LAMIALES NEWSLETTER
LA MIA LE S
Issue number 6
August 1998
ISSN 1358-2305
EDITORIAL
CONTENTS
Ray Harley, Alan Paton, Tivvy Harvey
Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK
A belated welcome, we are afraid,
to this the 6th issue of the Lamiales
Newsletter. Apart from perennial
excuses due to pressure of work,
one of us, who was taking a year in
Brazil, until mid-1997, is still trying to catch up, and is currently
very busy bringing together and
editing contributions for a Lamiales
part for Kubitzki’s Families and
Genera of Vascular Plants. This is
proving challenging, but he is very
grateful for all the support he has
received. In spite of all this, we still
hope to be on time for the next
issue, and have already received
contributions! Please continue to
send your valuable contributions to
y.harvey@rbgkew. org.uk.
The Newsletter is not intended for
papers which could be more appropriately published in reviewed scientific journals but is rather intended for preliminary research reports
especially on new or interesting topics, reports on the activities of
research groups, requests for information, news and comments. We
try to vary the content, so that there
will be something for everyone, so
please do not be down-hearted if
your paper does not appear. The
present issue starts with a topic that
could affect us all: phylogenetic
nomenclature, using Labiatae as a
case study, by Phil Cantino. This is
a highly contentious issue, so we
hope you will send us your views.
Dick Olmstead points out that in
line with molecular evidence, the
Lamiales includes not only the Ver-
benaceae and Labiatae, but also
other families such as Scrophulariaceae, Acanthaceae, Bignoniaceae,
Myoporaceae etc. Should the
Newsletter be more inclusive or
should we restrict it to just the Labiatae? Not wishing to either exclude
anyone already in our group, nor
wishing to take on a lot more work,
the editors have taken a unilateral
decision to keep things as they are
for the present, but your views
would be welcome.
Unfortunately, the hoped-for Lamiales meeting in Mexico has not
materialized, due to unforeseen difficulties, so we have suggested postponing Mexico for a future meeting.
As a lot has happened since our last
meeting, we look forward to suggestions for a venue and local support
for a meeting around the year 2001.
Our directory of research in the
Labiatae and Verbenaceae is now
showing its age, and needs updating.
Please can we ask you to complete
the accompanying form?
So far we have been fortunate to
have the newsletter production
funded by R.B.G. Kew. Due to current financial restraints and to
ensure continuity, we would like to
ask for a voluntary contribution of
£3 Sterling (cash or bank draft) or
$5 US (cash only), per issue. This
will cover postage and part of production costs and you may find it
worthwhile to pay for a few years
in advance. ❑
1
Editorial
1
Oxera, Faradaya and Hosea,
systematics and conservation
1
Labiatae as a case study in
Phylogenetic Nomenclature
3
Studies on Colombian
Labiatae lectins
6
Confirmation of a monophyletic
Chloanthoideae (Lamiaceae)
comprising tribes Chloantheae
and Prostanthereae
7
Ethnobotany and anticonvulsant
properties of Lamiaceae from
Rio Grande do Sul (Brazil)
10
Phlomic acid in Lamioideae
seed oils
13
Bibliography
18
OXERA, FARADAYA
AND HOSEA,
SYSTEMATICS AND
CONSERVATION
Rogier P. J. De Kok
Centre for Plant Biodiversity
Research,
Australia National Herbarium,
GPO Box 1600, Canberra, ACT
2601, Australia
The genera Oxera Labill., Faradaya
F. Muell. and Hosea Ridl. were
revised as part of a D.Phil thesis at
the Department of Plant Sciences,
Oxford (de Kok 1997). A cladistic
analysis of the whole group, based
on morphological, flavonoid and ➣
Table 1. Generic differences between Oxera, Faradaya and Hosea and some close relatives
Number of
fully matured
stamens
Mature
fruit
elongated
Nr. of
corolla
lobes
Flower
resupinate
Corolla wall
fleshy
Orientation
of the
vascular
bundles in
the petiole
Ovule
position
Oxera
2(-4)
Yes
4
No
Some
a
p
Faradaya
4
Yes
4
No
Yes
a
p
Hosea
4
Yes
4
No
No
b
p
Clerodendrum s.s.
4
No
5
Yes
No
b
c+m
Rotheca
4
No
5
No
No
b
c+m
Huxleya
4
?
5
?
No
?
c+m
Kalaharia
4
No
5
Yes
No
b
c
Key to symbols: a. interrupted full circle, b. half circle, c. basal, m. middle position & p. pendulous
nrDNA characters, was also part of
the project. The revisions, their
flavonoid contents (Grayer & de
Kok, in press) and the cladistic
analyses will be published elsewhere. In this article a brief
overview of the genera studied is
given, and the conservation status of
two rare Oxera taxa are discussed.
In the genus Oxera, 21 species and
six subspecies are recognised. The
genus is mainly restricted to New
Caledonia, but a new species which
is only known from cultivation is
recognised from Vanuatu. The
genus is subdivided into five informal groups, based on a number of
floral and fruit characters. In the
genus Faradaya only three species
are recognised: Faradaya amicorum (Seem.) Seem., F. lehuntei
(Horne ex Baker) A.C. Smith and
F. splendida F. Muell. The genus is
found from Samoa and Tonga to
New Guinea, North Queensland
and Sabah. The monotypic genus
Hosea is recognised as distinct
with its one species, Hosea lobbii
(C. B. Clarke) Ridl., confined to
Sarawak and Brunei.
Oxera, Faradaya and Hosea differ
from the larger genus Clerodendrum
s.s. and Rotheca and the monotypic
genera Huxleya and Kalaharia by
having a tetramerous corolla and an
elongated mature fruit (Table 1). This
morphological character is supported
for Clerodendrum s. s. and Rotheca
by a cladistic analysis of cp- and
nrDNA characters (Steane 1995).
Like Clerodendrum, Rotheca and
Huxleya, Hosea has two pairs of
equal stamens and a fragile corolla. It
also shares with Clerodendrum and
Rotheca the character of having its
vascular bundles in the cross-section
of the petiole in a half circle, rather
than in an interrupted full circle as in
Oxera and Faradaya. The general
shape of the corolla reminds one
strongly of Faradaya, but it differs in
the texture of the wall. Faradaya
corollas are thick and waxy compared
with Hosea flowers. The sharply
pointed shape of the fruit is one of the
main differences between Hosea and
its close relatives.
Faradaya has two pairs of equal stamens and a large fleshy corolla. It
shares with Oxera the character of
the vascular bundles in the crosssection of the petiole in an interrupted full circle. Oxera usually has one
pair of stamens, but sometimes may
have two pairs. The flowers of Faradaya are tetramerous and actinomorphic, unlike those of Clerodendrum
which have five corolla lobes, and
2
those of Oxera and Hosea, which
although having four corolla lobes,
are zygomorphic, as one of the
lobes is a well developed lip.
tion of its habitat for further development of Nouméa City. The last
remaining undamaged forest (Point
Lascalle) on the peninsula is earmarked for development as a housing estate (Gramon 1995). A road
was pushed through the forest in
1995, and has already severely damaged the main population of the
plant in this forest. The taxon is also
found in two smaller protected areas
on the peninsula: Parc Forestier M.
Corbasson (35 ha) and Parc Provincial du Ouen-Toro (44 ha). Both
parks are city parks rather than biodiversity reserves (Bouchet et al.
1995), and no special measures are
taken to safeguard the plants.
Specimens of both O. rugosa and
O. pulchella subsp. grandiflora
should be taken into cultivation to
safeguard the survival of the taxa.
Plants of O. pulchella which are
already in cultivation should be
properly identified as to which subspecies they belong and maintained
in case reintroduction of O. pulchella subsp. grandiflora into New
Caledonia becomes necessary. One
way to ensure the survival of O.
pulchella subsp. grandiflora is to
promote its use as a garden plant in
New Caledonia itself (Gramon
1995), but efforts will have to be
made to prevent hybridisation with
the northern subspecies.
References
Bouchet, P., Jaffré, T. & Veillon, J.
M. (1995). Plant extinction in New
Caledonia: protection of sclerophyll
forest urgently needed. Biodiversity
and Conservation 4: 415-428.
Gramon, A. (1995). Que sont les
belles fleurs sauvages devenues?
Les Nouvelles Calédoniens.
21.1.1995. Nouméa.
Grayer, R. J. & de Kok, R. P. J. (in
press). Flavonoids and verbascoside
as chemotaxonomic characters in
the genera Oxera and Faradaya
(Labiatae). Accepted by Biochemical Systematics and Ecology.
de Kok, R. P. J. (1997). The biology
and systematics of Oxera, Faradaya
and Hosea (Labiatae). D.Phil thesis,
University of Oxford, copies at L,
K, NSW & OXF.
Lucas, G. & Synge, H. (1978). The
IUCN plant red data book. Gresham
Press, Old Woking.
Steane, D. A. (1995). Molecular
Systematics of Clerodendrum L. s.l.
(Lamiaceae). D.Phil thesis. University of Oxford. ❑
Conservation
LABIATAE AS A CASE STUDY IN PHYLOGENETIC NOMENCLATURE
Two taxa of Oxera can be considered endangered according to the
IUCN status categories (Lucas &
Synge 1978: 25); both plants are
restricted to the sclerophyll forest of
New Caledonia which is the vegetation type most at risk on this Island
(Bouchet et al. 1995: 418-420).
Philip D. Cantino
Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701, U.S.A.
Oxera rugosa is known from only a
few localities in the south of the
island. There is one population
(which I have not seen) which grows
in the protected area of the Parc Territorial de la Thy. The other localities
are along the southern slopes of the
Mourange mountains. These areas
are unprotected and are in constant
danger of being burned (Bouchet et
al. 1995). In 1995, one of the best
known populations was accidentally
set on fire by the French army during
an exercise with live ammunition on
a nearby shooting range.
Oxera pulchella subsp. grandiflora
is restricted to the Nouméa peninsula, and is threatened by the destruc-
Because the journal Systematic
Biology (formerly Systematic Zoology) is not widely read in botanical
circles, many readers of this
newsletter may not be aware of a
recent paper (Cantino et al., 1997)
in which the classification of Labiatae is used to compare the current
system of nomenclature (as represented in the ICBN) to an alternative phylogenetic system proposed
by de Queiroz and Gauthier (1992,
1994, 1996). This case study suggests that phylogenetic nomenclature functions better than the current system when one’s objective is
to provide unambiguous and stable
names for clades.
For example, Teucrioideae (Cantino et al., 1992) is a novel and well
supported clade of 23 genera that
transcends the traditional boundary
between Labiatae and Verbenaceae.
One would like a newly discovered
clade to have a name that is both
unique (not previously applied to
another group) and stable. Teucrioideae, the name that had priority under the ICBN when the clade
was discovered, is not unique but at
least had not been used in recent
times and therefore was unlikely to
be associated with another grouping
in anyone’s mind. However, the
subsequent finding that Ajuga is a
member of this clade (Wagstaff and
Olmstead, 1997; Wagstaff et al.,
1998) requires that the taxon be
renamed Ajugoideae, which has
priority under the ICBN, an undesirable change because this name
has been applied already to at least
three other groupings in well
known or recent classifications.
Under the phylogenetic system of
nomenclature, this would not have
occurred; once a clade is named, it
3
retains that name permanently. The
greatest difficulties with the current system were encountered with
infrafamilial taxon names based on
the type of the family. For example, the ICBN has required me to
apply the name Lamioideae to
three different groups in 11 years
(Cantino and Sanders, 1986; Cantino et al., 1992; Cantino et al.,
1997), since any subfamily-level
taxon that includes Lamium must
bear this name.
The Phylogenetic System of
Nomenclature
Phylogenetic nomenclature differs
fundamentally from the current system in lacking mandatory ranks
above the species level and in the
way that taxon names are defined.
In the current system, plant names ➣
at or below the family level are
operationally defined as the taxon
assigned to rank X that contains
type Y (de Queiroz and Gauthier,
1994). For example, Verbenaceae is
the taxon of family rank that contains the type of the genus Verbena.
This kind of definition is also frequently used above the family level,
but this is not required by the
ICBN. Phylogenetic nomenclature
is entirely different in that names
are defined in terms of phylogenetic
relationships (de Queiroz and Gauthier, 1990; Schander and Thollesson, 1995). For example, Lamiaceae might be defined as the least
inclusive clade that contains Lamium purpureum, Glechoma hederacea, and Vitex agnus-castus
(although many more species were
cited in my published definition
[Cantino et al., 1997] to reduce the
likelihood of subsequent changes in
taxon membership due to future
phylogenetic findings).
Phylogenetic nomenclature is the
logical culmination of a revolution
that began with Darwin and was
advanced by Hennig. De Queiroz
(1997) dubbed this the “evolutionization of taxonomy.” As the principle of common descent has
assumed a progressively more
prominent position in the philosophical framework of systematics,
taxon names have come to have
implicit phylogenetic meanings.
The final step in this process is
replacement of our current system
of nomenclature, which is based on
preDarwinian principles, with one
that defines taxon names in explicit
phylogenetic terms.
A particularly desirable feature of
phylogenetic nomenclature is that it
makes it possible to name newly
discovered clades without changing
the names of other taxa. A disadvantage of the current system is that
the ranks of taxa, and hence their
names, are dependent on their position relative to other taxa. Thus
naming a newly discovered clade
can cause a cascade of name
changes elsewhere in the hierarchy
as taxa shift in rank (Kron, 1997;
Hibbett and Donoghue, 1998). By
analogy, if chemists were operating
under a system that required the
names of some elements to change
when a new one is discovered, surely the scientific community would
rebel! This serious drawback of the
current system discourages systematists from naming clades as they
are discovered (Hibbett and
Donoghue, 1998). As a result, our
classifications are falling farther
and farther behind our knowledge
of relationships.
Although phylogenetic nomenclature is “rankless” in that it lacks
mandatory ranks, there is no prohibition against using ranks if one wishes to (de Queiroz, 1997). The important thing is that the taxon name is
not dependent on rank and thus does
not change if its position in the hierarchy changes. This is one way in
which the phylogenetic system promotes nomenclatural stability.
berg and Brouillet, 1994; Olmstead,
1995), and it has been argued that
the concepts of monophyly and
paraphyly don’t even apply to
species because the relationships
among the individuals composing a
species are tokogenetic (reticulating) rather than phylogenetic
(Nixon and Wheeler, 1990). However, species are still objective entities if they are conceived of as segments of population level evolutionary lineages (de Queiroz, in press)
whose cohesiveness is due to natural processes such as interbreeding
(de Queiroz and Donoghue, 1988).
In contrast, a paraphyletic supraspecific taxon is subjective in that it has
no existence outside the mind of the
taxonomist. The claim that species
are sometimes paraphyletic is not a
valid argument for formal recognition of paraphyletic taxa above the
species level. Species need not be
monophyletic to be objective natural entities, but monophyly is necessary above the species level unless
one can demonstrate another natural
process that conveys objective existence to supraspecific entities.
Species and Clades
The Binominal System
In the absence of mandatory ranks,
the principal kinds of taxa in the
phylogenetic system are clades and
species. The acceptance of the
species category in a rankless system may seem like a logical contradiction, but it is not if one views
“species” not as a rank but as a fundamentally different kind of entity
than a clade. If systematics is a science (and I hope this is no longer at
issue), named taxa should correspond to objective natural entities,
i.e., products of evolution that exist
outside the human mind and whose
existence is hypothesized based on
evidence. Such entities are discovered rather than created. Species and
clades qualify as objective entities
for different reasons. Clades, being
monophyletic, are complete systems
of common descent. Species are not
necessarily monophyletic (Riese4
Because phylogenetic taxonomy
lacks formal ranks, there is no such
thing as a genus, thus binominal
nomenclature is incompatible with
phylogenetic taxonomy. Although
people may object to abandoning a
convention that has been in use for
over 200 years, a critical examination of binominal nomenclature
reveals serious drawbacks (Cantino, 1998). One problem is that
t h e binominal system cannot
accommodate uncertainty about
generic relationships. This can lead
to a taxonomic dilemma when a
genus (“X”) is found to be para- or
polyphyletic as currently delimited,
but phylogenetic resolution is too
poor to refer all of its species to
smaller monophyletic genera. The
most satisfactory approach would
be to name the well supported
clades within former genus X as
genera but designate the species
that lie outside these clades as
“incertae sedis” with regard to
genus. However, this cannot be
done within the binominal system.
A second problem with the binominal system is that it is a major
cause of nomenclatural instability
at the species level, because
changes in generic circumscription
necessitate species-level name
changes. This is not true at any
other rank (e.g., splitting a family
into two families does not require
that genus names change).
A viable alternative to binominal
nomenclature is a system of hyphenated uninomials (Michener, 1964).
Thus, for example, Lamium purpureum would become known as
Lamium-purpureum and would retain
this name even if subsequent research
demonstrated that it is not a member
of a clade called Lamium. The transition to such a system would be relatively painless if the hyphenated uninomials were based on currently
accepted binomials and retained the
same type. As Michener (1964)
noted, decoupling species names
from genus names would have two
advantages: It would greatly increase
nomenclatural stability at the species
level, where stability is most critical,
by preventing changes in species
names due to generic recircumscription. Secondly, it would facilitate
reclassification above the species
level in response to new research, as
systematists would no longer have to
weigh the implications for specieslevel nomenclature when deciding
whether to translate their findings
into classification.
The Future of Phylogenetic
Nomenclature
Any major change in the naming of
organisms is bound to involve a difficult transitional period, thus it
should only be undertaken if the end
product will be a substantial
improvement. For this reason, I
question the BioCode concept
(Greuter et al., 1996) which I view
as having a poor cost/benefit ratio.
Why impose a new system of rules
on the systematic community when
it suffers the same fundamental
drawbacks as the current system?
On the other hand, the phylogenetic
system has significant advantages. It
would facilitate the naming of newly
discovered clades without forcing
name changes elsewhere in the classification, and it would substantially
improve nomenclatural stability - at
least at the species level and perhaps
elsewhere. The jury is still out on
the latter point. My case study using
the Labiatae suggests that nomenclatural stability will improve above
the species level as well, but other
taxa should be examined.
been noted by both opponents (e.g.,
Brummitt, 1996, 1997; Sosef, 1997)
and proponents of the view that all
taxa above the species level should
be monophyletic. Even if one accepts
paraphyletic supraspecific taxa, phylogenetic nomenclature appears to
improve nomenclatural stability and
facilitate the naming of clades. If
these advantages are corroborated by
additional case studies, the systematic community should give serious
consideration to adopting phylogenetic nomenclature. Classification is
the principal way in which knowledge of relationships is imparted to
other biologists and the general public. Tradition alone is not sufficient
reason to retain a set of conventions
that impedes the communication of
scientific progress.
References
What is needed now is a draft code
of phylogenetic nomenclature, which
could be applied experimentally to a
wide variety of organisms with different levels of phylogenetic resolution. There is no reason why the two
nomenclatural systems cannot coexist, so long as names bear some kind
of designation to show which system
they pertain to. For example, if
names in the phylogenetic system
were to end with a standard symbol
such as the Greek letter φ (phi, for
phylogenetic), they would be immediately recognizable. Users would
know that a taxon called Scutellarioideaeφ represented a hypothesized
clade, whereas no such assumption
could be made about a taxon called
Scutellarioideae. Alternatively, a
uniform ending might be used for all
names in the phylogenetic system
(e.g., -ina; Kron, 1997), but this
would require the abandonment of
many unambiguous names such as
Asteraceae and Poaceae that already
apply (although not explicitly) to
clades.
The incompatibility of the traditional
“Linnaean” system of classification
(and nomenclature) with the goals of
modern phylogenetic systematics has
5
Brummitt, R. K. (1996). In defense
of paraphyletic taxa. Pp. 371-384.
In Maesen, L. J. G. van der, Burgt,
X. M. van der and Medenbach de
Rooy, J. M. van (eds.), The Biodiversity of African Plants. Proceedings XIVth AETFAT Congress, 2227 August 1994, Wageningen,
Netherlands. Dordrecht.
Brummitt, R. K. (1997). Taxonomy
versus cladonomy, a fundamental
controversy in biological systematics. Taxon 46: 723-734.
Cantino, P. D. 1998. Binomials,
hyphenated uninomials, and phylogenetic nomenclature. Taxon 47 (2):
425-429
Cantino, P. D. and Sanders, R. W.
(1986). Subfamilial classification of
Labiatae. Syst. Bot. 11: 163-185.
Cantino, P. D., Harley, R. M. and
Wagstaff, S. J. (1992). Genera of
Labiatae: status and classification.
Pp. 511-522. In Harley, R. M. and
Reynolds, T. (eds.), Advances in
Labiate Science. Royal Botanic
Gardens, Kew.
Cantino, P. D, Olmstead, R. G. and
Wagstaff, S. J. (1997). A comparison of phylogenetic nomenclature
with the current system: a botanical
case study. Syst. Biol. 46: 313-331. ➣
De Queiroz, K. (1996). A phylogenetic approach to biological nomenclature as an alternative to the Linnaean systems in current use. In
Reveal, J. L. (ed.), Proceedings of a
Mini-symposium on Biological
Nomenclature in the 21st Century.
[http://www.inform.umd.edu/PBIO/
nomcl/dequ.html].
De Queiroz, K. (1997). The Linnaean hierarchy and the evolutionization of taxonomy, with emphasis
on the problem of nomenclature.
Aliso 15: 125-144.
De Queiroz, K. In press. The general
lineage concept of species, species
criteria, and the process of speciation:
a conceptual unification and terminological recommendations. In Howard,
D. J. and Berlocher, S. H. (eds.), Endless Forms: Species and Speciation.
Oxford University Press, New York.
De Queiroz, K. and Donoghue, M.
J. (1988). Phylogenetic systematics
and the species problem. Cladistics
4: 317-338.
De Queiroz, K. and Gauthier, J.
(1990). Phylogeny as a central principle in taxonomy: phylogenetic
definitions of taxon names. Syst.
Zool. 39: 307-322.
De Queiroz, K. and Gauthier, J.
(1992). Phylogenetic taxonomy.
Annu. Rev. Ecol. Syst. 23: 449-480.
De Queiroz, K. and Gauthier, J.
(1994). Toward a phylogenetic system of biological nomenclature.
Trends Ecol. Evol. 9: 27-31.
Greuter, W., Hawksworth, D. L.,
McNeill, J., Mayo, M. A., Minelli,
A., Sneath, P. H. A., Tindall, B. J.,
Trehane, P., and Tubbs P. (1996).
Draft BioCode: the prospective
international rules for the scientific
names of organisms. Taxon 45:
349-372.
Hibbett, D. S. and Donoghue, M. J.
(1998). Integrating phylogenetic
analysis and classification in fungi.
Mycologia 90: 347-356.
Kron, K. A. (1997). Exploring alternative systems of classification.
Aliso 15: 105-112.
Michener, C. D. (1964). The possible use of uninominal nomenclature
to increase the stability of names in
biology. Syst. Zool. 13: 182-190.
Nixon, K. C. and Wheeler, Q. D.
(1990). An amplification of the phylogenetic species concept. Cladistics
6: 211-223.
Olmstead, R. G. (1995). Species
concepts and plesiomorphic species.
Syst. Bot. 20: 623-630.
Rieseberg, L. H. and Brouillet, L.
(1994). Are many plant species
paraphyletic? Taxon 43: 21-32.
Schander, C. and Thollesson, M.
(1995). Phylogenetic taxonomy—
some comments. Zool. Scripta 24:
263-268.
Sosef, M. S. M. (1997). Hierarchical models, reticulate evolution
and the inevitability of paraphyletic supraspecific taxa. Taxon 46:
75-85.
Wagstaff, S. J. and Olmstead, R.
G. (1997). Phylogeny of Labiatae
and Verbenaceae inferred from
rbcL sequences. Syst. Bot. 22:
165-179.
Wagstaff, S. J., Hickerson, L.,
Spangler, R. E., Reeves, P. A., and
Olmstead, R. G. (1998). Phylogeny
of Labiatae s.l. inferred from
cpDNA sequences. Plant Syst.
Evol. (in press). ❑
Gerardo Pérez*, Nohora Vega* & José Luis Fernández-Alonso**
*Biochemistry Laboratory, Chemistry Department, Universidad Nacional de Colombia, Bogotá, COLOMBIA
(E-mail: bioquim@ciencias.ciencias.unal.edu.co)
**Herbario Nacional Colombiano, Instituto de Ciencias Naturales, Ap 7495 Universidad Nacional de Colombia,
Bogotá, COLOMBIA
Recently the Laboratory of Biochemistry at the Chemistry Department and the Instituto de Ciencias
Naturales, both at the National
University, Bogotá, Colombia,
have joined forces to work on
lectins from Labiatae. Due to their
many interesting properties, two of
them being the specific recognition
of carbohydrates and the agglutination of cells, the lectins have
attracted much attention. Leguminosae and some Gramineae are the
commonest source of lectins; in
Labiatae only the Salvia species
from temperate zones have been
investigated in this respect (Bird &
Wingham, 1974, 1976, 1977, 1982)
6
Our second goal is to purify and
characterise the lectin from S.
bogotensis subsp. bogotensis, a
taxon which is quite abundant
near Bogotá. We have devised an
extraction scheme and are currently
working on the isolation and characterisation of the lectin. We are interested in establishing contacts with
groups working on Labiatae
lectins/proteins so we can discuss
results and eventually collaborate.
References
Bird, G. W. G. & Wingham, J. (1974).
Haemagglutinins from Salvia. VoxSang. 26: 163-166.
Bird, G. W. G. & Wingham, J. (1976).
More Salvia agglutinins. Vox-Sang.
30: 217-219.
Bird, G. W. G. & Wingham, J. (1977).
Yet more Salvia agglutinins. VoxSang. 32: 121-122.
Bird, G. W. G. & Wingham, J.
(1982). More Tn-specific lectins
from seeds of genus Labiatae: Hyptis
sp. Chan. Salvia lyrata and Marrubi-
um velutinum. Clin. Lab. Haematol.
4: 403-404.
Fernández-Alonso, J. L. (1990).
Notas sobre Scutellaria ( Labiatae )
en Colombia y Ecuador. Anales
Jard. Bot. Madrid 47 (1): 105-123.
Fernández-Alonso, J. L. (1995).
Estudios en Labiatae de Colombia,
novedades en los géneros Salvia e
Hyptis. Rev. Acad. Col. Ciencias.
19: 469-480.
Piller, V., Piller, F. & Cartron, J.
(1986). Isolation and characterisation of N-acetylgalactosamine specific lectin from Salvia sclarea
seeds. J. Biol. Chem. 261: 1406914075.
Wood, J. R. I. (1988). The genus
Lepechinia ( Labiatae ) in Colombia. Kew Bulletin 43: 291-301.
Wood, J. R. I. & Harley, R. M.
(1989). The genus Salvia (Labiatae)
in Colombia. Kew Bulletin 44: 211279. ❑
CONFIRMATION OF A MONOPHYLETIC CHLOANTHOIDEAE
(LAMIACEAE) COMPRISING TRIBES CHLOANTHEAE
AND PROSTANTHEREAE
STUDIES ON COLOMBIAN LABIATAE LECTINS
During the last decade some genera
from Labiatae in Colombia, have
been reviewed by Wood (1988),
Wood and Harley (1989) and Fernández-Alonso (1990, 1995). In the
course of this work many endemic
species have been described.
in Salvia bogotensis Benth., S. paliifolia Kunth, S. pauciserrata
Benth., S. rubescens Kunth, S.
amethystina J. E. Smith, and Lepechinia conferta (Benth.) Epling. In
Salvia scutellarioides Kunth, S. sordida Benth., S. uribei Wood &
Harley, S. gachantivana FernándezAlonso and Lepechinia salvifolia
(Kunth) Epling, no lectin has been
detected so far. Some species in
which lectins have been detected,
such as Salvia bogotensis ( section
Angulatae ), S. pauciserrata ( section Flexuosae ) and S. rubescens
(section Rubescentes ), are very
diverse in Colombia with several
subspecies.
and in Salvia sclarea L. a lectin
with very interesting properties
and potential medical applications,
has been characterised (Piller et
al. 1986).
Our first purpose is to screen the
numerous (62 taxa, including subspecies) neotropical Colombian
Salvias as well as some Lepechinia,
Hyptis and Minthostachys species
for seed lectins. Our preliminary
results show that a lectin is present
R. G. Olmstead*, P. A. Reeves* and B. J. Lepschi**
*Department of Botany, University of Washington, Seattle, WA 98195, USA
**Western Australian Herbarium, Department of Conservation and Land Management
Locked Bag 104, Bentley Delivery Centre, WA 6983, Australia
As part of our ongoing molecular
systematic research into the phylogeny of the Lamiales, we have
conducted an investigation of the
hypothesis put forward by Cantino
(1992) that the Labiate tribe
Prostanthereae and the Verbenaceous tribe Chloantheae (sensu
Bentham 1876; Labiatae subfam.
Prostantheroideae and Verbenaceae subfam. Chloanthoideae
sensu Briquet 1895) together form
a monophyletic group, which also
included Tectona (Viticoideae
sensu Briquet). The evidence for
this postulated relationship comes
from a cladistic analysis of morphological and anatomical characters, in which members of these
two groups come out adjacent to
each other in an unrooted tree (Cantino 1992). However, alternative
prospective rootings of the tree differed in whether the groups formed
a clade, or a paraphyletic group
with the root attaching among them.
Prior to Cantino’s (1992) suggestion, these groups generally had
been assigned to separate families.
However, Junell (1934) included
Chloantheae and much of the rest of
the Verbenaceae, excluding subfamily Verbenoideae, in the Lamiaceae. In contrast, Hutchinson
(1959) elevated the Chloantheae to
Chloanthaceae, a move accepted in
a series of papers on the group by
Munir (Munir 1976, 1977, 1978a,
1978b, 1978c, 1979). Based on the
study cited above, Cantino et al.
7
(1992) suggested a provisional
classification for the Labiatae that
included Prostanthereae and Tectona in the subfamily Chloanthoideae, but excluded Spartothamnella from the subfamily and
excluded Acharitea and Nesogenes
from the Labiatae. The latter three
genera were included in Chloanthoideae by Briquet (1895) and
Nesogenes and Spartothamnella
were included in Chloantheae by
Bentham (1876). This study builds
upon previous studies using chloroplast DNA (cpDNA) sequences of
the gene ndhF for phylogenetic
inference in the Lamiales (Olmstead & Reeves 1995; Scotland et
al. 1995; Steane et al. 1997;
Wagstaff et al. 1998).
➣
Materials and Methods
DNA of most ingroup species was
obtained from either silica-gel-dried
material (collected by BJL) or from
herbarium specimens. Sequences of
ndhF were determined by PCRamplification of the region encompassing the first 2135 nucleotides
(nt) of tobacco ndhF (2086 nt
excluding PCR primers at each end),
followed by either manual or automated dideoxy sequencing (Olmstead & Reeves 1995). A total of 45
sequences were analysed, including
23
previously
unpublished
sequences mostly representing
Chloantheae and Prostanthereae and
22 previously published sequences
representing species from throughout the Lamiaceae and related families. Sequences were aligned by eye
with gaps inserted to accommodate
variation in length and to optimize
positional homology among species
throughout the sequence. The phylogeny was inferred by parsimony
with all changes equally weighted
using the program PAUP* (test version 4.0d61 kindly provided by D.
Swofford). Analyses were run with
100 replicate random-order entries
of the taxa using a heuristic search
strategy (TBR swapping; MULPARS). Bootstrap analysis was done
with 500 replicates, each with a single random-order entry of the taxa
and a heuristic search with TBR
swapping, but with MULPARS off.
Results and Discussion
The sequences ranged in length
from 2062 nt in Pityrodia atriplicina to 2113 nt in Stachytarpheta
dichotoma, with a typical length of
2101 nt. Several of the sequences
that were derived from herbarium
specimens needed to be amplified in
smaller fragments than typically
used and in a few cases some
regions as long as ca. 500 nt were
not able to be sequenced. More
commonly this only resulted in
scoring the PCR primer sites as
(Wagstaff & Olmstead 1997), in
which Petrea did not form a clade
with the other four genera. These
results provide strong evidence that
tribe Chloantheae does not belong
with the Verbenaceae.
Fig. 1: Strict consensus tree of 72 equally parsimonious trees. The classification
of the Chloanthoideae into tribes Chloantheae and Prostanthereae is indicated by
brackets. “C” and “P” indicate Junell’s (1934) tribes Chloantheae and Physopsideae. An asterisk highlights the placement of Spartothamnella, assigned to
Chloanthoideae by Bentham (1876), Briquet (1895) and Munir (1979).
missing (typically ca. 25 nt each for
2-3 primer sites). The insertion of
24 gaps were required for the alignment. Most gaps were unique to a
single taxon and were not used for
phylogenetic inference.
The phylogenetic analysis yielded
72 equally parsimonious trees
(length = 2258; Consistency Index =
0.56; Retention Index = 0.60). The
strict consensus of these trees is
shown in Fig. 1 along with bootstrap
values for clades and a suggested
classification based on our results.
8
The results concur with previous
studies that indicate that the Verbenaceae s.s. do not form a clade with
the Lamiaceae s.l. (Olmstead et al.
1993; Wagstaff & Olmstead 1997)
and in finding a monophyletic
Lamiaceae sensu Cantino et al.
(1992) with the inclusion of Congea
(Symphorematoideae) (Wagstaff et
al. 1998). In this study five species
of Verbenaceae s.s., representing
the four remaining tribes, are
included and together form a clade.
This result contrasts with the results
from a previous rbcL-based study
Sampling here includes fewer representatives of each of the other subfamilies of Lamiaceae than in a previous study of the family (Wagstaff
et al. 1998), but includes all genera
of Chloantheae, except the monotypic Hemiphora, and all genera of
Prostanthereae, except the monotypic Wrixonia. In a previous study,
combined analysis of rbcL and ndhF
sequences provided weak indication
that Tectona did not belong with the
Australian Prostanthereae (represented by one species of Prostanthera only; tribe Chloantheae was
not represented), but ndhF
sequences were unable, by themselves, to resolve the question of
whether Tectona belonged with
Prostanthera (Wagstaff et al. 1998).
Sampling was insufficient to
address the greater question of
chloanthoid monophyly. The results
presented here provide strong evidence (99% bootstrap support) of a
monophyletic Chloanthoideae,
excluding Tectona. In addition,
these results find strong support for
monophyletic tribes Chloantheae
(94%) and Prostanthereae (100%).
There is modest support (58%) for a
relationship of this group with Callicarpa (subfamily Viticoideae). Tectona does not appear closely related
to any other group and fits the pattern of belonging to a viticoid grade
that is more or less basal to the other
subfamilies of Lamiaceae (except
Symphorematoideae, represented
here by Congea, which may be
basal to the rest of the family,
Wagstaff et al. 1998).
Relationships within tribes
Chloantheae and Prostanthereae are
incompletely resolved, but some
preliminary conclusions seem warranted. Briquet (1895) elevated Ben-
tham’s (1876) tribe Chloantheae to
subfamily Chloanthoideae and split
it into three tribes, Achariteae
(Acharitea, Nesogenes, Pityrodia
and Spartothamnella), Chloantheae
(Chloanthes, Denisonia, Cyanostegia and Hemiphora) and Physopsideae (Dicrastylis, Lachnostachys,
Mallophora, Newcastelia and
Physopsis). Subsequently, on the
basis of gynoecial morphology,
Junell (1934) suggested the transfer
Chloanthoideae to the Labiatae at
the tribal level, with two subtribes,
Chloanthinae and Physopsidinae,
and dispersed the elements of tribe
Achariteae to other groups in and
out of the Labiatae. He transferred
Pityrodia to Chloanthinae, Spartothamnella to tribe Viticeae (Labiatae), subtribe Ajuginae, near Teucridium, and Acharitea and Nesogenes to Stilbaceae (later segregated
into Nesogenaceae by Marais,
1981). Munir (1978b, 1979) maintained two tribes within the Chloanthaceae, Chloantheae and Physopsideae, similar to Junell’s (1934) circumscriptions, but maintained the
Australian endemic Spartothamnella
in tribe Chloantheae. Additional evidence for the relationship of Spartothamnella with the Ajugeae (Teucrioideae sensu Cantino et al. 1992),
close to Teucrium, comes from
pollen morphology (Abu-Asab and
Cantino 1992; Cantino 1992) and is
confirmed by our results.
Neither of the tribes Chloantheae
and Physopsideae, as described by
Briquet (1895) and modified by
Junell (1934), is monophyletic in
any of the most parsimonious trees.
Subsequent searches in which these
groups were constrained to monophyly individually and simultaneously resulted in trees four steps
longer for a monophyletic Physopsideae, 11 steps longer for
Chloantheae (sensu Junell 1934),
and 13 steps longer for both to be
monophyletic on the same tree.
These results concur with Cantino
(1992) in finding Pityrodia to be
9
polyphyletic and having P. halganiacea as sister to the rest of the
group. These results also concur
with Cantino in finding a clade
comprising Dicrastylis and Mallophora and in finding a close relationship between Lachnostachys
and Newcastelia (a clade here, but
unresolved in Cantino 1992).
Within tribe Prostanthereae, Cantino
(1992) and Conn (1992a) concurred
in recognizing two lineages, one
comprising Prostanthera, Eichlerago and Wrixonia, and the other comprising Hemiandra, Hemigenia,
Microcorys and Westringia. Our
results are congruent with their findings. Cantino (1992) and Conn
(1992a) also agreed that Eichlerago
was derived from within Prostanthera and should belong to that
genus (Conn 1992b), a conclusion
that is consistent with our results.
Both authors also concurred on the
close relationship between Microcorys and Westringia based on the
presence of a reduced number of fertile anthers. However, our results
suggest a closer relationship
between Microcorys and Hemigenia,
suggesting an independent loss of
one pair of fertile stamens in Microcorys and Westringia. A reduction in
fertile stamen number also occurs in
Wrixonia.
Before this project is completed, it
is hoped that Hemiphora and
Wrixonia can be added to the analysis to complete sampling of recognized genera in the subfamily.
Some preliminary conclusions that
can be drawn include 1) neither
Spartothamnella nor Tectona
belong in subfamily Chloanthoideae, 2) two tribes can be recognized within Chloanthoideae,
Chloantheae and Prostanthereae
(sensu Bentham 1876), and 3) Briquet’s (1895) tribes Achariteae,
Chloantheae, and Phypsopsideae,
and Junell’s (1934) tribes
Chloantheae and Physopsideae
should not be recognized.
➣
Note added in proof: Hemiandra sp.
= H. pungens; Microcorys sp = M.
exserta; Hemigenia sp. = M. obovata.
References
Abu-Asab, M. S. and Cantino, P. D.
(1992). Pollen morphology in subfamily Lamioideae (Labiatae) and
its phylogenetic implications. In
Harley, R. M. & Reynolds, T.
(Eds.). Advances in Labiate Science: 97-112. Royal Botanic Gardens, Kew, London.
Bentham, G. (1876). Verbenaceae
and Labiatae. In Bentham, G. and
Hooker, J. D. (editors). Genera
Plantarum, 2: 1131-1223. Reeve and
Co., London.
Briquet, J. (1895). Verbenaceae. In
Engler, A. & Prantl, K. (editors). Die
Natürlichen Pflanzenfamilien, 4 (part
3a): 132-375. W. Engelmann,
Leipzig.
Cantino, P. D. (1992). Evidence for
a polyphyletic origin of the Labiatae. Ann. Missouri Bot. Gard. 79:
361-379.
Cantino, P. D., Harley, R. M. &
Wagstaff, S. J. (1992). Genera of
Labiatae: Status and Classification.
In Harley, R. M. & Reynolds, T.
(Eds.). Advances in Labiate Science: 511-522. Royal Botanic Gardens, Kew, London.
Conn, B. J. (1992a). Relationships
within the tribe Prostanthereae
(Labiatae). In Harley, R. M. &
Reynolds, T. (Eds.). Advances in
Labiate Science: 55-64. Royal
Botanic Gardens, Kew, London.
Conn, B. J. (1992b). Status of the
genus Eichlerago (Labiatae). Telopea 44: 649-651.
Hutchinson, J. (1959). The families
of flowering plants, 2nd ed. Vol. 1.
Oxford University Press, London.
Junell, S. (1934). Zur Gynäceummorphologie und Systematik der
Verbenaceen und Labiaten. Symbolae Botanicae Upsalienses 4: 1-219.
Marais, W. (1981). Two new
gamopetalous families, Cyclocheilaceae and Nesogenaceae, for extraAustralian Dicrastylidaceae. Kew
Bull. 35: 797-812.
Munir, A. A. (1976). A taxonomic
revision of the genus Spartothamnella (Chloanthaceae). J. Adelaide
Bot. Gard. 1: 3-25.
Munir, A. A. (1977). A taxonomic
revision of the genus Chloanthes
(Chloanthaceae). J. Adelaide Bot.
Gard. 1: 83-106.
Munir, A. A. (1978a). A taxonomic
revision of the genus Cyanostegia
(Chloanthaceae). Brunonia 1: 45-67.
Munir, A. A. (1978b). Taxonomic
revision of Chloanthaceae tribe
Physopsideae. Brunonia 1: 407-692.
Munir, A. A. (1978c). A taxonomic
revision of the genus Hemiphora
(Chloanthaceae). J. Adelaide Bot.
Gard. 1: 161-166.
Munir, A. A. (1979). A taxonomic
revision of the genus Pityrodia
(Chloanthaceae). J. Adelaide Bot.
Gard. 2: 1-138.
Olmstead, R. G., Bremer, B., Scott,
K., and Palmer, J. D. (1993). A parsimony analysis of the Asteridae sensu
lato based on rbcL sequences. Ann.
Missouri Bot. Gard. 80: 700-722.
Olmstead, R. G. and Reeves, P. A.
(1995). Evidence for the polyphyly
of the Scrophulariaceae based on
chloroplast rbcL and ndhF
sequences. Ann. Missouri Bot. Gard.
82: 176-193.
Scotland, R. W., Sweere, J. S.,
Reeves, P.A. and Olmstead, R. G.
(1995). Higher level systematics of
Acanthaceae determined by chloroplast DNA sequences. Amer. J. Bot.
82: 266-275.
Steane, D. A., Scotland, R.W., Mabberley, D. J., Wagstaff, S. J.,
Reeves, P.A. and Olmstead, R. G.
(1997). Phylogenetic relationships
of Clerodendrum s. l. (Lamiaceae)
inferred from chloroplast DNA.
Syst. Bot. 22: 229-244.
Wagstaff, S. J. and Olmstead, R. G.
(1997). Phylogeny of the Labiatae
and Verbenaceae inferred from rbcL
sequences. Syst. Bot. 22: 165-179.
Wagstaff, S. J., Reeves, P. A.,
Hickerson, L., Spangler, R. E., and
Olmstead, R. G. (1998). Phylogeny
of Labiatae s. l. inferred from
cpDNA sequences. Pl. Syst. Evol.
209: 265-274. ❑
10
of chemical compounds with recognized sedative properties. Samples
were collected and voucher specimens were deposited at the ICN
herbarium.
ETHNOBOTANY AND
ANTICONVULSANT
PROPERTIES OF
LAMIACEAE FROM
RIO GRANDE DO SUL
(BRAZIL).
Coelho de Souza, G.P.1 and
Elisabetsky, E.1,2
Curso de Pós-graduação em Ciências Biológicas - Botânica, IB;
2 Laboratório de Etnofarmacologia,
ICBS, Universidade Federal do Rio
Grande do Sul, CP 5072, 90041-970,
Porto Alegre, RS, Brazil
1
Introduction
The Lamiaceae is one of the most
diverse and widespread plant
families in terms of ethnomedicine. The medicinal value of
Lamiaceae species is largely
based on their volatile oils
(Moerman, 1991). Regarding its
effects on the central nervous
system (SNC), studies proved
anticonvulsant activities in at
least four genera, among which
Aeollanthus (Elisabetsky et al.,
1995a), Leonurus (Chauhan,
1988), and Salvia (González,
1990) are represented in the state
of Rio Grande do Sul (RS), Brazil
(Fig. 1). According to Ab’Sáber
(1971) there are three vegetation
domains in this region, Undulating grasslands, Atlantic Forest
and Araucaria Mixed Forest. In
RS the genera Ocimum, Hyptis,
Cunila and Salvia are the best
represented. The genus Ocimum
with about 65 species, six endemic to South America, has its center of diversity in Tropical Africa
(Pereira, 1979). Hyptis has some
350 neotropical species between
the south of USA and Argentina,
with 17 species in RS (Bordignon, 1990). Cunila has two
centers of diversity, Mexico and
Subtropical South America with
11 species in RS. These are often
Fig. 1 - Localization of Rio Grande do Sul (27°- 34° S latitude; 49°40′- 57° 30′ W
longitude).
very aromatic and are frequently
used in traditional medicine (Bordignon, 1997). Salvia is the best
represented genus in RS (Epling
& Toledo, 1943) with 23 species
(Coelho de Souza, 1997).
This present paper contributes to the
ethnobotanical knowledge of Lamiaceae from RS. Following an
ethnopharmacological study of Aeollanthus suaveolens, which is used by
Amazonian “caboclos” against
epilepsy, and which led to the identification of two volatile substances
(linalool and g-decanolactone) active
as anticonvulsants (Elisabetsky et al.,
1995a; Coelho de Souza et al., 1997),
the present study especially focused
on species from the Subfamily
Nepetoideae used as sedatives/anticonvulsants.
Methods
Ethnobotany: An inventory of Lamiaceae species in RS was made in the
herbaria of Porto Alegre Colégio
Anchieta - PACA, Instituto de Ciências Naturais - ICN of the Universidade Federal do Rio Grande do Sul
and Universidade Federal de Pelotas
- PEL. Ethnobotanical information,
as well as chemical and pharmacological data available for the species
listed in the inventory were collected
from scientific and lay literature.
The analysis of these data guided the
selection of species for pharmacological tests. The criteria for selection were: 1) belonging to the
Nepetoideae, 2) being easily available and showing broad distribution
in RS, 3) being traditionally used as
a sedative, 4) known to have classes
11
Phytochemistry: Methods for
extraction of essential oils and pharmacological analysis have been
detailed elsewhere (Elisabetsky et
al., 1995a, Coelho de Souza, 1997).
In short, essential oils were extracted from 50-100g samples (leaves,
inflorescence and terminal buds) by
hydrodistillation (Clevenger apparatus). Essential oils were extracted
from Cunila galioides, Cunila menthoides, Cunila microcephala,
Cunila spicata, Glechon thymoides,
Hyptis mutabilis, Leonurus sibiricus
[Editorial note: This almost certainly refers to the species Leonurus
japonicus Houtt.], Nepeta cataria,
Ocimum selloi, Rosmarinus officinalis, Salvia officinalis. The pharmacological method was based on
that of Gladding et al. (1985),
focussing on pentylenetetrazol
(PTZ, 90 mg/kg, sc.)-induced convulsions (in mice, thirty minutes
after i.p. treatment, with doses ranging from 100 -700 mg/kg (i.p.) of
essential oils). Diazepam (1.0
mg/kg i.p.) and phenobarbital (30
mg/kg i.p.) were used as reference
drugs. Animals were observed for
60 min. for the presence of clonic
convulsions lasting more than 3 seconds. Results were analysed by
means of Fisher exact test.
Results
Through the herbarium survey, 125
species were listed, 49 being exotic
to and 76 native in RS. Native
species are distributed in 17 genera
including Aegiphila (4 spp.), Cunila
(10 spp.), Glechon (5 spp.), Hedeoma (3 spp.), Hesperozygis (2 spp.),
Hoehnea (2 spp.), Hyptis (17 spp.),
Marsypianthes (1 sp.), Ocimum (4
spp.), Peltodon (1 sp .), Rhabdocaulon (5 spp.), Salvia (14 spp.),
Satureja (1 spp.), Scutellaria (2 ➣
spp.), Stachys (2 spp.), Teucrium (2
spp.), and Vitex (1 spp.). The ethnobotanical survey shows that
53.4% of the species do not have
recorded traditional uses, while
36.4% are considered of medicinal
value. The medicinal uses are
diverse, mainly focussing on digestive (14.2%), respiratory (11.9%),
central nervous system ailments
(16.2%) and topically applied for
wound healing (11.5%). Lamiaceae
species are also used as seasonings
(15.3%), ornamentals (20.8%),
species used by bees for honey production (13.9%) or for other industrial purposes (13.9%), mainly as
cosmetics and personal care products. Among native species, 28%
are utilized as medicines.
500mg/kg, leading to convulsions
and death. Cunila galioides essential oil induced a dose related protection, reaching a maximum at
300mg/kg. The ED50 (95% confidence limit) was estimated to be
217.7 (14.0-367.6) mg/kg and the
LD50 (95%) was 398 (280.23057.2) mg/kg. Oils from Cunila
menthoides, Cunila microcephala,
Hyptis mutabilis, Leonurus sibiricus [Editorial note: This almost
certainly refers to the species
Leonurus japonicus Houtt.], Ocimum selloi, Rosmarinus officinalis,
were ineffective up to 700mg/kg
(data not shown).
Regarding phytochemical and
pharmacological data, we found
that 71.6% of species had not been
the subject of previous studies.
Phytochemical and/or pharmacological data are available for 20.7%
of the exotic and only 7.8% of
native species.
Cunila galioides is called “poejinho” due to its likeness with
Mentha pulegium and C. microcephala (known as “poejos”),
being used traditionally for the
same purposes (expectorant, tranquilizer and laxative) (Lopes et
al., 1988). Few studies are available for C. galioides, which is
restricted to southern Brazil. Preliminary phytochemical studies of
its essential oil shows the presence of neral, geranial and
linalool (Bordignon, 1997).
Linalool proved to be active as an
anticonvulsant in mice (Elisabetsky et al., 1995a) with relevant
neurochemical properties (Elisabetsky et al., 1995b). Citral has
sedative activity (Duke, 1992)
while neral and geranial are citral
isomers with no reported pharmacological studies. Therefore, the
abundant presence of linalool,
neral and geranial is likely to be
related to C. galioides essential
oil anticonvulsant activity.
Cunila spicata essential oil was
effective at 400 and 500mg/kg producing 66.7% of protection. Glechon thymoides essential oil was
effective at 500mg/kg producing
60% of protection while the Nepeta
cataria essential oil was effective at
700mg/kg producing 66.7% of protection. Salvia officinalis essential
oil was found to be toxic at
Salvia officinalis is considered as a
“cure-all” and, accordingly, indicated
for a myriad of uses (Font Quer,
1992; De Feo & Senatore, 1993).
When administered i.p. to mice S.
officinalis essential oil induced lethal
convulsions; this pattern of neurotoxicity is also found with opioids and
local anesthetics. Although phytochemical and pharmacological stud-
The exotic species are mostly cultivated and show a larger number of
ethnobotanical uses, reflecting the
influx of European immigrants and
culture to the State. It is likely that
some of the medicinal uses of native
species are related to their morphological and/or aromatic resemblance
to exotic species with medicinal uses
diffused throughout the world. Of the
exotic species, 86.7% (39) are cultivated while 13.3% (7) are adventives; among the cultivated species
84.6% (33) belong to the subfamily
Nepetoideae.
12
ies of this species are abundant
(Ribeiro et al., 1986, Masterova et
al., 1989, Rutherford et al., 1992,
Asanov et al., 1994, Tada et al.,
1994), the toxic effect revealed by
this study needs further investigation.
Conclusion
Lamiaceae species have been used
all over the world for seasoning,
perfumes and/or medicines (TomasBarberán, 1986; Heinrich, 1992;
Lawrence, 1992). Recently, aromatic species extracts and distillates
have become important raw material for the cosmetic, food and personal care industries (Lawrence,
1992). This study of Lamiaceae of
RS showed the importance of the
family for local communities and
revealed the paucity of phytochemical and pharmacological studies of
native species. Analysis of data
gives a rational basis for some traditional uses of Lamiaceae species
and suggests that this family is a
source of yet unknown medicinal
resources. It is hoped that these
conclusions can add value to the
flora of Rio Grande do Sul and its
associated traditional culture, and
therefore contribute to its conservation.
Acknowledgements
This work was supported by grants
from CNPq, FAPERGS and PREBELAC. The authors are grateful
to Sérgio Bordignon (UFRGS) for
essential oil samples and assistance
with botanical issues, Fernanda
Gaieski for assistance with pharmacological experiments and José
P. Coelho de Souza for editorial
assistance.
Note: A list of the labiate species
(both native and introduced) occuring in RS is available from Elaine
Elisabetsky (elisasky@vortex.
ufrgs. br) on request.
References and Bibliography
Ab’Sáber, A. N. (1971). A organização das paisagens inter e subtropicais brasileiras. Anais do III simpósio sobre o cerrado. Editora Edgard
Blücher Ltda, São Paulo, 1-14.
Asanov, E. B., Gaevskii, A. V.,
Tareeva, N. V. & Glyzin, N. I.
(1994). Spectrophotometric method
for analysing the amount of
royleanones in oil Salvia officinalis
root extract. Khim.-Farm. Zh.
28(2): 32-33.
Bordignon, S. A. L. (1990). O
gênero Hyptis Jacq. (Labiatae) no
Rio Grande do Sul. Universidade
Federal do Rio Grande do Sul,
Porto Alegre. M.Sc. Thesis.
Bordignon, S. A. L. (1997). Estudo
botânico e químico de espécies de
Cunila Royen ex L. (Labiatae) nativas do sul do Brasil. Universidade
Federal do Rio Grande do Sul,
Porto Alegre. Ph.D Thesis.
Chauan, A. K., Dobhal, M. P. &
Joshi, B. (1988). A review of medicinal plants showing anticonvulsant
activity. J. Ethnopharmacol. 22: 1-23.
Coelho de Souza, G. P., Elisabetsky, E., Nunes, D. S., Rabelo, S. K.
L. & Nascimento da Silva, M.
(1997). Anticonvulsant properties
of g-decanolactone in mice. J.
Ethnopharmacol. 58: 175-181.
Coelho de Souza, G. P. (1997). Estudo etnobotânico da família Lamiaceae no Rio Grande do Sul, com
ênfase na busca de espécies com propriedades nticonvulsivantes. Universidade Federal do Rio Grande do
Sul, Porto Alegre. M.Sc. Thesis.
De Feo, V. & Senatore, F. (1993).
Medicinal plants and phytotherapy
in the Amalfitan Coast, Salerno
Province, Campania, Southern Italy.
J. Ethnopharmcol. 39: 39-51.
Duke, J. A. (1992). Handbook of
biologically active phytochemicals
and their activities, CRC Press,
Boca Ranton, Florida, pp. 3-183.
Elisabetsky, E., Coelho de Souza,
G. P., Siqueira, I. R., Amador, T. A.
& Correa, M. A. (1995a). Sedative
properties of linalool. Fitoterapia
66(5): 407-414.
Elisabetsky, E., Marschner, J. &
Souza, D. O. (1995b). Effects of
linalool on glutamatergic system in
the rat cerebral cortex. Neurochemical Research 20(4): 461-465.
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Paulo, Brasil, V XLVIII, 107p.
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13 a ed. Editorial Labor, S.A.
Barcelona.
Gladding, G. D., Kupferberg, H. J.
& Swinyard, E. A. (1985). In H. H.
Frey and D. Janz (Editors). Antiepileptic Drugs, Handbook of
Experimental Pharmacology,
Springer-Verlag: 74: 341-347.
González, A. G., Luis, J. G. & Ravelo, A. G. (1990). IV Nepeta. Estudio fitoquímico. In: Plantas
iberoamericanas. Fuentes de moleculas bioactivas II Lamiaceae. vol.2,
Tenerife, Espanha, pp.195-197.
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botany of american Labiatae. In
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T.(Editors). Advances in Labiate
Sciences, pp. 475-488. Royal
Botanic Gardens, Kew.
Lawrence, B. M.(1992). Chemical
components of Labiatae oils and
their exploitation. In Harley, R. M.
and Reynolds, T.(Editors).
Advances in Labiate Sciences, pp.
399-436. Royal Botanic Gardens,
Kew.
Lopes, A. M. V., Eisinge, S. M.,
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M. H. C., Grassioli, D., Dellaméa,
V. L. & Siqueira, V. L. E. (1988).
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V. E. & Suchy, V. (1989). Phytochemical study of Salvia officinalis
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1-42.
13
Pereira, C. (1979). As espécies do
gênero Ocimum L. (Labiatae) da
América do Sul. Universidade Federal do Rio de Janeiro, Rio de
Janeiro, M.Sc. Thesis.
Ribeiro, R. A, Melo, M. R. F., Barros, F., Gomes, C., & Trolin, G.
(1986). Acute antihypertensive
effect in conscious rats produced by
some medicinal plants used in state
of São Paulo. J. Ethnopharmacol.
15(3): 261-269.
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C., Hansen, S. K., Witt, M. R.,
Bergendorff, O. & Sterner, O.
(1992). Isolation and identification
from Salvia officinalis of two diterpenes which t-butyl bicyclophosphoro [sulphur-35] thionate binding
to chloride channel of rat cerebrocortical membranes in vitro. Neurosci. lett. 135(2): 224-226.
Tada, M., Okuno, K., Chiba, K.,
Ohnishi, E. & Yoshii, T. (1994).
Antiviral diterpenes from Salvia
officinalis. Phytochem. 35(2):
539-541.
Tomás-Barberán, F. A. (1986). The
flavonoid compounds from the Labiatae. Fitoterapia, 57(2): 67-95. ❑
PHLOMIC ACID IN
LAMIOIDEAE SEED
OILS
Kurt Aitzetmüller1 and
Nanzad Tsevegüren2
1Institute for Chemistry and Physics
of Lipids, BAGKF, Piusallee 76, D48147 Münster, Germany
2Laboratory for Lipid Chemistry,
Chemistry Institute, Mongolian
Academy of Sciences, Shukov Str.
54, Ulan Bator 51, Mongolia
Products with an allene structure
(C=C=C) are very rare in nature
because their synthesis and biosynthesis is rather difficult. The only
major natural allene known to be
widely occurring is neoxanthin, one
of the yellow pigments in green
leaves. On the other hand, allenes ➣
were known to occur in a number of
seed oils since the early investigations of Bagby, Hagemann and coworkers (Bagby et al. 1965, Hagemann et al., 1967). The allenes
which were found by infrared spectroscopy were later characterized as
a fatty acid with 18 carbon atoms
and a ∆5,6allene structure (Fig. 1).
Fig. 1: Structures of laballenic, phlomic and lamenallenic acids and of the
more significant 20:1 fatty acids,
20:1D9cis and 20:1D11cis.
This fatty acid was thoroughly
investigated and characterized, and
was called laballenic acid (Bagby et
al., 1965), because its occurrence
was known only from plants belonging to the family Labiatae. With
very few exceptions, the early work
by Hagemann et al. (1967) showed
that laballenic acid occurred only in
members of what is now considered
as the subfamily Lamioideae by
Cantino et al. (1992).
In addition to laballenic acid, another allenic fatty acid was later discovered which had an additional
double-bond in position ∆16 (Mikolajczak et al., 1967). This fatty acid
was called lamenallenic acid,
because it was found only in the
seed oils of species belonging to the
genus Lamium (Mikolajczak et al.,
1967). This fatty acid so far has
never been observed in any other
Labiate seed oil outside genus
Lamium. We too found it only in
Lamium (L. maculatum being the
only species investigated here), but
not in other genera of Labiatae except possibly for traces (see Table
1) requiring confirmation.
Our own work (Aitzetmüller, 1997;
Tsevegüren et al., 1997;
Aitzetmüller et al., 1997) on seed
oil fatty acid fingerprints
(Aitzetmüller, 1993) so far confirmed the occurrence of laballenic
acid in seed oils of a number of
species belonging to subfamily
Lamioideae as defined by Cantino
et al. (1992). With the exception of
only one sample, obtained from
Bulgaria and labelled as Stachys
alpina, laballenic acid was present
in all members of the Lamioideae
investigated by us (about 20 spp.),
and at the same time it was absent
in all members of other Labiatae
subfamilies. So we currently believe
that the presence of allenic fatty
acids is a highly characteristic feature of this one subfamily. (The single sample of S. alpina which
proved the exception, may perhaps
be due to a misidentification.)
Figure 2 shows a number of typical
fatty acid fingerprints (Aitzetmüller,
1993) as obtained from a few members of the Labiatae plant family. As
usual we have labelled only those
fatty acids in the gas chromatograms of Figure 2, which are
“unusual” in the sense that they do
not occur in every normal seed oil.
We believe that these “fatty acid
fingerprints” are of chemotaxonomic value (Aitzetmüller, 1993;
Aitzetmüller, 1995) and can be used
in much the same way as one would
use “leaf shape” or “pollen surface
structure” in taxonomic work in
botany. The presence or absence of
these unusual fatty acids, at levels
above a few tenths of a percent at
least, is genetically determined (or,
to be more exact, the presence or
14
absence of the enzymes necessary
for the biosynthesis of this particular fatty acid depends on the presence and expression of the appropriate genes coding for them). The
presence of fatty acids with unusual
structures in the seed oils of closely
related genera can therefore also be
considered to be an important indicator of phylogenetic evolution
(Aitzetmüller, 1996).
Recently, phlomic acid was discovered and described (Aitzetmüller et
al., 1997) as a minor additional
allenic fatty acid occurring in certain representatives of the subfamily
Lamioideae.
Phlomic
acid
Table 1: Occurrence of laballenic acid (18:2 ∆5,6allene), phlomic acid (20:2 ∆7,8allene) and related fatty acids
(in % of total fatty acids) in seed oils of selected members of Labiatae: Lamioideae (this investigation).
Plant species
fat in
seed %
laballenic
acid
lamen-allenic
acid
11.8
28.5
12.5
32.1
34.0
34.5
16.0
30.4
32.9
41.1
22.7
31.6
33.1
25.1
18.1
13.0
12.0
11.8
11.7
10.3
10.1
9.1
8.6
7.6
7.2
6.1
[0.2]1
[0.2]1
-
3.3
1.3
0.7
0.4
0.2
0.7
1.4
0.6
0.3
0.6
0.3
0.4
0.7
2.9
0.3
0.5
[0.2]1
0.1
0.2
0.2
0.1
[0.1]1
0.3
0.2
[tr.]1
0.4
32.0
24.3
35.0
30.8
-
4.5
3.9
2.7
2.7
2.6
8.8
-
0.2
0.2
0.2
0.2
[tr.]1
0.3
[tr.]1
[0.1]1
Phlomis tuberosa
*Leonurus sibiricus
Phlomis fruticosa
Marrubium vulgare
Sideritis hyssopifolia
Leonurus cardiaca
Panzerina canescens
Ballota nigra
Molucella laevis
Physostegia virginiana var. alba
Stachys recta
Stachys palustris
Lamiastrum galeobdolon
Stachys byzantina
Phomis samia hort.2
Lamium maculatum
Galeopsis tetrahit
Galeopsis speciosa
1tentative
20:1∆9c
phlomic
acid
identification
P. russelliana
2presumably
[*Editorial note: Leonurus sibiricus, mentioned above, almost certainly refers to Leonurus japonicus Houtt.]
Fig. 2: Capillary gas chromatographic
fatty acid fingerprints showing the
presence of phlomic acid next to laballenic acid and 20:1D9cis in Phlomis
and Ballota, and of lamenallenic acid
in Lamium. (Unusual fatty acids only
are labelled).
(20:2∆7,8allene) seems to be the
chain-elongation product of laballenic acid, and it may have been
synthesized by the addition of a C2unit to the carboxyl end of laballenic acid - i.e., as usual in the
chain-elongation of normal fatty
acids. However, the chain-elongation of fatty acids which contain
double-bonds in position ∆5 or ∆6
of the fatty chain so far was not
known in plants (Aitzetmüller and
Tsevegüren, 1994; Aitzetmüller et
al., 1997). The finding is also significant for another reason: The
presence of small amounts of
phlomic acid seems to be correlated
with the occurrence of an unusual
20:1 fatty acid, 20:1∆9cis or 20:1n11. In most other seed oils, including those of most Labiatae,
20:1∆11cis (or 20:1n-9) is the
usual, and often the only, 20:1 fatty
acid present. Only in those species,
where small amounts of phlomic
acid occur, the gas chromatographic
peak representing 20:1∆9cis is larg-
er than the neighbouring peak of
20:1∆11cis. In species where
phlomic acid is absent, the peak size
ratio of the two 20:1 fatty acids is
always the other way round, or
20:1∆9cis is not present at all.
The chemotaxonomic significance
of the prescence or absence of
phlomic acid in the Lamioideae is
not yet known. This fatty acid was
present - at very low levels - in several species examined (Table 1). It
has not been found in Lamium and
in a large number of other genera,
where the seed oils contain laballenic acid. It was found, however, in
one sample of Lamiastrum galeobdolon (Aitzetmüller et al., 1997).
Cantino et al. (1992), in their revision of the Labiatae made no
attempt to subdivide their subfamily
Lamioideae into tribes or other subunits. The seed oil fatty acid patterns, however, may suggest this.
15
Acknowledgements
The authors gratefully acknowledge
experimental support by Gisela
Werner and Dr. K. Vosmann. N. T.
also acknowledges financial support
by the Alexander-von-HumboldtFoundation, Bonn, Germany.
References
Aitzetmüller, K. (1993). Capillary
GLC Fatty Acid Fingerprints of
Seed Lipids - A Tool in Plant
Chemotaxonomy? J. High Resol.
Chromatogr., 16: 488-490.
Aitzetmüller, K. (1995). Fatty
Acid Patterns of Ranunculaceae
Seed Oils: Phylogenetic Relationships. Plant Syst. Evol. [Suppl.] 9:
229-240.
Aitzetmüller, K. (1996). Seed Fatty
Acids, Chemotaxonomy and
Renewable Resources. In: OilsFats-Lipids 1995: Proceedings of
the 21st World Congress of the
International Society for Fat ➣
Research. P. J. Barnes & Associates, High Wycombe, pp. 117-120.
Aitzetmüller, K. (1997). Seed oil
fatty acids in the Labiatae. Lamiales
Newsletter (Kew/London) 5: 3-5.
Aitzetmüller, K. & Tsevegüren, N.
(1994). Seed Fatty Acids, “FrontEnd” - Desaturases and Chemotaxonomy - A Case Study in the
Ranunculaceae. J. Plant Physiol.
143: 538-543.
Aitzetmüller, K., Tsevegüren, N. &
Vosmann, K. (1997). A new allenic
fatty acid in Phlomis (Lamiaceae)
seed oil. Fett - Lipid 99: 74-78.
Bagby, M. O., Smith, C. R. &
Wolff, I. A. (1965). Laballenic acid.
A new allenic acid from Leonotis
nepetaefolia Seed Oil. J. Org.
Chem. 30: 4227-4229.
Cantino, P. D., Harley, R. M. &
Wagstaff, S. J. (1992). Genera of
Labiatae: Status and Classification.
In Harley, R. M. & Reynolds, T.
(Editors), Advances in Labiate Science. Royal Botanic Gardens, Kew,
pp. 511-522.
Hagemann, J. M., Earle, F. R.,
Wolff, I. A. & Barclay, A. S.
(1967). Search for New Industrial
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Lipids 2: 371-380.
Mikolajczak, K. L., Rogers, M. F.,
Smith, C. R. & Wolff, I. A. (1967).
An octadecatrienoic acid from
Lamium purpureum L. seed oil containing 5,6-allenic and trans-olefinic
unsaturation. Biochem. J. 105:
1245-1249.
Tsevegüren, N., Aitzetmüller, K. &
Werner, G. (1997). Fatty Acids of
some Central Asian Labiatae. Lamiales Newsletter (Kew/London) 5:6-8.
Note added: If one of our readers
could supply us with authentic
Lamium galeobdolon, L. amplexicaule and Stachys alpina seed from
an independent source, and/or with
seeds of very close other relatives
of Lamium, e. g. another Lamiastrum spp. or variety, we should like
to investigate this, too:Please contact us at our E-mail
address: aitzetm@uni-muenster.de ❑
BIBLIOGRAPHY OF RECENT TAXONOMIC
PUBLICATIONS ON THE LAMIALES
The following list of publications
has been abstracted from the Kew
Record of Taxonomic Literature
and from the Kew Economic
Botany Bibliographic Database
(November 1996 - February 1998),
and we are, again, extremely grateful to the editors and compilers for
their assistance in preparing this
bibliography. Regrettably our selection from the Economic Botany
Database is not complete due to
space constraints (the original list
ran to c. 80 pages!). As was the case
for the last newsletter, some authors
have sent us notification of publications not listed on the “Kew
Record” and these have also been
included. Where possible articles
are listed under the applicable
genus, or occasionally, tribe or family - and are arranged alphabetically. Where a number of references to
different taxa in the Lamiales are
made in a single paper they have
tended to be listed under the “General” heading. Any title completely
enclosed within brackets is the English translation of a title written in a
“symbol” font such as Arabic, Chinese, Japanese or Russian. All diacritical marks have been removed to
facilitate editing.
GENERAL
Zakirova, RO, Nafanailova, II,
Abdulina, SA. Kariologicheskoe
izuchenie nekotorykh predstavitelei
semeistva Dipsacaceae Lindl., sem.
Lamiaceae Lindl., sem. Rutaceae
Juss. Izv. Natsion. Akad. Nauk
Resp. Kazakhstan, Ser. Biol.
4(190): 96-98 (1995-1996).
Arreguin Sanchez, M de la L,
Palacios Chavez, R, Quiroz Garcia, DL. Morfologia de los granos
de polen de la familia Verbenaceae
del Valle de Mexico. Phytologia
80(5): 329-342 (1996).
16
Budantsev, AL, Lobova, TA.
Fruit morphology, anatomy and
taxonomy of tribe Nepeteae (Labiatae). Edinburgh J. Bot. 54(2):
183-216 (1997).
Correa, AM da S, Esteves, LM.
Flora polinica da Reserva do Parque Estadual das Fontes do Ipiranga (Sao Paulo, Brasil). Familia 144
Labiatae. (Pollen flora of the
“Reserva do Parque Estadual das
Fontes do Ipiranga” (Sao Paulo,
Brazil). Family 144 Labiatae).
Hoehnea 23(1): 141-145 (1996).
Hervas Serrano JL, Penafiel
Trueba, MS, Fernandez Ocana,
AM, Fernandez Lopez, C. Labiadas del Alto Guadalquivir hasta
1.995. Blancoana no.14: 93-103
(1997).
Ibarra Manriquez, G, Sinaca
Colin, S. Estacion de Biologia
Tropical “Los Tuxtlas”, Veracruz,
Mexico: lista floristica commentada
(Mimosaceae a Verbenaceae). Rev.
Biol. Trop. 44(1): 41-60 (1996).
Marin, PD. Nutlet sculpturing of
selected species from Ajugoideae,
Scutellarioideae and Stachyoideae
(Lamiaceae). Bull. Inst. Jard. Bot.
Univ. Beograd 26-27: 21-29 (19921993 publ. 1994).
Martinez, S, Botta, S, Mulgura,
ME. Morfologia de las inflorescencias en Verbenaceae-Verbenoideae: 1. Tribu Verbeneae.
(Morphology of inflorescences in
Verbenaceae-Verbenoideae: 1.
Tribe Verbeneae). Darwiniana
34(1-4): 1-17 (1996).
Orfila, EN, Farina, EL. Flora del
valle de Lerma: Lamiaceae Barnh (
=Labiatae Juss.). Aportes Bot. Salta
Ser. Flor. 4(2): 67p. (1996).
Vasic, O. A survey of the Mediterranean species of Lamiaceae family
in the flora of Serbia. Lagascalia
19(1-2): 263-270 (1997).
Von Poser, GL, Toffoli, ME,
Sobral, M, Henriques, AT. Iridoid
glucosides substitution patterns in
Verbenaceae and their taxonomic
implication. Pl. Syst. Evol. 205(34): 265-287 (1997).
Wagstaff, SJ, Olmstead, RG. Phylogeny of Labiatae and Verbenaceae inferred from rbcL
sequences. Syst. Bot. 22(1): 165179 (1997).
AEGIPHILA
Lopez Palacios, S. Novedades en
Verbenaceae de Venezuela: Aegiphila. Pittieria no. 5: 9-49 (1973).
Wallnöfer, B. New or noteworthy
species of Aegiphila, Styrax and
Zamia from Peru. Linz. Biol. Beitr.
28(2): 1053-1060 (1996).
AEOLLANTHUS
Souza, GPC de, and others. Anticonvulsant properties of ydecanolactone in mice. J. Ethnopharmacol. 58(3): 175-181 (1997).
AJUGA
Munguti, K. Indigenous knowledge
in the management of malaria and
visceral leishmaniasis among the
Tugen of Kenya. Indig. Knowl.
Devel. Monit. 5(1): 10-12 (1997).
Ruiz de Clavijo, E. A comparative study of the reproductive biology of two Ajuga species (Lamiaceae) in the southwest of the Iberian Peninsula. Int. J. Pl. Sci.
158(3): 282-291 (1997).
ARCHBOLDIA
De Kok, RPJ, Atkins, S. The
genus Archboldia E. Beer and H.J.
Lam is put into the synonymy of
Clerodendrum L. (Labiatae). Kew
Bull. 52(2): 503-504 (1997).
AVICENNIA
Parani, M, Lakshmi, M, Elango,
S, Ram, N, Anuratha, CS & Parida, A. Molecular phylogeny of
mangroves: 2. Intra- and inter-specific variation in Avicennia revealed
by RAPD and RFLP markers.
Genome 40(4): 487-495 (1997).
AVICENNIACEAE
Sanders, RW. The Avicenniaceae
in the southeastern United States.
Harvard Pap. Bot. no.10: 81-92
(1997).
BRAZORIA
Turner, MW. Systematic study of
the genus Brazoria (Lamiaceae),
and Warnockia (Lamiaceae) a new
genus from Texas. Pl. Syst. Evol.
203(1-2): 65-82 (1996).
CALAMINTHA
Karousou, R, Kokkini, S,
Bessiere, JM, Vokou, D.
Calamintha cretica (Lamiaceae), a
Cretan endemic: distribution and
essential oil composition. Nordic J.
Bot. 16(3): 247-252 (1996).
Morales, R, Luque, MN. El genero
Calamintha Mill. (Labiatae) en la
Peninsula Iberica e Islas Baleares.
(The genus Calamintha Mill. (Labiatae) from the Iberian Peninsula and
the Balearic Islands). An. Jard. Bot.
Madrid 55(2): 261-276 (1997).
CALCHAS
Heath, PV. A new generic name in
Lamiaceae: part 1. Calyx 5(4): 160
(1997).
CALLICARPA.
Atkins, S. Two new species of Callicarpa (Verbenaceae) from Brunei.
Kew Bull. 52(1): 227-230 (1997).
CITHAREXYLUM
Aymard, GA, Rueda, RM. A new
species of Citharexylum (Verbenaceae) from Ecuador. Novon 7(2):
95-97 (1997).
CLERODENDRUM
De Kok, RPJ, Atkins, S. The
genus Archboldia E. Beer and H.J.
Lam is put into the synonymy of
Clerodendrum L. (Labiatae). Kew
Bull. 52(2): 503-504 (1997).
Inoue, K, Hasegawa, M,
Kobayashi, S. A new species of
Clerodendrum (Verbenaceae) from
the Izu Islands. J. Jap. Bot. 72(2):
117-124 (1997).
17
Roy, R, and others. Antifungal
activity of the flavonoids from
Clerodendron infortunatum roots.
Fitoterapia 67(5): 473-474 (1996).
Singh, RS, and others. A new
hydroxyketone from Clerodendron
colebrookianum leaves. Fitoterapia
6: 548-549 (1997).
Steane, DA, Scotland, RW, Mabberley, DJ, Wagstaff, SJ, Reeves,
PA, Olmstead, RG. Phylogenetic
relationships of Clerodendrum s.l.
(Lamiaceae) inferred from chloroplast DNA. Syst. Bot. 22(2): 229243 (1997).
Tian, J, and others. New
cleroindicins from Clerodendron
indicum. J. Nat. Prod. 60(8): 766769 (1997).
Zhu, M, and others. Application of
radioligand receptor binding assays
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