treatments-xml/data/F7/B0/35/F7B035B5CAA150E9B59B166914AA2CF5.xml

<document id="B52E1EFDCCCF088F5C229BF02021D59A" ID-DOI="10.3897/phytokeys.244.124080" ID-publisher-id="124080" URI-arpha="BF915544-99E3-52A0-B16F-2DB1AFAE0098" article-type="review-article" checkinTime="1721762591914" checkinUser="pensoft" docAuthor="Suwanmala, Orawanya, Villarreal A., Juan Carlos, Li, Fay-Wei &amp; Chantanaorrapint, Sahut" docDate="2024" docId="F7B035B5CAA150E9B59B166914AA2CF5" docLanguage="en" docName="PhytoKeys 244: 271-283" docOrigin="PhytoKeys 244" docSource="https://phytokeys.pensoft.net/article/124080/download/xml/" docStyle="DocumentStyle:PensoftTaxPub.0000.journal_article.phytokeys" docStyleName="PensoftTaxPub.0000.journal_article.phytokeys" docTitle="Phaeoceros perpusillus var. scabrellus Suwanmala &amp; Chantanaorr. 2024, var. nov." docType="treatment" docVersion="1" dtd-version="3.0" lastPageNumber="283" masterDocId="BF91554499E352A0B16F2DB1AFAE0098" masterDocTitle="Phaeoceros perpusillus var. scabrellus (Notothyladaceae, Anthocerotophyta), a new taxon from northern Thailand" masterLastPageNumber="283" masterPageNumber="271" pageNumber="271" updateTime="1721762658523" updateUser="pensoft">
<mods:mods id="F43C55DA0D4855DC0C35D65553609C55" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="A96495A2266FA26C2830C175C3782CA2">
<mods:title id="58E120DC3416042911AA5464CF5D2C37">Phaeoceros perpusillus var. scabrellus (Notothyladaceae, Anthocerotophyta), a new taxon from northern Thailand</mods:title>
</mods:titleInfo>
<mods:name id="33B44CE9D9B9E9E69689FA6513D2A087" type="personal">
<mods:role id="A4205F35C46C7D53ADBC1F8D7A466A1E">
<mods:roleTerm id="3FA0909D4415999F64E53B90BD697717">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="E690C612128EB949EEA988F1FAC19343">Suwanmala, Orawanya</mods:namePart>
<mods:nameIdentifier id="02C99DD5CAA667B33EAB6B10C7B74ABE" type="ORCID">0000-0002-1113-7614</mods:nameIdentifier>
<mods:affiliation id="E7E248DE241DF5538560A1B4C7FF69EE">PSU Herbarium, Division of Biological Science, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90110, Thailand</mods:affiliation>
</mods:name>
<mods:name id="50DE4834C50C67B02B41CD1FF2534435" type="personal">
<mods:role id="D8648E525C6AE08EA00571BC87D110D7">
<mods:roleTerm id="11E8B4F0C48801DC6FF49EFC41855BD3">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="9B4BEFB31FBC2655967698F35EF0BEAF">Villarreal A., Juan Carlos</mods:namePart>
<mods:nameIdentifier id="17E024A03C9FA0F0F26B68C90F85D385" type="ORCID">0000-0002-0770-1446</mods:nameIdentifier>
<mods:affiliation id="86DE1B651286978215F3040B287F9AC1">Département de Biologie, Pavillon C. - E. Marchand Université Laval, Québec, Canada</mods:affiliation>
</mods:name>
<mods:name id="D0A304C5D22A773168B5CF7637795A65" type="personal">
<mods:role id="B768B3BBD997739FB8CE9DCAC9BB53C2">
<mods:roleTerm id="912EC223D5E1FBEBE3EAEFAE2F1B8AFC">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="520D8308803A92EF373C25D516E241C2">Li, Fay-Wei</mods:namePart>
<mods:nameIdentifier id="773C14D2AF550253825B8AC3241FB520" type="ORCID">0000-0002-0076-0152</mods:nameIdentifier>
<mods:affiliation id="54651DFCE94C72669E2C9602FE4D243D">Boyce Thompson Institute, Ithaca, New York, USA &amp; Plant Biology Section, Cornell University, Ithaca, New York, USA</mods:affiliation>
</mods:name>
<mods:name id="1E2BBF1A3D5E8ACD565F8EA17244247C" type="personal">
<mods:role id="CDF00D3DF045627D393CBF32EEC0CD54">
<mods:roleTerm id="A43EF81470FEAF92CEABC1CA071C54BD">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="A065D74B1F3559EF1CC106A1ADE94B2A">Chantanaorrapint, Sahut</mods:namePart>
<mods:nameIdentifier id="BD31FB0BA7BD4DC22D429C4F888CC458" type="ORCID">0000-0002-9739-0994</mods:nameIdentifier>
<mods:affiliation id="23C983A01EB82D577CDA6CCF14196065">PSU Herbarium, Division of Biological Science, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90110, Thailand</mods:affiliation>
</mods:name>
<mods:typeOfResource id="CF1A74A08A6D4BD1D9FB96CE52F95706">text</mods:typeOfResource>
<mods:relatedItem id="4B61701D44D62CB33EA746D9622D5CCF" type="host">
<mods:titleInfo id="56ACC6BA75719E5C9D97787AA42C78D3">
<mods:title id="287991B5E7E267CC7081536226ADA720">PhytoKeys</mods:title>
</mods:titleInfo>
<mods:part id="0FBE5C30AF696C65126C6D1C87EC72D9">
<mods:date id="E7FB795B5A411A7F965813FBE72088EA">2024</mods:date>
<mods:detail id="9F33853DF35FF3EED7A9EC275B4AA2A2" type="pubDate">
<mods:number id="A7DDB63CB43401EEF71A9414028D0DAA">2024-07-23</mods:number>
</mods:detail>
<mods:detail id="BB2F95B713610FFC5425B9BEFBA28525" type="volume">
<mods:number id="C15F6E33D0849B95582D066A823B05F8">244</mods:number>
</mods:detail>
<mods:extent id="0EA23E08B88C80F615C98AD1F64DC672" unit="page">
<mods:start id="6523458621328F3AD408E1FDCD9C7FE6">271</mods:start>
<mods:end id="552757F1859558791A8C152D4E0BC6F3">283</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification id="A616EEE84110C4D1B8D2678B76734575">journal article</mods:classification>
<mods:identifier id="6C35A00520BF29799002949BAB1F4766" type="DOI">10.3897/phytokeys.244.124080</mods:identifier>
</mods:mods>
<treatment id="F7B035B5CAA150E9B59B166914AA2CF5" ID-arpha="F7B035B5-CAA1-50E9-B59B-166914AA2CF5" LSID="urn:lsid:plazi:treatment:F7B035B5CAA150E9B59B166914AA2CF5" httpUri="http://treatment.plazi.org/id/F7B035B5CAA150E9B59B166914AA2CF5">
<subSubSection id="23B07614DE515A2F84054E4C7FC7D14C" type="nomenclature">
<paragraph id="584FD6B4092F6A628835A0E6744B392F">
<taxonomicName id="5234B669CE2660563B7C179A036D97C0" authority="Suwanmala &amp; Chantanaorr." authorityName="Suwanmala &amp; Chantanaorr." authorityYear="2024" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" status="var. nov." variety="scabrellus">
<emphasis id="EADC3E4FAE42C84347B20A08F1D90DA1" italics="true">Phaeoceros perpusillus</emphasis>
Chantanaorr. 
<emphasis id="D6D65ECB510D11DFB19ED88794B7F748" italics="true">var. scabrellus</emphasis>
Suwanmala &amp; Chantanaorr.
</taxonomicName>
<taxonomicNameLabel id="8F9A1D92DDE0738FB7B576E7E0631258" rank="variety">var. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="E12ADFA557CEF4822EA33B0B498D33E7" type="description">
<paragraph id="9A18430414D3D184CF60A6674247B670">
<figureCitation id="BD7B6B43C613683309DC31012D48E52E" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Phaeoceros perpusillus var. scabrellus A plants in natural habitat B dorsal view of thallus showing marginal tubers (arrow) C ventral view of thallus showing ventral tubers (arrows) D cross-section of thallus showing the large dark lump of Nostoc colony (NC = Nostoc colony) E dorsal epidermal cells of thallus showing a single chloroplast with pyrenoid (arrows) per cell F spores and pseudoelaters. Photos by O. Suwanmala (A from S. Chantanaorrapint &amp; O. Suwanmala 4116 B – F from S. Chantanaorrapint &amp; O. Suwanmala 4077)." figureDoi="10.3897/phytokeys.244.124080.figure2" httpUri="https://binary.pensoft.net/fig/1098338">Figs 2</figureCitation>
, 
<figureCitation id="50107F3D2EDBB4449201361CB9D9829B" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Phaeoceros perpusillus var. scabrellus A gametophyte forming half-rosettes with sporophyte (arrow indicate tuber) B ensiform thalli and sporophytes C gametophyte showing ventral tuber (arrow) D dorsal epidermal cells of thallus E, F cross sections of thalli G cross section of sporangium (AS = assimilative tissue, EP = epidermal cell of capsule, IN = inner most sporangium wall) H inner most cells of sporangium wall I epidermal cells of capsule with stoma J proximal view of spore K distal view of spore L pseudoelaters. All from holotype and drawings by O. Suwanmala. (All drawing from S. Chantanaorrapint &amp; O. Suwanmala 4116)." figureDoi="10.3897/phytokeys.244.124080.figure3" httpUri="https://binary.pensoft.net/fig/1098339">3</figureCitation>
, 
<figureCitation id="D4B3573D9071A276C0C38C4C97C68E97" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Scanning electron micrographs of spores A – F Phaeoceros perpusillus var. scabrellus A distal view of spore showing a central hump-like projection B proximal view of spore with a distinct triradiate mark C close-up of distal surface showing packed vermiculae D close-up of proximal surface showing loosely arranged vermiculae E proximal surface showing trilete mark and loosely arranged vermiculae F spores and pseudoelaters G – I P. perpusillus var. perpusillus G distal view of spore H proximal view of spore I close-up of distal surface showing pluripapillae. Photos by O. Suwanmala. (A – F images from S. Chantanaorrapint &amp; C. Promma 3129 G – I images from S. Chantanaorrapint &amp; O. Suwanmala 3883)." figureDoi="10.3897/phytokeys.244.124080.figure4" httpUri="https://binary.pensoft.net/fig/1098340">4</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EYOAG" type="material">
<paragraph id="12503E95812F149F1F90D8F799A30405">
<heading id="D43CEE38A2E452DAE450403AA3485395" reason="title">
<typeStatus id="C4F5461CC035E57A700CE72A4BC9A9C1">Type</typeStatus>
.
</heading>
</paragraph>
<paragraph id="FBCD78C6275857084D215A7EAB68F3E8">
<collectingCountry id="30F3EB22B4224E79EE3F95979BB21549" name="Thailand">Thailand</collectingCountry>
. 
<collectingRegion id="311606A785E4373ACF7592F447F401CE" country="Thailand" name="Chiang Mai">Chiang Mai Province</collectingRegion>
: Doi Suthep-Pui, Bhu Bing Palace, 
<quantity id="8679D057330DEC53E9ED21057C3BCE64" metricMagnitude="3" metricUnit="m" metricValue="1.4" unit="m" value="1400.0">1400 m</quantity>
, 
<date id="0B2CD0409BCA19F90BCEF3A0E0F73A43" value="2020-10-18">18 October 2020</date>
, 
<emphasis id="517ADF7E97A4B2708D772847C0EDEBD5" italics="true">
<collectionCode id="4B0AC8ABABDADDA6ED7BF38852765F37" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" type="Herbarium">S</collectionCode>
. Chantanaorrapint &amp; 
<collectionCode id="F6E07311CB37CC70ECF78D7A2A29D1D3" country="Norway" lsid="urn:lsid:biocol.org:col:13093" name="Botanical Museum - University of Oslo" type="Herbarium">O</collectionCode>
. Suwanmala 4077
</emphasis>
(
<typeStatus id="58C75E2B9E7BB9A395A810215BA2AF9C" type="holotype">holotype</typeStatus>
: 
<collectionCode id="E6386445BBD330A717956AC40D70A000" country="USA" httpUri="http://grbio.org/cool/5mg9-yyc1" name="Portland State University, Vertebrate Biology Museum" type="Museum">PSU</collectionCode>
!; 
<typeStatus id="971AF198C9A6F2BAD55DC750EC7A52E2" type="isotype">isotype</typeStatus>
: 
<collectionCode id="3D8A418A52D0A14CFFA395178E575754" country="Thailand" httpUri="http://grbio.org/cool/d4ij-e7pz" name="National Park, Wildlife and Plant Conservation Department" type="Herbarium">BKF</collectionCode>
!, 
<collectionCode id="2B3E49044C1BDE13CDA2ACE4D3469519" country="Japan" httpUri="http://biocol.org/urn:lsid:biocol.org:col:13202" lsid="urn:lsid:biocol.org:col:13202" name="Hattori Botanical Laboratory" type="Herbarium">NICH</collectionCode>
!).
</paragraph>
<caption id="8F1746E4179CB1EDE15F20B73DBB0616" ID-DOI="10.3897/phytokeys.244.124080.figure2" ID-arpha="8805D135-7231-51CF-8C58-EBBC04845C3E" httpUri="https://binary.pensoft.net/fig/1098338" startId="F2">
<paragraph id="149A9759E3F7ED3ECC40A63A5926DDC3">
<label id="4A699E4FE2C457CA33C03E3C7ED4446E">Figure 2.</label>
</paragraph>
<paragraph id="682585E57CCC08CC612588D3F3AE7A95">
<taxonomicName id="A7B5F3D715615C3E00088E3A7B614EA6" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" variety="scabrellus">
<emphasis id="62D1ACBE9833CFEF2A196F1FD139BA37" italics="true">Phaeoceros perpusillus var. scabrellus</emphasis>
</taxonomicName>
<emphasis id="26381E29B91992A7B4CA5C17E9E6A537" bold="true">A</emphasis>
plants in natural habitat 
<emphasis id="D2985D55CD0555100EE8000356FAEFBD" bold="true">B</emphasis>
dorsal view of thallus showing marginal tubers (arrow) 
<emphasis id="81CE0DC80ADFA7DD2A2DECA177760887" bold="true">C</emphasis>
ventral view of thallus showing ventral tubers (arrows) 
<emphasis id="E2CB99F222C5F5167C4BA788686F657A" bold="true">D</emphasis>
cross-section of thallus showing the large dark lump of 
<taxonomicName id="423A2C01C45CCEFA8436EB7847E59D56" authorityName="Vaucher" authorityYear="1888" class="Cyanophyceae" family="Nostocaceae" genus="Nostoc" kingdom="Bacteria" order="Nostocales" phylum="Cyanobacteria" rank="genus">
<emphasis id="9A893F49344FD1F0A5F9C40A8C15B755" italics="true">Nostoc</emphasis>
</taxonomicName>
colony (NC = 
<taxonomicName id="925E53E2BD7CDB14BC4316F868BD9845" authorityName="Vaucher" authorityYear="1888" class="Cyanophyceae" family="Nostocaceae" genus="Nostoc" kingdom="Bacteria" order="Nostocales" phylum="Cyanobacteria" rank="genus">
<emphasis id="CEE3995D568A33D654E95CB775779012" italics="true">Nostoc</emphasis>
</taxonomicName>
colony) 
<emphasis id="7E0E53184ADC631DF9BD1422F8E70AD0" bold="true">E</emphasis>
dorsal epidermal cells of thallus showing a single chloroplast with pyrenoid (arrows) per cell 
<emphasis id="738E6AADAFCE368651262BE944CB2E1F" bold="true">F</emphasis>
spores and pseudoelaters. Photos by O. Suwanmala (
<emphasis id="4C905B94173E3B7A72DA3D166425F7CF" bold="true">A</emphasis>
from 
<emphasis id="B9B47DF1FAF7B0E04B25A04874559CFC" italics="true">S. Chantanaorrapint &amp; O. Suwanmala 4116</emphasis>
<emphasis id="D90635A0D5869FFF63A8ACC6F1721533" bold="true">B – F</emphasis>
from 
<emphasis id="F9E55AB02817D669772A9F2D64326360" italics="true">S. Chantanaorrapint &amp; O. Suwanmala 4077</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0E5QAG" type="Diagnostic">
<paragraph id="41CF2D1FFC5C789CD93FF1AA8726987A">
<heading id="D79765984249EF720A3AF4D3D1747756" reason="title">Diagnostic.</heading>
</paragraph>
<paragraph id="0F932C9B0694CAA7F47706B9BD2DA08A">
<taxonomicName id="0FE5FE75DDCE7D87BAE5DA8CB6F8F62A" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" variety="scabrellus">
<emphasis id="7A2F8FD1951157DA177932A3D9EED5E8" italics="true">Phaeoceros perpusillus var. scabrellus</emphasis>
</taxonomicName>
is similar to the autonimic variety but differs in nearly smooth spores under light microscope (or vermiculate under 
<collectionCode id="58238A63A19B908AFF07C38BC3518A65">SEM</collectionCode>
), whereas the autonimic variety have pluripapillae on the distal surface and vermiculate on the proximal.
</paragraph>
<caption id="F30C3F2DB0CDC42D95883087BDB5866F" ID-DOI="10.3897/phytokeys.244.124080.figure3" ID-arpha="59F96907-38DD-512C-ACD1-6E65BDB77EC0" httpUri="https://binary.pensoft.net/fig/1098339" startId="F3">
<paragraph id="6031264985B7CE7F5F9661EAF192A052">
<label id="0FB063A000CC45CE2EF48C771BDC202A">Figure 3.</label>
</paragraph>
<paragraph id="196E5BEE2FAC0ADBC705D287031223FE">
<taxonomicName id="C017AD8C0D3227F252C56478E64B8DC4" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" variety="scabrellus">
<emphasis id="05D76D71817A9203834CB0244535E078" italics="true">Phaeoceros perpusillus var. scabrellus</emphasis>
</taxonomicName>
<emphasis id="08BF345246953ECDA074AD36E871D774" bold="true">A</emphasis>
gametophyte forming half-rosettes with sporophyte (arrow indicate tuber) 
<emphasis id="6843FBB1CA59D35D21D70DEB6ED9D935" bold="true">B</emphasis>
ensiform thalli and sporophytes 
<emphasis id="3F30649CE828563316DFEA8B2EB48152" bold="true">C</emphasis>
gametophyte showing ventral tuber (arrow) 
<emphasis id="E3C2AEC974003B5A8C58259ECFFD88BF" bold="true">D</emphasis>
dorsal epidermal cells of thallus 
<emphasis id="C7C224EA675554EFB8AEF705F7799738" bold="true">E, F</emphasis>
cross sections of thalli 
<emphasis id="8F3AF8861F128FB270663D01F5BB1B7C" bold="true">G</emphasis>
cross section of sporangium (AS = assimilative tissue, EP = epidermal cell of capsule, IN = inner most sporangium wall) 
<emphasis id="6EC3043E29A15DA0C8695C3520FF43A9" bold="true">H</emphasis>
inner most cells of sporangium wall 
<emphasis id="7AD11B14F27E590A828C6A1D8994E62F" bold="true">I</emphasis>
epidermal cells of capsule with stoma 
<emphasis id="19E834278CE77A43AA6C0A8D03B78AA9" bold="true">J</emphasis>
proximal view of spore 
<emphasis id="122F11EDC6B1D7602689D50F8C4780AA" bold="true">K</emphasis>
distal view of spore 
<emphasis id="47DB094B497ADCF0D90A0DC8554E13D9" bold="true">L</emphasis>
pseudoelaters. All from holotype and drawings by O. Suwanmala. (All drawing from 
<emphasis id="0A0E0510435BFFD710CF87993D7AA9C1" italics="true">S. Chantanaorrapint &amp; O. Suwanmala 4116</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0ELTAG" type="description">
<paragraph id="731C7A08B457113D35024078CFCFD3B0">
<heading id="73B84804971094D12FAAB6D590686E0A" reason="title">Description.</heading>
</paragraph>
<paragraph id="D1DAB76E104D292A1CC8FAB6D3D52716">
<emphasis id="47EBF8E84EC0C5358FBCC66B487F5F7A" bold="true" italics="true">Thallus</emphasis>
yellowish-green to dark green in fresh material, dull green to blackish- brown in dry material, prostrate or moderately adhering to the substratum, solid, ecostate, orbicular to sub-orbicular, dichotomously branched into several lobes, with a smooth dorsal surface; lobes ensiform or sometimes fan-shaped, up to 
<quantity id="ACC1E31C4F999E0896659550E841425E" metricMagnitude="-4" metricUnit="m" metricValue="8.0" unit="mm" value="0.8">0.8 mm</quantity>
long, 
<quantity id="EB23F116132946E936CD094CAE2BF379" metricMagnitude="-3" metricUnit="m" metricValue="2.0" metricValueMax="3.0" metricValueMin="1.0" unit="mm" value="2.0" valueMax="3.0" valueMin="1.0">1–3 mm</quantity>
wide; margins wavy, nearly entire to shallowly crenulate; apex flat, rarely ascending, occasionally tapering into apical tubers; tubers sometimes present on ventral surface. Thallus in cross section plano-convex to concave-convex, 4–10 cells thick in the middle region, without mucilage cavities. Dorsal epidermal cells rectangular to heptagonal, 28–75 × 25–50 µm, thin-walled, smooth. Chloroplast one per cell, large, occupying almost entire cell, variable in shape; pyrenoid present. 
<emphasis id="786478306D6CEA0980365B5A0A48062A" bold="true" italics="true">Nostoc colonies</emphasis>
scattered through the ventral side of thallus, appearing as dark dots. 
<emphasis id="FE9DB5B6A4D469218B69EC170C23B282" bold="true" italics="true">Rhizoids</emphasis>
hyaline or pale brown along ventral surface, inner wall smooth or tuberculate. 
<emphasis id="6E83154A5DDE2CC1832DCFB9EA341C46" bold="true" italics="true">Sexuality</emphasis>
monoicous. Androecia scattered and slightly raised over the dorsal surface of thallus, 2–3 antheridia per chamber; antheridia subglobose to globose, exposed at maturity, irregularly arranged jacket cells, shortly stalked, stalk with quadriseriate cells. Archegonia embedded in thallus, connected to the upper surface, scattered near the lobe of thallus. 
<emphasis id="CF4BF433737946DFE1F7E1625B89E5F2" bold="true" italics="true">Involucre</emphasis>
solitary, conical-cylindrical, up to 
<quantity id="08EC7BC80B093099971A0443E1674CB7" metricMagnitude="-3" metricUnit="m" metricValue="2.0" unit="mm" value="2.0">2 mm</quantity>
long, 2–4 cells thick, mouth smooth to crenulate. 
<emphasis id="200292D91CB2CC820414BD60C71C72EF" bold="true" italics="true">Sporophytes</emphasis>
capsule somewhat inclined, stout to narrowly cylindrical, 0.5–1 (– 1.2) cm long, yellow at apex, dehiscing from top toward base, bivalves rarely twisted when dry; epidermal cells of capsule elongate-rectangular, 68–200 × 12–30 µm, thick-walled, stomata present with two reniform guard cells, surrounded by 5–8 epidermal cells; assimilative layers 3–6 cells thick in cross section; the innermost capsule cells dark brown, subquadrate to rectangular; 27–67 × 22–53 µm; columella well-developed, red-brown, consisting of 16 cells (4 × 4 lines of cells) in cross section. 
<emphasis id="ACF0BA714BD6B73053FD8A6B2F258424" bold="true" italics="true">Spores</emphasis>
unicellular, yellow, rounded-triangular in polar view, equatorial diameter 32–50 µm in diameter, nearly smooth under light microscope (
<collectionCode id="5E2601A474EBA69D3AD264CD73473ECC" country="Mozambique" lsid="urn:lsid:biocol.org:col:13988" name="Secçáo de Botânica e Ecologia" type="Herbarium">LM</collectionCode>
), proximal surface with a distinct trilete mark, bordered by vermiculate strip on each side of trilete mark, each facet covered with fine vermiculate pattern; distal surface with a slightly dome-like region at the center, more densely vermiculate than proximal surface, sometimes with minute granules. 
<emphasis id="CB0A58BFA3E124F9C1C345C8E51160A8" bold="true" italics="true">Pseudoelaters</emphasis>
light brown or yellowish-brown at maturity, thin-walled, occasionally branched; pseudoelater cells subquadrate to short rectangular, 30–45 × 25–30 μm, without helicoidal band.
</paragraph>
<caption id="8319348E2315B66B794AB804028873A6" ID-DOI="10.3897/phytokeys.244.124080.figure4" ID-arpha="5E3933AF-2396-536F-B977-8B06F1A33B98" httpUri="https://binary.pensoft.net/fig/1098340" startId="F4">
<paragraph id="05AC9A616D3147F26EE658D9DC7181D8">
<label id="688E4BF31513B430BCCB0C297D83913D">Figure 4.</label>
</paragraph>
<paragraph id="69356A290E8FC1CBE556934433591E43">
Scanning electron micrographs of spores 
<emphasis id="BE22108C98ADACF2FCCBD3EC90E5DBFD" bold="true">A – F</emphasis>
<taxonomicName id="EF180D4439984ADD0F25CD544E44B91F" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" variety="scabrellus">
<emphasis id="13E1E268A989BD3F30A3FF043BAF1705" italics="true">Phaeoceros perpusillus var. scabrellus</emphasis>
</taxonomicName>
<emphasis id="EAAF54CEADB70575D6674E77006661C9" bold="true">A</emphasis>
distal view of spore showing a central hump-like projection 
<emphasis id="8E8435803029AAA4FD55E4949DDBD7C1" bold="true">B</emphasis>
proximal view of spore with a distinct triradiate mark 
<emphasis id="99963491414362DB74435626173B666A" bold="true">C</emphasis>
close-up of distal surface showing packed vermiculae 
<emphasis id="11352B81A9913193F789B91B5F6CF3B3" bold="true">D</emphasis>
close-up of proximal surface showing loosely arranged vermiculae 
<emphasis id="06A5D1FA8F8DA4A184B028410EBCF7A1" bold="true">E</emphasis>
proximal surface showing trilete mark and loosely arranged vermiculae 
<emphasis id="1E75D9124F5FF93E6CF1EDC8E29EA6B5" bold="true">F</emphasis>
spores and pseudoelaters 
<emphasis id="095C4C687AB778FB6537C66AB141663A" bold="true">G – I</emphasis>
<taxonomicName id="234085C13C9F4BD03359F6D8B70C9529" authority="var. perpusillus" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" variety="perpusillus">
<emphasis id="D52D74B3DAC9CF36A9BFEE9644FDE13D" italics="true">P. perpusillus var. perpusillus</emphasis>
</taxonomicName>
<emphasis id="7537A26E1B74DC7A36676068C055EAF8" bold="true">G</emphasis>
distal view of spore 
<emphasis id="55B4F98B5DA1D41000320BA7877A2CFE" bold="true">H</emphasis>
proximal view of spore 
<emphasis id="81EDE07942BEC502B54CB99C7CA3D57B" bold="true">I</emphasis>
close-up of distal surface showing pluripapillae. Photos by O. Suwanmala. (
<emphasis id="B12F3AFFCDAC195374E1F65080023F4B" bold="true">A – F</emphasis>
images from 
<emphasis id="50ABC5C2EF38D99F0AC1B09D0599B957" italics="true">S. Chantanaorrapint &amp; C. Promma 3129</emphasis>
<emphasis id="BDEF5FBD3CB5C179A842AA0DC398BE18" bold="true">G – I</emphasis>
images from 
<emphasis id="54C311B5AF6D1DE3E56C03B935BCC7CA" italics="true">S. Chantanaorrapint &amp; O. Suwanmala 3883</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0EYWAG" type="etymology">
<paragraph id="6DC0BA9CBFB3865555055C5302677BA0">
<heading id="8D8FBF28AECD2A25B9132DB1ACA0962F" reason="title">Etymology.</heading>
</paragraph>
<paragraph id="AF067F26B459914C851FC90A25C98212">The epithet of the variety refers to scabrate ornamentation observed under light microscope.</paragraph>
</subSubSection>
<subSubSection id="SECID0E4WAG" type="habitat">
<paragraph id="070E6AD406EBBD1AA3737B9E070E4A7F">
<heading id="FD12882AAD22D5FAFD48433B0C829431" reason="title">Habitat and distribution.</heading>
</paragraph>
<paragraph id="082579C7A7D7E0E709DAA00B7C0AAE1F">
<taxonomicName id="2634DF3242A28A2DD928B9FF59FE6A1E" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" variety="scabrellus">
<emphasis id="29F7A4D5996627B0A949F706A08DE1AC" italics="true">Phaeoceros perpusillus var. scabrellus</emphasis>
</taxonomicName>
is currently known only from northern 
<collectingCountry id="830B74DDC9AAA1A39F69121595BC4B57" name="Thailand">Thailand</collectingCountry>
. It grows on disturbed soil and sandstone in open site in grassland, pine-oak mixed montane deciduous forests at elevation of 
<quantity id="AEE0A5F74D164F559CBE8C60455D0D32" metricMagnitude="3" metricUnit="m" metricValue="1.7449999999999999" metricValueMax="2.1" metricValueMin="1.39" unit="m" value="1745.0" valueMax="2100.0" valueMin="1390.0">1390–2100 m</quantity>
. It may grow associated with other bryophytes such as 
<taxonomicName id="B4279DDB440525C7703B0A672AA88BD4" authority="Steph." authorityName="Steph." class="Anthocerotopsida" family="Anthocerotaceae" genus="Anthoceros" kingdom="Plantae" order="Anthocerotales" phylum="Anthocerotophyta" rank="species" species="subtilis">
<emphasis id="B23C2457D2CC4C8BD49F24CD31DBDA73" italics="true">Anthoceros subtilis</emphasis>
Steph.
</taxonomicName>
, 
<taxonomicName id="E17C8C07E598B87F3CD9C37A3B621566" class="Anthocerotopsida" family="Notothyladaceae" genus="Notothylas" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="species" species="levieri">
<emphasis id="C89071CF8852B4ECF34F43D57CF5AE28" italics="true">Notothylas levieri</emphasis>
</taxonomicName>
, 
<taxonomicName id="C4621BB21005D1025E7630FE043B5777" authority="(Schwein.) Sull. ex A. Gray" authorityName="A. Gray" baseAuthorityName="Schwein." class="Anthocerotopsida" family="Notothyladaceae" genus="Notothylas" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="species" species="orbicularis">
<emphasis id="A0F46F6BA0C4D212AB60FEA874727DB5" italics="true">N. orbicularis</emphasis>
(Schwein.) Sull. ex A. Gray
</taxonomicName>
, and 
<taxonomicName id="A0EB26449F58DC9CDEB7DD4EAFFB608B" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="species" species="carolinianus">
<emphasis id="EA68C57A09F4657A2A0341B767DBBA70" italics="true">P. carolinianus</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EAZAG" type="conservation status">
<paragraph id="5E4F32BAE2FFA422831F738A6BCF674A">
<heading id="BB18CE65CB7EC26207AABE98C511D401" reason="title">Conservation status.</heading>
</paragraph>
<paragraph id="445638B5DFB49D8FEE2F13405716FCAC">
This variety is currently known from three subpopulations, which are in protected areas (Chiang Dao Wildlife Sanctuary and Doi Suthep-Pui National Park). One of the subpopulations is located in a camping area, which is a common visiting site for tourists and dominated by 
<taxonomicName id="6FF9F5FCDAF5D4FF6A6CE4FE6B0E3241" authority="(Spreng.) R. M. King &amp; H. Rob." authorityName="R. M. King &amp; H. Rob." baseAuthorityName="Spreng." class="Magnoliopsida" family="Asteraceae" genus="Ageratina" kingdom="Plantae" order="Asterales" phylum="Tracheophyta" rank="species" species="adenophora">
<emphasis id="CB0829B3AEDF66693F186F9DBA7FEE5B" italics="true">Ageratina adenophora</emphasis>
(Spreng.) R. M. King &amp; H. Rob.
</taxonomicName>
(invasive species). Therefore, habitat quality is threatened by trampling and other destructive activities potentially caused by regular visits by tourists to the area, and invasive plant species. Together, these have the potential to cause a population reduction. The other subpopulation is also somewhat disturbed by human activities such as shifting cultivation. The extent of occurrence (
<abbrev id="ABBRID0ERZAG" xlink_title="extent of occurrence">
<collectionCode id="C0FF199C6447D88B1A56752B1A6823C5">EOO</collectionCode>
</abbrev>
) of 
<taxonomicName id="4E37DEF447EEC7B88538585E3CCEF6D4" class="Anthocerotopsida" family="Notothyladaceae" genus="Phaeoceros" kingdom="Plantae" order="Notothyladales" phylum="Anthocerotophyta" rank="variety" species="perpusillus" variety="scabrellus">
<emphasis id="B0D160551EC1BEB686B4CB235EB47424" italics="true">P. perpusillus var. scabrellus</emphasis>
</taxonomicName>
is estimated to be 
<quantity id="4A82F82E9CCF77D86C844BD454E81A74" metricMagnitude="5" metricUnit="m" metricValue="2.62925" unit="km" value="262.925">262.925 km</quantity>
<superScript id="6A1FF3F534F11429CB7EA254B2DD2267">2</superScript>
and its area of occupancy (
<abbrev id="ABBRID0EI1AG" xlink_title="area of occupancy">
<collectionCode id="A4C5D8F367934F304506FE649379198F">AOO</collectionCode>
</abbrev>
) is estimated to be 
<quantity id="06CF38CD11CF60F8B639E5F5DCCC12B5" metricMagnitude="4" metricUnit="m" metricValue="1.2" unit="km" value="12.0">12 km</quantity>
<superScript id="A82521E68742BC22F41AE25320C7BE6A">2</superScript>
, which falls within the limits for Endangered status under criterion 
<collectionCode id="4476CD69FB8CA39DDFE13B3BADC887CD" country="Germany" lsid="urn:lsid:biocol.org:col:15534" name="Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universitaet" type="Herbarium">B</collectionCode>
1 and 
<collectionCode id="5068B9D0751EE822C6F047C74F8B9F17" country="Germany" lsid="urn:lsid:biocol.org:col:15534" name="Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universitaet" type="Herbarium">B</collectionCode>
2 of 
<abbrev id="ABBRID0EO1AG" xlink_title="International Union for Conservation of Nature">
<collectionCode id="544C62751ED7929ED5D79EE0CA57D003">IUCN</collectionCode>
</abbrev>
Red List Categories and Criteria. Conservation efforts should focus on implementing strict regulations to reduce the impact of human activity and controlling invasive species, while also raising awareness among local communities about the importance of protecting the habitat.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ES1AG" type="material">
<paragraph id="687F7A879CF35329F0E1BA5DB51EC9C9">
<heading id="56CE3A75C192652E96FC64F6F18EAE55" reason="title">Additional specimens examined.</heading>
</paragraph>
<paragraph id="2E6076CB2F87D0D702F9F40515452072">
<emphasis id="F1190954DA3BBC1EF7421ACF3B02465F" bold="true">
<collectingCountry id="5141DBE0569C4A5BDF94300FA6D30789" name="Thailand">Thailand</collectingCountry>
</emphasis>
. 
<collectingRegion id="177D8C93DC6C5AD9979C17D97621D142" country="Thailand" name="Chiang Mai">Chiang Mai Province</collectingRegion>
: Chiang Dao Wildlife Sanctuary, 
<quantity id="41CCA6027FA07FE2FBF2FE8762E6C349" metricMagnitude="3" metricUnit="m" metricValue="1.85" metricValueMax="2.0" metricValueMin="1.7" unit="m" value="1850.0" valueMax="2000.0" valueMin="1700.0">1700–2000 m</quantity>
, 
<date id="CB1A79C6BAD517D9D3B3A241A8B87F55" value="2013-11-01">1 November 2013</date>
, 
<emphasis id="952447F4D7DF275F2A6D77C054276108" italics="true">
<collectionCode id="EF6CF49C5E01829F2BE14DC63B8800B5" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" type="Herbarium">S</collectionCode>
. Chantanaorrapint &amp; 
<collectionCode id="689FF38E15A44C00DFEB0A015540FF42" country="Denmark" name="University of Copenhagen" type="Herbarium">C</collectionCode>
. Promma 3125 
<collectionCode id="9E6E03C75DDC56E59AC27A18D16985BF" country="Germany" lsid="urn:lsid:biocol.org:col:15534" name="Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universitaet" type="Herbarium">B</collectionCode>
, 3129, 3216
</emphasis>
(
<collectionCode id="B412634D4CF94E9C5DA2618E8BF6B1DB" country="USA" httpUri="http://grbio.org/cool/5mg9-yyc1" name="Portland State University, Vertebrate Biology Museum" type="Museum">PSU</collectionCode>
); Doi Suthep-Pui National Park, Doi Mon Long Viewpoint, 
<quantity id="93582D52372D6A8A20F0194D12D4D088" metricMagnitude="3" metricUnit="m" metricValue="1.39" unit="m" value="1390.0">1390 m</quantity>
, 
<date id="35BCE841027DCEF9A5B6EC9AE86939A4" value="2015-11-04">4 November 2015</date>
, 
<emphasis id="0A3C9FD8D5947E8E88A94F61880633E5" italics="true">
<collectionCode id="436B75B350C43B19F43F2C7FDF919732" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" type="Herbarium">S</collectionCode>
. Chantanaorrapint &amp; 
<collectionCode id="BA71BE60428181DAA72AB9DD628F02E9" country="Austria" lsid="urn:lsid:biocol.org:col:15588" name="Naturhistorisches Museum Wien" type="Herbarium">W</collectionCode>
. Juengprayoon 143 
<collectionCode id="57488E400A00B8B14D566387AA586D58" country="Germany" lsid="urn:lsid:biocol.org:col:15534" name="Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universitaet" type="Herbarium">B</collectionCode>
</emphasis>
(
<collectionCode id="0C0B3BB4AAEA1FB568C2C985081FC447" country="USA" httpUri="http://grbio.org/cool/5mg9-yyc1" name="Portland State University, Vertebrate Biology Museum" type="Museum">PSU</collectionCode>
); 
<date id="0333289B95F841E29C9BD8E7BBBDD5B0" value="2020-11-15">15 November 2020</date>
, 
<emphasis id="31131F9A765ED378AF9E3B074075C390" italics="true">
<collectionCode id="FC89E07CA60B1C0E5A974D44B353356A" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" type="Herbarium">S</collectionCode>
. Chantanaorrapint &amp; 
<collectionCode id="ADD9D93CF5C84C70761097AB95E77949" country="Norway" lsid="urn:lsid:biocol.org:col:13093" name="Botanical Museum - University of Oslo" type="Herbarium">O</collectionCode>
. Suwanmala 4089, 4090
</emphasis>
(
<collectionCode id="24500A084F0FAF0FD00647B88EAAE9B2" country="USA" httpUri="http://grbio.org/cool/5mg9-yyc1" name="Portland State University, Vertebrate Biology Museum" type="Museum">PSU</collectionCode>
); Bhu Ping Palace, 
<quantity id="2B27A68A6562D6F02257CE6EEDB373B0" metricMagnitude="3" metricUnit="m" metricValue="1.4" unit="m" value="1400.0">1400 m</quantity>
, 
<date id="97C0E840EE8924C7A03700EF354CE784" value="2013-09-08">8 September 2013</date>
, 
<emphasis id="1AB4D3D55FBD24F23ED3527E851DE142" italics="true">
<collectionCode id="B643287CF7DE92FF503734A20C2EB66B" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" type="Herbarium">S</collectionCode>
. Rattanamanee 3
</emphasis>
(
<collectionCode id="D631A635A03061AF7F68EC66AF98982B" country="USA" httpUri="http://grbio.org/cool/5mg9-yyc1" name="Portland State University, Vertebrate Biology Museum" type="Museum">PSU</collectionCode>
); 
<date id="75DF537256D33F9D6AA3106B3938DB6A" value="2020-10-18">18 October 2020</date>
, 
<emphasis id="B460C93BC8A0B38F6ABBB70F6CB4EC77" italics="true">
<collectionCode id="96B92A90F57548AD9B63CD60277774CE" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" type="Herbarium">S</collectionCode>
. Chantanaorrapint &amp; 
<collectionCode id="9AA340DC652F6AAFA50B4EA421DA0FA3" country="Norway" lsid="urn:lsid:biocol.org:col:13093" name="Botanical Museum - University of Oslo" type="Herbarium">O</collectionCode>
. Suwanmala 4077
</emphasis>
(
<collectionCode id="F45144615388D7FD30A835656FC4EAFF" country="USA" httpUri="http://grbio.org/cool/5mg9-yyc1" name="Portland State University, Vertebrate Biology Museum" type="Museum">PSU</collectionCode>
); 
<date id="99249F30008BADE71BDC739E79BCA3A3" value="2021-10-05">5 October 2021</date>
, 
<emphasis id="69598260BFCE01BA3CA6E4986212D43E" italics="true">
<collectionCode id="2422B5865776B73A61E172CC1464B19D" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" type="Herbarium">S</collectionCode>
. Chantanaorrapint &amp; 
<collectionCode id="40AE2BB358F088FCAD6DE54D620E4BA2" country="Norway" lsid="urn:lsid:biocol.org:col:13093" name="Botanical Museum - University of Oslo" type="Herbarium">O</collectionCode>
. Suwanmala 4116
</emphasis>
(
<collectionCode id="9526F03154934550C8CBC989CD5506DE" country="USA" httpUri="http://grbio.org/cool/5mg9-yyc1" name="Portland State University, Vertebrate Biology Museum" type="Museum">PSU</collectionCode>
).
</paragraph>
</subSubSection>
</treatment>
</document>