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<document ID-DOI="http://dx.doi.org/10.3897/phytokeys.144.46700" ID-GBIF-Dataset="cd28da41-78ef-4362-bc21-5e114c6b2d05" ID-PMC="PMC7148385" ID-Pensoft-Pub="1314-2003-144-81" ID-Pensoft-UUID="B3D8E8B6CF515155A212E307E6A41405" ID-PubMed="32296283" ModsDocID="1314-2003-144-81" checkinTime="1585985472313" checkinUser="pensoft" docAuthor="McPartland, John M. &amp; Small, Ernest" docDate="2020" docId="69AF8A9C1FAF533EB01086732B3C9DF2" docLanguage="en" docName="PhytoKeys 144: 81-112" docOrigin="PhytoKeys 144" docSource="http://dx.doi.org/10.3897/phytokeys.144.46700" docTitle="Cannabis sativa subsp. var. var. himalayensis McPartl. &amp; E. Small" docType="treatment" docVersion="4" id="B3D8E8B6CF515155A212E307E6A41405" lastPageNumber="81" masterDocId="B3D8E8B6CF515155A212E307E6A41405" masterDocTitle="A classification of endangered high-THC cannabis (Cannabis sativa subsp. indica) domesticates and their wild relatives" masterLastPageNumber="112" masterPageNumber="81" pageNumber="81" updateTime="1668139750943" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>A classification of endangered high-THC cannabis (Cannabis sativa subsp. indica) domesticates and their wild relatives</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>McPartland, John M.</mods:namePart>
<mods:affiliation>College of Medicine, University of Vermont, Burlington, Vermont, 05405 USA &amp; GW Pharmaceuticals, Sovereign House, Histon, Cambridge, CB 24 9 BZ, UK</mods:affiliation>
<mods:nameIdentifier type="orcid">https://orcid.org/0000-0003-3385-2827</mods:nameIdentifier>
<mods:nameIdentifier type="email">mcpruitt@myfairpoint.net</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Small, Ernest</mods:namePart>
<mods:affiliation>Science and Technology Branch, Agriculture and Agri-Food Canada, Saunders Building, Central Experimental Farm, Ottawa, Ontario K 1 A 0 C 6, Canada</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>PhytoKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="volume">
<mods:number>144</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>81</mods:start>
<mods:end>112</mods:end>
</mods:extent>
</mods:part>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/phytokeys.144.46700</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/phytokeys.144.46700</mods:identifier>
<mods:identifier type="Pensoft-Pub">1314-2003-144-81</mods:identifier>
<mods:identifier type="Pensoft-UUID">B3D8E8B6CF515155A212E307E6A41405</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="163433677" LSID="urn:lsid:plazi:treatment:69AF8A9C1FAF533EB01086732B3C9DF2" httpUri="http://treatment.plazi.org/id/69AF8A9C1FAF533EB01086732B3C9DF2" lastPageNumber="81" pageId="0" pageNumber="81">
<subSubSection pageId="0" pageNumber="81" type="nomenclature">
<paragraph pageId="0" pageNumber="81">
<taxonomicName LSID="69AF8A9C-1FAF-533E-B010-86732B3C9DF2" authority="(Cazzuola) McPartl. &amp; E. Small" class="Magnoliopsida" family="Cannabaceae" genus="Cannabis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Cannabis sativa subsp. var. var. himalayensis" order="Rosales" pageId="0" pageNumber="81" phylum="Tracheophyta" rank="variety" species="sativa" subSpecies="var." variety="himalayensis">Cannabis sativa subsp. indica var. himalayensis (Cazzuola) McPartl. &amp; E.Small</taxonomicName>
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Two varieties of C. sativa subsp. indica from South Asia. On left a var. indica. On right b var. himalayensis." figureDoi="10.3897/phytokeys.144.46700.figure4" httpUri="https://binary.pensoft.net/fig/390914" pageId="0" pageNumber="81">Figure 4b</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="reference_group">
<paragraph pageId="0" pageNumber="81">
<taxonomicName class="Magnoliopsida" family="Cannabaceae" genus="Cannabis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Cannabis sativa subsp. var. var. hymalaiensis" order="Rosales" pageId="0" pageNumber="81" phylum="Tracheophyta" rank="variety" species="sativa" subSpecies="var." variety="hymalaiensis">Cannabis sativa var. hymalaiensis</taxonomicName>
Cazzuola, Il Regno vegetale tessili e tintoriale, p. 49, 1875 (misspelling corrected
<emphasis italics="true" pageId="0" pageNumber="81">apud</emphasis>
ICN Article 60.1) Basionym.
</paragraph>
<paragraph pageId="0" pageNumber="81">
<taxonomicName family="Cannabaceae" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="C. sativa subsp. var. var. hymalaiensis" order="Rosales" pageId="0" pageNumber="81" rank="variety" species="sativa" subSpecies="var." variety="hymalaiensis">C. sativa var. hymalaiensis</taxonomicName>
Cazzuola, Nuovo Giornale Botanico Italiano 5: 262, 1873, nomen nudum.
</paragraph>
<paragraph pageId="0" pageNumber="81">
<taxonomicName family="Cannabaceae" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="C. sativa subsp. var. var. himalayensis" order="Rosales" pageId="0" pageNumber="81" rank="variety" species="sativa" subSpecies="var." variety="himalayensis">C. sativa var. himalayensis</taxonomicName>
Cazzuola, Dizionario di botanica, p. 105, 1876 (later homonym).
</paragraph>
<paragraph pageId="0" pageNumber="81">
=
<taxonomicName family="Cannabaceae" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="C. sativa subsp. var. var. himalayensis" order="Rosales" pageId="0" pageNumber="81" rank="variety" species="sativa" subSpecies="var." variety="himalayensis">C. sativa var. himalayensis</taxonomicName>
Koch, Annales des Sciences Naturelles Botanique (Series 4) 1: 352, 1854, nomen nudum.
</paragraph>
<paragraph pageId="0" pageNumber="81">
=
<taxonomicName family="Cannabaceae" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="C. sativa" order="Rosales" pageId="0" pageNumber="81" rank="species" species="sativa">C. sativa</taxonomicName>
<normalizedToken originalValue="β">β</normalizedToken>
<emphasis italics="true" pageId="0" pageNumber="81">vulgaris</emphasis>
de Candolle, Prodromus 16(1):31, 1869 (
<emphasis italics="true" pageId="0" pageNumber="81">en part</emphasis>
, based on plants growing spontaneous in northern India and Burma).
</paragraph>
<paragraph pageId="0" pageNumber="81">
=
<taxonomicName family="Cannabaceae" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="C. sativa" order="Rosales" pageId="0" pageNumber="81" rank="species" species="sativa">C. sativa</taxonomicName>
<normalizedToken originalValue="α">α</normalizedToken>
<taxonomicName family="Cannabaceae" form="montana" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="indica subsp. f." order="Rosales" pageId="0" pageNumber="81" rank="form" species="indica" subSpecies="f.">indica f. montana</taxonomicName>
Fristedt, Upsala
<normalizedToken originalValue="Läkareförenings">Laekarefoerenings</normalizedToken>
<normalizedToken originalValue="Förhandlingar">Foerhandlingar</normalizedToken>
5: 507, 1869- 1870, nomen nudum.
</paragraph>
<paragraph pageId="0" pageNumber="81">
=
<taxonomicName family="Cannabaceae" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="C. himalyana" order="Rosales" pageId="0" pageNumber="81" rank="species" species="himalyana">C. himalyana</taxonomicName>
Zinger, Flora oder Allgemeine Botanische Zeitung 85: 207, 1898, nomen nudum.
</paragraph>
<paragraph pageId="0" pageNumber="81">
=
<taxonomicName authority="var. spontanea" family="Cannabaceae" higherTaxonomySource="treatment-meta" kingdom="Plantae" lsidName="C. sativa subsp. var. var. spontanea" order="Rosales" pageId="0" pageNumber="81" rank="variety" section="spontanea" species="sativa" subSpecies="var." variety="spontanea">C. sativa subsp. indica sect. spontanea var. spontanea</taxonomicName>
Clarke,
<taxonomicName class="Magnoliopsida" family="Cannabaceae" genus="Cannabis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Cannabis" order="Rosales" pageId="0" pageNumber="81" phylum="Tracheophyta" rank="genus">Cannabis</taxonomicName>
Evolution p. 224, 1987, nomen invalidum.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="neotype">
<paragraph pageId="0" pageNumber="81">Neotype.</paragraph>
<paragraph pageId="0" pageNumber="81">
Designated herein, INDIA: Himachal Pradesh, Shimla or Kinnaur (&quot;Himalaya Boreal. Occident., Regio Temp.&quot;),
<emphasis italics="true" pageId="0" pageNumber="81">T. Thompson</emphasis>
, 1847 (GH). No
<taxonomicName class="Magnoliopsida" family="Cannabaceae" genus="Cannabis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="himalayensis" order="Rosales" pageId="0" pageNumber="81" phylum="Tracheophyta" rank="species" species="himalayensis">
<emphasis italics="true" pageId="0" pageNumber="81">himalayensis</emphasis>
</taxonomicName>
specimens exist in the herbaria of Cazzuola or Koch (pers. communications, Lucia Amadei, herb. PI; Robert Vogt, herb. B).
<normalizedToken originalValue="Thompsons">Thompson's</normalizedToken>
specimen was designated as neotype because it represents the best of several collections he made in the Himalaya. It was distributed as an exsiccatum, with duplicates at several herbaria, providing isoneotypes (BM! K! LE! US!).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="diagnosis">
<paragraph pageId="0" pageNumber="81">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="81">Plants with THC% ≥0.3% in inflorescence and a THC/CBD ratio often ≥7, sometimes less; central leaflet length:width ratio ≥6 in fan leaves near the base of inflorescences; mature achenes usually &lt;3.6 mm long, with a persistent perianth and a protuberant base, and readily disarticulating from plant by a well-developed abscission zone.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="morphology">
<paragraph pageId="0" pageNumber="81">Morphology.</paragraph>
<paragraph pageId="0" pageNumber="81">Plants 1.0-3.0 m tall. Central stem (stalk) internodes relatively long (often&gt;10 cm, shorter in shorter plants), somewhat hollow (up to 1/2 stem diameter). Branches flexible, diverging from the stalk at relatively acute angles (around 45°). Leaf palmately compound, larger leaves usually with at least 7 leaflets, leaflet edges not overlapping. Central leaflet long and narrow, lanceolate in shape; margins with moderately coarse serrations, and rare secondary serrations. Female inflorescence (and infructescence) elongated and somewhat diffuse, with relatively obscure sugar leaves (a high perigonal bract-to-leaf index). Sugar leaves with CSGTs limited to the proximal half. Perigonal bract covered with a moderate density of CSGTs. Perianth membranous, hyaline with pigmented areas (brown and mottled or marbled in appearance); always persistent. Achene usually &lt;3.6 mm long, exocarp green-brown; with an elongated base and abscission zone that is relatively narrow.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="phytochemistry">
<paragraph pageId="0" pageNumber="81">Phytochemistry.</paragraph>
<paragraph pageId="0" pageNumber="81">
Dried female inflorescences: THC ≥0.3% (although two studies report plants with THC &lt;0.3%); weighted x¯ = 1.49%, range between 0.06% and 9.3%. THC/CBD ratios vary; two studies (those with THC &lt;0.3%), who shared accessions, reported ratios of only 1.28 and 1.56; these accessions may represent East Asian fiber-type domesticates that reacquired wild-type traits. Ratios in other studies are&gt;10, even&gt;100. THC content and THC/CBD ratios are skewed by THCV%+CBDV%, which is higher than any other variety: x¯ = 0.90% (
<bibRefCitation DOI="https://doi.org/10.3732/ajb.91.6.966" author="Hillig, KW" journalOrPublisher="American Journal of Botany" pageId="0" pageNumber="81" pagination="966 - 975" refId="B44" refString="Hillig, KW, Mahlberg, PG, 2004. A chemotaxonomic analysis of cannabinoid variation in Cannabis (Cannabaceae). American Journal of Botany 91 (6): 966 - 975, DOI: https://doi.org/10.3732/ajb.91.6.966" title="A chemotaxonomic analysis of cannabinoid variation in Cannabis (Cannabaceae)." url="https://doi.org/10.3732/ajb.91.6.966" volume="91" year="2004">Hillig and Mahlberg 2004</bibRefCitation>
). The terpenoid profile is similar to that of var.
<taxonomicName class="Magnoliopsida" family="Cannabaceae" genus="Cannabis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="var. indica" order="Rosales" pageId="0" pageNumber="81" phylum="Tracheophyta" rank="variety" species="himalayensis" variety="indica">
<emphasis italics="true" pageId="0" pageNumber="81">indica</emphasis>
</taxonomicName>
, except for higher levels of
<emphasis italics="true" pageId="0" pageNumber="81">
<normalizedToken originalValue="β">β</normalizedToken>
</emphasis>
-myrcene,
<emphasis italics="true" pageId="0" pageNumber="81">cis</emphasis>
-ocimene, and
<emphasis italics="true" pageId="0" pageNumber="81">
<normalizedToken originalValue="β">β</normalizedToken>
</emphasis>
-caryophyllene.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="genetics">
<paragraph pageId="0" pageNumber="81">Genetics.</paragraph>
<paragraph pageId="0" pageNumber="81">
Allozyme analysis (
<bibRefCitation DOI="https://doi.org/10.1007/s10722-003-4452-y" author="Hillig, KW" journalOrPublisher="Genetic Resources and Crop Evolution" pageId="0" pageNumber="81" pagination="161 - 180" refId="B42" refString="Hillig, KW, 2005a. Genetic evidence for speciation in Cannabis (Cannabaceae). Genetic Resources and Crop Evolution 52 (2): 161 - 180, DOI: https://doi.org/10.1007/s10722-003-4452-y" title="Genetic evidence for speciation in Cannabis (Cannabaceae)." url="https://doi.org/10.1007/s10722-003-4452-y" volume="52" year="2005 a">Hillig 2005a</bibRefCitation>
) partially segregated wild-type accessions from South Asian domesticates. He proposed that wild-type accessions from the Himalaya represented the ancestral source of South Asian domesticates.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="description">
<paragraph pageId="0" pageNumber="81">Other characters.</paragraph>
<paragraph pageId="0" pageNumber="81">
Generally late maturing; achenes fall from plant at maturity. Bast fiber content (as a percent of stalk dry weight) in Himalayan plants is higher than plants grown exclusively for drugs in southern India (
<bibRefCitation DOI="https://doi.org/10.1007/BF00710361" author="Bredemann, G" journalOrPublisher="Der Zuechter" pageId="0" pageNumber="81" pagination="257 - 269" refId="B6" refString="Bredemann, G, 1952. Weitere Beobachtungen bei Zuechtung des Hanfes auf Fasergehalt. Der Zuechter 22: 257 - 269, DOI: https://doi.org/10.1007/BF00710361" title="Weitere Beobachtungen bei Zuechtung des Hanfes auf Fasergehalt." url="https://doi.org/10.1007/BF00710361" volume="22" year="1952">Bredemann 1952</bibRefCitation>
;
<bibRefCitation author="de Meijer, EPM" journalOrPublisher="Critical Reviews in Plant Sciences" pageId="0" pageNumber="81" refId="B18" refString="de Meijer, EPM, 1994. Diversity in Cannabis. Doctoral thesis, Wageningen Agricultural University, Wageningen, The Netherlands." title="Diversity in Cannabis. Doctoral thesis, Wageningen Agricultural University, Wageningen, The Netherlands." year="1994">de Meijer 1994</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="provenance and uses">
<paragraph pageId="0" pageNumber="81">Provenance and uses.</paragraph>
<paragraph pageId="0" pageNumber="81">
Wild-growing (possibly indigenous) populations occur throughout montane India, Nepal, and Bhutan, where they are harvested for bast fiber (stalks),
<emphasis italics="true" pageId="0" pageNumber="81">
<normalizedToken originalValue="bhāng">bhang</normalizedToken>
</emphasis>
(leaves), hand-rubbed
<emphasis italics="true" pageId="0" pageNumber="81">charas</emphasis>
(
<emphasis italics="true" pageId="0" pageNumber="81">hashīsh</emphasis>
), or achenes (seeds). Achenes in some herbarium specimens from the Himalaya were relatively large with a reduced abscission mechanism, indicating the presence of genes from domesticated plants.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="81" type="basionym notes">
<paragraph pageId="0" pageNumber="81">Basionym notes.</paragraph>
<paragraph pageId="0" pageNumber="81">
Cazzuola spelled the epithet
<taxonomicName class="Magnoliopsida" family="Cannabaceae" genus="Cannabis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="himalayensis" order="Rosales" pageId="0" pageNumber="81" phylum="Tracheophyta" rank="species" species="himalayensis">
<emphasis italics="true" pageId="0" pageNumber="81">himalayensis</emphasis>
</taxonomicName>
variously between 1873 and 1876. His earliest publication did not provide a clear diagnosis, a nomen nudum, not validly published (
<emphasis italics="true" pageId="0" pageNumber="81">ICN</emphasis>
Art. 38.2,
<bibRefCitation author="Turland, NJ" journalOrPublisher="Systematic Botany" pageId="0" pageNumber="81" refId="B88" refString="Turland, NJ, 2018. International Code of Nomenclature for Algae, Fungi, and Plants (Shenzhen Code). Koeltz Scientific Books, Koenigstein, Germany." title="International Code of Nomenclature for Algae, Fungi, and Plants (Shenzhen Code). Koeltz Scientific Books, Koenigstein, Germany." year="2018">Turland 2018</bibRefCitation>
). Koch also proposed a taxon
<taxonomicName class="Magnoliopsida" family="Cannabaceae" genus="Cannabis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="himalayensis" order="Rosales" pageId="0" pageNumber="81" phylum="Tracheophyta" rank="species" species="himalayensis">
<emphasis italics="true" pageId="0" pageNumber="81">himalayensis</emphasis>
</taxonomicName>
without a clear diagnosis, and he equated it with the South Asian domesticate - an erroneous concept.
</paragraph>
</subSubSection>
</treatment>
</document>