ArticlePDF Available

Cannabis Taxonomy: The "sativa" vs. "indica" debate



No caption available
Content may be subject to copyright.
arijuana and hemp
(Cannabis) and the
closely related hop genus
(Humulus) are the only widely
known genera included in the
small, but economically valu-
able, Cannabaceae family. Swed-
ish botanist Carl Linnaeus, the
“father of modern taxonomy,
first published the scientific name
Cannabis sativa in his seminal
Species Plantarum of 1753. The
Latin name Cannabis derives
from Greek (kannabis) and may
have been originally derived from
Scythian. The term sativa simply
means “cultivated” and describes
the common hemp plant that was
widely grown across Europe in
Linnaeus’ time. We, the authors,
consider C. sativa to be native
to western Eurasia and especially
Europe, where, for millennia, the
plant has been grown for its strong
fibers and nutritious seeds, and
from where it was introduced to
the New World multiple times
during early European coloniza-
tion. Cannabis sativa plants also
produce very small amounts of
the compound delta-9-tetrahydro-
cannabinol (THC), the medically
valuable and primary psychoac-
tive cannabinoid found only in
Cannabis. Since C. sativa evolved
within the geographical limits of
western Eurasia, it represents only
a small portion of the genetic
diversity seen in the genus Canna-
bis worldwide.1
In 1785, European natu-
ralist Jean-Baptiste Lamarck
described and named a second
species, Cannabis indica, mean-
ing “Cannabis from India,” after
the origin of the first samples of
this highly psychoactive plant that
By Robert C. Clarke and
Mark D. Merlin, PhD
The ‘Sativa’ Vs.
Indica’ Debate
44 ISSUE 110 2016
Ripe female marijuana flowers
are covered with glandular
trichomes or resin glands.1
Photo ©2016 Todd McCormick 2016 ISSUE 110 45
reached Europe. Cannabis indica has the genetic potential
to produce relatively large amounts of THC. The species is
used for marijuana and hashish production, but in many
regions of eastern Asia it also has a long history of cultiva-
tion for fiber and seed. Humans make cloth out of C. indica
fibers and eat the seeds, but this native eastern Eurasian
species is more commonly used today as a drug plant with
widespread social and medicinal importance reaching well
beyond its original geographical range.1
While Karl Hillig, PhD, was a doctoral student at
Indiana University, he used morphological and chemical
characteristics to investigate the diversity of the Cannabis
genus and proposed taxonomic groupings (subspecies) that
support the original two-species concept.2 -5 Hillig recog-
nized European cultivated Cannabis as a separate species
(C. sativa). Because this species typically has narrow leaflets
and is primarily used for hemp fiber and seed production,
we refer to it as narrow leaflet hemp (NLH). European C.
sativa NLH populations are much less genetically diverse
than those found in many other regions.
Hillig assigned the remainder of the world’s cultivated
varieties to C. indica and divided them into three subspe-
cies. One of these subspecies, C. indica subsp. indica,
includes varieties that span the Indian subcontinent from
Southeast Asia to western India and into Africa. These
traditional drug varieties produce abundant amounts of
THC with little if any cannabidiol (CBD). CBD is the
second most common cannabinoid, and is non-psychoac-
tive, but it has been shown to be medicinally effective for
a variety of indications. By the 19th century, high-THC
C. indica subsp. indica reached the Caribbean region and
steadily spread throughout Central and South America.
Since the 1960s, most of the drug Cannabis that reached
North America and Europe was C. indica subsp. indica.
Marijuana users commonly call domestically grown plants
of these varieties “sativas” because their leaflets are relatively
narrow, and therefore exhibit a superficial resemblance to
those of European NLH plants. However, in our construct,
this is a misnomer as C. sativa plants produce little if any
THC. Based on Hillig’s research, we now refer to members
of C. indica subsp. indica as narrow leaf let drug (NLD)
varieties. Although they have relatively narrow leaf lets like
NLH (C. sativa) plants, the NLD plants can produce an
abundance of THC and are most commonly used for their
psychoactive effects. Based on taxonomic tradition, these
plants are properly called “indicas” rather than “sativas.”
A second C. indica subspecies originated in Afghanistan
where crops were traditionally grown to manufacture sieved
hashish, a mechanically concentrated Cannabis drug. From
1974 , when descriptions and photos of Afghan Cannabis
were published by Harvard professor Richard E. Schultes,
PhD, it became readily apparent that it represented a type
of drug Cannabis previously unknown outside of Eurasia,
belonging neither to Linnaeus’ C. sativa nor Lamarck’s C.
indica.6 Its shorter, more robust stature, and broad, dark
green leaves easily distinguish it from the taller, lighter
green, and more laxly branched NLD varieties. Because
of its limited geographic range and restricted usage, the
Afghan genome is less diverse than the NLD genome. By
the late 1970s, seeds of Afghan hashish varieties reached
Europe and North America and were rapidly disseminated
among marijuana growers. At this time, all Cannabis vari-
Cannabis sativa
Photo ©2016 Steven Foster
46 ISSUE 110 2016
eties were commonly considered to be members of a single
species, C. sativa, and the familiar NLD marijuana varieties
were called “sativas” to differentiate them from the newly
introduced and quite different looking Afghan varieties
commonly called “indicas.” Hillig named them C. indica
subsp. afghanica, which we now refer to as broad leaf-
let drug (BLD) varieties to differentiate them from NLD
varieties. On average, populations of BLD plants contain
approximately equal amounts of THC and CBD. Although
BLD varieties are also considered by us to be members of C.
indica, it is more correct to distinguish them from subspe-
cies indica from India by calling them subspecies afghanica,
or simply “Afghans.”
Hillig’s third grouping within C. indica is subspecies
chinensis, which comprises the traditional East Asian fiber
and seed varieties and associated feral populations. We
refer to this group as broad leaf let hemp (BLH). Like other
subspecies of C. indica, varieties of C. indica subsp. chinensis
possess the genetic potential to produce psychoactive THC,
but East Asian cultural traditions, such as Confucian-
ism, have long encouraged the selection of these varieties
for their economically valuable fiber and seed, rather than
their psychoactive potential. As a result, total cannabinoid
production is lower than in subspecies indica and afghanica.
Evolutionary theory predicts that, at some point in time,
there must have been a putative ancestor of the two modern
species, C. sativa and C. indica. This ancient ancestor is
often referred to as C. ruderalis, which may have origi-
nated somewhere in Central Asia. However, by now it is
probably extinct, and seemingly ancestral populations are
more likely descendants of feral plants that escaped from
cultivation long ago. Evolutionary hypotheses based on
plant distribution studies, paleoclimate modeling, archaeo-
logical evidence, and the historical record propose that C.
sativa NLH most likely originated in a temperate region of
western Eurasia, possibly in the foothills of the Caucasus
Mountains, from a putative hemp ancestor with diminished
biosynthetic potential to produce THC. Cannabis indica
likely originated in the Hengduan Mountains, in present-
day southwestern China, from a putative drug ancestor that
had evolved an enhanced ability to produce THC. Early C.
indica populations diversified as they were introduced by
humans to different geographical regions where they may
have further evolved into the three subspecies, all of which
produce THC.1
Cultivated plant varieties are called cultivars, and when
cultivars are grown and maintained by local farmers over
generations, we refer to them as landrace cultivars, or land-
races. Landraces evolve in a balance between natural selec-
tive pressures exerted by the local environment favoring
survival, and human selections favor-
ing a cultivar’s ability to both thrive
under cultivation and produce particu-
lar culturally preferred products. Early
humans spread Cannabis into many new
regions as they moved, and at each new
camp or settlement they selected seed
from superior plants that were suited
to their own uses and processing meth-
ods. By sowing seeds from the most
favorable individuals, traditional farm-
ers developed and maintained the land-
races upon which present-day hybrid
hemp and drug cultivars were founded.
Hemp cultivars were derived from
crosses between different European
NLH landraces and East Asian BLH
Table 1. Twenty-first century Canna-
bis taxonomy
Cannabis is presently subdivided into
two species, C. sativa and C. indica.
Cannabis indica is further divided into
three subspecies, C. indica subsp. chinen-
sis, C. indica subsp. indica, and C. indica
subsp. afghanica. Cannabis sativa and
C. indica subsp. chinensis are hemp
cultivars most commonly grown for fiber
and seed. Cultivars of C. indica subsp.
indica and C. indica subsp. afghanica are
most commonly grown for their drug
content, and most modern sinsemilla
drug Cannabis cultivars are hybrids of C.
indica subsp. indica and C. indica subsp.
afghanica landraces. 2016 ISSUE 110 47www.herbalgram.org2016 ISSUE 110 • 47
Cannabis sativa
Photo ©2016 Steven Foster
48 ISSUE 110 2016
landraces. Traditional Asian,
African, and New World drug
landraces were, until relatively
recently, all pure NLD types.
Before the introduction of BLD
landraces from Afghanistan in
the late 1970s, hybrids between
imported NLD landraces formed
the core genome of domestically
produced drug Cannabis in both
North America and Europe. It is
through crossing NLD and BLD
landraces from such geographi-
cally isolated populations that
modern hybrid sinsemilla (Span-
ish for “seedless”) cultivars were
Unfortunately, we cannot
return today to a region previ-
ously known for its fine Canna-
bis and expect to find the same
landraces that were growing there
decades before. Cannabis is open-
pollinated, with male and female flowers borne on sepa-
rate plants, and, therefore, two plants are usually required
to produce a seed. Random combinations of alleles (forms
of a gene) and accompanying variation are to be expected.
Cannabis landrace varieties are best maintained by repeated
natural and human selection in situ — nature selecting
for survival and humans selecting for beneficial traits.
Without persistent human
selection and maintenance,
these landrace varieties will
tend to drift back to their
atavistic, naturally selected
survival mode.
The Western world began
using imported marijuana
and hashish in the 1960s,
and all of the remarkable
imported varieties avail-
able then were tradition-
ally maintained landra-
ces. Within a decade, the
demand for quality drug
Cannabis exceeded tradi-
tional supplies, and mass
production in the absence
of selection became the
rule. Rather than plant-
ing only select seeds, farm-
ers began to sow all their
seeds in an effort to supply
market demand, and the quality of commercially avail-
able drug Cannabis began to decline. In addition, travel-
ers returned to the supplying nations and introduced seeds
of “improved” Western sinsemilla varieties that interbred
with the local landraces and thus contaminated the local
genomes. Landraces can no longer be replaced; they can
only be preserved. The few remaining pure landrace variet-
Cannabis resin glands secrete an aromatic essential oil
that contains medicinally important cannabinoids and
terpenoids.1 Photo ©2016 Marcus Richardson
Figure 1. Present-day distribution of Cannabis subspecies
Humans spread Cannabis worldwide for a variety of uses. The putative ancestor (PA) of all Cannabis originated some-
where in Central Asia. Our evolutionary hypothesis proposes that as Cannabis spread into new geographical regions and
cultural contexts, it evolved into four major gene pools and taxonomic groups: C. sativa narrow leaflet hemp (NLH), C. indica
subsp. chinensis broad leaflet hemp (BLH), C. indica subsp. indica narrow leaflet drug (NLD), and C. indica subsp. afghanica
broad leaflet drug (BLD) landraces. These four groups also include feral escapes from cultivation and “wild” populations.1 2016 ISSUE 110 49
ies in existence now, some kept alive for decades as seeds and
cuttings, are the keys to future developments in drug Canna-
bis breeding and evolution. It will be a continuing shame to
lose the best results of hundreds of years of selection by local
farmers. After all, our role should be as caretakers preserving
the legacy of traditional farmers for the future benefit of all.
Cannabis research is a work in progress, and not all
researchers agree on a single taxonomy.7 DNA sequencing
is currently being used to characterize the diversity of many
plant and animal groups, including Cannabis. While our
knowledge grows and the evolutionary history of Cannabis is
revealed, changes in taxonomic nomenclature will continue
to reflect our deepening understanding of this medically
valuable, yet controversial, plant. More broadly, whether
we discover that Cannabis plants belong to one or more
species, we can be sure that humans have long known, used,
dispersed, cultivated, and artificially selected these plants to
perpetuate a truly wide range of diversity.
Robert C. Clarke is the author of several Cannabis science
books and has traveled extensively throughout Eurasia docu-
menting traditional Cannabis production and use. His breed-
ing interests include selection and preservation of landrace
varieties, and developing narrow leaflet drug varieties and
hashish cultivars. Clarke is the co-founder and director of
BioAgronomics Group, an international cannabis industry
consultancy, serves as projects manager for the International
Hemp Association, and holds a seat on the Phylos Bioscience
Cannabis Evolution Project scientific advisory board. He may
be contacted at
Mark D. Merlin, PhD, is a professor in the
botany department of the University of Hawai`i
at Mānoa and an elected member of the Linnean
Society of London. A large part of his long-term
biological and historical research has involved drug
plants and their past and present uses by people.
He has authored or co-authored a number of books
on this general subject, including Man and Mari-
juana: Some Aspects of their Ancient Relationships
(Fairleigh Dickinson University Press, 1972), On
the Trail of the Ancient Opium Poppy (Associated
University Presses, 1984), Kava: The Pacific Drug
(co-authored with Vincent Lebot and Lamont
Lindstrom; Yale University Press, 1992), and, most
recently, Cannabis: Evolution and Ethnobotany
(co-authored with Robert C. Clarke; University
of California Press, 2013). He may be contacted at
Modern sinsemilla crops are vegetatively
reproduced in glass houses to ensure uniformity
and quality.1
Photo ©2016 Todd McCormick
1. Clarke RC, Merlin MD. Cannabis: Evolution and
Ethnobotany. Berkeley, CA: University of California
Press; 2013.
2. Hillig KW. A chemotaxonomic analysis of terpe-
noid variation in Cannabis. Biochem Syst Ecol.
3. Hillig KW. A multivariate analysis of allozyme
variation in 93 Cannabis accessions from the
VIR Germplasm Collection. J Indust Hemp.
4. Hillig KW. Genetic evidence for speciation in
Cannabis (Cannabaceae). Genet Resour Crop Ev.
5. Hillig KW, Mahlberg PG. A chemotaxonomic
analysis of cannabinoid variation in Cannabis
(Cannabaceae). Am J Bot. 2004;91(6):966-975.
6. Schultes RE, Klein WM, Plowman T, Lockwood
TE. Cannabis: An example of taxonomic neglect.
Bot Mus Leafl Harv Univ. 1974;23(9):337-364.
7. Small E. Evolution and classification of Cannabis
sativa (marijuana, hemp) in relation to human
utilization. Bot Rev. 2015;81(3):189-294.
... kafiristanica). In last Clarke and Merlin (2016). Abbreviation: NLH = narrow leaflet hemp; BLH = broad leaflet hemp; NLD = narrow leaflet drug; BLD = broad leaflet drug; CBD = cannabidiol, and THC = delta-9-tetrahydrocannabinol two decades, various new hybrid varieties have been also developed like 'Supersativa', 'Bedrocan', 'Bedrobinol', and 'Bediol' etc. (Clarke and Watson 2002;de Meijer 2004;Flemming et al. 2007). ...
... sativa and C. indica), originated from central Asia. The recent taxonomy of Cannabis was given by Clarke and Merlin (2016) as presented in Fig. 2.1. ...
... Graphical distribution of Cannabis subspecies(Clarke and Merlin 2016) into new geographical areas and evolved into 4 taxonomic groups along with gene pools as Cannabis sativa narrow leaflet hemp (NLH), C. indica broad leaflet hemp (BLH), C. indica narrow leaflet drug (NLD), Cannabis indica spp. afghanica broad leaflet drug (BLD). ...
Cannabis sativa (hemp) as multifunctional crop have traditional application as fiber, food, paper, textile and pharmaceutical potential as inflorescences and seed as sources of exciting bioactive secondary metabolites. The Genus Cannabis is the only producer of phytocannabinoids. Extensive studied have been made to describe the origin history, geographical ranges and genetic identity of the Cannabis species but it remains obscured to date. Various high through put genetic marker have been studied to assess the genetic diversity in hemp varieties. Studies also indicated that domestication origin affects the genetic groups of hemp which further consequences on the chemical diversity of the cannabis. Chemotaxonomy using chemical markers also played a crucial role in differencing and allocating the Cannabis taxa. Cannabinoids ratio and terpene composition are the major marker to play an important role in studying chemical diversity of Cannabis sp. Cannabis genus is the only source of phytocannabinoids the dominant chemical class. Other than cannabinoids terpene and non-cannabinoid phenolic compounds also contribute in the chemical diversity of the species. The vast array of phytochemicals presents in the genus have potential application in pharmaceutical industries. However, due to its legalization status very limited study on its chemical and genetic diversity have been done. Therefore, the species needs attention to explore its commercial value.
... However, vernacular "Sativa," "Indica," and "Ruderalis" threaten to subsume formal ICN nomenclature. Many authors have derided the inaccuracy of vernacular taxonomy (e.g., Hazekamp & Fischedick, 2012;Russo, 2016;Clarke & Merlin, 2016). ...
Full-text available
Debates over Cannabis sativa L. and C. indica Lam. center on their taxonomic 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 vernacular “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.
Full-text available
The formal botanical taxonomy of Cannabis sativa Linnaeus and C. indica Lamarck has become entangled and subsumed by a new vernacular taxonomy of “Sativa” and “Indica.” The original protologues (descriptions, synonymies, and herbarium specimens) by Linnaeus and Lamarck are reviewed. The roots of the vernacular taxonomy are traced back to Vavilov and Schultes, who departed from the original concepts of Linnaeus and Lamarck. The modified concepts by Vavilov and Schultes were further remodeled by underground Cannabis breeders in the 1980s and 1990s. “Sativa” refers to plants of Indian heritage, in addition to their descendants carried in a diaspora to Southeast Asia, South- and East Africa, and even the Americas. “Indica” refers to Afghani landraces, together with their descendants in parts of Pakistan (the northwest, bordering Afghanistan). Phytochemical and genetic research supports the separation of “Sativa” and “Indica.” But their nomenclature does not align with formal botanical C. sativa and C. indica based on the protologues of Linnaeus and Lamarck. Furthermore, distinguishing between “Sativa” and “Indica” has become nearly impossible because of extensive cross-breeding in the past 40 years. Traditional landraces of “Sativa” and “Indica” are becoming extinct through introgressive hybridization. Solutions for reconciling the formal and vernacular taxonomies are proposed.
Full-text available
Cannabis: Evolution and Ethnobotany is a comprehensive, interdisciplinary exploration of the natural origins and early evolution of this famous plant, highlighting its historic role in the development of human societies. Cannabis has long been prized for the strong and durable fiber in its stalks, its edible and oil-rich seeds, and the psychoactive and medicinal compounds produced by its female flowers. The culturally valuable and often irreplaceable goods derived from cannabis deeply influenced the commercial, medical, ritual, and religious practices of cultures throughout the ages, and human desire for these commodities directed the evolution of the plant toward its contemporary varieties. As interest in cannabis grows and public debate over its many uses rises, this book will help us understand why humanity continues to rely on this plant and adapts it to suit our needs.
Full-text available
Sample populations of 157 Cannabis accessions of diverse geographic origin were surveyed for allozyme variation at 17 gene loci. The frequencies of 52 alleles were subjected to principal components analysis. A scatter plot revealed two major groups of accessions. The sativa gene pool includes fiber/seed landraces from Europe, Asia Minor, and Central Asia, and ruderal populations from Eastern Europe. The indica gene pool includes fiber/seed landraces from eastern Asia, narrow-leafleted drug strains from southern Asia, Africa, and Latin America, wide-leafleted drug strains from Afghanistan and Pakistan, and feral populations from India and Nepal. A third putative gene pool includes ruderal populations from Central Asia. None of the previous taxonomic concepts that were tested adequately circumscribe the sativa and indica gene pools. A polytypic concept of Cannabis is proposed, which recognizes three species, C. sativa, C. indica and C. ruderalis, and seven putative taxa.
Cannabis sativa has been employed for thousands of years, primarily as a source of a stem fiber (both the plant and the fiber termed “hemp”) and a resinous intoxicant (the plant and its drug preparations commonly termed “marijuana”). Studies of relationships among various groups of domesticated forms of the species and wild-growing plants have led to conflicting evolutionary interpretations and different classifications, including splitting C. sativa into several alleged species. This review examines the evolving ways Cannabis has been used from ancient times to the present, and how human selection has altered the morphology, chemistry, distribution and ecology of domesticated forms by comparison with related wild plants. Special attention is given to classification, since this has been extremely contentious, and is a key to understanding, exploiting and controlling the plant. Differences that have been used to recognize cultivated groups within Cannabis are the results of disruptive selection for characteristics selected by humans. Wild-growing plants, insofar as has been determined, are either escapes from domesticated forms or the results of thousands of years of widespread genetic exchange with domesticated plants, making it impossible to determine if unaltered primeval or ancestral populations still exist. The conflicting approaches to classifying and naming plants with such interacting domesticated and wild forms are examined. It is recommended that Cannabis sativa be recognized as a single species, within which there is a narcotic subspecies with both domesticated and ruderal varieties, and similarly a non-narcotic subspecies with both domesticated and ruderal varieties. An alternative approach consistent with the international code of nomenclature for cultivated plants is proposed, recognizing six groups: two composed of essentially non-narcotic fiber and oilseed cultivars as well as an additional group composed of their hybrids; and two composed of narcotic strains as well as an additional group composed of their hybrids.
The Vavilov Institute (VIR) in Russia maintains the world's largest collection of Cannabis sativa hemp germplasm through periodic seed regeneration. Sample populations of 93 accessions from the VIR collection were assayed for allozyme variation at 17 gene loci by starch-gel electrophoresis. The frequencies of 47 alleles were subjected to principal components (PC) analysis. A scatter plot on the first two PC axes showed little tendency for the accessions to cluster according to country of origin or their assignment to eco-geographical groups. Three Chinese accessions previously assigned to the hemp biotype of C. indica clustered separately. Eleven other Chinese accessions clustered with accessions from Europe, Asia Minor, and central Asia, and are likely to be hybridized. Conservation ofC. indica hemp landraces is of vital importance to the hemp industry.
To determine whether the terpenoid composition of the essential oil of Cannabis is useful for chemotaxonomic discrimination, extracts of pistillate inflorescences of 162 greenhouse-grown plants of diverse origin were analyzed by gas chromatography. Peak area ratios of 48 compounds were subjected to multivariate analysis and the results interpreted with respect to geographic origin and taxonomic affiliation. A canonical analysis in which the plants were pre-assigned to C. sativa or C. indica based on previous genetic, morphological, and chemotaxonomic studies resulted in 91% correct assignment of the plants to their pre-assigned species. A scatterplot on the first two principal component axes shows that plants of accessions from Afghanistan assigned to the wide-leaflet drug biotype (an infraspecific taxon of unspecified rank) of C. indica group apart from the other putative taxa. The essential oil of these plants usually had relatively high ratios of guaiol, isomers of eudesmol, and other unidentified compounds. Plants assigned to the narrow-leaflet drug biotype of C. indica tended to have relatively high ratios of trans-β-farnesene. Cultivars of the two drug biotypes may exhibit distinctive medicinal properties due to significant differences in terpenoid composition.