506 lines
58 KiB
XML
506 lines
58 KiB
XML
<document ID-DOI="http://doi.org/10.5281/zenodo.6038818" ID-GBIF-Dataset="285c353d-8675-419a-b12b-8ea887ee64ac" ID-GBIF-Taxon="126106552" ID-Zenodo-Dep="6038818" checkinTime="1484152462690" checkinUser="plazi" docAuthor="Peter Wilf, Mónica R. Carvalho, María A. Gandolfo & N. Rubén Cúneo" docDate="2017" docId="03D67237FF81FF92B72CFB40FCE3E7F8" docLanguage="en" docName="aaag2737.full.pdf.imf" docOrigin="Science 355 (6320)" docStyle="DocumentStyle{}" docTitle="Physalis infinemundi Wilf, sp. nov." docType="treatment" docVersion="16" lastPageId="3" lastPageNumber="4" masterDocId="FFEF0A4FFF81FF91B239FF89FFEFE12C" masterDocTitle="Eocene lantern fruits from Gondwanan Patagonia and the early origins of Solanaceae" masterLastPageNumber="75" masterPageNumber="71" pageId="0" pageNumber="1" updateTime="1644520740356" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Eocene lantern fruits from Gondwanan Patagonia and the early origins of Solanaceae</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Peter Wilf</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Mónica R. Carvalho</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>María A. Gandolfo</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>N. Rubén Cúneo</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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||
<mods:title>Science</mods:title>
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||
</mods:titleInfo>
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||
<mods:part>
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<mods:date>2017</mods:date>
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<mods:detail type="volume">
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<mods:number>355</mods:number>
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</mods:detail>
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<mods:detail type="issue">
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<mods:number>6320</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>71</mods:start>
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<mods:end>75</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1126/science.aag2737</mods:identifier>
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<mods:identifier type="GBIF-Dataset">285c353d-8675-419a-b12b-8ea887ee64ac</mods:identifier>
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<mods:identifier type="Zenodo-Dep">239112</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6038818" ID-GBIF-Taxon="126106552" ID-Zenodo-Dep="6038818" LSID="urn:lsid:plazi:treatment:03D67237FF81FF92B72CFB40FCE3E7F8" httpUri="http://treatment.plazi.org/id/03D67237FF81FF92B72CFB40FCE3E7F8" lastPageId="3" lastPageNumber="4" pageId="0" pageNumber="1">
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||
<subSubSection pageId="0" pageNumber="1" type="nomenclature">
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||
<paragraph blockId="0.[1048,1487,135,1908]" pageId="0" pageNumber="1">
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<taxonomicName authority="Wilf" authorityName="Wilf" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="infinemundi" status="sp. nov.">
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<emphasis box="[1301,1484,1225,1244]" italics="true" pageId="0" pageNumber="1">Physalis infinemundi</emphasis>
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Wilf
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</taxonomicName>
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<taxonomicNameLabel box="[1094,1164,1251,1270]" pageId="0" pageNumber="1" rank="species">sp. nov.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="1" type="etymology">
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<paragraph blockId="0.[1048,1487,135,1908]" pageId="0" pageNumber="1">
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<emphasis bold="true" box="[1171,1285,1251,1270]" pageId="0" pageNumber="1">Etymology:</emphasis>
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Latin,
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<emphasis box="[1352,1481,1251,1270]" italics="true" pageId="0" pageNumber="1">in fine mundi</emphasis>
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, “at the end of the world,” reflecting provenance from the last stages of the Gondwanan supercontinent and from Patagonia, popularly known as “the end of the world.”
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="1" type="materials_examined">
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<paragraph blockId="0.[1048,1487,135,1908]" pageId="0" pageNumber="1">
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<emphasis bold="true" box="[1302,1452,1357,1376]" pageId="0" pageNumber="1">Type material:</emphasis>
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<materialsCitation ID-GBIF-Occurrence="1424526298" collectingDate="2002-12-07" collectionCode="Museo Paleontológico Egidio Feruglio, Trelew, Argentina, S" country="Argentina" location="Laguna del Hunco, Huitrera Formation, early Eocene" pageId="0" pageNumber="1" specimenCode="MPEF-Pb 6434a,b" specimenCount="1" stateProvince="Chubut Province" typeStatus="holotype">
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repository
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<collectionCode pageId="0" pageNumber="1">Museo Paleontológico Egidio Feruglio, Trelew, Argentina</collectionCode>
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(MPEF-Pb);
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<typeStatus box="[1354,1446,1411,1430]" pageId="0" pageNumber="1">holotypus</typeStatus>
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hic designatus
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<specimenCode box="[1158,1329,1437,1456]" pageId="0" pageNumber="1">MPEF-Pb 6434a,b</specimenCode>
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(
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<figureCitation box="[1345,1399,1437,1456]" captionStart="Fig. 1" captionStartId="1.[98,130,248,267]" captionTargetBox="[583,1563,249,1897]" captionTargetId="figure@1.[573,1485,249,1901]" captionTargetPageId="1" captionText="Fig. 1. Physalis infinemundi sp. nov. and selected herbarium specimens of Physalis calyces. (A to C) P. infinemundi holotype, MPEF-Pb 6434 a, b. (A) MPEF-Pb 6434 a, preserving pedicel attachment, venation compressed into the matrix behind the pedicel, four of the presumed five lobe tips, and the coalified berry revealed by cleavage through the calyx. (B) MPEF-Pb 6434 b, preserving a slen- der pedicel segment. (C) MPEF-Pb 6434 a, showing the secondary vein narrowly bifurcating at the arrow, before the lobe sinus visible in (A). (D and E) P. angustifolia, BH 0 0 0 0 79053, showing general features and preservation similar to the fossil holotype and secondary vein bifurcation at the arrow (berry dried and shrunken). (F) P. glutinosa, MICH 1295968, showing exposure of the (dried and shrunken) berry via calyx breakage as in (A) and (B). Scale bar, 1 cm. (G and H) P. infinemundi paratype, MPEF-Pb 6435 a (G) and 6435 b (H), showing invaginated, angled calyx with all five lobes preserved [arrows in (G)], intersecondary veins, and numerous veins compressed into the matrix behind the base. (I) P. angustifolia, MICH 1 5 1 4 1 1 8, showing rounded aspect similar to (G) and (H) versus elongate aspect of the same species in (D); compression of veins behind the pedicel (righthand specimen) similar to both fossils; and distortion effects of flattening under a stem (left-hand specimen) similar to the compression preservation in (G) and (H)." httpUri="https://zenodo.org/record/239113/files/figure.png" pageId="0" pageNumber="1">
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Fig.
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<bibRefCitation author="W. G. D ’ Arcy" box="[1390,1399,1437,1456]" editor="Solanaceae Ill" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="0" pageNumber="1" pagination="75 – 137" part="1991" refString="1. W. G. D ' Arcy, in Solanaceae Ill: Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds. (Royal Botanic Gardens, Kew, UK, 1991), pp. 75 - 137." type="book chapter" volumeTitle="Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds">1</bibRefCitation>
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</figureCitation>
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, A to C) (lowercase letters indicate part and counterpart), from
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<location LSID="urn:lsid:plazi:treatment:03D67237FF81FF92B72CFB40FCE3E7F8:8EA095FAFF81FF91B67EFA5BFBB7E72C" country="Argentina" name="Laguna del Hunco, Huitrera Formation, early Eocene" pageId="0" pageNumber="1" stateProvince="Chubut Province">Laguna del Hunco, Huitrera Formation, early Eocene</location>
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,
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<collectingRegion box="[1121,1269,1517,1536]" country="Argentina" name="Chubut" pageId="0" pageNumber="1">Chubut Province</collectingRegion>
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,
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<collectingCountry box="[1278,1367,1517,1536]" name="Argentina" pageId="0" pageNumber="1">Argentina</collectingCountry>
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,
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<location LSID="urn:lsid:plazi:treatment:03D67237FF81FF92B72CFB40FCE3E7F8:8EA095FAFF81FF91B766FA64FA22E72C" box="[1375,1485,1517,1536]" country="Argentina" name="quarry LH 13" pageId="0" pageNumber="1" stateProvince="Chubut Province">quarry LH13</location>
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(
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<bibRefCitation author="P. Wilf" box="[1055,1074,1543,1562]" journalOrPublisher="Science" pageId="0" pageNumber="1" pagination="122 - 125" part="300" refString="16. P. Wilf et al., Science 300, 122 - 125 (2003)." title="et al" type="journal article" year="2003">
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<emphasis box="[1055,1074,1543,1562]" italics="true" pageId="0" pageNumber="1">16</emphasis>
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</bibRefCitation>
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), collected
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<collectingDate box="[1179,1340,1543,1562]" pageId="0" pageNumber="1" value="2002-12-07">
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<date box="[1179,1340,1543,1562]" pageId="0" pageNumber="1" value="2002-12-07">7 December 2002</date>
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</collectingDate>
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(fig.
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<collectionCode box="[1388,1400,1543,1562]" pageId="0" pageNumber="1">S</collectionCode>
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<bibRefCitation author="M. A. Gandolfo & et al." box="[1400,1422,1543,1562]" journalOrPublisher="PLOS ONE" pageId="0" pageNumber="1" pagination="21084" part="6" refString="18. M. A. Gandolfo et al., PLOS ONE 6, e 21084 (2011)." type="journal article" year="2011">18</bibRefCitation>
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);
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</materialsCitation>
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<materialsCitation ID-GBIF-Occurrence="1424526299" collectingDate="2006-11-21" location="Laguna del Hunco quarry LH 04" pageId="0" pageNumber="1" specimenCode="MPEF-Pb 6435a,b" specimenCount="1" typeStatus="paratype">
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<typeStatus pageId="0" pageNumber="1">paratype</typeStatus>
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<specimenCode box="[1092,1254,1570,1589]" pageId="0" pageNumber="1">MPEF-Pb 6435a,b</specimenCode>
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(
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<figureCitation box="[1266,1314,1570,1589]" captionStart="Fig. 1" captionStartId="1.[98,130,248,267]" captionTargetBox="[583,1563,249,1897]" captionTargetId="figure@1.[573,1485,249,1901]" captionTargetPageId="1" captionText="Fig. 1. Physalis infinemundi sp. nov. and selected herbarium specimens of Physalis calyces. (A to C) P. infinemundi holotype, MPEF-Pb 6434 a, b. (A) MPEF-Pb 6434 a, preserving pedicel attachment, venation compressed into the matrix behind the pedicel, four of the presumed five lobe tips, and the coalified berry revealed by cleavage through the calyx. (B) MPEF-Pb 6434 b, preserving a slen- der pedicel segment. (C) MPEF-Pb 6434 a, showing the secondary vein narrowly bifurcating at the arrow, before the lobe sinus visible in (A). (D and E) P. angustifolia, BH 0 0 0 0 79053, showing general features and preservation similar to the fossil holotype and secondary vein bifurcation at the arrow (berry dried and shrunken). (F) P. glutinosa, MICH 1295968, showing exposure of the (dried and shrunken) berry via calyx breakage as in (A) and (B). Scale bar, 1 cm. (G and H) P. infinemundi paratype, MPEF-Pb 6435 a (G) and 6435 b (H), showing invaginated, angled calyx with all five lobes preserved [arrows in (G)], intersecondary veins, and numerous veins compressed into the matrix behind the base. (I) P. angustifolia, MICH 1 5 1 4 1 1 8, showing rounded aspect similar to (G) and (H) versus elongate aspect of the same species in (D); compression of veins behind the pedicel (righthand specimen) similar to both fossils; and distortion effects of flattening under a stem (left-hand specimen) similar to the compression preservation in (G) and (H)." httpUri="https://zenodo.org/record/239113/files/figure.png" pageId="0" pageNumber="1">
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Fig.
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<bibRefCitation author="W. G. D ’ Arcy" box="[1305,1314,1570,1589]" editor="Solanaceae Ill" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="0" pageNumber="1" pagination="75 – 137" part="1991" refString="1. W. G. D ' Arcy, in Solanaceae Ill: Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds. (Royal Botanic Gardens, Kew, UK, 1991), pp. 75 - 137." type="book chapter" volumeTitle="Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds">1</bibRefCitation>
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||
</figureCitation>
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||
, G and H),
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<location LSID="urn:lsid:plazi:treatment:03D67237FF81FF92B72CFB40FCE3E7F8:8EA095FAFF81FF91B7B3F9ABFB1EE77C" name="Laguna del Hunco quarry LH 04" pageId="0" pageNumber="1">Laguna del Hunco quarry LH04</location>
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||
(
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||
<bibRefCitation author="P. Wilf" box="[1277,1296,1597,1616]" journalOrPublisher="Science" pageId="0" pageNumber="1" pagination="122 - 125" part="300" refString="16. P. Wilf et al., Science 300, 122 - 125 (2003)." title="et al" type="journal article" year="2003">
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<emphasis box="[1277,1296,1597,1616]" italics="true" pageId="0" pageNumber="1">16</emphasis>
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||
</bibRefCitation>
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||
),
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<collectingDate box="[1313,1483,1597,1616]" pageId="0" pageNumber="1" value="2006-11-21">
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<date box="[1313,1483,1597,1616]" pageId="0" pageNumber="1" value="2006-11-21">21 November 2006</date>
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</collectingDate>
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||
.
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</materialsCitation>
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||
</paragraph>
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||
</subSubSection>
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||
<subSubSection lastPageId="3" lastPageNumber="4" pageId="0" pageNumber="1" type="description">
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||
<paragraph blockId="0.[1048,1487,135,1908]" lastBlockId="1.[98,537,135,208]" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1">
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<emphasis bold="true" box="[1048,1170,1623,1642]" pageId="0" pageNumber="1">Description:</emphasis>
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based on two dispersed, apparently mature, compressed fruiting calyces; only the holotype preserves berry and pedicel remains. Pedicel preserved length 18.8 mm, width 1 mm at inflection point of insertion. Calyx basally invaginated, highly inflated to completely surround the berry, five-lobed, angled, partly open at apex, length by width 2 5.2 mm by 1 3.2 mm (
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<bibRefCitation author="W. G. D ’ Arcy" box="[1320,1328,1809,1828]" editor="Solanaceae Ill" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="0" pageNumber="1" pagination="75 – 137" part="1991" refString="1. W. G. D ' Arcy, in Solanaceae Ill: Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds. (Royal Botanic Gardens, Kew, UK, 1991), pp. 75 - 137." type="book chapter" volumeTitle="Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds">1</bibRefCitation>
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||
.
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<bibRefCitation author="M. Millan & W. Crepet" box="[1332,1343,1809,1828]" journalOrPublisher="Bot. Rev." pageId="0" pageNumber="1" pagination="3 - 106" part="80" refString="9. M. Millan, W. Crepet, Bot. Rev. 80, 7 3 - 106 (2014)." title="9" type="journal article" year="2014">9</bibRefCitation>
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||
:
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<bibRefCitation author="W. G. D ’ Arcy" box="[1349,1357,1809,1828]" editor="Solanaceae Ill" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="0" pageNumber="1" pagination="75 – 137" part="1991" refString="1. W. G. D ' Arcy, in Solanaceae Ill: Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds. (Royal Botanic Gardens, Kew, UK, 1991), pp. 75 - 137." type="book chapter" volumeTitle="Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds">1</bibRefCitation>
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) on holotype, 21.6 mm by 17.6 mm (ratio 1.2:
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<bibRefCitation author="W. G. D ’ Arcy" box="[1347,1354,1836,1855]" editor="Solanaceae Ill" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="0" pageNumber="1" pagination="75 – 137" part="1991" refString="1. W. G. D ' Arcy, in Solanaceae Ill: Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds. (Royal Botanic Gardens, Kew, UK, 1991), pp. 75 - 137." type="book chapter" volumeTitle="Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds">1</bibRefCitation>
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) on paratype. Calyx lobes equally sized, sinuses angular and incised one-quarter to half the total calyx length, tips acute triangular. Venation with one robust primary meridional vein per lobe, terminating at lobe apex and alternating with secondary veins.
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</paragraph>
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<footnote pageId="0" pageNumber="1">
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<paragraph blockId="0.[98,537,1760,1907]" pageId="0" pageNumber="1">
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1Department of Geosciences, Pennsylvania State University, University Park,PA 16802, USA. 2L. H. Bailey Hortorium, Plant Biology Section, School of Integrative Plant
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<docJournal box="[401,456,1805,1822]" pageId="0" pageNumber="1">Science</docJournal>
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||
, Cornell University, Ithaca, NY 14853,USA. 3Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Museo PaleontolÓgico Egidio Feruglio,9100 Trelew, Chubut, Argentina. *Corresponding author. Email: pwilf@psu.edu
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</paragraph>
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</footnote>
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<paragraph blockId="1.[573,1012,135,208]" lastBlockId="3.[98,537,135,1748]" lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2">
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Secondary veins arising near the base, visually distinct from the primaries and other vein orders, and bifurcating close to the lobe sinuses (
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<figureCitation box="[975,1056,135,208]" captionStart="Fig. 1" captionStartId="1.[98,130,248,267]" captionTargetBox="[583,1563,249,1897]" captionTargetId="figure@1.[573,1485,249,1901]" captionTargetPageId="1" captionText="Fig. 1. Physalis infinemundi sp. nov. and selected herbarium specimens of Physalis calyces. (A to C) P. infinemundi holotype, MPEF-Pb 6434 a, b. (A) MPEF-Pb 6434 a, preserving pedicel attachment, venation compressed into the matrix behind the pedicel, four of the presumed five lobe tips, and the coalified berry revealed by cleavage through the calyx. (B) MPEF-Pb 6434 b, preserving a slen- der pedicel segment. (C) MPEF-Pb 6434 a, showing the secondary vein narrowly bifurcating at the arrow, before the lobe sinus visible in (A). (D and E) P. angustifolia, BH 0 0 0 0 79053, showing general features and preservation similar to the fossil holotype and secondary vein bifurcation at the arrow (berry dried and shrunken). (F) P. glutinosa, MICH 1295968, showing exposure of the (dried and shrunken) berry via calyx breakage as in (A) and (B). Scale bar, 1 cm. (G and H) P. infinemundi paratype, MPEF-Pb 6435 a (G) and 6435 b (H), showing invaginated, angled calyx with all five lobes preserved [arrows in (G)], intersecondary veins, and numerous veins compressed into the matrix behind the base. (I) P. angustifolia, MICH 1 5 1 4 1 1 8, showing rounded aspect similar to (G) and (H) versus elongate aspect of the same species in (D); compression of veins behind the pedicel (righthand specimen) similar to both fossils; and distortion effects of flattening under a stem (left-hand specimen) similar to the compression preservation in (G) and (H)." httpUri="https://zenodo.org/record/239113/files/figure.png" pageId="1" pageNumber="2">
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||
Fig.
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<bibRefCitation author="W. G. D ’ Arcy" box="[1048,1056,135,154]" editor="Solanaceae Ill" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="1" pageNumber="2" pagination="75 – 137" part="1991" refString="1. W. G. D ' Arcy, in Solanaceae Ill: Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds. (Royal Botanic Gardens, Kew, UK, 1991), pp. 75 - 137." type="book chapter" volumeTitle="Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds">1</bibRefCitation>
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||
</figureCitation>
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||
C). Intersecondary veins arising near the base, visually distinct from secondaries and tertiaries, dichotomizing into the random, irregular reticulum of tertiary through at least quinternary veins that fill most of the vein field. Some basal veins compressed into partial detachment from the calyx body, the remnants visible against the matrix. Berry round, flattened and coalified from fossilization, filling the full width of the calyx and appearing by surface relief to extend apically almost to the lobe sinuses; seeds not preserved.
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</paragraph>
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</subSubSection>
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||
<caption httpUri="https://zenodo.org/record/239113/files/figure.png" pageId="1" pageNumber="2" targetBox="[583,1563,249,1897]" targetPageId="1">
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||
<paragraph blockId="1.[98,538,248,1037]" pageId="1" pageNumber="2">
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||
<emphasis bold="true" pageId="1" pageNumber="2">
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Fig. 1.
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<taxonomicName box="[160,342,248,267]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="infinemundi" status="sp. nov.">
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<emphasis bold="true" box="[160,342,248,267]" italics="true" pageId="1" pageNumber="2">Physalis infinemundi</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[348,416,248,267]" pageId="1" pageNumber="2" rank="species">sp. nov.</taxonomicNameLabel>
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and selected herbarium specimens of
|
||
<taxonomicName box="[330,405,275,294]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="genus">
|
||
<emphasis bold="true" box="[330,405,275,294]" italics="true" pageId="1" pageNumber="2">Physalis</emphasis>
|
||
</taxonomicName>
|
||
calyces.
|
||
</emphasis>
|
||
(
|
||
<emphasis bold="true" box="[498,512,275,294]" pageId="1" pageNumber="2">A</emphasis>
|
||
to
|
||
<emphasis bold="true" box="[98,112,301,320]" pageId="1" pageNumber="2">C</emphasis>
|
||
)
|
||
<taxonomicName box="[127,247,301,320]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="infinemundi">
|
||
<emphasis box="[127,247,301,320]" italics="true" pageId="1" pageNumber="2">P. infinemundi</emphasis>
|
||
</taxonomicName>
|
||
holotype, MPEF-Pb 6434a,b. (A) MPEF-Pb 6434a, preserving pedicel attachment, venation compressed into the matrix behind the pedicel, four of the presumed five lobe tips, and the coalified berry revealed by cleavage through the calyx. (B) MPEF-Pb 6434b, preserving a slen- der pedicel segment. (C) MPEF-Pb 6434a, showing the secondary vein narrowly bifurcating at the arrow, before the lobe sinus visible in (A). (
|
||
<emphasis bold="true" box="[482,497,514,533]" pageId="1" pageNumber="2">D</emphasis>
|
||
and
|
||
<emphasis bold="true" box="[98,110,540,559]" pageId="1" pageNumber="2">E</emphasis>
|
||
)
|
||
<taxonomicName box="[126,250,540,559]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="angustifolia">
|
||
<emphasis box="[126,250,540,559]" italics="true" pageId="1" pageNumber="2">P. angustifolia</emphasis>
|
||
</taxonomicName>
|
||
, BH 0 0 0 0 79053, showing general features and preservation similar to the fossil holotype and secondary vein bifurcation at the arrow (berry dried and shrunken). (
|
||
<emphasis bold="true" box="[407,418,620,639]" pageId="1" pageNumber="2">F</emphasis>
|
||
)
|
||
<taxonomicName box="[430,530,620,639]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="glutinosa">
|
||
<emphasis box="[430,530,620,639]" italics="true" pageId="1" pageNumber="2">P. glutinosa</emphasis>
|
||
</taxonomicName>
|
||
, MICH 1295968, showing exposure of the (dried and shrunken) berry via calyx breakage as in (A) and (B). Scale bar, 1 cm. (
|
||
<emphasis bold="true" box="[327,342,700,719]" pageId="1" pageNumber="2">G</emphasis>
|
||
and
|
||
<emphasis bold="true" box="[385,400,700,719]" pageId="1" pageNumber="2">H</emphasis>
|
||
)
|
||
<taxonomicName box="[412,536,700,719]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="infinemundi">
|
||
<emphasis box="[412,536,700,719]" italics="true" pageId="1" pageNumber="2">P. infinemundi</emphasis>
|
||
</taxonomicName>
|
||
paratype, MPEF-Pb 6435a (G) and 6435b (H), showing invaginated, angled calyx with all five lobes preserved [arrows in (G)], intersecondary veins, and numerous veins compressed into the matrix behind the base. (
|
||
<emphasis bold="true" box="[258,264,832,851]" pageId="1" pageNumber="2">I</emphasis>
|
||
)
|
||
<taxonomicName box="[277,399,832,851]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="angustifolia">
|
||
<emphasis box="[277,399,832,851]" italics="true" pageId="1" pageNumber="2">P. angustifolia</emphasis>
|
||
</taxonomicName>
|
||
, MICH 1 5 1 4 1 1 8, showing rounded aspect similar to (G) and (H) versus elongate aspect of the same species in (D); compression of veins behind the pedicel (righthand specimen) similar to both fossils; and distortion effects of flattening under a stem (left-hand specimen) similar to the compression preservation in (G) and (H).
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/344619/files/figure.png" pageId="2" pageNumber="3" targetBox="[147,963,178,1904]" targetPageId="2">
|
||
<paragraph blockId="2.[1048,1487,181,493]" pageId="2" pageNumber="3">
|
||
<emphasis bold="true" pageId="2" pageNumber="3">
|
||
Fig. 2. Phylogenetic relationships of
|
||
<taxonomicName class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="2" pageNumber="3" phylum="Tracheophyta" rank="species" species="infinemundi" status="sp. nov.">
|
||
<emphasis bold="true" italics="true" pageId="2" pageNumber="3">Physalis infinemundi</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[1167,1241,208,227]" pageId="2" pageNumber="3" rank="species">sp. nov.</taxonomicNameLabel>
|
||
and selected
|
||
<taxonomicName box="[1377,1485,208,227]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="2" pageNumber="3" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
species.
|
||
</emphasis>
|
||
Strict consensus of 2835 most parsimonious trees of 3510 steps (CI = 0.438, RI = 0.726), based on a total evidence analysis using morphology and five gene partitions (ITS,
|
||
<emphasis box="[1333,1375,314,333]" italics="true" pageId="2" pageNumber="3">waxy</emphasis>
|
||
,
|
||
<emphasis box="[1385,1431,314,333]" italics="true" pageId="2" pageNumber="3">matK</emphasis>
|
||
,
|
||
<emphasis box="[1441,1481,314,333]" italics="true" pageId="2" pageNumber="3">ndhF</emphasis>
|
||
, and
|
||
<emphasis box="[1087,1136,341,360]" italics="true" pageId="2" pageNumber="3">trnL-F</emphasis>
|
||
; 7070 total characters). Decay indices are included above each branch (
|
||
<emphasis box="[1353,1376,367,386]" italics="true" pageId="2" pageNumber="3">20</emphasis>
|
||
) (higher indices represent stronger support). Labeled major clades follow (
|
||
<emphasis box="[1171,1191,421,440]" italics="true" pageId="2" pageNumber="3">10</emphasis>
|
||
). See the text and supplementary materials (
|
||
<emphasis box="[1143,1166,447,466]" italics="true" pageId="2" pageNumber="3">20</emphasis>
|
||
) for details and fig. S1 for the maximum likelihood tree.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection pageId="3" pageNumber="4" type="diagnosis">
|
||
<paragraph blockId="3.[98,537,135,1748]" pageId="3" pageNumber="4">
|
||
The fossils preserve several features that, together, are diagnostic of the
|
||
<taxonomicName box="[377,478,401,420]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
clade of
|
||
<taxonomicName box="[126,230,428,447]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
(
|
||
<emphasis box="[248,271,428,447]" italics="true" pageId="3" pageNumber="4">20</emphasis>
|
||
,
|
||
<bibRefCitation author="B. Axelius" box="[286,306,428,447]" journalOrPublisher="Am. J. Bot." pageId="3" pageNumber="4" pagination="118 - 124" part="83" refString="21. B. Axelius, Am. J. Bot. 83, 118 - 124 (1996)." title="21" type="journal article" year="1996">
|
||
<emphasis box="[286,306,428,447]" italics="true" pageId="3" pageNumber="4">21</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="E. Estrada & M. Martinez" box="[320,342,428,447]" editor="M. Nee" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="3" pageNumber="4" pagination="139 - 159" part="1999" refString="23. E. Estrada, M. Martinez, in Solanaceae IV: Advances in Biology and Utilization, M. Nee, D. E. Symon, R. N. Lester, J. P. Jessop, Eds. (Royal Botanic Gardens, Kew, UK, 1999), pp. 139 - 159." title="23" type="journal article" volumeTitle="Solanaceae IV: Advances in Biology and Utilization" year="1996">
|
||
<emphasis box="[320,342,428,447]" italics="true" pageId="3" pageNumber="4">23</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="M. Whitson & P. S. Manos" box="[358,380,428,447]" journalOrPublisher="Syst. Bot." pageId="3" pageNumber="4" pagination="216 - 230" part="30" refString="25. M. Whitson, P. S. Manos, Syst. Bot. 30, 216 - 230 (2005)." title="25" type="journal article" year="2005">
|
||
<emphasis box="[358,380,428,447]" italics="true" pageId="3" pageNumber="4">25</emphasis>
|
||
</bibRefCitation>
|
||
). These include the highly inflated, lantern-shaped, pedicellate, five-parted, angled, regular calyces covering a large, fleshy berry, along with the combination of (i) invaginated base, (ii) robust primary veins that terminate in the lobe tips (not the lobe sinuses), and (iii) secondary veins that fork before the sinus. Nonphysaline species of
|
||
<taxonomicName class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
with ICS lack some or all of these three features (e.g., those in
|
||
<taxonomicName box="[290,352,667,686]" class="Magnoliopsida" family="Solanaceae" genus="Deprea" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[290,352,667,686]" italics="true" pageId="3" pageNumber="4">Deprea</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName box="[361,444,667,686]" class="Magnoliopsida" family="Solanaceae" genus="Nicandra" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[361,444,667,686]" italics="true" pageId="3" pageNumber="4">Nicandra</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName box="[453,531,667,686]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[453,531,667,686]" italics="true" pageId="3" pageNumber="4">Solanum</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName box="[98,181,693,712]" class="Magnoliopsida" family="Solanaceae" genus="Withania" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[98,181,693,712]" italics="true" pageId="3" pageNumber="4">Withania</emphasis>
|
||
</taxonomicName>
|
||
, and the
|
||
<taxonomicName box="[263,367,693,712]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="tribe" tribe="Juanulloeae">Juanulloeae</taxonomicName>
|
||
clade) (figs. S
|
||
<bibRefCitation author="K. Kindscher" box="[486,496,693,712]" journalOrPublisher="Econ. Bot." pageId="3" pageNumber="4" pagination="298 - 310" part="66" refString="5. K. Kindscher et al., Econ. Bot. 66, 298 - 310 (2012)." title="et al" type="journal article" year="2012">5</bibRefCitation>
|
||
, S
|
||
<bibRefCitation author="R. G. Olmstead" box="[515,532,693,712]" journalOrPublisher="Bot. J. Linn. Soc." pageId="3" pageNumber="4" pagination="8" part="171" refString="11. R. G. Olmstead, Bot. J. Linn. Soc. 171, 8 0 - 102 (2013)." title="11" type="journal article" year="2013">11</bibRefCitation>
|
||
, and S
|
||
<bibRefCitation author="M. P. Zamora-Tavares & M. Martinez & S. MagallOn & L. Guzman-Davalos & O. Vargas-Ponce" box="[151,172,720,739]" journalOrPublisher="Mol. Phyl. Evol." pageId="3" pageNumber="4" pagination="41 - 50" part="100" refString="13. M. P. Zamora-Tavares, M. Martinez, S. MagallOn, L. Guzman-Davalos, O. Vargas-Ponce, Mol. Phyl. Evol. 100, 41 - 50 (2016)." title="13" type="journal article" year="2016">13</bibRefCitation>
|
||
) (
|
||
<bibRefCitation author="E. Estrada & M. Martinez" box="[192,214,720,739]" editor="M. Nee" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="3" pageNumber="4" pagination="139 - 159" part="1999" refString="23. E. Estrada, M. Martinez, in Solanaceae IV: Advances in Biology and Utilization, M. Nee, D. E. Symon, R. N. Lester, J. P. Jessop, Eds. (Royal Botanic Gardens, Kew, UK, 1999), pp. 139 - 159." title="23" type="journal article" volumeTitle="Solanaceae IV: Advances in Biology and Utilization" year="1996">
|
||
<emphasis box="[192,214,720,739]" italics="true" pageId="3" pageNumber="4">23</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="G. E. Barboza" box="[225,248,720,739]" journalOrPublisher="Kurtziana" pageId="3" pageNumber="4" pagination="9 - 79" part="28" refString="24. G. E. Barboza, Kurtziana 28, 6 9 - 79 (2000)." title="24" type="journal article" year="2000">
|
||
<emphasis box="[225,248,720,739]" italics="true" pageId="3" pageNumber="4">24</emphasis>
|
||
</bibRefCitation>
|
||
). Within
|
||
<taxonomicName box="[338,441,720,739]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
, there are several problematic species of
|
||
<taxonomicName box="[385,464,746,765]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus" sensu="lato">
|
||
<emphasis box="[385,464,746,765]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[469,495,746,765]" pageId="3" pageNumber="4" sensu="lato">s.l.</taxonomicNameLabel>
|
||
and other genera that diverge below the core
|
||
<taxonomicName box="[462,536,773,792]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[462,536,773,792]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
clade (
|
||
<figureCitation box="[154,202,800,819]" captionStart="Fig. 2" captionStartId="2.[1048,1080,181,200]" captionTargetBox="[147,963,178,1904]" captionTargetId="figure@2.[538,571,134,1913]" captionTargetPageId="2" captionText="Fig. 2. Phylogenetic relationships of Physalis infinemundi sp. nov. and selected Solanaceae species. Strict consensus of 2835 most parsimonious trees of 3510 steps (CI = 0.438, RI = 0.726), based on a total evidence analysis using morphology and five gene partitions (ITS, waxy, matK, ndhF, and trnL-F; 7070 total characters). Decay indices are included above each branch (20) (higher indices represent stronger support). Labeled major clades follow (10). See the text and supplementary materials (20) for details and fig. S 1 for the maximum likelihood tree." httpUri="https://zenodo.org/record/344619/files/figure.png" pageId="3" pageNumber="4">
|
||
Fig.
|
||
<bibRefCitation author="A. T. Hunziker" box="[191,202,800,819]" journalOrPublisher="The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein" pageId="3" pageNumber="4" refString="2. A. T. Hunziker, The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein, 2001)." title="2" type="book" year="2001">2</bibRefCitation>
|
||
</figureCitation>
|
||
) (equivalent to
|
||
<emphasis box="[336,353,800,819]" italics="true" pageId="3" pageNumber="4">P.</emphasis>
|
||
subgenus
|
||
<emphasis box="[443,529,800,819]" italics="true" pageId="3" pageNumber="4">Rydbergis</emphasis>
|
||
) of relatively homogeneous species (
|
||
<bibRefCitation author="M. P. Zamora-Tavares & M. Martinez & S. MagallOn & L. Guzman-Davalos & O. Vargas-Ponce" box="[436,456,826,845]" journalOrPublisher="Mol. Phyl. Evol." pageId="3" pageNumber="4" pagination="41 - 50" part="100" refString="13. M. P. Zamora-Tavares, M. Martinez, S. MagallOn, L. Guzman-Davalos, O. Vargas-Ponce, Mol. Phyl. Evol. 100, 41 - 50 (2016)." title="13" type="journal article" year="2016">
|
||
<emphasis box="[436,456,826,845]" italics="true" pageId="3" pageNumber="4">13</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<emphasis box="[468,524,826,845]" italics="true" pageId="3" pageNumber="4">
|
||
<bibRefCitation author="E. Estrada & M. Martinez" box="[468,490,826,845]" editor="M. Nee" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="3" pageNumber="4" pagination="139 - 159" part="1999" refString="23. E. Estrada, M. Martinez, in Solanaceae IV: Advances in Biology and Utilization, M. Nee, D. E. Symon, R. N. Lester, J. P. Jessop, Eds. (Royal Botanic Gardens, Kew, UK, 1999), pp. 139 - 159." title="23" type="journal article" volumeTitle="Solanaceae IV: Advances in Biology and Utilization" year="1996">23</bibRefCitation>
|
||
–
|
||
<bibRefCitation author="M. Whitson & P. S. Manos" box="[502,524,826,845]" journalOrPublisher="Syst. Bot." pageId="3" pageNumber="4" pagination="216 - 230" part="30" refString="25. M. Whitson, P. S. Manos, Syst. Bot. 30, 216 - 230 (2005)." title="25" type="journal article" year="2005">25</bibRefCitation>
|
||
</emphasis>
|
||
). Among these taxa,
|
||
<taxonomicName box="[276,487,853,872]" class="Magnoliopsida" family="Solanaceae" genus="Alkekengi" higherTaxonomySource="CoL" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="species" species="officinarum">
|
||
<emphasis box="[276,487,853,872]" italics="true" pageId="3" pageNumber="4">Alkekengi officinarum</emphasis>
|
||
</taxonomicName>
|
||
calyces are the most similar to the fossils, but they have a much larger size, consistently rounded shape, and nearly closed apex. Otherwise, each of these basal physaline species structurally differs from the fossils and core
|
||
<taxonomicName box="[360,432,986,1005]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[360,432,986,1005]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<emphasis box="[445,466,986,1005]" italics="true" pageId="3" pageNumber="4">20</emphasis>
|
||
,
|
||
<bibRefCitation author="G. E. Barboza" box="[477,498,986,1005]" journalOrPublisher="Kurtziana" pageId="3" pageNumber="4" pagination="9 - 79" part="28" refString="24. G. E. Barboza, Kurtziana 28, 6 9 - 79 (2000)." title="24" type="journal article" year="2000">
|
||
<emphasis box="[477,498,986,1005]" italics="true" pageId="3" pageNumber="4">24</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="M. Whitson & P. S. Manos" box="[509,529,986,1005]" journalOrPublisher="Syst. Bot." pageId="3" pageNumber="4" pagination="216 - 230" part="30" refString="25. M. Whitson, P. S. Manos, Syst. Bot. 30, 216 - 230 (2005)." title="25" type="journal article" year="2005">
|
||
<emphasis box="[509,529,986,1005]" italics="true" pageId="3" pageNumber="4">25</emphasis>
|
||
</bibRefCitation>
|
||
) (figs. S
|
||
<bibRefCitation author="A. T. Hunziker" box="[156,167,1012,1031]" journalOrPublisher="The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein" pageId="3" pageNumber="4" refString="2. A. T. Hunziker, The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein, 2001)." title="2" type="book" year="2001">2</bibRefCitation>
|
||
to S
|
||
<bibRefCitation box="[211,231,1012,1031]" pageId="3" pageNumber="4" refString="17. P. Wilf, N. R. Cuneo, I. H. Escapa, D. Pol, M. O. Woodburne, Annu. Rev. Earth Planet. Sci. 41, 561 - 603 (2013)">17</bibRefCitation>
|
||
). All these observations highlight the close relationship of the fossils with core
|
||
<taxonomicName box="[98,177,1065,1084]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[98,177,1065,1084]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="3" pageNumber="4" type="discussion">
|
||
<paragraph blockId="3.[98,537,135,1748]" lastBlockId="3.[573,1012,135,1748]" pageId="3" pageNumber="4">
|
||
Both the MP and ML results (
|
||
<figureCitation box="[384,433,1092,1111]" captionStart="Fig. 2" captionStartId="2.[1048,1080,181,200]" captionTargetBox="[147,963,178,1904]" captionTargetId="figure@2.[538,571,134,1913]" captionTargetPageId="2" captionText="Fig. 2. Phylogenetic relationships of Physalis infinemundi sp. nov. and selected Solanaceae species. Strict consensus of 2835 most parsimonious trees of 3510 steps (CI = 0.438, RI = 0.726), based on a total evidence analysis using morphology and five gene partitions (ITS, waxy, matK, ndhF, and trnL-F; 7070 total characters). Decay indices are included above each branch (20) (higher indices represent stronger support). Labeled major clades follow (10). See the text and supplementary materials (20) for details and fig. S 1 for the maximum likelihood tree." httpUri="https://zenodo.org/record/344619/files/figure.png" pageId="3" pageNumber="4">
|
||
Fig.
|
||
<bibRefCitation author="A. T. Hunziker" box="[422,433,1092,1111]" journalOrPublisher="The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein" pageId="3" pageNumber="4" refString="2. A. T. Hunziker, The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein, 2001)." title="2" type="book" year="2001">2</bibRefCitation>
|
||
</figureCitation>
|
||
and fig. S
|
||
<bibRefCitation author="W. G. D ’ Arcy" box="[522,529,1092,1111]" editor="Solanaceae Ill" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="3" pageNumber="4" pagination="75 – 137" part="1991" refString="1. W. G. D ' Arcy, in Solanaceae Ill: Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds. (Royal Botanic Gardens, Kew, UK, 1991), pp. 75 - 137." type="book chapter" volumeTitle="Taxonomy, Chemistry, Evolution, J. G. Hawkes, R. N. Lester, M. Nee, N. Estrada, Eds">1</bibRefCitation>
|
||
) support the fossils’ affinity with
|
||
<taxonomicName box="[397,494,1118,1137]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
and
|
||
<taxonomicName box="[98,174,1145,1164]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[98,174,1145,1164]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
. The newly identified species is placed in
|
||
<taxonomicName box="[125,224,1172,1191]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
with strong support in both MP and ML analyses and at a basal polytomy of (MP, weak support) or within (ML, strong support) the crown of core
|
||
<taxonomicName box="[238,317,1251,1270]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[238,317,1251,1270]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
. Both the MP and ML topologies are generally consistent with and reproduced most major clades from (
|
||
<bibRefCitation author="T. Sarkinen & L. Bohs & R. G. Olmstead & S. Knapp & BMC Evol" box="[454,474,1304,1323]" journalOrPublisher="Biol" pageId="3" pageNumber="4" pagination="214" part="13" refString="10. T. Sarkinen, L. Bohs, R. G. Olmstead, S. Knapp, BMC Evol. Biol. 13, 214 (2013)." title="10" type="journal article" year="2013">
|
||
<emphasis box="[454,474,1304,1323]" italics="true" pageId="3" pageNumber="4">10</emphasis>
|
||
</bibRefCitation>
|
||
), with robust support of the critical
|
||
<taxonomicName box="[376,475,1331,1350]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
clade. However, each tree recovered the closely related (
|
||
<bibRefCitation author="T. Sarkinen & L. Bohs & R. G. Olmstead & S. Knapp & BMC Evol" box="[156,176,1384,1403]" journalOrPublisher="Biol" pageId="3" pageNumber="4" pagination="214" part="13" refString="10. T. Sarkinen, L. Bohs, R. G. Olmstead, S. Knapp, BMC Evol. Biol. 13, 214 (2013)." title="10" type="journal article" year="2013">
|
||
<emphasis box="[156,176,1384,1403]" italics="true" pageId="3" pageNumber="4">10</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
<taxonomicName box="[189,308,1384,1403]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Iochrominae">Iochrominae</taxonomicName>
|
||
and
|
||
<taxonomicName box="[354,420,1384,1403]" class="Magnoliopsida" family="Solanaceae" genus="Deprea" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[354,420,1384,1403]" italics="true" pageId="3" pageNumber="4">Deprea</emphasis>
|
||
</taxonomicName>
|
||
as collapsed into a single clade that also included
|
||
<taxonomicName box="[444,530,1411,1430]" class="Magnoliopsida" family="Solanaceae" genus="Nicandra" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[444,530,1411,1430]" italics="true" pageId="3" pageNumber="4">Nicandra</emphasis>
|
||
</taxonomicName>
|
||
, which has
|
||
<taxonomicName box="[197,264,1437,1456]" class="Magnoliopsida" family="Solanaceae" genus="Deprea" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[197,264,1437,1456]" italics="true" pageId="3" pageNumber="4">Deprea</emphasis>
|
||
</taxonomicName>
|
||
-like calyx morphology but is not closely related to that genus. Nevertheless, the overall agreement of the MP and ML tree topologies with (
|
||
<bibRefCitation author="T. Sarkinen & L. Bohs & R. G. Olmstead & S. Knapp & BMC Evol" box="[262,282,1517,1536]" journalOrPublisher="Biol" pageId="3" pageNumber="4" pagination="214" part="13" refString="10. T. Sarkinen, L. Bohs, R. G. Olmstead, S. Knapp, BMC Evol. Biol. 13, 214 (2013)." title="10" type="journal article" year="2013">
|
||
<emphasis box="[262,282,1517,1536]" italics="true" pageId="3" pageNumber="4">10</emphasis>
|
||
</bibRefCitation>
|
||
) is notable in light of the limitations of the analysis. Notably, no species from outside
|
||
<taxonomicName box="[222,319,1570,1589]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
were misplaced among
|
||
<taxonomicName box="[98,196,1597,1616]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
, and no
|
||
<taxonomicName box="[275,372,1597,1616]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
species were misplaced into other clades. Considering the unknown additional organs of this extinct species, the incomplete knowledge of calyx morphology in extant
|
||
<taxonomicName box="[226,329,1703,1722]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
, and problematic resolution among extant basal species of
|
||
<taxonomicName box="[439,536,1729,1748]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="M. P. Zamora-Tavares & M. Martinez & S. MagallOn & L. Guzman-Davalos & O. Vargas-Ponce" box="[581,601,135,154]" journalOrPublisher="Mol. Phyl. Evol." pageId="3" pageNumber="4" pagination="41 - 50" part="100" refString="13. M. P. Zamora-Tavares, M. Martinez, S. MagallOn, L. Guzman-Davalos, O. Vargas-Ponce, Mol. Phyl. Evol. 100, 41 - 50 (2016)." title="13" type="journal article" year="2016">
|
||
<emphasis box="[581,601,135,154]" italics="true" pageId="3" pageNumber="4">13</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="M. Whitson & P. S. Manos" box="[614,636,135,154]" journalOrPublisher="Syst. Bot." pageId="3" pageNumber="4" pagination="216 - 230" part="30" refString="25. M. Whitson, P. S. Manos, Syst. Bot. 30, 216 - 230 (2005)." title="25" type="journal article" year="2005">
|
||
<emphasis box="[614,636,135,154]" italics="true" pageId="3" pageNumber="4">25</emphasis>
|
||
</bibRefCitation>
|
||
), we suggest that the newly identified species can be used, conservatively, to constrain the divergence of
|
||
<taxonomicName box="[742,843,189,208]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
to a minimum of 52.2 Ma.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="3" pageNumber="4" type="discussion">
|
||
<paragraph blockId="3.[573,1012,135,1748]" pageId="3" pageNumber="4">
|
||
<taxonomicName box="[594,801,242,261]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="species" species="infinemundi" status="sp. nov.">
|
||
<emphasis box="[594,801,242,261]" italics="true" pageId="3" pageNumber="4">Physalis infinemundi</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[810,885,242,261]" pageId="3" pageNumber="4" rank="species">sp. nov.</taxonomicNameLabel>
|
||
represents a derived lineage of
|
||
<taxonomicName authority="in Gondwanan" authorityName="in Gondwanan" box="[755,1011,268,287]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae in Gondwanan</taxonomicName>
|
||
South America at 52.2 Ma, pushing back considerably the evolutionary timing of the family. Compared with recent molecular divergence estimates (
|
||
<bibRefCitation author="T. Sarkinen & L. Bohs & R. G. Olmstead & S. Knapp & BMC Evol" box="[680,701,374,393]" journalOrPublisher="Biol" pageId="3" pageNumber="4" pagination="214" part="13" refString="10. T. Sarkinen, L. Bohs, R. G. Olmstead, S. Knapp, BMC Evol. Biol. 13, 214 (2013)." title="10" type="journal article" year="2013">
|
||
<emphasis box="[680,701,374,393]" italics="true" pageId="3" pageNumber="4">10</emphasis>
|
||
</bibRefCitation>
|
||
), these fossils are considerably older than the ~30 Ma crown for the entire
|
||
<taxonomicName class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
family as well as the ~11 Ma divergence of
|
||
<taxonomicName box="[598,704,454,473]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
. A second recent study similarly placed the
|
||
<taxonomicName box="[712,818,481,500]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="subTribe" subTribe="Physalinae">Physalinae</taxonomicName>
|
||
divergence at only ~9 Ma (
|
||
<bibRefCitation author="M. P. Zamora-Tavares & M. Martinez & S. MagallOn & L. Guzman-Davalos & O. Vargas-Ponce" box="[652,673,507,526]" journalOrPublisher="Mol. Phyl. Evol." pageId="3" pageNumber="4" pagination="41 - 50" part="100" refString="13. M. P. Zamora-Tavares, M. Martinez, S. MagallOn, L. Guzman-Davalos, O. Vargas-Ponce, Mol. Phyl. Evol. 100, 41 - 50 (2016)." title="13" type="journal article" year="2016">
|
||
<emphasis box="[652,673,507,526]" italics="true" pageId="3" pageNumber="4">13</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="3.[573,1012,135,1748]" pageId="3" pageNumber="4">
|
||
Our results reinforce the emerging pattern wherein numerous fossil plant taxa from Gondwanan Patagonia and Antarctica are substantially older than their corresponding molecular dates (
|
||
<bibRefCitation author="P. Wilf & I. H. Escapa" box="[632,653,640,659]" journalOrPublisher="New Phytol" pageId="3" pageNumber="4" pagination="283 - 290" part="207" refString="26. P. Wilf, I. H. Escapa, New Phytol. 207, 283 - 290 (2015)." title="26" type="journal article" year="2015">
|
||
<emphasis box="[632,653,640,659]" italics="true" pageId="3" pageNumber="4">26</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="V. D. Barreda" box="[663,682,640,659]" journalOrPublisher="Proc. Natl. Acad. Sci. U. S. A." pageId="3" pageNumber="4" pagination="10989 - 10994" part="112" refString="27. V. D. Barreda et al., Proc. Natl. Acad. Sci. U. S. A. 112, 10989 - 10994 (2015)." title="et al" type="journal article" year="2015">
|
||
<emphasis box="[663,682,640,659]" italics="true" pageId="3" pageNumber="4">27</emphasis>
|
||
</bibRefCitation>
|
||
), demonstrating Gondwanan history for groups conjectured to have post-Gondwanan origins under entirely different paleogeographic and paleoclimatic scenarios. Likewise, the derived position of the newly identified fossil species shows that the origins and diversification of
|
||
<taxonomicName box="[597,699,800,819]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
must have taken place at a much earlier time than previously thought, considerably before final Gondwanan breakup. Other regions of Gondwana are also likely to have played prominent roles in
|
||
<taxonomicName box="[840,944,906,925]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
evolution, especially Antarctica, which has produced other important asterid fossils (
|
||
<bibRefCitation author="V. D. Barreda" box="[876,897,959,978]" journalOrPublisher="Proc. Natl. Acad. Sci. U. S. A." pageId="3" pageNumber="4" pagination="10989 - 10994" part="112" refString="27. V. D. Barreda et al., Proc. Natl. Acad. Sci. U. S. A. 112, 10989 - 10994 (2015)." title="et al" type="journal article" year="2015">
|
||
<emphasis box="[876,897,959,978]" italics="true" pageId="3" pageNumber="4">27</emphasis>
|
||
</bibRefCitation>
|
||
). Moreover, the newly identified fossils directly help to resolve temporal inconsistencies between the evolutionary timing of
|
||
<taxonomicName box="[791,895,1039,1058]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
|
||
and its herbivores and mutualists (
|
||
<bibRefCitation author="I. J. GarzOn-Orduna & K. L. Silva-Brandao & K. R. Willmott & A. V. L. Freitas & A. V. Z. Brower" box="[814,837,1065,1084]" journalOrPublisher="Syst. Biol." pageId="3" pageNumber="4" pagination="752 - 767" part="64" refString="28. I. J. GarzOn-Orduna, K. L. Silva-Brandao, K. R. Willmott, A. V. L. Freitas, A. V. Z. Brower, Syst. Biol. 64, 752 - 767 (2015)." title="28" type="journal article" year="2015">
|
||
<emphasis box="[814,837,1065,1084]" italics="true" pageId="3" pageNumber="4">28</emphasis>
|
||
</bibRefCitation>
|
||
). The large fossil berry strongly implicates trophic associations with animals, as seen in extant
|
||
<taxonomicName box="[882,961,1118,1137]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[882,961,1118,1137]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="M. F. Willson" box="[976,999,1118,1137]" journalOrPublisher="Oikos" pageId="3" pageNumber="4" pagination="159 - 176" part="67" refString="29. M. F. Willson, Oikos 67, 159 - 176 (1993)." title="29" type="journal article" year="1993">
|
||
<emphasis box="[976,999,1118,1137]" italics="true" pageId="3" pageNumber="4">29</emphasis>
|
||
</bibRefCitation>
|
||
). Today,
|
||
<taxonomicName box="[646,725,1145,1164]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[646,725,1145,1164]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
inhabits South, Central, and North America, and Mexico is its center of diversity (
|
||
<bibRefCitation author="A. T. Hunziker" box="[651,662,1198,1217]" journalOrPublisher="The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein" pageId="3" pageNumber="4" refString="2. A. T. Hunziker, The Genera of Solanaceae (A. R. G. Gantner Verlag K. - G., Ruggell, Lichtenstein, 2001)." title="2" type="book" year="2001">
|
||
<emphasis box="[651,662,1198,1217]" italics="true" pageId="3" pageNumber="4">2</emphasis>
|
||
</bibRefCitation>
|
||
). Thus, the fossils establish a rare link to extant New World floras from late-Gondwanan Patagonian assemblages, whose living relatives are mostly concentrated in the Old World tropics and subtropics.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="3" pageNumber="4" type="biology_ecology">
|
||
<paragraph blockId="3.[573,1012,135,1748]" pageId="3" pageNumber="4">
|
||
The discovery of 52.2-Ma fossil inflated calyces demonstrates an ancient history for ICS and implies that the Eocene world was already populated with derived solanaceous reproductive structures. The potential adaptive functions of the inflated calyx are scarcely discussed in the literature (
|
||
<bibRefCitation author="J. Zhang & Y. Tian & L. Wang & C. He & J. Syst" box="[677,700,1490,1509]" journalOrPublisher="Evol" pageId="3" pageNumber="4" pagination="4 - 101" part="48" refString="30. J. Zhang, Y. Tian, L. Wang, C. He, J. Syst. Evol. 48, 9 4 - 101 (2010)." title="30" type="journal article" year="2010">
|
||
<emphasis box="[677,700,1490,1509]" italics="true" pageId="3" pageNumber="4">30</emphasis>
|
||
</bibRefCitation>
|
||
), but the lakeside rainforest paleoenvironment led us to consider flotation dispersal and protective drying for the berry. Simple kitchen-sink and stream experiments on several
|
||
<taxonomicName box="[573,652,1597,1616]" class="Magnoliopsida" family="Solanaceae" genus="Physalis" kingdom="Plantae" order="Solanales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[573,652,1597,1616]" italics="true" pageId="3" pageNumber="4">Physalis</emphasis>
|
||
</taxonomicName>
|
||
species confirmed that intact calyces hold stable air pockets around the berry, enabling flotation for several days in water and preventing wetting of the berry during rains. These insights suggest potential origins of ICS in ancient humid, riparian environments.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |