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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">A co-evolutionary relationship exists between <italic>Endoraecium</italic>
(<italic>Pucciniales</italic>
) and its <italic>Acacia</italic>
hosts in Australia</title>
<author><name sortKey="Mctaggart, A R" sort="Mctaggart, A R" uniqKey="Mctaggart A" first="A. R." last="Mctaggart">A. R. Mctaggart</name>
<affiliation><nlm:aff id="A1"> Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia;</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2"> Department of Agriculture, Fisheries and Forestry, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia.</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A3"> Department of Microbiology and Plant Pathology, Tree Protection Co-operative Programme (TPCP), Forestry and Agricultural Biotechnology Institute (FABI), Private Bag X20, University of Pretoria, Pretoria, 0028, South Africa.</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Doungsa Ard, C" sort="Doungsa Ard, C" uniqKey="Doungsa Ard C" first="C." last="Doungsa-Ard">C. Doungsa-Ard</name>
<affiliation><nlm:aff id="A1"> Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia;</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A4"> Plant Biosecurity Cooperative Research Centre, LPO Box 5012, Bruce 2617, Australia.</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Geering, A D W" sort="Geering, A D W" uniqKey="Geering A" first="A. D. W." last="Geering">A. D. W. Geering</name>
<affiliation><nlm:aff id="A1"> Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Aime, M C" sort="Aime, M C" uniqKey="Aime M" first="M. C." last="Aime">M. C. Aime</name>
<affiliation><nlm:aff id="A5"> Department of Botany and Plant Pathology, Purdue University, 915 W. State Street, West Lafayette, IN 47907, USA.</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Shivas, R G" sort="Shivas, R G" uniqKey="Shivas R" first="R. G." last="Shivas">R. G. Shivas</name>
<affiliation><nlm:aff id="A2"> Department of Agriculture, Fisheries and Forestry, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia.</nlm:aff>
</affiliation>
</author>
</titleStmt>
<publicationStmt><idno type="wicri:source">PMC</idno>
<idno type="pmid">26823628</idno>
<idno type="pmc">4713111</idno>
<idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4713111</idno>
<idno type="RBID">PMC:4713111</idno>
<idno type="doi">10.3767/003158515X687588</idno>
<date when="2015">2015</date>
<idno type="wicri:Area/Pmc/Corpus">000071</idno>
<idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000071</idno>
</publicationStmt>
<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">A co-evolutionary relationship exists between <italic>Endoraecium</italic>
(<italic>Pucciniales</italic>
) and its <italic>Acacia</italic>
hosts in Australia</title>
<author><name sortKey="Mctaggart, A R" sort="Mctaggart, A R" uniqKey="Mctaggart A" first="A. R." last="Mctaggart">A. R. Mctaggart</name>
<affiliation><nlm:aff id="A1"> Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia;</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2"> Department of Agriculture, Fisheries and Forestry, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia.</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A3"> Department of Microbiology and Plant Pathology, Tree Protection Co-operative Programme (TPCP), Forestry and Agricultural Biotechnology Institute (FABI), Private Bag X20, University of Pretoria, Pretoria, 0028, South Africa.</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Doungsa Ard, C" sort="Doungsa Ard, C" uniqKey="Doungsa Ard C" first="C." last="Doungsa-Ard">C. Doungsa-Ard</name>
<affiliation><nlm:aff id="A1"> Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia;</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A4"> Plant Biosecurity Cooperative Research Centre, LPO Box 5012, Bruce 2617, Australia.</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Geering, A D W" sort="Geering, A D W" uniqKey="Geering A" first="A. D. W." last="Geering">A. D. W. Geering</name>
<affiliation><nlm:aff id="A1"> Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Aime, M C" sort="Aime, M C" uniqKey="Aime M" first="M. C." last="Aime">M. C. Aime</name>
<affiliation><nlm:aff id="A5"> Department of Botany and Plant Pathology, Purdue University, 915 W. State Street, West Lafayette, IN 47907, USA.</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Shivas, R G" sort="Shivas, R G" uniqKey="Shivas R" first="R. G." last="Shivas">R. G. Shivas</name>
<affiliation><nlm:aff id="A2"> Department of Agriculture, Fisheries and Forestry, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia.</nlm:aff>
</affiliation>
</author>
</analytic>
<series><title level="j">Persoonia : Molecular Phylogeny and Evolution of Fungi</title>
<idno type="ISSN">0031-5850</idno>
<idno type="eISSN">1878-9080</idno>
<imprint><date when="2015">2015</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc><textClass></textClass>
</profileDesc>
</teiHeader>
<front><div type="abstract" xml:lang="en"><p><italic>Endoraecium</italic>
is a genus of rust fungi that infects several species of <italic>Acacia</italic>
in Australia, South-East Asia and Hawaii. This study investigated the systematics of <italic>Endoraecium</italic>
from 55 specimens in Australia based on a combined morphological and molecular approach. Phylogenetic analyses were conducted on partitioned datasets of loci from ribosomal and mitochondrial DNA. The recovered molecular phylogeny supported a recently published taxonomy based on morphology and host range that divided <italic>Endoraecium digitatum</italic>
into five species. Spore morphology is synapomorphic and there is evidence <italic>Endoraecium</italic>
co-evolved with its <italic>Acacia</italic>
hosts. The broad host ranges of <italic>E. digitatum</italic>
, <italic>E. parvum</italic>
, <italic>E. phyllodiorum</italic>
and <italic>E. violae-faustiae</italic>
are revised in light of this study, and nine new species of <italic>Endoraecium</italic>
are described from Australia based on host taxonomy, morphology and phylogenetic concordance.</p>
</div>
</front>
<back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Aime, Mc" uniqKey="Aime M">MC Aime</name>
</author>
</analytic>
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<biblStruct><analytic><author><name sortKey="Berkeley, Mj" uniqKey="Berkeley M">MJ Berkeley</name>
</author>
<author><name sortKey="Broome, Ce" uniqKey="Broome C">CE Broome</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Berndt, R" uniqKey="Berndt R">R Berndt</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Brown, Gk" uniqKey="Brown G">GK Brown</name>
</author>
<author><name sortKey="Murphy, Dj" uniqKey="Murphy D">DJ Murphy</name>
</author>
<author><name sortKey="Kidman, J" uniqKey="Kidman J">J Kidman</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Castresana, J" uniqKey="Castresana J">J Castresana</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Cummins, Gb" uniqKey="Cummins G">GB Cummins</name>
</author>
<author><name sortKey="Hiratsuka, Y" uniqKey="Hiratsuka Y">Y Hiratsuka</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Dietel, P" uniqKey="Dietel P">P Dietel</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Dietel, P" uniqKey="Dietel P">P Dietel</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Doungsa Ard, C" uniqKey="Doungsa Ard C">C Doungsa-ard</name>
</author>
<author><name sortKey="Mctaggart, Ar" uniqKey="Mctaggart A">AR McTaggart</name>
</author>
<author><name sortKey="Geering, Adw" uniqKey="Geering A">ADW Geering</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Gardes, M" uniqKey="Gardes M">M Gardes</name>
</author>
<author><name sortKey="Bruns, Td" uniqKey="Bruns T">TD Bruns</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Gardner, De" uniqKey="Gardner D">DE Gardner</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Hodges, Cs" uniqKey="Hodges C">CS Hodges</name>
</author>
<author><name sortKey="Gardner, De" uniqKey="Gardner D">DE Gardner</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Katoh, K" uniqKey="Katoh K">K Katoh</name>
</author>
<author><name sortKey="Asimenos, G" uniqKey="Asimenos G">G Asimenos</name>
</author>
<author><name sortKey="Toh, H" uniqKey="Toh H">H Toh</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Laundon, Gf" uniqKey="Laundon G">GF Laundon</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Mains, Eb" uniqKey="Mains E">EB Mains</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Maslin, Br" uniqKey="Maslin B">BR Maslin</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Mcalpine, D" uniqKey="Mcalpine D">D McAlpine</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Mctaggart, Ar" uniqKey="Mctaggart A">AR McTaggart</name>
</author>
<author><name sortKey="Geering, Adw" uniqKey="Geering A">ADW Geering</name>
</author>
<author><name sortKey="Shivas, Rg" uniqKey="Shivas R">RG Shivas</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Miller, Jt" uniqKey="Miller J">JT Miller</name>
</author>
<author><name sortKey="Murphy, Dj" uniqKey="Murphy D">DJ Murphy</name>
</author>
<author><name sortKey="Ho, Syw" uniqKey="Ho S">SYW Ho</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Murphy, Dj" uniqKey="Murphy D">DJ Murphy</name>
</author>
<author><name sortKey="Brown, Gk" uniqKey="Brown G">GK Brown</name>
</author>
<author><name sortKey="Miller, Jt" uniqKey="Miller J">JT Miller</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Nylander, Jaa" uniqKey="Nylander J">JAA Nylander</name>
</author>
<author><name sortKey="Wilgenbusch, Jc" uniqKey="Wilgenbusch J">JC Wilgenbusch</name>
</author>
<author><name sortKey="Warren, Dl" uniqKey="Warren D">DL Warren</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Ono, Y" uniqKey="Ono Y">Y Ono</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Ronquist, F" uniqKey="Ronquist F">F Ronquist</name>
</author>
<author><name sortKey="Huelsenbeck, Jp" uniqKey="Huelsenbeck J">JP Huelsenbeck</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Savile, Dbo" uniqKey="Savile D">DBO Savile</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Sawada, K" uniqKey="Sawada K">K Sawada</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Scholler, M" uniqKey="Scholler M">M Scholler</name>
</author>
<author><name sortKey="Aime, Mc" uniqKey="Aime M">MC Aime</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Stamatakis, A" uniqKey="Stamatakis A">A Stamatakis</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Stevens, Fl" uniqKey="Stevens F">FL Stevens</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Tamura, K" uniqKey="Tamura K">K Tamura</name>
</author>
<author><name sortKey="Peterson, D" uniqKey="Peterson D">D Peterson</name>
</author>
<author><name sortKey="Peterson, N" uniqKey="Peterson N">N Peterson</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Vialle, A" uniqKey="Vialle A">A Vialle</name>
</author>
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</author>
<author><name sortKey="Allaire, M" uniqKey="Allaire M">M Allaire</name>
</author>
</analytic>
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<biblStruct><analytic><author><name sortKey="Vilgalys, R" uniqKey="Vilgalys R">R Vilgalys</name>
</author>
<author><name sortKey="Hester, M" uniqKey="Hester M">M Hester</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Walker, J" uniqKey="Walker J">J Walker</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="White, Tj" uniqKey="White T">TJ White</name>
</author>
<author><name sortKey="Bruns, Td" uniqKey="Bruns T">TD Bruns</name>
</author>
<author><name sortKey="Lee, S" uniqKey="Lee S">S Lee</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Winter, G" uniqKey="Winter G">G Winter</name>
</author>
</analytic>
</biblStruct>
</listBibl>
</div1>
</back>
</TEI>
<pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Persoonia</journal-id>
<journal-id journal-id-type="iso-abbrev">Persoonia</journal-id>
<journal-id journal-id-type="publisher-id">Persoonia</journal-id>
<journal-title-group><journal-title>Persoonia : Molecular Phylogeny and Evolution of Fungi</journal-title>
</journal-title-group>
<issn pub-type="ppub">0031-5850</issn>
<issn pub-type="epub">1878-9080</issn>
<publisher><publisher-name>Naturalis Biodiversity Center & Centraallbureau voor Schimmelcultures</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">26823628</article-id>
<article-id pub-id-type="pmc">4713111</article-id>
<article-id pub-id-type="doi">10.3767/003158515X687588</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject>
</subj-group>
</article-categories>
<title-group><article-title>A co-evolutionary relationship exists between <italic>Endoraecium</italic>
(<italic>Pucciniales</italic>
) and its <italic>Acacia</italic>
hosts in Australia</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>McTaggart</surname>
<given-names>A.R.</given-names>
</name>
<xref ref-type="aff" rid="A1"><sup>1</sup>
</xref>
<xref ref-type="aff" rid="A2"><sup>2</sup>
</xref>
<xref ref-type="aff" rid="A3"><sup>3</sup>
</xref>
<xref ref-type="corresp" rid="COR1"></xref>
</contrib>
<contrib contrib-type="author"><name><surname>Doungsa-ard</surname>
<given-names>C.</given-names>
</name>
<xref ref-type="aff" rid="A1"><sup>1</sup>
</xref>
<xref ref-type="aff" rid="A4"><sup>4</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Geering</surname>
<given-names>A.D.W.</given-names>
</name>
<xref ref-type="aff" rid="A1"><sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Aime</surname>
<given-names>M.C.</given-names>
</name>
<xref ref-type="aff" rid="A5"><sup>5</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Shivas</surname>
<given-names>R.G.</given-names>
</name>
<xref ref-type="aff" rid="A2"><sup>2</sup>
</xref>
</contrib>
</contrib-group>
<aff id="A1"><label>1</label>
Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia;</aff>
<aff id="A2"><label>2</label>
Department of Agriculture, Fisheries and Forestry, Ecosciences Precinct, GPO Box 267, Brisbane, Queensland 4001, Australia.</aff>
<aff id="A3"><label>3</label>
Department of Microbiology and Plant Pathology, Tree Protection Co-operative Programme (TPCP), Forestry and Agricultural Biotechnology Institute (FABI), Private Bag X20, University of Pretoria, Pretoria, 0028, South Africa.</aff>
<aff id="A4"><label>4</label>
Plant Biosecurity Cooperative Research Centre, LPO Box 5012, Bruce 2617, Australia.</aff>
<aff id="A5"><label>5</label>
Department of Botany and Plant Pathology, Purdue University, 915 W. State Street, West Lafayette, IN 47907, USA.</aff>
<author-notes><corresp id="COR1">corresponding author e-mail: <email>alistair.mctaggart@gmail.com</email>
.</corresp>
</author-notes>
<pub-date pub-type="epub"><day>13</day>
<month>2</month>
<year>2015</year>
</pub-date>
<pub-date pub-type="ppub"><month>12</month>
<year>2015</year>
</pub-date>
<volume>35</volume>
<fpage>50</fpage>
<lpage>62</lpage>
<history><date date-type="received"><day>10</day>
<month>6</month>
<year>2014</year>
</date>
<date date-type="accepted"><day>18</day>
<month>12</month>
<year>2014</year>
</date>
</history>
<permissions><copyright-statement>© 2015 Naturalis Biodiversity Center & Centraalbureau voor Schimmelcultures</copyright-statement>
<copyright-year>2015</copyright-year>
<license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode"><license-p>You are free to share - to copy, distribute and transmit the work, under the following conditions:</license-p>
<license-p>Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work).</license-p>
<license-p>Non-commercial: You may not use this work for commercial purposes.</license-p>
<license-p>No derivative works: You may not alter, transform, or build upon this work.</license-p>
<license-p>For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode">http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode</ext-link>
. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights.</license-p>
</license>
</permissions>
<abstract abstract-type="executive-summary"><p><italic>Endoraecium</italic>
is a genus of rust fungi that infects several species of <italic>Acacia</italic>
in Australia, South-East Asia and Hawaii. This study investigated the systematics of <italic>Endoraecium</italic>
from 55 specimens in Australia based on a combined morphological and molecular approach. Phylogenetic analyses were conducted on partitioned datasets of loci from ribosomal and mitochondrial DNA. The recovered molecular phylogeny supported a recently published taxonomy based on morphology and host range that divided <italic>Endoraecium digitatum</italic>
into five species. Spore morphology is synapomorphic and there is evidence <italic>Endoraecium</italic>
co-evolved with its <italic>Acacia</italic>
hosts. The broad host ranges of <italic>E. digitatum</italic>
, <italic>E. parvum</italic>
, <italic>E. phyllodiorum</italic>
and <italic>E. violae-faustiae</italic>
are revised in light of this study, and nine new species of <italic>Endoraecium</italic>
are described from Australia based on host taxonomy, morphology and phylogenetic concordance.</p>
</abstract>
<kwd-group><kwd><italic>Atelocauda</italic>
</kwd>
<kwd>endocyclic rusts</kwd>
<kwd><italic>Mimosoideae</italic>
</kwd>
<kwd><italic>Racospermyces</italic>
</kwd>
<kwd><italic>Raveneliaceae</italic>
</kwd>
<kwd><italic>Uredinales</italic>
</kwd>
</kwd-group>
</article-meta>
</front>
<body><sec id="s1"><title>INTRODUCTION</title>
<p><italic>Endoraecium</italic>
(<italic>Pucciniales</italic>
, <italic>Raveneliaceae</italic>
) was established for two endocyclic species of rust fungi with pedicellate spores, <italic>E. acaciae</italic>
(the type species) and <italic>E. hawaiiense</italic>
, which were found on <italic>Acacia koa</italic>
(<italic>Fabaceae</italic>
, subfamily <italic>Mimosoideae</italic>
) in Hawaii (<xref rid="R12" ref-type="bibr">Hodges & Gardner 1984</xref>
). There are over 1 000 species of <italic>Acacia</italic>
s.str. (hereafter referred to as <italic>Acacia</italic>
) known in Australia (<xref rid="R20" ref-type="bibr">Murphy et al. 2010</xref>
, <xref rid="R16" ref-type="bibr">Maslin 2013</xref>
). The classification of <italic>Acacia</italic>
has moved from a traditional morphological one based on sections, to a molecular phylogenetic one based on clades (<xref rid="R20" ref-type="bibr">Murphy et al. 2010</xref>
, <xref rid="R19" ref-type="bibr">Miller et al. 2013</xref>
). At the commencement of this study, 20 species of <italic>Acacia</italic>
were recorded as hosts of seven species of <italic>Endoraecium</italic>
in Australia (<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
). These 20 host species fall within three clades of the plurinerved, uninerved and Botrycephalae (p.u.b.) group of <italic>Acacia</italic>
(<xref rid="R20" ref-type="bibr">Murphy et al. 2010</xref>
), namely the i) Botrycephalae subclade (<xref rid="R20" ref-type="bibr">Murphy et al. 2010</xref>
); ii) Juliflorae p.p. clade (<xref rid="R19" ref-type="bibr">Miller et al. 2013</xref>
); and iii) Plurinerves p.p. clade (<xref rid="R19" ref-type="bibr">Miller et al. 2013</xref>
), hereafter referred to as Botrycephalae, Juliflorae and Plurinerves, respectively.</p>
<p>Species of <italic>Endoraecium</italic>
produce a range of symptoms on <italic>Acacia</italic>
that include bullate swellings or pulvinate sori on phyllodes, or galls and witches’ brooms on stems. These rust fungi produce spores of three types: i) spermogonia; ii) pedicellate spores, which have reticulate to foveolate ornamentations and were variously called aecial urediniospores (<xref rid="R32" ref-type="bibr">Walker 2001</xref>
), uredo-like aeciospores (<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
) or teliospores in the endocyclic species (<xref rid="R12" ref-type="bibr">Hodges & Gardner 1984</xref>
); iii) teliospores, which are single celled, subhyaline and smooth walled. The teliospores are morphologically similar to those of <italic>Uromyces</italic>
s.l. and several species of <italic>Endoraecium</italic>
were at one time classified in that genus, including <italic>U. digitatus</italic>
(<xref rid="R34" ref-type="bibr">Winter 1886</xref>
), <italic>U. bicintus</italic>
(<xref rid="R17" ref-type="bibr">McAlpine 1906</xref>
), <italic>U. phyllodiorum</italic>
(<xref rid="R17" ref-type="bibr">McAlpine 1906</xref>
), <italic>U. hyalosporus</italic>
(<xref rid="R25" ref-type="bibr">Sawada 1913</xref>
) and <italic>U. koae</italic>
(<xref rid="R28" ref-type="bibr">Stevens 1925</xref>
). On the basis of morphology, some species currently considered as <italic>Endoraecium</italic>
were previously placed in other genera, including <italic>Pileolaria</italic>
(<xref rid="R7" ref-type="bibr">Dietel 1921</xref>
), <italic>Maravalia</italic>
(<xref rid="R8" ref-type="bibr">Dietel 1924</xref>
), <italic>Poliotelium</italic>
(<xref rid="R15" ref-type="bibr">Mains 1939</xref>
), <italic>Atelocauda</italic>
(<xref rid="R6" ref-type="bibr">Cummins & Hiratsuka 1983</xref>
, <xref rid="R22" ref-type="bibr">Ono 1984</xref>
, <xref rid="R11" ref-type="bibr">Gardner 1991</xref>
), and <italic>Racospermyces</italic>
(<xref rid="R32" ref-type="bibr">Walker 2001</xref>
).</p>
<p><xref rid="R11" ref-type="bibr">Gardner (1991)</xref>
proposed that <italic>Endoraecium</italic>
may be closely related to <italic>Uromycladium</italic>
(<italic>Pileolariaceae</italic>
), another genus of rust that occurs on <italic>Acacia</italic>
in Australia. However, in combined analyses of the Large Subunit (LSU) and Small Subunit (SSU) regions of ribosomal DNA (rDNA), the systematic position of <italic>Endoraecium</italic>
was shown by <xref rid="R26" ref-type="bibr">Scholler & Aime (2006)</xref>
and <xref rid="R1" ref-type="bibr">Aime (2006)</xref>
to have an affinity with mimosoid rusts in the <italic>Raveneliaceae</italic>
rather than with the <italic>Pileolariaceae.</italic>
</p>
<p>In this study, the systematic relationships of <italic>Endoraecium</italic>
on <italic>Acacia</italic>
in Australia were investigated with freshly collected field material and herbarium specimens. Currently, <italic>Endoraecium</italic>
contains 13 species, all on <italic>Acacia</italic>
, with seven endemic to Australia, one to South-East Asia and five to Hawaii (<xref rid="R17" ref-type="bibr">McAlpine 1906</xref>
, <xref rid="R25" ref-type="bibr">Sawada 1913</xref>
, <xref rid="R12" ref-type="bibr">Hodges & Gardner 1984</xref>
, <xref rid="R3" ref-type="bibr">Berndt 2011</xref>
). The species are <italic>E. acaciae</italic>
(type on <italic>A. koa</italic>
, Hawaii), <italic>E. angustiphyllodium</italic>
(type on <italic>A. koa</italic>
, Hawaii), <italic>E. bicinctum</italic>
(type on <italic>A. fasciculifera</italic>
, Australia), <italic>E. digitatum</italic>
(type on <italic>A. no-tabilis</italic>
, Australia), <italic>E. hawaiiense</italic>
(type on <italic>A. koa</italic>
, Hawaii), <italic>E. hyalo-sporum</italic>
(type on <italic>A. confusa</italic>
, Taiwan), <italic>E. kauaianum</italic>
(type on <italic>A. koa</italic>
, Hawaii), <italic>E. koae</italic>
(type on <italic>A. koa</italic>
, Hawaii), <italic>E. parvum</italic>
(type on <italic>A. leiocalyx</italic>
, Australia), <italic>E. phyllodiorum</italic>
(type on <italic>Acacia</italic>
sp.<italic>,</italic>
Australia), <italic>E. tierneyi</italic>
(type on <italic>A. harpophylla</italic>
, Australia), <italic>E. violae-faustiae</italic>
(type on <italic>Acacia</italic>
sp., Australia) and <italic>E. walkerianum</italic>
(type on <italic>A. penninervis</italic>
, Australia). The SSU, Internal Transcribed Spacer (ITS) and LSU regions of rDNA, and cytochrome c oxidase subunit 3 (CO3) of mitochondrial DNA, were used for phylogenetic reconstruction to determine whether the current taxonomy of Australian species was supported by molecular data and if further diversity existed within <italic>Endoraecium</italic>
.</p>
</sec>
<sec sec-type="materials|methods" id="s2"><title>MATERIALS AND METHODS</title>
<sec id="s2a"><title>Taxon selection</title>
<p>Fresh specimens were collected from New South Wales, the Northern Territory, Queensland, Victoria and Western Australia, representing a broad diversity of <italic>Acacia</italic>
(<xref ref-type="table" rid="T1">Table 1</xref>
). Species from representative genera of <italic>Raveneliaceae</italic>
, namely <italic>Diorchidium</italic>
, <italic>Kernkampella</italic>
, <italic>Ravenelia</italic>
and <italic>Sphaerophragmium</italic>
, were selected as outgroup taxa for the phylogenetic analyses. <italic>Uromycladium simplex</italic>
(<italic>Pileolariaceae</italic>
) was included as an outgroup to the <italic>Raveneliaceae</italic>
, as it represented another Australian endemic rust on <italic>Acacia</italic>
.</p>
</sec>
<sec id="s2b"><title>Morphology</title>
<p>Spores were scraped from leaf material, mounted in lactic acid and gently heated to boiling. Preparations were examined with a Leica DMLB microscope and images were taken with a Leica DFC500 camera. Measurements were made digitally from photographed spores. All previous taxonomic studies of <italic>Endoraecium</italic>
have adopted an ontogenic approach to spore classification (<xref rid="R12" ref-type="bibr">Hodges & Gardner 1984</xref>
, <xref rid="R32" ref-type="bibr">Walker 2001</xref>
, <xref rid="R26" ref-type="bibr">Scholler & Aime 2006</xref>
, <xref rid="R3" ref-type="bibr">Berndt 2011</xref>
). Aeciospores, urediniospores and in some cases teliospores, are often indistinguishable with this approach. For this reason, we used a morphological approach to classify spores (<xref rid="R14" ref-type="bibr">Laundon 1967</xref>
), which recognises urediniospores of <italic>Endoraecium</italic>
as spores with reticulate ornamentation, pedicels and germ pores.</p>
</sec>
<sec id="s2c"><title><italic>DNA extraction</italic>
, <italic>PCR and sequencing</italic>
</title>
<p>DNA was extracted from fresh or archived material. Spores were obtained from leaf material according to the protocol in <xref rid="R18" ref-type="bibr">McTaggart et al. (2014)</xref>
, and then extracted with the UltraClean Microbial DNA Isolation Kit (MoBio Laboratories Inc., Solana Beach, CA, USA).</p>
<p>The ITS region was amplified with ITS1F/ITS4B (<xref rid="R10" ref-type="bibr">Gardes & Bruns 1993</xref>
). The ITS2-LSU region was amplified with Rust2inv (<xref rid="R1" ref-type="bibr">Aime 2006</xref>
) / LR7 (<xref rid="R31" ref-type="bibr">Vilgalys & Hester 1990</xref>
) and nested with LROR/LR6 (<xref rid="R31" ref-type="bibr">Vilgalys & Hester 1990</xref>
). The SSU region was amplified with NS1 (<xref rid="R33" ref-type="bibr">White et al. 1990</xref>
) / Rust 18S-R (<xref rid="R1" ref-type="bibr">Aime 2006</xref>
) according to the protocol by <xref rid="R1" ref-type="bibr">Aime (2006)</xref>
. CO3 was amplified with the primers CO3_F1/CO3_R1 (<xref rid="R30" ref-type="bibr">Vialle et al. 2009</xref>
). All PCRs were performed with high fidelity Phusion enzyme (New England Biolabs Inc.) according to the manufacturer’s instructions. The PCRs were performed with the following annealing temperatures: SSU, ITS and nested LSU at 62 °C, the initial LSU at 60 °C, and CO3 at 55 °C. PCR products were sent to Macrogen Korea for direct sequencing. Sequences were uploaded to GenBank under the accession numbers listed in <xref ref-type="table" rid="T1">Table 1</xref>
.</p>
</sec>
<sec id="s2d"><title>Phylogenetic analyses</title>
<p>The SSU (19), ITS (47), LSU (51) and CO3 (29) sequences were aligned separately in MAFFT (<xref rid="R13" ref-type="bibr">Katoh et al. 2009</xref>
) (available: <ext-link ext-link-type="uri" xlink:href="http://www.ebi.ac.uk/Tools/msa/mafft/">http://www.ebi.ac.uk/Tools/msa/mafft/</ext-link>
). Non-homologous regions in the ITS alignment were removed using GBlocks (<xref rid="R5" ref-type="bibr">Castresana 2000</xref>
). The sequences from each locus were concatenated in a supermatrix and run as partitioned datasets with maximum likelihood and Bayesian inference as phylogenetic criteria. CO3 was translated and included as amino acid coding sequence. JTT was selected as a protein coding model of evolution in MEGA5 (<xref rid="R29" ref-type="bibr">Tamura et al. 2011</xref>
). Maximum likelihood was implemented as a search criterion in RAxML (<xref rid="R27" ref-type="bibr">Stamatakis 2014</xref>
). GTRGAMMA was specified as the model of evolution for nucleotide sequence data. The RAxML analyses were run with a rapid Bootstrap analysis (command -f a) using a random starting tree and 1 000 maximum likelihood bootstrap replicates. A Markov Chain Monte Carlo (MCMC) search in a Bayesian analysis was conducted with MrBayes (<xref rid="R23" ref-type="bibr">Ronquist & Huelsenbeck 2003</xref>
). Four runs, each consisting of four chains, were implemented for 5 million generations. The cold chain was heated at a temperature of 0.25. Substitution model parameters were sampled every 500 generations and trees were saved every 1 000 generations. Convergence of the Bayesian analysis was confirmed using AWTY (<xref rid="R21" ref-type="bibr">Nylander et al. 2008</xref>
) (available at: ceb.csit.fsu.edu/awty/) and a burn-in of 18 000 generations was calculated. The ML and Bayesian analyses were run three times to test accuracy. Alignments and trees were uploaded to TreeBASE (<ext-link ext-link-type="uri" xlink:href="http://purl.org/phylo/treebase/phylows/study/TB2:S15410">http://purl.org/phylo/treebase/phylows/study/TB2:S15410</ext-link>
).</p>
</sec>
</sec>
<sec id="s3"><title>RESULTS</title>
<p>The topologies recovered by Bayesian inference and maximum likelihood were identical (<xref ref-type="fig" rid="F1">Fig. 1</xref>
). No differences were recovered in the tree topologies between nucleotide and amino acid coding sequences of CO3. <italic>Endoraecium</italic>
was recovered as a monophyletic group sister to other members of the <italic>Raveneliaceae</italic>
.</p>
<p>Three major clades of <italic>Endoraecium</italic>
corresponded to three clades in <italic>Acacia</italic>
, namely Plurinerves, Juliflorae and Botrycepha-lae. The clade of <italic>Endoraecium</italic>
that diversified on Plurinerves contained the Hawaiian species of <italic>Endoraecium</italic>
and the Australian species, <italic>E. tierneyi</italic>
. Rust taxa formed well-supported clades corresponding to host species. This is contrary to the current broad host ranges proposed for <italic>E. digitatum</italic>
, <italic>E. parvum</italic>
, <italic>E. phylliodorum</italic>
and <italic>E. violae-faustiae</italic>
(<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
).</p>
<p>Morphology of the urediniospores was a synapomorphic character for the two large clades on Juliflorae and Botrycephalae (<xref ref-type="fig" rid="F1">Fig. 1</xref>
). Urediniospores of species on Juliflorae were golden brown with reticulate to foveolate ornamentation. Urediniospores of those on Botrycephalae were subhyaline to pale yellow with reticulate ornamentation.</p>
</sec>
<sec id="s4"><title>TAXONOMY</title>
<p><bold><italic>Endoraecium auriculiforme</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808980; <xref ref-type="fig" rid="F2">Fig. 2</xref>
</p>
<p><italic>Etymology</italic>
. Name refers to the host, <italic>Acacia</italic>
<italic>auriculiformis</italic>
, on which it was found.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Northern Territory, Darwin, Howard Springs (-12.4564, 131.0522), on <italic>A. auriculiformis</italic>
, 15 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
(holotype BRIP 56548).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, subepidermal, erumpent, on bullate swellings up to 1 cm long, or pulverulent on phyllode surface, linear to round, yellowish brown; paraphyses intrasoral, cylindrical, apex 6–9-digitate, subhyaline to yellow, 35–70 × 10–14 μm, thickened at apex, pedicel up to 40 μm. <italic>Urediniospores</italic>
obovoid, oval to fusiform, apex acute, yellowish brown, 31–46 × 16–20 μm; wall 3–5 μm thick at sides and 5–9 μm thick at apex, foveolate, with 3–4 equatorial germ pores; pedicel 3–5 μm, or absent. <italic>Telia</italic>
formed from uredinia, erumpent, reddish brown. <italic>Teliospores</italic>
cylindrical to fusiform, apex rounded, 2–4-digitate, hyaline to subhyaline, 36–54 × 13–25 μm; wall 1.5–2.5 μm thick at sides, 6–8 μm thick at apex; pedicel persistent, up to 38 μm. <italic>Basidia</italic>
cylindrical, 3-septate, up to 35–40 × 8–12 μm. <italic>Basidiospores</italic>
globose, hyaline, 4–6 μm, smooth-walled.</p>
<p>On phyllodes of <italic>Acacia auriculiformis</italic>
(Juliflorae).</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, Northern Territory, Howard Springs, on <italic>A. auriculiformis</italic>
, 9 May 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
, BRIP 55609; 15 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>R.G. Shivas</italic>
, BRIP 56550; Northern Territory, Darwin, Bronzewing Ave, Howard Springs, 15 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
& <italic>R.G. Shivas</italic>
, BRIP 56549.</p>
<p>Notes — <italic>Endoraecium auriculiforme</italic>
has foveolate uredinio-spores with an acute and thickened apex, and digitate paraphyses that resemble narrow teliospores. <italic>Endoraecium violae-faustiae</italic>
and <italic>E. peggii</italic>
also produce similar paraphyses. These three species all occur on hosts in the Juliflorae, with <italic>E. auriculiforme</italic>
restricted to <italic>A. auriculiformis</italic>
; <italic>E. violae-faustiae</italic>
to <italic>A. aula-cocarpa</italic>
and <italic>A. crassicarpa</italic>
, and <italic>E. peggii</italic>
to <italic>A. holosericea</italic>
. In the phylogenetic analyses, <italic>E. auriculiforme</italic>
was recovered as sister to <italic>E. peggii</italic>
(<xref ref-type="fig" rid="F1">Fig. 1</xref>
).</p>
<p><bold><italic>Endoraecium bicinctum</italic>
</bold>
(McAlpine) M. Scholler & Aime, Myco-science 47: 163. 2006</p>
<p><italic>Basionym</italic>
. <italic>Uromyces bicinctus</italic>
McAlpine, Rusts of Australia: 93. 1906.</p>
<p>≡ <italic>Pileolaria bicincta</italic>
(McAlpine) Dietel, Ann. Mycol. 19: 302. 1921.</p>
<p>≡ <italic>Atelocauda bicincta</italic>
(McAlpine) Cummins & Y. Hirats., Illustr. Gen. Rust Fungi, rev. edn (St. Paul): 147. 1983.</p>
<p>≡ <italic>Racospermyces bicinctus</italic>
(McAlpine) J. Walker, Australas. Mycol. 20: 16. 2001.</p>
<p><italic>Type.</italic>
A<sc>USTRALIA</sc>
, Queensland, Rockhampton, on <italic>A. fasciculifera</italic>
, 30 Nov. 1866, <italic>G.H. Robinson</italic>
(holotype VPRI 5751).</p>
<p>Notes — <italic>Endoraecium bicinctum</italic>
is known from four collections, which were examined by <xref rid="R32" ref-type="bibr">Walker (2001)</xref>
. It is only known to occur on <italic>A. fasciculifera</italic>
in Botrycephalae (<xref rid="R20" ref-type="bibr">Murphy et al. 2010</xref>
, <xref rid="R16" ref-type="bibr">Maslin 2013</xref>
).</p>
<p><bold><italic>Endoraecium carnegiei</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808981; <xref ref-type="fig" rid="F3">Fig. 3</xref>
</p>
<p><italic>Etymology</italic>
. Named after the Australian forest pathologist and fungal taxonomist, Dr Angus J. Carnegie, who generously assisted this study by collecting many specimens of rust on <italic>Acacia</italic>
, including the type specimen on which this taxon is based.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, New South Wales, Ando, Glen Allen State Forest, Poddy Hut Road (-36.7150, 149.3486), on <italic>A. dealbata</italic>
, 6 Oct. 2012, <italic>A.J. Carnegie</italic>
(holotype BRIP 57924).</p>
<p><italic>Uredinia</italic>
forming galls on stems, irregular, up to 3 cm, yellowish brown. <italic>Urediniospores</italic>
ellipsoidal to ovoid, subhyaline to yellow, 34–40 × 21–25 μm; wall 4.0–4.5 μm thick, uniform or slightly thicker at sides, with 6–8 equatorial germ pores. <italic>Telia</italic>
on leaves, pulverulent, erumpent, elliptical, less than 1 mm, yellowish brown. <italic>Teliospores</italic>
cylindrical to fusiform, apex rounded, 1–5-digitate, hyaline to subhyaline, 41–76 × 10–17 μm; wall 1.0–1.5 μm thick at sides, 6–19(–32) μm thick at apex; pedicel persistent, up to 51 μm.</p>
<p>On stems and leaves of <italic>A. dealbata</italic>
(Botrycephalae).</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, New South Wales, Mila, Bondi State Forest (-37.0847, 149.1078), on <italic>A. dealbata</italic>
, 4 Oct. 2012, <italic>A.J. Carnegie</italic>
, BRIP 57926; Victoria, Kergunyah, Murramurrangbong Range, Simpson Road (-36.3142, 146.9819), on <italic>A. dealbata</italic>
, 13 May 2013, <italic>C. Doungsa-ard</italic>
, <italic>W. Khem-muk</italic>
& <italic>A.D.W. Geering</italic>
, BRIP 59218.</p>
<p>Notes — <italic>Endoraecium carnegiei</italic>
is one of three species that forms galls on the stems of its host. The other two species, <italic>E. digitatum</italic>
and <italic>E. irroratum</italic>
, also infect species of <italic>Acacia</italic>
in the Botrycephalae<italic>.</italic>
<italic>Endoraecium carnegiei</italic>
is only known to occur on <italic>A. dealbata</italic>
in south-eastern Australia.</p>
<p><bold><italic>Endoraecium digitatum</italic>
</bold>
(G. Winter) M. Scholler & Aime, Mycoscience 47: 163. 2006.</p>
<p><italic>Basionym</italic>
. <italic>Uromyces digitatus</italic>
G. Winter, Rev. Mycol. (Toulouse) 8: 209. 1886.</p>
<p>≡ <italic>Coeomurus digitatus</italic>
(G. Winter) Kuntze (as ‘<italic>Caeomurus</italic>
’), Revis. Gen. Pl. 3: 450. 1898.</p>
<p>≡ <italic>Atelocauda digitata</italic>
(G. Winter) Cummins & Y. Hirats., Illustr. Gen. Rust Fungi, rev. edn (St. Paul): 147. 1983.</p>
<p>≡ <italic>Racospermyces digitatus</italic>
(G. Winter) J. Walker, Australas. Mycol. 20: 13. 2001.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, South Australia, near Gawler, on <italic>A. notabilis</italic>
, 1 July 1885, <italic>J.G.O. Tepper</italic>
(holotype SF35352); South Australia, west of Gawler, along road to Mallala (-34.5665, 138.7184), on <italic>A. notabilis</italic>
, 16 Oct. 2009, <italic>V. Faust-Berndt & R. Berndt</italic>
(epitype SF35352).</p>
<p>Notes — Our study shows that the known host range of <italic>E. digitatum</italic>
s.str. is restricted to <italic>A. notabilis</italic>
in Botrycephalae<italic>.</italic>
<xref rid="R32" ref-type="bibr">Walker (2001)</xref>
adopted a broad host range for <italic>E. digitatum</italic>
, but suspected it was a complex of closely related taxa, confined to one or a small group of hosts. <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
divided <italic>E. digitatum</italic>
into five species in Australia. He accepted <italic>E. phyllodiorum</italic>
as distinct from <italic>E. digitatum</italic>
, and further described three new taxa, <italic>E. parvum</italic>
, <italic>E. violae-faustiae</italic>
and <italic>E. walkerianum</italic>
. <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
designated an epitype for <italic>E. digitatum</italic>
on <italic>A. notabilis</italic>
, and listed six species of <italic>Acacia</italic>
as hosts. We found four of these hosts, <italic>A. dealbata</italic>
, <italic>A. falciformis</italic>
, <italic>A. irrorata</italic>
and <italic>A. podalyriifolia</italic>
, were infected by novel species of <italic>Endoraecium</italic>
. The remaining two hosts, <italic>A. deanei</italic>
and <italic>A. oshanesii</italic>
, most likely represent two new hosts of <italic>Endoraecium</italic>
. The urediniospores of <italic>E. digitatum</italic>
were not examined in this study, however, the description of the epitype (<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
) is similar to other species of <italic>Endoraecium</italic>
with subhyaline to golden yellow urediniospores and reticulate ornamentation on species of <italic>Acacia</italic>
in Botrycephalae.</p>
<p><bold><italic>Endoraecium disparrimum</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808982; <xref ref-type="fig" rid="F4">Fig. 4</xref>
</p>
<p><italic>Etymology</italic>
. Name refers to the host, <italic>Acacia disparrima</italic>
, on which it was found.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Queensland, Babinda (-17.3397, 145.8675), on <italic>A. dis-parrima</italic>
, 3 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
(holotype BRIP 55626).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, forming bullate swellings up to 1 cm long, or pulverulent, linear to round, up to 1 mm diam, yellowish brown. <italic>Urediniospores</italic>
subglobose to ovoid, apex obtuse, yellowish brown, 29–41 × 20–26 μm; wall 2–4 μm thick at sides and 5.5–9.0 μm thick at apex, reticulate, with 2–5 equatorial germ pores; pedicel 2 μm or absent. <italic>Telia</italic>
formed from uredinia, yellowish brown. <italic>Teliospores</italic>
cylindrical to fusiform, apex acuminate to rounded, hyaline to subhyaline, 31–50 × 20–26 μm; wall 1–2 μm thick at sides, 4–14 μm thick at apex; pedicel persistent, up to 33 μm.</p>
<p>On phyllodes of <italic>A. disparrima</italic>
(Juliflorae).</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, Herberton, Silver Valley Drive, on <italic>A. disparrima</italic>
, 10 Apr. 2012, <italic>R. Berndt</italic>
& <italic>V. Faust-Berndt</italic>
, BRIP 55659; Queensland, near Yungaburra, Danbulla State Forest, at the Chimney’s Rest Area, on <italic>A. disparrima</italic>
, 9 Apr. 2012, <italic>R. Berndt</italic>
& <italic>V. Faust-Berndt</italic>
, BRIP 55632.</p>
<p>Notes — <italic>Endoraecium disparrimum</italic>
occurs on <italic>A. disparrima</italic>
in Juliflorae. It was recovered as sister to <italic>E. violae-faustiae</italic>
, which occurs on closely related species of <italic>Acacia</italic>
. It differs from <italic>E. violae-faustiae</italic>
in that the urediniospores have an obtuse apex with reticulate rather than foveolate ornamentation, and paraphyses are absent.</p>
<p><bold><italic>Endoraecium falciforme</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808983; <xref ref-type="fig" rid="F5">Fig. 5</xref>
</p>
<p><italic>Etymology</italic>
. Name refers to the host, <italic>Acacia falciformis</italic>
, on which it was found.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Queensland, Mt Tibrogargan (-26.9283, 152.9494), on <italic>A. falciformis</italic>
, 15 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
(holotype BRIP 57583).</p>
<p><italic>Spermogonia</italic>
on fruit or phyllodes, amphigenous, subepidermal, erumpent, c. 100 μm. <italic>Uredinia</italic>
on fruit or on both phyllode surfaces, causing distortion, surrounding spermogonia, subepidermal, erumpent, pulverulent, linear, round to ellipsoidal, greater than 10 mm, reddish brown. <italic>Urediniospores</italic>
ellipsoidal to ovoid, subhyaline to yellow, 34–43 × 21–25 μm; wall unevenly thickened at equator, 3.0–6.5 μm and 3–5 μm thick at apex, reticulate, with 4–9 equatorial germ pores; pedicel 2 μm. <italic>Telia</italic>
on both phyllode surfaces, formed separately from uredinia, erumpent, c. 1 mm, orange to yellowish brown. <italic>Teliospores</italic>
cylindrical to clavate, apex rounded, 2–5-digitate, subhyaline, 47–63 × (13–)15–23 μm; wall 1.5–2.0 μm thick at sides, 12–24 μm thick at apex; pedicel persistent, up to 64 μm long × 6–8 μm thick.</p>
<p>On phyllodes and fruit of <italic>A. falciformis</italic>
(Botrycephalae).</p>
<p><italic>Additional specimen examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, Mt Tibrogargan, 4 Sept. 2012, <italic>K.M. Thomson</italic>
, BRIP 57643.</p>
<p>Notes — <italic>Endoraecium falciforme</italic>
is morphologically similar to <italic>E. walkerianum</italic>
, which both occur on species of <italic>Acacia</italic>
species in Botrycephalae. <italic>Endoraecium falciforme</italic>
has distinctively thicker walls at the equator of the urediniospores (3.0–6.5 μm) than <italic>E. walkerianum</italic>
(3.0–4.5 μm). <italic>Endoraecium falciforme</italic>
is recorded from south-east Queensland, whereas <italic>E. walkerianum</italic>
is known from southern Australia. Urediniospores from a paratype specimen of <italic>E. walkerianum</italic>
(BRIP 14205) were found to have thickened apices, whereas those of <italic>E. falciforme</italic>
were thickened at the equator, or the wall was of uniform thickness.</p>
<p><bold><italic>Endoraecium irroratum.</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808984; <xref ref-type="fig" rid="F6">Fig. 6</xref>
</p>
<p><italic>Etymology</italic>
. Name refers to the host, <italic>Acacia irrorata</italic>
, on which it was found.</p>
<p><italic>Type.</italic>
A<sc>USTRALIA</sc>
, Queensland, Main Range National Park, on <italic>A. irrorata</italic>
, 11 June 2012, <italic>A.D.W. Geering</italic>
(holotype BRIP 57286).</p>
<p><italic>Uredin ia</italic>
forming galls on stems, up to 2 cm long, yellowish brown. <italic>Urediniospores</italic>
ovoid, apex obtuse, subhyaline, yellow to yellowish brown, 32–42 × 17–24 μm; wall 2.0–3.5 μm thick at sides, apex sometimes thickened (up to 4 μm), reticulate, with 4–10 equatorial germ pores. <italic>Telia</italic>
formed from uredinia, yellowish brown. <italic>Teliospores</italic>
cylindrical to subfusiform, apex acuminate to rounded, 2–7-digitate, subhyaline to yellow, 47–73 × 15–22 μm; wall 1.0–1.5 μm thick at sides, 11–21 μm thick at apex; pedicel persistent, up to 46 μm.</p>
<p>On stems and phyllodes of <italic>A. irrorata</italic>
(Botrycephalae).</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, New South Wales, Warrumbungle National Park, on <italic>A. irrorata</italic>
, 20 Mar. 2012, <italic>R. Berndt</italic>
& <italic>V. Faust-Berndt</italic>
, BRIP 55671; Queensland, Mt Mee, Mt Mee State Forest, on <italic>A. irrorata</italic>
, 3 June 2012, <italic>C. Doungsa-ard</italic>
& <italic>R.G. Shivas</italic>
, BRIP 57279; Queensland, Mt Glorious, Wivenhoe Dam lookout area (-27.3077, 152.7136), <italic>A. irrorata</italic>
, 30 Nov. 2012, <italic>A.D.W. Geering</italic>
, BRIP 58054.</p>
<p>Notes — <italic>Endoraecium irroratum</italic>
occurs on <italic>A. irrorata</italic>
in Botrycephalae. <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
considered that the rust on <italic>A. irrorata</italic>
was <italic>E. digitatum</italic>
based on morphology. There are no clear morphological differences that separate <italic>E. irroratum</italic>
from the epitype of <italic>E. digitatum</italic>
(<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
). However, the results from the molecular phylogenetic analysis in this study indicate the rusts on Botrycephalae are each restricted to a single host species.</p>
<p><bold><italic>Endoraecium maslinii.</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808985; <xref ref-type="fig" rid="F7">Fig. 7</xref>
</p>
<p><italic>Etymology</italic>
. Named after the Australian botanist Bruce R. Maslin, an expert on <italic>Acacia</italic>
taxonomy, who has published approximately 250 <italic>Acacia</italic>
taxa and kindly identified many specimens in this study.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Western Australia, Midlands Rd, 10 km SE of Mingenew (-29.2847, 115.5319), on <italic>A. daphnifolia</italic>
, 28 Sept. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
(holotype BRIP 57872).</p>
<p><italic>Uredinia</italic>
forming bullate swellings on fruit and both phyllode surfaces, round, up to 1 cm, yellowish brown. <italic>Urediniospores</italic>
ellipsoidal, ovoid to fusiform, apex obtuse, subhyaline to yellowish brown, 37–46 × 20–26 μm; wall 2.5–5.0 μm thick at side, apex slightly thickened, 3.5–6.0 μm, reticulate, with 6–8 equatorial germ pores; pedicel absent or up to 2.5 μm.</p>
<p>On phyllodes of <italic>A. daphnifolia</italic>
(Botrycephalae).</p>
<p>Notes — <italic>Endoraecium maslinii</italic>
occurs on <italic>A. daphnifolia</italic>
in Botrycephalae and is the only species of <italic>Endoraecium</italic>
known from Western Australia. Only urediniospores have been found, which are yellowish brown and reticulate, and morphologically similar to other species on <italic>Acacia</italic>
in Botrycephalae.</p>
<p><bold><italic>Endoraecium parvum</italic>
</bold>
Berndt, Mycol. Progr. 10: 510. 2011</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Queensland, Caloundra, on <italic>A. leiocalyx</italic>
, 25 Aug. 1932, <italic>S.T. Blake</italic>
(holotype BRIP 7543!).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, subepidermal, erumpent, pulverulent, linear to round, yellowish brown. <italic>Urediniospores</italic>
globose, subglobose to obovoid, apex obtuse, yellowish brown to reddish brown, 25–38 × 16–22 μm; wall 3–4 μm thick at equator, hardly thickened at apex, reticulate, with 2–4 equatorial germ pores, pedicel 1.5–3.0 μm. <italic>Telia</italic>
formed from uredinia, erumpent, linear to dome-shaped, reddish brown. <italic>Teliospores</italic>
oval, apex rounded, 1–3-digitate, hyaline to subhyaline, 31–60 × 13–27 μm; wall 1–2 μm thick, mostly thickened at apex (5–14 μm); pedicel persistent, up to 66 μm.</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, Caloundra, on <italic>A. leiocalyx</italic>
, 25 Aug. 1932, <italic>S.T. Blake</italic>
, BRIP 7543; Queensland, Mt Coolum (-26.5656, 153.0969), on <italic>A. leiocalyx</italic>
, 28 July 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57512; Queensland, Mt Coolum (-26.5622, 153.0942), on <italic>A. leiocalyx</italic>
, 28 July 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57514; Queensland, Cunningham Highway (-28.0317, 152.4697), on <italic>A. leiocalyx</italic>
, 1 Aug. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>A.D.W. Geering</italic>
& <italic>R.G. Shivas</italic>
, BRIP 57524; Queensland, Lake Moogerah Road (-28.0247, 152.5189), on <italic>A. leiocalyx</italic>
, 1 Aug. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>A.D.W. Geering</italic>
& <italic>R.G. Shivas</italic>
, BRIP 57527; Queensland, West Haldon (-27.7722, 152.0903), on <italic>A. leiocalyx</italic>
, 1 Aug. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>A.D.W. Geering</italic>
& <italic>R.G. Shivas</italic>
, BRIP 57534; Queensland, Brisbane, Geebung (-27.3375, 153.0511), on <italic>A. leiocalyx</italic>
, 2 Sept. 2012, <italic>R.G.</italic>
& <italic>M.D.E. Shivas</italic>
, BRIP 57631; New South Wales, Shark Creek (-29.5664, 153.2003), on <italic>A. leiocalyx</italic>
, 13 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57568.</p>
<p>Notes — <italic>Endoraceium parvum</italic>
is distinguished from other species by the comparatively short (25–38 μm) urediniospores (<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
). <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
described <italic>E. parvum</italic>
from <italic>A. leiocalyx</italic>
(as <italic>A. concurrens</italic>
) and <italic>A. mangium</italic>
. Specimens of rust on <italic>A. mangium</italic>
were not included in this study and these warrant further study.</p>
<p><bold><italic>Endoraecium peggii</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808986; <xref ref-type="fig" rid="F8">Fig. 8</xref>
</p>
<p><italic>Etymology</italic>
. Named after the Australian forest pathologist and rust specialist, Dr Geoff S. Pegg, who has discovered several new plant pathogens in Australia.</p>
<p><italic>Type.</italic>
A<sc>USTRALIA</sc>
, Northern Territory, Darwin, 8 km from airport, on <italic>A. holosericea</italic>
, 7 May 2012, <italic>R.G. Shivas</italic>
(holotype BRIP 55602).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, pulverulent, subepidermal, erumpent, linear to round, up to 2 mm, yellowish brown; paraphyses intrasoral, cylindrical, yellow, with pedicel, 37–84 × 9–10 μm, thickened at apex, digitate. <italic>Urediniospores</italic>
ovoid, apex obtuse, yellowish brown to reddish brown, 31–45 × 20–25 μm; wall 3–5 μm thick at sides, apex 4–10 μm thick, foveolate to reticulate, with 2–3 equatorial germ pores; pedicel 3–6 μm. <italic>Telia</italic>
rare, on both surfaces of phyllode, darker than uredinia, erumpent, linear, brown to black. <italic>Teliospores</italic>
oval, apex rounded, no digitations, hyaline to subhyaline, 45–60 × (15–)19–27 μm; wall 1.5–2.0 μm thick at sides, sometimes thickened at apex, 7–17 μm; pedicel persistent, up to 52 μm.</p>
<p>On phyllodes of <italic>A. holosericea</italic>
(Juliflorae).</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, Dimbulah, Bourke Developmental Road, on <italic>A. holosericea</italic>
, 6 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
, BRIP 55631; Queensland, Brisbane, Mt Coot-tha Botanic Gardens (-27.477655, 152.972270), on <italic>A. holosericea</italic>
, Nov. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 58324.</p>
<p>Notes — <italic>Endoraecium peggii</italic>
is known from <italic>A. holosericea</italic>
in Juliflorae. This species of <italic>Acacia</italic>
is native to northern Australia (<xref rid="R16" ref-type="bibr">Maslin 2013</xref>
). The rust was also collected in south-east Queensland on a plant in the Mt Coot-tha Botanic Gardens. The teliospores of <italic>E. peggii</italic>
lack the apical digitations found in all other species of <italic>Endoraecium</italic>
, although this ornamentation is present on the paraphyses.</p>
<p><bold><italic>Endoraecium phyllodiorum</italic>
</bold>
(Berk. & Broome) Berndt, Mycol. Progr. 10: 503. 2011</p>
<p><italic>Basionym</italic>
. <italic>Melampsora phyllodiorum</italic>
Berk. & Broome, Trans. Linn. Soc. London 2, ser. 2: 67. 1883.</p>
<p>≡ <italic>Uromyces phyllodiorum</italic>
(Berk. & Broome) McAlpine, The Rusts of Australia: 95. 1906. (<xref rid="R17" ref-type="bibr">McAlpine (1906)</xref>
described teliospores from the type specimen, which were overlooked by <xref rid="R2" ref-type="bibr">Berkeley & Broome (1883)</xref>
. Under previous rules of nomenclature, <italic>U. phyllodiorum</italic>
was considered the name of a new species with a teleomorphic type (<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
). Under the current system of nomenclature, the name is cited as originally published, and is typified by the type of the basionym.)</p>
<p>≡ <italic>Pileolaria phyllodiorum</italic>
(Berk. & Broome) Dietel, Ann. Mycol. 19: 302. 1921.</p>
<p>= <italic>Uromyces phyllodii</italic>
Cooke & Massee (as ‘<italic>phyllodiae</italic>
’), in Cooke, Grevillea 17: 70. 1889.</p>
<p>≡ <italic>Coeomurus</italic>
<italic>phyllodii</italic>
(Cooke & Massee) Kuntze (as <italic>‘Caeomurus’</italic>
), Revis. Gen. Pl. 3: 450. 1898.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Queensland, Brisbane, on <italic>Acacia</italic>
sp., <italic>F.M. Bailey</italic>
no. 269, K(M) 146703.</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, subepidermal, erumpent, pulverulent or on bullate swellings, linear, round to irregular, yellowish brown. <italic>Urediniospores</italic>
subglobose to oval, apex obtuse, yellowish brown, (28–)30–52 × 17–26 μm; wall 2.5–4.0 μm thick at sides, 5–10 μm thick at apex, reticulate, with 3–7 equatorial germ pores; pedicel absent or 1.5–3.5 μm. <italic>Telia</italic>
form-ed from uredinia, erumpent, linear, reddish brown. <italic>Teliospores</italic>
cylindrical to oval, apex rounded, 1–4-digitate, hyaline to subhyaline, 38–58 × 17–28 μm; wall 1–2 μm thick at sides, 6–17 μm thick at apex; pedicel persistent, up to 30 μm.</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, Mount Coolum (-26.5611, 153.0839), on <italic>A. aulacocarpa</italic>
, 28 July 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57310; Queensland, Mt Coolum (-26.5611, 153.0839), on <italic>A. aulacocarpa</italic>
, 28 July 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57516; New South Wales, Woombah, at the beginning of Iluka Road (-29.3544, 153.2492), on <italic>A. aulacocarpa</italic>
, 11 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57578; New South Wales, Woombah, at the beginning of Iluka Road (-29.3544, 153.2492), on <italic>A. aulacocarpa</italic>
, 11 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57580; New South Wales, Woombah, Iluka Bluff (-29.3967, 153.3722), on <italic>A. aulacocarpa</italic>
, 11 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57579; New South Wales, Shark Creek (-29.5664, 153.2003), on <italic>A. aulacocarpa</italic>
, 13 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57569; New South Wales, Shark Creek (-29.5664, 153.2003), on <italic>A. aulacocarpa</italic>
, 13 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57570; Queensland, Mt Tibrogargan (-26.9264, 152.9417), on <italic>A. aulacocarpa</italic>
, 15 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57588; Queens-land, Mt Tibrogargan (-26.9258, 152.9508), on <italic>A. aulacocarpa</italic>
, 15 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57589; Queensland, Mt Tibrogargan (-26.9278, 152.9378), on <italic>A. aulacocarpa</italic>
, 15 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
, BRIP 57590.</p>
<p>Notes — <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
discussed the taxonomy of <italic>E. phyllodiorum,</italic>
and listed the hosts as <italic>A. aulacocarpa</italic>
, <italic>A. crassicarpa, A. holosericea</italic>
, <italic>A. mangium</italic>
and tentatively <italic>A. auriculiformis</italic>
. The type specimens of <italic>Melampsora</italic>
<italic>phyllodiorum</italic>
and <italic>Uromyces</italic>
<italic>phyllodii</italic>
were collected on unidentified species of <italic>Acacia</italic>
in Brisbane, Australia. <italic>Acacia</italic>
<italic>aulacocarpa</italic>
is the only host species listed by <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
that occurs naturally in or near to Brisbane. The other four species of <italic>Acacia</italic>
are restricted to northern Queensland and the Northern Territory (<xref rid="R16" ref-type="bibr">Maslin 2013</xref>
). Herbarium records and field observations by the authors show that <italic>Endoraecium</italic>
spp. occur on four species, <italic>A. aulacocarpa</italic>
, <italic>A. irrorata</italic>
, <italic>A. leiocalyx</italic>
and <italic>A. podalyriifolia</italic>
, in the Brisbane region. The rust on <italic>A. aulacocarpa</italic>
forms bullate swellings on phyllodes, very similar to those seen in the type specimen of <italic>E. phyllodiorum</italic>
and illustrated by <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
. Consequently, we suggest that <italic>A. aulacocarpa</italic>
is the host of the type of <italic>E. phyllodiorum</italic>
. <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
proposed that <italic>E. phyllodiorum</italic>
had a wide host range, including <italic>A. aulacocarpa</italic>
, <italic>A. auriculiformis</italic>
, <italic>A. holosericea</italic>
and <italic>A. mangium</italic>
. The rusts on <italic>A. auriculiformis</italic>
and <italic>A. holosericea</italic>
are now identified as <italic>E. auriculiforme</italic>
and <italic>E. peggii</italic>
, respectively. The rust on <italic>A. mangium</italic>
is unknown. <italic>Acacia aulacocarpa</italic>
is the only species of <italic>Acacia</italic>
known to host two species of <italic>Endoraecium</italic>
, <italic>E. phyllodiorum</italic>
and <italic>E. violae-faustiae</italic>
.</p>
<p><bold><italic>Endoraecium podalyriifolium</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808987; <xref ref-type="fig" rid="F9">Fig. 9</xref>
</p>
<p><italic>Etymology</italic>
. Name refers to the host, <italic>Acacia podalyriifolia</italic>
, on which it was found.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, New South Wales, Maclean, Wharf Street (-29.4589, 153.2111), on <italic>A. podalyriifolia</italic>
, 12 Aug. 2012, <italic>C. Doungsa-ard</italic>
& <italic>A.R. McTaggart</italic>
(holotype BRIP 57576).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, subepidermal, erumpent, pulverulent, round, 0.5–10.0 mm, yellow to yellowish brown. <italic>Urediniospores</italic>
cylindrical to ovoid, apex obtuse, subhyaline to yellow, (30–)33–44 × 23–27 μm; wall 3–4 μm thick, reticulate, with 4–6 germ pores. <italic>Telia</italic>
formed from uredinia, subepidermal, erumpent, yellow. <italic>Teliospores</italic>
cylindrical to fusiform, apex rounded, 2–5-digitate, subhyaline to yellow, 44–60 × 12–19 μm; wall 1 μm thick at sides, 6–12 μm thick at apex; pedicel persistent, over 60 μm.</p>
<p>On phyllodes of <italic>A. podalyriifolia</italic>
(Botrycephalae).</p>
<p><italic>Additional specimen examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, South Ripley, Ripley Road, on <italic>A. podalyriifolia</italic>
, 17 July 2012, <italic>A.D.W. Geering</italic>
, BRIP 57294.</p>
<p>Notes — <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
considered the rust on <italic>A. podalyriifolia</italic>
was <italic>E. digitatum</italic>
based on morphology and the close relationships of host species within this group. However, the results from the molecular phylogenetic analysis in this study indicate the rusts on Botrycephalae are each restricted to a single host species.</p>
<p><bold><italic>Endoraecium tierneyi</italic>
</bold>
(J. Walker & R.G. Shivas) M. Scholler & Aime, Mycoscience 47: 163. 2006</p>
<p><italic>Basionym</italic>
. <italic>Racospermyces tierneyi</italic>
J. Walker & R.G. Shivas, in Walker, Australas. Mycol. 20: 23. 2001.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Queensland, Tambo, near Castlevale, on <italic>A. harpophylla</italic>
, 6 June 2000, <italic>G.S. Pegg</italic>
(holotype BRIP 27071!).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, subepidermal, erumpent, linear, up to 500 μm, yellowish brown. <italic>Urediniospores</italic>
cylindrical, oval to broadly fusiform, apex obtuse, reddish brown, 26–36 × 13–18 μm; wall 2.0–2.5 μm thick at sides, 4.0–4.5 μm thick at apex, reticulate, with 4–8 equatorial germ pores. <italic>Telia</italic>
on both phyllode surfaces, erumpent. <italic>Teliospores</italic>
subglobose, ovoid to obovoid, apex rounded, reddish brown, darker at the apex, 27–41 × 20–26 μm; wall 1.0 μm thick at sides, 7–13 μm thick at apex, smooth; pedicel persistent, 50–55 μm long.</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, Springsure, on <italic>A. harpophylla</italic>
, 27 Feb. 2001, <italic>G.S. Pegg</italic>
, BRIP 27887; Queensland, Caldervale, on <italic>A. harpophylla</italic>
, 1 Mar. 2001, <italic>G.S. Pegg</italic>
, BRIP 27880.</p>
<p>Notes — <italic>Endoraecium tierneyi</italic>
occurs on <italic>A. harpophylla</italic>
in Plurinerves. It was recovered in the phylogenetic analyses as sister to three species of <italic>Endoraecium</italic>
in Hawaii that occur on <italic>A. koa</italic>
, also in Plurinerves. The teliospores are subglobose to obovoid, while other species of <italic>Endoraecium</italic>
in Australia have clavate teliospores.</p>
<p><bold><italic>Endoraecium tropicum</italic>
</bold>
McTaggart & R.G. Shivas, <italic>sp. nov.</italic>
— MycoBank MB808988; <xref ref-type="fig" rid="F10">Fig. 10</xref>
</p>
<p><italic>Etymology</italic>
. Name refers to the host, <italic>Acacia tropica</italic>
, on which it was found.</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Northern Territory, Gregory, Victoria Highway (-15.6003, 131.2136), on <italic>A. tropica</italic>
, 20 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
(holotype BRIP 56557).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, subepidermal, pulverulent, up to 2 mm, brown. <italic>Urediniospores</italic>
ovoid to fusiform, apex obtuse, yellowish brown, 30–40 × 17–22 μm; wall 3–4 μm thick at sides, apex usually 3–4 μm thick, sometimes slightly thickened (4–5 μm), reticulate, with 3–4, equatorial germ pores; pedicel 3–4 μm. <italic>Telia</italic>
formed from uredinia, erumpent. <italic>Teliospores</italic>
clavate, apex rounded, papillate rather than digitate, 40–53 × 16–23 μm; wall 1.5–2.0 μm thick at sides, 8–11 μm thick at apex; pedicel persistent, 5 μm.</p>
<p>On phyllodes of <italic>A. tropica</italic>
(Juliflorae).</p>
<p><italic>Additional specimen examined</italic>
. A<sc>USTRALIA</sc>
, Northern Territory, Gregory, Victoria Highway (-15.6014, 131.2136), on <italic>A. tropica</italic>
, 20 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
, BRIP 56555.</p>
<p>Notes — <italic>Endoraecium tropicum</italic>
was closely related to <italic>E. par-vum</italic>
in the phylogenetic analysis. Both species have urediniospores with uniformly thickened spore walls. The distribution of <italic>A. tropica</italic>
is restricted to the Northern Territory and north-west Queensland. <italic>Acacia leiocalyx</italic>
, the host of <italic>E. parvum</italic>
, occurs in eastern Australia (<xref rid="R16" ref-type="bibr">Maslin 2013</xref>
).</p>
<p><bold><italic>Endoraecium violae-faustiae</italic>
</bold>
Berndt, (as ‘<italic>violae-faustae</italic>
’) Mycol. Progr. 10: 513. 2011</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, Queensland, Cairns, Barron Gorge, MacDonald’s trail, on <italic>A. crassicarpa</italic>
(as <italic>Acacia</italic>
sp.), 18 Aug. 2006, <italic>V. Faust-Berndt & R. Berndt</italic>
(holotype BRIP 53388!).</p>
<p><italic>Uredinia</italic>
on both phyllode surfaces, subepidermal, erumpent, pulverulent, linear to round, yellowish brown, not forming bullate sori; paraphyses cylindrical with digitate heads, subhyaline to yellow, 35–88 × 6–15 μm thick at sides, thickened at apex. <italic>Urediniospores</italic>
globose, subglobose, obovoid to oval, apex acute, yellowish brown, 28–53 × 17–26 μm; wall 2.5–4.5 μm thick at sides, apex mostly thickened, 5.0–10.5 μm thick, foveolate, with 3–5 equatorial germ pores; pedicel 3–5 μm. <italic>Telia</italic>
formed from uredinia, erumpent, reddish brown. <italic>Teliospores</italic>
obovoid to oval, apex rounded, 1–4-digitate, hyaline to subhyaline, 41–53 × 20–28 μm; wall 1.5–3.0 μm thick at sides, 5–14 μm thick at apex; pedicel persistent, up to 43 μm.</p>
<p><italic>Additional specimens examined</italic>
. A<sc>USTRALIA</sc>
, Queensland, Coen, Peach No. 7, on <italic>A. crassicarpa</italic>
18 July 1999, <italic>R.G. Shivas & M. Gunther</italic>
, paratype BRIP 25816; Queensland, Julatten, Abbatoir Swamp (-16.6074, 145.3428), on <italic>A. aulacocarpa</italic>
, 5 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
, BRIP 55601; Queensland, Mareeba (-16.9414, 145.5411), on <italic>A. difficilis</italic>
, 6 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. Mc-Taggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
, BRIP 55611; Queensland, Innisfail (-17.5003, 146.0756), on <italic>A. difficilis</italic>
, 7 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
, BRIP 55616; Queensland, Dimbulah (-17.2967, 144.9736), on <italic>A. aula-cocarpa</italic>
, 6 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
, <italic>V. Faust-Berndt</italic>
, <italic>M.D.E.</italic>
& <italic>R.G. Shivas</italic>
, BRIP 55629; Northern Territory, Humpty Doo (-12.5964, 131.2083), on <italic>A. difficilis</italic>
, 22 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
& <italic>R.G. Shivas</italic>
, BRIP 56539; Northern Territory, Humpty Doo (-12.5964, 131.2083), on <italic>A. difficilis</italic>
, 22 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
& <italic>R.G. Shivas</italic>
, BRIP 56540; Northern Territory, Humpty Doo (-12.5964, 131.2083), on <italic>A. difficilis</italic>
, 22 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
& <italic>R.G. Shivas</italic>
, BRIP 56545; Northern Territory, Humpty Doo (-12.5964, 131.2083), on <italic>A. difficilis</italic>
, 22 Apr. 2012, <italic>C. Doungsa-ard</italic>
, <italic>A.R. McTaggart</italic>
, <italic>R. Berndt</italic>
& <italic>R.G. Shivas</italic>
, BRIP 56547.</p>
<p>Notes — <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
described the urediniospores of <italic>E. violae-faustiae</italic>
as foveolate, with a thickened and acute apex. The most distinctive character was the variable paraphyses, which resembled thin teliospores, similar to those found in <italic>E. auriculiforme</italic>
and <italic>E. peggii</italic>
. <italic>Endoraecium violae-faustiae</italic>
was reported on <italic>A. aulacocarpa</italic>
and <italic>A. crassicarpa</italic>
, also in Juli-florae, as well as <italic>A. flavescens</italic>
in Plurinerves (<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
). The host of the holotype (BRIP 53388), was re-identified as <italic>A. crassicarpa</italic>
, which is closely related to <italic>A. aulacocarpa</italic>
(<xref rid="R16" ref-type="bibr">Maslin 2013</xref>
). The host plant of the specimen on <italic>A. flavescens</italic>
(BRIP 25816) was re-identified as <italic>A. crassicarpa</italic>
. The distribution of <italic>E. violae-faustiae</italic>
is restricted to the northern parts of Australia on <italic>A. aulacocarpa</italic>
, <italic>A. crassicarpa</italic>
and <italic>A. difficilis</italic>
.</p>
<p><bold><italic>Endoraecium walkerianum</italic>
</bold>
Berndt, Mycol. Progr. 10: 509. 2011</p>
<p><italic>Type</italic>
. A<sc>USTRALIA</sc>
, New South Wales, Cavan Gap near Yass, on <italic>A. penninervis</italic>
, 20 Nov. 1951, <italic>E. Gauba</italic>
(holotype SF35354).</p>
<p><italic>Additional specimen examined</italic>
. A<sc>USTRALIA</sc>
, New South Wales, Bald Rock, on <italic>A. penninervis</italic>
, 26 Feb. 1984, <italic>J.W. Tierney</italic>
, paratype BRIP 14205.</p>
<p>Notes — <italic>Endoraecium walkerianum</italic>
was reported from <italic>A. penninervis</italic>
and <italic>A. obliquinervia</italic>
(<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
) in Botrycephalae (<xref rid="R16" ref-type="bibr">Maslin 2013</xref>
).</p>
<sec id="s4a"><title>A KEY TO THE KNOWN SPECIES OF ENDORAECIUM IN AUSTRALIA</title>
<p>1. Urediniospores hyaline to pale yellow (on Botrycephalae) .................................................. 2</p>
<p>1. Urediniospores golden or reddish brown ....................... 7</p>
<p>2. Urediniospore wall up to 5 μm or thicker at sides ................ 3</p>
<p>2. Urediniospore wall 2.0–4.5 μm thick at sides ......................... 4</p>
<p>3. Forming galls on stems of <italic>A. notabilis ......................... E. digitatum</italic>
</p>
<p>3. Forming hypertrophied lesions on <italic>A. falciformis ................................................. E. falciforme</italic>
</p>
<p>3. Forming hypertrophied lesions on <italic>A. daphnifolia ..........................................E. maslinii</italic>
</p>
<p>4. Forming hypertrophied lesions on phyllodes ......................... 5</p>
<p>4. Forming galls on stem ......................... 6</p>
<p>5. On <italic>A. fasciculifera ......................... E. bicinctum</italic>
</p>
<p>5. On <italic>A. podalyriifolia ......................... E. podalyriifolium</italic>
</p>
<p>5. On <italic>A. penninervis ......................... E. walkerianum</italic>
</p>
<p>6. On <italic>A. irrorata ......................... E. irroratum</italic>
</p>
<p>6. On <italic>A. dealbata ......................... E. carnegiei</italic>
</p>
<p>7. Teliospores globose to ellipsoid on <italic>A. harpophylla .................................................. E. tierneyi</italic>
</p>
<p>7. Teliospores clavate to obovoid or absent (on Juliflorae) 8</p>
<p>8. Urediniospore wall uniform ......................... 9</p>
<p>8. Urediniospore with thickened apex ......................... 10</p>
<p>9. Teliospores digitate, on <italic>A. leiocalyx ......................... E. parvum</italic>
</p>
<p>9. Teliospores not digitate, on <italic>A. tropica .........................E. tropicum</italic>
</p>
<p>10. Urediniospores foveolate, paraphyses present ......................... 11</p>
<p>10. Urediniospores reticulate, paraphyses absent ......................... 13</p>
<p>11. Urediniospores with an acute apex ......................... 12</p>
<p>11. Urediniospores not acute, on <italic>A. holosericea ......................... E. peggii</italic>
</p>
<p>12. On <italic>A. auriculiformis ......................... E. auriculiforme</italic>
</p>
<p>12. On <italic>A. aulacocarpa</italic>
, <italic>A. crassicarpa</italic>
or <italic>A. difficilis ............................................... E. violae-faustiae</italic>
</p>
<p>13. On <italic>A. disparrima ......................... E. disparrimum</italic>
</p>
<p>13. On <italic>A. aulacocarpa ......................... E. phyllodiorum</italic>
</p>
</sec>
</sec>
<sec id="s5"><title>DISCUSSION</title>
<p>Phylogenetic analyses of loci from nuclear rDNA and mitochondrial DNA showed that species of <italic>Endoraecium</italic>
in Australia have narrow host ranges. <xref rid="R24" ref-type="bibr">Savile (1971)</xref>
hypothesised that rusts speciated either by divergence with their hosts or by host jumps. The close relationship between <italic>Endoraecium</italic>
and species/subclades of <italic>Acacia</italic>
observed in this study indicates that <italic>Endoraecium</italic>
diversified by co-evolution with its hosts. For example, the five species of <italic>Endoraecium</italic>
on Botrycephalae, namely, <italic>E. carnegiei</italic>
, <italic>E. falciforme</italic>
, <italic>E. irroratum</italic>
, <italic>E. maslinii</italic>
and <italic>E. podalyriifolium</italic>
, were closely related with few differences in the studied genes and short branch lengths between species in the recovered phylogenetic trees. Species of <italic>Acacia</italic>
in Botrycephalae diversified c. 3.4–3.8 million years ago (<xref rid="R19" ref-type="bibr">Miller et al. 2013</xref>
), which would represent the maximum age of the corresponding <italic>Endoraecium</italic>
species.</p>
<p><italic>Endoraecium tierneyi</italic>
on <italic>A. harpophylla</italic>
in Australia was recovered as sister to the Hawaiian species, <italic>E. acaciae</italic>
, <italic>E. hawaiiense</italic>
and <italic>E. koae</italic>
, on <italic>A. koa</italic>
. These Australian and Hawaiian rusts all occur on species of <italic>Acacia</italic>
in Plurinerves. <xref rid="R12" ref-type="bibr">Hodges & Gardner (1984)</xref>
and <xref rid="R32" ref-type="bibr">Walker (2001)</xref>
proposed that the Australian species of <italic>Endoraecium</italic>
were the ancestor of the Hawaiian rusts. Our study supports this view by showing the rusts on Plurinerves had an Australian ancestor (plesiomorphic state). Further, the three rusts on <italic>Acacia</italic>
in Hawaii have differences in life cycle and morphology, but are identical in the LSU region of rDNA. These rusts have diversified on <italic>A. koae</italic>
after its relatively recent split from Australian species of <italic>Acacia</italic>
(<xref rid="R4" ref-type="bibr">Brown et al. 2012</xref>
). Endocyclic species occur in Hawaii but not in Australia, which shows that a reduced life cycle is a derived character in <italic>Endoraecium</italic>
.</p>
<p><italic>Endoraecium hyalosporum</italic>
from South-East Asia infects <italic>A. confusa</italic>
, which is the only example of a species of <italic>Endoraecium</italic>
on a host outside of the p.u.b. clade of <italic>Acacia</italic>
(<xref rid="R4" ref-type="bibr">Brown et al. 2012</xref>
). This may represent an example of a host jump. However the systematic position of <italic>E. hyalosporum</italic>
is unknown, and it is possible that it does not have a close phylogenetic relationship to other species of <italic>Endoraecium</italic>
. It is noteworthy that <italic>E</italic>
. <italic>hyalosporum</italic>
has been assigned to other genera at various times, namely, <italic>Maravalia</italic>
(<xref rid="R8" ref-type="bibr">Dietel 1924</xref>
), <italic>Poliotelium</italic>
(<xref rid="R15" ref-type="bibr">Mains 1939</xref>
) and <italic>Atelocauda</italic>
(<xref rid="R22" ref-type="bibr">Ono 1984</xref>
).</p>
<p>The morphology of urediniospores may be synapomorphic for the three clades of <italic>Endoraecium</italic>
. For example, the rusts on Juliflorae have reticulate to foveolate ornamentation on golden-brown urediniospores. The rusts on Botrycephalae have a raised reticulate ornamentation on subhyaline to pale yellow urediniospores. <italic>Endoraecium tierneyi</italic>
was the only rust examined on Plurinerves, and it had reticulate, reddish brown urediniospores. <xref rid="R3" ref-type="bibr">Berndt (2011)</xref>
noted the urediniospores of species of <italic>Endoraecium</italic>
on <italic>A. koa</italic>
have similar morphology to the rusts on Juliflorae, such as <italic>E. phyllodiorum</italic>
. We cannot propose any apomorphies for the rusts on Plurinerves at this stage. The rusts on Juliflorae, namely <italic>E. auriculiforme</italic>
, <italic>E. disparrimum</italic>
, <italic>E. peggii</italic>
, <italic>E. phyllodiorum</italic>
and <italic>E. violae-faustiae</italic>
, have urediniospores with thickened apices, whereas their sister rusts <italic>E. parvum</italic>
and <italic>E. tropicum</italic>
, also in Juliflorae, have uniformly thickened spore walls.</p>
<p>Other morphological characters were less useful for species delimitation in <italic>Endoraecium</italic>
. In general, the teliospores of rusts on species of <italic>Acacia</italic>
in Botrycephalae had numerous (up to 15) apical digitations and were clavate. The teliospores of rusts on species of <italic>Acacia</italic>
in Juliflorae had fewer (up to four) apical digitations and were obovoid. The presence of a gall or bullate swelling is not a useful character for species delimitation. For example, the sori of <italic>E. auriculiforme</italic>
and <italic>E. phyllodiorum</italic>
may occur on bullate swellings or on undistorted phyllodes (<xref rid="R3" ref-type="bibr">Berndt 2011</xref>
).</p>
<p><italic>Endoraecium</italic>
is likely to show further diversity. Only 14 species of <italic>Acacia</italic>
were sampled in this study. Other species such as <italic>A. glaucocarpa</italic>
(Botrycephalae), <italic>A. juliflorae</italic>
(Juliflorae) and <italic>A. viscidula</italic>
(Plurinerves) were observed by the authors as hosts for potentially new species of <italic>Endoraecium</italic>
. New taxa will need to be classified by a combined morphological, ecological (host range) and molecular approach. We predict that many new species of <italic>Endoraecium</italic>
, which have diversified by coevolution with their <italic>Acacia</italic>
hosts, remain to be found in Australia.</p>
</sec>
</body>
<back><ack><p>This work was funded by the Australian Biological Resources Study, grant number RFL212-33. CD would like to acknowledge the support of the Australian Government’s Cooperative Research Centres Program (Project No. PBCRC62081). Thank you to the anonymous reviewers whose suggestions helped improve this manuscript.</p>
</ack>
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<floats-group><table-wrap id="T1" orientation="portrait" position="float"><label>Table 1</label>
<caption><p>Specimens of <italic>Endoraecium</italic>
included in this study.</p>
</caption>
<table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">Taxon</th>
<th align="left" rowspan="1" colspan="1">Accession number</th>
<th align="left" rowspan="1" colspan="1">Subclade<xref ref-type="table-fn" rid="tfn2"><sup>1</sup>
</xref>
of <italic>Acacia</italic>
s.str.</th>
<th align="left" rowspan="1" colspan="1">Host</th>
<th colspan="4" align="center" rowspan="1">GenBank accession<hr></hr>
</th>
</tr>
<tr><th align="left" rowspan="1" colspan="1"></th>
<th align="left" rowspan="1" colspan="1"></th>
<th align="left" rowspan="1" colspan="1"></th>
<th align="left" rowspan="1" colspan="1"></th>
<th align="left" rowspan="1" colspan="1">LSU</th>
<th align="left" rowspan="1" colspan="1">ITS</th>
<th align="left" rowspan="1" colspan="1">SSU</th>
<th align="left" rowspan="1" colspan="1">CO3</th>
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<tbody><tr><td align="left" rowspan="1" colspan="1"><italic>E. acaciae</italic>
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<td align="left" rowspan="1" colspan="1">BPI 871098</td>
<td align="left" rowspan="1" colspan="1">Plurinerves</td>
<td align="left" rowspan="1" colspan="1"><italic>A. koa</italic>
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<td align="left" rowspan="1" colspan="1">DQ323916<xref ref-type="table-fn" rid="tfn3"><sup>2</sup>
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<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">DQ323917<xref ref-type="table-fn" rid="tfn3"><sup>2</sup>
</xref>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
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<tr><td align="left" rowspan="1" colspan="1"><italic>E. auriculiforme</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 55609</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. auriculiformis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862296</td>
<td align="left" rowspan="1" colspan="1">KJ862353</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862430</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56550</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. auriculiformis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862297</td>
<td align="left" rowspan="1" colspan="1">KJ862354</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862431</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56548<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. auriculiformis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862298</td>
<td align="left" rowspan="1" colspan="1">KJ862355</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862432</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56549</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. auriculiformis</italic>
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<td align="left" rowspan="1" colspan="1">KJ862299</td>
<td align="left" rowspan="1" colspan="1">KJ862356</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862433</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. carnegiei</italic>
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<td align="left" rowspan="1" colspan="1">BRIP 57926</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. dealbata</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862300</td>
<td align="left" rowspan="1" colspan="1">KJ862357</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862434</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57924<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. dealbata</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862301</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862435</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 59218</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. dealbata</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862302</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. disparrimum</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 55659</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. disparrima</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862303</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862402</td>
<td align="left" rowspan="1" colspan="1">KJ862436</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 55626<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. disparrima</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862304</td>
<td align="left" rowspan="1" colspan="1">KJ862358</td>
<td align="left" rowspan="1" colspan="1">KJ862403</td>
<td align="left" rowspan="1" colspan="1">KJ862437</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 55632</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. disparrima</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862305</td>
<td align="left" rowspan="1" colspan="1">KJ862359</td>
<td align="left" rowspan="1" colspan="1">KJ862404</td>
<td align="left" rowspan="1" colspan="1">KJ862438</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. falciforme</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 57583<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. falciformis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862306</td>
<td align="left" rowspan="1" colspan="1">KJ862360</td>
<td align="left" rowspan="1" colspan="1">KJ862405</td>
<td align="left" rowspan="1" colspan="1">KJ862439</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57643</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. falciformis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862307</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. hawaiiense</italic>
</td>
<td align="left" rowspan="1" colspan="1">BPI 871064</td>
<td align="left" rowspan="1" colspan="1">Plurinerves</td>
<td align="left" rowspan="1" colspan="1"><italic>A. koa</italic>
</td>
<td align="left" rowspan="1" colspan="1">DQ323920<xref ref-type="table-fn" rid="tfn3"><sup>2</sup>
</xref>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. irroratum</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 55671</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. irrorata</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862310</td>
<td align="left" rowspan="1" colspan="1">KJ862364</td>
<td align="left" rowspan="1" colspan="1">KJ862406</td>
<td align="left" rowspan="1" colspan="1">KJ862442</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57279</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. irrorata</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862311</td>
<td align="left" rowspan="1" colspan="1">KJ862365</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862443</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57286<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. irrorata</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862312</td>
<td align="left" rowspan="1" colspan="1">KJ862366</td>
<td align="left" rowspan="1" colspan="1">KJ862407</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 58054</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. irrorata</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862313</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. koae</italic>
</td>
<td align="left" rowspan="1" colspan="1">BPI 871071</td>
<td align="left" rowspan="1" colspan="1">Plurinerves</td>
<td align="left" rowspan="1" colspan="1"><italic>A. koa</italic>
</td>
<td align="left" rowspan="1" colspan="1">DQ323918<xref ref-type="table-fn" rid="tfn3"><sup>2</sup>
</xref>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">DQ323919<xref ref-type="table-fn" rid="tfn3"><sup>2</sup>
</xref>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. maslinii</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 57872<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. daphnifolia</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862314</td>
<td align="left" rowspan="1" colspan="1">KJ862367</td>
<td align="left" rowspan="1" colspan="1">KJ862408</td>
<td align="left" rowspan="1" colspan="1">KJ862444</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. parvum</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 57514</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862315</td>
<td align="left" rowspan="1" colspan="1">KJ862368</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57524</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862316</td>
<td align="left" rowspan="1" colspan="1">KJ862369</td>
<td align="left" rowspan="1" colspan="1">KJ862409</td>
<td align="left" rowspan="1" colspan="1">KJ862445</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57527</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862317</td>
<td align="left" rowspan="1" colspan="1">KJ862370</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57534</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862318</td>
<td align="left" rowspan="1" colspan="1">KJ862371</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57568</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862319</td>
<td align="left" rowspan="1" colspan="1">KJ862372</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57631</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862320</td>
<td align="left" rowspan="1" colspan="1">KJ862373</td>
<td align="left" rowspan="1" colspan="1">KJ862410</td>
<td align="left" rowspan="1" colspan="1">KJ862446</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57512</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862321</td>
<td align="left" rowspan="1" colspan="1">KJ862374</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 53616</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. leiocalyx</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862322</td>
<td align="left" rowspan="1" colspan="1">KJ862375</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. peggii</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 55602<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. holosericia</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862308</td>
<td align="left" rowspan="1" colspan="1">KJ862361</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862440</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 55631</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. holosericia</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862309</td>
<td align="left" rowspan="1" colspan="1">KJ862362</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862441</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 58324</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. holosericia</italic>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862363</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. phyllodiorum</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 57310</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862323</td>
<td align="left" rowspan="1" colspan="1">KJ862377</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57516</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862324</td>
<td align="left" rowspan="1" colspan="1">KJ862378</td>
<td align="left" rowspan="1" colspan="1">KJ862411</td>
<td align="left" rowspan="1" colspan="1">KJ862447</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57569</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862325</td>
<td align="left" rowspan="1" colspan="1">KJ862379</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57570</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862326</td>
<td align="left" rowspan="1" colspan="1">KJ862380</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57578</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862327</td>
<td align="left" rowspan="1" colspan="1">KJ862381</td>
<td align="left" rowspan="1" colspan="1">KJ862412</td>
<td align="left" rowspan="1" colspan="1">KJ862448</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57579</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862328</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862413</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57580</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862329</td>
<td align="left" rowspan="1" colspan="1">KJ862382</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57588</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862330</td>
<td align="left" rowspan="1" colspan="1">KJ862383</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57589</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862331</td>
<td align="left" rowspan="1" colspan="1">KJ862384</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57590</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862332</td>
<td align="left" rowspan="1" colspan="1">KJ862385</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. podalyriifolium</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 57294</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. podalyriifolia</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862333</td>
<td align="left" rowspan="1" colspan="1">KJ862386</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 57576<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. podalyriifolia</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862334</td>
<td align="left" rowspan="1" colspan="1">KJ862387</td>
<td align="left" rowspan="1" colspan="1">KJ862414</td>
<td align="left" rowspan="1" colspan="1">KJ862449</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. tierneyi</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 27071<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Plurinerves</td>
<td align="left" rowspan="1" colspan="1"><italic>A. harpophylla</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862335</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862415</td>
<td align="left" rowspan="1" colspan="1">KJ862450</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 27880</td>
<td align="left" rowspan="1" colspan="1">Plurinerves</td>
<td align="left" rowspan="1" colspan="1"><italic>A. harpophylla</italic>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862388</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 27887</td>
<td align="left" rowspan="1" colspan="1">Plurinerves</td>
<td align="left" rowspan="1" colspan="1"><italic>A. harpophylla</italic>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862389</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. tropicum</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 56555</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. tropica</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862336</td>
<td align="left" rowspan="1" colspan="1">KJ862390</td>
<td align="left" rowspan="1" colspan="1">KJ862416</td>
<td align="left" rowspan="1" colspan="1">KJ862451</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56557<xref ref-type="table-fn" rid="tfn1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. tropica</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862337</td>
<td align="left" rowspan="1" colspan="1">KJ862391</td>
<td align="left" rowspan="1" colspan="1">KJ862417</td>
<td align="left" rowspan="1" colspan="1">KJ862452</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>E. violae-faustiae</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 55601</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862338</td>
<td align="left" rowspan="1" colspan="1">KJ862392</td>
<td align="left" rowspan="1" colspan="1">KJ862418</td>
<td align="left" rowspan="1" colspan="1">KJ862453</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 55629</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862339</td>
<td align="left" rowspan="1" colspan="1">KJ862393</td>
<td align="left" rowspan="1" colspan="1">KJ862419</td>
<td align="left" rowspan="1" colspan="1">KJ862454</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 55660</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. aulacocarpa</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862340</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862420</td>
<td align="left" rowspan="1" colspan="1">KJ862455</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56544</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. difficillis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862341</td>
<td align="left" rowspan="1" colspan="1">KJ862395</td>
<td align="left" rowspan="1" colspan="1">KJ862421</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 55616</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. difficillis</italic>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862396</td>
<td align="left" rowspan="1" colspan="1">KJ862422</td>
<td align="left" rowspan="1" colspan="1">KJ862456</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 55611</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. difficillis</italic>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862397</td>
<td align="left" rowspan="1" colspan="1">KJ862423</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56539</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. difficillis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862342</td>
<td align="left" rowspan="1" colspan="1">KJ862398</td>
<td align="left" rowspan="1" colspan="1">KJ862424</td>
<td align="left" rowspan="1" colspan="1">KJ862457</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56540</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. difficillis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862343</td>
<td align="left" rowspan="1" colspan="1">KJ862399</td>
<td align="left" rowspan="1" colspan="1">KJ862425</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56545</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. difficillis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862344</td>
<td align="left" rowspan="1" colspan="1">KJ862400</td>
<td align="left" rowspan="1" colspan="1">KJ862426</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56547</td>
<td align="left" rowspan="1" colspan="1">Juliflorae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. difficillis</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862345</td>
<td align="left" rowspan="1" colspan="1">KJ862401</td>
<td align="left" rowspan="1" colspan="1">KJ862427</td>
<td align="left" rowspan="1" colspan="1">KJ862458</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>Kernkampella breyniae</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 56909</td>
<td align="left" rowspan="1" colspan="1">NA</td>
<td align="left" rowspan="1" colspan="1"><italic>Breynia cernua</italic>
(<italic>Euphorbiaceae</italic>
)</td>
<td align="left" rowspan="1" colspan="1">KJ862346</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862428</td>
<td align="left" rowspan="1" colspan="1">KJ862459</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>Ravenelia neocaledoniensis</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 56907</td>
<td align="left" rowspan="1" colspan="1">NA</td>
<td align="left" rowspan="1" colspan="1"><italic>Vachellia farnesiana</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862347</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">BRIP 56908</td>
<td align="left" rowspan="1" colspan="1">NA</td>
<td align="left" rowspan="1" colspan="1"><italic>V. farnesiana</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ862348</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862460</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>Ravenelia</italic>
sp.</td>
<td align="left" rowspan="1" colspan="1">BRIP 56904</td>
<td align="left" rowspan="1" colspan="1">NA</td>
<td align="left" rowspan="1" colspan="1"><italic>Cassia</italic>
sp.</td>
<td align="left" rowspan="1" colspan="1">KJ862349</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862461</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>Sphaerophragmium</italic>
sp.</td>
<td align="left" rowspan="1" colspan="1">BRIP 56910</td>
<td align="left" rowspan="1" colspan="1">NA</td>
<td align="left" rowspan="1" colspan="1"><italic>Albizia</italic>
sp.</td>
<td align="left" rowspan="1" colspan="1">KJ862350</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">KJ862429</td>
<td align="left" rowspan="1" colspan="1">KJ862462</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><italic>Uromycladium simplex</italic>
</td>
<td align="left" rowspan="1" colspan="1">BRIP 57571</td>
<td align="left" rowspan="1" colspan="1">Botrycephalae</td>
<td align="left" rowspan="1" colspan="1"><italic>A. pycnantha</italic>
</td>
<td align="left" rowspan="1" colspan="1">KJ632990<xref ref-type="table-fn" rid="tfn4"><sup>3</sup>
</xref>
</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
<td align="left" rowspan="1" colspan="1">N/A</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="tfn1"><p>* Holotype</p>
</fn>
<fn id="tfn2"><p><sup>1</sup>
Botrycephalae = Botrycephalae subcade (<xref rid="R20" ref-type="bibr">Murphy et al. 2010</xref>
); Juliflorae = Juliflorae p.p. clade (<xref rid="R19" ref-type="bibr">Miller et al. 2013</xref>
); Plurinerves = Plurinerves p.p. clade (<xref rid="R19" ref-type="bibr">Miller et al. 2013</xref>
).</p>
</fn>
<fn id="tfn3"><p><sup>2</sup>
<xref rid="R26" ref-type="bibr">Scholler & Aime (2006)</xref>
.</p>
</fn>
<fn id="tfn4"><p><sup>3</sup>
<xref rid="R9" ref-type="bibr">Doungsa-ard et al. (2014)</xref>
.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<fig id="F1" orientation="portrait" position="float"><label>Fig. 1</label>
<caption><p>Phylogram obtained in a maximum likelihood search in RAxML of the SSU, ITS, LSU and CO3 gene regions. Bootstrap support (≥ 70 %) values from 1 000 replicates above nodes. Posterior probabilities (≥ 0.95) from 4 200 trees in a Bayesian search are shown below nodes. Synapomorphic characters, namely host subclade and morphology of urediniospores, are mapped on to the topology.</p>
</caption>
<graphic xlink:href="per-35-050-g001"></graphic>
</fig>
<fig id="F2" orientation="portrait" position="float"><label>Fig. 2</label>
<caption><p><italic>Endoraecium auriculiforme</italic>
. a. Bullate sori on leaf (BRIP 56550); b, c. urediniospores (BRIP 55609); d. teliospores (BRIP 56550); e. germinating teliospore (BRIP 56549). — Scale bars: a = 1 cm; b–e = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g002"></graphic>
</fig>
<fig id="F3" orientation="portrait" position="float"><label>Fig. 3</label>
<caption><p><italic>Endoraecium carnegiei</italic>
. a. Telia on leaves (BRIP 59218); b. gall (BRIP 57924); c. urediniospores (BRIP 57924); d, e. teliospores (BRIP 57924). — Scale bars: b = 1 cm; c–e = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g003"></graphic>
</fig>
<fig id="F4" orientation="portrait" position="float"><label>Fig. 4</label>
<caption><p><italic>Endoraecium disparrimum</italic>
. a. Host (BRIP 55659); b, c. urediniospores (BRIP 55659); d. teliospores (BRIP 55626). — Scale bars: a = 1 cm; b–d = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g004"></graphic>
</fig>
<fig id="F5" orientation="portrait" position="float"><label>Fig. 5</label>
<caption><p><italic>Endoraecium falciforme</italic>
(BRIP 57583). a. Uredinia and telia surrounding spermogonia; b. uredinium; c. teliospores; d, e. urediniospores. — Scale bars: b = 1 cm; c–e = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g005"></graphic>
</fig>
<fig id="F6" orientation="portrait" position="float"><label>Fig. 6</label>
<caption><p><italic>Endoraecium irroratum</italic>
. a. Stem gall (BRIP 55671); b, c. urediniospores (BRIP 55671); d. teliospore (BRIP 57286). — Scale bars: a = 1 cm; b–d = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g006"></graphic>
</fig>
<fig id="F7" orientation="portrait" position="float"><label>Fig. 7</label>
<caption><p><italic>Endoraecium maslinii</italic>
(BRIP 57872). a–c. Bullate swellings on leaves; d, e. urediniospores. — Scale bars: c = 1 cm; d, e = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g007"></graphic>
</fig>
<fig id="F8" orientation="portrait" position="float"><label>Fig. 8</label>
<caption><p><italic>Endoraecium peggii</italic>
a. Uredinia (BRIP 55631); b. teliospores (BRIP 55602); c, d. urediniospores (BRIP 55631). — Scale bars: a = 1 cm; b–d = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g008"></graphic>
</fig>
<fig id="F9" orientation="portrait" position="float"><label>Fig. 9</label>
<caption><p><italic>Endoraecium podalyriifolium</italic>
(BRIP 57576). a, b. Pulverulent sori on leaves and stems; c. teliospores; d, e. urediniospores. — Scale bars: b = 1 cm; c–e = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g009"></graphic>
</fig>
<fig id="F10" orientation="portrait" position="float"><label>Fig. 10</label>
<caption><p><italic>Endoraecium tropicum</italic>
. a. Pulverulent sori (BRIP 56557); b, c. urediniospores (BRIP 56557); d. teliospores (BRIP 56555). — Scale bars: a = 1 cm; b–d = 10 μm.</p>
</caption>
<graphic xlink:href="per-35-050-g010"></graphic>
</fig>
</floats-group>
</pmc>
</record>
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