A m arambarri, s e freire, m n colares, n d bayón, m c novoa, c monti & s a stenglein leaf anatomy o

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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

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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.

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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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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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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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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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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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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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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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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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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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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.


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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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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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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 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.


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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, 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


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.


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


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.


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;


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

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55


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

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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


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).


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

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Recibido el 28 de Agosto de 2007, aceptado el 12 de Mayo de2008.

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