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Caracteres epidermicos de las especies del genero Baccharis (Asteraceae) usadas en la medicina popular.

RESUMEN

Se realizo un estudio de caracteres epidermicos de las hojas de las 38 especies de Baccharis usadas en la medicina popular. El analisis de los caracteres revela: 1) siete tipos de tricomas: conicos, flageliforme aseptados, filiformes flagelados, 1-armados, 2-4-armados, bulbiferos 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 estan presentes: anomociticos, anisociticos, ciclociticos, actinociticos, tetraciticos y estaurociticos; 4) que algunos tricomas muestran un alto valor diagnostico, por ejemplo, los tricomas 2-4-armados son exclusivos de B. dracunculifolia y los flagelados-aseptados ramificados estan presentes solo en B. trinervis; 5) que los tipos de estomas permiten la diferenciacion de especies con igual tipo de tricoma, (por ejemplo B. trimera de B. articulata). Se incluyen ilustraciones de los caracteres diagnosticos.

Palabras clave. Baccharis, Compositae, especies medicinales, aparatos entomaticos, tricomas.

EPIDERMAL CHARACTERS OF BA CCHARIS (ASTERACEAE) SPECIES USED IN TRADITIONAL MEDICINE

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.

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), Perez-Garcia 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), Muller (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 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, Mfller 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.

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 Muller (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 Willer (2006), sub "flagellate filiform" and "filiform hair" respectively.

(2) (b) Branched. three to five whip trichomes (with stalk 1-4-celled) appearing stellately branched from the only stalk cell. This trichomes type is present isolated only in B. trinervis (Fig. 113).

(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. mult fora, 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--armedtrichomes: 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, 4-5-celled, 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). Willer (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.

[FIGURE 1 OMITTED]

(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 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).

Subtype b: body with cells usually enlarging above, resulting cuneate in shape. Terminal cell short with dense contents, broadly triangular (broadly triangular inB. 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 1, 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. Milller (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; 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 stomatainB. 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 pm long. In only three species the stomata are more than 60 pm long, i.e. Baccharis articulata and B. gaudichaudiana, between 60 to 75 pm long, and B. illinita with stomata between 75 to 105 [micro]m 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. multiora, 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 [micro]m.

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 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.

[FIGURE 2 OMITTED]

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. multi. ora; 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.

[ILUSTRACION OMITIR]

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: Canada de Contreras, Hernandez l (LP).

B. coridifolia. ARGENTINA. Buenos Aires: San Nicolas, Cabrera 7161 (LP).

B. crispa. ARGENTINA. Cordoba: Tulumba, San Pedro Norte (Chico), Luti 4148 (LP).

B. decussata. COLOMBIA. Cundinamarca: Pacho-Paime, Hought 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: Apostoles, 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, Irmao Edesio 121 (LP).

B. latifolia. ARGENTINA. Salta: Los Yacones, Zardini 1197 (LP).

B. linearis. ARGENTINA. Rio Negro: Bariloche, Cabrera 5827 (LP).

B. megapotamica. PARAGUAY. Caaguazu: Caaguazu, Schinini 9140 (LP).

B. microcephala. ARGENTINA. Misiones: Posadas, Ekman 213 (LP).

B. multiora. MEXICO. Guerrero: Near Chilpancingo, Nelson 2205 (GH).

B. nitida. PERU. Amazonas: Leimebamba. Sagdstegui 7448 (LP).

B. notosergila. ARGENTINA. Buenos Aires: Bavio, Zardini 593 (LP).

B. odorata. PERU. Ancash: 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: Marro County, Solbrig 1927 (LP).

B. salicifolia. ARGENTINA. Corrientes: Gral. Paz, Rincon 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. Narino: Macizo del volcan Galeras, Cuatrecosas & Mora 26940 (US).

B. tola. ARGENTINA. Jujuy: Tumbaya, Trancas, Cabrera et al. 22513 (LP).

B. tricuneata. COLOMBIA. Antioquia: Santa Helena, Astragala 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 Crit6bal de Las Casas, Mendez Ton & Lopez 9546 (GH).

B. wrightii. MEXICO. Chihuahua: Valley Ortiz, Pringle 1658 (LP).

ACKNOWLEDGEMENTS

Thanks are given to Jochen Muller 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 Victor H. Calvetti for inking our 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 Cientificas y Tecnicas (CONICET),Argentina, is gratefully acknowledged.

Recibido: 22/09/2006

Aceptado: 18/04/2007

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SUSANA E. FREIRE

ESTRELLA URTUBEY

DANIEL A. GIULIANO

Division Plantas Vasculares, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina. freire@museoftnym.unlp.e

du.ar
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.

Species                   Distribution   Vernacular name

Baccharis anomala DC.     AR, BR, PA,    --
Pingraea anomala (DC.)    UR
F.H. Hellwig

Baccharis artenisioides   AR, UR         "mio-mio blanco",
Hook. & Arn.                             "pichana blanca",
                                         "romerillo", "romerillo
                                         blanco", "romerillo
                                         malo" (Argentina).

Baccharis articulata      AR, BO, BR,    "caape guasu",
(Lam.) Pers.              PA, UR         "carqueija", "carqueja",
Conyza articulata Lam;                   "carqueja amarga",
Molina articulata                        "carqueja blanca",
(Lam.) Less.;                            "carqueja cenicienta",
Pingraea articulata                      "carqueja crespa",
(Lam.) F.H. Hellwig                      "carqueja gris",
                                         "carquejilla", "charca
                                         melosa" (Paraguay),
                                         "cola de yacare",
                                         "planta del yaguarete",
                                         "yaguareta-caa"
                                         (Argentina); "carqueja",
                                         "carqueja-do-morro",
                                         "vassoura" (Brazil).

Baccharis boliviensis     AR, BO,        "chijua", "romero",
(Wedd.) Cabrera           CH, PE         "tola" (Argentina).
Heterothalamus
boliviensis
Wedd.; Pseudobaccharis
boliviensis (deba.)
Cabrera; boliviensis
(Wedd.) Cabrera

B. conferta Knuth         ME             "quauhizquiztli"
Baccharis xalapensis                     (Mexico).
Kunth;
Baccharis orizabaensis
Sch. Bip. ex Hemsl.

Baccharis coridifolia     AR, BO, BR,    "ajenjo del campo",
DC. Eupatorium            PA, UR         "mio", "mio-mio",
montevidense Spreng.                     "neo-neo", "neomo",
                                         "nia", "me", "niyo",
                                         "Nio-Nio", "romerillo"
                                         (Argentina); "mio-mio"
                                         (Brazil).

Baccharis crispa          AR, BR, PA,    "carqueija",
Spreng. Molina crispa     UR             "carqueja", "carqueja
(Sprung.) Less; M.                       crespa", "carquejilla"
cylindrica Less.;                        ka(Argentina), uyaguarete-
Baccharis cylindrica                     ka "a" (Paraguay).
(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

Baccharis decussata       CO, EC, PE,    "almanga", "armanga",
(Klatt) Hieron.           VE             "machimbi"
Pluchea decussate                        (Colombia).
Klatt; Baccharis
moritziana Hieron.

Baccharis douglasii DC.   ME, US         --
Baccharis haenkei DC.

Baccharis                 AR BO, BR,     "caape guasu",
aracunculipona DC.        PA, UR         "chilca", "chilca-y",
Baccharis                                "chilca mata ojo",
leptospermoides DC.;                     "suncho" (Argentina);
B. bracteata Hook                        "alecrim-de-vassoura",
& Arn.; B.                               "alecrim-do-campo",
dracunculifolia var.                     "vassourinha" (Brazil).
integerrima Kuntze f.
subviscosa Kuntze; B.
tandilensis Speg.; B.
dracunculifolia
f. spectabilis

Baccharis                 AR, BR, PA,    "carqueja" (Argentina,
gaudichaudiana DC.        UR             Brazil).
Baccharis articulata
var. gaudichaudiana
(DC.) Baker

Baccharis                 BO, CH, CO,    "charara", "jatun kimsa
genistelloides            EC, PE         kuchu", "nuchu-kuchu",
(Lam.) Pers.                             "quimsa-"kuchu", "tres
Conyza genistelloides                    espigas" (Bolivia);
Lam.; Molina reticulata                  "chilca" (Colombia);
Ruiz & Pav.; Baccharis                   "cuchu-cuchu", "muqu-
reticulata (Ruiz &                       muqu", "nudo-nudo",
Pav.) Pers.                              "qimsa-kuchu" (Peru).

Baccharis genistifolia    AR, BR, UR     --
DC.

Baccharis grisebachii                    "chilca mamil",
Hieron. Baccharis                        "quinchamal",
abietina Kuntze; B.       AR, BO         "quinchamali",
rosmarinifolia var.                      "romerillo", "tancha"
andicola Hauman                          (Argentina).

Baccharis heterophylla    GU, ME         --
Knuth Baccharis cuneata
DC.; Baccharis
spathulata S. Schauer

Baccharis illinita DC.    BR, PA         --

Baccharis latifolia       AR, BO, CO,    "chilca" (Argentina);
(Ruiz & Pav.) Pers.       EC, PE, VE     "aurauchu","yurak-
Molina latifolia Ruiz                    chilka" (Bolivia);
& Pav.; Baccharis                        "algodoncillo",
polyantha Kunth; B.                      "buentsamo",
floribunda Kunth; B.                     "buetsemo", "chilca",
riparia Kunth; Pluchea                   "chilca blanca", "chilca
glabra Griseb.;                          negra", "chilca rucia",
Baccharis polyantha f.                   "chilco", "chilco
genuina Hieron.; B.                      blanco", "chilco negro",
polyantha ver.                           "chilquilla negra
macrophylla Hieron;                      teoidora", "chilca",
Pingraea latifolia                       "chilco", "gurrubo"
(Ruiz & Pav.) F.H.                       (Colombia); "chilca",
Hellwig                                  "chilca negra",
                                         "chiza", "yana chilca"
                                         (Ecuador); "chilka"
                                         (Peru); "botonera",
                                         "niquitao" (Venezuela).

Baccharis linearis        AR, CH         "romerillo" (Argentina).
(Ruiz & 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

Baccharis megapolamica    AR, BR, PA,    --
Spreng.                   UR
Baccharis pyramidales
Gardner

Baccharis microcephala    AR, BR, PA,    "carqueija", "carqueja"
(Less.) DC.               UR             (Argentina); "carqueja",
Molina microcephala                      cambara" (Brazil),
Less.; B. microptera                     "yaguarete-ka'a"
Baker                                    (Paraguay).

Baccharis multiflora      ME             --
Kunth
Neomolina multiflora
(Kunth) F.H. Hellw.

Baccharis nitida          BO CO, EC,     "chilca blanca", "chilco
(Ruiz & Pav.) Pers.       PE, VE         blanco", "chilco negro",
Molina nitida Ruiz &                     "chilquilla arenilla",
Pav.; Baccharis                          "espadero", "humadero"
prinoides Kunth;                         (Colombia).
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 nolosergila     AR, BR, PA,    "carqueija", "carqueja"
Griseb.                   UR             (Argentina).
Baccharis curtifolia
S. Moore

Baccharis odorata Kunth   CO, PE         --

Baccharis pedunculata     AN, BO,        "chilca", "chilca
(Mill.) Cabrera           CO, CR, EC,    teoidos" (Colombia).
Conyza pedunculata        GU, HO,
Mill.; Eupatorium         ME, NI, PN,
cotinifolium Willd.;      PE, VE,
Baccharis cotinifolia
(Willd.) Urb.;
Pseudobaccharis
cotinifolia (Willd.)
I.L. Teodoro; Baccharis
cinnamomifolia Kunth;
Psila cinnamomifolia
(Kunth) Baccharis
venusta Kunth; B.
speciosa DC.;
Eupatorium braunii
Pol.; Baccharis braunii
(Poi.) Standl.; B.
splendens Heering; B.
ibaguensis Cuatree.

Baccharis pentlandii      BO, PE         --
DC. Baccharis fallax
Kuntze; B. rubricaulis
Rusby

Baccharis pilularis DC.   ME, US         "coyote brush" (US).
Baccharis congesta DC.;
B. consanguinea DC.;
B. pilularis var.
consanguinea (DC.)
Kuntze; B. pilularis
subsp. consanguinea
(DC.) C.B. Wolf

Baccharis salicifolia     US, Central    "chilca", "chilca
(Ruiz & Pav.) Pers.       and South      amarga", "chilca
Molina salicifolia Ruiz   America        blanca", "chilca dulce",
& Pav.; M. striata Ruiz                  "chilca ho", "chilca",
& Pav.; M. viscosa Ruiz                  "jarilla", "jarilla
& Pav.; M. parviora                      del rio", "junco",
Ruiz & Pav.; Baccharis                   "romerillo", "suncho",
glutinosa Pers.; B.                      "vara dulce", "yuno"
parviora (Ruiz & Pav.)                   (Argentina); "chilca"
Pers.; B. lanceolata                     (Bolivia); "chilca",
Knuth, B. iresinoides                    "chilco" (Colombia);
Kunth; B. farinosa                       "axixtlacotl" (Mexico);
B. longipes DC.; B.                      "chilca" (Peru);
coerulescens DC.; B.                     "mulefat", "seep
alamanii DC.; B.                         willow", "water wally"
longifolia DC.; B.                       (US).
marginales DC.; B.
chilquilla DC.; B.
longifolia DC.; B.
viscosa (Ruiz & Pav.)
Kuntze; B. viscosa
var. nigricans Kuntze;
B. marginales var.
longipes Heering;
B. lanceolata
Kunth emend. Heering;
B. purpurascens
Heering; Pingraea
salicifolia (Ruiz &
Pav.) F.H. Hellwig;
P. viscosa (Ruiz &
Pav.) F.H. Hellwig

Baccharis sarothroides    ME, US         "desert broom" (US).
A. Gray Baccharis
sarothroides var.
pluricephala Jeps.;
B. arizonica Eastw.

Baccharis scandens        AR, CH, PE     "chascoma", "chilca",
(Ruiz & Pav.) Pers.                      "chilca dulce", "fia
Molina scanderas Ruiz                    mate", "palo blanco",
& Pav.; Baccharis                        "suncho blanco"
adscendens Pers.; B.                     (Argentina).
fevillei DC.; B.
petiolata DC.; B.
calliprinos Griseb.; B.
mirabilis Heering;
Pingraea scanderas
(Ruiz & Pav.) F.H.
Hellwig

Baccharis serrifolia      GU, HO,        --
DC. Baccharis             ME, NI
kellermanii Greenm.;
Archibaccharis
prorepens S.F. Blake;
Baccharis prorepens
(S.F. Blake) J.D.
Jacks.

Baccharis teindalensis    CO, EC         --
Knuth
Baccharis capitata
Kunth

Baccharis tola Phil.      AR, BO,        "baila buen", "lejia",
Baccharis microphylla     CH, PE         "tola", "tola lejia"
var. incarum Wedd.;                      (Argentina).
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
Hooting; B. incarum
var. lejia (Phil.)
Cabrera; B. tola var.
incarum (Wedd.) Joch.
Muller

Baccharis tricaneata      BO, CO, EC     "china-thula", "chilca",
(L. f.) Pers.             PE, VE         "hirwa-kowa" (Bolivia);
Erigeron tricuneatum                     "anisillo", "sanalotodo"
L. f.; Baccharis                         (Colombia); "tayanqa"
microphylla Knuth                        (Pam); "chilca",
                                         "sanalotodo"
                                         (Venezuela).

Baccharis trimera                        "carqueja" (Argentina
(Lass.) DC.               AR, BR, BO,    Brazil); "nuchu-kuchu,"
Molina trimera Less.;     PA, PE, UR     "quimsa-kuchu"
Baccharis                                (Bolivia), "yaguarete-
genistelloides var.                      ka'a" (Paraguay).
trimera (Less.) Baker

Baccharis trinervis       AR, BO, BR,    "chihi kimsa kuchu"
Pers. Conyza trinervis    CO, EC, GU,    (Bolivia); "arnica",
Lam.; Baccharis laxa      HO, ME, NI,    "barzalito", "barzalito
Gardner; Heterothalamus   PA, PE, PN,    da monte", "chaparral
trinervis (Pers.)         UR, VE         da bestia", "chilca",
Hook. & Arn.;                            "chilquita", "chipolo",
Pseudobaccharis                          "gavilana", "lengua
trinervis (Pers.)                        de gate", "machucha",
Badillo; P. trinervis                    "mandaguasca",
(Pers.) I. L. Teodoro;                   "mamcha", "varejon"
Psila trinervis (Pers.)                  (Colombia); "alcotan"
Cabrera                                  (Ecuador); "Santa
                                         Maria" (Panama);
                                         "canaciche",
                                         "chilquita", "niquitaito"
                                         (Venezuela).

Baccharis vaccinioides    ES GU, HO,     --
Kunth                     ME
Baccharis confertoides
G.L. Nesom

Baccharis wrightii A.     ME             --
Gray

Species                   Folk medicinal         Tested biological
                          use                    activity

Baccharis anomala DC.     vagina and             --
Pingraea anomala (DC.)    wounds wash,
F.H. Hellwig              general infections
                          (Souza, 2004)

Baccharis artenisioides   antirrheumatic,        antifeedant
Hook. & Arn.              for treating           (Verdi et al., 2005)
                          traumatisms
                          (Zardini 1984)

Baccharis articulata      digestive,             antioxidant
(Lam.) Pers.              hepatoprotective,      (Verdi et al., 2005)
Conyza articulata Lam;    diuretic,
Molina articulata         antirrheumatic,
(Lam.) Less.;             antiseptic, for
Pingraea articulata       male sexual
(Lam.) F.H. Hellwig       impotence, for
                          female sterility
                          (Zardini, 1984)

Baccharis boliviensis     medicinal (Erizo       antibacterial
(Wedd.) Cabrera           et al., 2002)          (Erizo et al., 2002)
Heterothalamus                                   allelopathic
boliviensis                                      (Verdi et al., 2005)
Wedd.; Pseudobaccharis
boliviensis (deba.)
Cabrera; boliviensis
(Wedd.) Cabrera

B. conferta Knuth         for stomachache,       spasmolytic,
Baccharis xalapensis      laxative,              antibacterial
Kunth;                    stornutatory,          (Verdi et al. 2005)
Baccharis orizabaensis    for baldness,
Sch. Bip. ex Hemsl.       for insect bites
                          (Heinrich 1996)

Baccharis coridifolia     digestive, sudorific   antiviral,
DC. Eupatorium            (Amat, 1983)           allelochemical,
montevidense Spreng.                             antioxidant,
                                                 citotoxic,
                                                 insecticide
                                                 (Verdi et al., 2005)

Baccharis crispa          digestive,             antibiotic (Gutkind
Spreng. Molina crispa     hepatoprotective,      et al., 1981)
(Sprung.) Less; M.        antispasmodic,         antifeedant,
cylindrica Less.;         against fever,         antimicrobial,
Baccharis cylindrica      antirrheumatoc         antioxidant
(Less.) DC.; B.           antiseptic             (Verdi et al., 2005)
genistelloides var.       (Zardini, 1984)
crispa (Spreng.) Baker;
B. genistelloides var.
cylindrica (Less.)
Baker; B. perplexa I.L.
Teodoro & J.E. Vidal;
Pingraea crispa
(Spreng.) F.H. Hellwig

Baccharis decussata       antiinflammatory,      --
(Klatt) Hieron.           for favouring
Pluchea decussate         scaring (in cows;
Klatt; Baccharis          Correa & Yesid
moritziana Hieron.        Bernal, 1990)

Baccharis douglasii DC.   for treating sores     --
Baccharis haenkei DC.     and wounds
                          (Boldt, 1989)

Baccharis                 abortive (Iharlegui    --
aracunculipona DC.        & Hurrell, 1992)
Baccharis
leptospermoides DC.;
B. bracteata Hook
& Arn.; B.
dracunculifolia var.
integerrima Kuntze f.
subviscosa Kuntze; B.
tandilensis Speg.; B.
dracunculifolia
f. spectabilis

Baccharis                 digestive,             citotoxic
gaudichaudiana DC.        hepatoprotective,      (Verdi et al., 2005)
Baccharis articulata      antiseptic
var. gaudichaudiana       (Zardini, 1984)
(DC.) Baker

Baccharis                 astringent,            antiviral,
genistelloides            diuretic,              gastroprotective
(Lam.) Pers.              antirrheumatic,        (Verdi et al., 2005)
Conyza genistelloides     for paludism, for
Lam.; Molina reticulata   gonorrhea, for
Ruiz & Pav.; Baccharis    chest-ache, for
reticulata (Ruiz &        stomach-ache
Pav.) Pers.               (Correa & Yesid
                          Bernal, 1990)

Baccharis genistifolia    antispasmodic          --
DC.                       (Zardini, 1984)

Baccharis grisebachii     for treating           antimicrobial,
Hieron. Baccharis         wounds                 citotoxic, protein
abietina Kuntze; B.       (Zardini, 1984)        synthesis
rosmarinifolia var.                              (Verdi et al., 2005)
andicola Hauman

Baccharis heterophylla    for gastrointestinal   antimicrobial,
Knuth Baccharis cuneata   disorders              spasmolytic
DC.; Baccharis            (Rojas et al., 1999)   (Verdi et al., 2005)
spathulata S. Schauer

Baccharis illinita DC.    for gastric            gastroprotective
                          disorders (Baggio      (Verdi et al.,
                          et al., 2003)          2005)

Baccharis latifolia       tonic, antidiabetic,   --
(Ruiz & Pav.) Pers.       antirrheumatic,
Molina latifolia Ruiz     vulnerary,
& Pav.; Baccharis         emolient,
polyantha Kunth; B.       disinfectant, for
floribunda Kunth; B.      diarrhoea, for
riparia Kunth; Pluchea    inflammation,
glabra Griseb.;           for pulmonar
Baccharis polyantha f.    affections (Correa
genuina Hieron.; B.       & Yesid Bernal,
polyantha ver.            1990)
macrophylla Hieron;
Pingraea latifolia
(Ruiz & Pav.) F.H.
Hellwig

Baccharis linearis        for arterial blood,    insect repelent,
(Ruiz & Pav.) Pers.       for pyorrhea (San      allelopathic,
Molina linearis Ruiz      Martin, 1983)          citotoxic,
& Pav.; Baccharis         antispasmodic,         antifeedant
rosmarinifolia Hook. &    antirrheumatic,        (Verdi et al., 2005)
Arn.; B. lingulata        vulnerary, for
Less.; B. linifolia       inflammations,
Meyen; B.                 for wounds, for
rosmarinifolia var.       catarrh (Zin, 1922)
subsinuata DC.; B.
callistemoides Walp.;
B. subandina Phil.;
B. rosmarinifolia var.
callistemoides (Walp.)
Heering; B.
rosmarinifolia var.
subandina (Phil.)
Heering; B. holmbergii
Hicken

Baccharis megapolamica    potentially            allelocnemical,
Spreng.                   medicinal              cerebral toxicity,
Baccharis pyramidales                            antileukemic,
Gardner                                          antibiotic,
                                                 antiinflammatory,
                                                 analgesic
                                                 (Verdi et al., 2005)

Baccharis microcephala    digestive,             --
(Less.) DC.               hepatoprotective,
Molina microcephala       antiseptic (Zardini,
Less.; B. microptera      1984)
Baker

Baccharis multiflora      against catarrhs,
Kunth                     for urinaryproblems    --
Neomolina multiflora      (Heinrich, 1996)
(Kunth) F.H. Hellw.

Baccharis nitida          for cough, for         --
(Ruiz & Pav.) Pers.       bronchitis,
Molina nitida Ruiz &      for pulmonar
Pav.; Baccharis           affections
prinoides Kunth;          (Girault, 1987)
B. cassinoides auct.
non DC.; B. oronocensis
DC.; B. popayanensis
Hieron.; B. oronocensis
var. perua Cuatrec.;
B. nitida f.
angustifolia Cuatrec.;
B. lepidota Gilli
                                                 antiltiotic (Gutkind
Baccharis nolosergila     antirrheumatic         et al., 1981)
Griseb.                   (Zardini, 1984)        antimicrobial
Baccharis curtifolia                             (Verdi et al., 2005)
S. Moore

Baccharis odorata Kunth   for influenza,         --
                          for colds, for
                          bone fractures
                          and dislocations
                          (Girault, 1987)

Baccharis pedunculata     potentially            antifungal
(Mill.) Cabrera           medicinal              (Verdi et al., 2005)
Conyza pedunculata
Mill.; Eupatorium
cotinifolium Willd.;
Baccharis cotinifolia
(Willd.) Urb.;
Pseudobaccharis
cotinifolia (Willd.)
I.L. Teodoro; Baccharis
cinnamomifolia Kunth;
Psila cinnamomifolia
(Kunth) Baccharis
venusta Kunth; B.
speciosa DC.;
Eupatorium braunii
Pol.; Baccharis braunii
(Poi.) Standl.; B.
splendens Heering; B.
ibaguensis Cuatree.

Baccharis pentlandii      antirrheumatic,        gastroprotective,
DC. Baccharis fallax      antiseptic,            antiviral
Kuntze; B. rubricaulis    disinfectant,          (Verdi et al., 2005)
Rusby                     for cough, for
                          pneumonia,
                          for sprains, for
                          bone fractures
                          and dislocations
                          (Girault, 1987)

Baccharis pilularis DC.   for treating hay       --
Baccharis congesta DC.;   fever, sinusitis,
B. consanguinea DC.;      frontal headaches,
B. pilularis var.         to prevent
consanguinea (DC.)        baldness, as a
Kuntze; B. pilularis      female hygienic
subsp. consanguinea       agent (Harding-
(DC.) C.B. Wolf           Barlow, ined.)

Baccharis salicifolia     diuretic,              --
(Ruiz & Pav.) Pers.       against fevers,
Molina salicifolia Ruiz   diaphoretic, for
& Pav.; M. striata Ruiz   headache, for,
& Pav.; M. viscosa Ruiz   inflammation,,
& Pav.; M. parviora       for diarrhoea,
Ruiz & Pav.; Baccharis    for dysentery
glutinosa Pers.; B.       (Heinrich, 1996)
parviora (Ruiz & Pav.)    antisyphilnac
Pers.; B. lanceolata      (Zardini, 1984)
Knuth, B. iresinoides
Kunth; B. farinosa
B. longipes DC.; B.
coerulescens DC.; B.
alamanii DC.; B.
longifolia DC.; B.
marginales DC.; B.
chilquilla DC.; B.
longifolia DC.; B.
viscosa (Ruiz & Pav.)
Kuntze; B. viscosa
var. nigricans Kuntze;
B. marginales var.
longipes Heering;
B. lanceolata
Kunth emend. Heering;
B. purpurascens
Heering; Pingraea
salicifolia (Ruiz &
Pav.) F.H. Hellwig;
P. viscosa (Ruiz &
Pav.) F.H. Hellwig

Baccharis sarothroides    for colds, for         citotoxic
A. Gray Baccharis         relieving sore         (Verdi et al., 2005)
sarothroides var.         muscles (Heinrich,
pluricephala Jeps.;       1996)
B. arizonica Eastw.

Baccharis scandens        laxative,
(Ruiz & Pav.) Pers.       expectorant,
Molina scanderas Ruiz     for bronchitis,
& Pav.; Baccharis         for pulmonar
adscendens Pers.; B.      affections             --
fevillei DC.; B.          (Girault, 1987)
petiolata DC.; B.         antirrheumatic,
calliprinos Griseb.; B.   for wounds (Zen,
mirabilis Heering;        1922)
Pingraea scanderas
(Ruiz & Pav.) F.H.
Hellwig

Baccharis serrifolia      for gastrointestinal   antispasmodic
DC. Baccharis             illnesses              (Verdi et al., 2005)
kellermanii Greenm.;      (Heinrich, 1996)
Archibaccharis
prorepens S.F. Blake;
Baccharis prorepens
(S.F. Blake) J.D.
Jacks.

Baccharis teindalensis    antiinflammatory,      antiviral
Knuth                     analgesic,             (Verdi et al., 2005)
Baccharis capitata        antimicrobial          antiulceral,
Kunth                     (Vidari et al.,        antidiarrhoeic
                          2003)                  (Vidari et al.,
                                                 2003)

Baccharis tola Phil.      medicinal (Perez-      protein synthesis,
Baccharis microphylla     Garcia et al.,         antibacterial,
var. incarum Wedd.;       2001)                  antiinflammatory
B. lejia Phil.; B.                               (Verdi et al., 2005)
magellanica var.
subviscosa Kuntze;
B. magellanica var.
subviscossima Kuntze;
B. tola var. lejia
(Phil.) Reiche; B.
incarum (Wedd.)
Perkins; B. tafiensis
Hooting; B. incarum
var. lejia (Phil.)
Cabrera; B. tola var.
incarum (Wedd.) Joch.
Muller

Baccharis tricaneata      vulnerary,             --
(L. f.) Pers.             antidiabetic,
Erigeron tricuneatum      antianemic,
L. f.; Baccharis          antirrheumatic,
microphylla Knuth         antiseptic,
                          disinfectant,
                          for wounds, for
                          headache, for
                          bronchitis (Correa
                          & Yesid Bernal,
                          1990)

Baccharis trimera         for renal              vasorelaxnt,
(Lass.) DC.               affections, for        moluscicide,
Molina trimera Less.;     varix (Correa &        antimutagenic,
Baccharis                 Yesid Bemal,           hepatoprotective
genistelloides var.       1990)                  (Verdi et al., 2005)
trimera (Less.) Baker     digestive,
                          hepatoprotective,
                          antiseptic, for
                          female sterility
                          (Zardini, 1984)

Baccharis trinervis       antihemorrhoidal,      antiviral,
Pers. Conyza trinervis    antiiflammatory,       antiinflammatory,
Lam.; Baccharis laxa      antirrheumatic,        antioxidant
Gardner; Heterothalamus   for waist-ache, for    (Verdi et al., 2005)
trinervis (Pers.)         jaundice, for male
Hook. & Arn.;             sexual impotence,
Pseudobaccharis           for female sterility
trinervis (Pers.)         (Correa & Yesid
Badillo; P. trinervis     Bernal, 1990)
(Pers.) I. L. Teodoro;    for treating fever,
Psila trinervis (Pers.)   oedemas, sores,
Cabrera                   muscle cramps,
                          gastrointestinal
                          disorders
                          (Heinrich, 1996)

Baccharis vaccinioides    for gastrointestinal   --
Kunth                     disorders
Baccharis confertoides    (Heinrich, 1996)
G.L. Nesom

Baccharis wrightii A.     for head disorders     --
Gray                      emetic (Boldt, 1989)

Table 2: Epidermal characters

Trichomes isolated or paired

  Trichomes conical       Trichomes aseptate
                              flagellate

Trichomes   Trichomes   branched       simple
glandular   glandular
 present     absent

   B.          B.          B.            B.
decussata    anomala    trinervis   artemisioides

                                         B.
                                     grisebachii

Trichomes filiform
flagellate

  Trichomes       Trichomes
with apical         with
    cell           apical
  pear-like      cell tail-
                    like

     B.              B.
megapotarnica     multiora

                     B.
                 serrifolia

                     B.
                  wrightii

Trichomes in tufts

Trichomes 2-4-armed   Trichomes 1-armed

        B.                   B.
  dracunculifolia          crispa

Trichomes bulbiferous flagellate subtype b

    Apical cell triangular          Apical cell tail-like

   Stomata                         Stomata         Stomata
  cyclocytic                      cyclocytic     actinocytic

      B.                              B.             B.
  articulata                     heterophylla     conferta

      B.                                             B.
gaudichaudiana                                    pilularis

      B.                                             B.
 notosergila                                    teindalensis

                                                     B.
                                                 tricuneata

                                                     B.
                                                vaccinioides

Trichomes bulbiferous flagellate subtype c

   Stomata        Stomata medium: (35) 40-60 [micro]un long
    large:
75-100[micro]m
     long

      B.            Stomata        Stomata         Stomata
   illinita       anomocytic     actinocytic     cyclocytic

                      B.              B.             B.
                   douglasii       linearis     enistelloides

                      B.              B.
                 genistifolia       nitida

                      B.              B.
                 microcephala      odorata

                      B.              B.
                  pedunculata    sarothtoides

                      B.
                     tola

Trichomes bulbiferous flagellate subtype c

     Stomata small: 20-30 [micro]m long

  Anticlinal      Anticlinal epidermal cell
  epidermal             walls straight
  cell walls
   sinuate

      B.            Stomata        Stomata
  entlandii       anomocytic     actinocytic

                      B.              B.
                   latifolia     salicifolia

                      B.
                   scandens
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Title Annotation:Botanica
Author:Freire, Susana E.; Urtubey, Estrella; Giuliano, Daniel A.
Publication:Caldasia
Date:Jan 1, 2007
Words:6703
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