Models Cannabis Taxonomy

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Models of Cannabis Taxonomy, Cultural Bias, and Conﬂicts between Scientiﬁc

and Vernacular Names
Article in The Botanical Review · June 2017

DOI: 10.1007/s12229-017-9187-0
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Bot. Rev. (2017) 83:327–381
DOI 10.1007/s12229-017-9187-0
Models of Cannabis Taxonomy, Cultural Bias,

and Conflicts between Scientific and Vernacular Names
John M. McPartland 1,2 & Geoffrey W. Guy 1

1
GW Pharmaceuticals, Sovereign House, Histon, Cambridge CB24 9BZ, UK
2
Author for Correspondence; e-mail: mcpruitt@myfairpoint.net
Published online: 22 June 2017
# The New York Botanical Garden 2017
Abstract Debates over Cannabis sativa L. and C. indica Lam. center on their taxo-
nomic circumscription and rank. This perennial puzzle has been compounded by the
viral spread of a vernacular nomenclature, “Sativa” and “Indica,” which does not
correlate with C. sativa and C. indica. Ambiguities also envelop the epithets of wild-
type Cannabis: the spontanea versus ruderalis debate (i.e., vernacular “Ruderalis”), as
well as another pair of Cannabis epithets, afghanica and kafirstanica. To trace the rise
of vernacular nomenclature, we begin with the protologues (original descriptions,
synonymies, type specimens) of C. sativa and C. indica. Biogeographical evidence
(obtained from the literature and herbarium specimens) suggests 18th–19th century
botanists were biased in their assignment of these taxa to field specimens. This skewed
the perception of Cannabis biodiversity and distribution. The development of vernac-
ular “Sativa,” “Indica,” and “Ruderalis” was abetted by twentieth century botanists,
who ignored original protologues and harbored their own cultural biases. Predominant
taxonomic models by Vavilov, Small, Schultes, de Meijer, and Hillig are compared and
critiqued. Small’s model adheres closest to protologue data (with C. indica treated as a
subspecies). “Sativa” and “Indica” are subpopulations of C. sativa subsp. indica;
“Ruderalis” represents a protean assortment of plants, including C. sativa subsp. sativa
and recent hybrids.
Keywords Cannabis sativa . Cannabis indica . Taxonomy . Nomenclature
Introduction
Controversies regarding the taxonomy and nomenclature of C. sativa have generated

four recent publications in The Botanical Review. This present review is intended for a
general audience, i.e., botanists not specializing in taxonomy and nomenclature. We
address three issues considered opaque to generalists; specifically, how taxonomists: 1.
circumscribe units into groups; 2. assign these groups to particular ranks; 3. assign
names with due consideration to priority and other rules of nomenclature.
The first two points are subject to judgment: 1. Group circumscription may vary
depending upon various taxonomic characters and species concepts. 2. Taxonomic
ranks are relative levels within a hierarchy, and notoriously subjective. Darwin (1859)
328 J. M. McPartland, G. W. Guy
could not reconcile the continuous process of evolution with the discrete concept of
taxonomic ranks. “I was much struck how entirely vague and arbitrary is the distinction
between species and varieties.”
The third point is governed by a legalistic document, the International Code of
Nomenclature for Algae, Fungi, and Plants (ICN, McNeill, 2012). The ICN is period-
ically updated, and has become rather complicated. As our mentor Von Arx (1987)
said, “nonspecialists have difficulties in understanding the code and adhering to its
provisions.” To be considered for formal recognition, a botanical name must meet
several technical ICN provisions, e.g., effective and valid publication, typification (with
herbarium specimens), and priority. The ICN is available online for consultation
regarding these provisions. The names Cannabis sativa L. (Linnaeus, 1753) and
C. indica Lam. (Lamarck, 1785) have met these provisions.
Despite C. sativa and C. indica holding nomenclatural legitimacy, their taxonomic
acceptance remains a subjective decision, and depends on one’s species concept. For
example, many biologists (especially zoologists such as Ernst Mayr) conceptualize a
species as all potentially interbreeding populations of an organism. C. sativa and
C. indica are often presumed to have no physiological barriers to breeding (Small,
1972).
If C. sativa and C. indica are a single species according to Mayr’s species concept,
then only one name can be regarded as correct. The ICN decides this by priority, where
the correct name for a taxon is its earliest legitimate epithet. “Correct” and “legitimate”
have technical definitions in the ICN. So does “earliest.” The ICN designates 1753—
the publication year of Linnaeus’s Species Plantarum—as the starting date of botanical
nomenclature.
A review and discussion of Cannabis epithets in the scientific literature is urgently
needed. A vernacular taxonomy of drug-type plants, “Sativa” and “Indica,” has
entangled and subsumed formal C. sativa and C. indica. Thousands of websites
generalize about the morphological, phytochemical, organoleptic, and clinical proper-
ties of “Sativa” and “Indica.” Tracing these collective generalizations to published
sources is elusive; our search traces them back to Corral (2001) and Black and Capler
(2003): “Sativa” is recommended for treating depression, headaches, nausea, and loss
of appetite; it causes a stimulating and energizing type of psychoactivity. “Indica” is
recommended for treating insomnia, pain, inflammation, muscle spasms, epilepsy, and
glaucoma; it causes a relaxing and sedating psychoactivity. “Sativa” plants are said to
produce much more Δ9-tetrahydrocannabinol (THC) than cannabidiol (CBD), and a
terpenoid profile that smells “herbal” or “sweet.” “Indica” plants are said to produce
more CBD than “Sativa,” and a THC-to-CBD ratio closer to 1:1. The terpenoid profile
imparts an acrid or “skunky” aroma.
Small (2007) noted that “Sativa” and “Indica” were “quite inconsistent” with formal
nomenclature, because C. sativa subsp. sativa should apply to non-intoxicant plants.
Conflating formal and vernacular taxonomy has muddled otherwise excellent studies
(e.g., Sawler et al., 2015; Lynch et al., 2015; Belendiuk et al., 2015). It even appears in
Nature (Gould, 2015). Please stay alert to the fact that “Sativa” and “Indica” written in
quote marks mean different things than C. sativa and C. indica written in italics.
The purpose of this paper is to provide historical context to these debates, beginning
with the original protologues of C. sativa and C. indica. To this we add the protologues
of several epithets that touch on the vernacular debate: C. ruderalis, C. sativa var.
Models of Cannabis taxonomy 329
spontanea, C. indica var. kafirstanica, and C. indica var. afghanica. Lastly we review
six prominent post-1970 treatments of these taxa, by Schultes, Anderson, Small, de
Meijer, Hillig, and Gilmore. Our intent is to compare and contrast these existing
taxonomic models, rather than to propose a new model.
Methods
This review utilized three search engines to collect literature (Google Scholar, PubMed,
and Web of Science). We followed PRISMA guidelines for systematic reviews,
including inclusion and exclusion criteria (details provided upon request by the corre-
sponding author).
The ICN defines a protologue as everything associated with a taxonomic name at its
first valid publication. It includes the taxon’s diagnosis or description, synonymy,
discussion, illustrations, references, and type specimen (McNeill, 2012). A type spec-
imen is a physical reference point linked to a taxonomic name: a dried plant stored in a
herbarium. Type specimens authenticate a taxon’s description, but they may not serve
as exemplars of a taxon. Characters exhibited by type specimens may results from
environmental modifications.
Fifteen herbaria were consulted, designated by herbarium acronyms in Index
Herbariorum: B (Berlin), BM (British Museum, London), BPI (Beltsville, MD),
CUP (Cornel University), F (Field Museum, Chicago), ECON (Economic botany,
Harvard), GH (Gray, Harvard), IND (Bloomington, IN), K (Kew, London), LE (“Le-
ningrad,” St. Petersburg), LINN (Linnaeus, London), NY (Bronx, New York), P
(Paris), PH (Philadelphia), US (Smithsonian, Washington DC), WIR (Vavilov Institute,
St. Petersburg).
Results
The search algorithm identified many potential articles: Google Scholar, 1490 hits;
PubMed, 362 hits; Web of Science, 16 hits. Screening revealed the majority of these
were irrelevant (details provided upon request by the corresponding author). Our yield
was greatly enhanced by screening retrieved articles for supporting citations, and
retrieving these antecedent sources. The results of this systematic search are presented
in a narrative review, structured from an historical perspective. To maintain narrative
flow, fine details in this review are relegated to four Appendices.
Cannabis sativa L. Protologue
C. sativa L. 1753 holds priority under the ICN—the first Cannabis taxon appearing
after the starting date of botanical nomenclature. Botanists prior to Species Plantarum
also used the binomial, such as Caspar Bauhin and Leonhart Fuchs. The unrecognized
author who first coined C. sativa was Ermolao Barbaro, in a text published 23 years
after his death (Barbaro, 1516).
The description of C. sativa by Linnaeus (1753) was exceptionally brief: a generic
account of flower parts, applicable to the genus Cannabis. Linnaeus described the seed
(Latin: semen) as a nut (nux). Edwards and Vavasseur (1829) first referred to the seed as
an achene. We shall refer to the seed as a seed, unless quoting someone else.
Linnaeus listed five facultative synonyms and their five authors. All synonyms and
authors cited by Linnaeus came from northern Europe. Linnaeus notably excluded
Asian taxa from the C. sativa synonymy. This departs from his earlier taxonomic
synopsis of Cannabis, where Linnaeus (1737) synonymized several taxa assigned to
psychoactive Asian Cannabis. These Asian taxa appeared in a 1746 draft of Species
Plantarum (manuscript at the Linnean Society of London), but they were deleted from
the final version.
Linnaeus’s herbarium specimens also came from northern Europe. The type speci-
men, a seeded pistillate plant, is deposited at LINN (Fig. 1). Stearn (1974) provided
evidence that Linnaeus collected it in Sweden. Its morphology is consistent with a
northern European fiber-type landrace. The inflorescences are loose, not dense;
subtending floral leaves have a sparse covering of capitate-sessile glandular trichomes;
perigonal bracts have a relatively sparse covering of capitate-stalked glandular tri-
chomes. Seeds are large (4.8 × 2.5 mm), oblong in outline, pale green with a fine
reticulated pattern, with a short and pointed base with a simple articulation. Other
C. sativa specimens collected by Linnaeus (at BM) are consistent with “the old
cultivated hemp stock of northern Europe” (Stearn, 1974). We provide a full account
of Linnaeus’s description, synonymy, and references in Appendix 1a.
Fig. 1 Type specimens of C. sativa L. (left), and C. indica Lam. (right), photographs courtesy of McPartland
and Herb. P, respectively
Cannabis indica Lam. Protologue
Lamarck (1785) coined C. indica for plants with provenance from India, Southeast
Asia, and South Africa. He cited six synonyms and their authors. For a full account of
his protologue see Appendix 1b. Lamarck’s description of C. indica differed from his
description of C. sativa by eight “very distinct” morphological characters in stalks,
branching habitus, leaflets, and flowers. Lamarck also described chemotaxonomic
differences: C. indica produced a strong odor, and was psychoactive, “The principal
effect of this plant consists of going to the head, disrupting the brain, where it produces
a sort of drunkenness that makes one forget one’s sorrows, and produces a strong
gaiety.”
Lamarck’s type specimen at P is annotated “Chanvre rapporté de l’Inde par M.
Sonnerat.” Pierre Sonnerat made extensive botanical collections around Pondicherry.
The specimen’s unpollinated pistillate (“sinsemilla”) morphology is representative of a
South Asian landrace. It shows vigorous branching, with shorter internodes than
Linnaeus’s specimen (Fig. 1). Its inflorescences are somewhat compact; subtending
floral leaves have an abundant covering of capitate-sessile glandular trichomes;
perigonal bracts express a moderate density of capitate-stalked glandular trichomes.
The styles and stigmas are prominent, agglutinated with trichome exudate, and light
brown in color.
Historical Bias in Usage of C. sativa and C. indica
Botanists soon rendered subjective decisions regarding Lamarck’s concept of a

polyspecific genus. Willdenow (1805) reduced C. indica to a synonym C. sativa, rather
than a separate variety. He argued that no diagnostic differences existed between them,
because both species showed alternate branching (Lamarck erroneously claimed
C. indica was unique in this regard). Willdenow ignored Lamarck’s seven other
morphological differences. He also ignored Lamarck’s phytochemical differences,
possibly because Linnaeus (1751) rejected chemotaxonomic characters, such as fra-
grance and taste. Similarly, most British botanists in India cited C. sativa and ignored
C. indica.
We find evidence of cultural bias influencing these taxonomic decisions, arising
from personality cults surrounding Linnaeus and Lamarck. Today it is difficult to
fathom Linnaeus’s renown among botanists in the 18th–19th centuries, and the enmity
provoked by Lamarck’s deviation from Linnaean orthodoxy. See Appendix 2 for
attestations. For example, Willdenow was a disciple of Linnaeus, and updated
Species Plantarum after Linnaeus died. Willdenow (1805) rejected over half of
Lamarck’s new taxa (most have been reinstated by modern taxonomists). Early British
botanists working in India were tutored by Johann König, a student of Linnaeus.
William Roxburgh became the most influential member of König’s “United Bretheren”
in India. Roxburgh (1832) wrote, “I perfectly agree with Willdenow in thinking all the
varieties, if even such they can be called, centre in one species.”
It is expected that the distributions of C. sativa and C. indica (at species or
subspecies rank) would occupy separate floristic regions. The distribution of plants
that field botanists identified as C. sativa or C. indica in the 18th–19th centuries is
presented in Fig. 2. The map shows no hint of endemic distribution, for either C. sativa
Fig. 2 Taxon names applied by field botanists. Locations of “C. sativa” labeled alphanumerically, s1, s2, etc.
Locations of “C. indica” labeled alphanumerically, i1, i2, etc. Locations of other Cannabis taxa labeled
alphanumerically, ×1, ×2, etc. Base map shows boundaries of ten floristic regions by Takhtajan (1986). For full
citations of alphanumerical sites see Appendix 3
or C. indica, within any single floristic region. Even allowing for zoochory (animal
transport of germplasm, including humans) the distributions of C. sativa and C. indica
randomly overlap. The penetration of “C. sativa” deep into Asia is particularly striking.
As field botanists explored Asia and Africa, they applied the names C. sativa or
C. indica to plants inconsistent with the protologues of Linnaeus and Lamarck. In many
locations, both names were applied, by different botanists. In general, botanists from
Scandinavia and Great Britain used “C. sativa.” French and Francophone Russian
botanists used “C. indica” or coined new taxa, such as such as C. chinensis and
C. gigantea. Some botanists deposited herbarium specimens, and our examination of
their specimens verified erroneous determinations.
Wild-Type Cannabis
Reports of Cannabis growing in two phases of domestication—wild and cultivated—

trace back to Scythia, now Ukraine, in 440 BC (Herodotus, 2007). We prefer the phrase
“wild-type,” a term that includes a spectrum of plants. This spectrum was delineated by
Small (1984, 2015), ranging from truly wild (i.e., native, indigenous, aboriginal), feral
(i.e., weedy, naturalized), ruderal (either wild or weedy), to spontaneous (escapes of
cultivated plants). Russian naturalists encountered wild-type Cannabis near present-day
Saratov, and debated whether they were truly wild or escapes of cultivated plants
(Lepechin, 1774; Pallas, 1793).
Zinger (1898) first described wild-type characteristics (Fig. 3). These included small
seed size, a protuberant-and-tapered base with a prominent abscission zone, and a
persistent perianth (seed covering). The perianth provides camouflage coloring—a
matte surface (rather than a shiny pericarp) with irregular dark spots, called marbling,
mottling, or mosaic by various authors. The protuberant base has been described as a
horseshoe, circular torus, callus-ring, caruncle, basal constricted zone, or elaiosome.
Two botanists in Saratov, Dmitry Janischevsky and Nikolay Vavilov, studied wild-
type Cannabis. Vavilov (1922) coined the taxon C. sativa var. spontanea. His descrip-
tion is brief and limited to wild-type seed morphology. He added a physiological trait:
rapid seed disarticulation from the plant, due to a prominent abscission zone. Vavilov
Fig. 3 The wild-type phenotype. Left: domesticated versus wild-type Cannabis fruits, illustrated by Zinger
(1898). Right: type specimen of C. sativa var. spontanea Vav. (McPartland photo, herb. WIR)
described C. sativa var. spontanea in greater detail in subsequent publications. For his
full protologue and later descriptions, see Appendix 1c.
Vavilov’s lectotype specimen at WIR is illustrated in Fig. 3. The specimen consists
of the top 42 cm of what appears to be a tall plant, with internode spacing similar to
Linnaeus’s type specimen. Leaves have 3–5 leaflets, petioles short, leaflets narrow
lanceolate, up to 130 × 10 mm. Inflorescences are loose; subtending floral leaves have a
sparse covering of sessile glandular trichomes; perigonal bracts have few capitate-
stalked glandular trichomes and many cystolith trichomes. Seeds are medium-sized
(3.8–4.0 mm long), with reticulate venation between irregular dark marbling, and a
weakly protuberant base.
Janischevsky (1924) erected C. ruderalis, plus an alternative taxonomic rank:
C. sativa var. ruderalis. “I am inclined to consider it a well marked variety.” Janischevsky
penned a 13-page protologue (Appendix 1d), including a Latin diagnosis and two plates
of illustrations (all of seeds). He described a moderately tall plant, usually 0.7–1.1 m but
up to 2.1 m, branchier than domesticated plants, with wild-type seed characteristics.
Janischevsky’s lectotype specimen of C. sativa var. ruderalis was stored at LE (Small &
Cronquist, 1976). It may be lost (McPartland, pers. observ., St. Petersburg, 2010).
Janischevsky’s bibliography is extensive; he cited literature by 22 authors.
Two taxonomic names that represent contaxic entities, based on different type
specimens, are facultative synonyms. Vavilov’s taxon has priority at the rank of variety,
Janischevsky’s taxon has priority at the rank of species. Soviet botanists in the 1930s
set precedence by choosing C. ruderalis over C. sativa var. spontanea. We find
evidence of cultural bias influencing their decisions, during the era of Lysenkoism.
See Appendix 1d for attestations.
Historical Bias, Part II
Vavilov traveled to Afghanistan in 1924, and encountered all phases of domestica-

tion—wild, weedy, and cultivated. He described Afghani farmers growing Cannabis
exclusively for gashisha (“a grinded gray-green powder,” i.e., sieved hashīsh). He
assigned these plants to C. sativa (Vavilov & Bukinich, 1929), a concept that departed
from Linnaeus’s protologue of C. sativa as a European fiber-type plant.
Vavilov’s error likely derived from Tatiana Serebriakova, his assistant in Russia.
Vavilov sent seeds to Serebriakova, and she grew the accessions. Vavilov and Bukinich
(1929) presented her data, and then made a statement that seems to quote her, “By all
attributes it [Afghani plants] is linked directly with the Central Asian cultivated hemp, not
with C. indica, which is distinguished by its small leaves, small fruits, and low height (up
to 1 meter).” Serebriakova’s concept of C. indica as a diminutive plant with small leaves
is faulty—plants from India are relatively tall with large leaflets. Serebriakova and
Vavilov likely never saw a specimen of C. indica from India. The herbarium collection
at WIR lacks specimens from India (McPartland, pers. observ., St. Petersburg, 2010).
Vavilov discovered feral and wild-type plants in Afghanistan, and coined two new
taxa for them. Vavilov (1926) assigned C. sativa f. afghanica to feral plants with
obovate leaflets (a shape he said was not seen in European plants), medium height, and
profuse branching. The plants “constitute a morphological link between the wild and
the cultivated races of hemp.” Subsequently he transferred the taxon to C. indica, as
C. indica f. afghanica, and mentioned the alternative rank C. indica var. afghanica
(Vavilov & Bukinich, 1929). Some botanists argue that afghanica designates a wild-
type plant. But Vavilov’s descriptions, illustrations, and other evidence indicate that
afghanica was an escape of cultivated plants (Appendix 1e).
Vavilov and Bukinich (1929) coined C. indica var. kafirstanica for wild-type plants,
with very small seeds, dark-colored and mottled, with a “horseshoe” (Appendix 1f).
They muddied taxonomic waters by assigning afghanica to a third alternative rank,
C. indica var. kafiristanica forma afghanica.
Vavilov (1931) extended the range of C. sativa var. spontanea to Central Asia,
where he encountered wild-type plants. However, specimens deposited at WIR and LE
show that Vavilov assigned C. sativa var. spontanea to plants that departed from his
own protologue of the taxon—they are extremely short (≤0.61 m), unbranched, with
broad leaflets.
A new cultural bias appeared in the 1970s. Cannabis taxonomy became entangled in
the USA legal system, due to perceived threats to society regarding increased Cannabis
use. Whether Cannabis is polytypic (one species with many kinds) or polyspecific
(more one than one species) became a polarized debate in court cases. The primary
adversaries in these legal wrangles were Richard Evans Schultes, a Harvard ethnobot-
anist, and Ernest Small, a Canadian taxonomist who specialized in Cannabis.
Small, an expert witness on behalf of plaintiffs, argued for a single species, C. sativa.
“This arrangement is in harmony with social needs and significance” (Small &
Cronquist, 1976). He backed his legal testimony with a lot of data. Small and
colleagues conducted cross-breeding experiments, and they assessed many taxonomic
characters—morphological, biogeographical, developmental, and phytochemical (can-
nabinoid content, seed oil chemistry)—summing to 20 publications (Appendix 4).
Small and Cronquist (1976) stayed true to Linnaeus’s concept of sativa, Lamarck’s
concept of indica (albeit as a subspecies), and Vavilov’s concept of C. sativa var.
spontanea. They treated C. indica f. afghanica as a feral escape of cultivated plants,
separate from C. indica var. kafiristanica. They departed from Vavilov’s concept of
C. indica var. kafiristanica by extending the taxon to wild-type plants with narrow
lanceolate leaflets, growing outside of Afghanistan, in India, Nepal, China, South

Africa, and Columbia.
Small’s taxonomic concept is relatively simple: a two-step hierarchic classification
system (Fig. 4). The first step recognizes two subspecies based on THC content in
dried female flowering tops (with 0.3% THC as the dividing point). The second step
recognizes two varieties within each subspecies, based on their domestication phase.
However, analysis of THC/CBD ratios revealed three chemovars, Type I: THC
>0.3%, CBD <0.5%; Type II: THC >0.3%, CBD >0.5%; or Type III: THC <0.3%,
CBD >0.5%.
Type I chemovars were drug-type plants, mostly with provenance from countries
below the 30th parallel (30° N runs through Morocco, Iran, northern India, and
southern China; 30° S runs through South Africa, South America, and Australia). Type
III chemovars were fiber-type plants, mostly with provenance north of 30°N. Type II
chemovars were a mix of drug- and fiber-type plants; Small stated that they may reflect
hybridization between Type I and III.
Small’s taxonomic concept is cited and accepted by many botanists, and attested on
botanical websites (Appendix 4). Nevertheless his model has been criticized for being
“not natural at all” and ignoring morphological diversity (De Meijer, 1999), and for
methodological flaws (Hillig & Mahlberg, 2004). Small provided “passport data”
regarding the provenance of his accessions, and deposited voucher specimens of each
accession in several herbaria. Vouchers are critical for authenticating the identification
of a taxon; they allow other researchers to retrospectively analyse accessions (Culley,
2013). Several accessions came from botanical gardens, of questionable provenance
(e.g., three indica accessions with no measurable THC).
Schultes served an expert witness on behalf of defendants, whose attorneys argued
that narcotics laws cited C. sativa, whereas the accused possessed C. indica, a species
that was statutorily overlooked and technically legal. Schultes argued in favor of three
Cannabis species by reversing his opinion on Cannabis, which he initially considered a
monotypic genus (Schultes, 1970). Subsequently he recognized C. sativa, C. indica,
and C. ruderalis, and summed his research in a single paper (Schultes et al., 1974). He
changed his opinion based on analyses of herbarium specimens, a survey of the
Mississippi Cannabis plantation, and “preliminary field work” in Afghanistan.
Fig. 4 Classification of Cannabis sativa, according to Small & Cronquist, 1976

A former student explained that Schultes’s “intense dedication” to libertarian beliefs

led him to employ this “obscure taxonomic argument” (Davis, 1996). “In truth, the
evidence for Schultes’s position was somewhat dubious.” Schultes et al. (1974)
departed from the concepts of Lamarck, Vavilov, and Janischevsky:
& C. indica becomes narrowly typified to plants Schultes saw in Afghanistan, with
broad, oblanceolate leaflets, densely branched, more or less conical in shape, and
very short (< 1.3 m). Designating these plants as C. indica is faulty; Lamarck was
entirely unfamiliar with Afghani Cannabis. Lamarck’s protologue of C. indica
describes plants that are relatively tall, laxly branched, with narrow leaflets.
& By assigning cultivated Afghani plants to C. indica, Schultes departs from
Vavilov’s concept of cultivated Afghani plants as C. sativa.
& Schultes eschews Vavilov’s spontanea in favor of Janischevsky’s ruderalis. He
departs from Janischevsky’s concept of C. ruderalis by applying the taxon to
extremely short (≤0.61 m), unbranched plants with broad leaflets from Central
Asia, instead of Janischevsky’s relatively tall, laxly branched plants with narrow
leaflets from southeastern Europe.
Schultes’s concepts were embraced by two other defense witnesses, William

Emboden and Loran Anderson. Emboden (1972) originally considered Cannabis a
monotypic genus. Emboden (1974) reversed his opinion. He recognized C. sativa
(specimens from Afghanistan, Russia, Kansas), C. indica (specimens from Afghani-
stan, Turkey, India), and C. ruderalis (a specimen from Russia).
Anderson (1980) typified C. indica with plants Schultes collected in Afghanistan.
He assigned C. sativa to plants consistent with Lamarck’s C. indica—relatively tall,
laxly branched, narrow leaflets, with provenance from India. His concept of
C. ruderalis was consistent with Schultes, not Janischevsky. Anderson illustrated these
concepts in a line drawing (Fig. 5) This illustration has become pervasive on the web as
the poster child of vernacular nomenclature.
The Rise of Vernacular Taxonomy
Cannabis germplasm from Afghanistan was smuggled into the USA by the early
1970s. Differences were readily apparent between Afghani plants and drug plants
previously grown in the USA (with provenance from below 30° N, e.g., Mexico,
Columbia, Thailand). Stevens (1975) illustrated morphological differences between
Thai and “Afghanistani” plants. Frank and Rosenthal (1978) explained Schultes’s
three-species concept, and conflated it with Small’s three-chemotype concept.
After the publication of Anderson’s line drawing (Fig. 5), Clarke (1981) referred to
plants from Afghanistan “as type examples for Cannabis indica.” He noted the unique
smell of Afghani plants, an acrid odor rather than a sweet fragrance. Clarke (1987) first
described the unique organoleptic properties of Afghani plants, a “slow flat dreary
high.”
Cherniak (1982) applied the name “Cannabis indica” to plants from Afghanistan.
Their morphology, odor, and potency differed from “Cannabis sativa” strains. He
assigned “Cannabis sativa” to plants of South Asian heritage (Nepal, Burma, Thai-
land), and their descendants in Columbia, Jamaica, and Mexico. However, Cherniak
Fig. 5 Line drawing by Anderson (1980), courtesy of the Harvard University Herbaria and Botany Libraries
also categorized plants from India as “Cannabis indica.” The earliest consistent use of
“Sativa” and “Indica” we could find was in a Dutch seed catalog (Watson, 1985).
Schoenmakers (1986) collected “Ruderalis” germplasm near the Hungary-Ukraine
border. His photos of “Ruderalis” show plants with strong apical dominance and little
branching. These traits are consistent with a spontaneous escape of cultivated hemp,
and depart from concepts by Vavilov and Janischevsky. Alternatively, Frank (1988)
said that U.S. marijuana growers assigned “Ruderalis” to plants with “wide indica-like
leaf blades, and these might be hybrids.” In today’s vernacular taxonomy, “Ruderalis”
is applied to an increasingly protean assortment of plants. They exhibit one to three
characteristics: CBD ≅ THC, wild-type morphology, or early flowering (sometimes
called “autoflowering,” i.e., day-neutral, flowering not induced by light cycle).
Within “Sativa,” “Indica,” and “Ruderalis,” plant breeders have designated “strain”
names. Watson (1985) coined several well-known strain names, such as “Original
Haze” and “Skunk #1”. Cannabis breeders equate “strains” with “cultivars,” which
flouts another code, the International Code of Nomenclature for Cultivated Plants
(ICNCP, Brickell, 2009). The ICNCP has rules for designating cultivars, such as valid
publication, typification (“Nomenclatural Standard”), and priority.
Small (2015) pointed out that strains are conceptually identical to cultivars, but
almost no Cannabis strains have met ICNCP requirements for cultivar recognition.
Seedfinder (2015) listed 6510 strain names. Doyle (2007) referred to these names as
ganjanyms. We elaborate upon this controversy in Appendix 4.
Since the courtroom days of Schultes and Small, other botanists have addressed
C. sativa, C. indica, “Sativa,” and “Indica.” Google Scholar computes citation metrics
for authors and their publications. Taxonomic publications garnering the greatest
number of citations are by Etienne de Meijer, Karl Hillig, and Simon Gilmore.
De Meijer and colleagues analyzed cannabinoid profiles, morphological characters,
developmental traits, host-parasite characters, inheritance of chemical phenotypes and
SCAR markers, and sequence heterogeneity in cannabinoid synthase genes—summing
to 11 publications (Appendix 4). De Meijer and van Soest (1992) introduced vernacular
taxonomy to peer-reviewed literature: “Indica” referred to plants with broad leaflets,
compact habit, and early maturation, typified by plants from Afghanistan. “Sativa”
referred to plants with narrow leaflets, slender and tall habit, and late maturation,
typified by plants from India and their descendants in Thailand, South- and East Africa,
Colombia, and Mexico.
De Meijer (1999) argued for a single species, C. sativa, noting a lack of breeding
barriers and morphological discontinuities within the genus. He subdivided the species
into populations, using criteria from Schultes and Anderson. He disregarded priority by
naming C. sativa var. spontanea a synonym of C. ruderalis. De Meijer considered
cultivated and ruderal forms as extremes along a gradient, so he did not recognize wild-
type plants as a separate taxon (de Meijer, pers. commun., 2010). De Meijer provided
passport data regarding his accessions, but did not deposit voucher specimens in a
herbarium. His model has been criticized for methodological flaws (Hillig & Mahlberg,
2004), and a faulty inheritance model (Weiblen et al., 2015).
Hillig and colleagues analyzed genetic (allozyme) variation, cannabinoid profiles,
terpenoid profiles, morphological characters, developmental traits, and host-parasite
characters—summing to ten publications (Appendix 4). Hillig sidestepped vernacular
nomenclature. He applied the name “narrow-leaflet drug (NLD) biotype” to plants
corresponding to “Sativa,” and applied the name “wide-leaflet drug (WLD) biotype” to
plants corresponding to “Indica.” Hillig’s use of NLD/WLD is reminiscent of
Linnaeus’s use of ♀ and ♂ symbols to sort the confusing gender-based names applied
by earlier botanists.
Hillig provided passport data and voucher specimens. He made efforts to identify
and exclude hybrids, such as NDL x WLD drug crosses, whose recent breeding
histories are documented (e.g., De Meijer, 1999). He analyzed landraces, rather than
hybrid “strains,” (see example at end of Appendix 4) to enhance phylogenetic distinc-
tions. Although de Meijer (pers. commun., 2010) derides the exclusion of hybrids as
“cherry-picking,” it is a standard procedure (e.g., Kaplan & Fehrer, 2004). New
software for inferring population structure makes it easier to identify and exclude
hybrids (e.g., Cornille et al., 2013).
Hillig (2004, 2005) recognized two species, C. sativa and C. indica. The first species
segregated into two biotypes—a domesticated phase that corresponded with Linnaeus’s
sativa, and a wild-type phase that corresponded with Vavilov’s spontanea. He segre-
gated C. indica into four biotypes. The NLD biotype corresponded with Lamarck’s
indica. The WLD biotype corresponded with Vavilov’s afghanica. The “C. indica feral
biotype” referred to wild-type plants from India and Nepal—accessions assigned to

kafiristanica by Small and Cronquist (1976). The “C. indica hemp biotype” referred to
fiber-type plants from East Asia. Hillig noted that some nineteenth century botanists
segregated Chinese hemp from European C. sativa, reflecting their putative domesti-
cation from different gene pools. He traced the nomenclature to C. chinensis Delile,
1849, although this is a homonym of C. chinensis Fischer, 1810 (Appendix 3).
Hillig also proposed a third species, C. ruderalis. His concept of C. ruderalis as
Central Asian in origin corresponded with the concepts of Schultes and Anderson,
rather than Janischevsky. De Meijer (2014) criticized Hillig’s model for an over-
reliance on characters subjected to human selection, such as cannabinoid profiles,
although Hillig initially based his model on genetic data. Hillig’s nomenclature has
gained traction (e.g., McPartland & Guy, 2014; Russo, 2007; Lynch et al., 2015).
Clarke and Merlin (2013) objected to biotype names based on leaflet shape. They
erected a new biotype nomenclature, for example substituting Hillig’s “wide-leaflet
drug” with their “broad leaf drug” biotype. Botanically, “leaflet” is more accurate than
“leaf.”
Gilmore and colleagues conducted genetic studies, summing to six publications
(Appendix 4). Gilmore et al. (2007) sequenced five polymorphic loci (cpDNA and
mtDNA) from 76 CPRO accessions. Parsimony identified six haplogroups, which
nested into three clades: Clade A comprised mostly European fiber plants. Clade B
comprised a mix of Eurasian fiber plants, Central Asian drug plants, and hybrids. Clade
C comprised drug plants from South Asia (India, Nepal) and their putative descendants.
Gilmore noted that clade C corresponded with “drug-breeders taxon Sativa.”
Some authors have tried to reconcile “Sativa” and “Indica” with formal C. sativa and
C. indica. McPartland et al. (2000) noted that Afghani plants were mislabelled
“Indica.” They reassigned “Indica” to Vavilov’s taxon, at species rank (C. afghanica)
or varietal rank (C. sativa var. afghanica). McPartland and Guy (2014) and McPartland
(2014) proposed reconciling vernacular and scientific nomenclature: “Sativa” is really
indica, and “Indica” is actually afghanica, and “Ruderalis” is usually sativa.
The initial reaction to this proposition by recreational users was negative. An
editorial in High Times characterized the reconciled nomenclature as “undoubtedly a
little kooky” (Sirius, 2015). Researchers, however, are starting to take it on board (e.g.,
Henry, 2015). Clarke and Merlin (2016) erected a nearly identical reconciliation,
without citing precedent publications. Two table headings in their respective taxonomic
tables are exampled:
& Indica—Wrongly called “sativa” (Clarke & Merlin, 2016)

& Indica (formerly “Sativa”) (McPartland, 2014)
Rather than reconciling nomenclature, some authors call for a new approach. Arno
Hazekamp and colleagues analyzed cannabinoid and terpenoid profiles in hundreds of
herb samples obtained from Dutch coffeeshops and other sources. Their methods
segregated “Hemp,” “Sativa,” and “Indica.” But they propose jettisoning all vernacular
names in favor of a metabolomics approach, “from cultivar to chemovar” (Hazekamp
& Fischedick, 2012; Hazekamp et al., 2016).
Other taxonomic projects have analyzed “Sativa” and “Indica” herb samples, rather
than C. sativa and C. indica plants cultivated in a common garden experiment. Elzinga
et al. (2015) measured THC and CBD content in 35 samples obtained from
“chemotypical medicinal cannabis dispensaries.” They assigned strains to “Indica,”
“Sativa,” or “Hybrid” based on reports by the Leafly website. Their analysis “does not
support the classification between indica and sativa as it is commonly presented.”
Sawler et al. (2015) genotyped 43 fiber-type cultivars and 81 drug-type strains.
Drug-type strains were classified along a gradient of ancestry (percent “Sativa” vs.
percent “Indica”) reported in online strain databases. Population stratification using
SNP analysis with PLINK segregated clusters of fiber-type and drug-type samples,
whereas clusters of “Sativa” and “Indica” overlapped. They concluded that “Sativa”
and “Indica,” as well as strain names, did not reflect a meaningful genetic identity.
Lynch et al. (2015) conducted a genotype-chemotype study with 195 samples
obtained from medical dispensaries, breeders, or “by donation.” Haplotypes based on
SNP data were analyzed with FLOCK. This algorithm identifies the optimal number of
clusters (K) to divide a population, and their data optimized at K = 3. They character-
ized the three groups as “broad leaflet drug” (e.g., “Afghan Kush,” “Chemdawg”),
“narrow leaflet drug” (e.g., “Durban Poison,” “Easy Sativa”), and a polyphyletic
“hemp” group (e.g., ‘Finola,’ “AC/DC,” Chinese hemp, Dagestan plants). They ob-
tained THC/CBD data for 54 of the 195 samples via the Strain Fingerprint™ database,
developed by Steep Hill Labs. They presented histograms of mean THC% and CBD%,
from which cannabinoid ratios can be estimated: BLD: 16.5/0.2 = 82.5, NLD: 14.2/
2.2 = 6.45. Thus the latest THC/CBD ratios of twenty-first century ganjanyms show a
stunning reversal compared to their corresponding 1970s–1990s landraces. See
McPartland (2017) for an elaboration of this phenomenon.
Discussion
Considering the controversy over Cannabis taxonomy, it seems surprising that no one
has yet presented full protologues of C. sativa and C. indica for comparison. The
protologues by Linnaeus and Lamarck show taxonomic acumen for their time. We
cannot know why Linnaeus omitted Asian material from the C. sativa protologue, but
in retrospect it was a good decision. Lamarck chose taxonomic characters that modern
botanists still use to demarcate subpopulations within the genus.
Linnaeus harbored a “Eurocentric bias” in taxonomy and nomenclature, and taxon-
omists following him showed “secret conservative sympathies” (Walters, 1986). We
see this in the use of C. sativa and C. indica by 18th–19th century field botanists, which
showed no bioregional endemicity (Fig. 2). Linnaeus’s followers applied the name
C. sativa to plants in Asia, whereas botanists unconnected to Linnaeus used C. indica
or coined new taxa (Appendix 3). C. sativa became a metonym rather than a taxon.
Various types of socio-economic biases are known to skew floristic distribution maps
(Yang et al., 2014). Understanding patterns of Cannabis biogeography requires im-
provements in systematics, as well as hermeneutics (texual criticisms). We also see
cultural bias in Soviet botanists’s use of C. ruderalis during the Lysenko era, and
judicial concerns in the 1970s skewed concepts of Cannabis as a monospecific versus
polytypic genus.
We return to three issues in the first paragraph of our introduction: Taxonomists have
circumscribed Cannabis populations using morphology, chemotaxonomy, and genetics.
These characters have not provided robust taxonomic boundaries in recent studies. We
attribute this to the inclusion of recently hybridized accessions, which makes it difficult
to detect taxonomic dichotomies. Hillig did his best to exclude hybrids from analysis,
which enabled him to differentiate WLD and NLD populations within C. indica.
Hybridization threatens Cannabis biodiversity, which has become endangered by
widespread crossbreeding over the past 40 years. Wiegand (1935) first described
extinction through introgressive hybridization. Introgression refers to the infiltration
of genes between taxa through the bridge of F1 hybrids. Fertile offspring from these
crosses may display hybrid vigor (enhanced fitness), and replace one or both parental
populations (Ellstrand, 2003). Nearly 30 years ago, unhybridized plants of Indian
heritage and Afghani landraces had already become difficult to obtain (Clarke, 1987).
Small (2017) agrees that hybridization has largely obliterated population differences,
“especially between the two kinds of fiber forms and between the two kinds of
marijuana forms.”
The issue of taxonomic rank continues to bedevil Cannabis taxonomy. Do C. sativa
and C. indica segregate at the rank of species, subspecies, or varieties? Gilmore et al.
(2007) found little sequence variation in Cannabis (approximately 1 polymorphism per
1 kb sequenced cpDNA), which suggested their haplotypes segregated at a rank below
that of species. McPartland and Guy (2014) used similar loci (rbcL, matK, trnH-psbA,
trnL-trnF, ITS) to measure divergence between plant populations. The mean divergence
between C. sativa and C. indica equaled 0.41%. In comparison, a mean divergence of
0.43% existed between five pairs of plants considered different varieties (e.g., Camellia
sinensis var. sinensis and C. sinensis var. assamica). A mean divergence of 3.0%
separated five pairs of plants considered different species (e.g., Humulus lupulus and
H. japonicus).
Sawler et al. (2015) calculated the fixation index (FST) between groups. FST values
range from 0 to 1; a zero value indicates the groups freely interbreed; a 1 value
indicates the groups are completely isolated from one another. They calculated a FST
of 0.156 between fiber- and drug-type groups. This is similar to the degree of genetic
differentiation between human populations in Europe and East Asia. Lynch et al.
(2015) calculated a FST value of 0.099 between fiber- and drug-type groups, and a
FST of 0.036 between WLDs and NLDs.
The third issue—assigning names—is strictly governed by rules of nomenclature in
the ICN and ICNCP. However, vernacular “Sativa,” “Indica,” and “Ruderalis” threaten
to subsume formal ICN nomenclature. Many authors have derided the inaccuracy of
vernacular taxonomy (e.g., McPartland et al., 2000; Small, 2007; Hazekamp &
Fischedick, 2012; Russo, 2016; Clarke & Merlin, 2016).
In conclusion, this review traces the taxonomic histories of C. sativa and C. indica,
beginning with protologues by Linnaeus and Lamarck. Disregard of these protologues
and taxonomic bias by 18th–19th century botanists impacted upon reports of Cannabis
biodiversity and distribution. Further disregard by twentieth century botanists laid the
groundwork for “Sativa,” “Indica,” and “Ruderalis” concepts. These names do not
align with formal botanical nomenclature, but they do represent population segregates
that are threatened by extinction through introgressive hybridization. Even “the old
cultivated hemp stock of northern Europe” (Stearn, 1974) is threatened by hybridiza-
tion. Perhaps the first step in their preservation is taxonomic recognition, a subject for
future publications.
Acknowledgements We thank Karl Hillig, Patricia Pruitt, and Ernest Small for reviews of the manuscript.
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Appendix 1: Protologues
Appendix 1a. Cannabis sativa L.
Diagnosis or description
Linnaeus (1753) classified C. sativa in Class Dioecia (dioecious plants), and Order
Tetrandria (male plants have four stamens). These aspects of morphology are the only
descriptive elements he provided in Species Plantarum. Even by Linnaeus’s standards,

this is an extremely brief description. For genera with only one species, such as
Cannabis, Linnaeus provided a longer description in Genera Plantarum (Linnaeus,
1754). For this reason, the ICN explicitly links taxa coined in Species Plantarum with
their descriptions in the fifth edition of Genera Plantarum. The Cannabis description in
Genera Plantarum is limited to flower and seed morphology. Translated from Latin:
MALE. Calyx: Perianth five-parted: leaves oblong, acuminate-obtuse, concave.
Corolla: absent. Stamens: filaments five in number, thread-like, short. Anthers: oblong,
quadrangular.
FEMALE. Calyx: Perianth single-leaved, oblong, acuminate, opposite at the base,
dehiscing longitudinally, persistent. Corolla: absent. Pistil: Ovary minute, Style divid-
ed, sharply pointed, long. Stigma acute. Perianth minimal. Calyx tightly closed. Seed:
Nut globose-depressed, bivalvate.
Note a discrepancy: Species Plantarum Linnaeus (1753) described males with four
stamens, whereas Genera Plantarum (Linnaeus, 1754) described plants with five
stamens. Five stamens is correct. The correct version was passed down verbatim from
the first edition of Genera Plantarum, written in 1737. Hence the 1753 error was
typographic. Linnaeus corrected his mistake in the second edition of Species Plantarum
in 1763.
Synonymy and references

Linnaeus (1753) cited five C. sativa synonyms by five authors, presented in his typical
telegraphic style. We provide a translation of Linnaeus’s abbreviations in Box 1. Note a
typographic error: Linnaeus said C. foliis digitatis appeared in Materia Medica on page
475. In fact the taxon appears in Materia Medica on page 162 (Linnaeus, 1749).
Box 1 Translation of author and citation abbreviations by Linnaeus (1753)
Cannabis foliis digitatis Linnaeus, Hortus Cliffortianus (1738: p. 457); Linnaeus, Hortus Upsaliensis (1748:
p. 297); Linnaeus, Materia Medica (1749: p. 162, no. 457); Dalibard, Florae Parisiensis Prodromus (1749:
p. 200); van Royen, Florae Leydensis prodromus (1740: p. 221).
Cannabis sativa Bauhin, Pinax Theatri Botanici (1623: p. 320). ♀
Cannabis mas Daléchamps (d’Aléchamps), Historia generalis Plantarum (1587: p. 497). ♀
Cannabis erratica Bauhin, Pinax Theatri Botanici (1623, p. 320). ♂
Cannabis femina Daléchamps (d’Aléchamps), Historia generalis Plantarum (1587: 497). ♂
Linnaeus notably excluded taxa of Asian Cannabis from the C. sativa protologue.
Sixteen years earlier, in his previous synopsis of Cannabis, Linnaeus (1737) discussed
two taxa assigned to South Asian Cannabis by van Rheede tot Drakestein (1690), and
two taxa assigned to East Asian fiber by Kaempfer (1712). Furthermore, Linnaeus
(1737) cited three other authors, Bauhin, Ray, and Morison, who also coined taxa for
psychoactive Asian Cannabis, distinct from European hemp. Linnaeus knew about
those taxa—he cited taxa that Bauhin, Ray, and Morison assigned to European
hemp—but he did not address them. In his Materia Medica, Linnaeus (1749) listed
narcotica and inebrians as some of the attributes of Cannabis, which are adjectives
used by the aforementioned authors who described Asian Cannabis.
Type specimen
Linnaeus’s type specimen of C. sativa is stored in the Linnaean Herbarium (herb.

LINN), specimen no. 1177.2, a lectotype designated by Villiers (1973). As noted by
Jarvis (2007), Villiers holds a one-year priority over Schultes et al. (1974), who
designated a different lectotype, Hortus siccus Cliffortianus (herb. BM) p. 457. Stearn
(1974) dedudced from the numbering system that Linnaeus collected 1177.2 in 1753.
No type location is given, but in 1753 Linnaeus lived in Uppsala.
Lectotype 1177.2 consists of the uppermost 28 cm of a fruiting staminate
plant. Leaves branch alternately from the stalk, palmately compound, light
green, mostly with three leaflets, and with long petioles. Central leaflets are
narrowly lanceolate, acuminate, with sharp serrations, up to 78–85 × 7–9.5 mm.
Inflorescences are loose, not dense; subtending floral leaves have a sparse
covering of capitate-sessile glandular trichomes; perigonal bracts (i.e., bracteole,
calyx) have a relatively sparse covering of capitate-stalked glandular trichomes.
Styles and stigmas are not prominent, pale green in color. The fruit includes a
non-persistent perianth, achene large (4.8 × 2.5 mm), oblong in outline, pale
green with a fine reticulated pattern, base short and pointed with a simple
articulation.
The previously-recognized lectoype, Clifford Herbarium p. 457.1, is illustrated by
Stearn (1974). The flowering top is somewhat denser than 1172.2, with slightly larger
leaves, 3–5 leaflets per leaf, leaflets broadly lanceolate, and with larger achenes
(5.0 × 3.5 mm). The specimen likely dates to Linnaeus’s time in Holland, and
represents “the old cultivated hemp stock of northern Europe” (Stearn, 1974).
Appendix 1b. Cannabis indica Lam.
Lamarck (1785) provided a full description of C. sativa (not just its flower
parts, like Linnaeus). Lamarck then separated C. indica from C. sativa by “very
distinct” morphological and phytochemical differences. We translate from
French: “It is smaller, more ramifications of the stalks, which are tough
[woody] and more cylindrical, and distinguished particularly by alternating
branches. Their leaflets are very narrow, linear-lanceolate, and very acuminate.
The male individuals bear five to seven leaflets, but those that are female may
exhibit as few as three for each petiole, and leaves at the top [of the plant] are
even simple [not compound]. The female flowers have a velous calyx, and long
styles that are alike.”
We interpret “vellous calyx” to mean “velvety perigonal bract,” due to a dense pubes-
cence of capitate-stalked glandular trichomes. Lamarck noted that C. indica does not
provide a good source of fiber, and it produces a strong odor “resembling somewhat that
of tobacco.” He mentioned the intoxicating properties of C. indica when smoked in a pipe.
“The principal effect of this plant consists of going to the head, disrupting the brain, where it
produces a sort of drunkenness that makes one forget ones sorrows, and produces a strong
gaiety.”
Lamarck cited six synonyms and their authors:
1. Cannabi similis exotica Bauhin (Pinax Theatri Botanici 1623: p. 320); Bauhin
cited descriptions of Bangue by da Orta and Acosta, in Goa, India;
2. Kalengi-cansjava Rheede (Hortus Malabaricus 1690; 10: 119); assigned to male
plants in Kochi, India;
3. Tsjeru-cansjava Rheede (Hortus Malabaricus 1690; 10: 121); assigned to male
plants in Kochi, India;
4. Cannabis peregrina gemmis fructuum longioribus Morison (Plantarum historiæ
universalis Oxoniensis 1699; 3:433); Morison cited Bauhin and Rheede;
5. Cannabis indica Rumph (Herbarium Amboinense 1747; 5: 208), a “pre-Linnaean”
(pre-1753) taxon assigned to plants in Indonesia;
6. Dakka ou Bangua Prévost (1768), an editorialized version of Kolb (1719) who
described plants in South Africa.
Four authors in Lamarck’s synonymy (1, 2, 3, 4) previously appeared in Linnaeus

(1737). Lamarck omitted two synonyms cited by Linnaeus: Bangue cannabi Ray 1686,
and Bangue cannabi simile J. Bauhin and Cherler 1651–2. Both Linnaeus and Lamarck
omitted other “pre-Linnean” (pre-1753) authors who differentiated European hemp
from psychoactive Cannabis. Going back in time: Linder (1739), Burman (1737),
Drury (1729), Hooke (1726), Kaempfer (1712), Petiver (1703), Plukenet (1696), Fryer
(1698), Ray (1693), Chardin (1686), Knox (1681), de Flacourt (1661), Parkinson
(1640), Sennert (1629), dos Santos (1609), de Gouvea (1606), Van Linschoten
(1601), Alpini (1591), Acosta (1578), de L’Ecluse (1567), da Orta (1563), Fuchs
(1542), and Bock (1539). The first European to assign separate names to European
and Asian Cannabis was Ibn-al-Baitār, around 1240, in Arabic. He differentiated
qinnab (plants he knew in Spain) from plants he encountered in Egypt, qinnab hindī,
“Indian hemp” (Ibn al-Baitār, 1985).
Type specimen
Lamarck’s type specimen of C. indica is deposited at the Muséum National

d’Histoire Naturelle in Paris (herb. P). It consists of the uppermost 30 cm of a
flowering, seedless pistillate plant. Lamarck’s specimen shows more compact
branching with shorter internodes than Linnaeus’s specimen. Branching density
may reflect differences in cultivation: Lamarck’s specimen was presumably
open-grown, and Linnaeus’s specimen grown in typically dense hemp fields.
However, the effects of sowing density upon branching density are minimal in
the uppermost 30 cm of a flowering top. Leaves branch alternately from the
stalk, palmately compound, medium green, mostly with three leaflets, and short
petioles. Central leaflets are narrowly linear-lanceolate, acuminate, with sharp
serrations, 55–58 × 5–7 mm. Inflorescence are somewhat compact; subtending
floral leaves have an abundant covering of capitate-sessile glandular trichomes;
perigonal bracts express a moderate density of capitate-stalked glandular
trichomes. Styles and stigmas are prominent, agglutinized with trichome

exudate, and light brown. The type specimen is a sample of sinsemilla,
lacking fruits.
No type location is given. Lamarck (1785) wrote, “This plant, which Mr. Sonnerat
gave us pieces he brought back from India…” According to Schultes et al. (1974), “we
are at a loss to indicate a definite area, partly because of vagueness of geographical
terminology in that period.” In fact, we know when and where Pierre Sonnerat made his
collections. He explored India and China between 1774 and 1781. In India he made
extensive collections around Pondicherry (Ly-Tio-Fane, 1976), so Sonnerat likely
collected C. indica near Pondicherry. Unfortunately he did not mention Cannabis in
his travelogue (Sonnerat, 1782).
Discussion
Lamarck erroneously described C. indica with “alternately branching leaves” and

C. sativa with opposite branches (i.e., decussate, the succeeding pairs turned 180°).
In fact, both C. indica and C. sativa do both—opposite branching during vegetative
growth, switching to alternate branching during flowering. Lamarck may have erred
because the specimen he received from Sonnerat was a flowering top, with alternate
branching. Lamarck cited authors who illustrations of Asian Cannabis also depicted
flowering tops, with alternate branching habits (Acosta 1619, Rheede 1690, Rumpf
1747). Interestingly, the same year that Lamarck (1785) described C. indica, his co-
worker André Thouin was growing Chinese hemp. Thouin (1785) described chanvre
de Chine with alternate branching.
Note that we cite 1785 as the year that Lamarck coined C. indica. See Breistroffer
(1948) for evidence that the commonly cited date of 1783 is inapplicable. Each volume
of Encyclopédie Méthodique was printed in two parts; the second part of each was
printed later than the date on the title page. We use the date specified by Stafleu and
Cowan (1979).
Appendix 1c. Cannabis sativa var. spontanea Vav.
Vavilov (1922) collected wild-type germplasm near Saratov, Samara, Astrakhan, and
Tsaritsyn (Volograd). “The study of this hemp in our laboratory by Varvara F.
Antropova compels one to consider it to be, without doubt, wild.” He described a
series of forms that transition from the cultivated variety to the wild-type variety,
“differing in the coloration of the seeds and in the size and form.”
Plants at the wild end of the spectrum expressed anthocyanin in their stalks, and
produced seeds that were dull (not shiny), elongated (not rounded), and smaller than
seed from cultivated plants. Seed color ranged from light or white forms with a mosaic,
to dark gray and light brown with marbling.
Vavilov provided a brief diagnosis, translated from Russian: “At the bases of the
seeds there are formations that resemble ‘horseshoes’ (analogous to wild oats), along
which there occurs a breaking off and a shedding of seeds at maturation. Frequently this
horseshoe is clearly defined and undoubtedly is a morphological and biological feature

clearly distinguishing it from the cultivated form. At maturation, the wild variety with
the horseshoe is easily distinguishable fro the ordinary cultivated variety by the
shedding of the seeds which have not yet reached full maturity. As was mentioned
by Professor D. E. Janischevsky, wild forms of hemp appear primarily as dark-seeded
marbled forms.”
Vavilov (1922) cited no synonyms. He apparently overlooked Czernajew (1859), who

assigned the name C. sativa var. spontanea to wild-type plants near Kharkiv, Ukraine.
Czernajew’s taxon has no standing, because he did not describe the variety, and
therefore did not meet a basic provision of the ICN (Small & Cronquist, 1976). Vavilov
mentioned earlier work on wild hemp by Janischevsky, but did not cite a publication.
Type specimen
Several specimens labeled C. sativa var. spontanea are stored in Vavilov’s herbarium at
WIR. Small and Cronquist (1976) chose one as the lectotype, specimen Antropova 121.
The label states the plant was grown in 1925 at the Kamenno-Stepnaya experiment
station in Voronezh, from seeds collected near Saratov in 1921, by Vavilov’s assistant,
Varvara F. Antropova.
The specimen consists of the top 42 cm of what appears to be a large plant, with
internode spacing similar to Linnaeus’s type specimen. Leaves with 3–5 leaflets,
petioles short, leaflets narrow lanceolate, up to 130 × 10 mm. Inflorescences are loose;
subtending floral leaves have a sparse covering of capitate-sessile glandular trichomes;
perigonal bracts have relatively few capitate-stalked glandular trichomes and many
cystolith trichomes. Achenes are medium-sized (3.8–4.0 mm long), oblong in outline,
pale green with a fine reticulated pattern overlaid by irregular dark marbling, and a
weakly protuberant base.
Discussion
Vavilov (1926) debated whether C. sativa var. spontanea was truly wild, or a spontane-
ous escape of formerly cultivated plants. This question was also raised by Janischevsky
regarding Cannabis ruderalis (see below). Both botanists were unaware that this debate
arose earlier, regarding plants in the same location—Saratov. Three naturalists travelling
together in 1768 encountered wild-type plants about 10 km downriver from Saratov.
Lepechin (1774) said the plants did not differ from cultivated plants. He considered them
feral escapes, sown by former inhabitants. “This Indian plant species can not be native
here.” Nearby lay the ruins of Uvek, a trading center of the Kipchak Khanate, destroyed
by Tīmūr in 1395. Pallas (1793) said the wild-type hemp looked like cultivated plants,
which he attributed to former inhabitants. Falck (1786) wrote, “On the Volga one finds
the wild hemp especially on the sites of former cities.” He noted branchiness in the wild-
type plants, “These give no straight and uniform fibers.”
Vavilov and Janischevsky both decided that plants around Saratov were truly wild.
Small (1975a) throws a variable into the debate. He discovered that domesticated
Cannabis reverted to a wild-type phenotype within 50 generations (years) of prohibi-

tion in Canada. This plasticity makes it difficult to distinguish truly wild plants from
formerly cultivated plants that have reverted to wild-type phenotypes. Doebley et al.
(2006) also emphasized that domesticated C. sativa easily escapes cultivation and
reverts to a wild-type phenotype. Small and Cronquist (1976) suggested that truly
indigenous, never-domesticated, wild-type Cannabis may be extinct. “If unaltered wild
populations exist (which we doubt) they cannot be clearly distinguished from those
contaminated by the influence of domestication.”
Was there any reproductive isolation between wild-type and cultivated plants? Their
geographic distributions overlapped (sympatry). Hybridization and introgression was
limited by temporal (allochronic) isolation—Janischevsky noted that ruderalis matured
in mid-June, while cultivars were still in the vegetative stage. There may have been
some ecological inviability as well—a condition where hybrid offspring are normal but
suffer lower viability because they cannot find an appropriate ecological niche. How-
ever, Vavilov and Janischevsky noted intermediate forms, which suggested some
hybridization occurred between the populations.
Appendix 1d. Cannabis ruderalis Janisch.
Janischevsky (1924) coined the species name C. ruderalis, but added parenthetically an
alternative taxonomic rank: C. sativa var. ruderalis. “I am inclined to consider it a well
marked variety.” We provide key details of his large protologue—a 13 page article.
Translated from Latin: “Fruits rather small, rather hard, narrowly ovate, base attenuate,
perianth marbled-spotted, persistent and enclosing [the fruit] at maturity, [the fruit]
quickly dropping off [the plant].”
In addition to the Latin diagnosis, Janischevsky provided additional characters: Male
flower produced shorter perianths and anthers than those in cultivated plants. Average
seed size was 3.5 mm long × 2.5 mm wide, and rarely reached 4.5 × 3.0 mm. The
female flower’s perianth was persistent and stuck to the surface of the seed—not unlike
hops (Humulus lupulus). The seed had an elongated base in the form of a short pedicle.
Within the pedicle Janischevsky detected cells with oily inclusions—so he considered
the structure an elaiosome—an oil-rich body that attracts insects, which acts as a seed
dispersal agent.
Vegetative characters included relatively short height, usually 0.7–1.1 m but up to
2.1 m, with a strongly branched habitus. Stalks were usually ribbed, with thick xylem at
the base (i.e., woody), and with the upper stalk colored by anthocyanin. Leaves had 5–7
leaflets, colored deep green on the top side, and gray-white on the underside due to
cystolith trichomes.
Janischevsky conducted common garden experiments, which showed that wild-type
plants matured more rapidly than cultivars. They came into flower mid-June, and first
seeds were collected July 10th (while cultivars were still in vegetative stage). Wild-type
plants compared to cultivars showed greater drought tolerance, and more shade toler-
ance in forested areas. Wild-type seeds readily disarticulated from plants. The seeds
demonstrated a prolonged period of dormancy and slow germination, whereas seeds of

cultivated hemp had no period of dormancy.
Janischevsky described mutualism between wild hemp and an insect, Pyrrhocoris
apterus. The insect sucked oil out of the elaiosome. The rest of the seed was left intact
and capable of germination. In the process of feeding, the bug carried the seed “far
distances,” and facilitated the spread of wild hemp. The following year, Janischevsky
(1925) wrote a whole article about wild hemp and P. apterus.
Janischevsky did not list any explicit synonyms under C. ruderalis. He did not mention
C. sativa var. spontanea Czernajew (1859), or C. sativa var. spontanea Vavilov (1922).
He noted that taxa other than C. sativa were coined by other Russian botanists, such as
Messerschmidt (in Amman, 1739), Sievers (1796), Ledebour (1847,–49), and
Korshinskii (1898). Yet others suggested that C. sativa, of Asiatic origin, may also
be indigenous to Europe—but they expressed this opinion with great caution.
Janischevsky took five pages to recount observations by 22 authors: Erndtel, 1730,
Messerschmidt (in Amman, 1739), Sievers, 1796, Rochel, 1835, Claus, 1838,
Ledebour (1847–49), Christison, 1850, de Candolle, 1855, Becker, 1858, Heuffel,
1858, Dukerley 1866, de Candolle 1869, Kanitz, 1881, Zinger, 1885, Shmal’gauzen,
1897, Korshinskii, 1898, Velenovský 1898, Prain, 1904, Adamović, 1907, Bogdan,
1908, Ascherson & Graebner, 1911, and Vavilov 1922.
Type specimen and illustrations
Janischevsky’s lectotype specimen of C. sativa var. ruderalis, collected near Saratov,

was stored at the Komarov herbarium (LE) in St. Petersburg (Small & Cronquist 1976).
It may be lost (McP., pers. observ., St. Petersburg, 2010), and no suitable isolectotype
was identified at LE. Small (1975b) provided a photograph of the lectotype.
Janischevsky provided two plates of line drawings, reproduced here as third-
generation photocopies (Fig. 6, figure legend translated from Russian).
Fig. 6Line drawings of C. ruderalis by Janischevsky (1924))
Fig. 6 Line drawings of C. ruderalis by Janischevsky (1924)

Рис. 1. A fruit of hemp collected in the vicinity of Saratov (Dudakov ravine).

Рис. 2. Two diagrammatic drawings of the longitudinal section of the fruits. Рис. 2a.
A section in the plane that goes through the edge of the fruit. Рис. 2b. A section in the
plane that goes between the edges of the fruit. I- perianth; II- pericarp, e- epicarp, E-
endocarp, 5- point of insertion of perianth, 6-receptical transformed into elaiosome
which juts out into the space among the carpels (see diagram B) in the form of rounded
mases of tissue, v- vascular bundle.
Рис. 3. A bug Pyrrhocoris apterus on a Carex leaf carrying a hemp fruit.
Discussion
Janischevsky (1924) said he found hemp growing in “wild circumstances” in 1897,

near Buzuluk (Orenburg Oblast). It flourished along the Tok River, a right tributary of
the Samara, in Bashkir tribal land where hemp was not cultivated. Janischevsky added
that he began growing specimens of wild hemp in the botanical garden at Saratov
University in 1914. Vavilov arrived at Saratov University in 1917, joining his senior
colleague. Janischevsky and Vavilov took field trips together down the Volga
(Korotkova, 1978). It seems likely that Janischevsky pointed Vavilov’s attention to
wild hemp. In Vavilov’s archives, the subject of wild hemp first appeared in a 1920
letter to his assistant Evdokia A. Teplykh, “Collect wild or feral hemp on the borders
and the ravines near Saratov from selected plants, at least 500 plants, in separate bags.
Keep in mind that wild hemp seed falls off easily, so it must be carefully collected”
(Korotkova, 1978).
Vavilov (1926) elaborated upon C. sativa var. spontanea. His review of the literature
borrowed from Janischevsky (1924) in a few plagiarized sentences. Vavilov cited six of
Janischevsky’s 22 references (Erndtel, 1730, Ledebour, 1847–49, Korshinskii, 1898,
Velenovský, 1898, Adamović, 1907, Bogdan, 1908). However, Vavilov added infor-
mation about wild-type hemp from Mak & Regel, 1862, Boissier 1879, Hooker, 1890,
Britton 1899, Dimo & Keller, 1907, Semenov (in Krylov, 1909), Knorring and
Minkvitz Knorring & Minkvits, 1912, Vysotsky, 1915, Oganovsky, 1922, Sinskaya
1925, and Khrebtov, 1925.
Vavilov (1926) details work on wild-type hemp by his assistant Tatiana Ya.
Serebriakova. She compared germplasm collected by Antropova in Saratov with
germplasm collected by Eugeniya N. Sinskaya in the Altai (Sinskaya, 1925). Plants
from both accessions grew 60–150 cm tall, with moderately thick stalks that were
strongly branched. Leaves were medium sized, with 5–9 narrow leaflets. Seeds were
small (2.7–3.0 mm long), dark, orbicular-elongated in shape, with horseshoes and a
persistent, marbled perianth. Plants matured before cultivated plants, seeds rapidly
dehisced from plants, and seed germination was usually slow and unequal.
Vavilov (1931) extended the range of C. sativa var. spontanea to Central Asia.
While traversing the Tiān Shān to Lake Issyk Kul via the Bedel Pass, he wrote,
“Cannabis sativa var. spontanea is a very common plant in northern Tiān Shān,
especially on its north-facing slopes and valley.” Herbarium specimens collected on
the north-facing slopes of the Tiān Shān look very different than Vavilov’s type
specimen (shorter, with more branches, and broad oblanceolate leaflets). Zhukovsky
(1962) proposed that C. ruderalis had not been sufficiently explored in Central Asia,
and some populations may be undescribed forms.
Lastly we come to the question of priority. Vavilov (1922) scooped his senior
collegue by coining C. sativa var. spontanea. Janischevsky (1924) erected another
Latin name for the same wild-type hemp growing near Saratov. The taxonomic name
became a point of contention between the two authors: Janischevsky politely men-
tioned Vavilov’s research, but always used the epithet ruderalis (Janischevsky, 1924,
Janischevsky, 1925). Vavilov politely mentioned Janischevsky’s work, but always used
spontanea (Vavilov, 1926, 1931; Vavilov & Bukinich, 1929).
Two taxonomic names that represent contaxic entities, based on different type
specimens, are facultative synonyms. Vavilov’s earlier epithet has priority at the rank
of variety. Janischevsky’s taxon has priority at the rank of species.
Since then, botanists have altered the ranks of spontanea and ruderalis.
Serebriakova and Sizov (1940) elevated spontanea from a variety to a subspecies.
The subspecies rank has been adopted by several authors (e.g., Bócsa and Karus Bòcsa
& Karus, 1997; Hanelt, 2001; McPartland & Guy, 2004, Clarke & Merlin, 2013). Chu
(1959) reduced ruderalis to a sub-varietal form, as C. sativa f. ruderalis. Liou (1988)
brought Chu’s taxon back to a variety, C. sativa var. ruderalis. Liou mistakenly
believed he coined a var. novo. Soják (1960) used Janischevsky’s taxon to describe
hybrids between wild hemp and cultivated hemp, which he named X C. intersita. He
later changed the taxon to C. sativa subsp. intersita (Soják, 1980).
A troika of influential Soviet texts set precedence by choosing C. ruderalis over
C. sativa var. spontanea: Nekrasova (1934), Yarmolenko (1936), and Mal’tsev (1939).
Soviet science at that time writhed under the rise of T. D. Lysenko. Lysenko became the
pet scientist of Joseph Stalin. He fabricated genetic theories using the rhetoric of Marx
and Michurin, and methodically annihilated his academic opponents. Lysenko labeled
Vavilov a Trotskyite, which led to Vavilov’s arrest (Popovsky, 1984). After Vavilov was
arrested, his assistant Tatiana Serebriakova coauthored her final Cannabis paper with
Ivan A. Sizov. He was a Lysenkoite who “began energetically to liquidate the remnants
of Vavilov traditions” (Medvedev, 1969). Serebriakova and Sizov (1940) elevated
Vavilov’s taxon from a variety to a subspecies, but without his name in the basionym:
C. sativa subsp. spontanea Serebriakova. C. ruderalis was synonymized under that
taxon.
Schultes et al. (1974) treated C. ruderalis as a species separate from C. sativa. They
expanded Janischevsky’s taxon to Central Asia, and erroneously typified it with a
specimen from Tajikistan. Furthermore they described ruderalis as a very short,
unbranched plant with broad leaflets. This departs from the concepts of Vavilov and
Janischevsky. Schultes et al. (1974) cited Schultes, and described ruderalis as a short
plant. Anderson (1980) cited Schultes, and described ruderalis as a short plant with
broad, oblanceolate leaflets.
Small and Cronquist (1976) kept Vavilov’s taxon at the rank of variety. They
recombined it as C. sativa ssp. sativa var. spontanea, typified by a Vavilov specimen
from Saratov. They synonymized C. ruderalis under that taxon. They extended the
range of spontanea to wild plants from Central Asia.
Schoenmakers (1986) introduced “Ruderalis” to underground Cannabis
breeders. He collected germplasm near the Hungary-Ukraine border. His photos
of “roadside ruderalis” illustrated an escape from cultivation, and not a wild-
type plant. “The effect after smoking was stoney, but not high.” Other Canna-
bis breeders assigned the name “ruderalis” to feral plants in North
America—obviously escapes from cultivation, and not wild, indigenous plants

(e.g., Panik, 2012. Frank (1988) stated that potent, psychoactive “Ruderalis”
invariably turned out to be misidentified “Indica.” This confusion persists: a
2013 web video about “Charlotte’s Web” by CNN medical correspondent
Sanjay Gupta portrayed “Ruderalis” plants that clearly have Afghanistan
heritage.
Appendix 1e. Cannabis sativa f. afghanica Vav.
Vavilov encountered wild and weedy plants in Afghanistan. First he described

C. sativa f. afghanica (Vavilov, 1926). His description is transcribed below,
where we combine the best of two English translations (Vavilov, 1926, 1992):
“We observed wild hemp… in districts where hemp cultivation is entirely

unknown. Belts of black hemp follow the sowings of corn and other cereals
along the Kunar River (on the border between Afghanistan and India [now
Pakistan]) from Chekosarai to Jalālābād, a distance of 160–200 km. In between
Chekosarai and Jalālābād we discovered among the wild hemp a peculiar race
with light-coloured, small seeds, and with a thin perianth easily taken away (f.
afganica Vav.). The seeds of this race were very small (1000 seeds weighed
2.1–2.7 g), ten times smaller than the large-fruited races of the Far East (1000
seeds 26.0 g), the common Central Russian races (Orel, Kurst) show a weight
of 17–19 g.
“The wild hemp collected on the Kunar River approached the cultivated type
with respect to seed color and the slightly splitting pericarp, but is distin-
guished by readily shedding its seeds and in the development of “horse-shoes.”
When sown, the seeds germinate very slowly and unequally, i.e., the plants
show features of a typical wild plant. The wild hemp displayed other new
characteristics. The leaflets of these plants were distinguished their obovate
narrow shape, not observed by us among the European, Siberian and Turkestani
forms… The wild Afghani races found along the frontier with Pakistan, with
light-colored and easily splitting pericarps, have spread into Pakistan as well.
The plants constitute a morphological link between the wild and the cultivated
races of hemp with respect to the most important differentiating characteristics.”
In an accompanying table, Vavilov provided results of a common-garden
experiment of C. sativa f. afghanica by his assistant Serebriakova. Plants
matured late in the season, and seed germination was slow and unequal. Plants
grew 60–150 cm tall, with moderately thick stalks, and strongly branched.
Leaves were small, with 5–9 leaflets, and narrowly obovate in shape. Seed
was described in an either-or duality: Seed size was either small (2.7–3.0 mm)
or medium (3.0–4.0 mm), seed color was either dark or light, seed shape was
either orbicular-elongate or orbicular, the seed perianth was either persistant or
easily removed, the seed base either with or without a horseshoe. Vavilov
seemed to be describing two kinds of plants. He wrote, “the plants show
features of a typical wild plant,” followed by “the plants constitute a morpho-

logical link between the wild and the cultivated races of hemp.”
Vavilov provided no synonymy and cited no references.
Type specimen
No specimen labeled afghanica is deposited in Vavilov’s herbarium at WIR. Serebriakova

grew germplasm of Afghan provenance at the Kamenno-Stepnaya Experiment Station
(Voronezh Oblast); these herbarium specimens are labelled C. sativa var. spontanea, and
annotated “like ruderal” (WIR specimens 4031, 4032, 4034, 4036, 4038, 4044, 4046). All
are immature, without seeds, and could be either afghanica or kafirstanica.
Discussion
The either-or duality in Vavilov (1926) got sorted 3 years later, when Vavilov and
Bukinich (1929) named a second taxon of Afghan plants: C. indica var. kafirstanica.
The either-or descriptions in Vavilov (1926) referred to either afghanica or
kafirstanica. Vavilov and Bukinich described kafirstanica as truly wild, with tiny
achenes that were dark-colored and marbled, with an elaiosome. They described and
illustrated C. indica f. afghanica, with larger achenes that were light colored (“white”),
and a colorless involcre (perianth?), with little or no “horseshoe” (Fig. 7).
Fig. 7 Seeds illustrated by Vavilov and Bukinich (1929). Original caption: Left to right: 1. from northern
Afghanistan—cultivated form—Cannabis sativa L.; 2. ordinary Russia hemp from Orel; 3. wild hemp from
Saratov; 4., Cannabis indica f. kafiristanica Vav.; 5. Cannabis indica var. afghanica Vav. The upper row
enlarged 6 times, the lower row showing the bases of achenes enlarged 10 times
Fig. 7 Seeds illustrated by Vavilov and Bukinich (1929). Original caption: Left to right: 1. from northern
Afghanistan—cultivated form—Cannabis sativa L.; 2. ordinary Russia hemp from Orel; 3. wild hemp from
Saratov; 4., Cannabis indica f. kafiristanica Vav.; 5. Cannabis indica var. afghanica Vav. The upper row
enlarged 6 times, the lower row showing the bases of achenes enlarged 10 times
Vavilov and Bukinich considered afghanica a “specialized form” of kafirstanica,

“transitional” between wild kafiristanica and cultivated C. indica. Their nomenclature
was not consistent—they used C. indica var. afghanica on page 380 (see caption in
Fig. 7), f. afghanica on page 381, and C. indica var. kafirstanica f. afghanica on page
382. Small and Cronquist (1976) argue that the caption (Fig. 7) erroneously reversed
the ranks of two taxa, Cannabis indica f. kafiristanica and C. indica var. afghanica, so
they are typographical errors.
We propose that Vavilov erred by describing afghanica as “transitional” between wild
and cultivated plants. The afghanica was probably domesticated—a feral escape, or
recently naturalized. Its seed lacked a “horseshoe” (elongated base in the form of a short
pedicle), and lacked a persistent perianth. It resembled a small version of the Afghani
“cultivated form” (Fig. 7). The size of the cultivated seed would have been augmented by
irrigation and fertilizer. Vavilov and Bukinich described afghanica growing in “aban-
doned lots and neglected plots of land, in fertilized agricultural soil, and between
agricultural fields.” Arable land in Afghanistan is rarely abandoned and neglected.
Emboden (1974) assigned a varietal rank to the taxon, as C. indica var. afghanica. He
gave seed size as 2.8 × 1.9 mm, without stating the provenance of his material. Small and
Cronquist (1976) synonymized C. sativa f. afghanica Vav. under Lamark’s taxon—a
domesticated variety. Vavilov’s taxon at the rank of species, as C. afghanica, was
employed by McPartland (1996), McPartland et al. (2000), and Clarke and Watson
(2002). Clarke (1998) used a varietal rank, C. indica var. afghanica. The taxon
C. sativa var. afghan appeared in a U.S. patent (No. 6,403,530). Cervantes (2006) offered
two ranks, C. afghanica and C. sativa var. afghanica. A subspecies rank, C. indica subsp.
afghanica, was employed by McPartland and Hillig (2004) and Clarke and Merlin (2013).
Recently the species rank, C. afghanica, has been applied by others (Tennstedt & Saint,
2009, Brownjohn & Ashton, 2012, Macedo et al., 2013, Henry, 2015, Laursen, 2015).
Appendix 1 f. Cannabis indica var. kafiristanica Vav.
The description by Vavilov and Bukinich (1929) is translated from Russian: “Races of
wild hemp in eastern Afghanistan have extremely small fruits with mosaic (1000 fruits
weigh 2.1-2.7 g), i.e., 6-8 times smaller than small-seeded Central Russian cultivated
hemp (Orel and Kursk hemp weighs 17-19 g). Characteristic for them is ready
shattering of fruits due to the presence of a horseshoe, slow and uneven germination,
i.e., the usual attributes of a wild plant. As regards to vegetative features, Afghan wild-
weedy hemp is distinguished by small leaves with obovate leaflets of narrowed shape.
In general, it is characterized by short stature, profuse branching from the first
internode, and by short internodes.”
Vavilov and Bukinich also mention its early ripening (90–100 days in Voronezh
Oblast). They provided a drawing of the seed (Fig. 8). Although Vavilov and Bukinich
described kafiristanica plants as short in stature, a photograph of plants identified as
C. indica var. kafirstanica (Fig. 267 in Vavilov and Bukinich) shows plants that were
equal in height to maize plants with tassels.
Fig. 8Type specimen of C. sativa var. kafiristanica Vav. (McPartland photo, taken at herb. WIR)
Fig. 8 Type specimen of C. sativa var. kafiristanica Vav. (McPartland photo, taken at herb. WIR)
Vavilov provided no synonymy and cited no references.
Type specimen
Several specimens labeled C. sativa var. kafiristanica are deposited in Vavilov’s

herbarium at WIR in St. Petersburg. Small and Cronquist (1976) chose one as the
lectotype: WIR 3952, germplasm collected by Vavilov at Chekhosarai (now Asadābād)
in 1924 and cultivated in 1927 at Pushkin Experiment Station (Detskoye Selo, St.
Petersburg). A photograph of the lectotype appeared in Vavilov and Bukinich (1929),
and appears here (Fig. 8). The entire plant is mounted on the herbarium sheet, about
30 cm tall. It is an immature pistillate plant, with tight internode spacing; nine pairs of
opposite branches and only three alternate branches near the apex. Leaves with 5–7
overlapping leaflets, petioles long and thick, leaflets broad, oblanceolate, dark green,
with coarse serrations, up to 46 × 18 mm. The inflorescences are immature but
nevertheless compact and leafy, with an abundant covering of capitate-sessile glandular
trichomes.
Discussion
Vavilov said afghanica and kafirstanica grew in the Kunar River valley—the border
between Afghanistan and Pakistan—between Chighasaray (now Asadābād) and
Jalālābād. We note a geographical contradiction: The Kunar River valley is not part
of Kāfiristān (present-day Nuristān Province), it lies in Kunar Province, populated by
Pushtan Afghanis, not ethnic Kāfirs. The place where Vavilov collected kafirstanica
was not in Kāfiristān.
Serebriakova and Sizov (1940) made no mention of afghanica and kafirstanica; they
placed drug plants from Afghanistan in the taxon C. sativa subsp. culta prol. Asiatica
var. narcotica. Emboden (1974) gave seed size of C. indica var. kafiristanica as
3.0 × 2.2 mm (larger than var. afghanica), without stating the provenance of his material.
Small and Cronquist (1976) recombined Vavilov’s taxon as C. sativa subsp. indica
var. kafiristanica. They considered it the wild-type associate of domesticated C. sativa
subsp. indica var. indica. As such, they expanded the taxon’s concept beyond Afghan-
istan to wild-type plants from India, Nepal, China, South Africa, and Colombia. Hillig
and Mahlberg (2004) also extended kafiristanica beyond Afghanistan to feral popula-
tions from India and Nepal. The rank of kafiristanica has bounced around a bit, like that
of afghanica. Chrtek (1981) elevated it to a species rank, C. kafiristanica. A subspecies
rank, C. indica subsp. kafiristanica, was employed by McPartland and Guy (2004) and
Clarke and Merlin (2013).
Appendix 2: Taxonomic bias and personality cults surrounding Linnaeus and

Lamarck in the 18th–19th centuries
Carl Linnaeus (1707–1778) is the most famous naturalist of all time. “Linnaeus
became the subject of hero worship after his death to an extent previously unknown
in botany” (Stafleu, 1971a). Linnaeus’s impact on biological thought and his renown
among peers is difficult to convey today (Blunt, 2001). Linnaeus’s four-point “revolu-
tion” developed in stages: the sexual system (Linnaeus, 1735), hierarchical taxonomic
ranks (Linnaeus, 1735), generic reform as a set of theoretical axioms (Linnaeus, 1737a,
and their practical application Linnaeus (1737b), and lastly binomial nomenclature
(Linnaeus, 1753).
Linnaeus’s genius was soon recognized. He became a Full Professor at age 34. The
King of Sweden knighted him at age 46. A generation of devoted students trained
under Linnaeus, and he served as supervisor (“praeses”) for 186 Uppsala doctorates.
Seventeen of his most committed scholars became known as the “apostles.” Thanks to
Linnaeus’s connections with the Swedish East India Company, his apostles explored
the world and spread Linnaean taxonomy. Seven apostles died on expedition. Five
wrote about Cannabis (Hasselquist, Forsskål, Falk, Sparrman, Thunberg).
Linnaeus’s innovations competed with other taxonomic systems at that time, by Ray,
Morison, Rivinus, Tournefort, Herman, Boerhaave, Ludwig, and Magnol. The Linnae-
an system was embraced quickly in Holland (where he published 14 of his early
works), and of course Sweden. Next came Great Britain. English literature soon
dominated botany—particularly concerning Cannabis in India—so the reasons why
British botanists became the staunchest supporters of Linnaeus are worth reviewing:
Linnaeus visited England in 1736, and impressed four leading botanists—Hans
Sloane, Philip Miller, Peter Collinson, and John Dillenius. Dillenius (1741) introduced
Linnaeus’s work to English literature. Collinson organized the election of Linnaeus as a
Fellow of the Royal Society in 1754. He asked Linnaeus to send an apostle to London;
Linnaeus sent Daniel Solander, who gained a post at the British Museum. Solander
soon proselytized Joseph Banks, and they botanized together with James Cook on the
Endeavour.
Stillingfleet (1759) translated into English six essays from Linnaeus’s “unrivalled
school of natural history.” Coxe (1811) referred to Stillingfleet as “one of the body-
guards of Linnaeus.” Stillingfleet encouraged Hudson (1762) to organize Flora Anglica
along Linnaean lines, which “marks the establishment of Linnean principles of botany
in England” (Smith, 1824). Philip Miller, the influential author of Gardeners Dictio-
nary and keeper of the Chelsea Physic Garden, announced that he “adopted in a great
measure the system of Linnaeus” (Miller, 1768). James Edward Smith founded the
Linnaean Society of London in 1788. Four years earlier, Smith had purchased
Linnaeus’s herbarium and library from his widow. Smith (1821) published a tome of
Linnaeus’s correspondence, and cemented London as the center of Linnaean studies.
The anti-Linnaeans
Stafleu (1971a) states that Linnaeus’s authoritative pragmatism appealed strongly to the
practical and bourgeois British, whereas French philosopher-naturalists, under sway of
the Enlightenment, considered Linnaeus an authoritarian of the past.
Comte de Buffon (1707–1788) was a French naturalist and a proponent of the
Enlightenment. Buffon became Linnaeus’s primary scientific rival. He criticized three
aspects of Linnaeus’s four-point revolution. Buffon mocked Linnaeus’s sexual system
for being artificial. He sharply criticized Linnaeus for overthrowing Tournefort’s
generic reform based on arbitrary principles.
Linnaeus’s hierarchical system of taxonomic ranks particularly rankled compte de

Buffon (1749). He was under the influence of John Locke, an Enlightenment author,
who argued against essentialism in favor of nominalism. Linnaeus’s taxonomic system
was essentialist to its core—he treated species as typological and immutable entities
created by God. Buffon argued that we name a creature by its collection of observed
qualities (nominalism), rather than classify a creature by its similarity to an eternal
essence or prototype. Buffon repeated the argument in Locke (1690) that individuals
can be observed, but “species” and “genera” are unobservable abstractions, unless they
consist of a single individual.
Buffon clashed with Linnaeus over “species concepts”—an argument that continues
today regarding Cannabis. Linnaeus saw sharp-cut delineations between species.
Buffon argued that “Nature progresses by unknown gradations,” and organisms were
“not tied up in neat and orderly parcels.” compte de Buffon (1749) asserted, “In Nature
there actually exist only individuals; genera and orders and classes exist only in our
imaginations.” Subsequently, compte de Buffon (1753) recognized species, as a group
that could interbreed, “a constant succession of similar individuals that can reproduce
together.” This concept makes him the founder of the biological species concept,
usually attributed to Mayr (1942).
Jean-Baptiste Lamarck (1744–1829) was a protégé of Buffon. Lamarck’s
supporters hailed him as “the French Linnaeus.” He deviated from Linnaean
orthodoxy in many ways. Lamarck (1778) published a flora that departed from
Linnaeus’s approach in two ways—it was written in vernacular French instead
of Latin, and it used dichotomous keys for plant identification. Subsequently,
Lamarck (1783) launched a new format for writing flora texts. He devised the
new format as an alternative to the telegraphic Linnaean style. The format
remains in use today (Frodin, 2001).
Lamarck criticized three aspects of Linnaeus’s four-point revolution. Lamarck
(1783) critiqued Linnaeus’s systéme sexuel as defective. Lamarck (1788) presented a
lengthy critique of Linnaeus’s generic reform, “the so-called axioms and extremely
laconic maxims with which he filled his Philosophia botanica and Critica botanica,
rather than by solid proof which alone could convince those who were not impressed
by mere authority.” Lamarck (1778) repeated Buffon’s nominalist stance that all
hierarchical ranks above species were subjective creations of the human mind. He
argued that the local European flora was too riddled with species gaps to enable a valid
perception of higher ranks. Lamarck (1791) again asserted that higher ranks (Class,
Family, Genera) were surely not the work of Nature, but artificial, the result of arbitrary
human judgment. Lamarck (1792) enlarged upon this critique as a response to
pushback from Linnaeans.
Lamarck’s theory of evolution totally alienated Linnaeans. Lamarck (1809)
recognized evolution as a gradual process. This made it difficult to find well-
established, concrete lines of separation between taxonomic groups. In contrast,
Linnaeus treated species as entities with fixed forms given to them by God.
“Every genus is natural, created as such in the beginning, hence not to be rashly
split up or stuck together by whim or according to anyone’s theory” (Linnaeus,
1735). “We count as many species as there were forms created in the beginning”
(Linnaeus, 1751). This dispute continues to echo in the modern quarrel between
creationists and evolutionists.
The anti-Lamarckians
Despite Lamarck’s departures from Linnaean orthodoxy, he praised Linnaeus as the

greatest botanist in history. Late in his career he even regarded Linnaeus’s artificial
classification system as a useful device for identifying plants (Lamarck & Mirbel,
1803). Nevertheless, Lamarck’s criticisms “were neither forgotten nor forgiven” by
other botanists (Williams, 2001). “Those who had enjoyed Buffon’s support and
patronage, and those who accepted his model of natural history, underwent several
institutional setbacks and bore the brunt of a massive theoretical offensive” (Corsi,
1988).
In Lamarck among the Anglos, Hull (1984) wrote about Lamarck’s critics. “Few
British or American scientists got their views of Lamarck by reading translations of
Lamarck, let alone Lamarck’s original works in French. In most instances, Lamarck’s
reputation was formed by descriptions of his views in secondary sources.” Primary
among those secondary sources was Georges Cuvier, an opponent of Buffon. Cuvier
(1836) wrote a biting, ironic eulogy of Lamarck that was translated into English.
Carl Ludwig Willendow, a disciple of Linnaeus, updated Species Plantarum after
Linnaeus died. Willendow did his best to dismantle Lamarckian taxonomy and no-
menclature. A perusal of volume one (in two parts) reveals that Willdenow accepted 55
of Lamarck’s taxa, but rejected 61 as synonyms. Willdenow synonymized Lamarck’s
taxa for the flimsiest reasons. Sometimes he just annotated a question mark after them
(e.g., Lobelia nummularia). Willdenow reduced Coffea mauritiana to a variety of
Coffea arabica, “it seems to me hardly different.” He rejected several Eriocaulon
species because they weren’t illustrated. He rejected Fagara heterophylla because “it
seems kind of mixed.” He rejected Jasione perennis because it was “an imperfectly
dried specimen.”
Willdenow (1805) rejected Lamarck’s C. indica on one morphological
character—Lamarck described C. indica with alternate branching. Because C. sativa
also showed alternate branching, Willdenow argued no differences existed between
them. He ignored Lamarck’s other morphological and phytochemical differences.
Willdenow found one distinction in C. indica, “leaflets have a more tapered base,
which is known to be very inconsistent.” Willdenow incorporated C. indica into
C. sativa, rather than name it a separate variety.
Willdenow’s six-volume Species Plantarum was the most influential botanical work
of its day, although it came under “severe criticisms for many manifest errors” (Long,
1843). Wight and Walker-Arnott (1834) initiated the process of “clearing up many
doubtful synonyms in Willdenow,” although they neglected C. indica. Botanists have
reinstated nearly half of Willdenow’s 61 rejected species names (as well as two genera,
Canthium and Nuxia), many reinstated as basionyms.
British botanists in India displayed a strong Linnaean bias. Their opinions were
shaped by Johann König, a pupil of Linnaeus. König organized a botanical society in
India, the “United Brotherhood,” which included two influential British botanists,
Fleming and Roxburgh. Fleming (1810) wrote, “Dela Marck [de Lamarck] is of the
opinion that the Indian Gánja is a different species of Cannabis from the Cannabis
sativa. But Willdenow assures us that…he could not perceive any difference between
them.” Roxburgh (1832) confirmed, “I perfectly agree with Willdenow in thinking all
the varieties, if even such they can be called, centre in one species.” In Flora Indica,
Hooker and Thomson (1855) wrote about “bhang and chirris” obtained from Cannabis
sativa. “With regard to nomenclature, we shall not alter names established by Linnae-
us.” They degregate “hair-splitters,” and criticize Lamarck’s theory of evolution.
Lamarck’s phytochemical character—C. indica caused intoxication—was dismissed
by British botanists. Linnaeans adhered to Linnaeus (1751), who rejected phytochem-
ical characters, such as fragrance and taste. Lacking knowledge of Mendelian inheri-
tance, most Linnaeans believed that phytochemical traits depended entirely upon the
environment—either climate or cultivation. Fiber-type plants “degenerated” into drug-
type plants, and vice-versa, depending on how and where they were grown.
O’Shaughnessy (1838–1840) stated, “difference of climate seems to me more than
sufficient to account for the absence of the resinous secretion, and consequent want of
narcotic power… the Cannabis sativa and Indica are identical.” A cultivation argument
was first put forward by Fleming (1801). He believed phenotypic variation was due to
crop spacing—plants in Bengal were grown very far apart, often nine or ten feet,
compared to closely-sown European hemp fields. Royle (1840) attributed differences
between Indian hemp and European hemp to “openness of planting.” The Indian
method of growing plants 9–10 ft apart caused a loss in fiber softness and flexibility,
and gave rise to “a full secretion of the principles.”
Appendix 3: Details regarding alphanumerical sites in Fig. 2
Figure 2 illustrates the distribution of plants that field botanists in the 18th–19th
centuries identified as C. sativa, C. indica, or other Cannabis species. Many more
naturalists wrote about “hemp,” but did not assign a Latin name to the plants.
Our narrative is divided in two parts: Part A. Plants assigned the name Cannabis
sativa and segregates, and Part B. Plants assigned the name Cannabis indica and
segregates. Within each part, the narrative is organized by floristic regions.
The map in Fig. 2 shows boundaries of ten floristic regions by Good (1964) and
Takhtajan (1986): ❶ Euro-Siberian region, west; ❷ Euro-Siberian region, east; ❸
Mediterranean region; ❹ West and Central Asia (Irano-Turanian); ❺ Sino-Japanese
(East Asiatic); ❻ African-Indian desert (Saharo-Arabian); ❼ Sudano-Zambezian-
Sindhia; ❽ South Asia (Indian); ❾ Continental Southeast Asia (Indochina). ❿
Malayian (Malesian).
Part A . Cannabis sativa and segregates
❶ Euro-Siberian region, west:

Western Europe was blanketed by botanists who wrote scores of local floras. Many
mentioned C. sativa growing wild, ruderal, or spontaneous. For this region, we present
a geographic sample, rather than an exhaustive list. Early mentions include England
(s1, Hill, 1760), France and Switzerland (s2, Lamarck, 1785), Germany (s3,
Willdenow, 1787), Bohemia (s4, Presl & Presel, 1819), Poland (s5, Wulff, 1765),
Hungary (s6, Heuffel, 1858), Galicia (s7, Zawadzki, 1835), Lithuania and Volhynia
(s8, Eichwald, 1830), and the Moscow region (s9, Stephan, 1792).
Catherine the Great commissioned six physician-botanists to explore her new
dominions—Pallas, Lepechin, Falck, Georgi, Gmelin, and Güldenstädt. Three of them
studied under Linnaeaus (Lepechin, Falck, Georgi); Gmelin corresponded with him;
Güldenstädt had no connection with him; and Pallas was anti-Linnaean—Pallas (1766)
refuted Linnaeus’s scala naturae, and Pallas (1774) laid out an animal classification
meant to compete with Linnaeus’s Systema Naturae. Güldenstädt and Pallas departed
from Linnaean orthodoxy by coining new Cannabis species.
Pallas, Falck, and Lepechin wrote about wilder hanf near Chetverosvyatsky mon-
astery on the Volga, about 10 km downriver from Saratov (s10). A stone’s throw from
the monastery laid the ruins of Uvek, a trading center of the Tartars (i.e., Kipchak
Khanate) until 1395. Pallas (1793) said wilder hanf looked like cultivated plants, and he
attributed its presence to former inhabitants—the Tartars. Lepechin (1774) said the
plants were feral escapes of crops sown by the Tartars. Falck (1786) assigned the plants
to C. sativa, but noted a wild-type characteristic, “…only branched stems. These give
no straight and uniform fibers.”
Falck (1786) assigned C. sativa to feral plants along the Terek River in Northern
Caucasus (s11). Georgi (1800) wrote of C. sativa, “We have indigenous or self-growing
hemp in the Taurus mountains (s11), on the Terek (s11), along the Don, and the Dnieper
in Novorossiya (s12).” Güldenstädt (1791) encountered wilder hanf in Ukraine and
referred to it as “Cannabis” or a new name, Cannabis vulgaris (×1).
Russian exploration continued into the nineteenth century. Marschall von
Bieberstein (1808) wrote a flora of the Caucasus and Crimea, and listed ruderal
C. sativa, without naming a specific location (s13). Meyer (1831) reported C. sativa
growing wild around villages below Mount Elbrus (s11). von Besser (1822) said
C. sativa grew spontaneously near Kremenets in northwest Ukraine (s14).
Botanists started using variety names: Czernajew (1859) named wild hemp C. sativa
var. spontanea, near Kharkiv in northeast Ukraine (s15). Lindemann (1881) applied
Czernajew’s taxon to wild hemp growing near Kherson (s12). Korshinskii (1898)
applied C. sativa var. vulgaris to wild-type plants in southern Russia (s16). Vavilov
and Janischevsky at Saratov (s10), assigned two taxa to the same population of wild-
type plants: C. sativa var. spontanea (Vavilov, 1922) and C. sativa var. ruderalis
(Janischevsky, 1924).
❷ Euro-Siberian region, east:
Daniel Messerschmidt and Johann Gmelin explored Siberia, 1720–1727 and 1733–
1743, respectively. Messerschmidt, not a Linnaean, coined a prolix polynomial for
plants in Transbaikal, east of Lake Baikal: “Cannabis erratica, montana, procera,
daurica, folio minore semine lupulino similes, paruulo, guttato” (×2) (Amman, 1739).
Gmelin (1768) was a Linnaean, and used Linnaeus’s pre-1753 name, Cannabis foliis
digitatis, for wild hemp between Krasnoyarsk and Irkutsk (s17).
Falck (1786) assigned C. sativa to feral plants in Bashkiria (s18). Georgi (1800)
wrote of C. sativa, “We have indigenous or self-growing hemp… in the Urals (s19),
and the Ufa (s18), in Siberia between the rivers Yenisei and Irkutsk (s17).”
Pallas (1776) used Messerschmidt’s polynomial for “a small wild hemp, which has a
strange appearance” near Kyakhta (×2). Sievers (1796) assigned Cannabis erratica to
wild-type plants growing in Transbaikal near Kyakhta (×2). Ledebour (1847–49)
synonymized C. erratica by Messerschmidt and Sievers under a new varietal name,
but tentatively, with a question mark: Cannabis sativa β? davurica. Turczaninow
(1856) synonymized Messerschmidt’s taxon under C. sativa. Herder (1892) synony-
mized C. sativa β davurica and C. erratica under C. sativa.
von Ledebour et al. (1833) assigned C. sativa to wild-type plants growing along the
Bukhtarma River in the eastern Kazakh steppe (s20) and along the Ulagan River in the
Altai Mountains (s21). Nearly a century later, Sinskaya (1925) discovered a full
spectrum of Cannabis morphologies in the Altai, from wild-type forms to fully
domesticated plants (s21).
❸ Mediterranean region:
The headwaters region of the Po River in Italy is known as Piemonte (in Italian) and
Piémont (in French). Lamarck (1785) described C. sativa as presque naturalisée in
Piémont (s22). Other botanists assigned new names to a unique landrace in that region.
D’Andrieux (1771) coined Cannabis sativa gigantea for chanvre de Piémont (×3). His
son-in-law Vilmorin added cachet to chanvre du Piémont by giving it a new binomial
name, C. maxima (×3) (Sénac, 1826). Rey (1835) imported chanvre du Piémont from
Carmagnola, and coined the taxon C. gigantea (×3). Vilmorin (1837) countered Rey’s
new binomial by coining his own, C. gigantea (×3). His son, Vilmorin (1892) assigned
C. sativa excelsior to chanvre du Piémont (×3).
Elsewhere in Italy, Parlatore (1867) simply used C. sativa for feral hemp (s23).
Velenovský (1898) described spontaneous C. sativa in Bulgaria (s24). Across the
Mediterranean in Algeria, Guyon (1842) wrote about hhachiche. He knew Lamarck’s
C. indica from the literature, but stated that the plants in Algeria were “none other than
our common hemp, C. sativa” (s25). According to de Candolle (1869), Friedrich Noé, a
German botanist working in Istanbul (s26), used the name Cannabis orientalis for
plants in his herbarium collection, but did not publish the taxon. Noé’s collaborator
Boissier (1879) reduced C. orientalis (along with C. indica and C. chinensis) to a
synonym of C. sativa.
❹ West and Central Asia (Irano-Turanian):
Samuel Gmelin corresponded with Linneaus, like his aforementioned uncle. He
collected C. sativa near Lankarān and Astārā (s27) in present-day Azerbaijan (Herder,
1892). Hohenacker (1838) also collected ruderal specimens of C. sativa in Azerbaijan,
close to Lankarān and Astara on the border with Persia (s27).
Falck (1786) assigned C. sativa to feral plants in “Bukhārā” (the closest he actually
got to Bukhārā was Orsk, s28) and “Soongaria” (a.k.a., Dzungaria—Falck’s assistant
Bardanes skirted the edge of Dzungaria in Semey, s29). Becker (1873) collected wild
C. sativa near Kasumkent in Dagestan.
Chardin (1811), or rather his editor, assigned the taxon C. sativa to bueng plants in
Isfahān, Persia (s30). Claus (1838) reported “abundant” C. sativa on islands dotting the
Volga River delta (s31). Karelin and Kirilov (1841) assigned C. sativa to plants “in
pratensibus ad fl. Irtysch frequens [frequently in meadows near Irtysh River, s29], nec
non in deserti Soongoro-Kirghisici [and certainly on the Kirghiz steppe, s32], arenosis
ad lacum Noor-Saissan [growing on sand near Lake Zaysan, s29].”
Schrenk (in Trautvettero, 1867) collected C. sativa at three places in eastern
Kazakhstan in 1840–1841: on Lake Alakol near the Dzungarian Gate (s32); the
Kyskatsch mountains, between Lake Balkhash and Lake Zaysan (between s29 and
s32); and near Khantau, between Lake Balkhash and Bishkek (s33). Griffith (1847)
found Cannabis (no species name) at Jegdalek between Kabul and Jalalabad in
Afghanistan (s34).
Basiner (1848) assigned C. sativa to a plant called kender in Khiva (s35).
Bunge (1851) collected C. sativa near Kulagin on the Ural River, just north of
the Caspian Sea (north of s31). Becker (in Herder, 1892) collected wild-type
C. sativa along the lower Volga near Sarepta, Bogdo, Saratov, and Astrakhan
(between s10 and s31). Semenov (1998) reported wild C. sativa growing near
Kurmenty Pass northeast of Lake Issyk-Kul in 1857 (just east of s33). Seminov
wasn’t the best botanist—at herb. LE we saw a specimen he called “C. sativa”
that was actually Humulus lupulus.
Valikhanov (1865) said hashīsh was extracted from C. sativa in Kāšḡar (s36). Shaw
(1880) wrote a Turkic-English dictionary in Kāšḡar, and translated kaindir as the hemp
plant, C. sativa (s36). Aitchison (1888) applied the name C. sativa to plants at Rui-Kauf
in Persia (s37). Meyer (in USDA, 1912) collected C. sativa at Sanju oasis (s38), “a
small-seeded variety of hemp… used for seed oil and hashish made from the young
tops.”
❺ Sino-Japanese (East Asiatic)
Thunberg (1784), a student of Linnaeus, extended the C. sativa taxon to Nagasaki,
Japan (s39). von Bunge (1833) found C. sativa growing “quasi-feral” in northern China
(s40). Martius (1832) gave the name Cannabis sativa gigantea to plants growing 6 m
tall in China. He did not give a location, but Canton (i.e., Guǎngzhōu, s41) was the only
Chinese port open to foreigners at that time. Bretschneider (1882), a Russian physician
in Běijīng (s42), assigned Chinese hemp to C. sativa.
❻ African-Indian desert (Saharo-Arabian)
Forsskål (1775), a student of Linneaus, assigned C. sativa to plants in Egypt (s43).
Gastinel (1849) did not recognize C. indica as a species separate from C. sativa in
Cairo (s43).
❼ Sudano-Zambezian-Sindhia
Griffith (1847) found “Cannabis” (no species name) in places that later became
famous for charas: Dādur Nāla in Sindh (s44), and Burhan in Punjab (s45). Jacquemont
(1861) assigned C. sativa to plants in Rajputana and Punjab (between s44 and s45).
Aitchison (1864) described C. sativa in Punjab. Duthie (1898) reported wild C. sativa
growing in three Chitral locations—Drosh, Dir, and Mirga (s46).
❽ South Asia (Indian)
Many botanists who knew Lamarck’s taxon nevertheless adhered to the Linnaean
party line, and applied C. sativa to plants throughout India—in Bengal (s47, Fischer,
1810, Roxburgh, 1832, Butter, 1839), in southern India (s48, Buchanan, 1807), and in
the Himalaya (s49, Hardwicke, 1801, Royle, 1839, Hooker and Thomson, 1855).
Wallich (1828–1849) applied C. sativa to plants all over South Asia—Calcutta (Ben-
gal), Sudallapur (Karnataka), Bareilly (Uttar Pradesh), Hyderbad (Andhra Pradesh),
and Kathmandu (Nepal).
❾ Continental Southeast Asia (Indochina)
de Loureiro (1790) assigned C. sativa to plants cultivated for fiber and drugs in
“Cochinchina,” southern Vietnam (s50). Wallich (1828–1849) collected C. sativa in the
Toong Dong mountains near Innwa, Burma (s51). Griffith (1847) reported “C. sativa is
found here” in Sassi (south of s51). Crévost (1917) coined the taxon Cannabis gigantea
for plants grown for fiber and seed oil in the Tonkin highlands (×5).
❿ Malayian (Malesian)
Stickman (1754), a student of Linnaeus, assigned C. sativa to drug plants cultivated
in the Moluccas islands, Indonesia (s52). Thunberg (1796), another student, wrote
about C. sativa, known as ginje in Jakarta (3000 km west of s52).
Part B . Cannabis indica and segregates
❶ Euro-Siberian region, west

No plants native to this region were named C. indica by field botanists.
❷ Euro-Siberian region, east
No plants native to this region were named C. indica by field botanists.
❸ Mediterranean region
von Maltzan (1869) said hashīsh and kif in Algiers was made from C. indica (i1),
whereas De Courtive (1848) and Dukerley (1866) equivocated between C. indica and
C. sativa for plants from Algiers. Mongeri (1865) described C. indica cultivation at
Izmit and Bursa, south of Istanbul (i2).
❹ West and Central Asia (Irano-Turanian)
Regel (1880) identified plants growing near Lake Issyk-Kul as a new variety,
C. sativa γ asperrima (i3). Polak (1865) assigned C. indica to medicinal plants at
Tehran (i4). Vámbéry (1868) described beng or bengis as “the poison produced from
the Cannabis indica” in Bukhārā and Khoqand (i5).
Henderson and Hume (1873) assigned C. indica to drug-type plants cultivated in
Yarkand (i6). However, at the Kew Herbarium, a Yarkand specimen collected by
Henderson is labeled “Cannabis sinensis.” Vavilov and Bukinich (1929) found feral
and wild Cannabis in Afghanistan, and named them C. indica f. afghanica and
C. indica var. kafirstanica, respectively (i7).
❺ Sino-Japanese (East Asiatic)
Fischer (1810) erected a new taxon for Chinese hemp, Cannabis chinensis. He did
not state provenance for the germplasm, but Canton (i.e., Guǎngzhōu, ×6) was the only
Chinese port open to foreigners at that time. von Humboldt (1811) considered Chinese
hemp a type of C. indica (i7). von Siebold (1827) described C. sativa b indica in Japan
(i8). Boitard (1839) suggested that Chinese hemp was “probably the same species as
Indian bangue.”
Tatarinov (1858), a Russian physician in Běijīng, assigned C. indica to local medical
plants (i9). Hedde (1848) assigned C. indica to germplasm obtained from Shanti
district, Guǎngzhōu (i7). Itier (1846) assigned C. indica to germplasm obtained from
Huángtián, 170 km N.E. of Guǎngzhōu (i7).
Itier gave germplasm to Delile (1849), who grew it in Montpellier. Delile was
unaware of Fisher’s publication, and re-coined Cannabis chinensis (×6). Koch (1854)
noted that C. chinensis resembled drug plants from India. Vilmorin (1851) gave
Chinese hemp a new name, “Cannabis gigantea Delile.” This Latin name was an
unfortunate choice, because it had previous been applied to chanvre de Piémont (×3)
(Rey, 1835; Vilmorin, 1837).
Jomard (1852) wrote about Chinese hemp, to which he assigned Vilmorin’s taxon
C. gigantea. Itier (1853) also reassigned lo-má to C. gigantea. Heuzé (1860) synony-
mized Vilmorin’s taxon with “Cannabis indica Lam.” Alefeld (1866) assigned Can-
nabis sativa gigantea to “Chinese, Oberländer, or Piedmontese giant hemp”—in other
words, to both C. gigantea Vilmorin, 1837 (Piedmontese hemp) and C. gigantea Delile
ex Vilmorin, 1851 (Chinese hemp).
Pabst (1887) erected C. sativa var. chinensis, and described plants as “rich and
protruding branches, 3-6 m high.” Hoffmann (1944) coined Cannabis var. indica
subvar. Gigantea for Chinese hemp, indicating its kinship with indica. Zhang (1990)
noted relatively high levels of THC in some Yúnnán landraces, which he named
Cannabis sativa ssp. indica var. yunnanica (i10). These landraces may trace back to
Sayyid Ajjal Šams al-Din Omar (1211–1279), a Muslim from Bukhārā, appointed as
Yúnnán provincial governor by Kublai Khan. Hillig (2005) and Clarke and Merlin
(2013) considered Chinese hemp a biotype of indica.
❻ African-Indian desert (Saharo-Arabian)
Ibn al-Baitār, a Spanish Moor who moved to Cairo around 1240, identified qinnab
hindī (Indian hemp) growing in Egypt (i11), distinct from plants he knew in Spain,
qinnab and qinnab barrī (Ibn al-Baitār, 1985). Godard (1867) and Mackenzie (1893)
describe hashīsh made from C. indica in Egypt (i11).
Hasselquist (1766), a student of Linnaeus, broke from Linnaean orthodoxy and
assigned Cannabis vulgaris to plants cultivated in Palestine for chashis (×4). Stokes
(1812) synonymized Hasselquist’s taxon under C. indica (i12). Mongeri (1865) de-
scribed C. indica cultivation at Mosul in present-day Iraq (i13).
❼ Sudano-Zambezian-Sindhia
von Maltzan (1873) lived in Aden, Yemen, where he encountered hashīsh made
from C. indica (i14). Honigberger (1852) identified plants in Punjab and Kashmir as
C. indica, due to their unique chemistry compared to C. sativa (i15).
❽ South Asia (Indian)
Rheede (1690) gave the names Kalengi-cansjava and Tsjeru-cansjava to male and
female Cannabis, respectively, at Kochi on the Malabar coast (×7). O’Shaughnessy
(1838–1840) employed the taxon C. indica, yet he considered “the Cannabis sativa and
indica are identical” (i16). Kerr (1877) a British-trained Bengali botanist, referred to
Bengal gañjā as C. indica, distinct from C. sativa (i16).
Johann Friedrich Metz was a German missionary who collected plants on the
Malabar coast (i17). His collection of “Cannabis indica” at herb. LE is a dense,
lightly-seeded bud—the oldest specimen of manicured gañjā we have seen. Rottler
(1836–7), a Danish missionary at Tranquebar and Madras (i17), translated Tamil
kañcam as Cannabis indica. A few botanists in the Himalaya referred to local plants
as C. indica (i18, Cleghorn, 1866, Aitchison, 1869, Lawrence, 1895).
❾ Continental Southeast Asia (Indochina)
Hedde (1848) assigned C. indica to germplasm obtained from Tourane, now Đà
Nẵng, Vietnam (i19).
❿ Malayian (Malesian)
Rumpf (1747) assigned C. indica to drug plants cultivated in the Moluccas islands,
Indonesia (i20). Blume (1825), a German botanist who moved to Jakarta, used the
taxon Cannabis sativa var. indica for plants with the local name ginji (3000 km west of
i20).
Appendix 4: Taxonomic models in the 20th-21st century
The “species debate” became a cause célèbre in 1970s court cases. The primary
adversaries in this debate were Richard Evans Schultes (1915–2001), a Harvard
ethnobotanist, and Ernest Small (1940-), a Canadian taxonomist who specialized in
Cannabis. Their taxonomic debates in courtrooms often involved “an appreciable
irrational emotive component” (Small, 1979). One taxonomic paper from that era made
reference to “the Honorable Harry Anslinger,” and the “menace” posed by Cannabis
(Quimby et al., 1973).
Schultes’s taxonomic work is limited to two publications. Initially, Schultes (1970)
considered Cannabis a monotypic genus. Subsequently, Schultes et al. (1974) recog-
nized C. sativa, C. indica, and C. ruderalis. Schultes changed his opinion based on an
analysis of herbarium specimens, a survey of the Mississippi Cannabis plantation, and
“preliminary field work” in Afghanistan.
Schultes collaborated with Loren C. Anderson at Florida State University. Anderson
published two papers. Anderson (1974) compared wood anatomy in C. sativa (a feral
hemp plant in Kansas, likely of Chinese provenance) and C. indica (a plant that
Schultes collected in Afghanistan). Anderson (1980) compared leaf variation in four
populations of Cannabis: 1. C. sativa (fiber varieties—tall, laxly branched plants), 2.
C. sativa SS (Small-Seeded drug plants from India and Pakistan—relatively tall, with
narrow leaflets), 3. C. indica (drug plants from Afghanistan—short, densely branched,
with broad leaflets), and 4. C. ruderalis (wild plants, probably from Central Asia—very
short and unbranched). Anderson illustrated his species concept in a line drawing
(Fig. 5).
Small began Cannabis taxonomic research in 1971, under common-garden condi-
tions on a three-acre farm near Ottawa, Canada. He grew nearly 400 Cannabis
accessions obtained from the USDA, the UN, the University of Mississippi, and from
botanical gardens around the world. Small and colleagues conducted cross-breeding
experiments (Small, 1972, 1984). They assessed many taxonomic characters: cannabi-
noid content (Small & Beckstead, 1973a, b; Small et al., 1975; Small & Marcus, 2003),
seed chemistry (Small et al., 1976), morphological variation in seeds (Small 1975),
vegetative characters (Small et al., 1976), flower characters (Small & Naraine, 2015a,
b), developmental characters (Small et al., 2003), and biogeographical aspects (Small,
2015a, b).
This work was summarized (Small and Cronquist, 1976), and elaboratored in a two-
volume text (Small, 1979). His work continues to evolve—Small (2015a) has taken on
aspects of Hillig’s work regarding Chinese hemp. Small’s classification and nomencla-
ture is widely accepted around the world (e.g., Hanelt, 2001; Yang, 2003; Kojoma
et al., 2006; Mukherjee et al., 2008; Mabberley, 2008; Shipunov, 2010; Chandra et al.,
2013). Small’s taxonomic scheme is attested on high-profile websites. Examples:
http://frps.eflora.cn, ru.wikipedia.org/wiki
http://plants.usda.gov/java
www.ars-grin.gov
www.theplantlist.org
www.tropicos.org
www.ncbi.nlm.nih.gov/taxonomy
Google Scholar (https://scholar.google.com) computes citation metrics for authors

and their publications. We applied the “advanced search” algorithm, using the words
“taxonomy,” “cannabis,” and author name. After screening for appropriate hits, we
obtained the following results: Small: 18 publications, cited collectively by 675; Hillig:
7 publications, cited collectively by 299; de Meijer: 5 publications, cited collectively by
207; Schultes: 2 publications, cited collectively by 193; Gilmore: 5 publications, cited

collectively by 179. No other authors were cited more than 150 times.
Hillig and colleagues cultivated 157 accessions under common-garden conditions at
Indiana University. They obtained accessions from Small, the CPRO, Dutch seed
banks, the Vavilov Institute, law enforcement agencies, field stations, and botanical
gardens across Eurasia. Hillig analyzed genetic characters (Hillig, 2004a, 2005),
cannabinoid profiles (Hillig & Mahlberg, 2004), terpenoid profiles Hillig, 2004b, seed
morphology (Hillig, 2005b, c), leaflet characters (Hillig, 2005b, c), stalk morphology
(Hillig, 2005b, c), developmental characters (Hillig, 2005b, c), host-parasite characters
(Hillig, 2005b, c, McPartland & Hillig, 2003, 2004, 2006), and biogeographical aspects
(Hillig, 2005a, b).
De Meijer and colleagues systematically investigated Cannabis under common-
garden conditions at Wageningen University. They collected over 150 Cannabis
accessions—now the CPRO collection. They obtained germplasm from the Vavilov
Institute, IPK-Gatersleben Institute, Dutch seed banks, European breeders of fiber-type
plants, and botanical gardens across Eurasia. De Meijer has continued Cannabis studies
at GW Pharmaceuticals in the United Kingdom. De Meijer analyzed cannabinoid
content (De Meijer & van Soest, 1992; De Meijer & Keizer, 1996), the inheritance
of cannabinoid phenotypes (De Meijer et al., 2003; Mandolino et al., 2003; De Meijer
& Hammond, 2005. De Meijer et al., 2009a, b), stalk morphology and chemistry (De
Meijer, 1993a, 1994b, 1995), seed morphology and chemistry De Meijer & Keizer,
1996, leaflet characters (De Meijer et al., 1992; De Meijer & Keizer, 1996), host-
parasite relationships (De Meijer, 1993b; De Meijer & Keizer, 1996), developmental
characters (de Meijer & Keizer1996), sequence heterogeneity in THCA-synthase genes
(Onofri et al., 2015), and biogeographical aspects (De Meijer, 1999, De Meijer, 2004,
2014).
Gilmore and colleagues researched Cannabis genetics at Australian National Uni-
versity. Their early work analyzed police-confiscated materials of unknown origin,
lacking taxonomic inferences. They developed primers for microsatellite DNA loci,
also called short tandem repeats (STRs), and validated their work for forensic purposes
(Gilmore & Peakall, 2003; Gilmore et al., 2003, 2007; Howard et al., 2008, Howard
et al., 2009). Gilmore et al. (2007) switched to polymorphic cpDNA and mtDNA loci,
which they sequenced from known CPRO accessions.
“Strain” names
The International Code of Nomenclature for Cultivated Plants (ICNCP, Brickell, 2009)
regulates the naming of plants whose recent evolution has been influenced by human
selection. The ICNCP’s basic unit of classification is the cultivar, “an assemblage of
plants that (a) has been selected for a particular character or combination of characters,
(b) is distinct, uniform, and stable in these characters, and (c) when propagated by
appropriate means, retains those characters.”
The ICNCP’s rules for naming and describing cultivars involve valid publication,
typification (“Nomenclatural Standard”), and priority. The ICNCP is available online
for consultation regarding these provisions.
Cultivar names meeting ICNCP provisions are placed in single quotation marks. The
best known Cannabis cultivar, judging from the number of Google search hits, is
‘FINOLA’. The ICNCP also recognizes “Group” names, “All members of a Group
must share the character(s) by which that Group is defined.” Small (2015a) proposed
six Group names for cultivated kinds of Cannabis and their hybrids.
Article 2.2 in ICNCP stipulates that the words “variety,” “form,” and “strain” must
not be used for the word “cultivar” (Brickell, 2009). Notwithstanding Article 2.2, some
national and international plant registries use “variety” interchangeably with “cultivar.”
The words “variety” and “form” also designate subspecies ranks in the ICN. “Strain” is
not formally recognized by the ICN or ICNCP.
Segregates within “Sativa” and “Indica” are called “strains.” Watson (1985) coined
several well-known “strain” names, such as “Original Haze”, “Skunk #1”, “California
Orange”, “Afghani #1”, and “Early Girl”. Small (2015a) pointed out that strains are
conceptually identical to cultivars, but almost no strains have met ICNCP requirements
for cultivar recognition.
Clarke and Merlin (2015) took umbrage, and argued that “the bigger picture”
justifies using “strain” and “cultivar” as equivalents. Small (2015b) retorted that they
showed insufficient respect for codes of nomenclature, which hold pragmatic and moral
status. “The botanical codes are by and large adhered to by scientists, commercial
interests and editors because they provide stability and reliability to names that
otherwise would result in confusion.”
“Strain” names have proliferated exponentially. Watson (1985) named 10 strains.
Within 15 years, Dutch seed companies offered 150 strains for sale (Clarke, 2001). A
decade later, the number of strain names reached 900 (Cannabis Strain Database, 2010).
Most recently, Leafly (2015) listed 1535 strain names, and Seedfinder (2015) listed
6510 strain names. Doyle (2007) referred to strains as ganjanyms. Most strains are
recognized hybrids, characterized as “Sativa-dominant” or “Indica-dominant.” In to-
day’s largely illicit market, strain names are swapped and counterfeited, and generally
unreliable (Lee, 2013; Sawler et al., 2015; Pierson, 2016).
The strain “AK-47” examples the arbitrariness of vernacular classification: “AK-47”
won “best Sativa” in the 1999 Cannabis Cup, and won “best Indica” 4 years later. The
ancestry of “Super Silver Sour Diesel Haze” offers a window into this warren of
questionable pedigrees (Box 2).
Box 2 Putative ancestry of “Super Silver Sour Diesel Haze”, from Seedfinder (2015)
Super Silver Sour Diesel Haze, a cross of Super Silver Haze x Sour Diesel
Super Silver Haze, a cross of {(Haze x Haze) x Skunk #1} x {(Haze x Haze) x Northern
Lights #5}
o Haze, a cross of landraces from Colombia, Mexico, Thailand, and southern India
o Skunk #1, a cross of (Afghani x Colombian Gold) x Acapulco Gold
o Northern Lights #5, a murky pedigree, either pure Afghani, or a Thailand x Afghani
hybrid
Sour Diesel, a cross of Original Diesel x DNL
o Original Diesel, a cross of Chemdawg x (MassSuperSkunk x SensiNL)
Chemdawg, of unknown lineage
MassSuperSkunk, a cross of Skunk #1 x Afghani
SensiNL, a cross of three Northern Lights strains of shared pedigree, NL #1,
NL #2, and NL #5
o DNL, a cross of (RFK Skunk x Hawaiian) x Northern Lights
RFK Skunk, likely derived from Skunk #1
Hawaiian, a “Sativa/Indica hybrid”
Northern Lights, as described above
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Models of Cannabis Taxonomy, Cultural Bias, and Conﬂicts between Scientiﬁc

Article in The Botanical Review · June 2017

John McPartland Geoffrey W Guy

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Models of Cannabis Taxonomy, Cultural Bias,

John M. McPartland 1,2 & Geoffrey W. Guy 1

Keywords Cannabis sativa . Cannabis indica . Taxonomy . Nomenclature

Controversies regarding the taxonomy and nomenclature of C. sativa have generated

Cannabis sativa L. Protologue

Cannabis indica Lam. Protologue

Historical Bias in Usage of C. sativa and C. indica

Botanists soon rendered subjective decisions regarding Lamarck’s concept of a

Reports of Cannabis growing in two phases of domestication—wild and cultivated—

Historical Bias, Part II

Vavilov traveled to Afghanistan in 1924, and encountered all phases of domestica-

lanceolate leaflets, growing outside of Afghanistan, in India, Nepal, China, South

Fig. 4 Classification of Cannabis sativa, according to Small & Cronquist, 1976

A former student explained that Schultes’s “intense dedication” to libertarian beliefs

Schultes’s concepts were embraced by two other defense witnesses, William

The Rise of Vernacular Taxonomy

biotype” referred to wild-type plants from India and Nepal—accessions assigned to

& Indica—Wrongly called “sativa” (Clarke & Merlin, 2016)

Appendix 1 literature cited

Appendix 2 literature cited

Linnaeus C. 1737a. Critica Botanica. Conrad Wishoff, Leiden.

Appendix 3 literature cited

Appendix 4 literature cited

Appendix 1a. Cannabis sativa L.

descriptive elements he provided in Species Plantarum. Even by Linnaeus’s standards,

Synonymy and references

Box 1 Translation of author and citation abbreviations by Linnaeus (1753)

Linnaeus’s type specimen of C. sativa is stored in the Linnaean Herbarium (herb.

Appendix 1b. Cannabis indica Lam.

Synonymy and references

Lamarck cited six synonyms and their authors:

Four authors in Lamarck’s synonymy (1, 2, 3, 4) previously appeared in Linnaeus

Lamarck’s type specimen of C. indica is deposited at the Muséum National

trichomes. Styles and stigmas are prominent, agglutinized with trichome

Lamarck erroneously described C. indica with “alternately branching leaves” and

Appendix 1c. Cannabis sativa var. spontanea Vav.

horseshoe is clearly defined and undoubtedly is a morphological and biological feature

Synonymy and references

Vavilov (1922) cited no synonyms. He apparently overlooked Czernajew (1859), who

Cannabis reverted to a wild-type phenotype within 50 generations (years) of prohibi-

Appendix 1d. Cannabis ruderalis Janisch.

demonstrated a prolonged period of dormancy and slow germination, whereas seeds of

Synonymy and references

Type specimen and illustrations

Janischevsky’s lectotype specimen of C. sativa var. ruderalis, collected near Saratov,

Fig. 6 Line drawings of C. ruderalis by Janischevsky (1924)

Рис. 1. A fruit of hemp collected in the vicinity of Saratov (Dudakov ravine).

Janischevsky (1924) said he found hemp growing in “wild circumstances” in 1897,

America—obviously escapes from cultivation, and not wild, indigenous plants

Appendix 1e. Cannabis sativa f. afghanica Vav.

Vavilov encountered wild and weedy plants in Afghanistan. First he described

“We observed wild hemp… in districts where hemp cultivation is entirely

features of a typical wild plant,” followed by “the plants constitute a morpho-

Synonymy and references

Vavilov provided no synonymy and cited no references.

No specimen labeled afghanica is deposited in Vavilov’s herbarium at WIR. Serebriakova

Vavilov and Bukinich considered afghanica a “specialized form” of kafirstanica,