a m arambarri, s e freire, m n colares, n d bayón, m c novoa, c monti & s a stenglein leaf...
DESCRIPTION
ÂTRANSCRIPT
31
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
Leaf anatomy of medicinal shrubs and trees from Misionesforest 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 fromParanaense province. Forty five species of shrubs and trees belonging to 29 families inhabiting Misionesforest 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 differentialtraits, 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 peltatetrichomes in Tabebuia heptaphylla); midvein transection outlines (e.g. midvein convex and keel-shapedon the adaxial side in Schinus weinmanniifolia); presence and types of crystals (e.g. crystal sand inCordia ecalyculata, raphides in Psychotria carthagenensis). This paper also gives an ecologicalinterpretation 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, abundantsclerenchyma, multilayered epidermis, mesophyll formed exclusively by palisade parenchyma, multilayeredhypodermis, 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 treesfrom the Paranaense biogeographic province (Part 1: Gallery forests and Part 2: Misiones forest) thatpermit identified species using anatomy leaf characteristics.
Key words: anatomy, Argentina, Misiones forest, leaves, medicinal plants, Paranaense biogeographicprovince.
Resumen: Anatomía foliar de arbustos y árboles medicinales de la Selva Misionera de la provinciaParanaense (Argentina). Parte 2. El presente trabajo corresponde a la segunda entrega del estudio dearbustos y árboles medicinales de la provincia biogeografica Paranaense. En esta parte, se analizaron45 especies contenidas en 29 familias que habitan la Selva Misionera y han sido citadas con propiedadesmedicinales. El trabajo se acompaña con ilustraciones y se brinda una conclusión de los caracteresdiagnósticos, 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. escamosospeltados en Tabebuia heptaphylla); el contorno de la vena media (e.g. convexa y aquillada en la caraadaxial en Schinus weinmanniifolia); la presencia y tipo de cristales (e.g. arena cristalina en Cordiaecalyculata, rafidios en Psychotria carthagenensis). También se provee una interpretación ecológica delas especies estudiadas cuyas hojas presentaron predominantemente una combinación de caracteresmesomórficos (e.g. hojas hipostomáticas, mesofilo dorsiventral) y xeromórficos (e.g. cutícula gruesa,esclerénquima abundante, epidermis pluriseriada, mesofilo indiferenciado en empalizada, hipodermispluriseriada, esclereidas idioblásticas). Solamente dos especies, Ilex paraguariensis y Manihot grahamii,presentaron combinados caracteres mesomórficos (entre otros, hojas hipostomáticas y mesofilodorsiventral) e higromórficos (epidermis glabra). Finalmente, se provee una clave para la diferenciaciónde 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 caracteresanatómicos de la hoja.
Palabras clave: anatomía, Argentina, hoja, plantas medicinales, provincia biogeográfica Paranaense,Selva Misionera.
1Área de Botánica, Facultad de Ciencias Agrarias y Forestales, Calle 60 y 119, C.C. 31, 1900 La Plata, UniversidadNacional de La Plata, Argentina.2Instituto de Botánica Darwinion, Labardén 200, C.C.22, B1642 HYD San Isidro, Buenos Aires, Argentina.3Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina (CONICET)*Corresponding author. Tel: +54-221-423-6758 int. 461; Fax: +54-221-425-2346. E-mail: [email protected];[email protected]
ISSN 0373-580 XBol. Soc. Argent. Bot. 43 (1-2): 31 - 60. 2008
Bol. Soc. Argent. Bot. 43 (1-2) 2008
32
This paper is the second part of the study ofmedicinal shrubs and trees from Paranaense province.The first part including the medicinal shrubs and treesof the Gallery forests, was published by Arambarri etal. (2006). Misiones forest is situated in thenortheastern Argentina, extending from Misiones tonortheastern Corrientes political provinces. From abiogeographic point of view, Misiones and Galleryforests belong to Paranaense province within theAmazonian dominion of Neotropical region (Cabrera& Willink, 1973). In this region, more than 100 nativeshrubs and trees have been cited as used for food,dressing, religious and magic ceremonies, and in folkmedicine 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 theknowledge of native shrubs and trees with medicinalproperties, we studied the leaf anatomy of 45 speciesbelonging to 42 genera which are contained into 29families from Misiones forest.
Introduction
* For methods refer to Part 1: Arambarri et al. (2006).
On the basis of leaf anatomical features of 62species from Gallery forests and 45 species from theMisiones forest the following key was generated:
Results
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.
33
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.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
34
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.
35
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
Bol. Soc. Argent. Bot. 43 (1-2) 2008
36
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
37
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
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
Bol. Soc. Argent. Bot. 43 (1-2) 2008
38
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.
39
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
Bol. Soc. Argent. Bot. 43 (1-2) 2008
40
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.
41
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
Bol. Soc. Argent. Bot. 43 (1-2) 2008
42
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
43
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
dorsiventral with palisade parenchyma 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
with palisade parenchyma2-layered.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
44
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
45
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
Bol. Soc. Argent. Bot. 43 (1-2) 2008
46
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
47
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
Fig. 1. Leaf characteristics in transection and surface view. A-D: Araucaria angustifolia. A, leaf blade showing isobilateralmesophyll; compartmented cells and resiniferous ducts among vascular bundles, and idioblastic sclereids; B, thickness ofepidermal 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 cellsnearly 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 andsome disposed adaxially; J, petiole; K, adaxial epidermis, one cystolith in transection; L, adaxial epidermis showing thepeduncle of one cystolith in surface view. M-P: Schinus weinmanniifolia. M, midvein keel-shaped adaxially; N, petioletransection; 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.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
48
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, abaxialepidermis crystalliferous, thin anticlinal cell walls. E-H: Aspidosperma polyneuron. E, midvein, collenchyma absent and abicollateral vascular bundle; F, petiole, numerous clusters of idioblastic sclereids; G, adaxial epidermis showing cuticularornamentation; H, abaxial epidermis with an anomocytic stoma. I-L: Hennecartia omphalodes. I, midvein with adaxialchlorenchyma 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 ringsof 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 = 50μm; F, J, M, N, O = 1 mm.
49
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 andabaxial; mesophyll indifferentiated (palisade parenchyma); B, portion of the lobate petiole and vascular tissue; C, adaxialepidermis in surface view, showing a conical secretory structure; D, adaxial epidermis multilayered with a secretory structurein 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 astoma, in surface view. I-M: Quillaja brasiliensis. I, midvein, one collateral vascular bundles with xylematic and phloematiccups of sclerenchyma; J, petiole pubescent; K, adaxial epidermis and hypodermis in transection, showing the thickness of outerpericlinal 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, subcircularoutline, a ring of vascular tissue and medullary bundles, and numerous secretory ducts; P, petiolule; Q, abaxial epidermis withcuticular 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.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
50
Fig. 4. Leaf characteristics in transection and surface view. A-D: Rheedia brasiliensis. A, midvein showing chlorenchymaadaxial and the vascular tissue surrounded by a ring of sclerenchyma; B, petiole; C, adaxial epidermis with thick anticlinal cellwalls, 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 wallsthickened; I, epidermis transection showing enlarged epidermal cells in anticlinal direction, and palisade parenchyma with anidioblastic cell containing a druse. J-N: Jacaratia spinosa. J, midvein concave and without collenchyma, adaxially; K, portionof petiole showing the cortex and then a ring of sclerenchyma surrounding the vascular tissue; L, adaxial epidermis in surfaceview showing a secretory structure; M, adaxial epidermis exhibiting a secretory structure in transection; N, abaxial epidermisconspicuously papillose, in transection. O-R: Coutarea hexandra f. pubescens. O, midvein flattened adaxially; P, petioleconspicuously 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.
51
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
Conclusions
Character value
Epidermal features and leaf anatomy have highdiagnostic value at specific level and/or at familylevel: (1) crystaliferous epidermis characterized twospecies Maytenus ilicifolia and Trixis divaricatasubsp. divaricata; (2) cystolith-like structure intrichomes are present in Aloysia virgata var. virgataand Cordia curassavica. This structure in trichomesconfirms the relationships among the familiesVerbenaceae and Boraginaceae to which the speciesare belong respectively; (3) the ciclocytic stomatatype is present in Araucaria angustifolia andPilocarpus pennatifolius; (4) Cordia ecalyculatahas cystoliths in its uniseriate epidermis; (5)Schefflera morototoni can be distinguished by itsmultilayered and colenquimatous hypodermisdistributed on both surfaces, and the presence oftwo cicles of vascular bundles in the petiole; (6)Cecropia pachystachya is characterized by havingthe leaf blade mesophyll formed by palisadeparenchyma exclusively, and lobed petiole outline;(7) scale peltate trichomes are diagnostic epidermaltraits in Styrax leprosus (Styracaceae) and Tabebuiaheptaphylla (Bignoniaceae), and located in epidermaldepressions in Esenbeckia febrifuga (Rutaceae); (8)laticiferous tubes recognized by their coloured latexare present in Sapotaceae and Apocynaceae; (9)bicollateral vascular bundles are diagnostic leaf traitsof the families: Apocynaceae, Myrtaceae andSolanaceae; 10) secretory ducts into the phloematicparenchyma are conspicuous and a definitemicrocharacter in family Anacardiaceae; 11) epidermalheteromorphic cells over the secretory cavities are adiagnostic trait in Myrtaceae; 12) paracytic stomataand oil cells characterized the family Lauraceae; 13)very small druses uniquely were found inVerbenaceae, and 14) large secretory cavities arediagnostic features relating the families Bombacaceaeand Malvaceae.
Ecological implications
The species studied show predominantly acombination of mesomorphic and xeromorphic leafstructures, e.g. Aspidosperma polyneuron, Ceibaspeciosa, Gleditsia amorphoides var. amorphoides,Hexaclamys edulis. As mesomorphic characters canbe mentioned: (1) the majority of studied species havehypostomatic leaves, and the stomata lie at epidermislevel; (2) thirty eight of the total forty five analyzedtaxa have dorsiventral leaves; (3) most of speciesexhibit a well developed spongy parenchyma rich inintercellular spaces. Within xeromorphic features maybe mentioned: (1) anticlinal cell walls straight tocurved; (2) anticlinal and outer periclinal epidermalcell walls thick; (3) cuticle thick and/or ornamented;(4) epidermis multilayered or when 1-layered formedby large cells; (5) hypodermis on one or both leafsurfaces. However, twenty six species have also somehygromorphic features such as epidermis glabrous,anticlinal cell walls undulate, leaves amphistomaticor/and stomata elevated, e.g. Aloysia virgata var.virgata, Chrysophyllum spp., Cordia spp., Guaduachacoensis, Jacaratia spinosa, Myrcianthespungens, Sebastiania brasiliensis,Tabernaemontana catharinensis, Trixis divaricatasubsp. divaricata. Only two species, Ilexparaguariensis and Manihot grahamii havemesomorphic and hygromorphic traits. Most ofxeromorphic features / sun leaf type, such as a thickand/or ornamented cuticle, e.g. Pilocarpuspennatifolius (Rutaceae); thick cuticle and abundantsclerenchyma e.g. Gleditsia amorphoides var.amorphoides (Fabaceae); multilayered epidermis,e.g. Cecropia pachystachya (Cecropiaceae);presence of hypodermis 2-layered e.g. Hexaclamysedulis (Myrtaceae); collenchymatous andmultilayered hypodermis e.g. Schefflera morototoni(Araliaceae); mesophyll with palisade parenchymamultilayered e.g. Patagonula americana(Boraginaceae); mesophyll formed exclusively bypalisade parenchyma e.g. Cecropia pachystachya(Cecropiaceae); parenchyma with idioblasticsclereids e.g. Aspidosperma polyneuron(Apocynaceae), seems to be an adaptation of thebig trees to exposed situations with high level ofsolar irradiance.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
52
Tabl
e 1.
Lis
t of s
tudi
ed sp
ecie
s and
eco
logi
cal l
eaf c
hara
cter
istic
s ela
bora
ted
on th
e ba
sis o
f crit
eria
of t
he x
erom
orph
y, m
esom
orph
y, h
ygro
mor
phy,
sun
leaf
type
, med
ium
leaf
type
and
shad
e le
af ty
pe o
f Rot
h (1
984)
. Abb
revi
atio
ns: A
cellw
s: a
ntic
linal
cel
l wal
ls; a
bx: a
baxi
al; a
dx: a
daxi
al; C
: cut
icle
; E: e
pide
rmis
; Ecw
: epi
cutic
ular
wax
es; H
:hy
pode
rmis
; L: l
eave
s; M
: mes
ophy
ll; P
cellw
s: p
eric
linal
cel
l wal
ls; P
p: p
alis
ade
pare
nchy
ma;
Scl
: scl
eren
chym
a; S
t: st
omat
a.
Tax
aX
erom
orph
ic /
sun
leaf
type
Mes
omor
phic
/ m
ediu
m le
af ty
peH
igro
mor
phic
/ sh
ade
leaf
type
Alo
ysia
vir
gata
H
1-la
yere
d (a
dx a
ndA
cellw
s st
raig
ht to
cur
ved;
St e
leva
ted;
Scl
redu
ced
/ abs
ent
var.
vir g
ata
disc
ontin
uous
) ; E
pub
esce
ntL
hypo
stom
atic
; M
dor
sive
ntra
l to
indi
ffer
entia
ted
Ara
ucar
ia a
ngus
tifol
iaA
cellw
s th
ick
and
stra
ight
;L
amph
isto
mat
ic; M
isob
ilate
ral
Ecw
con
spic
uous
; E a
nd H
scl
erop
hyllo
us;
E gl
abro
usH
mul
tilay
ered
(bot
h fa
ces)
; St s
unke
n;M
with
idio
blas
tic s
cler
eids
Asp
idos
perm
a po
lyne
uron
Ecw
con
spic
uous
; C th
ick;
Ace
llws
curv
ed; L
hyp
osto
mat
ic;
Pp 1
-2-la
yere
d; S
cl a
bund
ant;
M d
orsi
vent
ral;
E su
bgla
brou
sId
iobl
astic
scl
erei
ds p
rese
ntA
stro
nium
bal
ansa
eC
stri
ate
(abx
face
)A
cellw
s cu
rved
; L h
ypos
tom
atic
;M
dor
sive
ntra
l; Pp
1-2
-laye
red
(sho
rt ce
lls);
Scl +
/- ab
unda
nt; E
sub
glab
rous
Cec
ropi
a pa
chys
tach
yaE
mul
tilay
ered
with
sec
reto
ry c
ells
;A
cellw
s st
raig
ht to
slig
htly
Scl r
educ
ed /
abse
ntM
indi
ffer
entia
ted
(tot
ally
Pp)
;cu
rved
; L h
ypos
tom
atic
E pu
besc
ent
Ced
rela
fiss
ilis
E pu
besc
ent;
Pp (l
arge
cel
ls)
Ace
llws
curv
ed (a
dx fa
ce);
Ace
llws
undu
late
(abx
face
)L
hypo
stom
atic
; M
dor
sive
ntra
l;Pp
1-la
yere
d; S
cl +
/- ab
unda
ntC
eiba
spe
cios
aE
2-la
yere
d; C
stri
ate
(abx
face
);A
cellw
s cu
rved
; L h
ypos
tom
atic
;Sc
l abu
ndan
t; Pp
1-2
-laye
red
M d
orsi
vent
ral;
E su
bgla
brou
sC
hrys
ophy
llum
gon
ocar
pum
Ecw
con
spic
uous
; C th
ick;
L hy
post
omat
ic; M
dor
sive
ntra
l;A
cellw
s un
dula
te;
Scl a
bund
ant
Pp 1
-2-la
yere
d (s
hort
cells
)E
glab
rous
Chr
ysop
hyllu
m m
argi
natu
mC
stri
ate
(abx
face
)L
hypo
stom
atic
; M d
orsi
vent
ral;
Ace
llws
undu
late
;Pp
1-la
yere
d (s
hort
cells
);E
glab
rous
Scl +
/- ab
unda
ntC
ordi
a cu
rass
avic
aPp
aren
chym
a 1-
2-la
yere
d;A
cellw
s cu
rved
; L h
ypos
tom
atic
;A
cellw
s un
dula
te (a
bx);
E pu
besc
ent
M d
orsi
vent
ral
Scl r
educ
ed /
abse
ntC
ordi
a ec
alyc
ulat
aPp
(lar
ge c
ells
)L
hypo
stom
atic
; M d
orsi
vent
ral;
Ace
llws
undu
late
;
Pp 1
-laye
red;
Scl
+/-
abun
dant
E gl
abro
us
Cou
tare
a he
xand
ra f
. pub
esce
nsC
stri
ate;
E p
ubes
cent
L hy
post
omat
ic; M
dor
sive
ntra
l;A
cellw
s un
dula
te;
Pp (l
arge
cel
ls)
Pp 1
-laye
red
Scl r
educ
ed /
abse
ntE
senb
ecki
a fe
brifu
gaEc
w c
onsp
icuo
us; C
stri
ate;
Ace
llws
curv
ed; L
hyp
osto
mat
ic;
Scl r
educ
ed /
abse
ntE
pube
scen
tM
dor
sive
ntra
l; Pp
1-la
yere
d(s
hort
cells
)
53
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and treesTa
ble
1. C
ont.
Gle
dits
ia a
mor
phoi
des
Ecw
con
spic
uous
; C th
ick;
Ace
llws
curv
ed; L
hyp
osto
mat
ic;
var.
amor
phoi
des
Scl a
bund
ant;
E pu
besc
ent
M d
orsi
vent
ral;
Pp 1
-laye
red
(sho
rt ce
lls)
Goc
hnat
ia p
olym
orph
aA
cellw
s st
raig
ht (a
dx);
C th
ick;
Ace
llws
curv
ed (a
bx);
subs
p. c
eano
thi fo
liaSt
with
sto
mat
al ri
m;
L hy
post
omat
ic; M
dor
sive
ntra
l;Pp
mul
tilay
ered
; E p
ubes
cent
Scl +
/- ab
unda
ntG
uadu
a ch
acoe
nsis
Ace
llws
stra
ight
and
thic
k;E
subg
labr
ous
L am
phis
tom
atic
;Ec
w c
onsp
icuo
us; C
thic
k;M
indi
ffer
entia
ted
form
ed b
ySc
l abu
ndan
tis
odia
met
ric c
ells
Hen
neca
rtia
om
phal
andr
aA
cellw
s th
ick;
Ecw
con
spic
uous
;A
cellw
s cu
rved
; L h
ypos
tom
atic
;C
thic
k; H
1-la
yere
d (a
dx);
M d
orsi
vent
ral 1
-laye
red
(sho
rt ce
lls);
E pu
besc
ent
Scl +
/- ab
unda
ntH
exac
lam
ys e
dulis
Ace
llws
thic
k; E
cw c
onsp
icuo
us;
Ace
llws
curv
ed; L
hyp
osto
mat
ic;
C th
ick;
H 2
-laye
red
(adx
);M
dor
sive
ntra
l; Sc
l +/-
abun
dant
;Pp
2-la
yere
dE
subg
labr
ous
Ilex
par
agua
rien
sis
Ace
llws
curv
ed; L
hyp
osto
mat
ic;
E gl
abro
us M
dor
sive
ntra
l; Pp
1-la
yere
d(s
hort
cells
); Sc
l +/-
abun
dant
Jaca
ratia
spi
nosa
Ace
llws
stra
ight
; E s
ecre
tory
cel
ls;
L hy
post
omat
ic ;
M d
orsi
vent
ral;
Scl r
educ
ed /
abse
nt;
Ecw
con
spic
uous
(abx
); Pp
(lar
ge c
ells
)Pp
1-la
yere
dE
glab
rous
Man
ihot
gra
ham
iiL
hypo
stom
atic
; M d
orsi
vent
ral;
Ace
llws
undu
late
;Pp
1-la
yere
d (s
hort
cells
)Sc
l red
uced
/ ab
sent
; E g
labr
ous
May
tenu
s ili
cifo
liaA
cellw
s st
raig
ht; E
cw c
onsp
icuo
us;
L hy
post
omat
ic ;
M is
obila
tera
l;C
thic
kSc
l +/-
abun
dant
E gl
abro
usM
yrci
anth
es p
unge
nsC
thic
k; H
1-la
yere
d (a
dx);
L hy
post
omat
ic
Ace
llws
undu
late
; M is
obila
tera
l;Sc
l abu
ndan
tE
glab
rous
Myr
ocar
pus
fron
dosu
sEc
w c
onsp
icuo
us; H
1-la
yere
d;A
cellw
s st
raig
ht to
cur
ved;
Pp m
ultil
ayer
edL
hypo
stom
atic
; M d
orsi
vent
ral;
Scl +
/- ab
unda
nt; E
sub
glab
rous
Nec
tand
ra la
nceo
lata
Ace
llws
stra
ight
; Scl
abu
ndan
t;L
hypo
stom
atic
;Pp
2-la
yere
d; E
pub
esce
ntM
dor
sive
ntra
lO
cote
a di
ospy
rifo
liaA
cellw
s st
raig
ht; P
p 1(
-2)-
laye
red;
L hy
post
omat
ic; M
dor
sive
ntra
l;Sc
l abu
ndan
tE
subg
labr
ous
Oco
tea
pube
rula
Ace
llws
stra
igh
(adx
) and
thic
k;A
cellw
s cu
rved
(abx
);Pp
2-la
yere
d; S
cl a
bund
ant;
L hy
post
omat
ic; M
dor
sive
ntra
lE
pube
scen
tPa
rapi
ptad
enia
rig
ida
E pu
besc
ent;
Pp (l
arge
cel
ls)
Ace
llws
curv
ed; L
hyp
osto
mat
ic;
M d
orsi
vent
ral;
Pp 1
-laye
red;
Scl +
/- ab
unda
ntPa
tago
nula
am
eric
ana
Pcel
lws
thic
k;A
cellw
s st
raig
ht to
cur
ved;
Pp m
ultil
ayer
ed (c
ells
not
so
long
)L
hypo
stom
atic
; M d
orsi
vent
ral;
Bol. Soc. Argent. Bot. 43 (1-2) 2008
54
Tabl
e 1.
Con
t.
Scl +
/- ab
unda
nt; E
sub
glab
rous
Picr
asm
a cr
enat
aC
stri
ate
(abx
face
);L
hypo
stom
atic
; M d
orsi
vent
ral ;
Ace
llws
undu
late
;Pp
(lar
ge c
ells
)Pp
1-la
yere
d; S
cl +
/- ab
unda
ntE
glab
rous
Pilo
carp
us p
enna
tifol
ius
Ace
llws
stra
ight
and
thic
k;L
hypo
stom
atic
; M d
orsi
vent
ral;
Ecw
con
spic
uous
; C s
triat
e;Pp
1-la
yere
d; S
cl +
/- ab
unda
ntPp
(lar
ge c
ells
); E
pube
scen
tPs
ycho
tria
car
thag
enen
sis
Ace
llws
stra
ight
; Ecw
con
spic
uous
;L
hypo
stom
atic
; M d
orsi
vent
ral;
E m
ultil
ayer
ed w
ith s
ecre
tory
cel
ls;
Pp 1
-laye
red;
Scl
+/-
abun
dant
;Pp
(lar
ge c
ells
)E
subg
labr
ous
Qui
llaja
bra
silie
nsis
H 1
-laye
red
(adx
and
abx
);A
cellw
s cu
rved
; L h
ypos
tom
atic
;M
isob
ilate
ral t
o in
diff
eren
tiate
d;Sc
l +/-
abun
dant
; E s
ubgl
abro
usPp
mul
tilay
ered
(adx
and
abx
)R
auvo
lfia
sello
wii
Ace
llws
stra
ight
;L
hypo
stom
atic
; M d
orsi
vent
ral;
Scl r
educ
ed /
abse
nt;
Pp (l
arge
cel
ls)
Pp 1
-laye
red;
E gl
abro
usR
heed
ia b
rasi
liens
isA
cellw
s st
raig
ht a
nd th
ick;
L hy
post
omat
ic; M
dor
sive
ntra
l;E
glab
rous
C v
ery
thic
k; S
cl a
bund
ant
Pp 1
-2 -l
ayer
ed (s
hort
cells
)R
upre
chtia
sal
icifo
liaA
cellw
s st
raig
ht; P
p m
ultil
ayer
ed;
L hy
post
omat
ic; M
dor
sive
ntra
l;Sc
l abu
ndan
tE
subg
labr
ous
Sche
ffle
ra m
orot
oton
iA
cellw
s st
raig
ht; H
mul
tilay
ered
L hy
post
omat
ic; M
dor
sive
ntra
l; (b
oth
face
s); S
t sun
ken
with
Scl +
/- ab
unda
ntpe
risto
mat
al ri
m; P
p 2-
laye
red;
E pu
besc
ent
Schi
nus
wei
nman
niifo
liaA
cellw
s st
raig
ht a
nd th
ick;
L hy
post
omat
ic w
ith s
ome
M is
obila
tera
l;C
stri
ate
(abx
face
); E
pube
scen
tst
omat
a lo
cate
d ad
axia
llySc
l red
uced
/ ab
sent
Seba
stia
nia
bras
ilien
sis
C s
triat
e; P
p (la
rge
cells
)M
dor
sive
ntra
l; Pp
1-la
yere
dA
cellw
s un
dula
te; L
am
phis
tom
atic
;Sc
l red
uced
/ ab
sent
; E g
labr
ous
Sb
ldi
ii
lld
dl
55
A. M. Arambarri et al., Leaf anatomy of medicinal shrubs and trees
Appendix 1. List of studied species with specimenvouchers and ethnobotany use. The numbers withinparentheses indicate the references listed at the end.Abbreviation: Jardín Botánico y Arboretum «C.Spegazzini», Facultad de Ciencias Agrarias yForestales, UNLP: FCAF.
Aloysia virgata (Ruiz & Pav.) Juss. var. virgata(Verbenaceae). Misiones, Parque Nac. Iguazú, 4-XI-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, 8-IX-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). SantaFe, General Obligado, 10-I-1937, Ragonese 2413(LP). Leaves: analgesic, febrifuge, in treatment ofinsolation (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-IX-1970, Dimitri & Amorín 1181 (LPAG). Leaves:expectorant, antiasthmatic, bronchitis, sedative,diuretic, cardiotonic (oficial drug monographiedin 5th edition of the Farmacopea NacionalArgentina) (7). Leaves: tonic, pectoral, cardiotonic,diuretic, anticatarrhal, antiasthmatic (2, 5, 8).Leaves: are useful to control cough (1). Trunk juiceis used against cutaneous bores and the twig juicehas properties as antidiarrheic (5). Leaves andcork: expectorant, tonic and anti-inflammatory (9).Fruits: edible (9-11). This species is in 6th editionof the Farmacopea Nacional Argentina and it wasincorporated in the Program for Atención Primariade 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). Cortexand wood: febrifuge, anti-inflammatory,antimalarial (2, 3, 8). Cortex: febrifuge, emetic,astringent, hepatic. Leaves: to treat urinary
Scl r
educ
ed /
abse
nt; E
gla
brou
sSo
roce
a bo
npla
ndii
Ecw
con
spic
uous
;A
cellw
s cu
rved
to u
ndul
ate;
E pu
besc
ent
L hy
post
omat
ic; M
dor
sive
ntra
l;Pp
1-2
–la
yere
d (s
hort
cells
);Sc
l +/-
abun
dant
Styr
ax le
pros
usA
cellw
s st
raig
ht a
nd th
ick;
L hy
post
omat
ic; M
dor
sive
ntra
l;Sc
l red
uced
/ ab
sent
Ecw
con
spic
uous
; Epi
derm
al c
ells
Pp 1
-laye
red
(sho
rt ce
lls)
larg
e; E
pub
esce
ntTa
bebu
ia h
epta
phyl
laEc
w c
onsp
icuo
us; C
stri
ate;
Ace
llws
curv
ed; L
hyp
osto
mat
ic;
E 2-
laye
red;
Pp
2-la
yere
d (la
rge
cells
);M
dor
sive
ntra
l; Sc
l +/-
abun
dant
E pu
besc
ent
Tabe
rnae
mon
tana
cat
hari
nens
isC
stri
ate
Ace
llws
curv
ed; L
hyp
osto
mat
ic to
Scl r
educ
ed /
abse
nt;
amph
isto
mat
ic; M
dor
sive
ntra
l; E
glab
rous
Pp 1
-laye
red
(sho
rt ce
lls)
Trix
is d
ivar
icat
aE
pube
scen
tL
hypo
stom
atic
;A
cellw
s un
dula
te;
subs
p. d
ivar
icat
aM
dor
sive
ntra
l;Sc
l red
uced
/ ab
sent
Pp 1
-laye
red
(sho
rt ce
lls)
Vite
x m
egap
otam
ica
Ecw
con
spic
uous
; Pp
mul
tilay
ered
;A
cellw
s cu
rved
; L h
ypos
tom
atic
;E
pube
scen
tM
dor
sive
ntra
l; Sc
l +/-
abun
dant
Tabl
e 1.
Con
t.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
56
illnesses (13).Ceiba speciosa (A. St.-Hil.) Ravenna (Bombacaceae).
Buenos Aires, La Plata, 17-VIII-2004, Stenglein19 (LPAG); 30-IX-2005, Arambarri 252 (LPAG).Flowers and thorns: analgesic and diuretic (2, 14).Seeds have 15-20% of oil with industrialapplications and edible. Cortex sap: to treat hernia.Thorns: analgesic, antiasthmatic and to treatalcoholism (4).
Chrysophyllum gonocarpum (Mart. & Eichler) Engl.(Sapotaceae). Buenos Aires, La Plata, FCAF, 8-XI-2005, Arambarri 254 (LPAG). Corrientes, SantaTeresa, 22-III-1954, Petersen 2644 (LP). Fruits:eaten directly or as candy. It were used by theJesuits as medicine (3, 15).
Chrysophyllum marginatum (Hook. & Arn.) Radlk.(Sapotaceae). Buenos Aires, La Plata, FCAF, 8-XI-2005, Arambarri 253 (LPAG). Corrientes,Concepción, 4-V-1974, Petersen 1072 (LP). Leavesand 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 leafystems: vermifuge, antispasmodic, digestive,antitoxic, to cure cough (1). Leaves: anti-inflammatory (16).
Cordia ecalyculata Vell. (Boraginaceae). Misiones,Eldorado, 30-III-2003, Martínez 59 (LPAG); ParqueNac. Iguazú, 6-XII-1971, Volkart s.n. (LPAG 4449).Leaves: carditonic, tonic (2). Fruits: with a viscidsubstance which is used as glue. Fruits are alsopalatable for wild animals (3).
Coutarea hexandra (Jacq.) K. Schum. f. pubescens(Pohl) Steyerm. (Rubiaceae). Buenos Aires, LaPlata, Jardín Sistemático de la Facultad deAgronomía, I-1969, Volkart s.n. (LPAG 4195).Misiones, Parque Nac. Iguazú, 9-I-1971, Dimitris.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, substituteof quinina (Peruvian bark) (2); and with aphrodisiacproperties (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
and roots: astringent (18). Cortex: with propertiesto reduce cholesterol (19). Fruits: irritant whichmake sneeze (5, 15); to treat asthma (1). Fruits:produce a gum used in sweets industry, pharmacyand cosmetic (3, 20).
Gochnatia polymorpha (Less.) Cabrera subsp.ceanothifolia (Less.) Cabrera (Asteraceae).Misiones, Candelaria, Loreto, Montes 11154 (LP);Corrientes, 6-II-1972, Santo Tomé, Krapovickaset al. 21091 (LP). Leaves and flowers: pectoral(2).
Guadua chacoensis (Rojas Acosta) Londoño & P.M.Peterson (Poaceae). Corrientes, Ituzaingó, 8-XII-1974, Krapovickas et al. 23851 (SI). Misiones,Montecarlo, I-1944, Porta s.n. (LP 900486); ParqueNac. Iguazú, 14-IX-1971, Dimitri 11246 (LPAG).Rhizomes: urinary antiseptic, laxative, analgesic(1, 21). Water acummulated in internodes is usedto 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, 15-IX-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, 23-III-2005, Martínez 83 (LPAG). Leaves: tonic,stimulant, diuretic, digestive, sudorific (3, 24);leaves macerated to treat dandruff; cephalalgia;anti-inflammatory; diuretic (1, 25). This specieswas included in the 1st edition of the FarmacopeaNacional Argentina (13).
Jacaratia spinosa (Aubl.) A. DC. (Caricaceae).Misiones, Parque Nac. Iguazú, 6-XI-1971, Dimitris.n. (LPAG); Facultad de Ciencias Forestales, 23-III-2005, Martínez 42 (LPAG). Latex: vermifuge andcathartic (2, 26); the plant has properties to controlgastrointestinal worms (5). Fruits: edible (3).
Manihot grahamii Hook. (Euphorbiaceae). BuenosAires, 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,
57
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 inexternal use as antiseptic (2, 6, 7, 29). Leaves:antispasmodic, digestive and antiseptic (8). Thisspecies would has antitumoral properties (21).Extract from leaves would have anti-ulcer activity(30). This species is included in the FarmacopeaBrasileña and it was introduced in the Program forAtención Primaria de la Salud en la provincia deMisiones (13).
Myrcianthes pungens (O. Berg) D. Legrand(Myrtaceae). Buenos Aires, La Plata, JardínBotánico de la Facultad de Agronomía, UNLP, 24-III-1966, Amorín 148 (LPAG); Capital Federal, 7-I-1991, 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 isused as substitute of the true incense (3); and inperfume and tincture industry (4).
Nectandra lanceolata Nees (Lauraceae). BuenosAires, La Plata, FCAF, IV-2004, Monti 24a, 24b(LPAG). Misiones, Puerto Iguazú, 21-X-1970,Amorín & Dimitri s.n. (LPAG 1387). This specieshas 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 toextract thorns or spines (5).
Parapiptadenia rigida (Benth.) Brenan (Fabaceae).Buenos Aires, La Plata, FCAF, 5-V-1966, Amorín352 (LPAG). Misiones, Parque Nac. Iguazú, 18-XI-1969, Hualde 122 (LPAG). Cortex: anti-inflammatory and in external use to wash ulcers(33). Cortex and leaves: astringent (2).
Patagonula americana L. (Boraginaceae). BuenosAires, La Plata, FCAF, 1969, Hualde s.n. (LPAG4461); XI-2005, Monti 28 (LPAG). Misiones,Parque Nac. Iguazú, XI-1972, Dimitri s.n. (LPAG4451). Cortex and leaves: vulnerary, cicatrizant,antisyphilitic (2).
Picrasma crenata (Vell.) Engl. (Simaroubaceae).Misiones, Pozo Azul, 28-III-2005, Martínez 56
(LPAG). Wood: antimalarial, antisyphilitic andinsecticide properties. It has action on centralnervous system (2, 21), and to control stomachicparasites (5).
Pilocarpus pennatifolius Lem. (Rutaceae). BuenosAires, 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 andleaves: contains the drug «folia jaborandi» withdiaphoretic, 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, Volkart148 (LPAG); 8-XI-1970, Dimitri s.n. (LPAG 4805).Fruits: psychotropic and toxic (2, 35). Leaves: usedby ants of genus Atta (17).
Quillaja brasiliensis (A. St.-Hil. & Tul.) Mart.(Rosaceae). Misiones, Candelaria-Loreto, 19-II-1951, Montes 12343 (LP); Candelaria, Santa Ana,10-I-1946, Montes 1746 (LP). Cortex: astringent(2); used as vegetable soap to wash hair andclothes (36).
Rauvolfia sellowii Müll. Arg. (Apocynaceae).Misiones, Eldorado, 29-III-2005, Martínez 60(LPAG). Cortex: hypotensor (2, 37). Cortex: withbitter taste and antimalarial properties (3).
Rheedia brasiliensis (Mart.) Planch. & Triana(Clusiaceae). Misiones, Parque Nac. Iguazú, 17-XI-1969, Hualde 16 (LPAG); Pozo Azul, 28-III-2005, Martínez 62 (LPAG). Cortex: medicinal (2).Fuits: edible. Seeds: to cure mange (5).
Ruprechtia salicifolia (Cham. & Schltdl.) C.A. Mey.(Polygonaceae). Corrientes, Perugorría, 16-XII-1948, Cabrera 10569 (LP). Misiones, Eldorado,22-III-2005, Martínez 40 (LPAG). Cortex: containsa bitter tonic (2).
Schefflera morototoni (Aubl.) Decne. & Planch.(Araliaceae). Buenos Aires, La Plata, calle 2 e/51 y53, 15-XI-2005, Martínez 97 (LPAG). Misiones,Eldorado, 24-III-2005, Martínez 69 (LPAG). Totreat 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, pruriginoussymptoms (1).
Bol. Soc. Argent. Bot. 43 (1-2) 2008
58
Sebastiania brasiliensis Spreng. (Euphorbiaceae).Buenos Aires. La Plata, FCAF, 19-IV-2004, Bayón639 (LPAG). Misiones. Eldorado, 22-III-2005,Martínez 34 (LPAG). Latex: is used externally tocure 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); PozoAzul, 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-IX-1971, Dimitri 11259 (LPAG). Cortex: produces anaromatic resin to elaborate pomade used incutaneous treatments (5). This species belongsthe same genus that some Asiatic species fromtheir cortex is extracted a whitish resin giving anaromatic essence known as «benjui» used inmedicine and perfumery (20).
Tabebuia heptaphylla (Vell.) Toledo (Bignoniaceae).Buenos Aires, La Plata, FCAF, 15-X-1992, Volkarts.n. (LPAG 2941). Misiones, Eldorado, 23-III-2003,Martínez 41 (LPAG). Wood: antirheumatic (1).Flowers: have properties against cough. Woodand cortex: to treat injure of the skin (3). Cortexand 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 toothdecay (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 andantimenorrhagic properties (2, 42).
Vitex megapotamica (Spreng.) Moldenke(Verbenaceae). Buenos Aires, La Plata, III-1984,Volkart s.n. (LPAG); FCAF, III-2006, Arambarri257 (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:
Mandrile, 2005; 10: Dimitri, 1978; 11: Rotman, 1987; 12:Chifa & Ricciardi, 2001; 13: Alonso & Desmarchelier, 2005;14: Filipov, 1994; 15: Martínez Crovetto, 1965; 16: Marinhoet 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: MateuAmengual, 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.
ALONSO, J & C. DESMARCHELIER. 2005. Plantasmedicinales autóctonas de la Argentina. Bases científicaspara su aplicación en atención primaria de la salud. Ed.L.O.L.A., Buenos Aires.
AMAT, A. G. & M. E. VAJÍA. 1991. Plantas medicinales yetnofarmacología en la provincia de Misiones (Argentina).Acta Farm. Bonaerense 10: 153-159.
AMAT, A. G. & M. E. YAJÍA. 1998. Plantas vascularesutilizadas en la fitoterapia de la provincia de Misiones(Argentina). Farmacobotánica y farmacognosia enArgentina. 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 fromGallery forests of the paranaense province (Argentina).Part 1. Bol. Soc. Argent. Bot. 41: 233-268.
ARIAS, M. E. & M. F. PARRADO. 1996. Estudiomorfoanatómico de hoja y fruto de Psychotriacarthagenensis Jacq. (Rubiaceae). Parodiana 9: 19-24.
BARBOZA, G. E., J. J. CANTERO, C. O. NÚÑEZ & L. ARIZAESPINAR (eds). 2006. Flora medicinal de la provinciade Córdoba (Argentina). Museo Botánico de Córdoba.Gráficamente ediciones, Córdoba. Argentina.
BERNARDI, L. 1985. Contribución a la dendrologíaparaguaya. II. Boissiera 37: 1-123. EdicionesConservatoire et jardin botaniques de la Ville de Genéve.
BILONI, J. S. 1990. Árboles autóctonos argentinos.
We appreciate the fine technical assistance ofSantiago M. Martínez in collecting material. Thecurators of herbaria (ERA, LP, LPAG, SI) for loaningthe specimens, María A. Migoya for inking our penciloriginal illustrations, and anonymous reviewers forsuggestions are acknowledged. Support wasprovided by the Comisión de Incentivos a losdocentes-investigadores, Decreto 2427/93 (Secretaríade Política Universitaria, Ministerio de Educación dela Nación Argentina).
Acknowledgements
Bibliography
59
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. Investigaciónexperimental preliminar en especies vegetales para aislarsustancias reductoras del colesterol y demás lípidossanguíneos. 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. ColecciónCientífica 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 deAmérica Latina. Programa regional de desarrollo científicoy tecnológico. OEA, Washington, D.C.
CHIFA, C. & A. I. A. RICCIARDI. 2001. Plantas de uso enmedicina 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 lafamilia Myrtaceae. Acta Farm. Bonaerense 15: 109-123.
CRISTIANI, L. Q. & J. L. AMORÍN. 1972. Estudio preliminardel Sarandí blanco Phyllanthus sellowianus Mueller Arg.(Euphorbiaceae). Farmacobotánica – publicación técnicaNº6. Págs. 11.
DIMITRI, M. J. 1978. Enciclopedia argentina de agriculturay jardinería. 3ra. edición. Acme S.A.C.I., Buenos Aires 1:1-644.
DOMÍNGUEZ, J. A. 1928. Contribución a la materia médicaargentina. Trabajo del Instituto de Botánica y Farmaciade 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. DISTASI. 2002. Especies forestales nativas de la selvaparanaense. 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 centralChaco. 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 ofMissouri Botanical Garden 89: 524-550.
GONZÁLEZ, M., A. LOMBARDO & A. J. VALLARINO.1939. Plantas de la medicina vulgar de Uruguay. TalleresGráficos S.A. Cerrito 580, Uruguay.
HIERONYMUS, J. 1882. Plantae diaphoricas floraeargentinae. Bol. Acad. Cienc. Córdoba 4: 199-598.
KATINAS, L. 1996. Revisión de las especies sudamericanasdel género Trixis (Asteraceae, Mutisieae). Darwiniana 34:27-108.
LAHITTE, H. B. & J. A. HURRELL. 1994. Los árboles de laIsla Martín García. Comisión de Investigaciones
Científicas (CIC). Buenos Aires.
LAHITTE, H. B. & J. A. HURRELL. 1999. Árbolesrioplatenses. Ed. L.O.L.A., Buenos Aires.
LEONARDI, D., O. DI SAPIO, M. GATTUSO & S.GATTUSO. 2002. Caracteres morfoanatómicos de lacorteza y hojas de Tabebuia impetiginosa y T. heptaphylla(Bignoniaceae). Bol. Soc. Argent. Bot. 37: 51-61.
LEONARDIS, R. F. J. 1975. Libro del árbol. Esenciasforestales 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 delParaguay. Cuerpo de Paz, Colección e Intercambio deInformación, 2da. Edición. Editorial Gráfica MercurioS.A., Paraguay.
LORCA, G. G., A. G. AMAT & C. GONZÁLEZ. 1995. Análisiscomparativo de caracteres diagnósticos para laidentificación de tres especies argentinas de Myrtaceaeempleadas en la medicina popular. Acta Farm. Bonaerense14: 81-86.
MANDRILE, E. L. 2005. Plantas medicinales que sedispensan en Argentina. Tomo 1. (2da edición). Colegiode 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: 54ºCongreso Nacional de Botanica, Belém, Brasil.
MARTÍNEZ CROVETTO, R. 1963. Esquema fitogeográficode 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 Tobasdel este del Chaco. Bonplandia 1: 279-333.
MARTÍNEZ CROVETTO, R. 1965. Estudios etnobotánicos.II. Nombres de plantas y su utilidad, según los indios Vilelasdel Chaco. Bonplandia 2: 1-23.
MARTÍNEZ CROVETTO, R. 1981. Las plantas utilizadas enmedicina popular en el noroeste de Corrientes. FundaciónMiguel Lillo. Miscelánea 69: 69-70.
MARTINS, A. G., S. S. GUTERRES & G. GONZÁLEZORTEGA. 2003. Anti-ulcer activity of spray-died powdersprepared from leaf extracts of Maytenus ilicifolia Martiusex Reiss. Acta Farm. Bonaerense 22: 39-44.
MARZOCCA, A. 1997. Vademécum de malezas medicinalesde la argentina, indígenas y exóticas. Orientación GráficaEditora SRL.
MASTROBERTI, A. A. & J. E. A. MARIATH. 2003. Leafanatomy of Araucaria angustifolia (Bertol.) Kuntze(Araucariaceae). Rev. Bras. Bot. 26: 343-353.
MATEU AMENGUAL, B. M. 1974. Catálogo bibliográficofitoquímico Argentino III. Fundación Miguel Lillo,Miscelánea 53, Tucumán.
PEREA, M., G. PEDRAZA & J. LUCEROS. 2007.Relevamiento de flora arbórea autóctona en la provinciade Catamarca. Consejo Federal de Inversiones, Ciudadautónoma de Buenos Aires y Gobierno de la provincia deCatamarca.
Bol. Soc. Argent. Bot. 43 (1-2) 2008
60
RATERA, E. & M. RATERA. 1980. Plantas de la floraargentina empleadas en medicina popular. Ed.Hemisferio Sur, Buenos Aires.
RODRÍGUEZ, J. M. & C. A. O’DONELL. 1943. Plantasmedicinales 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 reconocidaspropiedades medicinales o tóxicas. Base de Datos,CYTED-OEA.
ROTH, I. 1984. Stratification of tropical forests as seen inleaf structure. Junk Publ., The Hague, Boston Lancaster.
ROTMAN, A. 1978. Moraceae. En: A. BURKART (ed.), Florailustrada de Entre Ríos. Vol. 6: 27-28. Colección CientíficaINTA, Buenos Aires.
ROTMAN, A. 1987. Cecropiaceae. En: A. BURKART (ed.),Flora ilustrada de Entre Ríos. Vol. 6: 31.ColecciónCientífica INTA, Buenos Aires.
ROTMAN, A. 1995. Las especies argentinas del géneroEugenia (Myrtaceae). Bol. Soc. Argent. Bot. 31: 69-93.
SAWCHUK, B. & H. GALEANO. 1981. Descripción deárboles medicinales de la Selva Misionera. En: Manualpara la agricultura en la provincia de Misiones, RepúblicaArgentina. Parte 6: 1-168. (B. Sawchuk, E. J. Domínguez& A. A. Piekun), Programa Argentino-Alemán para el
fomento del desarrollo agrícola de la provincia de Misiones.
SORARÚ, S. & A. BANDONI. 1978. Plantas de la medicinapopular argentina. Ed. Albatros, Buenos Aires.
SPEGAZZINI, E. D. 1999. Análisis foliar por micrografíaanalítica cuali-cuantitativa, de los adulterantescogenéricos argentinos de Ilex paraguariensis St. Hil. var.paraguariensis –Aquifoliaceae- nv. «yerba mate». TesisDoctoral, Facultad de Ciencias Exactas, UniversidadNacional de La Plata.
SPEGAZZINI, E. D., M. T. CASTRO, J. ROSSI, S. M.CARPANO & M. T. NÁJERA. 2002. Micrografía foliarcuali-cuantitativa de las especies de Pilocarpus (Rutaceae):P. jaborandi Holmes, P. microphyllus Stape exWardleworth y P. pennatifolius Lem. Dominguezia 18:5-16.
TOURSARKISSIAN, A. 1980. Plantas medicinales de laArgentina. 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 compuestasargentinas 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 de2008.