205,1521,1546]" pageId="6" pageNumber="7" refString="Thomsen, E. (2001) Brachiopoda in the Faroe Island area. Frodskaparrit, 49, 109 - 126." type="journal article" year="2001">Thomsen, 2001</bibRefCitation>
). In the 19th century
<taxonomicName id="4C3B4D6B5D3AFF82FA865077FF4F7462" box="[151,254,1557,1582]" class="Rhynchonellata" family="Kraussinidae" genus="Megerlia" kingdom="Animalia" order="Terebratulida" pageId="6" pageNumber="7" phylum="Brachiopoda" rank="genus">
<emphasis id="B94FEAFA5D3AFF82FA865077FF4F7462" box="[151,254,1557,1582]" italics="true" pageId="6" pageNumber="7">Megerlia</emphasis>
</taxonomicName>
species were not recognized along the British coasts (Forbes &amp; Hanley 1850, 1853). The original Linnaean specimens may have come from another area in the eastern Atlantic or even from the Mediterranean Sea. The current genus description provided by Lee &amp; MacKinnon (2006, p. H2245) stresses the large foramen which is often abraded (considered submesothyrid to amphithyrid), the disjunct deltidial plates and the tuberculate internal valve floors. No cardinal process is developed. Crural bases are present and attached to the inner sides of socket ridges. Descending branches join the cardinalia with the posterior part of the ring of the loop. The brachidium is typically bifurcate (see MacKinnon &amp; Lee, 2006, fig.1312) and a complete ring is built; this is unique among
<taxonomicName id="4C3B4D6B5D3AFF82FA865173FE8A7566" box="[151,315,1809,1834]" class="Rhynchonellata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Terebratulida" pageId="6" pageNumber="7" phylum="Brachiopoda" rank="superFamily" superFamily="Kraussinoidea">Kraussinoidea</taxonomicName>
.
</paragraph>
<paragraph id="8B8436E85D3AFF83FAD65154FB2C7054" blockId="6.[151,1437,1302,2014]" lastBlockId="7.[151,1437,151,717]" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">
SEM observations show in detail the development in
<taxonomicName id="4C3B4D6B5D3AFF82F92A5155FC727502" box="[827,963,1846,1870]" class="Insecta" family="Cicadellidae" genus="Anomia" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="truncata">
<emphasis id="B94FEAFA5D3AFF82F92A5155FCE37502" box="[827,850,1847,1870]" italics="true" pageId="6" pageNumber="7">M</emphasis>
.
<emphasis id="B94FEAFA5D3AFF82F9755155FC727502" box="[868,963,1847,1870]" italics="true" pageId="6" pageNumber="7">truncata</emphasis>
</taxonomicName>
and interpretation of the earliest growth stages is here discussed again. The smallest specimen collected from Anzio has a width of
<quantity id="4CC39B0D5D3AFF82FE8A5138FB40753D" box="[1179,1265,1882,1906]" metricMagnitude="-3" metricUnit="m" metricValue="1.4" pageId="6" pageNumber="7" unit="mm" value="1.4">1.4 mm</quantity>
(Pl.1, Figs. 1a, 1b). This specimen is slightly biconvex. The tip of the beak is abraded but the submesothyrid foramen is still intact on its dorsal side. The dorsal valve is smooth and the ventral valve presents some rare tubercles near the anterior commissure. The second specimen (Pl. 1, Figs. 2a, 2b) has a width of
<quantity id="4CC39B0D5D3AFF82F9BB51A4FBA07591" box="[938,1041,1990,2014]" metricMagnitude="-3" metricUnit="m" metricValue="1.73" pageId="6" pageNumber="7" unit="mm" value="1.73">1.73 mm</quantity>
. The shell becomes ventribiconvex with a smooth dorsal valve whereas the external surface of the ventral valve presents a radial series of tubercles. Noteworthy is that the outer shell surface between the radial ornamentation is smooth. The beak is more abraded and the dorsal valve is affected as the foramen seems amphithyrid. This gradual change is quite continuous during growth as seen on Pl. 1, Figs. 37. It appears that the foramen is more and more abraded due to the shortness of the pedicle and the high energy environment where
<taxonomicName id="4C3B4D6B5D3BFF83F8C9574BFCD3730C" box="[728,866,297,320]" class="Insecta" family="Cicadellidae" genus="Anomia" kingdom="Animalia" order="Hemiptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="truncata">
<emphasis id="B94FEAFA5D3BFF83F8C9574BFCD3730C" box="[728,866,297,320]" italics="true" pageId="7" pageNumber="8">M. truncata</emphasis>
</taxonomicName>
is generally living. Another development is the appearance of radial costae with tubercles on the external side of the dorsal valve. This tuberculation becomes more important during growth (Pl. 2, Figs. 13) although the ventral tubercles always remain more developed in size. An underestimated character in
<taxonomicName id="4C3B4D6B5D3BFF83F81357F7FD3673E0" box="[514,647,405,428]" class="Insecta" family="Cicadellidae" genus="Anomia" kingdom="Animalia" order="Hemiptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="truncata">
<emphasis id="B94FEAFA5D3BFF83F81357F7FD3673E0" box="[514,647,405,428]" italics="true" pageId="7" pageNumber="8">M. truncata</emphasis>
</taxonomicName>
is the absence, at all growth stages, of secondary tubercles, spines or scales in the “intercostal” zones. In these zones the shell remains smooth and only concentric growth lines are perceptible (Pls.15). This is not the case in
<taxonomicName id="4C3B4D6B5D3BFF83F88757BFFCAF73B9" box="[662,798,476,501]" class="Rhynchonellata" family="Kraussinidae" genus="Megerlia" kingdom="Animalia" order="Terebratulida" pageId="7" pageNumber="8" phylum="Brachiopoda" rank="species" species="echinata">
<emphasis id="B94FEAFA5D3BFF83F88757BFFCAF73B9" box="[662,798,476,501]" italics="true" pageId="7" pageNumber="8">M. echinata</emphasis>
</taxonomicName>
(Pl. 10, Figs. 1, 2ad) as strong spines cover the whole external surface of the ventral valve (see also Fischer &amp; Oehlert, 1891, pl.7, figs 13a and b).
</paragraph>
<paragraph id="8B8436E85D3BFF83FAD65446FF5D7081" blockId="7.[151,1437,151,717]" pageId="7" pageNumber="8">
Examination of many specimens of
<taxonomicName id="4C3B4D6B5D3BFF83F8785447FD417070" box="[617,752,549,572]" class="Insecta" family="Cicadellidae" genus="Anomia" kingdom="Animalia" order="Hemiptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="truncata">
<emphasis id="B94FEAFA5D3BFF83F8785447FD417070" box="[617,752,549,572]" italics="true" pageId="7" pageNumber="8">M. truncata</emphasis>
</taxonomicName>
from the whole Mediterranean Sea also clearly shows that both valves are provided with radial tuberculation, the ventral tuberculation always being stronger than the dorsal one. Specimens with abraded dorsal tubercles are found but an entirely smooth dorsal valve without any tuberculation has not been observed. This feature is important for distinguishing
<taxonomicName id="4C3B4D6B5D3BFF83FE0D54F3FB2E70E4" box="[1052,1183,657,680]" class="Insecta" family="Cicadellidae" genus="Anomia" kingdom="Animalia" order="Hemiptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="species" species="truncata">
<emphasis id="B94FEAFA5D3BFF83FE0D54F3FB2E70E4" box="[1052,1183,657,680]" italics="true" pageId="7" pageNumber="8">M. truncata</emphasis>
</taxonomicName>
from
<taxonomicName id="4C3B4D6B5D3BFF83FEF254F3FAD870E4" box="[1251,1385,655,680]" class="Rhynchonellata" family="Kraussinidae" genus="Megerlia" kingdom="Animalia" order="Terebratulida" pageId="7" pageNumber="8" phylum="Brachiopoda" rank="species" species="echinata">
<emphasis id="B94FEAFA5D3BFF83FEF254F3FAD870E4" box="[1251,1385,655,680]" italics="true" pageId="7" pageNumber="8">M. echinata</emphasis>
</taxonomicName>
(see below).
</paragraph>
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