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<document id="60EB19B7CF290F167A72F040D5C824A6" ID-DOI="10.17348/jbrit.v15.i2.1206" ID-Zenodo-Dep="14111022" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_requiresApprovalFor="plazi" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1634280027544" checkinUser="donat" docAuthor="Fawcett, Susan &amp; Smith, Alan R." docDate="2021" docId="03B787F6FFDC9B7C62627B16FCE9FE0C" docLanguage="en" docName="2021_ThelypteridaceaeFerns_OACopy3.pdf.imf" docOrigin="Fort Worth, Texas, USA: BRIT Press" docTitle="Goniopteris" docType="treatment" docVersion="4" lastPageNumber="45" masterDocId="FF8EFF8EFFE99B4B602B7D49FFA7FFBB" masterDocTitle="A Generic Classification of the Thelypteridaceae" pageNumber="43" updateTime="1731444927364" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-SA-4.0">
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<mods:titleInfo id="6174C0AD86B25EF099E951682D7C555E">
<mods:title id="721AC0A2162A83459BFDED1F3AAD186D">A Generic Classification of the Thelypteridaceae</mods:title>
</mods:titleInfo>
<mods:name id="E75F72FF403CB08186032CD84AF697A4" type="personal">
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<mods:namePart id="CFDF6D7D4390783FFD233CD46D740D38">Fawcett, Susan</mods:namePart>
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<mods:namePart id="C2F5AA8FD0D2D2A2190B99B1007F7493">Smith, Alan R.</mods:namePart>
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<treatment id="03B787F6FFDC9B7C62627B16FCE9FE0C" ID-DOI="http://doi.org/10.5281/zenodo.14076336" ID-Zenodo-Dep="14076336" LSID="urn:lsid:plazi:treatment:03B787F6FFDC9B7C62627B16FCE9FE0C" httpUri="http://treatment.plazi.org/id/03B787F6FFDC9B7C62627B16FCE9FE0C" lastPageId="55" lastPageNumber="45" pageId="53" pageNumber="43">
<subSubSection id="C304656BFFDC9B7E62627B16FCB7F9C2" box="[585,784,1631,1657]" pageId="53" pageNumber="43" type="nomenclature">
<paragraph id="8BA136E0FFDC9B7E62627B16FCB7F9C2" blockId="53.[121,1251,1631,1745]" box="[585,784,1631,1657]" pageId="53" pageNumber="43">
<heading id="D0E9818CFFDC9B7E62627B16FCB7F9C2" allCaps="true" bold="true" box="[585,784,1631,1657]" centered="true" fontSize="11" level="2" pageId="53" pageNumber="43" reason="0">
<taxonomicName id="4C1E4D63FFDC9B7E62627B16FCB7F9C2" ID-CoL="4Q6Q" box="[585,784,1631,1657]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="53" pageNumber="43" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDC9B7E62627B16FCB7F9C2" bold="true" box="[585,784,1631,1657]" pageId="53" pageNumber="43">GONIOPTERIS</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDC9B7E60527BC1FCC7F90C" pageId="53" pageNumber="43" type="reference_group">
<paragraph id="8BA136E0FFDC9B7E60527BC1FCC7F90C" blockId="53.[121,1251,1631,1745]" pageId="53" pageNumber="43">
<taxonomicName id="4C1E4D63FFDC9B7E60527BC1FE18F926" authority="C. Presl, Tent. Pterid." authorityName="C. Presl, Tent. Pterid." box="[121,447,1671,1693]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="53" pageNumber="43" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDC9B7E60527BC1FF55F926" bold="true" box="[121,242,1672,1693]" pageId="53" pageNumber="43">Goniopteris</emphasis>
C. Presl, Tent.Pterid.
</taxonomicName>
181183, pl. 7, f.
<date id="FFA01020FFDC9B7E62717BC1FD67F926" box="[602,704,1672,1693]" pageId="53" pageNumber="43" value="1836-11-09">911.1836</date>
.—
<taxonomicName id="4C1E4D63FFDC9B7E62F67BC3FB23F927" authority="(C. Presl) Duek, Adansonia" authorityName="Duek, Adansonia" baseAuthorityName="C. Presl" box="[733,1156,1674,1692]" class="Polypodiopsida" family="Thelypteridaceae" genus="Thelypteris" kingdom="Plantae" order="Polypodiales" pageId="53" pageNumber="43" phylum="Tracheophyta" rank="subGenus" subGenus="Goniopteris">
<emphasis id="B96AEAF2FFDC9B7E62F67BC3FC88F920" box="[733,815,1674,1691]" italics="true" pageId="53" pageNumber="43">Thelypteris</emphasis>
subg.
<emphasis id="B96AEAF2FFDC9B7E63767BC2FC14F920" box="[861,947,1675,1691]" italics="true" pageId="53" pageNumber="43">Goniopteris</emphasis>
(C.Presl) Duek,Adansonia
</taxonomicName>
,n.s., 11:720. 1971.—
<treatmentCitation id="0ABF10F1FFDC9B7E60D87BECFCFBF90C" author="C. V. Morton" box="[243,860,1701,1719]" issue="4" journal="Amer. Fern J." page="154" pageId="53" pageNumber="43" score="85.37874681426794" volume="53" year="1963">
<taxonomicName id="4C1E4D63FFDC9B7E60D87BECFD43F90E" ID-CoL="8H87" authority="(C. Presl) C. V. Morton, Amer. Fern J." authorityName="C. V. Morton, Amer. Fern J." baseAuthorityName="C. Presl" box="[243,740,1701,1719]" class="Polypodiopsida" family="Thelypteridaceae" genus="Thelypteris" kingdom="Plantae" order="Polypodiales" pageId="53" pageNumber="43" phylum="Tracheophyta" rank="section" section="Goniopteris">
<emphasis id="B96AEAF2FFDC9B7E60D87BECFEE2F90D" box="[243,325,1701,1718]" italics="true" pageId="53" pageNumber="43">Thelypteris</emphasis>
sect.
<emphasis id="B96AEAF2FFDC9B7E615B7BECFE61F90E" box="[368,454,1701,1717]" italics="true" pageId="53" pageNumber="43">Goniopteris</emphasis>
(C. Presl) C.V. Morton, Amer. Fern J.
</taxonomicName>
53(4):154. 1963
</treatmentCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDC9B7E63757BEFFD1CF96A" pageId="53" pageNumber="43" type="materials_examined">
<paragraph id="8BA136E0FFDC9B7E63757BEFFD1CF96A" blockId="53.[121,1251,1631,1745]" pageId="53" pageNumber="43">
<typeStatus id="54A58842FFDC9B7E63587BECFC1AF90E" box="[883,957,1701,1717]" pageId="53" pageNumber="43" type="lectotype">LECTOTYPE</typeStatus>
(designated by
<collectionCode id="ED0FAE25FFDC9B7E641E7BECFB9AF90E" box="[1077,1085,1701,1717]" country="South Africa" lsid="urn:lsid:biocol.org:col:14647" name="University of the Witwatersrand" pageId="53" pageNumber="43" type="Herbarium">J</collectionCode>
.
<collectingRegion id="49DAF802FFDC9B7E646F7BEFFBD4F90C" box="[1092,1139,1702,1719]" country="Bermuda" name="Smith's" pageId="53" pageNumber="43">Smith</collectingRegion>
, Hist. Fil. 191. 1875):
<taxonomicName id="4C1E4D63FFDC9B7E60C07B89FE45F96A" authority="(Sw.) C. Presl" authorityName="C. Presl" baseAuthorityName="Sw." box="[235,482,1728,1745]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="53" pageNumber="43" phylum="Tracheophyta" rank="species" species="crenata">
<emphasis id="B96AEAF2FFDC9B7E60C07B89FEDCF96B" box="[235,379,1728,1744]" italics="true" pageId="53" pageNumber="43">Goniopteris crenata</emphasis>
(Sw.) C.Presl
</taxonomicName>
[=
<taxonomicName id="4C1E4D63FFDC9B7E61D27B89FD11F96A" authority="(Bory) Ching" authorityName="Ching" baseAuthorityName="Bory" box="[505,694,1728,1745]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="53" pageNumber="43" phylum="Tracheophyta" rank="species" species="poiteana">
<emphasis id="B96AEAF2FFDC9B7E61D27B89FDEBF96B" box="[505,588,1728,1744]" italics="true" pageId="53" pageNumber="43">G. poiteana</emphasis>
(Bory) Ching
</taxonomicName>
]
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDC9B7E60B27BACFCBEF94C" box="[153,793,1765,1783]" pageId="53" pageNumber="43" type="reference_group">
<paragraph id="8BA136E0FFDC9B7E60B27BACFCBEF94C" blockId="53.[153,793,1765,1783]" box="[153,793,1765,1783]" pageId="53" pageNumber="43">
For additional generic synonymy, see
<bibRefCitation id="EF8F4B11FFDC9B7E619F7BAFFDE6F94C" author="Christensen" box="[436,577,1766,1783]" firstAuthor="Christensen" journalOrPublisher="Selsk. Skr., Naturvidensk. Math. Afd." pageId="53" pageNumber="43" pagination="55 - 282" part="10" refId="ref107711" refString="CHRISTENSEN, C. 1913. A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 55 - 282." title="A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk" type="journal article" year="1913">Christensen (1913</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFDC9B7E626D7BAFFDCAF94C" author="Christensen" box="[582,621,1766,1783]" firstAuthor="Christensen" journalOrPublisher="Selsk. Skr., Naturvidensk Math. Afd." pageId="53" pageNumber="43" pagination="3 - 132" part="6" refId="ref107760" refString="CHRISTENSEN, C. 1920. A monograph of the genus Dryopteris, part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk Math. Afd., ser. 8, 6: 3 - 132." title="A monograph of the genus Dryopteris, part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk" type="journal article" year="1920">1920</bibRefCitation>
),
<bibRefCitation id="EF8F4B11FFDC9B7E62527BACFD45F94C" author="Proctor" box="[633,738,1765,1783]" firstAuthor="Proctor" journalOrPublisher="British Museum (Natural History), London, UK" pageId="53" pageNumber="43" refId="ref113097" refString="PROCTOR, G. R. 1985. Ferns of Jamaica. British Museum (Natural History), London, UK." title="Ferns of Jamaica" type="book" year="1985">Proctor (1985</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFDC9B7E62C37BAFFCA8F94C" author="Proctor" box="[744,783,1766,1783]" firstAuthor="Proctor" journalOrPublisher="Mem. New York Bot. Gard." pageId="53" pageNumber="43" pagination="1 - 389" part="53" refId="ref113120" refString="PROCTOR, G. R. 1989. Ferns of Puerto Rico and the Virgin Islands. Mem. New York Bot. Gard. 53: 1 - 389." title="Ferns of Puerto Rico and the Virgin Islands" type="journal article" year="1989">1989</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDF9B7D604A7DE8FF45FF4C" pageId="54" pageNumber="44" type="etymology">
<paragraph id="8BA136E0FFDF9B7D604A7DE8FF45FF4C" blockId="54.[97,1226,161,1783]" pageId="54" pageNumber="44">
<emphasis id="B96AEAF2FFDF9B7D604A7DE8FF6EFF0D" box="[97,201,161,182]" italics="true" pageId="54" pageNumber="44">Etymology.</emphasis>
—Gr.
<emphasis id="B96AEAF2FFDF9B7D61287DEBFE92FF0C" box="[259,309,162,183]" italics="true" pageId="54" pageNumber="44">gonia</emphasis>
, angle +
<emphasis id="B96AEAF2FFDF9B7D61A37DEBFE1BFF0C" box="[392,444,162,183]" italics="true" pageId="54" pageNumber="44">pteris</emphasis>
, fern, in reference to the angle of often acutely anastomosing veins below the sinus or these producing an excurrent vein joined by other connivent veins; this contrasts to the often obliquely united veins of
<taxonomicName id="4C1E4D63FFDF9B7D60537DABFF7BFF4C" box="[120,220,226,247]" class="Polypodiopsida" family="Thelypteridaceae" genus="Meniscium" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDF9B7D60537DABFF7BFF4C" box="[120,220,226,247]" italics="true" pageId="54" pageNumber="44">Meniscium</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDF9B7D60BB7C4BFC67FC6C" pageId="54" pageNumber="44" type="description">
<paragraph id="8BA136E0FFDF9B7D60BB7C4BFC67FC6C" blockId="54.[97,1226,161,1783]" pageId="54" pageNumber="44">
Plants terrestrial or epilithic, small to medium-sized (
<quantity id="4CE69B05FFDF9B7D62A57C4BFD42FEAC" box="[654,741,258,279]" metricMagnitude="-1" metricUnit="m" metricValue="4.25" metricValueMax="8.0" metricValueMin="0.5" pageId="54" pageNumber="44" unit="cm" value="42.5" valueMax="80.0" valueMin="5.0">580 cm</quantity>
);
<emphasis id="B96AEAF2FFDF9B7D62DE7C4BFCF4FEAC" bold="true" box="[757,851,258,279]" pageId="54" pageNumber="44">rhizomes</emphasis>
typically stout, short-creeping, ascending, or erect, scales generally with stellate or furcate hairs on surfaces and/or margins;
<emphasis id="B96AEAF2FFDF9B7D63A27C6BFC6BFE8C" bold="true" box="[905,972,290,311]" pageId="54" pageNumber="44">fronds</emphasis>
erect, arching, pendent, or prostrate, monomorphic, or weakly dimorphic (e.g.,
<taxonomicName id="4C1E4D63FFDF9B7D62657C0BFCB5FEEC" box="[590,786,322,343]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="tetragona">
<emphasis id="B96AEAF2FFDF9B7D62657C0BFCB5FEEC" box="[590,786,322,343]" italics="true" pageId="54" pageNumber="44">Goniopteris tetragona</emphasis>
</taxonomicName>
), typically pinnate-pinnatifid, rarely simple or pinnate-pinnatisect;
<emphasis id="B96AEAF2FFDF9B7D61077C2BFECFFECC" bold="true" box="[300,360,354,375]" pageId="54" pageNumber="44">stipes</emphasis>
stramineous to dull brown, grooved adaxially;
<emphasis id="B96AEAF2FFDF9B7D63147C2BFC11FECD" bold="true" box="[831,950,353,375]" pageId="54" pageNumber="44">stipe scales</emphasis>
brown, deltate-ovate to linear-lanceolate, typically with furcate or stellate hairs on surfaces and/or margins;
<emphasis id="B96AEAF2FFDF9B7D63567CC8FC18FE2D" bold="true" box="[893,959,385,406]" pageId="54" pageNumber="44">blades</emphasis>
membranaceous to thickly coriaceous, often drying dark green, apex conform or gradually reduced, base of blade unreduced, or, if reduced, without many pairs of gradually reduced pinnae, proliferous buds sometimes present adaxially along rachises, in axils of pinnae, sometimes also along costae (e.g.,
<taxonomicName id="4C1E4D63FFDF9B7D61DA7CABFD9DFE4D" box="[497,570,481,503]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="alata">
<emphasis id="B96AEAF2FFDF9B7D61DA7CABFD9DFE4D" box="[497,570,481,503]" italics="true" pageId="54" pageNumber="44">G. alata</emphasis>
</taxonomicName>
);
<emphasis id="B96AEAF2FFDF9B7D62667CABFD34FE4C" bold="true" box="[589,659,482,503]" pageId="54" pageNumber="44">pinnae</emphasis>
petiolulate, sessile, or narrowly adnate (especially distally), margins simple, crenate to deeply lobed, bases dilated, truncate, rounded, hastate, or with acroscopic auricles;
<emphasis id="B96AEAF2FFDF9B7D64BF7F4BFB6EFDAC" bold="true" box="[1172,1225,514,535]" pageId="54" pageNumber="44">veins</emphasis>
anastomosing, at an angle with veins from adjacent segments, forming an excurrent veinlet to the sinus, and creating one to a series of many areoles between costae and pinna margins (e.g.,
<taxonomicName id="4C1E4D63FFDF9B7D63247F0BFCD8FDEC" box="[783,895,578,599]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="tetragona">
<emphasis id="B96AEAF2FFDF9B7D63247F0BFCD8FDEC" box="[783,895,578,599]" italics="true" pageId="54" pageNumber="44">G. tetragona</emphasis>
</taxonomicName>
), or veins simple, free or forked in smaller 1-pinnate spp. (e.g.,
<taxonomicName id="4C1E4D63FFDF9B7D615A7F2BFE61FDCD" box="[369,454,609,631]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="abdita">
<emphasis id="B96AEAF2FFDF9B7D615A7F2BFE61FDCD" box="[369,454,609,631]" italics="true" pageId="54" pageNumber="44">G. abdita</emphasis>
</taxonomicName>
);
<emphasis id="B96AEAF2FFDF9B7D61F17F28FDECFDCD" bold="true" box="[474,587,609,630]" pageId="54" pageNumber="44">aerophores</emphasis>
absent, rarely tuberculate at pinna bases (
<taxonomicName id="4C1E4D63FFDF9B7D63CC7F2BFBD9FDCD" box="[999,1150,609,631]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="lugubriformis">
<emphasis id="B96AEAF2FFDF9B7D63CC7F2BFBD9FDCD" box="[999,1150,609,631]" italics="true" pageId="54" pageNumber="44">G. lugubriformis</emphasis>
</taxonomicName>
);
<emphasis id="B96AEAF2FFDF9B7D64B87F2BFE2FFD2C" bold="true" pageId="54" pageNumber="44">indument abaxially and adaxially</emphasis>
of stipitate or sessile, branched hairs, these furcate (T- or Y-shaped) or stellate (
<figureCitation id="13252A65FFDF9B7D645C7FCBFB1CFD2C" box="[1143,1211,642,663]" captionStart="FIG" captionStartId="22.[97,105,1615,1635]" captionTargetBox="[144,1177,161,1569]" captionTargetId="figure-8@22.[142,1177,161,1570]" captionTargetPageId="22" captionText="FIG. 2. Morphology of the Thelypteridaceae.A. Coryphopteris kolombangarae, SITW07554 (TAIF), glandular indusia.B. Goniopteris yaucoensis, Proctor 43584 (US), stellatehairs along adaxialcosta.C.Pneumatopterisglandulifera,SITW11069 (TAIF),peg-like aerophoresat pinna bases.D.Sphaerostephanospolycarpos,SITW11070 (TAIF),marginally glandular indusia.E.Sphaerostephanos doodioides,SITW05702(TAIF),setoseindusia,spherical glands on indusia,sporangia,and laminar tissue. F. Goniopteris reptans, Wright 813 (US), sori with shriveled, setose indusia, some hairs furcate. G. Macrothelypteris polypodioides, SITW11122 (TAIF), stipe scales. H. Hoiokula pendens, Hobdy 2664 (UC), setulose sporangia. I. Plesioneuron imbricatum, SITW05370 (TAIF), exindusiate sori, sporangia with spherical glands. J. Christella dentata, SITW10525 (TAIF), shriveled,setulose indusia.K. Sphaerostephanos heterocarpos SITW11079 (TAIF),abruptly reduced proximal pinnae,adaxial surface with acicular hairs.L. Coryphopteris subbipinnata, SITW11667 (TAIF),appressed stipe base scales,these marginally setulose.All photos by Cheng-Wei Chen except B,F,and H by Susan Fawcett." figureDoi="http://doi.org/10.5281/zenodo.14111046" httpUri="https://zenodo.org/record/14111046/files/figure.png" pageId="54" pageNumber="44">
<emphasis id="B96AEAF2FFDF9B7D645C7FCBFB1CFD2C" bold="true" box="[1143,1211,642,663]" pageId="54" pageNumber="44">Fig. 2B</emphasis>
</figureCitation>
), hairs occasionally bifurcate or multifurcate, branched hairs sometimes co-occurring with unicellular or pluricellular hyaline acicular hairs typical of other genera, branched hairs most easily observed along the adaxial groove of rachis, but sometimes also present on all axes and laminar tissue adaxially and abaxially, glands generally absent on axes and lamina in most spp. (except
<taxonomicName id="4C1E4D63FFDF9B7D61597E48FE72FCAD" box="[370,469,769,790]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="redunca">
<emphasis id="B96AEAF2FFDF9B7D61597E48FE72FCAD" box="[370,469,769,790]" italics="true" pageId="54" pageNumber="44">G. redunca</emphasis>
</taxonomicName>
), scales usually absent, except along costae abaxially in a few South American species;
<emphasis id="B96AEAF2FFDF9B7D60847E68FEA1FC8D" bold="true" box="[175,262,801,822]" pageId="54" pageNumber="44">pustules</emphasis>
rarely present (e.g.,
<taxonomicName id="4C1E4D63FFDF9B7D61EB7E68FD8AFC8C" box="[448,557,801,823]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="juruensis">
<emphasis id="B96AEAF2FFDF9B7D61EB7E68FD8AFC8C" box="[448,557,801,823]" italics="true" pageId="54" pageNumber="44">G. juruensis</emphasis>
</taxonomicName>
);
<emphasis id="B96AEAF2FFDF9B7D62147E6BFDC0FC8C" bold="true" box="[575,615,802,823]" pageId="54" pageNumber="44">sori</emphasis>
round, typically medial, indusiate or exindusiate, discrete, indusia bearing hairs, these either acicular or variously branched (
<figureCitation id="13252A65FFDF9B7D629D7E0BFD5DFCEC" box="[694,762,834,855]" captionStart="FIG" captionStartId="22.[97,105,1615,1635]" captionTargetBox="[144,1177,161,1569]" captionTargetId="figure-8@22.[142,1177,161,1570]" captionTargetPageId="22" captionText="FIG. 2. Morphology of the Thelypteridaceae.A. Coryphopteris kolombangarae, SITW07554 (TAIF), glandular indusia.B. Goniopteris yaucoensis, Proctor 43584 (US), stellatehairs along adaxialcosta.C.Pneumatopterisglandulifera,SITW11069 (TAIF),peg-like aerophoresat pinna bases.D.Sphaerostephanospolycarpos,SITW11070 (TAIF),marginally glandular indusia.E.Sphaerostephanos doodioides,SITW05702(TAIF),setoseindusia,spherical glands on indusia,sporangia,and laminar tissue. F. Goniopteris reptans, Wright 813 (US), sori with shriveled, setose indusia, some hairs furcate. G. Macrothelypteris polypodioides, SITW11122 (TAIF), stipe scales. H. Hoiokula pendens, Hobdy 2664 (UC), setulose sporangia. I. Plesioneuron imbricatum, SITW05370 (TAIF), exindusiate sori, sporangia with spherical glands. J. Christella dentata, SITW10525 (TAIF), shriveled,setulose indusia.K. Sphaerostephanos heterocarpos SITW11079 (TAIF),abruptly reduced proximal pinnae,adaxial surface with acicular hairs.L. Coryphopteris subbipinnata, SITW11667 (TAIF),appressed stipe base scales,these marginally setulose.All photos by Cheng-Wei Chen except B,F,and H by Susan Fawcett." figureDoi="http://doi.org/10.5281/zenodo.14111046" httpUri="https://zenodo.org/record/14111046/files/figure.png" pageId="54" pageNumber="44">
<emphasis id="B96AEAF2FFDF9B7D629D7E0BFD5DFCEC" bold="true" box="[694,762,834,855]" pageId="54" pageNumber="44">Fig. 2F</emphasis>
</figureCitation>
);
<emphasis id="B96AEAF2FFDF9B7D63257E0BFCD4FCEC" bold="true" box="[782,883,834,855]" pageId="54" pageNumber="44">sporangia</emphasis>
glabrous or with setulae (simple or branched);
<emphasis id="B96AEAF2FFDF9B7D60E47E28FEB6FCCD" bold="true" box="[207,273,865,886]" pageId="54" pageNumber="44">spores</emphasis>
brown, consistently ornamented with reticulate crests (
<bibRefCitation id="EF8F4B11FFDF9B7D631A7E2BFC7EFCCC" author="Patel" box="[817,985,866,887]" etAl="et al." firstAuthor="Patel" journalOrPublisher="Int. J. Pl. Sci." pageId="54" pageNumber="44" pagination="1016 - 1035" part="180" refId="ref112729" refString="PATEL, N., S. FAWCETT, M. SUNDUE, &amp; J. BUDKE. 2019 a. Evolution of perine morphology in Thelypteridaceae. Int. J. Pl. Sci. 180: 1016 - 1035." title="Evolution of perine morphology in Thelypteridaceae" type="journal article" year="2019" yearSuffix="a">Patel et al. 2019a</bibRefCitation>
);
<emphasis id="B96AEAF2FFDF9B7D63C17E2BFC52FCCC" box="[1002,1013,866,887]" italics="true" pageId="54" pageNumber="44">x</emphasis>
= 36, with 25 species counted, diploids and tetraploids, and a single triploid (
<taxonomicName id="4C1E4D63FFDF9B7D62B97EC8FCAEFC2D" box="[658,777,897,918]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="obliterata">
<emphasis id="B96AEAF2FFDF9B7D62B97EC8FCAEFC2D" box="[658,777,897,918]" italics="true" pageId="54" pageNumber="44">G. obliterata</emphasis>
</taxonomicName>
) known. No intergeneric hybrids have been reported, but intrageneric hybrids are quite common, especially among the species of the calciphilic Antillean radiation (
<bibRefCitation id="EF8F4B11FFDF9B7D604C7E8BFF44FC6C" author="Smith" box="[103,227,962,983]" firstAuthor="Smith" journalOrPublisher="Biogeogr." pageId="54" pageNumber="44" pagination="255 - 264" part="20" refId="ref114507" refString="SMITH, A. R. 1993 b. Phytogeographic principles and their use in understanding fern relationships. J. Biogeogr. 20: 255 - 264." title="Phytogeographic principles and their use in understanding fern relationships. J" type="journal article" year="1993" yearSuffix="b">Smith 1993b</bibRefCitation>
), where more than a dozen have been reported in
<collectingCountry id="F3097670FFDF9B7D62927E8BFD4BFC6C" box="[697,748,962,983]" name="Cuba" pageId="54" pageNumber="44">Cuba</collectingCountry>
alone (
<bibRefCitation id="EF8F4B11FFDF9B7D63047E8BFC12FC6C" author="Sanchez" box="[815,949,962,983]" firstAuthor="Sanchez" journalOrPublisher="Brittonia" pageId="54" pageNumber="44" pagination="482 - 503" part="69" refId="ref113773" refString="SANCHEZ, C. 2017. Lista de los helechos y licofitos de Cuba. Brittonia 69: 482 - 503." title="Lista de los helechos y licofitos de Cuba" type="journal article" year="2017">Sánchez 2017</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDF9B7D60BA7EABFD7AFB6C" pageId="54" pageNumber="44" type="diagnosis">
<paragraph id="8BA136E0FFDF9B7D60BA7EABFD7AFB6C" blockId="54.[97,1226,161,1783]" pageId="54" pageNumber="44">
<emphasis id="B96AEAF2FFDF9B7D60BA7EABFF55FC4C" box="[145,242,994,1015]" italics="true" pageId="54" pageNumber="44">Diagnosis.</emphasis>
—The nearly invariable presence of stellate or branched hairs somewhere on the plant, especially along the axes and on stipe base and rhizome scales, is the most diagnostic feature of the genus, readily distinguishing it from all other genera in the family. The paleotropical
<taxonomicName id="4C1E4D63FFDF9B7D620A7968FD3CFB8D" box="[545,667,1057,1078]" class="Polypodiopsida" family="Thelypteridaceae" genus="Ampelopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDF9B7D620A7968FD3CFB8D" box="[545,667,1057,1078]" italics="true" pageId="54" pageNumber="44">Ampelopteris</emphasis>
</taxonomicName>
is similar in bearing proliferous buds and stellate hairs (see illustrations in
<bibRefCitation id="EF8F4B11FFDF9B7D60C4790BFE03FBEC" author="Holttum" box="[239,420,1090,1111]" etAl="et al." firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="54" pageNumber="44" pagination="195 - 215" part="43" refId="ref110127" refString="HOLTTUM, R. E., U. SEN, &amp; D. MITTRA. 1970. Studies in the family Thelypteridaceae. II. A comparative study of the type-species of Thelypteris Schmidel, Cyclosorus Link, and Ampelopteris Kunze. Blumea 43: 195 - 215." title="Studies in the family Thelypteridaceae. II. A comparative study of the type-species of Thelypteris Schmidel, Cyclosorus Link, and Ampelopteris Kunze" type="journal article" year="1970">Holttum et al. 1970</bibRefCitation>
), but the hairs are easily abraded, and less conspicuous than in
<emphasis id="B96AEAF2FFDF9B7D63C47908FB6FFBEC" box="[1007,1224,1089,1111]" italics="true" pageId="54" pageNumber="44">
<taxonomicName id="4C1E4D63FFDF9B7D63C47908FBF9FBED" box="[1007,1118,1089,1110]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">Goniopteris</taxonomicName>
.
<taxonomicName id="4C1E4D63FFDF9B7D644F790BFB6FFBEC" box="[1124,1224,1090,1111]" class="Polypodiopsida" family="Thelypteridaceae" genus="Meniscium" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">Meniscium</taxonomicName>
</emphasis>
differs in having exindusiate sori that are frequently coalescent, many more pairs of anastomosing veins, usually entire or only serrate pinnae, and often longer-creeping rhizomes.
<taxonomicName id="4C1E4D63FFDF9B7D62B479C8FCAFFB2D" box="[671,776,1153,1174]" class="Polypodiopsida" family="Thelypteridaceae" genus="Steiropteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDF9B7D62B479C8FCAFFB2D" box="[671,776,1153,1174]" italics="true" pageId="54" pageNumber="44">Steiropteris</emphasis>
</taxonomicName>
often differs in the presence of well-developed aerophores at the pinna (and sometimes costule) bases, and in most cases, a distinct cartilaginous, non-vascularized keel at the bases of the sinuses; it also lacks stellate or furcate hairs.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDF9B7D60BA79A8FF73FA4C" pageId="54" pageNumber="44" type="biology_ecology">
<paragraph id="8BA136E0FFDF9B7D60BA79A8FF73FA4C" blockId="54.[97,1226,161,1783]" pageId="54" pageNumber="44">
<emphasis id="B96AEAF2FFDF9B7D60BA79A8FE22FB4D" box="[145,389,1249,1270]" italics="true" pageId="54" pageNumber="44">Biogeography and ecology.</emphasis>
—Numbering 138 species,
<taxonomicName id="4C1E4D63FFDF9B7D62A779A8FD5FFB4D" box="[652,760,1249,1270]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDF9B7D62A779A8FD5FFB4D" box="[652,760,1249,1270]" italics="true" pageId="54" pageNumber="44">Goniopteris</emphasis>
</taxonomicName>
is strictly neotropical, ranging north to Florida,
<collectingCountry id="F3097670FFDF9B7D604A784BFF0DFAAC" box="[97,170,1282,1303]" name="Mexico" pageId="54" pageNumber="44">Mexico</collectingCountry>
, through Mesoamerica and the Antilles, throughout Amazonia, and the foothills of the Andes, south to
<collectingCountry id="F3097670FFDF9B7D64AD784BFB6EFAAC" box="[1158,1225,1282,1303]" name="Bolivia" pageId="54" pageNumber="44">Bolivia</collectingCountry>
and
<taxonomicName id="4C1E4D63FFDF9B7D60A2786BFF4CFA8C" box="[137,235,1314,1335]" class="Polypodiopsida" family="Thelypteridaceae" genus="Amauropelta" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="argentina">Argentina</taxonomicName>
, and east through
<collectingCountry id="F3097670FFDF9B7D61B2786BFE74FA8C" box="[409,467,1314,1335]" name="Brazil" pageId="54" pageNumber="44">Brazil</collectingCountry>
, the Guianas and
<collectingCountry id="F3097670FFDF9B7D6250786BFD47FA8C" box="[635,736,1314,1335]" name="Venezuela" pageId="54" pageNumber="44">Venezuela</collectingCountry>
. They are predominantly plants of lower elevations, ranging to
<quantity id="4CE69B05FFDF9B7D60E0780BFEB3FAED" box="[203,276,1345,1367]" metricMagnitude="3" metricUnit="m" metricValue="1.85" pageId="54" pageNumber="44" unit="m" value="1850.0">1850 m</quantity>
in the Andes (
<bibRefCitation id="EF8F4B11FFDF9B7D618A780BFDB5FAEC" author="Smith" box="[417,530,1346,1367]" firstAuthor="Smith" journalOrPublisher="Fieldiana, Bot." pageId="54" pageNumber="44" pagination="1 - 80" part="29" refId="ref114420" refString="SMITH, A. R. 1992. Pteridophyta of Peru, Part. III, 16. Thelypteridaceae. Fieldiana, Bot. 29: 1 - 80." title="Pteridophyta of Peru, Part. III, 16. Thelypteridaceae" type="journal article" year="1992">Smith 1992</bibRefCitation>
), and to
<quantity id="4CE69B05FFDF9B7D624E780BFD09FAED" box="[613,686,1345,1367]" metricMagnitude="3" metricUnit="m" metricValue="1.7" pageId="54" pageNumber="44" unit="m" value="1700.0">1700 m</quantity>
in the Antilles. They are typically plants of shady forest understories, but a diverse clade of calciphiles, numbering ~30 species, has radiated in the karst regions of the Greater Antilles (
<bibRefCitation id="EF8F4B11FFDF9B7D609378CBFE9AFA2C" author="Fawcett" box="[184,317,1410,1431]" firstAuthor="Fawcett" journalOrPublisher="Amer. Fern. J." pageId="54" pageNumber="44" pagination="183 - 192" part="110" refId="ref108269" refString="FAWCETT, S. 2020. A new species of Goniopteris (Thelypteridaceae) from Hispaniola. Amer. Fern. J. 110: 183 - 192." title="A new species of Goniopteris (Thelypteridaceae) from Hispaniola" type="journal article" year="2020">Fawcett 2020</bibRefCitation>
). Two serpentine specialists,
<taxonomicName id="4C1E4D63FFDF9B7D627E78C8FD0BFA2C" box="[597,684,1409,1431]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="crypta">
<emphasis id="B96AEAF2FFDF9B7D627E78C8FD0BFA2C" box="[597,684,1409,1431]" italics="true" pageId="54" pageNumber="44">G. crypta</emphasis>
</taxonomicName>
and
<emphasis id="B96AEAF2FFDF9B7D62F078C8FCF7FA2D" box="[731,848,1409,1430]" italics="true" pageId="54" pageNumber="44">
<collectionCode id="ED0FAE25FFDF9B7D62F078C8FD49FA2D" box="[731,750,1409,1430]" country="Switzerland" lsid="urn:lsid:biocol.org:col:15706" name="Conservatoire et Jardin botaniques de la Ville de Genève" pageId="54" pageNumber="44" type="Herbarium">G</collectionCode>
. brittoniae
</emphasis>
, have likely independently arisen from calciphilic ancestors on
<collectingCountry id="F3097670FFDF9B7D616D78EBFEDCFA0C" box="[326,379,1442,1463]" name="Cuba" pageId="54" pageNumber="44">Cuba</collectingCountry>
, and
<collectingCountry id="F3097670FFDF9B7D618678E8FD87FA0C" box="[429,544,1441,1463]" name="Puerto Rico" pageId="54" pageNumber="44">Puerto Rico</collectingCountry>
, respectively. As with many members of the family, they respond favorably to disturbance, may become weedy, and may be especially abundant on open roadcuts and in coffee and cacao plantations.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFDF9B7C60BA7B4BFCE9FE0C" lastPageId="55" lastPageNumber="45" pageId="54" pageNumber="44" type="discussion">
<paragraph id="8BA136E0FFDF9B7C60BA7B4BFD0EFF6C" blockId="54.[97,1226,161,1783]" lastBlockId="55.[121,1251,162,503]" lastPageId="55" lastPageNumber="45" pageId="54" pageNumber="44">
<emphasis id="B96AEAF2FFDF9B7D60BA7B4BFE47F9AD" box="[145,480,1537,1559]" italics="true" pageId="54" pageNumber="44">Taxonomic and phylogenetic studies.</emphasis>
—Historically treated as a subgenus of
<taxonomicName id="4C1E4D63FFDF9B7D637D7B4BFC1EF9AC" box="[854,953,1538,1559]" class="Polypodiopsida" family="Dryopteridaceae" genus="Dryopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDF9B7D637D7B4BFC1EF9AC" box="[854,953,1538,1559]" italics="true" pageId="54" pageNumber="44">Dryopteris</emphasis>
</taxonomicName>
, the circumscription of species of
<taxonomicName id="4C1E4D63FFDF9B7D608B7B68FEABF98D" box="[160,268,1569,1590]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDF9B7D608B7B68FEABF98D" box="[160,268,1569,1590]" italics="true" pageId="54" pageNumber="44">Goniopteris</emphasis>
</taxonomicName>
has remained essentially unchanged for more than a century (
<bibRefCitation id="EF8F4B11FFDF9B7D637D7B6BFBA6F98C" author="Christensen" box="[854,1025,1570,1591]" firstAuthor="Christensen" journalOrPublisher="Selsk. Skr., Naturvidensk. Math. Afd." pageId="54" pageNumber="44" pagination="55 - 282" part="10" refId="ref107711" refString="CHRISTENSEN, C. 1913. A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 55 - 282." title="A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk" type="journal article" year="1913">Christensen 1913</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFDF9B7D64237B6BFB9DF98C" author="Christensen" box="[1032,1082,1570,1591]" firstAuthor="Christensen" journalOrPublisher="Selsk. Skr., Naturvidensk Math. Afd." pageId="54" pageNumber="44" pagination="3 - 132" part="6" refId="ref107760" refString="CHRISTENSEN, C. 1920. A monograph of the genus Dryopteris, part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk Math. Afd., ser. 8, 6: 3 - 132." title="A monograph of the genus Dryopteris, part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk" type="journal article" year="1920">1920</bibRefCitation>
). Christensens treatment was the first to rely on close observation of microscopic features, and he united the species of
<taxonomicName id="4C1E4D63FFDF9B7D64777B08FB6EF9ED" box="[1116,1225,1601,1622]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDF9B7D64777B08FB6EF9ED" box="[1116,1225,1601,1622]" italics="true" pageId="54" pageNumber="44">Goniopteris</emphasis>
</taxonomicName>
largely on the basis of their striking stellate or branched hairs. Although the hairs have been secondarily lost on rare occasions (e.g.,
<taxonomicName id="4C1E4D63FFDF9B7D60DE7BC8FEF9F92C" box="[245,350,1665,1687]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="species" species="macrotis">
<emphasis id="B96AEAF2FFDF9B7D60DE7BC8FEF9F92C" box="[245,350,1665,1687]" italics="true" pageId="54" pageNumber="44">G. macrotis</emphasis>
</taxonomicName>
), the few taxa without them were still included by
<bibRefCitation id="EF8F4B11FFDF9B7D63637BCBFBA7F92C" author="Christensen" box="[840,1024,1666,1687]" firstAuthor="Christensen" journalOrPublisher="Selsk. Skr., Naturvidensk. Math. Afd." pageId="54" pageNumber="44" pagination="55 - 282" part="10" refId="ref107711" refString="CHRISTENSEN, C. 1913. A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 55 - 282." title="A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk" type="journal article" year="1913">Christensen (1913)</bibRefCitation>
on the basis of other morphological similarities; their placement has now been supported by molecular data (Fawcett et al. in press).
<bibRefCitation id="EF8F4B11FFDF9B7D604A7B8BFEB2F96C" author="Christensen" box="[97,277,1730,1751]" firstAuthor="Christensen" journalOrPublisher="Selsk. Skr., Naturvidensk. Math. Afd." pageId="54" pageNumber="44" pagination="55 - 282" part="10" refId="ref107711" refString="CHRISTENSEN, C. 1913. A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 55 - 282." title="A monograph of the genus Dryopteris, part I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk" type="journal article" year="1913">Christensen (1913</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFDF9B7D61357B8BFEE9F96C" author="Christensen" box="[286,334,1730,1751]" firstAuthor="Christensen" journalOrPublisher="Selsk. Skr., Naturvidensk Math. Afd." pageId="54" pageNumber="44" pagination="3 - 132" part="6" refId="ref107760" refString="CHRISTENSEN, C. 1920. A monograph of the genus Dryopteris, part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk Math. Afd., ser. 8, 6: 3 - 132." title="A monograph of the genus Dryopteris, part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk" type="journal article" year="1920">1920</bibRefCitation>
) divided
<taxonomicName id="4C1E4D63FFDF9B7D61807B8BFD1DF96D" box="[427,698,1729,1751]" class="Polypodiopsida" family="Dryopteridaceae" genus="Dryopteris" kingdom="Plantae" order="Polypodiales" pageId="54" pageNumber="44" phylum="Tracheophyta" rank="subGenus" subGenus="Goniopteris">
<emphasis id="B96AEAF2FFDF9B7D61807B8BFDA8F96C" box="[427,527,1730,1751]" italics="true" pageId="54" pageNumber="44">Dryopteris</emphasis>
subg.
<emphasis id="B96AEAF2FFDF9B7D62657B88FD1DF96D" box="[590,698,1729,1750]" italics="true" pageId="54" pageNumber="44">Goniopteris</emphasis>
</taxonomicName>
into two sections—
<emphasis id="B96AEAF2FFDF9B7D63547B88FBA6F96D" box="[895,1025,1729,1750]" italics="true" pageId="54" pageNumber="44">Asterochlaena</emphasis>
and
<emphasis id="B96AEAF2FFDF9B7D64197B8BFB16F96C" box="[1074,1201,1730,1751]" italics="true" pageId="54" pageNumber="44">Eugoniopteris</emphasis>
— recognized by apices gradually reduced, or by a conform terminal pinna, respectively. Phylogenetic analyses have shown these subdivisions to be non-monophyletic, and therefore artificial. Necessary combinations have been made for nearly all known members of the genus (
<bibRefCitation id="EF8F4B11FFDE9B7C61D17D8BFD38FF6C" author="Salino" box="[506,671,194,215]" etAl="et al." firstAuthor="Salino" journalOrPublisher="PhytoKeys" pageId="55" pageNumber="45" pagination="11 - 50" part="57" refId="ref113700" refString="SALINO, A., T. E. ALMEIDA, &amp; A. R. SMITH. 2015. New combinations in neotropical Thelypteridaceae. PhytoKeys 57: 11 - 50." title="New combinations in neotropical Thelypteridaceae" type="journal article" year="2015">Salino et al. 2015</bibRefCitation>
).
</paragraph>
<paragraph id="8BA136E0FFDE9B7C60827DABFC2EFE2C" blockId="55.[121,1251,162,503]" pageId="55" pageNumber="45">
Molecular phylogenetic data support the monophyly of
<taxonomicName id="4C1E4D63FFDE9B7C62EE7DABFC96FF4C" box="[709,817,226,247]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="55" pageNumber="45" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDE9B7C62EE7DABFC96FF4C" box="[709,817,226,247]" italics="true" pageId="55" pageNumber="45">Goniopteris</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF8F4B11FFDE9B7C63157DABFC5AFF4C" author="ALMEIDA, T. E &amp; S. HENNEQUIN &amp; H. SCHNEIDER &amp; A. R. SMITH &amp; J. A. N. BATISTA &amp; A. J. RAMALHO &amp; K. PROITE &amp; K. &amp; A. SALINO" box="[830,1021,226,247]" journalOrPublisher="Molec. Phylogen. Evol." pageId="55" pageNumber="45" pagination="688 - 700" part="94" refId="ref106225" refString="ALMEIDA, T. E., S. HENNEQUIN, H. SCHNEIDER, A. R. SMITH, J. A. N. BATISTA, A. J. RAMALHO, K. PROITE, &amp; K. &amp; A. SALINO. 2016. Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera. Molec. Phylogen. Evol. 94: 688 - 700." title="Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera" type="journal article" year="2016">Almeida et al. 2016</bibRefCitation>
;
<bibRefCitation id="EF8F4B11FFDE9B7C642D7DABFB0BFF4C" author="Patel" box="[1030,1196,226,247]" etAl="et al." firstAuthor="Patel" journalOrPublisher="Int. J. Pl. Sci." pageId="55" pageNumber="45" pagination="1016 - 1035" part="180" refId="ref112729" refString="PATEL, N., S. FAWCETT, M. SUNDUE, &amp; J. BUDKE. 2019 a. Evolution of perine morphology in Thelypteridaceae. Int. J. Pl. Sci. 180: 1016 - 1035." title="Evolution of perine morphology in Thelypteridaceae" type="journal article" year="2019" yearSuffix="a">Patel et al. 2019a</bibRefCitation>
), and show strong biogeographical signal, with distinct radiations in the Antilles, the Andes, and
<collectingCountry id="F3097670FFDE9B7C63D67C4BFB91FEAC" box="[1021,1078,258,279]" name="Brazil" pageId="55" pageNumber="45">Brazil</collectingCountry>
(Fawcett et al. in press). The genus is in an early-diverging clade of cyclosoroids, and sister to a clade that includes the neotropical genus
<taxonomicName id="4C1E4D63FFDE9B7C60527C0BFF7AFEEC" box="[121,221,322,343]" class="Polypodiopsida" family="Thelypteridaceae" genus="Meniscium" kingdom="Plantae" order="Polypodiales" pageId="55" pageNumber="45" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDE9B7C60527C0BFF7AFEEC" box="[121,221,322,343]" italics="true" pageId="55" pageNumber="45">Meniscium</emphasis>
</taxonomicName>
, the pantropical
<taxonomicName id="4C1E4D63FFDE9B7C61507C08FE79FEED" box="[379,478,321,342]" class="Polypodiopsida" family="Thelypteridaceae" genus="Cyclosorus" kingdom="Plantae" order="Polypodiales" pageId="55" pageNumber="45" phylum="Tracheophyta" rank="genus" sensu="stricto">
<emphasis id="B96AEAF2FFDE9B7C61507C08FE79FEED" box="[379,478,321,342]" italics="true" pageId="55" pageNumber="45">Cyclosorus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2595789FFDE9B7C61C97C0BFE5AFEEC" box="[482,509,322,343]" pageId="55" pageNumber="45" sensu="stricto">
<emphasis id="B96AEAF2FFDE9B7C61C97C0BFE50FEEC" box="[482,503,322,343]" italics="true" pageId="55" pageNumber="45">s.s</emphasis>
.
</taxonomicNameLabel>
, the African and Southeast Asian
<taxonomicName id="4C1E4D63FFDE9B7C63107C08FC13FEED" box="[827,948,321,342]" class="Polypodiopsida" family="Thelypteridaceae" genus="Ampelopteris" kingdom="Plantae" order="Polypodiales" pageId="55" pageNumber="45" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDE9B7C63107C08FC13FEED" box="[827,948,321,342]" italics="true" pageId="55" pageNumber="45">Ampelopteris</emphasis>
</taxonomicName>
, and the Malesian
<taxonomicName id="4C1E4D63FFDE9B7C64487C08FB46FEED" baseAuthorityName="Holttum" baseAuthorityYear="1971" box="[1123,1249,321,342]" class="Polypodiopsida" family="Thelypteridaceae" genus="Mesophlebion" kingdom="Plantae" order="Polypodiales" pageId="55" pageNumber="45" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDE9B7C64487C08FB46FEED" box="[1123,1249,321,342]" italics="true" pageId="55" pageNumber="45">Mesophlebion</emphasis>
</taxonomicName>
(Fawcett et al. in press). In contrast, in the less densely-sampled, mostly plastid dataset of
<bibRefCitation id="EF8F4B11FFDE9B7C63EA7C2BFBCAFECC" author="Patel" box="[961,1133,354,375]" etAl="et al." firstAuthor="Patel" journalOrPublisher="Int. J. Pl. Sci." pageId="55" pageNumber="45" pagination="1016 - 1035" part="180" refId="ref112729" refString="PATEL, N., S. FAWCETT, M. SUNDUE, &amp; J. BUDKE. 2019 a. Evolution of perine morphology in Thelypteridaceae. Int. J. Pl. Sci. 180: 1016 - 1035." title="Evolution of perine morphology in Thelypteridaceae" type="journal article" year="2019" yearSuffix="a">Patel et al. (2019a)</bibRefCitation>
,
<taxonomicName id="4C1E4D63FFDE9B7C645F7C2BFB47FECC" box="[1140,1248,354,375]" class="Polypodiopsida" family="Thelypteridaceae" genus="Goniopteris" kingdom="Plantae" order="Polypodiales" pageId="55" pageNumber="45" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFDE9B7C645F7C2BFB47FECC" box="[1140,1248,354,375]" italics="true" pageId="55" pageNumber="45">Goniopteris</emphasis>
</taxonomicName>
is inferred be sister to the aforementioned genera, plus the remaining cyclosoroids.
</paragraph>
<paragraph id="8BA136E0FFDE9B7C608C7CEBFCE9FE0C" blockId="55.[121,1251,162,503]" box="[167,846,418,439]" pageId="55" pageNumber="45">
Most necessary combinations were made by
<bibRefCitation id="EF8F4B11FFDE9B7C62647CEBFCA6FE0C" author="Salino" box="[591,769,418,439]" etAl="et al." firstAuthor="Salino" journalOrPublisher="PhytoKeys" pageId="55" pageNumber="45" pagination="11 - 50" part="57" refId="ref113700" refString="SALINO, A., T. E. ALMEIDA, &amp; A. R. SMITH. 2015. New combinations in neotropical Thelypteridaceae. PhytoKeys 57: 11 - 50." title="New combinations in neotropical Thelypteridaceae" type="journal article" year="2015">Salino et al. (2015)</bibRefCitation>
, except:
</paragraph>
</subSubSection>
</treatment>
</document>