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<document id="5DA891D43EBA52A070E7373F95C48155" 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="03B787F6FFBC9B1D62257DEAFBF8FCAC" docLanguage="en" docName="2021_ThelypteridaceaeFerns_OACopy3.pdf.imf" docOrigin="Fort Worth, Texas, USA: BRIT Press" docTitle="Pseudophegopteris Ching" docType="treatment" docVersion="4" lastPageNumber="76" masterDocId="FF8EFF8EFFE99B4B602B7D49FFA7FFBB" masterDocTitle="A Generic Classification of the Thelypteridaceae" pageNumber="75" updateTime="1732279345664" updateUser="felipe" zenodo-license-document="CC-BY-NC-SA-4.0">
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<mods:titleInfo id="37E314A3CD9A22E01147E7B18D6D2928">
<mods:title id="463804B3699B8A4BCD94EE3251AC149D">A Generic Classification of the Thelypteridaceae</mods:title>
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<mods:namePart id="FB9DFBFD6521B35ACF6112C7A0F0F397">Fawcett, Susan</mods:namePart>
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<mods:namePart id="7D95F94418BBBDD667680EEA66BD838C">Smith, Alan R.</mods:namePart>
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<treatment id="03B787F6FFBC9B1D62257DEAFBF8FCAC" LSID="urn:lsid:plazi:treatment:03B787F6FFBC9B1D62257DEAFBF8FCAC" httpUri="http://treatment.plazi.org/id/03B787F6FFBC9B1D62257DEAFBF8FCAC" lastPageId="86" lastPageNumber="76" pageId="85" pageNumber="75">
<subSubSection id="C304656BFFBC9B1E62257DEAFCECFF06" box="[526,843,163,189]" pageId="85" pageNumber="75" type="nomenclature">
<paragraph id="8BA136E0FFBC9B1E62257DEAFCECFF06" blockId="85.[121,1248,163,251]" box="[526,843,163,189]" pageId="85" pageNumber="75">
<heading id="D0E9818CFFBC9B1E62257DEAFCECFF06" box="[526,843,163,189]" level="2" pageId="85" pageNumber="75">
<taxonomicName id="4C1E4D63FFBC9B1E62257DEAFCECFF06" ID-CoL="8R5Z4" authority="Ching" authorityName="Ching" box="[526,843,163,189]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">PSEUDOPHEGOPTERIS</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<treatmentCitationGroup id="AB0E11CEFFBC9B1E60527D82FEFAFF40" pageId="85" pageNumber="75">
<subSubSection id="C304656BFFBC9B1E60527D82FD19FF5B" box="[121,702,203,224]" pageId="85" pageNumber="75" type="reference_group">
<paragraph id="8BA136E0FFBC9B1E60527D82FD19FF5B" blockId="85.[121,1248,163,251]" box="[121,702,203,224]" pageId="85" pageNumber="75">
<treatmentCitation id="0ABF10F1FFBC9B1E60527D82FD1DFF5B" author="Pseudophegopteris Ching" box="[121,698,203,224]" journal="Acta Phytotax. Sin." page="313" pageId="85" pageNumber="75" score="3.863100345872848" volume="8" year="1963">
<taxonomicName id="4C1E4D63FFBC9B1E60527D82FED8FF5B" ID-CoL="8R5Z4" authority="Ching" authorityName="Ching" box="[121,383,203,224]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E60527D82FE9DFF5B" bold="true" box="[121,314,203,224]" pageId="85" pageNumber="75">Pseudophegopteris</emphasis>
Ching
</taxonomicName>
,
<bibCitation id="5E343E8CFFBC9B1E61A27D82FD1DFF5B" box="[393,698,203,224]" journal="Acta Phytotax. Sin." page="313" pageId="85" pageNumber="75" score="3.863100345872848" volume="8" year="1963">Acta Phytotax. Sin. 8:313. 1963</bibCitation>
</treatmentCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBC9B1E62967D87FEFAFF40" pageId="85" pageNumber="75" type="type_taxon">
<paragraph id="8BA136E0FFBC9B1E62967D87FEFAFF40" blockId="85.[121,1248,163,251]" pageId="85" pageNumber="75">
<typeStatus id="54A58842FFBC9B1E62E47D87FD48FF65" box="[719,751,206,222]" pageId="85" pageNumber="75">
<collectionCode id="ED0FAE25FFBC9B1E62E47D87FD48FF65" box="[719,751,206,222]" pageId="85" pageNumber="75">TYPE</collectionCode>
</typeStatus>
:
<taxonomicName id="4C1E4D63FFBC9B1E62D27D87FBC9FF5B" ID-CoL="8RBHX" authority="(Kunze) Ching" authorityName="Ching" authorityYear="1963" baseAuthorityName="Kunze" box="[761,1134,206,224]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="pyrrhorhachis">
<emphasis id="B96AEAF2FFBC9B1E62D27D87FC52FF64" box="[761,1013,206,223]" italics="true" pageId="85" pageNumber="75">Pseudophegopteris pyrrhorhachis</emphasis>
(Kunze) Ching
</taxonomicName>
[=
<taxonomicName id="4C1E4D63FFBC9B1E64AC7D87FEFEFF40" ID-CoL="BCCKW" authority="Kunze" authorityName="Kunze" class="Polypodiopsida" family="Thelypteridaceae" genus="Polypodium" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="pyrrhorhachis">
<emphasis id="B96AEAF2FFBC9B1E64AC7D87FE84FF41" italics="true" pageId="85" pageNumber="75">Polypodium pyrrhorhachis</emphasis>
Kunze
</taxonomicName>
]
</paragraph>
</subSubSection>
</treatmentCitationGroup>
<subSubSection id="C304656BFFBC9B1E60B27C46FDFDFE80" pageId="85" pageNumber="75" type="reference_group">
<paragraph id="8BA136E0FFBC9B1E60B27C46FDE1FEA5" blockId="85.[153,602,270,315]" box="[153,582,270,288]" pageId="85" pageNumber="75">
<emphasis id="B96AEAF2FFBC9B1E60B27C46FF45FEA4" box="[153,226,271,287]" italics="true" pageId="85" pageNumber="75">Toppingia</emphasis>
<collectionCode id="ED0FAE25FFBC9B1E60CE7C46FF53FE9B" box="[229,244,271,288]" country="Norway" lsid="urn:lsid:biocol.org:col:13093" name="Botanical Museum - University of Oslo" pageId="85" pageNumber="75" type="Herbarium">O</collectionCode>
. Deg.,
<collectionCode id="ED0FAE25FFBC9B1E610F7C47FE8BFEA5" box="[292,300,270,286]" country="Romania" lsid="urn:lsid:biocol.org:col:14415" name="&amp;quot;Alexandru Ioan Cuza&amp;quot; University" pageId="85" pageNumber="75" type="Herbarium">I</collectionCode>
. Deg. &amp;
<collectionCode id="ED0FAE25FFBC9B1E61477C46FEDDFE9B" box="[364,378,271,288]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="85" pageNumber="75" type="Herbarium">A</collectionCode>
.
<collectionCode id="ED0FAE25FFBC9B1E61567C46FE2DFE9B" box="[381,394,271,288]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="85" pageNumber="75">R</collectionCode>
.Sm. ex
<collectionCode id="ED0FAE25FFBC9B1E61EE7C46FE73FE9B" box="[453,468,271,288]" country="Norway" lsid="urn:lsid:biocol.org:col:13093" name="Botanical Museum - University of Oslo" pageId="85" pageNumber="75" type="Herbarium">O</collectionCode>
. Deg.&amp;
<collectionCode id="ED0FAE25FFBC9B1E623F7C47FDBBFEA5" box="[532,540,270,286]" country="Romania" lsid="urn:lsid:biocol.org:col:14415" name="&amp;quot;Alexandru Ioan Cuza&amp;quot; University" pageId="85" pageNumber="75" type="Herbarium">I</collectionCode>
. Deg.
</paragraph>
<paragraph id="8BA136E0FFBC9B1E60B27C60FDFDFE80" blockId="85.[153,602,270,315]" box="[153,602,297,315]" pageId="85" pageNumber="75">
For additional generic synonymy, see
<bibRefCitation id="EF8F4B11FFBC9B1E619F7C63FD83FE80" author="Holttum" box="[436,548,298,315]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="85" pageNumber="75" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">Holttum (1969</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBC9B1E62007C63FDF6FE80" author="Holttum" box="[555,593,298,315]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="85" pageNumber="75" pagination="17 - 52" part="19" refId="ref109463" refString="HOLTTUM, R. E. 1971. Studies in the family Thelypteridaceae III. A new system of genera in the Old World. Blumea 19: 17 - 52." title="Studies in the family Thelypteridaceae III. A new system of genera in the Old World" type="journal article" year="1971">1971</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBC9B1E60527C19FBA7FEDE" box="[121,1024,336,358]" pageId="85" pageNumber="75" type="etymology">
<paragraph id="8BA136E0FFBC9B1E60527C19FBA7FEDE" blockId="85.[121,1250,336,1797]" box="[121,1024,336,358]" pageId="85" pageNumber="75">
<emphasis id="B96AEAF2FFBC9B1E60527C19FF46FEDE" box="[121,225,336,357]" italics="true" pageId="85" pageNumber="75">Etymology.</emphasis>
—Gr.
<emphasis id="B96AEAF2FFBC9B1E61307C19FEC6FEDE" box="[283,353,336,357]" italics="true" pageId="85" pageNumber="75">pseudo,</emphasis>
false +
<emphasis id="B96AEAF2FFBC9B1E618E7C19FE45FEDE" box="[421,482,336,357]" italics="true" pageId="85" pageNumber="75">phegos</emphasis>
, beech +
<emphasis id="B96AEAF2FFBC9B1E62117C19FDC8FEDE" box="[570,623,336,357]" italics="true" pageId="85" pageNumber="75">pteris</emphasis>
= fern; a distinctive relative of
<taxonomicName id="4C1E4D63FFBC9B1E63BB7C19FC5DFEDE" box="[912,1018,336,357]" class="Polypodiopsida" family="Thelypteridaceae" genus="Phegopteris" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E63BB7C19FC5DFEDE" box="[912,1018,336,357]" italics="true" pageId="85" pageNumber="75">Phegopteris</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBC9B1E60817C38FCF0FB3D" pageId="85" pageNumber="75" type="description">
<paragraph id="8BA136E0FFBC9B1E60817C38FCF0FB3D" blockId="85.[121,1250,336,1797]" pageId="85" pageNumber="75">
Plants terrestrial, fronds of determinate growth (except in
<taxonomicName id="4C1E4D63FFBC9B1E62D97C39FC34FE3E" box="[754,915,368,389]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="keraudreniana">
<emphasis id="B96AEAF2FFBC9B1E62D97C39FC34FE3E" box="[754,915,368,389]" italics="true" pageId="85" pageNumber="75">P. keraudreniana</emphasis>
</taxonomicName>
, where growth is indeterminate), mostly medium-sized to very large, 50300+ cm tall;
<emphasis id="B96AEAF2FFBC9B1E62457CD8FD6BFE1D" bold="true" box="[622,716,401,422]" pageId="85" pageNumber="75">rhizomes</emphasis>
short- to long-creeping, or ascending, or erect, with thin scales, these brown to tan, lanceolate, with scattered hairs along margins and sometimes sparingly on surface;
<emphasis id="B96AEAF2FFBC9B1E64B57CF8FB46FE7D" bold="true" box="[1182,1249,433,454]" pageId="85" pageNumber="75">fronds</emphasis>
clustered, or remote, monomorphic;
<emphasis id="B96AEAF2FFBC9B1E61CB7C98FDBBFE5D" bold="true" box="[480,540,465,486]" pageId="85" pageNumber="75">stipes</emphasis>
<quantity id="4CE69B05FFBC9B1E62097C98FD21FE5E" box="[546,646,464,486]" metricMagnitude="-1" metricUnit="m" metricValue="4.5" metricValueMax="6.0" metricValueMin="3.0" pageId="85" pageNumber="75" unit="cm" value="45.0" valueMax="60.0" valueMin="30.0">3060 cm</quantity>
long, not grooved adaxially, stramineous to red-brown, often glossy;
<emphasis id="B96AEAF2FFBC9B1E60947CB8FE95FDBE" bold="true" box="[191,306,496,518]" pageId="85" pageNumber="75">stipe scales</emphasis>
thin, lanceolate to ovate, light brown,
<quantity id="4CE69B05FFBC9B1E62897CB8FD54FDBE" box="[674,755,496,518]" metricMagnitude="-3" metricUnit="m" metricValue="4.0" metricValueMax="6.0" metricValueMin="2.0" pageId="85" pageNumber="75" unit="mm" value="4.0" valueMax="6.0" valueMin="2.0">26 mm</quantity>
long, typically with superficial hairs;
<emphasis id="B96AEAF2FFBC9B1E64777CB9FB39FDBE" bold="true" box="[1116,1182,496,517]" pageId="85" pageNumber="75">blades</emphasis>
herbaceous to chartaceous, lacking buds or proliferations, pinnate-pinnatifid to bipinnate-pinnatifid or slightly more divided, with blade apex gradually tapering and pinnatifid; proximal pinnae of well-developed fronds not reduced or with 12 lowermost pairs somewhat reduced (sometimes to less than half the length of longest pinnae), but blades lacking greatly reduced, glanduliform pinnae; rachises adaxially not grooved, bearing simple and sometimes forked acicular hairs, some spp. glabrescent, typically lacking scales at maturity (except
<taxonomicName id="4C1E4D63FFBC9B1E62D67FD9FCF7FD1E" box="[765,848,656,677]" class="Polypodiopsida" family="Thelypteridaceae" genus="Phegopteris" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="dianae">
<emphasis id="B96AEAF2FFBC9B1E62D67FD9FCF7FD1E" box="[765,848,656,677]" italics="true" pageId="85" pageNumber="75">P. dianae</emphasis>
</taxonomicName>
, which is densely and persistently scaly);
<emphasis id="B96AEAF2FFBC9B1E60527FF8FF18FD7D" bold="true" box="[121,191,689,710]" pageId="85" pageNumber="75">pinnae</emphasis>
usually opposite to subopposite, sessile or increasingly adnate, especially more distal pinna (e.g., as in
<taxonomicName id="4C1E4D63FFBC9B1E64A07FF9FF1FFD5E" class="Polypodiopsida" family="Thelypteridaceae" genus="Phegopteris" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="rectangularis">
<emphasis id="B96AEAF2FFBC9B1E64A07FF9FF1FFD5E" italics="true" pageId="85" pageNumber="75">P.rectangularis</emphasis>
</taxonomicName>
), spreading or obliquely spreading, costae not grooved adaxially, truncate at bases, acute at tips, to ca. 15(20) × 2(3) cm wide, pinnatifid or pinnate-pinnatifid with pinnules strongly adnate and often connected at their bases, in larger species free, sessile or nearly so, with or without acroscopic and/or basiscopic, more lobed basal auricles;
<emphasis id="B96AEAF2FFBC9B1E64877E58FB46FC9D" bold="true" box="[1196,1249,785,806]" pageId="85" pageNumber="75">veins</emphasis>
free, often forking in ultimate segments, ± prominent, at least readily visible on both sides, vein ends clavate adaxially and not reaching segment margins;
<emphasis id="B96AEAF2FFBC9B1E61E27E19FD9DFCDE" bold="true" box="[457,570,848,869]" pageId="85" pageNumber="75">aerophores</emphasis>
absent at pinna bases;
<emphasis id="B96AEAF2FFBC9B1E63247E18FC74FCDD" bold="true" box="[783,979,849,870]" pageId="85" pageNumber="75">indument abaxially</emphasis>
, if present, of sparse to moderately dense unicellular hyaline acicular hairs, lacking scales, blades often glabrescent with age; short-stipitate glands sometimes present along costae and costules;
<emphasis id="B96AEAF2FFBC9B1E62117ED8FCA3FC1D" bold="true" box="[570,772,913,934]" pageId="85" pageNumber="75">indument adaxially</emphasis>
of hyaline acicular hairs to ca.
<quantity id="4CE69B05FFBC9B1E64117ED8FBD3FC1E" box="[1082,1140,912,934]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="85" pageNumber="75" unit="mm" value="1.0">1 mm</quantity>
long along costae, sometimes also with hairs on costules and ultimate veins, occasionally on laminar tissue between veins;
<emphasis id="B96AEAF2FFBC9B1E60527E99FF77FC5E" bold="true" box="[121,208,976,997]" pageId="85" pageNumber="75">pustules</emphasis>
absent on abaxial surfaces between veins;
<emphasis id="B96AEAF2FFBC9B1E625A7E98FD3EFC5D" bold="true" box="[625,665,977,998]" pageId="85" pageNumber="75">sori</emphasis>
medial to supramedial, round to oblong (length 2 × width), exindusiate, not confluent at maturity;
<emphasis id="B96AEAF2FFBC9B1E61DD7EB8FDFCFBBD" bold="true" box="[502,603,1009,1030]" pageId="85" pageNumber="75">sporangia</emphasis>
short-stalked, capsules glabrous or with short setulae 0.10.2 mm and/or yellowish short-stipitate glands ca. 0.1 mm adjacent to annulus;
<emphasis id="B96AEAF2FFBC9B1E63017959FCCBFB9E" bold="true" box="[810,876,1040,1061]" pageId="85" pageNumber="75">spores</emphasis>
tan to brown, with numerous shallow and narrow ridges forming a reticulate network of polygonal areoles having a smooth or papillate surface (
<bibRefCitation id="EF8F4B11FFBC9B1E64A77978FF0DFBDD" author="Holttum" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="85" pageNumber="75" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">Holttum 1969</bibRefCitation>
;
<bibRefCitation id="EF8F4B11FFBC9B1E60997919FE3CFBDD" author="Tryon &amp; Lugardon" box="[178,411,1104,1126]" firstAuthor="Tryon" journalOrPublisher="Springer-Verlag, New York, U. S. A" pageId="85" pageNumber="75" refId="ref115218" refString="TRYON, A. F. &amp; B. LUGARDON. 1991. Spores of Pteridophyta. Springer-Verlag, New York, U. S. A." title="Spores of Pteridophyta" type="book" year="1991">Tryon &amp; Lugardon 1991</bibRefCitation>
;
<bibRefCitation id="EF8F4B11FFBC9B1E61887918FDE5FBDD" author="Patel" box="[419,578,1104,1126]" etAl="et al." firstAuthor="Patel" journalOrPublisher="Int. J. Pl. Sci." pageId="85" pageNumber="75" 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>
), lacking broad wings;
<emphasis id="B96AEAF2FFBC9B1E63307919FC81FBDE" box="[795,806,1104,1125]" italics="true" pageId="85" pageNumber="75">x</emphasis>
= 31, diploids and tetraploids known, about 10 spp. counted. No hybridization with any other genus has been demonstrated.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBC9B1E608279D9FB98FAFD" pageId="85" pageNumber="75" type="diagnosis">
<paragraph id="8BA136E0FFBC9B1E608279D9FB98FAFD" blockId="85.[121,1250,336,1797]" pageId="85" pageNumber="75">
<emphasis id="B96AEAF2FFBC9B1E608279D9FEADFB1E" box="[169,266,1168,1189]" italics="true" pageId="85" pageNumber="75">Diagnosis.</emphasis>
—Characters used by
<bibRefCitation id="EF8F4B11FFBC9B1E61C879D8FDDCFB1D" author="Holttum" box="[483,635,1169,1190]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="85" pageNumber="75" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">Holttum (1969)</bibRefCitation>
to separate
<taxonomicName id="4C1E4D63FFBC9B1E62DB79C6FC3CFB1F" box="[752,923,1167,1188]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E62DB79C6FC3CFB1F" box="[752,923,1167,1188]" italics="true" pageId="85" pageNumber="75">Pseudophegopteris</emphasis>
</taxonomicName>
from
<taxonomicName id="4C1E4D63FFBC9B1E63FD79C6FBD4FB1F" box="[982,1139,1167,1188]" class="Polypodiopsida" family="Thelypteridaceae" genus="Macrothelypteris" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E63FD79C6FBD4FB1F" box="[982,1139,1167,1188]" italics="true" pageId="85" pageNumber="75">Macrothelypteris</emphasis>
</taxonomicName>
include: 1) thinner, brownish stipe base and rhizome apex scales on
<taxonomicName id="4C1E4D63FFBC9B1E628379E6FCF5FB7F" box="[680,850,1199,1220]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E628379E6FCF5FB7F" box="[680,850,1199,1220]" italics="true" pageId="85" pageNumber="75">Pseudophegopteris</emphasis>
</taxonomicName>
(vs. pale thickened scales); 2) absence of septate hairs on axes and blades in
<taxonomicName id="4C1E4D63FFBC9B1E61EF7986FDC9FB5F" box="[452,622,1231,1252]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E61EF7986FDC9FB5F" box="[452,622,1231,1252]" italics="true" pageId="85" pageNumber="75">Pseudophegopteris</emphasis>
</taxonomicName>
(vs. often septate); and exindusiate sori (vs. often with small indusia). In addition, ultimate segments and distal pinnae in
<taxonomicName id="4C1E4D63FFBC9B1E62B379A6FCE5FABF" box="[664,834,1263,1284]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E62B379A6FCE5FABF" box="[664,834,1263,1284]" italics="true" pageId="85" pageNumber="75">Pseudophegopteris</emphasis>
</taxonomicName>
are more pronouncedly adnate, the pinnae are opposite (vs. alternate), and there are differences in spore ornamentation. These two genera have retained their rank in publications on the family by
<bibRefCitation id="EF8F4B11FFBC9B1E61817878FD86FAFD" author="Smith" box="[426,545,1329,1350]" editor="Kubitzki, K." firstAuthor="Smith" journalOrPublisher="K. U. Kramer &amp; P. S. Green. Springer-Verlag, Berlin, Germany. Pp" pageId="85" pageNumber="75" pagination="263 - 272" refId="ref114359" refString="SMITH, A. R. 1990. Thelypteridaceae. In: Kubitzki, K., ed. The families and genera of vascular plants. I. Pteridophytes and gymnosperms. Vol. eds. K. U. Kramer &amp; P. S. Green. Springer-Verlag, Berlin, Germany. Pp. 263 - 272." title="Thelypteridaceae" type="book chapter" volumeTitle="The families and genera of vascular plants. I. Pteridophytes and gymnosperms. Vol" year="1990">
<collectingRegion id="49DAF802FFBC9B1E61817878FE43FAFD" box="[426,484,1329,1350]" country="Bermuda" name="Smith's" pageId="85" pageNumber="75">Smith</collectingRegion>
(1990
</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBC9B1E62017878FDFAFAFD" author="Smith" box="[554,605,1329,1350]" firstAuthor="Smith" journalOrPublisher="Taxon" pageId="85" pageNumber="75" pagination="705 - 731" part="55" refId="ref114696" refString="SMITH, A. R., K. M. PRYER, E. SCHUETTPELZ, P. KORALL, H. SCHNEIDER, &amp; P. G. WOLF. 2006. A classification for extant ferns. Taxon 55: 705 - 731." title="A classification for extant ferns" type="journal article" year="2006">2006</bibRefCitation>
) and in
<bibRefCitation id="EF8F4B11FFBC9B1E62837878FC84FAFD" author="PPG I" box="[680,803,1328,1350]" firstAuthor="PPG I" journalOrPublisher="J. Syst. Evol." pageId="85" pageNumber="75" pagination="563 - 603" part="54" refId="ref112966" refString="PPG I. 2016. A community-derived classification for extant lycophytes and ferns. J. Syst. Evol. 54: 563 - 603." title="A community-derived classification for extant lycophytes and ferns" type="journal article" year="2016">PPG I (2016)</bibRefCitation>
, as well as many recent floras.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBC9B1E60827806FEA3F97E" pageId="85" pageNumber="75" type="biology_ecology">
<paragraph id="8BA136E0FFBC9B1E60827806FEA3F97E" blockId="85.[121,1250,336,1797]" pageId="85" pageNumber="75">
<emphasis id="B96AEAF2FFBC9B1E60827806FDF9FADF" box="[169,606,1359,1380]" italics="true" pageId="85" pageNumber="75">
Biogeography and ecology.—
<taxonomicName id="4C1E4D63FFBC9B1E61987806FDF9FADF" box="[435,606,1359,1380]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">Pseudophegopteris</taxonomicName>
</emphasis>
comprises 28 species and is widely distributed from tropical West Africa (and offshore island endemic species on
<collectingCountry id="F3097670FFBC9B1E62177838FD3EFA3D" box="[572,665,1393,1414]" name="Sao Tome and Principe" pageId="85" pageNumber="75">São Tomé</collectingCountry>
and
<collectingCountry id="F3097670FFBC9B1E62EC7838FC8DFA3D" box="[711,810,1393,1414]" name="Saint Helena, Ascension and Tristan da Cunha" pageId="85" pageNumber="75">St. Helena</collectingCountry>
),
<collectingCountry id="F3097670FFBC9B1E63127838FC0AFA3D" box="[825,941,1393,1414]" name="Madagascar" pageId="85" pageNumber="75">Madagascar</collectingCountry>
,
<collectingCountry id="F3097670FFBC9B1E639D7838FBADFA3D" box="[950,1034,1393,1414]" name="Réunion" pageId="85" pageNumber="75">Réunion</collectingCountry>
,
<collectingCountry id="F3097670FFBC9B1E64387838FBEFFA3D" box="[1043,1096,1393,1414]" name="India" pageId="85" pageNumber="75">India</collectingCountry>
, southeast Asia, and
<collectingCountry id="F3097670FFBC9B1E608978D9FF7DFA1E" box="[162,218,1424,1445]" name="Japan" pageId="85" pageNumber="75">Japan</collectingCountry>
, through
<collectingCountry id="F3097670FFBC9B1E611D78D8FE27FA1D" box="[310,384,1425,1446]" name="Malaysia" pageId="85" pageNumber="75">Malesia</collectingCountry>
to
<collectingCountry id="F3097670FFBC9B1E61B578D8FE19FA1D" box="[414,446,1425,1446]" name="Fiji" pageId="85" pageNumber="75">Fiji</collectingCountry>
and Hawaii (
<bibRefCitation id="EF8F4B11FFBC9B1E621778D8FD60FA1D" author="Holttum" box="[572,711,1425,1446]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="85" pageNumber="75" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">Holttum 1969</bibRefCitation>
). The greatest diversity in the genus is in
<collectingCountry id="F3097670FFBC9B1E647B78D8FB29FA1D" box="[1104,1166,1425,1446]" name="China" pageId="85" pageNumber="75">China</collectingCountry>
, with 12 species (including four endemics), one with two varieties (
<bibRefCitation id="EF8F4B11FFBC9B1E62B778F8FC83FA7D" author="Lin" box="[668,804,1456,1478]" etAl="et al." firstAuthor="Lin" journalOrPublisher="Science Press, Beijing, China; Missouri Botanical Garden Press, St. Louis, U. S. A." pageId="85" pageNumber="75" pagination="319 - 396" refId="ref111240" refString="LIN, Y. X., Z. Y. LI, K. IWATSUKI, &amp; A. R. SMITH. 2013. Thelypteridaceae. In Z. Y. Wu, P. H. Raven, &amp; D. Y. Hong, eds., Flora of China, Vol. 2 - 3 (Pteridophytes). Science Press, Beijing, China; Missouri Botanical Garden Press, St. Louis, U. S. A. Pp. 319 - 396." title="Thelypteridaceae" type="book chapter" volumeTitle="Z. Y. Wu, P. H. Raven, &amp; D. Y. Hong" year="2013">Lin et al. 2013</bibRefCitation>
). One of the oceanic island endemics,
<taxonomicName id="4C1E4D63FFBC9B1E64A278F9FB7DFA7F" box="[1161,1242,1455,1477]" class="Polypodiopsida" family="Thelypteridaceae" genus="Phegopteris" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="dianae">
<emphasis id="B96AEAF2FFBC9B1E64A278F9FB7DFA7F" box="[1161,1242,1455,1477]" italics="true" pageId="85" pageNumber="75">P.dianae</emphasis>
</taxonomicName>
, from
<collectingCountry id="F3097670FFBC9B1E60857898FEB4FA5D" box="[174,275,1489,1510]" name="Saint Helena, Ascension and Tristan da Cunha" pageId="85" pageNumber="75">St. Helena</collectingCountry>
, is remarkable in its very dense, persistent, light brown, glabrous, ovate scales along the rachis. The other five Atlantic and Indian Ocean island species,
<taxonomicName id="4C1E4D63FFBC9B1E624A78B9FCA7F9BF" box="[609,768,1519,1541]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="andringitrensis">
<emphasis id="B96AEAF2FFBC9B1E624A78B9FCA7F9BF" box="[609,768,1519,1541]" italics="true" pageId="85" pageNumber="75">P.andringitrensis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1E4D63FFBC9B1E632178B9FCC1F9BF" box="[778,870,1519,1541]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="aubertii">
<emphasis id="B96AEAF2FFBC9B1E632178B9FCC1F9BF" box="[778,870,1519,1541]" italics="true" pageId="85" pageNumber="75">P. aubertii</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1E4D63FFBC9B1E635B78B9FC77F9BE" box="[880,976,1520,1541]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="cruciata">
<emphasis id="B96AEAF2FFBC9B1E635B78B9FC77F9BE" box="[880,976,1520,1541]" italics="true" pageId="85" pageNumber="75">P.cruciata</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1E4D63FFBC9B1E63F178B9FBF3F9BF" box="[986,1108,1519,1541]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="henriquesii">
<emphasis id="B96AEAF2FFBC9B1E63F178B9FBF3F9BF" box="[986,1108,1519,1541]" italics="true" pageId="85" pageNumber="75">P. henriquesii</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C1E4D63FFBC9B1E64AE78B9FF3CF99F" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="rammelooi">
<emphasis id="B96AEAF2FFBC9B1E64AE78B9FF3CF99F" italics="true" pageId="85" pageNumber="75">P.rammelooi</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EF8F4B11FFBC9B1E60E17B58FEFAF99D" author="Holttum" box="[202,349,1553,1574]" firstAuthor="Holttum" journalOrPublisher="Allertonia" pageId="85" pageNumber="75" pagination="169 - 234" part="1" refId="ref109882" refString="HOLTTUM, R. E. 1977 b. The family Thelypteridaceae in the Pacific and Australasia. Allertonia 1: 169 - 234." title="The family Thelypteridaceae in the Pacific and Australasia" type="journal article" year="1977" yearSuffix="b">Holttum 1977b</bibRefCitation>
;
<bibRefCitation id="EF8F4B11FFBC9B1E614E7B58FD84F99D" author="Pichi Sermolli" box="[357,547,1553,1574]" firstAuthor="Pichi Sermolli" journalOrPublisher="I. Bull. Jard. Bot. Natl. Belg." pageId="85" pageNumber="75" pagination="177 - 284" part="53" refId="ref112883" refString="PICHI SERMOLLI, R. E. G. 1983. A contribution to the knowledge of the Pteridophyta of Rwanda, Burundi, and Kivu (Zaire). I. Bull. Jard. Bot. Natl. Belg. 53: 177 - 284." title="A contribution to the knowledge of the Pteridophyta of Rwanda, Burundi, and Kivu (Zaire)" type="journal article" year="1983">Pichi Sermolli 1983</bibRefCitation>
), are much more similar to the larger, bipinnate-pinnatifid species such as
<taxonomicName id="4C1E4D63FFBC9B1E60527B79FEADF9FF" authorityName="Ching" authorityYear="1963" baseAuthorityName="Ching" box="[121,266,1583,1605]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="yunkweiensis">
<emphasis id="B96AEAF2FFBC9B1E60527B79FEADF9FF" box="[121,266,1583,1605]" italics="true" pageId="85" pageNumber="75">P. yunkweiensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1E4D63FFBC9B1E611D7B79FE3AF9FF" box="[310,413,1583,1605]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="paludosa">
<emphasis id="B96AEAF2FFBC9B1E611D7B79FE3AF9FF" box="[310,413,1583,1605]" italics="true" pageId="85" pageNumber="75">P.paludosa</emphasis>
</taxonomicName>
, from southeast Asia and
<collectingCountry id="F3097670FFBC9B1E62BF7B79FD79F9FE" box="[660,734,1584,1605]" name="Malaysia" pageId="85" pageNumber="75">Malesia</collectingCountry>
(
<bibRefCitation id="EF8F4B11FFBC9B1E62C27B79FCDAF9FE" author="Holttum" box="[745,893,1584,1605]" firstAuthor="Holttum" journalOrPublisher="J. S. African Bot." pageId="85" pageNumber="75" pagination="123 - 168" part="40" refId="ref109577" refString="HOLTTUM, R. E. 1974 a. Thelypteridaceae of Africa and adjacent islands. J. S. African Bot. 40: 123 - 168." title="Thelypteridaceae of Africa and adjacent islands" type="journal article" year="1974" yearSuffix="a">Holttum 1974a</bibRefCitation>
). The sole species in Hawaii,
<taxonomicName id="4C1E4D63FFBC9B1E64BB7B79FF6CF9DF" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="species" species="keraudreniana">
<emphasis id="B96AEAF2FFBC9B1E64BB7B79FF6CF9DF" italics="true" pageId="85" pageNumber="75">P.keraudreniana</emphasis>
</taxonomicName>
, is quite rare and localized; it differs from all others in the genus in having indeterminate growth, with dormant tips growing intermittently after lower pinnae mature (
<bibRefCitation id="EF8F4B11FFBC9B1E62AC7B39FCA2F93E" author="Palmer" box="[647,773,1648,1669]" firstAuthor="Palmer" journalOrPublisher="University of Hawaii Press, Honolulu, U. S. A" pageId="85" pageNumber="75" refId="ref112628" refString="PALMER, D. D. 2003. Hawai'i's ferns and fern allies. University of Hawaii Press, Honolulu, U. S. A." title="Hawai'i's ferns and fern allies" type="book" year="2003">Palmer 2003</bibRefCitation>
). Species of
<taxonomicName id="4C1E4D63FFBC9B1E63537B26FB85F93F" box="[888,1058,1647,1668]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E63537B26FB85F93F" box="[888,1058,1647,1668]" italics="true" pageId="85" pageNumber="75">Pseudophegopteris</emphasis>
</taxonomicName>
occur mainly along streams, in thickets, and in rock crevices in dense lowland and montane forests, up to
<quantity id="4CE69B05FFBC9B1E63827BD9FC56F91E" box="[937,1009,1680,1701]" metricMagnitude="3" metricUnit="m" metricValue="3.1" pageId="85" pageNumber="75" unit="m" value="3100.0">3100 m</quantity>
; most species occur from
<quantity id="4CE69B05FFBC9B1E60527BF9FEA7F97E" box="[121,256,1712,1733]" metricMagnitude="3" metricUnit="m" metricValue="1.75" metricValueMax="2.5" metricValueMin="1.0" pageId="85" pageNumber="75" unit="m" value="1750.0" valueMax="2500.0" valueMin="1000.0">10002500 m</quantity>
.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBC9B1D60827B99FBF8FCAC" lastPageId="86" lastPageNumber="76" pageId="85" pageNumber="75" type="discussion">
<paragraph id="8BA136E0FFBC9B1D60827B99FF78FD8C" blockId="85.[121,1250,336,1797]" lastBlockId="86.[97,1226,161,791]" lastPageId="86" lastPageNumber="76" pageId="85" pageNumber="75">
<emphasis id="B96AEAF2FFBC9B1E60827B99FDABF95F" box="[169,524,1743,1765]" italics="true" pageId="85" pageNumber="75">Taxonomic and phylogenetic studies.—</emphasis>
Both
<taxonomicName id="4C1E4D63FFBC9B1E62147B86FD4EF95F" box="[575,745,1743,1764]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E62147B86FD4EF95F" box="[575,745,1743,1764]" italics="true" pageId="85" pageNumber="75">Pseudophegopteris</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1E4D63FFBC9B1E633C7B86FC14F95F" box="[791,947,1743,1764]" class="Polypodiopsida" family="Thelypteridaceae" genus="Macrothelypteris" kingdom="Plantae" order="Polypodiales" pageId="85" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBC9B1E633C7B86FC14F95F" box="[791,947,1743,1764]" italics="true" pageId="85" pageNumber="75">Macrothelypteris</emphasis>
</taxonomicName>
were first recognized at generic rank by
<bibRefCitation id="EF8F4B11FFBC9B1E60E37BB9FEEEF8BE" author="Ching" box="[200,329,1776,1797]" firstAuthor="Ching" journalOrPublisher="Acta Phytotax. Sin." pageId="85" pageNumber="75" pagination="289 - 335" part="8" refId="ref107523" refString="CHING, R. C. 1963. A reclassification of the family Thelypteridaceae from the mainland of Asia. Acta Phytotax. Sin. 8: 289 - 335." title="A reclassification of the family Thelypteridaceae from the mainland of Asia" type="journal article" year="1963">Ching (1963)</bibRefCitation>
. Holttums studies further clarified their distinction and differences (
<bibRefCitation id="EF8F4B11FFBC9B1E63F37BB9FBC5F8BE" author="Holttum" box="[984,1122,1776,1797]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="85" pageNumber="75" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">Holttum 1969</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBC9B1E64417BB9FB3DF8BE" author="Holttum" box="[1130,1178,1776,1797]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="85" pageNumber="75" pagination="17 - 52" part="19" refId="ref109463" refString="HOLTTUM, R. E. 1971. Studies in the family Thelypteridaceae III. A new system of genera in the Old World. Blumea 19: 17 - 52." title="Studies in the family Thelypteridaceae III. A new system of genera in the Old World" type="journal article" year="1971">1971</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBC9B1E64887BB9FB79F8BE" author="Holttum" box="[1187,1246,1776,1797]" firstAuthor="Holttum" journalOrPublisher="J. S. African Bot." pageId="85" pageNumber="75" pagination="123 - 168" part="40" refId="ref109577" refString="HOLTTUM, R. E. 1974 a. Thelypteridaceae of Africa and adjacent islands. J. S. African Bot. 40: 123 - 168." title="Thelypteridaceae of Africa and adjacent islands" type="journal article" year="1974" yearSuffix="a">1974a</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBF9B1D604A7DEBFF39FF0C" author="Holttum" box="[97,158,162,183]" firstAuthor="Holttum" journalOrPublisher="Allertonia" pageId="86" pageNumber="76" pagination="169 - 234" part="1" refId="ref109882" refString="HOLTTUM, R. E. 1977 b. The family Thelypteridaceae in the Pacific and Australasia. Allertonia 1: 169 - 234." title="The family Thelypteridaceae in the Pacific and Australasia" type="journal article" year="1977" yearSuffix="b">1977b</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBF9B1D608C7DEBFF7EFF0C" author="Holttum" box="[167,217,162,183]" firstAuthor="Holttum" journalOrPublisher="Flora Malesiana" pageId="86" pageNumber="76" pagination="334 - 560" part="1" refId="ref109955" refString="HOLTTUM, R. E. 1982. Thelypteridaceae. Flora Malesiana, Ser. II, 1 (5): 334 - 560." title="Thelypteridaceae" type="journal article" year="1982">1982</bibRefCitation>
).
<taxonomicName id="4C1E4D63FFBF9B1D60C27DE8FE34FF0D" box="[233,403,161,182]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D60C27DE8FE34FF0D" box="[233,403,161,182]" italics="true" pageId="86" pageNumber="76">Pseudophegopteris</emphasis>
</taxonomicName>
is most closely related to
<taxonomicName id="4C1E4D63FFBF9B1D62A47DE8FC8CFF0D" box="[655,811,161,182]" class="Polypodiopsida" family="Thelypteridaceae" genus="Macrothelypteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D62A47DE8FC8CFF0D" box="[655,811,161,182]" italics="true" pageId="86" pageNumber="76">Macrothelypteris</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1E4D63FFBF9B1D63707DE8FC63FF0D" box="[859,964,161,182]" class="Polypodiopsida" family="Thelypteridaceae" genus="Phegopteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D63707DE8FC63FF0D" box="[859,964,161,182]" italics="true" pageId="86" pageNumber="76">Phegopteris</emphasis>
</taxonomicName>
, which together constitute the subfamily
<taxonomicName id="4C1E4D63FFBF9B1D60C37D8BFDBEFF6C" authority="(PPG I 2016)" baseAuthorityName="PPG I" baseAuthorityYear="2016" box="[232,537,194,215]" class="Polypodiopsida" family="Thelypteridaceae" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="subFamily" subFamily="Phegopteridoideae">
Phegopteridoideae (
<bibRefCitation id="EF8F4B11FFBF9B1D618F7D8BFDB4FF6C" author="PPG I" box="[420,531,194,215]" firstAuthor="PPG I" journalOrPublisher="J. Syst. Evol." pageId="86" pageNumber="76" pagination="563 - 603" part="54" refId="ref112966" refString="PPG I. 2016. A community-derived classification for extant lycophytes and ferns. J. Syst. Evol. 54: 563 - 603." title="A community-derived classification for extant lycophytes and ferns" type="journal article" year="2016">
PPG
<date id="FFA01020FFBF9B1D61FD7D8BFDB4FF6C" box="[470,531,194,215]" pageId="86" pageNumber="76" value="2016-01">I 2016</date>
</bibRefCitation>
)
</taxonomicName>
. In our analyses (Fawcett et al. in press), and also those of He and Zhang (2012),
<bibRefCitation id="EF8F4B11FFBF9B1D608C7DABFEDCFF4C" author="Schneider" box="[167,379,226,247]" etAl="et al." firstAuthor="Schneider" journalOrPublisher="J. Syst. Evol." pageId="86" pageNumber="76" pagination="61 - 70" part="51" refId="ref113867" refString="SCHNEIDER, H., L-J. HE, J. MARQUARDT, L. WANG, J. HEINRICHS, S. HENNEQUIN, &amp; X-C. ZHANG. 2013. Exploring the origin of the latitudinal diversity gradient: contrasting the sister fern genera Phegopteris and Pseudophegopteris. J. Syst. Evol. 51: 61 - 70." title="Exploring the origin of the latitudinal diversity gradient: contrasting the sister fern genera Phegopteris and Pseudophegopteris" type="journal article" year="2013">Schneider et al. (2013)</bibRefCitation>
, and
<bibRefCitation id="EF8F4B11FFBF9B1D61817DABFDC8FF4C" 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="[426,623,226,247]" journalOrPublisher="Molec. Phylogen. Evol." pageId="86" pageNumber="76" 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>
,
<taxonomicName id="4C1E4D63FFBF9B1D625D7DA8FC87FF4D" box="[630,800,225,246]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D625D7DA8FC87FF4D" box="[630,800,225,246]" italics="true" pageId="86" pageNumber="76">Pseudophegopteris</emphasis>
</taxonomicName>
is monophyletic and sister to a monophyletic
<taxonomicName id="4C1E4D63FFBF9B1D604A7C48FF6DFEAD" box="[97,202,257,278]" class="Polypodiopsida" family="Thelypteridaceae" genus="Phegopteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D604A7C48FF6DFEAD" box="[97,202,257,278]" italics="true" pageId="86" pageNumber="76">Phegopteris</emphasis>
</taxonomicName>
, and this combined clade is in turn sister to
<taxonomicName id="4C1E4D63FFBF9B1D625B7C48FCACFEAD" box="[624,779,257,278]" class="Polypodiopsida" family="Thelypteridaceae" genus="Macrothelypteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D625B7C48FCACFEAD" box="[624,779,257,278]" italics="true" pageId="86" pageNumber="76">Macrothelypteris</emphasis>
</taxonomicName>
. The close relationship of these three genera is evidenced by the fact they are all free-veined, often with forked veins that end before reaching the segment margins, and stipes, rachises, and costae not grooved adaxially. This separates them from all other genera of
<taxonomicName id="4C1E4D63FFBF9B1D63F07C0BFB25FEEC" baseAuthorityName="Holttum" baseAuthorityYear="1976" box="[987,1154,322,343]" class="Polypodiopsida" family="Thelypteridaceae" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="family">Thelypteridaceae</taxonomicName>
, except
<taxonomicName id="4C1E4D63FFBF9B1D604A7C28FF56FECD" box="[97,241,353,374]" class="Polypodiopsida" family="Thelypteridaceae" genus="Metathelypteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D604A7C28FF56FECD" box="[97,241,353,374]" italics="true" pageId="86" pageNumber="76">Metathelypteris</emphasis>
</taxonomicName>
and some
<taxonomicName id="4C1E4D63FFBF9B1D617C7C2BFE70FECC" box="[343,471,354,375]" class="Polypodiopsida" family="Thelypteridaceae" genus="Leptogramma" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D617C7C2BFE70FECC" box="[343,471,354,375]" italics="true" pageId="86" pageNumber="76">Leptogramma</emphasis>
</taxonomicName>
, which are not closely related. The phegopteroids were clearly understood and delineated by
<bibRefCitation id="EF8F4B11FFBF9B1D60CE7CCBFEDCFE2C" author="Holttum" box="[229,379,386,407]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="86" pageNumber="76" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">Holttum (1969)</bibRefCitation>
, who provided a revision/synopsis of these genera, prior to their recognition as subfamily
<taxonomicName id="4C1E4D63FFBF9B1D604A7CEBFE2CFE0C" authority="(PPG I 2016)" baseAuthorityName="PPG I" baseAuthorityYear="2016" box="[97,395,418,439]" class="Polypodiopsida" family="Thelypteridaceae" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="subFamily" subFamily="Phegopteridoideae">
Phegopteridoideae(
<bibRefCitation id="EF8F4B11FFBF9B1D61317CEBFE22FE0C" author="PPG I" box="[282,389,418,439]" firstAuthor="PPG I" journalOrPublisher="J. Syst. Evol." pageId="86" pageNumber="76" pagination="563 - 603" part="54" refId="ref112966" refString="PPG I. 2016. A community-derived classification for extant lycophytes and ferns. J. Syst. Evol. 54: 563 - 603." title="A community-derived classification for extant lycophytes and ferns" type="journal article" year="2016">
PPG
<date id="FFA01020FFBF9B1D61617CEBFE22FE0C" box="[330,389,418,439]" pageId="86" pageNumber="76" value="2016-01">I2016</date>
</bibRefCitation>
)
</taxonomicName>
.In part,because of the greater blade dissection in
<taxonomicName id="4C1E4D63FFBF9B1D637F7CE8FC59FE0D" box="[852,1022,417,438]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D637F7CE8FC59FE0D" box="[852,1022,417,438]" italics="true" pageId="86" pageNumber="76">Pseudophegopteris</emphasis>
</taxonomicName>
and
<emphasis id="B96AEAF2FFBF9B1D640C7CE8FB6EFE0D" box="[1063,1225,417,438]" italics="true" pageId="86" pageNumber="76">
<taxonomicName id="4C1E4D63FFBF9B1D640C7CE8FB62FE0D" box="[1063,1221,417,438]" class="Polypodiopsida" family="Thelypteridaceae" genus="Macrothelypteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">Macrothelypteris</taxonomicName>
,
</emphasis>
but also because of a belief in the relationship of
<taxonomicName id="4C1E4D63FFBF9B1D62077C8BFD76FE6C" baseAuthorityName="Holttum" baseAuthorityYear="1976" box="[556,721,450,471]" class="Polypodiopsida" family="Thelypteridaceae" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="family">Thelypteridaceae</taxonomicName>
to Cyatheaceae (
<bibRefCitation id="EF8F4B11FFBF9B1D63447C8BFC50FE6C" author="Holttum" box="[879,1015,450,471]" firstAuthor="Holttum" journalOrPublisher="Gardens' Bull. (Singapore)" pageId="86" pageNumber="76" pagination="267 - 298" part="11" refId="ref109375" refString="HOLTTUM, R. E. 1947. New species of vascular plants from the Malay Peninsula. Gardens' Bull. (Singapore) 11: 267 - 298." title="New species of vascular plants from the Malay Peninsula" type="journal article" year="1947">Holttum 1947</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBF9B1D63D57C8BFB88FE6C" author="Holttum" box="[1022,1071,450,471]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="86" pageNumber="76" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">1969</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBF9B1D641C7C8BFBCFFE6C" author="Holttum" box="[1079,1128,450,471]" firstAuthor="Holttum" journalOrPublisher="Flora Malesiana" pageId="86" pageNumber="76" pagination="334 - 560" part="1" refId="ref109955" refString="HOLTTUM, R. E. 1982. Thelypteridaceae. Flora Malesiana, Ser. II, 1 (5): 334 - 560." title="Thelypteridaceae" type="journal article" year="1982">1982</bibRefCitation>
), Holttum thought these three genera showed the most primitive frond-form in the family. Although they are sister to all other taxa of
<taxonomicName id="4C1E4D63FFBF9B1D60517F4BFEB8FDAC" baseAuthorityName="Holttum" baseAuthorityYear="1976" box="[122,287,514,535]" class="Polypodiopsida" family="Thelypteridaceae" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="family">Thelypteridaceae</taxonomicName>
in molecular analyses, all evidence suggests that they are not closely related to, or derived from, Cyatheaceae.
</paragraph>
<paragraph id="8BA136E0FFBF9B1D60BA7F0BFBE2FD4C" blockId="86.[97,1226,161,791]" pageId="86" pageNumber="76">
<emphasis id="B96AEAF2FFBF9B1D60BA7F0BFF6CFDEC" box="[145,203,578,599]" italics="true" pageId="86" pageNumber="76">Notes.</emphasis>
—Based on a sample of five species, spores of
<taxonomicName id="4C1E4D63FFBF9B1D62AB7F08FC8DFDED" box="[640,810,577,598]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D62AB7F08FC8DFDED" box="[640,810,577,598]" italics="true" pageId="86" pageNumber="76">Pseudophegopteris</emphasis>
</taxonomicName>
are very distinctive and consistently sculptured: low ridges forming a polygonal reticulate network, smooth papillose intra-areolar surfaces, and a gemmulate exospore (
<bibRefCitation id="EF8F4B11FFBF9B1D60EF7FCBFE14FD2C" author="Tryon &amp; Lugardon" box="[196,435,642,663]" firstAuthor="Tryon" journalOrPublisher="Springer-Verlag, New York, U. S. A" pageId="86" pageNumber="76" refId="ref115218" refString="TRYON, A. F. &amp; B. LUGARDON. 1991. Spores of Pteridophyta. Springer-Verlag, New York, U. S. A." title="Spores of Pteridophyta" type="book" year="1991">Tryon &amp; Lugardon 1991</bibRefCitation>
). This is somewhat similar to
<taxonomicName id="4C1E4D63FFBF9B1D62FF7FC8FC9AFD2D" box="[724,829,641,662]" class="Polypodiopsida" family="Thelypteridaceae" genus="Phegopteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D62FF7FC8FC9AFD2D" box="[724,829,641,662]" italics="true" pageId="86" pageNumber="76">Phegopteris</emphasis>
</taxonomicName>
, which has spores that differ in having a relatively unsculptured surface or a more coarsely reticulate network with a tuberculate intra-areolar surface. Spores of
<taxonomicName id="4C1E4D63FFBF9B1D604A7F88FF5AFD6D" box="[97,253,705,726]" class="Polypodiopsida" family="Thelypteridaceae" genus="Macrothelypteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D604A7F88FF5AFD6D" box="[97,253,705,726]" italics="true" pageId="86" pageNumber="76">Macrothelypteris</emphasis>
</taxonomicName>
have a finer reticulate network of ridges with smaller, more rounded areoles (similar to spores of some species of
<taxonomicName id="4C1E4D63FFBF9B1D60EA7FA8FE91FD4D" box="[193,310,737,758]" class="Polypodiopsida" family="Thelypteridaceae" genus="Amauropelta" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D60EA7FA8FE91FD4D" box="[193,310,737,758]" italics="true" pageId="86" pageNumber="76">Amauropelta</emphasis>
</taxonomicName>
), or a coarser, more irregular network of perforate folds (
<bibRefCitation id="EF8F4B11FFBF9B1D63647FABFB9DFD4C" author="Tryon &amp; Lugardon" box="[847,1082,738,759]" firstAuthor="Tryon" journalOrPublisher="Springer-Verlag, New York, U. S. A" pageId="86" pageNumber="76" refId="ref115218" refString="TRYON, A. F. &amp; B. LUGARDON. 1991. Spores of Pteridophyta. Springer-Verlag, New York, U. S. A." title="Spores of Pteridophyta" type="book" year="1991">Tryon &amp; Lugardon 1991</bibRefCitation>
).
</paragraph>
<paragraph id="8BA136E0FFBF9B1D60A47E4BFBF8FCAC" blockId="86.[97,1226,161,791]" box="[143,1119,769,791]" pageId="86" pageNumber="76">
All necessary combinations except one have been made in
<taxonomicName id="4C1E4D63FFBF9B1D62977E48FCC2FCAD" box="[700,869,769,790]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFBF9B1D62977E48FCC2FCAD" box="[700,869,769,790]" italics="true" pageId="86" pageNumber="76">Pseudophegopteris</emphasis>
</taxonomicName>
, by
<bibRefCitation id="EF8F4B11FFBF9B1D63A77E4BFBBDFCAC" author="Holttum" box="[908,1050,770,791]" firstAuthor="Holttum" journalOrPublisher="Blumea" pageId="86" pageNumber="76" pagination="5 - 32" part="17" refId="ref109433" refString="HOLTTUM, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5 - 32." title="Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris" type="journal article" year="1969">Holttum (1969</bibRefCitation>
,
<bibRefCitation id="EF8F4B11FFBF9B1D64087E4BFBF2FCAC" author="Holttum" box="[1059,1109,770,791]" firstAuthor="Holttum" journalOrPublisher="Flora Malesiana" pageId="86" pageNumber="76" pagination="334 - 560" part="1" refId="ref109955" refString="HOLTTUM, R. E. 1982. Thelypteridaceae. Flora Malesiana, Ser. II, 1 (5): 334 - 560." title="Thelypteridaceae" type="journal article" year="1982">1982</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBF9B1D604A7E1BFCE0FCDD" box="[97,839,849,871]" pageId="86" pageNumber="76" type="nomenclature">
<paragraph id="8BA136E0FFBF9B1D604A7E1BFCE0FCDD" blockId="86.[97,1227,812,897]" box="[97,839,849,871]" pageId="86" pageNumber="76">
<heading id="D0E9818CFFBF9B1D604A7E1BFCE0FCDD" box="[97,839,849,871]" level="2" pageId="86" pageNumber="76">
<taxonomicName id="4C1E4D63FFBF9B1D604A7E1BFD70FCDC" ID-CoL="8RTV4" authority="(Pic. Serm.) A. R. Sm. &amp; S. E. Fawc." authorityName="A. R. Sm. &amp; S. E. Fawc." baseAuthorityName="Pic. Serm." box="[97,727,850,871]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pseudophegopteris" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="species" species="rammelooi" status="comb. nov.">
<emphasis id="B96AEAF2FFBF9B1D604A7E1BFE34FCDC" bold="true" box="[97,403,850,871]" pageId="86" pageNumber="76">Pseudophegopteris rammelooi</emphasis>
(Pic.Serm.) A.R. Sm. &amp; S.E. Fawc.
</taxonomicName>
,
<emphasis id="B96AEAF2FFBF9B1D62F67E18FCE0FCDD" bold="true" box="[733,839,849,870]" pageId="86" pageNumber="76">
<taxonomicNameLabel id="A2595789FFBF9B1D62F67E18FCE0FCDD" box="[733,839,849,870]" pageId="86" pageNumber="76" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFBF9B1D636C7E1CFDA1FC3A" pageId="86" pageNumber="76" type="reference_group">
<paragraph id="8BA136E0FFBF9B1D636C7E1CFDA1FC3A" blockId="86.[97,1227,812,897]" pageId="86" pageNumber="76">
<taxonomicName id="4C1E4D63FFBF9B1D63717E1CFB6CFCDC" authority="Pic. Serm., Bull. Jard." box="[858,1227,853,871]" class="Polypodiopsida" family="Thelypteridaceae" genus="Macrothelypteris" kingdom="Plantae" order="Polypodiales" pageId="86" pageNumber="76" phylum="Tracheophyta" rank="species" species="rammelooi">
<emphasis id="B96AEAF2FFBF9B1D63717E1CFB8BFCDD" box="[858,1068,853,870]" italics="true" pageId="86" pageNumber="76">Macrothelypteris rammelooi</emphasis>
Pic.Serm.,Bull.Jard.
</taxonomicName>
Bot.Natl.Belg.53(1/2):270(272),fig.16.1983.
</paragraph>
</subSubSection>
</treatment>
</document>