www.unal.edu.co/icn/publicaciones/caldasia.htm
Freire et
al.
Caldasia 29(1):23-38.
2007
EPIDERMAL CHARACTERS OF BACCHARIS
(ASTERACEAE) SPECIES USED IN TRADITIONAL
MEDICINE
Caracteres epidérmicos de las especies del género Baccharis
(Asteraceae) usadas en la medicina popular
SUSANA E. FREIRE
ESTRELLA URTUBEY
DANIEL A. GIULIANO
División Plantas Vasculares, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina.
freire@museo.fcnym.unlp.edu.ar
ABSTRACT
A morphological study of 38 species of Baccharis used in traditional medicine
was carried out to provide some epidermal characters that will contribute to the
knowledge of the genus. The present study revealed: 1) seven different types of
trichomes: conical, aseptate flagellate, filiform flagellate, 1-armed, 2-4-armed,
bulbiferous flagellate, and glandular biseriate; 2) that 28 of the total of 38 species
have trichomes in tufts; 3) six different types of stomata: anomocytic, anisocytic,
cyclocytic, actinocytic, tetracytic, and staurocytic; 4) that some trichome types,
such as 2-4-armed (B. dracunculifolia) and aseptate flagellate branched (B. trinervis), show a high diagnostic value; 5) that the stomata types can be used to differentiate species with similar trichomes type (e.g. B. trimera and B. articulata).
Illustrations of the studied characters are provided.
Key words. Baccharis, Compositae, medicinal species, stomata, trichomes.
RESUMEN
Se realizó un estudio de caracteres epidérmicos de las hojas de las 38 especies
de Baccharis usadas en la medicina popular. El análisis de los caracteres revela:
1) siete tipos de tricomas: cónicos, flageliforme aseptados, filiformes flagelados,
1-armados, 2-4-armados, bulbíferos flagelados y glandulares biseriados; 2) que
28 especies del total de 38 especies medicinales estudiadas presentan tricomas
en “nidos pilosos”; 3) que seis tipos de estomas están presentes: anomocíticos,
anisocíticos, ciclocíticos, actinocíticos, tetracíticos y estaurocíticos; 4) que algunos
tricomas muestran un alto valor diagnóstico, por ejemplo, los tricomas 2-4-armados
son exclusivos de B. dracunculifolia y los flagelados-aseptados ramificados están
presentes sólo en B. trinervis; 5) que los tipos de estomas permiten la diferenciación
de especies con igual tipo de tricoma, (por ejemplo B. trimera de B. articulata). Se
incluyen ilustraciones de los caracteres diagnósticos.
Palabras clave. Baccharis, Compositae, especies medicinales, aparatos entomáticos,
tricomas.
23
Medicinal Baccharis
INTRODUCTION
Baccharis L. is one of the largest genera
of the tribe Astereae (Nesom, 1994). It
comprises c. 400 American species of shrubs
or subshrubs, occasionally small trees and
herbs, nearly all dioecious.
According to Zin (1922), Amat (1983),
Zardini (1984), Girault (1987), Boldt
(1989), Correa & Bernal (1990), Iharlegui
& Hurrell (1992), Soria (1993), Heinrich
(1996), Rojas et al. (1999), Pérez-García
et al. (2001), Erazo et al. (2002), Baggio
et al. (2003), Vidari et al. (2003), Souza
(2004), and a recent webpage (HardingBarlow), 38 medicinal species of
Baccharis with folk medicinal use (or
mentioned at least as medicinal herbs)
are recognized (Table 1); many of these
species can be distinguished by their leaf
or winged stem morphology (Appendix 1).
In 22 of these species biological activity
was tested (Gutkind et al. 1981, Vidari
et al. 2003, Verdi et al. 2005). Baccharis
megapotamica and B. pedunculata are also
included in this study since they can be
considered promisory medicinal species
judging from their tested biological
activity (Verdi et al. 2005).
Heering (1899), Volkens (1890), Quentin
(1911), Ariza Espinar (1973), Barroso
(1976), Pertusi (1987), Hellwig (1990,
1992), Müller (2006) and, are among authors
who have contributed most to the solving
taxonomic problems through the analysis
of micromorphology of leaf surface in
Baccharis.
Epidermal traits, i.e. epidermal cells,
stomata, and hairs, have proven to be
an important tool in taxa delimitation in
many plant families (e.g. Lackey 1978,
Metcalfe & Chalk 1950-1979, Sinclair &
Sharma 1971, Uphof et al. 1962), and also
in distinguishing medicinal species since
24
drugs of pharmaceutical use are made up of
dried and bruised parts in which the different
macroscopical characteristics of the species
are not generally distinguishable (e.g. Amat,
1988; Rapisarda et al., 1997). Within this
context, this study pretend to contribute to
the knowledge of the medicinal species of
the genus Baccharis.
MATERIAL AND METHODS
The study was performed using dried leaves
or winged stems (if only bract-like leaves are
present) taken from herbarium specimens
(Appendix 2). Accepted names, synonyms,
vernacular names, distribution, medicinal
uses, and biological activity for each studied
species are given in Table 1.
The epidermal microcharacters were
studied in samples cleared using the
technique of Dizeo de Strittmatter (1973),
and stained ussing safranin in 80% ethanol.
Measurements of stomata (length) were
taken using a Nikon light microscope
equipped with an ocular micrometer. The
average size of stomata was determined
based on measurements performed on 1520 replicates per sample.
Descriptive terminology for the trichomes
basically follows Ramayya (1962).
Whenever possible, additional synonyms
of trichome terminology were added
(Ariza Espinar 1973, Metcalfe & Chalk
1989, Müller 2006). The classification
of anticlinal epidermal cell wall patterns
follows Stace (1965). Stomata types were
classified according to Stace (1965), Van
Cotthem (1970), Metcalfe & Chalk (1979).
The nomenclature of the included species
follows Matuda (1957), Cuatrecasas (1967),
Ariza Espinar (1973), Barroso (1976),
Giuliano (2000) and Oliveira et al. (2006).
Drawings were made by the authors using
a microscope Leitz SM Lux with camera
lucida.
Freire et al.
Table 1. Distribution, vernacular names, folk medicinal uses and tested biological activity of
Baccharis species. AN = Antilles; AR = Argentina; BO = Bolivia; BR = Brazil; CH = Chile;
CO = Colombia; CR = Costa Rica; EC = Ecuador; ES = El Salvador; GU = Guatemala; HO
= Honduras; ME = Mexico; NC = Nicaragua; PA = Paraguay; PE = Peru; PN = Panama. UR
= Uruguay; US = United States; VE = Venezuela.
Folk medicinal
use
vagina and
wounds wash,
general infections
(Souza, 2004)
Tested biological
activity
Species
Distribution
Vernacular name
Baccharis anomala DC.
Pingraea anomala (DC.) F.H.
Hellwig
AR, BR, PA,
UR
--
AR, UR
“mío-mío blanco”,
“pichana blanca”,
“romerillo”, “romerillo
blanco”, “romerillo
malo” (Argentina).
antirrheumatic,
for treating
traumatisms
(Zardini, 1984)
antifeedant (Verdi et
al., 2005)
Baccharis articulata (Lam.)
Pers.
Conyza articulata Lam;
Molina articulata (Lam.) Less.;
Pingraea articulata (Lam.) F.H.
Hellwig
AR, BO, BR,
PA, UR
“caápé guasú”,
“carqueija”, “carqueja”,
“carqueja amarga”,
“carqueja blanca”,
“carqueja cenicienta”,
“carqueja crespa”,
“carqueja gris”,
“carquejilla”, “chirca
melosa” (Paraguay),
“cola de yacaré”,
“planta del yaguareté”,
“yaguaretá-caá”
(Argentina); “carqueja”,
“carqueja-do-morro”,
“vassoura” (Brazil).
digestive,
hepatoprotective,
diuretic,
antirrheumatic,
antiseptic, for
male sexual
impotence, for
female sterility
(Zardini, 1984)
antioxidant (Verdi et
al., 2005)
Baccharis boliviensis (Wedd.)
Cabrera
Heterothalamus boliviensis
Wedd.; Pseudobaccharis
boliviensis (Wedd.) Cabrera;
Psila boliviensis (Wedd.)
Cabrera
AR, BO,
CH, PE
“chijua”, “romero”,
“tola” (Argentina).
medicinal (Erazo
et al., 2002)
antibacterial (Erazo et
al., 2002)
allelopathic (Verdi et
al., 2005)
ME
“quauhizquiztli”
(Mexico).
for stomachache, laxative,
stornutatory,
for baldness,
for insect bites
(Heinrich, 1996)
spasmolytic,
antibacterial (Verdi et
al., 2005)
AR, BO, BR,
PA, UR
“ajenjo del campo”,
“mío”, “mío-mío”,
“neo-neo”, “neomo”,
“nía”, “nío”, “niyo”,
“ñío-ñío”, “romerillo”
(Argentina); “mio-mio”
(Brazil).
digestive, sudorific
(Amat, 1983)
antiviral,
allelochemical,
antioxidant, citotoxic,
insecticide (Verdi et
al., 2005)
AR, BR, PA,
UR
“carqueija”,
“carqueja”, “carqueja
crespa”, “carquejilla”
(Argentina), “yaguaretéka’á” (Paraguay).
digestive,
hepatoprotective,
antispasmodic,
against fever,
antirrheumatic,
antiseptic (Zardini,
1984)
antibiotic (Gutkind et
al., 1981)
antifeedant,
antimicrobial,
antioxidant (Verdi et
al., 2005)
Baccharis artemisioides Hook.
& Arn.
B. conferta Kunth
Baccharis xalapensis Kunth;
Baccharis orizabaensis Sch.
Bip. ex Hemsl.
Baccharis coridifolia DC.
Eupatorium montevidense
Spreng.
Baccharis crispa Spreng.
Molina crispa (Spreng.)
Less; M. cylindrica Less.;
Baccharis cylindrica (Less.)
DC.; B. genistelloides var.
crispa (Spreng.) Baker; B.
genistelloides var. cylindrica
(Less.) Baker; B. perplexa I.L.
Teodoro & J.E. Vidal; Pingraea
crispa (Spreng.) F.H. Hellwig
--
25
Medicinal Baccharis
Folk medicinal
use
antiinflammatory,
for favouring
scaring (in cows;
Correa & Yesid
Bernal, 1990)
for treating sores
and wounds
(Boldt, 1989)
Tested biological
activity
Species
Distribution
Vernacular name
Baccharis decussata (Klatt)
Hieron.
Pluchea decussata Klatt;
Baccharis moritziana Hieron.
CO, EC, PE,
VE
“almanga”, “armanga”,
“machimbí”
(Colombia).
Baccharis douglasii DC.
Baccharis haenkei DC.
ME, US
--
AR, BO, BR,
PA, UR
“caápé guasú”,
“chilca”, “chilca-y”,
“chilca mata ojo”,
“suncho” (Argentina);
“alecrim-de-vassoura”,
“alecrim-do-campo”,
“vassourinha” (Brazil).
abortive (Iharlegui
& Hurrell, 1992)
--
AR, BR, PA,
UR
“carqueja” (Argentina,
Brazil).
digestive,
hepatoprotective,
antiseptic (Zardini,
1984)
citotoxic (Verdi et al.,
2005)
Baccharis genistelloides
(Lam.) Pers.
Conyza genistelloides Lam.;
Molina reticulata Ruiz & Pav.;
Baccharis reticulata (Ruiz &
Pav.) Pers.
BO, CH, CO,
EC, PE
“charara”, “jatun kimsa
kuchu”, “nuchu-kuchu”,
“quimsa-kuchu”, “tres
espigas” (Bolivia);
“chilca” (Colombia);
“cuchu-cuchu”, “muqumuqu”, “nudo-nudo”,
“qimsa-kuchu” (Peru).
astringent,
diuretic,
antirrheumatic,
for paludism, for
gonorrhea, for
chest-ache, for
stomach-ache
(Correa & Yesid
Bernal, 1990)
antiviral,
gastroprotective
(Verdi et al., 2005)
Baccharis genistifolia DC.
AR, BR, UR
--
antispasmodic
(Zardini, 1984)
--
Baccharis grisebachii Hieron.
Baccharis abietina Kuntze; B.
rosmarinifolia var. andicola
Hauman
AR, BO
“chilca mamil”,
“quinchamal”,
“quinchamalí”,
“romerillo”, “tancha”
(Argentina).
for treating
wounds (Zardini,
1984)
antimicrobial,
citotoxic, protein
synthesis (Verdi et al.,
2005)
Baccharis heterophylla Kunth
Baccharis cuneata DC.;
Baccharis spathulata S.
Schauer
GU, ME
--
for gastrointestinal
disorders (Rojas et
al., 1999)
antimicrobial,
spasmolytic (Verdi et
al., 2005)
Baccharis illinita DC.
BR, PA
--
for gastric
disorders (Baggio
et al., 2003)
gastroprotective
(Verdi et al., 2005)
AR, BO, CO,
EC, PE, VE
“chilca” (Argentina);
“aurauchu”, “yurakchilka” (Bolivia);
“algodoncillo”,
“buéntsamo”,
“buétsemo”, “chilca”,
“chilca blanca”, “chilca
negra”, “chilca rucia”,
“chilco”, “chilco
blanco”, “chilco negro”,
“chilquilla negra
teñidora”, “chirca”,
“chirco”, “gurrubo”
(Colombia); “chilca”,
“chilca negra”,
“chiza”, “yana chilca”
(Ecuador); “chilka”
(Peru); “botonera”,
“niquitao” (Venezuela).
tonic, antidiabetic,
antirrheumatic,
vulnerary,
emolient,
disinfectant, for
diarrhoea, for
inflammation,
for pulmonar
affections (Correa
& Yesid Bernal,
1990)
--
Baccharis dracunculifolia DC.
Baccharis leptospermoides
DC.; B. bracteata Hook.
& Arn.; B. dracunculifolia
var. integerrima Kuntze
f. subviscosa Kuntze;
B. tandilensis Speg.; B.
dracunculifolia f. spectabilis
Heering
Baccharis gaudichaudiana
DC.
Baccharis articulata var.
gaudichaudiana (DC.) Baker
Baccharis latifolia (Ruiz &
Pav.) Pers.
Molina latifolia Ruiz & Pav.;
Baccharis poliantha Kunth; B.
floribunda Kunth; B. riparia
Kunth; Pluchea glabra Griseb.;
Baccharis polyantha f. genuina
Hieron.; B. polyantha var.
macrophylla Hieron; Pingraea
latifolia (Ruíz & Pav.) F.H.
Hellwig
26
--
--
Freire et al.
Species
Baccharis linearis (Ruíz &
Pav.) Pers. Molina linearis
Ruiz & Pav.; Baccharis
rosmarinifolia Hook. &
Arn.; B. lingulata Less.;
B. linifolia Meyen; B.
rosmarinifolia var. subsinuata
DC.; B. callistemoides
Walp.; B. subandina Phil.;
B. rosmarinifolia var.
callistemoides (Walp.)
Heering; B. rosmarinifolia var.
subandina (Phil.) Heering; B.
holmbergii Hicken
Distribution
AR, CH
Vernacular name
Folk medicinal
use
Tested biological
activity
“romerillo” (Argentina).
for arterial blood,
for pyorrhea (San
Martín, 1983)
antispasmodic,
antirrheumatic,
vulnerary, for
inflammations,
for wounds, for
catarrh (Zin, 1922)
insect repelent,
allelopathic, citotoxic,
antifeedant (Verdi et
al., 2005)
Baccharis megapotamica
Spreng.
Baccharis pyramidalis Gardner
AR, BR, PA,
UR
--
potentially
medicinal
Baccharis microcephala
(Less.) DC.
Molina microcephala Less.; B.
microptera Baker
AR, BR, PA,
UR
“carqueija”, “carqueja”
(Argentina); “carqueja”,
“cambará” (Brazil),
“yaguareté-ka’á”
(Paraguay).
allelochemical,
cerebral toxicity,
antileukemic,
antibiotic,
antiinflammatory,
analgesic (Verdi et al.,
2005)
digestive,
hepatoprotective,
antiseptic (Zardini,
1984)
--
ME
--
against catarrhs,
for urinary
problems
(Heinrich, 1996)
--
BO, CO, EC,
PE, VE
“chilca blanca”, “chilco
blanco”, “chilco negro”,
“chilquilla arenilla”,
“espadero”, “humadero”
(Colombia).
for cough, for
bronchitis,
for pulmonar
affections (Girault,
1987)
--
AR, BR, PA,
UR
“carqueija”, “carqueja”
(Argentina).
antirrheumatic
(Zardini, 1984)
antibiotic (Gutkind et
al., 1981)
antimicrobial (Verdi
et al., 2005)
--
antifungal (Verdi et
al., 2005)
Baccharis multiflora Kunth
Neomolina multiflora (Kunth)
F.H. Hellw.
Baccharis nitida (Ruiz & Pav.)
Pers.
Molina nitida Ruiz & Pav.;
Baccharis prinoides Kunth;
B. cassinoides auct. non
DC.; B. oronocensis DC.;
B. popayanensis Hieron.;
B. oronocensis var. perua
Cuatrec.; B. nitida f.
angustifolia Cuatrec.; B.
lepidota Gilli
Baccharis notosergila Griseb.
Baccharis curtifolia S. Moore
Baccharis odorata Kunth
CO, PE
--
Baccharis pedunculata (Mill.)
Cabrera
Conyza pedunculata Mill.;
Eupatorium cotinifolium
Willd.; Baccharis cotinifolia
(Willd.) Urb.; Pseudobaccharis
cotinifolia (Willd.) I.L.
Teodoro; Baccharis
cinnamomifolia Kunth; Psila
cinnamomifolia (Kunth)
Aristeg.; Baccharis venusta
Kunth; B. speciosa DC.;
Eupatorium braunii Pol.;
Baccharis braunii (Pol.)
Standl.; B. splendens Heering;
B. ibaguensis Cuatrec.
for influenza,
for colds, for
bone fractures
and dislocations
(Girault, 1987)
AN, BO,
CO, CR, EC,
GU, HO,
ME, NI, PN,
PE, VE
“chilca”, “chilca
teñidora” (Colombia).
potentially
medicinal
27
Medicinal Baccharis
Species
Distribution
Vernacular name
Baccharis pentlandii DC.
Baccharis fallax Kuntze; B.
rubricaulis Rusby
BO, PE
--
Baccharis pilularis DC.
Baccharis congesta DC.; B.
consanguinea DC.; B. pilularis
var. consanguinea (DC.)
Kuntze; B. pilularis subsp.
consanguinea (DC.) C.B. Wolf
ME, US
“coyote brush” (US).
US, Central
and South
America
“chilca”, “chilca
amarga”, “chilca
blanca”, “chilca dulce”,
“chilca hû”, “chirca”,
“jarilla”, “jarilla
del río”, “junco”,
“romerillo”, “suncho”,
“vara dulce”, “yuno”
(Argentina); “chilca”
(Bolivia); “chilca”,
“chilco” (Colombia);
“axixtlacotl” (Mexico);
“chilca” (Peru);
“mulefat”, “seep
willow”, “water wally”
(US).
Baccharis salicifolia (Ruíz &
Pav.) Pers.
Molina salicifolia Ruíz &
Pav.; M. striata Ruíz & Pav.;
M. viscosa Ruíz & Pav.;
M. parviflora Ruíz & Pav.;
Baccharis glutinosa Pers.;
B. parviflora (Ruíz & Pav.)
Pers.; B. lanceolata Kunth, B.
iresinoides Kunth; B. farinosa
Spreng.; B. longipes DC.; B.
coerulescens DC.; B. alamanii
DC.; B. longifolia DC.; B.
marginalis DC.; B. chilquilla
DC.; B. linifolia DC.; B.
viscosa (Ruíz & Pav.) Kuntze;
B. viscosa var.nigricans
Kuntze; B. marginalis var.
longipes Heering; B. lanceolata
Kunth emend. Heering;
B. purpurascens Heering;
Pingraea salicifolia (Ruíz &
Pav.) F.H. Hellwig; P. viscosa
(Ruíz & Pav.) F.H. Hellwig
Baccharis sarothroides A. Gray
Baccharis sarothroides var.
pluricephala Jeps.; B. arizonica
Eastw.
Baccharis scandens (Ruíz &
Pav.) Pers.
Molina scandens Ruíz & Pav.;
Baccharis adscendens Pers.;
B. fevillei DC.; B. petiolata
DC.; B. calliprinos Griseb.; B.
mirabilis Heering; Pingraea
scandens (Ruíz & Pav.) F.H.
Hellwig
Baccharis serrifolia DC.
Baccharis kellermanii Greenm.;
Archibaccharis prorepens S.F.
Blake; Baccharis prorepens
(S.F. Blake) J.D. Jacks.
Baccharis teindalensis Kunth
Baccharis capitata Kunth
28
Folk medicinal
use
antirrheumatic,
antiseptic,
disinfectant,
for cough, for
pneumonia,
for sprains, for
bone fractures
and dislocations
(Girault, 1987)
for treating hay
fever, sinusitis,
frontal headaches,
to prevent
baldness, as a
female hygienic
agent (HardingBarlow, ined.)
diuretic,
against fevers,
diaphoretic, for
headache, for
inflammation,
for diarrhoea,
for dysentery
(Heinrich, 1996)
antirrheumatic,
antisyphilitic
(Zardini, 1984)
for colds, for
relieving sore
muscles (Heinrich,
1996)
laxative,
expectorant,
for bronchitis,
for pulmonar
affections (Girault,
1987)
antirrheumatic,
for wounds (Zin,
1922)
Tested biological
activity
gastroprotective,
antiviral (Verdi et al.,
2005)
--
--
citotoxic (Verdi et al.,
2005)
ME, US
“desert broom” (US).
AR, CH, PE
“chascoma”, “chilca”,
“chilca dulce”, “fía
mate”, “palo blanco”,
“suncho blanco”
(Argentina).
GU, HO,
ME, NI
--
for gastrointestinal
illnesses
(Heinrich, 1996)
antispasmodic (Verdi
et al., 2005)
--
antiinflammatory,
analgesic,
antimicrobial
(Vidari et al.,
2003)
antiviral (Verdi et al.,
2005)
antiulceral,
antidiarrhoeic (Vidari
et al., 2003)
CO, EC
--
Freire et al.
Species
Distribution
Vernacular name
Folk medicinal
use
Tested biological
activity
Baccharis tola Phil.
Baccharis microphylla var.
incarum Wedd.; B. lejia Phil.;
B. magellanica var. subviscosa
Kuntze; B. magellanica var.
subviscossima Kuntze; B. tola
var. lejia (Phil.) Reiche; B.
incarum (Wedd.) Perkins; B.
tafiensis Heering; B. incarum
var. lejia (Phil.) Cabrera; B.
tola var. incarum (Wedd.) Joch.
Müller
AR, BO,
CH, PE
“baila buen”, “lejía”,
“tola”, “tola lejía”
(Argentina).
medicinal (PérezGarcía et al.,
2001)
protein synthesis,
antibacterial,
antiinflammatory
(Verdi et al., 2005)
“china-thula”, “chilka”,
“hirwa-kowa” (Bolivia);
“anisillo”, “sanalotodo”
(Colombia); “tayanqa”
(Peru); “chilca”,
“sanalotodo”
(Venezuela).
vulnerary,
antidiabetic,
antianemic,
antirrheumatic,
antiseptic,
disinfectant,
for wounds, for
headache, for
bronchitis (Correa
& Yesid Bernal,
1990)
--
AR, BR, BO,
PA, PE, UR
“carqueja” (Argentina,
Brazil); “nuchu-kuchu”,
“quimsa-kuchu”
(Bolivia), “yaguaretéka’á” (Paraguay).
for renal
affections, for
varix (Correa &
Yesid Bernal,
1990)
digestive,
hepatoprotective,
antiseptic, for
female sterility
(Zardini, 1984)
vasorelaxant,
moluscicide,
antimutagenic,
hepatoprotective
(Verdi et al., 2005)
Baccharis trinervis Pers.
Conyza trinervis Lam.;
Baccharis laxa Gardner;
Heterothalamus trinervis (Pers.)
Hook. & Arn.; Pseudobaccharis
trinervis (Pers.) Badillo; P.
trinervis (Pers.) I. L. Teodoro;
Psila trinervis (Pers.) Cabrera
AR, BO, BR,
CO, EC, GU,
HO, ME, NI,
PA, PE, PN,
UR, VE
“chiñi kimsa kuchu”
(Bolivia); “árnica”,
“barzalito”, “barzalito
de monte”, “chaparral
de bestia”, “chilca”,
“chilquita”, “chipolo”,
“gavilana”, “lengua
de gato”, “machucha”,
“mandaguasca”,
“marucha”, “varejón”
(Colombia); “alcotan”
(Ecuador); “Santa
María” (Panama);
“canaciche”,
“chilquita”, “niquitaíto”
(Venezuela).
antihemorrhoidal,
antiinflammatory,
antirrheumatic,
for waist-ache, for
jaundice, for male
sexual impotence,
for female sterility
(Correa & Yesid
Bernal, 1990)
for treating fever,
oedemas, sores,
muscle cramps,
gastrointestinal
disorders
(Heinrich, 1996)
antiviral,
antiinflammatory,
antioxidant (Verdi et
al., 2005)
Baccharis vaccinioides Kunth
Baccharis confertoides G.L.
Nesom
ES, GU, HO,
ME
--
for gastrointestinal
disorders
(Heinrich, 1996)
--
Baccharis wrightii A.Gray
ME
--
for head disorders
emetic (Boldt,
1989)
--
Baccharis tricuneata (L. f.)
Pers.
Erigeron tricuneatum L. f.;
Baccharis microphylla Kunth
Baccharis trimera (Less.) DC.
Molina trimera Less.;
Baccharis genistelloides var.
trimera (Less.) Baker
BO, CO, EC,
PE, VE
29
Medicinal Baccharis
RESULTS AND DISCUSSION
Epidermal characters (Table 2)
Leaf pubescence. All studied species of
Baccharis are pubescent (only a few, e.g. B.
dracunculifolia and B. trinervis are subglabrous
at maturity). Two major groups of indumentum
can be distinguished within the medicinal
species, one consisting of isolated trichomes
and the other with trichomes in tufts (= “nidos
pilosos”, Ariza Espinar, 1973; = “Haarnester”,
Hellwig 1992).
Seven different types of trichomes were
found:
(1) Conical trichomes: uniseriate, 5-6-celled
with body gradually narrowed to a sharp
point. Cells longer distally, often nodulose
at the joints. Lateral and cross walls slightly
thickened. This trichome type is present in
B. anomala and B. decussata. (Fig. 1 A).
Similiar trichomes were illustrated by Müller
(2006) for B. decussata subsp. jelskii (sub
“pedestal hair”) and by Ariza Espinar (1973)
for the non-medicinal B. pulchella.
(2) Aseptate flagellate trichomes (or whip
trichomes, Metcalfe & Chalk 1989):
uniseriate, 2-3-celled with body differentiated
in stalk and a long, whip-like terminal cell.
Stalk 1-2-celled, cells usually isodiametrical
or broader than long.
(2) (a) Simple: terminal cell very long,
flagellate, tubular, as wide as the cells of
the stalk. This trichome type is present in
B. artemisioides and B. grisebachii (Fig.
1 E), which have discolorous leaves with
white-lanate abaxial surface. Trichomes of
B. artemisioides were previously analyzed
and illustrated by Ariza Espinar (1973)
and Pertusi (1987). The aseptate flagellate
trichomes of B. grisebachii were illustrated
by Hellwig (1992) and by Müller (2006),
sub “flagellate filiform” and “filiform hair”
respectively.
(2) (b) Branched: three to five whip
trichomes (with stalk 1-4-celled) appearing
30
stellately branched from the only stalk cell.
This trichomes type is present isolated only
in B. trinervis (Fig. 1 B).
(3) Filiform flagellate trichomes: uniseriate,
many-celled, body filiform to cilindrical
and slightly tapering above. Cells usually
broader than long, isodiametrical, or slightly
longer than broad, with lateral walls convex.
Terminal cell relatively short, tail-like and
pointed (B. multiflora, B. serrifolia and B.
wrightii, in the latter the trichomes are only
found on the stem, Fig. 1 C) or pear-like
and rounded at the apex (B. megapotamica,
Fig. 1 D); those with tail-like apical cell
can be straight or incurved. Dense contents
have been seen in the terminal cell of these
trichomes, which probably have secretory
function.
(4) 1-armed trichomes: uniseriate, 3-4-celled,
with body differentiated into stalk and head.
Stalk 2-3-celled, cells usually broader than
long, lateral walls straight. Head 1-celled
(occasionally branched at the base), with
thick lateral and cross walls and pointed or
rounded at the apex. This trichome type is
present forming tufts or pilose nests only in B.
crispa (Fig. 1 F). It was previously analyzed
and illustrated by Ariza Espinar (1973) for
B. crispa. Hellwig (1992) found trichomes
1-armed in a non-medicinal species, B.
rhomboidalis and also in B. macraei and B.
pilcensis (Hellwig, 1990).
(5) 2-4-armed trichomes: uniseriate, 4-5celled, with body differentiated into stalk
and head. Stalk 3-4-celled, cells broader
than long with lateral walls convex. Head
1-celled, cell 2, 3 or 4-branched (falls/
collapsing at maturity). This trichome type
is present forming tufts or pilose nests
(associated or not to glandular trichomes)
only in B. dracunculifolia. (Fig. 1 G). Müller
(2006) described and illustrated these
trichomes of Baccharis dracunculifolia as
“uniseriate hairs with branched terminal
cells”. Hellwig (1992) found 4-armed
trichomes in a non-medicinal species B.
erioclada.
Table 2: Epidermal characters
Trichomes isolated or paired
Trichomes conical
Trichomes aseptate
flagellate
Trichomes
glandular
present
Trichomes
glandular
absent
branched
simple
B.
decussata
B.
anomala
B.
trinervis
B.
artemisioides
B.
grisebachii
Trichomes filiform flagellate
Trichomes
with apical cell
pear-like
B.
megapotamica
Trichomes
with apical
cell taillike
B.
multiflora
B.
serrifolia
B.
wrightii
Trichomes in tufts
Trichomes 2-4-armed
Trichomes 1-armed
B.
dracunculifolia
B.
crispa
Trichomes bulbiferous flagellate subtype b
Apical cell triangular
Apical cell tail-like
Stomata
cyclocytic
Stomata
actinocytic
B.
articulata
B.
gaudichaudiana
B.
notosergila
B.
heterophylla
B.
conferta
B.
pilularis
B.
teindalensis
B.
tricuneata
B.
vaccinioides
B.
illinita
Stomata medium: (35) 40-60 μm long
Stomata
anomocytic
Stomata
actinocytic
Stomata
cyclocytic
B.
douglasii
B.
genistifolia
B.
microcephala
B.
pedunculata
B.
tola
B.
linearis
B.
nitida
B.
odorata
B.
sarothtoides
B.
genistelloides
Stomata small: 20-30 μm long
Anticlinal
epidermal
cell walls
sinuate
B.
pentlandii
Anticlinal epidermal cell
walls straight
Stomata
anomocytic
B.
latifolia
B.
scandens
Stomata
actinocytic
B.
salicifolia
31
Freire et al.
Stomata
cyclocytic
Trichomes bulbiferous flagellate subtype c
Stomata
large:
75-100μm
long
Medicinal Baccharis
longer than broad or isodiametrical, terminal
cells of the stalk 2, swollen, sphaerical or
oblong-ovoid in shape. Head 1-celled, long,
flagellate. This trichome type is present
only in B. boliviensis. (Fig. 1 H). Similar
trichomes are also present in the unrelated
non-medicinal B. pingraea (Hellwig, 1992).
Figure 1. Trichomes.- A-E, Isolated
trichomes. A: conical, B. anomala, B: aseptate
flagellate branched, B. trinervis; C and D,
Filiform flagellate. C: B. multiflora; D: B.
megapotamica; E: aseptate flagellate simple,
B. artemisioides; F-L, Tufted trichomes
(“pilose nest”) of eglandular trichomes and
biseriate glandular trichomes. F: 1-armed
trichome, B. crispa; G: 2-4-armed trichomes,
B. dracunculifolia; H: bulbiferous flagellate,
subtype a, B. boliviensis; I and J, Bulbiferous
flagellate, subtype b. I: B. notosergila; J: B.
teindalensis; K and L, Bulbiferous flagellate,
subtype c. K: B. pedunculata; L: B. tola.
(6) Bulbiferous flagellate trichomes:
uniseriate, 5-6-celled, usually forming tufts
or pilose nests with glandular trichomes.
According to the number of subapical
cells and the lenght of the apical cell, three
subtypes can be distinguished:
Subtype a: body differentiated into stalk
and head. Stalk 2-3-celled, cells slightly
32
Subtype b: body with cells usually enlarging
above, resulting cuneate in shape. Terminal
cell short with dense contents, broadly
triangular (broadly triangular in B. notosergila
or narrowly triangular in B. articulata, B.
gaudichaudiana, and B. trimera) or more
often tail-like, i.e. sharply delimited from the
subapical cell (B. conferta, B. heterophylla,
B. pilularis, B. teindalensis, B. tricuneata,
B. vaccinioides). (Fig. 1 I, J). Trichomes of
B. articulata and B. trimera were previously
analyzed and illustrated by Pertusi (1987).
Hellwig (1992) found trichomes with tail-like
apical cell in the unrelated non-medicinal B.
paniculata.
Subtype c: this type represents a modification
of the subtype a with a differentiated head
constituted by a flagellate cell, equal or longer
than body, tapering above. This trichome
type is present as isolated trichomes in three
species, B. anomala, B. coridifolia, and B.
trimera, and forms tufts or pilose nests in
16 of the 38 studied species. (Fig. 1 K, L).
Bulbiferous trichomes with flagellate apical
cell of the medicinal species B. linearis and
B. salicifolia were previously illustrated by
Ariza Espinar (1973) and Hellwig (1992)
respectively. Müller (2006) illustrated
for B. coridifolia two types of trichomes,
i.e. bulbiferous flagellate subtype b (sub
“uniseriate hairs”) and aseptate flagellate
(sub “filiform hairs”).
(7) Biseriate glandular trichomes: such
trichomes are constituted by 2 rows of cells
in the body. Biseriate glandular trichomes
are widespread in many taxa studied. They
form tufts with non-glandular trichomes;
Freire et al.
occasionally, tufts are constituted exclusively
by two or three glandular trichomes (e.g. B.
vaccinioides).
Stomata (Fig.2): eleven species (B. boliviensis,
B. conferta, B. linearis, B. nitida, B. odorata,
B. pilularis, B. salicifolia, B. sarothroides,
B. teindalensis, B. tola, and B. tricuneata)
have actinocytic stomata (Fig. 2 A, B), with
five to seven subsidiary cells radially. One
species (B. crispa) has anisocytic stomata
(Fig. 2 C), with three subsidiary cells, of
which one is smaller than the other two. Six
species (B. articulata, B. gaudichaudiana,
B. genistelloides, B. heterophylla, B. illinita,
and B. notosergila) have cyclocytic stomata
(Fig. 2 E, F), with four to seven subsidiary
cells forming a narrow ring. In three species
(B. boliviensis, B. conferta, B. salicifolia)
with predominantly actinocytic stomata, a
few tetracytic stomata are present (Fig. 2 B),
which have four subsidiary cells, two lateral
and two polar. Only one of this species (B.
conferta) has also staurocytic stomata (Fig. 2
A), with four subsidiary cells with anticlinal
walls arranged crosswise to its guard cells.
The remaining species have anomocytic
stomata (Fig. 2 D, G).
Cyclocytic stomata in B. articulata, anisocytic
stomata in B. trimera, and anomocytic
stomata in B. artemisioides were previously
reported by Pertusi (1987). Ariza Espinar
(1973) analyzed and illustrated actinocytic
stomata in B. salicifolia (anomocytic sensu
Ariza Espinar), anomocytic stomata in B.
coridifolia, and anisocytic stomata in B.
crispa (anomocytic sensu Ariza Espinar).
The majority of the species have stomata
between 20 to 60 μm long. In only three species
the stomata are more than 60 μm long, i.e.
Baccharis articulata and B. gaudichaudiana,
between 60 to 75 μm long, and B. illinita with
stomata between 75 to 105 μm long.
Most of the species analyzed have
amphistomatic leaves, only twelve species,
i.e. Baccharis anomala, B. decussata,
B. dracunculifolia, B. grisebachii, B.
heterophylla, B. illinita, B. latifolia, B.
megapotamica, B. multiflora, B. pedunculata,
B. serrifolia, and B. trinervis.
Anticlinal epidermal cell walls (on abaxial
surface): only three species (B. anomala, B.
decussata, and B. pentlandii) amongst the
medicinal species, have sinuate anticlinal wall
(Stace’s types 5-6). The remaining species
have straight to slightly undulate anticlinal
walls (Stace’s types 1-2).
CONCLUSIONS
Certain anatomical characters allow to the
distinction of medicinal species of Baccharis.
For example: 1) B. trinervis is recognized
from other medicinal species by its branched
aseptate flagellate trichomes that are unique
within the whole genus; 2) filiform trichomes
with pear-like apical cell are exclusive of B.
megapotamica; 3) only two species, among
medicinal species, B. artemisioides and B.
grisebachii, have aseptate flagellate trichomes;
4) B. boliviensis and B. dracunculifolia can be
recognized among medicinal species by its
bulbiferous trichomes with two subapical cell
and 2-4-armed trichomes, respectively; 5) B.
illinita is distinguished from other medicinal
species by its large stomata of 75-100 μm.
Other characters, such as stomata type
and epidermal cell walls, can be used to
differentiate species with similar type of
trichomes, as a secondary feature. In B.
articulata, B. gaudichaudiana, B. notosergila,
and B. trimera, the trichomes are bubiferous
flagellate with triangular apical cell;
however, B. trimera can be distinguished by
its anisocytic stomata. In B. heterophylla the
trichomes are bulbiferous flagellate with taillike apical cell, as well as in B. conferta, B.
pilularis, B. teindalensis, B. tricuneata, and
B. vaccinioides; however, B. heterophylla
can be differentiated from them by its
cyclocytic stomata. Baccharis pentlandii has
33
Medicinal Baccharis
Figure 2. Stomata.- A: actinocytic and staurocytic stomata, B. conferta; B: actinocytic and
tetracytic stomata, B. boliviensis; C: anisocytic stomata, B. crispa; D: anomocytic stomata,
B. trimera; E: cyclocytic stomata, B. notosergila; F: cyclocytic stomata, B. illinita; G:
anomocytic stomata, B. pentlandii.
bulbiferous flagellate trichomes with long
apical cell, which is a common type within
medicinal species, but it can be distinguished
by its sinuate anticlinal epidermal cell walls
on abaxial surface.
pencil illustrations and Anabela S. de Oliveira
and Arturo Granda Paucar for providing
relevant bibliography. Support for this study
by the Consejo Nacional de Investigaciones
Científicas y Técnicas (CONICET), Argentina,
is gratefully acknowledged.
ACKNOWLEDGEMENTS
LITERATURE CITED
Thanks are given to Jochen Müller and two
anonimous reviewers for their valuable
comments on the manuscript. Special thanks
to the curators of the following herbaria: F,
GH, LP, and US for the loan of specimens. We
also thank Víctor H. Calvetti for inking our
34
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Freire et al.
Appendix 1. Baccharis species used in traditional medicine
A-C: Leaves. A, B. anomala; B, B. boliviensis; C, B. artemisioides; D, E: Stems. D, B.
articulata; E, B. crispa; F-J: Leaves. F, B. decussata; G, B. conferta; H, B. coridifolia; I, B.
dracunculifolia; J, B. douglasii; K, L: Stems. K, B. gaudichaudiana; L, B. genistelloides;
M-T: Leaves. M, B. genistifolia; N, B. grisebachii; O, B. heterophylla; P, B. illinita; Q, B.
tola; R, B. latifolia; S, B. linearis; T, B. megapotamica; U: Stem. B. microcephala; V-h:
Leaves. V, B. tricuneata; W, B. multiflora; X, B. nitida; Y. B. notosergila; Z, B. odorata;
a, B. pentlandii; b, B. pilularis; c, B. pedunculata; d, B. salicifolia; e, B. sarothroides; f, B.
scandens; g, B. serrifolia; h, B. teindalensis; i: Stem. B. trimera; j,k: Leaves. j, B. trinervis;
k, B. vaccinioides; l, B. wrightii.
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Medicinal Baccharis
Appendix 2. Representative specimens examined of Baccharis (one specimen per species
is cited).
Baccharis anomala. BRASIL. Santa Catarina: Capinzal, Smith et Klein 11917 (LP).
B. artemisioides. ARGENTINA. Buenos Aires: Pedro Luro, Cabrera 4514 (LP).
B. articulata. ARGENTINA. Buenos Aires: Elizalde, Cabrera 1799 (LP).
B. boliviensis. ARGENTINA. Jujuy: Yavi, Quebrada del Lecho, Tolaba et al. 1591 (LP).
B. conferta. MEXICO. Distrito Federal: Cañada de Contreras, Hernández 1 (LP).
B. coridifolia. ARGENTINA. Buenos Aires: San Nicolás, Cabrera 7161 (LP).
B. crispa. ARGENTINA. Córdoba: Tulumba, San Pedro Norte (Chico), Luti 4148 (LP).
B. decussata. COLOMBIA. Cundinamarca: Pacho-Paime, Haught 6060 (LP).
B. douglasii. UNITED STATES. California: Orange County, Raven 16861 (GH).
B. dracunculifolia. ARGENTINA. Jujuy: Valle Grande, Cabrera & Fabris 22678 (LP).
B. gaudichaudiana. ARGENTINA. Misiones: Apóstoles, Cabrera et al. 28469 (LP).
B. genistelloides. PERU. Amazonas: Chachapoyas, Wurdack 497 (LP).
B. genistifolia. ARGENTINA. Buenos Aires, Necochea, Cabrera 1289 (LP).
B. grisebachii. ARGENTINA. Jujuy: Humahuaca, Kiesling et al 614 (LP).
B. heterophylla. MEXICO. Michoacan: Lack Patzcuaro, T. C. Frye & E. M. Frye 2593
(GH).
B. illinita. BRASIL. Rio Grande do Sul: Criuva-San Francisco de Paula: Arroio das Muladas,
Irmão Edẽsio 121 (LP).
B. latifolia. ARGENTINA. Salta: Los Yacones, Zardini 1197 (LP).
B. linearis. ARGENTINA. Río Negro: Bariloche, Cabrera 5827 (LP).
B. megapotamica. PARAGUAY. Caaguazú: Caaguazú, Schinini 9140 (LP).
B. microcephala. ARGENTINA. Misiones: Posadas, Ekman 213 (LP).
B. multiflora. MEXICO. Guerrero: Near Chilpancingo, Nelson 2205 (GH).
B. nitida. PERU. Amazonas: Leimebamba. Sagástegui 7448 (LP).
B. notosergila. ARGENTINA. Buenos Aires: Bavio, Zardini 593 (LP).
B. odorata. PERU. Áncash: Bolognesi, Rumipuquio, Cerrate et al. 6508 (LP).
B. pedunculata. NICARAGUA. Jinotega: Between Matagalpa and Jinotega, Stevens &
Krukoff 10074 (F).
B. pentlandii. BOLIVIA. La Paz: Miraflores, Buchtien 8592 (LP).
B. pilularis. UNITED STATES. California: Marin County, Solbrig 1927 (LP).
B. salicifolia. ARGENTINA. Corrientes: Gral. Paz, Rincón de Vences, Pedersen 6983 (LP).
B. sarothroides. MEXICO. Sonora: Nacore Chico, Muller 3693 (GH).
B. scandens. ARGENTINA. San Juan: Iglesia, Cerro Antecristo, Kiesling et al. 9164 (LP).
B. serrifolia. GUATEMALA. Alta Verapaz, Hunnewell 17272 (GH).
B. teindalensis. COLOMBIA. Nariño: Macizo del volcán Galeras, Cuatrecasas & Mora
26940 (US).
B. tola. ARGENTINA. Jujuy: Tumbaya, Trancas, Cabrera et al. 22513 (LP).
B. tricuneata. COLOMBIA. Antioquia: Santa Helena, Astrágala 25 (LP).
B. trimera. ARGENTINA. Buenos Aires: La Plata, Cabrera 464 (LP).
B. trinervis. ARGENTINA. Jujuy: Ledesma, Valle Grande, Cabrera et al. 14516 (LP).
B. vaccinioides. MEXICO. Chiapas: San Critóbal de Las Casas, Méndez Ton & López 9546
(GH).
B. wrightii. MEXICO. Chihuahua: Valley Ortiz, Pringle 1658 (LP).
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