ISSN 0373-580 X A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees Bol. Soc. Argent. Bot. 43 (1-2): 31 - 60. 2008
Leaf anatomy of medicinal shrubs and trees from Misiones forest of the Paranaense Province (Argentina). Part 2. ANA M. ARAMBARRI1.*, SUSANA E. FREIRE1.2.3, MARTA N. COLARES1, NÉSTOR D. BAYÓN1, MARÍA C. NOVOA1, CLAUDIA MONTI1, and SEBASTIÁN A. STENGLEIN1.3 Summary: The present paper contains the study of the second part of medicinal shrubs and trees from Paranaense province. Forty five species of shrubs and trees belonging to 29 families inhabiting Misiones forest of the Paranaense biogeographic province (Argentina) have been cited with medicinal properties. The work provides illustrations of diagnostic characters and conclusions of the main botanical differential traits, such as the presence of crystaliferous epidermis (e.g. Trixis divaricata subsp. divaricata); stomata and trichomes types (e. g. ciclocytic stomata in Pilocarpus pennatifolius and scale peltate trichomes in Tabebuia heptaphylla); midvein transection outlines (e.g. midvein convex and keel-shaped on the adaxial side in Schinus weinmanniifolia); presence and types of crystals (e.g. crystal sand in Cordia ecalyculata, raphides in Psychotria carthagenensis). This paper also gives an ecological interpretation of the species studied which shows predominantly a combination of mesomorphic (e.g. hypostomatic leaves, dorsiventral mesophyll) and xeromorphic leaf traits (e.g. thick cuticle, abundant sclerenchyma, multilayered epidermis, mesophyll formed exclusively by palisade parenchyma, multilayered hypodermis, presence of sclereids). Only two species (Ilex paraguariensis and Manihot grahamii) have mesomorphic (e.g. hypostomatic leaves, dorsiventral mesophyll) and hygromorphic leaf characters (e.g. epidermis glabrous). Finally, the work provides a key to distinguish 107 medicinal shrubs and trees from the Paranaense biogeographic province (Part 1: Gallery forests and Part 2: Misiones forest) that permit identified species using anatomy leaf characteristics. Key words: anatomy, Argentina, Misiones forest, leaves, medicinal plants, Paranaense biogeographic province. Resumen: Anatomía foliar de arbustos y árboles medicinales de la Selva Misionera de la provincia Paranaense (Argentina). Parte 2. El presente trabajo corresponde a la segunda entrega del estudio de arbustos y árboles medicinales de la provincia biogeografica Paranaense. En esta parte, se analizaron 45 especies contenidas en 29 familias que habitan la Selva Misionera y han sido citadas con propiedades medicinales. El trabajo se acompaña con ilustraciones y se brinda una conclusión de los caracteres diagnósticos, tales como la presencia de epidermis cristalífera (e.g. Trixis divaricata subsp. divaricata); los tipos de estomas (e.g. ciclocíticos en Pilocarpus pennatifolius); y de tricomas (e.g. escamosos peltados en Tabebuia heptaphylla); el contorno de la vena media (e.g. convexa y aquillada en la cara adaxial en Schinus weinmanniifolia); la presencia y tipo de cristales (e.g. arena cristalina en Cordia ecalyculata, rafidios en Psychotria carthagenensis). También se provee una interpretación ecológica de las especies estudiadas cuyas hojas presentaron predominantemente una combinación de caracteres mesomórficos (e.g. hojas hipostomáticas, mesofilo dorsiventral) y xeromórficos (e.g. cutícula gruesa, esclerénquima abundante, epidermis pluriseriada, mesofilo indiferenciado en empalizada, hipodermis pluriseriada, esclereidas idioblásticas). Solamente dos especies, Ilex paraguariensis y Manihot grahamii, presentaron combinados caracteres mesomórficos (entre otros, hojas hipostomáticas y mesofilo dorsiventral) e higromórficos (epidermis glabra). Finalmente, se provee una clave para la diferenciación de 107 especies de arbustos y árboles medicinales, de la provincia biogeográfica Paranaense (Parte 1: Selvas en Galería y Parte 2: Selva Misionera) que permite identificar las especies utilizando caracteres anatómicos de la hoja. Palabras clave: anatomía, Argentina, hoja, plantas medicinales, provincia biogeográfica Paranaense, Selva Misionera. Área de Botánica, Facultad de Ciencias Agrarias y Forestales, Calle 60 y 119, C.C. 31, 1900 La Plata, Universidad Nacional de La Plata, Argentina. 2 Instituto de Botánica Darwinion, Labardén 200, C.C.22, B1642 HYD San Isidro, Buenos Aires, Argentina. 3 Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina (CONICET) * Corresponding author. Tel: +54-221-423-6758 int. 461; Fax: +54-221-425-2346. E-mail: anaramba@yahoo.com.ar; botgral@ceres.agro.unlp.edu.ar 1
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Introduction This paper is the second part of the study of medicinal shrubs and trees from Paranaense province. The first part including the medicinal shrubs and trees of the Gallery forests, was published by Arambarri et al. (2006). Misiones forest is situated in the northeastern Argentina, extending from Misiones to northeastern Corrientes political provinces. From a biogeographic point of view, Misiones and Gallery forests belong to Paranaense province within the Amazonian dominion of Neotropical region (Cabrera & Willink, 1973). In this region, more than 100 native shrubs and trees have been cited as used for food, dressing, religious and magic ceremonies, and in folk medicine by native population (Hieronymus, 1882; González et al., 1939; Domínguez, 1928; Rodríguez & O’Donell, 1943; Martínez Crovetto, 1963, 1964, 1965, 1981; Edwin & Reitz, 1967; Burkart, 1979, 1987; Cristiani & Amorín, 1972; Mateu Amengual, 1974;
Leonardis, 1975; Dimitri, 1978; Rotman, 1978, 1987, 1995; Sorarú & Bandoni, 1978; Ratera & Ratera, 1980; Toursarkissian, 1980; Sawchuk & Galeano, 1981; Xifreda, 1981; Zardini, 1984; Bernardi, 1985; Biloni, 1990; Amat & Vajía, 1991; Lahitte & Hurrell, 1994, 1999; Filipov, 1994; Marzocca, 1997; Amat & Yajía, 1998; Blariza et al., 1998; Chifa & Ricciardi, 2001; Eibl et al., 2002; Freire et al., 2002; López Villalba et al., 2002; Mastroberti & Mariath, 2003; Rondina et al., 2003; Alonso & Desmarchelier, 2005; Mandrile, 2005; Barboza et al., 2006). In order to increase the knowledge of native shrubs and trees with medicinal properties, we studied the leaf anatomy of 45 species belonging to 42 genera which are contained into 29 families from Misiones forest.
Results On the basis of leaf anatomical features of 62 species from Gallery forests and 45 species from the Misiones forest the following key was generated:
Key to 107 shrubs and trees with medicinal properties from Paranaense biogeographic province (Misiones and Gallery forests) 1. Epidermis in surface view: stomata arranged in parallel rows. 2. Hypodermis present. Ciclocytic or tetracytic stomata. 3. Ciclocytic stomata. Mesophyll isobilateral. Idioblastic sclereids present. 1. Araucaria angustifolia (Fig. 1) 3’. Tetracytic stomata. Mesophyll indifferentiated. Idioblastic sclereids absent. 2. Syagrus romanzoffiana 2’. Hypodermis absent. Paracytic stomata. 4. Stomatal apparatous of Dicotyledons type with reniform guard cells. Mesophyll with idioblastic sclereids containing crystals into their cavities. 3. Ligaria cuneifolia 4’. Stomatal apparatous of Poaceae (= Gramineae) type with osteocells as guard cells. Mesophyll without idioblastic sclereids. 4. Guadua chacoensis (Fig. 1) 1’. Epidermis in surface view: stomata arranged at random. 5. Indumentum of pilose nest type. 6. Leaves bract-like. Ciclocytic stomata.
* For methods refer to Part 1: Arambarri et al. (2006).
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A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees 7. Stem two winged. 5. Baccharis articulata 7’. Stem three winged. 6. Baccharis microcephala 6’. Leaves well developed. Anomocytic stomata. 7. Eupatorium laevigatum 5’. Indumentum of other type or absent. 8. Midvein vaulted (convex) on the adaxial side. 9. Cystoliths in enlarged epidermal cells (lithocysts) present. 10. Epidermis one layered. Parenchyma with crystal sand. 8. Cordia ecalyculata (Fig. 1) 10’. Epidermis multilayered. Parenchyma without crystal sand. 9. Ficus luschnathiana 9’. Cystoliths in enlarged epidermal cells (lithocysts) absent. 11. Phloem of vascular bundles with resiniferous ducts. 12. Midvein keel-shaped on the adaxial side. Mesophyll isobilateral. 10. Schinus weinmanniifolia (Fig. 1) 12’. Midvein not keel-shaped on the adaxial side. Mesophyll dorsiventral. 13. Petiole outline rounded, convex on the adaxial side. Vascular tissue formed by an arc of abaxial collateral vascular bundles and three or more inverse vascular bundles disposed adaxially. 11. Astronium balansae 13’. Petiole outline semicircular, flat on the adaxial side. Vascular tissue formed by an arc of abaxial collateral vascular bundles but without vascular bundles disposed adaxially. 12. Schinus longifolia var. longifolia 11’. Phloem of vascular bundles without resiniferous ducts. 14. Parenchyma with raphides. 13. Phytolacca dioica 14’. Parenchyma without raphides. 15. Petiole and/or leaf blade midvein with idioblastic sclereids in the parenchyma.
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Bol. Soc. Argent. Bot. 43 (1-2) 2008
16. Epidermis crystalliferous. Mesophyll isobilateral. 14. Maytenus ilicifolia (Fig. 2) 16’. Epidermis non crystalliferous. Mesophyll dorsiventral. 17. Midvein with one bicollateral vascular bundle. Collenchyma absent. Parenchyma with laticiferous tubes. 15. Aspidosperma polyneuron (Fig. 2) 17’. Midvein with one to several collateral vascular bundles. Collenchyma present. Parenchyma without laticiferous tubes. 18. Hypodermis present. Petiole outline rounded without winged lobes. 16. Hennecartia omphalandra (Fig. 2) 18’. Hypodermis absent. Petiole outline rounded with two winged lobes. 17. Inga verna subsp. affinis 15’. Petiole and/or leaf blade midvein without idioblastic sclereids in the parenchyma. 19. Parenchyma with crystal sand. 20. Midvein with bicollateral vascular bundles. 21. Epidermis pubescent, with stalked, multiangulate stellate trichomes. 18. Solanum granulosum-leprosum 21’. Epidermis glabrous or with other type of trichomes. 19. Nicotiana glauca 20’. Midvein with collateral vascular bundles. 22. Midvein with a collateral vascular bundle. Palisade parenchyma 1-layered. 20. Sambucus australis 22’. Midvein with an arc of collateral vascular bundles and inverse vascular bundles disposed adaxially. Palisade parenchyma 2-3 layered. 21. Patagonula americana 19’. Parenchyma without crystal sand.
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A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees 23. Mesophyll with multilayered and collenchymatous hypodermis under both leaf blade surfaces. Petiole with two rings of collateral vascular bundles with the xylem facing one another. 22. Schefflera morototoni (Fig. 2) 23’. Mesophyll without these characteristics. 24. Abaxial epidermis with woolly pubescence (densely covered with long tangled trichomes). 25. Adaxial epidermis 1-layered. Mesophyll dorsiventral. Petiole outline subcircular with a horseshoe-shaped vascular tissue. 23. Luehea divaricata 25’. Adaxial epidermis multilayered. Mesophyll indifferentiated, with two layers of palisade parenchyma. Petiole outline lobate with a lobate ring of vascular tissue. 24. Cecropia pachystachya (Fig. 3) 24’. Abaxial epidermis with trichomes or not, but never woolly pubescent. 26. Midvein and petiole with 3-9 separated collateral vascular bundles arranged in a row. 25. Tessaria integrifolia var. integrifolia 26’. Midvein and petiole with one or numerous collateral or bicollateral vascular bundles never distributed in a row. 27. Stomata anomocytic, anisocytic, diacytic and/or ciclocytic. 28. Midvein keel-shaped on the adaxial side. 29. Mesophyll dorsiventral. Trichomes glandular multicellular, and non-glandular 1-multicellular. 26. Allophylus edulis 29’. Mesophyll isobilateral. Trichomes T-shaped. 27. Pouteria salicifolia 28’. Midvein not keel-shaped on the adaxial side. 30. Leaf blade surfaces pubescent (trichomes glandular and non-glandular). 31. Trichomes T-shaped.
28. Pouteria gardneriana
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 31’. Trichomes of other types. 32. Stellate trichomes present. Parenchyma with conspicuous secretory cavities. 33. Leaf blade amphistomatic with a few stomata adaxially. Petiole with a ring of separated vascular bundles. 34. Glandular trichomes with 1-celled head. 29. Monteiroa glomerata 34’. Glandular trichomes with multicellular head. 30. Hibiscus striatus 33’. Leaf blade hypostomatic. Petiole with a continuous ring of vascular tissue. Glandular trichomes with multicellular head. 31. Pavonia sepium subsp. sepium 32’. Stellate trichomes absent. Parenchyma without (at least inconspicuous) secretory cavities. 35. Non-glandular and verrucose trichomes over pedestal. 36. Midvein with one abaxial collateral vascular bundle and two inverse bundles disposed adaxially. Leaf blade hypostomatic. 32. Lantana camara 36’. Midvein with one abaxial collateral vascular bundle and three or more inverse bundles disposed adaxially. Leaf blade amphistomatic. 33. Lippia alba 35’. Non-glandular and smooth or verrucose trichomes without pedestal. 37. Midvein with medullar vascular bundles. 34. Cedrela fissilis
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37’. Midvein without medullar vascular bundles. 38. Midvein with vascular tissue distributed in a horseshoe-shaped. Petiole with an arc of vascular bundles. 35. Terminalia australis 38’. Midvein with vascular tissue not distributed in a horseshoe-shaped. Petiole with a circle of numerous vascular bundles. 36. Ruprechtia salicifolia (Fig. 3) 30’. Leaf blade surfaces glabrous or glabrate. 39. Parenchyma with druses. 40. Epidermis 2-layered. Parenchyma with large secretory cavities. 37. Ceiba speciosa 40’. Epidermis 1-layered. Parenchyma without large secretory cavities. 41. Hypodermis present. 42. Hypodermis abaxial. Large styloids within the palisade cells. Petiole with three separated vascular bundles. 38. Phyllanthus sellowianus 42’. Hypodermis abaxial and adaxial Styloids absent. Petiole with one vascular bundle. 39. Quillaja brasiliensis (Fig. 3) 41’. Hypodermis absent. 43. Midvein with bicollateral vascular bundles. 44. Epidermis with glandular and non-glandular trichomes. Anticlinal palisade cell walls sinuous. 40. Cestrum parqui
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 44’. Epidermis with only glandular trichomes. Anticlinal palisade cell walls not sinuous. 41. Cestrum euanthes 43’. Midvein with collateral vascular bundles. 45. Midvein with a medullar secretory duct. Petiole and petiolule with medullar and perimedullar secretory ducts. 42. Picrasma crenata (Fig. 3) 45’. Midvein, petiole and petiolule without secretory ducts. 43. Gleditsia amorphoides var. amorphoides 39’. Parenchyma without druses. 46. Epidermis with T-shaped trichomes. 28. Pouteria gardneriana 46’. Epidermis without trichomes. 44. Chrysophyllum marginatum 27’. Stomata paracytic. 47. Porrect stellate trichomes and glandular trichomes with 1-celled head present. 45. Croton urucurana 47’. Porrect stellate trichomes and glandular trichomes with 1-celled head absent. 48. Leaf blade hypostomatic. 49. Petiole with two adaxial obtuse or winged lobes. 50. Petiole with winged lobes. Midvein with a bicollateral vascular bundle. 46. Rauvolfia sellowii 50’. Petiole with obtuse lobes. Midvein with collateral vascular bundles.
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A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees 51. Midvein with numerous collateral vascular bundles, disposed abaxially, laterally and adaxially. Parenchyma with secretory ducts. 47. Rheedia brasiliensis (Fig. 4) 51’. Midvein with one vascular bundle. Parenchyma without secretory ducts. 48. Senna corymbosa 49’. Petiole without adaxial lobes. 52. Petiole with a ring of vascular tissue. 53. Petiole with subepidermal parenchyma, and collenchyma disposed internally. Midvein with druses. 49. Manihot flabellifolia 53’. Petiole with subepidermal collenchyma, and aerenchyma disposed internally. Midvein with macles non druses. 50. Erythrina crista-galli 52’. Petiole without a ring of vascular tissue. 54. Druses absent. Parenchyma with oil cells. 55. Petiole with vascular tissue surrounded by fibres. 51. Ocotea diospyrifolia 55’. Petiole with vascular tissue not surrounded by fibres. 56. Parenchyma with styloids and microcrystals. 52. Ocotea acutifolia 56’. Parenchyma with solitary crystals and macles. 53. Nectandra angustifolia 54’. Druses present. Parenchyma without oil cells. 54. Casearia sylvestris
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 48’. Leaf blade amphistomatic. 57. Laticiferous tubes present. 58. Mesophyll dorsiventral. 59. Midvein with a bicollateral, V-shaped, vascular bundle. Latex with resinous appearance. 55. Tabernaemontana catharinensis 59’. Midvein with a collateral, never V-shaped, vascular bundle. Latex with translucent milky appearance. 56. Sebastiania brasiliensis 58’. Mesophyll isobilateral. 57. Sapium haematospermum 57’. Laticiferous tubes absent. 58. Salix humboldtiana 8’. Midvein flatened (flat or concave) on the adaxial side. 60. Epidermis with scale peltate trichomes. 61. Scale flattened with entire margin. 62. Adaxial epidermis 1-layered. 63. Petiole with secretory cavities in the cortex. 64. Mesophyll with druses. 59. Fagara hyemalis 64’. Mesophyll without druses. 65. Stellate trichomes present. Midvein with vascular tissue surrounded by sclerenchyma. 60. Fagara rhoifolia 65’. Stellate trichomes absent. Midvein with vascular tissue surrounded by parenchyma. 61. Esenbeckia febrifuga 63’. Petiole without secretory cavities in the cortex. 62. Myrsine laetevirens 62’. Adaxial epidermis 2-layered.
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A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees 63. Tabebuia heptaphylla 61’. Scale flattened with irregularly notched margin. Mesophyll with druses. 64. Styrax leprosus (Fig. 4) 60’. Epidermis glabrous or with other type of trichomes. 66. Trichomes with cystolith-like structures. 67. Mesophyll with crystal sand. 65. Cordia curassavica 67’. Mesophyll without crystal sand. 68. Petiole two lobed adaxially, each lobe with one vascular bundle. 66. Aloysia virgata var. virgata 68’. Petiole two lobed adaxially, each lobe with two vascular bundle. 67. Aloysia gratissima var. gratissima 66’. Trichomes without cystolith-like structures. 69. Presence of T-shaped trichomes (malpighiaceous). 70. Adaxial epidermis glabrous. 68. Gochnatia polymorpha subsp. ceanothifolia 70’. Adaxial epidermis pubescent. 69. Heteropteris glabra 69’. Absence of T-shaped trichomes (malpighiaceous). 71. Epidermis with candelabraform trichomes. 72. Glandular trichomes with 2-multicellular head. 70. Buddleja thyrsoides 72’. Glandular trichomes absent. 71. Buddleja brasiliense 71’. Epidermis without candelabraform trichomes. 73. Petiole with idioblastic sclereids in the cortex. 74. Epidermis pubescent with glandular and non-glandular trichomes. Petiole with a ring of vascular tissue. 72. Guarea macrophylla subsp. spicaeflora
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 74’. Epidermis glabrate with sparse pubescence of non-glandular trichomes. Petiole with an arc of separated vascular bundles. 73. Rollinia emarginata 73’. Petiole without idioblastic sclereids in the cortex. 75. Parenchyma with druses. 76. Mesophyll formed exclusively by palisade parenchyma. 74. Bauhinia forficata subsp. pruinosa 76’. Mesophyll differentiated (isobilateral or dorsiventral). 77. Mesophyll isobilateral. 78. Hypodermis present. Stomata anomocytic. Few non-glandular and unicellular trichomes over the abaxial (dorsal) face of the petiole. 75. Myrcianthes pungens 78’. Hypodermis absent. Stomata paracytic. Trichomes absent. 76. Blepharocalyx salicifolius 77’. Mesophyll dorsiventral. 79. Parenchyma of the leaf blade and petiole with secretory cavities. 80. Stomata ciclocytic. Trichomes glandular peltate, sunken in epidermal depressions. 77. Pilocarpus pennatifolius 80’. Stomata not ciclocytic. Trichomes of other type or absent. 81. Hypodermis present (adaxially). 82. Hypodermis 1-layered. 83. Mesophyll with palisade parenchyma 3-4-layered. Midvein with chlorenchyma adaxial. 78. Myrceugenia glaucescens 83’. Mesophyll with palisade parenchyma 1-layered. Midvein with collenchyma adaxial. 79. Eugenia uruguayensis 82’. Hypodermis 2-layered. Mesophyll
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A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees dorsiventral with palisade with parenchyma palisade parenchyma 2-layered. 2-layered. 80. Hexaclamys edulis 81’. Hypodermis absent. 84. Midvein concave adaxially. Mesophyll with palisade parenchyma 2-layered. 81. Eugenia involucrata 84’. Midvein flat adaxially. Mesophyll with palisade parenchyma 1-layered. 85. Petiole with vascular tissue surrounded by mechanical tissue not lignified. Druses numerous. 82. Eugenia uniflora 85’. Petiole with vascular tissue surrounded by mechanical tissue lignified. Druses scarce. 83. Eugenia pitanga 79’. Parenchyma of the leaf blade and petiole without secretory cavities. 86. Adaxial epidermis with secretory structures. Abaxial epidermis conspicuously papillose. 84. Jacaratia spinosa (Fig. 4) 86’. Adaxial and abaxial epidermis without these characteristics. 87. Petiole with adaxial winged lobes. 88. Epidermis crystaliferous. Midvein with an arc of 3-4 separated collateral vascular bundles. 85. Trixis divaricata subsp. divaricata 88’. Epidermis not crystaliferous. Midvein with one collateral vascular bundle. 86. Vernonia scorpioides 87’. Petiole with or without adaxial lobes, but never winged. 89. Petiole with two obtuse lobes. Vascular tissue dispossed in a horseshoe-shaped. 87. Ilex paraguariensis
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 89’. Petiole without lobes. 90. Hypodermis present, adaxially. Petiole outline rounded, with a ring of vascular bundles, and groups of phloem in medullar position. 88. Vitex megapotamica 90’. Hypodermis absent. Petiole outline semicircular, with only one vascular bundle, and without groups of phloem in medullar position. 89. Symplocos uniflora 75’. Parenchyma without druses. 91. Parenchyma with raphides. 90. Psychotria carthagenensis 91’. Parenchyma without raphides. 92. Parenchyma with crystal sand. 91. Coutarea hexandra f. pubescens (Fig. 4) 92’. Parenchyma without crystal sand. 93. Leaf blade with sclerenchymatic tissue under both epidermis and in the midvein around the vascular bundle. Petiole with sclerenchymatic cortex. 92. Berberis ruscifolia 93’. Leaf blade and petiole without these characteristics. 94. Trichomes 3-4-celled with a flagellate apical cell. 93. Baccharis coridifolia 94’. Trichomes, when present, without flagellate apical cell. 95. Stomata anomocytic, anisocytic and/or diacytic. 96. Petiole with adaxial winged lobes. 97. Midvein with one collateral vascular bundle. 98. Petiole with discontinuous subepidermal collenchyma. Epidermis only with glandular
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A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees trichomes. 94. Eupatorium ceratophyllum 98’. Petiole without collenchyma. Epidermis with glandular and non-glandular trichomes. 95. Eupatorium inulifolium 97’. Midvein with an arc of collateral vascular bundles disposed abaxially and a variable number of inverse vascular bundles adaxially. 96. Lantana megapotamica 96’. Petiole without adaxial winged lobes. 99. Epidermis pubescent. 100. Glandular trichomes present. Non-glandular trichomes over pedestals. 97. Lantana montevidensis 100’. Glandular trichomes absent. Non-glandular trichomes with other characteristics. 101. Epidermis papillose on both surfaces. Midvein located asymmetrically. 98. Enterolobium contortisiliquum 101’. Epidermis non-papillose. Midvein located symmetrically. 102. Leaf blade hypostomatic. Parenchyma with laticiferous tubes. 99. Sorocea bonplandii 102’. Leaf blade amphistomatic. Parenchyma without laticiferous tubes. 100. Sesbania punicea 99’. Epidermis glabrous. 103. Petiole with two adaxial lobes. Mesophyll with secretory cavities. 104. Leaf blade hypostomatic. Mesophyll dorsiventral with
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 palisade parenchyma multilayered. 101. Myrocarpus frondosus 104’. Leaf blade amphistomatic. Mesophyll non-dorsiventral. 102. Poiretia tetraphylla 103’. Petiole without lobes. Mesophyll without secretory cavities. 103. Chrysophyllum gonocarpum 95’. Stomata paracytic. 105. Midvein flat on both sides, adaxial and abaxial. Oil cells and cavities absent. 106. Epidermis abaxial papillose. Midvein located asymmetrically. 104. Parapiptadenia rigida 106’. Epidermis abaxial non-papillose. Midvein located symmetrically. 105. Calliandra parvifolia 105’. Midvein flat only on the adaxial side. Oil cells and cavities present. 107. Petiole pubescent. Mesophyll with microcrystals. 106. Nectandra lanceolata 107’. Petiole glabrous. Mesophyll without microcrystals.
107. Ocotea puberula
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Fig. 1. Leaf characteristics in transection and surface view. A-D: Araucaria angustifolia. A, leaf blade showing isobilateral mesophyll; compartmented cells and resiniferous ducts among vascular bundles, and idioblastic sclereids; B, thickness of epidermal anticlinal cell walls in surface view; C, a ciclocytic stoma with cuticular thickening of the guard cell polar regions; D, a schyzogenous duct. E-H: Guadua chacoensis. E, leaf blade transection, indifferentiated mesophyll, translucent cells nearly vascular bundles; F, sheath transection; G, hook and bicellular trichomes, large and short cells (suberose and silicose); H, abaxial epidermis slightly papillose. I-L: Cordia ecalyculata. I, midvein with an arc of abaxial separated vascular bundles and some disposed adaxially; J, petiole; K, adaxial epidermis, one cystolith in transection; L, adaxial epidermis showing the peduncle of one cystolith in surface view. M-P: Schinus weinmanniifolia. M, midvein keel-shaped adaxially; N, petiole transection; O, petiolule winged adaxially; P, abaxial epidermis showing anticlinal cell wall puntuations, cuticular ornamentation, and an anomocytic stoma. Bars: A, F = 300 μm; B-D, G, P = 50 μm; E, K, L = 100 μm; H = 30 μm; I, M = 500 μm; J, N, O = 1 mm.
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Fig. 2. Leaf characteristics in transection and surface view. A-D: Maytenus ilicifolia. A, midvein and mesophyll isobilateral; B, petiole, parenchyma with idioblastic sclereids; C, adaxial epidermis crystalliferous, thick anticlinal cell walls; D, abaxial epidermis crystalliferous, thin anticlinal cell walls. E-H: Aspidosperma polyneuron. E, midvein, collenchyma absent and a bicollateral vascular bundle; F, petiole, numerous clusters of idioblastic sclereids; G, adaxial epidermis showing cuticular ornamentation; H, abaxial epidermis with an anomocytic stoma. I-L: Hennecartia omphalodes. I, midvein with adaxial chlorenchyma under collenchyma; J, petiole outline rounded with a ring of collenchyma and clusters of idioblastic sclereids; K, adaxial epidermis and hypodermis in transection; L, abaxial epidermis showing a paracytic stoma in surface view. M-O: Schefflera morototoni. M, midvein with numerous secretory ducts; N, portion of petiole in transection, showing the two rings of vascular bundles with the xylem facing one another; O, petiolule. Bars: A, E, I = 500 μm; B, C = 100 μm; D, G, H, K, L = 50 μm; F, J, M, N, O = 1 mm.
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A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
Fig. 3. Leaf characteristics in transection and surface view. A-D: Cecropia pachystachya. A, midvein, collenchyma adaxial and abaxial; mesophyll indifferentiated (palisade parenchyma); B, portion of the lobate petiole and vascular tissue; C, adaxial epidermis in surface view, showing a conical secretory structure; D, adaxial epidermis multilayered with a secretory structure in transection. E-H: Ruprechtia salicifolia. E, midvein with numerous separated vascular bundles and abundant sclerenchyma; F, petiole with a circle of numerous vascular bundles; G, adaxial epidermis in surface view; H, abaxial epidermis exhibiting a stoma, in surface view. I-M: Quillaja brasiliensis. I, midvein, one collateral vascular bundles with xylematic and phloematic cups of sclerenchyma; J, petiole pubescent; K, adaxial epidermis and hypodermis in transection, showing the thickness of outer periclinal epidermal cell walls and a strong cuticle; L, adaxial epidermis in surface view; M, a non-glandular trichome. N-Q: Picrasma crenata. N, midvein, adaxial chlorenchyma lower collenchyma, one medular secretory duct; O, petiole, subcircular outline, a ring of vascular tissue and medullary bundles, and numerous secretory ducts; P, petiolule; Q, abaxial epidermis with cuticular ornamentation. Bars: A, B, F, O, P = 1 mm; C, D, G, H, K, L, Q = 50 μm; E = 300 μm; I, J, N = 500 μm; M = 25 μm.
49
Bol. Soc. Argent. Bot. 43 (1-2) 2008
Fig. 4. Leaf characteristics in transection and surface view. A-D: Rheedia brasiliensis. A, midvein showing chlorenchyma adaxial and the vascular tissue surrounded by a ring of sclerenchyma; B, petiole; C, adaxial epidermis with thick anticlinal cell walls, in surface view; D, abaxial epidermis with a paracytic stoma and cuticular ornamentation. E-I: Styrax leprosus. E, midvein flattened adaxially; F, petiole; G, a scale peltate trichome in surface view; H, adaxial epidermis with anticlinal cell walls thickened; I, epidermis transection showing enlarged epidermal cells in anticlinal direction, and palisade parenchyma with an idioblastic cell containing a druse. J-N: Jacaratia spinosa. J, midvein concave and without collenchyma, adaxially; K, portion of petiole showing the cortex and then a ring of sclerenchyma surrounding the vascular tissue; L, adaxial epidermis in surface view showing a secretory structure; M, adaxial epidermis exhibiting a secretory structure in transection; N, abaxial epidermis conspicuously papillose, in transection. O-R: Coutarea hexandra f. pubescens. O, midvein flattened adaxially; P, petiole conspicuously pubescent; Q, adaxial epidermis showing a non-glandular trichome; R, abaxial epidermis with a paracytic stoma. Bars: A, E, F, J, O, P = 500 μm; B, K = 1 mm; C, D, G-I, L = 50 μm; M, N, Q, R = 100 μm.
50
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
Conclusions Character value Epidermal features and leaf anatomy have high diagnostic value at specific level and/or at family level: (1) crystaliferous epidermis characterized two species Maytenus ilicifolia and Trixis divaricata subsp. divaricata; (2) cystolith-like structure in trichomes are present in Aloysia virgata var. virgata and Cordia curassavica. This structure in trichomes confirms the relationships among the families Verbenaceae and Boraginaceae to which the species are belong respectively; (3) the ciclocytic stomata type is present in Araucaria angustifolia and Pilocarpus pennatifolius; (4) Cordia ecalyculata has cystoliths in its uniseriate epidermis; (5) Schefflera morototoni can be distinguished by its multilayered and colenquimatous hypodermis distributed on both surfaces, and the presence of two cicles of vascular bundles in the petiole; (6) Cecropia pachystachya is characterized by having the leaf blade mesophyll formed by palisade parenchyma exclusively, and lobed petiole outline; (7) scale peltate trichomes are diagnostic epidermal traits in Styrax leprosus (Styracaceae) and Tabebuia heptaphylla (Bignoniaceae), and located in epidermal depressions in Esenbeckia febrifuga (Rutaceae); (8) laticiferous tubes recognized by their coloured latex are present in Sapotaceae and Apocynaceae; (9) bicollateral vascular bundles are diagnostic leaf traits of the families: Apocynaceae, Myrtaceae and Solanaceae; 10) secretory ducts into the phloematic parenchyma are conspicuous and a definite microcharacter in family Anacardiaceae; 11) epidermal heteromorphic cells over the secretory cavities are a diagnostic trait in Myrtaceae; 12) paracytic stomata and oil cells characterized the family Lauraceae; 13) very small druses uniquely were found in Verbenaceae, and 14) large secretory cavities are diagnostic features relating the families Bombacaceae and Malvaceae. Ecological implications
The species studied show predominantly a combination of mesomorphic and xeromorphic leaf structures, e.g. Aspidosperma polyneuron, Ceiba speciosa, Gleditsia amorphoides var. amorphoides, Hexaclamys edulis. As mesomorphic characters can be mentioned: (1) the majority of studied species have hypostomatic leaves, and the stomata lie at epidermis level; (2) thirty eight of the total forty five analyzed taxa have dorsiventral leaves; (3) most of species exhibit a well developed spongy parenchyma rich in intercellular spaces. Within xeromorphic features may be mentioned: (1) anticlinal cell walls straight to curved; (2) anticlinal and outer periclinal epidermal cell walls thick; (3) cuticle thick and/or ornamented; (4) epidermis multilayered or when 1-layered formed by large cells; (5) hypodermis on one or both leaf surfaces. However, twenty six species have also some hygromorphic features such as epidermis glabrous, anticlinal cell walls undulate, leaves amphistomatic or/and stomata elevated, e.g. Aloysia virgata var. virgata, Chrysophyllum spp., Cordia spp., Guadua chacoensis, Jacaratia spinosa, Myrcianthes pungens, Sebastiania brasiliensis, Tabernaemontana catharinensis, Trixis divaricata subsp. divaricata. Only two species, Ilex paraguariensis and Manihot grahamii have mesomorphic and hygromorphic traits. Most of xeromorphic features / sun leaf type, such as a thick and/or ornamented cuticle, e.g. Pilocarpus pennatifolius (Rutaceae); thick cuticle and abundant sclerenchyma e.g. Gleditsia amorphoides var. amorphoides (Fabaceae); multilayered epidermis, e.g. Cecropia pachystachya (Cecropiaceae); presence of hypodermis 2-layered e.g. Hexaclamys edulis (Myrtaceae); collenchymatous and multilayered hypodermis e.g. Schefflera morototoni (Araliaceae); mesophyll with palisade parenchyma multilayered e.g. Patagonula americana (Boraginaceae); mesophyll formed exclusively by palisade parenchyma e.g. Cecropia pachystachya (Cecropiaceae); parenchyma with idioblastic sclereids e.g. Aspidosperma polyneuron (Apocynaceae), seems to be an adaptation of the big trees to exposed situations with high level of solar irradiance.
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52
Acellws thick and straight; Ecw conspicuous; E and H sclerophyllous; H multilayered (both faces); St sunken; M with idioblastic sclereids Ecw conspicuous; C thick; Pp 1-2-layered; Scl abundant; Idioblastic sclereids present C striate (abx face)
Araucaria angustifolia
E 2-layered; C striate (abx face); Scl abundant; Pp 1-2-layered Ecw conspicuous; C thick; Scl abundant C striate (abx face)
Ceiba speciosa
C striate; E pubescent Pp (large cells) Ecw conspicuous; C striate; E pubescent
Coutarea hexandra f. pubescens
Esenbeckia febrifuga
Cordia ecalyculata
Pparenchyma 1-2-layered; E pubescent Pp (large cells)
Cordia curassavica
Chrysophyllum marginatum
Chrysophyllum gonocarpum
Cedrela fissilis
E multilayered with secretory cells; M indifferentiated (totally Pp); E pubescent E pubescent; Pp (large cells)
Cecropia pachystachya
Astronium balansae
Aspidosperma polyneuron
Xeromorphic / sun leaf type H 1-layered (adx and discontinuous) ; E pubescent
Taxa Aloysia virgata var. virgata
L hypostomatic; M dorsiventral; Pp 1-layered Acellws curved; L hypostomatic; M dorsiventral; Pp 1-layered (short cells)
Acellws curved (adx face); L hypostomatic; M dorsiventral; Pp 1-layered; Scl +/- abundant Acellws curved; L hypostomatic; M dorsiventral; E subglabrous L hypostomatic; M dorsiventral; Pp 1-2-layered (short cells) L hypostomatic; M dorsiventral; Pp 1-layered (short cells); Scl +/- abundant Acellws curved; L hypostomatic; M dorsiventral L hypostomatic; M dorsiventral; Pp 1-layered; Scl +/- abundant
Acellws straight to slightly curved; L hypostomatic
Acellws curved; L hypostomatic; M dorsiventral; Pp 1-2-layered (short cells); Scl +/- abundant; E subglabrous
Acellws curved; L hypostomatic; M dorsiventral; E subglabrous
Mesomorphic / medium leaf type Acellws straight to curved; L hypostomatic; M dorsiventral to indifferentiated
Acellws undulate; Scl reduced / absent Scl reduced / absent
Acellws undulate (abx); Scl reduced / absent Acellws undulate; E glabrous
Acellws undulate; E glabrous Acellws undulate; E glabrous
Acellws undulate (abx face)
Scl reduced / absent
L amphistomatic; M isobilateral E glabrous
Higromorphic / shade leaf type St elevated; Scl reduced / absent
Table 1. List of studied species and ecological leaf characteristics elaborated on the basis of criteria of the xeromorphy, mesomorphy, hygromorphy, sun leaf type, medium leaf type and shade leaf type of Roth (1984). Abbreviations: Acellws: anticlinal cell walls; abx: abaxial; adx: adaxial; C: cuticle; E: epidermis; Ecw: epicuticular waxes; H: hypodermis; L: leaves; M: mesophyll; Pcellws: periclinal cell walls; Pp: palisade parenchyma; Scl: sclerenchyma; St: stomata.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
Patagonula americana
Parapiptadenia rigida
Ocotea puberula
Ocotea diospyrifolia
Nectandra lanceolata
Myrocarpus frondosus
Myrcianthes pungens
Maytenus ilicifolia
Manihot grahamii
Jacaratia spinosa
Ilex paraguariensis
Hexaclamys edulis
Hennecartia omphalandra
Guadua chacoensis
Gleditsia amorphoides var. amorphoides Gochnatia polymorpha subsp. ceanothifolia
Table 1. Cont.
Pcellws thick; Pp multilayered (cells not so long)
Acellws straight; Scl abundant; Pp 2-layered; E pubescent Acellws straight; Pp 1(-2)-layered; Scl abundant Acellws straigh (adx) and thick; Pp 2-layered; Scl abundant; E pubescent E pubescent; Pp (large cells)
Acellws straight; Ecw conspicuous; C thick C thick; H 1-layered (adx); Scl abundant Ecw conspicuous; H 1-layered; Pp multilayered
Acellws straight; E secretory cells; Ecw conspicuous (abx); Pp (large cells)
Ecw conspicuous; C thick; Scl abundant; E pubescent Acellws straight (adx); C thick; St with stomatal rim; Pp multilayered; E pubescent Acellws straight and thick; Ecw conspicuous; C thick; Scl abundant Acellws thick; Ecw conspicuous; C thick; H 1-layered (adx); E pubescent Acellws thick; Ecw conspicuous; C thick; H 2-layered (adx); Pp 2-layered
Acellws curved; L hypostomatic; M dorsiventral; Pp 1-layered; Scl +/- abundant Acellws straight to curved; L hypostomatic; M dorsiventral;
Acellws straight to curved; L hypostomatic; M dorsiventral; Scl +/- abundant; E subglabrous L hypostomatic; M dorsiventral L hypostomatic; M dorsiventral; E subglabrous Acellws curved (abx); L hypostomatic; M dorsiventral
Acellws curved; L hypostomatic; M dorsiventral 1-layered (short cells); Scl +/- abundant Acellws curved; L hypostomatic; M dorsiventral; Scl +/- abundant; E subglabrous Acellws curved; L hypostomatic; M dorsiventral; Pp 1-layered (short cells); Scl +/- abundant L hypostomatic ; M dorsiventral; Pp 1-layered L hypostomatic; M dorsiventral; Pp 1-layered (short cells) L hypostomatic ; Scl +/- abundant L hypostomatic
Acellws curved; L hypostomatic; M dorsiventral; Pp 1-layered (short cells) Acellws curved (abx); L hypostomatic; M dorsiventral; Scl +/- abundant E subglabrous
Scl reduced / absent; E glabrous Acellws undulate; Scl reduced / absent; E glabrous M isobilateral; E glabrous Acellws undulate; M isobilateral; E glabrous
E glabrous
L amphistomatic; M indifferentiated formed by isodiametric cells
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
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54
S
b
l
dii
Sebastiania brasiliensis
Schinus weinmanniifolia
Schefflera morototoni
Ruprechtia salicifolia
Rheedia brasiliensis
Rauvolfia sellowii
Quillaja brasiliensis
Psychotria carthagenensis
Pilocarpus pennatifolius
Picrasma crenata
Table 1. Cont.
i
C striate (abx face); Pp (large cells) Acellws straight and thick; Ecw conspicuous; C striate; Pp (large cells); E pubescent Acellws straight; Ecw conspicuous; E multilayered with secretory cells; Pp (large cells) H 1-layered (adx and abx); M isobilateral to indifferentiated; Pp multilayered (adx and abx) Acellws straight; Pp (large cells) Acellws straight and thick; C very thick; Scl abundant Acellws straight; Pp multilayered; Scl abundant Acellws straight; H multilayered (both faces); St sunken with peristomatal rim; Pp 2-layered; E pubescent Acellws straight and thick; C striate (abx face); E pubescent C striate; Pp (large cells) ll
d
d l
L hypostomatic with some stomata located adaxially M dorsiventral; Pp 1-layered
L hypostomatic; M dorsiventral; Pp 1-layered; L hypostomatic; M dorsiventral; Pp 1-2 -layered (short cells) L hypostomatic; M dorsiventral; E subglabrous L hypostomatic; M dorsiventral; Scl +/- abundant
L hypostomatic; M dorsiventral; Pp 1-layered; Scl +/- abundant; E subglabrous Acellws curved; L hypostomatic; Scl +/- abundant; E subglabrous
Scl +/- abundant; E subglabrous L hypostomatic; M dorsiventral ; Pp 1-layered; Scl +/- abundant L hypostomatic; M dorsiventral; Pp 1-layered; Scl +/- abundant
M isobilateral; Scl reduced / absent Acellws undulate; L amphistomatic; Scl reduced / absent; E glabrous
Scl reduced / absent; E glabrous E glabrous
Acellws undulate; E glabrous
Bol. Soc. Argent. Bot. 43 (1-2) 2008
Acellws undulate; Scl reduced / absent
Scl reduced / absent; E glabrous
Appendix 1. List of studied species with specimen vouchers and ethnobotany use. The numbers within parentheses indicate the references listed at the end. Abbreviation: Jardín Botánico y Arboretum «C. Spegazzini», Facultad de Ciencias Agrarias y Forestales, UNLP: FCAF.
E pubescent
Ecw conspicuous; Pp multilayered; E pubescent Vitex megapotamica
Tabernaemontana catharinensis
Trixis divaricata subsp. divaricata
Acellws curved; L hypostomatic to amphistomatic; M dorsiventral; Pp 1-layered (short cells) L hypostomatic; M dorsiventral; Pp 1-layered (short cells) Acellws curved; L hypostomatic; M dorsiventral; Scl +/- abundant
Acellws curved; L hypostomatic; M dorsiventral; Scl +/- abundant
Acellws straight and thick; Ecw conspicuous; Epidermal cells large; E pubescent Ecw conspicuous; C striate; E 2-layered; Pp 2-layered (large cells); E pubescent C striate Styrax leprosus
Tabebuia heptaphylla
Ecw conspicuous; E pubescent Sorocea bonplandii
Table 1. Cont.
Acellws curved to undulate; L hypostomatic; M dorsiventral; Pp 1-2 –layered (short cells); Scl +/- abundant L hypostomatic; M dorsiventral; Pp 1-layered (short cells)
Scl reduced / absent
Scl reduced / absent; E glabrous
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
Aloysia virgata (Ruiz & Pav.) Juss. var. virgata (Verbenaceae). Misiones, Parque Nac. Iguazú, 4XI-1970, Dimitri & Amorín 58 (LPAG). Leaves, stems and flowers: to reduce menstruation. Leaves: external use as antimicotic (1). Araucaria angustifolia (Bertol.) Kuntze (Araucariaceae). Buenos Aires, La Plata, FCAF, 8IX-2006, Martínez 108 (LPAG 4424). Misiones, Eldorado, 2-III-2005, Martínez 35 (LPAG). Resin: antiseptic (2). Seeds: edible (3, 4). Aspidosperma polyneuron Müll. Arg. (Apocynaceae). Misiones, Parque Nac. Iguazú, 17-IX-1970, Dimitri & Amorín 68 (LPAG); Eldorado, 22-III-2005, Martínez 43 (LPAG). Expectorant, anticatarrhal (5). Astronium balansae Engl. (Anacardiaceae). Santa Fe, General Obligado, 10-I-1937, Ragonese 2413 (LP). Leaves: analgesic, febrifuge, in treatment of insolation (6); analgesic, to treat cephalalgia (2); purgative (1). Cecropia pachystachya Trécul (Cecropiaceae). Corrientes, Capital, 5-X-1973, sin colector, s.n. (LPAG 1004). Misiones, Eldorado, 23-III-2005, Martínez 46 (LPAG); Parque Nac. Iguazú, 16-IX1970, Dimitri & Amorín 1181 (LPAG). Leaves: expectorant, antiasthmatic, bronchitis, sedative, diuretic, cardiotonic (oficial drug monographied in 5 th edition of the Farmacopea Nacional Argentina) (7). Leaves: tonic, pectoral, cardiotonic, diuretic, anticatarrhal, antiasthmatic (2, 5, 8). Leaves: are useful to control cough (1). Trunk juice is used against cutaneous bores and the twig juice has properties as antidiarrheic (5). Leaves and cork: expectorant, tonic and anti-inflammatory (9). Fruits: edible (9-11). This species is in 6th edition of the Farmacopea Nacional Argentina and it was incorporated in the Program for Atención Primaria de la Salud in Misiones (12, 13). Cedrela fissilis Vell. (Meliaceae). Buenos Aires, FCAF, XI-2004, Monti 26 (LPAG). Misiones, Eldorado, 23-III-2005, Martínez 36 (LPAG). Cortex and wood: febrifuge, anti-inflammatory, antimalarial (2, 3, 8). Cortex: febrifuge, emetic, astringent, hepatic. Leaves: to treat urinary
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 illnesses (13). Ceiba speciosa (A. St.-Hil.) Ravenna (Bombacaceae). Buenos Aires, La Plata, 17-VIII-2004, Stenglein 19 (LPAG); 30-IX-2005, Arambarri 252 (LPAG). Flowers and thorns: analgesic and diuretic (2, 14). Seeds have 15-20% of oil with industrial applications and edible. Cortex sap: to treat hernia. Thorns: analgesic, antiasthmatic and to treat alcoholism (4). Chrysophyllum gonocarpum (Mart. & Eichler) Engl. (Sapotaceae). Buenos Aires, La Plata, FCAF, 8XI-2005, Arambarri 254 (LPAG). Corrientes, Santa Teresa, 22-III-1954, Petersen 2644 (LP). Fruits: eaten directly or as candy. It were used by the Jesuits as medicine (3, 15). Chrysophyllum marginatum (Hook. & Arn.) Radlk. (Sapotaceae). Buenos Aires, La Plata, FCAF, 8XI-2005, Arambarri 253 (LPAG). Corrientes, Concepción, 4-V-1974, Petersen 1072 (LP). Leaves and leafy stems: expectorant (1). Fruits: edible (3). Cordia curassavica (Jacq.) Roem. & Schult. (Boraginaceae). Corrientes, Mburucuyá, 9-X-1954, Cabrera 11680 (LP). Jujuy, Ledesma, 15-X-1964, Cabrera & Fabris 15971 (LP). Leaves and leafy stems: vermifuge, antispasmodic, digestive, antitoxic, to cure cough (1). Leaves: antiinflammatory (16). Cordia ecalyculata Vell. (Boraginaceae). Misiones, Eldorado, 30-III-2003, Martínez 59 (LPAG); Parque Nac. Iguazú, 6-XII-1971, Volkart s.n. (LPAG 4449). Leaves: carditonic, tonic (2). Fruits: with a viscid substance which is used as glue. Fruits are also palatable for wild animals (3). Coutarea hexandra (Jacq.) K. Schum. f. pubescens (Pohl) Steyerm. (Rubiaceae). Buenos Aires, La Plata, Jardín Sistemático de la Facultad de Agronomía, I-1969, Volkart s.n. (LPAG 4195). Misiones, Parque Nac. Iguazú, 9-I-1971, Dimitri s.n. (LPAG 4194). Cortex: febrifuge (17). Esenbeckia febrifuga (A. St.-Hil.) A. Juss. ex Mart. (Rutaceae). Argentina. Misiones. Puerto Iguazú. 13-XI-1969, Hualde 42 (LPAG 2386); 3-XI-1973 (LPAG 2389). Brasil, Paraná, 17-IV-1964, Hatschbach s.n. (LP). Cortex: febrifuge, substitute of quinina (Peruvian bark) (2); and with aphrodisiac properties (3). Gleditsia amorphoides (Griseb.) Taub. var. amorphoides (Fabaceae). Buenos Aires, La Plata, Jardín Botánico «C. Spegazzini», (FCAF) UNLP, 23-VIII-2004, Arambarri 246 (LPAG). Misiones, Eldorado, 29-III-2005, Martínez 47 (LPAG). Leaves
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and roots: astringent (18). Cortex: with properties to reduce cholesterol (19). Fruits: irritant which make sneeze (5, 15); to treat asthma (1). Fruits: produce a gum used in sweets industry, pharmacy and cosmetic (3, 20). Gochnatia polymorpha (Less.) Cabrera subsp. ceanothifolia (Less.) Cabrera (Asteraceae). Misiones, Candelaria, Loreto, Montes 11154 (LP); Corrientes, 6-II-1972, Santo Tomé, Krapovickas et al. 21091 (LP). Leaves and flowers: pectoral (2). Guadua chacoensis (Rojas Acosta) Londoño & P.M. Peterson (Poaceae). Corrientes, Ituzaingó, 8-XII1974, Krapovickas et al. 23851 (SI). Misiones, Montecarlo, I-1944, Porta s.n. (LP 900486); Parque Nac. Iguazú, 14-IX-1971, Dimitri 11246 (LPAG). Rhizomes: urinary antiseptic, laxative, analgesic (1, 21). Water acummulated in internodes is used to drink in wooded areas (3). Hennecartia omphalandra J. Poiss. (Monimiaceae). Misiones, Eldorado, 21-III-2003, Martínez 49 (LPAG). Plant have properties as digestive, antispasmodic (21). Hexaclamys edulis (O. Berg.) Kausel & Legrand (Myrtaceae). Buenos Aires, La Plata, FCAF, 15IX-1970, Hualde s.n. (LAPG); 19 VI-2006, Arambarri 259 (LPAG). Leaves: hepatic, digestive (1); antidiabetic (22, 23). Fruits: laxative (3). Ilex paraguariensis A. St.-Hil. (Aquifoliaceae). Buenos Aires, La Plata, FCAF, 29-III-2005, Arambarri s.n. (LPAG). Misiones, Eldorado, 23III-2005, Martínez 83 (LPAG). Leaves: tonic, stimulant, diuretic, digestive, sudorific (3, 24); leaves macerated to treat dandruff; cephalalgia; anti-inflammatory; diuretic (1, 25). This species was included in the 1st edition of the Farmacopea Nacional Argentina (13). Jacaratia spinosa (Aubl.) A. DC. (Caricaceae). Misiones, Parque Nac. Iguazú, 6-XI-1971, Dimitri s.n. (LPAG); Facultad de Ciencias Forestales, 23III-2005, Martínez 42 (LPAG). Latex: vermifuge and cathartic (2, 26); the plant has properties to control gastrointestinal worms (5). Fruits: edible (3). Manihot grahamii Hook. (Euphorbiaceae). Buenos Aires, Ensenada, 18-IV-2004, Bayón 636 (LPAG). Misiones, Eldorado, 29-III-2005, Martínez 37 (LPAG). Plant: drastic. (27), with toxic properties (28). Maytenus ilicifolia Mart. ex Reissek (Celastraceae). Buenos Aires, La Plata, FCAF, 20-VII-2006, Arambarri 262 (LPAG). Misiones, Montecarlo,
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees 26-III-05, Martínez 53 (LPAG). Vegetative parts: have properties as sialagogue, vulnerary, and in external use as antiseptic (2, 6, 7, 29). Leaves: antispasmodic, digestive and antiseptic (8). This species would has antitumoral properties (21). Extract from leaves would have anti-ulcer activity (30). This species is included in the Farmacopea Brasileña and it was introduced in the Program for Atención Primaria de la Salud en la provincia de Misiones (13). Myrcianthes pungens (O. Berg) D. Legrand (Myrtaceae). Buenos Aires, La Plata, Jardín Botánico de la Facultad de Agronomía, UNLP, 24III-1966, Amorín 148 (LPAG); Capital Federal, 7-I1991, Orfila s.n. (LPAG 4806). Fruits: edible (15, 31). Leaves: are tonic and astringent (2, 32). Myrocarpus frondosus Allemâo (Fabaceae). Misiones, Parque Nac. Iguazú, 20-XI-1969, Hualde 138 (LPAG); Eldorado, 21-III-2003, Martínez 48 (LPAG). Resin: antiseptic, cicatrizant, to control respiratory and urinary infections (2, 8). The aromatic resin extracted from the cortex is used as substitute of the true incense (3); and in perfume and tincture industry (4). Nectandra lanceolata Nees (Lauraceae). Buenos Aires, La Plata, FCAF, IV-2004, Monti 24a, 24b (LPAG). Misiones, Puerto Iguazú, 21-X-1970, Amorín & Dimitri s.n. (LPAG 1387). This species has properties to treat asthma (5). Ocotea diospyrifolia (Meisn.) Mez (Lauraceae). Misiones, Parque Nac. Iguazú, 18-XI-1969, Hualde s.n. (LPAG 1378). Leaves: carminative, emmenagogue, antirheumatic (1, 2). Ocotea puberula (Rich.) Nees (Lauraceae). Jujuy, Santa Bárbara, 6-I-1966, Cabrera 17367 (LP). Cortex: antidiarrhea (2); to apply as poultice to extract thorns or spines (5). Parapiptadenia rigida (Benth.) Brenan (Fabaceae). Buenos Aires, La Plata, FCAF, 5-V-1966, Amorín 352 (LPAG). Misiones, Parque Nac. Iguazú, 18XI-1969, Hualde 122 (LPAG). Cortex: antiinflammatory and in external use to wash ulcers (33). Cortex and leaves: astringent (2). Patagonula americana L. (Boraginaceae). Buenos Aires, La Plata, FCAF, 1969, Hualde s.n. (LPAG 4461); XI-2005, Monti 28 (LPAG). Misiones, Parque Nac. Iguazú, XI-1972, Dimitri s.n. (LPAG 4451). Cortex and leaves: vulnerary, cicatrizant, antisyphilitic (2). Picrasma crenata (Vell.) Engl. (Simaroubaceae). Misiones, Pozo Azul, 28-III-2005, Martínez 56
(LPAG). Wood: antimalarial, antisyphilitic and insecticide properties. It has action on central nervous system (2, 21), and to control stomachic parasites (5). Pilocarpus pennatifolius Lem. (Rutaceae). Buenos Aires, La Plata, FCAF, 12-VI-2006, Arambarri 258 (LPAG). Chaco, 11-VII-1943, Schultz 3948 (LP). Leaves: sudorific; expectorant, antirheumatic, febrifuge (8, 26). Leaves: diaphoretic, diuretic, sialagogue (2, 34). Inflorescence: capillary tonic, to cure insolation and dandruff (1). Cortex and leaves: contains the drug «folia jaborandi» with diaphoretic, expectorant, antirheumatic, febrifuge, excitant, and contraception properties (3). Psychotria carthagenensis Jacq. (Rubiaceae). Misiones, Eldorado, 21-III-2003, Martínez 52 (LPAG); Parque Nac. Iguazú, 20-XI-1969, Volkart 148 (LPAG); 8-XI-1970, Dimitri s.n. (LPAG 4805). Fruits: psychotropic and toxic (2, 35). Leaves: used by ants of genus Atta (17). Quillaja brasiliensis (A. St.-Hil. & Tul.) Mart. (Rosaceae). Misiones, Candelaria-Loreto, 19-II1951, Montes 12343 (LP); Candelaria, Santa Ana, 10-I-1946, Montes 1746 (LP). Cortex: astringent (2); used as vegetable soap to wash hair and clothes (36). Rauvolfia sellowii Müll. Arg. (Apocynaceae). Misiones, Eldorado, 29-III-2005, Martínez 60 (LPAG). Cortex: hypotensor (2, 37). Cortex: with bitter taste and antimalarial properties (3). Rheedia brasiliensis (Mart.) Planch. & Triana (Clusiaceae). Misiones, Parque Nac. Iguazú, 17XI-1969, Hualde 16 (LPAG); Pozo Azul, 28-III2005, Martínez 62 (LPAG). Cortex: medicinal (2). Fuits: edible. Seeds: to cure mange (5). Ruprechtia salicifolia (Cham. & Schltdl.) C.A. Mey. (Polygonaceae). Corrientes, Perugorría, 16-XII1948, Cabrera 10569 (LP). Misiones, Eldorado, 22-III-2005, Martínez 40 (LPAG). Cortex: contains a bitter tonic (2). Schefflera morototoni (Aubl.) Decne. & Planch. (Araliaceae). Buenos Aires, La Plata, calle 2 e/51 y 53, 15-XI-2005, Martínez 97 (LPAG). Misiones, Eldorado, 24-III-2005, Martínez 69 (LPAG). To treat respiratory affections (5). Schinus weinmanniifolia Engl. (Anacardiaceae). Corrientes, Santo Tomé, 14-II-1960, Pedersen 5436 (LP). Misiones, San José de Pindapoy, 9-I-1942, Bridarolli 2553 (LP). Vegetative parts: contraception (2); to treat asthmatic, pruriginous symptoms (1).
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 Sebastiania brasiliensis Spreng. (Euphorbiaceae). Buenos Aires. La Plata, FCAF, 19-IV-2004, Bayón 639 (LPAG). Misiones. Eldorado, 22-III-2005, Martínez 34 (LPAG). Latex: is used externally to cure warts and to calm tooth-ache from decays (36, 38, 39). The latex is also used to cure «huras» (caused by Dermatobia hominis) (5, 40). Cortex: is externally used as antiseptic (13). Sorocea bonplandii (Baill.) W.C. Burger, Lanj. & Wess. (Moraceae). Misiones, Parque Nac. Iguazú, 21-IX-1970, Dimitri & Amorín s.n. (LPAG); Pozo Azul, 28-III-2005, Martínez 63 (LPAG). Latex: toxic (2). Leaves and fruits: forage for animals (3). Styrax leprosus Hook. & Arn. (Styracaceae). Misiones. Parque Nac. Iguazú, 25-IX-1970, Gorgues & Brizuela s.n. (LPAG 4234); 11-IX1971, Dimitri 11259 (LPAG). Cortex: produces an aromatic resin to elaborate pomade used in cutaneous treatments (5). This species belongs the same genus that some Asiatic species from their cortex is extracted a whitish resin giving an aromatic essence known as «benjui» used in medicine and perfumery (20). Tabebuia heptaphylla (Vell.) Toledo (Bignoniaceae). Buenos Aires, La Plata, FCAF, 15-X-1992, Volkart s.n. (LPAG 2941). Misiones, Eldorado, 23-III-2003, Martínez 41 (LPAG). Wood: antirheumatic (1). Flowers: have properties against cough. Wood and cortex: to treat injure of the skin (3). Cortex and leaves: astringent, anti-blennorrhoeic, antiseptic. Wood: diuretic and abortive (41). Tabernaemontana catharinensis A. DC. (Apocynaceae). Misiones, Eldorado, 21-III-2005, Martínez 51 (LPAG); Puerto Iguazú, 18-IX-1970, Dimitri & Amorín 89 (LPAG). Latex: to treat tooth decay (1). Antidote for bites of snakes (2). Latex: to cure cutaneous affections (3). Trixis divaricata (Kunth) Spreng. subsp. divaricata (Asteraceae). Misiones, Loreto, 19-VIII-1954, Gròner 1444 (LP). Flowers: anti-inflammatory and antimenorrhagic properties (2, 42). Vitex megapotamica (Spreng.) Moldenke (Verbenaceae). Buenos Aires, La Plata, III-1984, Volkart s.n. (LPAG); FCAF, III-2006, Arambarri 257 (LPAG). Fruits: edible, diuretic, emmenagogic (2). Cortex: antisyphilitic (2) and antiaphrodisiac (3). References = 1: Martínez Crovetto, 1981; 2: Toursarkissian, 1980; 3: López Villalba et al., 2002; 4: Eibl et al., 2002; 5: Sawchuk & Galeano, 1981; 6: Martínez Crovetto, 1964; 7: Sorarú & Bandoni, 1978; 8: Ratera & Ratera, 1980; 9:
58
Mandrile, 2005; 10: Dimitri, 1978; 11: Rotman, 1987; 12: Chifa & Ricciardi, 2001; 13: Alonso & Desmarchelier, 2005; 14: Filipov, 1994; 15: Martínez Crovetto, 1965; 16: Marinho et al., 2003; 17: Bernardi, 1985; 18: Marzocca, 1997; 19: Blariza et al., 1998; 20: Leonardis, 1975; 21: Rondina et al., 2003; 22: Amat & Vajía, 1991; 23: Lorca et al., 1995; 24: Edwin & Reitz, 1967; 25: Spegazzini, 1999; 26: Mateu Amengual, 1974; 27: Domínguez, 1928; 28: Barboza et al., 2006; 29: Rodríguez & O’Donell, 1943; 30: Martins et al., 2003; 31: Perea et al., 2007; 32: Cortadi et al., 1996; 33: González et al., 1939; 34: Spegazzini et al., 2002; 35: Arias & Parrado, 1996; 36: Biloni, 1990; 37: Xifreda, 1981; 38: Lahitte & Hurrell, 1994; 39: Lahitte & Hurrell, 1999; 40: Amat & Yajía, 1998; 41: Leonardi et al., 2002; 42: Katinas, 1996.
Acknowledgements We appreciate the fine technical assistance of Santiago M. Martínez in collecting material. The curators of herbaria (ERA, LP, LPAG, SI) for loaning the specimens, María A. Migoya for inking our pencil original illustrations, and anonymous reviewers for suggestions are acknowledged. Support was provided by the Comisión de Incentivos a los docentes-investigadores, Decreto 2427/93 (Secretaría de Política Universitaria, Ministerio de Educación de la Nación Argentina).
Bibliography ALONSO, J & C. DESMARCHELIER. 2005. Plantas medicinales autóctonas de la Argentina. Bases científicas para su aplicación en atención primaria de la salud. Ed. L.O.L.A., Buenos Aires. AMAT, A. G. & M. E. VAJÍA. 1991. Plantas medicinales y etnofarmacología en la provincia de Misiones (Argentina). Acta Farm. Bonaerense 10: 153-159. AMAT, A. G. & M. E. YAJÍA. 1998. Plantas vasculares utilizadas en la fitoterapia de la provincia de Misiones (Argentina). Farmacobotánica y farmacognosia en Argentina. Edic. Cient. Americanas. ARAMBARRI, A. M., S. E. FREIRE, M. N. COLARES, N. D. BAYÓN, M. C. NOVOA, C. MONTI & S. A. STENGLEIN. 2006. Leaf anatomy of medicinal shrubs and trees from Gallery forests of the paranaense province (Argentina). Part 1. Bol. Soc. Argent. Bot. 41: 233-268. ARIAS, M. E. & M. F. PARRADO. 1996. Estudio morfoanatómico de hoja y fruto de Psychotria carthagenensis Jacq. (Rubiaceae). Parodiana 9: 19-24. BARBOZA, G. E., J. J. CANTERO, C. O. NÚÑEZ & L. ARIZA ESPINAR (eds). 2006. Flora medicinal de la provincia de Córdoba (Argentina). Museo Botánico de Córdoba. Gráficamente ediciones, Córdoba. Argentina. BERNARDI, L. 1985. Contribución a la dendrología paraguaya. II. Boissiera 37: 1-123. Ediciones Conservatoire et jardin botaniques de la Ville de Genéve. BILONI, J. S. 1990. Árboles autóctonos argentinos.
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees Tipográfica Editora Argentina. Buenos Aires. Argentina. BLARIZA, J. M., A. M. DE JENSEN, E. BAÑA Y SASSONE, D. FEKETE & M. LÓPEZ. 1998. Investigación experimental preliminar en especies vegetales para aislar sustancias reductoras del colesterol y demás lípidos sanguíneos. VI Simposio Argentino de Farmacobotánica. 3-7 de Agosto, Posadas, Misiones. Pág. 71. BURKART, A. 1979. Sapotaceae. En: A. BURKART (ed.), Flora ilustrada de Entre Ríos. Vol. 6: 43-47. Colección Científica INTA, Buenos Aires. BURKART, A. 1987. Leguminosae. En: A. BURKART (ed.), Flora ilustrada de Entre Ríos. Vol. 6: 465-466, 693-695. Colección Científica INTA, Buenos Aires. CABRERA, A. L. & A. WILLINK. 1973. Biogeografía de América Latina. Programa regional de desarrollo científico y tecnológico. OEA, Washington, D.C. CHIFA, C. & A. I. A. RICCIARDI. 2001. Plantas de uso en medicina vernácula del centro del Chaco Argentino. Miscelánea 117: 1-33. Fundación Miguel Lillo, Tucumán. CORTADI, A., O. DI SAPIO & M. GATTUSO. 1996. Caracteres anatómicos de tres especies medicinales de la familia Myrtaceae. Acta Farm. Bonaerense 15: 109-123. CRISTIANI, L. Q. & J. L. AMORÍN. 1972. Estudio preliminar del Sarandí blanco Phyllanthus sellowianus Mueller Arg. (Euphorbiaceae). Farmacobotánica – publicación técnica Nº6. Págs. 11. DIMITRI, M. J. 1978. Enciclopedia argentina de agricultura y jardinería. 3ra. edición. Acme S.A.C.I., Buenos Aires 1: 1-644. DOMÍNGUEZ, J. A. 1928. Contribución a la materia médica argentina. Trabajo del Instituto de Botánica y Farmacia de la Facultad de Ciencias Médicas, Buenos Aires. Pág. 44. EDWIN, G & P. R. REITZ. 1967. Aquifoliaceae. En: P. RAULINO REITZ (ed.). Flora ilustrada Catarinense. Parte 1. Fasc. AQUI, Itajaí, Santa Catarina, Brasil. EIBL, B., A. BOHREN, R. MENDEZ, G. SOSA & M. DI STASI. 2002. Especies forestales nativas de la selva paranaense. Treinta fichas de divulgación y anexos. PECOM Forestal S.A./Facultad de Ciencias Forestales, Universidad Nacional de Misiones (FCF-UNaM). FILIPOV, A. 1994. Medicinal plants of the Pilaga of central Chaco. Journal of Ethnopharmacology 44: 181-193. FREIRE, S. E., L. KATINAS & G. SANCHO. 2002. Gochnatia (Asteraceae-Mutisieae) and the Gochnatia complex: taxonomic implications from morphology. Annals of Missouri Botanical Garden 89: 524-550.
Científicas (CIC). Buenos Aires. LAHITTE, H. B. & J. A. HURRELL. 1999. Árboles rioplatenses. Ed. L.O.L.A., Buenos Aires. LEONARDI, D., O. DI SAPIO, M. GATTUSO & S. GATTUSO. 2002. Caracteres morfoanatómicos de la corteza y hojas de Tabebuia impetiginosa y T. heptaphylla (Bignoniaceae). Bol. Soc. Argent. Bot. 37: 51-61. LEONARDIS, R. F. J. 1975. Libro del árbol. Esencias forestales indígenas de la Argentina de aplicación industrial. T. 2, 2da. Edición. Celulosa Argentina S.A. LÓPEZ VILLALBA, J. A., E. L. LITTLE, G. F. RITZ, J. S. ROMBOLD & W. J. HAHN. 2002. Árboles comunes del Paraguay. Cuerpo de Paz, Colección e Intercambio de Información, 2da. Edición. Editorial Gráfica Mercurio S.A., Paraguay. LORCA, G. G., A. G. AMAT & C. GONZÁLEZ. 1995. Análisis comparativo de caracteres diagnósticos para la identificación de tres especies argentinas de Myrtaceae empleadas en la medicina popular. Acta Farm. Bonaerense 14: 81-86. MANDRILE, E. L. 2005. Plantas medicinales que se dispensan en Argentina. Tomo 1. (2da edición). Colegio de Farmacéuticos de la Provincia de Buenos Aires. MARINHO, C. R., M. C. VENTRELLA & R. M. S. A. MEIRA. 2003. Anatomía da lâmina foliar de Cordia curassavica (Jacq.) Roem. & Schult. (Boraginaceae). Resumo In: 54º Congreso Nacional de Botanica, Belém, Brasil. MARTÍNEZ CROVETTO, R. 1963. Esquema fitogeográfico de la provincia de Misiones (República Argentina). Bonplandia 1: 171-223. MARTÍNEZ CROVETTO, R. 1964. Estudios etnobotánicos. I. Nombres de plantas y su utilidad, según los indios Tobas del este del Chaco. Bonplandia 1: 279-333. MARTÍNEZ CROVETTO, R. 1965. Estudios etnobotánicos. II. Nombres de plantas y su utilidad, según los indios Vilelas del Chaco. Bonplandia 2: 1-23. MARTÍNEZ CROVETTO, R. 1981. Las plantas utilizadas en medicina popular en el noroeste de Corrientes. Fundación Miguel Lillo. Miscelánea 69: 69-70. MARTINS, A. G., S. S. GUTERRES & G. GONZÁLEZ ORTEGA. 2003. Anti-ulcer activity of spray-died powders prepared from leaf extracts of Maytenus ilicifolia Martius ex Reiss. Acta Farm. Bonaerense 22: 39-44. MARZOCCA, A. 1997. Vademécum de malezas medicinales de la argentina, indígenas y exóticas. Orientación Gráfica Editora SRL.
GONZÁLEZ, M., A. LOMBARDO & A. J. VALLARINO. 1939. Plantas de la medicina vulgar de Uruguay. Talleres Gráficos S.A. Cerrito 580, Uruguay.
MASTROBERTI, A. A. & J. E. A. MARIATH. 2003. Leaf anatomy of Araucaria angustifolia (Bertol.) Kuntze (Araucariaceae). Rev. Bras. Bot. 26: 343-353.
HIERONYMUS, J. 1882. Plantae diaphoricas florae argentinae. Bol. Acad. Cienc. Córdoba 4: 199-598.
MATEU AMENGUAL, B. M. 1974. Catálogo bibliográfico fitoquímico Argentino III. Fundación Miguel Lillo, Miscelánea 53, Tucumán.
KATINAS, L. 1996. Revisión de las especies sudamericanas del género Trixis (Asteraceae, Mutisieae). Darwiniana 34: 27-108. LAHITTE, H. B. & J. A. HURRELL. 1994. Los árboles de la Isla Martín García. Comisión de Investigaciones
PEREA, M., G. PEDRAZA & J. LUCEROS. 2007. Relevamiento de flora arbórea autóctona en la provincia de Catamarca. Consejo Federal de Inversiones, Ciudad autónoma de Buenos Aires y Gobierno de la provincia de Catamarca.
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Bol. Soc. Argent. Bot. 43 (1-2) 2008 RATERA, E. & M. RATERA. 1980. Plantas de la flora argentina empleadas en medicina popular. Ed. Hemisferio Sur, Buenos Aires. RODRÍGUEZ, J. M. & C. A. O’DONELL. 1943. Plantas medicinales del noreste Argentino. I. Instituto Miguel Lillo, Rev. Farm. 85: 53-65. RONDINA, R. V. D., A. L. BANDONI & J. D. COUSSIO (eds.). 2003. Plantas silvestres argentinas con reconocidas propiedades medicinales o tóxicas. Base de Datos, CYTED-OEA. ROTH, I. 1984. Stratification of tropical forests as seen in leaf structure. Junk Publ., The Hague, Boston Lancaster. ROTMAN, A. 1978. Moraceae. En: A. BURKART (ed.), Flora ilustrada de Entre Ríos. Vol. 6: 27-28. Colección Científica INTA, Buenos Aires. ROTMAN, A. 1987. Cecropiaceae. En: A. BURKART (ed.), Flora ilustrada de Entre Ríos. Vol. 6: 31.Colección Científica INTA, Buenos Aires. ROTMAN, A. 1995. Las especies argentinas del género Eugenia (Myrtaceae). Bol. Soc. Argent. Bot. 31: 69-93. SAWCHUK, B. & H. GALEANO. 1981. Descripción de árboles medicinales de la Selva Misionera. En: Manual para la agricultura en la provincia de Misiones, República Argentina. Parte 6: 1-168. (B. Sawchuk, E. J. Domínguez & A. A. Piekun), Programa Argentino-Alemán para el
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fomento del desarrollo agrícola de la provincia de Misiones. SORARÚ, S. & A. BANDONI. 1978. Plantas de la medicina popular argentina. Ed. Albatros, Buenos Aires. SPEGAZZINI, E. D. 1999. Análisis foliar por micrografía analítica cuali-cuantitativa, de los adulterantes cogenéricos argentinos de Ilex paraguariensis St. Hil. var. paraguariensis –Aquifoliaceae- nv. «yerba mate». Tesis Doctoral, Facultad de Ciencias Exactas, Universidad Nacional de La Plata. SPEGAZZINI, E. D., M. T. CASTRO, J. ROSSI, S. M. CARPANO & M. T. NÁJERA. 2002. Micrografía foliar cuali-cuantitativa de las especies de Pilocarpus (Rutaceae): P. jaborandi Holmes, P. microphyllus Stape ex Wardleworth y P. pennatifolius Lem. Dominguezia 18: 5-16. TOURSARKISSIAN, A. 1980. Plantas medicinales de la Argentina. Ed. Hemisferio Sur, Buenos Aires. XIFREDA C. C. 1981. El género Rauvolfia (Apocynaceae) en la Argentina. Parodiana 1: 119-137. ZARDINI, E. M. 1984. Etnobotánica de compuestas argentinas con especial referencia a su uso farmacológico. Acta Farm. Bonaerense 3: 77-99. Recibido el 28 de Agosto de 2007, aceptado el 12 de Mayo de 2008.