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<document id="F09FB2767FE7BDFDC6FA6EF9DE44D424" ID-DOI="10.11646/phytotaxa.324.3.1" ID-ISSN="1179-3163" ID-Zenodo-Dep="13698300" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1725570015657" checkinUser="felipe" docAuthor="Ertz, Damien, Poulsen, Roar S., Charrier, Maryvonne &amp; Søchting, Ulrik" docDate="2017" docId="F336CC7E5D62023BAEB4A37FFC03FE92" docLanguage="en" docName="phytotaxa.324.3.1.pdf" docOrigin="Phytotaxa 324 (3)" docSource="http://dx.doi.org/10.11646/phytotaxa.324.3.1" docStyle="DocumentStyle:96748F8F1B6C902996E134952A3A36B9.13:Phytotaxa.2014-.journal_article" docStyleId="96748F8F1B6C902996E134952A3A36B9" docStyleName="Phytotaxa.2014-.journal_article" docStyleVersion="13" docTitle="Henssenia Ertz, R. S. Poulsen &amp; Sochting 2017, gen. nov." docType="treatment" docVersion="2" lastPageNumber="224" masterDocId="0F0FB4065D74022CAE3CA20CFF9AFFFF" masterDocTitle="Taxonomy and phylogeny of the genus Steinera (Arctomiales, Arctomiaceae) in the subantarctic islands of Crozet and Kerguelen" masterLastPageNumber="238" masterPageNumber="201" pageNumber="223" updateTime="1725576277220" updateUser="ExternalLinkService" zenodo-license-figures="UNSPECIFIED" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="13034CE3405993351099A90922AFC287">Taxonomy and phylogeny of the genus Steinera (Arctomiales, Arctomiaceae) in the subantarctic islands of Crozet and Kerguelen</mods:title>
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<mods:namePart id="5AC8A33ED9D062B74AF9BDBFED8B2E0B">Ertz, Damien</mods:namePart>
<mods:affiliation id="791062F7225555F809E64A073CF72146">Botanic Garden Meise, Department of Research, Nieuwelaan 38, B- 1860 Meise, Belgium; &amp; Fédération Wallonie-Bruxelles, Direction Générale de lEnseignement non obligatoire et de la Recherche scientifique, Rue A. Lavallée 1, BE- 1080 Bruxelles, Belgium</mods:affiliation>
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<mods:namePart id="0D7AB78C01523F48D7FD040B607FF36A">Poulsen, Roar S.</mods:namePart>
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<mods:namePart id="3F69BF36F3D71B5B3BBA3594FD92011B">Charrier, Maryvonne</mods:namePart>
<mods:affiliation id="B8BF923B2B2F921B8FBE9E93A9ED6BB9">Université de Rennes 1, UMR CNRS 6553, avenue du Général Leclerc 263, 35042 Rennes, France;</mods:affiliation>
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<mods:namePart id="2AE01638E9A99233A516F26A350AA20E">Søchting, Ulrik</mods:namePart>
<mods:affiliation id="E8415480E267F177FD64475AE0E59ECC">University of Copenhagen, Department of Biology, Universitetsparken 15, DK- 2100 Copenhagen Ø, Denmark;</mods:affiliation>
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<mods:date id="73A33B8CFA4E610EEA59135DD142C050">2017</mods:date>
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<taxonomicName id="BC9F06EB5D62023AAEB4A37FFDC7FE71" ID-CoL="4VM9" ID-ENA="2045328" authority="Ertz, R.S. Poulsen &amp; Sochting" authorityName="Ertz, R. S. Poulsen &amp; Sochting" authorityYear="2017" box="[136,605,371,398]" class="Lecanoromycetes" family="Koerberiaceae" genus="Henssenia" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="genus" status="gen. nov.">
<emphasis id="49EBA17A5D62023AAEB4A37FFE9BFE72" bold="true" box="[136,257,371,397]" italics="true" pageId="22" pageNumber="223">Henssenia</emphasis>
Ertz, R.S. Poulsen &amp; Søchting
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<paragraph id="7B207D685D62023AAEB4A390FEE9FE4E" blockId="22.[136,1452,412,685]" box="[136,371,412,433]" pageId="22" pageNumber="223">MycoBank: MB 822683</paragraph>
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<subSubSection id="33852EE35D62023BAEB4A3CCFC03FE92" lastPageId="23" lastPageNumber="224" pageId="22" pageNumber="223" type="diagnosis">
<paragraph id="7B207D685D62023AAEB4A3CCFD21FDBE" blockId="22.[136,1452,412,685]" pageId="22" pageNumber="223">
<emphasis id="49EBA17A5D62023AAEB4A3CCFF73FE2A" box="[136,233,448,469]" italics="true" pageId="22" pageNumber="223">Diagnosis</emphasis>
: A genus of
<taxonomicName id="BC9F06EB5D62023AAF5CA3CCFE71FE2A" box="[352,491,448,469]" class="Lecanoromycetes" family="Koerberiaceae" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="family">Koerberiaceae</taxonomicName>
characterized by a subfoliose to placodioid rosette-shaped thallus up to
<quantity id="BC67D08D5D62023AAAAAA3CCFB4CFE2A" box="[1174,1238,448,469]" metricMagnitude="-2" metricUnit="m" metricValue="3.5" pageId="22" pageNumber="223" unit="cm" value="3.5">3.5 cm</quantity>
in diam., with distinct marginal lobes up to
<quantity id="BC67D08D5D62023AAFBBA3E8FE24FE06" box="[391,446,484,505]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" pageId="22" pageNumber="223" unit="mm" value="2.0">2 mm</quantity>
wide, an upper ±algal-free layer consisting of a ±paraplectenchymatous layer, a scytonemoid photobiont with filaments or clusters typically arranged in vertically elongated fascicles between the fan-shaped ascending hyphae, simple, small ascospores sometimes having a plasma-bridge.
</paragraph>
<paragraph id="7B207D685D62023AAEB4A05CFD67FD9A" blockId="22.[136,1452,412,685]" box="[136,765,592,613]" pageId="22" pageNumber="223">
<typeStatus id="A424C3CA5D62023AAEB4A05CFF21FD9A" box="[136,187,592,613]" pageId="22" pageNumber="223">Type</typeStatus>
:
<taxonomicName id="BC9F06EB5D62023AAEF8A05CFD67FD9A" authority="(Tuck.) Ertz, R. S. Poulsen &amp; Sochting" authorityName="Ertz, R. S. Poulsen &amp; Sochting" baseAuthorityName="Tuck." box="[196,765,592,613]" class="Lecanoromycetes" family="Koerberiaceae" genus="Henssenia" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="species" species="glaucella">
<emphasis id="49EBA17A5D62023AAEF8A05CFE1DFD9A" box="[196,391,592,613]" italics="true" pageId="22" pageNumber="223">Henssenia glaucella</emphasis>
(Tuck.) Ertz, R.S. Poulsen &amp; Søchting
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<paragraph id="7B207D685D62023AAEB4A078FE58FD52" blockId="22.[136,1452,412,685]" pageId="22" pageNumber="223">
<emphasis id="49EBA17A5D62023AAEB4A078FF75FD76" box="[136,239,628,649]" italics="true" pageId="22" pageNumber="223">Etymology</emphasis>
: The new genus is dedicated to the German lichenologist Aino Henssen (1925-2011) for her important contribution to the taxonomy of cyanolichens.
</paragraph>
<paragraph id="7B207D685D62023AAEB4A0D1FE96F8AD" blockId="22.[136,1452,733,2090]" pageId="22" pageNumber="223">
<emphasis id="49EBA17A5D62023AAEB4A0D1FF46FD09" box="[136,220,733,758]" italics="true" pageId="22" pageNumber="223">Thallus</emphasis>
subfoliose to placodioid, closely appressed to the substrate, rosette-shaped, up to
<quantity id="BC67D08D5D62023AAA91A0D1FB64FD09" box="[1197,1278,733,758]" metricMagnitude="-2" metricUnit="m" metricValue="3.5" pageId="22" pageNumber="223" unit="cm" value="3.5">3.5 cm</quantity>
diam., margin distinctly and ±regularly lobate, central parts sometimes becoming ±crustose-areolate; marginal lobes separated by deep fissures, radiating, ±flabellate, up to 1.5(2) mm wide. Upper surface smooth or with striae in distal parts, often becoming weakly roughened in central areas, not pruinose, without isidia. Areoles derived from secondary rimose cracking between the main fissures, complete or incomplete, angular, flat to slightly convex. Lower surface smooth, usually without rhizohyphae. Prothallus not developed.
<emphasis id="49EBA17A5D62023AACC1A19DFCF5FC55" box="[765,879,913,938]" italics="true" pageId="22" pageNumber="223">Apothecia</emphasis>
often abundant, discrete or sometimes contiguous, usually laminal and mostly restricted to the central areas of the thallus, developed on the upper surface of the thallus, up to
<quantity id="BC67D08D5D62023AAEF1A1D5FE8BFC0D" box="[205,273,985,1010]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="22" pageNumber="223" unit="mm" value="1.0">1 mm</quantity>
diam., immersed when young, eventually more or less level with the surface of the thallus or sessile; disc dull brown, reddish-brown or black, not pruinose; thalline margin visible in species having sessile ascomata, narrow, smooth, concolourous with thallus.
<emphasis id="49EBA17A5D62023AAC4DA62DFCA4FBC5" box="[625,830,1057,1082]" italics="true" pageId="22" pageNumber="223">Thalline anatomy:</emphasis>
thallus thin; hyphae in upper 1/33/4 of thallus running perpendicularly to surface and densely packed, getting short-celled towards upper surface; upper ±algal-free layer (or cortex) consists of a hyaline, ±paraplectenchymatous layer, which is overlaid by a very thin, uneven, amorphous, semi-translucent necrotic layer.
<emphasis id="49EBA17A5D62023AAFCDA681FDF7FB59" box="[497,621,1165,1190]" italics="true" pageId="22" pageNumber="223">Photobiont</emphasis>
a species of filamentous cyanobacteria, with cells often wider than long, lacking a thick gelatinous sheath (?), scytonemoid, mainly present in the upper half of the thallus with the exception of the thin upper cortical layer, heterocytes paler, filaments or clusters typically arranged in vertically elongated fascicles between the fan-shaped ascending hyphae, being usually pale orange in the upper layer and greenish in deeper parts of the thallus; hyphae below the algal layer more loosely arranged, either vertically aligned as in the central areoles, or ±horizontally radiating towards the apices of the marginal lobes, generally interspersed with simple chains of algal cells; hyphae in medulla
<emphasis id="49EBA17A5D62023AAC58A769FDEDFA81" box="[612,631,1381,1406]" italics="true" pageId="22" pageNumber="223">c.</emphasis>
25 μm wide. Lower surface pale brown to whitish in distal parts, with a ±paraplectenchymatous and irregularly developed thin cortical layer, sometimes with a few short hyphae attaching the thallus to the substrate.
<emphasis id="49EBA17A5D62023AAF8BA7A1FCD0FA39" box="[439,842,1453,1478]" italics="true" pageId="22" pageNumber="223">Apothecial anatomy: Proper exciple</emphasis>
thin or almost inconspicuous, hyaline, sometimes pale brownish near the surface, composed laterally of hyphae arranged in parallel with elongated cells, basally becoming paraplectenchymatous; thalline exciple visible in non-immersed ascomata, thin, concolourous with the thallus.
<taxonomicName id="BC9F06EB5D62023AAEB4A415FE9BF9CD" box="[136,257,1561,1586]" genus="Hymenium" higherTaxonomySource="GBIF" kingdom="Fungi" pageId="22" pageNumber="223" rank="genus">
<emphasis id="49EBA17A5D62023AAEB4A415FE9BF9CD" box="[136,257,1561,1586]" italics="true" pageId="22" pageNumber="223">Hymenium</emphasis>
</taxonomicName>
hyaline, not inspersed, I+ blue turning dark reddish brown, KI+ deep blue; epihymenium brown, I+ blue turning dark reddish brown, KI+ blue; paraphyses sparingly branched, mainly in upper part of hymenium; apical cells swollen.
<emphasis id="49EBA17A5D62023AAECDA46DFE1CF985" box="[241,390,1633,1658]" italics="true" pageId="22" pageNumber="223">Hypothecium</emphasis>
hyaline, consisting of densely short-celled hyphae above, becoming paraplectenchymatous below, I+ pale orange, KI+ pale orange with few dark blue spots.
<emphasis id="49EBA17A5D62023AAD55A489FC03F961" box="[873,921,1669,1694]" italics="true" pageId="22" pageNumber="223">Asci</emphasis>
narrowly clavate, usually with a ±distinct foot, with a ±well-developed tholus, with the outermost layer becoming distinctly I+ reddish, KI+ dark blue (rest of the wall I, KI); ascospores (47)8 per ascus, simple or with a more or less median plasma-bridge but without a true transverse septum, ellipsoid or subglobose, small, up to 15 × 7 μm, without a gelatinous sheath, I (except internal cytoplasm becoming dark orange).
<emphasis id="49EBA17A5D62023AAC2DA519FDEFF8D1" box="[529,629,1813,1838]" italics="true" pageId="22" pageNumber="223">Pycnidia</emphasis>
rarely present, immersed in the thallus,
<taxonomicName id="BC9F06EB5D62023AAA0FA519FB26F8D1" box="[1075,1212,1813,1838]" class="Lecanoromycetes" family="Umbilicariaceae" genus="Umbilicaria" kingdom="Fungi" order="Umbilicariales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="genus">
<emphasis id="49EBA17A5D62023AAA0FA519FB26F8D1" box="[1075,1212,1813,1838]" italics="true" pageId="22" pageNumber="223">Umbilicaria</emphasis>
</taxonomicName>
<typeStatus id="A424C3CA5D62023AAAFEA519FB6CF8D1" box="[1218,1270,1813,1838]" pageId="22" pageNumber="223">type</typeStatus>
; conidia simple, bacilliform.
</paragraph>
<paragraph id="7B207D685D62023AAE81A551FD6CF889" blockId="22.[136,1452,733,2090]" box="[189,758,1885,1910]" pageId="22" pageNumber="223">
<emphasis id="49EBA17A5D62023AAE81A551FEAAF889" box="[189,304,1885,1910]" italics="true" pageId="22" pageNumber="223">Chemistry</emphasis>
: No lichen substances detected by TLC.
</paragraph>
<paragraph id="7B207D685D62023BAE81A58DFCECFF22" blockId="22.[136,1452,733,2090]" lastBlockId="23.[136,1451,160,365]" lastPageId="23" lastPageNumber="224" pageId="22" pageNumber="223">
<emphasis id="49EBA17A5D62023AAE81A58DFF67F865" box="[189,253,1921,1946]" italics="true" pageId="22" pageNumber="223">Notes</emphasis>
:
<taxonomicName id="BC9F06EB5D62023AAF37A58DFE1AF865" authorityName="Ertz, R. S. Poulsen &amp; Sochting" authorityYear="2017" box="[267,384,1921,1946]" class="Lecanoromycetes" family="Koerberiaceae" genus="Henssenia" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="genus">
<emphasis id="49EBA17A5D62023AAF37A58DFE1AF865" box="[267,384,1921,1946]" italics="true" pageId="22" pageNumber="223">Henssenia</emphasis>
</taxonomicName>
differs from
<taxonomicName id="BC9F06EB5D62023AAC2DA58DFDF4F865" authorityName=", Henssen &amp; James" authorityYear="1982" box="[529,622,1921,1946]" class="Lecanoromycetes" family="Koerberiaceae" genus="Steinera" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="genus">
<emphasis id="49EBA17A5D62023AAC2DA58DFDF4F865" box="[529,622,1921,1946]" italics="true" pageId="22" pageNumber="223">Steinera</emphasis>
</taxonomicName>
mainly by the simple ascospores sometimes having a plasma bridge and in this case the ascospores appear as being 1-septate. The genus also differs from
<taxonomicName id="BC9F06EB5D62023AADC8A5A9FBCBF841" authorityName=", Henssen &amp; James" authorityYear="1982" box="[1012,1105,1957,1982]" class="Lecanoromycetes" family="Koerberiaceae" genus="Steinera" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="genus" sensu="stricto">
<emphasis id="49EBA17A5D62023AADC8A5A9FBCBF841" box="[1012,1105,1957,1982]" italics="true" pageId="22" pageNumber="223">Steinera</emphasis>
</taxonomicName>
<taxonomicNameLabel id="52D81C015D62023AAA64A5A9FB08F841" box="[1112,1170,1957,1982]" pageId="22" pageNumber="223" sensu="stricto">s. str.</taxonomicNameLabel>
by usually smaller thalli, a scytonemoid photobiont, asci with a thinner upper apical wall, and a different ascomata ontogeny (cf.
<bibRefCitation id="1F0E00995D62023AAB13A5C5FE8AF7F9" author="Henssen, A. M. &amp; James, P. W." pageId="22" pageNumber="223" pagination="227 - 256" refId="ref28484" refString="Henssen, A. M. &amp; James, P. W. (1982) The lichen genus Steinera. Bulletin of the British Museum (Natural History), Botany Series 10: 227 - 256." type="book chapter" year="1982">Henssen &amp; James 1982</bibRefCitation>
, p. 231, studied for
<taxonomicName id="BC9F06EB5D62023AAC3CA5E1FD25F7F9" box="[512,703,2029,2054]" class="Lecanoromycetes" family="Koerberiaceae" genus="Steinera" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="species" species="radiata">
<emphasis id="49EBA17A5D62023AAC3CA5E1FDC7F7F9" box="[512,605,2029,2054]" italics="true" pageId="22" pageNumber="223">Steinera</emphasis>
<emphasis id="49EBA17A5D62023AAC51A5E1FD25F7F9" box="[621,703,2029,2054]" italics="true" pageId="22" pageNumber="223">radiata</emphasis>
</taxonomicName>
).
<taxonomicName id="BC9F06EB5D62023AACE5A5E1FCD4F7F9" authorityName="Ertz, R. S. Poulsen &amp; Sochting" authorityYear="2017" box="[729,846,2029,2054]" class="Lecanoromycetes" family="Koerberiaceae" genus="Henssenia" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="genus">
<emphasis id="49EBA17A5D62023AACE5A5E1FCD4F7F9" box="[729,846,2029,2054]" italics="true" pageId="22" pageNumber="223">Henssenia</emphasis>
</taxonomicName>
might be related to
<taxonomicName id="BC9F06EB5D62023AAA0AA5E1FB32F7F9" box="[1078,1192,2029,2054]" class="Lecanoromycetes" family="Coccocarpiaceae" genus="Peltularia" kingdom="Fungi" order="Peltigerales" pageId="22" pageNumber="223" phylum="Ascomycota" rank="genus">
<emphasis id="49EBA17A5D62023AAA0AA5E1FB32F7F9" box="[1078,1192,2029,2054]" italics="true" pageId="22" pageNumber="223">Peltularia</emphasis>
</taxonomicName>
(a genus for which no sequences are available so far) because of similarities in anatomy and developmental morphology of the ascocarp (
<bibRefCitation id="1F0E00995D63023BAEACA2ACFE04FF46" author="Henssen, A. M. &amp; James, P. W." box="[144,414,160,185]" pageId="23" pageNumber="224" pagination="227 - 256" refId="ref28484" refString="Henssen, A. M. &amp; James, P. W. (1982) The lichen genus Steinera. Bulletin of the British Museum (Natural History), Botany Series 10: 227 - 256." type="book chapter" year="1982">Henssen &amp; James 1982</bibRefCitation>
), but the
<typeStatus id="A424C3CA5D63023BAC34A2ACFDA2FF46" box="[520,568,160,185]" pageId="23" pageNumber="224">type</typeStatus>
species of this genus differs notably by an umbilicate and more heteromerous thallus (e.g.
<bibRefCitation id="1F0E00995D63023BAF2EA2C8FD86FF22" author="Henssen, A. M. &amp; James, P. W." box="[274,540,196,221]" pageId="23" pageNumber="224" pagination="227 - 256" refId="ref28484" refString="Henssen, A. M. &amp; James, P. W. (1982) The lichen genus Steinera. Bulletin of the British Museum (Natural History), Botany Series 10: 227 - 256." type="book chapter" year="1982">Henssen &amp; James 1982</bibRefCitation>
,
<bibRefCitation id="1F0E00995D63023BAC1BA2C8FCF0FF22" author="Jorgensen, P. M. &amp; Galloway, D. J." box="[551,874,196,221]" pageId="23" pageNumber="224" pagination="189 - 196" refId="ref28629" refString="Jorgensen, P. M. &amp; Galloway, D. J. (1984) A new subantarctic species of the lichen genus Peltularia. The Lichenologist 16: 189 - 196. https: // doi. org / 10.1017 / S 0024282984000335" type="journal article" year="1984">Jørgensen &amp; Galloway 1984</bibRefCitation>
).
</paragraph>
<paragraph id="7B207D685D63023BAE81A2E4FC03FE92" blockId="23.[136,1451,160,365]" pageId="23" pageNumber="224">
The new genus
<taxonomicName id="BC9F06EB5D63023BAF46A2E4FE75FEFE" authorityName="Ertz, R. S. Poulsen &amp; Sochting" authorityYear="2017" box="[378,495,232,257]" class="Lecanoromycetes" family="Koerberiaceae" genus="Henssenia" kingdom="Fungi" order="Peltigerales" pageId="23" pageNumber="224" phylum="Ascomycota" rank="genus">
<emphasis id="49EBA17A5D63023BAF46A2E4FE75FEFE" box="[378,495,232,257]" italics="true" pageId="23" pageNumber="224">Henssenia</emphasis>
</taxonomicName>
is part of the
<taxonomicName id="BC9F06EB5D63023BACA1A2E4FCD9FEFE" box="[669,835,232,257]" class="Lecanoromycetes" family="Koerberiaceae" kingdom="Fungi" order="Peltigerales" pageId="23" pageNumber="224" phylum="Ascomycota" rank="family">Koerberiaceae</taxonomicName>
. Members of this family share as synapomorphy the presence of radiating lines caused by radiating-photobiont bundles just beneath the upper cortex separated by bundles of cyanobiont-free medullary hyphae (Spribille &amp; Muggia 2014). In
<taxonomicName id="BC9F06EB5D63023BADBEA33CFC6DFEB6" authorityName="Ertz, R. S. Poulsen &amp; Sochting" authorityYear="2017" box="[898,1015,304,329]" class="Lecanoromycetes" family="Koerberiaceae" genus="Henssenia" kingdom="Fungi" order="Peltigerales" pageId="23" pageNumber="224" phylum="Ascomycota" rank="genus">
<emphasis id="49EBA17A5D63023BADBEA33CFC6DFEB6" box="[898,1015,304,329]" italics="true" pageId="23" pageNumber="224">Henssenia</emphasis>
</taxonomicName>
the thin striae are clearly visible on the thallus surface in
<taxonomicName id="BC9F06EB5D63023BAF72A358FE5AFE92" authorityName="Ertz, R. S. Poulsen &amp; Sochting. An" authorityYear="2017" baseAuthorityName="Ertz, R. S. Poulsen &amp; Sochting. An" baseAuthorityYear="2017" box="[334,448,340,365]" class="Lecanoromycetes" family="Koerberiaceae" genus="Henssenia" kingdom="Fungi" order="Peltigerales" pageId="23" pageNumber="224" phylum="Ascomycota" rank="species" species="werthii">
<emphasis id="49EBA17A5D63023BAF72A358FE5AFE92" box="[334,448,340,365]" italics="true" pageId="23" pageNumber="224">H. werthii</emphasis>
</taxonomicName>
, but are less apparent in the other species.
</paragraph>
</subSubSection>
</treatment>
</document>