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<document id="BB972B84B0069F7F40CFFCFDA85ACB7F" 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="03B787F6FFD39B70626C7CA3FC0BFE4C" docLanguage="en" docName="2021_ThelypteridaceaeFerns_OACopy3.pdf.imf" docOrigin="Fort Worth, Texas, USA: BRIT Press" docTitle="Hoiokula" docType="treatment" docVersion="4" lastPageNumber="49" masterDocId="FF8EFF8EFFE99B4B602B7D49FFA7FFBB" masterDocTitle="A Generic Classification of the Thelypteridaceae" pageNumber="48" updateTime="1731444927364" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-SA-4.0">
<mods:mods id="FA878BC76FDDF8D28AE6A34B304A0D4B" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="C9676822FBD0AD64B6BD132B4E78FD30">
<mods:title id="229CC73376A7C8C8DB19BB025EADC042">A Generic Classification of the Thelypteridaceae</mods:title>
</mods:titleInfo>
<mods:name id="073D5F8E58CAB0B0375A31FCE43275B3" type="personal">
<mods:role id="0C125CD17CBD18AF0AFDD01B445C7B5A">
<mods:roleTerm id="23C7C6EC155F7E2B84A9D6597355BF57">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="078940AB05EA7824BC23D39D3A513B17">Fawcett, Susan</mods:namePart>
</mods:name>
<mods:name id="C5979BFA441631BAF4CB1083B2A43344" type="personal">
<mods:role id="AB8D7357E4EE999AB892511ECD60FF72">
<mods:roleTerm id="F85F916D351B2BC2B56F294A4380EB80">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="804F70DDC5CFBE47A9D4752716BD3C16">Smith, Alan R.</mods:namePart>
</mods:name>
<mods:typeOfResource id="B360879CA78D64F816AE44D8DF907EA6">text</mods:typeOfResource>
<mods:originInfo id="44FE62722469531CDFDEAFA8810714D7">
<mods:dateIssued id="09F1CE862B939DA87E4A90147F685A55">2021</mods:dateIssued>
<mods:publisher id="7FC37000483EE39EE8E21F8422D1DB52">BRIT Press</mods:publisher>
<mods:place id="D7332E266E302D74E44BC592C33ED14B">
<mods:placeTerm id="4967CE8B1E95C7C0FAFD342F17BA82CB">Fort Worth, Texas, USA</mods:placeTerm>
</mods:place>
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<mods:classification id="F5E4F22367E0787EFA08171A26993241">book</mods:classification>
<mods:identifier id="859568650BD331AB70D3325C80FDFF3A" type="DOI">10.17348/jbrit.v15.i2.1206</mods:identifier>
<mods:identifier id="555475C7A6F54F125C8BC8CBC80B7C6E" type="Zenodo-Dep">14111022</mods:identifier>
</mods:mods>
<treatment id="03B787F6FFD39B70626C7CA3FC0BFE4C" ID-DOI="http://doi.org/10.5281/zenodo.14076443" ID-Zenodo-Dep="14076443" LSID="urn:lsid:plazi:treatment:03B787F6FFD39B70626C7CA3FC0BFE4C" httpUri="http://treatment.plazi.org/id/03B787F6FFD39B70626C7CA3FC0BFE4C" lastPageId="59" lastPageNumber="49" pageId="58" pageNumber="48">
<subSubSection id="C304656BFFD39B71626C7CA3FD45FDBF" box="[583,738,490,516]" pageId="58" pageNumber="48" type="nomenclature">
<paragraph id="8BA136E0FFD39B71626C7CA3FD45FDBF" blockId="58.[97,1225,490,577]" box="[583,738,490,516]" pageId="58" pageNumber="48">
<heading id="D0E9818CFFD39B71626C7CA3FD45FDBF" allCaps="true" bold="true" box="[583,738,490,516]" centered="true" fontSize="11" level="2" pageId="58" pageNumber="48" reason="0">
<taxonomicName id="4C1E4D63FFD39B71626C7CA3FD45FDBF" ID-CoL="8RTZF" box="[583,738,490,516]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus" status="gen. nov.">
<emphasis id="B96AEAF2FFD39B71626C7CA3FD45FDBF" bold="true" box="[583,738,490,516]" pageId="58" pageNumber="48">HOIOKULA</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFD39B71604A7F5BFDA3FD9D" box="[97,516,529,551]" pageId="58" pageNumber="48" type="reference_group">
<paragraph id="8BA136E0FFD39B71604A7F5BFDA3FD9D" blockId="58.[97,1225,490,577]" box="[97,516,529,551]" pageId="58" pageNumber="48">
<taxonomicName id="4C1E4D63FFD39B71604A7F5BFE06FD9C" authority="S.E. Fawc. &amp; A.R. Sm." authorityName="S. E. Fawc. &amp; A. R. Sm." box="[97,417,530,551]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus" status="gen. nov.">
<emphasis id="B96AEAF2FFD39B71604A7F5BFF67FD9C" bold="true" box="[97,192,530,551]" pageId="58" pageNumber="48">Hoiokula</emphasis>
S.E. Fawc. &amp; A.R. Sm.
</taxonomicName>
,
<emphasis id="B96AEAF2FFD39B7161807F58FDA3FD9D" bold="true" box="[427,516,529,550]" pageId="58" pageNumber="48">
<taxonomicNameLabel id="A2595789FFD39B7161807F58FDA3FD9D" box="[427,516,529,550]" pageId="58" pageNumber="48" rank="genus">gen. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFD39B71622F7F5CFE49FDFA" pageId="58" pageNumber="48" type="type_taxon">
<paragraph id="8BA136E0FFD39B71622F7F5CFE49FDFA" blockId="58.[97,1225,490,577]" pageId="58" pageNumber="48">
—
<typeStatus id="54A58842FFD39B71623D7F5CFD92FD9E" box="[534,565,533,549]" pageId="58" pageNumber="48">
<collectionCode id="ED0FAE25FFD39B71623D7F5CFD92FD9E" box="[534,565,533,549]" pageId="58" pageNumber="48">TYPE</collectionCode>
</typeStatus>
:
<taxonomicName id="4C1E4D63FFD39B7162147F5CFC79FD9C" authority="(Brack.) S. E. Fawc. &amp; A. R. Sm." authorityName="S. E. Fawc. &amp; A. R. Sm." baseAuthorityName="Brack." box="[575,990,533,551]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B7162147F5CFD4DFD9D" box="[575,746,533,550]" italics="true" pageId="58" pageNumber="48">Hoiokula sandwicensis</emphasis>
(Brack.) S.E. Fawc. &amp; A.R. Sm.
</taxonomicName>
[=
<taxonomicName id="4C1E4D63FFD39B7163DA7F5CFE2AFDFA" authority="Brack., U. S. Expl. Exped., Filic." authorityName="Brack., U. S. Expl. Exped., Filic." class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B7163DA7F5CFB6EFD9D" box="[1009,1225,533,550]" italics="true" pageId="58" pageNumber="48">Stegnogramma sandwicensis</emphasis>
Brack.,U.S.Expl.Exped.,Filic.
</taxonomicName>
16:26.1854.]
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFD39B71604A7F1EFD24FD36" pageId="58" pageNumber="48" type="etymology">
<paragraph id="8BA136E0FFD39B71604A7F1EFD24FD36" blockId="58.[97,1226,599,1804]" pageId="58" pageNumber="48">
<emphasis id="B96AEAF2FFD39B71604A7F1EFE98FDD7" box="[97,319,599,620]" italics="true" pageId="58" pageNumber="48">Etymology.—Ho’i’o kula</emphasis>
is the common name for
<taxonomicName id="4C1E4D63FFD39B71621A7F1EFD64FDD7" box="[561,707,599,620]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B71621A7F1EFD64FDD7" box="[561,707,599,620]" italics="true" pageId="58" pageNumber="48">H. sandwicensis</emphasis>
</taxonomicName>
in the native Hawaiian language (
<bibRefCitation id="EF8F4B11FFD39B71642D7F11FB24FDD6" author="Palmer" box="[1030,1155,600,621]" firstAuthor="Palmer" journalOrPublisher="University of Hawaii Press, Honolulu, U. S. A" pageId="58" pageNumber="48" 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>
), and it is valued for its edible fiddleheads (
<bibRefCitation id="EF8F4B11FFD39B7161877F31FDDEFD36" author="Pukui &amp; Elbert" box="[428,633,632,653]" firstAuthor="Pukui" journalOrPublisher="University of Hawaii Press. Honolulu, U. S. A. Available at wehewehe. org. Accessed Feb" pageId="58" pageNumber="48" part="2020" refId="ref113151" refString="PUKUI, M. K. &amp; S. H. ELBERT. 2003. Hawaiian-English dictionary. University of Hawaii Press. Honolulu, U. S. A. Available at wehewehe. org. Accessed Feb 2020." title="Hawaiian-English dictionary" type="journal volume" year="2003">
<collectingRegion id="49DAF802FFD39B7161877F31FE4EFD36" box="[428,489,632,653]" country="Japan" name="Fukui" pageId="58" pageNumber="48">Pukui</collectingRegion>
&amp; Elbert 2003
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFD39B7160B97FD1FF1FFB16" pageId="58" pageNumber="48" type="description">
<paragraph id="8BA136E0FFD39B7160B97FD1FF1FFB16" blockId="58.[97,1226,599,1804]" pageId="58" pageNumber="48">
Plants terrestrial, cremnophilous or rheophytic, from
<quantity id="4CE69B05FFD39B7162847FD1FD4AFD17" box="[687,749,663,685]" metricMagnitude="-1" metricUnit="m" metricValue="4.0" pageId="58" pageNumber="48" unit="cm" value="40.0">40 cm</quantity>
to&gt;
<quantity id="4CE69B05FFD39B7163087FD1FCEDFD17" box="[803,842,663,685]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="58" pageNumber="48" unit="m" value="1.0">1 m</quantity>
tall;
<emphasis id="B96AEAF2FFD39B7163547FD1FC47FD16" bold="true" box="[895,992,664,685]" pageId="58" pageNumber="48">rhizomes</emphasis>
short-creeping;
<emphasis id="B96AEAF2FFD39B7164AF7FD1FB6EFD16" bold="true" box="[1156,1225,664,685]" pageId="58" pageNumber="48">fronds</emphasis>
once-pinnate, monomorphic, erect (
<taxonomicName id="4C1E4D63FFD39B7161907FFEFDEAFD77" box="[443,589,695,716]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B7161907FFEFDEAFD77" box="[443,589,695,716]" italics="true" pageId="58" pageNumber="48">H. sandwicensis</emphasis>
</taxonomicName>
) or pendent (
<taxonomicName id="4C1E4D63FFD39B7162F87FFEFC90FD77" authorityName=", Hobdy" authorityYear="2664" box="[723,823,695,716]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="pendens">
<emphasis id="B96AEAF2FFD39B7162F87FFEFC90FD77" box="[723,823,695,716]" italics="true" pageId="58" pageNumber="48">H. pendens</emphasis>
</taxonomicName>
);
<emphasis id="B96AEAF2FFD39B7163607FF1FC21FD76" bold="true" box="[843,902,696,717]" pageId="58" pageNumber="48">stipes</emphasis>
stramineous or dull brown, stipe bases and rhizome scales dull brown, glabrous or with short surficial hairs, ovate to linear-lanceolate;
<emphasis id="B96AEAF2FFD39B7164177F9EFBD9FD57" bold="true" box="[1084,1150,727,748]" pageId="58" pageNumber="48">blades</emphasis>
chartaceous, drying green, lanceolate to broadly deltate, apex gradually reduced, with distal pinnae somewhat decurrent, proximal pinnae not or only slightly reduced, basal pair sometimes somewhat deflexed, proliferous buds absent;
<emphasis id="B96AEAF2FFD39B7164A87E51FB6EFC96" bold="true" box="[1155,1225,792,813]" pageId="58" pageNumber="48">pinnae</emphasis>
crenate, dentate, shallowly lobed (&lt;halfway to costae), or subentire, typically straight, less commonly falcate, with acroscopic auricles, grooved adaxially;
<emphasis id="B96AEAF2FFD39B7161997E11FE40FCD6" bold="true" box="[434,487,856,877]" pageId="58" pageNumber="48">veins</emphasis>
prominent abaxially and adaxially, anastomosing with several pairs uniting below the sinus into a zig-zag excurrent vein, veins ending at pinna margins;
<emphasis id="B96AEAF2FFD39B7163787E3EFC63FC37" bold="true" box="[851,964,887,908]" pageId="58" pageNumber="48">aerophores</emphasis>
absent, or a small patch of darkened aerating tissue, sometimes slightly swollen or tuberculate;
<emphasis id="B96AEAF2FFD39B7162C07ED1FC16FC16" bold="true" box="[747,945,920,941]" pageId="58" pageNumber="48">indument abaxially</emphasis>
of broad-based (stout), tapering hyaline acicular hairs on veins and between veins, or restricted to veins,
<emphasis id="B96AEAF2FFD39B7163637EF1FBB7FC76" bold="true" box="[840,1040,952,973]" pageId="58" pageNumber="48">indument adaxially</emphasis>
of hyaline acicular hairs on and between veins, hairs on stipes and rachises short (&lt;
<quantity id="4CE69B05FFD39B7162E07E91FCA3FC56" box="[715,772,984,1005]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="58" pageNumber="48" unit="mm" value="1.0">1 mm</quantity>
) and sparse (
<taxonomicName id="4C1E4D63FFD39B7163AF7E9EFBB3FC50" box="[900,1044,982,1004]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B7163AF7E9EFBB3FC50" box="[900,1044,982,1004]" italics="true" pageId="58" pageNumber="48">H. sandwicensis</emphasis>
</taxonomicName>
), or long (&gt;
<quantity id="4CE69B05FFD39B7164A37E91FB65FC56" box="[1160,1218,984,1005]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="58" pageNumber="48" unit="mm" value="1.0">1 mm</quantity>
) and abundant (
<emphasis id="B96AEAF2FFD39B7160DA7EBEFEFAFBB0" box="[241,349,1014,1036]" italics="true" pageId="58" pageNumber="48">
<taxonomicName id="4C1E4D63FFD39B7160DA7EBEFEFEFBB0" authorityName=", Hobdy" authorityYear="2664" box="[241,345,1014,1036]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="pendens">H. pendens</taxonomicName>
,
</emphasis>
<figureCitation id="13252A65FFD39B71614B7EB1FE04FBB6" box="[352,419,1016,1037]" captionStart="FIG" captionStartId="24.[97,105,1752,1772]" captionTargetBox="[112,1215,163,1706]" captionTargetId="figure-8@24.[110,1215,160,1709]" captionTargetPageId="24" captionText="FIG.4.A.Hoiokula sandwicensis,Aborn s.n. (VT),pinna lobes,abaxial view.B.Hoiokula pendens,Hobdy2664 (UC),pinna lobes,adaxial view.C.Leptogrammapilosa, Diaz 6621 (UC), abaxial pinna lobes. D. Stegnogramma aspidioides, Palmer 1035 (UC), abaxial pinna. E. Stegnogramma wilfordii, Boufford 20159 (VT), proximal portionof lamina,abaxial view." figureDoi="http://doi.org/10.5281/zenodo.14111054" httpUri="https://zenodo.org/record/14111054/files/figure.png" pageId="58" pageNumber="48">
<emphasis id="B96AEAF2FFD39B71614B7EB1FE04FBB6" bold="true" box="[352,419,1016,1037]" pageId="58" pageNumber="48">Fig. 4B</emphasis>
</figureCitation>
), single-celled or septate; glands usually absent, but when present, spherical, translucent yellow-orange, on laminae, veins, and sporangia;
<emphasis id="B96AEAF2FFD39B716274795EFD11FB97" bold="true" box="[607,694,1047,1068]" pageId="58" pageNumber="48">pustules</emphasis>
absent on laminar tissue;
<emphasis id="B96AEAF2FFD39B7163817951FC75FB96" bold="true" box="[938,978,1048,1069]" pageId="58" pageNumber="48">sori</emphasis>
medial, round or elongate along veins, exindusiate;
<emphasis id="B96AEAF2FFD39B71617B7971FE12FBF6" bold="true" box="[336,437,1080,1101]" pageId="58" pageNumber="48">sporangia</emphasis>
abundantly to sparsely setulose (
<figureCitation id="13252A65FFD39B7162DB7971FC91FBF6" box="[752,822,1080,1101]" 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="58" pageNumber="48">
<emphasis id="B96AEAF2FFD39B7162DB7971FC91FBF6" bold="true" box="[752,822,1080,1101]" pageId="58" pageNumber="48">Fig. 2H</emphasis>
</figureCitation>
), rarely glabrous;
<emphasis id="B96AEAF2FFD39B7163CB797EFB85FBF7" bold="true" box="[992,1058,1079,1100]" pageId="58" pageNumber="48">spores</emphasis>
pale brown, with broad wings, in
<taxonomicName id="4C1E4D63FFD39B71612B791EFE31FBD0" box="[256,406,1110,1132]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B71612B791EFE31FBD0" box="[256,406,1110,1132]" italics="true" pageId="58" pageNumber="48">H. sandwicensis</emphasis>
</taxonomicName>
with secondary sculpturing of fimbriate microstructure (
<bibRefCitation id="EF8F4B11FFD39B7163E07911FB18FBD6" author="Tryon &amp; Lugardon" box="[971,1215,1112,1133]" firstAuthor="Tryon" journalOrPublisher="Springer-Verlag, New York, U. S. A" pageId="58" pageNumber="48" 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>
). Ploidy and hybrids are unknown, but the basic chromosome number is likely to be
<emphasis id="B96AEAF2FFD39B7163A8793EFC29FB37" box="[899,910,1143,1164]" italics="true" pageId="58" pageNumber="48">x</emphasis>
= 36, as is the case for its closest relatives.
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFD39B7160BA79FEFD2CFA57" pageId="58" pageNumber="48" type="diagnosis">
<paragraph id="8BA136E0FFD39B7160BA79FEFD2CFA57" blockId="58.[97,1226,599,1804]" pageId="58" pageNumber="48">
<emphasis id="B96AEAF2FFD39B7160BA79FEFF54FB77" box="[145,243,1207,1228]" italics="true" pageId="58" pageNumber="48">Diagnosis.</emphasis>
—In
<collectingRegion id="49DAF802FFD39B71610D79F1FEC9FB76" box="[294,366,1208,1229]" country="United States of America" name="Hawaii" pageId="58" pageNumber="48">Hawaii</collectingRegion>
,
<taxonomicName id="4C1E4D63FFD39B71615379FFFE6AFB70" box="[376,461,1206,1227]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71615379FFFE6AFB70" box="[376,461,1206,1227]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
is distinguished from
<taxonomicName id="4C1E4D63FFD39B71628679FFFC36FB70" box="[685,913,1206,1227]" class="Polypodiopsida" family="Thelypteridaceae" genus="Reholttumia" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="hudsoniana">
<emphasis id="B96AEAF2FFD39B71628679FFFC36FB70" box="[685,913,1206,1227]" italics="true" pageId="58" pageNumber="48">Reholttumia hudsoniana</emphasis>
</taxonomicName>
(formerly treated together with
<taxonomicName id="4C1E4D63FFD39B71604A799FFF12FB50" box="[97,181,1238,1259]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71604A799FFF12FB50" box="[97,181,1238,1259]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
in
<taxonomicName id="4C1E4D63FFD39B7160FE799EFEC1FB57" authorityName="sensu Holttum. In" authorityYear="1982" box="[213,358,1239,1260]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pneumatopteris" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B7160FE799EFEC1FB57" box="[213,358,1239,1260]" italics="true" pageId="58" pageNumber="48">Pneumatopteris</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF8F4B11FFD39B7161597991FDAFFB56" author="Holttum" box="[370,520,1240,1261]" firstAuthor="Holttum" journalOrPublisher="Allertonia" pageId="58" pageNumber="48" 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="EF8F4B11FFD39B7162397991FD2AFB56" author="Palmer" box="[530,653,1240,1261]" firstAuthor="Palmer" journalOrPublisher="University of Hawaii Press, Honolulu, U. S. A" pageId="58" pageNumber="48" 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>
)),
<taxonomicName id="4C1E4D63FFD39B71628E799FFD58FB50" box="[677,767,1238,1259]" class="Polypodiopsida" family="Thelypteridaceae" genus="Christella" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71628E799FFD58FB50" box="[677,767,1238,1259]" italics="true" pageId="58" pageNumber="48">Christella</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C1E4D63FFD39B71631D799EFC0DFB57" box="[822,938,1239,1260]" class="Polypodiopsida" family="Thelypteridaceae" genus="Menisciopsis" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71631D799EFC0DFB57" box="[822,938,1239,1260]" italics="true" pageId="58" pageNumber="48">Menisciopsis</emphasis>
</taxonomicName>
by having setulose sporangia and exindusiate sori.
<taxonomicName id="4C1E4D63FFD39B71610779BFFE27FAB0" box="[300,384,1270,1291]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71610779BFFE27FAB0" box="[300,384,1270,1291]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
is distinguished from
<taxonomicName id="4C1E4D63FFD39B71627F79BFFD74FAB0" box="[596,723,1270,1291]" class="Polypodiopsida" family="Thelypteridaceae" genus="Cyclogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71627F79BFFD74FAB0" box="[596,723,1270,1291]" italics="true" pageId="58" pageNumber="48">Cyclogramma</emphasis>
</taxonomicName>
by pinnae incised less than halfway to the costae (vs. more than half-way).Hamate hairs are absent in
<taxonomicName id="4C1E4D63FFD39B716201785FFDD9FA90" box="[554,638,1302,1323]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B716201785FFDD9FA90" box="[554,638,1302,1323]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
, but frequent in
<taxonomicName id="4C1E4D63FFD39B716331785FFC3EFA90" box="[794,921,1302,1323]" class="Polypodiopsida" family="Thelypteridaceae" genus="Cyclogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B716331785FFC3EFA90" box="[794,921,1302,1323]" italics="true" pageId="58" pageNumber="48">Cyclogramma</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1E4D63FFD39B7163EF785EFBEAFA97" box="[964,1101,1303,1324]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus" sensu="stricto">
<emphasis id="B96AEAF2FFD39B7163EF785EFBEAFA97" box="[964,1101,1303,1324]" italics="true" pageId="58" pageNumber="48">Stegnogramma</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2595789FFD39B716479785EFBC9FA97" box="[1106,1134,1303,1324]" pageId="58" pageNumber="48" sensu="stricto">
<emphasis id="B96AEAF2FFD39B716479785EFBC0FA97" box="[1106,1127,1303,1324]" italics="true" pageId="58" pageNumber="48">s.s</emphasis>
.
</taxonomicNameLabel>
Spores of most
<taxonomicName id="4C1E4D63FFD39B7160BF787EFEBAFAF7" box="[148,285,1335,1356]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B7160BF787EFEBAFAF7" box="[148,285,1335,1356]" italics="true" pageId="58" pageNumber="48">Stegnogramma</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1E4D63FFD39B716160787EFDB3FAF6" box="[331,532,1335,1357]" pageId="58" pageNumber="48">
<emphasis id="B96AEAF2FFD39B716160787EFE6CFAF7" box="[331,459,1335,1356]" italics="true" pageId="58" pageNumber="48">Leptogramma</emphasis>
species
</taxonomicName>
are echinate, rather than winged, (a feature shared by
<taxonomicName id="4C1E4D63FFD39B71643E787FFBCEFAF0" box="[1045,1129,1334,1355]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71643E787FFBCEFAF0" box="[1045,1129,1334,1355]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
and some species of
<taxonomicName id="4C1E4D63FFD39B7160EE781EFE9FFAD7" box="[197,312,1367,1388]" class="Polypodiopsida" family="Thelypteridaceae" genus="Menisciopsis" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B7160EE781EFE9FFAD7" box="[197,312,1367,1388]" italics="true" pageId="58" pageNumber="48">Menisciopsis</emphasis>
</taxonomicName>
).
<taxonomicName id="4C1E4D63FFD39B716167781FFE06FAD0" box="[332,417,1366,1387]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B716167781FFE06FAD0" box="[332,417,1366,1387]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
further differs from
<taxonomicName id="4C1E4D63FFD39B716247781EFD4AFAD7" box="[620,749,1367,1388]" class="Polypodiopsida" family="Thelypteridaceae" genus="Leptogramma" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B716247781EFD4AFAD7" box="[620,749,1367,1388]" italics="true" pageId="58" pageNumber="48">Leptogramma</emphasis>
</taxonomicName>
in having costae prominently grooved adaxially, vein endings reaching margins, multiple areoles below the sinuses (except in very small individuals), and presence of zig-zag excurrent veins. Until 2005, both species of
<taxonomicName id="4C1E4D63FFD39B71627578DFFD14FA10" box="[606,691,1430,1451]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71627578DFFD14FA10" box="[606,691,1430,1451]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
recognized herein were treated within
<taxonomicName id="4C1E4D63FFD39B71641D78DEFF7FFA70" class="Polypodiopsida" family="Thelypteridaceae" genus="Pneumatopteris" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B71641D78DEFF7FFA70" italics="true" pageId="58" pageNumber="48">Pneumatopteris sandwicensis</emphasis>
</taxonomicName>
;
<taxonomicName id="4C1E4D63FFD39B7160C878FFFE21FA70" authorityName=", Hobdy" authorityYear="2664" box="[227,390,1462,1483]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="pendens">
<emphasis id="B96AEAF2FFD39B7160C878FFFE21FA70" box="[227,390,1462,1483]" italics="true" pageId="58" pageNumber="48">Hoiokula pendens</emphasis>
</taxonomicName>
represents one of the most recently recognized native species in the Hawaiian fern flora (
<bibRefCitation id="EF8F4B11FFD39B71604C789EFF44FA57" author="Palmer" box="[103,227,1495,1516]" firstAuthor="Palmer" journalOrPublisher="Amer. Fern J." pageId="58" pageNumber="48" pagination="80 - 83" part="95" refId="ref112655" refString="PALMER, D. D. 2005. Pneumatopteris pendens (Thelypteridaceae), a new Hawaii endemic species of Pneumatopteris from Hawaii. Amer. Fern J. 95: 80 - 83." title="Pneumatopteris pendens (Thelypteridaceae), a new Hawaii endemic species of Pneumatopteris from Hawaii" type="journal article" year="2005">Palmer 2005</bibRefCitation>
;
<bibRefCitation id="EF8F4B11FFD39B7160C0789EFE6BFA57" author="Vernon &amp; Ranker" box="[235,460,1495,1516]" firstAuthor="Vernon" journalOrPublisher="Amer. Fern J." pageId="58" pageNumber="48" pagination="59 - 111" part="103" refId="ref115393" refString="VERNON, A. L. &amp; T. A. RANKER. 2013. Current status of the ferns and lycophytes of the Hawaiian Islands. Amer. Fern J. 103: 59 - 111." title="Current status of the ferns and lycophytes of the Hawaiian Islands" type="journal article" year="2013">Vernon &amp; Ranker 2013</bibRefCitation>
;
<bibRefCitation id="EF8F4B11FFD39B7161F8789EFD27FA57" author="Ranker" box="[467,640,1495,1516]" etAl="et al." firstAuthor="Ranker" journalOrPublisher="Amer. Fern J." pageId="58" pageNumber="48" pagination="54 - 72" part="109" refId="ref113269" refString="RANKER, T. A., C. T. IMADA, K. LYNCH, D. D. PALMER, A. L. VERNON, &amp; M. K. THOMAS. 2019. Taxonomic and nomenclatural updates to the fern and lycophyte flora of the Hawaiian Islands. Amer. Fern J. 109: 54 - 72." title="Taxonomic and nomenclatural updates to the fern and lycophyte flora of the Hawaiian Islands" type="journal article" year="2019">Ranker et al. 2019</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFD39B7160BA78BFFEC9F9D7" pageId="58" pageNumber="48" type="biology_ecology">
<paragraph id="8BA136E0FFD39B7160BA78BFFEC9F9D7" blockId="58.[97,1226,599,1804]" pageId="58" pageNumber="48">
<emphasis id="B96AEAF2FFD39B7160BA78BFFE49F9B0" box="[145,494,1526,1547]" italics="true" pageId="58" pageNumber="48">
Biogeography and ecology.—
<taxonomicName id="4C1E4D63FFD39B7161B278BFFE49F9B0" box="[409,494,1526,1547]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">Hoiokula</taxonomicName>
</emphasis>
is endemic to the Hawaiian Islands, distributed on all
<taxonomicName id="4C1E4D63FFD39B7163DD78BEFB89F9B7" box="[1014,1070,1527,1548]" genus="The" pageId="58" pageNumber="48" rank="species" species="major">major</taxonomicName>
islands (
<bibRefCitation id="EF8F4B11FFD39B7164A878BEFF34F997" author="Palmer" firstAuthor="Palmer" journalOrPublisher="University of Hawaii Press, Honolulu, U. S. A" pageId="58" pageNumber="48" 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>
,
<bibRefCitation id="EF8F4B11FFD39B7160B77B5EFF68F997" author="Palmer" box="[156,207,1559,1580]" firstAuthor="Palmer" journalOrPublisher="Amer. Fern J." pageId="58" pageNumber="48" pagination="80 - 83" part="95" refId="ref112655" refString="PALMER, D. D. 2005. Pneumatopteris pendens (Thelypteridaceae), a new Hawaii endemic species of Pneumatopteris from Hawaii. Amer. Fern J. 95: 80 - 83." title="Pneumatopteris pendens (Thelypteridaceae), a new Hawaii endemic species of Pneumatopteris from Hawaii" type="journal article" year="2005">2005</bibRefCitation>
). The two recognized species differ in habit and habitat, with the larger, erect
<taxonomicName id="4C1E4D63FFD39B7163937B5EFBECF990" box="[952,1099,1558,1580]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pneumatopteris" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="sandwicensis">
<emphasis id="B96AEAF2FFD39B7163937B5EFBECF990" box="[952,1099,1558,1580]" italics="true" pageId="58" pageNumber="48">H. sandwicensis</emphasis>
</taxonomicName>
occurring in forest understories and along stream margins, and the pendent
<taxonomicName id="4C1E4D63FFD39B7162E37B7EFC89F9F0" authorityName=", Hobdy" authorityYear="2664" box="[712,814,1590,1612]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="species" species="pendens">
<emphasis id="B96AEAF2FFD39B7162E37B7EFC89F9F0" box="[712,814,1590,1612]" italics="true" pageId="58" pageNumber="48">H. pendens</emphasis>
</taxonomicName>
occurring on damp rocks and cliffs, often near streams (
<bibRefCitation id="EF8F4B11FFD39B7160CD7B1EFEC4F9D7" author="Palmer" box="[230,355,1623,1644]" firstAuthor="Palmer" journalOrPublisher="Amer. Fern J." pageId="58" pageNumber="48" pagination="80 - 83" part="95" refId="ref112655" refString="PALMER, D. D. 2005. Pneumatopteris pendens (Thelypteridaceae), a new Hawaii endemic species of Pneumatopteris from Hawaii. Amer. Fern J. 95: 80 - 83." title="Pneumatopteris pendens (Thelypteridaceae), a new Hawaii endemic species of Pneumatopteris from Hawaii" type="journal article" year="2005">Palmer 2005</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C304656BFFD39B7060BA7B3EFC0BFE4C" lastPageId="59" lastPageNumber="49" pageId="58" pageNumber="48" type="discussion">
<paragraph id="8BA136E0FFD39B7060BA7B3EFC75FF6C" blockId="58.[97,1226,599,1804]" lastBlockId="59.[121,1249,162,503]" lastPageId="59" lastPageNumber="49" pageId="58" pageNumber="48">
<emphasis id="B96AEAF2FFD39B7160BA7B3EFDEEF930" box="[145,585,1654,1676]" italics="true" pageId="58" pageNumber="48">
Taxonomic and phylogenetic studies.—
<taxonomicName id="4C1E4D63FFD39B7161DE7B3FFDEEF930" box="[501,585,1654,1675]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">Hoiokula</taxonomicName>
</emphasis>
bears considerable resemblance to
<taxonomicName id="4C1E4D63FFD39B7163B27B3EFB85F937" box="[921,1058,1655,1676]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus" sensu="lato">
<emphasis id="B96AEAF2FFD39B7163B27B3EFB85F937" box="[921,1058,1655,1676]" italics="true" pageId="58" pageNumber="48">Stegnogramma</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2595789FFD39B71640C7B3FFBE7F937" box="[1063,1088,1654,1676]" pageId="58" pageNumber="48" sensu="lato">
<emphasis id="B96AEAF2FFD39B71640C7B3FFB9CF930" box="[1063,1083,1654,1675]" italics="true" pageId="58" pageNumber="48">s.l</emphasis>
.
</taxonomicNameLabel>
, and the similarities must have been apparent to
<bibRefCitation id="EF8F4B11FFD39B7161967BDEFD25F917" author="Brackenridge" box="[445,642,1687,1708]" firstAuthor="Brackenridge" journalOrPublisher="Botany. Cryptogamia. Filices, including Lycopodiaceae and Hydropterides" pageId="58" pageNumber="48" part="16" refId="ref106659" refString="BRACKENRIDGE, W. D. 1854. United Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U. S. N. Botany. Cryptogamia. Filices, including Lycopodiaceae and Hydropterides Vol. 16. C. Sherman, Philadelphia, PA, U. S. A." title="United Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U. S. N." type="journal volume" year="1854">Brackenridge (1854)</bibRefCitation>
, who described the
<typeStatus id="54A58842FFD39B71636D7BDEFCD6F917" box="[838,881,1687,1708]" pageId="58" pageNumber="48">type</typeStatus>
species in
<taxonomicName id="4C1E4D63FFD39B7163F77BDEFBC1F917" box="[988,1126,1687,1708]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B7163F77BDEFBC1F917" box="[988,1126,1687,1708]" italics="true" pageId="58" pageNumber="48">Stegnogramma</emphasis>
</taxonomicName>
and illustrated the elongate sori and setose sporangia in the protologue. Both of these features are shared by the three stegnogrammoid genera but are uncommon within
<taxonomicName id="4C1E4D63FFD39B7162777B9EFCA1F957" baseAuthorityName="Holttum" baseAuthorityYear="1976" box="[604,774,1751,1772]" class="Polypodiopsida" family="Thelypteridaceae" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="family">Thelypteridaceae</taxonomicName>
. Although
<bibRefCitation id="EF8F4B11FFD39B71635A7B9EFBB1F957" author="Holttum" box="[881,1046,1751,1772]" firstAuthor="Holttum" journalOrPublisher="Allertonia" pageId="58" pageNumber="48" 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>
treated the plants recognized here in
<taxonomicName id="4C1E4D63FFD39B71613E7BBFFECEF8B0" box="[277,361,1782,1803]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B71613E7BBFFECEF8B0" box="[277,361,1782,1803]" italics="true" pageId="58" pageNumber="48">Hoiokula</emphasis>
</taxonomicName>
as a species of
<taxonomicName id="4C1E4D63FFD39B7161DF7BBEFD23F8B7" authorityName="sensu Holttum. In" authorityYear="1982" box="[500,644,1783,1804]" class="Polypodiopsida" family="Thelypteridaceae" genus="Pneumatopteris" kingdom="Plantae" order="Polypodiales" pageId="58" pageNumber="48" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD39B7161DF7BBEFD23F8B7" box="[500,644,1783,1804]" italics="true" pageId="58" pageNumber="48">Pneumatopteris</emphasis>
</taxonomicName>
, he noted several characteristics that this “peculiar Hawaiian species” has in common with
<taxonomicName id="4C1E4D63FFD29B7061B67DEBFD81FF0C" box="[413,550,162,183]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B7061B67DEBFD81FF0C" box="[413,550,162,183]" italics="true" pageId="59" pageNumber="49">Stegnogramma</emphasis>
</taxonomicName>
, such as “…venation of the apical lamina and a few thick hairs between veins on the upper surface of pinnae, and … somewhat elongate sori” (
<bibRefCitation id="EF8F4B11FFD29B7063387D8BFC6FFF6C" author="Holttum" box="[787,968,194,215]" firstAuthor="Holttum" journalOrPublisher="Flora Malesiana" page="540" pageId="59" pageNumber="49" 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">Holttum 1982:540</bibRefCitation>
).
</paragraph>
<paragraph id="8BA136E0FFD29B70608C7DABFC0BFE4C" blockId="59.[121,1249,162,503]" pageId="59" pageNumber="49">
Based on both concatenated and coalescent analyses (Fawcett et al. in press), the phylogenetic position of
<taxonomicName id="4C1E4D63FFD29B7064A77DA8FB47FF4D" box="[1164,1248,225,246]" class="Polypodiopsida" family="Thelypteridaceae" genus="Hoiokula" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B7064A77DA8FB47FF4D" box="[1164,1248,225,246]" italics="true" pageId="59" pageNumber="49">Hoiokula</emphasis>
</taxonomicName>
is well-supported as sister to the stegnogrammoid ferns, which include the genera
<taxonomicName id="4C1E4D63FFD29B7063B07C48FBBEFEAD" box="[923,1049,257,278]" class="Polypodiopsida" family="Thelypteridaceae" genus="Cyclogramma" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B7063B07C48FBBEFEAD" box="[923,1049,257,278]" italics="true" pageId="59" pageNumber="49">Cyclogramma</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1E4D63FFD29B70640D7C4BFB08FEAC" box="[1062,1199,258,279]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B70640D7C4BFB08FEAC" box="[1062,1199,258,279]" italics="true" pageId="59" pageNumber="49">Stegnogramma</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C1E4D63FFD29B7060527C6BFE3CFE8C" authority="(Kuo et al. 2019)" baseAuthorityName="Kuo" baseAuthorityYear="2019" box="[121,411,290,311]" class="Polypodiopsida" family="Thelypteridaceae" genus="Leptogramma" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B7060527C6BFF5EFE8C" box="[121,249,290,311]" italics="true" pageId="59" pageNumber="49">Leptogramma</emphasis>
(
<bibRefCitation id="EF8F4B11FFD29B70612F7C6BFE32FE8C" author="Kuo" box="[260,405,290,311]" etAl="et al." firstAuthor="Kuo" journalOrPublisher="Cladistics doi. org" pageId="59" pageNumber="49" refId="ref110871" refString="KUO, L. Y., Y. H. CHANG, Y. H. HUANG, W. TESTO, A. EBIHARA, G. ROUHAN, L. G. QUINTANILLA, J. E. WATKINS, Y. M. HUANG, &amp; F. W. LI. 2019. A global phylogeny of Stegnogramma ferns (Thelypteridaceae): generic and sectional revision, historical biogeography and evolution of leaf architecture. Cladistics doi. org / 10.1111 / cla. 12399" title="A global phylogeny of Stegnogramma ferns (Thelypteridaceae): generic and sectional revision, historical biogeography and evolution of leaf architecture" type="book" year="2019">Kuo et al. 2019</bibRefCitation>
)
</taxonomicName>
. These results should be interpreted with caution, however, since the analysis assumes a bifurcating tree. Preliminary evidence from plastid data (L.
<collectionCode id="ED0FAE25FFD29B7062887C0BFD13FEEC" box="[675,692,322,343]" country="USA" lsid="urn:lsid:biocol.org:col:14792" name="Yale University" pageId="59" pageNumber="49" type="Herbarium">Y</collectionCode>
. Kuo, unpubl. data), and conflicting topologies among gene trees (Fawcett et al. in press) suggest the possibility that this lineage may be of hybrid origin involving
<taxonomicName id="4C1E4D63FFD29B7064137C2BFB1FFECC" box="[1080,1208,354,375]" class="Polypodiopsida" family="Thelypteridaceae" genus="Leptogramma" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B7064137C2BFB1FFECC" box="[1080,1208,354,375]" italics="true" pageId="59" pageNumber="49">Leptogramma</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1E4D63FFD29B7060527CCBFF4AFE2C" box="[121,237,386,407]" class="Polypodiopsida" family="Thelypteridaceae" genus="Menisciopsis" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B7060527CCBFF4AFE2C" box="[121,237,386,407]" italics="true" pageId="59" pageNumber="49">Menisciopsis</emphasis>
</taxonomicName>
, which occurs in Hawaii, Melanesia, and eastern Asia.
<taxonomicName id="4C1E4D63FFD29B7063227CCBFC2DFE2C" box="[777,906,386,407]" class="Polypodiopsida" family="Thelypteridaceae" genus="Leptogramma" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B7063227CCBFC2DFE2C" box="[777,906,386,407]" italics="true" pageId="59" pageNumber="49">Leptogramma</emphasis>
</taxonomicName>
is distributed throughout Asia and
<collectingCountry id="F3097670FFD29B7060527CEBFF09FE0C" box="[121,174,418,439]" name="India" pageId="59" pageNumber="49">India</collectingCountry>
, and is scattered in Africa, Europe, and North America, while the genera
<taxonomicName id="4C1E4D63FFD29B70634C7CE8FC41FE0D" box="[871,998,417,438]" class="Polypodiopsida" family="Thelypteridaceae" genus="Cyclogramma" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus">
<emphasis id="B96AEAF2FFD29B70634C7CE8FC41FE0D" box="[871,998,417,438]" italics="true" pageId="59" pageNumber="49">Cyclogramma</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1E4D63FFD29B70643F7CEBFB3AFE0C" box="[1044,1181,418,439]" class="Polypodiopsida" family="Thelypteridaceae" genus="Stegnogramma" kingdom="Plantae" order="Polypodiales" pageId="59" pageNumber="49" phylum="Tracheophyta" rank="genus" sensu="stricto">
<emphasis id="B96AEAF2FFD29B70643F7CEBFB3AFE0C" box="[1044,1181,418,439]" italics="true" pageId="59" pageNumber="49">Stegnogramma</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2595789FFD29B7064897CEBFB19FE0C" box="[1186,1214,418,439]" pageId="59" pageNumber="49" sensu="stricto">
<emphasis id="B96AEAF2FFD29B7064897CEBFB10FE0C" box="[1186,1207,418,439]" italics="true" pageId="59" pageNumber="49">s.s</emphasis>
.
</taxonomicNameLabel>
are both restricted to Southeast Asia and
<collectingCountry id="F3097670FFD29B7061C77C8BFD9DFE6C" box="[492,570,450,471]" name="Malaysia" pageId="59" pageNumber="49">Malesia</collectingCountry>
. No species of these three genera have been collected in Hawaii. The hybrid origin hypothesis is currently under investigation (Fawcett, Kuo et al. in prep.).
</paragraph>
</subSubSection>
</treatment>
</document>
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