Systematic Botany (2013), 38(3): pp.

818–829
© Copyright 2013 by the American Society of Plant Taxonomists
DOI 10.1600/036364413X670278

Molecular Phylogeny of the Jimsonweed Genus Datura (Solanaceae)

Robert Bye1,3 and Victoria Sosa2
1
Jardı́n Botánico del Instituto de Biologı́a, Universidad Nacional Autónoma de México, 04510 México,
Distrito Federal, México.
2
Biologı́a Evolutiva, Instituto de Ecologı́a, A. C., Carretera antigua a Coatepec 351, El Haya, 91070 Xalapa,
Veracruz, México.
3
Author for correspondence (rbyeunam@ibunam2.ibiologia.unam.mx)

Communicating Editor: Martin Wojciechowski

Abstract—Phylogenetic analyses were performed based on four plastid DNA sequences to determine if the Daturae and Datura are
monophyletic and to understand their relationships and classification. Species of Datura have been divided into three sections based upon
the position and type of the fruit: Ceratocaulis, Datura, and Dutra. Our analyses confirmed the monophyly of the Daturae clade with
two genera, Datura with herbaceous species and Brugmansia with woody species. The Datura clade consists of two distinct lineages, that
of D. ceratocaula, corresponding to the monospecific section Ceratocaulis, and one that gives rise to the remaining species of the genus.
The large clade readily divides into two groups but they do not correspond to the conventional classification of the genus. Species with
regularly dehiscent capsular fruits are present in both groups while species with irregularly dehiscent capsules are found in only one
group. Because section Dutra is polyphyletic, realignment of species within two traditional sections is necessary as follows: Datura
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19

containing D. arenicola, D. discolor, D. ferox, D. quercifolia, D. kymatocarpa, D. leichhardtii, and D. stramonium; Dutra containing D. inoxia,
D. lanosa, D. metel, D. reburra, and D. wrightii. The seed margin clearly distinguishes these sections, rather than the combination of fruit
position and dehiscence which has been used previously.
Keywords—Brugmansia, Daturae, infrageneric classification, toloache.
Copyright (c) American Society for Plant Taxonomists. All rights reserved.

Datura is one of the most conspicuous genera in the
Solanaceae, with D. wrightii Regel being the largest native
flower in the U. S. A. The attractively large, tubular flowers
have assured that “Angel’s trumpets” or “moon-flowers”
(Datura section Dutra) are cultivated in gardens around the
world. The patterns of morphological and chromosomal
variation of jimsonweed have attracted scientists interested
in species concepts (e.g. Avery et al. 1959) while the intri-
cate associations between insects and Datura have been the
subject of evolutionary ecological studies (e.g. Bello-Bedoy
and Núñez-Farfán 2011). For centuries, Datura species have
been employed by native peoples in southern North America
for their medicinal and psychotropic effects.
Most of the species of Datura have iconic cultural impor-
tance in North America that dates back to prehistoric times.
Archaeological evidence of ceramic pots with echinate orna-
mentation (Litzinger 1981) and rock art (Boyd 2003) attests
to the importance of bioactive constituents of Datura during
the pre-Hispanic period. The delirium induced by the
tropane alkaloids has been the neurophysiological basis of its
employment; more than 30 compounds from the tropane
group have been reported in Datura (Eich 2008). The dis-
orientation caused by the alkaloids and the guidance pro-
vided by elders during initiation ceremonies were critical
components of the passage ritual from adolescence to man-
hood among various indigenous groups in North America
(Safford 1922). The Spaniards interpreted such deliria as the
influence of inebriation allied with the devil (Bye and Linares
2000). Over time, jimsonweeds or thornapples have been the
basis of black magic in Mexico and other parts of the world
(Lewis and Elvin-Lewis 2003). Other names usually applied
to members of the genus Datura are “toloache” and “tlapatl”,
both derived from the Nahuatl language (Safford 1922). The
most commonly employed medicinal species of “toloaches”
that are sold regularly in markets of central Mexico for
treating wounds and inflammations are D. inoxia Mill. and
D. stramonium L. (Argueta Villamar et al. 1994). Poisonings
and deaths due to consumption of the plant occur regularly
(e.g. Russell et al. 2010). The international use of Datura as
818
a recreational drug is growing (Iglesias-Lepine et al. 2012)
and intentional anticholinergic plant poisonings due to its
ingestion in the U. S. A. increased 44% over the last 20 yr
(Vearrier and Greenberg 2010). The domesticated species,
D. metel L., has been used in initiation ceremonies in Africa
and India (Johnston 1972).
The genus Datura consists of herbs native to dry, tem-
perate and subtropical regions (Barclay 1959) although
many have become world-wide weeds (Haegi 1976; Fuentes
1980; Hammer et al. 1983). Lockwood (1973) interpreted
such characteristics as annual or short-lived perennial life
cycle, herbaceous habitat, self-compatibility, and dry, spiny,
dehiscent fruits as adaptations to xeric environments and
placed the center of origin and evolution of Datura in Mexico
and the southwestern United States. Of the 13 species, 12 are
native to Mexico with the greatest species richness occur-
ring in the Balsas Depression biogeographic province (Luna-
Cavazos and Bye 2011). For the purpose of this study, we
consider the taxa reported as endemics elsewhere to be con-
specific with Mexican taxa: 1) the Cuban D. velutinosa Fuentes
with D. inoxia (Hammer et al. 1983; Hokche et al. 2008); and
2) the Australian D. leichhardtii Benth. with D. pruinosa
Greenm. (Hammer et al. 1983; Symon and Haegi 1991).
Recently, the pre-Columbian presence of Datura in Eurasia
has been championed using iconographic, literature, and
linguistic sources. Based upon a revised interpretation of
ancient Latin and Greek texts, Touwaide (1998) argued that
Datura stramonium was known in the Old World prior to
the discovery of the New World in 1492. Using old Arabic
and Indic references as well as iconographic representa-
tions from southern India, Geeta and Gharaibeh (2007)
supported the hypothesis that D. metel was transferred to
the Old World at least a millennium ago. The pre-Columbian
distribution of D. ferox L. is uncertain; it is an infrequent
annual weed along the coasts of the Pacific and Atlantic
Oceans but is most common in drylands of northern
Argentina. The cosmopolitan taxa D. inoxia, D. metel, and
D. stramonium were introduced into other parts of the world
by Europeans (Symon and Haegi 1991).
 2013] BYE AND SOSA: MOLECULAR PHYLOGENY OF DATURA
Conventional taxonomy of Datura has recognized four sec-
tions: Brugmansia, Ceratocaulis, Datura, and Dutra (Bernhardi
1833; Safford 1921; Hammer et al. 1983). The first Datura
species published by Linnaeus (1753) were D. arborea L.,
D. metel, and D. stramonium; afterwards he added D. fastuosa
L., D. ferox, and D. tatula L. (Hadkins et al. 1997). Persoon
(1805) removed the woody daturas from Datura and raised
them to the genus Brugmansia. Later Bernhardi (1833) grouped
the nine published species of Datura and the three species
of Brugmansia among four sections: Brugmansia, Ceratocaulis,
Dutra, and Stramonium (recognized today as Datura). This
classification with few adjustments has been followed in
major taxonomic treatments (Dunal 1852; Safford 1921;
Satina and Avery 1959; Hammer et al. 1983) and floristic
studies in such regions as Europe, Cuba, Bahamas, Mexico,
Guatemala, Costa Rica, and Peru (Marzell et al. 1975;
Amshoff 1957; Correll and Correll 1982; Matuda 1952;
Gentry and Standley 1974; Standley and Morton 1938; and
Macbride 1962, respectively). Taxonomic revisions using
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
various data sets emphasized the need for realignment
of the sections in Datura. Certain morphological features
(e.g. woody habit, pendulous flowers, and elongated spine-
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
less fruits) and infertility of crosses between woody and
herbaceous daturas prompted Lockwood (1973) and Nee
(1986) to accept the generic recognition of Brugmansia.
Persson et al. (1999) separated Brugmansia in a group apart
from the other sections of Datura using features of the pollen.
Floristic treatments of Solanaceae have recognized sub-
sequently these two genera (e.g. D’Arcy 1973, 1993; Haegi
1976; Liberty Hyde Bailey Hortorium 1976; Hepper 1985;
Huxley et al. 1999a and 1999b; Shaw 2000; Hunziker
2001). Another section, the monotypic Ceratocaulis, was
raised to generic status under the names Apemon Raf. and
Ceratocaulos Spach during the 1830s although later authors
did not follow this separation (e.g. Dunal 1852). Because
D. ceratocaula Ortega shares various morphological features
with both Datura and Brugmansia, Safford (1921) regarded it
as “connecting” the two genera.
Contemporary taxonomy based upon morphological char-
acters recognizes three sections in Datura. Fruit characters
(e.g. pericarp ornamentation, position, dehiscence pattern,
and seed form) are diagnostic features used to separate these
sections. The fruit of the section Ceratocaulis is a spineless
pendant foraminicidal capsule (capsular fruits with a fleshy
pericarp that disintegrates irregularly; Knapp 2002) (Fig. 1
A–B). The nine species of section Dutra have spiny pendant
fruits. Fruits of four species are foraminicidal capsules
(Fig. 1 C–D) while those of the remaining species are
septifragal capsules (which dehisce upon drying into four
regular valves) (Fig. 1 E–G). All three species of section
Datura have spiny erect capsules (Fig. 1 H–K). In contrast
to the capsular fruits of Datura, the fruit of Brugmansia
is a berry (Fig. 1 L). The capsule is considered the
plesiomorphic state in the Solanaceae (Knapp 2002) and
its occurrence in D. stramonium represents a reversion
(Pabón-Mora and Litt 2011). To a lesser degree, the two seed
forms found in Datura (Gunn 1974) are partially correlated
with the sections of the genus. The C-shaped seed with a
triple ridge on the convex margin is found in some members
of the Dutra (i.e. D. inxoia, D. lanosa, D. metel, D. reburra, and
D. wrightii) while the seeds of other taxa of Dutra as well as
those of sections Ceratocaulis and Datura are C- or D-shaped
without the rim.
819
Allozymes were the first macromolecular markers uti-
lized to resolve the species relationships of Datura. Based
upon ten herbaceous species of Datura derived from garden
populations, Conklin and Smith (1971) subdivided the
classical three sections into five groups. The protein chemo-
taxonomy based upon amino acid sequences of ferredoxin
for seven cultivated species was interpreted as supporting
the conventional classification that included tree daturas
(Brugmansia) within the genus Datura (Mino 1994, 1995).
A study involving nuclear ITS-1 sequences, isozymes and
morphology, carried out on 12 species of garden-derived
Datura and Brugmansia (Mace et al. 1999) resulted: 1) in the
recognition of the arborescent species as a separate genus,
Brugmansia, apart from the herbaceous species in Datura,
and 2) in the placement of D. discolor in a section separate
from Datura and Dutra, even though no formal taxonomic
description was made. Jiao et al. (2002) using allozyme loci
of wild species of Datura demonstrated that members of
section Dutra were divided into two groups.
Previous phylogenetic analyses of the genus have been
limited to specific morphological sets of characters or to
a few species within the genus Datura. Even though the
phylogenetic analysis of pollen characters of eight species
of Datura and five of Brugmansia clearly separated these
two genera and placed the “putative linking taxon”,
D. ceratocaula, within the Datura clade, the species relation-
ships within the genus Datura were unresolved (Persson
et al. 1999). A cladistic analysis of-morphological characters
ranging from the whole plant to the seed for eight species in
section Dutra separated this section into two distinct clades
(Luna-Cavazos et al. 2009).
Few Datura species have been included in molecular
phylogenetic studies of Solanaceae. Olmstead et al. (1999)
used plastid molecular markers to demonstrate the mono-
phyly of the Daturae clade based upon only D. stramonium and
Brugmansia sp. in their analysis. Their later study (Olmstead
et al. 2008) with ndhF and trnL-F from D. stramonium,
D. leichhardtii, B. aurea, and B. sanguinea confirmed the
Daturae clade which consisted of Brugmansia, Datura and
Iochroma cardenasianum.
The objectives of this paper are to elucidate phyloge-
netic relationships among all the Datura species based
on four plastid DNA sequences and examine previous
infrageneric classification. The diagnostic morphological char-
acters of the fruit are discussed in the light of these results.
Materials and Methods
Taxon Sampling—A total of 25 terminal taxa were considered in the
analyses, 13 as the ingroup and 12 as the outgroup. Various accessions
were sequenced from different populations for every species in Datura
(included in Appendix 1). The ingroup included all species recognized
in Datura. Based on the results of Olmstead et al. (2008), the outgroup
taxa were representative of Nicotianoideae and Solanoideae clades. As
part of the outgroup we considered Brugmansia suaveolens (Humb. &
Bonpl. ex Willd.) Bercht. & C. Presl. and two morphological forms of
Brugmansia candida Pers., “pañuelo” (p) and “sencilla” (s) distinguished
by divided and tubular corolla, respectively. Nicotiana tabacum L. was
set as the functional outgroup (Appendix 1).
DNA Sequencing—DNA was isolated using either the modified
2x CTAB method (Rogers and Bendich 1985; Doyle and Doyle 1987)
or the DNeasy plant minikit (Quiagen, Valencia, California), following
the manufacturer’s instructions. Primers utilized for amplification of
matK were 1R_KIM and 3F_KIM following protocols of Dunning and
Savolainen (2010). Amplification of the ndhF region was carried out
using primers 32F-1101R and 1101F-2110R and sequenced following
 820 SYSTEMATIC BOTANY [Volume 38
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
Copyright (c) American Society for Plant Taxonomists. All rights reserved.

Fig. 1. Characteristic fruits of the conventional sections of Datura and of the genus Brugmansia. A–B. Section Ceratocaulis. C–G. Section Dutra
with pendant spiny septifragal and foraminicidal capsules. H–K. Section Datura with erect septifragal capsule dehiscing in four values. L. Brugmansia
with berry. A. D. ceratocaula with foraminicidal capsule prior to seed dispersal. B. D. ceratocaula with foraminicidal capsule after seed dispersal.
C. D. inoxia with foraminicidal capsule prior to seed dispersal. D. D. inoxia with foraminicidal capsule after seed dispersal. E. D. reburra with
septifragal capsule prior to seed dispersal. F. D. reburra with septifragal capsule dehiscing in four values for seed dispersal. G. D. pruinosa with
septifragal capsule dehiscing in four values for seed dispersal. H. D. stramonium with capsule prior to seed dispersal. I. D. stramonium with septifragal
capsule after seed dispersal. J. D. quercifolia with septifragal capsule prior to seed dispersal. K. D. quercifolia with septifragal capsule after seed dispersal.
L. B. sanguinea with berry.
 2013] BYE AND SOSA: MOLECULAR PHYLOGENY OF DATURA
protocols of Terry et al. (1997). The trnH-psbA region was amplified
and sequenced using primers trnH2 (Tate and Simpson 2003) and psbA
(Sang et al. 1997) based on protocols of Shaw et al. (2005). Amplifica-
tion of the trnL-F region was carried out using primer “C” and “F”
and sequenced based on the protocols of Taberlet et al. (1991). Ampli-
fied double-stranded DNA fragments were purified using QIAquick
columns (Qiagen) following the protocols provided by the manufacturer
and were sequenced at a commercial sequencing facility (Macrogen,
Inc., Seoul, Korea). Electropherograms were edited and assembled using
Sequencher 4.1 (Gene Codes, Ann. Arbor, Michigan). Sequences were
aligned manually using the program Se-Al v. 2.0a11 (Rambaut 2002);
they were deposited at GenBank (accession numbers are provided in
Appendix 1). Data matrices and trees were deposited in TreeBASE
(accession number 12499).
Phylogenetic Analyses—Parsimony and Bayesian inference was con-
ducted using TNT (Goloboff et al. 2003) and with MrBayes 3.2.1
(Huelsenbeck and Ronquist 2001; Ronquist and Huelsenbeck 2003),
respectively. Parsimony analyses were performed using a new tech-
nology search approach: the ratchet algorithm with 500 iterations.
Parsimony bootstrapping support for internal branches was estimated
with one thousand replicates using TBR branch swapping, with 100 itera-
tions saving one tree per replicate. With jModelTest.0.1.1 (Posada 2008)
the model of evolution was identified using the AICc selection criteria
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
and default search values for each one of the four plastid markers
(matK = TPM1uf, ndhF = TVM + 1, psbA-trnH = F81 + G, trnL-F =
TVM + g). The Bayesian Markov chain Monte Carlo (BMCMC) esti-
mate of phylogeny was inferred with default priors. Two indepen-
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
dent runs were carried out to assess the repeatability of stationarity.
For each run, one cold and three heated chains were set for
50,000,000 generations, sampling one tree every 1,000 generations.
Stationarity of the MCMC runs was based on convergence to a stable
value of the log likelihood score in separate runs, and trees sampled
prior to stationarity were excluded by “burnin” (20% of samples).
Fig. 2. Single most parsimonious tree (L= 698 steps, CI = 0.91, RI = 0.89) based on the combined molecular dataset. Bayesian topology of the
50% consensus tree after burn-in was similar and posterior probabilities are indicated below branches. Bootstrap support is indicated above branches.
821
Posterior probabilities for supported clades were determined by a 50%
majority-rule consensus of the trees retained after burn-in.
To understand the evolution of the diagnostic fruit and seed characters
(fruit position: pendant/erect; fruit type: berry/septifragal capsule/
foraminicidal capsule; convex seed margin with triple ridge: absent/
present), these features were optimized onto the resulting parsimony
tree using the program MacClade v.4.5 (Maddison and Maddison 2011)
with unordered characters and Fitch parsimony.
Results
The total length of the aligned sequences is 5,713 bases:
859 for matK, 2,128 for ndhF, 600 for trnH-psbA, and 2,126
for trnL-F. In the combined data matrix, 167 of 5,713 nucleo-
tide characters were parsimony informative and 589 vari-
able but parsimony uninformative. Parsimony ratchet search
retrieved a single most parsimonious tree (L = 696 steps,
CI = 0.91, RI = 0.89). Bootstrap (BS) values on the result-
ing tree ranged from low (50–74%), moderate (75 –84%)
and high (85 –100%). Parsimony and Bayesian analyses
retrieved similar topologies; Fig. 2 synthesizes results of
both analyses. Posterior probability (PP ³ 85%) from Bayesian
inference and bootstrap percentages from parsimony ana-
lyses are indicated. Datura species were retrieved in a well-
supported clade (BS = 99% and PP = 1), sister to the three
samples of Brugmansia. This inclusive clade received also
strong support (BS = 99% and PP = 1). Datura ceratocaula
is supported as the sister taxon to the rest of Datura species
 822 SYSTEMATIC BOTANY [Volume 38
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
Copyright (c) American Society for Plant Taxonomists. All rights reserved.

Fig. 3. Fruit and seed evolution in Datura. Three morphological characters (fruit position, fruit type and seed margin) were optimized onto the
resulting parsimony and Bayesian tree.
 2013] BYE AND SOSA: MOLECULAR PHYLOGENY OF DATURA 823
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
Copyright (c) American Society for Plant Taxonomists. All rights reserved.

Fig. 4. Seeds of Brugmanisa suaveolens and Datura species arranged according to their respective clades. A. B. suaveolens. B. D. ceratocaula. C. D. lanosa.
D. D. wrightii. E. D. metel. F. D. reburra. G. D. inoxia. H. D. ferox. I. D. quercifolia. J. D. stramonium. K. D. leichhardtii subsp. pruinosa. L. D. leichhardtii
subsp. leichhardtii. M. D. kymatocarpa. N. D. discolor. O. D. arenicola.

(BS = 62%, PP = 0.99). Datura inoxia, D. lanosa, D.metel,
D. reburra, and D. wrightii are in a well-supported clade
and within this group two pairs of sister species formed
well-supported groups: 1) D. lanosa and D. wrightii, and
2) D. metel and D. reburra. The group formed by the rest of
the species received moderate support (BS = 70%, PP = 0.99),
with D. arenicola as the sister taxon to the rest of species in
this group. Datura ferox, D. quercifolia, and D. stramonium were
in a small well-supported clade (BS = 82%, PP = 1); D. ferox
and D. quercifolia were sister species (BS = 82%, PP = 1).
 824 SYSTEMATIC BOTANY
With regard to fruit evolution there is a shift in position
from pendant to erect (Fig. 3A); this change occurred in
the clade formed by Datura stramonium, D. ferox, and
D. quercifolia. Fruit type changed from foraminicidal capsule
to septifragal capsule within Datura, while D. reburra is the
only species in its clade with a septifragal capsule (Fig. 3B).
Datura ceratocaula has seeds without a convex margin triple-
ridge (Figs. 3C, 4B). This character changed in the group
formed by D. lanosa, D. wrightii, D. metel, D. reburra, and
D. inoxia which possess seeds with a convex margin with a
triple-ridge (Figs. 3C, 4C–G). The species with seeds lacking
this convex margin were retained in the clade including
the rest of Datura taxa (Figs. 3C, 4H–O).
Discussion
Our results confirm the previous recognition of Brugmansia
as sister of Datura in the subtribe Datureae (Olmstead et al.
2008). Our analyses also support the recognition of Datura
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
as monophyletic, with the monospecific section Ceratocaulis
(represented by D. ceratocaula) as the sister species to the
rest of Datura. Our results indicate that section Dutra is
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
polyphyletic, thus the contemporary taxonomy of the sec-
tions Datura and Dutra requires reconsideration. The sec-
tion Datura as proposed by Bernhardi (1833; as section
Stramonium) now consists of: D. arenicola, D. discolor, D. ferox,
D. kymatocarpa, D. pruinosa, D. quercifolia, and D. stramonium.
The section Dutra is restricted to D. inoxia, D. lanosa,
D. metel, D. reburra, and D. wrightii. The ingroup clearly
depicts the Daturae clade with two subgroups: the two rep-
resentative species of Brugmansia and the 13 species of
Datura. Previous studies oriented at resolving intrafamilial
relationships (Olmstead et al. 2008) incorporated only
two species for each genus such that the divergence between
the genera was not clear nor was any subgeneric segrega-
tion obvious. The central focus of this study is to under-
stand the infrageneric classification of the species of Datura
based on groups retrieved by phylogenetic analyses. Previous
studies based on morphometrics (Luna-Cavazos et al. 2000)
and isozymes (Jiao et al. 2002) suggested the division of sec-
tion Dutra and the recognition of an additional section within
the genus Datura. Our results support two distinct lineages:
D. ceratocaula and a large clade formed by the rest of the
species and within this latter large clade two well-supported
groups. A clade with D. inoxia, D. lanosa, D. metel, D. reburra,
and D. wrightii constitutes section Dutra. The remaining
species of the conventional section Dutra (D. discolor,
D. kymatocarpa and D. leichhardtii) are now included in
another group along with members of the section Datura.
Hence, redefining the conventional section Dutra by the
members of this clade is justified by molecular data. How-
ever, no new section is needed for the removed species
because they are included in the other clade along with
members of section Datura. Consequently, the traditional
three infrageneric divisions of the genus Datura are retained.
Our studies indicate that the initial proposal for the
subgeneric division of Datura presented by Bernhardi (1833)
is appropriate for the herbaceous species. Our results cor-
roborate the earlier studies on the separation of the woody
Brugmansia from the genus Datura based upon morpho-
logical and hybridization studies. Previous molecular
studies suggested that Datura and Brugmansia were mem-
bers of separate clades but only three species of Datura
[Volume 38
were sequenced. Mace et al. (1999) performed cluster ana-
lyses of molecular data for seven species of Datura and
six species of Brugmansia and considered these genera
as sister groups. Within the Datura group, four clusters
were recognized. Apart from the three conventional sec-
tions, they accepted a fourth cluster that consisted of
D. inoxia, D. discolor and an unidentified species. The
inclusion of D. inoxia in the new section suggested that
section Discolor was polyphyletic because it was also
present in the Dutra cluster. The ancestries of the spe-
cies studied were uncertain because they were derived
from garden plants and may have been products of pre-
vious hybridization events.
Datura ceratocaula has been placed in its monotypic genus
(Apemon Raf., Ceratocaulos (Bernh.) Rchb., Ceratocaulos (Bernh.)
Spach) as well as transferred to Brugmansia (i.e. B. ceratocaula
(Ort.) Court ex Gaede). Bernhardi (1833) and Safford (1921)
maintained that D. ceratocaula was the “connecting link”
between Brugmansia and Datura based upon the features
of the flowers and fruit (Lockwood 1973). Even though
the spineless, pendant berry-like fruit and spathe-like
calyx may be similar to those of species of Brugmansia,
D. ceratocaula shares morphological features with those of
Datura such as herbaceous habit, dichasial inflorescence,
erect flower, circumscissile calyx, tetralocular fruit, and
small seed with caruncle. Our molecular data clearly
show that D. ceratocaula is a member of the genus Datura
rather than Brugmansia.
The inclusion of Datura discolor in section Dutra ques-
tioned the strength of the differences between section Datura
(cited as section Stramonium) and section Dutra according to
Fosberg (1959). He further stated that although the differen-
tiation between these two sections was weak, the mainte-
nance of the sections by Blakeslee and his associates (Avery
et al. 1959) was adequate for their genetic and cytological
work on the genus. Barclay (1959) indicated that the sec-
tional position of D. discolor was dubious and required fur-
ther work. Although our study returns D. discolor to section
Datura (formerly known as Stramonium) as Bernhardi (1833)
and Dunal (1852) originally classified it, the section Dutra
still is not marked by unique morphological features useful
in contemporary taxonomies. For instance, D. reburra with
regularly dehiscing septifragal capsules is in the clade that
also includes D. inoxia, D. lanosa, and D. wrightii with irregu-
larly dehiscing foraminicidal capsule.
Datura reburra, an endemic species of northwestern
Mexican Pacific coastal plains, was described by Barclay
(1959) from a single herbarium collection. He remarked on
its similarity to D. discolor and questioned the validity of the
sections Stramonium and Dutra without resolving the incon-
sistencies posed by this new taxon. The pendant fruit
with four well defined valves placed it in the section Dutra
in association with D. discolor rather than with D. inoxia
(Luna-Cavazos et al. 2000). Even though D. reburra does
not share the irregularly dehiscent fruits of other members
of its clade, its seed with a triple-ridge along the convex
margin is more similar to seeds of other members of the
D. lanosa clade than to those of the other group.
The fruit has been the critical structure in the conven-
tional classification of taxa within the genus Datura. How-
ever, our study indicates that any particular combination
of traditional diagnostic fruit characters is not exclusive
to each clade. Hence, there are discrepancies in species
 2013] BYE AND SOSA: MOLECULAR PHYLOGENY OF DATURA
composition in the sections of Datura among taxonomic
treatments. Foraminicidal capsules are found in D. ceratocaula
and in the D. lanosa clade (except D. reburra) and in none
of the species of the other clade. A particular fruit position
is not shared by all species within the D. arenicola clade
which has species with erect fruits (D. ferox, D. quercifolia,
and D. stramonium) as well as pendant fruits (D. arenicola,
D. discolor, D. kymatocarpa, and D. leinhhardtii subsp. pruinosa).
Hence, these fruit characters are limited in their usefulness
for the subgeneric classification of Datura. Similar discrep-
ancies were found in Solanum (Chiarini and Barboza 2007)
where closely related species in molecular phylogenies dif-
fered regarding fruit types.
The most recent monograph of Datura (Hammer et al.
1983) was based upon the material deposited in the German
seed bank. Two species endemic to western Mexico,
D. kymatocarpa and D. reburra, were placed in synonymy
with D. discolor. This situation reflected, in part, the con-
fusion surrounding the characterization of D. discolor
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
which is geographically widespread and highly variable
in its morphological expressions. Our molecular data sup-
port the recognition of these two species.
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
The first infrageneric classification of the genus Datura
presented by Bernhardi (1833) established four sections:
Stramonium (= Datura), Dutra, Ceratocaulis, and Brugmansia.
This classification was modified by Safford (1921) with the
shift of D. discolor from section Stramonium to section Dutra
based upon its nodding fruit. Since then, his concept of
sections for herbaceous species has been the basis of mono-
graphic and floristic treatments worldwide. During the last
fifty years, various authors (Jiao et al. 2002; Luna-Cavazos
et al. 2000; Mace et al. 1999; Persson et al. 1999) have ques-
tioned the taxonomic consistency of these sections using
macro- and micro-morphological characters and genetic
markers. Our study is the first to address the validity of
the sections using the wild species from the geographic
center of diversity of the genus. Datura is monophyletic
and has as its sister the genus Brugmansia. Datura is readily
divided into three sections that are supported by plastid
DNA data: Ceratocaulis, Datura, and Dutra. The monotypic
section Ceratocaulis, with the Mexican species, D. ceratocaula,
is sister to the remainder species of Datura. The clade
consisting of the redefined section Dutra with species
having pendant fruits; those with foraminicidal capsules
include D. inoxia, D. lanosa, D. metel, and D. wrightii while
one has regularly dehiscing capsules (D. reburra). The rest
of Datura species grouped in another clade corresponds
to section Datura which includes species with regularly
dehiscing septifragal capsules but with different fruit posi-
tions; those with pendant fruits are D. arenicola, D. discolor,
D. kymatocarpa, and D. leichhardtii and those with erect
fruits include D. ferox, D. quercifolia, and D. stramonium.
Our revision reincorporates D. discolor in this section where
Bernhardi originally placed it. Section Dutra consists of spe-
cies with pendant fruits and reniform seeds with triple-
ridged convex margin while those of section Datura lack
this feature. Phylogenetic analysis of plastid DNA data does
not support fruit position and pericarp dehiscence patterns
as diagnostic morphological characters for distinguishing
the sections. The presence or absence of the triple ridge
on the convex margin coincide with our molecular phy-
logeny and can differentiate between the sections Datura
and Dutra.
825
Taxonomic Treatment
BRUGMANSIA Pers., Syn. Pl. 1: 216. 1805. Datura sect.
Brugmansia (Pers.) Bernh., Neues J. Pharm. Aerzte 26:
120. 1833; Linnaea 8 (Litteratur-Bericht): 116. 1833.—
TYPE: Brugmansia candida Pers. Syn. Pl. 1: 216. 1805.
DATURA L., Sp. Pl. 1: 179. 1753. Stramonium [Tourn.] Scop.,
Fl. carniol., ed. 2, 1: 157. 1772. Ceratocaulos (Bernh.)
Rchb., Handb. Nat. Pfl.-Syst. 201. 1837. Apemon Raf., Fl.
tellur. 2: 11. 1837. Ceratocaulos (Bernh.) Rchb., Handb.
Nat. Pfl.-Syst. 201. 1837. Ceratocaulos (Bernh.) Spach,
Hist. Nat. Vég. 9: 68–69. 1840.—TYPE: Datura stramonium
L. Sp. Pl. 1: 179. 1753.
DATURA sect. CERATOCAULIS Bernh., Neues J. Pharm. Aerzte
26: 145. 1833; Linnaea 8 (Litteratur-Bericht): 135. 1833.—
TYPE: Datura ceratocaula Ortega Nov. Rar. Pl. Descr.
Dec. 11. 1797.
DATURA CERATOCAULA Ortega, Nov. Rar. Pl. Descr. Dec. 11.
1797. Brugmansia ceratocaula (Ortega) Dumort., Nouv.
Mém. de l’Acad. Roy. de Bruxelles 9: 134, t. 1,
f. 10. 1835. Brugmansia ceratocaula (Ortega) Court ex
Gaede, Ind. Pl. Hort. Bot. Leod. 29. 1828.—TYPE: SPAIN.
Real Jardı́n Botánico Madrid, without date, Ortega s. n.
(holotype: MA).
Datura macrocaulos Roth, Neue Beytr. Bot. 159. 1802.—TYPE:
lectotypification required.
Datura sinuata Sessé & Moc., Pl. nov. Hisp. 25. 1888; Fl. mexic.
40. 1893.—TYPE: MEXICO. Queretaro, near Queretaro,
1787–1804, Sessé & Mociño 5335 (lectotype: MA)
DATURA sect. DATURA. Stramonium Bernh., Neues J. Pharm.
Aerzte 26: 126. 1833; Linnaea 8 (Litteratur-Bericht): 121.
1833.—TYPE: Datura stramonium L. Sp. Pl. 1: 179. 1753.
Datura arenicola Gentry ex Bye & Luna, sp. nov.—TYPE:
MEXICO. Baja California Sur, ca. 30 miles south of
El Arco, Viscaino Desert area, lee side of sand dunes;
corolla lavender; eaten by large moth larvae, 17 Novem-
ber 1947 (fl., fr.) Howard Scott Gentry 7881 (holotype:
ARIZ 274295!; isotypes: ARIZ!, MICH!, SD!, UC!, US!).
Annual herb, first dichotomous branches prostrate, sec-
ondary branches prostrate to decumbent, 15–20 (–40) cm
tall, 25–40 (–120) cm wide, stems tomentose. Leaves simple,
alternate; blade ovate, 3.6–5 cm long, 2.4–4.2 cm wide, width
usually two-thirds the length, perimeter/silhouette ratio
1.01 –1.06, tomentose on both sides of blade and along
petiole, apex acute to obtuse, margin entire to shallowly
undulate lobed, unequal base; petiole 1.4–3.3 cm long.
Flower erect, borne in branch axil, single night blooming;
peduncle tomentose, 2–4 mm long, reflexed after anthesis;
calyx synsepalous, prismatic, 1.9–2.9 cm long, 4–6 mm
wide, apical teeth 5, lobe apex acuminate, lobes 2.4–6.2 mm
long, 1.8–2.6 mm wide at base, tomentose; corolla symptalous,
white to lavendar, funnelform, plicate, glabrous, 2.9–3.9 cm
long; acumenal lobes 5, each supported by 3 nerves,
2–6.1 mm long, 1.4–3.7 mm wide at base; stamens 5, adnate
at base to corolla tube, 1.2–1.6 cm long, free filament
1.1–1.3 cm long, anther 1.7–2.9 mm long; ovary with
minute spines, style 1.1–2.2 cm long, stigma 1.1–1.9 mm
long. Fruit pendant capsule dehiscing into four regular
valves, tetralocular due to septum and pseudoseptum,
 826 SYSTEMATIC BOTANY [Volume 38
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
Copyright (c) American Society for Plant Taxonomists. All rights reserved.

Fig. 5. Datura arenicola. A. Holotype: Gentry 7881 (ARIZ). B. Fruit and leaves (Bye 36326 & Linares). C. Lateral view of plant growing under
irrigation; note that the plant retains its prostrate habit; scale of 20 cm, marked in increments of 1 cm (Bye 36327 & Linares).

ovoid, 2.0–3.1 cm long, 1.8–2.1 cm wide; peducle reflexed, all-terrain vehicles have destroyed large tracts of its habitat
1.1–2.0 cm long; calyx base circumscissile, slightly accrescent by flattening the dunes within which it grows. None the
marginal frill, with 5 distinct ridges extending from peduncle; less, it appears to respond to human-caused disturbance by
pericarp puberulent, with stiff puberulent spines 4.6–6.1 mm invading cultivated fields and citrus orchards.
long. Seed reniform, black at maturity, with white caruncula, Conservation Status—Datura arenicola is endangered accord-
testa black at maturity, rugose, 3.2–4.2 cm long, 2.5–3.2 cm ing to the IUCN (2001) red list criteria in that its current
wide. Flowering November through January. Figure 5. geographic range is less than 20,000 km2 and it is known
Etymology—The specific epithet refers to the species’ from less than 10 populations. It has been seen only in the
association with sand. eastern margin of the Vizcaino Biosphere Reserve located
Distribution and Habitat—Originally named (but not in the central portion of the Baja California peninsula.
published) by H. S. Gentry, this taxon has been considered Additional Specimens Examined—MEXICO. Baja California Sur:
Municipio Mulegé, Vizcaino, 23 Jan 2010, R. Bye 36323, E. Linares &
a deformed specimen of D. discolor with which it is sym-
R. Watson (MEXU); 24 Jan 2010, R. Bye 36326 & E: Linares (MEXU),
patric. However, examination of living plants in dune fields R. Bye 36327 & E. Linares (MEXU).
as well as in irrigated orchards reveals that its charac-
teristic prostrate habit readily differentiates it from the
Table 1. Diagnostic characters of Datura arenicola compared to the
erect D. discolor (Table 1). Because of its primary branches
closely related D. discolor, with which it is sympatric.
are parallel the ground surface, the plant usually accumu-
lates blowing sand and debris to the point that it is Characters D. arenicola D. discolor
buried so that only the terminal portion of the branches
Habit Prostrate Erect
are exposed. Associated species growing on those dunes Corolla length Less than 4 cm Greater than 8 cm
include Mesembryanthemum crystallinum L. and Abronia Purple ring in throat of corolla Absent Present
gracialis Benth. This species is limited to the eastern margin Interacumen lobe of corolla Smaller Larger
of the Vizcaino Desert where it grows along arroyos, in relative to acumen lobe
Pubescence Tomentose Slightly pubescent
open dune fields as well as in agricultural areas. The
Leaf margin Undulate Dentate
expansion of hydroponic greenhouses and the forays of
 2013] BYE AND SOSA: MOLECULAR PHYLOGENY OF DATURA
DATURA DISCOLOR Bernh., Neues J. Pharm. Aerzte 26: 149. 1833;
Linnaea 8 (Litteratur-Bericht): 138. 1833.—LECTOTYPE:
without locality, date, or collector, Hermann, Parad. bat.
Stramonia corassavica humilior Hyoscyami folio Hermanni,
pp. 233–234, pl. 1689.
Datura thomasii Torr., Pacif. Railr. Rep. 5(2): 362–363. 1856.—
TYPE: U. S. A. California, Fort Yuma, without date,
Thomas s.n. (holotype: NY)
DATURA FEROX L., Amoen. Acad. 3: 403. 1756.—LECTOTYPE:
without locality, date, or collector, Boccone, Ic. Descr.
Rar. Pl. Stramonium ferox, t. 26. 1674.
DATURA KYMATOCARPA A. S. Barclay, Bot. Mus. Leafl. 18:
256–258, pl. 53 (lower), 54 (upper). 1959.—TYPE:
MEXICO. Estado de Mexico, Temascaltepec, 1935,
Hinton et al. 8173 (holotype: US).
DATURA LEICHHARDTII F. Muell. ex Benth. & F. Muell., Fl. Austral.
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
4: 468–469. 1868.—TYPE: AUSTRALIA. Queensland,
Suttor River, without date, D’Orsay s. n. (lectotype: K).
DATURA LEICHHARDII subsp. LEICHHARDII
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
DATURA LEICHHARDII subsp. PRUINOSA (Greenm.) A. S. Barclay
ex K. Hammer, Kulturpflanze 31: 30. 1983. Datura
pruinosa Greenm., Proc. Amer. Acad. Arts 33: 486.
1898.—TYPE: MEXICO. Oaxaca, Cuicatlan, 1895, Smith
943 (holotype: GH).
DATURA QUERCIFOLIA Kunth, Nov. gen. sp. (quarto ed.) 3:
7. 1818.—TYPE: MEXICO. Guanajuato, Zelaya, 1803,
Humboldt & Bonpland s. n. (holotype: P).
Datura villosa Fernald, Proc. Amer. Acad. Arts 35: 571.
1900.— TYPE: MEXICO. Jalisco, Bolaños, 1897, Rose
3680 (holotype: GH; isotype: US).
DATURA STRAMONIUM L., Sp. pl. 1: 179. 1753.—TYPE: without
locality or collector, specimen in Herb. Clifford 55,
Datura I (lectotype: BM).
DATURA STRAMONIUM f. STRAMONIUM
D. muricata Link, Enum. Hort. Berol. Alt. 1: 177. 1821.—TYPE:
lectotypification required.
D. bertolonii Parl. ex Guss., Fl. Sicul. Syn. 1: 267. 1842.—TYPE:
lectotypification required.
D. wallichii Dunal, Prodr. 13(1): 539. 1852.—TYPE: lecto-
typification required.
D. bernhardii Lundstr., Acta Horti Bergiana 5(3): 90–91, bild
40, tafel V-1. 1914.—TYPE: lectotypification required.
DATURA STRAMONIUM f. INERMIS (Juss. ex Jacq.) Hupke, Repert.
Spec. Nov. Regni Veg. Beih. 101: 135. 1938. D. inermis
Juss. ex Jacq., Hort. bot. vindob. 3: 44, pl. 82. 1776.
D. stramonium var. inermis (Juss. ex Jacq.) Fernald,
Rhodora 40: 184. 1938.—TYPE: lectotypification required.
DATURA STRAMONIUM f. TATULA (L.) B. Boivin, Naturaliste
Canad. 93: 1060. 1967. D. tatula L., Sp. pl., ed. 2, 1: 256.
1762. D. stramonium var. tatula (L.) Torr., Fl. n. middle
United States 232. 1824. D. stramonium var. tatula (L.)
Desc. ex Dunal, Prodr. 13(1): 540. 1852.—TYPE: without
locality, date or collector (lectotype: LINN 243.2).
827
DATURA sect. DUTRA Bernh., Neues J. Pharm. Aerzte 26: 134.
1833; Linnaea 8 (Litteratur-Bericht): 127. 1833.—TYPE:
Datura metel L., Sp. Pl. 1: 179. 1753.
DATURA INOXIA Mill., Gard. dict., ed. 8, no. 5. 1768. D. metel
sensu Sims, Bot. Mag. 35: plate 1440. 1812.—TYPE:
UNITED KINGDOM. Chelsea Physic Garden, without
date or collector, no. 1843 (neotype: BM).
D. guayaquilensis Kunth, Nov. gen. sp. (quarto ed.) 3: 8.
1818. D. meteloides sensu Dunal, Prodr. 13(1): 544.
1852.—TYPE: ECUADOR. Guayaquil, 1803, Humboldt &
Bonpland s. n. (holotype: P).
D. velutinosa V. R. Fuentes, Revista Jard. Bot. Nac. Univ.
Habana 1(2–3): 53. 1980.— TYPE: CUBA. La Habana,
San Antonio de los Baños, 1979, Fuentes 40646 (holo-
type: HAJB).
DATURA LANOSA A. S. Barclay ex Bye, Phytologia 61: 204.
1986.—TYPE: MEXICO. Sinaloa, Culiacan, 1944, Gentry
7052 (holotype: GH).
DATURA METEL L., Sp. Pl. 1: 179. 1753. —TYPE: without
locality or collector, Herb. Clifford 55, Datura 2a (lecto-
type: BM).
DATURA METEL var. METEL.
D. alba Rumph. ex Nees, Trans. Linn. Soc. London 17:
73. 1837. D. fastuosa L. var. alba (Rumph. ex Nees)
Clarke, Fl. Brit. India 4: 243. 1885.—TYPE: lectotypi-
fication required.
D. hummatu Bernh., Neues J. Pharm. Aerzte 26: 153. 1833;
Linnaea 8 (Litteratur-Bericht): 141. 1833.—TYPE: lecto-
typification required.
D. bojeri Delile, Sem. ann. 1836 Hort. Bot. Reg. Monspel. 23;
Annls. Sci. nat. 2(7): 286. 1837.—TYPE: lectotypification
required.
D. nilhummatu Dunal, Prodr. 13(1): 542. 1852.—TYPE: lecto-
typification required.
D. cornucopaea Wilks, Garden 46: 224–225, pl. 978. 1894.—
TYPE: lectotypification required.
DATURA METEL var. FASTUOSA (Bernh.) Danert, Pharmazie 9:
358. 1954. D. fastuosa L., Syst. nat., ed. 10, 2: 932 p.p.
1759.—TYPE: without locality, date, or collector
(lectotype: LINN 243.3 left-hand specimen).
DATURA METEL var. OBSCURA Danert, Pharmazie 9: 357. 1954.
D. fastuosa L. var. ᾳ L., Sp. pl., ed. 2, 1: 256. 1762.—TYPE:
lectotypification required.
D. hummatu Bernh.var. rubra Bernh. f. b Bernh., Neues J.
Pharm. Aerzte 26: 155. 1833; Linnaea 8 (Litteratur-
Bericht): 142. 1833.—TYPE: lectotypification required.
D. nilmummatu Dunal, Prodr. 13(1): 542, p.p. 1852.—TYPE:
lectotypification required.
DATURA METEL var. RUBRA (Bernh.) Danert, Pharmazie 9: 357.
1954. D. hummatu Bernh.var. rubra Bernh., Neues
J. Pharm. Aerzte 26: 155. 1833; Linnaea 8 (Litteratur-
Bericht): 142. 1833.—TYPE: lectotypification required.
D. hummatu Bernh.var. dubia Bernh., Neues J. Pharm.
Aerzte 26: 155. 1833; Linnaea 8 (Litteratur-Bericht):
141. 1833.—TYPE: lectotypification required.
 828 SYSTEMATIC BOTANY
D. nilhummatu Dunal, Prodr. 13(1): 542 p.p. 1852.—TYPE:
lectotypification required.
DATURA REBURRA A. S. Barclay, Bot. Mus. Leafl. 18: 258–260,
pl. 53 (upper), 54 (lower). 1959.—TYPE: MEXICO.
Sinaloa, Culiacan, 1904, Brandegee s. n. (holotype: UC).
DATURA WRIGHTII Regel, Gartenflora 8:193–194, plate 260. 1859.
D. meteloides Dunal f. wrightii (Regel) Danert, Pharmazie
9: 360. 1954.—NEOTYPE: U. S. A. Texas, Turkey Creek,
1849, Wright 526, (neotype: US).
Datura metel L. var. quinquecuspida Torr., Pacif. Railr. Rep. 7(3):
18. 1858.—TYPE: lectotypification required.
Acknowledgments. RB continues to appreciate Art Barclay who
bestowed upon him the never-ending challenges of Datura. Field assis-
tance provided by the following persons is greatly acknowledged:
Bruce Bartholomew, Sol Christians, David Gernandt, Les Landrum,
Edelmira Linares, Mario Luna, Gilberto Marquez, Ana Molina, Celerino
Montes of the Vizcaino Biosphere Reserve, R. Watson and the resi-
dences of Vizcaino, BCS. Virginia Evangelista, Susana Guzmán, Clarisa
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
Jiménez and Edelmira Linares provided technical assistance with speci-
mens and photographs. Julio Cesar Montero and Diana Martı́nez
Almaguer designed the color plates. Kanchi Gandhi, Fernando Chiang,
Georgina Ortega Leite and Judith Warnement supported our nomen-
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
clatural and bibliographic work. Arith Pérez conducted lab work
Instituto de Ecologı́a and Etelvina Gándara and Ismael G. Valdivieso
assisted in the phylogenetic analyses. The authors thank the curators
and staff of the herbaria cited for facilitating the examination of the
specimens as well as the reviewers and editors for their thoughtful
comments and recommendations.
Literature Cited
Amshoff, J. G. 1957. Datura. Pp. 369–373 in Flora de Cuba. Dicotyledoneas:
Melastomaceae a Plantaginaceae Vol. IV, eds. H. León and H. Alain.
La Habana: Imprenta P. Fernández y Cia.
Argueta Villamar, A., L. M. Cano Asseleih, and M. E. Rodarte. 1994.
Atlas de las plantas de la medicina tradicional mexicana. México, DF:
Instituto Nacional Indigenista.
Avery, A. G., S. Satina, and J. Rietsema. 1959. Blakeslee: The genus Datura.
New York: Ronald Press Co.
Barclay, A. S. 1959. New considerations in an old genus: Datura. Botanical
Museum Leaflets, Harvard University 18: 245–272.
Bello-Bedoy, R. and J. Núñez-Farfán. 2011. The effect of inbreeding on
defense against multiple enemies in Datura stramonium. Journal of
Evolutionary Biology 24: 518–530.
Bernhardi, J. J. 1833. Ueber die Arten der Gattung Datura. Neues Journal
der Pharmacie für Ärzte. Apotheker und Chemiker 26: 118–158.
Boyd, C. E. 2003. Rock Art of the Lower Pecos. College Station: Texas A&M
University Press.
Bye, R. and E. Linares. 2000. Mexico’s maddening plants of the Sixteenth
Century according to the Florentine Code. Pp. 15–29 in In
Chalchihuitl in Quetzalli, precious greenstone precious quetzal feather;
Mesoamerican studies in honor of Doris Heyden, ed. E. Quiñones Keber.
Lancaster: Labyrinthos.
Chiarini, F. and G. Barboza. 2007. Anatomical study of different fruit
types in Argentine species of Solanum subgen. Leptostemonum
(Solanaceae). Anales del Jardin Botanico de Madrid 64: 165–175.
Conklin, M. E. and H. H. Smith. 1971. Peroxidase isozymes: a mea-
sure of molecular variation in ten herbaceous species of Datura.
American Journal of Botany 58: 688–696.
Correll, D. S. and H. B. Correll. 1982. Datura. Pp. 1278–1281 in Flora of the
Bahama Archipelago. Vaduz: J. Cramer.
D’Arcy, W. G. 1973. Brugmansia, Datura in Flora of Panama. Annals of the
Missouri Botanical Garden 60: 582–586, 621–624.
D’Arcy, W. G. 1993. Brugmansia, Datura. Pp. 1100, 1104–1105 in Catalogue
of the flowering plants and gymnosperms of Peru / Catálogo de las
angiospermas y gimnospermas del Perú, eds. L. Brako and J. L. Zarucchi.
St. Louis, Missouri: Missouri Botanical Garden Press.
Doyle, J. J. and J. L. Doyle. 1987. A rapid DNA isolation procedure
for small quantities of fresh leaf material. Phytochemical Bulletin 19:
11–15.
Dunal, F. 1852. Datura. Prodromus systematis naturalis regni vegetabilis Prodromus
Systematis Universalis Regni Vegetabilis 13(part 1): 538–546, 690.
[Volume 38
Dunning, L. T. and V. Savolainen. 2010. Broad scale amplification of
matK for DNA barcoding plants, a technical note. Botanical Journal
of the Linnean Society 164: 1–9.
Eich, E. 2008. Solanaceae and Convolvulaceae– Secondary metabolites: biosyn-
thesis, chemotaxonomy, biological and economic significance. Berlin:
Springer-Verlag.
Fosberg, F. R. 1959. Nomenclatural notes on Datura L. Taxon 8: 52–57.
Fuentes, V. 1980. Solanaceas de Cuba I. Datura L. Revista del Jardı́n
Botánico Nacional 1: 61–81. (Cuba).
Geeta, R. and W. Gharaibeh. 2007. Historical evidence for a pre-
Columbian presence of Datura in the Old World and implications
for a first millennium transfer from the New World. Journal of
Biosciences 32: 1227–1244.
Gentry, J. L. and P. C. Standley. 1974. Datura in Flora of Guatemala.
Fieldiana: Botany 24: 37–42.
Goloboff, P. A., J. Farris, and K. Nixon. 2003. TNT: Tree analysis using
new technology. www.cladistics.com/aboutTNT.html.
Gunn, C. R. 1974. Seed characteristics of 42 economically important species
of Solanaceae in the United States. Agricultural Research Service
Technical Bulletin no. 1471. Washington, DC: U.S. Government
Printing Office.
Hadkins, E. S., R. Bye, W. A. Brandenburg, and C. E. Jarvis. 1997. Typifi-
cation of Linnaean Datura names (Solanaceae). Botanical Journal of
the Linnean Society 125: 295–308.
Haegi, L. 1976. Taxonomic account of Datura L. (Solanacaee) in Australia
with a note on Brugmansia Pers. Australian Journal of Botany 24:
415–435.
Hammer, K., A. Romeike, and C. Tittel. 1983. Vorarbeiten zur
monographischen Darstellung von wildpflanzensortimenten: Datura
L., sectiones Dutra Bernh, Ceratocaulis Bernh. et Datura. Die Kulturpflanze
31: 13–75.
Hepper, F. N. 1985. Brugmansia, Datura. Pp. 401–406 in A revised handbook
of the flora of Ceylon Volume VI, eds. M. D. Dassanayake and F. R.
Fosberg. New Dehli: Amerindi Publishing Company Private Limited.
Hokche, O., P. E. Berry, and O. Huber. 2008. Datura. P. 622 in Nuevo
catálogo de la flora vascular de Venezuela. Caracas: Fundación Instituto
Botánico de Venezuela.
Huelsenbeck, J. P. and F. Ronquist. 2001. MrBAYES: Bayesian inference
of phylogenetic trees. Bioinformatics 17: 754–755.
Hunziker, A. T. 2001. Genera Solanacearum – the genera of Solanaceae
illustrated, arranged according to a new system. Ruggell, Liechtenstein:
A. R. G. Gantner Verlag.
Huxley, A., M. Griffiths, and M. Levy, eds. 1999a. Brugmansia. P. 402 in
The new Royal Horticultural Society dictionary of gardening Vol. 1.
London: Macmillan Reference Limited.
Huxley, A., M. Griffiths, and M. Levy, eds. 1999b. Datura. Pp. 19–20 in
The new Royal Horticultural Society dictionary of gardening Vol. 2.
London: Macmillan Reference Limited.
Iglesias-Lepine, M. L., M. Mariñosa, M. D. Ferrer-Da Pena, and F. Epelde.
2012. El resurgir de las daturas y las brugmansias como drogas
recreacionales. Medicina Clı́nica 139: 90–91.
IUCN. 2001. Red list Categories and Criteria: Version 3.1. Gland, Switzerland
and Cambridge, U.K.: IUCN, Species Survival Commission.
Jiao, M., M. Luna-Cavazos, and R. Bye. 2002. Allozyme variation in
Mexican species and classification of Datura (Solanaceae). Plant
Systematics and Evolution 232: 155–166.
Johnston, T. 1972. Datura fastuosa: Its use in tsonga girls’ initiation.
Economic Botany 26: 340–351.
Knapp, S. 2002. Tobacco to tomatoes: a phylogenetic perspective on
fruit diversity in the Solanaceae. Journal of Experimental Botany 53:
2001–2022.
Lewis, W. H. and M. P. Elvin-Lewis. 2003. Medical botany, Plants affect-
ing human health. Hoboken: John Wiley & Sons, Inc.
Liberty Hyde Bailey Hortorium. 1976. Brugmansia, Datura. Pp. 184–185,
364–365 in Hortus third – A concise dictionary of plants cultivated in
the United States and Canada, eds. Staff of the Liberty Hyde Bailey
Hortorium. New York: Macmillan Publishing Co., Inc.
Linnaeus, C. 1753. Datura. P. 179 in Species Plantarum Vol. 1. Holmiae:
Impensis Laurentii Salvii.
Litzinger, W. J. 1981. Ceramic evidence for prehistoric datura use in
North America. Journal of Ethnopharmacology 4: 57–74.
Lockwood, T. E. 1973. Generic recognition of Brugmansia. Botanical
Museum Leaflets, Harvard University 23: 273–284.
Luna-Cavazos, M. and R. Bye. 2011. Phytogeographic analysis of the
genus Datura (Solanaceae) in continental Mexico. Revista Mexicana
de Biodiversidad 82: 977–988.
 2013] BYE AND SOSA: MOLECULAR PHYLOGENY OF DATURA
Luna-Cavazos, M., R. Bye, and M. Jiao. 2009. The origin of Datura metel
(Solanaceae): genetic and phylogenetic evidence. Genetic Resources
and Crop Evolution 56: 263–275.
Luna-Cavazos, M., M. Jiao, and R. Bye. 2000. Phenetic analysis of
Datura section Dutra (Solanaceae) in Mexico. Botanical Journal of
the Linnean Society 133: 493–507.
Macbride, J. F. 1962. Datura in Flora of Peru. Fieldiana: Botanical Series
13 (part 5B(no. 1)): 76–80.
Mace, E. S., C. G. Gebhart, and R. N. Lester. 1999. AFLP analysis of
genetic relationships in the tribe (Datureae) Solanaceae. Theoretical
and Applied Genetics 99: 634–641.
Maddison, D. R. and W. P. Maddison. 2011. MacClade. v. 4.08. http://
macclade.org/download.html.
Marzell, H., G. Bitter, and G. Hegi. 1975. Datura. Pp. 2612–2616 in
Illustrierte Flora von Mitteleuropa, Dicotyledones V Band, 4 Teil, ed.
G. Heigi. Berlin: Verlag Paul Parey.
Matuda, E. 1952. El género Datura en México. Boletı́n de la Sociedad
Botánica de México 14: 1–13.
Mino, Y. 1994. Amino acid sequence of ferredoxin from Datura arborea.
Phytochemistry 37: 429–431.
Mino, Y. 1995. Protein chemotaxonomy of genus Datura. IV. Amino
acid sequence of Datura ferredoxins depends not on the species
but the section of Datura plants from which it comes. Chemical &
Delivered by Publishing Technology to: Victoria Sosa IP: 187.157.168.78 on: Fri, 20 Sep 2013 14:07:19
Pharmaceutical Bulletin 43: 1186–1189.
Nee, M. 1986. Brugmansia, Datura. Pp. 12, 14–19, 69–76 in Solanaceae I.
Flora de Veracruz Fascı́culo 49. Xalapa: Instituto Nacional de
Investigaciones sobre Recursos Bióticos.
Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Olmstead, R. G., J. A. Sweere, R. E. Spangler, L. Bohs, and J. D. Palmer.
1999. Phylogeny and provisional classification of the Solanaceae
based on chloroplast DNA. Pp. 111–137 in Solanaceae IV: Advances
in biology and utilization, eds. M. Nee, D. E. Symon, R. N. Lester,
and J. P. Jessop. Richmond, Surrey: Royal Botanic Gardens, Kew.
Olmstead, R. G., L. Bohs, H. A. Migid, E. Santiago-Valentin, V. F. Garcia,
and S. M. Collier. 2008. A molecular phylogeny of the Solanaceae.
Taxon 57: 1159–1181.
Pabón-Mora, N. and A. Litt. 2011. Comparative anatomical and devel-
opmental analysis of dry and fleshy fruits of Solanaceae. American
Journal of Botany 98: 1415–1436.
Persoon, C. H. 1805. Brugmansia, Datura. Pp. 216–217 in Synopsis plantarum,
seu Enchiridium botanicum, complectens enumerationem systematicam
specierum hucusque cognitarum. Vol. 1. Paris: C.F. Cramerum.
Persson, V., S. Knapp, and S. Blackmore. 1999. Pollen morphology and
the phylogenetic analysis of Datura L. and Brugmansia Pers.
Pp. 171–187 in Solanaceae IV: Advances in biology and utilization,
eds. M. Nee, D. E. Symon, R. N. Lester, and J. P. Jessop. Richmond,
Surrey: Royal Botanic Gardens, Kew.
Posada, D. 2008. jModelTest: Phylogenetic model averaging. Molecular
Biology and Evolution 25: 1253–1256.
Rambaut, A. 2002. Se-Al Sequence Alignment Editor, v2.0a11. Oxford:
Department of Zoology, University of Oxford.
Rogers, S. O. and A. J. Bendich. 1985. Extraction of DNA from milli-
gram amounts of fresh, herbarium and mummified plant tissues.
Plant Molecular Biology 5: 69–76.
Ronquist, F. and J. P. Huelsenbeck. 2003. MrBayes 3: Bayesian phylo-
genetic inference under mixed models. Bioinformatics 19: 1572–1574.
Russell, J., C. Edwards, C. Jordan, A. Chu, D. Blythe, and J. Krick.
2010. Jimsonweed poisoning associated with a homemade stew –
Maryland, 2008. Morbidity and Mortality; Weekly Report 59: 102–104.
Safford, W. E. 1921. Synopsis of the genus Datura. Journal of the Washington
Academy of Sciences 11: 173–189.
Safford, W. E. 1922. Daturas of the Old World and New: an account of
their narcotic properties and their use in oracular and initiatory
ceremonies. Pp. 537–567 in Annual report of the Board of Regents
of the Smithsonian Institution 1920. Washington, DC: Government
Printing Office.
Sang, T., D. J. Crawford, and T. F. Stuessy. 1997. Chloroplast DNA phy-
logeny, reticulate evolution, and biogeography of Paeonia (Paeoniaceae).
American Journal of Botany 84: 1120–1136.
Satina, S. and A. G. Avery. 1959. A review of the taxonomic history
of Datura. Pp. 16–47 in Blakeslee: The genus Datura, eds. A. G. Avery,
S. Satina, and J. Rietsena. New York: Ronald Press.
Shaw, J., E. B. Lickey, J. T. Beck, S. B. Farmer, W. S. Liu, J. Miller, K. C.
Siripun, C. T. Winder, E. E. Schilling, and R. L. Small. 2005. The
tortoise and the hare. II: Relative utility of 21 noncoding chloro-
plast DNA sequences for phylogenetic analysis. American Journal of
Botany 92: 142–166.
829
Shaw, J. M. H. 2000. Brugmansia, Datura. Pp. 232–267 in The European
garden flora – Loganiaceae to Compositae Vol. VI, eds. J. Cullen and
S. M. Walters. Cambridge, UK: Cambridge University Press.
Standley, P. C. and C. V. Morton. 1938. Datura in Flora of Costa Rica.
Field Museum of Natural History Botanical Series 18: 1054–1056.
Symon, D. and L. A. R. Haegi. 1991. Datura (Solanaceae) is a New World
genus. Pp. 197–210 in Solanaceae III: Taxonomy, chemistry, evolution,
eds. J. G. Hawkes, R. N. Lester, M. Nee, and N. Estrada. Richmond,
Surrey: Royal Botanic Gardens, Kew.
Taberlet, P., L. Gielly, G. Pautou, and J. Bouvet. 1991. Universal primers
for amplification of three non-coding regions of chloroplast DNA.
Plant Molecular Biology 17: 1105–1109.
Tate, J. A. and B. B. Simpson. 2003. Paraphyly of Tarasa (Malvaceae)
and diverse origins of the polyploid species. Systematic Botany 28:
723–737.
Terry, R. G., G. K. Brown, and R. G. Olmstead. 1997. Examination
of subfamily phylogeny in Bromeliaceae using comparative
sequencing of plastid locus ndfF. American Journal of Botany 84:
664–670.
Touwaide, A. 1998. Datura stramonium L.: Old or New World? Delpinoa
39–40: 29–43.
Vearrier, D. and M. I. Greenberg. 2010. Anticholinergic delirium follow-
ing Datura stramonium ingestion: Implications for the internet age.
Journal of Emergencies, Trauma, and Shock 3: 303.
Appendix 1. Taxa considered in the phylogenetic study of Datura.
Vouchers, GenBank accession numbers for sequences obtained in
this project as well as for the previously published sequences of the
DNA sequences used in this paper are listed. GenBank accessions
correspond to matK, ndhF, psbA-trnH and trnL-trnF.
Brugmansia suaveolens (Humb & Bonpl. ex Willd.) Sweet. MEXICO.
Mexico: R. Bye 35848 (MEXU), KC146589, KC146618, KC146624, KC146642.
Brugmansia candida Pers. “pañuelo”. MEXICO. Mexico: Bye & Linares
Bc-p (MEXU), KC146590, KC146617, KC146625, KC146643. Brugmansia
candida Pers. “sencillo”. MEXICO. Mexico: Bye & Linares Bc-s (MEXU),
KC146591, KC146619, KC146626, KC146644. Capsicum minutiflorum
(Rusby) Hunz. BOLIVIA. Bohs & al. 3012 (UT),–, DQ667543,–, EU580970.
Datura arenicola Gentry ex Bye & Luna. MEXICO. Baja California Sur:
Bye & Linares 36323, 36326, 36327 (MEXU), KC146592, KC146608,
KC146627, KC146645. Datura ceratocaula Ortega. MEXICO. Guanajuato:
Bye & Luna 35879, 35884, 35888, 36081c (MEXU), KC146593,—-,—-,
KC146646. Datura discolor Bernh. MEXICO, Baja California Sur: Bye &
Linares 35960c, 36314bc1, 36328bc2, 36330bc3, 36333bc4, 36334bc5 (MEXU),
KC146594, KC146612, KC146628, KC146647. Datura ferox L. MEXICO.
Molina ARG2009 (MEXU), KC146595, KC146615, KC146629, KC146648.
Datura inoxia Mill. MEXICO. Bye & Linares 35876, 35988z, 36222p,
36178q, 20080717 (MEXU), KC146596, KC146611, KC146630, KC146649.
Datura kymatocarpa A. S. Barclay. MEXICO. Guerrero: Bye & Luna
36498 (MEXU), KC146597,—-, KC146631, KC146650. Datura lanosa
Barclay ex Bye. MEXICO. Chihuahua: Bye 35944, 35961, 35964, 35966
(MEXU), KC146598, KC146622, KC146632, KC146651. Datura metel
L. MEXICO. Distrito Federal: Bye & Linares TM (MEXU), KC146599,—-,
KC146633,—-. Datura leichhardtii F. Muell. ex. Benth. & F. Muell.
subsp. pruinosa (Greenm.) A.S. Barclay ex K. Hammer. MEXICO.
Puebla: Bye & Linares 36050 (MEXU), KC146600, KC146613, KC146634,
KC146652. Datura quercifolia Kunth. MEXICO. Guanajuato: Bye & Luna
35846, 35878, 35887, Bye, Gernandt & Chrisitans 35989z, 36114s (MEXU),
KC146601, KC146623, KC146635, KC146653. Datura reburra A. S.
Barclay. MEXICO. Sinaloa: Márquez S-1 (MEXU), KC146602, KC146609,
KC146636, KC146654. Datura stramonium L. MEXICO. Distrito Federal:
Bye 35845 (MEXU), KC146603, KC146621, KC146637, KC146655. Datura
wrightii Regel. U. S. A. Arizona: Landrum 11345 (MEXU), KC146604,
KC146610, KC146638, KC146656. Jaltomata procumbens (Cav.) J. L.
Gentry. MEXICO. Distrito Federal: Bye 35844 (MEXU), KC146605,
KC146614, KC146639, KC146657. Juanulloa mexicana (Schltdl.) Miers.
MEXICO: BIRM S.0411/69 (BIRM),–, EU580901,–, EU581008. Lycium
barbarum L. CHINA: Olmstead S-35 (WTU),–, EU580907,–, AB036572.
Nicotiana tabacum L. in cult., no voucher, L14953, Z00044. Physalis
alkekengi L. ASIA: D’Arcy 17707 (MO),–, U08927,–, DQ180420. Solandra
guttata D. Don. MEXICO. Mexico: Bye & Linares Sg (MEXU), KC146606,
KC146616, KC146640, KC146658. Solandra maxima (Sessé & Moc.) P.S.
Green. MEXICO. Distrito Federal: Bye & Linares 35837 (MEXU), KC146607,
KC146620, KC146641, KC146659. Solanum betaceum Cav. BOLIVIA: Bohs
2468 (UT),–, U47428,–, DQ180426.