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<document ID-DOI="http://dx.doi.org/10.3897/phytokeys.61.7420" ID-PMC="PMC4816979" ID-Pensoft-Pub="1314-2003-61-27" ID-Pensoft-UUID="FFD6FFE5611FFFA9FFEEFFCD13447221" ID-PubMed="27081347" ID-Zenodo-Dep="576347" ModsDocID="1314-2003-61-27" checkinTime="1456479453831" checkinUser="pensoft" docAuthor="Damian, Alexander &amp; Ormerod, Paul" docDate="2016" docId="433B2231821454259364C2986254D05B" docLanguage="en" docName="PhytoKeys 61: 27-35" docOrigin="PhytoKeys 61" docPubDate="2016-02-25" docSource="http://dx.doi.org/10.3897/phytokeys.61.7420" docTitle="Liparis aphylla G. A. Romero &amp; Garay 1999" docType="treatment" docVersion="3" id="FFD6FFE5611FFFA9FFEEFFCD13447221" lastPageNumber="30" masterDocId="FFD6FFE5611FFFA9FFEEFFCD13447221" masterDocTitle="Liparis aphylla (Malaxideae, Orchidaceae), a new leafless record from Peru" masterLastPageNumber="35" masterPageNumber="27" pageNumber="28" updateTime="1668141648867" updateUser="ExternalLinkService">
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<mods:title>Liparis aphylla (Malaxideae, Orchidaceae), a new leafless record from Peru</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Damian, Alexander</mods:namePart>
<mods:affiliation>Facultad de Ciencias Ambientales. Universidad Cientifica del Sur. Panamericana Sur Km. 19, Lima, Peru</mods:affiliation>
<mods:nameIdentifier type="email">adamian.pz@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ormerod, Paul</mods:namePart>
<mods:affiliation>P. O. Box 8210, Cairns 4870, Queensland, Australia</mods:affiliation>
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<mods:date>2016</mods:date>
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<mods:number>2016-02-25</mods:number>
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<mods:number>61</mods:number>
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<mods:start>27</mods:start>
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<mods:url>http://dx.doi.org/10.3897/phytokeys.61.7420</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/phytokeys.61.7420</mods:identifier>
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<treatment ID-GBIF-Taxon="182227294" LSID="urn:lsid:plazi:treatment:433B2231821454259364C2986254D05B" httpUri="http://treatment.plazi.org/id/433B2231821454259364C2986254D05B" lastPageId="4" lastPageNumber="30" pageId="1" pageNumber="28">
<subSubSection pageId="1" pageNumber="28" type="nomenclature">
<paragraph pageId="1" pageNumber="28">
<taxonomicName LSID="433B2231-8214-5425-9364-C2986254D05B" authority="G. A. Romero &amp; Garay" authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">Liparis aphylla G.A.Romero &amp; Garay</taxonomicName>
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Liparis aphylla. A Habit B Flower C Dissected perianth D Detail of the labellum disk base E Column withouh the anter cap, ventral view F Column lateral view, and anther cap. Drawn from A. Damian 0100 (MOL!)." figureDoi="10.3897/phytokeys.61.7420.figure1" httpUri="https://binary.pensoft.net/fig/79786" pageId="1" pageNumber="28">Figs 1</figureCitation>
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Liparis aphylla, A Habit B-C Two views of the of labellum. Arrows show the distinct elliptical concavity of the callus. Photographer: A. L. Damian." figureDoi="10.3897/phytokeys.61.7420.figure2" httpUri="https://binary.pensoft.net/fig/79787" pageId="1" pageNumber="28">, 2</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="28" type="reference_group">
<paragraph pageId="1" pageNumber="28">
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">Liparis aphylla</taxonomicName>
G.A.Romero &amp; Garay. Harvard Pap. Bot. 4(2): 483. 1999.
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="28" type="type">
<paragraph pageId="1" pageNumber="28">Type.</paragraph>
<paragraph pageId="1" pageNumber="28">
COLOMBIA.
<normalizedToken originalValue="Boyacá">Boyaca</normalizedToken>
: Sierra del Cocuy, 2800 m, &quot;terrestre, entre musgos asociada con
<taxonomicName class="Liliopsida" family="Orchidaceae" genus="Masdevallia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Masdevallia" order="Asparagales" pageId="1" pageNumber="28" phylum="Liliopsida" rank="genus">
<emphasis italics="true" pageId="1" pageNumber="28">Masdevallia</emphasis>
</taxonomicName>
sp., aparentemente
<normalizedToken originalValue="saprófita">saprofita</normalizedToken>
;
<normalizedToken originalValue="tépalos">tepalos</normalizedToken>
blanco-verdosos, labelo
<normalizedToken originalValue="púrpura">purpura</normalizedToken>
lila&quot; 20 July 1997,
<emphasis italics="true" pageId="1" pageNumber="28">
M. Ospina
<normalizedToken originalValue="Hernández">Hernandez</normalizedToken>
1487
</emphasis>
(Holotype: AMES!).
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="28" type="description">
<paragraph pageId="1" pageNumber="28">Description.</paragraph>
<paragraph pageId="1" pageNumber="28">
Herb, 4-10 cm tall, terrestrial. Rhizome and roots not seen. Pseudobulb subglobose, 2.5-5
<normalizedToken originalValue="×">x</normalizedToken>
2 mm, enveloped by a basal foliaceous green sheath 1.5 cm long. Leaves not seen. Inflorescence racemose, erect, successively (up to 6) flowered (usually two are open at a time), 7.8 cm. long; peduncle slender, 4 cm long; rachis weakly flexuous, distichous, 3.8 cm long; floral bracts lanceolate. acute, green, 5-7
<normalizedToken originalValue="×">x</normalizedToken>
3 mm. Flowers widely opening, resupinate, fragrance not detected, sepals and petals greenish, labellum rosy brown with a darker median stripe, column greenish suffused with rosy brown, pollinia yellow. Green, 4-6 long ovary with clavate, narrowly winged pedicel. Dorsal sepal oblong-lanceolate, obtuse, erect, 1-veined, 3-5.5
<normalizedToken originalValue="×">x</normalizedToken>
0.9-1.5 mm. Lateral sepals obliquely oblong-ovate, obtuse, midvein low carinate, parallell to each other under the labellum, 1-veined, 3-5
<normalizedToken originalValue="×">x</normalizedToken>
0.6-2 mm. Petals linear, obtuse, reflexed, 1-veined, 2.6-5
<normalizedToken originalValue="×">x</normalizedToken>
0.5-0.6 mm. Labellum subquadrate, distal margin serrate-denticulate, medially with a thickened glossy stripe, 3-5
<normalizedToken originalValue="×">x</normalizedToken>
3.5-5 mm; callus bilobed, each side with an erect, subquadrate lobe between which there is a distally thick-walled elliptical cavity. Column semiterete, thick basally but slender above, arcuate on its distal half, apex with small triangular wings on each side, 3 mm long; pollinia four in two pairs, waxy, triangular; anther cap ovoid. Capsule and seeds not seen.
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="28" type="ecology and distribution">
<paragraph pageId="1" pageNumber="28">Ecology and distribution.</paragraph>
<paragraph pageId="1" pageNumber="28">
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="1" pageNumber="28">Liparis aphylla</emphasis>
</taxonomicName>
is found in the Andes of Colombia, Ecuador and Peru, within an elevation range of 2600-3300 m. The distribution of this species, based on herbarium records, appears to be highly disjunct (Figure
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Known distribution of Liparis aphylla." figureDoi="10.3897/phytokeys.61.7420.figure3" httpUri="https://binary.pensoft.net/fig/79788" pageId="1" pageNumber="28">3</figureCitation>
). However, this extreme patchiness may be an artifact of limited collecting, and we suspect that
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="1" pageNumber="28">Liparis aphylla</emphasis>
</taxonomicName>
likely occurs throughout the Andean range, at climatically suitable locations ranging from the Cordillera Oriental/East Andes in Colombia to the northern Andes of Peru. Plants of
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="1" pageNumber="28">Liparis aphylla</emphasis>
</taxonomicName>
grow terrestrially among loose moss in wet, cold montane cloud forest with abundant bryophytes. Flowering period: May-July.
</paragraph>
<caption ID-Zenodo-Dep="976035" doi="10.3897/phytokeys.61.7420.figure1" httpUri="https://binary.pensoft.net/fig/79786" pageId="1" pageNumber="28" start="Figure 1" startId="F1">
<paragraph pageId="1" pageNumber="28">
<emphasis bold="true" pageId="1" pageNumber="28">Figure 1.</emphasis>
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="1" pageNumber="28">Liparis aphylla</emphasis>
</taxonomicName>
.
<emphasis bold="true" pageId="1" pageNumber="28">A</emphasis>
Habit
<emphasis bold="true" pageId="1" pageNumber="28">B</emphasis>
Flower
<emphasis bold="true" pageId="1" pageNumber="28">C</emphasis>
Dissected perianth
<emphasis bold="true" pageId="1" pageNumber="28">D</emphasis>
Detail of the labellum disk base
<emphasis bold="true" pageId="1" pageNumber="28">E</emphasis>
Column withouh the anter cap, ventral view
<emphasis bold="true" pageId="1" pageNumber="28">F</emphasis>
Column lateral view, and anther cap. Drawn from
<emphasis italics="true" pageId="1" pageNumber="28">A. Damian 0100</emphasis>
(MOL!).
</paragraph>
</caption>
<caption ID-Zenodo-Dep="976037" doi="10.3897/phytokeys.61.7420.figure2" httpUri="https://binary.pensoft.net/fig/79787" pageId="1" pageNumber="28" start="Figure 2" startId="F2">
<paragraph pageId="1" pageNumber="28">
<emphasis bold="true" pageId="1" pageNumber="28">Figure 2.</emphasis>
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="1" pageNumber="28">Liparis aphylla</emphasis>
</taxonomicName>
,
<emphasis bold="true" pageId="1" pageNumber="28">A</emphasis>
Habit
<emphasis bold="true" pageId="1" pageNumber="28">B-C</emphasis>
Two views of the of labellum. Arrows show the distinct elliptical concavity of the callus. Photographer: A. L. Damian.
</paragraph>
</caption>
<caption ID-Zenodo-Dep="976039" doi="10.3897/phytokeys.61.7420.figure3" httpUri="https://binary.pensoft.net/fig/79788" pageId="1" pageNumber="28" start="Figure 3" startId="F3">
<paragraph pageId="1" pageNumber="28">
<emphasis bold="true" pageId="1" pageNumber="28">Figure 3.</emphasis>
Known distribution of
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="1" pageNumber="28" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="1" pageNumber="28">Liparis aphylla</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
</subSubSection>
<subSubSection lastPageId="2" lastPageNumber="29" pageId="1" pageNumber="28" type="additional specimens examined">
<paragraph pageId="1" pageNumber="28">Additional specimens examined.</paragraph>
<paragraph lastPageId="2" lastPageNumber="29" pageId="1" pageNumber="28">
<emphasis bold="true" pageId="1" pageNumber="28">ECUADOR.</emphasis>
Prov. Azuay: East Cordillera, 4-6 km N. of Sevilla de Oro, 2745-3050 m, 16 August 1945, W.H. Camp E-4774
<pageBreakToken pageId="2" pageNumber="29" start="start">(</pageBreakToken>
AMES!, NY);
<emphasis bold="true" pageId="2" pageNumber="29">PERU.</emphasis>
Departamento Cajamarca, provincia Chota, Querocoto, entrance road to &quot;La Granja&quot;,
<geoCoordinate degrees="6" direction="south" minutes="20" orientation="latitude" precision="1" seconds="6.70" value="-6.3351946">6°20'6.70&quot;S</geoCoordinate>
,
<geoCoordinate degrees="79" direction="west" minutes="9" orientation="longitude" precision="1" seconds="24.49" value="-79.15681">79°9'24.49&quot;W</geoCoordinate>
; terrestrial, montane rainforest, 2600 m, 01 May 2014,
<emphasis italics="true" pageId="2" pageNumber="29">A. Damian 0100</emphasis>
(MOL!, ADP-spirit 3033).
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="29" type="conservation status">
<paragraph pageId="2" pageNumber="29">Conservation status.</paragraph>
<paragraph pageId="2" pageNumber="29">
This species is presently know only from three location worldwide; according to the IUCN Red List (
<bibRefCitation author="IUCN" journalOrPublisher="Standards and Petitions Subcommittee" pageId="5" pageNumber="32" refId="B7" refString="IUCN, 2014. Guidelines for using the IUCN Red List categories and criteria. Standards and Petitions Subcommittee" title="Guidelines for using the IUCN Red List categories and criteria" year="2014">IUCN 2014</bibRefCitation>
) and
<bibRefCitation author="Roque, J" editor="Leon, B" journalOrPublisher="Rev. peru. biol." pageId="6" pageNumber="33" pagination="759 - 878" refId="B13" refString="Roque, J, Leon, B, 2006. Orchidaceae endemicas del Peru. In: Leon, B, Pitman, N, Roque, J, Eds., El libro rojo de las plantas endemicas el Peru. Rev. peru. biol. 13 (2): 759 - 878" title="Orchidaceae endemicas del Peru." volume="13" volumeTitle="El libro rojo de las plantas endemicas el Peru." year="2006">Roque and Leon (2006)</bibRefCitation>
criteria, it should be listed as critically endangered (CR) B1ab(iii).
</paragraph>
</subSubSection>
<subSubSection lastPageId="4" lastPageNumber="31" pageId="2" pageNumber="29" type="discussion">
<paragraph pageId="2" pageNumber="29">Discussion.</paragraph>
<paragraph pageId="2" pageNumber="29">
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="2" pageNumber="29" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="2" pageNumber="29">Liparis aphylla</emphasis>
</taxonomicName>
was described from an individual plant collected in Sierra del Cocuy, Colombia, by Romero and Garay (1997) and from another record from Azuay, Ecuador, (1977). These two specimens, along with the Peruvian specimen reported in this paper, represent the only available material of this tiny rare orchid. The overall morphology observed in these three specimens, is quite uniform except for considerable variation in labellum shape, which ranges from quadrate in the specimen
<emphasis italics="true" pageId="2" pageNumber="29">Camp E-4774</emphasis>
to subquadrate in
<emphasis italics="true" pageId="2" pageNumber="29">
Ospina
<normalizedToken originalValue="Hernández">Hernandez</normalizedToken>
1487
</emphasis>
. Moreover, the elliptic concavity of the callus of our specimen did not appear to be present in other two specimens, although it is not clear if the absence of this concavity in the latter two specimens is an artifact of preservation.
</paragraph>
<paragraph lastPageId="3" lastPageNumber="30" pageId="2" pageNumber="29">
Unlike any others members of
<taxonomicName class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis" order="Asparagales" pageId="2" pageNumber="29" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="2" pageNumber="29">Liparis</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="2" pageNumber="29" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="2" pageNumber="29">Liparis aphylla</emphasis>
</taxonomicName>
appears as a leafless orchid with poor-developed root system. However, these two conditions need to be studied care
<pageBreakToken pageId="3" pageNumber="30" start="start">fully</pageBreakToken>
. Although we were unable to see any remnant roots or rhizome on the specimens examined, Ospina Hernandez sheet (1487) includes an interesting note cited as &quot;Plant tubers covered by fungal hyphae (...)&quot;. It is highly possible that
<normalizedToken originalValue="“tubers”">&quot;tubers&quot;</normalizedToken>
on this context actually refers to the pseudobulb and not to the presence of subterranean stems or shoots that resemble any kind of root-like system or rhizome as it occurs in many basal Epidendroids orchids (
<bibRefCitation author="Pridgeon, A" journalOrPublisher="Oxford University Press, Oxford" pageId="5" pageNumber="32" pagination="1 - 608" refId="B10" refString="Pridgeon, A, Cribb, PJ, Chase, MW, Rasmussen, FN, 2005. Genera Orchidacearum, volume 4: Epidendroideae (Part One). Oxford University Press, Oxford: 1 - 608" title="Genera Orchidacearum, volume 4: Epidendroideae (Part One)" year="2005">Pridgeon et al. 2005</bibRefCitation>
,
<bibRefCitation author="Campbell, FK" journalOrPublisher="The MIOS Journal" pageId="5" pageNumber="32" pagination="6 - 17" refId="B3" refString="Campbell, FK, 2014. A summary of Holomycotrophic orchids. The MIOS Journal 15 (4): 6 - 17" title="A summary of Holomycotrophic orchids." volume="15" year="2014">Campbell 2014</bibRefCitation>
). A closest analysis of the original drawing of
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="3" pageNumber="30" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="3" pageNumber="30">Liparis aphylla</emphasis>
</taxonomicName>
by Romero and Garay (1997) shows sort of filamentous structures emerging beneath the pseudobulb. Since the dimensions of these formations are indistinct (0.3
<normalizedToken originalValue="×">x</normalizedToken>
0.1 cm), is fairly accurate to attribute those filamentouse root-like structures to the &quot;fungal hyphae&quot; which Ospina was referring in the first place.
</paragraph>
<paragraph pageId="3" pageNumber="30">
Another strikingly feature on
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="3" pageNumber="30" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="3" pageNumber="30">Liparis aphylla</emphasis>
</taxonomicName>
is its leaflessness. As it happens with rhizome and roots, no remnants of withered or decomposed leaves were observed neither in the field nor in available herbarium specimens. As a result of this uncommon state within
<taxonomicName class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis" order="Asparagales" pageId="3" pageNumber="30" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="3" pageNumber="30">Liparis</emphasis>
</taxonomicName>
, Romero &amp; Garay decided to establish
<emphasis italics="true" pageId="3" pageNumber="30">
Sect.
<taxonomicName class="Magnoliopsida" family="Orchidaceae" genus="Aphylla" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Aphylla" order="Asparagales" pageId="3" pageNumber="30" phylum="Angiospermae" rank="genus">Aphylla</taxonomicName>
</emphasis>
(Romero and Garay 1997) to include this single species, which outstands essentially for its leafless habit, well-developed pseudobulb, plants of small size and muscicolous habitat. However, additional observations whether this set of characters, especially those referring to leaves and roots, are continuous or not along specimens were missing.
</paragraph>
<paragraph lastPageId="4" lastPageNumber="31" pageId="3" pageNumber="30">
Leaflessness is a feature that is present in many angiosperms (
<bibRefCitation author="Vincent, SFT" editor="Vincent, SFT" journalOrPublisher="Springer Science" pageId="6" pageNumber="33" pagination="19 - 101" refId="B17" refString="Vincent, SFT, Frendenstein, JV, Kissling, J, Christenhusz, JM, Stotler, ER, Crandal-Stotler, B, Wickett, N, Rudall, JP, Maas-van de Kamer, H, Maas, JM, 2013. Taxonomy and Classification. In: Vincent, SFT, Ed., Mycoheterotrophy: The Biology of Plants Living on Fungi. Springer Science: 19 - 101" title="Taxonomy and Classification." volumeTitle="Mycoheterotrophy: The Biology of Plants Living on Fungi." year="2013">Vicent et al. 2013</bibRefCitation>
,
<bibRefCitation author="Carlsward, BS" journalOrPublisher="Am. J. Bot." pageId="5" pageNumber="32" pagination="770 - 786" publicationUrl="10.1007/978-1-4614-5209-6_2" refId="B4" refString="Carlsward, BS, Whitten, WM, Williams, NH, Bytebier, B, 2006. Molecular Phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. Am. J. Bot. 93: 770 - 786, DOI: 10.1007/978-1-4614-5209-6_2" title="Molecular Phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness." url="10.1007/978-1-4614-5209-6_2" volume="93" year="2006">Calswards et al. 2006</bibRefCitation>
) and
<taxonomicName class="Magnoliopsida" family="Orchidaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Asparagales" pageId="3" pageNumber="30" phylum="Angiospermae" rank="family">Orchidaceae</taxonomicName>
is not the exception. At least 235 orchid species and 43 genera are leafless, most of them found in
<taxonomicName class="Magnoliopsida" family="Orchidaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Asparagales" pageId="3" pageNumber="30" phylum="Angiospermae" rank="subFamily" subFamily="Epidendroideae">Epidendroideae</taxonomicName>
<pageBreakToken pageId="4" pageNumber="31" start="start">(</pageBreakToken>
<bibRefCitation author="Freudenstein, JV" editor="Seberg, O" journalOrPublisher="Aarhus University Press, Arhus" pageId="5" pageNumber="32" pagination="25 - 37" refId="B5" refString="Freudenstein, JV, Barrett, CF, 2010. Mycoheterotrophy and diversity in Orchidaceae. In: Seberg, O, Petersen, G, Barfod, AS, Davis, JI, Eds., Diversity, phylogeny, and evolution in the monocotyledons. Aarhus University Press, Arhus: 25 - 37" title="Mycoheterotrophy and diversity in Orchidaceae." volumeTitle="Diversity, phylogeny, and evolution in the monocotyledons." year="2010">Freudenstein and Barrett 2010</bibRefCitation>
). For instance, within tribe
<taxonomicName class="Magnoliopsida" family="Orchidaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Asparagales" pageId="4" pageNumber="31" phylum="Angiospermae" rank="tribe" tribe="Malaxideae">Malaxideae</taxonomicName>
, two orchids have reported being leafless:
<taxonomicName authorityName="Tang &amp; F. T. Wang" authorityYear="1951" baseAuthorityName="King &amp; Pantl." class="Liliopsida" family="Orchidaceae" genus="Malaxis" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Malaxis aphylla" order="Asparagales" pageId="4" pageNumber="31" phylum="Magnoliophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="4" pageNumber="31">Malaxis aphylla</emphasis>
</taxonomicName>
(King &amp; Pantl.) T.Tang &amp; F.T.Wang and
<taxonomicName authorityName="Tang &amp; F. T. Wang" authorityYear="1951" baseAuthorityName="King &amp; Pantl." class="Liliopsida" family="Orchidaceae" genus="Malaxis" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Malaxis saprophyta" order="Asparagales" pageId="4" pageNumber="31" phylum="Magnoliophyta" rank="species" species="saprophyta">
<emphasis italics="true" pageId="4" pageNumber="31">Malaxis saprophyta</emphasis>
</taxonomicName>
(King &amp; Pantl.) T.Tang &amp; F.T.Wang (
<bibRefCitation author="Vincent, SFT" editor="Vincent, SFT" journalOrPublisher="Springer Science" pageId="6" pageNumber="33" pagination="19 - 101" refId="B17" refString="Vincent, SFT, Frendenstein, JV, Kissling, J, Christenhusz, JM, Stotler, ER, Crandal-Stotler, B, Wickett, N, Rudall, JP, Maas-van de Kamer, H, Maas, JM, 2013. Taxonomy and Classification. In: Vincent, SFT, Ed., Mycoheterotrophy: The Biology of Plants Living on Fungi. Springer Science: 19 - 101" title="Taxonomy and Classification." volumeTitle="Mycoheterotrophy: The Biology of Plants Living on Fungi." year="2013">Vincent et al. 2013</bibRefCitation>
). Most of these leafless orchid display any of the following arrangements or life-forms: (1) well-developed shoot system which forms the main body (e.g. leafless
<taxonomicName class="Liliopsida" family="Orchidaceae" genus="Vanilla" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Vanilla" order="Asparagales" pageId="4" pageNumber="31" phylum="Liliopsida" rank="genus">
<emphasis italics="true" pageId="4" pageNumber="31">Vanilla</emphasis>
</taxonomicName>
), (2) shoot system reduced, i.e. shootless orchids, roots forming the main body of the plant (e.g. Vandeae) (
<bibRefCitation author="Carlsward, BS" journalOrPublisher="Am. J. Bot." pageId="5" pageNumber="32" pagination="770 - 786" publicationUrl="10.1007/978-1-4614-5209-6_2" refId="B4" refString="Carlsward, BS, Whitten, WM, Williams, NH, Bytebier, B, 2006. Molecular Phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. Am. J. Bot. 93: 770 - 786, DOI: 10.1007/978-1-4614-5209-6_2" title="Molecular Phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness." url="10.1007/978-1-4614-5209-6_2" volume="93" year="2006">Carlsward et al. 2006</bibRefCitation>
), (3) roots fleshy, fasciculate, leaves basal but lacking at flowering time (e.g.
<taxonomicName lsidName="" pageId="4" pageNumber="31" rank="subTribe" subTribe="Spiranthinae">Spiranthinae</taxonomicName>
:
<taxonomicName family="Orchidaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Asparagales" pageId="4" pageNumber="31" phylum="Liliopsida" rank="tribe" tribe="Cranichideae">Cranichideae</taxonomicName>
) (
<bibRefCitation author="Salazar, GA" journalOrPublisher="Oxford University Press, Oxford" pageId="6" pageNumber="33" pagination="164 - 278" refId="B14" refString="Salazar, GA, 2003. Genera orchidacearum 3: orchidoideae part 2, Vanilloideae. Oxford University Press, Oxford: 164 - 278" title="Genera orchidacearum 3: orchidoideae part 2, Vanilloideae" year="2003">Salazar 2003</bibRefCitation>
), and (4) myco-heterotrophic orchids, achlorophyllous, roots reduced or absent, rhizome fleshy, coralloid, tuberlike or cylindric (e.g. Aphyllorchis, Gastrodia) (
<bibRefCitation author="Rasmussen, FN" editor="Wilson, KL" journalOrPublisher="CSIRO Melbourne" pageId="5" pageNumber="32" pagination="430 - 435" refId="B11" refString="Rasmussen, FN, 2000. Ins and Outs of Orchid Phylogeny. In: Wilson, KL, Morrison, DA, Eds., Monocots: Systematics and Evolution. CSIRO Melbourne: 430 - 435" title="Ins and Outs of Orchid Phylogeny." volumeTitle="Monocots: Systematics and Evolution." year="2000">Rasmussen 2000</bibRefCitation>
). A major question rise then among others, which life-form represents better to
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="4" pageNumber="31" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="4" pageNumber="31">Liparis aphylla</emphasis>
</taxonomicName>
?. Although Romero and Garay (1997) suggested it could be referred to as a
<normalizedToken originalValue="“saprophyte”">&quot;saprophyte&quot;</normalizedToken>
, this term proved to be inaccurate (
<bibRefCitation author="Leake, JR" journalOrPublisher="New Phytologist" pageId="5" pageNumber="32" pagination="171 - 216" publicationUrl="10.1111/j.1469-8137.1994.tb04272.x" refId="B8" refString="Leake, JR, 1994. The biology of Myco-heterotrophic ('saprophytic') plants. New Phytologist 127: 171 - 216, DOI: 10.1111/j.1469-8137.1994.tb04272.x" title="The biology of Myco-heterotrophic (' saprophytic') plants." url="10.1111/j.1469-8137.1994.tb04272.x" volume="127" year="1994">Leake 1994</bibRefCitation>
). We believed
<taxonomicName authorityName="G. A. Romero &amp; Garay" authorityYear="1999" class="Liliopsida" family="Orchidaceae" genus="Liparis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Liparis aphylla" order="Asparagales" pageId="4" pageNumber="31" phylum="Tracheophyta" rank="species" species="aphylla">
<emphasis italics="true" pageId="4" pageNumber="31">Liparis aphylla</emphasis>
</taxonomicName>
could represent a partially myco-heterotrophic (holo-mycotrophic) plant, i.e. clorophyllous plant that combines autotrophy and myco-heterotrophy to obtain carbon during at least one stage of its life cycle (
<bibRefCitation author="Rasmussen, HN" journalOrPublisher="Cambridge University Press, Cambridge" pageId="5" pageNumber="32" pagination="1 - 448" publicationUrl="10.1017/cbo9780511525452" refId="B12" refString="Rasmussen, HN, 1995. Terrestrial Orchids: From Seeds to Myco-trophic Plant. Cambridge University Press, Cambridge: 1 - 448, DOI: 10.1017/cbo9780511525452" title="Terrestrial Orchids: From Seeds to Myco-trophic Plant" url="10.1017/cbo9780511525452" year="1995">Rasmussen 1995</bibRefCitation>
). The nonexistence of a well-developed root system, leaflessness (= myco-heterotrophic species) and retainment of chlorophyll on its basal sheath, stem and bracts seem to confirm this hypothesis. Nonetheless, it is important to keep in mind that the myco-heterotrophic status is
<normalizedToken originalValue="“putative”">&quot;putative&quot;</normalizedToken>
on this species, and remains speculative until a careful physiological analysis has been carried out.
</paragraph>
</subSubSection>
</treatment>
</document>