259 lines
39 KiB
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259 lines
39 KiB
XML
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<mods:title id="300D5F71823B08059B656A2058FA1FE5">Revised systematics of Palaeozoic ‘ horseshoe crabs’ and the myth of monophyletic Xiphosura</mods:title>
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<mods:namePart id="42BB114D2D5FB5A0324D1BE99A303078">Lamsdell, James C.</mods:namePart>
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<mods:date id="F14F0F932FC1FA52DD7FB895F8F7270E">2013</mods:date>
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<mods:number id="8AA086A5533E4491B2572EE1C17E6CE9">167</mods:number>
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<subSubSection id="C325A3B11C6FA7299D442AFCFA57F9D0" pageId="3" pageNumber="4" type="nomenclature">
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<paragraph id="8B80F03A1C6FA7299D442AFCFB58FC03" blockId="3.[1069,1198,986,1007]" box="[1069,1198,986,1007]" pageId="3" pageNumber="4">
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<heading id="D0C847561C6FA7299D442AFCFB58FC03" box="[1069,1198,986,1007]" centered="true" fontSize="8" level="2" pageId="3" pageNumber="4" reason="8">
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<taxonomicName id="4C3F8BB91C6FA7299D442AFCFB58FC03" authorityName="Lamsdell" authorityYear="2013" box="[1069,1198,986,1007]" class="Xiphosura" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Arthropoda" rank="class">XIPHOSURA</taxonomicName>
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</heading>
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</paragraph>
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<paragraph id="8B80F03A1C6FA7299A532D26FA57F9D0" blockId="3.[826,1441,1024,1597]" pageId="3" pageNumber="4">
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The terminology for aspects of xiphosuran morphology largely follows
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<bibRefCitation id="EFAE8DCB1C6FA7299D5D2D39FA92FBD8" author="Siveter DJ & Selden PA" box="[1076,1380,1055,1077]" pageId="3" pageNumber="4" pagination="153 - 168" refId="ref18474" refString="Siveter DJ, Selden PA. 1987. A new, giant xiphosurid from the lower Namurian of Weardale, County Durham. Proceedings of the Yorkshire Geological Society 46: 153 - 168." type="journal article" year="1987">Siveter & Selden (1987)</bibRefCitation>
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and
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<bibRefCitation id="EFAE8DCB1C6FA7299A532D1BFBA2FBBE" author="Siveter DJ & Selden PA" box="[826,1108,1085,1107]" pageId="3" pageNumber="4" pagination="153 - 168" refId="ref18474" refString="Siveter DJ, Selden PA. 1987. A new, giant xiphosurid from the lower Namurian of Weardale, County Durham. Proceedings of the Yorkshire Geological Society 46: 153 - 168." type="journal article" year="1987">Selden & Siveter (1987)</bibRefCitation>
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. The prosoma comprises the entirety of the anterior tagma, including the dorsal carapace and the prosomal appendages. The prosomal appendages are denoted by roman numerals I–VI, appendage I being the chelicerae. In limulids the anterior cephalic region is termed the cephalothorax by
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<bibRefCitation id="EFAE8DCB1C6FA7299A352DD3FC00FAE6" author="Shultz JW" box="[860,1014,1269,1291]" pageId="3" pageNumber="4" pagination="283 - 303" refId="ref18307" refString="Shultz JW. 2001. Gross muscular anatomy of Limulus polyphemus (Xiphosura, Chelicerata) and its bearing on evolution in the Arachnida. Journal of Arachnology 29: 283 - 303." type="journal article" year="2001">Shultz (2001)</bibRefCitation>
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as a number of opisthosomal somites have been at least partially incorporated into the prosoma (Fig. 1D). The carapace/cephalothorax itself bears a number of structures; the cardiac lobe is located at the posterior of the carapace and is axially inflated, while the ophthalmic ridges extend either side of the cardiac lobe and dorsally shade the lateral compound eyes. Anterior to the cardiac lobe the median ocelli, simple eyes, can be identified in a number of taxa. Some species bear extraophthalmic ridges, transverse ridges that are arrayed on the
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</paragraph>
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</subSubSection>
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<subSubSection id="C325A3B11C6FA72D9CE72F50FD21FD0E" lastPageId="7" lastPageNumber="8" pageId="3" pageNumber="4" type="description">
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<paragraph id="8B80F03A1C6FA7299CE72F50FE25F880" blockId="3.[164,1441,1654,1902]" pageId="3" pageNumber="4">
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<emphasis id="B94B2C281C6FA7299CE72F50FEFAF94C" bold="true" pageId="3" pageNumber="4">· Figure 1.</emphasis>
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Schematic diagrams of a number of different arthropods showing examples of functional pseudotagmata compared with true tagmata. Somites are labelled 0–XVII; ‘tl’ indicates the telson, ‘m’ the mouth, ‘a’ the anus, and ‘g’ the gonopores. A, the oribatid mite
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<taxonomicName id="4C3F8BB91C6FA72998932FEEFC5BF936" authority="(Berlese, 1904)" baseAuthorityName="Berlese" baseAuthorityYear="1904" box="[506,941,1736,1755]" class="Arachnida" family="Epilohmanniidae" genus="Epilohmannia" kingdom="Animalia" order="Sarcoptiformes" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="cylindrica">
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<emphasis id="B94B2C281C6FA72998932FEEFCF5F936" box="[506,771,1736,1755]" italics="true" pageId="3" pageNumber="4">Epilohmannia cylindrica</emphasis>
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(
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<bibRefCitation id="EFAE8DCB1C6FA7299A7A2FEEFC50F936" author="Berlese A" box="[787,934,1736,1755]" pageId="3" pageNumber="4" pagination="10 - 32" refId="ref14537" refString="Berlese A. 1904. Acari nuovi III. Redia 2: 10 - 32." type="journal article" year="1904">Berlese, 1904</bibRefCitation>
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)
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</taxonomicName>
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. B, the trilobite
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<taxonomicName id="4C3F8BB91C6FA7299D0B2FEEFF16F914" authority="(Rominger, 1887)" baseAuthorityName="Rominger" baseAuthorityYear="1887" class="Trilobita" family="Dorypygidae" genus="Olenoides" kingdom="Animalia" order="Corynexochida" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="serratus">
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<emphasis id="B94B2C281C6FA7299D0B2FEEFAD0F936" box="[1122,1318,1736,1755]" italics="true" pageId="3" pageNumber="4">Olenoides serratus</emphasis>
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(
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<bibRefCitation id="EFAE8DCB1C6FA7299C5F2FEEFF2FF914" author="Rominger C" pageId="3" pageNumber="4" pagination="12 - 19" refId="ref17839" refString="Rominger C. 1887. Description of primordial fossils from Mount Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences of Philadelphia 39: 12 - 19." type="journal article" year="1887">Rominger, 1887</bibRefCitation>
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)
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</taxonomicName>
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. C, the solifuge
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<taxonomicName id="4C3F8BB91C6FA72998C12FC3FCDFF914" authority="Birula, 1929" authorityName="Birula" authorityYear="1929" box="[424,809,1765,1785]" class="Arachnida" family="Galeodidae" genus="Galeodes" kingdom="Animalia" order="Solifugae" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="armeniacus">
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<emphasis id="B94B2C281C6FA72998C12FC3FD7BF915" box="[424,653,1765,1784]" italics="true" pageId="3" pageNumber="4">Galeodes armeniacus</emphasis>
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<bibRefCitation id="EFAE8DCB1C6FA7299BF52FC3FCDFF914" author="Birula AA" box="[668,809,1765,1785]" pageId="3" pageNumber="4" pagination="273 - 282" refId="ref14553" refString="Birula AA. 1929. Uber Galeodes armeniacus n. sp. Zoologischer Anzeiger 84: 273 - 282." type="journal article" year="1929">Birula, 1929</bibRefCitation>
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</taxonomicName>
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. Solifuge arachnids possess opercula on the first three opisthosomal segments associated with the genital ducts and repiratory spiracles. These opercula could be considered homologous to the opercula of the thelyphonid
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<taxonomicName id="4C3F8BB91C6FA7299BFC2E07FBC3F8DE" authority="(Lucas, 1835)" baseAuthorityName="Lucas" baseAuthorityYear="1835" box="[661,1077,1824,1843]" class="Arachnida" family="Thelyphonidae" genus="Mastigoproctus" kingdom="Animalia" order="Uropygi" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="giganteus">
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<emphasis id="B94B2C281C6FA7299BFC2E07FC56F8DE" box="[661,928,1825,1843]" italics="true" pageId="3" pageNumber="4">Mastigoproctus giganteus</emphasis>
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(
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<bibRefCitation id="EFAE8DCB1C6FA7299AC62E06FBD9F8DE" author="Lucas H" box="[943,1071,1824,1843]" pageId="3" pageNumber="4" pagination="17 - 24" refId="ref17103" refString="Lucas H. 1835. Essai sur une monographie du genre Thelyphone. Magasin de Zoologie 8: 17 - 24." type="journal article" year="1835">Lucas, 1835</bibRefCitation>
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)
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</taxonomicName>
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which
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<bibRefCitation id="EFAE8DCB1C6FA7299DEA2E06FAE7F8DE" author="Shultz JW" box="[1155,1297,1824,1843]" pageId="3" pageNumber="4" pagination="335 - 365" refId="ref18269" refString="Shultz JW. 1993. Muscular anatomy of the giant whipscorpion Mastigoproctus giganteus (Lucas) (Arachnida: Uropygi) and its evolutionary significance. Zoological Journal of the Linnean Society 108: 335 - 365." type="journal article" year="1993">Shultz (1993)</bibRefCitation>
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showed to be opisthosomal appendages that had become completely sutured to the ventral body wall. D, the limulid xiphosurid
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<taxonomicName id="4C3F8BB91C6FA7299C202E1BFE39F880" authority="Linnaeus, 1758" authorityName="Linnaeus" authorityYear="1758" class="Merostomata" family="Limulidae" genus="Limulus" kingdom="Animalia" order="Xiphosurida" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="polyphemus">
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<emphasis id="B94B2C281C6FA7299C202E1BFED6F883" italics="true" pageId="3" pageNumber="4">Limulus polyphemus</emphasis>
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<bibRefCitation id="EFAE8DCB1C6FA72998402E7DFE39F880" author="Linnaeus C" box="[297,463,1882,1902]" pageId="3" pageNumber="4" pagination="1 - 824" refId="ref17061" refString="Linnaeus C. 1758. Systema Naturae 1: 1 - 824." type="journal article" year="1758">Linnaeus, 1758</bibRefCitation>
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</taxonomicName>
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.
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</paragraph>
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<paragraph id="8B80F03A1C69A72F99CD29E0FE07FE98" blockId="5.[164,779,198,1902]" pageId="5" pageNumber="6">surface of the carapace outside of the ophthalmic ridges. Interophthalmic ridges occur within the area demarcated by the ophthalmic ridges, and may correspond to internal apodemes for the attachment of extrinsic limb musculature, indicating arrangement of the prosomal appendages.</paragraph>
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<paragraph id="8B80F03A1C69A72F99D5285BFEE3FA8B" blockId="5.[164,779,198,1902]" pageId="5" pageNumber="6">
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The term opisthosoma refers to the post-prosomal segments, and is equivalent to the trunk region of other arthropods (see
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<bibRefCitation id="EFAE8DCB1C69A72F98F6289DFDBBFE3D" author="Hughes NC" box="[415,589,443,465]" pageId="5" pageNumber="6" pagination="185 - 206" refId="ref16497" refString="Hughes NC. 2003 a. Trilobite tagmosis and body patterning from morphological and developmental perspectives. Integrative and Comparative Biology 43: 185 - 206." type="journal article" year="2003">Hughes, 2003a</bibRefCitation>
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, b). The opisthosoma itself is not considered a single tagma, contrary to most recent treatments.
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<bibRefCitation id="EFAE8DCB1C69A72F988028DEFD46FDE3" author="Lankester ER" box="[489,688,504,526]" pageId="5" pageNumber="6" pagination="523 - 582" refId="ref16812" refString="Lankester ER. 1904. The structure and classification of the Arthropoda. Quarterly Journal of Microscopical Science 47: 523 - 582." type="journal article" year="1904">Lankester (1904)</bibRefCitation>
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considered chelicerates to possess three tagmata based on appendage differentiation: the prosoma, mesosoma, and metasoma. The prosoma comprises somites I–VI, the mesosoma somites VII–XIII, and the metasoma somites XIV–XVII (in xiphosurans, eurypterids, scorpions, and probably chasmataspidids the metasoma is formed from somites XIV–XIX).
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<bibRefCitation id="EFAE8DCB1C69A72F9B542BE8FF0FFCEF" author="van der Hammen L" pageId="5" pageNumber="6" pagination="3 - 52" refId="ref16253" refString="van der Hammen L. 1986 a. Comparative studies in Chelicerata IV. Apatellata, Arachnida, Scorpionida, Xiphosura. Zoologische Verhandelingen 226: 3 - 52." type="journal article" year="1986">van der Hammen (1986a)</bibRefCitation>
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termed the mesosoma and metasoma pseudotagmata and this has been followed by subsequent authors including
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<bibRefCitation id="EFAE8DCB1C69A72F98F82A0CFD3DFCAD" author="Siveter DJ & Selden PA" box="[401,715,810,832]" pageId="5" pageNumber="6" pagination="153 - 168" refId="ref18474" refString="Siveter DJ, Selden PA. 1987. A new, giant xiphosurid from the lower Namurian of Weardale, County Durham. Proceedings of the Yorkshire Geological Society 46: 153 - 168." type="journal article" year="1987">Selden & Siveter (1987)</bibRefCitation>
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and
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<bibRefCitation id="EFAE8DCB1C69A72F99CD2A6EFE7AFCB3" author="Rudkin DM & Young GA & Nowlan GS" box="[164,396,840,862]" pageId="5" pageNumber="6" pagination="1 - 9" refId="ref17932" refString="Rudkin DM, Young GA, Nowlan GS. 2008. The oldest horseshoe crab: a new xiphosurid from late Ordovician konservat-lagerstatten deposits, Manitoba, Canada. Palaeontology 51: 1 - 9." type="journal article" year="2008">
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Rudkin
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<emphasis id="B94B2C281C69A72F98682A6FFEC4FCB0" box="[257,306,840,862]" italics="true" pageId="5" pageNumber="6">et al</emphasis>
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. (2008)
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</bibRefCitation>
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, although van der Hammen considered the mesosoma to consist of somites VII–XIV and the metasoma somites XV–XIX (in scorpions). This in fact corresponds to the preabdomen and postabdomen; these are actually non-functional pseudotagmata defined by a dorsal constriction of the tergites (
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<figureCitation id="1304ECBF1C69A72F987D2D26FEAEFBFB" box="[276,344,1024,1046]" captionStart="Figure 2" captionStartId="5.[826,905,1164,1183]" captionTargetBox="[831,1438,197,1133]" captionTargetId="figure-567@5.[830,1439,196,1134]" captionTargetPageId="5" captionText="Figure 2. Schematic of a generalized synziphosurine arthropod, showing the distinction between the preabdominal and postabdominal non-functional pseudotagmata and the true tagmata of the prosoma, mesosoma, and metasoma." figureDoi="http://doi.org/10.5281/zenodo.5292233" httpUri="https://zenodo.org/record/5292233/files/figure.png" pageId="5" pageNumber="6">Fig. 2</figureCitation>
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). In xiphosurans the postabdomen comprises somites XV–XVII, while eurypterids frequently correspond to the condition in scorpions but sometimes have a preabdomen and postabdomen that corresponds to the true tagmata of mesosoma and metasoma. In chasmataspidids the situation is somewhat different, having somites VII–X fused together into a buckler and somites XI–XIX forming a freely articulating postabdomen. These are, however, considered to be functional pseudotagmata, as the fusion of the buckler places a functional constraint on the organism.
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</paragraph>
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<caption id="DF40A0B21C69A72F9A532DAAFC5BFAF8" ID-DOI="http://doi.org/10.5281/zenodo.5292233" ID-Zenodo-Dep="5292233" httpUri="https://zenodo.org/record/5292233/files/figure.png" pageId="5" pageNumber="6" startId="5.[826,905,1164,1183]" targetBox="[831,1438,197,1133]" targetPageId="5">
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<paragraph id="8B80F03A1C69A72F9A532DAAFC5BFAF8" blockId="5.[826,1441,1164,1301]" pageId="5" pageNumber="6">
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<emphasis id="B94B2C281C69A72F9A532DAAFC54FB72" bold="true" box="[826,930,1164,1183]" pageId="5" pageNumber="6">Figure 2.</emphasis>
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Schematic of a generalized synziphosurine arthropod, showing the distinction between the preabdominal and postabdominal non-functional pseudotagmata and the true tagmata of the prosoma, mesosoma, and metasoma.
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</paragraph>
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</caption>
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<paragraph id="8B80F03A1C69A72F99D52C49FBC0F997" blockId="5.[164,779,198,1902]" lastBlockId="5.[826,1441,1360,1902]" pageId="5" pageNumber="6">
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Somite VII is considered to be opisthosomal in origin, as is the conventional view (see
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<bibRefCitation id="EFAE8DCB1C69A72F9BE72CA8FF28FA2F" author="Snodgrass RE" pageId="5" pageNumber="6" pagination="1 - 363" refId="ref18535" refString="Snodgrass RE. 1952. Textbook of arthropod anatomy. New York: Cornell University Press, 1 - 363." type="book chapter" year="1952">Snodgrass, 1952</bibRefCitation>
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), and not prosomal as suggested by
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<bibRefCitation id="EFAE8DCB1C69A72F9BE52C8AFE85FA0C" author="Sturmer W & Bergstrom J" pageId="5" pageNumber="6" pagination="237 - 255" refId="ref18875" refString="Sturmer W, Bergstrom J. 1981. Weinbergina, a xiphosuran arthropod from the Devonian Hunsruck Slate. Palaontologische Zeitschrift 55: 237 - 255." type="journal article" year="1981">Stürmer & Bergström (1981)</bibRefCitation>
|
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(see also
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<bibRefCitation id="EFAE8DCB1C69A72F98922CEDFD2AFA0D" author="Haug C & Van Roy P & Leipner A & Funch P & Rudkin DM & Schollmann L & Haug JT" box="[507,732,1483,1504]" pageId="5" pageNumber="6" pagination="253 - 268" refId="ref16308" refString="Haug C, Van Roy P, Leipner A, Funch P, Rudkin DM, Schollmann L, Haug JT. 2012 a. A holomorph approach to xiphosuran evolution - a case study on the ontogeny of Euproops. Development Genes and Evolution 222: 253 - 268." type="journal article" year="2012">
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Haug
|
||
<emphasis id="B94B2C281C69A72F9B202CEDFD8FFA0D" box="[585,633,1483,1504]" italics="true" pageId="5" pageNumber="6">et al</emphasis>
|
||
., 2012a
|
||
</bibRefCitation>
|
||
for discussion on this subject). Evidence for this is shown in the dorsal expression of somite VII as a fully sclerotized tergite in synziphosurines (
|
||
<figureCitation id="1304ECBF1C69A72F9B1E2F01FD3DF9D0" box="[631,715,1575,1597]" captionStart="Figure 3" captionStartId="6.[144,223,1036,1055]" captionTargetBox="[303,1263,195,1006]" captionTargetId="figure-0@6.[303,1263,195,1006]" captionTargetPageId="6" captionText="Figure 3. A, Bunodes lunula d’Eichwald, 1854 from the late Silurian (Ludlow) of Oesel, Estonia. Specimen ELM G1:262:2, clearly showing the partially reduced pre-opercula tergite of somite VII (arrowed) between the downturned carapace and the hypertrophied tergite of somite VIII. Scale bar = 10 mm. B, ‘Eurypterus’ stoermeri Novojilov, 1959, a chasmataspidid from the early Devonian (Lochkovian) of Siberia, Russia. Specimen PIN 1138-1, exhibiting the microtergite of somite VII (arrowed) positioned between the carapace and the buckler. The microtergite curves anteriorly towards its lateral edges, and is particularly noticeable on the left-hand side. The occurrence of ridges associated with the lateral eye on the carapace is also labelled. Image courtesy of Dave Marshall. Scale bar = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.5292237" httpUri="https://zenodo.org/record/5292237/files/figure.png" pageId="5" pageNumber="6">Fig. 3A</figureCitation>
|
||
) and chasmataspidids (
|
||
<figureCitation id="1304ECBF1C69A72F981D2F63FE30F9B6" box="[372,454,1605,1627]" captionStart="Figure 3" captionStartId="6.[144,223,1036,1055]" captionTargetBox="[303,1263,195,1006]" captionTargetId="figure-0@6.[303,1263,195,1006]" captionTargetPageId="6" captionText="Figure 3. A, Bunodes lunula d’Eichwald, 1854 from the late Silurian (Ludlow) of Oesel, Estonia. Specimen ELM G1:262:2, clearly showing the partially reduced pre-opercula tergite of somite VII (arrowed) between the downturned carapace and the hypertrophied tergite of somite VIII. Scale bar = 10 mm. B, ‘Eurypterus’ stoermeri Novojilov, 1959, a chasmataspidid from the early Devonian (Lochkovian) of Siberia, Russia. Specimen PIN 1138-1, exhibiting the microtergite of somite VII (arrowed) positioned between the carapace and the buckler. The microtergite curves anteriorly towards its lateral edges, and is particularly noticeable on the left-hand side. The occurrence of ridges associated with the lateral eye on the carapace is also labelled. Image courtesy of Dave Marshall. Scale bar = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.5292237" httpUri="https://zenodo.org/record/5292237/files/figure.png" pageId="5" pageNumber="6">Fig. 3B</figureCitation>
|
||
) and in the way the tergite remains attached to the opisthosoma after disarticulation of the carapace (
|
||
<figureCitation id="1304ECBF1C69A72F98BB2FA4FDBDF975" box="[466,587,1666,1688]" captionStart="Figure 4" captionStartId="6.[144,223,1759,1778]" captionTargetBox="[303,1263,1288,1729]" captionTargetId="figure-122@6.[303,1263,1288,1729]" captionTargetPageId="6" captionText="Figure 4. A, Pasternakevia podolica Selden & Drygant, 1987 from the late Silurian (Ludlow) of Zalissia, Ukraine. Specimen ISEA I - F/MP/3/1499/08, isolated opsithosoma with articulated microtergite of somite VII (arrowed). Image courtesy of Ewa Krzemin´ ska. Scale bar = 10 mm. B, undescribed chasmataspidid from the Lower Devonian (Emsian) of Siberia, Russia. Specimen PIN 5116-6, disarticulated buckler with microtergite of somite VII (arrowed) still firmly attached to its anterior margin. Image courtesy of Dave Marshall. Scale bar = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.5292239" httpUri="https://zenodo.org/record/5292239/files/figure.png" pageId="5" pageNumber="6">Fig. 4A, B</figureCitation>
|
||
). This tergite is termed the microtergite in chasmataspidids, but this refers specifically to the heavily reduced condition seen in these taxa and the term pre-opercular tergite is used herein while microtergite is applied as a condition which the pre-opercular tergite can attain. The tergite of somite VIII may be called the opercular tergite and is hypertrophied in some synziphosurines. Embryological studies have shown that in limulids the tergite of somite VII and part of the tergite of somite VIII are incorporated into the prosoma (
|
||
<bibRefCitation id="EFAE8DCB1C69A72F9C3D2CABFC85FA2C" author="Scholl G" pageId="5" pageNumber="6" pagination="99 - 154" refId="ref17968" refString="Scholl G. 1977. Beitrage zur Embryonalen entwicklung von Limulus polyphemus L. (Chelicerata, Xiphosura). Zoomorphologie 86: 99 - 154." type="journal article" year="1977">Scholl, 1977</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFAE8DCB1C69A72F9AEC2C8AFAAEFA2F" author="Sekiguchi K & Yamamichi Y & Costlow JD" box="[901,1368,1452,1474]" pageId="5" pageNumber="6" pagination="53 - 73" refId="ref18119" refString="Sekiguchi K, Yamamichi Y, Costlow JD. 1982. Horseshoe crab developmental studies I. Normal embryonic development of Limulus polyphemus compared with Tachypleus tridentatus. In: Bonaventura J, Bonaventura C, Tesh S, eds. Physiology and biology of horseshoe crabs: studies on normal and environmentally stressed animals. New York: Liss, 53 - 73." type="book chapter" year="1982">Sekiguchi, Yamamichi & Costlow, 1982</bibRefCitation>
|
||
) with the lateral portions of the opercular tergite distinctly set off from the main body of the tergite, frequently deflected dorsally towards the outer margin, and these regions are termed free lobes. The term thoracetron is used for the fully fused dorsal opisthosomal shield in xiphosurids.
|
||
</paragraph>
|
||
<paragraph id="8B80F03A1C69A72D9A3A2FA4FD5CFE5F" blockId="5.[826,1441,1360,1902]" lastBlockId="7.[164,779,198,740]" lastPageId="7" lastPageNumber="8" pageId="5" pageNumber="6">
|
||
The opisthosomal appendages in aquatic chelicerates are modified into flattened opercula that bear the respiratory organs, termed book gills. In xiphosurans the opercula are not medially fused, although in modern xiphosurids they are connected by a thin membrane which would be unlikely to fossilize, and so its presence in extinct taxa is equivocal. The operculum of somite VIII is the genital operculum and bears the paired gonopores; in modern xiphosurids the genital operculum is devoid of respiratory structures, but they are present on the genital operculum of
|
||
<taxonomicName id="4C3F8BB91C6BA72D99CD2804FE35FEDA" authority="Orr et al., 2000" authorityName="Orr" authorityYear="2000" box="[164,451,290,311]" family="Offacolidae" genus="Offacolus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B94B2C281C6BA72D99CD2804FEF8FEDA" box="[164,270,290,311]" italics="true" pageId="7" pageNumber="8">Offacolus</emphasis>
|
||
<bibRefCitation id="EFAE8DCB1C6BA72D987C2804FE35FEDA" author="Orr PJ & Siveter DJ & Briggs DEG & Siveter DJ & Sutton MD" box="[277,451,290,311]" pageId="7" pageNumber="8" pagination="1497 - 1504" refId="ref17507" refString="Orr PJ, Siveter DJ, Briggs DEG, Siveter DJ, Sutton MD. 2000. A new arthropod from the Silurian Konservat- Lagerstatte of Herefordshire, UK. Proceedings of the Royal Society of London 267: 1497 - 1504." type="journal article" year="2000">
|
||
Orr
|
||
<emphasis id="B94B2C281C6BA72D982F2804FE80FEDA" box="[326,374,290,311]" italics="true" pageId="7" pageNumber="8">et al</emphasis>
|
||
., 2000
|
||
</bibRefCitation>
|
||
</taxonomicName>
|
||
and the condition is unclear in
|
||
<taxonomicName id="4C3F8BB91C6BA72D99A12866FEA3FEB8" authorityName="Richter & Richter" authorityYear="1929" box="[200,341,320,341]" class="Merostomata" family="Limulidae" genus="Weinbergina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Xiphosura" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B94B2C281C6BA72D99A12866FEA3FEB8" box="[200,341,320,341]" italics="true" pageId="7" pageNumber="8">Weinbergina</emphasis>
|
||
</taxonomicName>
|
||
. The appendages of somite VII are functionally incorporated into the prosoma and may be retained as fully pediform walking limbs or reduced to chilaria, which act in a masticatory function.
|
||
</paragraph>
|
||
<caption id="DF40A0B21C6AA72C99F92D2AFB3FFB22" ID-DOI="http://doi.org/10.5281/zenodo.5292237" ID-Zenodo-Dep="5292237" httpUri="https://zenodo.org/record/5292237/files/figure.png" pageId="6" pageNumber="7" startId="6.[144,223,1036,1055]" targetBox="[303,1263,195,1006]" targetPageId="6">
|
||
<paragraph id="8B80F03A1C6AA72C99F92D2AFB3FFB22" blockId="6.[144,1422,1036,1231]" pageId="6" pageNumber="7">
|
||
<emphasis id="B94B2C281C6AA72C99F92D2AFF0EFBF2" bold="true" box="[144,248,1036,1055]" pageId="6" pageNumber="7">Figure 3.</emphasis>
|
||
A,
|
||
<taxonomicName id="4C3F8BB91C6AA72C984E2D2AFD6FFBF2" authority="d'Eichwald, 1854" authorityName="d'Eichwald" authorityYear="1854" box="[295,665,1036,1055]" class="Merostomata" family="Bunodidae" genus="Bunodes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Xiphosura" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="lunula">
|
||
<emphasis id="B94B2C281C6AA72C984E2D2AFE25FBF2" box="[295,467,1036,1055]" italics="true" pageId="6" pageNumber="7">Bunodes lunula</emphasis>
|
||
<bibRefCitation id="EFAE8DCB1C6AA72C98892D2AFD6FFBF2" author="d'Eichwald CE" box="[480,665,1036,1055]" pageId="6" pageNumber="7" pagination="1 - 211" refId="ref15844" refString="d'Eichwald CE. 1854. Die Grauwackenschichten von Liev- und Esthland. Bulletin de la Societe Imperiale des Naturalistes de Moscou 27: 1 - 211." type="journal article" year="1854">d’Eichwald, 1854</bibRefCitation>
|
||
</taxonomicName>
|
||
from the late Silurian (Ludlow) of Oesel, Estonia. Specimen ELM G1:262:2, clearly showing the partially reduced pre-opercula tergite of somite VII (arrowed) between the downturned carapace and the hypertrophied tergite of somite VIII. Scale bar = 10 mm. B, ‘
|
||
<taxonomicName id="4C3F8BB91C6AA72C9A8B2D6EFBA0FBB7" box="[994,1110,1096,1114]" class="Merostomata" family="Eurypteridae" genus="Eurypterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eurypterida" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B94B2C281C6AA72C9A8B2D6EFBA0FBB7" box="[994,1110,1096,1114]" italics="true" pageId="6" pageNumber="7">Eurypterus</emphasis>
|
||
</taxonomicName>
|
||
’
|
||
<emphasis id="B94B2C281C6AA72C9D0D2D6EFB33FBB7" box="[1124,1221,1096,1114]" italics="true" pageId="6" pageNumber="7">stoermeri</emphasis>
|
||
<bibRefCitation id="EFAE8DCB1C6AA72C9DA62D61FA85FBB7" author="Novojilov NJ" box="[1231,1395,1095,1114]" pageId="6" pageNumber="7" pagination="241 - 258" refId="ref17450" refString="Novojilov NJ. 1959. Merostomes du Devonian inferieur et moyen de Siberie. Annales de la Societe Geologique du Nord 78: 241 - 258." type="journal article" year="1959">Novojilov, 1959</bibRefCitation>
|
||
, a chasmataspidid from the early Devonian (Lochkovian) of Siberia, Russia. Specimen PIN 1138-1, exhibiting the microtergite of somite VII (arrowed) positioned between the carapace and the buckler. The microtergite curves anteriorly towards its lateral edges, and is particularly noticeable on the left-hand side. The occurrence of ridges associated with the lateral eye on the carapace is also labelled. Image courtesy of Dave Marshall. Scale bar = 2 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF40A0B21C6AA72C99F92FF9FBD6F885" ID-DOI="http://doi.org/10.5281/zenodo.5292239" ID-Zenodo-Dep="5292239" httpUri="https://zenodo.org/record/5292239/files/figure.png" pageId="6" pageNumber="7" startId="6.[144,223,1759,1778]" targetBox="[303,1263,1288,1729]" targetPageId="6">
|
||
<paragraph id="8B80F03A1C6AA72C99F92FF9FBD6F885" blockId="6.[144,1422,1759,1896]" pageId="6" pageNumber="7">
|
||
<emphasis id="B94B2C281C6AA72C99F92FF9FF0EF91F" bold="true" box="[144,248,1759,1778]" pageId="6" pageNumber="7">Figure 4.</emphasis>
|
||
A,
|
||
<taxonomicName id="4C3F8BB91C6AA72C984F2FF9FCC1F91E" authority="Selden & Drygant, 1987" authorityName="Selden & Drygant" authorityYear="1987" box="[294,823,1759,1779]" class="Merostomata" genus="Pasternakevia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Xiphosura" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="podolica">
|
||
<emphasis id="B94B2C281C6AA72C984F2FF9FDE8F91F" box="[294,542,1759,1778]" italics="true" pageId="6" pageNumber="7">Pasternakevia podolica</emphasis>
|
||
<bibRefCitation id="EFAE8DCB1C6AA72C9B432FF9FCC1F91E" author="Selden PA & Drygant DM" box="[554,823,1759,1779]" pageId="6" pageNumber="7" pagination="537 - 542" refId="ref18222" refString="Selden PA, Drygant DM. 1987. A new Silurian xiphosuran from Podolia, Ukraine, USSR. Palaeontology 30: 537 - 542." type="journal article" year="1987">Selden & Drygant, 1987</bibRefCitation>
|
||
</taxonomicName>
|
||
from the late Silurian (Ludlow) of Zalissia, Ukraine. Specimen ISEA I - F/MP/3/1499/08, isolated opsithosoma with articulated microtergite of somite VII (arrowed). Image courtesy of Ewa Krzemin´ ska. Scale bar = 10 mm. B, undescribed chasmataspidid from the Lower Devonian (Emsian) of Siberia, Russia. Specimen PIN 5116-6, disarticulated buckler with microtergite of somite VII (arrowed) still firmly attached to its anterior margin. Image courtesy of Dave Marshall. Scale bar = 2 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8B80F03A1C6BA72D99D5289DFD21FD0E" blockId="7.[164,779,198,740]" pageId="7" pageNumber="8">
|
||
The institutional abbreviations for xiphosuran specimens figured in this study are as follows:
|
||
<materialsCitation id="3B57FA671C6BA72D99CD28DEFF08FDC1" collectionCode="ELM" country="Estonia" location="Estonian Museum of Natural History" municipality="Tallinn" pageId="7" pageNumber="8" specimenCount="1">
|
||
<collectionCode id="ED2E68FF1C6BA72D99CD28DEFF16FDE0" box="[164,224,504,525]" country="South Africa" httpUri="http://grbio.org/cool/9trn-613c" name="East London Museum" pageId="7" pageNumber="8">ELM</collectionCode>
|
||
,
|
||
<location id="8EE0A6E11C6BA72D998728DEFD52FDE3" LSID="urn:lsid:plazi:treatment:0396412C1C6FA72D9D442AFCFD21FD0E:8EE0A6E11C6BA72D998728DEFD52FDE3" box="[238,676,504,526]" country="Estonia" municipality="Tallinn" name="Estonian Museum of Natural History" pageId="7" pageNumber="8">Estonian Museum of Natural History</location>
|
||
,
|
||
<collectingMunicipality id="6BE46A401C6BA72D9BDB28DEFCF1FDE3" box="[690,775,504,526]" pageId="7" pageNumber="8">Tallinn</collectingMunicipality>
|
||
,
|
||
<collectingCountry id="F328B0AA1C6BA72D99CD2B30FF08FDC1" box="[164,254,534,556]" name="Estonia" pageId="7" pageNumber="8">Estonia</collectingCountry>
|
||
</materialsCitation>
|
||
;
|
||
<materialsCitation id="3B57FA671C6BA72D98622B31FD30FDA6" collectionCode="ISEA" country="Poland" county="Krakow" location="Museum" municipality="Evolution of Animals" pageId="7" pageNumber="8" specimenCount="1">
|
||
<collectionCode id="ED2E68FF1C6BA72D98622B31FEBDFDC1" box="[267,331,535,556]" pageId="7" pageNumber="8">ISEA</collectionCode>
|
||
,
|
||
<location id="8EE0A6E11C6BA72D983E2B31FE4CFDC1" LSID="urn:lsid:plazi:treatment:0396412C1C6FA72D9D442AFCFD21FD0E:8EE0A6E11C6BA72D983E2B31FE4CFDC1" box="[343,442,535,556]" country="Poland" county="Krakow" municipality="Evolution of Animals" name="Museum" pageId="7" pageNumber="8">Museum</location>
|
||
of the
|
||
<location id="8EE0A6E11C6BA72D9B642B30FF34FDA6" LSID="urn:lsid:plazi:treatment:0396412C1C6FA72D9D442AFCFD21FD0E:8EE0A6E11C6BA72D9B642B30FF34FDA6" country="Poland" county="Krakow" municipality="Evolution of Animals" name="Institute of Systematics" pageId="7" pageNumber="8">Institute of Systematics</location>
|
||
and
|
||
<collectingMunicipality id="6BE46A401C6BA72D986B2B13FE0AFDA6" box="[258,508,565,587]" pageId="7" pageNumber="8">Evolution of Animals</collectingMunicipality>
|
||
,
|
||
<collectingCounty id="62E188B61C6BA72D9B622B13FD93FDA6" box="[523,613,565,587]" pageId="7" pageNumber="8">Kraków</collectingCounty>
|
||
,
|
||
<collectingCountry id="F328B0AA1C6BA72D9B1D2B13FD30FDA6" box="[628,710,565,587]" name="Poland" pageId="7" pageNumber="8">Poland</collectingCountry>
|
||
</materialsCitation>
|
||
;
|
||
<materialsCitation id="3B57FA671C6BA72D9BBC2B13FD91FD87" collectionCode="MM" country="Canada" location="Manitoba" pageId="7" pageNumber="8" specimenCount="1" stateProvince="Manitoba">
|
||
<collectionCode id="ED2E68FF1C6BA72D9BBC2B13FCF0FDA7" box="[725,774,565,586]" country="France" name="University of Montpellier" pageId="7" pageNumber="8">MM</collectionCode>
|
||
,
|
||
<collectingRegion id="49FB3ED81C6BA72D99CD2B72FEE7FD87" box="[164,273,596,618]" country="Canada" name="Manitoba" pageId="7" pageNumber="8">Manitoba</collectingRegion>
|
||
Museum, Winnipeg,
|
||
<collectingCountry id="F328B0AA1C6BA72D9B652B72FD91FD87" box="[524,615,596,618]" name="Canada" pageId="7" pageNumber="8">Canada</collectingCountry>
|
||
</materialsCitation>
|
||
;
|
||
<materialsCitation id="3B57FA671C6BA72D9B1C2B72FDCBFD65" collectionCode="PIN" country="Russia" location="Paleontological Institute" pageId="7" pageNumber="8" specimenCount="1" stateProvince="Moscow">
|
||
<collectionCode id="ED2E68FF1C6BA72D9B1C2B72FD50FD84" box="[629,678,596,617]" country="Russia" httpUri="http://grbio.org/cool/kjjc-ziy5" name="Paleontological Institute, Russian Academy of Sciences" pageId="7" pageNumber="8">PIN</collectionCode>
|
||
,
|
||
<location id="8EE0A6E11C6BA72D9BDC2B72FE83FD65" LSID="urn:lsid:plazi:treatment:0396412C1C6FA72D9D442AFCFD21FD0E:8EE0A6E11C6BA72D9BDC2B72FE83FD65" country="Russia" name="Paleontological Institute" pageId="7" pageNumber="8" stateProvince="Moscow">Paleontological Institute</location>
|
||
,
|
||
<collectingRegion id="49FB3ED81C6BA72D98ED2B55FE28FD65" box="[388,478,627,648]" country="Russia" name="Moscow" pageId="7" pageNumber="8">Moscow</collectingRegion>
|
||
,
|
||
<collectingCountry id="F328B0AA1C6BA72D98842B54FDCBFD65" box="[493,573,626,648]" name="Russia" pageId="7" pageNumber="8">Russia</collectingCountry>
|
||
</materialsCitation>
|
||
;
|
||
<materialsCitation id="3B57FA671C6BA72D9B252B55FEE7FD29" collectionCode="PWL" country="Germany" county="Mainz" location="Landessammlung" municipality="Naturkunde" pageId="7" pageNumber="8" specimenCount="1" stateProvince="Rheinland-Pfalz">
|
||
<collectionCode id="ED2E68FF1C6BA72D9B252B55FD7EFD65" box="[588,648,627,648]" pageId="7" pageNumber="8">PWL</collectionCode>
|
||
,
|
||
<location id="8EE0A6E11C6BA72D9BFF2B54FEF5FD4A" LSID="urn:lsid:plazi:treatment:0396412C1C6FA72D9D442AFCFD21FD0E:8EE0A6E11C6BA72D9BFF2B54FEF5FD4A" country="Germany" county="Mainz" municipality="Naturkunde" name="Landessammlung" pageId="7" pageNumber="8" stateProvince="Rheinland-Pfalz">Landessammlung</location>
|
||
für
|
||
<collectingMunicipality id="6BE46A401C6BA72D98222BB7FE21FD4A" box="[331,471,657,679]" pageId="7" pageNumber="8">Naturkunde</collectingMunicipality>
|
||
<collectingRegion id="49FB3ED81C6BA72D98832BB7FD53FD4A" box="[490,677,657,679]" country="Germany" name="Rheinland-Pfalz" pageId="7" pageNumber="8">Rheinland-Pfalz</collectingRegion>
|
||
,
|
||
<collectingCounty id="62E188B61C6BA72D9BD52BB7FCF0FD4A" box="[700,774,657,679]" pageId="7" pageNumber="8">Mainz</collectingCounty>
|
||
,
|
||
<collectingCountry id="F328B0AA1C6BA72D99CD2B89FEE7FD29" box="[164,273,687,708]" name="Germany" pageId="7" pageNumber="8">Germany</collectingCountry>
|
||
</materialsCitation>
|
||
;
|
||
<materialsCitation id="3B57FA671C6BA72D98762B89FD25FD0E" collectionCode="SMF" country="Germany" location="Frankfurt am Main" pageId="7" pageNumber="8" specimenCount="1">
|
||
<collectionCode id="ED2E68FF1C6BA72D98762B89FEA0FD29" box="[287,342,687,708]" country="Germany" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34838" lsid="urn:lsid:biocol.org:col:34838" name="Forschungsinstitut und Natur-Museum Senckenberg" pageId="7" pageNumber="8">SMF</collectionCode>
|
||
, Naturmuseum und Forschungsinstitut Senckenberg,
|
||
<location id="8EE0A6E11C6BA72D981B2BE8FDACFD09" LSID="urn:lsid:plazi:treatment:0396412C1C6FA72D9D442AFCFD21FD0E:8EE0A6E11C6BA72D981B2BE8FDACFD09" box="[370,602,718,740]" country="Germany" name="Frankfurt am Main" pageId="7" pageNumber="8">Frankfurt am Main</location>
|
||
,
|
||
<collectingCountry id="F328B0AA1C6BA72D9B012BE8FD25FD0E" box="[616,723,718,739]" name="Germany" pageId="7" pageNumber="8">Germany</collectingCountry>
|
||
</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |