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<mods:title id="1824022FA0FD1AE7EBFBF304B3FA2D20">Chondrinidae taxonomy revisited: New synonymies, new taxa, and a checklist of species and subspecies (Mollusca: Gastropoda: Pulmonata) 2539</mods:title>
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<taxonomicName id="4C4D2391CE76EC5C7EEEFE39C50DC984" ID-CoL="7NZBY" authority="Reichenbach, 1828" authorityName="Reichenbach" authorityYear="1828" box="[151,524,418,444]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="genus">
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<emphasis id="B9398400CE76EC5C7EEEFE39C61AC984" bold="true" box="[151,283,418,444]" italics="true" pageId="25" pageNumber="26">Chondrina</emphasis>
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<typeStatus id="54F6E6B0CE76EC5C7EEEFE5EC7D0C9E7" box="[151,209,453,479]" pageId="25" pageNumber="26">Type</typeStatus>
species (design.
<bibRefCitation id="EFDC25E3CE76EC5C7FE0FE5EC5B2C9E7" author="Reichenbach, H. G. L." box="[409,691,453,479]" pageId="25" pageNumber="26" refId="ref36981" refString="Reichenbach, H. G. L. (1836) Allgemeine Taschenbibliothek der Naturwissenschaften 5. Theil. 2. Bandchen. Zoologie oder Naturgeschichte des Thierreichs. Nauwerck, Zittau, 110 - 272. [Not seen]." type="book" year="1836">Reichenbach 1836: 152</bibRefCitation>
):
<taxonomicName id="4C4D2391CE76EC5C7CBEFE5EC370C9E7" authority="Bruguiere, 1792" authorityName="Bruguiere" authorityYear="1792" box="[711,1137,453,479]" class="Gastropoda" family="Bulimulidae" genus="Bulimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="avenaceus">
<emphasis id="B9398400CE76EC5C7CBEFE5EC4A9C9E6" box="[711,936,453,478]" italics="true" pageId="25" pageNumber="26">Bulimus avenaceus</emphasis>
<bibRefCitation id="EFDC25E3CE76EC5C7DC9FE5EC370C9E7" author="Bruguiere, J. G." box="[944,1137,453,479]" pageId="25" pageNumber="26" pagination="345 - 758" refId="ref33944" refString="Bruguiere, J. G. (1792) Encyclopedie methodique ou par ordre de matieres. Histoire naturelle des vers, volume 1. Panckoucke, Paris, pp. 345 - 758." type="book chapter" year="1792">Bruguière, 1792</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF25812CE76EC5C7EBCFE77C37FCB5A" blockId="25.[151,1436,418,1871]" pageId="25" pageNumber="26">
Notes.—For this genus the results of a preliminary, molecular, phylogenetic analysis are available (
<bibRefCitation id="EFDC25E3CE76EC5C7B2BFE77C67ACA14" author="Kokshoorn, B." pageId="25" pageNumber="26" refId="ref35737" refString="Kokshoorn, B. (2008) Resolving Riddles and Presenting New Puzzles in Chondrinidae Phylogenetics. PhD thesis, Leiden University, 192 pp." type="book" year="2008">Kokshoorn 2008: 4158</bibRefCitation>
). This analysis (see fig. 4) made clear that in any case, a revision of the classification presented by
<bibRefCitation id="EFDC25E3CE76EC5C7F6BFDA2C6FFCA6B" author="Gittenberger, E." box="[274,510,569,595]" pageId="25" pageNumber="26" pagination="1 - 267" refId="ref34929" refString="Gittenberger, E. (1973) Beitrage zur Kenntnis der Pupillacea III. Chondrininae. Zoologische Verhandelingen, 127, 1 - 267." type="journal article" year="1973">Gittenberger (1973)</bibRefCitation>
is necessary. In particular the nominal taxa united by that author as a single, polytypic species, i.e.
<taxonomicName id="4C4D2391CE76EC5C7FDBFDFBC48ECA42" authority="(Des Moulins, 1835)" baseAuthorityName="Des Moulins" baseAuthorityYear="1835" box="[418,911,608,634]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="farinesii">
<emphasis id="B9398400CE76EC5C7FDBFDFBC58DCA41" box="[418,652,608,633]" italics="true" pageId="25" pageNumber="26">Chondrina farinesii</emphasis>
(
<bibRefCitation id="EFDC25E3CE76EC5C7CE4FDFBC487CA42" author="Des Moulins, C." box="[669,902,608,634]" pageId="25" pageNumber="26" pagination="142 - 165" refId="ref34355" refString="Des Moulins, C. (1835) Description de quelques mollusques terrestres et fluviatiles de la France, nouveau ou peu connus. Actes de la Societe Linneenne de Bordeaux, 7, 142 - 165." type="journal article" year="1835">Des Moulins, 1835</bibRefCitation>
)
</taxonomicName>
, should not all be lumped. Obviously, there are many more taxa, characterized by shells with more or less obsolete, apertural teeth. In some cases, shell dimensions, shape and sculpture, could be used under the direction of the molecular data, but many questions remain unanswered because of a lack of sufficient information. Since it is unlikely that this situation will improve substantially in the near future, we have already adapted the classification of the
<taxonomicName id="4C4D2391CE76EC5C7ABDFD61C242CB2B" authorityName="Reichenbach" authorityYear="1828" box="[1220,1347,762,787]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="genus">
<emphasis id="B9398400CE76EC5C7ABDFD61C242CB2B" box="[1220,1347,762,787]" italics="true" pageId="25" pageNumber="26">Chondrina</emphasis>
</taxonomicName>
taxa to the new insights. Since the dataset presented in figure 4 is inconclusive at some points in the tree, we refrain from revising taxa that show ambiguous positions in the phylogenetic reconstruction.
</paragraph>
<paragraph id="8BF25812CE76EC5C7EBCFCF5C6B8CCAF" blockId="25.[151,1436,418,1871]" pageId="25" pageNumber="26">
New taxa are formally described and named only when they can be recognized on the basis of molecular and (to some extent) conchological characteristics. For the time being, to promote further research, some taxa are referred to with numbers only. These taxa can only be defined on very limited DNA data, so that their geographical distribution and the variation in shell morphology remain unknown. More sequence data are needed to acquire more reliably classified material. Shell morphology should be studied again in search of hitherto unnoticed common characteristics. Inevitably, an additional increase of DNA data will dictate further changes in the following overview. The phylogenetic relationships within
<taxonomicName id="4C4D2391CE76EC5C7DE6FBCCC31FCC48" authorityName="Reichenbach" authorityYear="1828" box="[927,1054,1111,1136]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="genus">
<emphasis id="B9398400CE76EC5C7DE6FBCCC31FCC48" box="[927,1054,1111,1136]" italics="true" pageId="25" pageNumber="26">Chondrina</emphasis>
</taxonomicName>
are more complicated than may be assumed at first sight.
</paragraph>
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The apertural teeth do not vary randomly in
<taxonomicName id="4C4D2391CE76EC5C7CADFB3FC452CC85" authorityName="Reichenbach" authorityYear="1828" box="[724,851,1188,1213]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="genus">
<emphasis id="B9398400CE76EC5C7CADFB3FC452CC85" box="[724,851,1188,1213]" italics="true" pageId="25" pageNumber="26">Chondrina</emphasis>
</taxonomicName>
. There is a “bauplan I” with more than two palatals and a spiralis, next to a “bauplan II” with two palatals at most and no spiralis. Toothless forms may be considered derived from the latter bauplan. These two groups are not clades however. Some individual specimens or even species do not belong to one of the alternatives. In
<taxonomicName id="4C4D2391CE76EC5C7DF4FA83C30CCD09" authorityName="Altimira" authorityYear="1967" box="[909,1037,1304,1329]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="aguilari">
<emphasis id="B9398400CE76EC5C7DF4FA83C30CCD09" box="[909,1037,1304,1329]" italics="true" pageId="25" pageNumber="26">C. aguilari</emphasis>
</taxonomicName>
there may be an infrapalatalis but never a spiralis, whereas in
<taxonomicName id="4C4D2391CE76EC5C7F9BFAA4C5B8CD60" authorityName="Kokshoorn &amp; Gittenberger" authorityYear="2010" box="[482,697,1343,1368]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="pseudavenacea">
<emphasis id="B9398400CE76EC5C7F9BFAA4C5B8CD60" box="[482,697,1343,1368]" italics="true" pageId="25" pageNumber="26">C. pseudavenacea</emphasis>
</taxonomicName>
there are three palatals, again without a spiralis. According to
<bibRefCitation id="EFDC25E3CE76EC5C7EEEFAFEC6BBCD47" author="Gittenberger, E." box="[151,442,1381,1407]" pageId="25" pageNumber="26" pagination="1 - 267" refId="ref34929" refString="Gittenberger, E. (1973) Beitrage zur Kenntnis der Pupillacea III. Chondrininae. Zoologische Verhandelingen, 127, 1 - 267." type="journal article" year="1973">Gittenberger (1973: 232)</bibRefCitation>
there may be an infrapalatalis, a suprapalatalis and an obsolete spiralis in
<taxonomicName id="4C4D2391CE76EC5C7B65FAFEC295CD46" baseAuthorityName="Westerlund" baseAuthorityYear="1872" box="[1308,1428,1381,1406]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="calpica">
<emphasis id="B9398400CE76EC5C7B65FAFEC295CD46" box="[1308,1428,1381,1406]" italics="true" pageId="25" pageNumber="26">C. calpica</emphasis>
</taxonomicName>
. In this species bauplan II is the rule, but rarely specimens with apertural teeth according to bauplan I occur as well. We have to conclude that the apertural teeth became obsolete or were reduced completely several times independently, resulting in shells that misleadingly suggest a close relationship in various cases.Several authors have contributed to the currently accepted taxonomy of the group (
<bibRefCitation id="EFDC25E3CE76EC5C7A50F99BC3EACE22" author="Nordsieck, H." box="[1065,1259,1536,1562]" pageId="25" pageNumber="26" pagination="1 - 20" refId="ref36499" refString="Nordsieck, H. (1962) Die Chondrinen der Sudalpen. Achiv fur Molluskenkunde, 91, 1 - 20." type="journal article" year="1962">Nordsieck 1962</bibRefCitation>
,
<bibRefCitation id="EFDC25E3CE76EC5C7A82F99BC238CE22" author="Nordsieck, H." box="[1275,1337,1536,1562]" pageId="25" pageNumber="26" pagination="243 - 261" refId="ref36521" refString="Nordsieck, H. (1970) Die Chondrina-Arten der dinarischen Lander. Archiv fur Molluskenkunde, 100, 243 - 261." type="journal article" year="1970">1970</bibRefCitation>
;
<bibRefCitation id="EFDC25E3CE76EC5C7B30F99BC626CE78" author="Gittenberger, E." pageId="25" pageNumber="26" pagination="1 - 267" refId="ref34929" refString="Gittenberger, E. (1973) Beitrage zur Kenntnis der Pupillacea III. Chondrininae. Zoologische Verhandelingen, 127, 1 - 267." type="journal article" year="1973">Gittenberger 1973</bibRefCitation>
;
<bibRefCitation id="EFDC25E3CE76EC5C7F4AF9BDC54ACE78" author="Gomez, B. J. &amp; Angulo, E." box="[307,587,1574,1600]" pageId="25" pageNumber="26" pagination="29 - 40" refId="ref35101" refString="Gomez, B. J. &amp; Angulo, E. (1982) El Genero Chondrina (Mollusca, Gastropoda) en el Pais Vasco. Iberus, 2, 29 - 40." type="journal article" year="1982">Gómez &amp; Angulo 1982</bibRefCitation>
).
<taxonomicName id="4C4D2391CE76EC5C7C26F9BCC455CE78" baseAuthorityName="Bruguiere" baseAuthorityYear="1792" box="[607,852,1575,1600]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="avenacea">
<emphasis id="B9398400CE76EC5C7C26F9BCC455CE78" box="[607,852,1575,1600]" italics="true" pageId="25" pageNumber="26">Chondrina avenacea</emphasis>
</taxonomicName>
and
<taxonomicName id="4C4D2391CE76EC5C7DF7F9BCC36FCE78" baseAuthorityName="Westerlund" baseAuthorityYear="1883" box="[910,1134,1575,1600]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="subSpecies" species="arcadica" subSpecies="clienta">
<emphasis id="B9398400CE76EC5C7DF7F9BCC36FCE78" box="[910,1134,1575,1600]" italics="true" pageId="25" pageNumber="26">C. arcadica clienta</emphasis>
</taxonomicName>
have been used as model organisms in ecological and evolutionary studies (
<bibRefCitation id="EFDC25E3CE76EC5C7C88F9D6C309CE5F" author="Armbruster, G. F. J. &amp; Hofer, M. &amp; Baur, B." box="[753,1032,1613,1639]" pageId="25" pageNumber="26" pagination="325 - 333" refId="ref32724" refString="Armbruster, G. F. J., Hofer, M. &amp; Baur, B. (2007) Effect of cliff connectivity on the genetic population structure of a rockdwelling land snail species with frequent self-fertilization. Biochemical Systematics and Ecology, 35, 325 - 333." type="journal article" year="2007">Armbruster et al. 2007</bibRefCitation>
; Baur, A. &amp; Baur, B. 1991; Baur, B. &amp; Baur, A. 1990, 1994, 1995; Baur, B. et al. 1993, 1995; Baur, A. et al. 1992, 1994;
<bibRefCitation id="EFDC25E3CE76EC5C7ACBF9EFC296CEB6" author="Froberg, L. &amp; Baur, A. &amp; Baur, B." box="[1202,1431,1652,1678]" pageId="25" pageNumber="26" pagination="83 - 96" refId="ref34776" refString="Froberg, L., Baur, A. &amp; Baur, B. (1993) Differential herbivore damage to calcicolous lichens by snails. Lichenologist, 25, 83 - 96." type="journal article" year="1993">Fröberg et al. 1993</bibRefCitation>
;
<bibRefCitation id="EFDC25E3CE76EC5C7EEEF900C69ACE8D" author="Hesbacher, S. &amp; Baur, B. &amp; Baur, A. &amp; Proksch, P." box="[151,411,1691,1717]" pageId="25" pageNumber="26" pagination="233 - 246" refId="ref35282" refString="Hesbacher, S., Baur, B., Baur, A. &amp; Proksch, P. (1995) Sequestration of lichen compounds by three species of terrestrial snails. Journal of Chemical Ecology, 21, 233 - 246." type="journal article" year="1995">Hesbacher et al. 1995</bibRefCitation>
;
<bibRefCitation id="EFDC25E3CE76EC5C7FD1F900C5B1CE8D" author="Szarowska, M. &amp; Falniowski, A. &amp; Mazan-Mamczarz, K." box="[424,688,1691,1717]" pageId="25" pageNumber="26" pagination="79 - 89" refId="ref37522" refString="Szarowska, M., Falniowski, A. &amp; Mazan-Mamczarz, K. (2003) Genetic structure of isolated selfers: Chondrina clienta (Westerlund, 1888) (Gastropoda: Stylommatophora) in Krakow-Czestochowa Upland. Malakologische Abhandlungen, 21, 79 - 89." type="journal article" year="2003">Szarowska et al. 2003</bibRefCitation>
). However, a study devoted to the phylogeny of the
<taxonomicName id="4C4D2391CE76EC5C7B64F900C7EDCEE3" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B9398400CE76EC5C7B64F900C29DCE8C" box="[1309,1436,1691,1716]" italics="true" pageId="25" pageNumber="26">Chondrina</emphasis>
species
</taxonomicName>
was still not available.
<bibRefCitation id="EFDC25E3CE76EC5C7F84F95AC5E7CEE3" author="Gittenberger, E." box="[509,742,1729,1755]" pageId="25" pageNumber="26" pagination="1 - 267" refId="ref34929" refString="Gittenberger, E. (1973) Beitrage zur Kenntnis der Pupillacea III. Chondrininae. Zoologische Verhandelingen, 127, 1 - 267." type="journal article" year="1973">Gittenberger (1973)</bibRefCitation>
divided the genus in four subgroups. He regarded that classification rather artificial, probably reflecting only some morphological and geographical similarities rather than phylogenetic relationships. Since this subdivision does indeed not reflect the evolutionary history of the genus, it is no longer acceptable.
</paragraph>
<paragraph id="8BF25812CE75EC5E7EBFF82FC5D5C954" blockId="26.[151,1435,1972,2036]" lastBlockId="27.[151,1436,152,364]" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27">
Since
<taxonomicName id="4C4D2391CE75EC5F7F6BF82FC6BCCFF6" authorityName="Arrebola &amp; Gomez" authorityYear="1998" box="[274,445,1972,1998]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="26" pageNumber="27" phylum="Mollusca" rank="species" species="maginensis">C. maginensis</taxonomicName>
has to be considered the sister-group of all other
<taxonomicName id="4C4D2391CE75EC5F7A5AF82FC3A2CFF5" authorityName="Reichenbach" authorityYear="1828" box="[1059,1187,1972,1997]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="26" pageNumber="27" phylum="Mollusca" rank="genus">
<emphasis id="B9398400CE75EC5F7A5AF82FC3A2CFF5" box="[1059,1187,1972,1997]" italics="true" pageId="26" pageNumber="27">Chondrina</emphasis>
</taxonomicName>
taxa combined, that species is dealt with first. The remaining
<taxonomicName id="4C4D2391CE75EC5F7CF7F841C40FCFCB" authorityName="Reichenbach" authorityYear="1828" box="[654,782,2010,2035]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="26" pageNumber="27" phylum="Mollusca" rank="genus">
<emphasis id="B9398400CE75EC5F7CF7F841C40FCFCB" box="[654,782,2010,2035]" italics="true" pageId="26" pageNumber="27">Chondrina</emphasis>
</taxonomicName>
taxa are provisionally arranged in six species groups, guided by the results of the molecular analysis (
<bibRefCitation id="EFDC25E3CE74EC5E7CB7FF03C4DAC88A" author="Kokshoorn, B. &amp; Schoor, M. van &amp; Erkelens, I. &amp; Gittenberger, E." box="[718,987,152,178]" pageId="27" pageNumber="28" refId="ref35763" refString="Kokshoorn, B., Schoor, M. van, Erkelens, I. &amp; Gittenberger, E. (2010) Waves of dispersal in island-hopping Chondrina species (Gastropoda, Pulmonata, Chondrinidae). Zoologischer Anzeiger, in press." type="book" year="2010">Kokshoorn et al. 2010</bibRefCitation>
)(see also fig. 4). Since the taxonomic diversity is much larger than previously expected, the molecular analysis should be considered far from complete. Consequently, the arrangement of species based on it, must be considered hypothetical at most. As a final group, some nominal taxa are listed that have been synonymized or overlooked before, but might represent separate
<taxonomicName id="4C4D2391CE74EC5E7F2BFEB6C533C97F" box="[338,562,301,327]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="27" pageNumber="28" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B9398400CE74EC5E7F2BFEB6C6D2C97E" box="[338,467,301,326]" italics="true" pageId="27" pageNumber="28">Chondrina</emphasis>
species
</taxonomicName>
or subspecies.The geographical distributions of the
<taxonomicName id="4C4D2391CE74EC5E7AD2FEB6C22AC97E" authorityName="Reichenbach" authorityYear="1828" box="[1195,1323,301,326]" class="Gastropoda" family="Chondrinidae" genus="Chondrina" kingdom="Animalia" order="Stylommatophora" pageId="27" pageNumber="28" phylum="Mollusca" rank="genus">
<emphasis id="B9398400CE74EC5E7AD2FEB6C22AC97E" box="[1195,1323,301,326]" italics="true" pageId="27" pageNumber="28">Chondrina</emphasis>
</taxonomicName>
taxa discussed in this paper are shown in fig. 5, 6 and 7.
</paragraph>
</subSubSection>
</treatment>
</document>